Improvements in the use of Plant Identification Keys are achieved when the principles of Apical Taxonomy are Applied.

Improvements in the use of Plant Identification Keys are achieved when the principles of Apical Taxonomy are Applied.
Running Title: Apical Taxonomy
Author: Susan Dunlap

ACKNOWLEDGMENTS
    The author extends many thanks to the following individuals for giving their guidance and support along the way: Cleo Condoravdi PhD, David Dreyfuss PhD, Dr. Lewis J. Feldman, Brian Kemble, R. Paul Kiparsky PhD, Peter Kline, Dr. Manuela Kogon, Adina Magill, Tim Metcalf, Peter Midford PhD, Ernesto Sandoval, Martin Schneider, and Richard J. Waldinger PhD.

ABSTRACT
    This article presents research that explores the advantages of plant identification resources that concentrate on branch apices. A recent discovery, that the overhead view of branch apices proves to be especially relevant for distinguishing one species from another, provides impetus for this study. Studies of several genera from 205 plant families were conducted; the results demonstrate benefits gained by integrating this discovery into plant keys and constructing keys that include and concentrate on vegetative attributes, many that are contained within the plant’s apices. A model key for ribbed cacti was prepared that when compared to an existing key demonstrated dramatic improvement in its use. Five aspects of apical taxonomy support its broad application: 1) the presence of a suite of vegetative traits unique to the branch apex, 2) the efficiency achieved by directing individuals to attend to the plant’s apex for identification purposes, 3) the efficiency achieved in the use of keys that incorporate apical images and vegetative traits into them, 4) the efficiency achieved by the design and use of vegetative keys, and 5) improved accessibility provided to a diverse group of biologists who need plant identification resources including field workers, conservationists, bioinformatic specialists, horticulturalists, and taxonomists. The study includes discussions of strategies for applying apical taxonomy to a broad range of taxa and how to develop identification resources that make use of the discovery. The results confirm that apical taxonomy warrants further development, as it provides viable solutions for many difficulties associated with current identification and classification resources.
Key words: plant morphology; vegetative keys; character analysis; identification key; taxonomy; apical taxonomy

INTRODUCTION     
This article presents research that explores the advantages of plant identification resources that concentrate on branch apices and reference apical images. A 2001 discovery provided the impetus for this research: that a species-specific bundle of vegetative traits is clustered at plant apices and visible in a top-down view of the growing tip. The results demonstrate that the overhead apical view of the branch tip is especially relevant for distinguishing one species from another. Research shows that the plant’s apex contains a diagnostic and taxa-specific concentration of characters and that there is a taxonomically useful correlation between them. One goal of Dunlap’s eighteen-year study was to determine the range of taxa to which apical taxonomy applies. The study included a review of over 27,000 novel images that were assembled of nearly 6,000 species. It also included the construction of model vegetative keys unfettered from the constraints of plant hierarchies (heretofore referred to as vegetative keys); these keys represented genera from over 200 plant families and referenced both customary and novel characters found within the plant apices. Further, these keys were accessed for the benefits derived from their use most especially when compared to the use of a typical hierarchical key that originates from the top of a family or plant group. A novel vegetative key for all ribbed cacti was prepared that demonstrated dramatic improvement in its use. A review of an additional 200 plant families is warranted to further establish the benefits attained by the practice of apical taxonomy.
    Five aspects of apical taxonomy support its broad application: 1) the presence of a suite of vegetative traits unique to the branch apex, 2) the efficiency achieved by directing individuals to attend to and monitor the plant’s apex for identification purposes, 3) the efficiency achieved in the use of keys which incorporate apical images and vegetative traits into them, 4) the efficiency achieved by the design and use of vegetative keys, 5) and benefits and improved accessibility provided to those concerned about conservation, genetics, and the plant needs of pollinators and wildlife. The results of this study support the argument that apical taxonomy warrants further development, as it provides users with an efficient plant identification system that resolves many of the difficulties associated with the use of current identification and classification resources, such as those encountered in the use dichotomous keys and the insertion of ephemeral traits and technical terms within them. Demonstrating the benefits of apical taxonomy is the primary contribution of this work.
    The common starting point for those working to devise modern plant identification tools is traditional plant identification resources and keys. The armature of new tools, such as Delta, Lucid, and plantsystematics, was built upon plant traits extracted from historical resources and traditional keys. Many of those working in this area devised methods to limit the use of technical terminology such as those found in traditional dichotomous and hierarchical keys (Kirchoff (2011, 2014) and Waldchen 2018). Kirchoff argued that the steep learning curve required to master use of technical terms contributed to a shortage of skilled plant identification experts. The greater the technical burden associated with a plant identification system, the fewer number of people who will be able to use it. Several modern plant identification resources augment the plant keys with visual aids; doing so minimizes the need for a user to memorize technical terms and aligns more directly with the visual attitude and expertise of many users. Waldchen argued that the skills of individuals from diverse backgrounds needs to be considered, advocating that knowledge from individuals with visual expertise be integrated into plant identification resources. In 2008 Kirchoff states that “keys are not based on pattern recognition, the forte of visual experts (and) would be more effective if they were visually based.” While the use of visual aids has clear benefits, the manner in which these aids are used varies; inserting images into a key to illustrate traits used within it and to compress it are discussed. The need to normalize the definition and use of trait and nomenclature terminology continues to this day; a clear path to resolving the problems that emerge from this need are being tackled by Kilian, Balduzzi, and others (see Discussion below). Additional challenges to ease the use of plant keys remain and are addressed below.
    A survey of numerous plant-related publications (see Supplement Reference), concentrating on print books available to gardeners and horticulturalists, revealed that four main weaknesses hinder their use as aids in identifying an unknown specimen. First, while cultivation techniques and landscaping features are often fully described, identification attributes and plant keys are generally absent. Second, the images used did not clearly illustrate the unique physical trait(s) of a species. Third, the images fail to empower the reader to distinguish one taxa from another. Rather, images might reveal one attribute of a taxa, such its shape at maturity, and another attribute of a second taxa, such as a close-up of its flower. Fourth, when keys are present, progressing through them is difficult: the user is required to have either advanced knowledge of the name of a plant or a working knowledge of plant parts and attributes – including flower parts, roots, seeds, pollen, non-floral organs, structural elements, surface features, vegetative features, and internal processes. The difficulties encountered in the use of the key are caused by the use of technical terms, the dependence upon numerical sub-categories and descriptors, and the co-mingling of ephemeral attributes within it. For example, when seeking to understand the definition for “gymnosperm”, someone new to the study of botany sought the definition of 85 additional words (such as conifer, imbricated, and valvate) to fully understand the original term. While inserting such traits may in some instances be required to advance in a key, their ubiquitous use places burdens on the user. The user will encounter uncertainties while attempting to navigate such a key, as they may have limited knowledge of a plants’ internal processes or, if working with a vegetative sample, may lack the means to acquire additional information about a plant that is now needed to advance in the key, such as details about the roots or microscopic characteristics of either the flower or pollen grains. The results of this current study demonstrate that apical taxonomy enriches the volume of vegetative data that can be used in keys and by doing so significantly diminishes the need for them to be populated with technical terms and ephemeral traits.
    Two systems of resources aid those seeking to identify a plant or learn about its contribution to local wildlife. One system is oriented toward those with scientific interests while the other is oriented toward those with cultivation interests. These resources include local and online nurseries, specialized societies, botanical gardens, universities, and herbaria. Plant identification forums are helpful, such as that managed by the UBC Botanical Garden, as are regionally specific horticulture and interest groups. Local nursery personnel provide information as well, especially if the plant in question is cultivated. At a professional level, a physical sample or a clear visual representation of the specimen is preferred over a verbal description. At any level, other than expert to expert, the exchange of a sample of the plant in question is required. For instance, to identify a plant, international universities require a physical sample of a plant as images alone do not suffice. Individuals who need guidance on how to care for and select plants are impacted by the differences between these resources, primarily caused by the depleted resources available to gardeners who need information about plants that are no longer in cultivation. The difficulties encountered impact the care, use, and selection of plants.
    That a novel and diagnostic cluster of vegetative traits exists at the branch apex of plant species was described and submitted to the USPTO by Dunlap (2005, 2012) (S1); formal approval of this submission was obtained in 2012and 2013. Research to explore the utility of the discovery was conducted by Dunlap between 2001-2018 and is referenced below.  The research includes the development of model plant keys and describes the benefits derived from an apically-biased identification resource. The proposed apical system can be used to satisfy a broad range of plant identification needs. The research conducted has shown that the practice of apical identification is at a state that demonstrates it can be widely applied. That it is important to do so will be argued throughout this article.
    It is beyond the scope of this article to prove how apical taxonomy will apply to all 405 plant families, as additional work needs to be done to isolate plant characters for 205 families; however, the article explores potentially beneficial methods to overcome obstacles that may be encountered by those who review additional families.

METHODS: INITIAL COMPILATION OF THE DATA
    The apical complex of a plant is defined as “the novel cluster of traits contained in an overhead view of a plant’s growing tip”. (Dunlap 2005) Both dormant and active growth observed at the growing tips of plants was documented and reviewed as part of this study. Initially, succulent species were the focus of the study and apical images of them collected. As the benefit derived by implementing apical taxonomy upheld, the research expanded to include cultivated woody and herbaceous taxa. When feasible, apical images of several individuals were gathered, especially for those genera studied in depth, including the Euphorbia (L.), other succulents, cultivated taxa, and Cactaceae. Environmental factors affecting a plant’s growth were noted (such as drought and root stress). (S4) Excel databases for several plant groups were created, each one comprised of customary vegetative traits such as those found in many plant keys, including leaf arrangement and shape. The entries were then supplemented with data about unconventional vegetative traits found in the apical images, such as the geometric shape of the ‘empty’ cone formed by the apical leaf cluster, unique leaf arrangements (such as was found in Rhododendron (L.) discussed below), distinct leaf pigments, and descriptive relationships between various leaf components. The data, assembled for several plant groups and families, was sorted to isolate and identify taxonomically productive vegetative traits and to construct model keys making use of them; most of these traits are found at the plant apex. Details about the methods used for each plant group is described below.  
    Over 7,000 novel apical images of 2000 succulent species were gathered and examined during the early stages of the study (plant sources are mentioned below). Excel databases were compiled for several taxa; customary and unconventional vegetative traits were recorded for each one. Entries in the databases were monitored to access when a sufficient variety of traits had been recorded to enable one species to be isolated from its nearest relative. If vegetative traits were insufficient to divide a few taxa, the apical images of those few taxa were examined in tandem in an effort to find distinctive attributes. Such efforts were instrumental to help determine how to achieve the maximum benefits of apical taxonomy and monitor when those benefits had been realized.
    During the second stage of the study, the goal shifted from plant identification to plant selection (by emphasizing cultivated plants). At that time, 2001-2010, the majority of resources available for study that provided access to both text and live plants, were resources that applied to cultivated plants. These resources included those in print, those in the horticultural supply chain (from grower to retailer), and knowledge available from horticultural groups. Cultivation data for 10,000 taxa was compiled and, when feasible, novel images for them were gathered. (The database was subsequently reduced to the 3641 taxa for which images had been acquired.) Over 20,000 images were assembled (and digitized as needed); the images were gathered during field visits to over 60 regional growers and nurseries. The grower’s species and cultivar nomenclature for each taxon was reviewed and work done to update the nomenclature to standards found in the RHS Plant Finder; theplantlist currently serves as the primary source to verify nomenclature and plant family structure. Offline print resources used to compile the customary trait data for these plants included European Garden Flora, Volumes I-VI, Index of Garden Plants (IGP), and Hortus Third. Excel databases of customary traits were constructed for these plants, and gaps in the data addressed by examining live plants, the apical images, or online resources. The data entries were compiled in a manner that supported the execution of a multi-access key – which provides the user continuous access to each entry in the key. When multiple qualities were found – e.g. taxon containing both blunt and rounded leaf tips – both qualities were noted in the database. Quantity and dimension descriptors for traits was avoided so as to ascertain whether it was feasible to develop a multi-access identification tool wholly comprised of descriptive traits. However, quantitative data was used in limited ways: quantities used for identification purposes were avoided –  such as the number of pistils found in a flower – while those desired in horticultural settings were used but not for identification purposes – such as the mature height and width range of a plant. Dimensions particularly useful for cultivation were recorded. Model plant selection software was developed for 3641 taxa that presented a user with both data and images of each plant. (See Figure 1 below.) It contained data for over 200 plant families, including woody and herbaceous taxa, several varieties and cultivars, and numerous plant types including 300 annuals, 81 biennials, 48 ferns, 83 grasses, 23 palms, 1262 perennials, 1195 shrubs, 107 succulents, 410 trees, and 127 vines. The 2018 iteration of this selection software has been enhanced by the addition of nectar-plant data that applies to all US butterflies. A sample of the selection software can be found at aerulean.com.

    Succulent Plants
    Live succulent plants, including Cactaceae, were widely available for use in this study. Three examinations of succulents were conducted to develop proofs of the benefits to be gained by the use of apical taxonomy. These individual studies, described below, applied to succulent rosettes, Euphorbia, and Cactaceae.
    A database for cultivated succulent rosettes was developed to illustrate the basic utility of a visually-enhanced apical key. Prepared in 2001, this new apical key contained all 62 succulent rosettes listed in The American Horticultural Society Encyclopedia of Garden Plants.(S2) The user progresses in the key by selecting from a list of traits that isolate one plant type from another. Each selection made decreases the remaining traits and leads to a set of apical images. The key includes a variety of succulents including species with rosette-form vegetation and species with spiral leaf formation. This model vegetative key was successfully applied to a small group of plants and provided the impetus to continue the study. Further work is required to develop a tool that applies to all succulent rosettes found in the plant kingdom.
    The Euphorbia were studied in depth to establish that the benefits of the discovery applied to a large genus. The goal was to determine if it was possible to distinguish one taxa from another when focusing exclusively on vegetative traits. For several reasons, Euphorbia were selected for study: they are widely grown and collected, have been studied over time, and there are many publications about them. The Euphorbia were gathered into 35 groups, of which approximately 600 species are cultivated. Over 2200 images for 494 of these cultivated species were collected, digitized, and studied; multiple apical images of several individuals were collected during field visits to collectors and growers and so represented taxa grown in a variety of conditions. Apical images of several taxa are shown here. (S3) The expressions of specific character traits were noted, especially those affected by the age of the plant and its growing conditions. Pot-bound individuals contained instructive information about the expression of a trait – e.g., the position of the flowering eye remained constant while some traits were supple but reduced in size or atrophied on an individual grown in stressful conditions. Duplicate apical images of forty-three taxa were compared in depth and were found to be visually comparable, for example, new and mature apical images of a taxa contained a similar arrangement of traits but often dissimilar pigmentation. An Excel database for 494 Euphorbia species were created; the presence or absence of 25 vegetative traits was recorded for each taxon. (S4) The database, used in concert with apical images, was used to create sample plant identification software that could be used to satisfy an identification query. A positive selection of one trait, chosen at will, narrowed the results to an increasingly smaller subset of taxa – each selection made reduced the remaining traits and reduced the remaining set of apical images. As is customary with some multi-access keys, the user was provided continuous access to the list of 25 traits and was empowered to de-select a previous selection. Users were also given continuous access to an ever-decreasing display of apical images; this access empowered them to select and identify taxa from the remaining set of apical images and so reduce the need for them to navigate a more elaborate key. This feature is one of many that empowers those with broad familiarity to the plant kingdom and those with visual skills to advance an identification query efficiently. The study expanded to other plant groups as the study of Euphorbia upheld, and established that a diagnostic set of salient macro character trait(s) remained stable and visible. (S3, S4)
    Herbaceous and Woody Plants
    Apical images of cultivated plants were collected, digitized, and electronically organized. A subset of these plants was isolated for further study in order to access the feasibility of applying the methodology used to develop the Euphorbia software to a group of non-succulent herbaceous and woody plants. Most of the apical images were collected while taking field trips to regional growers, nurseries and botanical gardens. Sample keys were prepared for those plants known to be notoriously difficult to identify such as Arctostaphylos (Adanson), Rhododendron (L.), and Salvia (L.). See Figure 2 apical images. As with the studies of succulents, these databases were designed to facilitate identification by enabling a user to reduce a query to a subset of species and to empower them to narrow the results to a single plant. They were primarily comprised of traits that can be observe in an apical view of the branch tip.
    Apical images of these plants illustrated several unique attributes exhibited by the petiole including its visibility, pigmentation, scale, texture, attachment features at both ends, and angle off the stem. The images also captured vegetative details that apply to venation, adaxial pigmentation (e.g. contrast between leaf surface, margin, and veins), texture, features of the leaf margin (e.g. pigmented, rolled, flared, spiny, scalloped, hairy), the shape of the apical leaf cavity, and numerous, transient distinctions expressed by emergent and secondary leaf clusters including fundamental changes to their pigmentation, texture, scale, and shape. While long term studies will be required, it was found that multiple traits of each plant retained distinguishing attributes over the lifespan of the plant; when feasible both emergent and mature apices were recorded and reviewed for each plant.
    A sample of the Rhododendron key is shown on Table 1. Numerous diagnostic attributes were found in an apical view, including the petiole details itemized above. The apical view of the leaf cavity exposed basic attributes that included whether the ‘empty’ cavity formed a tight cone, quickly flattened, or quickly inverted. Further, apical views of several Rhododendron exposed an unusual, distinguishing characteristic: many taxon form an incomplete spiral. The results of this study are described below. (See Figure 3.)
    Cultivated Plants
    During the next phase of the study, apical taxonomy was applied to cultivated plants. A resource was prepared that could be used by anyone who needs basic information about growing cultivated plants. A database was compiled of those attributes particularly useful to determine a plant’s suitability to meet a variety of horticultural needs. These plant attributes include those of concern to horticulturalists and property owners, such as the likely dimensions of a mature plant and temperature and watering requirements. Over 20,000 novel images of cultivated taxa were collected during visits to the aforementioned California growers, nurseries, botanical gardens, and private collectors. These images represented a sampling of plants from over 200 families. Selection software was created for 3641 cultivated taxa for which images had been gathered. It enabled users to select from 392 attributes, with 2,000 discrete selections within those primary criteria. See (S5) for a sample of the attributes used in the introductory software. Many of these criteria were derived by examining apical images and extracting data from them. The data-extraction effort facilitated an expansion of the selection fields by correcting inconsistencies and omissions contained in the digitized text data referenced above (see The Index of Garden Plants). Working with large format and macro-lens image files facilitated the compilation and assignment of trait-data. A 2007 USDA-sponsored survey of master gardeners was conducted that showed a broad interest in this tool. Subsequently, it has been modified to also support the addition of pollinator data. Expanding the database to include a broader range of cultivated plants is warranted. The current model applies predominantly to Zone 9 plants that can also be grown in other zones. It would be beneficial to expand the database to include cultivated plants that grow in Zones 1-8 and 10-12.

    Additional Taxa
    Studies of Poaceae and large single-stemmed taxa were conducted to assess some of the challenges that may be encountered when apical taxonomy is applied to a broader range of plants. Poaceae were selected because, as became apparent during the review of apical images of cultivated grasses, distinguishing between grasses was found to be particularly challenging. Large single-stemmed taxa were reviewed because of the difficulties that would be encountered when attempting to gather apical images in the field. It is anticipated that in some instances either that gathering an apical image is too challenging, as is the case with tall, single-stemmed taxa such as some Arecaceae (Bercht & J. Presl) and Agave (L.). Diagnostic attributes were found for these plants while pursuing development of an apically biased resource. For example, reviewing the definitive key for weedy grasses of California revealed that the shape of the culm is diagnostic. A review of Agave reveals that the apical view of the terminal spine of each leaf is diagnostic for each taxon. A review of North American tree-form Arecaceae, including members of the Butia ((Becc.) Becc.), Sabal (Adans.), Brahea (Mart.ex Endl.), Livistona (R.Br.), Washingtonia (H.Wendl.), and Phoenix (L.) genera, reveals that attributes of the hastula proves to be diagnostic. It is encouraging that these attributes are generally visible from the ground. More work is needed to record the apical variety for the remaining half of the plant kingdom. The studies of these genera illustrate methods that can be applied to others. While apical images of these additional taxa may not clearly isolate them from a close relative, that there are a sufficient variety of vegetative attributes to develop a vegetative key has been successfully argued below.
    
Cactaceae
    The results of a comprehensive study of a single plant family, Cactaceae, was conducted to illustrate the benefits that can be achieved when the principles of apical taxonomy are used in the development of a key. A custom key for twenty-five percent of Cactaceae was prepared and then compared directly to an existing key; that study and its results are described below.
    Prior to initiating a study of the cacti family, a written description of visual data contained in the apical complex was prepared for Ferocactus recurvus ((Mill.) Borg.). That much has been written about F. recurvus allowed a direct comparison between historical descriptions of the plant and a written description that documented the breadth of vegetative data contained in an apical image of a single individual. Many of the attributes contained in this description had yet to be considered in the development of a plant key. The breadth of traits found and their taxonomic distinctions were then considered in the construction of a database so that their contribution to a cacti identification resource could be more fully considered. (S6) See Figure 4 Ferocactus (Britton & Rose) apical images.
    A comprehensive review of the cactus family was conducted to allow a direct comparison between an apical systematic key and a traditional one. Original apical images of 1800 cultivated cacti (S7) (several images shown here) were collected from live plants during numerous visits to growers, plant retailers, botanical gardens, and private individuals located in California and Arizona. Images of several individuals of taxa were gathered when feasible. An Excel database for 934 of these Cactaceae was prepared; both customary traits, as described in The Cactus Family (Anderson), and unconventional traits were documented, such as whether the spines cross at apex or whether the rib is grooved or un-grooved at the apex. The presence or absence of 76 vegetative traits were recorded. See Table 2 for a sample of these traits. Collecting and reviewing the apical images provided exposure to the expression and form of traits common to the family; the exposure was used productively in the development of the database. The traits compiled were selected for the subtle variability expressed by them that was observed in the apical images. The 76 traits divided the 934 taxa into manageable subsets, while the variability expressed by both these primary 76 traits and the remaining attributes contributed to the overall utility of the key by presenting apical images of taxa that contained a novel, diagnostic assembly of these attributes. The goal to reduce a query to one taxon dictated both the choices made and the subsequent addition of traits.
    A subset of the all Cactaceae was then isolated; ribbed cacti were chosen as a representative sample of the family, an attribute ascribed in the literature to 463 taxa, or twenty-five percent of the family (as in (IGP), (Anderson), and The New Cactus Lexicon (Lexicon)). This subset was chosen because several apical images of them were available to consider for this study and because this attribute had been reliably reported in the literature. The master 76-trait database was then methodically reduced to determine the most efficient subset of traits that would lead to identification of ribbed cacti. The aforementioned 76 traits were reduced to a subset of 23 omnipresent macro vegetative traits that applied to the 463 ribbed taxa. All data gaps were resolved. An ordered sort, such as that shown on Table 3, was made by selecting a trait that applied to the greatest number of taxa. See Table 4 for a sample of an ordered trait selection thread. Populating the key with an overabundance of traits provided guidance on which of them proved to be the most productive with respect to identification. The most productive and stable omnipresent traits were selected for this key. The results are described below.

