Infrageneric classification of Agave L. (Asparagaceae: Agavoideae / Agavaceae): a nomenclatural assessment and updated classification at the rank of section, with new combinations

Abstract

An updated infrageneric classification is provided for the genus Agave L. s.l., therefore including A. subg. Manfreda, i.e., the genera Manfreda Salisb., Polianthes L., and Prochnyanthes S.Watson (Asparagaceae: Agavoideae / Agavaceae), with emphasis on the rank of section. Nomenclatural assessments are provided for (1) all unranked infrageneric ‘group’ names introduced by Trelease (1913) for the Caribbean region, by (2) Gentry (1982) for continental North America, and (3) for all infrageneric taxa at sectional rank hitherto published in Agave by Salm-Dyck, Engelmann, Berger, Ullrich, Webb & Starr, Starr & Webb, and Hochstätter. We also show that Jacobi did not publish names at the rank of section. In addition, a revised infrageneric classification of Agave s.l. is provided, including new combinations and new names mainly at the rank of section, especially for those unranked group names used by Trelease (1913) and Gentry (1982) for which no corresponding formal name at sectional rank is available. Three new combinations are made in Agave for species recently described in Manfreda and Polianthes.

Full text

Summary: An updated infrageneric classification
is provided for the genus Agave L. s.l., therefore
including A. subg. Manfreda, i.e., the genera Man-
freda Salisb., Polianthes L., and Prochnyanthes
S.Watson (Asparagaceae: Agavoideae /
Agavaceae), with emphasis on the rank of section.
Nomenclatural assessments are provided for (1)
all unranked infrageneric ‘group’ names intro-
duced by Trelease (1913) for the Caribbean region,
by (2) Gentry (1982) for continental North Amer-
ica, and (3) for all infrageneric taxa at sectional
rank hitherto published in Agave by Salm-Dyck,
Engelmann, Berger, Ullrich, Webb & Starr, Starr
& Webb, and Hochstätter. We also show that Ja-
cobi did not publish names at the rank of section.
In addition, a revised infrageneric classification of
Agave s.l. is provided, including new combinations
and new names mainly at the rank of section, es-
pecially for those unranked group names used by
Trelease (1913) and Gentry (1982) for which no
corresponding formal name at sectional rank is
available. Three new combinations are made in
Agave for species recently described in Manfreda
and Polianthes.
Zusammenfassung: Für die Gattung Agave L. s.l.,
also einschließlich A. subg. Manfreda mit den
Gattungen Manfreda Salisb., Polianthes L., and
Prochnyanthes S.Watson (Asparagaceae:
Agavoideae / Agavaceae), wird eine aktualisierte
infragenerische Klassifikation mit Schwerpunkt
auf der Rangstufe der Sektion vorgeschlagen.
Nomenklatorische Überprüfungen werden
vorgenommen für (1) alle als ‘group’ (Gruppe)
veröffentlichten ranglosen infragenerischen
Namen, die von Trelease (1913) für die Karibis-
che Region, und von (2) Gentry (1982) für das kon-
tinentale Nordamerika eingeführt wurden, und
(3) für alle infragenerischen Taxa auf der
Rangstufe der Sektion, die bisher in der Gattung
Agave von Salm-Dyck, Engelmann, Berger, Ull-
rich, Webb & Starr, Starr & Webb und Hochstät-
ter veröffentlicht worden sind. Wir zeigen auch,
dass Jacobi keine Namen auf der Rangstufe der
Sektion veröffentlicht hat. Ergänzend wird eine
überarbeitete infragenerische Klassifikation von
Agave s.l. vorgestellt, einschließlich von Neukom-
binationen und neuen Namen vorwiegend auf der
Rangstufe der Sektion, insbesondere für die von
Trelease (1913) und Gentry (1982) verwendeten
ranglosen, als ‘group’ (Gruppe) veröffentlichten
Namen, für die kein entsprechender Name auf
Sektionsrang verfügbar ist. Drei neue Kombina-
tionen in der Gattung Agave werden für kürzlich
in den Gattungen Manfreda und Polianthes neu
beschriebene Arten veröffentlicht.
Introduction
For millennia, representatives of the genus
Agave L. (Asparagaceae: Agavoideae / Agavaceae)
have played a significant role in the indigenous
civilizations of North America, with several
species being of considerable economic importance
in ethnobotany, ecology, and horticulture (Fig-
ures1 &2). Plants of numerous species are wild-
harvested or cultivated as agricultural crops,
especially for the extraction of fibres from their
leaves (Figure 3), the production of fermented
(pulque) or distilled (mezcal, including tequila) al-
coholic beverages (Figures4 & 5), and as a source
of food (Gentry, 1982). Geographically, Agave s.l.
(incl. A.subg. Manfreda (Salisb.) Baker) ranges
from the east-central (A.virginica L.) and south-
ern United States of America throughout Mexico,
Mesoamerica, and the Caribbean to northern
South America (Colombia and Venezuela); the
240 Bradleya 37/2019
Bradleya 37/2019
pages 240–264
Infrageneric classification of Agave L. (Asparagaceae:
Agavoideae / Agavaceae): a nomenclatural assessment and
updated classification at the rank of section, with
new combinations
Joachim Thiede1, Gideon F. Smith2& Urs Eggli3
1. Schenefelder Holt 3, 22589 Hamburg, Germany. (email: joachim_thiede@gmx.de)
2. Department of Botany, P.O. Box 77000, Nelson Mandela University, Port Elizabeth, 6031
South Africa. (email: smithgideon1@gmail.com)
3. Sukkulenten-Sammlung Zürich / Grün Stadt Zürich, Mythenquai 88, CH-8002 Zürich,
Switzerland. (email: Urs.Eggli@zuerich.ch)

