Phylogeny and Circumscription of Cephalocereus (Cactaceae) Based on Molecular

and Morphological Evidence

Author(s): Héctor J. Tapia, María Luisa Bárcenas-Argüello, Teresa Terrazas, and Salvador Arias
Source: Systematic Botany, 42(4):1-15.
Published By: The American Society of Plant Taxonomists
URL: http://www.bioone.org/doi/full/10.1600/036364417X696546

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Systematic Botany (2017), 42(4): pp. 1–15
© Copyright 2017 by the American Society of Plant Taxonomists
DOI 10.1600/036364417X696546
Date of publication December 27, 2017

Phylogeny and Circumscription of Cephalocereus (Cactaceae) Based on

Molecular and Morphological Evidence

Héctor J. Tapia,1 Marı́a Luisa Bárcenas-Argüello,1 Teresa Terrazas,2

and Salvador Arias1,3
1
Jardin Botanico, Instituto de Biologia, Universidad Nacional Autonoma de Mexico, Apdo. Postal 70-614,
Mexico 04510 D. F., Mexico.
2
Departamento de Botanica, Instituto de Biologia, Universidad Nacional Autonoma de Mexico, Apdo. Postal
70-233, Mexico 04510 D. F., Mexico.
3
Author for correspondence (sarias@ib.unam.mx)

Communicating Editor: Susana Freire

Abstract—Cephalocereus, Neobuxbaumia, and Pseudomitrocereus (Cactaceae, Cactoideae, Echinocereeae) are related genera of columnar cacti
native to Mexico with current ambiguous circumscription. We applied maximum parsimony and Bayesian inference methods to reconstruct the
phylogeny of the Cephalocereus group using molecular data from seven chloroplast regions (petL-psbE, psbA-trnH, rpl16, rpl32-trnLUAG, trnL-F, trnQ-
rps16, and ycf1), simple coded indels, and 46 structural characters. The Cephalocereus group was recovered as monophyletic with high support values,
whereas Neobuxbaumia appeared as paraphyletic, due to the polyphyly of Cephalocereus and the derived position of Pseudomitrocereus within this
group. Topology was mostly congruent among the different phylogenetic methods explored, and three pervasive clades were observed. Two
structural characters were confirmed as synapomorphies for the Cephalocereus group: prismatic crystals in the dermal system and a perianth woody
cap persistent in the fruit. The derived position of Pseudomitrocereus suggests that new hypotheses must be explored regarding the possible hybrid
origin of this taxon. We propose the transfer of all species of Neobuxbaumia, Cephalocereus, and Pseudomitrocereus to a single genus, in which
Cephalocereus takes priority over the other names. Based on our results, a new circumscription for Cephalocereus is proposed, including a taxonomic
synthesis, two new combinations (Cephalocereus multiareolatus and Cephalocereus sanchezmejoradae), and a key for species.

Keywords—Combined analyses, morphology, Neobuxbaumia, Pseudomitrocereus, taxonomy.

Cephalocereus Pfeiff., Neobuxbaumia Backeb., and Pseudomi-
trocereus Bravo & Buxb. (Cactaceae, Cactoideae, Echinocereeae)
are three genera usually recognized in taxonomic treatments
for columnar cacti (Bravo-Hollis 1978; Guzmán-Cruz et al.
2003; Hunt et al. 2006; Arias et al. 2012). These genera are
endemic to Mexico, distributed from Tamaulipas in the
northeast to Chiapas in the southeast and to Jalisco in the west
(Guzmán-Cruz et al. 2003), where they inhabit ample envi-
ronmental gradients (e.g. precipitation regimes and edaphic
conditions) along their geographic ranges in the tropical de-
ciduous forest and thorn scrub vegetation (Bravo-Hollis et al.
1970, 1971a, b, 1973; Bravo-Hollis 1978; INEGI 2014).
Cephalocereus was erected by Pfeiffer (1838) as a group of
species characterized by having a differentiated flowering
region near the stem apices and naked flowers with tiny scales
(Table 1). The number of accepted species has changed be-
tween different taxonomic treatments, but currently five taxa
are accepted (Table 2). Neobuxbaumia was described by
Backeberg (1938) to include giant columnar species with
crown flowering, without a dense coverage of trichomes (non-
differentiated flowering regions), bare flowers, and edible
fruits with white pulp, segregating it from Cephalocereus
(Table 1). During the twentieth century, taxonomists extended
the circumscription of Neobuxbaumia by recognizing up to nine
species (Table 2). Hunt et al. (2006) accepted the existence of
two recently described species of Neobuxbaumia: N. squamulosa
Scheinvar & Sánchez-Mej. and N. sanchezmejoradae Lau, but
considered N. multiareolata (E.Y. Dawson) Bravo, Scheinvar &
Sánchez-Mej. as an infraspecific entity of N. mezcalaensis
(Bravo) Backeb., although the morphometric study by Arroyo-
Cosultchi et al. (2010) had brought sufficient evidence to
validate the idea that these taxa represent two distinct species.
Pseudomitrocereus is a name proposed by Bravo and Buxbaum
(in Buxbaum 1961) to replace Mitrocereus, which was con-
sidered by Backeberg (1938) as a subgenus of Cephalocereus
and subsequently elevated to genus (Backeberg 1960). This
1
monotypic genus is characterized by its imposing tree-size
(one of the largest taxa of southern Mexico), with a flowering
region differentiated at the apex of mature branches, and
flowers and fruits covered with abundant trichomes and
bristles (Table 1).
The relationships of these three genera have been subject to
different interpretations. Britton and Rose (1920) recognized
Cephalocereus as a large genus (48 species), relating species
from North and South America. However, other taxonomists
argued that this circumscription was artificial and divided
it into smaller genera, leaving the genus Cephalocereus s. s.
with only five species, all native to Mexico. Backeberg
(1960) related Cephalocereus with eight other genera distributed
in North America, including four genera erected by this author
(Table 2; Neodawsonia Backeb., Haseltonia Backeb., Neobuxbaumia,
Mitrocereus Backeb.). Dawson (1952a) based on morphological
similarities proposed Neobuxbaumia and Carnegiea Britton &
Rose as related taxa. Later, Buxbaum (1958, 1961) considered
Neobuxbaumia closely related to Pachycereus Britton & Rose,
Lemaireocereus Britton & Rose, and particularly to Cephalocereus
and Backebergia Bravo. This hypothesis was not supported by
Gibson and Horak (1978), who suggested that Cephalocereus is
the sister taxon of a lineage recognized as subtribe Pachy-
cereinae, and integrated by Pachycereus, Backebergia, Neo-
buxbaumia, and Pseudomitrocereus. Also, Gibson and Horak
(1978) hypothesized a relationship among Neobuxbaumia and
Carnegiea, based on their common growth form and other
morphological similarities of the stem, flower, and fruit.
Consequently, some authors proposed the inclusion of Neo-
buxbaumia in Carnegiea (Hunt and Taylor 1986, 1990), a view
shared in some taxonomic treatments (e.g. Barthlott and Hunt
1993); however, in other treatments, these genera were re-
tained as independent (Hunt 1999; Guzmán-Cruz et al. 2003;
Hunt et al. 2006).
Phylogenetic evidence based on molecular (rpl16, trnL-F,
and the ITS region of nuclear ribosomal DNA) and morphological
2 SYSTEMATIC BOTANY [Volume 42

Table 1. Comparison of diagnostic morphological characters between genera included in the Cephalocereus group.

Cephalocereus (Pfeiffer 1838; Neobuxbaumia (Backeberg 1938; Pseudomitrocereus (Backeberg 1938;

Character \ Taxon Bravo-Hollis 1978; Hunt et al. 2006) Bravo-Hollis 1978; Hunt et al. 2006) Buxbaum 1961; Bravo-Hollis 1978)
Branching pattern Unbranched / branched Unbranched / branched Branched
Differentiated flowering region Present Absent / present Present
Position of the flowering region Apical/lateral Apical Apical
Flower shape Tubular-campanulate / Tubular-campanulate / tubular-funnelform Shortly campanulate
infundibuliform-campanulate
Flower scales size Tiny Tiny Big
Flower areoles trichomes Present Present Present
Flower areoles bristles/spines Absent / present Absent Present
Pulp colour Red or white White White

characters showed that Cephalocereus, Neobuxbaumia, and Pseu-
domitrocereus form a monophyletic group (Cephalocereus group),
sister to the Pachycereus and Lemaireocereus groups and clearly
distinct from Carnegiea (Arias et al. 2003, 2005; Arias and
Terrazas 2006), although they did not find enough evidence
on its internal relationships. Morphological characters sup-
porting the relationships of the Cephalocereus group include
the occurrence of calcium oxalate crystals in the hypodermis
and epidermis (Terrazas and Loza-Cornejo 2002), a perianth
woody cap that is persistent in the fruit, white pulp, and
longitudinal dehiscence (Arias and Terrazas 2006). Neobuxbaumia
bears a non-differentiated flowering region (lacking a cepha-
lium), whereas Cephalocereus has a differentiated flowering
region, a lateral cephalium in subgen. Cephalocereus (Vázquez-
Sánchez et al. 2005, 2007) or an apical pseudocephalium in
subgen. Neodawsonia (Bravo-Hollis 1978; Bárcenas-Argüello
2006); bare flower is an attribute shared by species of both
genera. Pseudomitrocereus also has large trichomes and bristles
(modified spines) in the flower, in addition to a differentiated
flowering region, as defining characters (Gibson and Horak
1978; Terrazas and Loza-Cornejo 2002; Arias and Terrazas
2006; Bárcenas-Argüello 2006; Hunt et al. 2006).
Phylogenetic studies at the tribe or higher levels have shown
differing results due to the poor sampling of taxa and the
limited number of markers included. Phylogenetic analysis of
Cactaceae by Bárcenas et al. (2011) based on the trnK-matK
region provided resolution sufficient only to recover Cepha-
locereus and Neobuxbaumia as members of the tribe Echino-
cereeae, along with Pachycereus, Peniocereus (A.Berger) Britton &
Rose, Echinocereus Engelm., and Stenocereus (A.Berger) Riccob.,
among other genera. Hernández-Hernández et al. (2011)
recovered seven species from the Cephalocereus group as
monophyletic while offering no resolution regarding the re-
lationships among the three genera. Thus, the phylogenetic
relationships of these three genera remain unclear. In the
present study, we undertook a more comprehensive phylo-
genetic analysis of the Cephalocereus group using the following
two different sets of characters: a) DNA sequence data from
seven chloroplast markers (petL-psbE, psbA-trnH, rpl16, rpl32-
trnLUAG, trnL-F, trnQ-rps16, and ycf1) and b) 46 structural
characters. Our goals are to present a robust hypothesis of
phylogenetic relationships for the three genera within the
Cephalocereus group, to analyse the evolution of structural
characters, and to reassess the circumscription and classifi-
cation of these taxa. We expect that the inclusion of mor-
phological data will corroborate and strengthen the phylogeny
produced using molecular data alone.
Materials and Methods
Taxon Sampling—To test the monophyly of the genera Cephalocereus,
Neobuxbaumia, and Pseudomitrocereus, all the fifteen recognized species of
the group sensu Arias et al. (2003) were considered, including nine species
of Neobuxbaumia, five species of Cephalocereus, and the monotypic Pseu-
domitrocereus (Table 2; Hernández-Ledesma et al. 2015). We include 23
terminals as ingroup, considering more than one terminal for some species
of Neobuxbaumia (N. euphorbioides (Haw.) Buxb., N. mezcalaensis, N. scoparia
(Poselg.) Backeb., N. squamulosa, and N. tetetzo (F.A.C.Weber ex K.Schum.)
Backeb.) in order to represent the amplitude or disjunction of their geo-
graphic ranges and to test the delimitation of these species. Geographic
distributions were identified from herbarium specimens (MEXU, XAL) and
taxonomic revisions, and an extensive sampling was conducted to de-
lineate distributions for each species, from 2012 to 2015. The outgroup
includes 12 terminals representing the tribe Echinocereeae (subtribes

Table 2. Species listed in Cephalocereus, Neobuxbaumia, and Pseudomitrocereus, according to several authors.

