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Year: 2015
A taxonomic backbone for the global synthesis of species diversity in the
angiosperm order Caryophyllales
Hernández-Ledesma, Patricia ; Berendsohn, Walter G ; Borsch, Thomas ; Mering, Sabine Von ; Akhani,
Hossein ; Arias, Salvador ; Castañeda-Noa, Idelfonso ; Eggli, Urs ; Eriksson, Roger ; Flores-Olvera,
Hilda ; Fuentes-Bazán, Susy ; Kadereit, Gudrun ; Klak, Cornelia ; Korotkova, Nadja ; Nyffeler, Reto ;
Ocampo, Gilberto ; Ochoterena, Helga ; Oxelman, Bengt ; Rabeler, Richard K ; Sanchez, Adriana ;
Schlumpberger, Boris O ; Uotila, Pertti
Abstract: The Caryophyllales constitute a major lineage of flowering plants with approximately 12500
species in 39 families. A taxonomic backbone at the genus level is provided that reflects the current
state of knowledge and accepts 749 genera for the order. A detailed review of the literature of the
past two decades shows that enormous progress has been made in understanding overall phylogenetic
relationships in Caryophyllales. The process of re-circumscribing families in order to be monophyletic
appears to be largely complete and has led to the recognition of eight new families (Anacampserotaceae,
Kewaceae, Limeaceae, Lophiocarpaceae, Macarthuriaceae, Microteaceae, Montiaceae and Talinaceae),
while the phylogenetic evaluation of generic concepts is still well underway. As a result of this, the
number of genera has increased by more than ten percent in comparison to the last complete treatments
in the Families and genera of vascular plants” series. A checklist with all currently accepted genus names
in Caryophyllales, as well as nomenclatural references, type names and synonymy is presented. Notes
indicate how extensively the respective genera have been studied in a phylogenetic context. The most
diverse families at the generic level are Cactaceae and Aizoaceae, but 28 families comprise only one to six
genera. This synopsis represents a first step towards the aim of creating a global synthesis of the species
diversity in the angiosperm order Caryophyllales integrating the work of numerous specialists around the
world.
DOI: https://doi.org/10.3372/wi.45.45301
Posted at the Zurich Open Repository and Archive, University of Zurich
ZORA URL: https://doi.org/10.5167/uzh-122159
Journal Article
Published Version
Originally published at:
Hernández-Ledesma, Patricia; Berendsohn, Walter G; Borsch, Thomas; Mering, Sabine Von; Akhani,
Hossein; Arias, Salvador; Castañeda-Noa, Idelfonso; Eggli, Urs; Eriksson, Roger; Flores-Olvera, Hilda;
Fuentes-Bazán, Susy; Kadereit, Gudrun; Klak, Cornelia; Korotkova, Nadja; Nyffeler, Reto; Ocampo,
Gilberto; Ochoterena, Helga; Oxelman, Bengt; Rabeler, Richard K; Sanchez, Adriana; Schlumpberger,
Boris O; Uotila, Pertti (2015). A taxonomic backbone for the global synthesis of species diversity in the
angiosperm order Caryophyllales. Willdenowia, 45(3):281-383.
DOI: https://doi.org/10.3372/wi.45.45301
A taxonomic backbone for the global synthesis of species diversity in the
angiosperm order Caryophyllales
Author(s): Patricia Hernández-Ledesma, Walter G. Berendsohn, Thomas Borsch, Sabine Von Mering,
Hossein Akhani, Salvador Arias, Idelfonso Castañeda-Noa, Urs Eggli, Roger Eriksson, Hilda FloresOlvera, Susy Fuentes-Bazán, Gudrun Kadereit, Cornelia Klak, Nadja Korotkova, Reto Nyffeler,
Gilberto Ocampo, Helga Ochoterena, Bengt Oxelman, Richard K. Rabeler Adriana Sanchez, Boris O.
Schlumpberger & Pertti Uotila
Source: Willdenowia, 45(3):281-383.
Published By: Botanic Garden and Botanical Museum Berlin (BGBM)
DOI: http://dx.doi.org/10.3372/wi.45.45301
URL: http://www.bioone.org/doi/full/10.3372/wi.45.45301
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Willdenowia 45 – 2015
281
Patricia Hernández-LedeSMa1,2, WaLter G. BerendSOHn1, tHOMaS BOrScH1,3, SaBine VOn
MerinG1, HOSSein aKHani4, SaLVadOr ariaS5, ideLFOnSO caStañeda-nOa6, UrS eGGLi7, rOGer
eriKSSOn8, HiLda FLOreS-OLVera9, SUSY FUenteS-Bazán1, GUdrUn Kadereit10, cOrneLia
KLaK11, nadJa KOrOtKOVa1,3, retO nYFFeLer12, GiLBertO OcaMPO13, HeLGa OcHOterena9,
BenGt OXeLMan8, ricHard K. raBeLer14, adriana SancHez15, BOriS O. ScHLUMPBerGer16 &
Pertti UOtiLa17
A taxonomic backbone for the global synthesis of species diversity in the angiosperm
order Caryophyllales
Abstract
Hernández-Ledesma P., Berendsohn W. G., Borsch th., Mering S. von, akhani H., arias S., castañeda-noa i., eggli
U., eriksson r., Flores-Olvera H., Fuentes-Bazán S., Kadereit G., Klak c., Korotkova n., nyffeler r., Ocampo G.,
Ochoterena H., Oxelman B., rabeler r. K., Sanchez a., Schlumpberger B. O. & Uotila P.: a taxonomic backbone
for the global synthesis of species diversity in the angiosperm order Caryophyllales. – Willdenowia 45: 281 – 383.
2015. – Version of record first published online on 11 September 2015 ahead of inclusion in december 2015 issue;
iSSn 1868-6397; © 2015 BGBM Berlin.
dOi: http://dx.doi.org/10.3372/wi.45.45301
the Caryophyllales constitute a major lineage of flowering plants with approximately 12 500 species in 39 families.
a taxonomic backbone at the genus level is provided that reflects the current state of knowledge and accepts 749
genera for the order. a detailed review of the literature of the past two decades shows that enormous progress has
been made in understanding overall phylogenetic relationships in Caryophyllales. the process of re-circumscribing
families in order to be monophyletic appears to be largely complete and has led to the recognition of eight new families (Anacampserotaceae, Kewaceae, Limeaceae, Lophiocarpaceae, Macarthuriaceae, Microteaceae, Montiaceae
and Talinaceae), while the phylogenetic evaluation of generic concepts is still well underway. as a result of this, the
number of genera has increased by more than ten percent in comparison to the last complete treatments in the “Families and genera of vascular plants” series. a checklist with all currently accepted genus names in Caryophyllales, as
well as nomenclatural references, type names and synonymy is presented. notes indicate how extensively the respective genera have been studied in a phylogenetic context. the most diverse families at the generic level are Cactaceae
and Aizoaceae, but 28 families comprise only one to six genera. this synopsis represents a first step towards the aim
of creating a global synthesis of the species diversity in the angiosperm order Caryophyllales integrating the work of
numerous specialists around the world.
additional key words: flowering plants, Caryophyllales network, checklist, phylogeny, taxon concept, genus, World
Flora Online, edit Platform for cybertaxonomy
General e-mail address for correspondence: caryophyllales@bgbm.org
1 Botanic Garden and Botanical Museum Berlin (BGBM), Freie Universität Berlin, Königin-Luise-Str. 6 – 8, 14195 Berlin, Germany.
2 current address: Laboratorio de Genética Molecular y ecología evolutiva, Facultad de ciencias naturales, Universidad autónoma de Querétaro, campus aeropuerto, Querétaro, Qto. 76140, Mexico.
3 institut für Biologie, Systematische Botanik und Pflanzengeographie, Freie Universität Berlin, altensteinstr. 6, 14195 Berlin,
Germany.
4 department of Plant Sciences, School of Biology, college of Science, University of tehran, P.O. Box 14155-6455, tehran, iran.
5 Jardín Botánico, instituto de Biología, Universidad nacional autónomo de México (UnaM), circuito exterior s.n., ciudad
Universitaria, ap. postal 70-614, México d.F. 04510, Mexico.
6 Jardín Botánico de Villa clara, Universidad central “Marta abreu” de Las Villas, Facultad de ciencias agropecuarias, carretera
de camajuaní km 5½, Santa clara, cuba.
7 Sukkulenten-Sammlung zürich, Mythenquai 88, cH-8002 zürich, Switzerland.
8 department of Biological and environmental Sciences, University of Gothenburg, Box 461, Se-40530 Göteborg, Sweden.
9 departamento de Botánica, instituto de Biología, Universidad nacional autónoma de México (UnaM), circuito exterior s.n.,
ciudad Universitaria, ap. postal 70-614, México d.F. 04510, Mexico.
(addresses continued on next page)
282
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
Introduction
Background
recent years have yielded a wealth of new informatics
tools and infrastructures to facilitate working with taxonomic data. Searching and accessing the necessary literature and type specimens has become much faster and
easier, thus stimulating research in plant systematics.
Modern monographic work synthesizes knowledge on
a group of organisms and generates, manages, and publishes high quality data as needed for a variety of applications. to be biologically meaningful and to allow correct
identification especially at the species level, the entities
recognized such as species or genera should as much
as possible reflect the latest understanding provided by
phylogenetic and evolutionary approaches (Marhold &
al. 2013; Borsch & al. 2015; naciri & Linder 2015). in
order to achieve this, an integration of the ever-increasing
number of phylogenetic and evolutionary studies and the
data generated by them with formal monographic work
is imperative. this requires the research process to be organized in a way that explicitly links data on characters
and specimens with evolutionary results and taxon concepts, and that allows for continuous updating to reflect
the continuous generation of knowledge (Borsch & al.
2015). at the same time there is now an increased awareness for the need of a comprehensive assessment of the
species diversity on our planet as a basis for conservation and sustainable use (Lughada & Miller 2009; Paton
2009; Hendry & al. 2010).
The Caryophyllales Global Synthesis Initiative
We have started a joint initiative entitled “Global synthesis of species diversity in the angiosperm order Caryo
phyllales”. the idea was to develop a practical model for
integrative monographic work that is based on a sizable
group of world-wide occurring organisms. Our approach
is to develop a network and an internet portal based on
a collaborative approach of institutions and individual
researchers studying various aspects of the diversity
and evolution of the Caryophyllales. Major partners
will function as focal points with a long-term institutional commitment that ensures sustainability of the
initiative. at the moment the core partnership consists
of: the instituto de Biología, Universidad nacional autónoma de México – UnaM (Mexico); the instituto de
Botánica darwinion (argentina); and the Botanic Garden and Botanical Museum Berlin – BGBM (Germany).
the BGBM is committed to support the coordination of
the initiative and will provide the biodiversity informatics infrastructure. apart from aiming at satisfying the
general scientific and applied need for quality data, we
specifically envision the application of the Synthesis in
the context of plant conservation. One of the immediate
outputs of the Caryophyllales synthesis will be an up-todate taxonomic backbone for the World Flora Online as
called for by the convention on Biological diversity’s
conference of the Parties (2012). considering the enormous progress on understanding and describing Caryo
phyllales diversity that has been made in the past two
decades and will continue into the future, and also the
need to have full coverage of the diversity for the users,
the approach will entail a mechanism to integrate new
results as they become available (Borsch & al. 2015) and
therefore to present the best possible treatment for any
given taxon. a comprehensive review and treatment at
the generic level is an important step that will then be
extended to the species level and be complemented by
descriptive and other information.
Caryophyllales as a model group
reasons for choosing Caryophyllales as model group
are diverse. the group is one of the major lineages of
angiosperms with about 12 500 species. it is strongly
supported as monophyletic by several molecular phylogenetic studies (Savolainen & al. 2000; Soltis & al. 2000;
cuénoud & al. 2002; Hilu & al. 2003; Brockington & al.
2009; Schäferhoff & al. 2009; Qiu & al. 2010; Soltis &
al. 2011; crawley & Hilu 2012).
the Caryophyllales are of great ecological and evolutionary interest because they show multiple origins of
specialized morphological, anatomical, and biochemical
traits. the order for example comprises the highest diversity of species with c4 photosynthesis after the grasses
(Sage & al. 2011). Several lineages are highly specialized
with adaptations to extreme habitats such as xeric conditions, salinity, or nitrogen-poor soils, and thus the group
includes many succulent, halophytic, gypsophilous and
(addresses continued from previous page)
10 institut für allgemeine und Spezielle Botanik, Herbarium MJG, anselm-Franz-von-Bentzelweg 9a, Johannes Gutenberg-Universität Mainz, 55099 Mainz, Germany.
11 Bolus Herbarium, department of Biological Sciences, University of cape town, 7701 rondebosch, South africa.
12 institut für Systematische Botanik, Universität zürich, zollikerstrasse 107, cH-8008 zürich, Switzerland.
13 departamento de Biología, centro de ciencias Básicas, Universidad autónoma de aguascalientes, avenida Universidad 940,
ciudad Universitaria, c.P. 20131, aguascalientes, ags., Mexico.
14 University of Michigan Herbarium-eeB, 3600 Varsity drive, ann arbor, Mi 48108-2228, U.S.a.
15 Facultad de ciencias naturales y Matemáticas, Programa de Biología, Universidad del rosario, carrera 24 no. 63c-69, Bogotá
d.c., colombia.
16 Herrenhäuser Gärten, Herrenhäuser Str. 4, 30419 Hannover, Germany.
17 Botanical Museum, Finnish Museum of natural History, P.O. Box 7, Fi-00014 University of Helsinki, Finland.
Willdenowia 45 – 2015
carnivorous plants. the Caryophyllales are the order
with the highest number of halophytes containing more
than 21 % of all halophytic species (Flowers & al. 2010)
and with the evolutionary oldest halophyte lineages (e.g.
Kadereit & al. 2012a). the anatomy of Caryophyllales
is also interesting because there are many wood features
that are difficult to interpret (e.g. successive cambia, vessel elements perforation plates, ray anatomy, and raylessness; carlquist 2010). in several families, pollen has
evolved complex architectures and ultrastructures, based
on the tricolpate pollen of the eudicots (Skvarla & nowicke 1976; nowicke 1994) with several Amaranthaceae
exhibiting strongly derived metareticulate pollen with
the highest number of apertures known in angiosperms
(Borsch 1998; Borsch & Barthlott 1998). Caryophyllales
are characterized by a unique phenomenon of petal loss
and repeated reinvention (Brockington & al. 2012; ronse
de craene 2013).
Furthermore, the order is relevant in the context of
the Global Strategy for Plant conservation and citeS
by including groups of plants with many endangered
species (e.g. Hunt 1999), most importantly Cactaceae,
Droseraceae and Nepenthaceae. Species of economic
importance include cereals and green vegetables (e.g.
amaranth, quinoa, spinach, sugar beet), ornamentals (e.g.
many Cactaceae and Caryophyllaceae species, carnivorous groups), noxious weeds (e.g. Alternanthera phi
loxeroides (Mart.) Griseb., Amaranthus spinosus L. and
Mirabilis and Opuntia species), and of medical importance (mainly allergens; e.g. Amaranthus retroflexus L.,
Atriplex species, Kali turgidum (dumort.) Guterm.).
the rapidly increasing number of fully sequenced
genomes (currently five: two Chenopodiaceae, two
Amaranthaceae and one Caryophyllaceae; http://www.
ncbi.nlm.nih.gov/genome) and trancriptomes (66 species of Caryophyllales are included in the 1KP initiative; https://sites.google.com/a/ualberta.ca/onekp/home)
opens new exciting opportunities for evolutionary studies in this order.
Circumscription and phylogenetic relationships of
Caryophyllales
For many decades the order just included the taxa characterized by a free central placentation (= Centrosper
mae), perisperm and curved embryos (Bittrich 1993a).
Based on phylogenetic analyses, the Caryophyllales are
now understood in a wider sense as also including Po
lygonales, Nepenthales and smaller lineages that were
distantly placed in earlier classification systems, such as
Rhabdodendron or Simmondsia (aPG 1998; cuénoud &
al. 2002). this concept of the order is also basically followed here. We will summarize the changes in the classification of Caryophyllales and the different families
below. this will help to understand the changes during
the long transition phase from pre-cladistic to phylogeny-based taxonomy.
283
Several pre-cladistic classification systems were
proposed for the Caryophyllales (for a review until the
1990s see cronquist & thorne 1994). rodman & al.
(1984) were the first to evaluate the classification of
Caryophyllales based on a cladistic analysis of morphological characters. they reasserted the monophyly
of the group and produced one of the first classifications based on a phylogenetic hypothesis (table 1),
even though this study was questioned with respect to
its methodology and character selection (Gianassi & al.
1992). Subsequently, early molecular systematic studies
(i.e. rettig & al. 1992; downie & Palmer 1994; downie
& al. 1997; Lledó & al. 1998) indicated the close relationship of the members of subclass Caryophyllidae (i.e.
Caryophyllales, Plumbaginales and Polygonales sensu
cronquist 1981). Further studies (e.g. albert & al. 1992;
chase & al. 1993) showed close phylogenetic relationships of Caryophyllidae with the carnivorous lineages
Droseraceae and Nepenthaceae (Nepenthales sensu
cronquist 1981). Morton & al. (1997) found that the
Madagascan Asteropeiaceae (Theales sensu takhtajan
1987) and Physenaceae (described by takhtajan 1985,
but placed in Sapindales) both belong to Caryophyllales.
this placement of Asteropeiaceae was further supported
by a morphological cladistic analysis (Luna & Ochoterena 2004). Other studies (e.g. Fay & al. 1997) clarified
the placement of Rhabdodendraceae (Rosales sensu
cronquist 1981), Simmondsiaceae (previously placed in
either Euphorbiaceae or Buxaceae; tobe & al. 1992),
Tamaricaceae and Frankeniaceae (Violales sensu cronquist 1988). the suggested affinities of all these groups
to Caryophyllales were examined by nandi & al. (1998),
with respect to the fit of morphological characters, who
adopted the concept of “caryophyllids s.l.” for a clade
including Caryophyllales sensu cronquist (1981) plus
most of the taxa mentioned above. nandi & al. (1998)
further showed that the Dioncophyllaceae (Theaneae
sensu takhtajan 1987) and Ancistrocladaceae (Theales
sensu cronquist 1981) also belong to the carnivorous
clade within the caryophyllids.
Based on a review of published molecular phylogenetic studies, the angiosperm Phylogeny Group (aPG
1998) considered 26 families to constitute the Caryo
phyllales with an expanded taxon concept. in this concept the order included all the families of the caryophyllids s.l. (nandi & al. 1998) plus several family segregates
such as Achatocarpaceae and Stegnospermataceae
(segregated from Phytolaccaceae), Drosophyllaceae
(segregated from Droseraceae) and Sarcobataceae (segregated from Chenopodiaceae). a molecular study by
Savolainen & al. (2000) tested this circumscription and
retrieved a well-supported clade. On their trees the authors annotated the families Halophytaceae (segregated
from Chenopodiaceae) and Petiveriaceae (segregated
from Phytolaccaceae).
Since then, further studies have improved the understanding of the phylogenetic relationships within
284
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
the expanded Caryophyllales. the study by cuénoud &
al. (2002) based on 18S rdna, rbcL, atpB, and partial
matK sequences, was relevant in terms of its sampling,
which included most of the families treated by Kubitzki
& al. (1993) and Mabberley (1997), including Agdesti
daceae, Barbeuiaceae and Gisekiaceae (segregated
from Phytolaccaceae). cuénoud & al. (2002) retrieved
a well-supported Caryophyllales clade in most of their
analyses, and one of their most relevant results was the
detection of major subclades: the “core Caryophyllales”
and “non-core Caryophyllales”. the core Caryophyl
lales included the traditionally recognized Caryophyl
lales (cronquist 1981) and their segregated families;
within this clade two subclades were recovered, one is
the “lower core Caryophyllales” including Achatocar
paceae, Amaranthaceae s.l. (including Chenopodiace
ae), Asteropeiaceae and Caryophyllaceae, and the other
is the “higher core Caryophyllales” including the rest
of the traditional Caryophyllales and their segregated
families. Within the “higher core Caryophyllales”, Cor
bichonia and Lophiocarpus (rbcL+matK analysis) were
considered as separate linages within Molluginaceae
and Phytolaccaceae, respectively. the “non-core Cary
ophyllales” clade also included two major subclades:
one including Frankeniaceae, Plumbaginaceae, Polygo
naceae and Tamaricaceae, and the other consisting of
the carnivorous families Ancistrocladaceae, Dionco
phyllaceae, Droseraceae and Nepenthaceae. the analysis of cuénoud & al. (2002) resulted in inconclusive positions for Rhabdodendraceae and Simmondsiaceae. in
their combined tree, Rhabdodendraceae were recovered
as sister to all Caryophyllales (100 % Bootstrap; BS),
and Simmondsiaceae as sister to the core Caryophyllales
(moderate BS), while in the analysis of matK (low BS),
both taxa as sisters were recovered as sister to the core
Caryophyllales.
the study by Hilu & al. (2003) based on matK
also retrieved two moderately supported major clades:
“Caryophyllales i” and “Caryophyllales ii”, the former
including the core Caryophyllales plus Simmondsiaceae
and Rhabdodendraceae (expanded core Caryophylla
les). Within this clade two sister groups were recovered,
“higher core i” and “higher core ii”, one comprising
Aizoaceae, Nyctaginaceae and relatives and the other
Cactaceae, Portulacaceae, and relatives. the “Caryo
phyllales ii” corresponded to the non-core Caryophyl
lales of cuénoud & al. (2002).
Schäferhoff & al. (2009) employed sequence data
of the petD group ii intron and matK and recovered the
“caryophyllids” and “polygonids” as major clades with
high confidence. the caryophyllids include the expanded
core Caryophyllales, which in general correspond to the
“Caryophyllales i” of Hilu & al. (2003). the polygonids
correspond to the non-core Caryophyllales of cuénoud
& al. (2002) and Caryophyllales ii of Hilu & al. (2003).
Furthermore, Schäferhoff & al. (2009) described the Mi
croteaceae (segregated from Phytolaccaceae) with the
sole genus Microtea, which they sampled for the first
time in any molecular study. the study underscored the
importance of a representative taxon sampling because
Microtea was identified based on just two markers as an
isolated lineage that together with the Simmondsiaceae
is the successive sister to the rest of the caryophyllids.
Other recent authors mainly increased the number of
characters analysed from the chloroplast. Brockington
& al. (2009) using nine plastid genes from the singlecopy region, the inverted repeat, and two nuclear genes,
recovered the non-core Caryophyllales and core Caryo
phyllales clades with Rhabdodendraceae followed by
Simmondsiaceae plus the clade Asteropeiaceae–Physena
ceae as successive sisters of the rest of the core Caryo
phyllales. Within the core Caryophyllales, the authors
designated the “globular inclusion” clade as the clade
that corresponds to the “higher core Caryophyllales” of
cuénoud & al. (2002). Within this clade, they referred to
the clade containing Cactaceae, Portulacaceae, and relatives as the “portulacaceous cohort” (an earlier-suggested
name by rodman & al. 1984, “cohort Portulacares”) and
the lineage including Aizoaceae, Nyctaginaceae, and
most parts of Phytolaccaceae possessing raphides as the
“raphide clade”. Soltis & al. (2011) used 17 genes (representing the three plant genomes) and came to results very
similar to those of Schäferhoff & al. (2009) and Brockington & al. (2009).
Several phylogenetic studies have focused on the Por
tulacineae (= Cactineae/Portulacaceous cohort) (applequist & Wallace 2001; nyffeler 2007; nyffeler & al.
2008; Ocampo & columbus 2010). the most recent
study by nyffeler & eggli (2010a) resulted in the disintegration of Portulacaceae, recognizing eight monophyletic families including the newly described Anacampse
rotaceae (segregated from Portulacaceae), the concept
of Portulacaceae s.str. as a monotypic family, changes
of the circumscription of some families (Didiereaceae),
and the re-establishment and change of concept of others
(Montiaceae and Talinaceae).
the family names Limeaceae and Lophiocarpaceae
were published in 2005 (Hoogland & reveal 2005) and
2008 (doweld & reveal 2008), respectively, based on
phylogenetic data (e.g. cuénoud & al. 2002) that were
confirmed in later works (e.g. Schäferhoff & al. 2009;
Brockington & al. 2009). in a similar way the Kewaceae
were validated (christenhusz & al. 2014) to accommodate the second lineage of the biphyletic genus Hyper
telis that had been found outside of Molluginaceae s.str.
(Schäferhoff & al. 2009; Brockington & al. 2011; christin & al. 2011) but in an isolated position sister to the
raphide clade. the genus Macarthuria that was resolved
in an isolated position as sister to the remainder of the
core Caryophyllales (Brockington & al. 2011; christin & al. 2011) was accommodated in the new family
Macarthuriaceae (christenhusz & al. 2014).
in summary, our concept of Caryophyllales includes
39 families (Fig. 1; table 1, 2). it is in line with the fami-
Willdenowia 45 – 2015
285
Fig. 1. Summary of the current knowledge on phylogenetic relationships in the Caryophyllales. Based on cúenod & al. (2002),
Brockington & al. (2009) and Schäferhoff & al. (2009). Branch widths shown as triangles indicate species richness in these clades. –
lll = high support (95-100 BS/JK/PP), ll = medium support (75-94 BS/JK/PP), l = low support (50-74 BS/JK/PP).
286
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
table 1. circumscription of Caryophyllales in a phylogenetic context according to different authors. the names in bold represent
changes in comparison to the previous concept. * = not at family level in aPG; ** = different concept from aPG iii (2009) and
Stevens (2001 onwards).
Centrospermae
Caryophyllidae
(rodman & al. 1984) (chase & al. 1993;
Morton & al. 1997;
Fay & al. 1997)
Aizoaceae
Amaranthaceae
Expanded Caryophyllales Expanded Caryophyllales
(aPG 1998)
(Savolainen & al. 2000;
cuénoud & al. 2002;
aPG ii 2003)
Current Caryophyllales
(Brockington & al. 2009;
Schäferhoff & al. 2009;
aPG iii 2009; Stevens
2001 onwards; nyffeler
& eggli 2010a; christin
& al. 2011; christenhusz
& al. 2014)
Achatocarpaceae
Achatocarpaceae
Agdestidaceae*
Aizoaceae
Amaranthaceae**
Anacampserotaceae
Ancistrocladaceae
Asteropeiaceae
Barbeuiaceae
Basellaceae
Cactaceae
Caryophyllaceae
Chenopodiaceae**
Didiereaceae
Dioncophyllaceae
Droseraceae
Drosophyllaceae
Frankeniaceae
Gisekiaceae
Halophytaceae
Kewaceae
Limeaceae
Lophiocarpaceae
Macarthuriaceae
Microteaceae
Molluginaceae
Montiaceae
Nepenthaceae
Nyctaginaceae
Achatocarpaceae
Agdestidaceae*
Aizoaceae
Amaranthaceae
Aizoaceae
Amaranthaceae
Aizoaceae
Amaranthaceae
Ancistrocladaceae
Asteropeiaceae
Ancistrocladaceae
Asteropeiaceae
Basellaceae
Cactaceae
Caryophyllaceae
Chenopodiaceae
Didiereaceae
Basellaceae
Cactaceae
Caryophyllaceae
Chenopodiaceae
Didiereaceae
Dioncophyllaceae
Droseraceae
Drosophyllaceae
Frankeniaceae
Basellaceae
Cactaceae
Caryophyllaceae
Didiereaceae
Dioncophyllaceae
Droseraceae
Drosophyllaceae
Frankeniaceae
Didiereaceae
Dioncophyllaceae
Droseraceae
Drosophyllaceae
Frankeniaceae
Gisekiaceae
Halophytaceae
Molluginaceae
Molluginaceae
Molluginaceae
Molluginaceae
Nyctaginaceae
Nepenthaceae
Nyctaginaceae
Nepenthaceae
Nyctaginaceae
Physenaceae
Phytolaccaceae
Plumbaginaceae
Polygonaceae
Portulacaceae
Rhabdodendraceae
Physenaceae
Phytolaccaceae
Plumbaginaceae
Polygonaceae
Portulacaceae
Rhabdodendraceae
Sarcobataceae
Simmondsiaceae
Stegnospermataceae
Nepenthaceae
Nyctaginaceae
Petiveriaceae
Physenaceae
Phytolaccaceae
Plumbaginaceae
Polygonaceae
Portulacaceae
Rhabdodendraceae
Sarcobataceae
Simmondsiaceae
Stegnospermataceae
Tamaricaceae
Tamaricaceae
Phytolaccaceae
Portulacaceae
Simmondsiaceae
Tamaricaceae
lies recognized by the aPG iii (2009) and Stevens (2001
onwards) but separates Agdestidaceae from Phytolac
caceae and Chenopodiaceae from Amaranthaceae and is
updated by adding Kewaceae and Macarthuriaceae. in
aPG iii (2009) Agdestis was included within Agdesti
doideae (Phytolaccaceae) although its position as sister
of Sarcobataceae obtained by cuénoud & al. (2002) and
Ancistrocladaceae
Asteropeiaceae
Barbeuiaceae
Basellaceae
Cactaceae
Caryophyllaceae
Physenaceae
Phytolaccaceae
Plumbaginaceae
Polygonaceae
Portulacaceae
Rhabdodendraceae
Sarcobataceae
Simmondsiaceae
Stegnospermataceae
Talinaceae
Tamaricaceae
Schäferhoff & al. (2009) supports the acceptance of the
family described by nakai (1942). aPG iii (2009) also
recognized the Sarcobataceae. the Amaranthaceae are
treated in a very wide sense in aPG iii (2009) including
all Chenopodiaceae, merely reflecting that the two families form a monophyletic group (cuénoud & al. 2002;
Kadereit & al. 2003; Müller & Borsch 2005a), while the
Willdenowia 45 – 2015
287
table 2. comparison of the current treatment with the two volumes edited by Kubitzki & al. or Rhabdodendraceae along
(1993) and Kubitzki & Bayer (2003) representing the so far most inclusive generic treatment with the core Caryophyllales
of the Caryophyllales.
(= Centrospermae). the polyFamily
No. of genera (Kubitzki & al.
1993; Kubitzki & Bayer 2003)
Achatocarpaceae
Agdestidaceae
Aizoaceae
Amaranthaceae
Anacampserotaceae
Ancistrocladaceae
Asteropeiaceae
Barbeuiaceae
Basellaceae
Cactaceae
Caryophyllaceae
Chenopodiaceae
Didiereaceae
Dioncophyllaceae
Droseraceae
Drosophyllaceae
Frankeniaceae
Gisekiaceae
Halophytaceae
Kewaceae
Limeaceae
Lophiocarpaceae
Macarthuriaceae
Microteaceae
Molluginaceae
Montiaceae
Nepenthaceae
Nyctaginaceae
Physenaceae
Phytolaccaceae
Plumbaginaceae
Polygonaceae
Portulacaceae
Rhabdodendraceae
Sarcobataceae
Simmondsiaceae
Stegnospermataceae
Talinaceae
Tamaricaceae
incertae sedis
2
n.a.
127
69
n.a.
1
1
n.a.
4
98
87
98
4
3
3
1
2
n.a.
1
n.a.
n.a.
n.a.
n.a.
n.a.
13
n.a.
1
31
1
17
27
43
29
1
n.a.
1
1
n.a.
4
5
Total
675
relationships of the major groups of Chenopodiaceae are
still under debate. in this case, a merger resulting in a
shift of family assignment for a major lineage of plants
with many genera appearing in numerous studies in ecology, agriculture, and conservation had been promoted
without robust phylogenetic data (see also respective
family treatments).
For ease of recognition, we distinguish the two major
Caryophyllales clades as caryophyllids and polygonids
following Schäferhoff & al. (2009). the caryophyllids
are the larger clade and include Simmondsiaceae and/
No. of genera
(present publication)
2
1
125
79
3
1
1
1
4
139
101
104
6
3
3
1
1
1
1
1
1
2
1
1
10
13
1
31
1
12
29
55
1
1
1
1
1
3
5
1
gonids include the “carnivorous clade” with Ancistrocla
daceae, Dioncophyllaceae,
Droseraceae, Drosophyllace
ae and Nepenthaceae plus
the Frankeniaceae + Tama
ricaceae and Plumbaginace
ae + Polygonaceae subclades
(Fig. 1).
Rationale for a revised generic classification
More than twenty years have
passed since the publication
of the comprehensive treatment of the centrospermous
families of Caryophyllales
by several authors in “Families and genera of vascular
plants” (Kubitzki & al. [eds.]
1993). there, 15 families
are recognized in the order
(Achatocarpaceae, Aizoace
ae, Amaranthaceae, Basel
laceae, Cactaceae, Caryo
phyllaceae, Chenopodiaceae,
Didiereaceae, Halophytace
ae, Hectorellaceae, Mollugi
naceae, Nyctaginaceae, Phy
tolaccaceae, Portulacaceae
and
Stegnospermaceae
[=
Stegnospermataceae]).
ten years later the treatment
was completed with the publication by Kubitzki & Bayer
(2003), where the concept of
“expanded Caryophyllales”
was adopted, by now also
treating Ancistrocladaceae,
Asteropeiaceae, Dioncophyl
749
laceae, Droseraceae, Dro
sophyllaceae,
Frankenia
ceae, Nepenthaceae, Physenaceae, Rhabdodendraceae,
Simmondsiaceae, and Tamaricaceae. in addition to the
treatments of these families, cuénoud (2003) discussed
the circumscription of the expanded Caryophyllales including Plumbaginaceae and Polygonaceae previously
considered as separate orders by Kubitzki (1993b) and
Brandbyge (1993), respectively. the two volumes edited
by Kubitzki & al. (1993) and Kubitzki & Bayer (2003)
represented the most inclusive generic treatment of the
Caryophyllales with 675 genera in 27 families. in addition, there are even more comprehensive family-wide
288
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
treatments including all genera and even species for
the Aizoaceae (Hartmann & al. 2001a, b), Basellaceae
(eriksson 2007), Cactaceae (Hunt 2006) and Portu
lacaceae (eggli 2002).
the amount of new data accumulated in the past two
decades has considerably improved our understanding
about the Caryophyllales and marks a major transition
from a pre-phylogenetic to a largely phylogeny-based
classification. Several molecular phylogenetic studies
have evaluated the intrafamilial classifications adopted
by various authors in Kubitzki & al. (1993), Hartmann &
al. (2001a, b) and Hunt (2006), for example: Aizoaceae
(Hassan & al. 2005; Klak & Bruyns 2012; Klak & al.
2003a, b, 2007, 2013; Bohley & al. 2015), Amaranthaceae (Kadereit & al. 2003; Müller & Borsch 2005a, b;
Sánchez-del Pino & al. 2009; Masson & Kadereit 2013),
Cactaceae (arias & al. 2005; Butterworth 2006; ritz &
al. 2007, 2012; Butterworth & edwards 2008; Griffith
& Porter 2009; Ocampo & columbus 2010; Korotkova
& al. 2010, 2011; Bárcenas & al. 2011; calvente & al.
2011a, b; demaio & al. 2011; Hernández-Hernández
& al. 2011; Majure & al. 2012; Franck & al. 2013a, b;
Vázquez-Sánchez & al. 2013), Caryophyllaceae (Oxelman & al. 2001; Fior & al. 2006; Harbaugh & al. 2010;
Greenberg & donoghue 2011), Chenopodiaceae (Kadereit & al. 2003, 2006a, b, 2010; Kapralov & al. 2006;
akhani & al. 2007; zacharias & Baldwin 2010; Kadereit
& Freitag 2011; Fuentes-Bazán & al. 2012a, b), Didiereaceae (applequist & Wallace 2000), Nyctaginaceae
(Levin 2000; douglas & Manos 2007), Plumbaginaceae
(Lledó & al. 1998, 2001, 2005), Polygonaceae (Sanchez
& Kron 2008, 2009; Burke & al. 2010; Burke & Sanchez
2011; Sanchez & al. 2011; Schuster & al. 2011; Kempton 2012; Sun & zhang 2012).
these and other molecular phylogenetic studies have
resulted in the confirmation or rejection of monophyly
in several taxa, and consequently in changes of their circumscription or status. Some of the re-established taxa
are for example Lymanbensonia, Nyctocereus (Cacta
ceae, Korotkova & al. 2010, arias & al. 2005, respectively); Atocion, Eudianthe, Heliosperma (= Ixoca),
Viscaria (Caryophyllaceae, Oxelman & al. 2001);
Lipandra, Oxybasis (Chenopodiaceae, Fuentes-Bazán &
al. 2012b); and Afrobrunnichia (Polygonaceae, Sanchez
& Kron 2009).
taxa for which the circumscription had, or has, to be
changed in order to accept them as monophyletic groups
are for example: Arenaria (Caryophyllaceae, Harbaugh
& al. 2010), Atocion (Caryophyllaceae, Frajman &
al. 2009b), Atraphaxis (Polygonaceae, Schuster & al.
2011a, b), Austrocylindropuntya (Cactaceae, ritz &
al. 2012), Bassia (Chenopodiaceae, Kadereit & Freitag
2011), Beta (Chenopodiaceae, Kadereit & al. 2006b),
Brunnichia (Polygonaceae, Sanchez & Kron 2009),
Chenopodium (Chenopodiaceae, Fuentes-Bazán &
al. 2012b), Echinopsis (Cactaceae, Schlumpberger &
renner 2012), Ferocactus (Cactaceae, Vázquez-Sánchez
& al. 2013), Grayia (Chenopodiaceae, zacharias &
Baldwin 2010), Hatiora (Cactaceae, Korotkova & al.
2011), Limoniastrum (Plumbaginaceae, Lledó & crespo 2000), Lychnis (Caryophyllaceae, Oxelman & al.
2001; Popp & al. 2008), Mammillaria (Cactaceae, Bárcenas & al. 2011; Hernández-Hernández & al. 2011),
Mesembryanthemum (Aizoaceae, Klak & al. 2007),
Minuartia (Caryophyllaceae, dillenberger & Kadereit
2014), Moehringia (Caryophyllaceae, Fior & Karis
2007), Opuntia (Cactaceae, Majure 2012), Pachycereus
(Cactaceae, arias & terrazas 2009), Peniocereus (Cac
taceae, arias & al. 2005), Pfeiffera (Cactaceae, Korotkova & al. 2010), Polycarpon (Caryophyllaceae, Kool
& al. 2007), Silene (Caryophyllaceae, Oxelman & al.
2001), Suaeda (Chenopodiaceae, Schütze & al. 2003),
and Viscaria (Caryophyllaceae, Frajman & al. 2009b).
in addition, molecular phylogenies also have resulted in the identification and description of new taxa at all
levels, for example: Anacampserotaceae (nyffeler &
eggli 2010a); Microteaceae (Schäferhoff & al. 2009);
Didiereoideae, Portulacarioideae (Didiereaceae, applequist & Wallace 2003); Blossfeldieae (Cactaceae,
Butterworth 2006); Caribeae (Nyctaginaceae, douglas
& Spellenberg 2010); Eremogoneae (Caryophyllaceae,
Harbaugh & al. 2010); Gymnopodieae, Leptogoneae
(Polygonaceae, Burke & Sanchez 2011); Chenopodiastrum (Chenopodiaceae, Fuentes-Bazán & al. 2012b);
and Surreya (Amaranthaceae, Masson & Kadereit
2013).
the new data also demonstrate that developing a classification system for the order is a dynamic process. an
updated backbone at the generic level serves to present
the current state of knowledge. We believe that this is an
important step because many projects or researchers are
specifically dealing with certain genera. Building upon
a generic-level backbone will increase the efficiency of
implementing the next steps towards a synopsis at species level. For example, Oxelman & al. (2013) keep a
dynamically updated classification of Sileneae online.
the long-term aim is to provide a portal where taxonomic, chorologic, nomenclatural, and phylogenetic information can be retrieved, along with literature, dna
sequences and images. this resource can be a valuable
subproject for infrageneric and species-level taxonomy,
and also for various other biological research projects
where there is a strong need for a solid taxonomy based
on phylogenetic relationships in Sileneae (e.g. Bernasconi & al. 2009). Such initiatives will be strongly supported by the Caryophyllales network, also by providing
a sustained informatics infrastructure and a joint concept
for future monographic work (Borsch & al. 2015). the
published treatment of the genera of Caryophyllales,
produced directly from an edit-Platform database, will
provide a stepping stone for further refinement, also to
encourage further research and participation in the network. Members of the Caryophyllales network will be
able to correct and add to the information presented as it
Willdenowia 45 – 2015
289
Fig. 2. a: Achatocarpaceae: Phaulothamnus spinescens a. Gray, U.S.a., texas, 22 aug 2001, Borsch & al. 3446 (B, iSc), photo
by t. Borsch. – B – d: Aizoaceae: B: Tetragonia decumbens Mill., South africa, cape town, Muizenberg, 1 Mar 2015, photo by P.
Bruyns. – c: Cheiridopsis robusta n. e. Br., South africa, richtersveld, north of Port nolloth, Sep 1991, photo by W. Barthlott.
– d: Braunsia apiculata (Kensit) L. Bolus, South africa, Witteberg, 6 aug 2013, photo by P. Bruyns. – e & F: Amaranthaceae:
e: Gomphrena haenkeana Mart., Bolivia, Santa cruz, 6 apr 2003, Borsch & Ortuño 3627 (B, LPB), photo by t. Borsch. – F:
Pleuropetalum sprucei (Hook. f.) Standl., Germany, Botanischer Garten Berlin, photo by t. Borsch.
290
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
Fig. 3. a: Amaranthaceae: Tidestromia lanuginosa (nutt.) Standl., U.S.a., texas, Borsch & al. 3439 (B, iSc, MeXU), photo by t.
Borsch. – B – d: Cactaceae: B: Carnegiea gigantea (engelm.) Britton & rose, U.S.a., arizona, Organ Pipe cactus national Monument, 11 apr 1992, photo by W. Barthlott. – c: Opuntia ficusindica (L.) Mill., Spain, canarias, tenerife, near Orotava, Feb 1989,
photo by W. Barthlott. – d: Pereskia aculeata Mill., Monaco, Jardin exotique de Monaco, Jun 1998, photo by W. Barthlott. – e – G:
Caryophyllaceae: e: Arenaria cretica Spreng., Greece, Mt Olimbos, 29 Jul 2005, photo by n. turland. – F: Bolanthus creutzburgii
subsp. zaffranii Phitos & al., Greece, Kriti, Paleochora, 31 Mar 2009, Turland & al. 1841 (MO, PaL, UPa), photo by n. turland. –
G: Silene virginica L., U.S.a., tennessee, Great Smoky Mountains national Park, 25 Jun 2012, photo by n. turland.
Willdenowia 45 – 2015
291
Fig. 4. a: Caryophyllaceae: Dianthus androsaceus (Boiss. & Heldr.) Hayek, Greece, Mt taigetos, 14 Jul 2007, photo by n. turland.
– B–d: Chenopodiaceae: B: Allenrolfea occidentalis (S. Watson) Kuntze, U.S.a., texas, 23 aug 2001, Borsch & al. 3447 (B, iSc),
photo by t. Borsch. – c: Chenopodium quinoa Willd., Bolivia, departamento de La Paz, altiplano, 2010, photo by t. Borsch. – d:
Chenopodium vulvaria L., Germany, Botanischer Garten der Universität Mainz, aug 2015, photo by G. Kadereit. – e & F: Didieri
aceae: e: Alluaudia ascendens (drake) drake, Germany, Botanische Gärten der Universität Bonn, 2010, photo by n. Korotkova. – F:
Portulacaria namaquensis Sond., namibia, W of aussenkehr, 5 Jul 2013, photo by P. Bruyns. – G: Dioncophyllaceae: Triphyophyl
lum peltatum (Hutch. & dalziel) airy Shaw, côte d’ivoir, Parc national de tai, north of Mt niénokoué, 1998, photo by W. Barthlott.
292
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
Fig. 5. a & B: Droseraceae: a: Drosera cistiflora L., Germany, Botanische Gärten der Universität Bonn, Jan 2001, photo by W.
Barthlott. – B: Drosera cuneifolia L. f., South africa, cape town, table Mountain, 6 Jan 2008, photo by n. turland. – c: Halo
phytaceae: Halophytum ameghinoi (Speg.) Speg., ex Sukkulenten-Sammlung zürich, photo by t. Borsch. – d: Montiaceae: Clay
tonia virginica L., U.S.a., Missouri, rockpile Mountain Wilderness, 10 apr 2010, photo by n. turland. – e – G: Nyctaginaceae: e:
Bougainvillea spectabilis Willd., Germany, Botanische Gärten der Universität Bonn, Oct 1990, photo by W. Barthlott. – F: Guapira
rufescens (Heimerl) Lundell, cuba, Holguín, 1 Mar 2010, Borsch & al. 4273 (B, HaJB), photo by t. Borsch. – G: Pisonia aculeata
L., cuba, Holguín, 27 Feb 2010, Borsch & al. 4229 (B, HaJB), photo by t. Borsch.
Willdenowia 45 – 2015
293
Fig. 6. a & B: Phytolaccaceae: a: Rivina humilis L., Germany, Botanische Gärten der Universität Bonn, Borsch 3542 (BOnn),
photo by t. Borsch. – B: Trichostigma octandrum (L.) H. Walter, cuba, Villa clara, 1 Mar 2012, Borsch & al. 5265 (B, HaJB),
photo by t. Borsch. – c & d: Plumbaginaceae: c: Acantholimon androsaceum (Jaub. & Spach) Boiss., Greece, Kriti, Lefka Ori,
13 Jul 2006, photo by n. turland. – d: Armeria maritima (Mill.) Willd. subsp. maritima, U.K., devon, Foreland Point, 20 May
2008, photo by n. turland. – e – G: Polygonaceae: e: Coccoloba shaferi Britton, cuba, Holguín, 1 Mar 2010, Borsch & al. 4270
(B, HaJB) photo by t. Borsch. – F: Coccoloba uvifera (L.) L., cuba, Holguín, 1 Mar 2010, photo by t. Borsch. – G: Triplaris
americana L., Peru, San Martín, río Huallaga, 27 Jun 2009, photo by a. Sanchez.
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Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
Fig. 7. a: Sarcobataceae: Sarcobatus vermiculatus (Hook.) torr., Germany, Botanischer Garten Berlin, 29 aug 2015, photo by n.
turland. – B: Tamaricaceae: Tamarix ramosissima Ledeb., azerbaijan, Borsch & al. 5461 (B, BaK), photo by t. Borsch.
is databased. Once published, the continuously updated
dynamic treatment will also be available as a freely accessible online data portal (http://caryophyllales.org/).
Revised generic classification of Caryophyllales
Methodology and content
the names of genera listed follows the rules of nomenclature (Mcneill & al. 2012) and the family assignments
adhere to aPG iii (2009) and Stevens (2001 onwards),
if not noted otherwise. notes are added to many genera
providing information about the current state of knowledge in terms of monophyly or phylogenetic relationships. the data management is effected by means of the
edit Platform for cybertaxonomy software suite (Berendsohn 2010; Berendsohn & al. 2011). in a first step,
data from names in current Use 3 (Greuter & al. 1993)
were imported. additions and corrections were then incorporated particularly from Bittrich 1993b – d; Bittrich
& Kühn 1993; Brandbyge 1993; carolin 1993; endress
& Bittrich 1993; Kubitzki 1993a, b; Kühn 1993; rohwer
1993; Sperling & Bittrich 1993; townsend 1993; dickison 2003; Gaskin 2003; Kubitzki 2003a – e; Porembski
2003; Porembski & Barthlott 2003; Prance 2003, and
for Aizoaceae and Cactaceae, corrections were incorporated from Hartmann (2001a, b) and Hunt (2006) respectively; if not noted otherwise. data cited include the
generic name, its author citation and its nomenclatural
reference, the name of the type species, selected synonyms (including all names listed in ncU-3; Greuter &
al. 1993), and later publications with their respective nomenclatural data.
author names are abbreviated in conformity with
Brummitt & Powell (1992) and its updates online; titles
of serials in the nomenclatural reference citations are abbreviated in conformity with Bridson & al. (2004) and
the titles of monographs are abbreviated in conformity
with Stafleu & cowan (1976 – 1988) and their successors,
except that all components start with capital letters.
the name of the type species follows ncU-3 (Greuter & al. 1993); for genera not treated there, the names
were obtained from reviewing protologues, index
nominum Genericorum (inG; Farr & zijlstra 1996+),
tropicos (undated), or the international Plant names
index (iPni 2004+). to denote the taxon concept followed in the present publication, a “sec.” (secundum,
following, according to; Stearn 1992) reference is given
(see, e.g., Berendsohn 1997; Franz & cardona-duque
2013). this is a bibliographic citation of a (recent) paper or work giving the circumscription of the taxon (by
means of a description, synonymy and/or details of the
relationship to other taxa). in some cases, this is further
discussed in a note, particularly with reference to the
authors mentioned in the previous paragraph and later
publications. the text of the following section, classification, consists of direct output from the edit-Platform
database.
Willdenowia 45 – 2015
Classification
the families and genera are listed in alphabetical order,
with a single incertae sedis genus at the very end of the
list. each accepted name is given in bold and includes
the standardized information mentioned above. the homotypic and heterotypic synonyms are listed according
to the conventions in Willdenowia. Many names are followed by notes as mentioned above.
Achatocarpaceae Heimerl sec. aPG (2009).
a small family comprising two genera and 16 species
occurring in tropical america, from southeastern United
States to South america (Medina 2009). traditionally,
the family has been included in Phytolaccaceae s.l., but
its position as an independent lineage has been well supported by several molecular phylogenetic studies (Hilu &
al. 2003; Schäferhoff & al. 2009; Brockington & al. 2009,
2011), which also showed that the family is more closely related to the Amaranthaceae/Chenopodiaceae clade
rather than to Phytolaccaceae. Achatocarpaceae are characterized by having unisexual flowers, the gynoecium with
two connate carpels, unilocular ovaries with two styles and
a single ovule, berrylike fruits and pollen with obscure
pores (Martínez-García 1985; Lipscomb 2003).
Achatocarpus triana in ann. Sci. nat., Bot., ser. 4, 9:
45. 1858 sec. Bittrich (1993b). – type: Achatocarpus
nigricans triana
Phaulothamnus a. Gray in Proc. amer. acad. arts 20:
293. 1885 sec. Bittrich (1993b). – type: Phaulotham
nus spinescens a. Gray – Fig. 2a.
Agdestidaceae nakai sec. cuénoud & al. (2002).
a monotypic family distributed from southern United
States to nicaragua (rohwer 1993a), introduced and naturalized in Florida and the antilles and cultivated as ornamentals in South america (rzedowski & calderón 2000).
traditionally, Agdestis was placed in Phytolaccaceae, subfamily Agdestioideae (e.g. rohwer 1993a; Stevens 2001
onwards; nienaber & thieret 2003), but several molecular
phylogenetic studies have shown that it represents a wellsupported independent lineage (cuénoud & al. 2002; Hilu
& al. 2003; Schäferhoff & al. 2009). these studies also
showed a close but only moderately supported relationship of the family with Sarcobataceae. Agdestidaceae are
climbers and characterized by paniculate inflorescences,
semi-inferior ovaries and cypselas crowned by winglike
sepals (nienaber & thieret 2003).
Agdestis Moc. & Sessé ex dc., Syst. nat. 1: 511, 543.
1817 sec. rohwer (1993a). – type: Agdestis clemati
dea Moc. & Sessé ex dc.
Monotypic; southern United States, Mexico, and
central america.
Aizoaceae Martinov sec. aPG (2009).
the Aizoaceae have a worldwide distribution throughout the tropics and subtropics (Hartmann 2001a, b).
295
However, the centres of diversity are in the southwestern part of africa (Bittrich 1986; Jürgens 1986;
Hartmann 1991). relatively few genera occur outside
of southern africa, mainly those from subfamilies Aizo
oideae, Sesuvioideae and Tetragonioideae. in contrast,
Mesembryanthemoideae and Ruschioideae are largely
restricted to southern africa with few species found outside of this area (e.g. Mesembryanthemum crystallinum
L., M. nodiflorum L. and Carpobrotus, Delosperma, Di
sphyma and Sarcozona species) (Hartmann 2001a, b).
the family consists predominantly of succulent (mostly
leaf succulent), annual to perennial herbs, subshrubs or
shrubs, with undifferentiated perianth or biseriate with
petals of staminodial origin, with mostly hygrochastic
loculicidal fruits.
Acrodon n. e. Br. in Gard. chron., ser. 3, 81: 12. 1927
sec. Hartmann (2001a). – type: Acrodon bellidiflorus
(L.) n. e. Br.
Acrosanthes eckl. & zeyh. in enum. Pl. afric. austral. [ecklon & zeyher]: 328. 1837 sec. Hartmann
(2001a). – type: Acrosanthes anceps Sond.
= Aizoon subg. Acrosanthes (eckl. & zeyh.) d. dietr.,
Syn. Pl. 3: 130. 1842.
= Didaste e. Mey. ex Harv. & Sond., Fl. cap. 2: 472.
1862, nom. nud.
Aizoanthemum dinter ex Friedrich in Mitt. Bot. Staatssamml. München 2: 343. 1957 sec. Hartmann (2001a).
– type: Aizoanthemum membrumconnectens dinter
ex Friedrich
Aizoon L., Sp. Pl. 1: 488. 1753 sec. Hartmann (2001a) ≡
Veslingia Heist. ex Fabr., enum.: 201. 1759. – type:
Aizoon canariense L.
Aloinopsis Schwantes in z. Sukkulentenk. 2: 177. 1926
sec. Hartmann (2001a) ≡ Acaulon n. e. Br. in J. Bot.
66: 76. 1928 ≡ Aistocaulon Poelln. ex H. Jacobsen,
Succ. Pl.: 123. 1935. – type: Aloinopsis aloides
(Haw.) Schwantes
the Aloinopsis clade includes several small genera
(ranging from one to six species), i.e. Aloinopsis,
Deilanthe, Nananthus, Pleiospilos, Prepodesma,
Rabiea and Tanquana (Klak & al. 2013). the group
is found outside the winter-rainfall region of South
africa. the status and generic placement of numerous species in this group has been subject to many
changes. For example, the monotypic Prepodesma
has been included in five different genera by different taxonomic treatments. Aloinopsis, Nananthus and
Rabiea are particularly poorly known in terms of species delimitation.
Amphibolia L. Bolus in J. S. african Bot. 31: 169. 1965
sec. Hartmann (2001a). – type: Amphibolia maritima
L. Bolus
Antegibbaeum Schwantes ex c. Weber in Baileya 16: 10.
1968 sec. Hartmann (2001a). – type: Antegibbaeum
fissoides (Haw.) c. Weber
a monotypic genus, which is endemic to the Little
Karoo, South africa. the placement of this genus as
296
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
sister to Cylindrophyllum remains statistically poorly
supported (Klak & al. 2013).
Antimima n. e. Br. in Gard. chron., ser. 3, 87: 211. 1930
sec. Hartmann (2001a). – type: Antimima dualis (n.
e. Br.) n. e. Br.
a large genus of 96 species, which has never been
revised. Most species were previously placed in
Ruschia, but separated from the latter based on fruit
characters. Hartmann (2001a) recognized five subgenera within Antimima, but did not indicate which
species belong to which subgenus. the molecular
study by Klak & al. (2013) suggests that Antimima is
not monophyletic in its current circumscription. a detailed morphological and molecular study is needed
to establish generic boundaries within the Antimima
clade, in which several other smaller genera such as
Brausia, Hammeria, Smicrostigma and Zeuktophyl
lum take part (Klak & al. 2013).
Apatesia n. e. Br. in Gard. chron., ser. 3, 81: 12. 1927 sec.
Hartmann (2001a). – type: Apatesia pillansii n. e. Br.
Arenifera a. G. J. Herre in Sukkulentenk. 2: 35. 1948
sec. Hartmann (2001a). – type: Arenifera pillansii (L.
Bolus) a. G. J. Herre
there are four species included in Arenifera (Hartmann 2001a). Since this group was not sampled by
Klak & al. (2013), its phylogenetic position within
the tribe Ruschieae remains uncertain.
Argyroderma n. e. Br. in Gard. chron., ser. 3, 71: 92.
1922 sec. Hartmann (2001a). – type: Argyroderma
testiculare (aiton) n. e. Br.
= Roodia n. e. Br. in Fl. Pl. South africa 2: 78. 1922.
Astridia dinter in Gard. chron., ser. 3, 80: 430. 1926 sec.
Hartmann (2001a). – type: Astridia velutina dinter
Bergeranthus Schwantes in z. Sukkulentenk. 2: 179.
1926 sec. Hartmann (2001a). – type: Bergeranthus
scapigerus (Haw.) Schwantes
Bijlia n. e. Br. in J. Bot. 66: 267. 1928 sec. Hartmann
(2001a). – type: Bijlia cana (Haw.) n. e. Br.
= Bolusanthemum Schwantes in Gartenwelt 32: 514.
1928.
Braunsia Schwantes in Gartenwelt 32: 644. 1928 sec.
Hartmann (2001a). – type: Braunsia nelii Schwantes
– Fig. 2d.