genus is most diverse in central and southern
Mexico (García-Mendoza & Galván, 1995). The
number of species included in Agave varies de-
pending on how the genus is circumscribed and
how species are delimited, with estimates gener-
ally ranging from about 200 (García-Mendoza,
2002; excluding A.subg. Manfreda, i.e., = Agave
s.s.) to some 220 species (Thiede, 2001; Agave s.l.)
or about 265 species (Thiede, 2019; Agave s.l.).
More than 40 new species have been described
since Thiede (2001), arguably the most widely
cited recent treatment of Agave s.l., appeared in
print.
For Agave s.s., two subgenera are recognised:
A.subg. Agave with broadly paniculate inflores-
cences with much-branched part-inflorescences
(often wrongly termed ‘umbels’ or ‘umbellate’; Fig-
ures13–15, 17 & 21), and A.subg. Littaea (Tagl.)
Baker with narrowly elongated panicles with
part-inflorescences mostly carrying geminate
flowers (often wrongly termed ‘spicate’ or ‘spike-
241
Bradleya 37/2019
Figure 1. The comparative hardiness, ease of cultivation, and strong, visually pleasing symmetry of the
rosettes and leaves of species such as Agave utahensis (here: var. nevadensis), here photographed in an
open bed in the Denver Botanical Garden, Denver, Colorado, USA, 10 August 2017, contribute to the in-
creasing popularity of agaves in amenity and public gardening. Photograph: Gideon F. Smith.
Figure 2. Small-growing, variegated-leaved
plants, such as this specimen of Agave victoria-
reginae that sport longitudinal white sections
along the leaf margins, respond well to cultivation
in places where space is at a premium.
Photograph: Gideon F. Smith.