Britton & Rose Backeberg Buxbaum Bravo-Hollis Hunt & Taylor Hernández-Ledesma
Taxon (1920) (1938, 1960) (1958, 1961) (1978) (1986, 1990) et al. (2015)
Cephalocereus apicicephalium - Neodawsonia Cephalocereus Cephalocereus Cephalocereus Cephalocereus
C. columna-trajani Cephalocereus Haseltonia Cephalocereus Cephalocereus Cephalocereus Cephalocereus
C. nizandensis - Neodawsonia - Cephalocereus - Cephalocereus
C. senilis Cephalocereus Cephalocereus Cephalocereus Cephalocereus Cephalocereus Cephalocereus
C. totolapensis - Neodawsonia - Cephalocereus - Cephalocereus
Neobuxbaumia euphorbioides Cephalocereus Rooksbya Neobuxbaumia Neobuxbaumia Carnegiea Neobuxbaumia
N. macrocephala Cephalocereus Mitrocereus Neobuxbaumia Neobuxbaumia Carnegiea Neobuxbaumia
N. mezcalaensis - Neobuxbaumia Neobuxbaumia Neobuxbaumia Carnegiea Neobuxbaumia
N. multiareolata - Neobuxbaumia Neobuxbaumia Carnegiea Neobuxbaumia
N. polylopha Cephalocereus Neobuxbaumia Neobuxbaumia Neobuxbaumia Carnegiea Neobuxbaumia
N. sanchezmejoradae - - - - Neobuxbaumia
N. scoparia Cephalocereus Neobuxbaumia Neobuxbaumia Neobuxbaumia Carnegiea Neobuxbaumia
N. squamulosa - - - Neobuxbaumia - Neobuxbaumia
N. tetetzo - Neobuxbaumia Neobuxbaumia Neobuxbaumia Carnegiea Neobuxbaumia
Pseudomitrocereus fulviceps Pachycereus Mitrocereus Pseudomitrocereus Mitrocereus Pachycereus Pseudomitrocereus
2017] TAPIA ET AL.: PHYLOGENY OF CEPHALOCEREUS 3

Pachycereinae and Echinocereinae), three terminals from Hylocereeae, one
terminal from Leptocereeae, and one terminal from Trichocereeae (Hunt
et al. 2006; Arias et al. 2005). The included terminals belong to the subfamily
Cactoideae, and most of them specifically correspond to the ACHLP clade
referred by Nyffeler (2002). All newly sampled specimens belong to
Cephalocereus, Neobuxbaumia, and Pseudomitrocereus, whereas the remaining
vouchers were taken from previous studies (Arias et al. 2003, 2005;
Vázquez-Sánchez et al. 2013; Sánchez et al. 2014) (Appendix 1).
DNA Sequence Data—Total DNA was extracted from silica gel dried
samples using the EZ-10 spin column genomic plant DNA mini-prep kit
(BioBasic Inc., Ontario, Canada). The extraction conditions followed the
manufacturer’s instructions, except for the time of tissue incubation, which
was increased from 20 to 90 min (at 65°C) to overcome problems with
excessive mucilage. The presence of DNA was confirmed by agarose
gel electrophoresis and staining with GelRed™ (Biotium Inc., Fremont,
California). DNA extractions were stored at -20°C until required. Seven
chloroplast regions were selected for phylogenetic analysis (rpl16, trnL-F,
psbA-trnH, rpl32-trnL, trnQ-rps16, petL-psbE, and ycf1; Table 3) based on
previous studies on familial, tribal, or generic levels. Although the ITS
region has been used in Cactaceae and related plant families, it accounts
for a considerable paralogy and even more efforts are needed to improve
performance and take advantage of sequence informativeness in nuclear
markers (Hershkovitz and Zimmer 1997; Gorelick 2002; Nyffeler and Eggli
2010; Korotkova et al. 2011).
The PCR conditions, concentrations, and custom thermal cycling pro-
files were adjusted to achieve high-quality amplification for each region.
The extension step (72°C) was set to 90 s for long fragments (1,000–1,500 bp)
and to 60 s for short fragments (400–800 bp). The standard thermal cycling
profile comprised an initial denaturation step (94°C / 60 s) followed by 30
to 34 cycles of denaturation (94°C / 30 s), annealing (variable temperature /
30 s), and extension (72°C / variable time) followed by a final extension
step (72°C / 10 min). Target regions were amplified in 25 or 50 mL volumes,
depending on whether only external or both internal and external se-
quencing primers were required (Table 3). The standard 25 mL polymerase
chain reactions contained PCR Buffer [13], BSA [16 mg/mL], MgCl2
[1.5 mM], mixed dNTPs [200 mM], each primer [0.1 mM], Platinum® Taq
polymerase (0.625 U per reaction; Invitrogen Corp., Carlsbad, California),
and DNA template [0.5 mg]. Unpurified PCR products were submitted to
the High-Throughput Genomics Center at the University of Washington
(Seattle, Washington) for Sanger sequencing. Chromatograms were as-
sembled and edited in Sequencher® 4.8 (GeneCodes Corp., Ann Arbor,
Michigan). The software BioEdit (Hall 1999) was used to stack individual
sequences for each region; the sequences were primary aligned using
ClustalW tool (Thompson et al. 1994). Alignments were improved with
MAFFT (Katoh and Standley 2013) using standard parameters and then
corrected by eye, ends were trimmed to remove low-quality data as re-
quired, and DNA regions concatenated in Mesquite 3.04 (Maddison and
Maddison 2015). Concatenation of DNA matrices was justified given that
the seven markers used came from the chloroplast, which is uniparentally
inherited and lacks recombination. Simple indel coding was applied to
maximize the number of informative sites in the molecular data matrix
(Simmons and Ochoterena 2000; Ochoterena 2009). Data matrices are ar-
chived in the Dryad Digital Repository (Tapia et al. 2017).
Structural Data—A selection of structural characters was scored for a
reduced set of taxa comprising one terminal for each of the fifteen species of
the Cephalocereus group, ten species representatives of other genera from
the tribe Echinocereeae, and one species from Leptocereeae. The characters
included were generated in the present study (Table 4) by examining
specimens obtained through fieldwork (2012–2015), herbaria (MEXU,
XAL), living collections (Botanic Gardens of Instituto de Biologı́a, UNAM),
and literature (Gibson and Horak 1978; Terrazas and Loza-Cornejo 2002;
Flores-Ortiz 2004; Vázquez-Sánchez et al. 2005, 2007; Arias and Terrazas
2006; Bárcenas-Argüello 2006). The structural matrix included morpho-
logical (35), anatomical (9), and chemical (2) characters (Appendix 2). Two
complex characters referred as differentiated flowering region and nec-
tarial chamber diaphragm were disaggregated into simple characters
(Table 4; characters 11–15 and 19–21, respectively). Bracketed numbers in
the following sections refer to the characters listed in Table 4. The structural
matrix was coded with binary and multistate unordered characters and
edited in Mesquite (Maddison and Maddison 2015).
Phylogenetic Analyses—Two matrices were constructed to run separate
analyses; the first matrix included DNA sequences and gaps (indel coding),
and the second comprised DNA sequences, gaps, and structural data
(combined matrix, 26 taxa). Molecular and combined matrices were
analysed using maximum parsimony (MP) and Bayesian inference (BI). For
the two matrices, parsimony ratchet analysis was conducted using PAUP
(Swofford 2002) from a file generated using PRAP2 (Müller 2004); the
number of iterations was 10,000, the ’Multrees’ option was disabled, the
initial value of ’Maxtrees’ was set to 100 and was auto-increased by 100;
the branch-swapping algorithm used was the tree-bisection-reconnection
(TBR). The trees with the best score were held in memory at each step, and
the strict consensus was saved from the gathered trees filtered using the
optimality criterion; Bootstrap (bts) and Jack-knife (jck) support were then
calculated from 10,000 replicates. The best-fit substitution model was
determined using the Bayesian information criterion (BIC), as implemented
in jModelTest (Darriba et al. 2012). The model selected by this method
(GTR 1 I 1 G) was applied to BI analyses for the entire molecular dataset,
and the MK model was applied only for structural data in the combined BI
analysis. All BI analyses were implemented in MrBayes (Huelsenbeck and
Ronquist 2001). The molecular matrix was partitioned into eight character
datasets, permitting the probability rates of the substitution model to vary
across partitions; all parameters were unlinked, four chains were run si-
multaneously for 10,000,000 generations, sampling parameters and trees
each 1,000 generations, and branch lengths were saved. The trees and
their respective support values were edited to improve visualization in
TreeGraph 2 (Stöver and Müller 2010). Optimization of unambiguous
characters was conducted using Winclada (Nixon 2002) on a maximum
clade credibility tree (BI) to recognize the contribution of the selected
structural characters in the phylogeny and the synapomorphies and ho-
moplasies that explain each recovered clade.
Taxonomic Studies—Original material at K, MEXU, MO, RSA, and US
was consulted for typification. A complementary search included B, G, M,

Table 3. Plastid regions amplified and primer sequences. T(a) 5 annealing temperature.

Plastid region Reference Primer sequences (50 -30 ) Expected size (bp) T(a)
rpl16 (Hernández-Hernández et al. 2011) F(1F): GCT ATG CTT AGT GTG TGA CTC GTT 900–1,200 63°C
R(3R): CTT CTA TTT GTC TAG GCG TGA TCC
trnL-F (Taberlet et al. 1991) F(c): CGA AAT CGG TAG ACG CTA CG 700–1,100 55°C
R(f): ATT TGA ACT GGT GAC ACG AG
F-int(e): GGT TCA AGT CCC TCT ATC CC
R-int(d): GGG GAT AGA GGG ACT TGA AC
psbA-trnH (Sang et al. 1997) F(psbA): GTT ATG CAT GAA CGT AAT C 300–700 52°C
(Tate and Simpson 2003) R(trnH2): CGC GCA TGG TGG ATT CAC A
rpl32-trnL (Shaw et al. 2007) F(rpl32): CAG TTC CAA AAA AAC GTA CTT C 500–1,400 52°C
R(trnLUAG): CTG CTT CCT AAG AGC AGC GT
(Plume et al. 2013) F-int: GTA ACT CTT GAA ATC ATT ATT TC
R-int: GTT ATC TTA GGT TTC AAC AAA CC
trnQ-rps16 (Shaw et al. 2007) F(trnQUUG): GCG TGG CCA AGY GGT AAG GC 600–1,500 52°C
R(rps16X1): GTT GCT TTY TAC CAC ATC GTT T
petL-psbE (Shaw et al. 2007) F(petL): AGT AGA AAA CCG AAA TAA CTA GTT A 400–700 52°C
R(psbE): TAT CGA ATA CTG GTA ATA ATA TCA GC
ycf1 (Franck et al. 2012) F(4182): AAA TAY RRA TAG AAA ATA TTT KGA TT 900–1,100 44°C
R(5248): GAA TTC TYA ATT CTC TAC GAC G
4 SYSTEMATIC BOTANY [Volume 42

Table 4. Character and character states for the morphological data matrix. [*] 5 characters mapped, but excluded from phylogenetic analyses.

Character\character states State 0 State 1 State 2 State 3

1. Apical dominance Monopodial Sympodial — —
2. Position of the first branch N/A Basitonic Mesotonic Acrotonic
3. Areole length (mm) 3–4 5–6 9–10 12
4. Interareolar distance (mm) 2–8 10–12 15–18 20–25
5. Stem podaria Absent Present — —
6. Interareolar depression Absent Present — —
7. Number of radial spines 3–7 9–11 13–15 20
8. Medial to radial spine ratio Equal . 1.5 — —
9. Central to radial spine ratio Minor Equal Major —
10. Radial coverage , 180° 180–270° . 270° —
11. Flowering region Apical Subapical Lateral —
12. Areole size (relative to vegetative region) Equal Larger — —
13. Spine number modification Diminish No changes Increase —
14. Spine consistency Rigid Flexible — —
15. Trichome density Equal Higher — —
16. Chlorenchyma Absent Present — —
17. Podaria evident Absent Present — —
18. Cephalium [*] Absent Present — —
19. Pseudocephalium [*] Absent Present — —
20. Ovary form Obovate depressed Obovate Wide obovate Ovate
21. Projection of axial tissue delimiting nectarial chamber Absent Present — —
22. Bend at the base of the first stamen row Absent Present — —
23. First stamen rows fused Free Adnate — —
24. Flower shape Short tubular Campanulate Tubular infundibuliform Long tubular
25. Spines in the flower Absent Present — —
26. Scale shape (receptacular tube) N/A Acute Obtuse —
27. Scale shape (pericarpel) N/A Acute Obtuse —
28. Scale margin Entire Erose — —
29. Scale apex form Obtuse Mucronate Acuminate —
30. Pericarpel with trichomes Scarce Abundant — —
31. Pulp color White Red — —
32. Spines in the fruit Absent Present — —
33. Podaria in the fruit Absent Present — —
34. Woody cap persistence Deciduous Persistent — —
35. Periclinal wall microrelief Not evident Minutely warty Verrucose —
36. Periclinal wall form Plain Concave Convex —
37. Keel Absent Present — —
38. Terpenoids Absent Present — —
39. Alkaloids Absent Present — —
40. Cuticle texture Smooth Grooved — —
41. Cuticle thickness Thin Thick Very thick —
42. Prismatic crystals in dermal system Absent Present — —
43. Stomata position Superficial Sunken — —
44. Hypodermal cellular layers Scarce Numerous Very numerous —
45. Hypodermal thickness Thin Thick — —
46. Vessel element density Scarce Numerous Very numerous —
47. Vessel element proportion (free/aggregated) Majority free Majority aggregated — —
48. Mucilage cells Absent Present — —

NY, and P, to ensure and recognize the existence of original material
studied here. The protologues of each published name were reviewed and
compared with the taxonomic information provided by Bravo-Hollis
(1978) and Hunt et al. (2006).
Results
Data Matrices and Parsimony Statistics—A preliminary
analysis on single markers did not show strongly supported
conflicts among different DNA regions (Wiens 1998), thus
allowing the concatenation of single marker matrices. The full
molecular matrix comprised 6,918 characters, including 59 gap
characters (Table 5). Mutational hotspot regions (524 charac-
ters) were excluded from the molecular dataset before con-
ducting the MP and BI analyses as suggested by Borsch and
Quandt (2009); excluded characters were mainly in ycf1 (454
characters). The total number of parsimony-informative sites
for the molecular matrix was 675; complete parsimony sta-
tistics for the seven markers and gaps are shown in Table 5.
The region with the greatest number of informative sites was
ycf1 despite the difficulty of achieving PCR amplification for
this region, followed by the rpl32-trnLUAG, rpl16, and trnL-F
regions, which were less difficult to amplify. Missing data
represent 3.6% of the molecular matrix and is mainly con-
centrated in three taxa (Cephalocereus nizandensis (Bravo &
T.MacDoug.) Buxb., Marshallocereus aragonii (F.A.C.Weber)
Backeb., and Harrisia earlei Britton & Rose) and in the regions
ycf1 and trnQ-rps16. The resolution obtained using data from
single molecular markers was very low, generating un-
resolved or unsupported topologies (data not shown) due to
low molecular divergence and the closeness of the taxa in-
cluded. The incorporation of seven chloroplast DNA regions
and gaps allowed us to consistently improve the resolution and
support values of the different analyses. Markers with the
highest variability rates were also the longest (ycf1, rpl32, trnL-F,
and rpl16), and the shorter markers exhibited less variability and
were less informative (psbA-trnH, petL-psbE and trnQ-rps16).
2017] TAPIA ET AL.: PHYLOGENY OF CEPHALOCEREUS 5

Table 5. Parsimony statistics for molecular data single marker, gaps, concatenated molecular matrices (MM), and combined (molecular 1
morphological) matrices (CM).