= Echinus L. Bolus in Fl. Pl. South africa 7: 266. 1927,
nom. illeg.
Brianhuntleya chesselet, S. a. Hammer & i. Oliver in
Bothalia 33: 161. 2003 sec. chesselet & al. (2003). –
type: Brianhuntleya intrusa (Kensit) chesselet, S. a.
Hammer & i. Oliver
a monotypic genus, from the Worcester-robertson
Karoo (South africa). its sister relationship to Bijlia
(two species) remains poorly supported (Klak & al.
2013).
Calamophyllum Schwantes in z. Sukkulentenk. 3: 15,
28. 1927 sec. Hartmann (2001a). – type: Calamo
phyllum teretifolium (Haw.) Schwantes
a mysterious genus including three species. these
were described by Haworth between 1795 and 1812.
However, for two of the names no types have been
selected yet, whereas for the third a drawing by duncansan serves as a lectotype (Hartmann 2001a). the
distribution of the genus is uncertain.
Carpanthea n. e. Br. in Gard. chron., ser. 3, 78: 412.
1925 sec. Hartmann (2001a). – type: Carpanthea
pomeridiana (L.) n. e. Br.
= Macrocaulon n. e. Br. in Gard. chron., ser. 3, 81: 12.
1927.
Carpobrotus n. e. Br. in Gard. chron., ser. 3, 78: 433.
1925 sec. Hartmann (2001a) ≡ Abryanthemum neck.,
elem. Bot. 2: 82. 1790, nom. inval. – type: Carpo
brotus edulis (L.) L. Bolus
Carruanthus (Schwantes) Schwantes in z. Sukkulentenk. 3: 106. 1927 sec. Hartmann (2001a) ≡ Berge
ranthus subg. Carruanthus Schwantes in z. Sukkulentenk. 2: 180. 1926. – type: Carruanthus caninus
(Lam.) Schwantes
= Tischleria Schwantes in Sukkulentenk. 4: 78. 1951.
Cephalophyllum n. e. Br. in Gard. chron., ser. 3, 78:
433. 1925 sec. Hartmann (2001a). – type: Cephalo
phyllum tricolorum (Haw.) n. e. Br.
Cerochlamys n. e. Br. in J. Bot. 66: 171. 1928 sec. Hartmann (2001a). – type: Cerochlamys trigona n. e. Br.
Chasmatophyllum dinter & Schwantes in z. Sukkulentenk. 3: 14, 17. 1927 sec. Hartmann (2001a). – type:
Chasmatophyllum musculinum (Haw.) dinter &
Schwantes
Cheiridopsis n. e. Br. in Gard. chron., ser. 3, 78: 433.
1925 sec. Hartmann (2001a). – type: Cheiridopsis tu
berculata (Mill.) n. e. Br. – Fig. 2c.
Cheiridopsis was found to be closely related to Ih
lenfeldtia and Odontophorus (Klak & al. 2013). in
addition, one of the three subgenera of Cheiridopsis,
C. subg. Odontophoroides, could be more closely
related to Odontophorus than to the remainder of
Cheiridopsis (Hartmann 2001b). although Cheiri
dopsis and Odontophorus were revised at species level (Hartmann 1976; Hartmann & dehn 1987), their
generic limits need to be reinvestigated.
Circandra n. e. Br. in Gard. chron., ser. 3, 87: 126. 1930
sec. Hartmann (2001a). – type: Circandra serrata
(L.) n. e. Br.
a monotypic genus; its only species was already
known to Linnaeus as Mesembryanthemum serratum
L. the area where it was previously recorded has been
subject to extensive cultivation, so the species had
been thought to be extinct. However, it was rediscovered in 2007 and is currently considered as critically
endangered (Klak & Low 2007). the hypanthium
found in the flowers suggests a close relationship with
Erepsia, where this species had been included previously (as E. serrata (L.) L. Bolus).
Cleretum n. e. Br. in Gard. chron., ser. 3, 78: 412. 1925
sec. Klak & Bruyns (2012). – type: Cleretum papu
losum (L. f.) L. Bolus
Willdenowia 45 – 2015
= Dorotheanthus Schwantes in Möller’s deutsche
Gärtn.-zeitung 42: 283. 1927.
= Aethephyllum n. e. Br. in Möller’s deutsche Gärtn.zeitung 43: 400. 1928.
= Pherolobus n. e. Br. in Möller’s deutsche Gärtn.-zeitung 43: 400. 1928.
= Sineoperculum Van Jaarsv. in J. S. african Bot. 48: 5.
1982.
Conicosia n. e. Br. in Gard. chron., ser. 3, 78: 433. 1925
sec. Hartmann (2001a). – type: Conicosia pugioni
formis (L.) n. e. Br.
= Herrea Schwantes in Möller’s deutsche Gärtn.-zeitung 42: 436. 1927.
Conophytum n. e. Br. in Gard. chron., ser. 3, 71: 19.
1922 sec. Hammer (2001). – type: Conophytum
minutum (Haw.) n. e. Br.
= Derenbergia Schwantes in z. Sukkulentenk. 2: 137.
1925.
= Ophthalmophyllum dinter & Schwantes in Möller’s
deutsche Gärtn.-zeitung 42: 64. 1927.
= Herreanthus Schwantes in Gartenwelt 32: 514. 1928.
= Berresfordia L. Bolus, notes Mesembryanthemum 2:
313. 1930.
Corpuscularia Schwantes in z. Sukkulentenk. 2: 185.
1926 sec. Hartmann (2001a) ≡ Schonlandia L. Bolus
in Fl. Pl. South africa 7: 259. 1927. – type: Corpus
cularia lehmannii (eckl. & zeyh.) Schwantes
the genus includes eight species and is endemic to
the eastern cape, South africa. its phylogenetic position near Delosperma has been confirmed (Klak & al.
2013), but the genus lacks a taxonomic revision.
Cylindrophyllum Schwantes in z. Sukkulentenk. 3: 15,
28. 1927 sec. Hartmann (2001a). – type: Cylindro
phyllum calamiforme (L.) Schwantes
Cylindrophyllum includes five species (Hartmann
2001a), but lacks a taxonomic revision.
Cypselea turpin in ann. Mus. natl. Hist. nat. 7: 219. 1806
sec. Hartmann (2001a) ≡ Radiana raf., Specchio Sci.
1: 88. 1814. – type: Cypselea humifusa turpin
= Millegrana Juss. ex turpin in ann. Mus. natl. Hist.
nat. 7: 220. 1806, nom. nud.
Cypselea includes three species (Hartmann 2011a)
and is nested within Sesuvium (Bohley & al. 2015;
Hassan & al. 2005; thulin & al. 2012). two of the
species are endemic to Paraguay and cuba, respectively (Hartmann 2011a).
Deilanthe n. e. Br. in Gard. chron., ser. 3, 88: 278. 1930
sec. Hartmann (2001a). – type: Deilanthe peersii (L.
Bolus) n. e. Br.
Delosperma n. e. Br. in Gard. chron., ser. 3, 78: 412.
1925 sec. Hartmann (2001a). – type: Delosperma
echinatum (Lam.) Schwantes
a large genus of 142 species, which has never been
revised. the study by Klak & al. (2013) suggests that
Delosperma is not monophyletic in its current circumscription. a detailed morphological and molecular study is needed to establish generic boundaries
297
within the Delosperma clade, in which several other
smaller genera, including Corpuscularia, Ectotropis,
Frithia, Mestoklema and Trichidiadema take part
(Klak & al. 2013).
Dicrocaulon n. e. Br. in J. Bot. 66: 141. 1928 sec. ihlenfeldt (2001a). – type: Dicrocaulon pearsonii n.
e. Br.
Didymaotus n. e. Br. in Gard. chron., ser. 3, 78: 433.
1925 sec. Hartmann (2001a). – type: Didymaotus la
pidiformis (Marloth) n. e. Br.
the phylogenetic position of this monotypic genus
remains largely unresolved (Klak & al. 2013).
Dinteranthus Schwantes in z. Sukkulentenk. 2: 184.
1926 sec. Hartmann (2001a). – type: Dinteranthus
microspermus (dinter & derenb.) Schwantes
Diplosoma Schwantes in z. Sukkulentenk. 2: 179. 1926
sec. ihlenfeldt (2001b). – type: Diplosoma retrover
sum (Kensit) Schwantes
= Maughania n. e. Br. in J. cact. Succ. Soc. amer. 2:
389. 1931.
= Maughaniella L. Bolus in J. S. african Bot. 28: 264.
1962.
Disphyma n. e. Br. in Gard. chron., ser. 3, 78: 433. 1925
sec. Hartmann (2001a). – type: Disphyma crassifo
lium (L.) L. Bolus
Dracophilus dinter & Schwantes in Möller’s deutsche
Gärtn.-zeitung 42: 187. 1927 sec. Hartmann (2001a)
≡ Juttadinteria subg. Dracophilus Schwantes in z.
Sukkulentenk. 2: 183. 1926. – type: Dracophilus de
laetianus (dinter) dinter & Schwantes
Drosanthemum Schwantes in z. Sukkulentenk. 3: 14,
29. 1927 sec. Hartmann (2001a). – type: Drosanthe
mum hispidum (L.) Schwantes
a large genus of 107 species, which has only partly
been revised. With the exception of few misplaced
species, the genus is thought to be monophyletic (Klak
& al. 2003b; Klak & al. 2013). Hartmann (2007) recognized eight subgenera in Drosanthemum and also
provided a key to the subgenera with a list of species
included for each of them. although also distribution
maps were included for all eight subgenera, no vouchers were cited (Hartmann 2007), so that it remains uncertain on which material the maps were based. in addition, only one of the eight subgenera has so far been
revised in part (Hartmann 2008). Since many species
are threatened by agriculture or urban expansion, the
genus is in urgent need of revision.
Eberlanzia Schwantes in z. Sukkulentenk. 2: 189. 1926
sec. Hartmann (2001a). – type: Eberlanzia clausa
(dinter) Schwantes
Eberlanzia includes eight species (Hartmann 2001a).
However, the two species sampled by Klak & al.
(2013) do not group together, suggesting that the genus is not monophyletic in its current circumscription.
Ebracteola dinter & Schwantes in z. Sukkulentenk. 3:
15, 24. 1927 sec. Hartmann (2001a). – type: Ebracte
ola montismoltkei (dinter) dinter & Schwantes
298
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
Ectotropis n. e. Br. in Gard. chron., ser. 3, 81: 12. 1927
sec. Hartmann (2001a). – type: Ectotropis alpina n.
e. Br.
Enarganthe n. e. Br. in Gard. chron., ser. 3, 87: 71.
1930 sec. Hartmann (2001a). – type: Enarganthe oc
tonaria (L. Bolus) n. e. Br.
a monotypic genus, which is endemic to namqualand. relationships to other members in the Conophy
tum clade remain uncertain.
Erepsia n. e. Br. in Gard. chron., ser. 3, 78: 433. 1925
sec. Hartmann (2001a). – type: Erepsia inclaudens
(Haw.) Schwantes
= Piquetia n. e. Br. in Gard. chron., ser. 3, 78: 433.
1925, nom. illeg.
= Semnanthe n. e. Br. in Gard. chron., ser. 3, 81: 12.
1927.
= Kensitia Fedde in repert. Spec. nov. regni Veg. 48:
11. 1940.
Esterhuysenia L. Bolus in S. african J. Bot. 33: 308.
1967 sec. Hartmann (2001a). – type: Esterhuysenia
alpina L. Bolus
a small genus with five species endemic to the cape
region of South africa. See further notes under Ham
meria.
Faucaria Schwantes in z. Sukkulentenk. 2: 176. 1926
sec. Groen & Hartmann (2001). – type: Faucaria tig
rina (Haw.) Schwantes
Fenestraria n. e. Br. in Gard. chron., ser. 3, 78: 433.
1925 sec. Hartmann (2001b). – type: Fenestraria au
rantiaca n. e. Br.
a monotypic genus from namaqualand. See further
notes under Cephalophyllum.
Frithia n. e. Br. in Gard. chron., ser. 3, 78: 433. 1925
sec. Hartmann (2001b). – type: Frithia pulchra n.
e. Br.
this bitypic genus was found to be closely related to
Delosperma (Klak & al. 2013). See further remarks
under Delosperma.
Galenia L., Sp. Pl. 1: 359. 1753 sec. Hartmann (2001b).
– type: Galenia africana L.
= Kolleria c. Presl., Symb. Bot. 1: 23. 1831.
= Sialodes eckl. & zeyh., enum. Pl. afric. austral. 3:
329. 1837.
= Tephras e. Mey. ex Harv. & Sond., Fl. cap. 2: 477.
1862.
Gibbaeum Haw. ex n. e. Br. in Gard. chron., ser. 3, 71:
129. 1922 sec. Hartmann (2001b). – type: Gibbaeum
pubescens (Haw.) n. e. Br.
= Rimaria n. e. Br. in Gard. chron., ser. 3, 78: 413.
1925.
= Argeta n. e. Br. in Gard. chron., ser. 3, 82: 113.
1927.
= Imitaria n. e. Br. in J. Bot. 65: 348. 1927.
= Mentocalyx n. e. Br. in Gard. chron., ser. 3, 81: 251.
1927.
= Muiria n. e. Br. in Gard. chron., ser. 3, 81: 116.
1927.
the monotypic Muiria was placed in Gibbaeum, as
G. hortenseae (n. e. Br.) thiede & Klak, sec. Goldblatt & Manning (2000). the species was confirmed
to be closely related to Gibbaeum, but its relationship to other species in the genus remains unresolved
(Klak & al. 2013).
Glottiphyllum Haw. ex n. e. Br. in Gard. chron., ser. 3,
70: 311. 1921 sec. Hartmann (2001b). – type: Glot
tiphyllum linguiforme (L.) n. e. Br.
Gunniopsis Pax, nat. Pflanzenfam. 3(lb): 44. 1889 sec.
Hartmann (2001b) ≡ Aizoon subg. Gunniopsis Pax &
K. Hoffm. in J. S. african Bot. Soc. 25: 30. 1959. –
type: Gunniopsis quadrifaria Pax
= Gunnia F. Muell., rep. Pl. Babbage’s exped.: 9.
1859, nom. illeg.
= Neogunnia Pax & K. Hoffm., nat. Pflanzenfam. (ed.
2) 16c: 225. 1934.
the genus is endemic to australia.
Hallianthus H. e. K. Hartmann in Bot. Jahrb. Syst. 114:
167. 1992 sec. Hartmann (2001b). – type: Hallian
thus planus (L. Bolus) H. e. K. Hartmann
Hammeria Burgoyne in cact. Succ. J. (Los angeles)
70(4): 204. 1998 sec. Hartmann (2001b). – type:
Hammeria salteri (L. Bolus) Burgoyne
a small genus consisting of only three species. the
two species included in the molecular study by Klak
& al. (2013) were not resolved as sisters. However,
they were shown to group with other small genera
such as Braunsia and Esterhuysenia in the Antimima
clade (Klak & al. 2013).
Hartmanthus S. a. Hammer in Haseltonia 3: 79. 1995
sec. Hartmann (2001b). – type: Hartmanthus perga
mentaceus (L. Bolus) S. a. Hammer
Hereroa (Schwantes) dinter & Schwantes in z. Sukkulentenk. 3: 15, 23. 1927 sec. Hartmann (2001b) ≡
Bergeranthus subg. Hereroa Schwantes in z. Sukkulentenk. 2: 180. 1926. – type: Hereroa puttkammeri
ana (dinter & Berger) dinter & Schwantes
Hereroa includes 27 species but lacks a taxonomic
revision. the study by Klak & al. (2013) reveals
Rhombophyllum (five species) and Bergeranthus (ten
species) as its closest relatives. denser sampling may
in addition show that Hereroa is not monophyletic,
with Rhombophyllum likely to be nested within it.
On account of the close morphological resemblance
between these genera, generic limits need to be critically reinvestigated.
Hymenogyne Haw., revis. Pl. Succ.: 192. 1821 sec.
Hartmann (2001b). – type: Hymenogyne glabra (aiton) Haw.
= Thyrasperma n. e. Br. in Gard. chron., ser. 3, 78:
412. 1925.
Ihlenfeldtia H. e. K. Hartmann in Bot. Jahrb. Syst. 114:
47. 1992 sec. Hartmann (2001b). – type: Ihlenfeldtia
excavata (L. Bolus) H. e. K. Hartmann
the two species currently included in Ihlenfeldtia
were previously included in Cheiridopsis. However,
Willdenowia 45 – 2015
the two species were moved each into their own genus and thought to be closely related to Tanquana
(three species) and Vanheerdia (two species), based
on characters of the fruits (Hartmann 1992). However Klak & al. (2013) confirmed the previous position
of Ihlenfeldtia as a close relative of Cheiridopsis,
which is supported by characteristics of the leaves
(Hartmann 1992). See further notes under Cheiri
dopsis.
Jacobsenia L. Bolus & Schwantes, notes Mesembryanthemum 3: 255. 1954 sec. ihlenfeldt (2001c). – type:
Jacobsenia kolbei (L. Bolus) L. Bolus & Schwantes
= Anisocalyx L. Bolus, notes Mesembryanthemum 3:
385. 1958, nom. illeg.
= Drosanthemopsis rauschert in taxon 31: 555. 1982.
although Jacobsenia currently includes only three
species, they were shown not to be monophyletic
(Klak & al. 2013).
Jensenobotrya a. G. J. Herre in Sukkulentenk. 4: 79.
1951 sec. Hartmann (2001b). – type: Jensenobotrya
lossowiana a. G. J. Herre
Jordaaniella H. e. K. Hartmann in Biblioth. Bot. 136:
57. 1983 sec. Hartmann (2001b). – type: Jordaan
iella clavifolia (L. Bolus) H. e. K. Hartmann
Juttadinteria Schwantes in z. Sukkulentenk. 2: 182.
1926 sec. Hartmann (2001b). – type: Juttadinteria
kovisimontana (dinter & a. Berger) Schwantes
Khadia n. e. Br. in Gard. chron., ser. 3, 88: 279. 1930
sec. Hartmann & chesselet (2001). – type: Khadia
acutipetala (n. e. Br.) n. e. Br.
Knersia H. e. K. Hartmann & Liede in Bradleya 31: 126.
2013 sec. Hartmann & Liede-Schumann (2013). –
type: Knersia diversifolia (L. Bolus) H. e. K. Hartmann & Liede
a monotypic genus, which was recently erected to
accommodate a species previously misplaced in Dro
santhemum (Klak & al. 2013; Hartmann & LiedeSchumann 2013).
Lampranthus n. e. Br. in Gard. chron., ser. 3, 87: 71.
1930, nom. cons. sec. Hartmann (2001b). – type:
Lampranthus multiradiatus (Jacq.) n. e. Br.
= Aristanthus Schwantes in z. Sukkulentenk. 3: 28.
1827.
= Mesembryanthus necker ex rothm. in notizbl. Bot.
Gart. Berlin-dahlem 15: 413. 1941, nom. inval.
Lampranthus is a large genus of 194 species, which
has never been revised. a molecular study of the
Lampranthus group identified a core of closely related species, which makes up the current genus (Klak
& al. 2003a). Groups of species not closely related to
Lampranthus s.str. were placed in other genera, with
some placed in new genera (Klak 2005).
Lapidaria (dinter & Schwantes) n. e. Br. in Gard.
chron., ser. 3, 84: 472. 1928 sec. Hartmann (2001b)
≡ Dinteranthus subg. Lapidaria dinter & Schwantes
in Möller’s deutsche Gärtn.-zeitung 42: 223. 1927.
– type: Lapidaria margaretae (Schwantes) n. e. Br.
299
a monotypic genus, which was shown to be sister to
Dinteranthus (Klak & al. 2013), where it had been
placed previously. the two genera form a clade together with Lithops and Schwantesia (Klak & al.
2013).
Leipoldtia L. Bolus in Fl. Pl. South africa 7: t. 256. 1927
sec. Hartmann (2001b). – type: Leipoldtia constricta
(L. Bolus) L. Bolus
= Rhopalocyclus Schwantes in Gartenwelt 32: 599.
1928.
Lithops n. e. Br. in Gard. chron., ser. 3, 71: 44. 1922
sec. cole & cole (2001). – type: Lithops lesliei (n.
e. Br.) n. e. Br.
Lithops is one of the best-known genera among collectors of succulents. Species and subspecies are
largely distinguished by the colour and markings
present on the flattened leaf tops. the genus was
shown to be closely related to Dinteranthus, Lapi
daria and Schwantesia by Klak & al. (2013). in view
of the close morphological resemblance between the
four genera in terms of fruit and floral characters, it
needs to be reinvestigated whether all of the genera
should be maintained.
Machairophyllum Schwantes in Möller’s deutsche
Gärtn.-zeitung 42: 187. 1927 sec. Hartmann (2001b).
– type: Machairophyllum albidum (L.) Schwantes
= Perissolobus n. e. Br. in Gard. chron., ser. 3, 88: 278.
1930.
Malephora n. e. Br. in Gard. chron., ser. 3, 81: 12. 1927
sec. Hartmann (2001b). – type: Malephora mollis
(aiton) n. e. Br.
= Crocanthus L. Bolus in Fl. Pl. South africa 7: 255.
1927.
= Hymenocyclus Schwantes in Möller’s deutsche
Gärtn.-zeitung 42: 27. 1927.
the genus includes 16 species, but lacks a taxonomic
revision. Since the group is rather homogenous, further sampling is likely to confirm the monophyly of
the genus with the species currently included.
Marlothistella Schwantes in Gartenwelt 32: 599. 1928
sec. Hartmann (2001b). – type: Marlothistella union
dalensis Schwantes
Mesembryanthemum L., Sp. Pl. 1: 480. 1753, nom. cons.
sec. Klak & al. (2007). – type: Mesembryanthemum
nodiflorum L.
= Brownanthus Schwantes in z. Sukkulentenk. 3: 14, 20.
1827 ≡ Trichocyclus n. e. Br. in Bothalia 1(3): 151.
1922, nom. illeg.
= Aptenia n. e. Br. in Gard. chron., ser. 3, 78: 412. 1925.
= Aridaria n. e. Br. in Gard. chron., ser. 3, 78: 433.
1925.
= Aspazoma n. e. Br. in Gard. chron., ser. 3, 78: 413.
1925.
= Dactylopsis n. e. Br. in Gard. chron., ser. 3, 78: 413.
1925.
= Phyllobolus n. e. Br. in Gard. chron., ser. 3, 78: 413.
1925.
300
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
= Prenia n. e. Br. in Gard. chron., ser. 3, 78: 412. 1925.
= Psilocaulon n. e. Br. in Gard. chron., ser. 3, 78: 433.
1925.
= Sceletium n. e. Br. in Gard. chron., ser. 3, 78: 412.
1925.
= Synaptophyllum n. e. Br. in Gard. chron., ser. 3, 78:
412. 1925.
= Pseudobrownanthus ihlenf. & Bittrich in Bot. Jahrb.
Syst. 105: 319. 1985.
= Caulipsolon Klak, ill. Handb. Succ. Pl. Aizoaceae
a – e: 103. 2002.
= Volkeranthus Gerbaulet in Bradleya 30: 196. 2012.
a new infrageneric classification has been proposed by
Klak & Bruyns (2013). a broad generic circumscription for Mesembryanthemum has been reaffirmed and
Mesembryanthemum subdivided into five subgenera,
with all five shown to be monophyletic. two species
were recently reinstated and shown to form part of subgenus Volkeranthus, which is sister to the remainder of
Mesembryanthemum (Klak & al. 2014). thus, Mesem
bryanthemum currently includes 105 species.
Mestoklema n. e. Br. ex Glen in Bothalia 13: 454. 1981
sec. Hartmann (2001b). – type: Mestoklema tubero
sum (L.) n. e. Br. ex Glen
See notes under Delosperma.
Meyerophytum Schwantes in Möller’s deutsche
Gärtn.-zeitung 42: 436. 1927 sec. ihlenfeldt
(2001d). – type: Meyerophytum meyeri (Schwantes)
Schwantes
= Depacarpus n. e. Br. in Gard. chron., ser. 3, 87: 71.
1930.
Mitrophyllum Schwantes in z. Sukkulentenk. 2: 181.
1926 sec. Hartmann (2001b). – type: Mitrophyllum
mitratum (Marloth) Schwantes
= Conophyllum Schwantes in z. Sukkulentenk. 3: 321.
1928.
= Mimetophytum L. Bolus, notes Mesembryanthemum
3: 252. 1954.
Monilaria Schwantes in Gartenwelt 33: 69. 1929 sec.
ihlenfeldt (2001e). – type: Monilaria chrysoleuca
(Schltr.) Schwantes
Mossia n. e. Br. in Gard. chron., ser. 3, 87: 71. 1930 sec.
Hartmann (2001b). – type: Mossia intervallaris (L.
Bolus) n. e. Br.
Namaquanthus L. Bolus, notes Mesembryanthemum 3: 257. 1954 sec. Hartmann (2001b). – type:
Namaquanthus vanheerdei L. Bolus
Namibia dinter & Schwantes in z. Sukkulentenk. 3:
106. 1927 sec. Hartmann (2001b) ≡ Juttadinteria
subg. Namibia Schwantes in z. Sukkulentenk. 2: 184.
1926. – type: Namibia cinerea dinter & Schwantes
Nananthus n. e. Br. in Gard. chron., ser. 3, 78: 433.
1925 sec. Hartmann (2001b). – type: Nananthus vit
tatus (n. e. Br.) Schwantes
Nelia Schwantes in Möller’s deutsche Gärtn.-zeitung
43: 92. 1928 sec. Hartmann (2001b). – type: Nelia
meyeri Schwantes
= Sterropetalum n. e. Br. in Gard. chron., ser. 3, 83:
266. 1928.
Neohenricia L. Bolus in J. S. african Bot. 4: 51. 1938
sec. Hartmann (2001b) ≡ Henricia L. Bolus, notes
Mesembryanthemum 3: 39. 1936. – type: Neohen
ricia sibbettii (L. Bolus) L. Bolus
a small genus including only two species. See further
remarks under Stomatium.
Octopoma n. e. Br. in Gard. chron., ser. 3, 87: 72. 1930
sec. Hartmann (2001b). – type: Octopoma octojuge
(L. Bolus) n. e. Br.
Octopoma has been recognized by several authors
(Hartmann 2001b) and Klak & al. (2013). However,
the two infrageneric groups distinguished on account
of differences in fruit morphology (Hartmann 2001b)
were not confirmed by Klak & al. (2013).
Odontophorus n. e. Br. in Gard. chron., ser. 3, 81: 12.
1927 sec. Hartmann (2001b). – type: Odontophorus
marlothii n. e. Br.
See remarks under Cheiridopsis.
Oophytum n. e. Br. in Gard. chron., ser. 3, 78: 413.
1925 sec. ihlenfeldt (2001f). – type: Oophytum ovi
forme (n. e. Br.) n. e. Br.
Orthopterum L. Bolus in S. african Gard. 17: 281. 1927
sec. Hartmann (2001b). – type: Orthopterum wal
toniae L. Bolus
Orthopterum, comprising two species, is closely allied to Faucaria (Klak & al. 2013), from which it
mainly differs by the repeatedly opening and closing
fruits (Hartmann 2001b).
Oscularia Schwantes in Möller’s deutsche Gärtn.-zeitung 42: 187. 1927 sec. Hartmann (2001b). – type:
Oscularia deltoides (L.) Schwantes
Ottosonderia L. Bolus in notes Mesembryanthemum [H.
M. L. Bolus] 3: 292. 1958 sec. Hartmann (2001b). –
type: Ottosonderia monticola (Sond.) L. Bolus
a monotypic genus from namaqualand, which was
shown to be closely allied to Jordaaniella and to
Ruschia sandbergensis L. Bolus (Klak & al. 2013).
However, relationships to other members of the xeromorphic winter-rainfall clade remain in many parts
poorly resolved.
Peersia L. Bolus in Fl. Pl. South africa 7: t. 264. 1927
sec. Hartmann (2001b). – type: Peersia macradenia
(L. Bolus) L. Bolus
a small genus of only three species, which was shown
to be closely allied to Rhinephyllum (Klak & al. 2013),
where all three species were previously placed.
Phiambolia Klak in Bradleya 21: 112. 2003 sec. Klak
(2003). – type: Phiambolia hallii (L. Bolus) Klak
Pleiospilos n. e. Br. in Gard. chron., ser. 3, 78: 433.
1925 sec. Hartmann (2001b). – type: Pleiospilos bo
lusii (Hook. f.) n. e. Br.
= Punctillaria n. e. Br. in Gard. chron., ser. 3, 78: 433.
1925.
Plinthus Fenzl in nov. Stirp. dec.: 52. 1839 sec. Hartmann (2001b). – type: Plinthus cryptocarpus Fenzl
Willdenowia 45 – 2015
Polymita n. e. Br. in Gard. chron., ser. 3, 87: 72. 1930
sec. Hartmann (2001b). – type: Polymita pearsonii
n. e. Br.
a small genus including only two species. it is
closely allied to Schlechteranthus (Klak & al. 2013),
which also only incorporates two species. as indicated by the molecular analysis by Klak & al.
(2013), the generic limits need to be critically reinvestigated.
Prepodesma n. e. Br. in Gard. chron., ser. 3, 88: 279.
1930 sec. Hartmann (2001b). – type: Prepodesma
orpenii (n. e. Br.) n. e. Br.
Psammophora dinter & Schwantes in z. Sukkulentenk.
2: 188. 1926 sec. Hartmann (2001b). – type: Psam
mophora nissenii (dinter) dinter & Schwantes
Rabiea n. e. Br. in Gard. chron., ser. 3, 88: 279. 1930
sec. Hartmann (2001b). – type: Rabiea albinota
(Haw.) n. e. Br.
Rhinephyllum n. e. Br. in Gard. chron., ser. 3, 82: 92.
1927 sec. Hartmann (2001b). – type: Rhinephyllum
muirii n. e. Br.
= Neorhine Schwantes in Monatsschr. deutsch. Kakteen-Ges. 2: 22. 1930.
Rhombophyllum (Schwantes) Schwantes in z. Sukkulentenk. 3: 16, 23. 1927 sec. Hartmann (2001b) ≡
Bergeranthus subg. Rhombophyllum Schwantes in z.
Sukkulentenk. 2: 180. 1926. – type: Rhombophyllum
rhomboideum (Salm-dyck) Schwantes
Ruschia Schwantes in z. Sukkulentenk. 2: 186. 1926
sec. Hartmann (2001b). – type: Ruschia rupicola
(engl.) Schwantes
a large genus including 206 species, for which no
taxonomic revision has been compiled. dehn (1993)
recognized nine subgenera, of which only one has
been studied further, Ruschia subg. Spinosae (Salmdyck) dehn (Hartmann & Stüber 1993). However, it
has since been established that Ruschia is not monophyletic in its current circumscription (Klak & al.
2013). the clade in which species of Ruschia s.str.
are found is still poorly resolved, so that relationships of species groups of current Ruschia remain
uncertain. in addition, much denser sampling is required to establish monophyly and relationships of
the subgenera of Ruschia and their relationship to
other members of the xeromorphic winter-rainfall
clade (Klak & al. 2013).
Ruschianthus L. Bolus in J. S. african Bot. 27: 62. 1960
sec. Hartmann (2001b). – type: Ruschianthus falca
tus L. Bolus
a monotypic genus, which resolved within the Cono
phytum clade (Klak & al. 2013).
Ruschiella Klak in Bradleya 23: 100. 2005 sec. Hartmann (2001b). – type: Ruschiella argentea (L. Bolus) Klak
Saphesia n. e. Br. in Gard. chron., ser. 3, 91: 205. 1932
sec. Hartmann (2001b). – type: Saphesia flaccida
(Jacq.) n. e. Br.
301
Monotypic. Saphesia is an insufficiently known genus that needs further study to clarify its identity
(Klak & al. 2015).
Sarcozona J. M. Black in trans. & Proc. roy. Soc. South
australia 58: 176. 1934 sec. Hartmann (2001b). –
type: Sarcozona pulleinei (J. M. Black) J. M. Black
the genus consists of only two species, which are endemic to australia.
Schlechteranthus Schwantes in Monatsschr. deutsch.
Kakteen-Ges. 1: 16. 1929 sec. Hartmann (2001b). –
type: Schlechteranthus maximiliani Schwantes
a small genus of two species, which is endemic to
namaqualand. See further remarks under Polymita.
Schwantesia dinter in Möller’s deutsche Gärtn.-zeitung
42: 234. 1927 sec. Hartmann & zimmermann (2001).
– type: Schwantesia ruedebuschii dinter
Scopelogena L. Bolus in J. S. african Bot. 28: 9. 1962
sec. Hartmann (2001b). – type: Scopelogena ver
ruculata (L.) L. Bolus
a small genus with two species, which was placed in
a clade with two species of the polyphyletic Ruschia
(Klak & al. 2013). a comprehensive revision of
Ruschia should therefore also address the generic delimitation of Scopelogena.
Sesuvium L., Syst. nat., ed. 10: 1052, 1058, 1371. 1759
sec. Hartmann (2001b) ≡ Halimus rumph. ex Kuntze,
revis. Gen. Pl. 1: 263. 1891, nom. illeg. ≡ Halimum
Loef. ex Hiern. in cat. afr. Pl. 1: 411. 1898. – type:
Sesuvium portulacastrum (L.) L.
= Diplochonium Fenzl, nov. Stirp. dec.: 57. 1839.
= Pyxypoma Fenzl in ann. Wiener Mus. naturgesch. 2:
293. 1840.
= Psammanthe Hance in ann. Bot. Syst. 2: 659. 1851.
the genus includes about 15 species; the exact
number, however, is unknown and a taxonomic treatment is needed. Sesuvium contains an african clade
consisting of c4 species and an american clade consisting of Cypselea (also c4) and a c3 Sesuvium clade
(Bohley & al. 2015). Sesuvium portulacastrum (L.)
L., which belongs to the american clade, is found
along tropical and subtropical coasts.
Skiatophytum L. Bolus in S. african Gard. 17: 435. 1927
sec. Hartmann (2001b) ≡ Gymnopoma n. e. Br. in
Gard. chron., ser. 3, 83: 194. 1928. – type: Skiato
phytum tripolium (L.) L. Bolus
= Caryotophora Leistner, notes Mesembryanthemum
3: 289. 1958.
Skiatophytum forms part of the tribe Apatesieae,
which consists of only 11 species. the tribe is considered to be monophyletic (ihlenfeldt & Gerbaulet
1990; Klak & al. 2003b; Klak & al. 2015). Skiatophy
tum includes only three species, which are endemic
to the south-western cape region of South africa
(Klak & al. 2015). Based on a recent phylogenetic
study, Klak & al. (2015) proposed that the monotypic
Caryotophora Leistner should be considered part of
Skiatophytum. in addition, it was shown that the lec-
302
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
totype and protologue of Mesembryanthemum flacci
dum Jacq. did not correspond to the species currently
associated with this name, which was described as S.
flaccidifolium Klak (Klak & al. 2015). the type of
the monotypic Saphesia, which is M. flaccidum, was
found to be an insufficiently known species.
Smicrostigma n. e. Br. in Gard. chron., ser. 3, 87: 72.
1930 sec. Hartmann (1993). – type: Smicrostigma
viride (Haw.) n. e. Br.
a monotypic genus, which was shown to be closely
related to Zeuktophyllum (two species) and Octopo
ma p.p. (Klak & al. 2013). all three taxa are endemic
to the Little Karoo, South africa. the overall similarity between these taxa suggests that a broader generic
concept should be adopted for this group of species.
Stayneria L. Bolus in J. S. african Bot. 27: 47. 1960 sec.
Hartmann (2001b). – type: Stayneria littlewoodii L.
Bolus
a monotypic genus, which was found to be closely
allied to parts of the polyphyletic genus Ruschia
(Klak & al. 2013).
Stoeberia dinter & Schwantes in z. Sukkulentenk. 3: 14,
17. 1927 sec. Hartmann (2001b). – type: Stoeberia
beetzii (dinter) dinter & Schwantes
= Ruschianthemum Friedrich in Mitt. Bot. Staatssamml.
München 3: 563. 1960.
Hartmann (2001b) treated Ruschianthemum as a
distinct genus with R. gigas (dinter) Friedrich as
the only species. However, the species had already
previously placed in Stoeberia because of strong
similarities; it differs mostly in its fruit morphology,
which has traditionally played an important role
in delimiting genera in Aizoaceae. However, fruit
characters have recently been shown to be far more
homoplasious than previously expected (Klak & al.
2013), suggesting that fruit morphology on its own
does not justify the recognition as a distinct genus.
Given the large overall similarity in all other morphological characters to Stoeberia, this species has
been reinstated as a member of Stoeberia by chesselet & van Wyk (2002), based on very similar arguments.
Stomatium Schwantes in z. Sukkulentenk. 2: 175. 1926
sec. Hartmann (2001b). – type: Stomatium suaveo
lens Schwantes
= Agnirictus Schwantes in Monatsschr. deutsch. Kakteen-Ges. 2: 21. 1930, nom. inval.
Stomatium currently includes 39 species, but lacks a
taxonomic revision. it was shown to be closely related to Chasmatophyllum (eight species), Mossia (one
species), Neohenricia (two species), Peersia (three
species) and Rhinephyllum (11 species) by Klak &
al. (2013). Both Chasmatophyllum and Rhinephyllum
also lack a taxonomic revision. all of these genera
occur outside the winter-rainfall region of South africa. the group shares a similar floral morphology,
i.e. yellow or more rarely cream-coloured petaloid
staminodes, absence of filamentous staminodes and
a concavely shaped ovary wall. Over the past decades
species have been shifted between genera since generic boundaries are poorly circumscribed.
Tanquana H. e. K. Hartmann & Liede in Bot. Jahrb.
Syst. 106: 479. 1986 sec. Hartmann (2001b). – type:
Tanquana archeri (L. Bolus) H. e. K. Hartmann &
Liede
Based on differences in fruit morphology, Hartmann
& Liede (1986) excluded three species from Pleiospi
los and established a new genus for them, Tanquana.
However, its previously recognized close relationship
to Pleiospilos was confirmed by Klak & al. (2013),
and is also corroborated by leaf-morphological characters (Hartmann & Liede 1986).
Tetragonia L., Sp. Pl. 1: 480. 1753 sec. Hartmann
(2001b) ≡ Ludolfia adans., Fam. Pl. 2: 244. 1763
≡ Tetragonocarpus Hassk. in Flora 40: 99. 1857. –
type: Tetragonia fruticosa L. – Fig. 2B.
= Demidovia Pall., enum. Hort. demidof: 150. 1781.
= Tetragonella Miq. in Lehm. Pl. Preiss. 1: 245. 1845.
= Anisostigma Schinz in Bull. Herb. Boissier 5 app. 3:
78. 1897.
Titanopsis Schwantes in z. Sukkulentenk. 2: 178. 1926
sec. Hartmann (2001b). – type: Titanopsis calcarea
(Marloth) Schwantes
= Verrucifera n. e. Br. in Gard. chron., ser. 3, 88: 278.
1930.
Trianthema L., Sp. Pl. 1: 223. 1753 sec. Hartmann
(2001b) ≡ Reme adans., Fam. Pl. 2: 245. 1763 ≡ Por
tulacastrum Juss. ex Medik., Philos. Bot.: 99. 1789.
– type: Trianthema portulacastrum L.
= Papularia Forssk., Fl. aegypt.-arab.: 69. 1775.
= Meridiana L. f., Suppl. Pl.: 248. 1782.
= Ancistrostigma Fenzl in ann. Wiener Mus. naturgesch. 2: 293. 1840.
the genus belongs to Sesuvioideae and comprises
about 28 species in two monophyletic clades, T. subg.
Trianthema and T. subg. Papularia (Bohley & al.
2015). the latter has been revised by Hartmann & al.
(2011). nearly all species are c4 plants: an exception
is the c3 species T. ceratosepala Volkens & irmsch.
Tribulocarpus S. Moore in J. Bot. 59: 228. 1921 sec.
thulin & al. (2012). – type: Tribulocarpus dimor
phantha (Pax) S. Moore
Tribulocarpus belongs to the Sesuvioideae (Klak &
al. 2003; thulin & al. 2012) and is sister to the remaining genera of the subfamily, i.e. Sesuvium (incl.
Cypselea), Trianthema and Zaleya. it is the only genus in the Sesuvioideae that includes only c3 species.
Trichodiadema Schwantes in z. Sukkulentenk. 2: 187.
1926 sec. niesler (2001). – type: Trichodiadema stel
ligerum (Haw.) Schwantes
the genus includes 32 species and is divided into
two subgenera (Hartmann & niesler 2013). the latter
study as well as earlier studies appear to be largely
based on the types of Trichodiadema (niesler 1997),
Willdenowia 45 – 2015
since very little additional material (none from a
South african herbarium) is cited as the basis for their
taxonomic conclusions. distribution ranges for the
recognized species remain uncertain due to the lack
of cited vouchers. in addition, monophyly of the genus needs to be reinvestigated in view its having been
found nested among species of Delosperma (Klak &
al. 2013).
Vanheerdea L. Bolus ex H. e. K. Hartmann in Bradleya 10: 15. 1992 sec. Hartmann (2001b). – type:
Vanheerdea roodiae (n. e. Br.) L. Bolus ex H. e. K.
Hartmann
Vanzijlia L. Bolus in Fl. Pl. South africa 7. t. 256: 262.
1927 sec. Hartmann (2001b). – type: Vanzijlia annu
lata (a. Berger) L. Bolus
Vlokia S. a. Hammer in cact. Succ. J. (Los angeles) 66:
256. 1994 sec. Hartmann (2001b). – type: Vlokia ater
S. a. Hammer
Wooleya L. Bolus in J. S. african Bot. 27: 48. 1960 sec.
Hartmann (2001b). – type: Wooleya farinosa L. Bolus
a monotypic genus from the cost of namaqualand. its
phylogenetic position within the xeromorphic winterrainfall clade remains unresolved (Klak & al. 2013).
Zaleya Burm. f. in Fl. indica (n. L. Burman): 110. 1768
sec. Hartmann (2001b). – type: Zaleya decandra
Burm. f.
= Rocama Forssk., Fl. aegypt.-arab.: 71. 1775.
the genus is monophyletic and belongs to Sesuvio
ideae, where it is sister to Sesuvium (Bohley & al.
2015). Zaleya is a c4 genus and distributed in eastern
africa, southern asia and australia. it contains seven
species (Hartmann 2011b).
Zeuktophyllum n. e. Br. in Gard. chron., ser. 3, 81: 12.
1927 sec. Hartmann (2001b). – type: Zeuktophyllum
suppositum (L. Bolus) n. e. Br.
Amaranthaceae Juss. sec. Müller & Borsch (2005).
Amaranthaceae belong to a clade together with Chenopo
diaceae. Support for the monophyly of the “Amaran
thaceae–Chenopodiaceae alliance” is found consistently
in all molecular phylogenetic analyses (Manhart & rettig
1994; downie & al. 1997; cuénoud & al. 2002; Kadereit
& al. 2003; Müller & Borsch 2005a; Schäferhoff & al.
2009; Brockington & al. 2009). the family circumscription of the Amaranthaceae in the sense of Schinz (1893)
was upheld by townsend (1993) and confirmed as monophyletic with high statistical confidence by Kadereit
& al. (2003) and Müller & Borsch (2005a). Following
this concept the Amaranthaceae predominantly occur in
tropical and subtropical regions with most of the species
diversity in the neotropics, eastern and southern africa
and australia (Müller & Borsch 2005a, b; Sánchez-del
Pino & al. 2009). Subfamily Gomphrenoideae has been
revealed as monophyletic and nested within the Ama
ranthoideae and is characterized by unilocular anthers
(Sánchez-del Pino & al. 2009) and metareticulate pol-
303
len (Borsch & Barthlott 1998; in core Gomphrenoideae
except Irenella, Iresine and Woehleria). in contrast, subfamily Amaranthoideae is largely paraphyletic. the genera Bosea and Charpentiera were found as successive
sisters to the remainder of the Amaranthaceae (Müller &
Borsch 2005a). the Celosioideae (corresponding to the
celosioid clade) are the only natural tribe in the pre-phylogenetic classification of the family and further major lineages are constituted by the amaranthoid clade (Amaran
thus, Chamissoa and relatives), the aervoid clade (Aerva,
Ptilotus and relatives) and the achyranthoid clade (Achy
ranthes, Centemposis, Cyathula, Pupalia, Sericocoma
and many other african genera; Müller & Borsch 2005b).
the angiosperm Phylogeny Group (aPG 1998) proposed
to apply the name Amaranthaceae to the complete Ama
ranthaceae–Chenopodiaceae alliance, essentially adopting the family concept of Baillon (1887) and Mallingson
(1922). the broad family circumscription was also adopted in subsequent versions of the aPG classification (aPG
ii 2003; aPG iii 2009). However, since recent phylogenetic analyses rather indicate the monophyly of the core
Chenopodiaceae but are not yet conclusive about the position of the subfamily Polycnemoideae, the widely used
family name Chenopodiaceae is maintained (see introduction to the family Chenopodiaceae). the four genera
of the well-supported polycnemoid lineage (Hemichroa,
Nitrophila, Polycnemum, Surreya) that corresponds to
the subfamily Polycnemoideae share petaloid tepals, two
large bracteoles supporting the flower, an androecium that
is basally united into a tube and bilocular anthers with the
Amaranthaceae sensu Schinz (1893), Masson & Kadereit
(2013). We are therefore provisionally treating this subfamily under the Amaranthaceae along with endlicher
(1841), Moquin-tandon (1849) and Scott (1977).
Achyranthes L., Sp. Pl. 1: 204. 1753, nom. cons. prop.
sec. townsend (1993). – type: Achyranthes aspera L.
Achyropsis Benth. & Hook. f., Gen. Pl. 3(1): 36. 1880
sec. townsend (1993) ≡ Achyranthes sect. Achyropsis
Moq. in candolle, Prodr. 13(2): 310. 1849. – type:
not designated.
Aerva Forssk. in Fl. aegypt.-arab.: 170. 1775, nom.
cons. sec. townsend (1993). – type: Aerva tomentosa
Forssk.
the genus may not be monophyletic and includes two
principal lineages (thiv & al. 2006). One of these was
shown as sister to the remainder of the aervoid clade
(represented by A. javanica Juss.; Müller & Borsch 2005a) and the other (represented by A. leucura
Moq.; Müller & Borsch 2005b) as sister to Ptilotus.
Further study of the aervoid clade is needed to clarify
generic concepts.
Allmania r. Br. ex Wight in J. Bot. 1: 226. 1834 sec.
townsend (1993). – type: Allmania nodiflora (L.) r.
Br. ex Wight
Allmaniopsis Suess. in Mitt. Bot. Staatssamml. München
4. 1952 sec. townsend (1993). – type: Allmaniopsis
fruticulosa Suess.
304
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
Alternanthera Forssk., Fl. aegypt.-arab.: 28, 59. 1775
sec. townsend (1993). – type: Alternanthera achy
ranthes Forssk.
= Telanthera r. Br., Observ. congo. 1818.
= Brandesia Mart., nov. Gen. Sp. Pl. 2: 25. 1826.
= Buchholzia Mart., nov. Gen. Sp. Pl. 2: 49. 1826.
= Mogiphanes Mart., nov. Gen. Sp. Pl. 2: 29. 1826.
the genus Alternanthera is well supported as monophyletic in the current circumscription and is characterized by the presence of capitate stigmas and in
most species also distinct androecial appendages that
alternate with the filaments. the previously recognized genera do not represent natural entities except
Mogiphanes, which is nested within one of the two
major subclades of Alternanthera (Sánchez-del Pino
& al. 2012).
Amaranthus L., Sp. Pl. 1: 989. 1753 sec. townsend
(1993). – type: Amaranthus caudatus L.
= Acnida L., Sp. Pl. 2: 1027. 1753 ≡ Amaranthus subg.
Acnida (L.) aellen ex K. r. robertson in J. arnold
arbor. 62(3): 283. 1981.
= Albersia Kunth, Fl. Berol. 2: 144. 1838 ≡ Amaranthus
subg. Albersia (Kunth) Gren. & Godr., Fl. France 3:
3. 1856.
= Acanthochiton torr., rep. exped. zuñi & colorado
rivers: 170. 1853 ≡ Amaranthus sect. Acanthochi
ton (torr.) Mosyakin & K. r. robertson in ann. Bot.
Fenn. 33: 277. 1996.
= Goerziella Urb., repert Spec. nov. regni Veg. 20: 301.
1924 ≡ Amaranthus sect. Goerziella (Urb.) Mosyakin
& K. r. robertson in ann. Bot. Fenn. 33: 280. 1996.
the genus, with its more than 75 currently recognized species, is monophyletic and constitutes a
c4 lineage (Sage & al. 2007) within the otherwise
completely c3 amaranthoid clade (Müller & Borsch
2005b) of subfamily Amaranthoideae. the current
infrageneric system of the genus (Mosyakin & robertson 1996, 2003), recognizing three subgenera
(A. subg. Acnida (L.) aellen ex K. r. robertson,
A. subg. Albersia (Kunth) Gren. & Godr. and A.
subg. Amaranthus) and several sections, was developed before the advent of molecular phylogenetic
methods and is now in need of revision. Amaran
thus subg. Acnida, represented by dioecious species
currently placed in three sections, seems to be nonmonophyletic, since dioecy in Amaranthus probably
developed independently at least twice (Mosyakin
2005). Surprisingly, no comprehensive molecular
phylogenetic study of Amaranthus has been done
yet, despite the economic importance of the genus,
containing some pseudocereal and green crops, popular ornamentals, and noxious weeds.
Arthraerua (Kuntze) Schinz, nat. Pflanzenfam. 3(1a):
109. 1893 sec. townsend (1993) ≡ Aerva sect. Ar
thraerua Kuntze in Jahrb. Königl. Bot. Gart. Berlin 4:
272. 1886. – type: Arthraerua leubnitziae (Kuntze)
Schinz
Bosea L., Sp. Pl. 1: 225. 1753 sec. townsend (1993). –
type: Bosea yervamora L.
Calicorema Hook. f., Gen. Pl. 3(1): 34. 1880 sec.
townsend (1993). – type: Calicorema capitata
(Moq.) Hook. f.
the genus is not monophyletic as currently circumscribed because its two species, Calicorema capita
ta and C. squarrosa (Schinz) Schinz, appear in two
completely different lineages of the achyranthoid
clade (Müller & Borsch 2005a, b). correct generic
assignment has to await a comprehensive analysis of
the achyranthoid clade.
Celosia L., Sp. Pl. 1: 205. 1753 sec. townsend (1993). –
type: Celosia argentea L.
Centema Hook. f., Gen. Pl. 3(1): 31. 1880 sec. townsend
(1993). – type: Centema angolensis Hook. f.
Centemopsis Schinz in Vierteljahrsschr. naturf. Ges.
zürich 56: 242. 1911 sec. townsend (1993). – type:
not designated.
the genus is probably monophyletic considering
phylogenetic trees of Müller & Borsch (2005b) and
Sage & al. (2007).
Centrostachys Wall., Fl. ind., ed. 1820: 497. 1824 sec.
townsend (1993). – type: Centrostachys aquatica
(r. Br.) Wall.
Chamissoa Kunth in nov. Gen. Sp. [H. B. K. ] 2: 196, t.
125. 1818, nom. cons. sec. townsend (1993). – type:
Chamissoa altissima (Jacq.) Kunth
Charpentiera Gaudich. in Voy. Uranie, Bot.: 444, t. 48.
1826 sec. townsend (1993). – type: Charpentiera
obovata Gaudich.
Chionothrix Hook. f., Gen. Pl. 3(1): 33. 1880 sec.
townsend (1993). – type: Chionothrix somalensis
(S. Moore) Hook. f.
Cyathula Blume in Bijdr. Fl. ned. ind. 11: 548. 1826
sec. townsend (1993). – type: Cyathula prostrata
(L.) Blume
Cyphocarpa (Fenzl) Lopr. in Bot. Jahrb. Syst. 27: 42.
1899 sec. townsend (1993) ≡ Sericocoma subg. Ky
phocarpa Fenzl in Linnaea 17: 324. 1843. – type:
Cyphocarpa trichinioides (Fenzl) Lopr.
Dasysphaera Volkens ex Gilg, nat. Pflanzenfam. nachtr.
2-4, 1: 153. 1897 sec. townsend (1993). – type: not
designated.
Deeringia r. Br., Prodr. Fl. nov. Holland.: 413. 1810
sec. townsend (1993). – type: Deeringia celosioides
r. Br.
= Dendroportulaca eggli in adansonia, sér. 3, 19: 49.
1997.
Celosieae. Dendroportulaca (formerly placed in
Portulacaceae) has been shown to be referable to
Deeringia and the only species, Dendroportulaca
mirabilis eggli, has been transferred there (applequist & Pratt 2005).
Digera Forssk. in Fl. aegypt.-arab.: 65. 1775 sec.
townsend (1993). – type: Digera arvensis Forssk.
Eriostylos c. c. towns. in Kew Bull. 46: 237. 1991
Willdenowia 45 – 2015
sec. townsend (1993). – type: Eriostylos stefaninii
(chiov.) c. c. towns.
Froelichia Moench, Methodus: 50. 1794 sec. townsend
(1993). – type: Froelichia lanata Moench
Froelichiella r. e. Fr. in ark. Bot. 16(13): 3. 1921 sec.
townsend (1993). – type: Froelichiella grisea (Lopr.)
r. e. Fr.
Gomphrena L., Sp. Pl. 1: 224. 1753 sec. townsend
(1993). – type: Gomphrena globosa L. – Fig. 2e.
Gossypianthus Hook. in icon. Pl.: 251. 1840 sec. clemants (2003). – type: Gossypianthus rigidiflorus Hook.
close relationships between Gossypianthus and
Guilleminea and a merger of both genera were longdisputed (Mears 1967; eliasson 1988). Phylogenetic
analysis of plastid (Sánchez-del Pino & al. 2009) and
nuclear sequences (t. Ortuño & t. Borsch, unpubl.
data) show that they are more distantly related and do
not form sister groups.
Guilleminea Kunth, nov. Gen. Sp. (quarto ed.) 6: 40, pl.
518. 1823 sec. clemants (2003) ≡ Brayulinea Small,
Fl. S. e. U. S.: 394. 1903. – type: Guilleminea illece
broides Kunth
See notes under Gossypianthus.
Hebanthe Mart., Beitr. amarantac.: 96. 1825 sec. Borsch
& Pedersen (1997). – type: not designated.
the genus was resurrected based on its very distinctive flower and pollen morphology by Borsch & Pedersen (1997) and also appears to be monophyletic
based on molecular phylogenetic analysis (Sánchezdel Pino & al. 2009; Borsch & al. 2011).
Hebanthodes Pedersen in Bonplandia (corrientes) 10:
102. 2000 sec. Pedersen (2000). – type: Hebanthodes
peruviana Pedersen
Monotypic and known from a single historical specimen (Pedersen 2000). affinities are unclear but a
placement within the gomphrenoid clade of Gom
phrenoideae (Sánchez-del Pino & al. 2009) is certain, where it shares a pollen morphology similar to
Pfaffia.
Hemichroa r. Br., Prodr. Fl. nov. Holland.: 409. 1810
sec. Masson & Kadereit (2013). – type: Hemichroa
pentandra r. Br.
Polycnemoideae. Hemichroa consists of only one
species; two further species have been segregated as
Surreya (see there for details; Masson & Kadereit
2013). the succulent halophyte H. pentandra r. Br.
is endemic to australia. it is sister to Surreya (Masson
& Kadereit 2013).
Henonia Moq. in candolle, Prodr. 13(2): 237. 1849 sec.
townsend (1993). – type: Henonia scoparia Moq.
Herbstia Sohmer in Brittonia 28: 448. 1977 sec. townsend
(1993). – type: Herbstia brasiliana (Moq.) Sohmer
Hermbstaedtia rchb. in consp. regn. Veg.: 164. 1828
sec. townsend (1993). – type: Hermbstaedtia glauca
(J. c. Wendl.) Steud. ex rchb.
Indobanalia a. n. Henry & B. roy in Bull. Bot. Surv.
india 10: 274. 1969 sec. townsend (1993) ≡ Bana
305
lia Moq. in candolle, Prodr. 13(2): 278. 1849, nom.
illeg. – type: Indobanalia thyrsiflora (Moq.) a. n.
Henry & B. roy
Irenella Suess. in repert. Spec. nov. regni Veg. 35: 318.
1934 sec. townsend (1993). – type: Irenella chryso
tricha Suess.
Iresine P. Browne in civ. nat. Hist. Jamaica: 358. 1756,
nom. cons. sec. townsend (1993). – type: Iresine dif
fusa Humb. & Bonpl. ex Willd.
= Dicraurus Hook. f., Gen. Pl. 3(1): 42. 1880.
the genus is monophyletic (Sánchez-del Pino & al.