like’ inflorescences; Figures6 & 7). When Agave
is treated in a broad sense, the third subgenus,
A.subg. Manfreda, accounts for the three closely
related genera that were formerly upheld: Man-
freda Salisb., Polianthes L., and Prochnyanthes
S.Watson. Species included in A.subg. Manfreda
usually have true bulbs, mostly short-lived leaves
and elongated spikes or racemes with single or ge-
minate, often ± curved flowers (Figures18–20).
Authors disagree whether these three genera
(Manfreda, Polianthes, and Prochnyanthes)
should be upheld (see e.g., McVaugh, 1989: 247–
260 (Manfreda), Verhoek-Williams, 1975 (Po-
lianthes), Castro-Castro et al., 2010
(Prochnyanthes)) or included in Agave s.l., as
A.subg. Manfreda (Thiede, 2001; Govaerts et al.,
2018; JSTOR Global Plants, 2018).
Most phylogenetic studies based on molecular
(e.g., Bogler & Simpson, 1996; Bogler et al., 2006;
Flores-Abreu et al., 2019) or morphological data
(e.g., Hernández-Sandoval, 1995) showed that
Manfreda, Polianthes, and Prochnyanthes are
nested within Agave, thus rendering Agave s.s.
paraphyletic. Thiede (2001), for a taxonomic syn-
opsis, converted these phylogenetic results into a
classification that yielded a monophyletic re-cir-
cumscription of Agave: Manfreda, Polianthes, and
Prochnyanthes were included in Agave s.l. and to-
gether classified as A.subg. Manfreda (Salisb.)
Baker.
Given the comparatively large number of spe-
cies (c. 265; see above) included in the genus
Agave (s.l.), many infrageneric taxa below the
rank of subgenus (sections, subsections, series,
and subseries) as well as unranked, informal taxa
(Groups, Reihen, Unterreihen, etc., or even unde-
signated ones) have been published over the past
c. 200 years as part of efforts to facilitate species
identification. These were compiled into an anno-
tated checklist by Mottram (2015). Most widely
used are the rankless, informal ‘Groups’ introdu-
ced in the monographic treatments of: 1) Trelease
(1913) for the Caribbean species, and 2) Gentry
(1982) for the species of continental North Ame-
rica and slightly beyond. More recently, Ullrich
(1991b; 1992; 1993; 1995), Webb & Starr (2015),
Starr & Webb (2015), and Hochstätter (2015) com-
bined and gave formal recognition at the rank of
section to several unranked infrageneric group
names proposed by Trelease and Gentry, or pu-
blished new names at sectional rank. However,
names published earlier by Salm-Dyck (1859,
1861), Jacobi (1864, 1868), Engelmann (1875),
and Berger (1915) at sectional and other ranks,
including ‘unranked’, have not so far been for-
mally assessed.
Complementary to an updated synopsis of
Agave s.l. (Thiede, 2019), a revised infrageneric
classification is proposed here. This classification
is premised on the following:
- For infrageneric taxa below the rank of subge-
nus, the rank of section is consistently applied
throughout. Infrageneric names at series and
subseries ranks remain unconsidered unless
they serve as basionyms of names at sectional
rank (an exception is made for series publis-
hed by Hochstätter (2016) in Manfreda).
- The classification has its roots in the unranked
informal ‘groups’ of Trelease (1913) and Gen-
try (1982), and all names published at sectio-
nal rank by Salm-Dyck, Jacobi, Engelmann,
Berger, Ullrich, Webb & Starr, Starr & Webb,
and Hochstätter.
- For the unranked infrageneric group names in-
troduced by Trelease (1913) and Gentry
(1982), for which corresponding formal names
at sectional rank have not yet been published,
new combinations are published here. As far
as possible all typifications and new combina-
tions aim to retain the widely used unranked
group names of Trelease (1913) and Gentry
(1982) at formal sectional rank.
- Infrageneric unranked names as listed by Mot-
tram (2015) are not exhaustively assessed.
Such an assessment is not necessary as names
without a clear indication of its rank are “in-
operative in questions of priority except for ho-
monymy” (International Code of Nomenclature
for algae, fungi, and plants (ICN) Art. 37.3.;
Turland et al., 2018). Therefore, rankless
names not considered cannot affect the classi-
fication suggested here.
- This paper focuses on nomenclature, and cir-
cumscriptions and descriptive, especially mor-
phological, information are not provided for
242 Bradleya 37/2019
Figure 3. Twine produced from fibres extracted
from the leaves of Agave sisalana. The cut end of
a leaf of A. sisalana is visible in the background.
Photograph: Gideon F. Smith.

the infrageneric taxa. Such information can be
found in Trelease (1913), Gentry (1982),
Thiede (2001), Hochstätter (2015, 2016), and
Thiede (2019).
This paper sequentially provides:
- 1) A nomenclatural assessment, including typi-
fications, of the unranked infrageneric group
names used by Trelease (1913) and Gentry
(1982);
- 2) A nomenclatural assessment, including typi-
fications, of some rankless names as well as all
formal infrageneric taxa at sectional rank pu-
blished in Agave by Salm-Dyck (1859, 1861),
Jacobi (1864, 1868), Engelmann (1875), Ber-
ger (1915), Ullrich (1991b, 1992, 1993, 1995),
Webb & Starr (2015), Starr & Webb (2015),
and Hochstätter (2015); and
- 3) A formal, revised infrageneric classification of
Agave s.l. with an emphasis at the rank of
section, including the publication of new com-
binations at the rank of section for those in-
formal groups used by Trelease (1913) and
243
Bradleya 37/2019
Figure 4. The landscape around the town of Tequila (Jalisco, Mexico) where Agave tequilana 'Azul' is cul-
tivated in large plantations was declared as UNESCO World Heritage "Agave Landscape and Ancient In-
dustrial Facilities of Tequila" in 2006. Photograph: Joachim Thiede.
Figure 5. Tequila, a type of mescal, is consumed
in vast quantities across the world, very often as
the major component in cocktails. Tanks with dis-
tilled tequila at Tequila Orendain, Tequila,
Jalisco, Mexico, 5 November 2015.
Photograph: Joachim Thiede.