Dataset rpl16 trnL-F psbA-trnH rpl32-trnL trnQ-rps16 petL-psbE ycf1 gaps MM CM

Terminals 40 40 40 40 40 40 40 40 40 26
Length (bp) 1,159 1,096 550 1,380 603 573 1,498 59 6,918 6,964
Excluded characters 11 19 6 28 6 0 454 0 524 524
Indels per marker 4 9 4 23 1 3 15 — — —
Uninformative characters 100 87 46 137 36 38 179 0 623 549
Informative characters 60 44 34 159 16 11 292 59 675 448
Tree length (steps) 212 160 103 420 68 57 857 77 2011 1668
Consistency Index [CI] 0.840 0.850 0.883 0.824 0.838 0.947 0.699 0.805 0.757 0.691
Homoplasy Index [HI] 0.160 0.150 0.116 0.176 0.162 0.053 0.301 0.195 0.243 0.309
Retention Index [RI] 0.782 0.859 0.878 0.815 0.710 0.880 0.817 0.912 0.802 0.710
Rescaled Consistency Index [RC] 0.657 0.730 0.775 0.671 0.596 0.834 0.571 0.734 0.608 0.490
% informative 5.2 4.0 6.2 11.5 2.6 1.9 19.5 100 9.7 6.4

The combined matrix included also 46 structural characters, all
of which were parsimony-informative, but a reduction in the
number of informative characters it is observed in the entire
dataset, due to the reduced number of terminals retained in
this analysis (Table 5).
Phylogenetic Analyses—The MP analysis of the molecular
matrix resulted in three most-parsimonious trees, with a
length of 2,011 steps (Table 5; Consistency Index [CI] 5 0.7573,
Retention Index [RI] 5 0.8023, Rescaled Consistency Index
[RC] 5 0.6076). The strict consensus tree shows that Mar-
shallocereus, Carnegiea, and Bergerocactus Britton & Rose are
sister genera of the Cephalocereus group (Fig. 1A) with poor
support (MP: 52% bts, 60% jck). The monophyly of Cepha-
locereus group was recovered in the molecular analysis with
the highest support values (MP: 100% bts, 100% jck; Fig. 1A).
Within the Cephalocereus group, three clades show strong
support. Clade I includes N. euphorbioides and N. polylopha
(100% bts / 100% jck). Clade II includes nine species with low
support and two strongly supported subclades. Within sub-
clade II.1, Neobuxbaumia tetetzo is recovered as a sister species
of Cephalocereus senilis and C. columna-trajani (92% bts / 84%
jck); whereas in subclade II.2, N. mezcalaensis is recovered as a
sister species of N. squamulosa and N. multiareolata (98% bts /
95% jck). In addition, N. macrocephala is revealed as a sister
species of subclade II.2, whereas N. sanchezmejoradae and
Pseudomitrocereus fulviceps are sister species of both subclades
with low support. Clade III (subgen. Neodawsonia) includes C.
apicicephalium, C. nizandensis, and C. totolapensis (100% bts /
99% jck). The average standard deviation of split frequencies
(, 0.001) and the effective sample size (. 5,000) in the BI
analysis indicated convergence of the four independent runs.
The majority-rule consensus BI tree is congruent with the MP
tree and also recovered the Cephalocereus group and the three
clades that were observed in the MP analysis but added greater
support values (0.94–1.0 PP; Fig. 1B). Other relationships also
gained support; N. macrocephala appeared nested within clade
II as a sister of subclade II.2 (0.94 PP), and P. fulviceps appeared
as the first diversified taxon of clade II with high support
(1.0 PP).
The MP statistics for the combined data revealed slightly
lower values for consistency index, tree length, and number of
informative characters, compared with those produced for
molecular data alone (Table 5). Nonetheless, the topology for
this analysis (not shown) was mostly congruent with the other
molecular and combined analyses. The choice of the combined
BI tree for the optimization of the structural characters was
based on the better resolution offered by this analysis, and the
topological and supporting similarities for the shared clades
observed among the different data types and methods. The
optimization reveals two synapomorphies that support the
clade Cephalocereus: perianth woody cap [34], and prismatic
crystals in hypodermis and epidermis [42] (Fig. 2; Table 4).
Based on our results, the avoiding of para- or polyphyletic taxa
requires a new circumscription for the genus Cephalocereus,
which is given below.
Discussion
The results of our phylogenetic analyses employing mo-
lecular and combined data show that Cephalocereus is a
polyphyletic genus whereas Neobuxbaumia is paraphyletic,
because all species that comprise both genera are nested in a
single clade and mixed into the subclades. The present study
also reveals one deletion (49 bp) in the trnQ-rps16 region, one
deletion (11 bp) and one insertion (2 bp) in trnL-F, and several
base changes representing molecular synapomorphies for the
Cephalocereus group; moreover, crystals in the dermal system
[42] and the woody cap persistent in the fruit [34] are syna-
pomorphies, while the white pulp [31] is homoplastic (Fig. 2).
Therefore, we propose that Cephalocereus must be recognized
as a single natural genus and Neobuxbaumia and Pseudomi-
trocereus included within it (see Taxonomic Treatment). Cepha-
locereus (as recognized here) is a member of the Pachycereinae
subtribe, which also includes Bergerocactus, Carnegiea, Mar-
shallocereus Backeb., Pachycereus, Pterocereus Th. MacDoug. &
Miranda, Lemaireocereus, and Deamia Britton & Rose. The re-
lationship of Cephalocereus and Carnegiea is partially accepted
if it is recognized that Carnegiea is closely related to Pachycereus
and Marshallocereus (Fig. 1).
Cephalocereus and its Interspecific Relationships—The
combined analyses indicated that Neobuxbaumia scoparia is
sister to the remaining species of Cephalocereus group (Figs. 1, 2).
Distinctive attributes are: plants highly branched, sympodial
[1], subapical flowering region not differentiated [11], and
flower and fruit lacking spines [25, 32] (Fig. 3A–B). The dis-
tribution of this species is disjunct in Veracruz (near Jalapa)
and Oaxaca (Tehuantepec isthmus), growing over volcanic or
limestone rocks and inhabiting tropical dry forest (Dawson
1952b; Bravo-Hollis 1978). Three species of Cephalocereus
subgen. Neodawsonia form a well-supported clade in the mo-
lecular and combined analyses (clade III; Figs. 1, 2). This clade
contains an insertion of one triplet in ycf1 and several punctual
mutations as molecular synapomorphies; the morphological
synapomorphy is the apical pseudocephalium [19] (Fig. 3C–D;
6 SYSTEMATIC BOTANY [Volume 42

Fig. 1. Phylogenetic trees for molecular dataset based on: A. Maximum parsimony strict consensus for 3 most parsimonious trees from molecular
analysis, showing support values for bootstrap (above branches), and jackknife (below branches). B. Maximum clade credibility tree for BI analysis, posterior
probabilities are shown above branches.

Vázquez-Sánchez et al. 2005; Bárcenas-Argüello 2006). This
finding corroborates the monophyly of Neodawsonia that was
suggested by Bárcenas-Argüello (2006) based on morphological
data. Neodawsonia was erected as a genus by Backeberg (1960)
and was later transferred to a subgenus of Cephalocereus by
Bravo-Hollis (1974) as accepted by Heath (1992) and Arias et al.
(2012); however, other authors dismissed any infrageneric
classification (Gibson and Horak 1978; Anderson 2001; Hunt
et al. 2006), a conclusion we support based on our results. These
three species are distributed from Tehuantepec isthmus (Oaxaca)
to the Central Depression (Chiapas) in southeastern Mexico, and
partially cohabit with N. scoparia (e.g. Juchitan, Oaxaca). These
species inhabit tropical deciduous forest at elevations below
700 m, growing mainly over rocky and shallow substrates
(Bravo-Hollis 1978; Bárcenas-Argüello et al. 2010).
Neobuxbaumia euphorbioides and N. polylopha conform a clade
with strong support (clade I; Fig. 1). These two species have
three insertions in the psbA-trnH region as molecular syna-
pomorphies and two homoplastic characters: monopodial
stems [1] and a mucronate scale apex [29] (Fig. 2). The
monopodial growth form is shared with other taxa from
different clades (e.g. N. mezcalaensis), indicating some degree
of convergent evolution. Moreover, they show contrasting
variation: N. polylophus have up to 40 ribs and up to 7 radial
spines [7], whereas N. euphorbioides has only 8–10 ribs and
three radial spines in the vegetative areole (Tapia et al. 2016;
Fig. 3E–I). This association supports the proposal of Buxbaum
(1961), who suggested a relationship between N. euphorbioides
and N. polylopha based on morphology. These two species
inhabit the Sierra Madre Oriental from Guanajuato, Hidalgo,
Queretaro, and San Luis Potosı́ and extending to Veracruz and
Tamaulipas in association with tropical deciduous forests at
elevations between 900 and 1,500 m and are markedly asso-
ciated with rocky substrates (Bravo-Hollis 1978).
2017] TAPIA ET AL.: PHYLOGENY OF CEPHALOCEREUS 7

Fig. 2. BI tree from combined (molecular 1 morphological) data. Bold lines represent 1.0 posterior probabilities; the lower values appear annotated
above branches. Character optimization of unambiguous changes is represented over branches: characters are designated with numbers (according with
Table 4) and the character state is annotated below. Synapomorphies are highlighted with bold numbers and brackets.

Cephalocereus columna-trajani, C. senilis, Neobuxbaumia mac-
rocephala, N. mezcalaensis, N. multiareolata, N. squamulosa,
N. sanchezmejoradae, and N. tetetzo represent the most inclusive
and strongly supported clade that was identified within the
genus (clade II; Figs. 1, 2, 3J–Q). Molecular synapomorphies
correspond only to punctual mutations in ycf1, and mor-
phological synapomorphies are absent. Our results indicate
that C. columna-trajani, C. senilis, and N. tetetzo constitute an
unexpected subclade (II.1) that is strongly supported, because
this clade comprises the type species for Neobuxbaumia
(N. tetetzo) and Cephalocereus (C. senilis). Molecular synapo-
morphies for this subclade are only punctual mutations,
whereas the evident podaria in the reproductive region [17]
is a morphological synapomorphy. Otherwise, the lateral
cephalium [18] represents a synapomorphy for C. columna-
trajani and C. senilis (Fig. 3M–N). The hypotheses suggested by
Bárcenas-Argüello (2006) that relate subgenera Cephalocereus
and Neodawsonia have not been confirmed by this study.
Moreover, the differentiated flowering regions (cephalium
and pseudocephalium) show a pattern of convergent evolu-
tion based on the unique and distinctive attributes that define
each of these regions, as is found in Backebergia militaris and
other less-related taxa (Vázquez-Sánchez et al. 2016). Even
among the morphological characters evaluated here, none is
shared by these two subgenera. For example, the lateral
cephalium (only shared by species of subgen. Cephalocereus)
appeared as non-homologous to the apical pseudocephalium
of subgen. Neodawsonia (Vázquez-Sánchez et al. 2007). Whereas
C. columna-trajani and N. tetetzo inhabit the Tehuacan-Cuicatlan
Valley (the latter occurs even in the Central Valleys, Oaxaca),
only C. senilis inhabits a limited region in the Sierra Madre
Oriental (Hidalgo and Veracruz). In fact, putative naturally
hybridizing populations between C. columna-trajani and
N. tetetzo have been reported in the Tehuacan-Cuicatlan
Valley (Vite et al. 1996). These three species occur in sea-
sonal thorn and tropical deciduous forests at elevations be-
tween 900 and 2200 m (Bravo-Hollis 1978; Arias et al. 2012).
The other well-supported subclade comprises Neobuxbaumia
mezcalaensis, N. multiareolata, and N. squamulosa (subclade II.2;
Figs. 1, 2). Molecular synapomorphies consist of only punctual
mutations, whereas the presence of spines on pericarpel and
receptacular tube [25] is a homoplastic character that is shared
by members of this subclade. Historically, N. mezcalaensis and
N. multiareolata have been recognized as closely related taxa
(Dawson 1948; Bravo-Hollis et al. 1973; Arroyo-Cosultchi
et al. 2010); however, the molecular and combined analyses
performed here suggest that N. multiareolata and N. squamulosa
are sister species and that N. mezcalaensis is sister of both.
Neobuxbaumia squamulosa was described by Scheinvar and
Sánchez-Mejorada (1990) and has been recognized as the valid
taxon; nonetheless, during our previous study on the identi-
fication of Neobuxbaumia species (Tapia et al. 2016), we found
the same taxon in the Tehuantepec isthmus that had been
previously described in absence of flowers and fruits by
Dawson (1948) as Cephalocereus nudus (see Taxonomic treat-
ment; Fig. 3O–Q). Species of this subclade have a wider
8 SYSTEMATIC BOTANY [Volume 42