2009; Borsch, Flores Olvera, zumaya & Müller, in
review) with approximately 45 species all of which
are characterized by Iresine-type pollen (Borsch
1998). the two species formerly classified as Di
craurus on the base of alternate and not opposite
leaves are nested within the Iresine clade, confirming the merger by Henrickson & Sundberg (1986).
their dense indumentum with branched trichomes
appears to be an adaptation to the dry habitats of
northern Mexico.
Lagrezia Moq. in candolle, Prodr. 13(2): 252. 1849 sec.
townsend (1993). – type: Lagrezia madagascarien
sis (Poir.) Moq.
= Apterantha c. H. Wright in Bull. Misc. inform. Kew
1918: 202. 1918.
Leucosphaera Gilg, nat. Pflanzenfam. nachtr. 2 – 4,
1: 152. 1897 sec. townsend (1993). – type: Leu
cosphaera bainesii (Hook. f.) Gilg
Lithophila Sw., Prodr. [O. P. Swartz]: 1, 14. 1788 sec.
townsend (1993). – type: Lithophila muscoides Sw.
Lopriorea Schinz in Vierteljahrsschr. naturf. Ges. zürich
56: 251. 1911 sec. townsend (1993). – type: Loprio
rea ruspolii (Lopr.) Schinz
Marcelliopsis Schinz, nat. Pflanzenfam., ed. 2, 16c: 48.
1934 sec. townsend (1993) ≡ Marcellia Baill. in Bull.
Mens. Soc. Linn. Paris 1(79): 625. 1886, nom. illeg.
– type: Marcellia mirabilis Baill.
Mechowia Schinz in nat. Pflanzenfam. 3(1a): 110. 1893
sec. townsend (1993). – type: Mechowia grandiflora
Schinz
Nelsia Schinz in Vierteljahrsschr. naturf. Ges. zürich
56: 247. 1912 sec. townsend (1993). – type: Nelsia
quadrangula (engl.) Schinz
Neocentema Schinz in Vierteljahrsschr. naturf. Ges.
zürich 56: 248. 1911 sec. townsend (1993). – type:
not designated.
Nitrophila S. Watson in Botany [Fortieth Parallel]: 297.
1871 sec. Masson & Kadereit (2013) ≡ Banalia sect.
Idiopsis Moq. in candolle, Prodr. 13(2): 279. 1849. –
type: Nitrophila occidentalis (Moq.) S. Watson
Polycnemoideae. Nitrophila consists of four (to eight)
species distributed in western north america and
South america, and the genus represents a classical
example of an amphitropical desert disjunction (Masson & Kadereit 2013). Nitrophila shows leaf anatomical adaptations to physiological drought.
306
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
Nothosaerva Wight in icon. Pl. ind. Orient. [Wight] 6:
17. 1853 sec. townsend (1993). – type: Nothosaerva
brachiata (L.) Wight
Nototrichium W. F. Hillebr. in Fl. Hawaiian isl.: 372.
1888 sec. townsend (1993). – type: not designated.
Nyssanthes r. Br., Prodr. Fl. nov. Holland.: 418. 1810
sec. townsend (1993). – type: not designated.
Pandiaka Hook. f., Gen. Pl. 3(1): 35. 1880 sec. townsend
(1993) ≡ Achyranthes sect. Pandiaka Moq. in candolle, Prodr. 13(2): 310. 1849. – type: Pandiaka in
volucrata (Moq.) B. d. Jacks.
Pedersenia Holub in Preslia 70: 181. 1998 sec. Holub
(1998). – type: Pedersenia argentata (Mart.) Holub
the genus was resurrected by Pedersen (1997) under Trommsdorffia Mart., a later homonym of Trom
msdorffia Bernh. (Compositae; Holub 1998). Peder
senia is well supported as monophyletic (Borsch &
al. 2011).
Pfaffia Mart., Beitr. amarantac.: 103. 1825 sec. townsend
(1993). – type: Pfaffia glabrata Mart.
= Sertuernera Mart., nov. Gen. Sp. Pl. 2: 36. 1826.
Philoxerus r. Br., Prodr. Fl. nov. Holland.: 416. 1810
sec. Bao & al. (2003). – type: Philoxerus conicus
r. Br.
= Blutaparon raf., new Fl. 4: 45. 1838.
the genus name was lectotypified by Standley (1917)
using an australian species, P. conicus r. Br. (≡ Gom
phrena conica (r. Br.) Spreng.). Palmer (1998) accepted G. conica along with the other australian species of Gomphrena and indicated that this is a rare
species that grows in sandy soils close to coasts. considering this, Philoxerus would have to be a synonym
of Gomphrena. the problem is that Hooker (1880,
Genera plantarum) kept the genus name Philoxerus
separate from Gomphrena and, rather than using morphological characters, applied a genus concept for
Philoxerus to comprise Gomphrena species of coastal habitats in america, africa and australia. this is
practically upheld in the genus concept of Bluta
paron raf. (townsend 1993), with four coastal species, although townsend did not even cite the name
Philoxerus. Mears (1982a, b) argued that Philoxerus
had been used for the american coastal species, so
he actually looked for a name that would define a genus of coastal species based on the american coastal
plants originally described by Linnaeus as G. vermic
ularis. What Mears overlooked is that G. conica also
appears to be a coastal plant (Palmer 1998), so that
Bentham’s 1880 generic concept of a gomphrenoid
genus of coastal plants under the name Philoxerus
would actually have been correct with five and not
four species. Strictly applying such a genus concept
to formal nomenclature, Blutaparon is a synonym
of Philoxerus. However, in the course of analysing
evolutionary relationships it will have to be seen if
the adaptation to coastal habitats correlates with other
characters that could provide synapomorphies for cir-
cumscribing and maintaining a genus Philoxerus, and
if these synapomorphies are shared by P. conicus and
the other coastal species.
Pleuropetalum Hook. f. in London J. Bot. 5: 108. 1846
sec. townsend (1993). – type: Pleuropetalum darwi
nii Hook. f. – Fig. 2F.
Pleuropetalum is a member of Amaranthoideae. the
genus is unusual in the family because of a higher
stamen and carpel number (eliasson 1988; ronse decraene & al. 1999).
Pleuropterantha Franch., Sert. Somal.: 59 (t. 5). 1882
sec. townsend (1993). – type: Pleuropterantha
revoilii Franch.
Polycnemum L., Sp. Pl. 1: 35. 1753 sec. Masson & Kadereit (2013). – type: Polycnemum arvense L.
= Selago adans., Fam. Pl. 2: 268. 1763, nom. illeg.
= Rovillia Bubani, Fl. Pyren. 1: 182. 1897.
Polycnemoideae. the genus comprises six species
distributed in eurasia and northwestern africa. it is
sister to the rest of Polycnemoideae (Masson & Kadereit 2013).
Polyrhabda c. c. towns. in Kew Bull. 39: 775. 1984 sec.
townsend (1993). – type: Polyrhabda atriplicifolia
c. c. towns.
Pseudogomphrena r. e. Fr. in ark. Bot. 16(13): 17.
1920 sec. townsend (1993). – type: Pseudogom
phrena scandens r. e. Fr.
Pseudoplantago Suess. in repert. Spec. nov. regni Veg.
35: 334. 1934 sec. townsend (1993). – type: Pseudo
plantago friesii Suess.
Pseudosericocoma cavaco in Mém. Mus Hist. nat.,
Paris, Ser. Bot., 13: 66. 1962 sec. townsend (1993).
– type: Pseudosericocoma pungens (Fenzl) cavaco
Psilotrichopsis c. c. towns. in Kew Bull. 29 (3): 464.
1974 sec. townsend (1993). – type: Psilotrichopsis
curtisii (Oliv.) c. c. towns.
Psilotrichum Blume, Bijdr. Fl. ned. ind. 11: 544. 1826
sec. townsend (1993). – type: Psilotrichum trichoto
mum Blume
the genus is highly polyphyletic (Müller & Borsch
2005a, b). correct generic assignment has to await
a comprehensive analysis of the achyranthoid clade.
Ptilotus r. Br., Prodr. Fl. nov. Holland.: 415. 1810 sec.
townsend (1993). – type: not designated.
= ?Dipteranthemum F. Muell. in South Sc. record. 3:
281. 1883.
Ptilotus has been recovered as monophyletic in the
current circumscription, with the species diversity the
result of a rapid diversification in australia (Hammer
& al. 2015).
Pupalia Juss. in ann. Mus. natl. Hist. nat. 2: 132. 1803,
nom. cons. sec. townsend (1993). – type: Pupalia
lappacea (L.) Juss.
Quaternella Pedersen in Bull. Mus. natl. Hist. nat., B,
adansonia Sér. 4, 12: 92. 1990 sec. Pedersen (2000).
– type: Quaternella confusa Pedersen
Rosifax c. c. towns. in Kew Bull. 46: 101. 1991 sec.
Willdenowia 45 – 2015
townsend (1993). – type: Rosifax sabuletorum c. c.
towns.
Saltia r. Br. ex Moq. in candolle, Prodr. 13(2): 325.
1849 sec. townsend (1993). – type: Saltia papposa
(Forssk.) Moq.
= Psilodigera Suess. in Mitt. Bot. Staatssamml.
München 4: 109. 1952.
Sericocoma Fenzl in endlicher, Gen. Pl. Suppl. 2: 33.
1842 sec. townsend (1993). – type: Sericocoma tri
chinioides Fenzl
Sericocomopsis Schinz in Bot. Jahrb. Syst. 21: 184. 1895
sec. townsend (1993). – type: Sericocomopsis hilde
brandtii Schinz
Sericorema Lopr. in Bot. Jahrb. Syst. 27: 39. 1899 sec.
townsend (1993) ≡ Sericocoma sect. Sericorema
Hook. f., Gen. Pl. 3(1): 30. 1880. – type: Sericorema
remotiflora Lopr.
Sericostachys Gilg & Lopr. ex Lopr. in Bot. Jahrb. Syst.
27: 50. 1899 sec. townsend (1993). – type: not designated.
Siamosia K. Larsen & Pedersen in nordic J. Bot. 7: 271.
1987 sec. townsend (1993). – type: Siamosia thai
landica K. Larsen & Pedersen
Stilbanthus Hook. f. in Hooker’s icon. Pl. 13: 67. 1879
sec. townsend (1993). – type: Stilbanthus scandens
Hook. f.
Surreya r. Masson & G. Kadereit in taxon 62: 109.
2013 sec. Masson & Kadereit (2013). – type: Surreya
diandra (r. Br.) r. Masson & G. Kadereit
Polycnemoideae. the australian Surreya comprises
two species of subshrubs (Masson & Kadereit 2013).
Tidestromia Standl. in J. Wash. acad. Sci. 6: 70. 1916 sec.
townsend (1993) ≡ Alternanthera sect. Cladothrix
Moq. in candolle, Prodr. 13(2): 359. 1849 ≡ Cla
dothrix (Moq.) nutt. ex Benth. & Hook. f., Gen. Pl.
3(1): 37. 1880, nom. illeg. – type: Tidestromia lanugi
nosa (nutt.) Standl. – Fig. 3a.
the genus is monophyletic (Sánchez-del Pino & al.
2009).
Trichuriella Bennet in indian J. Forest. 8: 86. 1985 sec.
townsend (1993) ≡ Trichurus c. c. towns. in Kew
Bull. 29(3): 466. 1974, nom. illeg. – type: Trichuri
ella monsoniae (L. f.) Bennet
Volkensinia Schinz in Vierteljahrsschr. naturf. Ges.
zürich 57: 535. 1912 sec. townsend (1993). – type:
Volkensinia prostrata (Volkens ex Gilg) Schinz
Woehleria Griseb., abh. Königl. Ges. Wiss. Göttingen.
9: 11. 1860 sec. townsend (1993). – type: Woehleria
serpyllifolia Griseb.
Xerosiphon turcz. in Bull. Soc. imp. naturalistes Moscou 16: 55. 1843 sec. Pedersen (1990). – type:
Xerosiphon gracilis turcz.
a well-circumscribed monophyletic genus with two
species that was long treated as part of a widely circumscribed genus Gomphrena Mart. but resurrected
by Pedersen (1990) because of its morphological
distinctness (gamopetalous perianth, cauline leaves
307
reduced to scales). Molecular phylogenetic analyses
(Sage & al. 2007; Sánchez-del Pino & al. 2009) depicted Xerosiphon as an isolated lineage in the gomphrenoid clade of subfamily Gomphrenoideae.
Anacampserotaceae eggli & nyffeler sec. aPG
(2009).
a family with three genera and around 36 species mainly
distributed in the southern and eastern parts of africa,
but also found in north america, South america, and
australia (nyffeler & eggli 2010a). the species of this
family are traditionally considered members of Portu
lacaceae; however, molecular phylogenetic studies have
shown that the traditional Portulacaceae are not monophyletic (Hershkovitz & zimmer 1997; applequist &
Wallace 2001; nyffeler 2007; nyffeler & eggli 2010a;
Ocampo & columbus 2010). nyffeler & eggli (2010a)
proposed the segregation of the traditional Portulacace
ae into four families (Anacampserotaceae, Montiaceae,
Portulacaceae and Talinaceae) based on morphological
and molecular data. in this context, the Anacampsero
taceae are recognized by their capsules with loculicidal
dehiscence, endocarp valves forming a basket-like structure and seeds with testa layers separate from each other
(nyffeler & eggli 2010a).
Anacampseros L., Opera Var.: 232. 1758, nom. cons.
sec. nyffeler & eggli (2010a). – type: Anacampseros
telephiastrum dc.
= Talinaria Brandegee in zoe 5: 231. 1908.
= Xenia Gerbaulet in Bot. Jahrb. Syst. 113: 552. 1992.
= Avonia (e. Mey. ex Fenzl) G. d. rowley in Bradleya
12: 111. 1994.
Anacampseros with c. 34 herbaceous species distributed in africa, australia, north and South america, is
the most diverse genus of Anacampserotaceae (nyffeler & eggli 2010a). Phylogenetic analyses recover
this lineage as a derived monophyletic group with
moderate statistical support (nyffeler & eggli 2010a).
Grahamia Gillies ex Hook. & arn. in Bot. Misc. 3: 331.
1833 sec. carolin (1993). – type: Grahamia bracte
ata Gillies
Talinopsis a. Gray in Smithsonian contr. Knowl. 1: 14.
1852 sec. carolin (1993). – type: Talinopsis frutes
cens a. Gray
Phylogenetic analyses recover the north american
Talinopsis frutescens a. Gray, the only member of the
genus, as the most basal member of Anacampserota
ceae (nyffeler & eggli 2010a; Ocampo & columbus
2010).
Ancistrocladaceae Planch. ex Walp. sec. aPG (2009).
a monogeneric family comprising 18 species with a disjunct paleotropical distribution in western and central
africa and southeastern asia (rischer & al. 2005). the
family includes only non-carnivorous plants characterized
by having nuts, ruminate endosperm and a gynoecium
partly inferior with a single ovule (Heubl & al. 2006).
308
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
traditionally, the family was placed either in the order
Theales (e.g. thorne 1992) or Dilleniales (e.g. thorne
2000). However, the position of the family within Cary
ophyllales and its close relationship with the “partially
carnivorous” Dioncophyllaceae (see there) was shown
by the early molecular phylogenetic study of nandi &
al. (1998). these results were confirmed by subsequent
studies (e.g. Meimberg & al. 2000; cuénoud & al. 2002;
Hilu & al. 2003; Brockington 2009, 2011; Schäferhoff
& al. 2009; renner & Specht 2011), which have also
shown, with high support, that both Ancistrocladaceae
and Dioncophyllaceae are part of the “carnivorous clade”
of the Caryophyllales. Other studies focusing on the evolution of carnivory and relationships within this clade
(e.g. Heubl & al. 2006; renner & Specht 2011) suggest
that the absence of carnivory in Ancistrocladaceae can be
explained as a complete secondary loss of this character.
Ancistrocladus Wall., numer. List: 1052. 1829, nom.
cons. sec. Porembski (2003). – type: Ancistrocladus
hamatus (Vahl) Gilg
Asteropeiaceae takht. ex reveal & Hoogland sec.
aPG (2009).
a monogeneric family with eight species endemic to
Madagascar (Kubitzki 2003). the genus was traditionally placed in Theales, either in its own family (e.g.
takhtajan 1987; thorne 1992) or within Theaceae (e.g.
cronquist 1988). However, early molecular phylogenetic
studies have shown the affinities of Asteropeiacae within
Caryophyllales and the close relationship with Physena
ceae (e.g. Morton & al. 1997). these results were confirmed by subsequent studies (e.g. cuénoud & al. 2002;
Brockington 2009, 2011; Soltis & al. 2011). the clade
Asteropeiaceae–Physenaceae is also well supported by
wood-anatomical characters (e.g. Miller & dickison
1992; dickison & Miller 1993; carlquist 2006); some
member species (with small circular alternate pits on
vessels, vasicentric tracheids plus fibre tracheids, abaxial
confluent diffuse parenchyma and predominantly uniseriate rays) have been proposed as synapomorphies to the
family (e.g. carlquist 2006).
Asteropeia thouars, Hist. Vég. isles austral. afriq.: 5152, pl. 15. 1805 sec. Kubitzki (2003a). – type: As
teropeia multiflora thouars
Barbeuiaceae nakai sec. aPG (2009).
a monotypic family restricted to Madagascar (rohwer
1993). the family is characterized by ovaries consisting
of two united carpels with two locules and by capsules
(rohwer 1993). traditionally, the family was placed in
Phytolaccaceae subfamily Barbeuioideae, but its position as an independent lineage has been supported by
several molecular phylogenetic studies (cuénoud & al.
2002; Hilu & al. 2003; Schäferhoff & al. 2009).
Barbeuia thouars in Gen. nov. Madagasc.: 6. 1806 sec.
rohwer (1993a). – type: Barbeuia madagascariensis
Steud.
Basellaceae raf. sec. aPG (2009).
Basellaceae are a small tropical and subtropical family
native to the americas, southeastern africa, Madagascar and possibly asia. the centre of diversity is in the
andes of northwestern South america, but the centre of
origin may very well be in africa. at present, four genera
(Anredera, Basella, Tournonia, Ullucus) with a total of
19 species are recognized, most of them succulent vines
occurring in dry habitats. Some species are cultivated,
and one (Ullucus tuberosus caldas) is an important highandean crop grown for its edible tubers.
Anredera Juss., Gen. Pl.: 84. 1789 sec. eriksson (2007) ≡
Clarisia abat in Mem. acad. real Soc. Med. Sevilla
10: 418. 1792. – type: Anredera spicata J. F. Gmel.
= Boussingaultia Kunth, nov. Gen. Sp. (quarto ed.) 7:
194 t. 645. 1825.
= Tandonia Moq. in candolle, Prodr. 13(2): 226. 1849
≡ Boussingaultia sect. Tandonia (Moq.) Volkens, nat.
Pflanzenfam. 3(1a): 128. 1893 ≡ Anredera sect. Tan
donia (Moq.) Steenis, Flora Malesiana, ser. 1, 5: 302.
1957.
= Boussingaultia sect. Moquiniella Hauman in anales
Mus. nac. Buenos aires 33: 351. 1925.
= Boussingaultia sect. Euboussingaultia Volkens, nat.
Pflanzenfam. 3(1a): 128. 1893, nom. inval.
= Siebera c. Presl in isis (Oken) 21: 275. 1828, nom.
nud.
= Beriesa Steud., nomencl. Bot., ed. 2, 1: 199. 1840,
nom. nud.
a monophyletic group of species in Anredera corresponds to the previously recognized taxon Tandonia,
but a formal recognition of Tandonia would make the
remaining Anredera paraphyletic (eriksson 2007).
Basella L., Sp. Pl. 1: 272. 1753 sec. eriksson (2007). –
type: Basella rubra L.
= Gandola raf., Sylva tellur.: 60. 1838.
One species, B. paniculata Volkens, is morphologically deviating in Basella, and may be better placed in
a genus of its own. a phylogenetic analysis based on
morphological data gave inconclusive results regarding its placement (eriksson 2007).
Tournonia Moq. in candolle, Prodr. 13(2): 221, 225.
1849 sec. eriksson (2007). – type: Tournonia hooker
iana Moq.
Ullucus caldas in Seman. nuev. Granad.: 185. 1809 sec.
eriksson (2007). – type: Ullucus tuberosus caldas
= Melloca Lindl. in Gard. chron. 42: 685. 1847.
Cactaceae Juss. sec. aPG (2009).
Cactaceae comprise about 120 to 130 genera and some
1450 to 1870 species (Hunt 2006; nyffeler & eggli
2010b). Most species are highly modified perennial stem
succulents which conserve water to survive temporary
dry periods. Only some two dozen species of the genera
Pereskia, Pereskiopsis and Quiabentia have a shrubby or
tree-like habit with more or less fleshy leaves. all species of the family bear characteristic spine clusters (i.e.
Willdenowia 45 – 2015
areoles), representing short shoots with leaves transformed into spines already at the stage of primordia.
Some taxa are spineless and even lack areoles at maturity but all species bear areoles as seedlings. this characteristic is a true synapomorphy of the entire family.
cacti are native to the americas, except for the widely
distributed Rhipsalis baccifera (Sol.) Stearn that also
occurs in tropical africa, Madagascar, and on islands
in the indian Ocean. Several species from different lineages have been introduced worldwide as crop plants
or ornamentals and have become naturalized, and are
classified as invasive aliens in several areas, including australia, southern africa, and the Mediterranean.
For a long time in the past, the classification into genera
and suprageneric groups was based on form characteristics of vegetative and reproductive structures, culminating in the fine-grained classifications of Backeberg
(1958 – 1962, 1966) or Buxbaum (1962) and endler &
Buxbaum (1974). Many of the highly modified structural
features are associated with the succulent life strategy
(e.g. nyffeler & al. 2008), and hence provide particular
challenges in the interpretation of a classification based
on purported relationships. the consensus classification
initiative as reported by Hunt & taylor (1986) and subsequent papers helped to overcome the deviating systems
used in the second part of the 20th century, but also fell short
in not being based on further and expanded data sets of
comparative data for reconstructing relative relationships.
However, the molecular phylogenetic studies (see the introduction and nyffeler & eggli (2010b) provide the base
for an increasingly stable backbone classification for major suprageneric clades. at the same time, unexpected
novel placements are suggested by such studies for several species or genera, such as Blossfeldia (nyffeler 2002)
or Lymanbensonia (Korotkova & al. 2010), while longestablished genera, such as Echinocactus and Ferocactus
but also Mammillaria have been found to be polyphyletic
(Bárcenas & al. 2011; Hernández-Hernández & al. 2011;
Vázquez-Sánchez & al. 2013). to use these findings for
updating the generic classification of the family is a pronounced challenge (Hunt 2006; nyffeler & eggli 2010b).
Acanthocereus (engelm. ex a. Berger) Britton & rose
in contr. U. S. natl. Herb. 12: 432. 1909 sec. Hunt
(2006) ≡ Cereus subsect. Acanthocereus engelm. ex
a. Berger in rep. (annual) Missouri Bot. Gard. 16:
77. 1905. – type: Acanthocereus baxaniensis (Karw.)
Borg
= Monvillea Britton & rose, cactaceae 2: 21. 1920.
currently accepted as monotypic with A. tetragonus
(L.) Britton & rose (Hunt 2006), whereas all other
names suggested in this genus are of uncertain application or were wrongly assigned to Acanthocereus.
the molecular phylogenetic study of arias & al.
(2005) showed that Acanthocereus would need to be
expanded to include Peniocereus subg. Pseudoacan
thocereus Sánchez-Mej., but no new combinations
have yet been published.
309
Acharagma (n. P. taylor) Glass in Guía identif. cact.
amenazadas México 1: [ac/ag]. 1997 sec. VázquezSánchez & al. (2013) ≡ Escobaria sect. Acharagma
n. P. taylor in Kakteen and. Sukk. 34: 185. 1983.
– type: Acharagma roseanum (Boed.) e. F. anderson
Acharagma includes only two species that were found
well-supported as sister to each other by VázquezSánchez & al. (2013).
Ariocarpus Scheidw. in Bull. acad. roy. Sci. Bruxelles
5: 491. 1838 sec. Vázquez-Sánchez & al. (2013). –
type: Ariocarpus retusus Scheidw.
= Neogomesia castañeda in cact. Succ. J. (Los angeles) 13: 98. 1941.
= Roseocactus a. Berger in J. Wash. acad. Sci. 15: 45.
the monophyly of Ariocarpus was repeatedly confirmed (Butterworth & al. 2002; Bárcenas & al. 2011;
Hérnandez-Hérnandez & al. 2001; Vázquez-Sánchez
& al. 2013). recent traditional treatments by anderson & Fitz Maurice (1998) and Lüthy & Moser
(2002).
Armatocereus Backeb. in Blätt. Kakteenf. 1938(6): [21].
1938 sec. Hunt (2006). – type: Armatocereus laetus
(Kunth) Backeb.
Arrojadoa Britton & rose, cactaceae 2: 170. 1920 sec.
Hunt (2006). – type: Arrojadoa rhodantha (Gürke)
Britton & rose
= Pierrebraunia esteves in cact. Succ. J. (Los angeles)
69: 296. 1997.
= Arrojadoopsis Guiggi in cactology 1: 26. 2007.
recent floristic treatment by taylor & zappi (2004).
Arthrocereus a. Berger in Kakteen: 146, 337. 1929,
nom. cons. sec. Hunt (2006). – type: Cereus damazi
oi K. Schum.
Astrophytum Lem., cact. Gen. Sp. nov.: 3-6. 1839 sec.
Vázquez-Sánchez & al. (2013). – type: Astrophytum
myriostigma Lem.
= Digitostigma Velazco & nevárez in cact. Suc. Mex.
47: 79. 2002, nom. inval.
confirmed as monophyletic, including Digitostigma;
therefore the transfer of Digitostigma to Astrophytum,
as suggested by Hunt (2003), is justified.
Austrocactus Britton & rose, cactaceae 3: 44. 1922 sec.
Hunt (2006). – type: Austrocactus bertinii (cels)
Britton & rose
Austrocylindropuntia Backeb. in Blätt. Kakteenf. 6: 21.
1938 sec. ritz & al. (2012). – type: Austrocylindro
puntia exaltata (a. Berger) Backeb.
= Andinopuntia Guiggi, cactology 2(Suppl.): [1]. 2011.
= Banfiopuntia Guiggi, cactology 2(Suppl.): [1]. 2011.
= Peruviopuntia Guiggi, cactology 2(Suppl.): [1].
2011.
= Trichopuntia Guiggi, cactology 2(Suppl.): 2. 2011.
Austrocylindropuntia as originally treated in Hunt
(2006) was found as not monophyletic by ritz & al.
(2012). Austrocylindropuntia lagopus (K. Schum.)
F. ritter was found sister to the remaining species
of Austrocylindropuntia and Cumulopuntia and was
310
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
therefore segregated as a monotypic genus Punotia
d. r. Hunt; see also there.
Aylostera Speg. in anales Soc. ci. argent. 96: 75. 1923
sec. ritz & al. (2007). – type: Aylostera pseudomi
nuscula (Speg.) Speg.
= Mediolobivia Backeb. in Blätt. Kakteenf. 1934(2).
1934.
= Digitorebutia Frič & Kreuz. ex Buining in Succulenta
(netherlands) 22. 1940.
See notes under Rebutia. according to the molecular
phylogenetic study of Mosti & al. (2011), Aylostera
falls in two clades, Aylostera s.str. and Mediolobivia
(incl. A. einsteinii (Frič ex Kreuz. & Buining) Mosti &
Papini), which were recognized as subgenera by these
authors. the genus is an excellent example for the
notorious oversplitting prevalent in many cacti: Hunt
(2006) (as Rebutia subg. Rebutia) accepted ten species, while Mosti & al. (2011) argued for 110 species.
Aztekium Boed. in Monatsschr. deutsch. Kakteen-Ges.
1: 52. 1929 sec. Vázquez-Sánchez & al. (2013). –
type: Aztekium ritteri (Boed.) Boed.
the genus contains only three species; two of them
(A. ritteri and A. hintonii Glass & W. a. Fitz Maur.)
have been found well supported as sisters to each other (Vázquez-Sánchez & al. 2013).
Bergerocactus Britton & rose in contr. U. S. natl. Herb.
12: 435. 1909 sec. Hunt (2006). – type: Bergerocac
tus emoryi (engelm.) Britton & rose
Blossfeldia Werderm. in Kakteenkunde 1937: 162. 1937
sec. Hunt (2006). – type: Blossfeldia liliputana
Werderm.
the monotypic peculiar genus Blossfeldia is found as
sister to the rest of the Cactoideae (nyffeler 2002;
Butterworth 2006).
Borzicactus riccob. in Boll. reale Orto Bot. Palermo
8: 261. 1909 sec. Hunt (2012b). – type: Borzicactus
ventimigliae riccob.
= Clistanthocereus Backeb. in Cactaceae (Berlin)
1937(1): 24. 1937.
= Seticereus Backeb. in Kakt. and. Sukk. 1937: 37.
1937.
= Akersia Buining in Succulenta (netherlands) 1961: 25.
1961.
= Borzicactella H. Johnson ex F. ritter, Kakteen Südamerika 4: 1385. 1981.
Borzicactus is reinstated based on the results of
Schlumpberger & renner (2012). its circusmcription
has been the subject of some debate, as summarized
by Bregman (1992). the exact delimitation of Borzi
cactus and the genera currently included or considered related to it is still unclear.
Brachycereus Britton & rose, cactaceae 2: 120. 1920
sec. Hunt (2006). – type: Brachycereus nesioticus
(K. Schum.) Backeb.
Brasilicereus Backeb. in Blätt. Kakteenf. 1938(6): 22.
1938 sec. Hunt (2006). – type: Brasilicereus phaea
canthus (Gürke) Backeb.
= Bragaia esteves, Hofacker & P. J. Braun in Kakteen
and. Sukk. 60(12): 328. 2009.
recent floristic monograph by taylor & zappi (2004).
Brasiliopuntia (K. Schum.) a. Berger, entwicklungslin.
Kakt. 17, 18: 94. 1926 sec. Majure & al. (2012) ≡
Opuntia subg. Brasiliopuntia K. Schum., Gesamtbeschr. Kakt. 1898. – type: Brasiliopuntia brasilien
sis (Willd.) a. Berger
Originally monotypic with B. brasiliensis. Majure &
al. (2012) found good support for a sister-group relation of Opuntia schickendantzii F. a. c. Weber., and
transferred this species to Brasiliopuntia.
Browningia Britton & rose, cactaceae 2: 63. 1920 sec.
Hunt (2006). – type: Browningia candelaris (Meyen)
Britton & rose
= Gymnanthocereus Backeb. in Blätt. Kakteenf.
1937(8): nachtr. 15 [2]. 1937.
= Azureocereus akers & H. Johnson in cact. Succ. J.
(Los angeles) 21: 133. 1949.
= Gymnocereus Backeb., cactaceae Handb. Kakteen.
Pereskioideae Opuntioideae 2: 920. 1959, nom. illeg.
Calymmanthium F. ritter in Kakteen and. Sukk. 13: 25.
1962 sec. Hunt (2006). – type: Calymmanthium sub
sterile F. ritter
Monotypic; sampled by Korotkova & al. (2010) and
resolved as sister to Lymanbensonia.
Carnegiea Britton & rose in J. new York Bot. Gard. 9:
187. 1908 sec. Hunt (2006). – type: Carnegiea gi
gantea (engelm.) Britton & rose – Fig. 3B.
Castellanosia cárdenas in cact. Succ. J. (Los angeles)
23: 90. 1951 sec. Hunt (2006). – type: Castellanosia
caineana cárdenas
Cephalocereus Pfeiff. in allg. Gartenzeitung (Otto &
dietrich) 6: 142. 1838 sec. arias & al. (2012) ≡ Pi
locereus Lem., cact. Gen. Sp. nov.: 6-7. 1839, nom.
illeg. – type: Cephalocereus senilis (Haw.) K. Schum.
= Haseltonia Backeb. in Blätt. Sukkulentenk. 1: 3.
1949.
= Neodawsonia Backeb. in Blätt. Sukkulentenk. 1: 4.
1949.
See under Neobuxbaumia.
Cereus Mill. in Gard. dict. abr., ed. 4: [308]. 1754 sec.
Hunt (2006). – type: Cereus hexagonus (L.) Mill.
= Piptanthocereus (a. Berger) riccob. in Boll. reale
Orto Bot. Palermo 8: 225. 1909.
= Subpilocereus Backeb. in Blätt. Kakteenf. 1938(6).
1938.
= Mirabella F. ritter, Kakteen Südamerika 1: 108.
1979.
Cipocereus F. ritter in Kakteen Südamerika 1: 54. 1979
sec. Hunt (2006). – type: Cipocereus pleurocarpus
F. ritter
= Floribunda F. ritter, Kakteen Südamerika 1: 58.
1979.
recent floristic monograph by taylor & zappi (2004).
Cleistocactus Lem., ill. Hort. 8. 1861 sec. Hunt (2006). –
type: Cleistocactus baumannii (Lem.) Lem.
Willdenowia 45 – 2015
= Maritimocereus akers & Buining in Succulenta
(netherlands) 1950: 49. 1950.
= Bolivicereus cárdenas in cact. Succ. J. (Los angeles)
23: 91. 1951.
= Cephalocleistocactus F. ritter in Succulenta (netherlands): 108. 1959.
= Seticleistocactus Backeb. in descr. cact. nov. 3.
1963.
= Hildewintera F. ritter in Kakteen and. Sukk. 17: 11.
1966, nom. inval.
= Winterocereus Backeb., Kakteenlexikon 455. 1966.
the broad circumscription of Cleistocactus as employed by anderson (2001, 2005), and Hunt (2006)
goes back to the Cactaceae consensus classification
reported by Hunt & taylor (1986), where the predominantly ornithophilous floral syndrome was used as a
diagnostic character. Schlumpberger & renner (2012)
found that Cleistocactus s.l. is polyphyletic – the
monotypic Cephalocleistocactus was placed as sister
to Yungasocereus, with Cleistocactus s.str. as sister to
Vatricania next to Weberbauerocereus, and two terminals representing the former Borzicactus and Loxantho
cereus were placed in the Oreocereus clade, the former
next to Matucana and the latter next to Haageocereus.
deciding whether Cleistocactus s.l. should be retained
or split up is difficult, since sampling of the group and
its possible sister taxa is still inadequate. the affiliation of Loxanthocereus with Haageocereus was seen
earlier, and nyffeler & eggli (2010b) listed it as synonym of Haageocereus.
Coleocephalocereus Backeb. in Blätt. Kakteenf. 1938(6):
[22]. 1938 sec. Hunt (2006). – type: Coleocepha
locereus fluminensis (Miq.) Backeb.
= Buiningia Buxb. in Krainz, Kakteen: 46 – 47, c iV.
1971.
recent floristic monograph by taylor & zappi (2004).
Consolea Lem. in rev. Hort. (Paris) 1862: 174. 1862 sec.
Majure & al. (2012). – type: Consolea spinosissima
(Mill.) Lem.
Plastid and nuclear itS data so far provided inconclusive results for the placement of Consolea and
its separation from Opuntia. Consolea was found to
be imbedded in Opuntia by Griffith & Porter (2009)
based on combined nuclear and plastid data. the plastid and nuclear data of Majure & al. (2012) supported
the monophyly but were incongruent regarding the
placement of Consolea: while plastid data resolved
Consolea outside of Opuntia (BS=53%), nuclear itS
data resolve Consolea within Opuntia (BS=75%),
yet both these placements receive only weak support. Support for a placement outside of Opuntia
increased to 81% BS when only diploids were included in a combined nuclear and plastid analysis.
Majure & al. (2012) pointed out that evolution in
Opuntia and allies involves hybridization and allopolyploidization and that Consolea might be of allopolyploid origin, as indicated by the incongruent
311
plastid and nuclear trees. nevertheless, Majure & al.
(2012) argued for recognizing Consolea as a genus
distinct from Opuntia because of good support for its
monophyly, the placement by combined plastid and
nuclear data outside of Opuntia and unique morphological characteristics.
Copiapoa Britton & rose, cactaceae 3: 85. 1922 sec.
Hunt (2006). – type: Copiapoa marginata (Salmdyck) Britton & rose
= Pilocopiapoa F. ritter in Kakteen and. Sukk. 12: 20.
1961.
recent floristic treatment by Hoffmann & Walter
(2005).
Corryocactus Britton & rose, cactaceae 2: 66. 1920
sec. Hunt (2006). – type: Corryocactus brevistylus
(K. Schum. ex Vaupel) Britton & rose
= Erdisia Britton & rose, cactaceae 2: 104. 1920.
Coryphantha (engelm.) Lem. in cactées: 32. 1868, nom.
cons. prop. sec. Hunt (2006) ≡ Mammillaria subg.
Coryphantha engelm. in Proc. amer. acad. arts 3:
264. 1856. – type: Coryphantha sulcata (engelm.)
Britton & rose
= Lepidocoryphantha Backeb. in Blätt. Kakteenf.
1938(6): 22. 1938.
= Escobrittonia doweld in Sukkulenty 3: 17. 2000.
Found as highly polyphyletic by Bárcenas & al.
(2011), and as nested in Mammillaria. One core Co
ryphantha clade was resoved but only weakly supported as monophyletic (0.65 PP from Bayesian
inference). Vázquez-Sánchez & al. (2013) likewise
found Coryphantha as polyphyletic, but not nested
in Mammillaria; however, far fewer species were
sampled therein. One maximally supported group
was found that also contains Neolloydia matehualen
sis Backeb., while other Coryphantha species were
found close to Echinomastus and Escobaria. as in
the whole mammilloid clade, support for the relevant
nodes is still weak and generic limits of Coryphantha
need further evaluation. See also notes under Mam
millaria and Neollydia. recent traditional monograph
by dicht & Lüthy (2003).
Cumarinia (F. M. Knuth) Buxb. in Oesterr. Bot. z. 98:
61. 1951 sec. Vázquez-Sánchez & al. (2013). – type:
Cumarinia odorata (Boed.) Buxb.
Monotypic; segregated from Coryphantha based on
the results of Vázquez-Sánchez & al. (2013).
Cumulopuntia F. ritter in Kakteen Südamerika 2: 399.
1980 sec. ritz & al. (2012). – type: Cumulopuntia
ignescens (Vaupel) F. ritter
= Sphaeropuntia Guiggi in cactology 3 (Suppl. ii): 1.
2012.
Griffith & Porter (2009) found no support for a
monophyletic Cumulopuntia, but it was also not
contradicted. Cumulopuntia was then confirmed as
monophyletic by ritz & al. (2012). Cumulopuntia
falls in two clades in the molecular phylogeny, one
consisting of C. sphaerica (c. F. Först.) e. F. an-
312
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
derson and related taxa from the W andean slopes
of chile and Peru, characterized by forming dwarf
shrubs with easily detachable stem segments, and
another consisting of Cumulopuntia s.str., characterized by growth in often dense cushions, with firmly
attached stem segments (Griffith & Porter 2009; ritz
& al. 2012). For the C. sphaerica clade, the generic
name Sphaeropuntia was recently published, but its
circumscription is not yet fully resolved, and it this
thus better treated as synonym for the time being.
Cylindropuntia (engelm.) F. M. Knuth, nye Kaktusbog
102. 1930 sec. Hunt (2006) ≡ Opuntia subg. Cylin
dropuntia engelm. in Proc. amer. acad. arts 3: 302.
1856. – type: Cylindropuntia arborescens (engelm.)
F. M. Knuth
Griffith & Porter (2009) found no support for a monophyletic Cylindropuntia based on combined nuclear
and plastid markers, while Bárcenas & al. (2011)
found a monophyletic Cylindropuntia with high to
maximal support based on plastid data only.
Dendrocereus Britton & rose, cactaceae 2: 113. 1920
sec. Hunt (2006). – type: Dendrocereus nudiflorus
(engelm.) Britton & rose
Denmoza Britton & rose, cactaceae 3: 78. 1922 sec.
Hunt (2006). – type: Denmoza rhodacantha (Salmdyck) Britton & rose
Monotypic; distributed in argentina. Formal monograph by Leuenberger (1993).
Discocactus Pfeiff. in allg. Gartenzeitung (Otto & dietrich) 5: 241. 1837 sec. Hunt (2006). – type: Disco
cactus insignis Pfeiff.
recent floristic treatment by taylor & zappi (2004).
Disocactus Lindl., edwards’s Bot. reg. 31: t. 9. 1845 sec.
Hunt (2006). – type: Disocactus biformis (Lindl.)
Lindl.
= Aporocactus Lem. in ill. Hort. 7: misc. 67. 1860.
= Cereus subsect. Heliocereus a. Berger in rep. (annual) Missouri Bot. Gard. 16: 78. 1905 ≡ Heliocereus
(a. Berger) Britton & rose in contr. U. S. natl. Herb.
12: 433. 1909.
= Chiapasia Britton & rose, cactaceae 4: 203. 1923.
= Nopalxochia Britton & rose, cactaceae 4: 204.
1923.
= Bonifazia Standl. & Steyerm. in Publ. Field Mus. nat.
Hist., Bot. Ser. 23: 66. 1944.
= Pseudonopalxochia Backeb., cactaceae Handb. Kakteen. Pereskioideae Opuntioideae 1: 69. 1958.
Echinocactus Link & Otto, Verh. Vereins. Beförd. Gartenbaues Königl. Preuss. Staaten 3: 420. 1827 sec.
Vázquez-Sánchez & al. (2013). – type: Echinocactus
platyacanthus Link & Otto
= Echinofossulocactus Lawr. in Gard. Mag. & reg. rural domest. improv. 17: 317. 1841.
= Homalocephala Britton & rose, cactaceae 3: 181.
1922.
= Emorycactus doweld in Succulenta (netherlands)
75: 270. 1996.
= Meyerocactus doweld in Succulenta (netherlands)
75: 271. 1996.
= Kroenleinia Lodé in cact. avent. int. 102: 25. 2014.
Echinocactus turns out to be paraphyletic in recent
molecular studies (Bárcenas & al. 2011; HernándezHernández & al. 2011; Vázquez-Sánchez & al. 2013),
with E. grusonii Hildm. resolved in a separate clade
from the remaining four species, yet with only low
support (Vázquez-Sánchez & al. 2013) or in a polytomy (Bárcenas & al. 2011). Vázquez-Sánchez &
al. (2013) re-circumscribed Echinocactus to include
only four species, excluding E. grusonii and also E.
polycephalus engelm. & J. M. Bigelow but did not
suggest new generic assignment for these species.
the generic name Kroenleinia was recently erected
for E. grusonii, but it may be premature to accept this
monotypic genus in view of the numerous unresolved
or poorly supported topologies in the group.
Echinocereus engelm. in Wislizenus, Mem. tour n.
Mexico: 91. 1848 sec. Sánchez & al. (2014). – type:
Echinocereus viridiflorus engelm.
= Wilcoxia Britton & rose in contr. U. S. natl. Herb.
12: 434. 1909.
Echinocereus was studied in detail and found as
monophyletic by Sánchez & al. (2014) but excluding
E. pensilis J. a. Purpus, which was resolved distant
from Echinocereus and as as sister to the Stenocereus
group. Because E. pensilis had been regregated as a
monotypic genus Morangaya, its reinstatement was
suggested by Sánchez & al. (2014).
Echinopsis zucc. in abh. Math.-Phys. cl. Königl. Bayer.
akad. Wiss. 2: 675. 1837 sec. Hunt (2006). – type:
Echinopsis eyriesii (turpin) zucc. ex Pfeiffer & Otto
= Trichocereus (a. Berger) riccob. in Boll. reale Orto
Bot. Palermo 8: 236. 1909.
= Chamaecereus Britton & rose, cactaceae 3: 48.
1922.
= Lobivia Britton & rose, cactaceae 3: 49. 1922.
= Acanthocalycium Backeb., Kaktus aBc: 224, 412.
1935.
= Soehrensia Backeb. in Blätt. Kakteenf. 1938(6): [21].
1938.
= Setiechinopsis (Backeb.) de Haas in Succulenta
(netherlands) 22: 9. 1940.
= Acantholobivia Backeb. in cactaceae (Berlin)
1941(2): 76. 1942.
= Pseudolobivia (Backeb.) Backeb. in cactaceae (Berlin) 1941: 76. 1942.
= Reicheocactus Backeb. in cactaceae (Berlin) 1941(2):
76. 1942.
= Helianthocereus Backeb. in cact. Succ. J. Gr. Brit.
11: 53. 1949.
= Leucostele Backeb. in Kakt. and. Sukk. 4: 1. 1953.
the present wide circumscription of Echinopsis goes
back to the mid-1970s. First indications that this
broad Echinopsis is polyphyletic were found by Lendel & al. (2006) and ritz & al. (2007), and Schlump-
Willdenowia 45 – 2015
berger & renner (2012) in their deeply sampled study
indeed found vast polyphyly and paraphyly throughout most of the subtribe Trichocereinae. Species of
Echinopsis were scattered over eight different clades
and interspersed with species of Acanthocalycium,
Arthrocereus, Borzicactus, Cephalocleistocactus,
Cleistocactus, Denmoza, Espostoa, Haageocereus,
Harrisia, Matucana, Mila, Oreocereus, Oroya, Pyg
maeocereus, Rauhocereus, Samaipaticereus, We
berbauerocereus and Yungasocereus, all of which
are part of a highly supported clade (100% BS).
to transform their results into a formal classification of monophyletic genera is no easy task. it
would entail either to further broaden an already
very heterogeneous genus by including the genera mentioned above or to accept about a dozen segregates (valid generic names are at hand).
nevertheless, maintaining Echinopsis (sensu
Hunt 2006) is rather not an option, as it is clearly
polyphyletic and should be split up. the necessary
new combinations are already available (Schlumpberger 2012); a fully revised generic circumscription
is still to be published.
Epiphyllum Haw. in Syn. Pl. Succ.: 197. 1812 sec. Hunt
(2006) ≡ Phyllocactus Link, Handbuch 2: 10. 1829,
nom. illeg. – type: Epiphyllum phyllanthus (L.) Haw.
= Marniera Backeb. in cact. Succ. J. (Los angeles) 22.
1950.
Epithelantha F. a. c. Weber ex Britton & rose, cactaceae 3: 92. 1922 sec. Vázquez-Sánchez & al. (2013).
– type: Epithelantha micromeris (engelm.) F. a. c.
Weber ex Britton & rose
the number of species is in dispute, and the recent
work of donati & zanovello (2011) recognizes about
half a dozen species. So far only E. micromeris sampled in a phylogenetic study (Vázquez-Sánchez & al.
2013), and found in an isolated position within the
tribe Cacteae.
Eriosyce Phil. in anales Univ. chile 41: 721. 1872 sec.
Hunt (2006). – type: Eriosyce sandillon (Gay) Phil.
= Islaya Backeb. in Blätt. Kakteenf. 1834: [3]. 1834.
= Neoporteria Britton & rose, cactaceae 3: 94. 1922.
= Pyrrhocactus a. Berger, Kakteen: 215, 345. 1929.
= Horridocactus Backeb. in Blätt. Kakteenf. 1938(6):
[21]. 1938.
= Neochilenia Backeb. in repert. Spec. nov. regni
Veg. 51: 60. 1942.
= Thelocephala Y. itô, explan. diagr. austroechinocactinae: 292. 1957.
= Rimacactus Mottram in Bradleya 19: 75. 2001.
the diminutive Eriosyce laui Lüthy from northern
chile has been found to differ morphologically from
the remaining taxa of Eriosyce s.l. by nyffeler &
eggli (1997), and was subsequently segregated as the
monotypic genus Rimacactus. as long as this segregation is not backed up by molecular data, it appears
premature to accept the genus. recent treatments by
313
Kattermann (1994) and Hoffmann & Walter (2005;
chile).
Escobaria Britton & rose, cactaceae 4: 53. 1923
sec. Hunt (2006). – type: Escobaria tuberculosa
(engelm.) Britton & rose
= Neobesseya Britton & rose, cactaceae 4: 51. 1923.
= Cochiseia W. H. earle in Saguaroland Bull. 30: 65.
1976.
= Escocoryphantha doweld in Sukkulenty 1999(1): 10.
1999.
See notes under Mammillaria.
Escontria rose in contr. U. S. natl. Herb. 10: 125. 1906
sec. Hunt (2006). – type: Escontria chiotilla (F. a. c.
Weber ex K. Schum.) rose
recent monograph by Gibson (1988a).
Espostoa Britton & rose, cactaceae 2: 60. 1920 sec.
Hunt (2006). – type: Espostoa lanata (Kunth) Britton & rose
= Pseudoespostoa Backeb. in Blätt. Kakteenf. 1834:
gen. 104. 1834 ≡ Binghamia Britton & rose, cactaceae 2: 167. 1920.
= Thrixanthocereus Backeb. in Blätt. Kakteenf.
1937(8): nachtr. 15. 1937.
Espostoopsis Buxb. in Krainz, Kakteen: 38 – 39, c Va.
1968 sec. Hunt (2006). – type: Espostoopsis dy
bowskii (rol.-Goss.) Buxb.
Eulychnia Phil. in Fl. atacam.: 23. 1860 sec. Hunt
(2006). – type: Eulychnia breviflora Phil.
= Philippicereus Backeb. in cactaceae (Berlin) 1941(2):
75. 1942.
recent monograph by Hoffmann & Walter (2005).
Facheiroa Britton & rose, cactaceae 2: 173. 1920 sec.
Hunt (2006). – type: Facheiroa pubiflora Britton &
rose
= Zehntnerella Britton & rose, cactaceae 2: 176. 1920.
recent floristic treatment by taylor & zappi (2004).
Ferocactus Britton & rose, cactaceae 3: 123. 1922 sec.
Vázquez-Sánchez & al. (2013). – type: Ferocactus
wislizeni (engelm.) Britton & rose
= Bisnaga Orcutt in cactography 1. 1926 ≡ Ferocactus
sect. Bisnaga (Orcutt) n. P. taylor & J. Y. clark in
Bradleya 1: 6. 1983.
Vázquez-Sánchez & al. (2013) found Ferocactus in
its current circumscription to be vastly polyphyletic,
and the same is true for F. sect. Bisnaga. the Ferocac
tus clade found by Vázquez-Sánchez & al. (2013) also
includes the genera Glandulicactus, Leuchtenbergia,
Stenocactus and Thelocactus, corroborating the results of a much less dense sampling by HernándezHernández & al. (2011). the Ferocactus clade is morphologically characterized by pericarpels with scales
and ribbed stems, and Vázquez-Sánchez & al. (2013)
suggested expanding Ferocactus to embrace the genera just mentioned as the best taxonomic solution to
make Ferocactus monophyletic, yet Leuchtenbergia
is the oldest name of this assemblage and would have
priority, unless the name Ferocactus is conserved.
314
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
Frailea Britton & rose, cactaceae 3: 208. 1922 sec.
Hunt (2006). – type: Frailea cataphracta (dams)
Britton & rose
Geohintonia Glass & W. a. Fitz. Maur. in cact. Suc.
Mex. 37: 16. 1992 sec. Vázquez-Sánchez & al.
(2013). – type: Geohintonia mexicana Glass & W.
a. Fitz. Maur.
Monotypic; sampled by Vázquez-Sánchez & al.
(2013) and resolved as sister to Aztekium.
Glandulicactus Backeb. in Blätt. Kakteenf. 1938(6):
[22]. 1938 sec. Hunt (2006). – type: Glandulicactus
uncinatus (Galeotti ex Pfeiff.) Backeb.
See notes under Ferocactus.
Grusonia rchb. f. ex Britton & rose, cactaceae 1: 215.
1919 sec. Hunt (2006). – type: Grusonia bradtiana
(J. M. coult.) Britton & rose
= Corynopuntia F. M. Knuth, Kaktus aBc: 114, 410.
1936.
= Micropuntia daston in amer. Midl. naturalist 36:
661. 1946.
= Marenopuntia Backeb. in desert Pl. Life 22: 27. 1950.
Corynopuntia was included in Grusonia s.l. (Wallace & dickie 2002; anderson 2001, 2005; nyffeler
& eggli 2010b), then accepted as separate genus by
Hunt (2006). Griffith & Porter (2009) argued for recognizing Corynopuntia as a distinct genus, and Gru
sonia as monotypic, although support for the monophyly of Corynopuntia was only 67% BS in their
study. their data also suggest that Micropuntia could
be recognized as a separate genus. Bárcenas & al.
(2011) did not find support for treating Corynopuntia
separately from Grusonia, and in addition found no
support for a monophyletic Corynopuntia; therefore,
the circumscription of Corynopuntia still needs to be
clarified.
Gymnocalycium Pfeiff. ex Mittler, taschenb. cactuslieb.
2: 124. 1844 “Gymnocalicium” sec. demaio & al.
(2011). – type: Gymnocalycium gibbosum (Haw.)
Pfeiff. ex Mittler
Found as monophyletic at first by ritz & al. (2007),
then studied in more detail and confirmed as monophyletic by Meregalli & al. (2010). demaio & al.
(2011) conducted the most detailed phylogenetic
study by sampling almost the whole genus and again
confirmed the monophyly of Gymnocalycium with
maximal support. recent illustrated synopsis by
charles (2009).
Haageocereus Backeb. in Blätt. Kakteenf. 1934(6): [1].
1934 sec. Hunt (2006). – type: Haageocereus pseu
domelanostele (Werderm. & Backeb.) Backeb.
= Loxanthocereus Backeb. in cactaceae (Berlin)
1937(1): 24. 1937.
= Peruvocereus akers in cact. Succ. J. (Los angeles)
19: 67. 1947.
= Maritinocereus akers & Buining in Succulenta
(netherlands) 1950: 49. 1950.
recent monograph by calderón & al. (2007).
Harrisia Britton in Bull. torrey Bot. club 35: 561. 1909
sec. Franck & al. (2013a). – type: Harrisia gracilis
(Mill.) Britton
= Eriocereus riccob. in Boll. reale Orto Bot. Palermo
8: 238. 1909.
= Roseocereus Backeb. in Blätt. Kakteenf. 1938(6): 21.
1938.
= Estevesia P. J. Braun in Kakteen and. Sukk. 60(3):
64. 2009.
Harrisia was confirmed as monophyletic by Franck
(2012), with a revised infrageneric classification published shortly after (Franck & al. 2013a). the recently
described genus Estevesia P. J. Braun was not included in any molecular study so far. it was provisionally
placed in the synonymy of Harrisia by nyffeler &
eggli (2010b). For synopsis see Franck (2012); further phylogenetic studies see Franck & al. (2013b).
Hatiora Britton & rose in L. H. Bailey, Standard cycl.
Hort.: 1432. 1915 sec. Korotkova & al. (2011). –
type: Hatiora salicornioides (Haw.) Britton & rose
= Pseudozygocactus Backeb. in Blätt. Kakteenf.
1938(6): [5, 21]. 1938.
the circumscription of Hatiora has been clarified recently. Hatiora including Rhipsalidopsis as adopted
by Barthlott (1987), Barthlott & Hunt (1993), Barthlott & taylor (1995), Hunt (2006) and nyffeler &
eggli (2010b) was found to be polyphyletic (calvente
& al. 2011; Korotkova & al. 2011). Hatiora should
therefore be restricted to species with cylindrical
stems, terete pericarpels, and small yellow-orange
or magenta flowers, corresponding to Hatiora in the
traditional sense. accordingly, Rhipsalidopsis in its
traditional circumscription should again be accepted
at generic rank.
Hylocereus (a. Berger) Britton & rose in contr. U. S.
natl. Herb. 12: 428. 1909 sec. Hunt (2006) ≡ Cereus
subg. Hylocereus a. Berger in rep. (annual) Missouri Bot. Gard. 16: 72. 1905. – type: Hylocereus
triangularis (L.) Britton & rose
= Wilmattea Britton & rose, cactaceae 2: 195. 1920.
Hylocereus is morphologically very similar to Selen
icereus, and available phylogenetic studies as well as
morphological and anatomical data so far suggest that
the two genera could be merged (Hernández-Hernández & al. 2011; Bárcenas & al. 2011, Gómez-Hinostrosa & al. 2014), but they still need to be studied more
extensively before firm conclusions on their circumscription are possible.
Jasminocereus Britton & rose, cactaceae 2: 146. 1920
sec. Hunt (2006). – type: Jasminocereus thoarsii (F.
a. c. Weber) Backeb.
Lasiocereus F. ritter in Kakteen Südamerika 4: 1477.
1981 sec. Hunt (2006). – type: Lasiocereus rupicola
F. ritter
Lemaireocereus Britton & rose in contr. U. S. natl.
Herb. 12: 424. 1909 sec. arias & al. (2012) ≡ Pachyc
ereus subg. Lemaireocereus (Britton & rose) Bravo
Willdenowia 45 – 2015
in cact. Suc. Mex. 17: 119. 1972 ≡ Pachycereus sect.
Lemaireocereus (Britton & rose) P. V. Heath in calyx 2: 106. 1992. – type: Lemaireocereus hollianus
(F. a. c. Weber) Britton & rose
= Anisocereus Backeb. in Blätt. Kakteenf. 1938(6): 21.