RESULTS
    The general results that apply to all the plant groups included in this study are provided here (results for individual plant groups are inserted below). It was found that the tools developed mitigated the problems associated with defining botanical terms by illustrating those terms with images and demonstrated that it is feasible to construct vegetative keys for a broad range taxa. The plant’s apex proved to be especially diagnostic: the attributes and features of the stem can typically be observed during the entire growing season, the top-down view of the branch apex contains a suite of traits that are observable in one view or image of the plant, and the apical region presents itself with more reliability, as it is less influenced by the twisting effects of growth and other environmental effects. Efficiencies to apical taxonomy are gained by empowering users to select taxa from a decreasing set of apical images and by focusing their attention on the plant’s growing tip in natural settings. Additional benefits support the broad application of apical taxonomy: 1) the presence of a suite of vegetative traits unique to the branch apex, 2) the ease of acquiring identification of a plant by directing individuals to specific and generally omnipresent attributes, 3) the efficiency achieved by incorporating apical images and vegetative traits into plant keys, 4) reducing the number of traits needed to navigate a key, 5) avoiding, if not altogether omitting, the use of ephemeral traits (thus removing the uncertainty encountered by attempts to navigate keys that contain them), and 6) those provided to individuals who attend to the conservation and plant needs of wildlife (discussed below). These improvements alleviate many problems affecting the usability of plant keys that have been advocated by others (see Kirsch’s discussion above in the Introduction and Balduzzi’s work with mathematics cited below).
    A specialist on Euphorbia, while using the 25-trait key, identified an unknown sample by selecting as few as 6 traits, not even needing to refer to the remaining nineteen. Narrowing to a subset of images was sufficient to enable identification of taxa; allowing an individual to select from the remaining assortment of apical images replaced the need for a more elaborate key. Initially more than twenty-five traits were tracked in the database, as the final and most productive set was unknown in advance of its construction. The 25-trait key that applied to 494 taxa will need to be expanded as more Euphorbia are added.
    The study of the herbaceous and woody plant group exposed two attributes that contributed to the development of apical taxonomy and can be usefully inserted in plant keys. One pertained to the geometric cavity transcribed by the apical leaf cluster (see the Discussion for a complete description of the benefits derived by attending to this attribute). Another attribute, found in Rhododendron pertains to the incomplete spiral leaf arrangement, which provides an example of previously unrecognized leaf arrangement that is easily perceived in an apical view of the growing tip. This leaf arrangement, if used in a key, would quickly reduce a data-set to very few taxa. (See Figure 3) Fralish & Franklin (2002) described the leaf arrangement of Rhododendron as: “alternate; leaves usually clustered at twig tip.” The twig tip of the plant is indeed diagnostic, and provides even more diagnostic evidence that applies to Rhododendron leaf arrangement when one attends to the apical view. That there are numerous diagnostic attributes contained in the apical view of herbaceous and woody plants contributed to the pursuit of this study. It can be expected that many more such attributes will be found.
    The study of cultivated plants unearthed deficiencies in the literature and exposed a primary benefit found in the use of the software (some are contained in the Discussion section below). One deficiency found in the both the taxonomic and horticultural literature was that trait voids were present; many vegetative attributes remain unassigned to each plant to which the trait applies. It is possible a particular trait has not been established for a taxon but it is also possible that its potential contribution to taxonomy or horticultural has yet to be appreciated or valued. One benefit of the cultivation software was realized: that it is possible to blur the line between selection and identification tools. The initial approach, used in the construction of the woody & herbaceous, Euphorbia, and Cactaceae databases, was to retain a separate list of criteria for these two tasks, as there are many criteria important to only one of them. Identification usually has the goal of narrowing the list of candidate plants to a single species. Selection of a plant often results in a list of alternatives or a list of complementary plants, from which further selection can only be made by reference to information that the user has not so far considered (such as availability, plant characteristics, and other attributes that the user now browses). Methods were explored to merge these goals into a single resource. It had been assumed that plant selection and identification tasks were too distinct to find convergence between them, a convergence in part disguised by the numerous stylistic and content distinctions between plant keys and horticulture literature. Inserting leaf details into the selection tool was motivated by a desire to empower horticulturalists in new ways – for example by enabling them to construct a garden wholly comprised of plants with variegated leaves, with opposite leaves, or leaves of a particular color or shape. That such leaf details also empowered users to identify plants was an additional benefit. The leaf characters that proved to be particularly useful as identification aids included leaf shape, tip, arrangement, pigmentation, and retention. Thus, it was productive to explore how to create a selection tool that also functioned as an identification tool. The prototype selection tool comprised of 3641 taxa, modeled here, was implemented in essentially the same manner as a plant identification tool. Forty categories of vegetative attributes were considered during its construction (See Figure 1 and S5.)
    The studies of Agave, Poaceae and Arecaceae, as discussed above, illustrate techniques one might use to further develop apical taxonomy. Content in images and written descriptions that suggested which trait differentiated one species from another was assembled for these plant groups. Such diagnostic attributes can be used to construct a vegetative key. While the culm may not be visible in an apical view of a grass taxa, its presence or absence in and of itself is diagnostic. A review of existing keys may expose a short list of diagnostic macro vegetative attributes, as it did in studies of these taxa. A review of the literature may expose a biologist’s single observation of an intersection-trait that later proves to be diagnostic for an entire plant family. These intersection vegetative attributes will contribute to the development of apical taxonomy, as will the completion of data entries for known traits and a reasoned reduction of synonyms. The studies conducted reveal that more work needs to be done in this area, most especially when combined with an awareness of the abundant descriptive material contained in apical imagery. Apical images will serve multiple functions as they are a source of new macro vegetative data and their use in keys provides economy in its use. (see Euphorbia (S3, S4) and Ferocactus (S6))
    The study of Cactaceae produced several results. The study of Ferocactus recurvus duplicated the results found elsewhere: the apical complex is a data-rich region of the plant and is full of characteristics not typically translated into plant descriptions or keys. A test case prepared for ribbed cacti, which make up 25% of Cactaceae, demonstrated a fundamental improvement in the use of the key when compared to an existing key. This test case key was comprised of 23 omnipresent traits. The results show that 10 trait sorts were sufficient to reduce 75% of the ribbed taxa to 20 or fewer apical images. (See Table 3) Adding three additional omnipresent vegetative traits reduced the remaining search queries to the 20-image threshold. Populating the key with an overabundance of traits provided guidance on which of them proved to be the most productive with respect to identification. The data regarding those taxa with identical configurations of omnipresent traits were evaluated showing that sorts 1-17, arranged in descending order of occurrence, were extremely productive. For example, of the 438 ribbed taxa, 238 also contained tubercles; of those with tubercles, 122 taxa also contained radiate spines; of those with tubercles and radiate spines, 54 taxa contained a notch or furrow and then 21 of those contained curved spines. By referencing the additional traits in the key, the final 21 taxa in this thread divided into a subset of twenty or fewer taxa. Executing 23 sorts reduces all but four configurations to zero. Tracking 40 vegetative traits may sufficiently reduce the results for all 1816 recognized taxa in the cactus family to the recommended 20-image threshold. Further work is needed to apply apical taxonomy to the 1353 non-ribbed taxa that populate the family.
    The above key was then compared to a published resource to determine their comparative effectiveness and ease of use. The current dichotomous key of the Cactaceae family (Lexicon), which comprises 152 genera and 1816 species, contains 328 steps in the longest path that serves to guide the user to one of these 152 genera, and no lower; most of these steps also contain several numerically defined subcategories, making the actual total impossible to calculate. Identifying to the species level could double or triple those steps to 700 or more. The (Lexicon) key references 47 traits: 18 vegetative traits and 29 ephemeral traits, including flowers, pollen, roots, seeds, and fruit. A user is required to develop a working knowledge of the numerical subcategories in order to advance in the key. Thus, a user is hobbled by attempts to use the key, as the co-mingled references to 29 numerically defined ephemeral traits raises the level of uncertainty in its use to well over 60%. A user can advance to the definition of six taxa without encountering ephemeral traits. Two of these six taxa are members of monotypic genera; the insertion of ephemeral traits hinders access to the remaining 23 monotypic genera. Thus 1810 taxa cannot be isolated or identified without navigating through and encountering a broad assortment of ephemeral traits. It is reasonable to conclude that the sample key of ribbed Cactaceae improved identification by at least an order of magnitude. Actually, as it overcomes the ephemeral-related uncertainty that hobbles the use of the existing key, the proposed key illustrates a mechanism to create an infinitely better one.
    By comparison, tracking an estimate of 40 vegetative traits in the model apical key produces far better results. Removing uncertainty by deleting references to ephemeral traits is a major benefit that apical taxonomy can provide. That 23 traits are sufficient to divide 463 taxa, compared to the above 328-plus, illustrates the broad distinction between the two taxonomic systems. Removing references to ephemeral traits contributes to the improvement. The results confirm that, by integrating apical images and data into identification resources and, significantly, by embracing the discipline of developing a vegetative key, users can be provided with an efficient identification system. Apical taxonomy guides users to the branch apices and navigates them through a key while referencing a carefully selected assortment of vegetative attributes. The apically driven scheme enables the user to tell the software what is known and to narrow the results by selecting from a list of omnipresent traits. Each trait selection independently reduces the dataset and presents an increasingly narrow set of apical images from which the user can choose. Such improvements in the cacti key, that address both its efficiency and uncertainty, provide substantive proof that apical taxonomy warrants further development.

    Future Applications
    Apical taxonomy may prove to be useful to individuals in numerous plant-dependent communities. Its value to those with an interest in cultivation – members of one of the two systems of identification resources described above – can be realized by organizing and presenting the apical data in a manner that caters to their needs and by devising industry-appropriate resources. When using a complete dataset of vegetative traits, it was possible to merge selection and identification tools, as, by adding additional traits, a selection tool could be used to perform numerous identification tasks. Organizing a sub-set of apical images and related identification tools may also provide some benefits, as a narrow set of taxa – such as nectar plants or drought-tolerant shrubs –  may be of particular concern to an interest group or sector. A dedicated resource may provide benefits and advantages to an interest group.
    Implementing the proposed identification system would benefit those in interconnected fields of interest – such as climate scientists and conservationists – and would enhance the ability of ecologists to identify plant pollinator pairs. In a study of plant identification resources applied to the nectar-plant needs of US butterflies, data gaps were found that may also suggest improvements that can occur elsewhere. Bahlai 2016 suggested that crowd-sourced image content can be used “to create lists of plants most probable to attract the desired pollinator taxa.” Bahlai asserts that currently “much of the reasoning for including a plant in (pollinator) lists is based on anecdotal evidence.” There is an abundance of information available to establish much of this anecdotal evidence. It would be far better to provide landowners with reliable information of the interaction between specific nectar plants and insects. Two examples are offered here to illustrate how plant identification problems interfere with the understanding of climate-driven changes that may impact the food and nesting needs of wildlife.     Butterfliesandmoths.org(Butterflies) – a useful site that hosts a database compiled from several Lepidoptera collections – displays all manner of information that pertains to every US butterfly species including applicable regional maps, descriptions of species, larvae host plant information, and adult food preferences. While the botanical names for butterfly larvae host plants at this website are genuinely useful, very few botanical names are listed in their adult butterfly nectar food categories. The plant terms used on (Butterflies) to describe the adult food includes a few named taxa, but also includes insufficient descriptors that fail to guide a naturalist to a list of nectar plant taxa. These plant terms include: “unknown” (applied to 125 butterfly species), “flower nectar” (applied to 284 butterfly species), and common names of plants (applied to 391 butterflies). Those with an interest in supporting wildlife will benefit when plant identification tools are simpler to use so that the adult food category can be more fully developed. Inaturalist.org(inaturalist) – a depository of open-source scientific images of plant and animal life sponsored by the California Academy of Sciences – hosts over 2.5 million observations of birds and insects including over 700,000 observations of Lepidoptera, 300,000 observations of Superfamily Papilionoidea, 43,000 observations of Family Apidae, and over a million observations of birds. Very few of the observations of nectaring Lepidoptera have been correlated with the plants upon which they are feeding. Basically, (inaturalist) contains a vast compilation of data with much of the ecological use of the plant kingdom data yet to be developed. Mining the data in (inaturalist) to fill in the unknowns in (butterflies), or vice versa, would be simpler to achieve if plant identification processes were modernized. If those who collect observations of wildlife gathered a suite of images including information about the associated plants, a useful library of apical images would be available to generate vegetative trait data. Were simple plant identification resources available, then 1) the correlation between insects and the nectar plants upon which they depend would be more broadly known, 2) their use would enhance the ability to anticipate the plant needs of wildlife, 3) they would better empower naturalists to provide for plant dependent and nectar-feeding wildlife, and 4) they would enable those in the scientific community to better manage the impact of climate change.

DISCUSSION
    The earliest reference found that discussed a plant’s apical growth was written by Wright (1873) who wrote: “the limit to the Fibonacci series...was that no two leaves would ever precisely overlap when the stem was viewed from above.” While reference to the growing tip is found in subsequent literature, the angle of view was not described. The botanist Densmore (1920:17)wrote: “The spiral arrangement of leaves is more clearly evident at the apex of a branch, where the twisting effects of growth are less evident.” It is now pertinent to update these historic observations by stating that the top down view of the branch apex provides a concentrated view of a broad assortment of plant traits and trait clusters, including many uninfluenced by the effects of growth. That those observing a poplar cultivated plant, Rhododendron, failed to recognize that it’s leaf arrangement was semi-spiral, demonstrates that the angle of view of the growing tip clearly matters. Attending to it produces a body of evidence and diagnostic features that facilitate the development of robust vegetative keys. In 2009 Winsor stated that both Darwin and Agassiz acknowledged that (morphological) categories are both “artificial and natural.” This means that plant categories are natural in the sense that they are present, and artificial in the sense that they are categorized by taxonomists. Some natural categories, however artificial, have been overlooked, the apical complex being one. Recognizing the apex as a primary structural component of plants will facilitate the development of a vegetative key.
    Apical taxonomy, such as is modeled above, enables users to progress unimpeded through a key by incorporating a full range of omnipresent vegetative features into them and omitting references to micro-elements, seasonally delimited attributes, or non-binary features. The apex contains information about both customary and novel traits, and significantly, when taken in aggregate, contains a diagnostic cluster of traits that can be used to create efficient identification tools. Removing ephemeral attributes from keys, (such those found in the (Lexicon) and The Jepson Manual, Higher Plants of California) and thus the burden of uncertainty encountered in their use, is a major benefit. Producing a key that enables a user to navigate through salient features may require compiling an abundance of data, as was done during the development of both the Cactaceae and Euphorbia keys. In the Cactaceae sample, it was found that expanding the database by a few traits facilitated a reduction of the results to very few taxa, well below the 20-image threshold. A model key may either reduce a data set to a single result or reduce the burden of using it by narrowing a dataset to an increasingly smaller subset of apical images.
    A few teams have recently worked to develop automated plant identification tools that streamline and update customary keys (Pullan (2005), Victor (2011), Kilian (2015), Balduzzi (2017), and NRCS (2008)). Their work and the potential impact of apical taxonomy on their work is summarized below. Pullan et al (2005) developed the Prometheus Model. He states that the “expectations of the taxonomic community...are driven largely by the desire to make better and more efficient use of earlier research, mediated by the storage and exchange of taxonomic information in digital form.” His concerns over the use of and need to normalize the “natural language” used in historical keys have been advanced by the work of Kilian in 2015 and others. Pullan affirmed the need to normalize the terms used through the ages to describe each trait. Victor (2011) argued for the need to synthesize all the information about plants and make that information readily available and affirmed the need to synthesize and normalize historical data. Furthermore, these terms need to be examined to isolate those that may apply to the intersection between basic characters – such as between the leaf and petiole and the petiole and stem. Such intersection-traits are diagnostic in plants as diverse as grasses, palms, and Rhododendron. Some intersection-traits have been isolated and named, as is the case with culm in grasses and hastula in palms. This naming convention draws attention to the trait and serves to demonstrate its diagnostic and taxonomic utility. That said, the distinctive attributes and breadth of intersection-traits has yet to be fully explored, named and applied. Details of many of them can be captured in apical images. Naming the configurations that apply to the intersection between leaf and petiole – such as a petiole that terminates at or protrudes into the leaf base, or a protruding petiole that is either flush with or raised from the leaf face – will be shown to be taxonomically useful. While some of the petiole-to-stem attributes have been named – such as clasping – their taxonomic utility may not have been fully realized or applied. These intersection-traits can be used to fundamentally divide plant families. As it proves useful, their diagnostic utility can be used to justify elevating them to a superior position in a vegetative key and thus used to actively divide plant families. Kilian et al (2015), described a model key that produces numerous advantages over earlier systems, including the capacity to insert metadata, to generate novel “trees” (e.g. text keys and outlines that may be useful to those who study either regional flora or taxa that contain a particular trait), and to incorporate images into derivative resources. Kilian also affirms a need to normalize plant terminology. He states that a prerequisite to deriving the benefits of derivative resources is “that the data are structured and compatible;” and describes in depth the techniques used to generate these compatibilities. He stated that historical collections are “currently in a far-reaching process of transformation” to update their collections with the intent of making them more useful to researchers. In the future, adding apical images and data derived from them can be integrated into these systems as part of these needed transformations. Another model making use of SLIKS software is being used by biologists including those at Jepson and NRCS; one version of this software, developed by Dr. David Bogler in 2008 and found at the NRCS site, contains keys for five plant groups. Empowering a user at NRCS to reduce a field in the Eriacaceae key one attribute at a time in tandem is a true asset. While very promising, it will become even more useful when images are displayed with the text. The Ericaceae key contains 376 fields that apply to 153 taxa. Of these 376 fields, 137 referenced vegetation (13 of these 137 included numerical subcategories). Ideally, the ratio of fields to taxa will improve in future iterations; adding vegetative data, descriptive images, and apical images of each taxa will make this an even more powerful resource. New efforts are also underway to integrate mathematics into plant morphology resources. Balduzzi et al (2017) is working to normalize the language and systems used to create modern tools in an effort to overcome many of the difficulties that have plagued the construction and use of plant keys. His team acknowledges the challenges of developing tools that apply to organisms that may “transform over time.” The transformative quality of plants affects the ability of taxonomists to isolate a narrow set of characters that apply to an individual plant. This transformative quality impacts the ‘integrity’ of an attribute, as seemingly incongruous terms that have been applied to an individual may each be true. These arguments support the feasibility of integrating the morphological attributes of plant apices into the modern tools such as those modeled by Balduzzi. Apical taxonomy enables taxonomists to consider that there are intersection-traits with properties that are stable over time. The application of apical taxonomy may prove to mitigate and overcome the “incongruence between classifications based on different parts of the body or different life history stages.” In general, the traits compiled align with the current guidance advocated by Lindberg, namely that “the more characters on which it is based the better the given classification system will be” and that “every character is of equal weight.” He recognizes the need to use “more and better-described characters.” Thus, intersection-traits should be recognized and their value be equally weighted to all other vegetative attributes.
    These valuable new tools will not include information on apical taxonomy unless they’re specifically designed to do so. Thus, the specific novelty and value of the vegetative apical region of plants needs to be recognized so that both apical images and knowledge gleaned from them can be integrated into computerized resources.
    The study conducted and described in this article included preparation of written descriptions of attributes contained within the apical complex; these descriptions and the collection and study of apical images demonstrate that the apex contains a rich assortment of traits. The amount of exposed stem of an herbaceous plant or the density of its leaf coverage might be the equivalent of the omnipresent, vertical, fleshy valley between the ribs of cacti that is either grooved, smooth, or impressed with a line. The density of leaf coverage combined with information about intersection-traits proved to be equally diagnostic and reductive. Each of the vegetative categories, including characters and character traits, express themselves in a variety of ways, amounting to, minimally, 320 variations of character including macro leaf characters that apply to a leaf’s form, arrangement, primary shape, tip shape, margin shape, texture, distinctive qualities of new growth, primary color, secondary color, and finish. Twenty leaf venation patterns are included in this list. However, according to Andres-Hernandez 2012, there are seventeen leaf venation characters found in species of Bursera (Jacq. ex L.) alone. Further study is needed to assess the potential contribution to apical taxonomy of distinct leaf venation characters found throughout the plant kingdom, as the macro elements of them are unknown. (S5) This assortment of characters will be eclipsed when the full range of apically-derived data is added, as the work described supports the supposition that the list of traits can be greatly expanded. Fundamental attributes contained within the plant apices – such as the intersection-trait data described above – can be added to the list of 320 characters. Other novel apical attributes can be described in both negative and positive terms – e.g. the negative space described by the apical cluster and the positive density of visible leaves. The task of assigning traits to the appropriate taxa needs to be completed as well, for attributes unevenly assigned to each plant to which they apply form a barrier in the use of a key. Combining vegetative traits from all these sources will facilitate the development and use of modern tools. When fundamental plant attributes are added to this list – such as habit, form, type, location, and stem details – the possible combinations become even more meaningful and expands the character list exponentially. Vegetative traits can be used in a key to guide the user to a set of apical images, where the subtleties contained within them and the dynamics of the myriad possible combinations will be exposed to the advantage of the person seeking an identification. It is clear there are an abundance of vegetative traits; this abundance suggests that a review of the rationale used to undergird the insertion of ephemeral traits in plant keys is needed.
CONCLUSION
    Apical taxonomy presents an opportunity to develop a plant identification system that vigorously references a full range of both customary and novel vegetative attributes. It has been demonstrated that a broad range of benefits and efficiencies are realized in the use of keys unfettered from the constraints of plant hierarchies such as those modeled by Dunlap and NRCS. It has been demonstrated that vegetative attributes can be used successfully in the design of a key and that a key which directs attention to the branch apex reduces the burden of using it. It has been argued that apical images are an important component of identification tools as their use contribute numerous benefits: 1) empowering the collection of vegetative data, 2) providing efficiency by limiting the variety of vegetative attributes that need be isolated and described, and 3) accommodating the visual comprehension skills of users and thus capitalizing on the visual skills of the user to interpolate data contained in the apex without being obliged to translate that data into words.
    Providing individuals with the means to access information about a plant’s cultivation, monitoring, or care are additional benefits of apical taxonomy. By creating tools that are simpler to navigate, an interest-related level of mastery is conceivable; those who wish to acquire knowledge about plants – whether they be professionals, scientists, or novices – can more easily progress to productive levels of understanding, as they may more readily overcome the linguistic hurdles embedded in the current systems. Anticipating the requirements of wildlife for plant food, nectar, and seeds further necessitates the need for contemporary plant identification resources. Neither the text-based nor the visual systems available allow blind queries within the potential vegetative inventory. It is conceivable that overcoming the handicaps of the current systems will boost the diversity of plants in cultivation and enhance the collection of knowledge of the interdependencies between plants and wildlife.
    Apical taxonomy presents an opportunity to improve both formal and informal plant identification resources, expand knowledge about the vegetative characteristics of plants, and facilitate the development of conservation and pollinator pair data. The improvements occur by drawing attention to the plant apex, by enabling users to start with what is known, and by adding apical images to keys and automated tools. The proposed system can help individuals gain knowledge about plants and acquire the benefits associated with the stewardship, tending, and care of the natural world. Taxonomists, released from the burdens associated with systematics described above, can repurpose their mandate.

Compliance with Ethical Standards
Conflict of Interest: The author declares that there are no conflicts of interest.
Funding: This study received no funding.