Fig. 3. Vegetative and reproductive morphology of Cephalocereus (see Taxonomic Treatment). A. Sympodial growth of C. scoparius. B. Subapical flower
buds of C. scoparius. C. Monopodial growth of C. totolapensis. D. Fruits and apical pseudocephalium of C. totolapensis. E. Monopodial growth of C. polylophus.
F. Subapical flowers of C. polylophus. G. Longitudinal section of flower, C. polylophus. H. Monopodial growth of C. euphorbioides. I. Flowers of C. euphorbioides.
J. Sympodial growth of C. tetetzo. K. Apical flower buds and podaria of C. tetetzo. L. Basitonic branching of C. senilis. M. Lateral cephalium of C. senilis.
N. Lateral cephalium and flower of C. columna-trajani. O. Sympodial growth of C. nudus. P. Apical fruits of C. nudus. Q. Flower of C. nudus. R. Sympodial
growth of C. fulviceps. S. Apical pseudocephalium and flowers of C. fulviceps. T. Flower of C. fulviceps. Scale bar 5 5 cm.
2017] TAPIA ET AL.: PHYLOGENY OF CEPHALOCEREUS
geographic distribution, occupying the provinces of Rio Balsas
Basin and Costa Pacifica Mexicana (partially Colima, Jalisco,
Michoacan, Guerrero, Morelos, Puebla, and Oaxaca; Morrone
2005).
Neobuxbaumia macrocephala and N. sanchezmejoradae belong
to clade II but exhibit low affinity to other species in the
subclades II.1 and II.2, in both molecular and combined an-
alyses. Neobuxbaumia macrocephala has been previously linked
to N. tetetzo because they share growth form [1, 2], spination
pattern [7–10] (Tapia et al. 2016), and (partially) geographic
distribution (Bravo-Hollis 1978; Arias et al. 2012). The apical
flowering region of N. macrocephala is slightly modified with
pinkish-black flexible spines, spineless flowers, and fruits with
few flexible spines distinguish this species. Based on our re-
sults, this species is sister of subclade II.2, sharing only
punctual mutations. Molecular synapomorphies consist of
punctual mutations in ycf1, and no morphological synapo-
morphies were observed. These species are geographically
restricted to Mixteca Baja in Oaxaca (N. sanchezmejoradae) and
Tehuacan-Cuicatlan Valley, between Puebla and Oaxaca
(N. macrocephala) (Hunt et al. 2006; Arias et al. 2012).
Pseudomitrocereus fulviceps is resolved as a sister to the
remaining species in clade II, although support varied from
weak to strong (Figs. 1, 2). This species has a large number of
exclusive molecular and morphological characters, such as a
woolly apical differentiated flowering region [19], a flower
completely covered by large trichomes, and a wide pericarpel
(Arias et al. 2012; Fig. 3R–T). The significant number of
autapomorphies found in this taxon has led to an unstable
taxonomic assignment and to its transfer through several
genera including Mitrocereus (Backeberg 1938, 1960; Bravo-
Hollis 1954), Pseudomitrocereus (Buxbaum 1961; Hernández-
Ledesma et al. 2015), Cephalocereus (Moore 1975), Pachycereus
(Britton and Rose 1920; Hunt and Taylor 1991), and Carnegiea
(Heath 1992). Previously this species was certainly included as
an ally of the Cephalocereus group (Arias et al. 2003) based on
molecular data, this finding was supported also by morpho-
logical data (Terrazas and Loza-Cornejo 2002; Arias and
Terrazas 2006) and is confirmed by this study. Particular in-
terest has been directed to the extreme variation found in the
flower of Pseudomitrocereus fulviceps: flower shape and size,
trichomes long and abundant, and bristles in the flower are-
oles; these characters were interpreted in previous studies as
the retention of ancestral features that are also present in the
sister Pachycereus group. Such discordance expresses the ne-
cessity for alternative explanations for dichotomic speciation
concepts (Hausdorf 2011). Among others, the hypotheses of
reticulate evolution and hybridization remain to be tested, as
for other Cactaceae species and in species of other plant
families (Vite et al. 1996; Seehausen 2004). We suggest that a
particular study focusing on the hypothesis of non-dichotomous
evolutionary processes, in which its cohabiting related species
(Neobuxbaumia tetetzo, N. macrocephala, Cephalocereus columna-
trajani) are included, may help to explain the origin of such
distinctive features of P. fulviceps.
Contribution of the Molecular and Structural Data—The
assessment of combined data matrices (molecular 1 structural)
revealed an increase in resolution and support values for some
nodes over congruent topologies, relative to those produced
with molecular data alone. The overall consistency index was
higher in molecular than in combined dataset, indicating a
higher amount of homoplasy in the structural data, as has been
noted in previous studies (e.g. Ritz et al. 2007). The plastid
9
markers employed in this study were selected from a group of
regions reported in the literature (Butterwhorth et al. 2002;
Cúenoud et al. 2002; Nyffeler 2002; Arias et al. 2003, 2005;
Shaw et al. 2005, 2007; Korotkova et al. 2010, 2011; Bárcenas
et al. 2011; Hernández-Hernández et al. 2011; Franck et al.
2012; Vázquez-Sánchez et al. 2013; Sánchez et al. 2014), their
contribution to variability was ycf1.rpl32-trnL.rpl16.psbA-
trnH.trnL-F.trnQ-rps16.petL-psbE (in decreasing order);
and the order of informativeness was ycf1.rpl32-trnL.psbA-
trnH.rpl16.trnL-F.trnQ-rps16.petL-psbE. Here the psbA-
trnH region showed an important contribution other than its
length as reported by Lahaye et al. (2008) and the CBOL Plant
Working Group (2009) (Table 5). The eight diagnostic char-
acters referred by previous authors (Table 1), were recovered
in this analysis as unreliable or inconsistent with the dis-
tinction of three genera, because these characters cannot
produce a grouping unambiguously. The disaggregation of
complex characters, as cephalium [characters 11 to 17] or dia-
phragm [characters 21 to 23] (Table 4), provided a more ac-
curate and sharp depiction of species; nonetheless, this also has
failed to produce groups consistent with the three genera
scheme. Among the 46 structural characters incorporated, five
vegetative attributes persisted as homoplasious (e.g. interar-
eolar distance, number of radial spines, and coverage of radial
spines). This subset of morphological characters has been
noted several times as being unreliable for defining natural
groups among the tribe Echinocereeae (Gibson and Horak
1978) or entirely homoplasious (Terrazas and Loza-Cornejo
2002; Arias and Terrazas 2006; Bárcenas-Argüello 2006). Ten
other homoplasious characters were identified in the repro-
ductive region of the stems (e.g. the position of the flowering
region), flower (e.g. the presence of spines), fruit (e.g. pulp
colour), and seed (e.g. the presence of a keel). Indeed, some of
these characters also behaved as homoplasious in a phylo-
genetic analysis of Pachycereus by Arias and Terrazas (2006),
which includes species of Cephalocereus. Evident podaria
and chlorenchyma loss in the flowering region represent
synapomorphies for C. senilis 1 C. columna-trajani. These
characters represent obligate modifications in the flowering
region to be recognized as cephalium; these characters are
missing in the flowering regions called pseudocephalia,
indicating a non-homologous but convergent origin for these
complex characters, as already suggested by Vázquez-Sánchez
et al. (2005, 2016). Our results confirm the hypothesis of Buxbaum
(1964a, b; and supported by Terrazas and Loza-Cornejo 2002)
that cephalium yields no phylogenetic information because
it has evolved several times within Cactoideae. Prismatic
crystals in the dermal system and a woody cap derived from
perianth remnants have been confirmed as synapomorphies
for the genus Cephalocereus as defined here. These characters
were previously recognized as synapomorphies for some
species of Cephalocereus (Terrazas & Loza-Cornejo 2002), but
here we extend the observations to include all fifteen species
of this group and corroborate that finding. Further studies
on this group will explore speciation, estimate the di-
vergence time between taxa, and examine the geographic
patterns of speciation and the inheritance of geographic
ranges.
Taxonomic Treatment
The fifteen accepted species in genus Cephalocereus based
on the results presented here are listed below. Two new
10 SYSTEMATIC BOTANY
combinations are included; in addition four lectotypes, nine
neotypes, and a new synonym are proposed.
CEPHALOCEREUS Pfeiff., Allg. Gartenzeitung 6(18): 142. 1838.—
LECTOTYPE (designed by Britton & Rose, Contr. U.S.
Natl. Herb. 12: 415. 1909): Cactus senilis Haw.
Haseltonia Backeb., Blätt. Sukkulentenk. 1: 3. 1949.—TYPE:
Pilocereus hoppenstedtii Roezl ex Rümpler. Cephalocereus
hoppenstedtii (Roezl ex Rümpler) K. Schum. Cereus hoppenstedtii
(Roezl ex Rümpler) A. Berger. Haseltonia hoppenstedtii (Roezl
ex Rümpler) Backeb.
Neobuxbaumia Backeb., Blätt. Kakteenf. 6. s.p. 1938.—TYPE:
Cereus tetazo F.A.C. Weber ex J. M.Coult. Carnegiea tetazo
(F.A.C. Weber ex J.M. Coult.) P.V.Heath. Cephalocereus
tetetzo (F.A.C. Weber ex J.M. Coult.) Diguet. Pilocereus
tetetzo F.A.C. Weber ex K. Schum. Cephalocereus tetetzo
(F.A.C. Weber ex K. Schum.) Vaupel. Pachycereus tetetzo
(F.A.C. Weber ex K. Schum.) Ochot. Neobuxbaumia
tetetzo (F.A.C. Weber ex K. Schum.) Backeb.
Pilocereus Lem., Cact. Gen. Sp. Nov. 6. 1839.—TYPE: Cactus
senilis Haw. Cereus senilis (Haw.) DC. Cephalophorus senilis
(Haw.) Lem. Cephalocereus senilis (Haw.) Pfeiff. Echino-
cactus senilis (Haw.) Beaton. Pilocereus senilis (Haw.) Lem.
Cephalocereus senilis (Haw.) K. Schum. [non Pilocereus
K. Schum.]
Pseudomitrocereus Bravo & Buxb., Bot. Stud. 12: 49, 53. 1961.—
TYPE: Pilocereus fulviceps F.A.C. Weber ex K. Schum.
Cereus fulviceps (F.A.C. Weber ex K. Schum.) A. Berger.
Mitrocereus fulviceps (F.A.C. Weber ex K. Schum.) Backeb.
ex Bravo. Pseudomitrocereus fulviceps (F.A.C. Weber ex
K. Schum.) Bravo & Buxb. Cephalocereus fulviceps (F.A.C.
Weber ex K. Schum.) H.E. Moore. Pachycereus fulviceps
(F.A.C. Weber ex K. Schum.) D.R. Hunt. Carnegiea fulviceps
(F.A.C. Weber ex K. Schum.) P.V. Heath [non Cephalocereus
subgen. Mitrocereus Backeb. Mitrocereus (Backeb.) Backeb.]
Carnegiea subgen. Rooksbya Backeb., Cact. Succ. J. (Los
Angeles) 22(5): 154. 1950. Rooksbya (Backeb.) Backeb.,
Cactaceae (Backeberg) 4. 2165. 1960.—TYPE: Cereus
euphorbioides Haw. Cactus euphorbioides (Haw.) Spreng.
Pilocereus euphorbioides (Haw.) Rümpler. Cephalocereus
euphorbioides (Haw.) Britton & Rose. Lemaireocereus
euphorbioides (Haw.) Werderm. Carnegiea euphorbioides
(Haw.) Backeb. Neobuxbaumia euphorbioides (Haw.) Buxb.