1938.
in the second half of the 20th century, Lemaireocer
eus was referred to as a synonym of Pachycereus (see
there) by Buxbaum (1961), Bravo-Hollis (1978), and
Gibson & Horak (1978), based on similar floral morphology. Phylogenies based on molecular (arias &
al. 2003) and structural data (arias & terrazas 2006)
consistently have revealed that Lemaireocereus is
an early-diversified lineage within Pachycereinae.
Lemaireocereus should be restricted to species with
rounded ribs, terminal flowers with long hairs and
bristles, fruit with irregular dehiscence, and red pulp
(arias & terrazas 2009; arias & al. 2012).
Leocereus Britton & rose, cactaceae 2: 108. 1920 sec.
Hunt (2006). – type: Leocereus bahiensis Britton &
rose
recent floristic treatment by taylor & zappi (2004).
Lepismium Pfeiff. in allg. Gartenzeitung 3: 315. 1835
sec. Korotkova & al. (2011). – type: Lepismium com
mune Pfeiff.
= Nothorhipsalis doweld in Sukkulenty 4(1 – 2): 29.
2002.
= Ophiorhipsalis (K. Schumann) doweld in Sukkulenty 4(1 – 2): 39. 2002.
Several considerably different generic concepts have
been suggested for Lepismium in the past 80 years. it
was either recognized as monotypic for L. cruciforme
(Vell.) Miq., e.g. by Britton & rose (1923) or included into Rhipsalis (Schumann 1899; Vaupel 1925,
1926). Barthlott (1987) and Barthlott & taylor (1995)
redefined Lepismium based on the mesotonic branching as the main diagnostic character, but this circumscription was found to be polyphyletic by nyffeler
(2002) and Korotkova & al. (2010). consequently,
some of its species were transferred to Lymanbenso
nia and Pfeiffera by Korotkova & al. (2010). recent
monograph by Barthlott & taylor (1995).
Leptocereus (a. Berger) Britton & rose in contr. U. S.
natl. Herb. 12: 433. 1909 sec. Hunt (2006) ≡ Cereus
subg. Leptocereus a. Berger in rep. (annual) Missouri Bot. Gard. 16: 79. 1905. – type: Leptocereus
assurgens (c. Wright ex Griseb.) Britton & rose
= Neoabbottia Britton & rose in Smithsonian Misc.
collect. 72: 2. 1921.
Leuchtenbergia Hook. in Bot. Mag.: 4393. 1848 sec.
Hunt (2006). – type: Leuchtenbergia principis
Hook.
See notes under Ferocactus.
Leuenbergeria Lodé in cact. avent. int. 97: 26. 2012
sec. Lodé (2012) ≡ Pereskia subg. Leuenbergera G.
d. rowley in cactaceae Syst. init. 32: 7. 2014. –
type: Leuenbergia quisqueyana (alain) Lodé
315
Segregated from Pereskia to include the northern
clade; see note under Pereskia.
Lophocereus (a. Berger) Britton & rose in contr. U.
S. natl. Herb. 12: 426. 1909 sec. arias & al. (2012)
≡ Cereus subg. Lophocereus a. Berger in rep. (annual) Missouri Bot. Gard. 16: 62. 1905 ≡ Pachycereus
sect. Lophocereus (a. Berger) P. V. Heath in calyx
2: 106. 1992. – type: Lophocereus schottii (engelm.)
Britton & rose
= Marginatocereus (Backeb.) Backeb., cact. Jahrb.
deutsch. Kakt.-Ges. 77. 1941 (1942).
Lophocereus (including L. gates and L. schottii) was
strongly recognized as a separate genus, restricted
to the Sonoran desert, by e.g. Lindsay (1963) and
Bravo-Holis (1978). comparative studies carried out
by Gibson & Horak (1978) showed that those species
share morphological and anatomical characteristics
with Pachycereus marginatus (dc.) Britton & rose.
However, other taxonomists preferred to include this
genus and others (e.g. Backebergia, Lemaireocer
eus, Marginatocereus, Mitrocereus, Pterocereus)
in a broader genus Pachycereus (Barthlott & Hunt
1993; Hunt 2006). Phylogenetic studies based on
structural (anatomy + morphology) and molecular
data confirm that Lophocereus is monophyletic including three species (L. marginatus (dc.) S. arias
& terrazas as sister to the remaining species). the
genus represents a lineage within the subtribe Pach
ycereinae, but is not directly related to Pachycer
eus s.str. or Backebergia (see there; Hartmann S. &
al. 2001, 2002; arias & al. 2003; arias & terrazas
2006). a proposal to recognize this genus newly
circumscribed (now going also beyond the Sonoran desert) was conducted by arias & al. (2012).
Lophocereus now includes taxa characterized by cylindrical stems with basal branching, an apical fertile
zone with areoles, and spines larger than those of the
sterile zone, and two or more flowers per areole. the
flowering zone is conspicuously modified in all three
species, although in L. gatesii M. e. Jones and L.
schottii internodes are shorter and spines are longer
(arias & terrazas 2009; arias & al. 2012). Structural changes in the fertile zone exist between several
genera of Pachycereinae, including cephalium (e.g.
Backebergia and Cephalocereus species), pseudocephalium (e.g. Lophocereus and Neobuxbaumia species) and intermediate forms. However, those structures are highly homoplastic and occur within several
genera.
Lophophora J. M. coult. in contr. U. S. natl. Herb.
3: 131. 1894 sec. Vázquez-Sánchez & al. (2013). –
type: Lophophora williamsii (Lem. ex Salm-dyck)
J. M. coult.
Butterworth & al. (2002) found L. williamsii as sister
to Obregonia and L. diffusa (croizat) Bravo as sister to Acharagma, yet both with only moderate support. in contrast, Lophophora williamsii and L. dif
316
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
fusa were resolved as sisters with moderate support
in the study of Vázquez-Sánchez & al. (2013), who
also found high support for the sister relationship of
Lophophora and Obregonia, justifying generic rank
for both.
Lymanbensonia Kimnach in cact. Succ. J. (Los angeles) 56: 101. 1984 sec. Korotkova & al. (2010). –
type: Lymanbensonia micrantha (Vaupel) Kimnach
= Acanthorhipsalis Kimnach in cact. Succ. J. (Los angeles) 55: 177. 1983, nom. illeg.
Segregated from Acanthorhipsalis (Kimnach 1984),
but otherwise either assigned to Lepismium (Barthlott 1987; Barthlott & taylor 1995; anderson 2001,
2005) or to Pfeiffera (Hunt 2006). the molecular
phylogenetic study of Korotkova & al. (2010) unexpectedly found the three species now assigned to
Lymanbensonia to represent a highly supported isolated clade distant from either Lepismium or Pfeiffera.
as this new clade contained the nomenclatural type
of Lymanbensonia, this generic name was reinstated.
Maihuenia (Phil. ex F. a. c. Weber) K. Schum. in Gesamtbeschr. Kakt. 651: 754. 1898 sec. Hunt (2006)
≡ Pereskia subg. Maihuenia Phil. ex F. a. c. Weber
in Bois, dict. Hort. 2: 938. 1898. – type: Maihuenia
poeppigii (Otto ex Pfeiff.) F. a. c. Weber ex K. Schum.
recent monograph by Leuenberger (1997).
Maihueniopsis Speg. in anales Soc. ci. argent. 99: 86.
1925 sec. ritz & al. (2012). – type: Maihueniopsis
molfinoi Speg.
= Puna r. Kiesling in Hickenia 1: 289. 1982 ≡ Mai
hueniopsis subg. Puna (r. Kiesling) Stuppy in Succ.
Pl. res. 6: 50. 2002.
Griffith & Porter (2009) found Maihueniopsis
polyphyletic based on a combined analysis of nuclear
itS and plastid trnLF, but ritz & al. (2012) found
a monophyletic Maihueniopsis to be strongly supported by nuclear phyC and plastid trnK/matK. the
reasons for these deviating results are discussed in detail by ritz & al. (2012) and appear to result from peculiarities in the evolution of the itS sequences used
by Griffith & Porter (2009) that seem unsuitable to
adequately represent phylogenetic relationships.
Mammillaria Haw. in Syn. Pl. Succ.: 177. 1812, nom.
cons. sec. Hunt (2006) ≡ Cactus L., Sp. Pl. 1: 466.
1753 ≡ Neomammillaria Britton & rose, cactaceae
4: 65. 1923. – type: Mammillaria simplex Haw.
= Mammillaria subg. Cochemiea K. Brandegee, erythea 5: 113. 1897.
= Cochemiea (K. Brandegee) Walton in cact. J. (London) 2: 50. 1899.
= Bartschella Britton & rose, cactaceae 4: 57. 1923.
= Dolichothele (K. Schum.) Britton & rose, cactaceae
4: 61. 1923.
= Mamillopsis Britton & rose, cactaceae 4: 19. 1923.
= Phellosperma Britton & rose, cactaceae 4: 60. 1923.
= Solisia Britton & rose, cactaceae 4: 64. 1923.
= Chilita Orcutt in cactography 2. 1926.
= Porfiria Boed. in z. Sukkulentenk. 2. 1926.
= Krainzia Backeb. in Blätt. Kakteenf. 1938(6): [22].
1938.
= Mammilloydia Buxb. in Oesterr. Bot. z. 98: 64. 1951.
= Oehmea Buxb. in Sukkulentenk. 7: 17. 1951.
= Pseudomammillaria Buxb. in Oesterr. Bot. z. 98: 84.
1951.
= Leptocladodia Buxb. in Oesterr. Bot. z. 101. 1954.
= Escobariopsis doweld in Sukkulenty 3: 23. 2000.
Mammillaria is the largest genus within Cacta
ceae, and numerous suggestions for infrageneric
entities have been proposed, often then segregated
as different genera; the different taxonomic concepts were summarized by Butterworth & Wallace
(2004). although several phylogenetic studies dealing with the genus and allies have been published,
there are still many uncertainties that result from
insufficient phylogenetic resolution and support.
Mammillaria was studied in detail using data from the
plastid rpl16 intron and psbA-trnH intergenic spacer
by Butterworth & Wallace (2004), who sampled c. 4/5
of the accepted species, and Bárcenas & al. (2011) for
trnK/matK compiled an even more extensive sampling.
Mammillaria was also included in the phylogenetic studies of the tribe Cacteae by Butterworth & al. (2002) and Vázquez-Sánchez & al.
(2013), though with much fewer species sampled.
the first sequence data already hinted at a non-monophyly of Mammillaria (Butterworth & al. 2002), yet
without support. the results of Butterworth & Wallace
(2004), based on a detailed sampling, again suggested
polyphyly of Mammillaria. the genera Coryphantha,
Escobaria, Mammilloydia, Neolloydia, Ortegocactus
and Pelecyphora were found nested in a maximally
supported Mammillaria s.l. clade. Bárcenas & al.
(2011) did not find sufficient support for a monophyletic Mammillaria, and Coryphantha (likewise
polyphyletic), Escobaria and Ortegocactus were nested in different Mammillaria clades. Vázquez-Sánchez
& al. (2013) found that Coryphantha and Mammil
laria could be separate clades, yet Mammillaria was
supported as monophyletic only in the parsimony tree
(61% BS/78% JK), but not found as monophyletic
by Bayesian inference. a clade of Coryphantha incl.
Neolloydia was maximally supported in the parsimony and Bayesian trees, but C. macromeris (engelm.)
Lem. fell outside that clade, suggesting that Cory
phantha is likewise polyphyletic. Escobaria was found
polyphyletic as well (Vázquez-Sánchez & al. 2013),
but only few species have been sampled. the results
of Vázquez-Sánchez & al. (2013) also provided some
insights into generic limits in the whole assemblage, as
well as taxonomic changes by segregating Cochemiea
from Mammillaria, and Cumarinia from Coryphantha.
Mammilloydia was found nested in Mammillaria (Butterworth & al. 2002; Butterworth & Wallace 2004; Bárcenas & al. 2011; Vázquez-Sánchez & al. 2013), and
Willdenowia 45 – 2015
all authors argue Mammilloydia should therefore no
longer be recognized at generic rank. the Mammil
laria assemblage therefore remains one of the Cac
taceae groups that need further detailed study. Some
nodes were so far only weakly supported, and final
conclusions regarding the monophyly and generic
limits of Mammillara must await a more extensive
sampling, especially for Coryphantha and Escobaria;
only then will firm taxonomic and nomenclatural
conclusions be possible.
Matucana Britton & rose, cactaceae 3: 102. 1922 sec.
Hunt (2006). – type: Matucana haynei (Otto ex
Salm-dyck) Britton & rose
= Submatucana Backeb., cactaceae Handb. Kakteen.
Pereskioideae Opuntioideae 2: 1059. 1959.
= Eomatucana F. ritter in Kakteen and. Sukk. 16: 230.
1965.
recent monograph by Bregmann (1996).
Melocactus Link & Otto in Verh. Preuss. Ver. Gartenb.
3: 417. 1827, nom. cons. sec. Hunt (2006). – type:
Cactus melocactus L.
recent monograph by taylor (1991); recent floristic
treatment by taylor & zappi (2004; Brazil).
Micranthocereus Backeb. in Blätt. Kakteenf. 1938(6):
[22]. 1938 sec. Hunt (2006). – type: Micrantho
cereus polyanthus (Werderm.) Backeb.
= Austrocephalocereus Backeb. in Blätt. Kakteenf.
1938(6): [22]. 1938.
= Siccobaccatus P. J. Braun & esteves in Succulenta
(netherlands) 69: 6. 1990.
recent floristic treatment by taylor & zappi (2004).
Mila Britton & rose, cactaceae 3: 211. 1922 sec. Hunt
(2006). – type: Mila caespitosa Britton & rose
Miqueliopuntia Frič ex F. ritter in Kakteen Südamerika
3: 869. 1980 sec. Hunt (2006). – type: Miquelio
puntia miquelii (Monv.) F. ritter
Monotypic; sampled by Griffith & Porter (2009) and
not found nested in any other genus, justifying generic rank.
Morangaya G. d. rowley in ashingtonia 1: 44. 1944
sec. Sánchez & al. (2014). – type: Morangaya pensi
lis (K. Brandegee) G. d. rowley
See note for Echinocereus.
Myrtillocactus console in Boll. reale Orto Bot. Palermo
1: 10. 1897 sec. Hunt (2006). – type: Myrtillocactus
geometrizans (Mart. ex Pfeiff.) console
Neobuxbaumia Backeb. in Blätt. Kakteenf. 6: 17; 8, 12,
24. 1938 sec. arias & al. (2012). – type: Neobuxbaum
ia tetetzo (J. M. coult.) Backeb.
= Rooksbya (Backeb.) Backeb., cactaceae Handb. Kakteen. Pereskioideae Opuntioideae 4: 2165. 1960.
Phylogenetic studies so far resolved Neobuxbaumia
as closely related to Cephalocereus and Pseudomi
trocereus (arias & al. 2003; arias & terrazas 2006;
Hernández-Hernández & al. 2011). However, these
studies did not specifically focus on Neobuxbaumia,
and its generic limits are therefore not yet firmly estab-
317
lished. arias & al. (2003) found Neobuxbaumia in a
weakly supported polytomy with Cephalocereus and
Pachycereus fulviceps (F. a. c. Weber ex Schumann)
d. r. Hunt (= Pseudomitrocereus) as sister to both.
the two Cephalocereus species were well supported
as sister to each other, but could not be separated from
Neobuxbaumia in any tree (arias & al. 2003). Bárcenas & al. (2011) and Hernández-Hernández & al.
(2011) found Neobuxbaumia to be polyphyletic but
the relevant nodes were weakly supported, therefore
a monophyletic Neobuxbaumia is neither confirmed
not contradicted by the currently available data.
Neolloydia Britton & rose in Bull. torrey Bot. club 49:
251. 1922 sec. Hunt (2006). – type: Neolloydia co
noidea (dc.) Britton & rose
Found to be polyphyletic by Vázquez-Sánchez & al.
(2013), with the type species sister to the rest of the
mammilloid clade, but support <50%, while N. mate
hualensis Backeb. was nested in Coryphantha with
maximal support.
Neoraimondia Britton & rose, cactaceae 2: 181. 1920
sec. Hunt (2006). – type: Neoraimondia macrostibas
(K. Schum.) Britton & rose
= Neocardenasia Backeb. in Blätt. Sukkulentenk. 1: 2.
1949.
Neowerdermannia Frič in Kaktusár 1: 85. 1930 sec. Hunt
(2006). – type: Neowerdermannia vorwerkii Frič
Nyctocereus (a. Berger) Britton & rose in contr. U. S.
natl. Herb. 12: 423. 1909 sec. arias & al. (2005). –
type: Nyctocereus serpentinus (Lag. & rodr.) Britton
& rose
Monotypic; segregated from Peniocereus by arias &
al. (2005) because N. serpentinus was resolved distant from the Peniocereus clade.
Obregonia Frič in zivot v Prirod 29(2): 3. 1925 sec.
Vázquez-Sánchez & al. (2013). – type: Obregonia
denegrii Frič
See notes under Lophophora.
Opuntia Mill. in Gard. dict. abr., ed. 4: [974]. 1754 sec.
Majure & al. (2012). – type: Opuntia vulgaris Mill.
– Fig. 3c.
= Nopalea Salm-dyck, cact. Hort. dyck. (1849): 6364, 233. 1850.
Opuntia is the second-largest genus of the family
Cactaceae. as in all species-rich Cactaceae groups,
numerous different generic conceps with a varying
number of segregate genera have been suggested for
Opuntia. Both extensive splitting (e.g. Backeberg
1966) or lumping into a broadly defined Opuntia were
put forward (rowley 1958; Benson 1982). the first
phylogenetic study by Wallace & dickie (2002) based
on the rpl16 intron found Opuntia in the broad sense
to be polyphyletic. For the revised generic classification they presented based on their data, they argued for
splitting Opuntia, because otherwise further genera
(e.g. Pereskiopsis, Pterocactus) were also nested within Opuntia and merging those would make Opuntia a
318
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
highly heterogenous assemblage. therefore, Wallace
& dickie suggested reinstating the earlier-proposed
Opuntia segregates Austrocylindropuntia, Brasilio
puntia, Consolea, Corynopuntia, Cumulopuntia,
Cylindropuntia, Grusonia, Maihueniopsis, Miquelio
puntia, Nopalea, Tephrocactus, and Tunilla. Opuntia
s.str. was thus restricted to the taxa with flattened
stems and reticulate pollen. this concept was entirely
adapted by Hunt (2006), and largely by nyffeler &
eggli (2010b). Griffith & Porter (2009), using data
from plastid trnLF and nuclear itS, found Opuntia
in this restricted sense to additionally include Con
solea and Nopalea, the clade including all these genera received 100% support, and both Consolea and
Nopalea were also as monophyletic with 100%. the
tree resolution, however, did not allow an immediate
conclusion on the delimitation of these genera. Nopa
lea used to be separated from Opuntia s.str. because it
differs primarily in its hummingbird-syndrome flowers. nevertheless, it was repeatedly found to be nested
in Opuntia (Wallace & Gibson 2002; Griffith & Porter 2009; Bárcenas & al. 2011; Hernández-Hernández
& al. 2011; Majure & al. 2012) and is therefore no
longer maintained as separate genus. the relationship
of Consolea to Opuntia has remained more difficult
to resolve, but available data suggest it is not part of
Opuntia (see also notes under Consolea).
Oreocereus (a. Berger) riccob. in Boll. reale Orto Bot.
Palermo 8: 258. 1909 sec. Hunt (2006) ≡ Cereus subg.
Oreocereus a. Berger in rep. (annual) Missouri Bot.
Gard. 16: 64. 1905. – type: Oreocereus celsianus
(Lem. ex Salm-dyck) riccob.
= Arequipa Britton & rose, cactaceae 3: 100. 1922.
= Morawetzia Backeb. in Jahrb. deutsch. Kakteen-Ges.
1: 73. 1936.
= Arequipiopsis Kreuz. & Buining in repert. Spec.
nov. regni Veg. 50: 198. 1941.
Oroya Britton & rose, cactaceae 3: 102. 1922 sec. Hunt
(2006). – type: Oroya peruviana (K. Schum.) Britton
& rose
Ortegocactus alexander in cact. Succ. J. (Los angeles)
33: 39. 1961 sec. Vázquez-Sánchez & al. (2013). –
type: Ortegocactus macdougallii alexander
Merging Ortegocactus into Mammillaria was proposed by Hunt & taylor (1990) and Barthlott & Hunt
(1993). the sole species, O. macdougallii, was first
sampled by Butterworth & Wallace (2004) and found
nested in Mammillaria, so the authors argued future
transfer to Mammillaria may be justified, but must
await further clarification of generic limits in this
group. Vázquez-Sánchez & al. (2013) found O. mac
dougallii not nested in Mammillaria, but in a weakly
supported polytomy in the mammilloid clade, suggesting maintaining it as a separate genus for the time
being.
Pachycereus (a. Berger) Britton & rose in contr. U.
S. natl. Herb. 12: 420. 1909 sec. arias & terrazas
(2009) ≡ Cereus subg. Pachycereus a. Berger in rep.
(annual) Missouri Bot. Gard. 16: 63. 1905. – type:
Pachycereus pringlei (S. Watson) Britton & rose
= Backebergia Bravo in anales inst. Biol. Univ. nac.
México 24: 230. 1954.
= Pterocereus t. Macdoug. & Miranda in ceiba 4: 135.
1954.
Phylogenetic studies based on morphological and
molecular data show consistently that Pachycereus
s.str. is a monophyletic group with five species
(arias & al. 2003; arias & terrazas 2006, 2009;
arias & al. 2012). Other species previously considered in Pachycereus (Buxbaum 1961; Gibson &
Horak 1978; anderson 2001; Hunt 2006; nyffeler
& eggli 2010b) have been transferred to Lemaireo
cereus, Lophocereus, and Pseudomitrocereus. More
inclusive and robust new evidence may corroborate or refute the current delimitation of these last
genera. Pachycereus s.str. includes tree-like species, interareolar grooves on the stems, abundant
trichomes on the flower, and flexible spines on the
fruit. the genera Backebergia and Pterocereus (both
monotypic) remain inconclusive on molecular data
available (arias & al. 2003; Hernández-Hernández
2011); therefore their recognition as separate genera
remains premature. recent monograph by arias &
terrazas (2009).
Parodia Speg. in anales Soc. ci. argent. 96: 70. 1923
sec. Hunt (2006). – type: Parodia microsperma (F. a.
c. Weber) Speg.
= Malacocarpus Salm-dyck, cact. Hort. dyck. (1849):
24-25, 141. 1850 ≡ Wigginsia d. M. Porter in taxon
13: 210. 1964.
= Notocactus (K. Schum.) Frič in cacti Price-List 1928:
[3]. 1928.
= Acanthocephala Bakckeb. in Blätt. Kakteenf.
1938(6): [7]. 1938.
= Eriocephala Backeb. in Blätt. Kakteenf. 1938(6): [7,
21]. 1938 ≡ Eriocactus Backeb. in cactaceae (Berlin)
1941: 76. 1942.
= Brasilicactus Backeb. in cactaceae (Berlin) 1941: 76.
1942.
= Brasiliparodia F. ritter, Kakteen Südamerika 1: 144.
1979.
Pediocactus Britton & rose in ill. Fl. n. U. S. (Britton &
Brown) 2: 569. 1913 sec. Hunt (2006). – type: Pedio
cactus simpsonii (engelm.) Britton & rose
= Utahia Britton & rose, cactaceae 3: 215. 1922.
= Navajoa croizat in cact. Succ. J. (Los angeles) 15:
89. 1943.
= Pilocanthus B. W. Benson & Backeb. in Kakteen
and. Sukk. 8: 188. 1957.
= Neonavajoa doweld in Sukkulenty 1999(2): 24.
1999.
= Puebloa doweld in Sukkulenty 1999(1): 20. 1999.
recent monographs by Heil & al. (1981) and Hochstätter (2007).
Willdenowia 45 – 2015
Pelecyphora ehrenb. in Bot. zeitung (Berlin) 1: 737.
1843 sec. Vázquez-Sánchez & al. (2013). – type: Pe
lecyphora aselliformis ehrenb.
= Encephalocarpus a. Berger, Kakteen: 331. 1929.
the generic limits are not yet clarified, Pelecyphora
was found monophyletic by Vázquez-Sánchez & al.
(2013) and Bárcenas & al. (2011), who additionally
found Escobaria paraphyletic to Pelecyphora.
Peniocereus (a. Berger) Britton & rose in contr. U. S.
natl. Herb. 12: 428. 1909 sec. arias & al. (2005) ≡
Cereus subsect. Peniocereus a. Berger in rep. (annual) Missouri Bot. Gard. 16: 77. 1905. – type: Peni
ocereus greggii (engelm.) Britton & rose
= Neoevansia W. t. Marshall, cactaceae (Marshall &
Bock): 84. 1941.
= Cullmannia distefano in Kakteen and. Sukk. 7: 8.
1956.
the generic circumscription of Peniocereus was revised based on the molecular phylogenetic study
of arias & al. (2005). their study based on plastid
trnLF and rpl16 found Peniocereus polyphyletic,
its species resolved in three lineages. Peniocereus
subg. Pseudoacanthocereus Sánchez-Mej. was found
to be nested in Acanthocereus, yet both were also
paraphyletic. For a classification reflecting these relationships, Peniocereus subg. Pseudoacanthocereus
would need to be transferred to Acanthocereus. the
other major Peniocereus clade found by arias & al.
(2005) corresponds to Peniocereus subg. Peniocereus.
Peniocereus serpentinus (Lag. & rodr.) n. P. taylor
was resolved as a separate lineage. Since it is the type
species of the earlier-proposed genus Nyctocereus,
arias & al. (2005) suggested reinstating it as monotypic.
Pereskia Mill. in Gard. dict. abr., ed. 4: [1026]. 1754
sec. Hunt (2006). – type: Pereskia aculeata Mill. –
Fig. 3d.
= Pereskia sect. Rhodocactus a. Berger, Kakteen: 43.
1929 ≡ Rhodocactus (a. Berger) F. M. Knuth, nye
Kaktusbog: 102. 1930.
Pereskia has been repeatedly found to be paraphyletic by nyffeler (2002), edwards & al. (2005), and
Butterworth & edwards (2008). the genus forms
a grade at the base of the Cactaceae, with a northern clade including Mesoamerican and caribbean
pereskias as the first branching group followed by a
southern clade, with mainly the andean pereskias,
which also include the nomenclatural type of Pere
skia (Butterworth & Wallace 2005; edwards & al.
2005). no nomenclatural changes to reflect the paraphyly of Pereskia were proposed by edwards & al.
(2005), who preferred their results to be tested with
additional genes before suggesting a new classification for Pereskia. also, no generic name was readily
available for the northern Pereskia clade – the type
of the earlier-proposed segregate Rhodocactus was
in the southern clade together with the type of Pere
319
skia itself. Pereskia was accepted as polyphyletic to
reflect its morphological differences to the rest of the
Cactaceae. Both Pereskia clades have characters that
are interpreted as ancestral within Cactaceae, such
as a woody stem, the presence of true leaves, a flower
morphology that differs from the rest of the Cacta
ceae and c3 photosynthesis. Only recently, the northern pereskias were segregated as Leuenbergeria, yet
this segregation also received criticism because the
two clades are hard to distinguish morphologically
(http://www.mobot.org/MOBOt/research/edge/
apr13/apr13lit.shtml; Hunt 2013). Seeking a compromise between molecular phylogenetic hypotheses
and nomenclatural stability, rowley (2013) suggested a subgenus Leuenbergera (note the different spelling!) for the northern Pereskia clade. Monograph by
Leuenberger (1986).
Pereskiopsis Britton & rose in Smithsonian Misc. collect. 50: 331. 1907 sec. Hunt (2006). – type: Pereski
opsis brandegeei (K. Schum.) Britton & rose
Pfeiffera Salm-dyck in cact. Hort. dyck. 1844: 40. 1845
sec. Korotkova & al. (2010). – type: Pfeiffera cerei
formis Salm-dyck
= Rhipsalis subg. Acanthorhipsalis K. Schum., Gesamtbeschr. Kakt.: 615. 1898 ≡ Acanthorhipsalis (K.
Schum.) Britton & rose, cactaceae 4: 211. 1923.
the circumscription of Pfeiffera has undergone several radical changes in the past, and until the early
1980s, it was treated as a monotypic genus with P.
ianthothele (Monv.) F. a. c. Weber. Kimnach (1983)
subsumed Pfeiffera under Rhipsalis, while Barthlott (1987), Barthlott & taylor (1995) and anderson
(2001, 2005) synonymized it with Lepismium. in
the molecular phylogeny of nyffeler (2002), P. ian
thothele unexpectedly grouped together with two traditional Lepismium species, and widely distant from
either Rhipsalis or Lepismium. Hunt (2006) broadened the concept of Pfeiffera to include nine species.
this circumscription of Pfeiffera was evaluated and
clarified by Korotkova & al. (2010), who rejected the
circumscription of Hunt (2006), which also included
the species now segregated as Lymanbensonia (see
there). recent annotated checklist by Barthlott &
taylor (1995, as Lepismium subg. Pfeiffera (Salmdyck) Barthlott).
Pilosocereus Byles & G. d. rowley in cact. Succ. J. Gr.
Brit. 19: 66. 1957 sec. Hunt (2006). – type: Piloso
cereus leucocephalus (Poselg.) Byles & G. d. rowley
= Pseudopilocereus Buxb. in Beitr. Biol. Pflanzen 44:
249. 1968.
recent monograph by zappi (1994); recent floristic
treatment by taylor & zappi (2004).
Polaskia Backeb. in Blätt. Sukkulentenk. 1: 4. 1949 sec.
Hunt (2006) ≡ Chichipia Backeb. in Liste cact. Jard.
Bot. Les cèdres 12. 1950, nom. illeg. – type: Pola
skia chichipe (rol.-Goss.) Backeb.
320
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
= Heliabravoa Backeb. in cact. Succ. J. Gr. Brit. 18:
23. 1956.
recent monograph by Gibson (1988b).
Praecereus Buxb. in Beitr. Biol. Pflanzen 44: 273. 1968
sec. Hunt (2006). – type: Praecereus smithianus
(Britton & rose) Buxb.
Pseudoacanthocereus F. ritter in Kakteen Südamerika
1: 47. 1979 sec. Hunt (2006). – type: Pseudoacan
thocereus brasiliensis (Britton & rose) F. ritter
Pseudomitrocereus Bravo & Buxb. in Bot. Stud. 12: 53.
1961 sec. arias & al. (2012). – type: Pseudomitro
cereus fulviceps (F. a. c. Weber ex K. Schum.) Bravo
& Buxb.
= Cephalocereus subg. Mitrocereus Backeb. in Blätt.
Kakteenf. 1938(6). 1938 ≡ Mitrocereus (Backeb.)
Backeb. in cactaceae (Berlin) 2: 77. 1942.
Monotypic; Pseudomitrocereus fulviceps was previously included in Pachycereus or Cephalocereus,
later elevated to generic rank as Mitrocereus (Backeberg 1942) and later Pseudomitrocereus (Bravo &
Buxbaum, in Buxbaum 1961). arias & al. (2003)
found P. fulviceps to be unrelated to Pachycereus and
instead as sister sister to a clade of Cephalocereus and
Neobuxbaumia. therefore, Pseudomitrocereus was
reinstated by arias & al. (2012). Pseudomitrocereus
is characterized by having distinct fertile stem parts,
flowers completely covered with trichomes, and
thick axial tissue (pericarpel and receptacle; Buxbaum 1961). its inclusion in Pachycereus was supported by non-informative attributes (e.g. growth
form), shared by other members of Pachycereinae
(or Echinocereinae sensu nyffeler & eggli 2010b).
However, it is part of the clade “Cephalocereus”
according to arias & al. (2003), composed by Ce
phalocerus, Neobuxbaumia, and Pseudomitrocer
eus. the species of this clade share the presence of
prismatic crystals in the epidermis, inner stamens
and nectarial chamber, while the fruit is dehiscent and the pulp is white (arias & terrazas 2006).
Mitrocereus was based on the name Pilocereus chry
somallus Lem. as the type species, but this name
represents another species included in the synonymy
of Pachycereus militaris (audot) d. r. Hunt. consequently, Buxbaum and Bravo (Buxbaum 1961) proposed the name Pseudomitrocereus, with P. fulviceps
as the nomenclatural type.
Pseudorhipsalis Britton & rose, cactaceae 4: 213. 1923
sec. Hunt (2006). – type: Pseudorhipsalis alata (Sw.)
Britton & rose
= Wittia K. Schum. in Monatsschr. Kakteenk. 13: 117.
1903, nom. illeg.
= Wittiocactus rauschert in taxon 31: 558. 1982.
= Disisorhipsalis doweld in Sukkulenty 4(1 – 2): 40.
2002.
recent monographs by Kimnach (1993) and Bauer
(2003).
Pterocactus K. Schum. in Monatsschr. Kakteenk. 7: 6.
1897 sec. Griffith & Porter (2009). – type: Pterocac
tus kuntzei K. Schum.
confirmed as monophyletic with maximal support by
Griffith & Porter (2009) and ritz & al. (2012). recent
monograph by Kiesling (1982).
Punotia d. r. Hunt in cactaceae Syst. init. 25: 26. 2011
sec. ritz & al. (2012). – type: Punotia lagopus (K.
Schum.) d. r. Hunt
the sole species of this recently segregated genus,
Punotia lagopus, was formerly placed in Austro
cylindropuntia, but was recovered as sister to the
remaining species of Austrocylindropuntia and Cu
mulopuntia by ritz & al. (2012). it differs from
Austrocylindropuntia in several characters, especially
its growth form as flat, extensive cushions.
Pygmaeocereus H. Johnson & Backeb. in natl. cact. &
Succ. Journ. 12: 86. 1957 sec. Hunt (2006). – type:
Pygmaeocereus bylesianus andreae & Backeberg
Suggested as synonym of Haageocereus by nyffeler
& eggli (2010b).
Quiabentia Britton & rose, cactaceae 4: 252. 1923 sec.
Hunt (2006). – type: Quiabentia zehntneri (Britton &
rose) Britton & rose
Rapicactus Buxb. & Oehme in cactaceae (Berlin) 1942:
24. 1942 sec. Hunt (2006). – type: Rapicactus sub
terraneus (Backeb.) Buxb. & Oehme
= Lodia Mosco & zanovello in Bradleya 18: 44. 2000.
traditionally included in Turbinicarpus (see there);
separated from it by Vázquez-Sánchez & al. (2013)
after Turbinicarpus was found to be polyphyletic by
them and previously also by Bárcenas & al. (2011).
recent monograph by Lüthy (2003).
Rauhocereus Backeb. in descr. cact. nov. 5. 1957 sec.
Hunt (2006). – type: Rauhocereus riosaniensis Backeb.
Rebutia K. Schum. in Monatsschr. Kakteenk. 5: 102.
1895 sec. ritz & al. (2007). – type: Rebutia minu
scula K. Schum.
the circumscription of Rebutia s.l. vs a suite of proposed segregates (including Aylostera, Digitorebutia,
Mediolobivia, Sulcorebutia and Weingartia) has been
the subject of continued debate in the past 30 years.
the wide circumscription (including these taxa) was
adopted by anderson & al. (2001) and Hunt (2006),
but not by anderson (2005), who recognized Sul
corebutia and Weingartia. the broad concept goes
back to the consensus Cactaceae classification as
summarized by Hunt & taylor (1986), and some participants of the discussions at that time even argued
that Rebutia sensu latissimo should be placed in the
synonymy of an even more expanded Echinopsis.
recent molecular phylogenetic studies showed,
however, that Rebutia does not belong in the Echi
nopsis clade (ritz & al. 2007; Mosti & al. 2011;
Schlumpberger & renner 2012), and that the genus
in this broad concept is an untenable polyphyletic assemblage, as first noted by Lendel & al. (2006). in
Willdenowia 45 – 2015
the molecular phylogeny of ritz & al. (2007), three
independent clades with taxa of Rebutia s.l. are
found, namely “Rebutia i” (including the segregates
Aylostera, Digitorebutia and Mediolobivia), “Rebu
tia ii” (conforming to Rebutia s.str.) and Weingartia
(incl. Cintia and Sulcorebutia). While Rebutia s.str. is
placed as sister to Browningia, Aylostera is placed in
a clade with Cereus and Stetsonia (ritz & al. 2007;
Mosti & al. 2011). therefore it appears reasonable to
abandon the concept of Rebutia s.l., to restrict Rebu
tia to the “true” rebutias, and to accept both Aylostera
as well as Weingartia as separate genera. Most of the
necessary new combinations have been published
for Aylostera (Monti & al. 2011) and Weingartia
(Hentzschel & augustin 2008).
Rhipsalidopsis Britton & rose, cactaceae 4: 209. 1923
sec. Korotkova & al. (2011). – type: Rhipsalidopsis
rosea (Lagerh.) Britton & rose
as explained under Hatiora, the inclusion of Rhipsa
lidopsis in Hatiora is not supported by recent molecular phylogenies. calvente & al. (2011) found
the two traditional Rhipsalidopsis species (R. gaert
neri (regel) Moran, R. rosea) are sister to Schlum
bergera, but with moderate support. Korotkova & al.
(2011), however, found Hatiora s.str., Rhipsalidopsis
and Schlumbergera to form a grade, and even though
support for this topology is also moderate, support
for the monophyly of the three genera is maximal:
therefore, Rhipsalidopsis (easter cacti) is best kept
separate from Schlumbergera (christmas cacti). recent annotated checklist by Barthlott & taylor (1995,
as Hatiora subg. Rhipsalidopsis (Britton & rose)
Barthlott).
Rhipsalis Gaertn. in Fruct. Sem. Pl. 1: 137. 1788, nom.
cons. sec. Korotkova & al. (2011). – type: Rhipsalis
cassutha Gaertn.
= Erythrorhipsalis a. Berger in Monatsschr. Kakteenk.
30: 4. 1920.
the circumscription of Rhipsalis – one of the oldest genera of the family – has changed repeatedly
over time, and often Hatiora, Lepismium and Pseu
dorhipsalis, all now accepted at generic rank, were
variously subsumed under Rhipsalis. the morphology-based circumscription of Rhipsalis by Barthlott & taylor (1995) has been entirely confirmed as
monophyletic with maximal support in the molecular
phylogenetic study of Korotkova & al. (2011); the
same result was shown by calvente & al. (2011b),
though with a less comprehensive sampling.
Rhipsalis is notable since R. baccifera (Sol.) Stearn
is the only species of the family that naturally occurs
outside the new World. recent annotated checklist
by Barthlott & taylor (1995).
Salmiopuntia Frič ex Guiggi, cactology 2 (Suppl.): 2.
2011 sec. Majure & al. (2012). – type: Salmiopuntia
salmiana (J. Parm. ex Pfeiff.) Guiggi
this monotypic genus has been found in a polytomy
321
with Brasiliopuntia + Tacinga and Opuntia s.str. (i.e.
the platyopuntioids) by Griffith & Porter (2009). the
study of Majure & al. (2012) confirmed that Salmi
opuntia is not part of Opuntia s.str.
Samaipaticereus cárdenas in cact. Succ. J. (Los angeles) 24: 141. 1952 sec. Hunt (2006). – type: Samai
paticereus corroanus cárdenas
Schlumbergera Lem. in ill. Hort. 5: 24. 1858 sec. Korotkova & al. (2011). – type: Schlumbergera epiphyl
loides Lem.
= Zygocactus K. Schum., Fl. Bras. 4: 223. 1890.
= Epiphyllanthus a. Berger in rep. (annual) Missouri
Bot. Gard. 16: 84. 1905.
Schlumbergera (christmas cacti) is one of the bestknown and one of the morphologically best-defined
Cactaceae genera, recognizable by its flattened stems
and bright pink zygomorphic flowers. its monophyly
was confirmed by the molecular phylogenetic analysis of calvente & al. (2010) and Korotkova & al.
(2011). recent annotated checklist by Barthlott &
taylor (1995).
Sclerocactus Britton & rose, cactaceae 3: 212. 1922
sec. Vázquez-Sánchez & al. (2013). – type: Sclero
cactus polyancistrus (engelm. & J. M. Bigelow) Britton & rose
= Echinomastus Britton & rose, cactaceae 3: 147.
1922.
= Toumeya Britton & rose, cactaceae 3: 91. 1922,
nom. illeg.
= Ancistrocactus Britton & rose, cactaceae 4: 3. 1923.
= Coloradoa Boissev. & c. davidson in colorado cact.
54. 1941.
confirmed as monophyletic by Butterworth & al.
(2002) and Vázquez-Sánchez & al. (2013). the generic status and limits of Echinomastus need further
evaluation because it was found to be polyphyletic
by Vázquez-Sánchez & al. (2013). revisions/monographs by Heil & Porter (1994) and Hochstätter
(2005).
Selenicereus (a. Berger) Britton & rose in contr. U. S.
natl. Herb. 12: 429. 1909 sec. Hunt (2006) ≡ Cereus
subsect. Selenicereus a. Berger in rep. (annual)
Missouri Bot. Gard. 16: 76. 1905. – type: Seleni
cereus grandiflorus (L.) Britton & rose
= Cryptocereus alexander in cact. Succ. J. (Los angeles) 22: 164. 1950.
= Chiapasophyllum doweld in Sukkulenty 4(1 – 2): 32.
2002.
Stenocactus (K. Schum.) a. W. Hill in index Kew. Suppl.
8: 228. 1933 sec. Hunt (2006) ≡ Echinocactus subg.
Stenocactus K. Schum., Gesamtbeschr. Kakt.: 292.
1898. – type: Stenocactus coptonogonus (Lem.) a.
W. Hill ex a. Berger
See notes under Ferocactus.
Stenocereus (a. Berger) riccob. in Boll. reale Orto Bot.
Palermo 8: 253. 1909, nom. cons. sec. Hunt (2006) ≡
Cereus subg. Stenocereus a. Berger in rep. (annual)
322
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
Missouri Bot. Gard. 16: 66. 1905. – type: Steno
cereus stellatus (Pfeiff.) riccob.
= Rathbunia Britton & rose in contr. U. S. natl. Herb.
12: 414. 1909.
= Machaerocereus Britton & rose, cactaceae 2: 114.
1920.
= Lemaireocereus subg. Isolatocereus Backeb. in Blätt.
Kakteenf. 1938(6): 17. 1938 ≡ Isolatocereus Backeb.
in cactaceae (Berlin) 1941(2): 47, 76. 1942.
= Ritterocereus Backeb. in Jahrb. deutsch. KakteenGes. 1941: 76. 1942.
= Hertrichocereus Backeb. in cact. Succ. J. (Los angeles) 22: 153. 1950.
= Marshallocereus Backeb. in cact. Succ. J. (Los angeles) 22: 154. 1950.
= Griseocactus Guiggi in cactology 3(Suppl.): 1. 2012.
= Griseocereus Guiggi in cactology 3: 7. 2012, nom.
inval.
recent treatments by Gibson (1991) and arreola-nava & terrazas (2003).
Stephanocereus a. Berger in entwicklungslin. Kakt.: 97.
1926 sec. Hunt (2006). – type: Stephanocereus leu
costele (Gürke) a. Berger
= Coleocephalocereus subg. Lagenopsis Buxb. in
Krainz, Kakteen: 48 – 49, ciVb. 1972 ≡ Pilosocereus
subg. Lagenopsis (Buxb.) Braun in Bradleya 6: 89.
1988 ≡ Stephanocereus subg. Lagenopsis n. P. taylor & eggli in Bradleya 9: 91. 1991 ≡ Lagenocereus
doweld in turczaninowia 5: 8. 2002.
recent floristic treatment by taylor & zappi (2004).
Stetsonia Britton & rose, cactaceae 2: 64. 1920 sec.
Hunt (2006). – type: Stetsonia coryne (Salm-dyck)
Britton & rose
Strombocactus Britton & rose, cactaceae 3: 106. 1922
sec. Vázquez-Sánchez & al. (2013). – type: Strombo
cactus disciformis (dc.) Britton & rose
confirmed as monophyletic by Vázquez-Sánchez &
al. (2013).
Strophocactus Britton & rose in contr. U. S. natl. Herb.
16: 262. 1913 sec. Hunt (2006). – type: Strophocac
tus wittii (K. Schum.) Britton & rose
= Deamia Britton & rose, cactaceae 2: 212. 1920.
Tacinga Britton & rose, cactaceae 1: 39. 1919 sec. Hunt
(2006). – type: Tacinga funalis Britton & rose
recent floristic monograph by taylor & zappi
(2004).
Tephrocactus Lem. in cactées: 88. 1868 sec. ritz & al.
(2012). – type: Tephrocactus diademata (Lem.) Lem.
= Ursopuntia P. V. Heath, calyx 6(2): 41. 1999.
= Quasitephrocactus G. Popov, Kakt. Klub 15(1 – 2):
13, 2012, nom. illeg.
Tephrocactus was confirmed as monophyletic by ritz
& al. (2012). recent monographs by Kiesling (1984)
and Gilmer & thomas (1998).
Thelocactus (K. Schum.) Britton & rose in Bull. torrey
Bot. club 49: 251. 1922 sec. Hunt (2006) ≡ Echino
cactus subg. Thelocactus K. Schum., Gesamtbeschr.
Kakt.: 429. 1898. – type: Thelocactus hexaedropho
rus (Lem.) Britton & rose
= Hamatocactus Britton & rose, cactaceae 3: 104.
1922.
= Torreyocactus doweld in Sukkulenty 1998(1): 19.
1998.
See notes under Ferocactus.
Tunilla d. r. Hunt & iliff in cactaceae Syst. init. 9: 10.
2000 sec. Hunt (2006). – type: Tunilla soehrensii
(Britton & rose) d. r. Hunt & iliff
Turbinicarpus Buxb. & Backeb. in cactaceae (Berlin) 1937(1): 27. 1937 sec. Vázquez-Sánchez & al.
(2013). – type: Turbinicarpus schmiedickeanus
(Boed.) Buxb. & Backeb.
= Gymnocactus Backeb. in Blätt. Kakteenf. 1938(6):
[22]. 1938.
= Normanbokea Kladiwa & Buxb. in Krainz, Kakteen
40: 40, c Viiib. 1969.
= Bravocactus doweld in Sukkulenty 1998(1): 22. 1998.
= Kadenicarpus doweld in Sukkulenty 1998(1): 22.
1998.
Turbinicarpus has been found to be polyphyletic in
the molecular studies of Bárcenas & al. (2011) and
Hernández-Hernández & al. (2011). the most comprehensively sampled dataset of Vázquez-Sánchez
& al. (2013) showed Turbinicarpus to fall into three
separate clades. Turbinicarpus was re-circumscribed
restricted to 11 species, while species with a tuberous
root connected to the body with a long, thin neck are
now segregated as Rapicactus based on these results.
two further species (T. horripilus (Lem.) V. John &
Říha and T. pseudomacrochele (Backeb.) Buxb. &
Backeb.) are outside the main Turbinicarpus clade
(incl. Gymnocactus) and a new generic name would
be needed for them. recent treatments by Lüthy
(2002) and Lüthy & Moser (2002).
Uebelmannia Buining in Succulenta (netherlands) 46:
159. 1967 sec. Hunt (2006). – type: Uebelmannia
gummifera (Backeb. & Voll) Buining
recent works by nyffeler (1998), Lüthy & Moser
(2002), and taylor & zappi (2004).
Vatricania Backeb. in cact. Succ. J. (Los angeles) 22:
154. 1950 sec. Schlumpberger & renner (2012). –
type: Vatricania guentheri (Kupper) Backeb.
included in Espostoa s.l. by modern lexicographic
treatments such as anderson (2001, 2005) and Hunt
(2006), the genus was found to be distant from the Es
postoa in the Cleistocactus s.str. clade by Schlumpberger & renner (2012). consequently, the monotypic Vatricania was suggested to be reinstated.
Weberbauerocereus Backeb. in cactaceae (Berlin)
1941(2): 31, 75. 1942 sec. Hunt (2006). – type:
Weberbauerocereus fascicularis (Meyen) Backeb.
recent monograph by arakaki (2003).
Weberocereus Britton & rose in contr. U. S. natl. Herb.
12: 431. 1909 sec. Hunt (2006). – type: Webero
cereus tunilla (F. a. c. Weber) Britton & rose
Willdenowia 45 – 2015
= Werckleocereus Britton & rose in contr. U. S. natl.
Herb. 12: 432. 1909.
= Eccremocactus Britton & rose in contr. U. S. natl.
Herb. 16: 261. 1913.
Weingartia Werderm. in Kakteenkunde 1937: 20, 21.
1937 sec. ritz & al. (2007). – type: Weingartia fidai
ana (Backeb.) Werderm.
= Sulcorebutia Backeb. in cact. Succ. J. Gr. Brit. 13:
96. 1951.
= Cintia Kníže & Říha in Kaktusy (Brno) 31: 37. 1995.
= Gymnorebutia doweld in Sukkulenty 4(1 – 2): 24.
2002.
Weingartia and Sulcorebutia used to be merged in
Rebutia, e.g. by Barthlott & Hunt (1993), anderson
(2001), and Hunt (2006), but were recognized by
anderson (2005). the Rebutia s.l. assemblage was
found highly polyphyletic by ritz & al. (2007), and
was shown to be separated into three well-supported clades. One of these clades comprises species of
Cintia, Sulcorebutia and Weingartia and includes the
nomenclatural type of Weingartia. ritz & al. (2007)
suggested that all three could be merged into a single
genus, for which Weingartia is the oldest name.
Yavia r. Kiesling & Piltz in Kakteen and. Sukk. 52(3):
57. 2001 sec. Hunt (2006). – type: Yavia cryptocarpa
r. Kiesling & Piltz
Yungasocereus F. ritter in Kakteen Südamerika 2: 668.
1980 sec. Hunt (2006). – type: Yungasocereus inqui
sivensis (cárdenas) F. ritter
Caryophyllaceae Juss. sec. aPG (2009).
a family of chiefly opposite-leaved herbs comprising
about 100 genera and 3000 species. the family is widely distributed in north-temperate, montane and alpine
areas with a centre of diversity in the eastern Mediterranean and irano-turanean regions, while presence in
the tropics and the southern hemisphere is limited and
mostly at higher elevations (Bittrich 1993c; rabeler &
Hartman 2005a). Several taxa (especially species of
Dianthus, Gypsophila and Silene) are important in the
horticultural trade, while others (e.g. Stellaria media
(L.) Vill.) have become widely known weedy taxa. the
number of genera included here is over 10% higher than
most recent estimates (Bittrich 1993c; rabeler & Hartman 2005a; Harbaugh & al. 2010), reflecting the results
of recent molecular studies on large genera (especially
Minuartia; dillenberger & Kadereit 2014) as well as
retention of several genera (e.g. Myosoton, Velezia and
Xerotia) that may eventually disappear. the family is
monophyletic as circumscribed by Bittrich (1993c), although the “traditional” division into three subfamilies
(Bittrich 1993c; Pax & Hoffmann 1934) based on stipule, petal, sepal and fruit features does not provide
monophyletic groups and should be replaced with the
tribe-based scheme presented by Harbaugh & al. (2010)
and confirmed by subsequent studies (e.g. Greenberg &
donoghue 2011).
323
Acanthophyllum c. a. Mey. in Verz. Pfl. casp. Meer.:
210. 1831 sec. Pirani & al. (2014). – type: Acantho
phyllum mucronatum c. a. Mey.
= Ochotonophila Gilli in repert. Spec. nov. regni Veg.
59: 169. 1956.
= Kuhitangia Ovcz. in dokl. akad. nauk tadzh. SSr
10: 50. 1967.
= Scleranthopsis rech. f. in ann. naturhist. Mus. Wien
70: 37. 1967.
consists of about 60 cushion-forming subshrubby
species of the subalpine steppe region in central to
southwestern asia (Bittrich 1993b; Ghaffari 2004).
Pirani & al. (2014) showed that the genus is paraphyletic in this circumscription with Allochrusa, Di
aphanoptera p.p., Ochotonophila and Scleranthopsis
nested within it.
Achyronychia torr. & a. Gray in Proc. amer. acad. arts
7: 330. 1868 sec. Bittrich (1993c). – type: Achyrony
chia cooperi torr. & a. Gray
Monotypic genus; southwestern United States and
Mexico. Hartman (2005a) noted that seed and flower
characters suggest a close relationship to Scopulo
phila. Greenberg & donoghue (2011) showed a similar result from molecular data.
Agrostemma L., Sp. Pl. 1: 435. 1753 sec. Oxelman & al.
(2001) ≡ Githago adans., Fam. Pl. 2: 255. 1763. –
type: Agrostemma githago L.
two to three species, probably native in the Mediterranean region, but widely spread as agricultural weeds
and/or ornamentals. Several phylogenetic studies
(Oxelman & Lidén 1995; Oxelman & al. 1997; Fior
& al. 2006; Greenberg & donoghue 2011) strongly
support Agrostemma as a sister group to the rest of
the tribe Sileneae.
Allochrusa Bunge ex Boiss., Fl. Orient. 1: 559. 1867
sec. Bittrich (1993c). – type: Allochrusa versicolor
(Fisch. & c. a. Mey.) Boiss.
comprises seven species from southwestern asia that
are probably nested in Acanthophyllum (Pirani & al.
2014).
Ankyropetalum Fenzl in Bot. zeitung (Berlin) 1: 393.
1843 sec. Bittrich (1993c). – type: Ankyropetalum
gypsophiloides Fenzl
Four species in the eastern Mediterranean region east
to armenia. closely related to Gypsophila, but not yet
sampled for dna.
Arenaria L., Sp. Pl. 1: 423. 1753 sec. Sadeghian & al.
(2015). – type: Arenaria serpyllifolia L. – Fig. 3e.
= Spergulastrum Michx., Fl. Bor.-amer. 1: 275. 1803.
= Cernohorskya á. Löve & d. Löve in Preslia 46: 127.
1974.
= Willwebera á. Löve & d. Löve in Lagascalia 4: 9.
1974.
about 160 species, in north-temperate areas, the Mediterranean, and andean South america. Harbaugh &
al. (2010), Greenberg & donoghue (2011) and most
recently Sadeghian & al. (2015) have sampled Are
324
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
naria and, between their results, have removed about
one-half of the species into four segregate genera not
aligning in the same tribe as Arenaria. Sadeghian &
al. (2015) found that four of the five remaining subgenera that Mcneill (1962) recognized form Are
naria s.str., with the placement of A. subg. Dicranilla
(Fenzl) F. n. Williams still unknown. While a few
of the infrageneric groups recognized by Mcneill
(1962) are confirmed by molecular results (e.g. A.
subg. Leiosperma Mcneill, A. sect. Plinthine (rchb.)
Mcneill), most are not.
Atocion adans. in Fam. Pl. (adanson) 2: 254. 1763 sec.
Oxelman & al. (2001). – type: Atocion armeria (L.)
raf.
= Minjaevia tzvelev in novosti Sist. Vyssh. rast. 33:
102. 2001.
recently revised by Frajman & al. (2013), who recognized six, mostly european species. Well supported
as monophyletic by several unlinked dna sequence
regions, and also as sister to Viscaria (Frajman & al.
2009b; see under Viscaria).
Bolanthus (Ser.) rchb., deut. Bot. Herb.-Buch: 205.
1841 sec. Bittrich (1993c) ≡ Saponaria sect. Bolan
thus Ser. in candolle, Prodr. 1: 366. 1824. – type:
Bolanthus hirsutus (Labill.) Barkoudah – Fig. 3F.
about 15 species in the eastern Mediterranean region,
especially Greece and turkey (Koç & Hamzaoğlu
2015). closely related to Acanthophyllum and Gyp
sophila, but not yet sampled for dna.
Brachystemma d. don, Prodr. Fl. nepal.: 216. 1825 sec.
Bittrich (1993c). – type: Brachystemma calycinum
d. don
Monotypic; Himalayas, Se asia. Likely near Arena
ria and Moehringia; one rbcL sequence exists (SaslisLagoudakis & al. 2012), but has not been included in
a phylogeny including these genera.
Bufonia L., Sp. Pl. 1: 123. 1753 sec. Bittrich (1993c). –
type: Bufonia tenuifolia L.
about 20 species in the Mediterranean region. Greenberg & donoghue (2011) showed Bufonia as sister to
the remainder of Sagineae (except for Drypis), while
dillenberger & Kadereit (2014) found it was an unsupported sister to a clade containing Minuartia s.str.
and Mcneillia.
Calycotropis turcz. in Bull. Soc. imp. naturalistes Moscou 35: 327. 1862 sec. Bittrich (1993c). – type: Caly
cotropis minuartioides turcz.
Monotypic; Mexico. Listed as a “doubtful genus” in
the Caryophyllaceae by Bittrich (1993c).
Cardionema dc., Prodr. 3: 372. 1828 sec. Bittrich
(1993c). – type: Cardionema multicaule dc.
Six species found from western north america south
to chile. Sosa & al. (2006) found Cardionema and
Scopulophila clustered with Cerdia. Greenberg &
donoghue (2011) showed Cardionema belonging to
a poorly resolved group of genera in the tribe Poly
carpaeae.