Supplemental Material
S1-S11
S1 Background of discovery.
S2 Sample of Herbaceous plant traits – leaf details.
S3 Euphorbia apical photograph compilation – Jacobson.
S4 Euphorbia sort criteria.
S5 Euphorbia Photography Master – sample.
S6 Euphorbia comparison of photographs overview.
S7 Euphorbia – visual data in apices.
S8 Euphorbia – visual data in apices, descriptions by others.
S9 Ferocactus – visual data in apices.
S10 Master Photography List – other genera – cacti.
S11 Master Photography List – other genera – succulents (non-cacti).

S1 Dunlap, S. October 30, 2002
I. Background
II. Statement regarding Prominent Attributes of Apical System
III. Process Used to Identify Euphorbia
I. Background: The current state of identifying a plant.
Current methods used:
    1. Electronic and Traditional Libraries and Publications
    2. Human Resources
1. Electronic and Traditional Libraries and Publications
Someone seeking to identify an unknown plant specimen using literary resources will find that the easiest system to navigate is a visual key. The second choice is a verbal (written) key concentrating on a plant’s visual characteristics - one avoiding as much as possible the use of ephemeral characteristics and internal processes. Finding one’s way to this reality is not obvious.
Data about 50,000 U.S. native, commercial, and naturalized species have been documented and made available for research on the Web by the USDA. The plants are organized by scientific and common names. The data is designed for someone seeking information about a known specimen or plant group. For a novice and for someone seeking information about an unknown specimen, there are obstacles. For example, there are no keys at the site (visual or verbal), and some discretion must be used when inputting common names. Entering ‘conifers’ leads to a helpful site whereas entering ‘evergreen’ does not.
Photographs of many of the plants are available both at the USDA site and at others linked to it (e.g., one link is Cal Photo, an excellent user-friendly UCB site of California natives and other flora). However, the novice will encounter hurdles using this system as well. For example: the keys that include the California nutmeg indicate the leaves of the plant are flat and opposite; Cal Photo images show some species with leaves that spiral and two different cones. After researching three other web sites, information was found regarding the differences between the male and female plants; these differences have not been clearly stated with consistency across the system. There are photographs of +-25 percent of the species in the database. The photographs convey a general quality about the plant but rarely illustrate enough detail to identify a species. Those seeking to identify a specimen have to look elsewhere.  
On a smaller scale, one’s ability to navigate to a species improves provided the plant group is known by the seeker. Many states compile information on plants native to their region and many specialists compile monographs on a specific genus or a small plant group (e.g. conifers). Many of these agencies and authors include keyed identification systems – some visual, some verbal. The keys are typically limited in one of three ways: 1) in breadth (e.g. leaf/twig keys of regional deciduous trees), 2) ease of navigation (too obtuse), or 3) in depth (terminating at the genus level). These smaller concentrated keys can be useful when used in conjunction with keys of larger breadth. They frequently pick up where the larger keys leave off, or, add species specific details omitted in large volumes. Limitations aside, the smaller scale resources organized around a key are the easiest to navigate for someone seeking identity of a species within the group of plants contained in the database. These resources can frustrate a novice who may not have the ability to recognize the lack of breadth.
One dichotomous key of great breadth is in The Jepson Manual, 1993, a 1424-page volume of 8,000 California natives. This book, fundamentally comprised of text, is a major primary resource. Its editors have made every attempt to construct the key using vegetative characteristics (non-reproductive) and simple terminology, and advise that non-native species in an included genus may not fit into their key. A novice using this resource will have difficulty navigating to an unknown specimen. Two features limit the volume’s usefulness for that aim: 1) the keys do include ephemeral and linguistically challenging details and 2) the keys are specific to the plants covered in the volume.
The Flora of North America, 2002+, is a multi-volume set on native flora in North America. About 20 percent of the set have been published. This resource is similar to The Jepson Manual, with the same limitations. Like the Jepson, it uses keys at the beginning of each plant group to guide to a family. Within a family, there are keys to guide to a genus and within the genus to the species. In the gymnosperm section, one cannot navigate to three of the six families covered without superior linguistic skills, a reproductive sample, a microscope, and prior knowledge that the species in question is a native plant included in the group.
In Conifers of California, R.M. Lanner, 1999, the author came close to constructing a vegetative key. The book includes two dichotomous keys of California conifers. One is based on the seed-bearing structure. The other is based solely on leaf characteristics. In every instance, the leaf key leads to a genus in the text, and in some cases a specific species. Within the text, one finds additional leaf characteristics that isolate a species. However, this guidance is not systematically included (e.g. the position of leaf glands on cypresses can isolate a species; this information was not systematically discussed or added to the key). This text is organized based on vegetation, and nearly overcomes the suggestion that unique characteristics of a species are invisible or ephemeral. A novice with prior knowledge that a specimen is a California native conifer can get to the genus 100% of the time and the species approximately 60% of the time using this resource. Anyone with a specimen of a non-California native, cultivated, or hybrid (natural or bred) conifer will have to look elsewhere. One monograph, (Kussman, Gerd, Timber Press) in the library of a regional botanical garden, covers all conifers including clones, hybrids and cultivated species. Its sophisticated keys are designed for professional caretakers, breeders, and cultivators.
Another class of literature is available for those with horticultural interests. In this area the primary book is The American Horticultural A-Z Encyclopedia of Garden Plants, 1997. The 1095-page volume covers 15,000 cultivated plants in 2,000 genera (including some varieties and many cultivars) and has 6,000 photographs. The text is alphabetically arranged by genus using scientific names, supplemented by an index of common names. It is primarily a tool for gardeners and landscapers. In addition to instructions for planting and care, it includes basic educational text. It does not have keys. Those seeking the identity of a plant will be hampered by similar limitations to those found in the botanical literature.  
Those books available for the specialized collector cover species in limited groups of genera. The best of these are informative and some include photographs of every species in a genus. However, the tendency to under-represent a species in the photograph persists – in deference to floral characteristics, unique omnipresent features are not represented systematically. It is uncommon for these books to include keys.
Thus, three weaknesses hamper the use of the above resources for identifying an unknown specimen. One is that the keys are either nonexistent or incomplete. Secondly, the photographs where present do not clearly illustrate the unique physical trait(s) of a particular species. The data isolating a species from a close relative is not represented in a photograph. Thirdly, navigating to a species requires either advance knowledge of the name of a plant or a working knowledge of vegetative features including flower parts, non-floral organs, structural elements, and surface features, and even, in some cases internal processes. This is the case even though it is only the constant and visible details which are accessible to the nonprofessional. Too much is required of the novice for several reasons. Firstly, the novice is quickly overwhelmed by specialized terminology – e.g. eighty-five specialized botanical terms are encountered when a novice seeks to develop an understanding of the term “gymnosperm.” Secondly, there is typically no warning a key system may be incomplete. In addition, the visual quality of the plant – one of the primary qualities that aroused interest to begin with – is quickly lost or omitted. These weaknesses conspire to hamper the capacity and willingness of a novice to engage in a kingdom as omnipresent and essential as plants.

2. Human Resources, including personnel at: Local Nurseries, Specialized Societies, Botanical Gardens,
Universities, Herbariums: Regional and National, and World Experts.
There are two overlapping systems of human resources to engage when seeking the identity of a plant. One is oriented around scientific interests, the other around cultivation interests. As in a text-based query, it is not obvious to a novice this distinction exists. The author found that only at the expert level is one able to forego the necessity of exchanging a sample of the plant. In such an instance, the person seeking to identify a specimen would have to be skilled at communicating precise botanical details to the expert. At any level other than expert to expert, the exchange of a visual sample of the plant in question is required – either excellent photographs or a specimen. A university of international stature requests a physical sample of a plant is included with any identification query. Without an adequate photography standard in place, a specimen is required. Thus, even at a professional level, a physical sample or a clear visual representation of the specimen is preferred over a verbal description. This same courtesy needs to be offered to the novice.
In both resource systems the personnel prefer a plant sample that includes a reproductive organ – fruits, flowers, and/or seed pods. If a vegetative sample is provided to personnel in a scientific environment (e.g. the plant identification service provided by UCD), it is likely that they will be able to identify it to the genus level. When the reproductive organ of the plant is included, they can identify the species. Were that same sample given to the local nursery personnel, the success rate would likely depend upon the sample being from a cultivated species. Personnel at a specialized local nursery can typically isolate a cultivated vegetative specimen to a plant family or genus. To go further one must find experts.
Experts can sometimes be found locally, in societies or botanical gardens. Societies, organized around different plant groups, frequently attract individuals with expert knowledge about a genus within a particular plant group. These societies by no means cover all plant groups (approximately five plant groups are so represented in Northern California). There is no guarantee that an individual with complete expertise about a particular genus is a member. Where present these local experts can be an excellent resource. If further research is needed, they can point to a new resource. Some of these societies have excellent specialized libraries.
Local botanical gardens are another resource for finding specialists. Frequently there is a plant expert on the staff who may provide assistance. Botanical gardens tend to be specialized also – focusing on particular plant groups, local or native flora, or a large focused private collection made accessible to the public. It has been the author’s experience that regional botanical gardens tend to have the broadest collections while private ones are more specialized. If either the regional or the specialized garden has the genus in question, then the search may stop there, for someone on the staff is likely informed about all species in the genus. However, in the larger gardens the expert may not be readily available to answer questions. In such a case, the administrator’s policy may allow someone to leave a plant specimen for possible identification.
If the search has yet to yield results, one must pursue resources at a university or herbarium. If the searcher has established that the plant in question is a native, a query to a regional university or herbarium should produce good results. If the specimen is an imported species, one may have to query to a national or international herbarium. In a focused two-day search on the web, looking under ‘plant identification’ the author discovered two fee-based plant identification services. For a fee, they would identify a specimen to the level of family, perhaps to a lower level. After an additional two days of research in a related area, the author stumbled upon a free plant identification resource linked to a plant photo database. The personnel at this source (UCD), with 250,000 herbaria samples and an extensive library, expected to be able to identify a plant to the genus level when given a vegetative sample. They expected to reach the species when provided with a reproductive organ. A fee-based contract for their service is available for those with an on-going need for plant identification. Such contracts have been made with the United States Forest Service – who may provide them with as many as 300 samples at a time – and with an individual requesting identification of 20 plants or more per annum. If the sample in question is part of a particularly large and diverse genus, like Euphorbia, the quest may require input from a world expert. Finding the right one and approaching them can be a challenge.  

Summary:
    1. A keyed navigation system free of reproductive organs does not exist. The existing keys and literature suggest that in many instances such a key could be constructed to the species level.  
2. Separating two species within one genus may rely upon identifying a subtle difference in a single physical trait. Frequently that trait can be photographed and the related and similar species successfully compared. The utility of the existing system would improve were more rigorous observation and photographic standards established and implemented. Existing keys can be canvassed for persistent vegetative features and expanded to include all visible traits. The expanded keys would guide those collecting photographic data. Several sources, both professional and amateur, suggest using a 10x lens to examine specimens for details included in a key. This magnification is well within the range of photographic equipment.
3. The apex of a vegetative branch or stem contains a unique cluster of traits. It portrays the basic structure and scale of the stem, the type of growth activity contained in the center of the apex (the leading edge), and, where present, leaf characteristics such as leaf arrangement, leaf type, and leaf color. Frequently, as with cacti and succulents, the apex also reveals epidermal characteristics of the stem.
4. A majority of species studied contained a unique physical trait in the apex, enabling a direct visual comparison to close relatives.
5. The trait-cluster contained in apex of a plant provides an efficient view. When pattern recognition software improves, apical photographs can be used as a critical component of an identification system.
5. Plant trait data contained in apical photographs can be incorporated into dichotomous keys that becoming increasingly useful as more species are added. Keys containing apical details need to be devised.
II. Statement regarding Prominent Attributes of Apical System
    The apical view adds a classification tool for managing and navigating the plant kingdom. This tool may prove to be as helpful in distinguishing species as the flower is now. It can be used to make the existing plant identification process more efficient. Its efficiency is derived from the fact that it records multiple features: the overall, species-specific vegetative pattern and a concentrated view of several characteristics – a trait-cluster. Its effectiveness stems from the fact that its implementation creates useful photographic standards and retains the visual characteristics omnipresent in the plant kingdom.
In the genera studied as many as 10 characteristics were recorded in the apical view. Thus, ten steps in existing keys can be condensed. The efficiency of the apical view trait-cluster makes it possible to devise a manageable visual system - a photo-based key of the plant kingdom. A photo-based key could be used either independently or as a supplement to text-based resources. It could guide users to the scientific name thus making available to them the abundant text-based resources so arranged.  Even professionals use a photo-based resource to guide them to a scientific name (the plant itself).  
    The incorporation of an apical view into the current system will make it possible to delete many non-vegetative clues in the existing keys. The virtue of its visual attributes is that the universal characteristic exchanged in an encounter between humans and plants is retained. It aligns itself with the visual attributes of the plant kingdom.
    The photographic discipline required in order to generate apical data will contribute to the overall body of knowledge about plants. This will be achieved by the fact that this system generates intimate visual data about vegetation that has not yet been incorporated into the verbal keys. Thus, the secondary data collected will further contribute to the expansion of both verbal and visual keys.
Field collecting guidelines for this system can be simply described. Additionally, by overriding a reproductive focus, its vegetative focus loosens the seasonal constraints currently imposed on fieldwork.
    The apical system can be used to construct a novice-friendly identification system. Its visual attributes address the linguistic hurdles encountered in the current system. Once established and in place, this system can be used to guide a novice through the plant kingdom. Only with tremendous effort, time commitment, resources, professional connections, and persistence can one consistently navigate to a species using the existing system.  
    Both the professional and novice are affected by primary weaknesses in the current system: 1) the bias of keying off reproductive organs creating an environment where vegetative features have been overlooked, and 2) the lack of methodical and disciplined photography. Apical systematics, and its accompanying photographic discipline, begins to resolve the problems created by these weaknesses.

III. Process Used to Identify Euphorbia
The author pursued the identity of both native and cultivated plants. In both plant groups, the pursuit involved both human and literary resources. Described below is the approach taken to identify several species of cultivated Euphorbias.
Plants were purchased from local sources. Frequently, a plant was identified in broad terms, as either a cactus or succulent. Some plants lacked any identification labels. In other instances, an unidentified cutting was acquired from a friendly source. The plants on hand at all local vendors, both specialized (seven) and non-specialized (five), were examined for a match. This method was sometimes successful, especially if pursued over time.
If the plant was identified in broad terms on a label and purchased from a non-specialized vendor (e.g. a large-scale multi-purpose retailer), then the staff was not approached. Generally, staff at such a vendor has limited botanical knowledge and the incomplete labeling provided by their wholesaler betrays the limits of the wholesaler’s capacity. Subsequently, it has been born out that some seedsmen offer seeds bundled in family groups. These seeds are less expensive than those identified by a binomial. The plants so cultivated are labeled by a common name (at the family level), such as ‘cactus’ or ‘succulent’.
When dealing with specialized vendors, the author approached the staff to help identifying the species. One retail vendor in the San Francisco Bay Area has a small library of books accessible to their clients. These books are helpful but limited (discussed above). Most retail vendors have staff capable of identifying broad groups of plants. If one desires the scientific name, they offer to forward a request to their wholesaler. Most often, a query to the wholesaler was required. Several repeats of this sequence revealed a local wholesaler with a consistently thorough knowledge of his plants. Later this vendor was approached directly. As careful as he is, this grower too was happy to receive assistance identifying some of his mother plants and imported stock.
Plants falling outside the range of local retail sources were encountered. A visit to the library of the regional botanical garden – 35 miles north of San Francisco – proved helpful. This library had information on specific plant groups and about local and national societies, pointing the way to more specialists. A prominent advertisement in a publication found there, produced by the Cacti and Succulent Society of America, led the author to a specialized local garden with limited public access.
A visit to this garden, a 90-minute drive north, proved helpful. The founder and primary collector, Dr. Herman Schwartz, gave a tour of his cacti and succulent collection. He was the primary editor of several specialized books including a 10-volume set of books on Euphorbias. He agreed to allow the author to photograph and study his collection. He too was in need of identifying many plants in his extensive collection.
For the next sixteen months, this garden was visited on a weekly basis and 2000 photographs amassed of its collection. Between visits, the images were organized, printed, and studied. The plant’s identity was pursued by comparing vegetative cuttings and original photographs to the published images found in books – including unique out-of-print volumes on loan. Reference books were required in order to understand and make use of these texts. Both a botany dictionary and glossaries were sought and, where available, acquired. The best visual glossary on cacti (Buxbaum, Dr. Franz. Morphology of Cacti, 1950) has been out of print for over fifty years. Many plants remained to be identified.
During an overlapping seven-month period, on a semi-monthly basis, other regional resources were pursued. Other regional botanical gardens, collectors, and specialized growers were visited and their collections photographed and studied. Research was conducted at two university libraries (Stanford and UC Berkeley). Many unidentified plants remained.
During that same period, monthly meetings of a specialized society and its biannual national convention were attended. The local society meetings were of high caliber, informative, educational, and managed by botany professionals. Much was gained by listening to these professionals – most especially the curator of the Ruth Bancroft Garden, Brian Kemble. Specialists and specialized vendors attended the convention – including new growers, botanical experts, and book vendors. Researching these resources confirmed that the last published key to Euphorbias was generated in 1941. Many newly discovered species have been circulating since then, including several in Dr. Schwartz’s collection. A number of sources indicated that the principal U.S. grower for the genus was located in Arizona.
A ten-day visit to the grower was arranged. Without an historical relationship between Dr. Schwartz and the nursery owner, this visit would not have been possible. Among other things, the trip was complicated by 113-degree heat in the greenhouses. Nearly 700 photographs were taken of their Euphorbias. Over a subsequent three-week period, these images were organized, printed, and compared to those already printed from the original collection. Some unidentified plants remain. This last group can be identified by purchasing named plants from the group in question and comparing them to existing plants in the field. The unidentified list of plants has narrowed to a few within the Euphorbia medusa group – about a dozen plants.


S2 sample of Herbaceous Plant traits
Leaf details
Prepared by Susan Dunlap

Conifer
Retention
deciduous
evergreen
semi-evergreen

Form
compound
lobed
simple

Fragrant

Arrangement
tuft
spiral
alternate
rosette
opposite
2-ranked
4-ranked

Primary Shape
elliptic
hastate
lance
lanceolate
ligulate
linear
lorate
lyrate
oblanceolate
oblique
oblong
obovate
ovate
sagittate
scale
spathulate
Tip Shape
blunt
notched
pointed
rounded

Margin Shape
hairy
rough
scalloped
spiny
toothed
undulate

Texture
bloom
glandular
hairy
pubescent
rough
smooth
spiny
spotted
sticky
stiff hairs
textured
tubercles
veins folded
veins textured
waxy
woolly

Basic character
leathery
rigid
stiff
succulent
thick
thin

New growth distinct
color
form

Primary Color
cool
blue-green
dark green
gray-green
light green
medium blue-green
medium green
medium gray
olive
purple
varied
yellow-green
dark
black
brown
purple
varied
neutral
gray
medium gray
varied
white
warm
orange
pink
red
varied
yellow

Secondary Color
leaf face
mottled
spotted
striped
variegated
margin
veins

Size
extra-large
large
medium
medium-large
small
tiny
very small

Finish
glossy
matte
semi-glossy

S3 Euphorbia apical photograph compilation by Susan Dunlap
Groupings by Jacobsen, Hermann F.  Lexicon of Succulent Plants. London, Blandford Press Ltd., 1974.
Created Dec 6, 2001; Updates: Jan. 30, 2002; March 1, 2002; March 2, 2002, July 17, 2002, July 30, 2002

Group#            # of species        % of group    Total in Group       
    photographed         identified                      
MI    moved to Group 4
MII    moved to Group 2
MIII        2    =    33.3%        6        
MIV        0                2
MV        1        100%        1
MVI           2     =    11.11%        18
MVII        0    =            9    
MVIII        0                2
1        0                16
2        1    =    12.5%        8
3        0                8
4              1    =    1.49%        67        
5        0                4
6                2          =    40%        5
7        0                5
8        0                6
9             2          =    15.38%        13        
10        0                3
11             3       =    100%        3
12        1    =    100%        1
13         3     =    42.86%        7
14        10    =    52.63%        19
15         11    =    47.83%        23
16         1    =    50%        2
17         4    =    66.67%        6
18         9    =    90%        10
19         14    =    66.67%        21        
20        2        9.09%        22
21         5    =    62.5%        8
22        13    =    68.42%        19
23        0                3
24         62    =    54.39%        114
25a        1    =    33.33%        3
25b        1        100%        1
26         1    =    12.5%        8
27        4        66.67%        6
28         4        30.77%        13
Subtotal:    160     =    34.63%        of 462        

July 30, 2002 data:
Group 0        84    =    100%        of x# in Group*    
Grand total:    244    =    44.69%        of 546         Results Hold for 244 species.

*NIC 50 photographs of species that had yet to be identified.
Expected adjustment to total: 244+50=294, or 49.33% of 596 species in above groups.