Rooksbya euphorbioides (Haw.) Backeb.
1. CEPHALOCEREUS APICICEPHALIUM E.Y. Dawson, Allan Hancock
Found. Publ. Occas. Pap. 1: 10. 1948. Neodawsonia apici-
cephalium (E.Y. Dawson) Backeb., Blätt. Sukkulentenk.
1: 4. 1949.—TYPE: MEXICO. Oaxaca: from lava cliffs and
hills 17 km west of Tehuantepec, 27 Jan 1947, E. Y. Dawson
2993 (holotype: RSA! liquid).
Neodawsonia guengolensis Bravo, Anales Inst. Biol. Univ. Nac.
Mexico 27: 12. 1956.—TYPE: MEXICO. Oaxaca, cerro de
Gu[i]engola, en suelo de lava, H. Bravo s.n. (holotype:
MEXU, not preserved). LECTOTYPE (designated here):
[illustration] “Ejemplar de Neodawsonia guengolensis sp.
nov. del cerro Gu[i]engola” in Bravo, Anales Inst. Biol.
Univ. Nac. Mexico 27(1): 14, Figure 6. 1956. EPITYPE
(designated here): MEXICO. Oaxaca: cerro Guiengola,
1966, H. Bravo Hollis s.n. (MEXU 501508!)
[Volume 42
Neodawsonia nana Bravo, Anales Inst. Biol. Univ. Nac. Mexico
27: 15. 1956 —TYPE: MEXICO. Oaxaca, Agua Caliente, en
roca caliza, H. Bravo s.n. (holotype: MEXU, not preserved).
LECTOTYPE (designated here): [illustration] “Neo-
dawsonia nana sp. nov. Ejemplares creciendo en La Ventosa”
in Bravo, Anales Inst. Biol. Univ. Nac. Mexico 27(1): 15,
Figure 7. 1956. EPITYPE (designated here): Oaxaca, 15 km
al NW de Tehuantepec, creciendo en cantiles con Ficus,
1500 m, 19 Feb 1978, O. Téllez s.n. (MEXU 235436!).
2. CEPHALOCEREUS COLUMNA-TRAJANI (Karw. ex Pfeiff.) K. Schum.,
Gesamtbeschr. Kakt. 198. 1897. Cereus columna-trajani
Karw. ex Pfeiff., Enum. Diagn. Cact. 76. 1837. Cepha-
lophorus columna-trajani (Karw. ex Pfeiff.) Lem., Cact. Aliq.
Nov. Desc. p. 1831. Pilocereus columna-trajani (Karw. ex
Pfeiff.) Lem., Cact. Gen. Sp. Nov. 9. 1839. Cephalocereus
columna (Karw. ex Pfeiff.) K. Schum., Nat. Pflanzenfam.
3(6a): 182. 1894. Pachycereus columna-trajani (Karw. ex
Pfeiff.) Britton & Rose, Contr. U.S. Natl. Herb. 12(10): 421.
1909. Mitrocereus columna-trajani (Karw. ex Pfeiff.) Backeb.
ex E.Y. Dawson, Allan Hancock Found. Publ. Occas. Pap.
1: 4. 1948. Haseltonia columna-trajani (Karw. ex Pfeiff.)
Backeb., Cactaceae (Backeberg) 4: 2263. 1960.—TYPE:
MEXICO. [Puebla]: San Sebastian [Zinacatepec], W. F. von
Karwinsky (not preserved). NEOTYPE (designated here):
MEXICO. Oaxaca: Coixtlahuaca, Tepelmeme Villa de
Morelos, aproximadamente 1 km S del Rı́o Santa Lucı́a,
1285 m, 27 Mar 1994, U. Guzmán Cruz 1065 (MEXU
280206!).
Pilocereus hoppenstedtii Roezl ex Rümpler, Handb. Cacteenk.
667. 1886. Cephalocereus hoppenstedtii (Roezl ex Rümpler)
K. Schum., Nat. Pflanzenr. 3(6a): 181. 1894. Cereus hop-
penstedtii (Roezl ex Rümpler) A. Berger, Rep. (Annual)
Missouri Bot. Gard. 16: 70. 1905. Haseltonia hoppenstedtii
(Roezl ex Rümpler) Backeb., Sukkulentenkunde 1: 3.
1949.—TYPE: MEXICO. B. Roezl s.n. (not preserved).
NEOTYPE (designated here): MEXICO. Puebla, Mpio.
Coxcatlán, 1 km al noroeste de San José Tilapa, 780 m, 21
Jul 1993, S. Arias & U. Guzmán 1112 (MEXU 771928!).
3. CEPHALOCEREUS EUPHORBIOIDES (Haw.) Britton & Rose, Cact. 2:
33. 1920. Cereus euphorbioides Haw. Suppl. Pl. Succ. 75.
1819. Cactus euphorbioides (Haw.) Spreng., Syst. Veg.
[Sprengel] 2: 496. 1825. Pilocereus euphorbioides (Haw.)
Rümpler, Handb. Cacteenk. 658. 1886. Lemaireocereus
euphorbioides (Haw.) Werderm., Repert. Spec. Nov. Regni
Veg. Sonderbeih. C t. 84. 1934. Carnegiea euphorbioides
(Haw.) Backeb., Cact. Succ. J. (Los Angeles) 23(4): 121.
1951. Neobuxbaumia euphorbioides (Haw.) Buxb., Cactus
(Paris) 40: 52. 1954. Rooksbya euphorbioides (Haw.) Backeb.,
Cactaceae (Backeberg) 4: 2167. 1960 —TYPE: not cited
(not located). NEOTYPE (designated here): MEXICO.
Veracruz: Actopan, 7.9 km al NW de Trapiche del Rosario,
710 m, 3 Mar 2012, H. J. Tapia 01 (MEXU 1402251!).
Cereus olfersii Salm-Dyck, Hort. Dyck. 335. 1834. Rooksbya
euphorbioides var. olfersii (Salm-Dyck) Backeb., Cactaceae
(Backeberg) 4: 2170. 1960. Neobuxbaumia euphorbioides var.
olfersii (Salm-Dyck) Bravo, Las Cact. Méx. 1: 660. 1978.
Carnegieae euphorbioides var. olfersii (Salm-Dyck) P.V.
Heath, Calyx 2(3): 108. 1992 —NEOTYPE (designated
here): MEXICO. Veracruz: Palo Gacho, 25 Apr 1970,
H. Sánchez-Mejorada 70–0407 (MEXU 133842!).
2017] TAPIA ET AL.: PHYLOGENY OF CEPHALOCEREUS
4. CEPHALOCEREUS FULVICEPS (F.A.C. Weber ex K. Schum.) H.E.
Moore, Baileya 19: 165. 1975. Pilocereus fulviceps F.A.C.
Weber ex K. Schum., Gesamtbeschr. Kakt. 176. 1897.
Cereus fulviceps (F.A.C. Weber ex K. Schum.) A. Berger,
Rep. (Annual) Missouri Bot. Gard. 16: 64. 1905. Mitro-
cereus fulviceps (F.A.C. Weber ex K. Schum.) Backeb. ex
Bravo, Anales Inst. Biol. Univ. Nac. Mexico 24: 232. 1954.
Pseudomitrocereus fulviceps (F.A.C. Weber ex K. Schum.)
Bravo & Buxb., Bot. Stud. 12: 99. 1961; Pachycereus ful-
viceps (F.A.C. Weber ex K. Schum.) D.R. Hunt, Bradleya
9: 89. 1991. Carnegiea fulviceps (F.A.C. Weber ex K. Schum.)
P.V. Heath, Calyx 2(3): 108. 1992.—TYPE: MEXICO.
Puebla, Tehuacan, F.A.C. Weber s.n. (not located).
NEOTYPE (designated here): MEXICO. Puebla: Zapotitlán
Salinas, 11 km sobre la terracerı́a a San Luis Atolotitlán,
1800 m, 8 Aug 1990, S. Arias 863 (MEXU 1302119!;
isoneotypes, MEXU 1301778!, 1301777!).
5. CEPHALOCEREUS MACROCEPHALUS F.A.C. Weber ex K. Schum.,
Gesamtbeschr. Kakt. 197. 1897. Pilocereus macrocephalus
(F.A.C. Weber ex K. Schum.) F.A.C. Weber, Dict. Hort.
[Bois] 2: 966. 1898. Cereus macrocephalus (F.A.C. Weber ex
K. Schum.) A. Berger, Rep. (Annual) Missouri Bot. Gard.
16: 62. 1905. Neobuxbaumia macrocephala (F.A.C. Weber ex
K. Schum.) E.Y. Dawson, Cact. Succ. J. (Los Angeles)
24: 173. 1952. Carnegiea macrocephala (K.Schum.) P.V.
Heath, Calyx 2(3): 109. 1992 —TYPE: MEXICO. [Puebla]:
Tehuacan (not located). NEOTYPE (designated here):
MEXICO. Puebla: Zapotitlán Salinas, 15 km W de
Zapotitlán Salinas, 1575 m, 12 Apr 1970, H. Sánchez-
Mejorada 70–0401 (MEXU 133824!).
Pilocereus ruficeps F.A.C. Weber ex Rol.-Goss., Bull. Mus. Hist.
Nat. (Paris) 11(6): 509. 1905. Cereus ruficeps (F.A.C. Weber
ex Rol.-Goss.) Vaupel, Monatsschr. Kakteenk. 23: 27, 83.
1913. Pachycereus ruficeps (F.A.C. Weber ex Rol.-Goss.)
Britton & Rose, Cactaceae (Britton & Rose) 2: 75. 1920.
Mitrocereus ruficeps (F.A.C. Weber) Backeb., Cactaceae
(Backeberg) 4: 2244. 1960.—TYPE: MEXICO. [Puebla]:
Tehuacan, Cerro de San Jose Tochapa, L. Diguet s.n.
(isotype: US barcode 117101!).
6. CEPHALOCEREUS MEZCALAENSIS Bravo, Anales Inst. Biol. Univ.
Nac. Mexico, 3: 378. 1932. Pilocereus mezcalaensis (Bravo)
W.T. Marshall, Cactaceae 75. 1941. Neobuxbaumia mezca-
laensis (Bravo) Backeb., Beitr. Sukkulentenk. Sukku-
lentenpflege 3. 1941. Carnegiea mezcalaensis (Bravo) P.V.
Heath, Calyx 2(3): 109. 1992 —LECTOTYPE (designated
by Scheinvar & Olalde, 2008): [illustration] “Grupo de
Cephalocereus mezcalaensis sp. n. caracterı́stico del Ca~
nón
del Zopilote” in Bravo, Anales Inst. Biol. Univ. Nac.
Mexico, 3: 382. Figure 8. 1932. EPITYPE (designated here):
MEXICO. Guerrero, Mpio. Eduardo Neri, Casa Verde, a
2 km al O en carretera a Xochipala, 800 m, 04 Jun 1980,
H. Kruze 2930 (MEXU 1254733!)
Cephalocereus mezcalaensis Bravo var. robustus E.Y. Dawson,
Allan Hancock Found. Publ. Occas. Pap. 1: 9. 1948. Neo-
buxbaumia mezcalaensis var. robusta (E.Y. Dawson) Backeb.,
Cact. Succ. J. (Los Angeles) 23: 122. 1951. Neobuxbaumia
mezcalaensis var. robusta (E.Y. Dawson) E.Y. Dawson, Cact.
Succ. J. (Los Angeles) 24: 169. 1952.—TYPE: Mexico.
Puebla: from hillsides of the Ca~ non del Rio Atoyac, 6 Jan
1947, E. Y. Dawson 2998 (holotype: RSA 0068734!).
11
7. Cephalocereus multiareolatus (E.Y. Dawson) H.J. Tapia &
S. Arias comb. nov. Cephalocereus mezcalaensis Bravo var.
multiareolatus E.Y. Dawson, Allan Hancock Found. Publ.
Occas. Pap. 1: 9. 1948. Neobuxbaumia mezcalaensis var.
multiareolata (E.Y. Dawson) E.Y. Dawson, Cact. Succ.
J. (Los Angeles) 24: 169. 1952. Neobuxbaumia multiareolata
(E.Y. Dawson) Bravo, Scheinvar & Sánchez-Mej., Cact.
Suc. Mex. 17(4): 120. 1972. Carnegiea mezcalaensis var.
multiareolata (E.Y. Dawson) P.V. Heath, Calyx 2(3): 109.
1992.—TYPE: Mexico. Guerrero: from lava hills near
Acahuizotl[a], 1 Feb 1947, E. Y. Dawson 2996 (holotype:
RSA liquid!).
8. CEPHALOCEREUS NIZANDENSIS (Bravo & T. MacDoug.) Buxb.,
Kakteen and. Sukk. 16: 45. 1965. Neodawsonia nizandensis
Bravo & T. MacDoug., Anales Inst. Biol. Univ. Nac.
Mexico 29: 82. 1959.—TYPE: MEXICO. Oaxaca, Nizanda,
H. Bravo s.n. (not preserved). LECTOTYPE designated
here: [illustration] “Neodawsonia nizandensis, creciendo en
el cerro Nizanda” in Bravo & MacDougall, Anales Inst.
Biol. Univ. Nac. Mexico 29: 84. Figure 9. 1959. EPITYPE
(designed here): MEXICO. Oaxaca, Nizanda, 1958,
H. Bravo s.n. (MEXU 505145!).
9. CEPHALOCEREUS NUDUS E.Y. Dawson, Allan Hancock Found.
Publ. Occas. Pap. 1: 12. 1948. Cephalocereus tetetzo var.
nudus E.Y. Dawson, Desert Pl. Life 24(3): 54. 1952. Neo-
buxbaumia tetetzo (F.A.C. Weber ex K. Schum.) Backeb.
var. nuda (E.Y. Dawson) E.Y. Dawson, Cact. Succ. J. (Los
Angeles) 24: 169. 1952. Carnegiea tetetzo var. nuda (E.Y.
Dawson) P.V. Heath, Calyx 2(3): 108. 1992.—TYPE:
MEXICO. Oaxaca: Tehuantepec, 1947, E. Y. Dawson 3002
(holotype: RSA liquid!).
Neobuxbaumia squamulosa Scheinvar & Sánchez-Mej., Cact. Suc.
Mex. 35: 13. 1990 syn. nov.—TYPE: Mexico. Michoacan:
Coalcoman, 19 Oct 1987, H. Sánchez-Mejorada 4250 (holo-
type: MEXU barcode 474887!).
10. CEPHALOCEREUS POLYLOPHUS (DC.) Britton & Rose, Contr.
U.S. Natl. Herb. 12(10): 419. 1909. Cereus polylophus DC.,
Mém. Mus. Hist. Nat. 17: 115. 1828. Pilocereus polylophus
Salm-Dyck, Cact. Hort. Dyck. 24; ed. II. 1844. Neo-
buxbaumia polylopha (DC.) Backeb., Beitr. Sukkulentenk.
Sukkulentenpflege. 1941: 3. Carnegiea polylopha (DC.) D.R.
Hunt, Bradleya 6: 100. 1988.—TYPE: MEXICO. Hidalgo,
Thomas Coulter 15 (not located). NEOTYPE (designated
here): MEXICO. Hidalgo: Metztitlan, H. Sánchez-Mejorada
70–0406 (MEXU 133840!).
11. Cephalocereus sanchezmejoradae (A.B. Lau) H.J. Tapia &
S. Arias comb. nov. Neobuxbaumia sanchezmejoradae A.B.
Lau, Cact. Suc. Mex. 39: 3. 1994.—TYPE: MEXICO.
Oaxaca: Santiago Nuyoó, Tierra Azul, sobre formaciones
de roca volcánica erosionada, 1600 m, May 1976, A. B. Lau
1378 (holotype: MEXU 01291728!; isotype MEXU 1291729!).
Carnegiea laui P.V. Heath, Calyx 2(3): 109. 1992. Neobuxbaumia
laui (P.V. Heath) D.R. Hunt, Cactaceae Consensus Init.
4: 5. 1997.—TYPE: [illustration] “Stem of the new Neo-
buxbaumia from Teotila”, in Lau, Brit. Cact. Succ. J. 4(4):
101. Figure 3. 1986 (holotype).
Carnegiea nova P.V. Heath, Calyx 2(3): 109. 1992.—TYPE:
[illustration] cover illustration, in Brit. Cact. Succ. J. 4(4):
cover. 1986 (holotype).
12 SYSTEMATIC BOTANY [Volume 42