Cerastium L., Sp. Pl. 1: 437. 1753 sec. Greenberg &
donoghue (2011). – type: Cerastium arvense L.
includes 100 or, more likely, close to 200 northtemperate species, especially diverse in the eastern
Mediterranean. the genus is in need of monographic
study. the most recent infrageneric classification is
presented by Schischkin (1936); even with corrected
nomenclature and inclusion of extra-russian taxa, it
is not likely to be representative of relationships in
the genus. Greenberg & donoghue (2011) included
39 species of Cerastium in their study and found several interesting points. Cerastium subg. Dichodon
(Bartl. ex rchb.) Boiss. should be treated as a genus,
Dichodon (see there), being a sister to Holosteum. as
in Dianthus, resolution of the species was very poor,
most species falling into either a polytomy of 11 species or one of 23. they also found Cerastium formed
a clade within Stellaria. these genera are considered
quite distinct by nearly all workers, so this must be
investigated further.
Cerdia Moc. & Sessé ex dc., Prodr. 3: 377. 1828 sec.
Sosa & al. (2006). – type: not designated.
Monotypic; endemic to Mexico. Placement within
the Polycarpeae is probable (near Cardionema and
Scopulophila?), but Sosa & al. (2006) suggested that
further study is needed. in a broader survey using a
different voucher, Greenberg & donoghue (2011)
found Cerdia clustering near Drymaria.
Chaetonychia (dc.) Sweet in Hort. Brit., ed. 3: 263.
1839 sec. Bittrich (1993c) ≡ Paronychia sect. Chae
tonychia dc., Prodr. 3: 370. 1828. – type: Chaetony
chia cymosa (L.) Sweet
Monotypic; western Mediterranean. Probably a close
relative of Paronychia, but as yet not sampled for molecular phylogenetic analysis.
Cherleria L., Sp. Pl. 1: 425. 1753 sec. dillenberger &
Kadereit (2014). – type: Cherleria sedoides L.
= Wierzbickia rchb., deut. Bot. Herb.-Buch: 205; Syn.
red.: 106. 1841.
= Lidia á. Löve & d. Löve in Bot. not. 128: 510.
1976.
Originally including only C. sedoides found in mountains of europe, but dillenberger & Kadereit (2014)
proposed expanding it to 19 species of eurasia and
western north america; Mosyakin suggests 23 to account for some additional eastern european taxa not
yet transferred to Cherleria (S. Mosyakin, unpubl.
data). Formerly included (with Pseudocherleria)
in Minuartia sect. Spectabiles (Fenzl) Hayek, dillenberger & Kadereit (2014) found the two groups
segregated into different clades far from Minuartia
s.str., proposing the recognition of both Cherleria and
Pseudocherleria.
Colobanthus Bartl., Ord. nat. Pl.: 305. 1830 sec. Bittrich (1993c). – type: Colobanthus quitensis (Kunth)
Barthlott
comprises 20 species of cushion plants most diverse
Willdenowia 45 – 2015
in the southern hemisphere. the genus is monophyletic and a sister to Sagina.
Cometes L., Syst. nat., ed. 12 (2): [109, 127]. 1767 sec.
Bittrich (1993c). – type: Cometes surattensis L.
two species; deserts from nW india to ne africa.
Likely a member of Polycarpaeae, but the one available itS sequence has not been included in a broader
survey.
Corrigiola L., Sp. Pl. 1: 271. 1753 sec. Bittrich (1993c).
– type: Corrigiola litoralis L.
about 11 species. Harbaugh & al. (2010) and Greenberg & donoghue (2011) both confirmed placement
(with Telephium) in tribe Corrigioleae, near the base
of Caryophyllaceae.
Cyathophylla Bocquet & Strid, Mount. Fl. Greece 1:
175. 1986 sec. Bittrich (1993c). – type: Cyathophylla
chlorifolia (Poir.) Bocquet & Strid
Monotypic; mountains of Greece and turkey. closely
related to Saponaria, but not yet sampled for dna.
Dadjoua Parsa, Fl. iran 8: 248. 1960 sec. Bittrich (1993c).
– type: Dadjoua pteranthoidea Parsa
Monotypic; iran. Listed as a “doubtful genus” in the
Caryophyllaceae by Bittrich (1993c).
Dianthus L., Sp. Pl. 1: 409. 1753 sec. Bittrich (1993c). –
type: Dianthus caryophyllus L. – Fig. 4a.
With about 300 species, Dianthus is the second largest genus in the Caryophyllaceae. Dianthus is most
diverse in southeastern europe and southwestern
asia. no recent monographic work has been undertaken; the most comprehensive infrageneric classification is presented in Pax & Hoffmann (1934).
although Greenberg & donoghue (2011) included 37
species in their analysis, virtually no resolution was
found; 26 species formed a polytomy. May include
Velezia (see there).
Diaphanoptera rech. f. in repert. Spec. nov. regni Veg.
48: 41. 1940 sec. Bittrich (1993c). – type: Diaphano
ptera khorasanica rech. f.
a genus of six species according to Schiman-czeika
(1988), but recent molecular phylogenetic analyses indicate polyphyly, with some species nested in
Acanthophyllum (Pirani & al. 2014).
Dicheranthus Webb in ann. Sci. nat., Bot., ser. 3, 5: 28.
1846 sec. Bittrich (1993c). – type: Dicheranthus plo
camoides Webb
Monotypic; canary islands. a member of the Poly
carpaeae, clustering with Pteranthus (Greenberg &
donoghue 2011).
Dichodon (Bartl. ex rchb.) rchb., deut. Bot. Herb.-Buch:
205. 1841 sec. ikonnikov (1973) ≡ Stellaria [unranked] Dichodon Bartl. ex rchb., Fl. Germ. excurs.
24: 785. 1832. – type: Dichodon dubium (Bastard)
ikonn.
= Provancheria B. Boivin in naturaliste canad. 93:
644. 1967.
Five species of the arctic, central europe, and iran.
treated as Cerastium subg. Dichodon (Bartl. ex
325
rchb.) Boiss. in most recent works. Greenberg &
donoghue (2011) found that the two sampled species of Dichodon formed a clade sister to Holosteum,
and together formed a clade sister to Cerastium +
Moenchia.
Dolophragma Fenzl, ann. Wiener Mus. naturgesch. 1:
63. 1836 sec. Sadeghian & al. (2015). – type: Dolo
phragma globiflorum Fenzl
a genus of four or five Himalayan species. Most recently treated as a subgenus of Arenaria (Mcneill
1962). Sadeghian & al. (2015) suggested the genus
be again recognized after finding that the one sampled species clustered near Eremogone, either as a
sister to Silene or between Eremogone and Silene.
they also noted that the result reported by Greenberg
& donoghue (2011), showing Arenaria przewalskii
Maxim. clustering with members of Lepyrodiclis and
Pseudostellaria, suggests that Dolophragma may be
polyphyletic.
Drymaria Willd. ex Schult., Syst. Veg. ed. 15bis 5: 31,
406. 1819 sec. Bittrich (1993c). – type: Drymaria
arenarioides Humb. & Bonpl. ex Schult.
= Pinosia Urb. in ark. Bot. 23a(5): 70. 1930.
about 50 species, all but two found only in the new
World. Little is known about relationships within
Drymaria. duke’s (1962) preliminary revision,
in which he described but did not validly publish
17 series, is the only recent comprehensive study.
Greenberg & donoghue (2011) included all four
sampled taxa and show a poorly resolved, possibly
polyphyletic genus.
Drypis L., Sp. Pl. 1: 413. 1753 sec. Bittrich (1993c). –
type: Drypis spinosa L.
Monotypic; eastern Mediterranean. Formerly placed
in an isolated position within the Caryophylloideae.
Molecular studies, including Harbaugh & al. (2010),
Greenberg & donoghue (2011) and dillenberger &
Kadereit (2014), place Drypis as sister to all other
sampled taxa in tribe Sagineae.
Eremogone Fenzl in Vers. darstell. alsin.: 13. 1833
sec. rabeler & Wagner (2015). – type: Eremogone
graminifolia Fenzl
= Brewerina a. Gray in Proc. amer. acad. arts 8: 620.
1873.
about 90 species, most diverse in eastern asia and
western north america. Harbaugh & al. (2010) confirmed the wide separation from Arenaria that Fior &
al. (2006) reported. Broad sampling is still needed to
resolve infrageneric relationships; existing information (Sadeghian & al. 2015) is not consistent with the
extant classification (Mcneill 1962) erected for these
taxa in two subgenera under Arenaria.
Eudianthe (rchb.) rchb., deut. Bot. Herb.-Buch: 206.
1841 sec. Oxelman & al. (2001) ≡ Lychnis [unranked]
Eudianthe rchb., Fl. Germ. excurs. 24: 824. 1832 ≡
Pontinia Fries in Bot. not. (1843): 141. 1843. – type:
Eudianthe coelirosa (L.) Fenzl ex endl.
326
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
two western Mediterranean annual species, well supported as not belonging to the core Silene/Lychnis
group (Oxelman & Lidén 1995; Oxelman & al. 1997;
Oxelman & al. 2001).
Facchinia rchb., deut. Bot. Herb.-Buch: 204; Syn. red.:
63. 1841 sec. dillenberger & Kadereit (2014). – type:
Facchinia lanceolata (all.) rchb.
Five species found in high mountains of europe. dillenberger & Kadereit (2014) found Facchinia to be
both distant from Minuartia s.str. and a sister to the
clade containing Colobanthus and Sagina.
Gymnocarpos Forssk., Fl. aegypt.-arab.: 65. 1775 sec.
Oxelman & al. (2002). – type: Gymnocarpos decan
drus Forssk.
= Sclerocephalus Boiss., diagn. Pl. Orient., ser. 1, 3:
12. 1843.
= Lochia Balf. f. in Proc. roy. Soc. edinb. 12: 409.
1884.
ten species, occurring from the canary islands east
to Mongolia. Oxelman & al. (2002) found an expanded Gymnocarpos was monophyletic and sister to part
of Paronychia.
Gypsophila L., Sp. Pl. 1: 406. 1753 sec. Bittrich (1993c).
– type: Gypsophila repens L.
= Bolbosaponaria Bondarenko in Opred. rast. Sred.
azii 2: 327. 1971.
= Pseudosaponaria (F. n. Williams) ikonn. in novosti
Sist. Vyssh. rast. 15: 144. 1979.
Gypsophila includes about 150 species and is especially diverse in the eastern Mediterranean and southwestern asia. Most of the infrageneric classification
is derived from Barkoudah’s (1962) monograph of
Gypsophila and three related genera. Greenberg &
donoghue (2011) included 24 species in their analysis and found Gypsophila to be polyphyletic, with
most species forming a clade sister to Saponaria and
four species resolving close to Dianthus/Petrorhagia;
one of these species, G. muralis L., is here treated
as Psammophiliella. Pirani & al. (2014) found G.
cerastioides d. don nested within Acanthophyllum.
recognition of Bolbosaponaria seems likely; while
Greenberg & donoghue (2011) found B. bucharica
(B. Fedtsch.) Bondarenko clustered with two other
species of Gypsophila, Pirani & al. (2014) found it
to be a sister taxon to Diaphanoptera afghanica Podl.
Habrosia Fenzl in Bot. zeitung (Berlin) 1: 322. 1843 sec.
Bittrich (1993c). – type: Habrosia spinuliflora (Ser.)
Fenzl
Monotypic; southwestern asia. Smissen & al. (2003)
placed Habrosia as sister to Drypis. Greenberg &
donoghue (2011), citing the itS voucher from the
Smissen & al. (2003) study, reported H. spinuliflora
nested in Minuartia (Sabulina sec. dillenberger &
Kadereit 2014), sister to a clade of five north american species.
Haya Balf. f. in Proc. roy. Soc. edinb. 12: 408. 1884 sec.
Bittrich (1993c). – type: Haya obovata Balf. f.
Monotypic; Socotra island. Kool & al. (2007, 2012)
found H. obovata nested in a clade of Polycarpaea;
the genus is retained pending additional resolution of
the polyphyletic Polycarpaea.
Heliosperma (rchb.) rchb., deut. Bot. Herb.-Buch: 206.
1841, nom. cons. sec. Frajman & rabeler (2006) ≡
Silene [unranked] Heliosperma rchb., Fl. Germ.
excurs. 24: 817. 1832 ≡ Ixoca raf., autik. Bot.: 25.
1840. – type: Silene quadrifida (L.) L.
a chiefly central and southeastern european group
with four to 16 species depending on species delimitations (Frajman & Oxelman 2007). Heliosperma has
been conserved over its senior synonym Ixoca (Barrie 2011), as proposed by Frajman & rabeler (2006).
Frajman & al. (2009a) analysed several independent
nuclear and plastid loci showing strong support for
monophyly of the genus, although it appears to have
a complex history, possibly involving ancient hybridization events.
Herniaria L., Sp. Pl. 1: 218. 1753 sec. Bittrich (1993c).
– type: Herniaria glabra L.
= Heterochiton Graebn. & Mattf. in Syn. Mitteleur. Fl.
5: 870. 1919.
about 50 species, most of them narrowly distributed
endemics. Herniaria remains largely unsampled for
dna (four species in Greenberg & donoghue 2011)
and is likely to be closely related to Paronychia subg.
Anoplonychia (Fenzl) rchb.; see Oxelman & al.
(2002) and Greenberg & donoghue (2011).
Holosteum L., Sp. Pl. 1: 88. 1753 sec. Bittrich (1993c). –
type: Holosteum umbellatum L.
three to four species of temperate eurasia. While
Harbaugh & al. (2010) found that Holosteum and
Moenchia were sister taxa, Greenberg & donoghue
(2011) found Holosteum and Dichodon to be sisters,
with that clade a sister to the clade that include Cera
stium and Moenchia.
Honckenya ehrh. in neues Mag. aerzte 5: 206. 1783
sec. Bittrich (1993c). – type: Honckenya peploides
(L.) ehrh.
Monotypic; circumpolar in sandy coastal areas. Harbaugh & al. (2010) found Honckenya and Wilhelmsia
are sister to each other and both are the closest relatives to the Hawaiian Schiedea.
Illecebrum L., Sp. Pl. 1: 206. 1753 sec. Bittrich (1993c).
– type: Illecebrum verticillatum L.
Monotypic; native to the canary islands and the Mediterranean. Greenberg & donoghue (2011) showed
Illecebrum belonging to a poorly resolved group of
genera in the Polycarpaeae, closest to Cardionema as
shown by Kool & al. (2007).
Kabulia Bor & c. e. c. Fisch. in indian Forester 65: 611.
1939 sec. Bittrich (1993c). – type: Kabulia akhtarii
Bor & c. e. c. Fisch.
Monotypic; afghanistan. there is no new information to contradict Bittrich’s (1993) placement as incertae sedis in the Paronychioideae (?Paronychieae).
Willdenowia 45 – 2015
Krauseola Pax & K. Hoffm., nat. Pflanzenfam. (ed. 2)
16c: 308. 1934 sec. Bittrich (1993c). – type: Krau
seola mosambicina (Moss) Pax & Hoffm.
two species from tropical east africa. Likely included in the Polycarpaeae, but not yet sampled for dna.
Lepyrodiclis Fenzl in endlicher, Gen. Pl.: 966. 1840 sec.
Bittrich (1993c). – type: Lepyrodiclis holosteoides
(c. a. Mey.) Fisch. & c. a. Mey.
three species of central asia. Sadeghian & al.
(2015) found two species formed a clade sister to
one including Odontostemma and Pseudostellaria.
Greenberg & donoghue (2011) noted that L. ho
losteoides clustered with Stellaria monosperma
Buch.-Ham. ex d. don.
Loeflingia L., Sp. Pl. 1: 35. 1753 sec. Bittrich (1993c). –
type: Loeflingia hispanica L.
Seven species of the Mediterranean, southwestern
asia, and western north america. Fior & al. (2006)
and Harbaugh & al. (2010) both showed Loeflingia
and Polycarpon clustering together; a result not
shown in the Kool & al. (2007) study of Polycarpon.
Greenberg & donoghue (2011) found it clustered in
a poorly resolved clade including eleven other genera
of Polycarpaeae.
Lychnis L., Sp. Pl. 1: 436. 1753 sec. Oxelman & al.
(2001). – type: Lychnis chalcedonica L.
= Coronaria Guett. in Hist. acad. roy. Sci. Mém.
Math. Phys. (Paris 4to) 1750: 229. 1754.
= Hedona Lour., Fl. cochinch. 1: 286. 1790.
= Exemix raf., autik. Bot.: 27. 1840.
= Coccyganthe (rchb.) rchb., deut. Bot. Herb.-Buch:
206. 1841 ≡ Lychnis [unranked] Coccyganthe rchb.,
Fl. Germ. excurs. 24: 825. 1832.
= Uebelinia Hochst. in Flora 24: 664. 1841.
this circumscription, including around twenty species, is strongly supported as monophyletic (e.g. Popp
& al. 2008; Greenberg & donoghue 2011), with the
african Uebelinia nested within. However, its relationships to Silene are not fully resolved (see under
Silene).
Mcneillia dillenb. & Kadereit, taxon 63: 78. 2014 sec.
dillenberger & Kadereit (2014). – type: Mcneillia
graminifolia (ard.) dillenb. & Kadereit
Five species of southeastern europe and turkey. treated as Minuartia [sect. Lanceolatae (Fenzl) Graebn.]
ser. Graminifoliae Mattf. by Mcneill (1962), dillenberger & Kadereit (2014) found these taxa forming a
clade sister to Minuartia s.str.
Microphyes Phil., Fl. atacam.: 20, t. 1. 1860 sec. Bittrich
(1993c). – type: Microphyes litoralis Phil.
= Wangerinia c. Franz in Bot. Jahrb. Syst. 42(2 – 3, Beibl. 97): 11. 1908.
three species in chile. traditionally placed in Poly
carpaeae, but not yet sampled for dna.
Minuartia L., Sp. Pl. 1: 89. 1753 sec. dillenberger &
Kadereit (2014). – type: Minuartia dichotoma L.
= ?Queria L., Sp. Pl. 1: 90. 1753.
327
= Alsinanthe (Fenzl) rchb., deut. Bot. Herb.-Buch:
205. 1841.
= Tryphane (Fenzl) rchb., deut. Bot. Herb.-Buch: 205.
1841 ≡ Alsine [unranked] Tryphane Fenzl in endlicher, Gen. Pl.: 965. 1840.
= Alsinopsis Small, Fl. S. e. U. S.: 419. 1903.
= Lidia á. Löve & d. Löve in Bot. not. 128: 510. 1976.
= Minuopsis W. a. Weber in Phytologia 58(6): 383.
1985.
about 54 species, chiefly in Mediterranean europe and
eastward into south-central asia. While several molecular studies had shown Minuartia to be polyphyletic,
dillenberger & Kadereit’s (2014) study is the most
comprehensive to date, including the first sequences
for Minuartia sect. Minuartia. they found that the 96
species of Minuartia sampled belonged to ten different clades representing four different tribes. this circumscription restricts Minuartia to two of the twelve
sections of Minuartia subg. Minuartia recognized by
Mcneill (1962).
Minuartiella dillenb. & Kadereit, taxon 63: 78. 2014
sec. dillenberger & Kadereit (2014). – type: Minu
artiella acuminata (turill) dillenb. & Kadereit
Four species of the mountains of turkey and iran.
treated as Minuartia [sect. Lanceolatae (Fenzl)
Graebn.] ser. Dianthifoliae Mattf. by Mcneill (1962),
dillenberger & Kadereit (2014) found the sampled
taxa forming an isolated clade that could be interpreted as sister to a clade that included Colbanthus,
Facchinia, Sabulina and Sagina.
Moehringia L., Sp. Pl. 1: 359. 1753 sec. Fior & Karis
(2007). – type: Moehringia muscosa L.
a group of 25 north-temperate species. Fior & Karis
(2007) found Moehringia could be made monophyletic by transferring four iberian species to Arenaria.
Moenchia ehrh. in neues Mag. aerzte 5: 203. 1783,
nom. cons. sec. Bittrich (1993c). – type: Moenchia
quaternella ehrh.
three species found in western and central europe.
While Harbaugh & al. (2010) noted that Moenchia
and Holosteum were sister taxa, Greenberg & donoghue (2011) found that Moenchia was a sister to
Cerastium.
Mononeuria rchb., deut. Bot. Herb.-Buch: 205; Syn.
red.: 118. 1841 sec. dillenberger & Kadereit (2014).
– type: Mononeuria patula (Michx.) dillenb. & Kadereit
= Geocarpon Mack. in torreya 14: 67. 1914.
= ?Selleola Urb. in ark. Bot. 23a(5): 69. 1930.
= Porsildia á. Löve & d. Löve in Bot. not. 128: 509.
1976.
nine species of eastern north america. dillenberger & Kadereit (2014) found Geocarpon was nested
within a clade consisting of Minuartia sect. Uni
nerviae (Fenzl) Mattf.; that clade was sister to a clade
containing Triplateia and three species of Stellaria on
the basis of matK sequences.
328
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
Myosoton Moench, Methodus: 225. 1794 sec. Bittrich
(1993c). – type: Myosoton aquaticum (L.) Moench
= Malachium Fr. ex rchb., Fl. Germ. excurs. 24: 795.
1832, nom. illeg.
Monotypic; temperate eurasia. treatment of the species as Stellaria aquatica L. may be warranted pending a serious review of Stellaria. it was found clustering near species of Stellaria sect. Stellaria by both
Harbaugh & al. (2010) and by Greenberg & donoghue (2011) in a study that more densely sampled
Stellaria.
Odontostemma Benth. ex G. don in Gen. Syst. 1: 449.
1831 sec. Sadeghian & al. (2015). – type: Odonto
stemma glandulosum Benth. ex G. don
= Gooringia F. n. Williams in Bull. Herb. Boissier 5:
530. 1897.
about 65 species of the Himalayas and adjacent
southern china. considered as a subgenus of Are
naria by many (e.g. Mcneill 1962), Harbaugh & al.
(2010) proposed, and Sadeghian & al. (2015) confirmed, that Odontostemma should be treated as a
genus, clustering with Cerastium and Stellaria rather
than Arenaria. Work on new combinations necessary
for recognizing most species in Odontostemma is underway (r. rabeler & W. Wagner, unpubl. data).
Ortegia L., Sp. Pl. 1: 560. 1753 sec. Bittrich (1993c). –
type: Ortegia hispanica L.
Monotypic; italy and iberian Peninsula. a member
of tribe Polycarpaeae, but relationships vary in different studies. Fior & al. (2006) showed Ortegia in
a cluster with Loeflingia and Polycarpon; Kool & al.
(2007) reported Ortegia clustering with Cardionema
and Illecebrum. curiously, using the matK sequence
from the Fior & al. (2006) study, both Harbaugh &
al. (2010) and Greenberg & donoghue (2011) found
that Ortegia clustered with a Hawaiian collection of
Drymaria cordata (L.) Willd. ex Schult. var. pacifica
Mizush.
Paronychia Mill. in Gard. dict. abr., ed. 4: [1019]. 1754
sec. Bittrich (1993c). – type: Paronychia argentea
Lam.
= Anychia Michx., Fl. Bor.-amer. 1: 112 – 113. 1803.
= Siphonychia torr. & a. Gray, Fl. n. amer. 1: 173.
1838.
= Gibbesia Small in Bull. torrey Bot. club 25: 621.
1898.
= Anychiastrum Small, Fl. S. e. U. S.: 400. 1903.
= Odontonychia Small, Fl. S. e. U. S.: 401. 1903.
= Gastronychia Small, Man. S. e. Fl.: 480, f. 1933.
in a study mainly addressing Gymnocarpos, Oxelman & al. (2002) found Paronychia to be polyphyletic, with the subgenera Paronychia and Siphonychia
forming a strongly supported sister group to Gym
nocarpos, whereas species in P. subg. Anoplonychia
(Fenzl) rchb. were found to be more closely related
to Herniaria and Philippiella. this was confirmed by
Greenberg & donoghue (2011). the genus consists
of 110 (Hartman & al. 2005) or more than 150 species
(Bittrich 1993b). it is one of the large genera in the
family that has not yet been extensively studied with
dna sequence data, especially in P. subg. Anoplony
chia (Fenzl) rchb. (only two of 48 species sampled).
Pentastemonodiscus rech. f. in anz. Österr. akad.
Wiss., Math.-naturwiss. Kl. 102: 11. 1965 sec. Bittrich (1993c). – type: Pentastemonodiscus monoch
lamydeus rech. f.
Monotypic; afghanistan. Presumed to be close to
Scleranthus, but has not yet been sampled for dna.
Petrocoptis a. Braun ex endl. in endl. Gen. Suppl. 2: 78.
1842 sec. Oxelman & al. (2001). – type: Petrocoptis
pyrenaica (Bergeret) a. Braun ex Walp.
= Silenopsis Willk. in Bot. zeitung (Berlin) 5: 237.
1847.
endemic to the iberian Peninsula, in particular the
Pyrenees. Species-level taxonomy is controversial,
with anything between one and 12 species (cires &
Prieto 2015) recognized. Phylogenetically, it occupies a position distinctly outside of the core Silene/
Lychnis clade according to several putatively unlinked genes (e.g. Oxelman & Lidén 1995; Oxelman
& al. 1997; Popp & Oxelman 2004), but the exact
position varies, suggesting a possible ancient hybrid
origin (Frajman & al. 2009a). cires & Prieto (2015)
confirmed the genus was monophyletic but noted
that additional study was needed to resolve infrageneric relationships.
Petrorhagia (Ser.) Link in Handbuch 2: 235. 1829 sec.
rabeler & Hartman (2005b) ≡ Gypsophila sect.
Petrorahgia Ser. in candolle, Prodr. 1: 354. 1824. –
type: Petrorhagia saxifraga (L.) Link
= Tunica Ludw., inst. regn. Veg. (ed. 2): 129. 1757.
= Kohlrauschia Kunth, Fl. Berol. ed. 2. 1: 108. 1838.
= Fiedleria rchb., icon. Fl. Germ. Helv. 6: 42. 1844.
comprising 33 species, ranging from the canary islands east to Kashmir. Shown to cluster as sister to a
clade including Dianthus and Velezia by Harbaugh &
al. (2010), Greenberg & donoghue (2011) and Pirani
& al. (2014). the genus has not been widely sampled.
although kept separate by Bittrich (1993c), most recent treatments of the genus include Kohlrauschia as
a section in Petrorhagia following the monograph of
Ball & Heywood (1964). this may deserve further
investigation since Greenberg & donoghue (2011)
cited three samples in their study; a voucher of “P.
velutina Guss.” (a later name for P. dubia (raf.) G.
López & romo) was shown as a sister to a clade including P. saxifraga (L.) Link and a second voucher
of P. dubia; the identification of the vouchers should
be verified.
Philippiella Speg. in revista Fac. agron. Univ. nac. La
Plata 1897: 566. 1897 sec. Bittrich (1993c). – type:
Philippiella patagonica Speg.
Monotypic; Patagonia. Oxelman & al. (2002) and
Greenberg & donoghue (2011) found P. patagonica
Willdenowia 45 – 2015
was nested within Herniaria; the genus is retained
pending additional sampling in Herniaria.
Phrynella Pax & K. Hoffm., nat. Pflanzenfam. (ed. 2)
16c: 364. 1934 sec. Bittrich (1993c). – type: Phrynel
la ortegioides (Fisch. & c. a. Mey.) Pax & K. Hoffm.
Monotypic; turkey. Possibly related to Gypsophila,
but not yet sampled for dna.
Pirinia M. Král in Preslia 56: 161. 1984 sec. Bittrich
(1993c). – type: Pirinia koenigii M. Král
Monotypic; Bulgaria. Placed in the Polycarpaeae, but
not yet sampled for dna.
Pleioneura rech. f. in Bot. Jahrb. Syst. 75: 357. 1951
sec. Bittrich (1993c). – type: Pleioneura griffithiana
(Boiss.) rech. f.
Monotypic; central asia to Himalayas. Possibly related to either Psammosilene or Saponaria (Bittrich
1993c), but not yet sampled for dna.
Plettkea Mattf. in Schriften Vereins naturk. Unterweser
7: 11, 13, 17. 1934 sec. Bittrich (1993c). – type: not
designated.
Four species of the Peruvian andes. the single species that has been sequenced clustered among species
of Stellaria in both Harbaugh & al. (2010) and Greenberg & donoghue (2011).
Pollichia aiton in Hort. Kew. 1: 5. 1789 – 1789, nom. cons.
sec. Bittrich (1993c). – type: Pollichia campestris
aiton
Monotypic; eastern and southern africa. Kool & al.
(2012) placed P. campestris as sister to the monotypic Sphaerocoma; both genera form a clade that
is sister to a clade containing Polycarpaea and Poly
carpon.
Polycarpaea Lam. in J. Hist. nat. 2: 3, 5. 1792, nom.
cons. sec. Bittrich (1993c). – type: Polycarpaea tene
riffae Lam.
= Robbairea Boiss., Fl. Orient. 1: 735. 1867.
= Reesia ewart in Proc. roy. Soc. Victoria, n.s., 26: 9.
1913.
a paleotropical group of 50+ species. Kool & al.
(2007, 2012) found it to be polyphyletic; additional
sampling is required to treat the genus, resolve infrageneric relationships and decide how some small genera (e.g. Haya, Xerotia) should be treated.
Polycarpon L., Syst. nat., ed. 10: 859, 881, 1360. 1759
sec. Kool & al. (2007). – type: Polycarpon tetraphyl
lum (L.) L.
Monotypic; Mediterranean and western north
america. Kool & al. (2007) found Polycarpon was
polyphyletic with species distributed in three clades.
two of these included species of Polycarpaea and
were removed from Polycarpon. the third included
members of the P. tetraphyllum group; tight relations
in the remaining clade suggested reduction to one
polymorphic species.
Polytepalum Suess. & Beyerle in Bot. Jahrb. Syst. 69:
143. 1938 sec. Bittrich (1993c). – type: Polytepalum
angolense Suess. & Beyerle
329
Monotypic; angola. Placed in the Polycarpaeae, but
not yet sampled for dna.
Psammophiliella ikonn. in novosti Sist. Vyssh. rast.
11: 116. 1976 sec. ikonnikov (1976). – type: Psam
mophiliella muralis (L.) ikonn.
= Psammophila Fourr. ex ikonn. in novosti Sist. Vyssh.
rast. 8: 273. 1971, nom. illeg. ≡ Psammophila Fourr.
in ann. Soc. Linn. Lyon sér. 2. 16: 345. 1868, nom.
inval.
Four species of central asia. Most often treated as
Gypsophila subg. Macrorrhizaea, but both Greenberg & donoghue (2011) and Pirani & al. (2014)
showed P. muralis as sister to a clade of Dianthus/
Petrorhagia, clearly separate from the remainder of
Gypsophila.
Psammosilene W. c. Wu & c. Y. Wu in L. P. King, icon.
Pl. Medic. 1: [s.n.], t. 1. 1945 sec. Bittrich (1993c). –
type: Psammosilene tunicoides W. c. Wu & c. Y. Wu
Monotypic; in montane forests of Yunnan, china.
Oxelman & Lidén (1995) found Psammosilene to be
sister to subfamily Caryophylloideae, while Greenberg & donoghue (2011) found it to be a sister to
tribe Caryophylleae (Dianthus/Gypsophila/Sapo
naria, etc.).
Pseudocerastium c. Y. Wu & al. in acta Bot. Yunnan.
20: 395. 1998 sec. Lu & rabeler (2001). – type:
Pseudocerastium stellarioides X. H. Guo & X. P.
zhang
Monotypic; china. Presumed close to Cerastium, but
not yet sampled for dna.
Pseudocherleria dillenb. & Kadereit in taxon 63: 79.
2014 sec. dillenberger & Kadereit (2014). – type:
Pseudocherleria laricina (L.) dillenb. & Kadereit
comprises 12 species found in the caucasus region,
arctic asia and northwestern north america. Formerly included (with Cherleria) in Minuartia sect.
Spectabiles (Fenzl) Hayek, dillenberger & Kadereit
(2014) found the two groups segregated into different
clades far from Minuartia s.str., proposing the recognition of both genera.
Pseudostellaria Pax, nat. Pflanzenfam. (ed. 2) 16c: 318.
1934 sec. Bittrich (1993c). – type: Pseudostellaria
rupestris (turcz.) Pax
a group of about 20 species, mostly in central asia
east to Japan, with one species in europe and three
in western north america. the few species thus far
sampled cluster near Lepyrodiclis and Odontostem
ma. Greenberg & donoghue (2011) included four
species and found the american P. jamesiana (torr.)
W. a. Weber & r. L. Hartm. did not cluster with the
three asian species; their report showing Stellaria
jamesiana torr. (≡ P. jamesiana (torr.) W. a. Weber
& r. L. Hartm.) clustering among Cerastium is based
on a misidentified specimen of C. arvense L.
Pteranthus Forssk. in Fl. aegypt.-arab.: 36. 1775 sec.
Bittrich (1993c). – type: Pteranthus dichotomus
Forssk.
330
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
Monotypic; northern africa east to iran. a member
of the Polycarpaeae, clustering with Dicheranthus
(Greenberg & donoghue 2011).
Pycnophyllopsis Skottsb. in Kongl. Svenska Vetenskapsakad. Handl. 56(5): 216. 1916 sec. Bittrich (1993c).
– type: Pycnophyllopsis muscosa Skottsb.
Segregation of Pycnophyllopsis from Pycnophyl
lum has been confirmed (M. timaná, unpubl. data).
Plettkea may belong here.
Pycnophyllum remy in ann. Sci. nat., Bot., ser. 3, 6:
355. 1846 sec. Bittrich (1993c). – type: not designated.
a genus of 17 andean species that clusters close to
Drymaria, a result first reported by Smissen & al.
(2003) and confirmed in four further studies. this
contradicts the earlier placement (e.g. Bittrich 1993c)
as a member of subfamily Alsinoideae.
Reicheella Pax, nat. Pflanzenfam., nachtr. 2: 21. 1900
sec. Bittrich (1993c). – type: Reicheella andicola
(Phil.) Pax
Monotypic; chile. not yet sampled in a molecular
study.
Rhodalsine J. Gay in ann. Sci. nat., Bot., ser. 3, 4: 25.
1845 sec. Favarger & Monserrat (1991). – type:
Rhodalsine procumbens J. Gay
= Psammanthe rchb., deut. Bot. Herb.-Buch: 205;
Syn. red.: 94. 1841, nom. rej. prop. (Kool & thulin
2013).
Five mostly Mediterranean species. Formerly treated as Minuartia subg. Rhodalsine (J. Gay) Graebn.
(Mcneill 1962), Harbaugh & al. (2010), Greenberg
& donoghue (2011) and Kool (2012) each found
Rhodalsine to be sister to Spergula and Spergularia.
Sabulina rchb., Fl. Germ. excurs. 24: 785. 1832 sec.
dillenberger & Kadereit (2014). – type: Sabulina
tenuifolia (L.) Hiern.
comprising c. 65 species (possibly 70, including
some eastern european and western asian taxa not
yet transferred to Sabulina: S. Mosyakin, unpubl.
data), all but two found in the northern hemisphere
(europe, asia and north america). including members of six sections of Mcneill’s (1962) Minuartia
subg. Minuartia as well as Stellaria fontinalis (Short
& Peter) B. L. rob., these species form a clade that
is sister to a clade including Colobanthus, Facchinia
and Sagina. rabeler & al. (2014) suggested this clade
may be further subdivided, possibly recognizing four
other genera.
Sagina L., Sp. Pl. 1: 128. 1753 sec. Bittrich (1993c). –
type: Sagina procumbens L.
= Spergella rchb., Handb. Gewächsk., ed. 2, 1: 65.
1827.
a genus of about 30 species, most diverse in northtemperate and arctic areas with a few taxa found on
some tropical mountains. Sampling shows Sagina to
be monophyletic, although infrageneric relationships
have not been studied.
Sanctambrosia Skottsb. ex Kuschel in ark. Bot., ser. 2,
4: 418. 1962 sec. Bittrich (1993c). – type: Sanctam
brosia manicata (Skottsb.) Skottsb. ex Kuschel
Monotypic; San ambrosio island (desventurados
archipelago), chile. Kool (2012) reported it nested
within a Spergularia clade.
Saponaria L., Sp. Pl. 1: 408. 1753 sec. Bittrich (1993c).
– type: Saponaria officinalis L.
= Melandryum [unranked] Gastrolychnis Fenzl in endlicher, Gen. Pl.: 974. 1840.
= Spanizium Griseb., Spic. Fl. rumel. 1: 180. 1843.
about 40 species, most diverse in the Mediterranean and southwestern asia. the most comprehensive
monograph dates from 1910 (Simmler 1910), with
Shults (1989) providing an updated account for russian taxa. Up to now, sampling has been minimal and
offers no information on how related genera (Bol
bosaponaria, Cyathophylla, Pleioneura, etc.) may
best be treated.
Schiedea cham. & Schltdl. in Linnaea 1: 46. 1826 sec.
Wagner & al. (2005). – type: Schiedea ligustrina
cham. & Schltdl.
= Alsinidendron H. Mann in Proc. Boston Soc. nat.
Hist. 10: 311. 1866.
a monophyletic group of 34 species endemic to the
Hawaiian islands. See Wagner & al. (2005) for a
monographic/phylogenetic revision and Harbaugh &
al. (2010) for comments on the origin of Schiedea.
Scleranthus L., Sp. Pl. 1: 406. 1753 sec. Bittrich (1993c).
– type: Scleranthus annuus L.
= Mniarum J. r. Forst. & G. Forst, char. Gen. Pl., ed. 2:
[1]. 1776.
about 12 species native to eurasia and australasia.
Smissen & al. (2003) found Scleranthus to be monophyletic and to be treated as two subgenera: S. subg.
Scleranthus (three species, eurasia) and S. subg.
Mniarum (J. r. Forst. & G. Forst.) Pax) (nine species, southeastern australasia). dillenberger & Kadereit (2014) found Scleranthus was sister to one of ten
clades of Minuartia s.l., treated by them as Cherleria.
Scopulophila M. e. Jones in contr. W. Bot. 12: 5. 1908
sec. Bittrich (1993c). – type: Scopulophila nitrophi
loides Jones
two species; southwestern United States and Mexico. Hartman (2005b) noted seed and flower characters suggesting a close relationship to Achyronychia;
Greenberg & donoghue (2011) showed this for S.
rixfordii (Brandegee) Munz & i. M. Johnst., but S.
parryi (Hemsl.) i. M. Johnst. clustered with Sphaero
coma aucheri Boiss. (= S. hookeri t. anderson subsp.
aucheri (Boiss.) Kool & thulin).
Silene L., Sp. Pl. 1: 416. 1753, nom. cons. prop. sec. Oxelman & al. (2001) ≡ Viscago zinn, cat. Pl. Gott.:
188. 1757 ≡ Kaleria adans., Fam. Pl. 2: 506. 1763 ≡
Corone Hoffmanns ex Steud., nomencl. Bot., ed. 2,
1: 422. 1840 ≡ Oncerum dulac, Fl. Hautes-Pyrénées:
255. 1867. – type: Silene anglica L. – Fig. 3G.
Willdenowia 45 – 2015
= Cucubalus L., Sp. Pl. 1: 414. 1753 ≡ Scribaea Borkh.
in rhein. Mag. 1: 591. 1793.
= Oberna adans., Fam. Pl. 2: 255. 1763 ≡ Silene sect.
Behenantha Otth in candolle, Prodr. 1: 367. 1824
≡ Behenantha (Otth) Schur in Verh. naturf. Vereins
Brünn 15(2): 130. 1877.
= Otites adans., Fam. Pl. 2: 255. 1763.
= Lychnanthos S. G. Gmel. in novi comment. acad.
Sci. imp. Petrop. 14(1): 525. 1770.
= Melandrium röhl., deutschl. Fl. (ed. 2) Phanerog.
Gew. 2: 37, 274. 1812.
= Lychnis sect. Physolychnis Benth., ill. Bot. Himal.
Mts.: 80. 1834 ≡ Lychnis [unranked] Gastrolychnis
Fenzl in endlicher, Gen. Pl.: 974. 1840 ≡ Gastrolych
nis (Fenzl) rchb., deut. Bot. Herb.-Buch: 206. 1841
≡ Wahlbergella Fries in Bot. not. (1843): 143. 1843
≡ Physolychnis rupr. in Mém. acad. imp. Sci. SaintPétersbourg, Sér. 7, 14: 41. 1869.
= Alifiola raf., autik. Bot.: 24. 1840.
= Ebraxis raf., autik. Bot.: 29. 1840.
= Evactoma raf., autik. Bot.: 23. 1840.
= Pleconax raf., autik. Bot.: 24. 1840 ≡ Conosilene
(rohrb.) Fourr. in ann. Soc. Linn. Lyon sér. 2. 16:
344. 1868.
= Xamilenis raf., autik. Bot.: 24. 1840.
= Elisanthe (Fenzl) rchb., deut. Bot. Herb.-Buch: 206.
1841.
= Silenanthe Griseb. & Schenk in archiv für naturgeschichte 18: 300. 1852.
= Polyschemone Schott, nymann & Kotschy in Schott,
analecta Bot.: 55. 1854.
= Carpophora Klotzsch in Bot. ergebn. reise Waldemar: 139. 1862.
= Leptosilene Fourr. in ann. Soc. Linn. Lyon sér. 2. 16:
344. 1868.
= Muscipula Fourr. in ann. Soc. Linn. Lyon sér. 2. 16:
344. 1868.
= Petrosilene Fourr. in ann. Soc. Linn. Lyon sér. 2. 16:
344. 1868.
= Petrocoma rupr. in Mém. acad. imp. Sci. SaintPétersbourg, Sér. 7, 15(2): 200. 1869.
= Anotites Greene in Leafl. Bot. Observ. crit. 1: 97.
1904.
= Gastrocalyx Schischk. in izv. Kavkazsk. Muz. 12:
200. 1919 ≡ Schischkiniella Steenis in Blumea 15:
145. 1967.
= Charesia e. a. Busch in trudy Bot. Muz. 19: 182.
1926.
= Sofianthe tzvelev in novosti Sist. Vyssh. rast. 33:
97. 2001.
= Neoussuria tzvelev in novosti Sist. Vyssh. rast. 34:
299. 2002.
Generic delimitation has been notoriously controversial (see Oxelman & Lidén 1995 for a review). Some
authors have lumped all c. 850 species of the tribe
Sileneae (except Agrostemma) in Silene (e.g. Greuter
1995), whereas tzvelev (2001) recognized 23 gen-
331
era in europe alone. Molecular evidence clearly supports separation of Agrostemma, Atocion, Eudianthe,
Heliosperma, Petrocoptis and Viscaria (e.g. Oxelman
& Lidén 1995; Oxelman & al. 1997, 2001; Popp &
Oxelman 2004; Frajman & al. 2009a, b; Greenberg &
donoghue 2011). However, monophyly of Silene, in
the sense adopted here, is only rarely supported by individual gene trees. Several studies have identified two
major clades (S. subg. Behenantha (Otth) endl. and S.
subg. Silene; e.g. Oxelman & Lidén 1995; Oxelman &
al. 1997, 2001; eggens & al. 2007; erixon & Oxelman
2008; rautenberg & al. 2012; aydin & al. 2014), but
their relationship to Lychnis is ambiguous. Silene sect.
Atocion Otth, a small group of annuals from the eastern
Mediterranean, appears to be blurring the picture, possibly due to highly elevated substitution rates across
the genome (z. aydin & al., unpubl. data).
Solitaria (Mcneill) Sadeghian & zarre, Bot. J. Linn.
Soc. 178: 667. 2015 sec. Sadeghian & al. (2015) ≡
Arenaria subg. Solitaria Mcneill, notes roy. Bot.
Gard. edinburgh. 24: 128, 1962. – type: Solitaria ci
liolata (edgew.) Sadeghian & zarre
a genus of about seven Himalayan species. Sadeghian & al. (2015) found Solitaria clustering as a sister
to either Odontostemma or Pseudostellaria.
Spergula L., Sp. Pl. 1: 440. 1753 sec. Bittrich (1993c). –
type: Spergula arvensis L.
Five north-temperate species. While López González
(2010) suggested Spergularia should be included in
Spergula based on morphology, Kool (2012) demonstrated that both genera are monophyletic.
Spergularia (Pers.) J. Presl & c. Presl, Fl. cech.: 94.
1819, nom. cons. sec. Bittrich (1993c) ≡ Arenaria
subg. Spergularia Pers., Syn. Pl. 1: 504. 1805 ≡ Tissa
adans., Fam. Pl. 2: 507, 611. 1763. – type: Spergu
laria rubra (L.) J. Presl & c. Presl
= Delia dumort. in Fl. Belg. 1: 110. 1827.
about 60 species (Hartman & rabeler 2005), especially diverse in the Mediterranean and temperate
South america. the genus is monophyletic (Kool
2012), but infrageneric relationships are not defined.
Sphaerocoma t. anderson in J. Proc. Linn. Soc., Bot.
5: 16. 1861 sec. Kool & al. (2012). – type: Sphaero
coma hookeri t. anderson
Monotypic; in deserts from Somalia east to Pakistan.
Kool & al. (2012) noted that Sphaerocoma is sister to
the monotypic Pollichia and together they form a sister
clade to one including Polycarpaea and Polycarpon.
Stellaria L., Sp. Pl. 1: 421. 1753 sec. Bittrich (1993c). –
type: Stellaria holostea L.
= Alsine L., Sp. Pl. 1: 272. 1753.
= Tytthostemma nevski in trudy Bot. inst. akad. nauk
S. S. S. r., Ser. 1, Fl. Sist. Vyss. rast 4: 305. 1937.
= Mesostemma Vved. in Bot. Mater. Gerb. Bot. inst.
Uzbekistansk. Fil. akad. nauk S. S. S. r. 3: 4. 1941.
= Fimbripetalum (turcz.) ikonn. in novosti Sist. Vyssh.
rast. 14: 78. 1977.
332
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
a genus of about 150 to more likely 200 species of
eurasia and north america, most diverse in the mountains of central asia. Stellaria is in need of a monographic revision; the most recent infrageneric classification is that of Pax & Hoffman (1934). Greenberg
& donoghue (2011) conducted the most extensive
sampling of Stellaria to date, including 44 species.
Stellaria is clearly polyphyletic and in need of further study: S. obtusa engelm. appeared as a sister to a
clade including Honckenya, Schiedea and Wilhemsia;
three Mexican/caribbean species were sister to Minu
artia sect. Uninerviae (Fenzl) Mattf. (= Mononeuria
of dillenberger & Kadereit 2014); S. americana (Porter ex B. L. rob.) Standl. clustered with Pseudostel
laria jamesiana (torr.) W. a. Weber & r. L. Hartm.;
and S. holostea, the type of Stellaria, appeared as sister to the clade that includes Cerastium, Dichodon,
Holosteum, Moenchia and the majority of Stellaria
species sampled.
Stipulicida Michx., Fl. Bor.-amer. 1: 26, pl. 6. 1803 sec.
Bittrich (1993c). – type: Stipulicida setacea Michx.
Stipulicida is found only in the southeastern United
States and cuba. Long thought to be monotypic, a
recent morphological study (Poindexter & al. 2014)
proposed recognition of two species. Work is underway to confirm placement in the Polycarpaeae (K.
neubig & r. rabeler, unpubl. data).
Telephium L., Sp. Pl. 1: 271. 1753 sec. Bittrich (1993c).
– type: Telephium imperati L.
Five species of the Mediterranean and southwestern
asia. Harbaugh & al. (2010) and Greenberg & donoghue (2011) both placed Telephium in tribe Corrigi
oleae, near the base of Caryophyllaceae.
Thurya Boiss. & Balansa in Boissier, diagn. Pl. Orient., ser. 2, 5: 63. 1856 sec. Bittrich (1993c). – type:
Thurya capitata Boiss. & Balansa
Monotypic; SW asia. not yet sampled in a molecular
study.
Thylacospermum Fenzl in endlicher, Gen. Pl.: 967. 1840
sec. Bittrich (1993c). – type: Periandra caespitosa
cambess.
two species, central asia and Himalayas. the phylogenetic placement is still uncertain. Harbaugh &
al. (2010) placed it “tentatively” into Eremogoneae,
whereas Greenberg & donoghue (2011) and dillenberger & Kadereit (2014) both found Thylacosper
mum closely aligned with Spergula; Greenberg &
donoghue (2011) considered their placement “uncertain”.
Triplateia Bartl. in Ord. nat. Pl.: 305. 1830 sec. dillenberger & Kadereit (2014). – type: Triplateia diffusa
Bartl.
= Hymenella Ser. in candolle, Prodr. 1: 389. 1824, nom.
illeg.
Monotypic; endemic to central Mexico. treated by
Mcneill (1962) as Minuartia subg. Hymenella (Ser.)
Mcneill. Harbaugh & al. (2010) and Greenberg &
donoghue (2011) both reported it as sister to Geocar
pon minimum Mack., a species endemic to the Ozark
region of the United States. dillenberger & Kadereit
(2014) found that it was a sister taxon to three species of Stellaria from Mexico and the caribbean.
this clade was, depending on the gene chosen, either
sister to Mononeuria (Minuartia sect. Uninerviae
+ Geocarpon) (matK) or sister to a clade including
Honckenya, Schiedea and Wilhelmsia (itS).
Vaccaria Wolf, Gen. Pl.: 3. 1776 sec. Bittrich (1993c). –
type: Vaccaria pyramidata Medik.
One or four species, native to eurasia. While usually thought to be closely related to Saponaria, both
Harbaugh & al. (2010) and Greenberg & donoghue
(2011) found a potential relation with Gypsophila
based on different vouchers: sister to Gypsophila in
the former study, clustering near the base of a Gypso
phila clade in the latter.
Velezia L., Sp. Pl. 1: 332. 1753 sec. Bittrich (1993c). –
type: Velezia rigida L.
Six species occurring from the Mediterranean east to
afghanistan. May be included in Dianthus; Harbaugh
& al. (2010) and Greenberg & donoghue (2011) both
found V. rigida nested in Dianthus, while Pirani & al.
(2014) showed Velezia as a sister to Dianthus.
Viscaria Bernh. in Syst. Verz.: 261. 1800, nom. cons. sec.
Oxelman & al. (2001) ≡ Steris adans., Fam. Pl. 2:
255. 1763. – type: Viscaria vulgaris Bernh.
= Liponeurum Schott, nymann & Kotschy in Schott,
analecta Bot.: 55. 1854.
recently revised by Frajman & al. (2013) with three
chiefly european and north american species.
Wilhelmsia rchb., consp. regn. Veg.: 206. 1828 sec.
Bittrich (1993c). – type: Arenaria physodes Fisch.
ex Ser.
Monotypic; arctic northwestern north america and
eastern asia. Harbaugh & al. (2010) found Wilhelm
sia and Honckenya are sister to each other and both
are the closest relatives to the Hawaiian Schiedea.
Xerotia Oliv. in Hooker’s icon. Pl.: t. 2359. 1895 sec. Bittrich (1993c). – type: Xerotia arabica Oliv.
Monotypic; arabia. Found to be nested in one of the
clades of Polycarpaea by Kool & al. (2012); placement awaits further resolution of polyphyly in Poly
carpaea.
Chenopodiaceae Vent. sec. Müller & Borsch (2005).
the family Chenopodiaceae is cosmopolitan predominantly occurring in temperate and subtropical regions,
and especially in semi-arid or arid environments (Kühn
1993; Kadereit & al. 2003). Our delimitation of the
Chenopodiaceae follows the concept of Ulbrich (1934),
and Kühn (1993) with the exception of the Polycnemoi
deae (see Amaranthaceae). considering that the core of
Chenopodiaceae (composed of Betoideae, Camphoros
moideae, Chenopodioideae, Salicornioideae, Salsoloi
deae and Suaedoideae) is likely to be monophyletic, we
Willdenowia 45 – 2015
maintain the Chenopodiaceae as a family distinct from
the Amaranthaceae in line with a series of current taxonomic treatments and morphological, physiological and
phylogenetic studies (tzvelev & al. 1996; Welsh & al.
2003; zhu & al. 2003; Kadereit & al. 2005; Kapralov &
al. 2006; Voznesenskaya & al. 2007; akhani & al. 2007;
zacharias & Baldwin 2010; Kadereit & al. 2010; Sukhorukov 2010; Flores-Olvera & al. 2011; Sukhorukov &
Kushunina 2014). We believe that name stability is important as it facilitates the assignment of genera to the
respective major Amaranthaceae and Chenopodiaceae
clades in line with the vast literature on Chenopodiaceae.
the monophyletic core Chenopodiaceae had already been
found with maximum support based on matK-trnK sequence data (Müller & Borsch 2005a), although relationships of the six major subfamilies were not clear. Much
progress has been made in the last decade on the internal
relationships of Chenopodiaceae. Schütze & al. (2003)
found two major clades of Suaedoideae Ulbr., to which
Bienertia is sister. the Salicornioideae were clearly identified as monophyletic and are a lineage of about 90 species growing worldwide in coastal and inland saline habitats (Kadereit & al. 2006) with often succulent-articulated
stems. Phylogenetic analysis yielded good support for the
Camphorosmoideae that include several major lineages of
mostly steppe, semi-desert and desert plants (Kadereit &
Freitag 2011), but genera of the Salsoloideae such as Sal
sola L. were depicted as largely polyphyletic (akhani &
al. 2007; Kadereit & Freitag 2011). the Chenopodioideae
were confirmed as monophyletic, although the members
of the genus Chenopodium in its pre-phylogenetic circumscription appeared scattered across the subfamily,
leading to a re-circumscription at genus and tribal level
(Fuentes-Bazán & al. 2012a, b).
Acroglochin Schrad., Mant. 1: 69, 227. 1822 sec. Kühn
(1993). – type: Acroglochin chenopodioides Schrad.
the central asian genus Acroglochin represents an
ancient lineage in Chenopodiaceae and stands phylogenetically isolated from other subfamilies (Hohmann & al. 2006; Kadereit & al. 2012, online supplement). the genus was formerly included in Betoideae
and should probably be classified as a subfamily of
its own.
Agriophyllum M. Bieb. in Fl. taur.-caucas. 3: 6.
1819 – 1820 sec. Kühn (1993). – type: Agriophyllum
arenarium M. Bieb.
Agriophyllum comprises six western and central asian
species of annual herbs and belongs to the Corisper
moideae (Kadereit & al. 2003).
Allenrolfea Kuntze in revis. Gen. Pl. 2: 545. 1891 sec.
Kühn (1993). – type: Allenrolfea occidentalis (S.
Watson) Kuntze – Fig. 4B.
Allenrolfea comprises two or three species of stemsucculent halophytes distributed in the americas (Kadereit & al. 2006a).
Anabasis L., Sp. Pl. 1: 223. 1753 sec. Kühn (1993). –
type: Anabasis aphylla L.
333
= Brachylepis c. a. Mey. ex Ledeb., icon. Pl. 1: 12.
1829.
= Fredolia (coss. & durieu ex Bunge) Ulbr., nat.
Pflanzenfam. (ed. 2) 16c: 451, 578. 1934.
= Esfandiaria charif & aellen in Verh. naturf. Ges. Basel 63: 262. 1952.
a diverse genus within the Salsoleae s.str. (akhani
& al. 2007) distributed throughout northern african
and eurasian steppes, semi-deserts and deserts. the
genus evolved some extremely drought-tolerant species, e.g. A. calcarea (charif & aellen) Bokhari &
Wendelbo, showing anatomical and morphological
adaptations to drought such as stunted growth forms,
reduced leaves, central water storage tissues and a
multi-layered epidermis (Bokhari & Wendelbo 1978).
the fruit anatomy of the genus was studied by Sukhorukov (2008).
Anthochlamys Fenzl in endlicher, Gen. Pl.: 300. 1837
sec. Kühn (1993). – type: Anthochlamys polyga
loides (Fisch. & c. a. Mey.) Moq.
Anthochlamys comprises three southwestern asian
species of annual herbs and belongs to the Corisper
moideae (Kühn 1993a; Kadereit & al. 2003). carpologically it is closely related to Corispermum (Sukhorukov 2014).
Aphanisma nutt. ex Moq. in candolle, Prodr. 13(2): 43,
54. 1849 sec. Kühn (1993). – type: Aphanisma blito
ides nutt. ex Moq.
this monotypic genus is distributed in coastal habitats of california and according to Hohmann & al.
(2006) it belongs to Betoideae–Hablitzieae. together
with its sister genus Oreobliton, which is distributed
in northern africa, Aphanisma represents an interesting example of a western eurasian–western north
american disjunction (Kadereit & Baldwin 2012).
Archiatriplex G. L. chu in J. arnold arbor. 68: 461.
1987 sec. Kühn (1993). – type: Archiatriplex nanpin
ensis G. L. chu
this monotypic genus is only known from northern
Sichuan province, china, near nanping (chu 1987).
Archiatriplex is interpreted as an ancient lineage of
the Chenopodieae (formerly Atripliceae), based on
molecular phylogenetic and morphological evidence
(Kadereit & al. 2010).