Euphorbia Groups. Jacobsen, Hermann F. Group 0
actinoclada
adjurana
aff. Gorgonis
albipollinifera
amfolozieuzis
ammak
andanica
angustiflora
astrophora
atroflora
baoiensis
barnhartii
boranensis
brevitorta
bussei
chorisia
classenii
clavyii
clivicola
cummulate
cuprispina
Djibouti Lav.
dusimata
elegantissima
eyassiana
fluminis
fractiflexa
furcata
galgalana
gamkensis
giletti
greenwayii
groenwaldii
gymnocalycium
hermentian
holmesiae
horwoodii
hypogaea
ingens
kalisana
leistneri
lowii
magnacapsula
magnacapsularis
makallensis
malevola
marsabitensis
mayuranathani
meridonalis
micrantha
mitriformis
mosaica
mozambiquensis
Mrs. Ash
nubigena
nutrufirnus
ornithopus
persistifolia
petraea
pseudoburuana
quadrisoina
richardsiae
robusta
robustior
rubechii
rubrispinosa
s.n. Zimbabwe
saxorum
scizacantha
seibanica
septentrionalis
sepulta clone
serendipita
strangulata
subsala
supressa
taruensis
teixeirae
trapifolia
turbinicarpus
utoniensis
vajravehii
valaris
viduiflora
xylacantha
Group 1
anachoreta
atropupurea
balsamifera
barbicollis
berthelotii
bravoana
dendroides
lambii
mellifera
noxia
orthoclada
piscatoria
pseudograntii
punicea
quadrata
tuckeyana
Group 2
boissieri
commersonii
epiphylloides
hedyotoides
leuconeura
lophogona
neohumbertii
viguieri
Group 3
espinosa
frutescens
giumboensis
grosseri
geurichiana
sacchii
scheffleri
somalensis
Group 4
aequoris
alata
amarifontana
angrae
antisyphilitica
aphylla
arbuscula
arceuthobioides
aspericaulis
brachiata
burmanii
cameronii
cassythoides
carunculifera
caterviflora
carifera
caulis
brachiata
burmannii
cameronii
cassythoides
caruncunculifera
caterviflora
cerifera
fragilramosa
gentilis
gossypina
gregaria
geummifera
herrei
intisy
juttae
karroensis
lactiflua
laro
lateriflora
leucodendron
lignosa
macella
mauritanica
muricata
mundii
nubica
obtusifolia
oncoclada
paxiana
perpera
platyclada
pseudobrachiata
plagiantha
rectirama
rhombifolia
rudolfii
sarcostemmatoides
schimperi
silicicola
spartaria
spicata
spinea
stapelioides
stenoclada
stolonifera
tenax
tirucallii
transvaalensis
verruculosa
xylophylloides
Group 5
phosphorea
pteroneura
sipolisii
weberbaueri
Group 6
gariepina
halleri
hamata
peltigera
schaeferi
Group 7
gueinzii
ledermanniana
multifida
pseudotuberosa
trichadenia
Group 8
crispa
ecklonii
pseudohyopogaea
rubella
silenifolia
tuberosa
Group 9
bubalina
bupleurifolia
clandestina
clava
cuylindrica
grantii
hallii
longetuberculosa
montieri
oxystegia
platycephala
pubiglans
wildii
Group 10
eustacei
loricata
multifolia
Group 11
fasciculata
restituta
schoenlandii
Group 12
clavarioides
Group 13
ernestii
flanaganii
franksiae
gatbergensis
gorgonis
pugniformis
woodii
Group 14
bergeri
bolusii
caput-medusae
colliculina
confluens
davyi
duseimata
esculenta
fortuita
bypogaea
inermis
maleolens
marlothiana
muirii
pseudoduseimata
ramiglans
superans
tuberculata
tuberculatoides
Group 15
albertensis
argillicola
arida
baliola
bergii
brakdamensis
braunsii
brevirama
crassipes
decepta
filiflora
friedrichiae
hopetownensis
inornata
marientalensis
melanohydrata
multiceps
namibensis
nelii
orabensis
pentops
rangeana
rudis
Group 16
namaguensis
multiramosa
Group 17
globosa
ornithopus
planiceps
polycephala
tridentata
wilmanae
Group 18
jansenvillensis
juglans
meloformis
obesa
pseudoglobosa
susannae
symmetrica
tubiglans
turbiniformis
valida
Group 19
aggregata
anoplia
cereiformis
cucumerina
cumulata
enopla
ferox
fimbriata
heptagona
horrida
inconstantia
leviana
mammillaris
nesemannii
pentagona
pillansii
poissonii
polygona
pulvinata
stellaespina
submammillaris
Group 20
beharensis
boiteaui
brachphylla
capuronii
delphinensis
denisiana
duranii
fianarantsoae
genoudiana
guillemetii
horombensis
isaloensis
leandriana
mahafalensis
mangokyensis
milii
quartzicola
razafinjohanii
rossii
tardieuana
tzimbazazae
zakamenae
Group 21
desmondii
neriifolia
nivulia
royleana
subanica
teke
trapiifolia
undulatifolia
Group 22
aeruginosa
clavigera
clivicola
decidua
dekindtii
enormis
fanshawei
groenewaldii
imitata
knuthii
micracantha
persistens
restricta
schinzii
squarrosa
stellata
tortirama
tortistyla
vandermerwei
Group 23
alcicornis
dawei
ramipressa
Group 24
abyssinica
acrurensis
ambroseae
angularis
antiquorum
atrocarmesina
avasmontana
ballyi
barnardii
barghartii
baylissii
bothae
bougheyi
breviarticulata
buruana
cactus
canariensis
candelabrum
carterana
coerulescens
complexa
confertiflora
confinalis
conspicua
contorta
cooperi
curvirama
cussonioides
dawei
decliviticola
deightonii
disclusa
doinetiana
echinus
eduardoi
erlangeri
evansii
excelsa
fortissima
franckiana
golisana
gracilicaulis
grandialata
grandicornis
grandidens
graniticola
griseola
halipedicola
handiensis
heterochroma
hottentota
hubertii
inarticulata
inculta
ingenticapsa
intercedens
jubata
kamerunica
keithii
kibwezensis
knobelii
lactea
ledienii
lemaireana
letestui
lividiflora
longispina
lydenbergensis
macrogluypha
malveola
maluensis
memoralis
migiurtinorum
mlanjeana
neglecta
neutra
nigrispina
obovalifolia
officinarum
opuntioides
paganorum
parciramulosa
perangusta
persistentifolia
phillipsiae
polyacantha
proballyana
pseudocactus
qarad
quinquecostata
ramulosa
reinhardtii
resinifera
robecchii
rowlandii
semperflorens
sekukuniensis
seretii
spiralis
strangulata
tanaensis
tenuirama
tetragona
thi
tortilis
triangularis
trigona
venenata
virosa
volkmanae
wakefieldii
waterbergensis
williamsonii
winkleri
zoutpansbergensis
Group 25a
cryptospinosa
fruiticosa
multiclava
Group 25b
columnaris
Group 26
darbandensis
immersa
poissonii
sapinii
unicornis
unispina
venenifica
Group 27
ballyana
glochidiata
graciliramea
monacantha
schizacantha
triaculeata
Group 28
augustiflora
coerulans
ellenbeckii
inqequispina
isacantha
ndurumensis
nyassae
quadrangularis
susalsa
tetracantha
tetracanthoides
uhligiana
whellanii
Group MI - moved to Group 4
Group MII - moved to Group 2
Group MIII
caput-aureum
croizatii
didieroides
guillauminiana
pauliana
perrieri
Group MIV
biaculeata
pedilanthoides
Group V
pachypodioides
Group VI
beharensis
brachyphylla
capuronii
delphinensis
duranii
fianarantsoae
genoudiana
guillemetii
horombensis
leandriana
mangokyensis
milii
quartzicola
razafinjohanii
tardieuana
tsimabazazae
zakamenae
Group VII
anakarensis
boiteaui
cap-saintemariensis
cylindrifolia
decaryi
francoisii
milloitt
moratii
primulifolia
Group VIII
bosseri
platyclada


S4 Sample from Euphorbia Photo Master
Prepared by Susan Dunlap, 2003

Species                        Group            slide info     slide id

Euphorbia abyssinica            GR24        P    A20-06
Euphorbia abyssinica             GR24        P15    apices & brdr, Sl27/15-16, Sl31/10-13, apex,                             s/32-15, Sl25/19-20
Euphorbia abyssinica variegated         GR24        P    Sl32/16
Euphorbia acrurensis                   GR24            Sl8/12; no apex

Euphorbia actinoclada            GR0        P    A14-16
Euphorbia actinoclada            GR0        P22+    Ann Sl17/22+(3 P's)
Euphorbia adjurana                GR0        P24-5    A3-23-26 A14-2

Euphorbia aeruginosa            GR22            ctg apex, bdr33/3-4
Euphorbia aeruginosa            GR22        P33    A1-33,34
Euphorbia aeruginosa             GR22        P17    Ann SL30/16-18
Euphorbia aeruginosa nova            GR22?        P    A13-7
Euphorbia aff. Gorgonis            GR0        P    A10-9
Euphorbia aggregata                GR19        P    A3-4
Euphorbia albertensis            GR15        P    SL37/33

Euphorbia albipollinifera            GR0        P    A10-16
Euphorbia albipollinifera cockscomb    GR0        P    A22-16    
Euphorbia amfolozieuzis            GR0        P10    A16-10,11

Euphorbia ammak            GR0        P    A3-33
Euphorbia ammak            GR0            frost apex/spp/                                    P17-8    rib Sl11/16-19

Euphorbia ammak variagata        GR0        P35-6    A4-35-7
Euphorbia ammak variagata        GR0        P    A20-01
Euphorbia andanica                GR0        P1    A2-1,2
Euphorbia angularis               GR24        P    SL42/25    
Euphorbia angustiflora            GR0        P17    A11-16,17

Euphorbia anoplia            GR19        P    A9-30
Euphorbia anoplia             GR19        P    SL43/3

Euphorbia antiquorum            GR24        P28    A1-28-31                                    P29    A329-30
Euphorbia antiquorum            GR24        
Euphorbia antiquorum            GR24         P19    apices Sl26/16-20,31, Sl32/2-3    

Euphorbia arida GM 290            GR15        PPP    A22-31,33-4    
Euphorbia arida                GR15        P    A10-11    
Euphorbia arida                GR15        P34    SL40/33-35, Sl43/8
Euphorbia astrophora                GR0        P    A3-3
22. Euphorbia atroflora            GR0        P    A9-22

Euphorbia abyssinica            GR24        P    A20-06
Euphorbia abyssinica             GR24        P15    apices & brdr, Sl27/15-16, Sl31/10-13, apex,                             s/32-15, Sl25/19-20
Euphorbia abyssinica variegated         GR24        P    Sl32/16
Euphorbia acrurensis                   GR24            Sl8/12; no apex

Euphorbia actinoclada            GR0        P    A14-16
Euphorbia actinoclada            GR0        P22+    Ann Sl17/22+(3 P's)
Euphorbia adjurana                GR0        P24-5    A3-23-26 A14-2

Euphorbia aeruginosa            GR22            ctg apex, bdr33/3-4
Euphorbia aeruginosa            GR22        P33    A1-33,34
Euphorbia aeruginosa             GR22        P17    Ann SL30/16-18
Euphorbia aeruginosa nova            GR22?        P    A13-7
Euphorbia aff. Gorgonis            GR0        P    A10-9
Euphorbia aggregata                GR19        P    A3-4
Euphorbia albertensis            GR15        P    SL37/33

Euphorbia albipollinifera            GR0        P    A10-16
Euphorbia albipollinifera cockscomb    GR0        P    A22-16    
Euphorbia amfolozieuzis            GR0        P10    A16-10,11

Euphorbia ammak            GR0        P    A3-33
Euphorbia ammak            GR0            frost apex/spp/                                    P17-8    rib Sl11/16-19

Euphorbia ammak variagata        GR0        P35-6    A4-35-7
Euphorbia ammak variagata        GR0        P    A20-01
Euphorbia andanica                GR0        P1    A2-1,2
Euphorbia angularis               GR24        P    SL42/25    
Euphorbia angustiflora            GR0        P17    A11-16,17

Euphorbia anoplia            GR19        P    A9-30
Euphorbia anoplia             GR19        P    SL43/3

Euphorbia antiquorum            GR24        P28    A1-28-31                                    P29    A329-30
Euphorbia antiquorum            GR24        
Euphorbia antiquorum            GR24         P19    apices Sl26/16-20,31, Sl32/2-3    

Euphorbia arida GM 290            GR15        PPP    A22-31,33-4    
Euphorbia arida                GR15        P    A10-11    
Euphorbia arida                GR15        P34    SL40/33-35, Sl43/8
Euphorbia astrophora                GR0        P    A3-3
22. Euphorbia atroflora            GR0        P    A9-22

Euphorbia avasmontana         GR24           P3,16    Sl28/6, Sl29/3,16                                     P6,9    apex Sl6/06+, Sl19/34- 3, SL42/9-10
Euphorbia avasmontana            GR24        P    A22-01  
Euphorbia ballyana                GR27        P    A2-6
Euphorbia baoiensis                GR0        P29    A4-25-6,29

Euphorbia barnardii            GR24        P24    A13-24,5
Euphorbia barnardii            GR24        P28    A19-26-28?
Euphorbia barnardii             GR24         P17-8    SL29/17-20, A abbey garden, Transv                                                                        apex Sl27/19, 35/9-10
Euphorbia barnhartii                GR0        P35    A1-35-7
Euphorbia boranensis                GR0        P17    A9-16,17
Euphorbia bothae                GR24        P    A14-25
Euphorbia bougheyi                GR24        P18    A13-18,19
Euphorbia brevirama                GR15        P25        SL40/25, Sl43/37
Euphorbia brevitorta                GR0        P    A10-10
Euphorbia buruana                GR24        P13    A10-13,14
Euphorbia bussei                GR0        P    A15-8
Euphorbia cactus                GR24        P-22    A12-22,23

Euphorbia canariensis            GR24        P    A3-11                                        P19    A14-19,20, A15-21,23
Euphorbia canariensis             GR24        P1           Jim; apex, SL38/1+31
Euphorbia canariensis             GR24        P36-7    apex, Sl7

Euphorbia caput-medusae        GR14        P    A19-22, A19-11
Euphorbia caput-medusae        GR14         P43-4    apex Sl37/20+26,                                    P42-6    Sl42/6-7, Sl43/4
Euphorbia chorisia?                  GR0            Sl11; FLWR
Euphorbia classenii                GR0        P11,12    A12-11,12
Euphorbia clava                GR9        P    A3-10
Euphorbia clavarioides v. truncata        GR 12        P    A17-36

Euphorbia clavigera            GR22        P    A10-12                                        P    A16-15
Euphorbia clavigera?             GR22        P18        Sl16; FLWR
Euphorbia clavyii                GR0        P    A2-38
Euphorbia clivicola                GR0        P    A16-7

Euphorbia coerulescens        GR24        P    A8-2
Euphorbia coerulescens        GR24        P9    A4-9,10                                        P    A10-33 A13-15,16    
Euphorbia coerulescens             GR24                   P34,36    Apex, Sl11/34-6,                                           P28/8,14 Sl25/9-11, Sl28/8-14                                P25-11;                                             P26-12

Euphorbia colliculina            GR20        P    A10-7
Euphorbia colliculina            GR20        P    SL43-33

Euphorbia columnaris            GR25        P13    A4-13,14,18
Euphorbia columnaris            GR25            A17-x1

Euphorbia confinalis            GR24        P    A11-23
Euphorbia confinalis                GR24        PP         Ann; apices, Sl30/12-15
Euphorbia confinalis               GR24        PP    Jim; apex Sl38/16-18,36-7
Euphorbia confinalis             GR24        P4    H; apex Sl36/4-6

 Euphorbia confinalis rhodesica         GR24        P    A8-36 seedling
Euphorbia confinalis ssp rhodesica     GR24           P42/27       spp. Sl19/24-27, Sl42/27
Euphorbia cooperi ssp calidicola        GR24        P1    A4-1,2

Euphorbia cooperi            GR24                   P1         apex   Sl41/1-2
Euphorbia cooperi?            GR24            A19-31,32
Euphorbia cooperi-like            GR24        P    A3-34        

Euphorbia crassipes            GR15        P
Euphorbia crassipes            GR15        P19    ctg, SL33/19-22, Sl8    , seed 6/2/87  
Euphorbia croizattii                        P3    Sl6/FLWR
Euphorbia cummulate                GR0        P    A9-34,35
Euphorbia cuprisbina                GR0        P    A2-35
Euphorbia curvirama                 GR24                P3,5,36    apices, SL27/1-8, 35-6, rib, Sl27/33 apices, Sl7

Euphorbia decepta            GR15        P    A3-2
Euphorbia decepta            GR15        P    A22-29
Euphorbia decepta            GR15        P37    Sl40/36-7    
Euphorbia deightonii                GR24        P10    A2-9,10 A9-7,8
Euphorbia desmondii                GR21        P13    A3-13,14
Euphorbia didieroides            GrM III        P    Sl35/28
Euphorbia Djibouti Lav. 13176         GR0        P    A14-15
Euphorbia dusimata                GR0        P    A10-15

Euphorbia echinus        GR24        P    A7-38(OR33)
Euphorbia echinus        GR24        P    A4-11
Euphorbia echinus               GR24        P34    Ann; apex, Sl30/34-5
Euphorbia echinus               GR24        P23    H; apices, Sl28/22-4                                P17    Sl28/17&29
Euphorbia echinus                   GR24        P2    Jim; apex 38/2-6
Euphorbia elegantissima            GR0        P    A11-15

Euphorbia ellenbeckii            GR28        P    A10-38
Euphorbia ellenbeckii               GR28             WRONG LABEL

Euphorbia enopla            GR19        P    A8-3
Euphorbia enopla            GR19        P    A9-4
Euphorbia enopla             GR19        P28        Ann; apices, SL30/26-8
Euphorbia enopla            GR19                apex Sl36/28-                                    P2    29, SL42/1-2
 
Euphorbia enormis            GR22        P6    A10-6,8
Euphorbia enormis            GR22        P    Sl31/35
Euphorbia enormis             GR22        P    SL42/18-9            
Euphorbia erlangeri                GR24        P    A14-22
Euphorbia ernestii                GR13        P    A22-30    
Euphorbia esculenta               GR14           P30       flwr, apexSl5/29, Sl6/30+                                     P43-5    Sl7, Sl37/35, Sl43/5-6

Euphorbia evansii            GR24            Sl32    2 bad shots                                P30    Sl36-30
Euphorbia evansii            GR24        P    A15-18    
Euphorbia excelsa                GR21
Euphorbia eyassiana                GR0         P36    apex, brdr Sl31/36-7

Euphorbia fasciculata            GR11        P    A13-22
Euphorbia fasciculata            GR11        P33    apex Sl40/32,33

Euphorbia ferox                GR19        P34    A2-34 A9-3        
Euphorbia ferox                GR19
Euphorbia ferox                GR19        P36    apex/side, Sl33/33-6
Euphorbia ferox?        GR19        P    A7-34
Euphorbia fimbriata                GR19        P23    apex, Sl37/23-4, brdr/tip, SL31/22
Euphorbia flanigani                GR13        P    A19-11,12
Euphorbia fluminis                GR0        P1    A24-38; A5-1, A15-4,5

Euphorbia fortissima        GR24        P19-20    A2-19-20, A2-5 A8-32,33,34-5?, A9-6                    P25-6    A12-24,25,26
Euphorbia fortissima           GR24            wrong label                                        P9           apex/side, Sl8/9,10

Euphorbia fractiflexa        GR0        P6,4    A4-5,6
Euphorbia fractiflexa        GR0        P5           Sl31/4-7, Sl32/23-25
Euphorbia fractiflexa        GR0        P    Sl43-09
       
Euphorbia franckiana         GR21         P             apex/br, SL33/18-19
Euphorbia franckiana        GR21        P    A14-1
 
     Euphorbia friedrichiae        GR15        P    A10-20
Euphorbia friedrichiae?        GR15        P22    Sl35/21-22

Euphorbia fruticosa           GR25a         apex, Sl8/13,                                        P9,11    Sl43/9+11
       Euphorbia fruticosa        GR25a        P25    A1-24,25                                    P    A10-27                                        P    A9-15
Euphorbia fruticosa        GR25a        P    A17-14

Euphorbia fruticosa inermis       GR25a            home
Euphorbia fruticosa inermis       GR25a            P6    Sl4
   Euphorbia furcata                GR0        P    A9-38

Euphorbia galgalana        GR0        P    A13-9
Euphorbia galgalana         GR0        P2    SL34/1-2    vol9/231, B&L.#765
Euphorbia gamkensis                GR0        P1,35    A3-1, A17-34-5
Euphorbia gilettii ssp. gilettii             GR0           P34    apex/side Sl33/33-    34, Sl42/5

Euphorbia globosa        GR17        P    A16-9
Euphorbia globosa               GR17            P35-1    apex Sl35/1-2, SL34/39-9, Sl43/1
Euphorbia grandialata             GR24        P14    SL29/12+ Sl32/17-22      

Euphorbia grandicornis        GR24        P16    A3-15-18 A14-9,10    
Euphorbia grandicornis             GR24     doesn't compare to Sl28/2-5, SL27/34                                    spp, Sl25/16-18, Sl28/31-35                                P14    Sl26/13-15
Euphorbia grandicornis crest     GR24        P32    Sl9 Ann
 
Euphorbia grandidens        GR24        P21    A13-20,21
Euphorbia grandidens        GR24             SL33/23, Sl16; no apex                                 P34    bra tip    Sl7        

Euphorbia graniticola        GR24        P27    A3-27-28                                     P20-1    A12-20,21
Euphorbia graniticola               GR24        P7,8,9    Jim; apex SL38/7-9
Euphorbia graniticola        GR24        P3    H; ctg SL32/3
Euphorbia graniticola        GR24        P31    Ann/apices/rib, Sl30/1-4,31

Euphorbia greenwayii        GR0        P26-7    A1-26,27 A13-6
Euphorbia greenwayii           GR0        P31    E bed, ctg, SL32/28-31
Euphorbia greenwayii           GR0        P26    Jim; apex    SL38/26-7
Euphorbia griseola                GR24        P    A14-21
Euphorbia griseola v. waterbergensis errata    GR24         P7    apex, brdr, SL33/5-9, V9/4
Euphorbia griseola v. hoviea             GR24            P22    apex,    SL29/21-22
Euphorbia griseola v. mashonica         GR24         P23-4    apex,    SL29/23-24

Euphorbia groenwaldii         GR0        P5    A3-5,6 A10-1
Euphorbia groenwaldii?        GR0        P    A20-09    
       Euphorbia guillauminiana            GRMIII        P    A15-25

Euphorbia gymnocalycium    GR0        P    A22-36
Euphorbia gymnocalycium    GR0        P    (Miles)
Euphorbia hamata                 GR6             apex Sl40/29-                                                 P    42-31    31, Sl42/31

Euphorbia handiensis                P    A4-12
Euphorbia handiensis         GR24             P32    apex/brder, SL33/31-2
       Euphorbia hermentian               GR0            A19-36
 
Euphorbia heterochroma         GR24        P19    A9-5                                        P    A15-19,20
Euphorbia heterochroma     GR24        P    Sl7-28
Euphorbia holmesiae                GR0        P    A14-3

Euphorbia horrida        GR19        P    A7-37
Euphorbia horrida        GR19        P    Ann; apex Sl30/19

Euphorbia horrida striata        GR19
Euphorbia horrida v. striata     GR19        PP    apex Sl36/1,12, snowflake                                P32    apex & ribs Sl4, Sl40/4-5,                                P6    Sl5/6, Sl42/4    
Euphorbia horrida type            GR19
Euphorbia horrida noorsveldensis         GR19        P    A9-26
Euphorbia horwoodii;                GR0            
Euphorbia hypogaea                GR0        P15    A4-15

Euphorbia inarticulata        GR24               P12-14    A9-12-14
Euphorbia inarticulata        GR24         P16    apex/sd, SL33/15-17
Euphorbia inconstantia            GR19        P    A8-4
   Euphorbia inequispina            GR28        P    A16-12
Euphorbia inermis hutanae               GR14        P16    SL43/16-7

Euphorbia ingens            GR0        P    A9-37
Euphorbia ingens        GR0        P    Sl26-07
Euphorbia ingens chocolate drops     GR0            A19-37
Euphorbia jensenvillensis            GR18        P18    apex, Sl37/18-19
Euphorbia kalisana                GR0        P    A13-2,3, A15-2,3
Euphorbia keithii                GR24        P25    apex SL29/25, SL34/30-2, Sl42/14    
Euphorbia kibwezensis crest             GR24        P16    apex    Sl5, Sl6                                        P    Sl16 label wrong

Euphorbia knuthii        GR22        P    A12-3,4,5
Euphorbia knuthii        GR22            H; apex/sd                                    P15     SL33/13-15
Euphorbia knuthii        GR22        P    Jim; apex, s39/28-9

Euphorbia lactea            GR24        P14    A1-14,15 A4-7,8
Euphorbia lactea            GR24        P11    apex Sl37/10-15, SL42/33-34 label wrong
Euphorbia lactea crest         GR24        P10-4    Ann; Sl13/10-14                                    P11    Sl7

Euphorbia ledienii        GR24        PP    A12-16-18
Euphorbia ledienii         GR24               apices, rib, plants:                                       P24,26-7    SL27/23-31, s39/34-7                                P30    apices, side   Sl7/30
Euphorbia ledienii crest        GR24            Sl19
   Euphorbia leistneri                GR0        P19    A4-19,20
Euphorbia leucodendron                 GR4        P24    Sl6
 Euphorbia longispina                GR24        P16    A16-16-18
 Euphorbia lowii                 GR0        P2,3,4    A5-2-5 A11-10,11    
 