12. CEPHALOCEREUS SCOPARIUS (Poselg.) Britton & Rose, Contr. 14. CEPHALOCEREUS TETETZO (F.A.C. Weber ex J.M. Coult.)
U.S. Natl. Herb. 12(10): 419. 1905. Pilocereus scoparius Diguet, Cactac. util. Mex. (Archiv. Hist. Nat. Acclim.
Poselg., Allg. Gartenzeitung (Otto & Dietrich) 21: 126. France) 4. 370. 1928. Cereus tetazo F.A.C. Weber ex J. M.
1853. Cereus scoparius A. Berger, Rep. (Annual) Missouri Coult., Contr. U.S. Natl. Herb. 3(7): 409. 1896. Carnegiea
Bot. Gard. 16: 63. 1905. Neobuxbaumia scoparia (Poselg.) tetazo (F.A.C. Weber ex J.M. Coult.) P.V.Heath, Calyx 2
Backeb., Beitr. Sukkulentenk. Sukkulentenpflege 3. 1941. (3): 108. 1992.
Carnegiea scoparia (Poselg.) P.V. Heath, Calyx 2(3): 109.
1992.—TYPE: MEXICO. Veracruz, “Nahe bei La Sol- Pilocereus tetetzo F.A.C. Weber ex K. Schum., Gesamtbeschr.
edad”, Poselger s.n. (not located). NEOTYPE (designated Kakt. 175. 1897. Cephalocereus tetetzo (F.A.C. Weber ex
here): MEXICO. Veracruz: cerca de Jalapa, 1957, H. Bravo K. Schum.) Vaupel, Monatsschr. Kakteenk. 19. 73. 1909.
Hollis s.n. (MEXU 70513!). Pachycereus tetetzo (F.A.C. Weber ex K. Schum.) Ochot.,
Cactac. de Mexico 17. 1922. Neobuxbaumia tetetzo (F.A.C.
Lemaireocereus setispinus E.Y. Dawson, Allan Hancock Found. Weber ex K. Schum.) Backeb., Blätt. Kakteenf. 6. 24.
Publ. Occas. Pap. 1: 19. 1948.—TYPE: MEXICO. Oaxaca: 1938.—TYPES: MEXICO. Puebla: Mit Pilocereus Hop-
at base of volcanic hill a few kilometres east of Juchitán, penstedtii Web. bei Zapotitlan in der Nähe von Tehuacan,
26 Jan 1947, E. Y. Dawson 3007 (syntypes: RSA!). 1866, 1869, F. A. C. Weber s.n. (syntypes: MO 2212310!,
13. CEPHALOCEREUS SENILIS (Haw.) Pfeiff. Allg. Gartenzeitung 6: 2212312!).
142. 1838. Cactus senilis Haw., Philos. Mag. J. 63: 41. 1824. 15. CEPHALOCEREUS TOTOLAPENSIS (Bravo & T. MacDoug.) Buxb.,
Cereus senilis (Haw.) DC., Prodr. 3: 464. 1828. Cepha- Kakteen and. Sukk. 16: 45. 1965. Neodawsonia totolapensis
lophorus senilis (Haw.) Lem., Cact. aliq. nov. XII. 1838. Bravo & T. MacDoug., Anales Inst. Biol. Univ. Nac.
Echinocactus senilis (Haw.) Beaton, Gard. Mag. & Reg. Mexico 29: 74. 1959.—TYPE: MEXICO. Oaxaca, Totola-
Rural Domest. Improv. 15: 550. 1839. Pilocereus senilis pan (not preserved). LECTOTYPE (designated here): [il-
(Haw.) Lem., Cact. Gen. Sp. Nov. 7. 1839. Cephalocereus lustration] “Neodawsonia totolapensis, creciendo cerca de
senilis (Haw.) K. Schum., Nat. Pflanzenfam. 3 (6a): 181.
Tequisistlán” in Bravo and MacDougall, Anales Inst. Biol.
1894.—TYPE: MEXICO (not located). NEOTYPE (desig-
Univ. Nac. Mexico 29: 75. Figure 1. 1959. EPITYPE
nated here): MEXICO. Hidalgo: Metztitlan, 17 Jun 2005,
(designed here): MEXICO. Oaxaca, Mpio. San Pedro
G. Arroyo-Cosultchi 23 (MEXU 1439221!).
Totolapan, 600 m sobre el camino de San José de Gracia a El
Cactus bradypus Lehm., Ind. Sem. Hort. Hamburg. 17. 1826 —TYPE: Romedo, 861 m, 16 Mar 2012, S. Arias 2182 (MEXU
not preserved. 1404304!).