Arthrocnemum Moq. in chenop. Monogr. enum.: 111.
1840 sec. Kühn (1993). – type: Arthrocnemum glau
cum Ung.-Sternb.
Arthrocnemum belongs to Salicornioideae. in its current circumscription, the genus consists of two disjunctly distributed species, the eurasian and northern
african A. macrostachyum (Moric.) K. Koch and
the north american and Mesoamerican A. subtermi
nale (Parish) Standl. Both are stem-succulent hygrohalopyhtes (Kadereit & al. 2006a).
Arthrophytum Schrenk in Bull. cl. Phys.-Math. acad.
Pétersb. 3: 211. 1845 sec. Kühn (1993). – type: Ar
throphytum subulifolium Schrenk
334
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
the genus belongs to Salsoleae s.str. and comprises
c. nine species (akhani & al. 2007).
Atriplex L., Sp. Pl. 1: 1052. 1753 sec. Kadereit & al.
(2010). – type: Atriplex hortensis L.
= Obione Gaertn., Fruct. Sem. Pl. 2: 198. 1791.
= Atriplex [unranked] Theleophyton Hook. f. in London
J. Bot. 6: 279. 1847 ≡ Theleophyton (Hook. f.) Moq.
in candolle, Prodr. 13(2): 44, 115. 1849.
= Blackiella aellen in Bot. Jahrb. Syst. 68: 423. 1938.
= Morrisiella allen in Bot. Jahrb. Syst. 68: 422. 1938.
= Pachypharynx allen in Bot. Jahrb. Syst. 68: 429.
1938.
= Senniella aellen in Bot. Jahrb. Syst. 68: 416. 1938.
= Cremnophyton Brullo & Pavone in candollea 42:
622. 1987.
Atriplex is the most species-rich genus within
Chenopodiaceae with c. 300 species. this cosmopolitan genus comprises annual or perennial herbs,
subshrubs and shrubs that are often prominent floristic elements of steppes, semi-deserts and coastal habitats (Kadereit & al. 2010). Most species of Atriplex
are c4 plants that all belong to one large c4 lineage.
Many species of the genus are halophytes and possess salt glands. Ontogenetic studies showed that the
two more or less concrescent “bracteoles” that envelop the fruit and that are characteristic of Atriplex are
better interpreted as two tepals (Flores-Olvera & al.
2011). the circumscription of Atriplex has changed
over time, and several infrageneric classifications
have been proposed (Flores & davis 2001; Kadereit
& al. 2010). recent phylogenetic studies based on
molecular data (Kadereit & al. 2010; zacharias &
Baldwin 2010) show that Atriplex in its traditional
circumscription is not monophyletic and includes
several satellite genera that have been separated in
the past. a new infrageneric classification is needed.
Previously Atriplex was placed in the tribe Atripli
ceae. However, because the previous Chenopodieae
are paraphyletic to Atripliceae the tribes were merged
together by Fuentes-Bazán & al. (2012b). the accepted name of the tribe in the new, monophyletic
definition is Atripliceae.
Axyris L., Sp. Pl. 1: 979. 1753 sec. Kühn (1993). – type:
Axyris amaranthoides L.
Axyris, together with Ceratocarpus and Kraschenin
nikovia, constitutes the Axyrideae (Kadereit & al.
2010). the genus consists of six species mainly concentrated in the mountains of central asia and eastern
Siberia (Sukhorukov 2011); some species (especially
A. amaranthoides L.) occur as alien weeds in eurasia
and north america beyond their native range. investigated species of the genus show heterocarpy (Sukhorukov 2005, 2011).
Baolia H. W. Kung & G. L. chu in acta Phytotax. Sin.
16(1): 119. 1978 sec. Kühn (1993). – type: Baolia
bracteata H. W. Kung & G. L. chu
the phylogenetic position of this rare monotypic
genus from china is unknown. according to Kühn
(1993a) it belongs to the Chenopodioideae.
Bassia all. in Mélanges Philos. Math. Soc. roy. turin
3: 177. 1766 sec. Kadereit & Freitag (2011). – type:
Bassia muricata (L.) asch.
= Kochia roth in J. Bot. (Schrader) 1800(1): 307. 1801.
= Echinopsilon Moq., ann. Sci. nat. Bot., ser. 2, 2: 127.
1834, nom. illeg.
= Londesia Fisch. & c. a. Mey. in index Seminum [St.
Petersburg (Petropolitanus)] 2: 40. 1836.
= Panderia Fisch. & c. a. Mey. in index Seminum [St.
Petersburg (Petropolitanus)] 2: 21. 1836.
= Kirilowia Bunge in del. Sem. Hort. dorpat. 1843: 7.
1843.
= Chenoleoides (Ulbr.) Botsch. in Bot. zhurn. (Moscow & Leningrad) 61: 1408. 1976 ≡ Chenolea sect.
Chenoleoides Ulbr., nat. Pflanzenfam. (ed. 2) 16c:
530. 1934.
Bassia belongs to Camphorosmoideae–Camphoros
meae and in the circumscription adopted here consists of c. 20 c4 annuals or perennials. the genus is
distributed from the western Mediterranean to eastern
asia (Kadereit & Freitag 2011), with the main centre
of diversity in central asia. it represents an interesting
example of c4 leaf-type diversity (Freitag & Kadereit
2014) and multiple reduction of water-storage tissue
(akhani & Khoshravesh 2013; Kadereit & al. 2014).
Beta L., Sp. Pl. 1: 222. 1753 sec. Kadereit & al. (2006b).
– type: Beta vulgaris L.
Beta comprises seven species of annuals or biennial and perennial herbs with a storage root. Beta is
subdivided into two sections and is the only genus
of tribe Beteae. Beta sect. Procumbentes Ulbr. (≡ B.
[unranked] Patellares tranzschel) was excluded from
Beta on the basis of molecular phylogenetic and morphological results (see under Patellifolia; Hohmann
& al. 2006; Kadereit & al. 2006b). Beta vulgaris and
its various cultivated varieties (sugar beet, beetroot,
fodder beet and chard) are the economically most important crops within Caryophyllales (McGrath & al.
2011). For B. vulgaris the chloroplast genome (Li &
al. 2014) and the nuclear genome (dohm & al. 2013)
have been sequenced recently.
Bienertia Bunge ex Boiss. in Fl. Orient. [Boissier]: 945.
1879 – 1879 sec. Kühn (1993). – type: Bienertia cy
cloptera Bunge ex Boiss.
the genus comprises three species that grow in temporarily wet saline habitats in iran and surrounding
countries (akhani & al. 2005, 2012). the discovery
of akhani & al. (1997) and Freitag & Stichler (2002)
that B. cycloptera Bunge is a c4 plant without Kranz
tissues triggered a large number of physiological, biochemical and genetic studies investigating c4 photosynthesis in this genus (akhani & al. 2009).
Blitum L., Sp. Pl. 1753 1: 4. 1753 sec. Fuentes-Bazán &
al. (2012b) ≡ Morocarpus Boehm., def. Gen. Pl., ed.
3: 385. 1760, nom. illeg. ≡ Chenopodium sect. Bli
Willdenowia 45 – 2015
tum (L.) Benth. & Hook f., Gen. Pl. 3(1): 52. 1880 ≡
Chenopodium sect. Eublitum aellen in Verh. naturf.
Ges. Basel 41: 103. 1930 ≡ Chenopodium subg. Bli
tum (L.) Hiitonen, Suom. Kasvio: 307. 1933. – type:
Blitum capitatum L.
= Anserina dumort., Fl. Belg. 1: 21. 1827 ≡ Agatho
phytum Moq. in ann. Sci. nat., Bot., ser. 2, 1: 291.
1834, nom. illeg. ≡ Orthosporum subg. Agathophy
tum t. nees, Gen. Fl. Germ. [1]: ad. t. [57]. 1835 ≡
Chenopodium sect. Agathophytum (t. nees) Benth.
& Hook. f., Gen. Pl. 3(1): 52. 1880.
= Monolepis Schrad. in index Seminum Hort. acad.
Gotting. 1830: 4. 1830.
= Chenopodium [unranked] Californica Standl., n.
amer. Fl. 21(1): 30. 1916.
= Chenopodium sect. Atriplicina allen in Verh. naturf.
Ges. Basel 41: 99. 1930 ≡ Scleroblitum Ulbr., nat.
Pflanzenfam. (ed. 2) 16c: 495. 1934.
in the last century, the Linnaean Blitum, with its two
species, B. capitatum and B. virgatum L., was usually merged with Chenopodium s.l. (e.g. aellen 1929;
iljin & aellen 1936; aellen & Just 1943; aellen
1960 – 1961; Grubov 1966; Brenan & akeroyd 1993;
Mosyakin 1996; Uotila 1997, 2001a, b; clemants &
Mosyakin 2003), or rarely recognized in the original
Linnaean circumscription (Scott 1978a). However,
the resurrection of this genus based on phylogenetic reconstruction supports a monophyletic lineage
and a wide concept of Blitum including c. ten species (Fuentes-Bazán & al. 2012b), most of them in
the northern hemisphere and one (B. atriplicimum F.
Muell.) in australia. Blitum belongs to the tribe An
serineae.
Camphorosma L., Sp. Pl. 1: 122. 1753 sec. Kadereit &
Freitag (2011). – type: Camphorosma monspeliaca L.
Camphorosma belongs to Camphorosmoideae–Cam
phorosmeae and consists of four c4 annuals or perennials. the genus is distributed from the western Mediterranean to central asia (Kadereit & Freitag 2011).
Caroxylon thunb., nov. Gen. 2: 37. 1782 sec. akhani &
al. (2007) ≡ Salsola sect. Caroxylon (thunb.) Fenzl,
nov. Gen. 2: 37. 1782. – type: Caroxylon aphyllum
(L. f.) tzvelev
= Salsola sect. Cardiandra aellen in notes roy. Bot.
Gard. edinburgh 28: 32. 1967.
= Salsola sect. Malpighipila Botsch. in Bot. zhurn.
(Moscow & Leningrad) 54: 990. 1969.
= Salsola sect. Irania Botsch. in Bot. zhurn. (Moscow
& Leningrad) 71: 1400. 1986.
= Nitrosalsola tzvelev in Ukrayins’k. Bot. zhurn. 50:
80. 1993.
the genus Caroxylon was resurrected by tzvelev
(1993) and then confirmed and re-circumscribed
based on molecular and morphological evidence
(akhani & al. 2007). in that circumscription, it is
the most diverse genus in Salsoloideae with c. 100
species distributed in central and southwestern asia,
335
the Mediterranean region and northern and southern
africa (Feodorova 2011). Feodorova & Samigullin
(2014) revealed four clades within Caroxylon s.l.
and provisionally advocated further splitting of the
genus, with recognition of Caroxylon s.str., a recircumscribed Nitrosalsola, and possibly two other
segregate genera, based on molecular and morphological evidence.
Ceratocarpus L., Sp. Pl. 1: 969. 1753 sec. Kühn (1993).
– type: Ceratocarpus arenarius L.
= Ceratoides Gagnebin in acta Helv. Phys.-Math. 2:
59. 1755.
this monotypic genus comprises one annual widespread eurasian species and belongs to the Axyrideae
G. Kadereit & Sukhor. (Kadereit & al. 2010).
Chenolea thunb. in nov. Gen. Pl.: 9. 1781 sec. Kadereit
& Freitag (2011). – type: Chenolea diffusa thunb.
this genus is distributed in southern africa and comprises two perennial species (Snijman & Manning
2013; Kadereit & al. 2014). Other taxa previously
placed in Chenolea are now mainly included in Bas
sia s.l. or Chenoleoides (see discussion under Bas
sia).
Chenopodiastrum S. Fuentes & al. in Willdenowia 42:
14. 2012 sec. Fuentes-Bazán & al. (2012b) ≡ Cheno
podium subsect. Undata aellen & iljin ex Mosyakin
& clemants in novon 6: 400. 1996. – type: Chenopo
diastrum murale (L.) S. Fuentes & al.
= Chenopodium [unranked] Hybrida Standl., n. amer.
Fl. 21(1): 13. 1916 ≡ Chenopodium sect. Grossefo
veata aellen & iljin ex Mosyakin in Ukrayins’k.
Bot. zhurn. 50: 75. 1993 ≡ Chenopodiastrum sect.
Grossefoveata (Mosyakin) Mosyakin in Phytoneuron
2013-56: 6. 2013.
Chenopodiastrum is a widespread new genus with six
or seven species and segregated from Chenopodium
s.l. its recognition is based mainly on molecular phylogenetic studies (Fuentes-Bazán & al. 2012a, b). the
genus is subdivided into two groups, for which sectional rank was proposed (Mosyakin 2013).
Chenopodium L., Sp. Pl. 1: 218. 1753 sec. FuentesBazán & al. (2012b) ≡ Chenopodium sect. Leprophyl
lum dumort., Fl. Belg. 1: 21. 1827 ≡ Chenopodium
sect. Chenopodiastrum Moq. in candolle, Prodr.
13(2): 61. 1849 ≡ Vulvaria Bubani, Fl. Pyren. 1: 174.
1897, nom. illeg. – type: Chenopodium album L. –
Fig. 4c & d.
= Rhagodia r. Br., Prodr. Fl. nov. Holland.: 408. 1810.
= Einadia raf., Fl. tellur. 4: 121. 1838.
= Chenopodium ser. Cicatricosa aellen in Feddes repert. Spec. nov. regni Veg. 69: 69. 1964 ≡ Chenopo
dium subsect. Cicatricosa (aellen) Mosyakin & clemants in novon 6: 402. 1996.
= Chenopodium ser. Favosa aellen in Feddes repert.
Spec. nov. regni Veg. 69: 69. 1964 ≡ Chenopodium
subsect. Favosa (aellen) Mosyakin & clemants in
novon 6: 401. 1996.
336
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
= Chenopodium subsect. Fremontiana (Standl.)
Mosyakin & clemants in novon 6: 401. 1996 ≡
Chenopodium [unranked] Fremontiana Standl. in Fl.
Bor.-amer. (Michaux) 21: 18. 1916.
= Chenopodium subsect. Leptophylla (Standl.)
Mosyakin & clemants in novon 6: 400. 1996 ≡
Chenopodium [unranked] Leptophylla Standl. in Fl.
Bor.-amer. (Michaux) 21: 14. 1916.
= Chenopodium subsect. Standleyana Mosyakin & clemants in novon 6: 402. 1996.
Chenopodium has been considered one of the most
diverse genera within Chenopodiaceae with c. 150
species (Kühn 1993), or even up to 250 species (under a narrow species concept). the circumscription
has considerably changed over time, and several
infrageneric classifications have been proposed. in
a wide sense, Kühn (1993) and Mosyakin & clemants (1996) recognized three subgenera: C. subg.
Ambrosia a. J. Scott, C. subg. Blitum (L.) Hiitonen
and C. subg. Chenopodium, and this classification
was followed by several authors in recent treatments for the genera. However, it was proposed,
based initially on morphological data, to include C.
subg. Ambrosia into the re-circumscribed genus Dys
phania r. Br. (Mosyakin & clemants 2003, 2008;
clemants & Mosyakin 2003; zhu & al. 2003). recent phylogenetic studies based on molecular data
(Fuentes-Bazán & al. 2012a, b) have shown that
Chenopodium in its traditional circumscription is
not monophyletic and consists of six independent
lineages. Fuentes-Bazán & al. (2012b) also gave
the morphological descriptions of the segregates,
including Chenopodium s.str., which still remains
the most species-rich and most widespread genus
of the group. Chenopodium belongs to Atripliceae
(earlier Chenopodieae), which is monophyletic in
the circumscription by Fuentes-Bazán & al. (2012b).
the typification of the genus Chenopodium is debated. if the same solution is adopted for Chenopodium
as that proposed for Salsola by akhani & al. (2014),
i.e. the recognition of the lectotype proposed under
the “american code” (arthur & al. 1907) (C. rubrum
L. in our case), then the genus recognized here as
Oxybasis should be called Chenopodium s.str., and
the genus containing C. album L. (the lectotype of
Chenopodium as recognized here) should probably be called Rhagodia, which will have disastrous
consequences for taxonomy and nomenclature of
the group (see discussion in Mosyakin & clemants
1996; Fuentes-Bazán & al. 2012b).
Choriptera Botsch. in Bot. zhurn. (Moscow & Leningrad) 52: 804. 1967 sec. Kühn (1993). – type: Chori
ptera semhahensis (Vierh.) Botsch.
= Gyroptera Botsch. in Bot. zhurn. (Moscow & Leningrad) 52: 807. 1967.
this genus has not yet been included in any molecular phylogenetic study. it belongs to the african-
arabian subtribe Sevadinae, presumably included in
Salsoleaae (Botschantzev 1975).
Climacoptera Botsch. in Sborn. Geobot. akad. Sukachev: 111. 1956 sec. akhani & al. (2007). – type:
Climacoptera lanata (Pall.) Botsch.
Climacoptera s.str. represents a monophyletic c4 genus within Caroxyleae. the genus is distributed in
central and southwestern asia and comprises only annual species. Highly contradictory species numbers,
ranging from six to c. 42, are given (akhani & al.
2007; Pratov 1986).
Corispermum L., Sp. Pl. 1: 4. 1753 sec. Kühn (1993). –
type: Corispermum hyssopifolium L.
Corispermum comprises 60–65 annual psammophytic (rarely glareophytic) species naturally distributed mainly in eurasia, with fewer than ten species
native in north america (Mosyakin 1995). Species
delimitations and distribution are poorly understood
because of high morphological variability and possible recent explosive radiation of local races. there
is one molecular phylogenetic study of Corispermum
by Xue & zhang (2011) that is limited to chinese
species and shows a rather poor infrageneric resolution. the genus is in need of a taxonomic revision
based on comprehensive molecular phylogenetic and
morphological studies.
Cornulaca delile in Fl. egypte: 206. 1813-1814 sec.
Kühn (1993). – type: Cornulaca monacantha delile
Cornulaca comprises c. six species occurring in central and southwestern asia and northern africa. the
genus is presumably monophyletic (akhani & al.
2007).
Cyathobasis aellen in candollea 12: 160. 1949 sec. Kühn
(1993). – type: Cyathobasis fruticulosa (Bunge) aellen
a monotypic genus from anatolia, closely related
to Girgensohnia and Hammada of Salsoleae s.str.
(akhani & al. 2007).
Cycloloma Moq. in chenop. Monogr. enum.: 17. 1840
sec. Kühn (1993). – type: Cycloloma platyphyllum
(Michx.) Moq.
= Cyclolepis Moq. in ann. Sci. nat., Bot., sér. 2, 1: 203.
1834, nom. illeg.
a monotypic genus from north america that is probably phylogenetically nested within Dysphania (Dys
phanieae; G. Kadereit, unpubl. data).
Didymanthus endl. in nov. Stirp. dec.: 7. 1839 sec.
Kühn (1993). – type: Didymanthus roei endl.
this monotypic genus belongs to the australian Cam
phorosmeae and is closely related to Dissocarpus and
Eriochiton (cabrera & al. 2009). the genus is endemic to australia (Wilson 1984).
Dissocarpus F. Muell. in trans. Phil. inst. Vict. 2: 75.
1858 sec. Kühn (1993). – type: Dissocarpus biflorus
F. Muell.
Four species are currently recognized in Dissocarpus
(australian Camphorosmeae). the genus is endemic
Willdenowia 45 – 2015
to australia (Wilson 1984). it is closely related to Di
dymanthus and Eriochiton (cabrera & al. 2009).
Dysphania r. Br., Prodr. Fl. nov. Holland.: 411. 1810
sec. Fuentes-Bazán & al. (2012b). – type: Dysphania
littoralis r. Br.
= Chenopodium [unranked] Orthosporum r. Br., Prodr.
Fl. nov. Holland.: 407. 1810 ≡ Blitum [unranked]
Orthosporum (r. Br.) c. a. Mey. in Ledebour, Fl.
altaic. 1: 11. 1829 ≡ Orthosporum (r. Br.) t. nees,
Gen. Fl. Germ. [1]: ad t. [57] [!]. 1834 ≡ Dysphania
sect. Orthospora (r. Br.) Mosyakin & clemants in
Ukrayins’k. Bot. zhurn. 59: 382. 2002.
= Chenopodium [unranked] Botryoides c. a. Mey. in
Ledebour, Fl. altaic. 1: 410. 1829 ≡ Chenopodium
[unranked] Botrys rchb., Fl. Germ. excurs. 24: 580.
1832 ≡ Chenopodium sect. Botrys (rchb.) W. d. J.
Koch, Syn. Fl. Germ. Helv: 607. 1837 ≡ Ambrina sect.
Botryois Moq., chenop. Monogr. enum.: 36. 1840,
nom. illeg. ≡ Vulvaria sect. Botrys (rchb.) Bubani,
Fl. Pyren. 1: 177. 1897 ≡ Botrys (rchb.) nieuwl. in
amer. Midl. naturalist 3: 274. 1914 ≡ Chenopodium
subsect. Botrys aellen & iljin, Fl. UrSS 6: 46. 1936
≡ Neobotrydium Moldenke in amer. Midl. naturalist
35: 330. 1946 ≡ Chenopodium sect. Botryoides a. J.
Scott in Bot. Jahrb. Syst. 100: 212. 1978 ≡ Dysphania
sect. Botryoides (c. a. Mey.) Mosyakin & clemants
in Ukrayins’k. Bot. zhurn. 59: 383. 2002.
= Roubieva Moq. in ann. Sci. nat., Bot., ser. 2, 1: 292.
1834 ≡ Ambrina Spach, Hist. nat. Vég. 5: 295. 1836,
nom. illeg. ≡ Chenopodium sect. Roubieva (Moq.)
Volkens, nat. Pflanzenfam. 3(1a): 61. 1893 ≡ Dys
phania sect. Roubieva (Moq.) Mosyakin & clemants
in Ukrayins’k. Bot. zhurn. 59: 382. 2002.
= Botrydium Spach, Hist. nat. Vég. 5: 298. 1836.
= Ambrina Moq., chenop. Monogr. enum.: 36. 1840
≡ Ambrina sect. Adenois Moq., chenop. Monogr.
enum.: 39. 1840 ≡ Chenopodium sect. Ambrina
Benth. & Hook. f., Gen. Pl. 3(1): 51. 1880 ≡ Cheno
podium [unranked] Ambrosioidia Standl., n. amer.
Fl. 21(1): 26. 1916 ≡ Chenopodium subg. Ambrosia
a. J. Scott in Bot. Jahrb. Syst. 100: 211. 1978 ≡ Dys
phania sect. Adenois (Moq.) Mosyakin & clemants
in Ukrayins’k. Bot. zhurn. 59: 382. 2002.
= Chenopodium [unranked] Carinata Standl., n. amer.
Fl. 21(1): 27. 1916.
= Chenopodium [unranked] Incisa Standl., n. amer. Fl.
21(1): 25. 1916.
= Meiomeria Standl., n. amer. Fl. 21(1): 7. 1916.
= Chenopodium sect. Tetrasepala allen in Bot. Jahrb.
Syst. 63: 490. 1930 ≡ Dysphania sect. Tetrasepalae
(allen) a. J. Scott in Bot. Jahrb. Syst. 100: 218. 1978.
= Chenopodium sect. Margaritaria Brenan in Kew
Bull. 11: 166. 1956.
= Chenopodium sect. Nigrescentia allen in acta Bot.
acad. Sci. Hung. 19: 3. 1973.
= Dysphania sect. Caudatae a. J. Scott in Bot. Jahrb.
Syst. 100: 218. 1978.
337
the widespread genus Dysphania comprises c. 50
species native mostly to South america (D. sect. Ade
nois), eurasia and africa (D. sect. Botryoides) and
australia (D. sect. Dysphania, D. sect. Orthospora
and D. sect. Tetrasepalae). traditionally, only native
australian taxa were included in Dysphania (Scott
1978; Wilson 1983, 1984); later an expanded circumscription of the genus was proposed (Mosyakin &
clemants 2002, 2008) based on morphological evidence. Further molecular phylogenetic studies (Kadereit & al. 2003, 2010; Fuentes-Bazán & al. 2012a, b)
demonstrated that Dysphania is phylogenetically
distant from Chenopodium and forms the tribe Dys
phanieae together with the closely related Suckleya
and Teloxys. the latter was included in Dysphania
based on morphology (Mosyakin & clemants 2002,
2008; clemants & Mosyakin 2003; zhu & al. 2003),
but should be recognized as a separate genus based
on molecular results (Kadereit & al. 2010; FuentesBazán & al. 2012a, b).
Enchylaena r. Br., Prodr. Fl. nov. Holland.: 407. 1810
sec. Kühn (1993). – type: Enchylaena tomentosa r.
Br.
Enchylaena seems to be polyphyletic (cabrera & al.
2009). However, more molecular data for this genus
of australian Camphorosmeae are needed before taxonomic rearrangements can be done.
Eokochia Freitag & G. Kadereit in taxon 80: 72. 2011
sec. Kadereit & Freitag (2011). – type: Eokochia
saxicola (Guss.) Freitag & G. Kadereit
a rare endangered monotypic genus of Camphoros
meae growing on coastal cliffs in the central Mediterranean (iamonico & Kadereit 2013). Eokochia is sister to the north american genus Neokochia (Kadereit
& Freitag 2011), thus belonging to a clade showing
an ancient Mediterranean–north american disjunction.
Eremophea Paul G. Wilson in Fl. australiana 4: 326.
1984 sec. Kühn (1993). – type: Eremophea aggre
gata Paul G. Wilson
the genus is endemic to australia and belongs to the
Camphorosmeae. it is closely related to Neobassia
(cabrera & al. 2009).
Exomis Fenzl ex Moq. in chenop. Monogr. enum.: 49.
1840 sec. Kühn (1993). – type: Exomis axyrioides
Fenzl ex Moq.
a monotypic genus distributed in South africa. it
belongs to the Archiatriplex clade within Chenopo
dieae, formerly Atripliceae (Kadereit & al. 2010).
Extriplex e. H. zacharias in Syst. Bot. 35: 850. 2010
sec. zacharias & Baldwin (2010). – type: Extriplex
joaquinana (a. nelson) e. H. zacharias
the two species of Extriplex are endemic to the california Floristic Province. Extriplex belongs to the
Archiatriplex clade within Chenopodieae, formerly
Atripliceae (Kadereit & al. 2010; see zacharias &
Baldwin 2010 for detailed information on the genus).
338
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
Girgensohnia Bunge ex Fenzl, Fl. ross.: 835. 1851 sec.
Sukhorukov (2007). – type: Girgensohnia pallasii
Bunge
Girgensohnia comprises five annual species in central
asia and iran (Sukhorukov 2007). the genus belongs
to the Salsoleae s.str. and is closely related to Cyatho
basis and Hammada (akhani & al. 2007).
Grayia Hook. & arn. in Bot. Beechey Voy.: 387. 18401840 sec. zacharias & Baldwin (2010). – type:
Grayia polygaloides Hook. & arn.
= Zuckia Standl. in J. Wash. acad. Sci. 5: 58. 1915.
a small north american genus that belongs to the
Archiatriplex clade within Chenopodieae, formerly
Atripliceae (Kadereit & al. 2010; see zacharias &
Baldwin 2010 for detailed information on the genus).
Grubovia Freitag & G. Kadereit in taxon 80: 72. 2011
sec. Kadereit & Freitag (2011). – type: Grubovia
dasyphylla (Fisch. & c. a. Mey.) Freitag & G. Kadereit
a central asian genus comprising three annual c3
species previously included in Bassia and/or Kochia.
Grubovia is sister to the speciose clade of australian
Camphorosmeae (Kadereit & Freitag 2011).
Hablitzia M. Bieb. in Mém. Soc. imp. naturalistes Moscou 5: 24. 1817 sec. Kühn (1993). – type: Hablitzia
tamnoides M. Bieb.
the monotypic genus belongs to the Hablitzieae–Be
toideae (Hohmann & al. 2006; Kadereit & al. 2006).
Hablitzia tamnoides is one of the very few climbing species in Chenopodiaceae. annual shoots grow
from a fleshy root in this species, which is endemic to
caucasus and nW iran.
Halarchon Bunge in Mém. acad. imp. Sci. St.-Pétersbourg, Sér. 7, 4(11): 19, 75. 1862 sec. Kühn (1993).
– type: Halarchon vesiculosus (Moq.) Bunge
Phenotypically this monotypic genus from afghanistan fits into Halimocnemis s.l. However, several attempts to generate sequences for this species have
failed (H. akhani, pers. comm.).
Halimione aellen in Verh. naturf. Ges. Basel 49: 121.
1938 sec. Kadereit & al. (2010). – type: Halimione
pedunculata (L.) aellen
Halimione consists of three species (one annual,
two perennial), which are distributed in europe, the
Mediterranean and western asia. the genus is often
included in Atriplex. Molecular and morphological
data, however, support the generic status of Halimi
one (Kadereit & al. 2010), which is sister to the species-rich Atriplex in the tribe Chenopodieae, formerly
Atripliceae.
Halimocnemis c. a. Mey. in Ledebour, Fl. altaic. 1:
381. 1829 sec. akhani & al. (2007). – type: Halimo
cnemis sclerosperma (Pall.) c. a. Mey.
= Halanthium K. Koch in Linnaea 17: 313. 1844.
= Gamanthus Bunge in Mém. acad. imp. Sci. SaintPétersbourg, Sér. 7, 4(11): 19, 76. 1862.
= Halotis Bunge in Mém. acad. imp. Sci. Saint-Pétersbourg, Sér. 7, 4(11): 19, 73. 1862.
Halimocnemis is an aggregate of irano-turanian
annual species that is phylogenetically not well resolved. Based on phylogenetic studies (akhani & al.
2007), a broad concept was adopted in which Gaman
thus, Halanthium and Halotis are included. Further
phylogenetic studies are required for possible inclusion of genera such as Halarchon, Physandra and
Piptoptera.
Halocharis Moq. in candolle, Prodr. 13(2): 48, 201.
1849 sec. Kühn (1993). – type: Halocharis sulphu
rea (Moq.) Moq.
Halocharis comprises seven annual species and belongs to the Caroxyleae (akhani & al. 2007).
Halocnemum M. Bieb. in Fl. taur.-caucas. 3: 3.
1819 – 1820 sec. Kühn (1993). – type: Halocnemum
strobilaceum (Pall.) M. Bieb.
Halocnemum belongs to Salicornioideae and comprises two hygrohalophytic species of shrubs. the genus is distributed in the southern Mediterranean and
southern, western and west-central asia and is closely
related to Halopeplis and Halostachys (Kadereit & al.
2006).
Halogeton c. a. Mey. in icon. Pl. [Ledebour] 1: 10. 1829
sec. Kühn (1993). – type: Halogeton glomeratus (M.
Bieb.) c. a. Mey.
= Agathophora (Fenzl) Bunge in Mém. acad. imp. Sci.
Saint-Pétersbourg, Sér. 7, 4(11): 19, 92. 1862.
Halogeton belongs to Salsoleae s.str. and is likely
monophyletic (akhani & al. 2007). this eurasian genus, one species of which is also found in the southwestern and partly central United States as a widespread invasive alien, comprises c. five annual and
perennial species and is often found in saline habitats.
Halopeplis Bunge ex Ung.-Sternb. in Vers. Syst. Salicorn.: 102. 1866 sec. Kühn (1993). – type: Halopep
lis nodulosa (delile) Bunge ex Ung.-Sternb.
Halopeplis comprises three species of annual and
perennial hygrohalophytes distributed in the southern
Mediterranean, South africa and southern, western
and central asia. the genus belongs to Salicornieae
and is closely related to Halocnemum and Halosta
chys (Kadereit & al. 2006).
Halostachys c. a. Mey. ex Schrenk in Bull. cl. Phys.Math. acad. Pétersb. 1: 361. 1843 sec. Kühn (1993).
– type: Halostachys caspica (M. Bieb.) c. a. Mey.
ex Schrenk
this monotypic genus of Salicornieae is distributed
in central, southern and western asia and southern
and eastern europe. it is closely related to Halo
cnemum and Halopeplis (Kadereit & al. 2006).
nomenclatural note: Pfeiffer (1874) had chosen
Halostachys songarica Schrenk as the type of Ha
lostachys, but this species was by that time already
placed in the new genus Halopeplis (see Piirainen
2015 for details). a proposal has been published to
Willdenowia 45 – 2015
conserve the name Halostachys with H. caspica as its
conserved type (Piirainen 2015).
Halothamnus Jaub. & Spach, ill. Pl. Orient. 2: 50. 1845
sec. Kothe-Heinrich (1993). – type: Halothamnus
bottae Jaub. & Spach
= Aellenia Ulbr., nat. Pflanzenfam. (ed. 2) 16c: 567.
1934.
Halothamnus belongs to Salsoleae s.str. and is likely
monophyletic (akhani & al. 2007). the genus comprises 21 species, most of which are small shrubs or
subshrubs, only two species are annuals. it is found
from Somalia in the west to Kazakhstan in the east
in desert and semi-desert habitats (Kothe-Heinrich
1993).
Haloxylon Bunge, Fl. ross.: 292. 1852 sec. Kühn (1993).
– type: Haloxylon ammodendron (c. a. Mey.) Bunge
the molecular phylogenetic studies by akhani & al.
(2007) reject a wide interpretation of Haloxylon as
suggested by Hedge (1977).
Hammada iljin in Bot. zhurn. (Moscow & Leningrad)
33: 582. 1948 sec. akhani & al. (2007). – type: Ham
mada leptoclada (Popov) iljin
Generic circumscription of Hammada requires more
studies. the molecular phylogeny of three studied
species showed paraphyly of the studied samples
(akhani & al. 2007).
Heterostachys Ung.-Sternb., atti congr. Bot. Firenze
1874: 267, 268, 331. 1876 sec. Kühn (1993). – type:
Heterostachys ritteriana (Moq.) Ung.-Sternb.
this halophytic genus belongs to the Salicornioideae
and comprises two species in central and South
america. it is closely related to Allenrolfea (Kadereit
& al. 2006).
Holmbergia Hicken in apuntes Hist. nat. 1: 65. 1909
sec. zacharias & Baldwin (2010). – type: Holm
bergia exocarpa (Griseb.) Hicken
a monotypic South american genus of Chenopodieae
(incl. Atripliceae) (Kadereit & al. 2010; zacharias &
Baldwin 2010). it is one of the rare Chenopodiaceae
with berry-like fruits.
Horaninovia Fisch. & c. a. Mey. in enum. Pl. nov. 1: 10.
1841 sec. Kühn (1993). – type: Horaninowia ulicina
Fisch. & c. a. Mey.
= Eremochion Gilli in repert. Spec. nov. regni Veg.
62: 22. 1959.
a likely monophyletic genus within Salsoleae s.str.
comprising six annual species distributed in desert
habitats in central and western asia (akhani & al.
2007).
Iljinia Korovin ex Kom., Fl. UrSS 6: 309, 877. 1936 sec.
Kühn (1993). – type: Iljinia regelii (Bunge) Korovin
ex Kom.
this presumably monotypic genus likely belongs to
Salsoloideae, but has not yet been included in molecular studies.
Kali Mill., Gard. dict. abr., ed. 4: [unpaged]. 1754 sec.
akhani & al. (2007) ≡ Salsola sect. Kali (Mill.) du-
339
mort., Fl. Belg. (dumortier): 23. 1827. – type: Kali
turgidum (dumort.) Guterm.
the genus comprises c. 20 c4 annual species with
spiny leaf tip, except for the shrubby species Kali
griffithii (Bunge) akhani & roalson, an endemic of
southeastern iran, southern afghanistan and adjacent
Pakistan. the genus is native to asia, europe and the
Mediterranean basin and is also widely introduced in
australia, South africa and north america (Brullo &
al. 2015a, b). However, it is likely that some species
are native to australia and north america (see Hrusa
& Gaskin 2008; chinnock 2010). the typification
and nomenclature of Kali are discussed in akhani &
al. (2014). See also under Salsola.
Kalidium Moq. in candolle, Prodr. 13(2): 46, 146. 1849
sec. Kühn (1993). – type: Kalidium foliatum (Pall.)
Moq.
= Kalidiopsis aellen in notes roy. Bot. Gard. edinburgh 28: 31. 1967.
this genus belongs to the Salicornioideae and comprises five perennial halophytic species that are distributed in central and southwestern asia as well as
southern and southeasternmost europe. the monophyly of Kalidium is only weakly supported by molecular data (Kadereit & al. 2006).
Kaviria akhani & roalson in int. J. Pl. Sci. 168: 948.
2007 sec. akhani & al. (2007). – type: Kaviria to
mentosa (Moq.) akhani
= Salsola sect. Belanthera iljin in trudy Bot. inst. nauk
SSSr 1,3: 158. 1937.
this genus belongs to Caroxyleae and includes c. ten
xerohalophytic species mainly distributed in deserts
of central and southwestern asia (akhani & al. 2007).
Krascheninnikovia Gueldenst. in novi comment. acad.
Sci. imp. Petrop. 16: 551. 1772 sec. Kühn (1993). –
type: Krascheninnikovia ceratoides (L.) Gueldenst.
= Eurotia adans., Fam. Pl. 2: 260. 1763, nom. illeg.
Krascheninnikovia, according to a molecular study
(itS phylogeny only) by Heklau & röser (2008),
comprises only one widespread and highly polymorphic species with two subspecies (eurasian and north
american ones). However, considerable morphological diversity and wide geographical distribution (from
southern europe through central asia to southwestern
and west-central north america) of respresentatives
of the genus suggest recognition of several species
and/or subspecies (Grubov 1966; zhu & al. 2003).
the genus belongs to the Axyrideae (Kadereit & al.
2010).
Lagenantha chiov. in Fl. Somala 1: 292. 1929 sec. Kühn
(1993). – type: Lagenantha nogalensis chiov.
the genus likely belongs to Salsoloideae, but has not
yet been included in molecular studies.
Lipandra (Less.) Moq. in chenop. Monogr. enum.: 19.
1840 sec. Fuentes-Bazán & al. (2012b) ≡ Oligandra
Less. in Linnaea 9: 199. 1834, nom. illeg. ≡ Gan
driloa Steud., nomencl. Bot., ed. 2, 1: 662. 1840,
340
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
nom. illeg. ≡ Oliganthera endl., Gen. Pl., Suppl. 1:
1377. 1841, nom. illeg. – type: Lipandra atripli
coides (Less.) Moq.
= Chenopodium [unranked] Polysperma Standl., n.
amer. Fl. 21(1): 13. 1916 ≡ Chenopodium subsect.
Polysperma (Standl.) Kowal ex Mosyakin & clemants in novon 6: 400. 1996.
the isolated lineage of Chenopodium polyspermum
L., revealed in the phylogenetic study of FuentesBazán & al. (2012b), is well supported by the unique
morphological characters of that widespread eurasian species, which led to the creation of a monotypic
subsection within Chenopodium s.l. (Mosyakin &
clemants 1996).
Maireana Moq. in chenop. Monogr. enum.: 95. 1840
sec. Kühn (1993). – type: Maireana tomentosa Moq.
= Austrobassia Ulbr., nat. Pflanzenfam. (ed. 2) 16c:
532. 1934.
= Duriala (r. H. anderson) Ulbr., nat. Pflanzenfam.
(ed. 2) 16c: 537. 1934.
= Eriochiton (r. H. anderson) a. J. Scott in repert.
Spec. nov. regni Veg. 89: 119. 1978.
a species-rich genus of Camphorosmoideae, mostly
endemic to australia (Wilson 1975). Monophyly of
the genus is unclear (cabrera & al. 2009).
Malacocera r. H. anderson in Proc. Linn. Soc. new
South Wales, ser. 2, 51: 382. 1926 sec. Kühn (1993).
– type: Malacocera tricornis (Benth.) r. H. anderson
a small genus endemic to australia and belonging
to Camphorosmoideae (Wilson 1984; cabrera & al.
2009).
Manochlamys aellen in Bot. Jahrb. Syst. 70: 379. 1939
sec. Kadereit & al. (2010). – type: Manochlamys al
bicans (aiton) aellen
a monotypic genus distributed in South africa. it
belongs to the Archiatriplex clade within Chenopo
dieae, formerly Atripliceae (Kadereit & al. 2010).
Microcnemum Ung.-Sternb., atti congr. Bot. Firenze
1874: 268, 269, 280. 1876 sec. Kühn (1993). – type:
Microcnemum fastigiatum (Loscos & J. Pardo) Ung.Sternb.
a monotypic genus in Salicornioideae of rare hygrohalophytic herbs with two subspecies that show a
disjunct distribution in the western and eastern Mediterranean region to central iran (Kadereit & Yaprak
2008).
Microgynoecium Hook. f., Gen. Pl. 3(1): 56. 1880 sec.
Kühn (1993). – type: Microgynoecium tibeticum
Hook. f.
the phylogenetic position of the monotypic Himalayan genus Microgynoecium is in Atripliceae (earlier
Chenopodieae) based on morphological and strong
molecular evidence (Kadereit & al. 2010; FuentesBazan & al. 2012a, b).
Micromonolepis Ulbr., nat. Pflanzenfam. (ed. 2) 16c:
499. 1934 sec. Ulbrich (1934). – type: Micromono
lepis pusilla (torr. ex S. Watson) Ulbr.
a monotypic genus distributed in western north
america and likely closely related to Chenopodium
(Kadereit & al. 2010).
Nanophyton Less. in Linnaea 9: 197. 1834 sec. Kühn
(1993). – type: Nanophyton erinaceum (Pall.)
Bunge
a genus of c. ten closely related xerophytic species
that are distributed in central asia (Pratov 1985). Na
nophyton is related to Halocharis and Kaviria based
on phylogentic studies (akhani & al. 2007).
Neobassia a. J. Scott in Feddes repert. 89: 117. 1978
sec. Kühn (1993). – type: Neobassia astrocarpa (F.
Muell.) a. J. Scott
a small genus endemic to australia and belonging
to Camphorosmoideae (Wilson 1984, cabrera & al.
2009).
Neokochia (Ulbr.) G. L. chu & S. c. Sand. in Madroño
55: 255. 2009 sec. Kadereit & Freitag (2011) ≡ Ko
chia sect. Neokochia Ulbr., nat. Pflanzenfam. (ed.
2) 16c: 535. 1934. – type: Neokochia americana (S.
Watson) G. L. chu & S. c. Sand.
a north american genus of Camphorosmoideae
comprising two closely related species of subshrubs
or dwarf shrubs.
Noaea Moq. in candolle, Prodr. 13(2): 207. 1849 sec.
Kühn (1993). – type: Noaea mucronata (Forssk.)
asch. & Schweinf.
a small genus of three xerophytic c4 species distributed in northern africa eastwards to southwestern and
central asia (akhani & al. 2007).
Nucularia Batt. in Bull. Soc. Bot. France 50: 469. 1903
sec. Kühn (1993). – type: Nucularia perrinii Batt.
Ofaiston raf., Fl. tellur. 3: 46. “1836” [1837] sec. Kühn
(1993). – type: Ofaiston monandrum (Pall.) Moq.
a monotypic genus endemic to southeastern europe,
southwestern Siberia, and central asia. it is closely
related to Petrosimonia (akhani & al. 2007).
Oreobliton durieu in duch. rev. Bot. 2: 428. 1847 sec.
Kühn (1993). – type: Oreobliton thesioides durieu &
Moq. ex durieu
Hohmann & al. (2006) showed that this monotypic
genus belongs to Betoideae–Hablitzieae. Oreobliton
thesioides is a subshrub distributed on calcareous
rocks in algeria and tunisia. together with its sister
genus Aphanisma it represents an interesting example of a western eurasian–western north american
disjunction (Kadereit & Baldwin 2012).
Osteocarpum F. Muell. in trans. Phil. inst. Vict. 2: 77.
1858 sec. Mueller (1858). – type: Osteocarpum sal
suginosum F. Muell.
= Babbagia F. Muell., rep. Pl. Babbage’s exped.: 21.
1859.
a small genus endemic to australia and belonging to
Camphorosmoideae (cabrera & al. 2009).
Oxybasis Kar. & Kir. in Bull. Soc. imp. naturalistes Moscou: 738. 1841 sec. Fuentes-Bazán & al. (2012b). –
type: Oxybasis minutiflora Kar. & Kir.
Willdenowia 45 – 2015
= Blitum subg. Pseudoblitum (Gren. & Godr.) Schur,
enum. Pl. transsilv.: 571. 1866 ≡ Chenopodium sect.
Pseudoblitum (Gren. & Godr.) Syme, engl. Bot., ed.
3, 8: 20. 1868 ≡ Chenopodium [unranked] Rubra
Standl., n. amer. Fl. 21(1): 29. 1916.
= Chenopodium sect. Pseudoblitum Hook. f. in Benth. &
Hook. f., Gen. Pl. 3: 52. 1880 ≡ Blitum sect. Pseudo
blitum (Hook. f.) Mosyakin in Ukrayins’k. Bot. zhurn.
69(3): 394. 2012 ≡ Oxybasis sect. Pseudoblitum (Hook.
f.) Mosyakin in Phytoneuron 2013-56: 3. 2013.
= Chenopodium [unranked] Glauca Standl., n. amer.
Fl. 21(1): 28. 1916 ≡ Chenopodium subsect. Glauca
(Standl.) a. J. Scott in Bot. Jahrb. Syst. 100: 216. 1978
≡ Chenopodium sect. Glauca ignatov in Sosud. rast.
Sovet. dal’nego Vostoka 3: 22. 1988 ≡ Oxybasis sect.
Glaucae (Standl.) Mosyakin in Phytoneuron 201356: 4. 2013 ≡ Blitum sect. Glauca (Standl.) Mosyakin
in Ukrayins’k. Bot. zhurn. 69(3): 395. 2012.
= Chenopodium [unranked] Urbica Standl., n. amer. Fl.
21(1): 11. 1916 ≡ Chenopodium sect. Urbica (Standl.)
Mosyakin in Ukrayins’k. Bot. zhurn. 59: 700. 2002
≡ Oxybasis sect. Urbicae (Standl.) Mosyakin in Phytoneuron 2013-56: 5. 2013.
= Chenopodium sect. Degenia aellen in Magyar Bot.
Lapok 25: 56. 1927.
Oxybasis was described by Karelin & Kirilov
(1841) and included at that time only one species,
O. minutiflora Kar. & Kir. (= Oxybasis chenopodio
ides (L.) S. Fuentes & al.). the phylogenetic studies by Fuentes-Bazán & al. (2012b) and Sukhorukov & al. (2013) supported the monophyly of this
widespread genus as a member of Chenopodieae
and enlarged its circumscription with species segregated from Chenopodium s.l. at least ten species are currently known (some recently transferred
from Chenopodium: see Mosyakin 2013; Sukhorukov 2014), and some occur in saline habitats.
Since Oxybasis contains O. rubra (L.) S. Fuentes &
al., and its basionym, C. rubrum L., is considered by
some authors to be lectotype of Chenopodium, the
adoption of that lectotype would result in Oxybasis
becoming a synonym of Chenopodium (see there).
Patellifolia a. J. Scott & al. in taxon 26: 284. 1977 sec.
Kadereit & al. (2006b) ≡ Beta [unranked] Patellares
tranzschel in trudy Prikl. Bot. Selekts. 17: 205. 1927
≡ Patellaria J. t. Williams & al. in Feddes repert. 87:
289. 1976, nom. illeg. – type: Patellifolia webbiana
(Moq.) a. J. Scott & al.
according to Kadereit & al. (2006) Patellifolia is a
separate genus, more closely related to Habliztia than
to Beta. according to thulin & al. (2010) Patellifo
lia includes only one polymorphic species within a
wide Macaronesian–Mediterranean distribution and a
small disjunct eastern african population.
Petrosimonia Bunge in Mém. acad. imp. Sci. St.-Pétersbourg, Sér. 7, 4(11): 19, 52. 1862 sec. Kühn (1993).
– type: Petrosimonia monandra (Pall.) Bunge
341
a genus of c. 12 species distributed in saline soils of
central and southwestern asia, westwards to the eastern Mediterranean. Petrosimonia is a typical genus
with bifurcate hairs. it forms a monophyletic group
with Ofaiston within Caroxyleae (akhani & al. 2007).
Physandra Botsch. in Sborn. Geobot. akad. Sukachev:
114. 1956 sec. Botschantzev (1956). – type: Physan
dra halimocnemis (Botsch.) Botsch.
Physandra has not yet been included in any phylogentical study. it is presumably a member of Hali
mocnemis s.l. (akhani & al. 2007).
Piptoptera Bunge in trudy imp. S.-Peterburgsk. Bot.
Sada 5: 644. 1877 sec. Kühn (1993). – type: Pipto
ptera turkestana Bunge
See notes under Halimocnemis on the possible synonymy of Piptoptera.
Proatriplex (W. a. Weber) Stutz & G. L. chu in amer. J.
Bot. 77: 366. 1990 sec. zacharias & Baldwin (2010)
≡ Atriplex subg. Proatriplex W. a. Weber in Madroño
10: 189. 1950. – type: Proatriplex pleiantha (W. a.
Weber) Stutz & G. L. chu
Pyankovia akhani & roalson in int. J. Pl. Sci. 168(6):
949. 2007 sec. akhani & al. (2007). – type: Pyan
kovia brachiata (Pall.) akhani & roalson
Pyankovia is a recent segregate of Climacoptera and
Salsola s.l.; it was initially described as a monotypic
genus (akhani & al. 2007). Further studies showed
that the genus contains more than one species (Wen
& al. 2010). there are probably at least three species
distributed from southeasternmost europe through
the caspian area, the caucasus, and iran to central
asia (S. Mosyakin, unpubl. data).
Rhaphidophyton iljin in acta inst. Bot. acad. Sc.
UrSS, Ser. i, 3: 157. 1936 sec. Kühn (1993). – type:
Rhaphidophyton regelii (Bunge) iljin
a monotypic genus from central asia belonging to
the tribe Salsoleae of Salsoloideae (akhani & al.
2007).
Roycea c. a. Gardner in J. roy. Soc. Western australia
32: 77. 1948 sec. Kühn (1993). – type: Roycea pyc
nophylloides c. a. Gardner
a small genus comprising three species, endemic to
australia (Wilson 1984).
Salicornia L., Sp. Pl. 1: 3. 1753 sec. Kadereit & al.
(2006a). – type: Salicornia europaea L.
the genus is distributed worldwide (except for australia and South america) in salt marshes and saline
inland habitats and consists of annual species. it is
nested within the perennial Sarcocornia. For recent
molecular and morphological studies see Kadereit &
al. (2007, 2012), akhani (2008), teege & al. (2011),
Slenzka & al. (2013) and Steffen & al. (2015).
Salsola L., Sp. Pl. 1: 222. 1753. – type: Salsola soda L.
= Salsola sect. Coccosalsola Fenzl. in Ledeb. Fl. ross.
3,2: 802. 1851 ≡ Salsola sect. Coccosalsola Fenzl
subsect. Coccosalsola (Fenzl.) Botsch. p.p. in nov.
Sist. Vys. rast. 13: 94. 1976;
342
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
= Soda Fourr. in ann. Soc. Linn. Lyon sér. 2, 17: 145.
1869;
= Seidlitzia Bunge ex Boiss., Fl. Orient. 4: 950. 1879;
= Hypocylix Wol. in denkschr. Kaiserl. akad. Wiss.,
Wien. Math.-naturwiss. Kl. 51: 275. 1886;
= Darniella Maire & Weiller in Bull. Soc. Hist. nat. afrique n. 30: 301. 1939;
= Fadenia aellen & c. c. towns. in Kew Bull. 27: 501.
1972;
= Salsola sect. Obpyrifolia Botsch. & akhani in Bot.
zhurn. 74(11): 1664. 1989;
= Neocaspia tzvelev in Ukrayins’k. Bot. zhurn. 50(1):
81. 1993.
Salsola s.l. was a heterogenous and polyphyletic
complex, which has been split into at least ten lineages based on nuclear and chloroplast markers (akhani
& al. 2007; Pyankov & al. 2001; Kadereit & Freitag
2013). Caroxylon as the largest group, and Climaco
ptera, Kaviria and Pyankovia were transferred to the
tribe Caroxyleae (Caroxyloneae). Several other segregates have either been described as new genera or
were resurrected from existing names, including Kali,
Turania and Xylosalsola. three names were informally mentioned: “Canarosalsola”, “Collinosalsola” and
“Oreosalsola”, the last soon to be formally published
(akhani & Khoshravesh, in press). the two species,
S. webbii Moq. and S. genistoides Juss. ex Poir., are
sister of Salsoleae and therefore should be described
as separate genera (Voznesenskaya & al. 2013).
the typification of the genus Salsola is debated
(akhani & al. 2014; Mosyakin & al. 2014), and a
conserved type, S. kali L., is proposed instead of the
current type, S. soda (Mosyakin & al. 2014). if accepted, the name Salsola L. will replace Kali Mill.
and Salsola sensu akhani & al. will be Soda Fourr. in
its present circumscription accepted here, Salsola is
still a morphologically very diverse group that probably deserves further splitting into several more natural genera, following more comprehensive molecular
and morphological studies.
Sarcocornia a. J. Scott in J. Linn. Soc., Bot. 75: 366.
1978 sec. Kadereit & al. (2006a) ≡ Salicornia sect.
Perennes duval-Jouve ex Moss in J. Bot. 49: 178.
1911. – type: Sarcocornia perennis (Mill.) a. J. Scott
= Salicornia subg. Arthrocnemoides Ung.-Sternb. in
Versuch einer Systematik der Salicornieen: 54. 1866.
Sarcocornia belongs to Salicornieae and comprises
c. 28 species of perennial, stem-succulent halophytes
distributed worldwide (alonso & crespo 2008; Steffen & al. 2010; de la Fuente & al. 2013). the genus is
paraphyletic with respect to Salicornia. a wordwide
molecular phylogenetic study shows the multiple parallel evolution of prostrate, mat-forming habits (Steffen & al. 2015).
Sclerochlamys F. Muell. in trans. Phil. inst. Vict. 2: 76.
1858 sec. Mueller (1858). – type: Sclerochlamys
brachyptera F. Muell.
a small genus belonging to the Camphorosmoideae
and endemic to australia (Wilson 1984; cabrera &
al. 2009).
Sclerolaena r. Br., Prodr. Fl. nov. Holland.: 410. 1810
sec. Kühn (1993). – type: Sclerolaena uniflora r. Br.
= Cyrilwhitea ising in trans. roy. Soc. South australia
88: 1964. 1964.
= Stelligera a. J. Scott in repert. Spec. nov. regni Veg.
89: 114. 1978.
a species-rich genus of Camphorosmoideae, endemic to australia. Monophyly of the genus remains
unclear (Wilson 1980; cabrera & al. 2009); further
studies are needed.
Sedobassia Freitag & G. Kadereit in taxon 60: 72. 2011
sec. Kadereit & Freitag (2011). – type: Sedobassia
sedoides (Pall.) Freitag & G. Kadereit
this monotypic genus belongs to Camphorosmoide
ae, with an annual species distributed from Hungary
to southern Siberia and showing a c3/c4 intermediate
photosynthetic pathway (Kadereit & al. 2014). the illegitimate name Salsola sedoides Pall. (the basionym
of Sedobassia sedoides) was proposed for conservation against Salsola sedoides L. (Freitag & Sennikov
2014). if this proposal is accepted, the name Sedobas
sia sedoides (Pall.) Freitag & G. Kadereit will remain
in use.
Sevada Moq. in candolle, Prodr. 13(2): 47, 154. 1849
sec. Kühn (1993). – type: Sevada schimperi Moq.
a monotypic african genus.
Spinacia L., Sp. Pl. 1: 1027. 1753 sec. Kühn (1993). –
type: Spinacia oleracea L.
the small eurasian genus Spinacia is supported as
monophyletic and sister to Blitum, both genera belonging to the tribe Anserineae dumort. (FuentesBazán & al. 2012a).
Spirobassia Freitag & G. Kadereit in taxon 60: 71. 2011
sec. Kadereit & Freitag (2011). – type: Spirobassia
hirsuta (L.) Freitag & G. Kadereit
a monotypic genus in Camphorosmoideae comprising an annual species distributed from the northern
Mediterranean to southern Siberia.
Stutzia e. H. zacharias in Syst. Bot. 35: 851. 2010 sec.
zacharias & Baldwin (2010). – type: Stutzia dioica
(nutt.) e. H. zacharias
= Endolepis torr. in Pacif. rail. rep. 12: 47. 1860,
nom. illeg.
Suaeda Forssk. ex J. F. Gmel. in Onomat. Bot. compl. 8:
797. 1776, nom. cons. sec. Kapralov & al. (2006). –
type: Suaeda vera Forssk. ex J. F. Gmel.
= Alexandra Bunge in Linnaea 17: 120. 1843.
= Brezia Moq. in candolle, Prodr. 13(2): 47. 1849.
= Calvelia Moq. in candolle, Prodr. 13(2): 47. 1849.
= Helicilla Moq. in candolle, Prodr. 13(2): 47, 169.
1849.
= Borsczowia Bunge in trudy imp. S.-Peterburgsk.