Euphorbia magnacapsula        GR0        P20    (A4-3,), A1-20, A2-14
Euphorbia magnacapsula         GR0             apex/tip, SL31/23                                    P34    Sl27/34, Sl28/03
  Euphorbia magnacapsularis            GR0        P16    A1-16,17
 
Euphorbia makallensis        GR0        P    A14-12
Euphorbia makallensis?          GR0                   P25/6    Sl11/20-21, Sl28/25-6
Euphorbia malevola                GR0        P29    A2-29,30                                        P13    A12-13,14

Euphorbia mammillaris           GR19            P12,13,10   Jim; apex Sl38/10-11
Euphorbia mammillaris         GR19         P    SL42/8, SL43/18
Euphorbia mammillaris        GR19        P    A2-3
Euphorbia mammillaris        GR19        P2    A19-1-4
Euphorbia mammillaris variagata         GR19        P    A10-28
Euphorbia marlothiana            GR14            apex Sl37/36
Euphorbia marsabitensis            GR0        P36-7    A2-36-7                                            P29    A9-28-30

Euphorbia mayuranathani        GR0        P31    A3-31,32        
Euphorbia mayuranathani     GR0        P14-5    apex, side: Sl11/14-15, Sl32/5- 6, Sl40/26-8    
 Euphorbia melanohydrata            GR15        P    A10-21
 Euphorbia meloformis            GR18        P    A10-29
 Euphorbia meloformis falsa            GR18        P    A16-1
 
Euphorbia memoralis        GR24        P    A12-6, 7
Euphorbia memoralis        GR24        P13    Sl34-13 SL42/15
Euphorbia meridionalis            GR0        P    A13-17
Euphorbia micrantha                GR0        P    A3-5
Euphorbia miguritinorum            GR24        P4    A13-4,5
Euphorbia milii                   GRM.VI        P11     Sl4                                            P10    Sl5
Euphorbia milii bojeri                GRM.VI        PP    SL42/22-23
Euphorbia mitriformis                GR0        P37-8    A10-35-7; A16-2
Euphorbia monacantha            GR27        P9,10    A15-9,10

Euphorbia monteiri            GR9        P20    apex Sl40/22, Sl42/20
Euphorbia monteiri            GR9        P7    Sl16 Sl19/1-6 blossom
Euphorbia mosaica                GR0        P4-23    A1-1; A4-23
Euphorbia mozambiquensis            GR0        P    A13-23
Euphorbia Mrs. Ash                GR0        P    A4-24
Euphorbia multiceps                GR15        P    A9-20
Euphorbia multiramosa            GR16        P    apex Sl37/22
Euphorbia namibensis                GR15           P        
Euphorbia nelii                GR15        P    A9-21

Euphorbia nerifolia        GR21        P7,8    A2-7,8
Euphorbia nerifolia         GR21         P21    Sl25/21-22, new tip, spp @lf base                        P9,13    spp, br tip, Sl11/9-13
Euphorbia neriifolia        GR21        P    Jim; apex S39/27
Euphorbia nesemanii?            GR19        P    apex     Sl41/10
 Euphorbia nigrispina                GR24            A1-12,13                                        P27    A2-26-8
Euphorbia nivulia crest            GR21        P    A13-1
Euphorbia nubigena                GR0        P    A11-9
Euphorbia nutrufirnus?            GR0
Euphorbia obesa Ann             GR18        P22    apex, Sl30/22, see symmetrica, Sl30/23
Euphorbia odontophora            GR0
Euphorbia opuntioides            GR24        P    A11-24
Euphorbia ornithopus             GR0             apices,    SL33/37-8
Euphorbia pachypodiodes            GRV        PP

Euphorbia parciramulosa        GR24        P    A16-14
Euphorbia parciramulosa         GR24         P29    apices,    Sl29/26-7, Sl32/12, SL42/17
Euphorbia peltigera                        P    Sl43-22    
Euphorbia pentagona                GR19        P    A10-26
Euphorbia pentops                 GR15        P    A11-1
Euphorbia perangusta                GR24              P12,13    A15-11-14

Euphorbia persistentifolia     GR24                     Ann; apex SL30/31
Euphorbia persistifolia        GR24? spp    P    A9-11

Euphorbia petraea        GR0         P    ctg apex/brdr, Sl31/33-34
Euphorbia petraea        GR0        PP    A11-18,19
Euphorbia pettigera (S&Cv2)             GR0            Sl42/30, Sl43/22
Euphorbia pfersdorffii                        Sl49
Euphorbia phillipsae                GR24        P    A1-2
Euphorbia phillipsioides            GR0        P    A3-8
Euphorbia pillansii                GR19        P24    A10-23-5

Euphorbia polyacantha        GR24        P18    A1-18,19
Euphorbia polyacantha        GR24        P    A19-30
Euphorbia polyacantha v. rosenii         GR24              P9,12-3    apices, SL27/9-13                                        P37    rib, Sl27/32,37                                             P19,21-2    apices    Sl7

Euphorbia polygona        GR19        P    A2-11
Euphorbia polygona        GR19        P8-35    A7-35,36(orA8)
Euphorbia polygona         GR19        P4    Jim; apex s39/4,5
Euphorbia polygona        GR19        P    A9-23,24, /horrida snowflake
Euphorbia proballyana            GR24        P32    A9-32,33

Euphorbia pseudoburuana        GR0        P5    A2-4,5                                        P    A5-6
Euphorbia pseudoburuana     GR0            apex, brdrs                                    P17    31/17,18,33,34    

Euphorbia pseudocactus        GR24        P6    A1-6 A9-3
Euphorbia pseudocactus        GR24            Sl33/29; apices:                                    P38    Sl28/38,                                        P1,12    29/1,2,12, SL42/13,16
Euphorbia pseudocactus        GR24        P25    Jim; apex, S39/22-26

Euphorbia pseudocactus lyttoniana     GR24        P15-6    A15-15-17
Euphorbia pseudocactus lyttoniana   GR24        P     apex, SL38/15    
Euphorbia pseudoglobosa            GR18        P21    A4-21,22
Euphorbia pulvinata                        Sl49
Euphorbia quadrangularis            GR28        P33    A2-31,33 A11-12
Euphorbia quadrispina            GR0        P    A11-3
Euphorbia quinquecostata            GR24        P13    A13-12,13
Euphorbia ramiglans                GR14                P43-30    SL43
Euphorbia reinhardtii                GR24        P38    apex, Sl7/38, Sl8/1,2

Euphorbia resinifera        GR24        P-31    A1-31,32
Euphorbia resinifera        GR24        P11-2    A7-10-12
Euphorbia resinifera        GR24              P18    apex, SL28/18
Euphorbia restituta                GR11              P23    apex Sl40/22-3

Euphorbia restricta        GR22        P7    A1-7,8,9 A4-27,28, A8-31
Euphorbia restricta clone #1     GR22         P29    apex SL29/29
Euphorbia restricta?        GR22                   SL35/11-13
Euphorbia richardsae robustior            GR22        P6,7    A11-6,7        
Euphorbia richardsiae v. richardsiae         GR22           P38    brdr/tip Sl31/38-9
Euphorbia robecchii                GR24        P13    A14-13,14
Euphorbia robusta                GR0        P2    A12-1,    (2?)
   Euphorbia robustior                GR0            

Euphorbia royleana        GR21        P    A2-32
Euphorbia royleana        GR21        P    A8-1
Euphorbia royleana        GR21        P    A20-04
Euphorbia royleana         GR21        P30    apex s39/30, Sl40/28, Sl42/32
Euphorbia rubechii                GR0        P    A11-2    
Euphorbia rubrispinosa            GR0        P    A14-5
Euphorbia s.n. Zimbabwe            GR0        P10    A1-10,11
 
Euphorbia samburuensis        GR24        P6    A15-6,7
Euphorbia samburuensis        GR24            Sl24; flwr
   Euphorbia saxorum                GR0        P    A11-22
   Euphorbia schinzii                GR22        P5    A11-4,5

Euphorbia schoenlandii?        GR11        P    A19-33,34
Euphorbia schoenlandii           GR11        P    Ann; apex, Sl30/25
Euphorbia schoenlandii           GR11        P18    H; apex SL40/18-                                    P20    21, Sl43/20
Euphorbia sizacantha                GR0        P15    A2-15; A10-34
Euphorbia seibanica                GR0        P    A22-11

Euphorbia sekukuniensis        GR24        P21    A1-21-23
Euphorbia sekukuniensis        GR24        P25    Sl36/25
Euphorbia septentrionalis            GR0        P12,24    A2-12,13 A14-24

Euphorbia sepulta        GR0        P    A3-7 A4-16,17    
Euphorbia sepulta clone 1     GR0            SL28/29    looks like echinus
Euphorbia serendipita                GR0        P11    A13-11
Euphorbia shizacantha             GR0        P15    A2-15
Euphorbia spp. alverton, S.A.            GR0        P    A11-8
Euphorbia spp. angola            GR0        P    apex, SL29/28
Euphorbia spp. Nova                 GR0        P    Greystown Natal, squarrosa type                                            ctg SL33or34
Euphorbia Spp nova                GR0        P    SL34/36-7
Euphorbia spp. nova                 GR0        P    Sl35/5-6

Euphorbia squarrosa        GR22        P10-4    A9-18; A10-4,5
Euphorbia squarrosa         GR22            aff. fanshawii
Euphorbia squarrosa        GR22        P    Sl24-09
Euphorbia squarrosa        GR22         P38    Jim; apex Sl38/38, s39/1
Euphorbia ssp. riou-brevirama         GR0            apex Sl5/27

Euphorbia stellaespina        GR19        P    A9-25
Euphorbia stellaespina hyb    GR19         w/mammillaris Edward Hummell                                 A19-8-10
Euphorbia stellaespina         GR19             apex: SL29, Sl16/25, Sl36/                                P26    18-19, Sl42/21+26                                    P    Sl16

Euphorbia stellata, GM2005Mother    GR22        P    A10-2, A16-03
Euphorbia stellata        GR22            Sl21
Euphorbia stellata        GR22        P21    Ann apices, Sl30/20-21
Euphorbia strangulata                GR0        P22    A3-21,22                                         P32    A4-30-34; A14-18
Euphorbia submammilaris             GR19           2nd spp     SL33/1-2, apex SL32/34-35
   Euphorbia subsala                GR0        P13-4    A11-13,14
Euphorbia supernans                 GR14        P31    apex Sl37/31-32
   
Euphorbia supressa        GR0        P    A10-3
Euphorbia supressa        GR0        P    A22-28
Euphorbia supressa           GR0        P    SL43/7

Euphorbia susannae crest    GR18            Sl21, Sl22
Euphorbia suzannae crest    GR18
Euphorbia symmetrica            GR18        P    Ann apex SL30/23
Euphorbia taruensis ctg               GR0         P    apices SL32/34-5, marked obtusifolia
Euphorbia teixeirae    Ann            GR0        P7    apices,    Sl30/5-7
Euphorbia tirucalli?                GR4        P    Sl42/35

Euphorbia tortilis            GR24        P21    A2-21-24
Euphorbia tortilis            GR24        P    A4-4 A3-12 A10-32
Euphorbia tortilis            GR24        P5,7    A19-5,6,7
Euphorbia tortilis            GR24        P21    apex SL31, Sl43/21

Euphorbia tortirama        GR22        P    A16-8
Euphorbia tortirama        GR22             P10,13-4   apex Sl16/10-14    apex, spp; Sl20/10-14,                                                                    P11    Sl36/18-9, Sl42/11-12
Euphorbia tortirama        GR22        P29    Ann apices Sl3029-30
Euphorbia trapifolia                GR0        P35    A3-35,36,37 A11-25
Euphorbia triaculeata                GR27        P8-10    A12-8,9,10

Euphorbia triangularis        GR24        P    A13-10 A15-1
Euphorbia triangularis        GR24                   SL32/30-32, SL33/10-12                                Sl43/11?/ kamerunica apex?                                P6     Sl8/4-6
Euphorbia tridentata                 GR17            SL34 label wrong    ctg                                         poss friedrichiae

Euphorbia trigona             GR24        P21    Jim; apex s39/20-21
Euphorbia trigona         GR24            apex s39/32-                                            P43-13    33, SL43/12
Euphorbia trigona rubra        GR24        P    A19-35
Euphorbia trunk, big house, near back door, bursting pot        Sl20
Euphorbia tuberculata             GR14                 P42-29    Sl35/19-20, Sl42/28-9                                    P14    Sl5, SL43/13+ SL34                                    P4    Sl7
Euphorbia tuberculatoides               GR14        P24    Sl5/

Euphorbia tubiglans        GR18        P10    A9-9,10
Euphorbia tubiglans        GR18        P24-5    SL34-24,25
Euphorbia turbinicarpus            GR0        
Euphorbia turbinicarpus pseudo pectinatus     GR0
Euphorbia turbiniformis            GR18        P    A8-24
Euphorbia twinkle twirls            GR0            SL36/38, SL37/1
Euphorbia uhligiana                GR28        P23    A14-23
Euphorbia unispina                 GR26                  P20,24    apices, Sl16/20-24                                    P38    SL40/37,38, SL43/10
Euphorbia utoniensis             GR0           P27    SL33/26-28
Euphorbia vajravehii                GR0        P    A14-17

Euphorbia valaris        GR0        P9    Ann, apices, SL30/8-11
Euphorbia valaris            GR0        P    A13-14 A9-1,2

Euphorbia valida            GR18        PP    A3-9, A10-30
Euphorbia valida            GR18            Ann; female br1a; Sl30/24
Euphorbia valida?          GR18            P31         Sl43/26+31
Euphorbia vandermerwei             GR22        P    A16-13

Euphorbia venenata        GR24        P29    A8-26(27-9?)
Euphorbia venenata        GR24             P15    apex Sl40/14-15                                    P3    Sl41/3-9, Sl42/36
Euphorbia viduiflora                GR0        P    A12-19
Euphorbia viguieri vilanandrensis        GR2        P22    A15-22,24

Euphorbia virosa            GR24        P7    apex     Sl7
Euphorbia virosa    Ann        GR24            P32    apex; SL30/32-33
Euphorbia waterbergensis             GR24        P21    Sl35/20-4
Euphorbia wild spp                GR0        P14    A22-15
Euphorbia williamsonii                
Euphorbia wilmaniae                GR17        P    SL32-01
Euphorbia woodii             GR13                  P43-19    apex: SL28/19, Sl43/19
Euphorbia xylacantha (P. Bisseret)         GR0        P    A22-08
Euphorbia xylophylloides            GR4        P    Sl19-13

Euphorbia zoutpanbergensis    GR24        P6,7    A14-6,7,8
Euphorbia zoutpansbergensis    GR24         P10    apex SL28/10-13Sl8/7,8    


S5 Euphorbia sort criteria
Prepared by Susan Dunlap

Flower & Stem
Flower & Stem: flower residue/peduncle present
Flower & Stem: stem color green
Flower & Stem: stem color grey-green
Flower & Stem: stem color light green
Flower & Stem: stem color light grey
Flower & Stem: stem color light grey-green
Flower & Stem: stem color light red
Flower & Stem: stem color variegated
Leaf
Leaf: leaf dry state persistent
Leaf: leaf scar large & conspicuous
Leaf: leaf present
Leaf: leaf quickly deciduous
Spine
Spine: branching spine present
Spine: one prominent spine present
Spine: two spines present
Spine: three or more spines present
Spine: spine color distinct from spine shield
Spine: spine tip contrasting color
Spine: spineless
Rib
Rib: rib edge continuous horny
Rib: rib present
Rib: rib wavy
Rib or Tubercle: valley deep groove
Rib or Tubercle: valley impress line
Rib or Tubercle: valley smooth


S6 Euphorbia Comparison of photographs: overview
Prepared by Susan Dunlap

Spps from different collections w/marked similarities or accountable differences:

abyssinica
actinoclada
albipollinifera
ammak
antiquorum
arida
canariensis
coerulescens
cooperi
echinus
enopla
fruticosa
gamkensis
globosa
graniticola
greenwayii
gymnocalycium
handiensis
knuthii
lactea
makallensis
mammillaris
mammillaris var.
nerifolia
parciramulosa
polygona
pseudoburuana
pseudocactus
pseudocactus lyttoniana
resinifera
restricta
royleana
schoenlandii
sekukuniensis
squarrosa
stellaespina
supressa
tortilis
tortirama
triangularis
trigona rubra
valida
venenata


Species from different collections with unaccountable differences:
(issues with labeling, photograph, maturity of stem/branch, etc.)

avasmontana    labeling/variety
barnardii    labeling
colliculina    labeling
confinalis    impact of grower
evansii         labeling/photo
fractiflexa     labeling
griseola         br vs stem
groenwaldii     br vs stem
heterochroma    
horrida        male vs female/variety
ingens         br vs stem
ledienii         labeling
magnacapsula     br vs stem
petraea        photo
tuberculata    br vs stem
tubiglans    photo
valaris         br vs stem
virosa         br vs stem
zoutpanbergensis    br vs stem




S7 Description of general and apical traits belonging to Euphorbia pulvinata, E. canariensis, E. coerulescens, and E. galgalana.
Prepared by Susan Dunlap

Note:  A rib refers to a single growth cycle. A single growth cycle is also referred to as a plastochron, a term used in discussions of a plant’s meristem.

Euphorbia pulvinata                
Young plant was examined that had a primary stem with seven branches emerging around it at the base. The stem had 9 ribs with a prominent leaf scar (1.5mm wide) at the end of each tubercle. The stem diameter NIC ribs = 35mm; rib diameter = 55 mm. Tubercles emerged in roughly the following pattern: ribs 1,5,8 emerged nearly simultaneous with ribs 3,6,9, and 2,4,7.

The center of the apex was depressed 5mm from the top of the second growth cycle of tubercles. The leaf scar was comprised of dry white material with a brown scab forming a ring around the base of the scar. There were no spines. A thin, brown, woody, peduncle 1.5mm below the leaf scar was present.  It was 3-5mm long, emerged out of the flowering eye, and persisted irregularly. The rest of the plant is fleshy and is a medium hued green.

At the apex, the leaf scar was situated at the very top and center of a tubercle.  As the plant grows, the position of the leaf scar migrates 135 degrees.  Initially the leaf scar faces upward and migrates to face the ground within 4-5 growth cycles.

When viewing the apex, the leaf scar was visible for 3 growth cycles and created a concentration of white circles. The ribs grew in a symmetrical, radiating pattern.  At 25mm below the apex, the rib pattern shifted to a half spiral and then resumed the symmetrical radiating pattern. This shift, that occurred 5-6 growth cycles below the apex at a point at which 2 new ribs emerge, contributed to the complexity of what was visible at the apex.
 
At the apex, each rib was 10-11 mm deep and 4 mm wide. The leaf scar and prickle were positioned on edge of the rib. The spacing between the leaf scars was 8mm. At the center of the apex, the ribs were separated by 5mm. This gap was occupied by the emerging ribs and dominated by round white leaf scars. There is no fleshy green stem material visible in this gap.

The tip of the tubercle separates itself from the main body of the rib soon after it emerges at the apex. This separation persists as the plant grows, forming a distinct pattern down the ridge of the rib. Each tubercular unit becomes more distinct as the plant grows, forming an undulating pattern down the sides of the rib. This pattern is visible by the time a new rib emerges (in this case, 5-6 growth cycles). The ridge of the rib is where this distinct pattern occurs, containing the leaf scar, the peduncle, and the tubercle itself.

Euphorbia canariensis
This plant had one stem and two branches, is a recently rooted cutting. The cutting is 14" high; one branch is 5 1/2" high and the other is 2 1/4" high. The shorter branch is 31 mm diameter including the ribs and 13 mm diameter not including the ribs.  The stem is concave between the ribs. It is obscured from view at the apex.

New growth at the apex of smallest branch contains several features. Initially, the spine shield emerges from the top of the apical dome and separates itself from the depressed gap in the center of the apex. Most of the fleshy green epidermis between the ribs is hidden or obscured by spine shields. There is a slight depression at the center (4mm between the lowest point and the top of the youngest spine shield), and a gap between emerging spine shields (4mm). There are 6 ribs. The alternating rib pattern of emerging spines is rib #'s1,4,6, 2,5,3; 1,4,6,2,5,3, etc.

The spine shield is wider than the ridge of the ribs. The ridge is 2 mm wide. The largest spine shield is 4 mm wide. The ridge of the rib, between the spine shields, is "U" shaped.  Overall, the spine shield is somewhat triangular in shape. It attaches to the stem without any other markings. New growth of both the spines and spine shield is reddish brown. At the apex of old branches, the spine shield is tan and the spines are dark brown. As they age further, both the spine shields and spines become light gray. These color changes are not visible from an apical view of an actively growing branch. The profile of the ridge between the spine shields is "U" shaped.  There is a small slit or flowering-eye in-between the spine shields where, on a mature plant, a flower might emerge. This plant has never flowered.

An inward-pointing, semi-triangular leaf, attached to the spine shield, is visible in the first life cycle within the apical dome. It is pronounced for 2 growth cycles, shrivels by the third growth cycle, and disappears within 5 growth cycles. A tiny horizontal impress line remains at the original point of attachment.

A symmetrical pair of spines emerge almost fully grown, are nearly perpendicular to the rib (135-170 degrees apart), and are 7-mm long. For one growth cycle they are the same color as the spine shield. There is a small ridge between them where they meet in the center of the spine shield.


Euphorbia coerulescens apex:
Dormant stem, recently rooted, 4.5 cm diameter, 12 cm high. The concave portion of the stem is 1.9 cm in diameter. Top segment is 4.5 cm high.

The apical footprint forms a circle that is concave between the ribs; the ridge of the ribs in the apical region is 4-mm wide. There are 5 ribs on this plant. The fleshy part of the stem is a medium-hued grey-green; each rib is separated by a light brown dry border that runs the length of the rib.  This border exhibits a slight twist as it grows, but does not spiral. The fleshy part of the stem appears to swell very slightly along the ridge at the intersection between the fleshy part and the dry ridge. Actually, the fleshy part stays the same width as it is the dry elements that vary in width on top of it. The color of the dry ridge does not very along its vertical edges. It is the same color up to the point that it attaches to the fleshy stem. Within this border are pairs of protruding spines; the youngest pair of spines are 1-mm long at the center of the apex growing to 9-mm long at the periphery of the apical circumference.  The tips of the spines are a very slightly darker brown than the spine border.    

The dry border varies in width: below the spines the border narrows to 1-1.5-mm; above the spines it narrows to 3-mm and terminates at a horizontal row of fleshy grey-green colored ovals, 3 each 0.5-mm wide. As the plant matures, these ovals become surrounded by a thin, dry ring.  

Inserted 1.5-mm above the spines is a tiny pair of prickles, appearing as small brown dots. They protrude no more than .25-mm.

At the center of the apex, there is a 3.5-mm gap between the new sets of spines. It is divided into the same number of segments as there are ribs, a feature somewhat difficult to see with the naked eye. The material in this gap is distinct from the stem and the spine border. It appears dryer than the stem and fleshier than the spine border. Overall it is musky green. The gap in front of the next rib poised to grow a spine-pair is a slightly darker hue than other ribs contained in this area of the apex. The spines emerge in fixed sequence – every other rib: 1-3-5-2-4, 1-3-5-2-4, etc.