Key to the Species of CEPHALOCEREUS

1. Sympodial growth with basitonic or mesotonic branching . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Plants with basitone pattern of branching, lateral cephalium present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13. C. senilis
2. Plants with mesotone pattern of branching, variable degree of ramification, differentiated or undifferentiated flowering region . . . . . . . . . . . . 3
3. Undifferentiated flowering region . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4. High degree of ramification, slender plagiotropic branches, subapical flowering, floral tube pinkish . . . . . . . . . . . . . . . . . 12. C. scoparius
4. Low to moderate degree of ramification, orthotropic branches variable in thickness, flowering variable in position, floral tube green or
often reddish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5. Low degree of ramification, often simple stems, subapical flowering, papyraceous scales on flowers and fruits . . . . . . . . 9. C. nudus
5. Moderate degree of ramification, apical to subapical flowering, fleshy scales on flowers and fruits when present . . . . . . . . . . . . . . 6
6. Thin branches (, 12 cm diam), 25 ribs or more, reddish to brownish spines concentrated towards the stem apices, floral tube dark
red colored . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11. C. sanchezmejoradae
6. Thick stem and branches (.20 cm diam), stems with less than 17 ribs, stem apices densely covered with trichomes or spines,
floral tube green or reddish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7. Reproductive stem apices covered with brownish trichomes, podaria evident near the apex, apical flowering . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14. C. tetetzo
7. Whitish to reddish spines concentrated towards the apex, podaria not evident, subapical flowering . . . . 5. C. macrocephalus
3. Flowering region differentiated in an apical pseudocephalium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8. Permanent yellowish apical pseudocephalium, thick stems (. 20 cm diam), red flowers densely covered with trichomes and bristles,
lime-green stems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. C. fulviceps
8. Intermittent apical pseudocephalium leaving annular remnants from flowering events, thin stems (, 15 cm diam), green bare flowers, green
to dark green stems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9. Tall plants, from 3 to 5 m in height, few branches (up to 5), often simple stems (up to 12 cm diam), 24 to 28 ribs . 15. C. totolapensis
9. Short plants, up to 2 m in height, with more than five branches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10. Few branches (5 to 10), 8 to 10 cm diam, 24 to 26 ribs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. C. apicicephalium
10. Several branches (up to 20), up to 8 cm diam, 21 to 23 ribs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8. C. nizandensis
1. Monopodial growth, simple stems, branches are produced only after an injury . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11. Differentiated flowering region in a lateral cephalium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. C. columna-trajani
11. Undifferentiated flowering region . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12. Flowering along the stem, less than 20 ribs, 5 to 7 radial spines in stem areoles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13. Thick stems up to 15 m height, 60 cm diam, up to 16 ribs, floral tube green . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6. C. mezcalaensis
13. Thin stems up to 7 m height, 12 cm diam, up to 20 ribs, floral tube reddish green . . . . . . . . . . . . . . . . . . . . . . . . . . 7. C. multiareolatus
12. Flowering concentrated below subapical region but can emerge scarcely along the stem, pinkish internal tepals, up to 20 ribs . . . . . . . 14
14. Numerous ribs (up to 40), thick stem up to 40 cm diam, flower up to 6 cm in length, spines yellow, 7 to 9 radial spines in stem
areoles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10. C. polylophus
14. Few (8 to 10) prominent ribs, thin stem up to 12 cm diam, flower up to 10 cm in length, spines white to grey, 3 radial spines in stem
areoles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. C. euphorbioides
2017] TAPIA ET AL.: PHYLOGENY OF CEPHALOCEREUS
Acknowledgments. The present study is a fragment of the dissertation
of HT and is presented as partial requirement to opt for a PhD degree in
Biological Sciences at Posgrado en Ciencias Biológicas (PCB), UNAM. The
first author thanks to CONACyT for the scholarship granted (332595).
We are grateful to David Gernandt for the revisions to preliminary
manuscript, Lidia Cabrera (Instituto de Biologı́a, UNAM), Montserrat
Vazquez-Sánchez (CIIDIR, IPN), Gabriel Arroyo-Cosultchi (UAM-Xochimilco),
Daniel Sánchez (Universidad de Guadalajara), Isaura Rosas, Daniel Franco,
David Aquino, Miguel Angel Cruz, Marisela Ya~ nez, and Yolanda Morales
(Instituto de Biologı́a, UNAM) for supporting the laboratory and field work,
Boris O. Schlumpberger (Herrenhausen Gardens, Hannover) for providing
Strophocactus specimen, and Fernando Chiang for its nomenclatural support.
We also thank the curators of the herbaria MEXU and XAL for loaning
specimens critical for this study; as well as James R. Smith, Susana Freire, and
two anonymous reviewers for constructive criticisms that improved this
manuscript. This research was supported by the program UNAM-DGAPA-
PAPIIT ,IN208315..
Literature Cited
Anderson, E. F. 2001. The cactus family. Portland, Oregon: Timber Press.
Arias, S. and T. Terrazas. 2006. Análisis cladı́stico del género Pachycereus
(Cactaceae) con caracteres morfológicos. Brittonia 58: 197–216.
Arias, S., T. Terrazas, and K. M. Cameron. 2003. Phylogenetic analysis of
Pachycereus (Cactaceae, Pachycereeae) based on chloroplast and nu-
clear DNA sequences. Systematic Botany 28: 547–557.
Arias, S., T. Terrazas, H. J. Arreola-Nava, M. Vázquez-Sánchez, and
K. M. Cameron. 2005. Phylogenetic relationships in Peniocereus
(Cactaceae) inferred from plastid DNA sequence data. Journal of Plant
Research 118: 317–328.
Arias, S., S. Gama-López, U. Guzmán-Cruz, and B. Vázquez-Benı́tez. 2012.
Cactaceae. Pp. 1–246 in Flora del Valle de Tehuacan-Cuicatlan fascı́culo
95, ed. R. Medina-Lemos. Mexico D. F.: Instituto de Biologı́a, Uni-
versidad Nacional Autónoma de Mexico.
Arroyo-Cosultchi, G., T. Terrazas, S. Arias, and L. López-Mata. 2010.
Delimitación de Neobuxbaumia mezcalaensis y N. multiareolata (Cacta-
ceae) con base en análisis multivariados. Boletı́n de la Sociedad Botánica
de México 86: 53–64.
Backeberg, C. 1938. Systematische Übersichtsarbeit. Blätter für Kakteen-
forschung 6 sine pag.
Backeberg, C. 1960. Die Cactaceae Band IV. Jena: Veb Gustav Fischer Verlag.
Bárcenas-Argüello, M. L. 2006. Filogenia del género Cephalocereus Pfeiff.
(Cactaceae) sensu Anderson con base en caracteres estructurales. M.S.
thesis. Montecillo, Estado de Mexico: Colegio de Postgraduados.
Bárcenas-Argüello, M. L., M. C. Gutiérrez-Castorena, T. Terrazas, and
L. López-Mata. 2010. Rock-soil preferences of three Cephalocereus
(Cactaceae) species of tropical dry forests. Soil Science Society of America
Journal 74: 1374–1382.
Bárcenas, R. T., C. Yesson, and J. A. Hawkins. 2011. Molecular systematics
of the Cactaceae. Cladistics 27: 470–489.
Barthlott, W. and D. R. Hunt. 1993. Cactaceae. Pp. 161–197 in The families
and genera of vascular plants vol. 2, ed. K. Kubitzki, G. Rohwer, and
V. Bittrich. Berlin: Springer-Verlag.
Borsch, T. and D. Quandt. 2009. Mutational dynamics and phylogenetic
utility of noncoding chloroplast DNA. Plant Systematics and Evolution
282: 169–199.
Bravo-Hollis, H. 1954. Un nuevo género de la familia de las Cactáceas:
Backebergia. Anales del Instituto de Biologı́a. Universidad Nacional
Autónoma de Mexico 24: 215–232.
Bravo-Hollis, H. 1974. Nuevas combinaciones II. Cactáceas y Suculentas
Mexicanas 19: 47.
Bravo-Hollis, H. 1978. Las Cactáceas de Mexico vol. 1. Ed. 2. Mexico D. F.:
Universidad Nacional Autónoma de Mexico.
Bravo-Hollis, H., L. Scheinvar, and H. Sánchez-Mejorada. 1970. Estudio
comparativo del género Neobuxbaumia Backeberg. Cactáceas y Sucu-
lentas Mexicanas 15: 75–82.
Bravo-Hollis, H., L. Scheinvar, and H. Sánchez-Mejorada. 1971a. Estudio
comparativo del género Neobuxbaumia Backeberg II. Cactáceas y
Suculentas Mexicanas 16: 3–14.
Bravo-Hollis, H., L. Scheinvar, and H. Sánchez-Mejorada. 1971b. Estudio
comparativo del género Neobuxbaumia Backeberg III. Cactáceas y
Suculentas Mexicanas 16: 75–83.
Bravo-Hollis, H., L. Scheinvar, and H. Sánchez-Mejorada. 1973. Estudio
comparativo del género Neobuxbaumia Backeberg IV. Cactáceas y
Suculentas Mexicanas 18: 59–67.
13
Britton, N. L. and J. N. Rose. 1920. The Cactaceae: Descriptions and illustrations
of plants of the cactus family vol. II. Washington D. C.: The Carnegie
Institution of Washington.
Butterwhorth, C. A., J. H. Cota-Sánchez, and R. S. Wallace. 2002. Molecular
systematics of tribe Cacteae (Cactaceae: Cactoideae): A phylogeny
based on rpl16 intron sequence variation. Systematic Botany 27:
257–270.
Buxbaum, F. 1958. The phylogenetic division of the subfamily Cereoideae,
Cactaceae. Madro~ no 14: 177–206.
Buxbaum, F. 1961. Die Entwicklungslinien der Tribus Pachycereeae F.
Buxb. (Cactaceae-Cereoideae). Botanical Studies (Taipei, Taiwan) 12:
1–107.
Buxbaum, F. 1964a. Was ist ein Cephalium?. Kakteen und Andere Sukku-
lenten 15: 28–31.
Buxbaum, F. 1964b. Was ist ein Cephalium?. Kakteen und Andere Sukku-
lenten 15: 43–48.
CBOL Plant Working Group. 2009. A DNA barcode for land plants.
Proceedings of the National Academy of Sciences USA 106: 12794–12797.
Cúenoud, P., V. Savolainen, L. W. Chatrou, M. Powell, R. J. Grayer, and
M. W. Chase. 2002. Molecular phylogenetics of Caryophyllales based
on nuclear 18s rDNA and plastid rbcL, atpB, and matK sequences.
American Journal of Botany 89: 132–144.
Darriba, D., G. L. Taboada, R. Doallo, and D. Posada. 2012. jModelTest 2:
More models, new heuristics and parallel computing. Nature Methods
9: 772.
Dawson, E. Y. 1948. New cacti of southern Mexico. Allan Hancock Foun-
dation Publications: Occasional Papers 1: 1–69.
Dawson, E. Y. 1952a. Field observations on some cacti of Oaxaca and
Puebla, Mexico. Desert Plant Life 24: 53–58.
Dawson, E. Y. 1952b. Notes on Neobuxbaumia. Cactus and Succulent Journal
24: 167–173. (Los Ángeles).
Flores-Ortiz, C. M. 2004. Quimiotaxonomı́a del género Neobuxbaumia. Ph.D.
thesis. Mexico, D. F.: Universidad Nacional Autónoma de Mexico.
Franck, A. R., B. J. Cochrane, and J. R. Garey. 2012. Low-copy nuclear
primers and ycf1 primers in Cactaceae. American Journal of Botany 99:
e405–e407.
Gibson, A. C. and K. E. Horak. 1978. Systematic anatomy and phylogeny of
Mexican columnar cacti. Annals of the Missouri Botanical Garden 65:
999–1057.
Gorelick, R. 2002. DNA sequences and cactus classification — a short
review. Bradleya 20: 1–4.
Guzmán-Cruz, U., S. Arias, and P. Dávila. 2003. Catálogo de Cactáceas
Mexicanas. Mexico D. F.: Universidad Nacional Autónoma de Mexico /
CONABIO.
Hall, T. A. 1999. BioEdit: A user-friendly biological sequence alignment
editor and analysis program for Windows 95/98/NT. Nucleic Acids
Symposium Series 41: 95–98.
Hausdorf, B. 2011. Progress toward a general species concept. Evolution 65:
923–931.
Heath, P. V. 1992. The restoration of Rathbunia Britton & Rose. Calyx
(Toronto, Ont.) 2: 102–115.
Hernández-Hernández, T., H. M. Hernández, J. A. De-Nova, R. Puente,
L. E. Eguiarte, and S. Magallón. 2011. Phylogenetic relationships and
evolution of growth form in Cactaceae (Caryophyllales, Eudicotyle-
doneae). American Journal of Botany 98: 44–61.
Hernández-Ledesma, P., W. G. Berendsohn, T. Borsch, S. Von Mering,
H. Akhani, S. Arias, I. Casta~neda-Noa, U. Eggli, R. Eriksson, H. Flores-
Olvera, S. Fuentes-Bazán, G. Kadereit, C. Klak, N. Korotkova,
R. Nyffeler, G. Ocampo, H. Ochoterena, B. Oxelman, R. K. Rabeler,
A. Sanchez, B. O. Schlumpberger, and P. Uotila. 2015. A taxonomic
backbone for the global synthesis of species diversity in the angio-
sperm order Caryophyllales. Willdenowia 45: 281–383.
Hershkovitz, M. A. and E. A. Zimmer. 1997. On the evolutionary origins of
the cacti. Taxon 46: 217–232.
Huelsenbeck, J. P. and F. Ronquist. 2001. MrBayes: Bayesian inference of
phylogenetic trees. Bioinformatics 17: 754–755. (Oxford).
Hunt, D. R. 1999. CITES Cactaceae Checklist. Milborne Port: Royal Botanic
Gardens Kew / International Organization for Succulent Plant Study.
Hunt, D. R. and N. P. Taylor. 1986. The genera of Cactaceae: Towards a new
consensus. Bradleya 4: 65–78.
Hunt, D. R. and N. P. Taylor. 1990. The genera of Cactaceae: Progress towards
consensus report of the IOS Working Party. Bradleya 8: 85–107.
Hunt, D. R. and N. P. Taylor. 1991. Notes on miscellaneous genera of
Cactaceae. Bradleya 9: 81–93.
Hunt, D. R., N. P. Taylor, and G. Charles. 2006. The new cactus lexicon.
Milborne Port: DH Books.
14 SYSTEMATIC BOTANY
INEGI. 2014. Mapa digital de Mexico. Available at: http://gaia.inegi.org.
mx/mdm6/.
Katoh, K. and D. M. Standley. 2013. MAFFT multiple sequence alignment
software version 7: Improvements in performance and usability.
Molecular Biology and Evolution 30: 772–780.
Korotkova, N., T. Borsch, D. Quandt, N. P. Taylor, K. F. Müller, and
W. Barthlott. 2011. What does it take to resolve relationships and to
identify species with molecular markers? An example from the epiphytic
Rhipsalideae (Cactaceae). American Journal of Botany 98: 1549–1572.
Korotkova, N., L. Zabel, D. Quandt, and W. Barthlott. 2010. A phylogenetic
analysis of Pfeiffera and the reinstatement of Lymanbensonia as an
independently evolved lineage of epiphytic Cactaceae within a new
tribe Lymanbensonieae. Willdenowia 40: 151–172.
Lahaye, R.M. van der Bank, D. Bogarin, J. Warner, F. Pupulin, G. Gigot, O.