Bot. Sada 5: 643. 1877.
Molecular phylogenetic studies clearly show that
Willdenowia 45 – 2015
Alexandra and Borsczowia should be included in a
monophyletic Suaeda (Kapralov & al. 2006), despite
the arguments by Lomonosova & Freitag (2011), who
preferred a paraphyletic Suaeda by keeping Alexan
dra as a separate genus. the study by Schütze & al.
(2003) is currently the most comprehensive molecular and morphological study of the genus. the pollen
morphology of Suaeda was studied by dehghani &
akhani (2009).
Suckleya a. Gray in Proc. amer. acad. arts 11: 103. 1876
sec. Kühn (1993). – type: Suckleya petiolaris a. Gray
the monotypic north american genus Suckleya belongs to the Dysphanieae, in which it is sister to Cy
cloloma and Dysphania (Kadereit & al. 2010).
Sympegma Bunge in Bull. acad. imp. Sci. Saint-Pétersbourg 25: 351, 371. 1879 sec. Kühn (1993). – type:
Sympegma regelii Bunge
Tecticornia Hook. f., Gen. Pl. 3(1): 65. 1880 sec. Shepherd & Wilson (2007). – type: Tecticornia cinerea (F.
Muell.) Baill.
= Pachycornia Hook. f., Gen. Pl. 3(1): 65. 1880.
= Halosarcia Paul G. Wilson in nuytsia 3: 28. 1980.
= Sclerostegia Wilson in nuytsia 3: 17. 1980.
= Tegicornia Paul G. Wilson in nuytsia 3: 25. 1980.
a genus of Salicornioideae with c. 25 hygrohalophytic species, largely endemic to australia.
Teloxys Moq. in ann. Sci. nat., Bot., ser. 2, 1: 289.
1834 sec. Fuentes-Bazán & al. (2012b) ≡ Chenopo
dium sect. Teloxys (Moq.) Beck, icon. Fl. Germ.
Helv. (reichenbach) 24: 116. 1908 ≡ Chenopodium
[unranked] Aristata Standl., n. amer. Fl. 21(1): 25.
1916 ≡ Chenopodium subsect. Teloxys (Moq.) aellen
& iljin, Fl. UrSS 6: 47. 1936 ≡ Dysphania subsect.
Teloxys (Moq.) Mosyakin & clemants in Ukrayins’k.
Bot. zhurn. 59: 383. 2002. – type: Teloxys aristata
(L.) Moq.
Since the treatment of Beck (1907 – 1909), Teloxys
was included and mostly accepted in Chenopodium
subsect. Teloxys. For the Flora of north america,
Mosyakin & clemants (2002) transfered this species to Dysphania. However, the phylogenetic studies of Kadereit & al. (2010) and Fuentes-Bazán & al.
(2012a) recovered an isolated position of the monotypic Teloxys, supporting its first circumscription
(Moquin 1834) and also revealing its close relationship to Cycloloma, Dysphania and Suckleya.
Threlkeldia r. Br., Prodr. Fl. nov. Holland.: 409. 1810
sec. Kühn (1993). – type: Threlkeldia diffusa r. Br.
a small genus belonging to the Camphorosmoideae
and endemic to australia (Wilson 1984; cabrera &
al. 2009).
Traganopsis Maire & Wilczek in Bull. Soc. Hist. nat.
afrique n. 27: 67. 1936 sec. Kühn (1993). – type:
Traganopsis glomerata Maire & Wilczek
Traganum delile, descr. Égypte, Hist. nat. 2: 204. 18131814 sec. Kühn (1993). – type: Traganum nudatum
delile
343
Turania akhani & roalson, int. J. Pl. Sci. 168: 946.
2007 sec. akhani & al. (2007). – type: Turania sog
diana (Bunge) akhani
= Salsola sect. Androssowia rilke in Biblioth. Bot. 149:
77. 1999.
= Salsola sect. Sogdiana (iljin) rilke in Biblioth. Bot.
149: 69. 1999.
a small segregate genus of the Salsola s.l. complex,
with three species endemic to aralo-caspian sandy
deserts.
Xylosalsola tzvelev in Ukrayins’k. Bot. zhurn. 50: 81.
1993 sec. akhani & al. (2007). – type: Xylosalsola
arbuscula (Pall.) tzvelev
a segregate genus of Salsola s.l. consisting of small
or large shrubs occurring in sandy or gravelly habitats of the central asian and iranian deserts (tzvelev
1993; akhani & al. 2007).
Didiereaceae radlk. sec. aPG (2009).
a family with six genera and 20 species (Bruyns & al.
2014). traditionally, Didiereaceae included xerophytic
shrubs and trees endemic to Madagascar with short lateral shoots bearing spines or alternate leaves (Kubitzki
1993a; cuénoud 2003). However, molecular phylogenetic studies (applequist & Wallace 2001, 2003; nyffeler &
eggli 2010a; Bruyns & al. 2014) showed a well-supported clade including the traditional Didiereaceae plus the
african genera Calyptrotheca, Ceraria and Portulacaria,
previously placed in Portulacaceae. this expanded circumscription of the family is accepted here, which includes also much-branched plants with opposite leaves
and without spines. applequist & Wallace (2003) divided
the family into three subfamilies: Calyptrothecoideae,
Didiereoideae (= traditional Didiereaceae) and Portu
lacarioideae. the recent molecular phylogeny of Bruyns
& al. (2014) supports the monophyly of these subfamilies
and the inclusion of Ceraria within Portulacaria.
Alluaudia (drake) drake in Bull. Mus. Hist. nat. (Paris)
9: 37. 1903 sec. Kubitzki (1993a) ≡ Didierea sect.
Alluaudia drake in compt. rend. Hebd. Séances
acad. Sci. 133: 240. 1901. – type: Alluaudia procera
(drake) drake – Fig. 4e.
Alluaudiopsis Humbert & choux in compt. rend. Hebd.
Séances acad. Sci. 199: 1651. 1934 sec. Kubitzki
(1993a). – type: Alluaudiopsis fiherenensis Humbert
& choux
Calyptrotheca Gilg in Bot. Jahrb. Syst. 24: 307. 1897
sec. carolin (1993). – type: Calyptrotheca somalen
sis Gilg
Decarya choux in compt. rend. Hebd. Séances acad.
Sci. 188: 1620. 1929 sec. Kubitzki (1993a). – type:
Decarya madagascariensis choux
Didierea Baill. in Bull. Mens. Soc. Linn. Paris 1-2: 258.
1880 sec. Kubitzki (1993a). – type: Didierea mada
gascariensis Baill.
Portulacaria Jacq. in coll. 1: 160. 1787 sec. carolin
(1993). – type: Portulacaria afra Jacq. – Fig. 4F.
344
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
= Ceraria Pearson & Stephens in ann. S. african Mus.
9: 32. 1912.
Dioncophyllaceae airy Shaw sec. aPG (2009).
a small family of woody lianas comprising three monotypic genera endemic to the Guineo-congolian rainforest
(Poremski & Barthlott 2003). the family is characterized
by leaves with grapnels on branches or paired at the leaf
apex, elongated funicles and large discoid and winged
seeds (Heubl & al. 2006). the family is considered as
partially carnivorous because it includes both carnivorous
(Triphyophyllum) and non-carnivorous taxa (Dionco
phyllum and Habropetalum). the studies by Heubl & al.
(2006) and renner & Specht (2011) concluded that within
Dioncophyllaceae occurred a partial secondary loss of
carnivory. See also notes under Ancistrocladaceae.
Dioncophyllum Baill. in Bull. Mens. Soc. Linn. Paris
1–2: 870. 1890 sec. Porembski & Barthlott (2003). –
type: Dioncophyllum thollonii Baill.
Habropetalum airy Shaw in Kew Bull. 1951: 334. 1952
sec. Porembski & Barthlott (2003). – type: Habro
petalum dawei (Hutch. & dalziel) airy Shaw
Triphyophyllum airy Shaw in Kew Bull. 1951: 341.
1952 sec. Porembski & Barthlott (2003). – type: Tri
phyophyllum peltatum (Hutch. & dalziel) airy Shaw
– Fig. 4G.
Droseraceae Salisb. sec. aPG (2009).
the family includes perennial or annual carnivorous herbs
and sometimes submerged aquatics (Kubitzki 2003b)
characterized by having perception of tactile and chemical stimuli, leaf blade and tentacle movement and genetically by a loss of the rpl2 intron (Heubl & al. 2006). the
family comprises three genera, two of them monotypic:
Aldrovanda distributed in eurasia, southeastern africa
and northeastern australia, and Dionaea endemic to the
southeastern United States. Drosera is cosmopolitan and
comprises probably more than 100 species (Kubitzki
2003b; rivadavia & al. 2003). the family is well known
to attract, capture, retain and digest small prey animals
(mainly small arthropods) with active snap-traps (Aldro
vanda [waterwheel plant] and Dionaea [Venus flytrap])
or with active sticky flypaper traps (Drosera [= sundews])
and to absorb the resulting nutrients (Poppinga 2013).
the relationships of Droseraceae to the other carnivorous
families of the Caryophyllales remain unclear; the results
of several molecular phylogenetic studies resulted in three
main hypotheses: Droseraceae as sister of Nepenthaceae
(e.g. nandi & al. 1998: rbcL; cuénoud & al. 2000; Brockington & al. 2009: combined nuclear and plastid data;
Schäferhoff & al. 2009: petD); Droseraceae as sister of a
clade including Drosophyllaceae + [Ancistrocladaceae +
Dioncophyllacae] (e.g. Schäferhoff & al. 2009: petD) and
Droseraceae as sister of the rest of the carnivorous families (e.g. Meimberg & al. 2000: partial matK; Schäferhoff
& al. 2009: complete matK; renner & Specht 2011: combined nuclear, ribosomal and plastid data).
Aldrovanda L., Sp. Pl. 1: 281. 1753 sec. Kubitzki
(2003b). – type: Aldrovanda vesiculosa L.
Dionaea J. ellis in nova acta regiae Soc. Sci. Upsal.,
ser. 2, 1: 98. 1773 sec. Kubitzki (2003b). – type: Dio
naea muscipula J. ellis
Drosera L., Sp. Pl. 1: 281. 1753 sec. Kubitzki (2003b). –
type: Drosera rotundifolia L. – Fig. 5a & B.
= Sondera Lehm., nov. Stirp. Pug. 8: 44. 1844.
= Freatulina chrtek & Slavíková in Čas. nár. Mus.,
Odd. Přír. 165: 140. 1996.
Drosera has a worldwide distribution, but the majority of species are found in the southern hemisphere,
especially in southwestern australia and new zealand (Kubitzki 2003b; rivadavia & al. 2003). Several
classifications have been proposed for the genus; the
last one was that by Seine & Barthlott (1994), who
recognized three subgenera and 11 sections based on
morphological, anatomical, palynological and cytotaxonomical characters; the molecular phylogenetic
study that included the most representative subgenera and sectional sampling so far (i.e. rivadavia &
al. 2003) supported the monophyly of only some of
these groups.
Drosophyllaceae chrtek & al. sec. aPG (2009).
a monotypic family that includes carnivorous subshrubs distributed in Spain, Portugal and Morocco (Kubitzki 2003c). these are characterized by reverse circinate leaves, basal placentation, polyporate pollen and
a chromosome base number x = 6 (Heubl & al. 2006).
Historically, the single genus Drosophyllum was placed
within Droseraceae, but its position as an independent
lineage has been well supported by several molecular phylogenetic studies (e.g. Meimberg & al. 2000;
cuénoud & al. 2002; Hilu & al. 2003; Brockington &
al. 2009; Schäferhoff & al. 2009). these studies also
revealed the closer relationship of Drosophyllacae with
the clade Ancistrocladaceae + Dioncophyllaceae rather
than Droseraceae.
Drosophyllum Link in neues J. Bot. 1(2): 53. 1805 sec.
Kubitzki (2003c). – type: Drosophyllum lusitanicum
(L.) Link
Frankeniaceae desv. sec. aPG (2009).
a monogeneric family with 70–80 species of halophytic
and xerophytic shrubs, subshrubs and herbs (Whalen
1987; Kubitzki 2003d) distributed throughout the warmer dry regions of the world (Kubitzki 2003d). Kubitzki
(2003d) recognized two genera: Frankenia and the monotypic Hypericopsis; however in the same year Olson &
al. (2003) supported the inclusion of Hypericopsis within
Frankenia based on wood-anatomical characters. the position of Hypericopsis within the eurasian and australian
clade of Frankenia has also been well supported by the
molecular phylogenetic study of Gaskin & al. (2004).
Frankenia L., Sp. Pl. 1: 331. 1753 sec. Gaskin & al.
(2004). – type: Frankenia laevis L.
Willdenowia 45 – 2015
= Beatsonia roxb. in Beatson, tracts St. Helena: 300.
1816.
= Hypericopsis Boiss., diagn. Pl. Orient. 6: 25. 1846.
= Niederleinia Hieron. in Bol. acad. nac. ci. 3: 218.
1879.
= Anthobryum Phil. in anales Mus. nat. Santiago de
chile 1891: 51. 1891.
345
tenhusz & al. (2014). – type: Kewa salsoloides
(Burch.) christenh.
eight species, distributed in africa and Saint Helena;
checklist of species in christenhusz & al. (2014).
these species were formerly included in Hypertelis
(Molluginaceae), but have been shown to occupy an
isolated position in Caryophyllales (christin & al.
2011).
Gisekiaceae nakai sec. aPG (2009).
Gisekia was excluded from Aizoaceae and raised to family level by nakai (1942). recent molecular studies support the family status and show an isolated position of
Gisekiaceae within core-Caryophyllales (Brockington &
al. 2009; Schäfferhoff & al. 2009; crawley & Hilu 2012;
Bissinger & al. 2014).
Gisekia L., Mant. Pl.: 554, 562. 1771 sec. Gilbert (1993).
– type: Gisekia pharnacioides L.
Gilbert (1993) revised the genus and accepted seven
species; however, Bissinger & al. (2014) found all
species to be polyphyletic and suggested to treat them
as one polymorphic species or species complex, Gise
kia pharnaceoides agg. Gisekia pharnaceoides is a c4
species with atriplicoid Kranz anatomy and nad-Me
biochemical type. the lineage originated in South
africa and presumably migrated along arid areas
of eastern africa during the late Miocene/Pliocene
(Bissinger & al. 2014).
Halophytaceae a. Soriano sec. aPG (2009).
a monotypic family of succulent monoecious herbs,
endemic to semi-deserts of western and southwestern
argentina (Hunziker 1998; Bittrich 1993c; Pozner &
cocucci 2006). For many years the position of the only
species, Halophytum ameghinoi (Speg.) Speg. within
Caryophyllales was uncertain. When the species was
described, it was placed in Aizoaceae and later transferred to Chenopodiaceae (e.g. cronquist 1981). Several
molecular phylogenetic studies have shown that it represents a well-supported independent lineage within the
Portulacineae (Brockington & al. 2009, 2011; nyffeler
& eggli 2010a; Ocampo & al. 2010; arakaki & al. 2011),
but its relationships with the other families in this group
remain uncertain. the most recent phylogenetic study,
based on data from several nuclear and chloroplast markers, supports a close relationship between Halophytum
and Basellaceae and a close relationship of both with Di
diereaceae (anton & al. 2014).
Halophytum Speg. in anales Mus. nac. Buenos aires 7:
152. 1902 sec. Bittrich (1993d). – type: Halophytum
ameghinoi (Speg.) Speg. – Fig. 5c.
Limeaceae Shipunov ex reveal sec. aPG (2009).
a monogeneric family with c. 20 species, distributed
mainly in southern africa with a few species in Sudan,
ethiopia and southern asia (endress & Bittrich 1993).
traditionally, the single genus Limeum was placed in
Molluginaceae. However, the position of the genus as
an independent lineage and its distant placement from
Molluginaceae has been well supported by several molecular studies (Brockington & al. 2009; Schäferhoff &
al. 2009; christin & al. 2011). the family includes herbs
and subshrubs characterized by pseudomonomerous twochambered ovaries (endress & Bittrich 1993).
Limeum L., Syst. nat., ed. 10: 995. 1759 sec. endress &
Bittrich (1993). – type: Limeum africanum L.
Lophiocarpaceae doweld & reveal sec. aPG (2009).
Small family of about six species distributed in africa, mainly in the southwest, and southwestern asia
(endress & Bittrich 1993; rohwer 1993). the family includes the genus Lophiocarpus, previously placed
in Phytolaccaceae subfamily Microteoideae and the
genus Corbichonia, previously placed in Mollugi
naceae. the clade Lophiocarpus + Corbichonia was
first recovered and well supported in the molecular
phylogeny based on matK sequences by cuénoud &
al. (2002). the family was described by doweld and
reveal (2008) and the clade was later confirmed by
Schäferhoff & al. (2009) and Brockington & al. (2011).
the two genera included in Lophiocarpaceae are morphologically very different. While members of Lophiocarpus
are herbs and sometimes suffrutescent, characterized by
flowers in spikes (with five tepals and four stamens) and
achenes (rohwer 1993), members of Corbichonia are
herbs or subshrubs, characterized by flowers in cymes
(with five sepals and several petaloid staminodes and
stamens) and capsules (endress & Bittrich 1993; Boulos
1999; Sukhorukov & Kushunina 2015).
Corbichonia Scop. in intr. Hist. nat.: 264. 1777 sec. endress & Bittrich (1993). – type: Corbichonia decum
bens (Forssk.) exell
Lophiocarpus turcz. in Bull. Soc. imp. naturalistes
Moscou 16: 55. 1843 sec. rohwer (1993a). – type:
Lophiocarpus polystachyus turcz.
Kewaceae christenh. sec. christenhusz & al. (2014).
Monogeneric family segregated from Molluginaceae
(christenhusz & al. 2014) based on results from christin
& al. (2011).
Kewa christenh. in Phytotaxa 181: 240. 2014 sec. chris-
Macarthuriaceae christenh. sec. christenhusz & al.
(2014).
a monogeneric family restricted to australia. the poorly
known genus Macarthuria has been shown to be sister to
346
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
all core Caryophyllales (Brockington & al. 2011; christin & al. 2011) and a separate family Macarthuriaceae
was proposed (christenhusz & al. 2014).
Macarthuria Hügel ex endl., enum. Pl.: 11. 1837 sec.
endress & Bittrich (1993). – type: Macarthuria aus
tralis Hügel ex endl.
about ten species of rush-like shrubs from australia,
especially southwestern australia.
Microteaceae Schäferh. & Borsch sec. aPG (2009).
a monogeneric family restricted to the neotropics and
distributed from central america and the antilles to
South america (rohwer 1993; Schäferhoff & al. 2009).
Based mainly on the presence of single-ovuled ovaries,
nowicke (1969) placed Microtea, together with Lophio
carpus, in Phytolaccaceae subfamily Microteoideae.
However, Schäferhoff & al. (2009) showed that these two
genera are not closely related and the position of Micro
tea as an independent lineage was well supported, resulting in the description of the new family. these results
were later confirmed by Brockington & al. (2011).
Microtea Sw., Prodr. [O. P. Swartz]: 4, 53. 1788 sec.
rohwer (1993a). – type: Microtea debilis Sw.
a poorly studied genus of annual herbs from central
and South america and the antilles. the number of
species is estimated to c. 12 (Schäferhoff & al. 2011);
a modern monograph is lacking. Microtea was found
in an isolated phylogenetic position (Schäferhoff &
al. 2011; two species have been sampled).
Molluginaceae Bartl. sec. aPG (2009).
a family with nine genera and c. 90 species mainly distributed in southern africa, but also found in the tropics around the world. the circumscription has been
problematic and some of the taxa formerly assigned to
Molluginaceae are now considered as members of other
families (especially Aizoaceae and Phytolaccaceae) or
as independent families within the Caryophyllales (e.g.
Kewaceae, Limeaceae, Lophiocarpaceae) (endress &
Bittrich 1993; Schäferhoff & al. 2009; christin & al.
2011; christenhusz & al. 2014). the family as currently
circumscribed is characterized by an undifferentiated
perianth with alternitepalous stamens, except for Glinus,
which occasionally has small petals (Brockington & al.
2013).
Adenogramma rchb. in iconogr. Bot. exot. 2: 3. 1828
sec. endress & Bittrich (1993). – type: Adenogram
ma mollugo rchb.
Coelanthum e. Mey. ex Fenzl in ann. Wiener Mus.
naturgesch. 1: 353. 1836 sec. endress & Bittrich
(1993). – type: Coelanthum grandiflorum e. Mey. ex
Fenzl
Glinus L., Sp. Pl. 1: 463. 1753 sec. endress & Bittrich
(1993). – type: Glinus lotoides L.
Glischrothamnus Pilg. in Bot. Jahrb. Syst. 40: 396. 1908
sec. endress & Bittrich (1993). – type: Glischro
thamnus ulei Pilg.
Hypertelis e. Mey. ex Fenzl in ann. Wiener Mus. naturgesch. 1: 352. 1836 sec. endress & Bittrich (1993).
– type: Hypertelis spergulacea e. Mey. ex Fenzl
Monotypic genus (after segregation of Kewa, see
there), distributed in namibia. Might belong to an
expanded Mollugo (christin & al. 2011), but further
study is needed to clarify this.
Mollugo L., Sp. Pl. 1: 89. 1753 sec. endress & Bittrich
(1993). – type: Mollugo verticillata L.
recent phylogenetic analysis has shown that the genus is not monophyletic and that its species are scattered across the Molluginaceae phylogeny (christin
& al. 2011). a thorough re-evaluation of the circumscription of Mollugo is clearly needed.
Pharnaceum L., Sp. Pl. 1: 272. 1753 sec. endress & Bittrich (1993). – type: Pharnaceum incanum L.
Polpoda c. Presl in Polpoda: 1 – 2. 1829 sec. endress &
Bittrich (1993). – type: Polpoda capensis c. Presl
Psammotropha eckl. & zeyh. in enum. Pl. afric. austral. [ecklon & zeyher]: 286. 1836 sec. endress &
Bittrich (1993). – type: Psammotropha parvifolia
eckl. & zeyh.
Suessenguthiella Friedrich in Mitt. Bot. Staatssamml.
München 2: 60. 1955 sec. endress & Bittrich (1993).
– type: Suessenguthiella scleranthoides (Sond.)
Friedrich
Montiaceae raf. sec. aPG (2009).
a family with 13 genera and around 200 species distributed around the world (nyffeler & eggli 2010a). the species of this family are traditionally considered as members of Portulacaceae; however, molecular phylogenetic
studies have shown that the traditional Portulacaceae are
not monophyletic (Hershkovitz & zimmer 1997; applequist & Wallace 2001; nyffeler 2007; nyffeler & eggli
2010a; Ocampo & columbus 2010). nyffeler & eggli
(2010a) proposed the segregation of the traditional Por
tulacaceae into four families (Anacampserotaceae, Mon
tiaceae, Portulacaceae and Talinaceae) based on morphological and molecular data. the circumscription of
Montiaceae follows the proposal of Hershkovitz (1993,
2006) and Hershkovitz & zimmer (2000). Montiaceae
also includes Hectorellaceae (applequist & al. 2006;
Wagstaff & Hennion 2007; nyffeler & eggli 2010a).
Calandrinia Kunth, nov. Gen. Pl. (folio ed.) 6: 77. 1823,
nom. cons. sec. Hershkovitz (1993). – type: Calan
drinia caulescens Kunth
= Baitaria ruíz & Pav., Fl. Peruv. Prodr.: 63. 1823.
= Monocosmia Fenzl, nov. Stirp. dec.: 84. 1839.
Calyptridium nutt. in Fl. n. amer. 1: 198. 1838 sec. nyffeler & eggli (2010a). – type: Calyptridium monan
drum nutt.
Calyptridium is a north american genus with eight
species (Guilliams 2009). although Hershkovitz
(1990) treated Calyptridium as a section of Cistanthe,
phylogenetic analyses have shown that this consideration makes Cistanthe a non-monophyletic group
Willdenowia 45 – 2015
(Hershkovitz & zimmer 2000; applequist & Wallace
2001; Hershkovitz 2006).
Cistanthe Spach in Hist. nat. Vég. 5: 229. 1836 sec. nyffeler & eggli (2010a). – type: not designated.
= Spraguea torr. in Smithsonian contr. Knowl. 6(2): 4.
1853.
= Diazia Phil., Fl. atacam.: 22. 1860.
= Silvaea Phil., Fl. atacam.: 21. 1860.
= Philippiamra Kuntze, revis. Gen. Pl. 2: 58. 1891,
nom. illeg.
= Lewisiopsis Govaerts, World checkl. Seed Pl. 3: 21.
1999.
See under Calyptridium (Montiaceae).
Claytonia L., Sp. Pl. 1753 1: 204. 1753 sec. Miller &
chambers (2006) ≡ Claytonia [unranked] Euclayto
nia Walp., repert. Bot. Syst. 2: 237. 1843, nom. inval.
≡ Claytonia [unranked] Cormosae a. Gray in Proc.
amer. acad. arts 22: 278. 1887 ≡ Claytonia sect.
Cormosae a. Gray ex Poelln. in repert. Spec. nov.
regni Veg. 30: 281. 1932, nom. superfl. – type: Clay
tonia virginica L. – Fig. 5d.
= Limnia Haw., Syn. Pl. Succ.: 11. 1812 ≡ Claytonia
sect. Limnia (Haw.) torr. & a. Gray, Fl. n. amer. 1:
199. 1838 ≡ Montia sect. Limnia (Haw.) B. L. rob.,
Syn. Fl. n. amer. 1: 273. 1897 ≡ Claytonia subg.
Limnia (Haw.) Holub in Preslia 47: 328. 1975.
= Claytonia [unranked] Caudicosae a. Gray in Proc.
amer. acad. arts 22: 279. 1887.
= Claytonia [unranked] Rhizomatosae a. Gray in Proc.
amer. acad. arts 22: 280. 1887 ≡ Montia [unranked]
Rhizomatosae (a. Gray) B. L. rob., Syn. Fl. n. amer.
1: 272. 1897 ≡ Claytonia sect. Rhizomatosae (a.
Gray) Poelln. in repert. Spec. nov. regni Veg. 30:
281, 296. 1932.
= Claytonia sect. Chenopodinae Poelln. in repert.
Spec. nov. regni Veg. 30: 280. 1932.
Hectorella Hook. f. in Handb. n. zeal. Fl.: 27. 1864
sec. Philipson (1993). – type: Hectorella caespitosa
Hook. f.
Monotypic; endemic to new zealand (South island).
the taxonomic position of Hectorella remained controversial for a long time and was even treated in a
separate family along with Lyallia (Hectorellaceae;
Philipson & Skipworth 1961). However, phylogenetic
analyses have confirmed that this monotypic genus is
nested in Montiaceae (applequist & al. 2006; Wagstaff & Hennion 2007; nyffeler & eggli 2010a).
Lenzia Phil. in anales Univ. chile 23: 381. 1863 sec.
carolin (1993). – type: Lenzia chamaepitys Phil.
Lewisia Pursh, Fl. amer. Sept. 2: 368. 1814 sec. Hershkovitz & Hogan (2003). – type: Lewisia rediviva
Pursh
= Oreobroma Howell in erythea 1: 31. 1893.
= Erocallis rydb. in Bull. torrey Bot. club 33: 140.
1906.
Lyallia Hook. f., Fl. antarct. 2: 548, t. 122. 1847 sec.
Philipson (1993). – type: Lyallia kerguelensis Hook. f.
347
Monotypic; endemic to the subantarctic Kerguelen
islands. Lyallia kerguelensis was found to be sister to
Hectorella and both are nested in Montiaceae (Wagstaff & Hennion 2007; see also under Hectorella).
Montia L., Sp. Pl. 1: 87. 1753 sec. Miller (2004). – type:
Montia fontana L.
= Crunocallis rydb. in Bull. torrey Bot. club 33: 139.
1906.
= Naiocrene (torr. & a. Gray) rydb. in Bull. torrey
Bot. club 33: 139. 1906.
= Montiastrum rydb., Fl. rocky Mts.: 1061. 1917.
= Limnalsine rydb., n. amer. Fl. 21(4): 295. 1932.
= Mona Ö. nilsson in Bot. not. 119: 266. 1966.
= Neopaxia Ö. nilsson in Bot. not. 119: 469. 1966.
= Maxia Ö. nilsson in Palynol. 7: 359. 1967.
= Claytoniella Jurtzev in Bot. zhurn. (Moscow & Leningrad) 57: 644. 1972.
Montiopsis Kuntze, revis. Gen. Pl. 3(3): 14. 1898 sec.
Hershkovitz (1993). – type: Montiopsis boliviana
Kuntze
Parakeelya Hershk. in Phytologia 84: 101. 1998 sec.
Hershkovitz (1998). – type: Parakeelya ptychosper
ma (F. Muell.) Hershk.
= ?Rumicastrum Ulbr., nat. Pflanzenfam. (ed. 2) 16c:
519. 1934.
Based on phylogenetic analyses (Hershkovitz 1996),
Hershkovitz (1998) transferred 35 australian Calan
drinia species to the new genus Parakeelya. However, the relationships of the species of this genus
within Montiaceae are not well supported (Hershkovitz 1996; Hershkovitz & zimmer 2000), so further studies are needed to evaluate its affinities.
australian botanists still continue to use the name
Calandrinia for species assignable to Parakeelya.
the relationships of the australian genus Rumi
castrum are not clear. it was considered as a genus
closely related to Atriplex (Chenopodiaceae). carolin
(1987) and Hershkovitz (1993) used the name to represent the australian calandrinias (Montiaceae); however, Hershkovitz & zimmer (2000) opted to use the
name Parakeelya for those taxa. Further studies are
required to clarify the correct use of Rumicastrum.
Phemeranthus raf. in Specchio Sci. 1: 86. 1814 sec.
Kiger (2004) ≡ Talinum sect. Phemeranthus (raf.)
dc., Prodr. 3: 356. 1828. – type: Phemeranthus
teretifolius (Pursh) raf.
Species of Phemeranthus were considered as members of Talinum. However, morphological and molecular analyses have shown that c. 30 new World species that have terete to semi-terete leaves represent a
lineage different from Talinum (Talinaceae; carolin
1987; Hershkovitz & zimmer 2000; applequist &
Wallace 2000; nyffeler & eggli 2010a; Ocampo &
columbus 2010).
Schreiteria carolin in Palynol. 3: 330. 1985 sec. carolin (1993). – type: Schreiteria macrocarpa (Speg.)
carolin
348
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
an enigmatic monotypic genus, placed here with
doubts, and not included in any recent analysis.
Nepenthaceae dumort. sec. aPG (2009).
a monogeneric family comprising 120 – 138 species
(McPherson 2009, 2011) native to tropical asia, distributed from Madagascar through indo-Malesia to
new Guinea and new caledonia (Kubitzki 2003e;
Meimberg & Heubl 2006). the family includes
woody climbers or scrambling shrubs and some epiphytes (Kubitzki 2003e) widely known as the carnivorous “pitcher plants”. they are characterized by unisexual flowers, axilar placentation, filaments united
into a column, three- or four-locular ovaries and the
loss of vascularization in glands (Heubl & al. 2006).
the affinities of Nepenthaceae have long been discussed
(Meimberg & al. 2001). traditionally, the family was
placed in the order Nepenthales, either as a monofamilial
order (e.g. takhtajan 1980) or together with Droseraceae
and Sarraceniaceae (e.g. cronquist 1988). the placement
of the family within Caryophyllales was shown by the
early molecular phylogenetic study of nandi & al. (1998).
Several molecular phylogenetic studies have shown (although with moderate support) the close relationship
of Nepenthaceae and Droseraceae (nandi & al. 1998;
cuénoud & al. 2000; Brockington & al. 2009; Schäferhoff & al. 2009; further information under Droseraceae).
another study, based on parsimony analysis of combined
rbcL and matK shows with high support Nepenthes as
sister to the rest of the carnivorous families, whereas the
study of renner & Specht (2011), based on the ML and
Bayesian analysis of the combined data of nuclear, ribosomal and plastid dna, shows also with high support the
relationship of Nepenthaceae with the Drosophyllaceae
+ [Dioncophyllaceae + Ancistrocladaceae] clade.
Nepenthes L., Sp. Pl. 1: 955. 1753 sec. Kubitzki (2003e).
– type: Nepenthes distillatoria L.
= Anurosperma Hallier f. in Beih. Bot. centralbl. 39(2):
162. 1921.
Nyctaginaceae Juss. sec. aPG (2009).
this family comprises c. 30 genera and 300 – 400 species (Bittrich & Kühn 1993; Spellenberg 2003) of trees,
shrub and herbs. these are found in all warmer areas
of the world (douglas & Spellenberg 2010), but mostly in the americas, with two centres of distribution:
arid western north america (southwestern U.S.a. and
northern Mexico) and the neotropics (tropical and subtropical South america and the antilles). Some genera,
such as Boerhavia, Mirabilis and Pisonia, have some
species occurring in the Old World, but some of them
are introduced (Mirabilis), whereas Commicarpus, with
few american species, is most diverse in africa; Phaeo
ptilum is endemic to southwestern africa and Botswana
(Bittrich & Kühn 1993; douglas & Spellenberg 2010).
recently, douglas & Spellenberg (2010), based on the
molecular phylogeny of the family by douglas & Manos
(2007), made some adjustments to Bittrich and Kühn’s
classification of 1993, so that seven tribes were recognized: Boldoeae, Bougainvilleeae, Caribeeae, Colignoni
eae, Leucastereae, Nyctagineae and Pisonieae; the relationship of Caribeeae with the others is unknown since it
is known only from the type. Several genera, especially
those of north america that include the suffrutescent and
herbaceous taxa, have been the focus of interest of various studies. However, most of the taxa distributed in the
neotropics, including the trees and shrubs in the diverse
genera Guapira, Neea and Pisonia, are poorly known.
Abronia Juss., Gen. Pl.: 448. 1789 sec. Galloway (2003).
– type: Abronia californica J. F. Gmel.
Acleisanthes a. Gray in amer. J. Sci. arts ser. 2, 15: 259.
1853 sec. Levin (2002). – type: Acleisanthes crassi
folia a. Gray
= Selinocarpus a. Gray in amer. J. Sci. arts ser. 2, 15:
262. 1853.
= Ammocodon Standl. in J. Wash. acad. Sci. 6: 631.
1916.
Allionia L., Syst. nat., ed. 10, 2: 883, 890, 1361. 1759,
nom. cons. sec. turner (1994) ≡ Wedelia Loefl., iter.
Hispan.: 180. 1758 ≡ Wedeliella cockerell in torreya
9: 166. 1909, nom. illeg. – type: Allionia incarnata
L.
Andradea allemão, Pl. novas Brasil andradea. 1845
sec. Bittrich & Kühn (1993). – type: Andradea flo
ribunda allemão
Anulocaulis Standl. in contr. U. S. natl. Herb. 12: 374.
1909 sec. Hernández-Ledesma & al. (2010). – type:
Anulocaulis eriosolenus (a. Gray) Standl.
Belemia Pires in Bol. Mus. Paraense “emilio Goeldi”,
n. S., Bot. 52: 1. 1981 sec. Bittrich & Kühn (1993). –
type: Belemia fucsioides Pires
Boerhavia L., Sp. Pl. 1: 3. 1753 sec. Bittrich & Kühn
(1993). – type: Boerhavia erecta L.
Boerhavia, with c. 40 species, is distributed in warmtemperate and tropical regions worldwide (Spellenberg 2003) and has been recognized as a natural
group by douglas & Manos (2007). Several authors
(Fay 1980; Spellenberg 2001, 2003) have highlighted
that at the species level this is a taxonomically difficult group due to morphological variation. especially
among annuals of the Sonoran desert and the pantropical B. diffusa Vahl and B. coccinea Mill. complex (Spellenberg 2001, 2003), apparently factors
such as wide dispersal, hybridization and autogamy
have contributed to that variation (Fay 1980; Spellenberg 2001, 2003). the genus is in need of a critical
revision.
Boldoa cav. ex Lag. in Gen. Sp. Pl.: 9. 1816 sec. Bittrich
& Kühn (1993). – type: not designated.
the genus is monotypic, with B. purpurascens cav.
ex Lag. distributed from Mexico and the antilles to
northern South america. along with Cryptocarpus
and Salpianthus, Boldoa is placed within the tribe
Boldoeae (douglas & Spellenberg 2010), and in
Willdenowia 45 – 2015
several treatments (Standley 1911, 1918, 1931; Fay
1980; Pérez & al. 2000; Spellenberg 2001; Hernández-Ledesma & Flores 2003; González 2007) the
genus has been included in the wide concept of the
genus Salpianthus. Here we follow Bittrich & Kühn
(1993) and Harling (2010), who consider them as
separate genera because of differences of the perianth: Boldoa has a campanulate perianth (2 – 3.5 mm
long) with glandular and uncinate hairs, Salpianthus
has a tubular perianth (6 – 7 mm long) with straight
hairs, while Cryptocarpus has a pyriform perianth
(1.5 – 2 mm long). a revision and phylogenetic analysis including all the species of the tribe is necessary
to evaluate the circumscription of the genera.
Bougainvillea comm. ex Juss., Gen. Pl.: 91. 1789, nom.
cons. sec. Bittrich & Kühn (1993). – type: Bougain
villea spectabilis Willd. – Fig. 5e.
Standley and Steyermark (1946) state that Bou
gainvillea contains c. 14 species native to South
america, three of which were cultivated in tropical and subtropical regions of the world. according
to Fay (1980), the genus includes ten species, but
that author argued that artificial selection processes,
hybridization and the spread of clonal variants have
produced a complex pattern of variation only loosely related to any natural group. Gillis (1976) treated
the bougainvilleas of cultivation, considering three
species and one hybrid. the biology, artificial selection as well as the lack of a monographic treatment
make it difficult to determine the current number of
species.
Caribea alain in candollea 17: 113. 1960 sec. Bittrich &
Kühn (1993). – type: Caribea litoralis alain
an endemic genus from cuba that has a unique
morphology among the Nyctaginaceae (douglas &
Spellenberg 2010). Caribea includes compact bushforming taprooted perennials characterized by opposite leaves forming a stipulariform sheath at the
base (Bittrich & Kühn 1993; douglas & Spellenberg
2010). Because the genus is known only from the
type collection, the most recent classification system
for the family (douglas & Spellenberg 2010) included it in its own tribe, Caribeeae. it is awaiting its rediscovery in the field.
Cephalotomandra H. Karst. & triana in nuev. Jen. esp.:
23. 1855 sec. Bittrich & Kühn (1993). – type: Cepha
lotomandra fragrans H. Karst. & triana
Colignonia endl., Gen. Pl.: 311. 1837 sec. Bittrich &
Kühn (1993). – type: Colignonia parviflora (Kunth)
choisy
Commicarpus Standl. in contr. U. S. natl. Herb. 12: 373.
1909 sec. Bittrich & Kühn (1993). – type: Commi
carpus scandens (L.) Standl.
Cryptocarpus Kunth, nov. Gen. Sp. (folio ed.) 2: 150.
“1817” [1818] sec. Bittrich & Kühn (1993). – type:
Cryptocarpus pyriformis Kunth
the genus is monotypic with C. pyriformis restricted
349
to ecuador, Peru, and the Galapagos islands. For further information see notes under Boldoa.
Cuscatlania Standl. in J. Wash. acad. Sci. 13: 437. 1923
sec. Bittrich & Kühn (1993). – type: Cuscatlania vul
canicola Standl.
a monotypic genus, C. vulcanicola is a perennial
herb reported from el Salvador.
Cyphomeris Standl. in contr. U. S. natl. Herb. 13: 428.
1911 sec. Mahrt & Spellenberg (1995). – type: Cy
phomeris gypsophiloides (M. Martens & Galeotti)
Standl.
Grajalesia Miranda in anales inst. Biol. Univ. nac.
México 21: 299. 1951 sec. Bittrich & Kühn (1993). –
type: Grajalesia ferruginea Miranda
Guapira aubl. in Hist. Pl. Guiane: 308. 1775 sec. Bittrich & Kühn (1993). – type: Guapira guianensis
aubl. – Fig. 5F.
= Torrubia Vell., Fl. Flumin.: 139. “1825” [1829].
a neotropical genus with c. 70 species, distributed
from southern Florida to South america and the antilles. it is closely related to Neea, also being dioecious and having fleshy fruits. Both genera form a
complex and their distinctness has been questioned
by several authors (e.g. Standley 1931; Burger 1983;
Pool 2001; douglas & Manos 2007) because they are
distinguished only by the presentation of the stamens,
which are included in Neea and exserted in Guapira.
in the phylogenetic analysis by douglas & Manos
(2007), the two genera form a clade in which both
are paraphyletic; however those authors questioned if
this result was the effect of their sampling (Guapira,
two species; Neea, three species) or whether the paraphyly is due to the lack of resolution between both
genera. Guapira needs a taxonomic revision and also
needs to be evaluated in a phylogenetic analysis that
includes an extensive sampling along with Neea.
Leucaster choisy in candolle, Prodr. 13(2): 457. 1849
sec. Bittrich & Kühn (1993). – type: Leucaster cani
florus (Mart.) choisy
Mirabilis L., Sp. Pl. 1: 177. 1753 sec. Le duc (1995). –
type: Mirabilis jalapa L.
= Oxybaphus L’Hér. ex Willd., Sp. Pl. (ed. 4): 170, 185.
1797.
= Quamoclidion choisy in candolle, Prodr. 13(2): 429.
1849.
= Allioniella rydb. in Bull. torrey Bot. club 29: 687.
1902.
= Hesperonia Standl. in contr. U. S. natl. Herb. 12:
306, 360. 1909.
a genus with 50 – 60 american and one asiatic species. it includes herbs, suffrutescent herbs and subshrubs characterized by the presence of involucres of
accrescent bracts, often connate, which surround one
or more flowers. traditionally the genus was classified into six sections, some of them corresponding
to previously separated genera. Molecular phylogenetic studies, which have mainly been focused on the
350
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
north american species, support the monophyly of
the genus (Levin 2000; douglas & Manos 2007; P.
Hernández-Ledesma & al., unpubl. data) but not the
monophyly of the sections (P. Hernández-Ledesma &
al., unpubl. data). in order to achieve a natural subgeneric classification, the South american species
should be included in the sampling.
Neea ruiz & Pav., Fl. Peruv. Prodr.: 52. 1794 sec. Bittrich & Kühn (1993). – type: Neea verticillata ruiz
& Pav.
Neea shows extensive morphological variation in
habit, leaves, pubescence, inflorescences, flowers and
fruits (Burger 1983). Some authors (e.g. González
2007) have considered it the taxonomically least understood group in the neotropics. Neea seems to be
the most species-rich genus within Nyctaginaceae;
douglas & Spellenberg (2010) mentioned that the
genus has c. 80 species. However, the lack of a revision, along with the morphological variation and
dioecy, has generated many species names (c. 150),
whereas the actual number of species remains uncertain. For further information see notes under Gua
pira.
Neeopsis Lundell in Wrightia 5: 241. 1976 sec. Bittrich
& Kühn (1993). – type: Neeopsis flavifolia (Lundell)
Lundell
Nyctaginia choisy in candolle, Prodr. 13(2): 429. 1849
sec. Bittrich & Kühn (1993). – type: Nyctaginia capi
tata choisy
Okenia Schltdl. & cham. in Linnaea 5: 92. 1830 sec.
Bittrich & Kühn (1993). – type: Okenia hypogaea
Schltdl. & cham.
Phaeoptilum radlk. in abh. naturwiss. Vereins Bremen
8: 435. 1883 sec. Bittrich & Kühn (1993). – type:
Phaeoptilum spinosum radlk.
Pisonia L., Sp. Pl. 1753 1: 1026. 1753 sec. Bittrich &
Kühn (1993). – type: Pisonia aculeata L. – Fig. 5G.
= Ceodes J. r. Forst. & G. Forst., char. Gen. Pl., ed. 2:
141. 1776.
= Calpidia thouars, Hist. Vég. isles austral. afriq.: 37.
1805.
= Rockia Heimerl in Oesterr. Bot. z. 63: 289. 1913.
= Heimerlia Skottsb. in Svensk Bot. tidskr. 30: 738.
1936 ≡ Heimerliodendron Skottsb. in Svensk Bot.
tidskr. 35: 364. 1941.
this genus includes shrubs, trees and woody climbers characterized by stout spines on the stems and coriaceous fruits with stipitate glands. its distribution is
pantropical with a centre of diversity in the neotropics. Molecular phylogenetic studies (e.g. douglas &
Manos 2007; León de la Luz & Levin 2012) supported the monophyly of Pisonia, although the genus was
poorly sampled in both studies. Pisonia has not been
monographed, and the number of species is uncertain;
some treatments considered 40 species (e.g. Spellenberg 2001; deFilipps & Maina 2003; González 2007)
whereas others (e.g. Spellenberg 2003) considered a
range between 10 – 50 species; in the literature there
are numerous accepted and unresolved names.
Pisoniella (Heimerl) Standl. in contr. U. S. natl. Herb.
13: 385. 1911 sec. Bittrich & Kühn (1993) ≡ Pisonia
sect. Pisoniella Heimerl, nat. Pflanzenfam. 3(1b): 29.
1889. – type: Pisoniella arborescens (Lag. & rodr.)
Standl.
Ramisia Glaz. ex Baill. in Bull. Mens. Soc. Linn. Paris
1(88): 697. 1887 sec. Bittrich & Kühn (1993). – type:
Ramisia reclinata Glaz.
Reichenbachia Spreng. in Bull. Soc. Philom. 1823: 54.
1823 sec. Bittrich & Kühn (1993). – type: Reichen
bachia hirsuta Spreng.
Salpianthus Bonpl., Pl. aequinoct. 1(6): 154. 1807 sec.
Bittrich & Kühn (1993). – type: Salpianthus are
narius Bonpl.
the genus includes shrubs with alternate leaves, a
four- or five-lobed tubular petaloid perianth with
straight glandular hairs, three to four long-exserted
stamens and a linear style (Bittrich & Kühn 1993).
three species are recognized following this concept:
S. aequalis Standl., S. arenarius and S. macrodon
tus Standl., all of them with restricted distributions
in Mexico. Salpianthus was assumed to be monophyletic by douglas & Manos (2007); however, only
S. arenarius was included in their study. For further
information see notes under Boldoa.
Tripterocalyx (torr.) Hook. in Hooker’s J. Bot. Kew
Gard. Misc. 5: 261. 1853 sec. Galloway (2003) ≡
Abronia [unranked] Tripterocalyx torr., rep. exped.
rocky Mts.: 96. 1843. – type: Tripterocalyx micran
thus (torr.) Hook.
Physenaceae takht. sec. aPG (2009).
a monogeneric family with two species endemic to
Madagascar (dickison 2003). traditionally, the only
genus Physena was placed in Capparales/Capparaceae
(e.g. Pax & Hoffmann 1936) or Flacourtiaceae (e.g. Perrier de la Bâthie 1946). Later, it was considered as a family of its own and placed in the order Sapindales (e.g.
takhtajan 1980, 1987) and was then even transferred
to the separate order Physenales (e.g. takhtajan 1997).
However, already the early molecular phylogenetic studies of Morton & al. (1997) showed the affinities of Phy
senaceae with Caryophyllales and its close relationship
to Asteropeiaceae. these results were confirmed by subsequent molecular phylogenetic studies (e.g. cuénoud
& al. 2002; Brockington 2009, 2011; Soltis & al. 2011).
the relationship between Asteropeiaceae and Physena
ceae is also supported by wood-anatomical characters.
For further information see notes under Asteropeiaceae.
Physena noronha ex thouars in Gen. nov. Madagasc.:
6. 1806 sec. dickison (2003). – type: Physena mada
gascariensis thouars ex tul.
Phytolaccaceae r. Br. sec. aPG (2009).
this family comprises herbs, trees or lianas dis-
Willdenowia 45 – 2015
tributed mainly in the americas, including the antilles, but with some members distributed in australia and new caledonia. they are characterized by
styloids, elongate crystals, racemes or spikes and four
or five tepals (rohwer 1993a; Stevens 2001 onwards).
the circumscription of the family has long been controversial. Following the treatment by rohwer (1993a),
Phytolaccaceae have been disintegrated step by step according to the results of molecular phylogenetic studies
(e.g. cuénoud & al. 2002; Hilu & al. 2003; Schäferhoff
& al. 2009; Brockington & al. 2011), which have shown
that the subfamilies Agdestioideae, Barbeuioideae and
Microteoideae (sec. rohwer 1993a) are well-supported
independent lineages. therefore, these taxa are now
treated at family level (see further notes under those
families). these studies have also shown that Phytolac
caceae s.l. comprising the subfamilies Phytolaccoideae
and Rivinoideae (sec. rohwer 1993a) are not monophyletic. the most recent study by Brockington & al.
(2011) included most of the genera recognized in these
subfamilies and showed that the Phytolaccoideae (= Phy
tolaccaceae s.str.) represents a well-supported independent lineage, while the support for Rivinoideae is present
but weak. recent studies (J. Petersen, t. Borsch & P.
Hernández-Ledesma, unpubl. data) show that the latter is
probably more closely related to Nyctaginaceae than to
Phytolaccaceae s.str. Rivinaceae have been recognized
as an independent family within Caryophyllales by Stevens (2001 onwards). However, the correct family name
for a clade that includes the genera Petiveria and Rivina
would have to be Petiveriaceae c. agardh (1824) and not
Rivinaceae c. agardh (1824). Both family names were
published in the same work (agardh 1824) but Meissner (1836) included Rivina under Petiveriaceae separate
from Phytolaccaceae. this gives priority to Petiveriace
ae. the taxon has a complicated taxonomic history. in
some early treatments members were classified either
within Phytolaccaceae and distinct from Petiveriaceae
c. agardh (Lindley 1853), or vice versa (e.g. Hutchinson
1959; Brown & Varadarajan 1985), or at an infrafamiliar
or infrageneric level within Phytolaccaceae (e.g. Petiveri
eae, Rivineae, Rivinoideae) (including Petiveria and related genera) (e.g. Heimerl 1889, 1934; rohwer 1993a).
Anisomeria d. don in edinb. n. Phil. Journ. 13: 238.
1832 sec. rohwer (1993a). – type: Anisomeria co
riacea d. don
Ercilla a. Juss. in ann. Sci. nat. (Paris) 25: 11. 1832 sec.
rohwer (1993a). – type: Ercilla volubilis a. Juss.
Gallesia casar. in nov. Stirp. Bras. dec. 5: 43. 1843 sec.
rohwer (1993a). – type: Gallesia scorodendrum
casar.
Hilleria Vell. in Fl. Flumin.: 47. 1829 sec. rohwer
(1993a). – type: Hilleria elastica Vell.
= Mohlana Mart., nov. Gen. Sp. Pl. 3: 170. 1829.
Ledenbergia Klotzsch ex Moq. in candolle, Prodr. 13(2):
4, 14. 1849 sec. rohwer (1993a). – type: Ledenber
gia seguierioides Klotzsch ex Moq.
351
= Flueckigera Kuntze, revis. Gen. Pl. 2: 550. 1891.
Monococcus F. Muell., Fragm. 1: 46. 1858 sec. rohwer
(1993a). – type: Monococcus echinophorus F. Muell.
Petiveria L., Sp. Pl. 1: 342. 1753 sec. rohwer (1993a). –
type: Petiveria alliacea L.
Phytolacca L., Sp. Pl. 1: 441. 1753 sec. rohwer (1993a).
– type: Phytolacca americana L.
= Nowickea J. Martínez & J. a. Mcdonald in Brittonia
41: 399. 1989.
Phytolacca comprises 25 – 35 species of perennial
herbs, shrubs and trees distributed in north and South
america, eastern asia and new zealand. the genus Nowickea is here included; it was characterized
by a well-developed gynophore, green herbaceous
and often elongated tepals and obovoid or obpyriform fruits with narrowly ellipsoid seeds (Martínez
& Mcdonald 1989). Since its publication, the genus
was known only from the type and considered as distinct from Phytolacca. However, cruz & alcántara
(2000) described several anomalous characteristics
in P. icosandra L. and showed similarities with No
wickea. recently, ramírez-amezcua & Steinmann
(2013) showed that the Nowickea species correspond
to anomalous plants of P. icosandra: the evidence was
based on specimens showing the characteristic flowers of P. icosandra along with anomalous flowers (in
one plant) showing the distinctive characteristics of
Nowickea.
Rivina L., Sp. Pl. 1: 121. 1753 sec. rohwer (1993a). –
type: Rivina humilis L. – Fig. 6a.
Schindleria H. Walter in Bot. Jahrb. Syst. 37, Beibl. 85:
24. 1906 sec. rohwer (1993a). – type: not designated.
Seguieria Loefl., iter. Hispan.: 191. 1758 sec. rohwer
(1993a). – type: Seguieria americana L.
Trichostigma a. rich. in Hist. Fis. cuba 10: 306. 1845
sec. rohwer (1993a). – type: Trichostigma rivinoides
a. rich. – Fig. 6B.
= Villamilla ruiz & Pav. ex Moq. in candolle, Prodr.
13(2): 10. 1849.
Plumbaginaceae Juss. sec. aPG (2009).
a cosmopolitan family of perennial herbs or shrubs,
rarely climbers, mainly distributed in the temperate zones
of the northern hemisphere, especially in the Mediterranean and irano-turanian regions but also in southern africa, southern South america and Western australia. the
family comprises 25 – 30 genera and 650 – 1000 species,
which predominantly occur in arid and saline environments and often in coastal habitats. the family is characterized by flowers that have stamens opposite the petals
and a single basal anatropous ovule with a curled funicle.
Molecular studies based on different markers have shown
that Plumbaginaceae are well supported as monophyletic
family within Caryophyllales and sister to Polygonaceae
(e.g. cuénoud & al. 2002; Hilu & al. 2003). Lledó & al.
(1998, 2001) confirmed the classification of Plumbagi
352
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
naceae into two subfamilies, Limonioideae and Plumba
ginoideae, well differentiated by morphological, chemical
and molecular characters. Plumbaginoideae are mostly
distributed in the pantropical region and comprise four
genera; Plumbago with c. 20 species is the largest. Limo
nioideae have diversified in regions with a Mediterranean
climate and are morphologically more diverse. this subfamily is divided into two tribes: Aegialitideae (one genus
with two species) and Limonieae. Most species of Limo
nieae (> 85%) are grouped into three genera: Acantholi
mon, Armeria and Limonium, while the remaining species
belong to monotypic or small genera (Kubitzki 1993b)
mostly segregated from Acantholimon and Limonium.
the status of most of these genera is unclear; generic concepts and relationships are in need of revision.
Acantholimon Boiss. in diagn. Pl. Orient., ser. 1, 7:
69. 1846, nom. cons. sec. Kubitzki (1993b). – type:
Acantholimon glumaceum (Jaub. & Spach) Boiss. –
Fig. 6c.
a large genus of cushion-forming subshrubs;
150 – 200 species (including many narrow endemics) distributed from southeastern europe to central asia, centred in the mountainous regions of
turkey, iran and afghanistan (Kubitzki 1993b).
the study by Lledó & al. (2005) included only one
representative of Acantholimon, which was recovered
in a clade together with Cephalorhizum and Dictyoli
mon. Moharrek & al. (2014) studied 50 species Acan
tholimon from iran. due to the unresolved position
of Cephalorhizum turcomanicum Popov (found either
as sister to Acantholimon or nested within it), monophyly of Acantholimon is uncertain. Old sections of
Acantholimon were not found as monophyletic (Moharrek & al. 2014).
Aegialitis r. Br., Prodr. Fl. nov. Holland.: 426. 1810 sec.
Kubitzki (1993b). – type: Aegialitis annulata r. Br.
a genus of two woody mangrove species (shrubs or
small trees); one native to southeastern asia, the other
native to australia and Papua new Guinea. Sister to
the rest of Limonioideae (Lledó & al. 2001, 2005) and
placed in the monogeneric tribe Aegialitideae (Lledó
& al. 2001).
Afrolimon Lincz. in novosti Sist. Vyssh. rast. 16: 168.
1979 sec. Kubitzki (1993b). – type: Afrolimon pere
grinum (P. J. Bergius) Lincz.
a group of about ten species from the cape region of
South africa. two representatives sampled by Lledó
& al. (2005; A. peregrinum, A. purpuratum Lincz.)
were found nested within Limonium. according to
these results, the status as a distinct genus can not be
maintained.
Armeria Willd., enum. Pl.: 333. 1809, nom. cons. sec.
Kubitzki (1993b) = Statice L., Sp. Pl. 1: 274. 1753. –
type: Armeria vulgaris Willd. – Fig. 6d.
a genus of c. 90 species, found in temperate regions
of the northern hemisphere and in South america
(chile, tierra del Fuego); a centre of distribution is
the iberian Peninsula (nieto Feliner 1990). Monophyly of Armeria is supported by several studies (Lledó
& al. 2005; Moharrek & al. 2014). Fuertes aguilar &
nieto Feliner (2003) discussed the reticulate evolution in Armeria.