The center of the apex is slightly depressed. The youngest spine (spine pair 1a on rib 1) is more depressed than the older spine pair on the same rib (spine pair 1b on rib 1). The next oldest spine pair (2a) is level with the older spine pair on the same rib (2b). The next oldest spine pair (3a) is higher than the older spine pair on the same rib (3b), as is the case with ribs 4 and 5.  Rib 4a is higher than 4b, and 5a is higher than 5b. The overall appearance of the top, then, is relatively flat.


Euphorbia galgalana:
Plant is a recent cutting with a light green stem and two young branches. The stem and branches each have 5 ribs. The stem diameter is 24-mm including the spines, 3.5-mm at the apex not including the spines or ribs, and is 6-mm at 3 growth cycles including ribs. The spine tips are 60 degrees apart. The flowering eye is a protruding red dot above spine shield that is 1-mm wide and 0.5-mm high. The depression at the apex is insignificant.

The spine shield is a metallic grey color with a hair-line border of brown around its perimeter.  The spine shield is 5-7mm long and is wider at the top in the area where the spines emerge.  It is 2mm wide at the top and 0.8mm wide at the base.

The rib growth sequence is 1,4,2,5,3, 1,4,2,5,3, etc. at the apex.

The insignificant leaf scar disappears quickly. It reads as a dry line of material at the top edge of the spine shield. On a more robust and actively growing stem, the leaf is very slightly more conspicuous, but both these observations were made using a magnifying lens. The position of the leaf scar does not shift.

On this specimen the cross section of the stem is not perfectly cylindrical but undulates at insertion of each rib. On the main stem, the ribs spiral very slightly. In a span of three inches one rib migrated 90 degrees counterclockwise. This mild spiraling contributes to the number of spine pairs visible in a photograph of the apex. The rib profile forms an inverted “L” shape.  There is a 2-mm deep ridge at the top of the spine shield that tapers down the length of the spine shield.  There is 8-mm between the top of one spine shield and the top of the next, with a 2.5-mm gap between the bottom of one and the top of the next. In the apical view, this ridge contributes to the visibility of the flowering eye for more than one growth cycle.


S8 Euphorbia Descriptions, a small selection by others recorded in specialized publications.
Compiled by Susan Dunlap

Jacobsen on Euphorbia canariensis and E. coerulescens
Jacobsen, Hermann F.  Lexicon of Succulent Plants. London, Blandford Press Ltd., 1974.

Group 24. (Sect. Euphorbia. - 5. Trigonae Bgr. and 6. Polygonae Bgr.; Keys 18 and 19 according to White, Dyer and Sloane). - Shrubby and arborescent succulents 30 cm up to more than 3 m high; Br. developed at the base of a +_ shortened main St., with or without side-shoots, or with an erect trunk, often divided into two or more main St.; Br. with or without flowering Br. and lateral shoots which are 3-13 angled, trunk always many-angled, +_ roundish when old; Th. short or up to 5 cm long; Th.-shields with pairs of Th. solitary or confluent; L. large, later falling (Type s. Pl.63)
    E. canariensis L. v. canariensis (Pl. 67/3) (Gr 24). - Canary Is. - (shrub) branching from the base, up to 12 m tall; Br. numerous, ascending, (4-)5(-6)-angled, fresh green, sides flat, angles acute, sinuate-tuberculate, Th. pairs c. 14 cm apart, 4-5 mm long, thin.
    E. coerulescens Haw. (Pl. 69/1) (Gr. 24) (E. virosa Boiss. p. part., E. V. V. C. Bgr.). - Cape: Jansenville, Steytlerville, Uitenhage D. - Thorny, succulent (shrub), spreading below ground and with numerous erect ST. up to 1.5m tall, forming broad bushes; St. 3-4 cm thick, often branching above, segments round, 4-6 angled with very concave sides, grey-blue, angles sinuate-tuberculate with a brownish horny band.  Th. in pairs, 6-12 mm long, stiff, dark brown.

White, Dyer, & Sloane
White, Dyer, Sloane. Succulent Euphorbieae (Southern Africa), Vols. I & II. Pasadena, CA., Abbey Garden Press, 1941.

on Euphorbia coerulescens
Vol. I = keys
Vol. II=species description
Key 18
Shrub-like succulents with spine pairs
I. Branches slender, usually 1 in (2.5cm) or less thick; cymes single from each of the flowering eyes, consisting of 3 cyathia horizontally disposed:
A. Ovary exserted from the involucre:
        1. Branches not constricted into segments, 4-6 (often 5)-angled
griseola
2. branches slightly constricted at distant intervals, 4 or 5-angled.
heterochroma
B. Ovary sessile or subsessile:
1. spine shields united into a continuous margin along the angles; branches 4-angled, up to 0.6 in. (1.5cm) thick
lydenburgensis
2. spine shields separate, decurrent:
a.  spines 2-6 lin (4-12mm) long, with prickles 0.5-32 lin. (1-4mm) long; branches 4-angled, usually less than 0.4 in (1 cm) thick
complexa
b. spines 2-2.5 lin (4-5 mm) long, with prickles 1-1.2 lin. (2-2.5mm) long; branches apparently 4-6-angled, 0.8 to 1.2 in (2-3cm) thick when dried
subsalsa
II Branches over 1 in, (2.5 cm) thick and usually up to 2 in (5cm) or more thick; see also E. subsalsa:
A. Cymes single from each of the flowering eyes, consisting of 3 cyathia vertically disposed:
1.  Branch segments 5-8-angled, broadest about the middle part; capsule globose with a fleshy covering
virosa
2.  Branch segments 4-5-angled, broadest below the middle part; capsule subacutely 3-lobed, drying on maturity.
barnardii
B.  Cymes 3 (or rarely variably 1 to 4) from each of the flowering eyes, each normally consisting of 3 cyathia vertically disposed:   
1.  Branch segments up to about 2 in (5cm) thick at the widest portion, rarely more:
a. Spine shields united into a continuous margin up to about 1.5 lin (3mm) broad and more or less uniformly broad throughout their length:
i. Capsule sessile or subsessile; branch segments usually broadest below the middle part
    venenata
ii. Capsule exserted from the involucre on a slender pedicel:
+ Branches constricted into symmetrical segments, which are broadest about the middle part
avasmontana
++ Branches not or only very slightly constricted into segments
    hottentota
b.  Spine shields separate or irregularly united into a continuous margin, but when continuous then contracting from each spine pair to the flowering eye below and not uniformly broad:
i.  Capsule 3-3.5 lin. (6-7mm) in diameter, exserted on a slender pedicel; sides of branches with yellowish-green marking:
+ Branches 5-7-angled, with a slender central core and very thin wing-like angles; spine shields united into a continuous or rarely interrupted margin
    perangusta
++ Branches usually 5-angled, with compressed angles which are not deeply wing-like; spine shields separate, though sometimes so close together as to appear continuous.
    knobelii
ii.  Capsule about 6 lin (12mm) in diameter, subsessile; sides of branches green or green with yellowish-green markings
    pseudocactus
ii. Capsule 5 to 6 lin. (10-12mm) in diameter exserted on a slender pedicel:
+ Plants spreading by means of rhizomes; branches glaucous, with the segments usually rounded in outline and the margins often wavy
                    coerulescens

Vol. II text on Euphorbia coerulescens:
(Frequently quoted by Jacobsen; S. Carter subsequent world authority)

"Plant: a spiny succulent shrub, branching from the base and multiplying by means of rhizomes, which spread underground and give rise at intervals to stiffly erect young plants; branches numerous, 0.7 to 1.5 m. or more high, 3-5 cm. thick, markedly constricted into rounded oblong or elongated segments 3.7-10 cm. long, 4-6 angled, with slightly concave sides, more or less glaucous, dark green, generally sparingly branched, the spreading secondary branches produced in clusters or somewhat whorled; angles sinuate-tubercled, with continuous or rarely interrupted horny margins, which at first are pale brown, finally becoming grey;

Spines: in pairs, 6-12 mm. long, rather stout, diverging, dark brown;

Inflorescence: cymes  produced near the apex of the branches, 1 to 3 together from each flowering eye, peduncled; the cymes consisting of 3 cyathia vertically disposed, the central one male and the 2 lateral ones bisexual; peduncles 5-6 mm. long, forking at the apex into 2 cyme branches about 3 mm. long, bearing 2 bracts just below the sessile central involucre at the fork and 2 bracts at the apex of each cyme branch just below the lateral involucres; bracts at the fork of the peduncle short, broadly ovate, those at the apex of the cyme branches sub quadrate, all with a few minute cilia; involucre somewhat cup-shaped, 5-6 mm. in diameter across the glands, glabrous, with 5 glands and 5 sub quadrate, fimbriate lobes; glands about 2.5 mm. in their greater diameter, transversely oblong, bright canary yellow;

Pistillate flower: ovary pedicelled; styles 3 mm. long, united for half their length, free above, with bifid tips:

Capsule: exserted from the involucre, 3-angled."

S. Carter on Euphorbia galgalana
Euphorbia Journal, Vol 9 pg. 231
Schwartz, Herman, ed. The Euphorbia Journal, Volumes 1-10, Mill Valley, CA, Strawberry Press, 1983-1996.

Related to both Euphorbia nigrispina and E. geldorensis, E. galgalana is a succulent perennial, much-branched from the base, forming sprawling clumps 15-60cm high and to 1 m in diameter.  Branches spreading, terete, 4-5-angled, 1-1.5cm thick; angles very shallowly toothed, with tubercles 8-10mm apart.  Spine-shields quite separate, oblong-ovate 3.5-8x2-3.5mm; spines paired, mostly 1-1.5cm long, slightly recurved; prickles vestigial or apparently absent.  Leaves deltoid, 0.5x0.8mm, quickly deciduous.  Flowering eyes immediately above the spine-shields; cymes solitary, subsessile, 1-forked; bracts ovate, 1x1mm, margin minutely denticulate. Cyathia 2.5x3.5mm, funnel-shaped; glands 5, transversely oblong, 0.8x1.5mm, spreading touching, bright yellow; involucral lobes 5, rounded, 0.8x1.2mm, margin denticulate. E. galgalana differs from E. nigrispina in its spreading instead of erect branching, from E. geldorensis in its 4- or 5- instead of 6-angled branches, and from both in its very distinctly separated spine-shields and longer, curved spines of fairly uniform length. Inflorescence characters however, like E. geldorensis, are almost identical to those of E. nigrispina.  

Jacobsen on Euphorbia pulvinata (Group 19)
"Branching basally; BR. numerous of equal length, forming a low and lightly convex cushion up to 1.5m (0), 3-6 cm tall, 3-4 cm (0), with (6-)7(-8) slightly crenate angles 7-9 mm high; Ped. numerous along the angles, wine-red or purple-brown, becoming thorny, 10-15mm long."

Jacobsen’s statement about all species in Group 19:
"Dwarf or shrubby succulents, partly with tuberous roots; the main shoot as well as the Br. cylindrical with tubercles (L.-bases) arranged in longitudinal rows often forming areolate angles; angles 6-18, +- prominent; sterile Ped. persistent as Th., rarely falling."



S9 Ferocactus Descriptions
Compiled by Susan Dunlap

Britton and Rose
Anderson
Lindsey
Dunlap

Britton and Rose 1922
barrel cactus, Bisnaga
Bisnaga Orcutt 1926
Plants solitary or branched, often becoming large. Stems depressed globose to globose to cylindrical. Ribs few to many, often large and prominent. Areoles usually large, bearing flowers only when young, with nectar-secreting glands. Spines variable, usually heavy, sometimes hooked. Flowers borne near the stem tips, solitary, both female and male organs present, radially symmetrical, short funnel-form, funnel-form, or bell shaped, with conspicuous scales; areoles of pericarpels and floral tubes naked; perianth parts and stamens separated by a ring of hairs. Fruits globose to oblong, thick walled, dry or juicy at maturity, dehiscing by basal pores or irregular slits. Seeds oval, shiny blackish brown, flat to slightly concave to somewhat pitted, 1.4-2.4mm long. Distribution: arid and semiarid regions of the southwestern United States and northern and central Mexico, particularly well represented on the peninsula of Baja, California. Ferocactus is often divided into section Ferocactus, with dry fruits dehiscing basally, and section Bisnaga, with juicy and indehiscent or irregularly dehiscent fruits. First collected in early 18th C. (1700's). The most thorough analysis of Ferocactus was by George Lindsay in 1955. Two other important contributions have been made by Nigel Taylor (1984) and Hugo Cota and Robert Wallace (1997). Closely related to Echinocactus in having stem tips that are not densely woolly. The relationship of Stenocactus to Ferocactus has also been debated.  Arthur Gibson (1992, 67) suggested that Ferocactus might also be closely related to the North American columnar cacti. The research by Cota and Wallace, however, using DNA sequencing, has shown that Ferocactus is not closely related to the columnar cacti. The relationship to Echinocactus, however, is far less clear, and Cota and Wallace's conclusions is that both Ferocactus and Echinocactus evolved from an Echinocactus-like ancestor. Some realignment of species in the two genera was necessary and is reflected in the treatments of them here. Ferocactus comprises 29 species of globose top cylindrical cacti, sometimes branched, producing radially symmetrical bee-pollinated flowers that bear delta-shaped to rounded scales and that have the stamens separated from the perianth parts by a ring of hairs. Most also have reduced gland-like spines in the upper part of the areole. The cacti flower during the day in spring and summer.

Anderson typical entry for species:
Ferocactus latispinus (Haworth) Briton & Rose 1922
Pochas (referring to the fruits)
Cactus latispinus Haworth 1824, Bisnaga recurva subsp. latispina (Haworth) Doweld 1999.
Cactus recurvus P. Miller 1768, rejected name; Ferocactus recurvus (P. Miller) Borg 1937; Bisnaga recurva (P. Miller) Doweld 1999, not validly published (etc.). Plants solitary, depressed globose to flattened, light green to 30cm (12in) high and 40cm (16in) diameter. Ribs about 21, acute, tuberculate. Spines reddish to yellowish to whitish. Central spines 4, upper 3 straight, flattened, banded, ascending, to 4cm (1.6in) long and 4mm wide, lower one curved or hooked at the tip, flattened, banded, to 5cm (2in) long and 9mm (0.4in) wide. Radial spines 5-15, radiating, straight or slightly curved, most banded, some flattened. Flowers funnel-form, purplish pink or yellow, with densely imbricate, ciliate bracts, to 4cm (1.6in) long and in diameter. Fruits ovoid, to 2.5cm (1in) long, covered with scales. Distribution: central Mexico. Two subspecies of Ferocactus latispinus are recognized. Subspecies latispinus typically has 9-15 radial spines that vary from stout and dark to fine and white; it is the most widespread, occurring in southeastern
Durange, Zacatecas, Aquascalientes, western San Luis Potosi, eastern Jalisco, Guanajuato, Queretaro, Hidalgo, Puebla, and Mexico. Subspecies spiralis has 5-7 stout radial spines; it is found only in southern Puebla and southern Oaxaca.

George Lindsey Entry for Ferocactus latispinus:
Stem simple, globose or flattened, to 3dm tall and 4dm wide, light green. Ribs about 21, vertical, acute, tuberculate, with a large protuberance above each areole. Areoles large, oval truncated floriferous section above, this concealed in older areoles by the overlapping tubercle. Spines reddish to yellowish; central spines 4, the upper 3 straight, annulate, flattened, ascending, to 4cm long and 4mm wide; the lower central spine longer, curved or hooked at the tip, flattened, annulate, to 5cm long and 9mm wide, usually descending; radial spines 12-15, radiating, straight or very slightly recurved, some flattened and some terete, most annulate, the lowest directly below the central spine and shortest, the next two flattened more than the rest; gland-spines produced in flowering areoles, above the spine bundle, persistent but soon covered by the tubercle above. Flowers produced in the summer months, funnel-form, purple or yellow, to 4cm long and as broad; scales of ovary very densely imbricate, 3mm long and 2mm wide, sclerous, ciliate, with acute tip, intergrading with scales of tube and outer perianth segments; scales of tube with fleshy, ciliated basal portion about 5mm long and wide, with an attenuate ciliate sclerous tip extending about 5mm above; outer perianth segments lanceolate, 18mm long and 4mm wide, with a thickened basal portion and a ciliate-fimbriate sclerous aristate tip; inner perianth segments lanceolate, 16mm long and 3mm wide, margins entire but tip mucronate; inner walls of hypanthium very fleshy, to 6mm thick; filaments numerous, 3-10mm long, anthers minute; style 25mm long, the upper 5mm divided into about 16 unequal stigma loves; color of flowers varies from orchid to yellow, the most common being mahogany outer perianth segments, orchid inner perianth segments, red stamens, red style, and yellow stigma lobes, but on many plants, particularly those with yellow spines, the flower parts may be shades of yellow.  Fruit oval, to 2.5cm long and 1.75cm wide, with the withered perianth 4cm long; covered with densely imbricate acute sclerous scales. Seed, small, elongate-reniform, shiny dark brown, deeply pitted, to 1.5mm long, 1mm wide and 0.6mm thick.

4. Dunlap observations of Ferocactus latispinus.
Apex of a young plant: 2in h, 3in w (NIC spines) (5cm high x 8cm wide).
The ribs are separated by 4mm of wool at the center of the apex; the top edge of each rib will then emerge and become exposed above the wool. The wool in the apical areole is pale and long; as the areole ages the wool turns gray – loosing volume, length, and height.
The youngest spines are pale lime green; as they age the lime green initially retreats to the base of the spine and then disappears altogether. The primary spine and secondary spine-set acquire a light brown tip and then become buff-white with banding as they age. The circumference of the young spine-set is smaller than that of mature spine set, and the spines nearly converge at the broadest circumference of the plant. Variable areole spacing appears to reflect diverse growing conditions. The oldest spine set contains nine spines – a primary central spine with a slightly hooked tip surrounded by eight secondary spines. Seven of the secondary spines are equal in size; the eighth spine, in the 6 o'clock position, is smaller in diameter. The youngest spine set contains eleven spines – a primary central spine, a set of secondary spines (in this case five) emerging from within the areole around the primary spine, and a second set of secondary spines (in this case five) emerging out of the periphery of the areole. The base of all the spines form a flat ellipse in cross section. The primary hooked spine has a three-part a-symmetrical cross section; one half forms a flat triangle; the other half is forms half an ellipse.
This specimen has fifteen ribs. The terminus of four of these ribs is set back from the apex by 4-5mm; these ribs were added to the plant as it matured. The top of each rib is relatively sharp, only 2mm wide. The skin is green with a mid-range hue value; it leans toward the blue-green end of the spectrum. An apical view of the footprint of the plant produces a deeply ribbed circle. The ribs are 14mm deep and 17mm wide; its width swells slightly at the point at which an areole is inserted. Each rib infrequently undulates at the point at which an areole is inserted on an adjoining rib.  