Maurin, S. Duthoit, T. G. Barraclough, and V. Savolainen. 2008. DNA
barcoding the floras of biodiversity hotspots. Proceedings of the National
Academy of Sciences USA 105: 2923–2928.
Maddison, W. P. and D. R. Maddison. 2015. Mesquite: A modular system
for evolutionary analysis, v. 3.04. http://mesquiteproject.org.
Moore, H. E. 1975. Nomenclatural notes for hortus third. Cactaceae. Baileya
19: 165–166.
Morrone, J. J. 2005. Hacia una sı́ntesis biogeográfica de Mexico. Revista
Mexicana de Biodiversidad 76: 207–252.
Müller, K. 2004. PRAP-computation of Bremer support for large data sets.
Molecular Phylogenetics and Evolution 31: 780–782.
Nixon, K. C. 2002. WinClada v. 1.00.08. Ithaca, New York: published by the
author.
Nyffeler, R. 2002. Phylogenetic relationships in the cactus family (Cacta-
ceae) based on evidence from trnK/matK and trnL-trnF sequences.
American Journal of Botany 89: 312–326.
Nyffeler, R. and U. Eggli. 2010. Disintegrating Portulacaceae: A new fa-
milial classification of the suborder Portulacineae (Caryophyllales)
based on molecular and morphological data. Taxon 59: 227–240.
Ochoterena, H. 2009. Homology in coding and non-coding DNA se-
quences: A parsimony perspective. Plant Systematics and Evolution 282:
151–168.
Pfeiffer, L. 1838. Ueber Lemaire’s Beschreibung einiger neuen Cacteen.
Allgemeine Gartenzeitung 6: 142.
Plume, O., S. C. K. Straub, N. Tel-Zur, A. Cisneros, B. Schneider, and
J. J. Doyle. 2013. Testing a hypothesis of intergeneric allopolyploidy in
vine cacti (Cactaceae: Hylocereeae). Systematic Botany 38: 737–751.
Ritz, C. M., L. Martins, R. Mecklenburg, V. Goremykin, and F. H. Hellwig.
2007. The molecular phylogeny of Rebutia (Cactaceae) and its allies
demonstrates the influence of paleogeography on the evolution of
South American mountain cacti. American Journal of Botany 94:
1321–1332.
Sánchez, D., S. Arias, and T. Terrazas. 2014. Phylogenetic relationships in
Echinocereus (Cactaceae, Cactoideae). Systematic Botany 39: 1183–1196.
Sang, T., D. J. Crawford, and T. F. Stuessy. 1997. Chloroplast DNA phy-
logeny, reticulate evolution, and biogeography of Paeonia (Paeonia-
ceae). American Journal of Botany 84: 1120–1136.
Scheinvar, L. and H. Sánchez-Mejorada. 1990. Neobuxbaumia squamulosa
Scheinvar et Sánchez Mejorada sp. nov. Cactáceas y Suculentas Mexi-
canas 35: 13–18.
Seehausen, O. 2004. Hybridization and adaptive radiation. Trends in
Ecology & Evolution 19: 198–207.
Shaw, J., E. B. Lickey, J. T. Beck, S. B. Farmer, W. Liu, J. Miller, K. C. Siripun,
C. T. Winder, E. E. Schilling, and R. L. Small. 2005. The tortoise and the
hare II: Relative utility of 21 noncoding chloroplast DNA sequences
for phylogenetic analysis. American Journal of Botany 92: 142–166.
Shaw, J., E. B. Lickey, E. E. Schilling, and R. L. Small. 2007. Comparison of
whole chloroplast genome sequences to choose noncoding regions for
phylogenetic studies in angiosperms: The tortoise and the hare III.
American Journal of Botany 94: 275–288.
Simmons, M. P. and H. Ochoterena. 2000. Gaps as characters in sequence-
based phylogenetic analyses. Systematic Biology 49: 369–381.
Stöver, B. C. and K. F. Müller. 2010. TreeGraph 2: Combining and visu-
alizing evidence from different phylogenetic analyses. BCM Bio-
informatics 9: 1–9.
Swofford, D. L. 2002. PAUP* Phylogenetic analysis using parsimony (*and
other methods), v. 4. Sunderland: Sinauer Associates.
Taberlet, P., L. Gielly, G. Pautou, and J. Bouvet. 1991. Universal primers for
amplification of three non-coding regions of chloroplast DNA. Plant
Molecular Biology 17: 1105–1109.
[Volume 42
Tapia, H. J., S. Arias, L. Yá~
nez-Espinosa, and T. Terrazas. 2016. El uso de
espinas del tallo en la identificación de las especies de Neobuxbaumia
(Cactaceae). Revista Mexicana de Biodiversidad 87: 288–300.
Tapia, H. J., M. L. Bárcenas-Argüello, T. Terrazas, and S. Arias. 2017. Data
from: Phylogeny and circumscription of Cephalocereus (Cactaceae)
based on molecular and morphological evidence. Dryad Digital Re-
pository. http://dx.doi.org/10.5061/dryad.vg1g6.
Tate, J. A. and B. B. Simpson. 2003. Paraphyly of Tarasa (Malvaceae) and
diverse origins of the polyploid species. Systematic Botany 28: 723–737.
Terrazas, T. and S. Loza-Cornejo. 2002. Phylogenetic relationship of
Pachycereeae: A cladistic analysis based on anatomical-morphological
data. Pp. 66–86 in Columnar cacti and their mutualists: Evolution, ecology,
and conservation, eds. T. H. Fleming and A. Valiente-Banuet. Tucson:
The University of Arizona Press.
Thompson, J. D., D. G. Higgins, and T. J. Gibson. 1994. CLUSTAL W:
Improving the sensitivity of progressive multiple sequence alignment
through sequence weighting, position-specific gap penalties and
weight matrix choice. Nucleic Acids Research 22: 4673–4680.
Vázquez-Sánchez, M., T. Terrazas, and S. Arias. 2005. Morfologı́a y
anatomı́a del cefalio de Cephalocereus senilis (Cactaceae). Anales del
Jardin Botanico de Madrid 62: 153–161.
Vázquez-Sánchez, M., T. Terrazas, and S. Arias. 2007. Morphology and
anatomy of the Cephalocereus columna-trajani cephalium: Why tilting?
Plant Systematics and Evolution 265: 87–99.
Vázquez-Sánchez, M., T. Terrazas, and S. Arias. 2016. Comparative
morphology and anatomy of Backebergia militaris (Echinocereeae–
Cactaceae) cephalium. Plant Systematics and Evolution 302: 245–256.
Vázquez-Sánchez, M., T. Terrazas, S. Arias, and H. Ochoterena. 2013.
Molecular phylogeny, origin and taxonomic implications of the tribe
Cacteae (Cactaceae). Systematics and Biodiversity 11: 103–116.
Vite, F., E. Portilla, A. Zavala-Hurtado, P. L. Valverde, and A. Dı́az-Solı́s.
1996. A natural hybrid population between Neobuxbaumia tetetzo and
Cephalocereus columna-trajani (Cactaceae). Journal of Arid Environments
32: 395–405.
Wiens, J. J. 1998. Combining data sets with different phylogenetic histories.
Systematic Biology 47: 568–581.
APPENDIX 1. Taxon, source, collector, collection number, (Herbarium ac-
ronym), GenBank accession numbers (rpl16, trnL-F, psbA-trnH, rpl32-trnL,
trnQ-rps16, petL-psbE, and ycf1). The sequences lacking for a locus/
specimen was marked with dash (—), N.A.: no data.
Acanthocereus tetragonus (L.) Hummelinck, MEXICO: Nuevo León, Arias
1450 (MEXU), KU598021, KU598074, KU597969, KY624736, KU598127,
KY624806, KU598178; Bergerocactus emoryi Britton & Rose, MEXICO: Baja
Cal., Arias 1307 (CHAPA), DQ099994, DQ099925, KF783478, KY624730,
KF783697, KY624800, KY624834; Carnegiea gigantea Britton & Rose, U. S. A.:
Arizona, NYGB N.A. (cult.), AY181591, AY181619, KY624692, KY624725,
KY624762, KY624795, KY624830; Cephalocereus apicicephalium E.Y.Dawson,
MEXICO: Oaxaca, Arias 1257 (MEXU), DQ099996, DQ099927, KY624684,
KY624717, KY624756, KY624787, KY624825; C. columna-trajani (Karw.)
K.Schum., MEXICO: Puebla, Arias 1377 (CHAPA), AY181599, AY181648,
KY624685, KY624718, KY624757, KY624788, —; C. nizandensis (Bravo &
T.MacDoug.) Buxb., MEXICO: Oaxaca, Terrazas 633 (CHAPA), DQ099997,
DQ099928, KY624686, KY624719, —, KY624789, —; C. senilis Pfeiff., MEXICO:
Hidalgo, Terrazas 529 (CHAPA), AY181616, AY181638, KY624687, KY624720,
KY624758, KY624790, KY624826; C. totolapensis (Bravo & T.MacDoug.)
Buxb., MEXICO: Oaxaca, Terrazas 628 (CHAPA), KY624647, KY624661,
KY624688, KY624721, KY624759, KY624791, KY624827; Deamia testudo
(Karw. ex Zucc.) Britton & Rose, MEXICO: Oaxaca, Ya~ nez 01 (MEXU),
KY624648, KY624662, KY624695, KY624731, KY624765, KY624801,
KY624835; D. testudo (Karw. ex Zucc.) Britton & Rose, GUATEMALA:
Izabal, Veliz 19628 (BIGU), KY624649, KY624663, KY624696, KY624732,
KY624766, KY624802, KY624836; Escontria chiotilla Rose, MEXICO: Puebla,
Arias 891 (MEXU), AY181608, AY181622, KY624693, KY624726, KY624763,
KY624796, KY624831; Harrisia earlei Britton & Rose, U. S. A.: Saint Croix
Botanical Garden, Fleming 118 (cult.), DQ100008, DQ099939, KY624665,
KY624698, —, KY624768, —; Lemaireocereus hollianus Britton & Rose,
MEXICO: Puebla, Arias 1373 (CHAPA), KU598012, KU598064, KU597959,
KY624735, KU598117, KY624805, KU598169; Leptocereus quadricostatus
Britton & Rose, PUERTO RICO: Cabo Rojo, Arias 1464 (MEXU), KF783620,
KF783690, AY851582, KY624737, KF783768, KY624807, KY624837; Mar-
shallocereus aragonii (F.A.C.Weber) Backeb., NICARAGUA: The Huntington
Botanical Gardens 66864 (cult.), AY181611, AY181630, KY624689, KY624722, —,
KY624792, —; M. eichlamii (Britton and Rose) Buxb., MEXICO: Yucatán, Arias
1363 (MEXU), AY181610, AY181629, KY624690, KY624723, KY624760,
2017] TAPIA ET AL.: PHYLOGENY OF CEPHALOCEREUS
KY624793, KY624828; Myrtillocactus geometrizans Console, MEXICO: Quere-
taro, Terrazas 557 (CHAPA), DQ100012, DQ099943, KY624694, KY624728,
KY624764, KY624798, KY624833; Neobuxbaumia euphorbioides (Haw.) Buxb. Ex
Bravo, MEXICO: Veracruz, Tapia 01 (MEXU), KY624637, KY624651,
KY624666, KY624699, KY624738, KY624769, KY624808; N. euphorbioides
(Haw.) Buxb. Ex Bravo, MEXICO: Tamaulipas, Hamman N.A., (cult.),
AY181595, AY181635, KY624667, KY624700, KY624739, KY624770, KY624809;
N. macrocephala (F.A.C.Weber ex K.Schum.) E.Y. Dawson, MEXICO: Puebla,
Arias 1048 (MEXU), DQ100013, DQ099944, KY624668, KY624701, KY624740,
KY624771, KY624810; N. mezcalaensis (Bravo) Backeb., MEXICO: Puebla, Tapia
44 (MEXU), KY624638, KY624652, KY624669, KY624702, KY624741,
KY624772, KY624811; N. mezcalaensis (Bravo) Backeb., MEXICO: Morelos,
Tapia 42 (MEXU), KY624639, KY624653, KY624670, KY624703, KY624742,
KY624773, KY624812; N. mezcalaensis (Bravo) Backeb., MEXICO: Guerrero,
Terrazas 533 (CHAPA), AY181600, AY181645, KY624671, KY624704,
KY624743, KY624774, KY624813; N. multiareolata (Daws.) Bravo Scheinvar &
Sánchez-Mej., MEXICO: Guerrero, Terrazas 531 (CHAPA), AY181594,
AY181636, KY624672, KY624705, KY624744, KY624775, KY624814; N. poly-
lopha (DC) Backeb., MEXICO: Hidalgo, Terrazas 530 (CHAPA), AY181597,
AY181644, KY624673, KY624706, KY624745, KY624776, KY624815; N. san-
chezmejoradae Lau, MEXICO: Oaxaca, Tapia 39 (MEXU), KY624640,
KY624654, KY624674, KY624707, KY624746, KY624777, KY624816;
N. scoparia (Poselg.) Backeb., MEXICO: Oaxaca, Hamman N.A. (cult.),
AY181596, AY181625, KY624675, KY624708, KY624747, KY624778,
KY624817; N. scoparia (Poselg.) Backeb., MEXICO: Veracruz, Tapia 02
(MEXU), KY624641, KY624655, KY624676, KY624709, KY624748,
KY624779, KY624818; N. squamulosa Scheinvar & Sánchez-Mej., MEX-
ICO: Oaxaca, Tapia 24 (MEXU), KY624642, KY624656, KY624677,
KY624710, KY624749, KY624780, KY624819; N. squamulosa Scheinvar &
Sánchez-Mej., MEXICO: Oaxaca, Tapia 11 (MEXU), KY624643, KY624657,
KY624678, KY624711, KY624750, KY624781, KY624820; N. squamulosa
Scheinvar & Sánchez-Mej., MEXICO: Jalisco, Tapia 30 (MEXU), KY624644,
KY624658, KY624679, KY624712, KY624751, KY624782, KY624821; N.
tetetzo (F.A.C.Weber ex K.Schum.) Backeb., MEXICO: Puebla, Arias 1376
(CHAPA), AY181592, AY181632, KY624680, KY624713, KY624752,
KY624783, —; N. tetetzo (F.A.C.Weber ex K.Schum.) Backeb., MEXICO:
Oaxaca, Tapia 20 (MEXU), KY624645, KY624659, KY624681, KY624714,
KY624753, KY624784, KY624822; N. tetetzo (F.A.C.Weber ex K.Schum.)
Backeb., MEXICO: Oaxaca, Tapia 41 (MEXU), KY624646, KY624660,
KY624682, KY624715, KY624754, KY624785, KY624823; Pachycereus
pringlei Britton & Rose, MEXICO: Baja Cal. Sur, Arias 1348 (CHAPA),
AY181589, AY181642, KF783549, KY624729, JX111635, KY624799, —;
Pseudomitrocereus fulviceps (F.A.C.Weber) Bravo & Buxb., MEXICO:
15
Puebla, Tapia 29 (MEXU), AY181602, AY181621, KY624683, KY624716,
KY624755, KY624786, KY624824; Pterocereus gaumeri (Britton & Rose) Th.
MacDougall & Miranda, MEXICO: Yucatán, Arias 1360 (MEXU),
AY181606, AY181626, KY624691, KY624724, KY624761, KY624794,
KY624829; Selenicereus grandiflorus Britton & Rose, MEXICO: Veracruz,
Guzmán 1365 (MEXU), DQ100039, DQ099970, KU597971, KY624734,
KU598129, KY624804, KU598180; Stenocereus stellatus Riccob., MEXICO:
Puebla, Arias 1375 (MEXU), AY181590, AY181640, KF783555, KY624727,
KF783766, KY624797, KY624832; Strophocactus wittii (K.Schum.) G.D.
Rowley, Hannover, Herrenhäuser Gärten B05X138, (cult.), KY624650,
KY624664, KY624697, KY624733, KY624767, KY624803, —.
Appendix 2. Structural character matrix according to Table 4.
Characters
Species 000000000111111111122222222223333333333444444444
123456789012345678901234567890123456789012345678
N. euphorbioides 000111002021210000010000100010010110100011001?01
N. macrocephala 122110102111211000021100021101001100100011011101
N. mezcalaensis 001111002121100000001100121020011101101001001?01
N. multiareolata 000111011120110000021100021000001101101??1??????
N. polylopha 000011112220110000011110011010001110100??1??????
N. sanchezmejoradae 120011010110110000001100022020001102100??1??????
N. scoparia 120211002110210000011110000000001100101011001101
N. squamulosa 122111002111111000011100021020011111100??1??????
N. tetetzo 123111112101011000011100021010001100101011001?01
P. fulviceps 122200112201011010121111122001011110100021021?01
C. apicicephalium 111010101101101010100110022000000101000101000000
C. columna-trajani 001010202211111101020110000000000100101011011201
C. nizandensis 11111020120110101010011002200000010?000??1??????
C. senilis 112010302111111101020110000000000110100??1??????
C. totolapensis 00011010110110101010011002200000010?000011020100
P. pringlei 1221001011210010000300010220211110000000101?1?01
P. gaumeri 11020010102010000000010002202010101?1010100?0?01
S. stellatus 123210101121100000001011122000111012010000011?01
E. chiotilla 122000101101100000000001011020101022010010001010
M. eichlamii 1211100011111000000110110221000110021010100?1?10
C. gigantea 12230020220100100000000102202010101000111011110?
B. emoryi 112000301220100000000001122000111010000010001201
L. hollianus 112310102200100000000002122011111000101010000?11
M. geometrizans 122300001020100000000001000000100002010120021000
A. tetragonus 122310000120100000030003111010110000000??0??????
L. quadricostatus 122310100200100000000000111000110010000??0??????