Bakerolimon Lincz. in novosti Sist. Vyssh. rast. 1968:
175. 1968 sec. Kubitzki (1993b). – type: Bakeroli
mon plumosum (Phil.) Lincz.
two species, distributed in the deserts of chile and
Peru. One sampled species was found in a clade together with Armeria–Psylliostachys, Myriolimon and
Saharanthus (Lledó & al. 2005).
Bamiania Lincz. in Bot. zhurn. (Moscow & Leningrad)
56: 1634. 1971 sec. Kubitzki (1993b). – type: Bami
ania pachycorma (rech. f.) Lincz.
Monotypic; from afghanistan. no sequence data are
available for this species yet.
Bukiniczia Lincz. in Bot. zhurn. (Moscow & Leningrad)
56: 1634. 1971 sec. Kubitzki (1993b) ≡ Aeoniopsis
rech. f., Fl. iran. 108: 24. 1974. – type: Bukiniczia
cabulica (Boiss.) Lincz.
Monotypic; B. cabulica is distributed in afghanistan
and Pakistan. not yet included in molecular studies.
Cephalorhizum Popov & Korovin in trudy turkestansk.
naucn. Obsc. 1: 39. 1923 sec. Kubitzki (1993b). –
type: not designated.
two(?) species from afghanistan and central asia.
One species was sampled (C. coelicolor (rech. f.)
rech. f.) and found in a clade together with Acan
tholimon acerosum (Willd.) Boiss. and Dictyolimon
macrorrhabdos (Boiss.) rech. f. (Lledó & al. 2005).
Sequence data for C. turcomanicum Popov were generated by akhani & al. (2013).
Ceratolimon M. B. crespo & Lledó in Bot. J. Linn. Soc.
132: 169. 2000 sec. crespo & Lledó (2000). – type:
Ceratolimon feei (Girard) M. B. crespo & Lledó
= Bubania Girard in Mém. Sect. Sci. acad. Sci.
Montpellier 1: 182. 1848, nom. illeg.
= Limoniastrum sect. Bubania Batt., Fl. algkr. (dicot.): 726. 1890 ≡ Limoniastrum subg. Bubania
(Batt.) Maire in Bull. Soc. Hist. nat. afrique n. 27:
247. 1936.
Ceratolimon, a segregate from Limoniastrum, includes four species of dwarf shrubs with disjunct
distributions on the atlantic and indian Ocean edges
of the Sahara desert (crespo & Lledó 2000). three
species sampled by Lledó & al. (2000) formed a
well-supported clade, that is sister to Limoniastrum.
Ceratostigma Bunge, enum. Pl. china Bor.: 55. 1833
sec. Kubitzki (1993b). – type: Ceratostigma plum
baginoides Bunge
= Valoradia Hochst. in Flora 25(1): 239. 1842.
a genus of about eight species; distributed in asia,
especially in china and the Himalayas; one species
in eastern africa.
Chaetolimon (Bunge) Lincz. in trudy tadzikisk. Bazy
8: 586. 1940 sec. Kubitzki (1993b) ≡ Acantholimon
Willdenowia 45 – 2015
sect. Chaetolimon Bunge in Mém. acad. imp. Sci.
Saint-Pétersbourg, Sér. 7, 18(2): 68. 1872. – type:
Chaetolimon sogdianum Lincz.
Dictyolimon rech. f. in Fl. iran. 108: 21. 1974 sec. Kubitzki (1993b). – type: Dictyolimon macrorrhabdos
(Boiss.) rech. f.
Four species distributed in afghanistan, Pakistan, and
india. One representative was sampled (D. macro
rrhabdos) and found in a clade together with Acan
tholimon acerosum (Willd.) Boiss. and Cephalorhi
zum coelicolor (rech. f.) rech. f. (Lledó & al. 2005).
Dyerophytum Kuntze in revis. Gen. Pl. 2: 394. 1891
sec. Kubitzki (1993b) ≡ Vogelia Lam., tabl. encycl.
2: 147. 1792, nom. illeg. – type: Dyerophytum afri
canum (Lam.) Kuntze
three species of shrubs or subshrubs; from india,
arabia, Socotra and southern africa.
Ghaznianthus Lincz. in novosti Sist. Vyssh. rast. 16:
167. 1979 sec. Kubitzki (1993b). – type: Ghaznian
thus rechingeri (Freitag) Lincz.
Monotypic; from afghanistan. no sequence data are
available yet.
Gladiolimon Mobayen in revis. taxon. acanthol.: 296.
1964 sec. Kubitzki (1993b). – type: Gladiolimon
speciosissimum (aitch. & Hemsl.) Mobayen
Monotypic; distributed in afghanistan. no sequence
data are available yet.
Goniolimon Boiss. in candolle, Prodr. 12: 632. 1848 sec.
Kubitzki (1993b). – type: not designated.
Ikonnikovia Lincz. in Fl. UrSS 18: 745. 1952 sec. Kubitzki (1993b). – type: Ikonnikovia kaufmanniana
(regel) Lincz.
Monotypic; distributed in central asia. no sequence
data are available yet.
Limoniastrum Fabr. in enum. Meth. Pl. Hort. Helmstad:
25. 1759 sec. crespo & Lledó (2000). – type: Limo
niastrum articulatum Moench
Only two species, Limoniastrum guyonianum Boiss.
and L. monopetalum (L.) Boiss., distributed in the
Mediterranean region. narrow circumscription (see
crespo & Lledó 2000) based on results from Lledó
& al. (2000).
Limoniopsis Lincz. in Fl. UrSS 18: 744. 1952 sec. Kubitzki (1993b). – type: Limoniopsis owerinii (Boiss.)
Lincz.
two species, Limoniopsis davisii Bokhari and L. ow
erinii, distributed in eastern turkey and caucasia, respectively. not yet sampled in any molecular studies.
Limonium Mill., Gard. dict. abr., ed. 4: [1328]. 1754,
nom. cons. sec. Kubitzki (1993b). – type: Limonium
vulgare Mill.
= Eremolimon Lincz. in novosti Sist. Vyssh. rast. 22:
200. 1985.
the largest genus of the family with an estimated c. 350
species with a preference for coastal habitats; distributed worldwide but mainly in the Mediterranean region.
Afrolimon was shown to be nested in Limonium and
353
related to L. vulgare, the type of Limonium (Lledó &
al. 2005). Limonium is divided into two major clades
corresponding to subgenera, but otherwise the current infrageneric classification proved to be artificial
(Lledó & al. 2005). akhani & al. (2013) studied the
irano-turanian taxa of Limonium. they stated that
segregation of Eremolimon is not supported by morphology or molecular data (akhani & al. 2013). evolutionary studies of this group are complicated by hybridization, many microspecies and apomictic taxa.
Muellerolimon Lincz. in Bot. zhurn. (Moscow & Leningrad) 67: 675. 1982 sec. Kubitzki (1993b). – type:
Muellerolimon salicorniaceum (F. Muell.) Lincz.
Monotypic genus; halophytic M. salicorniaceum distributed in Western australia. related to Goniolimon
(Lledó & al. 2005).
Myriolimon Lledó & al. in taxon 54: 811. 2005 sec. aPG
(2009) ≡ Statice sect. Myriolepis Boiss. in candolle,
Prodr. 12: 667. 1848 ≡ Limonium sect. Myriolepis
(Boiss.) Sauvage & Vindt, Fl. Maroc 1: 47, 74. 1952
≡ Limonium subg. Myriolepis (Boiss.) Pignatti in
Bot. J. Linn. Soc. 64: 361. 1971 ≡ Myriolepis (Boiss.)
Lledó & al. in taxon 52: 71. 2003, nom. illeg. – type:
Myriolimon ferulaceum (L.) Lledó & al.
two species distributed along the central and western
coasts of the Mediterranean region.
Neogontscharovia Lincz. in Bot. zhurn. (Moscow &
Leningrad) 56: 1633. 1971 sec. Kubitzki (1993b). –
type: Neogontscharovia miranda (Lincz.) Lincz.
Plumbagella Spach in Hist. nat. Vég. 10: 333. 1841 sec.
Kubitzki (1993b). – type: Plumbagella micrantha
(Ledeb.) Boiss.
Monotypic; distributed in central asia.
Plumbago L., Sp. Pl. 1: 151. 1753 sec. Kubitzki (1993b).
– type: Plumbago europaea L.
a genus of 10 – 20 species (“leadworts”) with pantropical distribution.
Popoviolimon Lincz. in Bot. zhurn. (Moscow & Leningrad) 56: 1633. 1971 sec. Kubitzki (1993b). – type:
Popoviolimon turcomanicum (Popov ex Lincz.)
Lincz.
Monotypic; distributed in turkmenistan. no sequence
data are available yet.
Psylliostachys (Jaub. & Spach) nevski in trudy Bot. inst.
akad. nauk S. S. S. r., Ser. 1, Fl. Sist. Vyss. rast 4:
314. 1937 sec. Kubitzki (1993b) ≡ Statice subg. Psyl
liostachys Jaub. & Spach, ill. Pl. Orient. 1: 158. 1844.
– type: Psylliostachys spicata (Willd.) nevski
two or three species; distributed in asia (former Soviet central asia, iran, afghanistan). Psylliostachys
species formed a well-supported clade (Moharrek &
al. 2014) and were shown to be sister to representatives of Armeria (Lledó & al. 2001, 2005; Moharrek
& al. 2014).
Saharanthus M. B. crespo & Lledó in Bot. J. Linn. Soc.
132: 169. 2000 sec. crespo & Lledó (2000) ≡ Cabal
leroa Font Quer in cavanillesia 7: 150. 1935, nom.
354
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
inval. ≡ Lerrouxia caball. in trab. Mus. nac. ci. nat.,
Ser. Bot., 28: 13. 1935, nom. illeg. – type: Saharan
thus ifniensis (caball.) M. B. crespo & Lledó
Monotypic genus; distributed in the western Sahara
desert. Segregated from Limoniastrum based on results from Lledó & al. (2000); was found in a clade
together with Armeria–Psylliostachys, Bakerolimon
and Myriolimon (Lledó & al. 2005).
Vassilczenkoa Lincz. in novosti Sist. Vyssh. rast. 16:
166. 1979 sec. Kubitzki (1993b). – type: Vassil
czenkoa sogdiana (Lincz.) Lincz.
Polygonaceae Juss. sec. aPG (2009).
the Polygonaceae are a morphologically diverse clade
containing more than 50 genera and 1200 species. the
family is a monophyletic group with the morphological
synapomorphies of an ocrea, orthotropous ovules, (usually) trigonous achenes and quincuncial aestivation (Judd
& al. 2007). Polygonaceae are distributed worldwide and
are present in almost all ecosystems ranging from tropical
rainforests to alpine regions and tundra (Brandbyge 1993;
Sanchez & al. 2009). Burke & Sanchez (2011), based on
phylogenetic data, recognized three subfamilies: Eriogo
noideae, Polygonoideae and Symmerioideae. Polygonoi
deae were considered non-monophyletic in previous studies (Lamb Frye & Kron 2003; Sanchez & Kron 2008), but
a new circumscription by Sanchez & al. (2011) supported
a monophyletic subfamily including the type genus Poly
gonum and other genera such as Atraphaxis, Fagopyrum,
Fallopia, Koenigia, Muehlenbeckia, Oxyria, Persicaria,
Rheum and Rumex; whereby Eriogonoideae was expanded to include currently recognized Antigonon, Coccoloba,
Ruprechtia, Triplaris and other members of the woody
genera previously included in Polygonoideae (Sanchez &
Kron 2009; Burke & al. 2010). it is important to mention
that much work is still needed within the tribe Eriogoneae
(or Eriogonoideae s.str.), since most of the recognized
genera have no support as being monophyletic (Kempton
2012). Symmerioideae is monotypic and the only species
recognized is Symmeria paniculata Benth.; this subfamily
shows a unique trans-atlantic disjunction, in the amazon Basin and western africa, which needs further study
(Burke & Sanchez 2011).
Acanthoscyphus Small in Bull. torrey Bot. club 25: 53.
1898 sec. reveal (2005). – type: Acanthoscyphus pa
rishii (Parry) Small
Aconogonon (Meisn.) rchb. in Handb. nat. Pfl.-Syst.:
236. 1837 sec. Brandbyge (1993). – type: Acono
gonon alpinum (all.) Schur
Afrobrunnichia Hutch. & dalziel in Fl. W. trop. afr.
[Hutchinson & dalziel] 1: 118. 1927 sec. Sanchez &
Kron (2009). – type: Afrobrunnichia erecta (asch.)
Hutch. & dalziel
Antigonon endl., Gen. Pl.: 310. 1837 sec. Brandbyge
(1993) ≡ Corculum (endl.) Stuntz in Bull. Bur. Pl.
industr. U. S. d. a. 282: 86. 1913. – type: Antigonon
leptopus Hook. & arn.
the genus Antigonon, with three to six species, consists of woody or herbaceous perennial lianas that
grow in Mexico and central america, with the exception of A. leptopus Hook. & arn., which is widely cultivated as an ornamental (Brandbyge 1993).
Sanchez & Kron (2009), Sanchez & al. (2009), Burke
& al. (2010) and Burke & Sanchez (2011), based on
consistent and highly supported molecular data, proposed that Antigonon and Brunnichia, two genera
with suffrutescent habit and tendril-bearing lianas, are
clearly distinguished from the rest of the subfamily
Eriogonoideae. according to Brandbyge (1993), the
described species are poorly defined and a taxonomic
revision is needed.
Aristocapsa reveal & Hardham in Phytologia 66: 84.
1989 sec. Brandbyge (1993). – type: Aristocapsa in
signis (curran) reveal & Hardham
Atraphaxis L., Sp. Pl. 1: 333. 1753 sec. Schuster & al.
(2011a). – type: Atraphaxis spinosa L.
Bistorta adans., Fam. Pl. 2: 277, 525. 1763 sec. Brandbyge (1993). – type: Bistorta major Gray
Brunnichia Banks ex Gaertn. in Fruct. Sem. Pl. 1: 213.
1788 sec. Sanchez & Kron (2009). – type: Brun
nichia cirrhosa Gaertn.
Calligonum L., Sp. Pl. 1: 530. 1753 sec. Brandbyge
(1993). – type: Calligonum polygonoides L.
Centrostegia a. Gray ex Benth. in candolle, Prodr. 14:
27. 1856 sec. Brandbyge (1993). – type: Centroste
gia thurberi a. Gray ex Benth.
Chorizanthe r. Br. ex Benth. in trans. Linn. Soc. London 17: 405, 416. 1836 sec. reveal (2005). – type:
Chorizanthe virgata Benth.
= Acanthogonum torr. in Pacif. rail. rep. 4: 132. 1856.
= Eriogonella Goodman in ann. Missouri Bot. Gard.
21: 90. 1934.
Coccoloba P. Browne in civ. nat. Hist. Jamaica: 209.
1756, nom. cons. sec. Brandbyge (1993). – type:
Coccoloba uvifera (L.) L. – Fig. 6e & F.
= Guaiabara Mill., Gard. dict. abr., ed. 4: [590]. 1754.
Coccoloba includes c. 120 neotropical species,
which are grouped in four areas with distinguished
endemism: the antilles, central america, northern
South america and the amazon region of Brazil
(Stohr 1982; Brandbyge 1993). the presence of an
ocrea (also ochrea), flowers with five tepals and eight
stamens and the globose or trigonous achenes are the
fundamental characteristics that support the relationships among Coccoloba, Neomillspauhia and Po
dopterus (Sanchez & Kron 2009; Burke & al. 2010;
Burke & Sanchez 2011). the particular ecological
conditions and ecological isolation of the antilles allow inferring a radiation, mainly in cuba and Hispaniola, with c. 40 endemic species; however, there is
no biogeographic hypothesis for the genus. currently,
Coccoloba is classified in several sections, which
have not been phylogenetically evaluated.
Dedeckera reveal & J. t. Howell in Brittonia 28: 245.
Willdenowia 45 – 2015
1976 sec. Brandbyge (1993). – type: Dedeckera eu
rekensis reveal & J. t. Howell
Dodecahema reveal & Hardham in Phytologia 66: 86.
1989 sec. Brandbyge (1993). – type: Dodecahema
leptoceras (a. Gray) reveal & Hardham
Duma t. M. Schust. in int. J. Plant. Sci. 172: 1053. 2011
sec. Schuster & al. (2011b). – type: Duma florulenta
(Meisn.) t. M. Schust.
a new genus segregated form Muehlenbeckia, based
on the molecular study by Schuster & al. (2011). the
genus comprises three species restricted to australia,
and is characterized by erect shrubs with thornlike
branches. this habit it not found in any other Mueh
lenbeckia (as studied by Schuster & al. 2011). in addition, Duma does not possess extrafloral nectaries at
the petiole base, which are present in most species of
Fallopia, Muehlenbeckia and Reynoutria.
Emex campd. in Monogr. rumex: 56. 1819, nom. cons.
sec. Brandbyge (1993). – type: Emex spinosa (L.)
campd.
Eriogonum Michx., Fl. Bor.-amer. 1: 246. 1803 sec.
Brandbyge (1993). – type: Eriogonum tomentosum
Michx.
= Pterogonum H. Gross in Bot. Jahrb. Syst. 49: 239.
1913.
= Sanmartinia M. Buchinger in com. inst. nac. invest.
cienc. nat., Buenos aires, cienc. Bot. 1: 5. 1950.
Eskemukerjea Malick & Sengupta in Bull. Bot. Surv.
india 11: 433. 1972 sec. Brandbyge (1993). – type:
Eskemukerjea nepalensis Malick & Sengupta
this genus has been considered as a synonym of
Fagopyrum, based on pollen morphology (Hong
1988). However, recent molecular analyses did not
place it in Fagopyrum (Ohsako & al. 2001; Galasso
& al. 2009; Sanchez & al. 2011). Since there are
morphological characters that suggest placement in
Fagopyrum, but no molecular evidence for that relationship, Eskemukerjea was left as incertae sedis by
Sanchez & al. (2011).
Fagopyrum Mill., Gard. dict. abr., ed. 4: [495]. 1754,
nom. cons. sec. Brandbyge (1993). – type: Fago
pyrum esculentum Moench
= Harpagocarpus Hutch. & dandy in Bull. Misc. inform. Kew 1926: 364. 1926.
= Parapteropyrum a. J. Li in acta Phytotax. Sin. 19:
330. 1981.
Fallopia adans., Fam. Pl. 2: 277, 557. 1763 sec. Brandbyge (1993). – type: Polygonum scandens L.
= Bilderdykia dumort., Fl. Belg. 1: 18. 1827.
= Pleuropterus turcz. in Bull. Soc. imp. naturalistes
Moscou 21: 587. 1848.
Gilmania coville in J. Wash. acad. Sci. 26: 210. 1936
sec. Brandbyge (1993). – type: Gilmania luteola
(coville) coville
Goodmania reveal & ertter in Brittonia 28: 427. 1977
sec. Brandbyge (1993). – type: Goodmania luteola
(Parry) reveal & ertter
355
Gymnopodium rolfe in Hooker’s icon. Pl.: t. 2699. 1901
sec. Brandbyge (1993). – type: Gymnopodium flori
bundum rolfe
= Millspaughia B. L. rob. in Bot. Jahrb. Syst. 36(3,
Beibl. 80): 13. 1905.
the genus Gymnopodium was originally described
with three species, growing as shrubs or small trees
on limestone soils in Belize, Guatemala, and the Yucatán Peninsula in Mexico (Blake 1921; Brandbyge
1993). Sanchez & al. (2009) and Burke & al. (2010)
showed that Gymnopodium is strongly supported as
monophyletic in the subfamily Eriogonoideae (sec.
Burke & Sanchez 2011); based on leaf shape and
pubescence characters, the genus should be recognized with only one polymorphic species (Burke &
Sanchez 2011).
Harfordia Greene & Parry in Proc. davenport acad. nat.
Sci. 5: 27. 1886 sec. Brandbyge (1993). – type: Har
fordia macroptera (Benth.) Greene & Parry
Hollisteria S. Watson in Proc. amer. acad. arts 14: 296.
1879 sec. Brandbyge (1993). – type: Hollisteria la
nata S. Watson
Johanneshowellia reveal in Brittonia 56: 299. 2004 sec.
aPG (2009). – type: not designated.
Knorringia (czukav.) tzvelev in nov. Syst. Pl. Vas. 24:
76. 1987 sec. Sanchez & al. (2011) ≡ Polygonum sect.
Aconogonon Meisn., Monogr. Polyg.: 43, 55. 1826
≡ Polygonum sect. Knorringia czukav. in novosti
Sist. Vyssh. rast. 3: 92-93. 1966 ≡ Aconogonon sect.
Knorringia (czukav.) Soják in Preslia 46: 151. 1974.
– type: Knorringia sibirica (Laxm.) tzvelev
the genus was segregated from Polygonum s.l. (or
from Aconogonum) and placed in the Coccolobeae
by Hong (1989). Later on it was included within Per
sicaria (Persicarieae) by Brandbyge (1993) and after
that considered as incertae sedis within Polygoneae
by Galasso & al. (2009). its isolated position from
Persicaria was statistically well supported by molecular data in Sanchez & al. (2011). the taxon is
sister to the remaining members of the Polygoneae,
with which it shares characters such as tepal nervature, structure of the exocarp and pollen morphology
(Galasso & al. 2009).
Koenigia L., Syst. nat., ed. 12, 2: 3, 35. 1767 sec. Brandbyge (1993). – type: Koenigia islandica L.
Lastarriaea J. rémy, Fl. chil. 5: 289. 1851-1852 sec.
Brandbyge (1993). – type: Lastarriaea chilensis J.
rémy
Leptogonum Benth., Gen. Pl. 3(1): 103. 1880 sec.
Brandbyge (1993). – type: Leptogonum domingensis
Benth.
Leptogonum is an interesting genus of small trees
or shrubs, endemic to Hispaniola (Liogier 1983;
Brandbyge 1989). in Burke & al. (2010), this genus
was placed in the subfamily Eriogonoideae and recognized as its own subtribe Leptogoneae (Burke &
Sanchez 2011), based on the lack of accrescent tepals
356
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
in the fruit, the reduction to three stamens, and the
leaves clustered at the stem apices.
Magoniella adr. Sanchez in Syst. Bot. 36: 708. 2011 sec.
Sanchez & al. (2011). – type: Magoniella obidensis
(Huber) adr. Sanchez
this recently published genus comprises two species
distributed in Brazil, Bolivia and Venezuela, and it
was segregated from Ruprechtia based on molecular and morphological characters (Sanchez & Kron
2011). Magoniella is characterized by a strict lianaceous habit, and it shares with Salta and Triplaris the
presence of a scar at the base of the perianth in the
fruit.
Mucronea Benth. in trans. Linn. Soc. London 17: 405,
419. 1836 sec. Brandbyge (1993). – type: Mucronea
californica Benth.
Muehlenbeckia Meisn., Pl. Vasc. Gen. 1: 227. 1841,
nom. cons. sec. Brandbyge (1993). – type: Muehlen
beckia australis (G. Forst.) Meisn.
= Homalocladium (F. Muell.) L. H. Bailey in Gentes
Herb. 2: 56. 1929.
Nemacaulis nutt. in Proc. acad. nat. Sci. Philadelphia 4:
18. 1848 sec. Brandbyge (1993). – type: Nemacaulis
denudata nutt.
Neomillspaughia S. F. Blake in Bull. torrey Bot. club
48: 84. 1921 sec. Brandbyge (1993). – type: Neomill
spaughia paniculata (donn. Sm.) S. F. Blake
With only two recognized species, the genus is restricted to the dry forests of Mexico and central
america (Brandbyge 1993; Burke & al. 2010). Previous to their assignment to a new genus by Blake
(1921), species of Neomillspaughia had been placed
in either Campderia Benth. (donnell Smith 1899) or
Podopterus (Gross 1913). roberty & Vautier (1964)
included Neomillspaughia in the genus Podopterus;
however, based on molecular data, Neomillspaughia
receives strong support as sister to Coccoloba
(Sanchez & al. 2009; Burke & al. 2010).
Oxygonum Burch. ex campd. in Monogr. rumex: 18.
1819 sec. Brandbyge (1993). – type: Oxygonum ala
tum Burch.
Oxygonum comprises c. 35 species and is confined to
the african continent and Madagascar (Graham 1957;
Ortiz & Paiva 1999). Several studies have placed the
genus in Polygoneae (Haraldson 1978; Brandbyge
1993; Hong & al. 1998; Galasso & al. 2009); however, Haraldson (1978) suggested a close affinity to
Fagopyrum and genera in Rumiceae. ronse decraene
& akeroyd (1988) suggested an affinity with Poly
gonum. Oxygonum has not been sampled in any molecular study; therefore it was left as incertae sedis by
Sanchez & al. (2011).
Oxyria Hill in Veg. Syst. 10: 24. 1765 sec. Brandbyge
(1993). – type: Oxyria digyna (L.) Hill
Oxytheca nutt. in Proc. acad. nat. Sci. Philadelphia 4:
16. 1848 sec. Brandbyge (1993). – type: Oxytheca
dendroidea nutt.
Persicaria Mill., Gard. dict. abr., ed. 4: [1054]. 1754
sec. Brandbyge (1993). – type: Persicaria maculosa
Gray
= Tovara adans., Fam. Pl. 2: 276, 612. 1763.
= Antenoron raf., Fl. Ludov.: 28. 1817.
= Echinocaulon (Meisn.) Spach, Hist. nat. Vég. 10:
521. 1841.
= Cephalophilum Meisn. ex Börner in Bot. Syst. not.
276. 1912.
= Physopyrum Popov in ind. Sem. Hort. Bot. almaat.
acad. Sci. UrSS. 2: 23. 1935.
Podopterus Bonpl., Pl. aequinoct. 2: 89. 1812 sec. Brandbyge (1993). – type: Podopterus mexicanus Bonpl.
Podopterus includes three species restricted to Mexico and Guatemala (Brandbyge 1993). the genus has
strong morphological affinities to Neomillspaugia
and Coccoloba, based on habit and the presence of
five tepals (Burke & al. 2010). although the placement of Podopterus is not well supported, Burke &
Sanchez (2011) included the genus in the tribe Coc
colobeae alongside Coccoloba and Neomillspaughia.
Neomillspaughia and Podopterus share the presence
of accrescent and membranous inner tepals (Blake
1921; roberty & Vautier 1984).
Polygonum L., Sp. Pl. 1: 359. 1753, nom. cons. sec. Schuster & al. (2011a). – type: Polygonum aviculare L.
= Polygonella Michx., Fl. Bor.-amer. 2: 240. 1803.
Pteropyrum Jaub. & Spach, ill. Pl. Orient. 2: 7. 1844 sec.
Brandbyge (1993). – type: Pteropyrum aucheri Jaub.
& Spach
Pterostegia Fisch. & c. a. Mey. in index Seminum [St.
Petersburg] 2: 23. 1835 sec. Brandbyge (1993). –
type: Pterostegia drymarioides Fisch. & c. a. Mey.
Pteroxygonum dammer & diels in Bot. Jahrb. Syst. 36:
36. 1905 sec. Brandbyge (1993). – type: Pteroxy
gonum giraldii dammer & diels
a monotypic genus found in china. the genus was
considered part of Fagopyrum (Haraldson 1978;
ronse decraene & akeroyd 1988) but molecular
studies do not support this placement (Sun & al. 2008;
Sanchez & al. 2009; tavakkoli & al. 2010). Sun & al.
(2008) suggested that this genus should be placed in
Persicarieae, but in Sanchez & al. (2009) the position
is unresolved. in tavakkoli & al. (2010) there is conflicting placement of Pteroxygonum depending on the
gene region. therefore, Sanchez & al. (2011) decided
to leave this genus as incertae sedis.
Reynoutria Houtt., nat. Hist. 2(8): 639. 1777 sec. Brandbyge (1993). – type: Reynoutria japonica Houtt.
Rheum L., Sp. Pl. 1: 371. 1753 sec. Brandbyge (1993). –
type: Rheum rhaponticum L.
Rubrivena M. Král in Preslia 57(1): 65. 1985 sec. Brandbyge (1993) ≡ Persicaria sect. Rubrivena (M. Král)
S. P. Hong in Pl. Syst. evol. 186: 112. 1993. – type:
Rubrivena polystachya (Wall. ex Meisn.) M. Král
a monotypic genus distributed in afghanistan, Pakistan, india and china (Qaiser 2001). the taxonomy of
Willdenowia 45 – 2015
Rubrivena is complex; its members have been included in Polygonum (P. polystachyum; Li & al. 2003)
and Persicaria (P. wallichii; Freeman 2005), and both
names are accepted by tropicos (undated). However,
based on molecular studies, the placement of Ru
brivena is strongly supported as sister to Aconogonon
and Koenigia (Sanchez & al. 2011).
Rumex L., Sp. Pl. 1: 333. 1753 sec. Brandbyge (1993). –
type: Rumex patientia L.
= Acetosella (raf.) Fourr. in ann. Soc. Linn. Lyon ser.
2, 17: 145. 1869.
= Bucephalophora Pau in not. Bot. Fl. españ. 1: 24.
1887.
= Acetosa Mill., Gard. dict. abr., ed. 4: [unpaged].
1754.
Ruprechtia c. a. Mey. in Mém. acad. imp. Sci. StPétersbourg, Sér. 6, Sci. Math., Seconde Pt. Sci. nat.
6: 148. 1840 sec. Brandbyge (1993). – type: Ru
prechtia ramiflora (Jacq.) c. a. Mey.
= Enneatypus Herzog in Meded. rijks-Herb. 46: 3.
1922.
Salta adr. Sanchez in Syst. Bot. 36: 708. 2011 sec.
Sanchez & al. (2011). – type: Salta triflora (Griseb.)
adr. Sanchez
a new monotypic genus described in Sanchez &
Kron (2011), based on morphological and molecular
data. this genus is commonly found in argentina,
Bolivia, Brazil and Paraguay, and is characterized by
a pronounced development of brachyblasts and the
short axis of the inflorescences borne on a short shoot
(Pendry 2004; Sanchez & Kron 2011). Molecular
studies have strongly supported the placement of this
genus as sister of a clade that includes Magoniella,
Ruprechtia and Triplaris (Burke & al. 2010; Sanchez
& Kron 2011).
Sidotheca reveal in Harvard Pap. Bot. 9: 211. 2004 sec.
aPG (2009) ≡ Oxytheca sect. Neoxytheca ertter. in
Brittonia 32: 92. 1980. – type: not designated.
Sidotheca was established as a new name, replacing
Oxytheca sect. Neoxytheca.
Stenogonum nutt. in Proc. acad. nat. Sci. Philadelphia
4: 19. 1848 sec. Brandbyge (1993). – type: Stenogo
num salsuginosum nutt.
Symmeria Benth. in London J. Bot. 4: 630. 1845 sec.
Brandbyge (1993). – type: Symmeria paniculata
Benth.
Systenotheca reveal & Hardham in Phytologia 66: 85.
1989 sec. Brandbyge (1993). – type: Systenotheca
vortreidei (Brandegee) reveal & Hardham
Triplaris Loefl., iter. Hispan.: 256. 1758 sec. Brandbyge
(1993). – type: Triplaris americana L. – Fig. 6G.
Portulacaceae Juss. sec. nyffeler & eggli (2010a).
a monogeneric family with c. 100 species mainly distributed in tropical and subtropical areas of the world.
Portulaca L., Sp. Pl. 1: 445. 1753 sec. nyffeler & eggli
(2010a). – type: Portulaca oleracea L.
357
= Lemia Vand., Fl. Lusit. Brasil. Spec. 35. 1788.
= Sedopsis (engl. ex Legrand) exell & Mendonça,
consp. Fl. angol. 1: 116. 1937.
= Merida neck., elem. Bot. 2: 382. 1790, nom. inval.
= Lamia Vand. ex endl., Gen. Pl.: 949. 1840, nom. inval.
although the circumscription of the genus has been
relatively stable, the infrageneric classification remains controversial. Previous proposals (von Poellnitz 1934; Legrand 1958; Geesink 1969) are only in
part consistent with the results of a recent phylogenetic analysis (Ocampo & columbus 2012). the genus is monophyletic and has two main lineages: one
whose members have opposite leaves (OL clade) and
are distributed in africa, asia and australia (except
P. quadrifida L., which is a pantropical weed), and a
second lineage whose species have alternate to subopposite leaves (aL clade), are more widespread and
originated in the new World. these major clades and
their subclades have anatomical and morphological
features (Ocampo & columbus 2012; Ocampo & al.
2013) that will be used to amend the classification of
Portulaca.
Rhabdodendraceae Prance sec. aPG (2009).
a monogeneric family comprising three species distributed in tropical South america, the Guyanas, the amazonian region and northeastern Brazil (Prance 2003). the
family has had a complicated taxonomic history. traditionally, species that are now placed in Rhabdodendron
were included in different families of Rutales, in the family Chrysobalanaceae (as the genus Lecostemon dc.)
(Bentham 1853) or in Rutaceae (Gilg & Pilger 1905;
Huber 1909; takhtajan 1980). in other systems (e.g.
cronquist 1981), Rhabdodendron was placed within Ro
sales (for a detailed taxonomic history until the 1970s
see Prance 1972). Based on morphological, palynological and anatomical characters, Prance (1972) considered
Rhabdodendron in its own family, and suggested for the
first time some affinities with Caryophyllales, specifically with Phytolaccaceae. Later on, the early molecular
phylogenetic study of Fay & al. (1997) confirmed the affinities of Rhabdodendraceae with Caryophyllales. Since
then, the position of the family within the order was also
confirmed by subsequent studies (e.g. cuénoud & al.
2002; Hilu & al. 2003; Schäferhoff & al. 2009; Brockington & al. 2009, 2011; Qiu & al. 2010; Soltis & al. 2011),
although there are several hypothesis about its internal
position.
Rhabdodendron Gilg & Pilg. in Verh. Bot. Vereins Prov.
Brandenburg 47: 152. 1905 sec. Prance (2003). –
type: Rhabdodendron columnare Gilg & Pilg.
Sarcobataceae Behnke sec. aPG (2009).
a monogeneric family with two species distributed
in north america (Kühn 1993; Hils & al. 2003), from
the western United States to northwestern Mexico. the
358
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
family includes shrubs characterized by having thorny
branches, ebracteolate and unisexual flowers, staminate
flowers arranged in spikes, whereas the pistillate ones
are solitary (Wels & al. 2003). traditionally, the only genus, Sarcobatus, was placed in Chenopodiaceae (for a
detailed taxonomic history until the 1990s see Behnke
1997). the early molecular phylogenetic study by downie & al. (1997) supported the position of Sarcobatus as an
independent lineage. in this study, Sarcobatus showed a
close relationship with members of Nyctaginaceae and
Phytolaccaceae rather than Chenopodiaceae. Based
on these results in addition to characters of the sieveelement plastids and some morphological characters,
Behnke (1997) described the new family; nevertheless,
some authors continued to treat the genus as part of
Chenopodiaceae (e.g. Hils & al. 2003). the position of
Sarcobatus as an independent lineage was confirmed by
other molecular phylogenetic studies (e.g. cuénoud & al.
2002; Hilu & al. 2003; Brockington 2009, 2011; Soltis &
al. 2011; Schäferhoff & al. 2009), which showed a close
but only moderately supported relationship of the family
with Agdestidaceae.
Sarcobatus nees, reise nord-america 2: 446. 1841 sec.
Hils & al. (2003). – type: Sarcobatus maximilianii
nees – Fig. 7a.
= Fremontia torr., rep. exped. rocky Mts.: 95. 1843.
Simmondsiaceae tiegh. ex reveal & Hoogland sec.
aPG (2009).
a monotypic family native to the Sonoran desert of northwestern Mexico and to neighbouring regions in arizona
and southern california (Vázquez Yanes & al. 1999); it
is also introduced in some countries of South america,
europe, asia and africa. the family includes evergreen
dioecious shrubs with opposite and thick leaves, clearly
articulated near the stem; the staminate flowers are small
and borne in terminal inflorescences, while the pistillate
flowers are single and axillary; the calyx is much enlarged in fruit (Stevens 2001 onwards; Köhler 2003).
traditionally, the family was placed in Hamamelidales
(sensu takhtajan 1980), Euphorbiales (sensu cronquist
1988) or in its own order Simmondsiales (sensu takhtajan 1997), in some cases within Buxaceae or close to
it. However, the early molecular phylogenetic study
by Fay & al. (1997) showed the affinities of Simmond
siaceae with Caryophyllales: this agrees also with several morphological characters of the stylodia, calyces
and secondary growth (Köhler 2003). the affinities of
the family with Caryophyllales were confirmed by subsequent molecular phylogenetic studies (e.g. cuénoud
& al. 2002; Brockington 2009, 2011; Soltis & al. 2011),
which showed that Simmondsiaceae are closer to Rhab
dodendraceae and/or to the remainder of the caryophyllid clade. For further information see notes under Rhab
dodendraceae.
Simmondsia nutt. in London J. Bot. 3: 400. 1844 sec.
Köhler (2003). – type: Simmondsia californica nutt.
the only species, Simmondsia chinensis c. K.
Schneid., is known as a dominant shrub in its native
distribution area. the species is well appreciated
for the liquid wax, extracted from the seeds, which
is used mainly in the cosmetic industry (jojoba;
Vázquez Yanes & al. 1999).
Stegnospermataceae nakai sec. aPG (2009).
a monogeneric family with three species occurring from northwestern Mexico to nicaragua and the
antilles (rohwer 1992). the family includes small
trees and shrubs characterized by bisexual flowers with a two-whorled perianth, one whorl consisting of five free green sepals, and the other whorl of
five white narrow-based petals adherent to the alternisepalous stamens at the base. the fruits are
capsules and the seeds are arillate (rohwer 1993).
When the only genus, Stegnosperma, was described in
1844, it was placed in Phytolaccaceae and accepted
by other authors (e.g. Heimerl 1934). nakai (1942)
elevated the genus to the family level. recognition as
a family was also supported by morphological, palynological and wood-anatomical characters (e.g. nowicke 1969; Bell 1980; carlquist 1999). For a detailed
taxonomic history until the 1980s see Bell (1980).
the early molecular phylogenetic studies of downie &
al. (1997) and Fay & al. (1997) showed the position of
Stegnosperma as an independent lineage. However, both
classifications, the recognition of Stegnospermataceae
(e.g. rohwer 1993; takhtajan 1997) and Stegnosperma
within Phytolacacceae (e.g. Stevens 2001), continued
to be used. Subsequent phylogenetic studies (e.g. Savolainen & al. 2000; cuénoud & al. 2002; Schäferhoff
& al. 2009; Qiu & al. 2010; Brockington 2009, 2011;
Soltis & al. 2011) confirmed the findings of downie &
al. (1997) and Fay & al. (1997), resulting in the wide
recognition of Stegnospermataceae as a separate family.
Stegnosperma Benth., Bot. Voy. Sulphur: 17. 1844 sec.
rohwer (1993b). – type: Stegnosperma halimifolium
Benth.
Talinaceae doweld sec. nyffeler & eggli (2010a).
a family with three genera and around 28 species
mainly distributed in africa, but with a few taxa in the
americas and the tropics around the world (nyffeler &
eggli 2010a). the species of this family are traditionally considered as members of Portulacaceae; however, molecular phylogenetic studies have shown that
the traditional Portulacaceae are not monophyletic
(Hershkovitz & zimmer 1997; applequist & Wallace
2001; nyffeler 2007; nyffeler & eggli 2010a; Ocampo
& columbus 2010). nyffeler & eggli (2010a) proposed
the segregation of the traditional Portulacaceae into
four families (Anacampserotaceae, Montiaceae, Portu
lacaceae and Talinaceae) based on morphological and
molecular data.
Willdenowia 45 – 2015
359
Amphipetalum Bacigalupo in candollea 43: 409. 1988
sec. nyffeler & eggli (2010a). – type: Amphipetalum
paraguayense Bacigalupo
Talinella Baill. in Bull. Mens. Soc. Linn. Paris 1(69):
569. 1886 sec. applequist (2005). – type: Talinella
boiviniana Baill.
= Sabouraea Leandri in adansonia sér. 2, 2: 224. 1962.
Unique in the suborder Portulacineae in having berry-like fruits. Molecular phylogenies show Talinella
embedded in Talinum (Hershkovitz & zimmer 1997;
applequist & Wallace 2001; nyffeler 2007; nyffeler
& eggli 2010a), but nyffeler & eggli (2010a) suggested to accept the genus pending further research
towards a deeply sampled phylogeny of Talinum.
recent treatments by eggli (1997) and applequist
(2005).
Talinum adans., Fam. Pl. 2: 245, 609. 1763, nom. cons.
sec. nyffeler & eggli (2010a). – type: Talinum trian
gulare (Jacq.) Willd.
Molecular and phylogenetic analyses have shown
that the new World species with terete to semi-terete
leaves formerly treated as members of Talinium form
a monophyletic genus within Montiaceae (Phemeran
thus; carolin 1987; Hershkovitz & zimmer 2000;
applequist & Wallace 2000; nyffeler & eggli 2010a;
Ocampo & columbus 2010).
genus or as a synonym of Myricaria (Yang & Gaskin
2007) or Tamarix (Baum 1978).
Reaumuria L., Syst. nat. (ed. 10) 2: 1081. 1759 sec.
Gaskin (2003). – type: Reaumuria vermiculata L.
Reaumuria is a xerohalophytic genus with c. 13
shrubby and rarely annual species occurring in
deserts and semi-deserts of southwestern and central
asia (Bobrov 1966; zohary & danin 1970). except
for one polymorphic species group (R. alternifolia
(Labill.) Britten), Reaumuria species are characterized by cylindrical succulent leaves.
Tamarix L., Sp. Pl. 1: 270. 1753 sec. Gaskin (2003). –
type: Tamarix gallica L. – Fig. 7B.
Tamarix with c. 60 species is most diversified in saline
and wet habitats of the Old World and is naturalized in
australia and the americas, sometimes as aggressive
invasive plants. it is one of the few lineages in Caryo
phyllales that contain large trees and shrubs with a
significant role in carbon sequestration and vegetation
under harsh and salty conditions. the taxonomy and
phylogenetic reconstruction of Tamarix are challenging due to the absence of reliable constant characters
and the occurrence of hybridization even among morphologically very different species (Gaskin & Kazmer 2009; Mayonde & al. 2015; Samadi & al. 2013;
H. akhani & t. Borsch, unpubl. data).
Tamaricaceae Link sec. aPG (2009).
Five genera and c. 80 species occurring in africa, asia and
europe with major distribution in the irano-turanian and
Mediterranean regions (Gaskin 2003). Phylogenetic studies support the monophyly of the genera. three well-supported clades have been recovered: Hololachna and Reau
muria; Myricaria and Myrtama; and Tamarix. Tamarix is
sister to the clade comprising Myricaria and Myrtama,
and this group is sister to Hololachna and Reaumuria
(Gaskin & al. 2004). the main feature in most genera of
Tamaricaceae is the presence of salt glands, which enable
successful growth in salty and riparian habitats.
Hololachna ehrenb. in Linnaea 2: 273. 1827 sec. Gaskin
(2003). – type: Hololachna songarica (Pall.) ehrenb.
this is a monotypic genus restricted to central asia
and Mongolia. Hololachna is sister to Reaumuria
(Gaskin & al. 2004).
Myricaria desv. in ann. Sci. nat. (Paris) 4: 349. 1825
sec. Gaskin (2003). – type: Myricaria germanica
(L.) desv.
Myricaria is a hygrophytic genus with c. 13 species
occuring in europe and central asia. Molecular phylogenetic studies support a sister group relationship
between Myrtama and Myricaria (Wang & al. 2009).
Myrtama Ovcz. & Kinzik. in dokl. akad. nauk tadzh.
SSr 20(7): 55. 1977 sec. Gaskin (2003) ≡ Tamari
caria Qaiser & ali in Blumea 24: 153. 1978. – type:
Myrtama elegans (royle) Ovcz. & Kinzik.
the monotypic genus Myrtama is variously interpreted in taxonomic references, both as an independent
Incertae sedis
Jorena adans. in Fam. Pl. (adanson) 2: 249. 1763 sec.
Bittrich (1993c). – type: not designated.
Listed as a “doubtful genus” in the Caryophyllaceae
by Bittrich (1993c).
Summary: current knowledge, trends, gaps
Phylogenetic sampling as a basis for classification
the synopsis of the genera currently accepted in Caryo
phyllales along with a discussion on the recent work dealing with these genera provides a comprehensive source
of information on the current knowledge of this group of
plants. in the context of global undertakings, such as the
World Flora Online (WFO; cBd-SBStta 2012), this
study forms the basis for a gap analysis on the availability of treatments for a major group of flowering plants.
the results indicate that there is a substantial taxonomic
turnover when comparing the current classification with
generic concepts available in the complete treatment of
the order in Kubitzki’s “Families and genera of vascular
plants” (FGVP; Kubitzki & al. 1993; Kubitzki & Bayer
2003; table 2). the number of families has increased
substantially (27 vs 39 families), reflecting changes
necessary because families were not monophyletic (e.g.
Portulacaceae). in addition, several isolated lineages
have been recovered that were consequently elevated
to family rank (e.g. Kewaceae, Macarthuriaceae). the
most diverse families in terms of numbers of genera are
360
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
the Cactaceae, Aizoaceae, Chenopodiaceae and Caryo
phyllaceae (all with over 100 genera), while 28 families
comprise only one to six genera (table 2). at the generic
level, the numbers have increased by more than ten percent in comparison to the last complete treatments in the
FGVP volumes (table 2). While the number of genera
has remained equal (or nearly so) in 18 families, generic
boundaries have changed dramatically in some families,
especially in Cactaceae and Caryophyllaceae.
it is also clear that sampling at the species level is far
from complete, so that many genera or entire tribes lack
data needed to assert their monophyly and/or their exact
position in the families, while others are already known
to be polyphyletic but are insufficiently sampled to be reclassified. in addition, for many taxa no taxonomic revision is available or the existing one is clearly outdated.
For example, in the Aizoaceae of South africa, 55 %
of taxa are in need of revision, 52 % of the recognized
taxa in the family have not been treated in any revision,
with an additional 12 % of taxa revised prior to 1970
(von Staden & al. 2013). in the Ruschioideae the five
largest genera, Ruschia (206 species), Lampranthus (194
species), Delosperma (142 species), Drosanthemum (107
species) and Antimima (96 species) have never been comprehensively revised at species level (i.e. there is no key
to the species). the same is true for numerous smaller
genera such as Stomatium (39 species), Hereroa (27 species) and Malephora (16 species). in addition, a recent
extensive phylogenetic study of Ruschieae, the most speciose clade in Aizoaceae, showed that numerous genera
are not monophyletic, including the large genus Ruschia
(Klak & al. 2013). despite the lack of resolution in parts
of the tree due to the lack of variable gene regions, the
many cases of polyphyly detected in the phylogeny
were an indication of misplaced taxa and narrow generic concepts upheld by traditional taxonomists (Klak
& al. 2013). in particular, mono- and bitypic genera in
Ruschieae, which were found to be nested within larger
genera, need critical re-evaluation (Klak & al. 2013).
in contrast, for the Mesembryanthemoideae a phylogeny is available with an almost complete sampling of
species (Klak & al. 2007) as well as detailed morphological studies and revisions published for most clades
over the last 30 years (e.g. Bittrich 1986; Klak & Linder
1998; Klak & al. 2006; Gerbaulet 1995, 1996a – c, 1997,
2001). However, a conflict in genus delimitaton has
erupted between taxonomists with regard to the number
of genera that should be recognized in Mesembryanthe
moideae. Whereas Klak & Bruyns (2013) favoured a
generic concept based on monophyly, Gerbaulet (2012)
supported the traditional system of “many genera”,
which upholds also genera shown to be paraphyletic
(e.g. Phyllobolus). no detailed phylogeny is available for the Aizooideae, which include c. 108 species
in seven genera. Finally, a further phylogeny including
18 species from Tetragonioideae indicated that several
genera, such as Aizoanthemum, Aizoon and Gunniop
sis may not be monophyletic (c. Klak, pers. comm.).
in contrast, phylogenetic relationships of the smallest
subfamily, Sesuvioideae, which is sister to the remaining Aizoaceae (Klak & al. 2003a, b), are resolved and
generic concepts were clarified recently (thulin & al.
2012; Bohley & al. 2015).
For Basellaceae, eriksson (2007) recognized four
genera and 19 species in comparison to four genera and
17 – 2 2 species accepted by Sperling & Bittrich (1993). in
his phylogenetic analysis based on morphological data,
three of the genera are supported as monophyletic, while
the monophyly of the fourth genus (Basella) is more uncertain. this analysis is well sampled (all taxa), but the
resolution is rather poor. no analysis based on molecular
data has been done yet.
Available treatments in modern floras are patchy on
a global level
Monographic work provides the in-depth synthetic information, and the checklist and gap analysis presented here
is aimed at defining part of the baseline for such analysis
in the Caryophyllales where it is still missing. However,
for the aim of creating a global synthesis of knowledge in
the Caryophyllales it is indispensable to consider also the
information published in floras.
it is difficult to know in how many floras or related
works the Caryophyllales have been treated in the past,
especially in regions with a long history of botanical activity such as c and W europe. in fact, if we take the
establishment of the Linnaean classification system and
naming as a starting point, we can commence right away
in the 18th century, for example with Linnaeus’s own Flo
ra suecica (Linné 1745). Flora treatments are numerous;
setting aside the numerous works of mostly historical interest, Frodin (2001) in the second edition of “Guide to
standard floras of the world” gave information on nearly
1000 general floras distributed in ten major regions of the
world. Only in a few cases is there specific information
about the families or groups treated in each flora (e.g.
Flora of Nigeria: Caryophyllales by Ghazanfar 1991);
for the other floras it is necessary to review each flora
individually in order to identify works of significance for
a global synthesis.
Our approach for uniting the information available
for the global synthesis is partly based on taking advantage of information technologies, and fortunately
floras are increasingly published on the World Wide
Web. an initial review of such publication has revealed
that many historical floras that include treatments of
Caryophyllales are already available online, for example the pre-1900 floras of the alps, australia, Barbados,
Brazil, india, Jamaica, niger, Sri Lanka and Syria, and
pre-1990 treatments from chile, costa rica, Fiji, Guatemala, Japan, Madagascar, Panama, South africa and
taiwan. the bibliographic references of these floras are
cited in Frodin (2001), but can also be accessed through
Willdenowia 45 – 2015
the Biodiversity Heritage Library (BHL 2005+), JStOr
(JStOr 2000+), Gallica (1997+) and Google Books
(2015). More recent floras including the Caryophyl
lales are those from china, nicaragua, the Malesian
region (indonesia, Malaysia, Singapore, Brunei darussalam, the Philippines, and Papua new Guinea) and the
zambesi river basin (Botswana, Malawi, Mozambique,
zambia, zimbabwe and the caprivi Strip), in which
the last treatments for some families of Caryophylla
les were printed in the 2000s. incomplete floras (and
incomplete for Caryophyllales so far) treat argentina,
the Hawaiian islands, north america north of Mexico,
the Marquesas islands, Mesoamerica, Madagascar, the
neotropics, Pakistan, Palestine and tasmania.
Most of these are simply digitized print treatments
(representing images of the actual print work, which, depending on their quality, may or may not be searchable
after optical character recognition – Ocr). in contrast to
this, very few “true” e-floras exist, i.e. floras produced
with the online publication as their principal output and
making full use of existing biodiversity informatics techniques. an example of the latter is the Flora of Western
australia (Western australian Herbarium 1998+).
However, increasingly various intermediates between
electronic representations of print media and true e-floras
are becoming available, partly as a result of the computerized editing process of the print publication, and partly
because printed floras are “marked up” in order to database their content, for example the treatments of Flora
Malesiana (see Hamann & al. 2014).
another important source of information on Caryo
phyllales are checklists, which are mostly available online, because most of them were developed over the past
two decades. Some of them refer to taxa treated in previously printed floras, some of them are continuously
updated and others are in progress. Such checklists are
available for africa, argentina, australia, Bolivia, Botswana, Brazil, central africa, cono Sur (argentina, southern Brazil, chile, Paraguay and Uruguay), costa rica,
croatia, cyprus, ecuador, europe plus the Mediterranean
region, Germany, the Guiana Shield (Guyana, Suriname,
French Guiana and part of Venezuela), iran, ireland, israel, Lesotho, Madagascar, Mexico, Micronesia, Mongolia,
Myanmar, namibia, nepal, new zealand, the pan-arctic
region, Paraguay, Peru, Portugal, the Philippines, Singapore, South africa, southern africa, Suriname, Swaziland, Switzerland, taiwan and the United States.
all of these floristic projects have generated valuable
information that has increased our knowledge about the
Caryophyllales. an online bibliography of such sources
of information focussing on Caryophyllales is in preparation, and we envision using this as the base of a comprehensive gap analysis for the order, and also as the basis
for an analysis of regional differences in taxon concepts.
it became clear from the preliminary survey that such
gaps exist, and that there is a lack of synchronization
of taxon concepts, partly due to the state of knowledge
361
at the time of the production of the treatment, but often
also caused by a specific local perspective that needs to
be placed into a wider geographic context. this was one
of the reasons for the decision to use the edit Platform
for cybertaxonomy for data management, because this
is currently the only taxonomic software system natively
supporting different classifications, taxonomic concepts
and taxon-concept relationships. it indicates also the
need for increased efforts to share and integrate the information generated and to promote the filling of gaps in
both geographic and taxonomic coverage. this will be
facilitated by the application of information technology,
making the information openly available in electronic
form and thus furthering the process of future revision
and dissemination. additionally, it enables new kinds of
links to current data, including those available only in
virtual form, which has not readily been possible in the
past (Frodin 2001).
Conclusions and future work
While the published version of this treatment only includes citable publications as its basic reference, there
will be a dynamic online version of this generic synopsis
that will not only be continuously updated but also become more extensive. to facilitate both interaction in the
scientific community and to inspire further research on
the Caryophyllales, key data to relevant current projects
and research underway will be presented. One of the key
steps on the way to a synthesis of Caryophyllales will be
identifying specialists who are working at the species level; some of them are those who contributed to this generic synopsis, but others have already been identified and
agreed to collaborate. Within the network, we then have
to organize the work on taxonomic groups with several
specialists and to develop a format, as standardized as
possible, for the species-level taxonomic treatments. in
addition, directories of specialists, of electronic resources
and an online bibliography for the Caryophyllales will be
developed. Starting with the Caryophyllales 2015 conference in Berlin (September 2015), regular meetings of
the Caryophyllales community will drive this process.
Role of authors
the draft of the generic checklist and the initial data
entry was the work of PH, who also provided the treatments of Achatocarphaceae, Agdestidaceae, Ancistro
cladeceae, Asteropeiaceae, Barbeuiaceae, Didiereace
ae, Dioncophyllaceae, Droseraceae, Drosophyllacae,
Frankeniaceae, Halophytaceae, Limeaceae, Lophiocar
paceae, Microteaceae, Nepenthaceae, Nyctaginaceae,
Physenaceae, Rhabdodendraceae, Sarcobataceae, Sim
mondsiaceae and Stegnospermataceae and collaborated
in some notes of Chenopodiaceae, Phytolaccaceae and
362
Hernández-Ledesma & al.: a taxonomic backbone for Caryophyllales
Polygonaceae. the following groups were revised by
specific authors: Aizoaceae: cK, with contributions by
GK (Sesuvioideae); Amaranthaceae: tB, with contributions by GK (Polycnemoideae); Anacampserotaceae,
Molluginaceae and Portulacaceae: GO; Montiaceae and
Talinaceae: GO, with contributions by Ue; Cactaceae:
Sa, Ue, nK, rn, BOS; Caryophyllaceae: rr, BO (Si
leneae), with contributions by SvM; Basellaceae: re;
Chenopodiaceae: Ha, HFO, SFB, GK, PU; Gisekiaceae:
GK; Plumbaginaceae, Kewaceae, Macarthuriaceae and
contributions to other families (e.g. Deeringia, Hyperte
lis, Microtea): SvM; Polygonaceae: icn, aS; Tamari
caceae: Ha. WGB extensively rechecked the nomenclatural references and standardization of database entries.
SvM edited entries and updated the database. introduction and summary were prepared as a draft by PH, tB
and WGB. comments from co-authors were incorporated, and the final text edited by WGB, SvM, nK and tB.
Acknowledgements
We would like to acknowledge the technical support by
Katja Luther, andreas Müller and cherian Mathew at the
BGBM during work with the edit Platform software
and the production of the generic list directly from the
database. there was a productive exchange with James
Solomon (Missouri Botanical Garden) while checking
our nomenclatural data against the tropicos database.
Werner Greuter, nicholas turland, and Wolf-Henning
Kusber are acknowledged for advice on complicated cases of nomenclature. demet töre provided literature on
Plumbaginaceae.
Wilhelm Barthlott, Peter Bruyns and nicholas turland are thanked for granting permission to use their photographs. david Hunt, John Mcneill, Sergei Mosyakin,
Kai Müller, Louis ronse de craene, nigel taylor and one
anonymous reviewer are thanked for their valuable comments on an earlier version of the manuscript.
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