S10 Master List Other Genera – Cacti
Prepared by Susan Dunlap

Updated July, Aug, Dec, 2002     
Updated Jan 5, 2003, Added SL66-SL87
    
Acanthocalycium griseum            A17-3
Acinocereus spp. Ann                sl45
Ariocarpus fissuratus v. lloydii            sl23
Ariocarpus fissuratus        P        sl23
Ariocarpus retusus         Phiz         80 or 83    1
Armeria caespitosa             home     sl85    26
Armeria    spp sea pink         Home    SL85    2
Asclepias speciosa Yerba Buena         SL82    1 & 7    
Astelia hervosa v. chathamica             SL52
Astroloba aspera major            sl48
Astroloba bicarinata                sl51
Astrophytum asterias x            sl22
Astrophytum capricorne v. aureum         SL54
Astrophytum capricorne v. niveum         SL54
Astrophytum coahuilense         home     sl73    22
Astrophytum myriostigma v. columnari 'huboki'    A17-32
Astrophytum myriostigma            sl22
Astrophytum myriostigma            sl48
Astrophytum myriostigma    P        A7-18
Astrophytum ornatum                sl48
Atztekium ritteri (George)
Atztekium ritteri                P37    sl9
Blossfeldia campaniflora            sl13
Borzicactus celsiamis                SL61
Brizocactus hendriksenianus v. densilanta    SL61
Brois’s bush                 home     sl80 or 83    8-10
Brucii hybrid                 SL55
Bulbine latifolia             home     sl81        15-17
Caralluma foetida                A11-20,21
Caralluma pachycymbium keithii        A18-34,35
Carnegea gigantia    (spp)            A17-20
Cephalocereus senilis                A7-17,18
Cereus peruvianis                A19-16,17
Cereus tetragonus
Cheiridopsis herrei                SL50
Chileorebutia esmeraldana    Ann        P26,28    sl14
Chileorebutia esmeraldana    Ann        P9    sl10
Cleistocactus straosii                SL62
Cleosia spp             home         sl85    5
Coaliuilese tricost another spp/v.
Coaloiuiles tricost
Cochemiea maratima                sl48
Cochemiea poselgeri                sl48
Copiapoa alticostata                SL63-36 syn coquimbana
Copiapoa areospina                SL63-5 not found
Copiapoa atacamensis v calderana    Ann     SL66        3-4
Copiapoa barquitensis                sl21
Copiapoa barquitensis                SL62-24 syn hypogaea
Copiapoa borealis                SL64-15    echinata syn of C. fiedleriana   echinata v. borealis
Copiapoa bridgesii         Ann        SL66        1-2
Copiapoa brunescens – dura? echinoides? Ann     SL66    13-14
Copiapoa brunescens            SL63-12     syn megarhiza
Copiapoa calderana            sl48
Copiapoa carrizalensis         SL63-38     syn C. malletiana            
Copiapoa cinarescens         Ann     SL66        26-7
Copiapoa cinarescens            SL63-30
Copiapoa cinera v. albispina         SL63-7
Copiapoa cinera            SL63-6
Copiapoa columna-alba        SL63-4     syn cinerea
Copiapoa cuprea            SL63-22     syn echinoides v. cuprea
Copiapoa cupreata            SL63-14     syn echinoides C. cupreatus syn echinoides
Copiapoa dealbata crest    Ann    P33-4    sl14/33+syn of malletiana
Copiapoa dealbata longispina P    A17-31     syn of malletiana
Copiapoa dealbata            SL63-15     syn malletiana
Copiapoa dealbata    Ann    P15    sl9     syn of malletiana
Copiapoa desertica            SL62-32-3 not f    ound
Copiapoa desertica            SL62-32-3 not found
Copiapoa desertorum            SL63-13     syn taltalensis
Copiapoa desertorum            SL63-9     syn taltalensis
Copiapoa domeykoensis         SL62-25-6 syn pendulina; pendulina=syn of C. coquimbana   
Copiapoa dumetorum            SL63-18 not found
Copiapoa dumetorum            SL63-18 not found
Copiapoa dura            sl48    syn echinoides v. dura
Copiapoa dura            SL63-25     syn echinoides v. dura
Copiapoa dura            SL63-26    syn echinoides v. dura
Copiapoa echinata side?    Ann    SL66        25
Copiapoa echinata            SL63-34-35 syn fielderiana
Copiapoa echinoides            SL63-32-33
Copiapoa eremophila            SL63-8     syn haseltoniana
Copiapoa esmeraldana        SL62—31
Copiapoa ferox            SL63-17     syn solaris v. ferox
Copiapoa fiedleriana         Ann     SL66        31-33
Copiapoa fiedleriana            SL64-21
Copiapoa gigantea            sl48     syn of haseltoniana
Copiapoa goldii Ann            sl45 not found
Copiapoa goldii            SL64-10 not found
Copiapoa goldii            SL64-10 not found    rare
Copiapoa grandiflora            SL62-27    
Copiapoa humilie         Ann     SL66        29-30
Copiapoa humilis            SL62-29
Copiapoa hypogaea            SL62-28   barquitensis=syn?
Copiapoa imbricata            SL63-37 catalogue name=? C.dura lapshin.org
Copiapoa imbricata            SL64-3 as above
Copiapoa krainziana v. bruinispina    SL63-2-3
Copiapoa krainziana            SL63-10
Copiapoa laui         Ann    SL66        5-6, 12
Copiapoa longispina            SL63-31     syn of mollicula
Copiapoa longistamii            sl48 spp? longistaminea?
Copiapoa longistaminea     Ann     SL66        19-20
Copiapoa longistaminea     Ann     SL66        9
Copiapoa macrocarpa            SL63-23-24 not found
Copiapoa macrocarpa            SL63-23-24 not found
Copiapoa maleolata            SL62—30 not found
Copiapoa maleolata            SL62—30 not found
Copiapoa malletiana         Ann     SL66        34-35
Copiapoa malletiana            SL64-4-5
Copiapoa marginata         Ann     SL66        23
Copiapoa marginata            SL63-16
Copiapoa megarhzia            SL63-27    
Copiapoa minima            SL64-13-14 not found
Copiapoa minima            SL64-13-14 not found
Copiapoa minuta            SL64-11-12 not found
Copiapoa minuta            SL64-11-12 not found
Copiapoa mollicula         Ann     SL66        10
Copiapoa mollicula         Ann     SL66        28
Copiapoa mollicula Ann P14,19,15,16    SL12-14-19
Copiapoa multispina            SL64-16-17 not found
Copiapoa multispina            SL64-16-17 not found
Copiapoa pseudocoquimbana         SL62-34-5 syn of coquimbana    
Copiapoa pseudocoquimbana v. valgata SL62-36-7
Copiapoa rupestris         Ann     SL66        24
Copiapoa rupestris            SL63-19     syn of taltalensis
Copiapoa scopalina            SL63-11     syn of krainziana(lapshin)
Copiapoa serpentissulcata Ann     SL66        11
Copiapoa serpentisulcata v. castanea     SL63-21 spp castanea syn=serpentisulcata
Copiapoa solaris         Ann     SL66        21-22
Copiapoa solaris v. longispina         SL62-38-9 spp
Copiapoa ssp    Ann    P21        sl12
Copiapoa tenuispina            SL63-20 aka tenuissima?
Copiapoa tenuissima         Ann     SL66        7-8
Copiapoa tigrillo             SL64-8-9 not found Mesa gardens sells tigillensis
Copiapoa tigrillo             SL64-8-9 not found Mesa gardens sells tigillensis
Copiapoa tocoplana            SL63-28-29   aka tocopillana?
Copiapoa ubligiana            SL64-6-7 not found
Copiapoa ubligiana            SL64-6-7 not found cinerea v.?
Copiapoa valienarensis         SL64-18 not found; vallenarensis?
Copiapoa valle de Huasco        SL64-19    not found
Copiapoa valle de Huasco        SL64-19    not found
Copiapoa vallenarensis        SL64-20 vallenarensis=syn coquimbana
Copiapoa varispinata (conglomerata)     SL66    15-18 Ann
Copiapoa wagenknechtii        SL63-39, SL64-2 syn coquimbana
Copiapoa wagenkneditii P29,32    SL14     syn of coquimbana
Cordyline australi albertii         SL55
Coryphantha durangensis        sl10
Coryphantha elephantidens    Ann    P23,25,26    sl13
Coryphantha greenwoodii        sl48
Coryphantha greenwoodii P        A7-5-9
Coryphantha greenwoodii Ann P4    sl10/4
Coryphantha pseudoechinus    P13    sl6
Coryphantha radians    Ann P17,18,20-1    sl13/17+
Coryphantha sulcata    Ann    P20    sl9
Discocactus bueneckeri         home     sl73    28-9
Discocactus cueneckeri         home     sl73    18,19
Discocactus fuiicornis deflexispinus AnnP37    sl14
Dolicothele camptotricha            sl48
Dorstenia foetida
Dorstenia foetida                SL51
Echinocactus grusonii cristat   Ann        P34    sl18
Echinocactus imgens                sl23 no apex
Echinocactus texansis                A17-4
Echinocactus texansis            P    A17-24
Echinocactus visnaga                 SL55
Echinocereus engelmanii v. acicularis         SL54
Echinocereus enneacanthus v. stramineus    A7-29,30
Echinocereus luteus            sl48
Echinocereus moracallii        SL51
Echinocereus pectinatus        sl22
Echinocereus purpureus Ann        sl45
Echinocereus reichenbachii
Echinocereus rigidisimis        A17-26
Echinocereus rigidisimus v. rubispinus    A17-21
Echinocereus rosanthus (form of)    A17-10
Echinocereus rosanthus diff. Clone    A17-18
Echinocereus v. dauasii (red-tipped spp)
Echinofossulocactus albatas        sl48
Echinofossulocactus coptonogonus     sl48
Echinofossulocactus tegel bergii Ann    sl14/7+     P7,13
Echinopsis "Pacific Sunset"
Echinopsis hybrid Pastel Peach        A17-5,6,7
Echinopsis pentlandii             SL55
Echinopsis Stars and Stripes
Echinopsis subdenudata
Echinopsis subdenudata        sl48
Echionocereus pectinatus v. rupispinus    A19-23
Epithelantha unquispina     home     sl73    17
Epiphilium spp    Phiz         80 or 83    2-5
Epithalanthus angispina    P    sl23/4-7
Epithelantha bokei        P    sl23/12-14
Epithelantha micro v. unquispina Ann    P    sl45
Epithelantha micromeris    P10    sl23/8-11
Epithelantha unquispina 2/4/03 home     sl73    23
Epthelantha micromeris
Erica longifolia            SL60
Erigeron glaucus ‘sea breeze’ seaside daisy                                         Yerba Buena     SL82    10
Eriocereus                sl24/20
Eriocyce ceratistes    Ann        P27    sl12
Eriocyce rhodentiophila        P29-30    sl12
Eriogeron compact form Seaside daisy     Yerba Buena                                             SL82    8
Eriogonium latifolium Yerba Buena     SL82    5-6
Eriogonium umbellatum v pallyanthum – dwarf     Yerba Buena                                         SL82    2-3
Escobaria chichuahuensis Ann        sl45
Escobaria chihuahuensis
Escobaria dasyacantha        sl23
Escobaria rigida            sl23
Espostoa lanata            SL48
Euphorbia clandestina     home         sl80 or 83    16-17
Euphorbia flanniganii            SL51
Euphorbia haworthio. home         sl80 or 83    23-24
Euphorbia obesa home         sl80 or 83    18-19
Euphorbia purpurasum blossom home     sl81 or 84    23
Fern spp natrlz’d 7/27/03 home     sl85    33
Ferocactus alamosanus v. platygonus    A7-24-5
Ferocactus chrysacanthus        sl48
Ferocactus echidne v. victoriensis    SL61
Ferocactus glaucescens         SL55
Ferocactus glaucescens        sl48
Ferocactus glaucescens        SL61
Ferocactus gracilis        P    A7-13-16
Ferocactus histrix            SL61
Ferocactus latispinus            sl48
Ferocactus microdiscus        sl48
Ferocactus pottsii v. alamosanus     sl55
Ferocactus pringlii hybrid        A17-13
Ferocactus recktispinus    P    A17-33
Ferocactus recurvus        P    sl24/25
Ferocactus wizlizenii    Ann    P24,26    sl18/24+
Ferraria crispa            SL60
Fragaria chiddensis beach strawberry     SL82    9
Fragaria vesca woodland strawberry     SL82        4
Frailea cataphracta
Furcata seloa             SL55
Glandulicactus mathssonii        SL51
Gymnali artigas            sl21
Gymnocalycium albertensis     home     sl81 or 84
Gymnocalycium andreae-baldianum x andreaei                                                P    21/5+
Gymnocalycium asterium Ann    P        sl45
Gymnocalycium asterium            sl48        
Gymnocalycium baldianum    P31        A7-31,32
Gymnocalycium bruchii             SL54
Gymnocalycium bruchii     home         sl73    15
Gymnocalycium capillaense             SL54
Gymnocalycium cardenasiunum    P14    sl14
Gymnocalycium carminanthum    P32,34    sl12
Gymnocalycium castelianosii    home         sl73    21
Gymnocalycium castellanosii     P17,19        sl10
Gymnocalycium custelianosii     home         sl73    26
Gymnocalycium denudatum     home         sl73    27
Gymnocalycium gibbosum             SL54
Gymnocalycium griseopallidum Ann    P    sl45
Gymnocalycium horstii        P    A19-21
Gymnocalycium mazanense             SL54
Gymnocalycium mehlianum     home         sl73    16
Gymnocalycium mihanovichii friedrichii    sl48
Gymnocalycium monvillei             sl48
Gymnocalycium nigoum nin    P        sl21/18-23
Gymnocalycium pflanzii             SL54
Gymnocalycium pflanzii (g. marguezii)         SL54
Gymnocalycium pflanzii v. riograndense     SL54
Gymnocalycium pflanzii            sl48
Gymnocalycium quehlianum         home     sl73    25
Gymnocalycium ragonesii Ann    P        sl45
Gymnocalycium rotundulum     home         sl81 or 84
Gymnocalycium rotundulum            sl48
Gymnocalycium saglionis             SL54
Gymnocalycium schickendantzii (sierra valasco)     SL54
Gymnocalycium schickendantzii delaetii     SL54
Gymnocalycium spp (nuevo mundo Bolivia)     SL54
Gymnocalycium spp (sierra medina)         SL54
Gymnocalycium spigazinni        P    A17-11
Gymnocalycium tricanthum        P    A8/16
Gymnocalycium tucavocense            sl48
Gymnocalycium valnicekianum         SL54
Gymnocalycium weissianum             SL54
Gymnocalycium x                sl48
Hamatocactus hamatocanthus            sl48
Hatiora salicorinoides home             sl80 or 83    25
Hatiora salicorinoides                SL51
Hatiora salicorionides tail end of Morgan Territory sl73?    34-37
Horridocactus horridus Ann            sl45
Horridocactus horridus            SL49
Horridocactus lissocarpus            SL49
Lategan Rauch oudshorn            sl17
Leuchtenbergia principis 3-4 ea.        SL50
Lobilia elongata                sl18
Lobivia famatinensis Ann            sl45
Lobivia glauca Ann                sl45
Lobivia versicolor x culpinensis        SL51
Loopocereus schottii monstrosum        A17-23
Lophooereus schottii monstrosus    P24-6    sl11/24-31
Maihuenia poeppigii (chili) bed 167        SL61
Mamillaria luethyii graft
Mammilaria duwei    Ann        P34    sl22/
Mammilaria spp        Phiz         80 or 83    6
Mammillaria aureicentra            SL50
Mammillaria bombycina
Mammillaria carmenae            sl18
Mammillaria celsiana                SL50
Mammillaria celsiana                SL51
Mammillaria compacticalus        P27    sl18
Mammillaria duwei Ann            sl45
Mammillaria ebayone        
Mammillaria elongata                SL49
Mammillaria geminispina
Mammillaria hahniana            SL50
Mammillaria hejapensis            sl24
Mammillaria hejapensis            sl24
Mammillaria heteropodum    
Mammillaria huitzilogochtlii            sl24
Mammillaria huitzilogochtlii            sl24
Mammillaria isotensis
Mammillaria jaliscana form            sl18
Mammillaria krassuckal Ann    P16,17,18    sl18
Mammillaria leptacantha Ann            sl45
Mammillaria magnimamma            SL49
Mammillaria melanocentra            SL49
Mammillaria microhelia            SL49
Mammillaria nejapensis
Mammillaria pottsii                sl24
Mammillaria satorii                sl24
Mammillaria silvatica                SL50
Mammillaria spp                SL50
Mammillaria spp                SL50
Mammillaria spp        Ann         SL66       36
Mammillaria stampferi            SL49
Mammillaria stapelioides            SL35/23-24
Matts pick 2, only one in cultivation        sl11/32,33,37
Matt's pick    from Java    flower
Matucana aurantiaca                SL61
Matucana aureiflora                SL49
Matucana madisoniorum        P15-6    sl13
Matucana violaceus
Melocactus galucescens         home     sl73    20
Melocactus glaucescens         home     sl73    24
Melocactus glaucescens            SL62
Melocactus neryi    Ann        P28-9    sl9
Melocactus violaceus ssp margaritaceus        SL49
Monteiroi brandbergensis            A1-4,5
Mystery 1 1/4"H x 1" dia 4" pot
mystery pitted red edge, mild glaucous, branching; lf=1 ½ cm w                                home            sl84    36
Mystery reddish brown
Mystery w/red stem extended tubercles + spine shield on tip or tubercle
Neobuxbaumia polylopha            sl23
Neochilenia floccosa Ann            sl45
Neochilenia gracilis Ann            sl45
Neochilenia imitans        P3        sl12
Neochilenia napina    Ann    P18        sl9/18+
Neochilenia occulata    Ann    P14        sl22
Neochilenia residua    Ann    P22-6        sl12/22+
Neochilenia scoparia        P19/24        sl14/19-24
Neochilenia simulans    Ann    P3,4        sl14/3+                                        P36        sl13
Neochilenia subikii        P6,9,11        sl12/+
Neoparodia castanea
Neoporteria mitis
Neoporteria nidus v. senilis            SL49
Notocactus leninghausii             SL54
Notocactus magnificus             SL54
Notocactus scopa                SL49
Notocactus submammulosus            SL61
Notocactus submamulosus Ann    P6    sl18
Notocactus x                    SL51
Obregonia denegrii        (G.)    P
Opuntia ssp                    sl22
Opuntia (articulated)                 SL51
Opuntia (paper)                 SL51
Opuntia articulata    
Opuntia articulata v. inermis
Opuntia engelmannii                sl24
Opuntia ficus-indica                sl24
Opuntia fulgida (juvenilEuphorbia mutant)
Opuntia linquiformis                SL50
Opuntia phaeacerntha or phaeacantha        sl24
Opuntia spp no name
Opuntia ssp. cholla                sl24
Opuntia subulata monstronse            SL50
Opuntia tuncata (took broken piece)        SL61
Opuntia variagata maverick            SL50
Orbeopsis melanacantha            sl22
Oreocereus celsianus                sl48
Oreocereus trollii                sl48
Oreocereus varicolor v. facnaensis        sl24 fruit
Oreocereus x (longer hair)    wrong=hendron    sl48
Oroya caespitosa    
Parodia (notocactus) schlossor            A19-18,19,20
Parodia herzogii                SL49
Parodia maasii         Ann         sl67    4
Parodia maasii                SL62
Parodia maxima         Ann         SL66        37
Parodia maxima                SL49
Parodia microsperma v serenana Ann        SL67     2-3
Parodia nivosa                        
Parodia pencilata                SL51
Parodia penicilata                SL50
Parodia schwebsiana     Ann             SL66        38
Parodia subterranea    
Pediocactus simpsonii    6 spp            A17-25    
Pelecyphora aselliformis            sl23
Pelecyphora valdeziana            sl23
Perergonium spp        Phiz         sl79 or 82    11
Peumus boldus (monimiaceae) (Chili; looks like Puya)                                        GG Park     sl71
Pygmaecereus bylesianus            sl21
Pyrrhocactus chorosensis Ann        P11,13    sl10/11+
Rebutia densipectinata    Ann    P11    sl18    
Rebutia heliosa    Ann        P10    sl10
Rebutia sinilis                SL50
Rebutia graessnerii                sl18
Sensevaria s0p FKH                SL51
Soehrensia bruchii                SL62
Stenocactus capespitosus Ann    P        sl45
Stenocactus erectocentrus            sl48
Stenocactus multicostata Ann        P16-7    sl9
Stenocactus parksiensis        P    A17-22
Stenocactus pruinosis            P    
Stenocactus spp        Ann        SL67         1
Stenocactus ssp. lau                sl48
Stenocereus griseous                A17-2
Submatucana madisoniorum
Sulocorebutia rauschii    Ann        P7    sl18
Tephrocactus molinensis            sl22
Teucadendron discolor (lrg bush in S Af Section GG Park sl71
Thelocactus flavidispinus            sl21
Thelocactus hexadrophorus x rinconensis
Thelocactus lloydii                sl14
Thelocactus phymatothele            sl13
Thelocactus rinconensis    Ann    P223    sl9
Toumeya paperacantha            A17-27
Trichocaulon columnaris    Ann    P9,11    sl17
Trichocereus grandiflorious
Trichocereus    taquenbalensis, tacanensis, tacaquirensis                                         Stanford     sl86    18
Turbinicarpus polaskii Ann            sl45
Turbinicarpus schmiedikunus v. dickisonias Ann    sl45
Ubelmania buiningii                A19-29
Ubelmania pectinifera            A19-24,25
Ulbelmannia pseudopectinifera Ann        sl45
Weingartia lanata                SL51
Wild bees Kuil
winterizing             Stanford     sl86    10
Xanthorrhoea preissei             SL55




S11 Master List Other Genera - primarily Succulents
Prepared by Susan Dunlap

Abromietiella chlorantha    
Acanthus mollus – lrg shing lf’d plant @ front path GG Park     sl71
Adromischus maximus globular ctg took in Oct.            sl16
Adromiscus spp in ground             7/27/03    home     sl85    36
Adromiscus undulatum             Phiz         sl79 or 82    18
Aeonium (poss same as #11; fewer markings)     home         sl84    15
Aeonium (x…mimo)  from Rogers Weld            109 ucd
Aeonium algonica                        SL60
Aeonium arborescens v acho purpurea         Stanf         sl86    31
Aeonium arboreum                 home         sl84    9
Aeonium canariense v. palmense                SL60
Aeonium front yd spp             home         sl84    21
Aeonium gomerence                 home         sl84    20
Aeonium kiwi                 home         sl84    17
Aeonium lancerottense                    SL60
Aeonium pseudotabuleforme             home     sl85    37
Aeonium pseudotabuleforme             home     sl84    34
Aeonium sedifolium  TSG             home     sl85    17
Aeonium sedifolium  whole plant         home     sl85    18
Aeonium simii                 Stanf     sl86    27
Aeonium simsii    107ucd
Aeonium spp                 home     sl84    23
Aeonium spp (Russell’s)             home     sl84    6
Aeonium spp bending to light             home     sl84    3
Aeonium spp H. grn w/markings; solitary     home     sl84    11
Aeonium spp 2 diff rosettes on same plant     home     sl84    12
Aeonium spp tallest = ?             Stanf     sl86    32
Aeonium spp under cast smiling moon         home     sl84    8
Aeonium spp                     home     sl84    4
Aeonium spp                    GGPark sl71
Aeonium spp                    home     sl84    13
Aeonium spp                    home     sl84    5    
Aeonium spp                    Stanf     sl86    30
Aeonium spp? #8 may be parent         home     sl84    30
Aeonium spp? sticky, lt grn, caespitose?,    home     sl84 28
Aeonium spp? sunburned, 6” clay pot         home     sl84    39
Aeonium tabuliforme ctg             GGPark sl70
Aeonium v swartzkoff             home     sl84    10
Aeonium variagate                 home     sl84    16                
Aeoonium spp (sticky)             home     sl84    18
Agave americana sml silver             Stanf     sl86    21
Agave angustifolia                 Stanf     sl86    22
Agave attenuata                 SL55
Agave colorqata                 SL54
Agave fernandi regis                 SL52
Agave filifera                106ucd
Agave filifera?                 sl44
Agave geminiflora                 SL55
Agave grey blk tip                 SL54
Agave lechuguilla                 SL55
Agave macroculmis                SL61
Agave Mckelveyana                SL61
Agave monster                 SL55
Agave ocahui                SL61
Agave palmeri                SL61
Agave parryi v. hauchncensis            SL51
Agave polianthiflora                SL59
Agave pumila                104 ucd
Agave purpusorum                 SL55
Agave schidigera                SL61
Agave schottii                 SL55
Agave shawii bab.092                106ucd
Agave spp b64.016                106ucd
Agave spp in bloom            Stanf     sl86    12
Agave spp                    SL55
Agave stricta                 SL54
Agave stricta                 SL55
Agave stricta red                 SL55
Agave stricta                    SL59
Agave toumeyana v. bella            SL60
Agave univittata                 SL54
Agave utahensis ssp. kaibabensis        SL59
Agave vexans                SL61
Agave victoria-reginae v.             SL55
Agave vilmoriana                 SL52
Agave victoria-reginae            106ucd
Agave x                    sl44
Aloe abyssicola                 SL53
Aloe aculeata                sl23
Aloe aculeata                SL60
Aloe adigratana                sl46
Aloe allwyn                    SL60
Aloe arenicola                SL49-6
Aloe aristata v. parvifolia            SL60
Aloe bakeri                    SL51
Aloe berhana                    SL60
Aloe branddraaiensis                SL60
Aloe brevifolia w/Aeoniums in bkgrnd     Stanf     sl86    17
Aloe brevifolia                SL61
Aloe camperi (confirm)            SL60
Aloe camperi                sl46
Aloe capitata (#2)                SL60
Aloe capitata                    SL60
Aloe ciliaris                     sl52
Aloe claviflora                SL60
Aloe comptonii                SL61
Aloe comptonil                 SL53
Aloe cryptopoda                SL60
Aloe decoingsii                107ucd
Aloe deltoidiodenta v brevifolia         SL53
Aloe dichotoma             Stanf     sl86    6
Aloe distans                 Stanf     sl86    15
Aloe distans                    sl46
Aloe dortheae – not frost tolerant     Stanf     sl86    26
Aloe erinacea  plant #2            110 ucd
Aloe erinacea B92.944            110 ucd
Aloe ferox                 Stanf     sl86    24
Aloe fragilis  BAB.017            109 ucd
Aloe fulleri  B2000.186            109 ucd
Aloe globuligemma x (speciosa?)         SL53
Aloe grandidentata    #10?         Stanf     sl86    8
Aloe humilis  B2002.138            109 ucd
Aloe jucunda                    A8-6,7
Aloe juvena                    110 ucd
Aloe karasbergensis                SL53
Aloe mabendiensis                 SL53
Aloe maristata                 SL52
Aloe marlothii – toothy, lt grn, 24” Stanf     sl86    4
Aloe marlothii                 SL52
Aloe melanacantha v. erinacea   E bed 12"h    SL42/3
Aloe melanacantha                110 ucd
Aloe millotii                    110 ucd
Aloe mitriformis                SL60
Aloe mubendiensis                SL60
Aloe nobilis hybrid  #11         Stanf     sl86    9
Aloe peglerae                SL60
Aloe pfinslooi                SL60
Aloe pilansii                    sl22
Aloe plicatilis                SL60
Aloe polyphylla                 SL52 Stanf     sl86    14
Aloe polyphylla                SL60
Aloe reitzii                    SL60
Aloe sapoirania                sl46
Aloe sinkatana                 SL53
Aloe speciosus                SL60
Aloe spp                     SL52
Aloe striata                 Stanf     sl86    3
Aloe striata                    SL61
Aloe striatula hardy to 0        Stanf     sl86    1
Aloe tenuior – smaller than striatula    Stanf     sl86    7
Aloe tenuior                    sl46
Aloe ukambensis                SL60
Aloe vanbalenii                sl46
Aloe vera                     SL52
Aloe vera yellow flower =        Stanf     sl86    2
Aloes +++                     SL55
Aloes 6-19                    SL62
Aloinopsis rubrolineata            A19-26,27
Anacampseros marlothii  B94.424        109 ucd
Anacampseros spp            home     sl85    10
Anacampseros spp            Phiz     sl79 or 82    12
Anacempsceros telephiastrum  baa.807        107ucd
Ancistrocactus sheeri                103 ucd
Arctostaphylos cruzensis             Yerba  Buena SL82    15
Arctostaphylos densiflora             Yerba  Buena SL82    17
Arctostaphylos edmunsii v parvifolia         Yerba  Buena SL82    13-14
Arctostaphylos franciscana             Yerba  Buena SL82    12
Arctostaphylos manzanita            Yerba  Buena SL82    21-22
Arctostaphylos sunset manzanita         Yerba  Buena SL82    16
Arctostaphylos uva-ursi ‘radiant’         Yerba  Buena SL82    11
Arequipa efectocylindrica            104 ucd
Arequipa weingartiana            102 ucd
Armeria caespitosa     7/27/03    home         sl85    26
Armeria    spp sea pink         Home    SL85    2
Asclepias speciosa Yerba  Buena         SL82    1 & 7    
Astelia hervosa v. chathamica             SL52
Astroloba aspera major            sl48
Astroloba bicarinata                sl51
Avonia quinaria                    105 ucd
Brois’s bush                 home     sl80 or 83    8-10
Bulbine latifolia         5/19/03 home         sl81    15-17
Caralluma foetida                A11-20,21
Caralluma pachycymbium keithii        A18-34,35
Ceanothus ‘yankee point’, carmel creep Yerba  Buena         SL82    24-25
Ceanothus gloriosus porrectus        Yerba  Buena         SL82    20
Ceanothus griseus v horizontous     Yerba  Buena         SL82    18
Ceanothus impressus             Yerba  Buena         SL82    19
Cleosia spp                 home             sl85    5
Coaliuilese tricost another spp/v.
Coaloiuiles tricost
Coleus aromaticus         home         sl85    13
Colyledron rubrovenosa            sl17
Comphry home                 sl80 or 83    35-37
Conophytum maughanii - big!!         sl10
Coral bells            Home        SL85    3
Cordyline australi albertii             SL55
Coteledon?  (Tylecodon?)     Home        SL85    5    
Cotyledon ladismithensis     home         sl84    33
Cotyledon ladismithensis variagate?home     sl84    34
Cotyledon orbicularis v oophylla     home     sl84    32
Cotyledon orbiculata                sl46
Cotyledon rubrovenosa   Ann    P28        sl17
Cotyledon sdlg. H                 SL54
Crassula austrailiensis (air)     Phiz     sl79 or 82    21-22
Crassula barbata            sl17
Crassula barkleyi     Phiz         sl79 or 82    4
Crassula columnella            sl49-7
Crassula columnella Ann     P15    sl17
Crassula compacta (jade-like)     home     sl84    25
Crassula corymbulosa     Phiz     sl79 or 82        23-26
Crassula falcatta            SL51
Crassula golum monsytrose        sl49
Crassula higgensii  B94.425        110 ucd
Crassula ivory pagoda            sl49
Crassula merchandii            sl49
Crassula moonglow            sl49
Crassula Morgan's beauty        sl49
Crassula multiclava     7/27/03    home     sl85    30
Crassula multiclava home         sl80 or 83    29-30
Crassula obiculata flower (#13) Phiz     sl79 or 82    20
Crassula obvallata  plant #2     home     sl85    4
Crassula obvallata         home     sl85    3
Crassula orbiculata     Phiz         sl79 or 82    13-14
Crassula pallida         home     sl80 or 83    15
Crassula perfoliata v. perfoliata    SL61
Crassula picturata            sl22
Crassula plegmatoides        sl49
Crassula spp (reseda)             SL54
Crassula spp        Phiz         sl79 or 82    17
Crassula spp        Phiz         sl79 or 82    19
Crassula spp        Phiz         sl79 or 82    9-10
Crassula susannae            sl49
Cycods revoluta             SL52
Cynanchum marnieranum Ann        sl45
Cyphostemma juttae            SL60
Dasylirion                 SL55
Dasylirion glaucophyllum  bab.084    106ucd
Dasylirion longissima, Gig Harbor sept 2002
Dasylirion spp             SL55
Dasylirion spp             SL55
Dasylirion wheeleri   b92.874        106ucd
Dasylirion wheeleri             SL55
Dasylirion wheeleri apex Gig Harbor sept 2002
Delosperma nubigenum
Denmoza formosa            SL61
Dioon spinulosum             SL61
Dolicothele camptotricha        sl48
Dorstenia foetida            SL51
Dudleya ‘cymosa’    at edge of brick Home    SL85    38
Dudleya ‘snata cruz is’ at edge of brick Home    SL85    34
Dudleya abramsii     home             sl83    26-7
Dudleya brittonii                 SL55
Dudleya brittonii?     at edge of brick Home    SL85    35
Dudleya c.2.6                SL85    13
Dudleya C.3.1  pebble beach            SL85    19
Dudleya c.3.1                SL85    11
Dudleya c.3.2 (SI                SL85    12
Dudleya c.3.3 (s.3                SL85    114
Dudleya c.3.4 sl4                SL85    15
Dudleya c.3.5 s.1                SL85    17
Dudleya c.4.2  one site            SL85    18
Dudleya c.5.1 one site                SL85    19
Dudleya c.5.2  one site            SL85    20
Dudleya c.6/ s.2.1  pac grove            SL85    22
Dudleya c.6/s.1.1  marina            SL85    21
Dudleya c.6/s.3.b.1 highlands inn        SL85    23
Dudleya c.6/s.4.1 bixby creek            SL85    24
Dudleya c.6/s.4.2 bixby creek            SL85    25
Dudleya c.6/s.4.3                SL85    26
Dudleya c.6/s.5.1 so of bixby            SL85    27
Dudleya c.6/s.6.1 big sur ranch            SL85    28
Dudleya c.6/s.6.3 s of coast gallery        SL85    29
Dudleya c.6/s.7.1  s of coast gallery        SL85    30
Dudleya c.6/s.8.1 n of coast gallery        SL85    31
Dudleya densiflora 11/25/02, Sun         sl64? 11
Dudleya edulis                SL59
Dudleya edulis    at edge of brick Home    SL85    37
Dudleya edulis? wegmans 6”     home         sl83    24
Dudleya greenei    at edge of brick Home    SL85    36
Dudleya greenii    Succ Garden     home     sl85    1
Dudleya pulverulenta     5/19/03    home         sl81    11-12
Dudleya pulverulenta ssp arizonica    Home    SL85    33
Dudleya pulverulenta                SL59
Dudleya Santa cruz Is     at edge of brick Home    SL85    39
Dudleya setchelli (yerba buena) Home        SL85    32
Dudleya spp     home                 sl81 or 84
Dudleya spp #2 source = ?    home         sl83    29
Dudleya spp Bear Valley         sl80 or 83    11-14
Dudleya spp C.1.1             sl83    31
Dudleya spp C.1.2             sl83    332
Dudleya spp C.1.3             sl83    33
Dudleya spp C.2.1             sl83    34
Dudleya spp C.2.2             sl83    35
Dudleya spp C.2.3             sl83    36
Dudleya spp c.2.4             sl84    1
Dudleya spp c.2.5             sl84    2
Dudleya spp herman sml glaucous, linear oval     home     sl83    25
Dudleya spp I.1.1  hw 20             sl83    21
Dudleya spp I.1.2                 sl83    22
Dudleya spp I.2.1 Behemian hey         sl83    30
Dudleya spp Lucielle D virens uppermost apex     home     sl83    23
Dudleya spp Lucille?                 sl83    28
Duvalia parriflora         Phiz         sl79 or 82    28-29
Dykia spp            Phiz         sl79 or 82    6
Dykia marnier-lapostollei            107ucd
Ebelmanmnia buiningii
Echeveria agavoides     GG Park         sl71
Echeveria agavoides                sl44
Echeveria albicans                 SL55
Echeveria albicans"?                sl44
Echeveria alpina (Lone Pine)     home         sl85    8
Echeveria amoena         home         sl85    19
Echeveria derenbergii     home         sl84    37
Echeveria derenbergii        home         sl85    11
Echeveria elegans                SL62
Echeveria fimbriata ?  stem (broken 2 wks ago) home     sl85    7
Echeveria fimbriata    TSG        Home        SL85    9
Echeveria fimbriata?  TSG  label lost     home         sl85    6
Echeveria johnsonii  Portulacaceae        109 ucd
Echeveria laui  B98.199            109 ucd
Echeveria multicaulis     home             sl84    27
Echeveria multicaulis                SL59
Echeveria pallida                SL51
Echeveria peacockii                104 ucd
Echeveria pruinose? hairy, alt., gray-grn  home     sl84    26
Echeveria pulidonis                SL59
Echeveria simulans                SL59
Echeveria spp             home     sl85    20
Echeveria spp        Stanf         sl86    29
Echeveria spp?  Anacampseros?     home     sl85    9
Echeveria spp? brn tip, fuzzy,         home     sl84    38
Echeveria topsy-turvy     Phiz             sl79 or 82    1
Echeveria?   E bed 23', 31'            SL43/2
Echium fastuosum                 SL52
Echium wildpretii                SL52
Echium wildprettii Wegmans     7/27/03    home     sl85    27
Edithcolea grandis                A1-3
Erica longifolia                SL60
Erigeron glaucus ‘sea breeze’ seaside daisy Yerba  Buena    SL82    10
Eriocereus                        sl24/20
Eriogeron compact form Seaside daisy     Yerba  Buena     SL82    8
Eriogonium latifolium         Yerba  Buena     SL82    5-6
Eriogonium umbellatum v pallyanthum    Yerba  Buena     SL82    2-3
Euphorbia clandestina     home             sl80 or 83    16-17
Euphorbia flanniganii                SL51
Euphorbia haworthio.. home             sl80 or 83    23-24
Euphorbia obesa home             sl80 or 83    18-19
Euphorbia purpurasum blossom  home         sl81 or 84    23
Fern spp  natrlz’d      7/27/03    home         sl85    33
Ferraria crispa                SL60
Fragaria chiddensis beach strawberry     Yerba  Buena     SL82    9
Fragaria vesca woodland strawberry     Yerba  Buena     SL82    4
Furcata seloa             SL55
Gasteira batesiana            110 ucd
Gasteria     spp        Stanf     sl86    23
Gasteria ‘frosti’     5/19/03 home     sl81        13-14
Gasteria acinacifolia     5/19/03 home     sl81        10
Gasteria acinacifolia             SL62
Gasteria acinacifolia         Stanf     sl86    28
Gasteria acinacifolia            SL65
Gasteria acinacifolia        Stanf     sl86    13
Gasteria angustifolia            110 ucd
Gasteria batesiana             SL65
Gasteria batesiana            SL64
Gasteria batsialia            SL51
Gasteria baylissiana            SL64
Gasteria baylissiana            110 ucd
Gasteria beckeri  BASA.267        109 ucd
Gasteria bicolor v. bicolor        SL65
Gasteria bicolor v. liliputana        SL65
Gasteria bicolor            SL50
Gasteria bicolor            SL64
Gasteria brachyphylla            SL65
Gasteria brevifolia            SL61
Gasteria brivifolia B84.172        109 ucd
Gasteria caespitosa            110 ucd
Gasteria carinata v verrucosa        110 ucd
Gasteria carinata v. carinata        SL65
Gasteria carinata v. retusa        SL65
Gasteria carinata v. verrucosa        SL65
Gasteria carinata            110 ucd
Gasteria carinata            SL65
Gasteria croucherii            SL64
Gasteria decepiens            SL65 out
Gasteria disticha            110 ucd
Gasteria disticha            SL64
Gasteria ellaphieae            SL65
Gasteria ellephiae            SL64
Gasteria excelsa            SL60
Gasteria excelsa            SL65
Gasteria frostii            SL51
Gasteria gigantea            SL51
Gasteria gigantia            SL51
Gasteria glomerata            SL64
Gasteria green ice            SL50
Gasteria hilotica            SL64 out
Gasteria humilis            SL65 out
Gasteria liliputana            SL61
Gasteria liliputana            110 ucd
Gasteria maculata            sl22
Gasteria madagascarensis        SL64 out
Gasteria nigricans marmorata        SL51
Gasteria nitida v armstrongii=Gasteira armstrongii  B93.382    109 ucd
Gasteria nitida v. armstrongii        SL61
Gasteria nitida v. armstrongii        SL65
Gasteria nitida v. armstrongii        sl22
Gasteria nitida v. nitida        SL65
Gasteria nitida            SL65
Gasteria obtusa            SL51
Gasteria obtusifolia            SL64 out
Gasteria pillansii + variagate        SL65
Gasteria pillansii v. ernest-ruschii    SL65
Gasteria pillansii v. pillansii        SL65
Gasteria pillansii varigate        SL64
Gasteria pillansii            110 ucd
Gasteria pillansii            SL61
Gasteria pulchra            SL65
Gasteria rawlinsonii + spiral form    SL64
Gasteria sugared            SL51
Gasteria vlokii            SL65
Gasteria vlokii\Sensevaria pinquicula Ann    sl45
Gasteria white wing                 SL51
Geranium spp        Phiz         sl79 or 82    33-34
Geranium x oronianum “Walter’s gift’ home     sl85    32
Geranium-like next to apple     home         sl80 or 83    27-8
Glottiphylum latifolium                SL60
Graptopetalum macdougalii     home         sl80 or 83    26
Graptopetalum opendandrum            SL51
Graptopetalum saxifragoides     home         sl81 or 84    24
Graptopetalum saxifragoides     home         sl84    22
Haworthia angustifolia  b97.239        107ucd
Haworthia arachnopidea  BAB.324        109 ucd
Haworthia attenuata (growing in less light)     110 ucd
Haworthia attenuata v caespitosa        108ucd
Haworthia attenuata                108ucd
Haworthia bayeri  b98.020            107ucd
Haworthia bolusii v blackbeardiana        108ucd
Haworthia bolusii                108ucd
Haworthia cassytha                SL50
Haworthia coarctata ssp coarctata        108ucd
Haworthia coarctata v coarctata forma greeni    108ucd
Haworthia comptoniana            109 ucd
Haworthia cooperi                108ucd
Haworthia cymbiformis cv. variegatum        SL50
Haworthia cymbiformis v obesa    108ucd
Haworthia cymbiformis v planifolia    108ucd
Haworthia cymbiformis v transiens    108ucd
Haworthia cymbiformis        SL50
Haworthia cymbiformis        SL51
Haworthia decipiens  b91.533        108ucd
Haworthia emelyea            107ucd
Haworthia fasciata forma browneana    108ucd
Haworthia fasciata            108ucd
Haworthia glauca v herrei forma armstrongii    108ucd
Haworthia herbacea                108ucd
Haworthia hybrid Bev's Wonder            
Haworthia joeyae cymbiformis form  b98.019    107ucd
Haworthia lemondonut Ann            sl45
Haworthia limifolia v limifolia  b85.294    107ucd
Haworthia limifolia v schultiana    108ucd
Haworthia limnifolia    
Haworthia longiana v albinota        108ucd
Haworthia magnifica usplendens    108ucd
Haworthia magnifica v acuminata    108ucd
Haworthia magnifica v notabilis    108ucd
Haworthia magnifica            SL50
Haworthia magnificameiringii        SL50
Haworthia mantelii            SL50
Haworthia maraisii v meiringii        108ucd
Haworthia marumiana v batesiana    108ucd
Haworthia mirabilis v paradoxa    108ucd
Haworthia mutica            108ucd
Haworthia nigra            108ucd
Haworthia odonoghieana AnnP31    sl21/
Haworthia parksiana B94.202        109 ucd
Haworthia puelinitziana        SL50
Haworthia radula            108ucd
Haworthia reinwardii            SL50
Haworthia reinwardtii v riebeekensis    110 ucd
Haworthia reticulata v subregularis    108ucd
Haworthia retusa v. multilineata or mirabilis v beukmanii        109 ucd
Haworthia retusa            SL50
Haworthia rycroftiana            108ucd
Haworthia scraba            108ucd
Haworthia soridida Ann        sl45
Haworthia springbockvalkensis  b92.023107ucd
Haworthia spp            SL50
Haworthia spp        Phiz     sl79 or 82    3
Haworthia spp        Stanf     sl86    16
Haworthia starhiana            SL61
Haworthia starkiana v starkiana    108ucd
Haworthia tessalata x gasteria            
Haworthia truncata v. crassa Ann P12    sl17
Haworthia truncata            SL50
Haworthia ttenuata            SL61
Haworthia turgida v pallidifolia    108ucd
Haworthia turgida v suberecta        107ucd
Haworthia turgida v suberecta        108ucd
Haworthia turgida            108ucd
Haworthia unicolor v venteri        108ucd
Haworthia v. longibracteata        107ucd
Haworthia venosa ssp tessalata     SL51
Haworthia venosa ssp tessellata    108ucd
Haworthia venosa v tessallata        108ucd
Haworthia venosa v. recurva        sl61
Haworthia viscosa            SL50
Haworthia viscosa            110 ucd
Haworthia wooleyi            SL50
Hereroa tugwelliae            SL60
Hoodia pilifera            A17-16
Horrisia justbertii            A17-8
Huerchera next to natrlzd fern     7/27/03    home     sl85    31
Huernia distincta         Phiz         sl79 or 82    30-31
Huernia kenneayana spp?    Ann        sl22
Huernia kennedyana     Phiz             sl79 or 82    2    
Huernia kennedyana    Ann    P36        sl18
Huernia pillansii  B82.073            109 ucd
Huernia plowesii    Ann    P28-30        sl15
Huernia schneideriana                SL50
Ibiris                Home    SL85    4
Iislaya grandiflorens II            sl21
inside concave glaucous fleshy stem divides @ base & branches     home sl84    24
Islaya islayensis minor Ann            sl45
Jatropha cuneata        P24        sl4
Jatropha cuneata?orange blossom,         sl4, sl5
Kalanchoe beharensis                sl44
Kalanchoe longiflora                sl44
Kalanchoe rhombopilosa  B75.063        109 ucd
Kalanchoe spp?         home         sl85    2
Kleinia obesa                A10-31
Lamb’s ear            Home        SL85        8
Leachia cactiformis                110 ucd
Larryleachia cactiforme             sl17
Monadenium arborescens        P    A9-27
Monadenium guentherii
Monadenium heteropodium            sl48
Monadenium heteropodum        P    A10-17
Monadenium magnificum
Monadenium magnificum        P    A10-22
Monadenium reflexum    Ann    P    sl45
Monadenium renneyi            P    A10-18,19
Monadenium richii            P    A17-9
Monadenium sp. Tanzania        P    A2-16-18;A9-19
Monadenium spectabile         P            sl30/36-7
Nolian microsirpa                 SL54
Nolina                     SL55
Nolina             Stanf         sl86    19
Nolina interrata                 SL55
Nolina perryi  11/25/02, Sun             sl64? 12
Orchid? spp        Phiz             sl79 or 82    5
Oscularia deltoides     home             sl84    31
Pachyphytum compactum     5/19/03 home     sl81        8-9
Pachyphytum fittkaui                 SL55
Pachyphytum gulutinicaule            SL60
Pachyphytum oviforum
Pachyphytum oviforum  home        P16    sl4flwr
Pachypodium brevicaule  B2001.157  Apocynace109 ucd
Pachypodium gayi    big housespp,        sl19/21-23
Pachypodium lamerei
Pachypodium namaquanum  B98.276 profile    109 ucd
Pachypodium namaquanum  B98.276        109 ucd
Pachypodium namaquanum Ann    P17    sl17
Pachyveria 'aphra'                 SL55
Pectinaria longipes            P32    sl17
Penstemon                Home    SL85        7
Perergonium spp            Phiz     sl79 or 82    11
Peumus boldus (monimiaceae) (Chili; looks like Puya)GG Park     sl71
Poelinitzia rubrifolia  B82.193            110 ucd
Polycyphora aselliformis SIB 804
Portulaca Molokiniensis  B2000.092        109 ucd
Pseudolithos migiurtinus            110 ucd
Pterodiscus aurianticus            109 ucd
Puya berteroniana in Chili    GG Park     sl71
Puya berteroniana again (in succ/cact section GG Park     sl71
Puya spp ex B.Kemble                 SL55
Rhus integrifolia lemonade berry Yerba Buena     SL82    27
Rhus ovata         Yerba  Buena         SL82    26
Red and Mike w/ cover     Stanf         sl86    5
Rosularia pallida  b59.092            107ucd
Salvia argentea (siler sape) 7/27/03    home     sl85    28
Saponaria x striata                sl46
Sarcocanlan burmannu            sl24
Sarcocanlan penciilinum            sl24
Scopellogena verruculata            sl46
Sedum  Gramma’s gray-grn 7/27/03 home     sl85    35
Sedum furfuraceum    home             sl84    29
Sedum hintonii  B70.112            109 ucd
Sedum liveforever vera james homeP34,37    sl15
Sedum ozxacanum                SL59
Sedum palmeri                SL59
Sedum spp  flower of         home         sl85    22
Sedum spp (front yard) home             sl80 or 83    20-22
Sedum spp (Praire Crk trip;         home     sl85    21
Sedum spp lt grn natrlzd    7/27/03    home     sl85    34
Sedum spp or Crassula     7/27/03    home     sl85    24
Sedum spp        home             sl84    7
Sedum spp        Phiz             sl79 or 82    27
Sedum spp        Phiz             sl79 or 82    15-16
Sedum spp        Phiz             sl79 or 82    35-36
Sempervivum ‘oddity’ hs leek 7/27/03home         sl85    25
Sempervivum arachnoideum  spp #2 7/27/03    home     sl85    29
Sempervivum brachnoideum – lucille         home     sl85    12
Sempervivum sos +    ?  & jade     Stanf         sl86    25
Senecio ballyi   b83.010                107ucd
Senecio ficoides                    SL61
Senecio haworthiodes                SL62
Senecio mandraliscae (snip; grey fingers) GG Park     sl71
Senecio mandraliscae                SL61
Senecio mweroensis ssp saginatus  b97.142    107ucd
Senecio mweroensis ssp saginatus        SL51
Senecio mweroensis ssp saginatus        SL51
Senecio scaposus                SL51
Senecio scaposus                SL51
Senecio spp  TSG         home         sl85    16
Senecio spp TSG         home         sl85    23
Sensevaria s0p FKH                SL51
sml grey Dudleya – farinosa?     home         sl81 or 84
sml grn pachyphytum-like; 2” pot     home     sl84    14
Stapelia flavirostrus                sl22
Stapeliopsis neronis     Phiz             sl79 or 82    32
Stetsonia coryne
Teucadendron discolor  (lrg bush in S Af Section GG Park     sl71
Tylecodon buchholziana    Ann        P5    sl17
Tylecodon cacalioides                sl22
Tylecodon cacaloides    Ann            sl45        
Tylecodon decepience    Ann        P4    sl18
Tylecodon ellaphieae rosijnfijiecberg AnnP30-1    sl17
Tylecodon reticulata    Ann         P34,38    sl17
Yucca                     SL55
Yucca aloeifolia variagata     Stanf         sl86    20
Yucca baccata                SL61
Yucca filifera (variagata)     Stanf         sl86    11
Yucca filifera                SL61
Yucca rostata                 SL52
Yucca rostrata                SL59
Yucca whipplei                SL52