Gentry (1982) for which no corresponding for-
mal name at sectional rank is available.
The infrageneric classification published here
provides a nomenclatural framework of formal,
ranked names for all infrageneric groupings in
Agave. We are aware that evidence for the mono-
phyly of most sections remains wanting; however,
the system we propose is sufficiently robust to en-
able and facilitate communication about the sug-
gested clustering of species.
1) Nomenclatural assessment of the unran-
ked informal infrageneric group names in-
troduced by Trelease (1913) and Gentry
(1982)
All unranked informal groups introduced by
Trelease (1913) and Gentry (1982) are treated
here as published by either Trelease or Gentry if
no earlier rankless homonym with the explicit de-
nomination “group” could be found in original
sources or in Mottram (2015).
Trelease (1913), Caribbean region
In his treatment of the agaves (all in A.subg.
Agave) of the Caribbean region, Trelease (1913)
classified the recognised species into seven un-
ranked infrageneric “Groups” (Trelease, 1913: 8–
10, 15–16), each with a brief English description.
Trelease did not indicate types in the diagnoses of
the individual groups. However, we interpret the
statement towards the end of p. 8: “The charac-
ters and affinities of the groups are shown more
clearly in a contrast of the species taken as typical
of each”, and the subsequent indication of the
“species taken as typical” towards the top of p. 9,
as a clear indication of the types.
Agave [infragen. unranked, “group”] Vivipa-
rae Baker, Gard. Chron. ser. nov. 8: 780, 808.
1877. Type: A.vivipara L.
Trelease (1913: 8–9, 16, 17) did not explicitly
refer to Baker’s work and treated this group as
newly published by himself. However, it had been
published earlier by Baker (1877; see page refe-
rences above). As neither of these two names was
published with a clear indication of rank, neither
can be considered superfluous under ICN Art.
52.1 (Turland et al. 2018), as priority is rank-li-
mited (ICN Art. 11.2) and for an unranked taxon
there cannot be any name that ought to have been
adopted. Baker (1877) did not indicate a type. The
designation of A. vivipara as type by Trelease
(1913: 9) was not required, since that species
[A.vivipara] is automatically the type according
to ICN Art. 10.8. and Ex. 12 (Turland et al., 2018).
244 Bradleya 37/2019
Figure 6. Agave geminiflora (part of the inflores-
cence) is the type of A. subg. Littaea and of A. sect.
Littaea. Photograph: Gideon F. Smith.
Figure 7. Agave striata (here: subsp. striata; cul-
tivated plant with inflorescences) is the type of A.
sect. Juncineae [A. subg. Littaea]. Cultivated,
Planten un Blomen, Hamburg, Germany, 4 July
2018. Photograph: Joachim Thiede.

Agave [infragen. unranked, “group”] Cari-
baeae Trel., Mem. Natl. Acad. Sci. 11: 9, 16,
21. 1913. Type: A. karatto Mill. The designa-
tion of A. karatto as type by Trelease (1913: 9)
is in accordance with ICN Art. 10.8 (Turland et
al., 2018).
Agave [infragen. unranked, “group”] Antil-
lanae Trel., Mem. Natl. Acad. Sci. 11: 9, 16,
29. 1913. Type: A. antillarum Descourt.
Agave [infragen. unranked, “group”] Baha-
manae Trel., Mem. Natl. Acad. Sci. 11: 9, 16,
38. 1913. Type: A. bahamana Trel.
Agave [infragen. unranked, “group”] Antil-
lares Trel., Mem. Natl. Acad. Sci. 11: 9, 16, 42.
1913. Type: A. willdingii Tod.
Agave [infragen. unranked, “group”] Ina-
guenses Trel., Mem. Natl. Acad. Sci. 11: 9, 16,
45. 1913. Type: A. nashii Trel. The designation
of A. nashii Trel. as type by Trelease (1913: 9)
is in accordance with ICN Art. 10.8. (Turland
et al., 2018).
Agave [infragen. unranked, “group”] Sisala-
nae Trel., Mem. Natl. Acad. Sci. 11: 9, 16, 47.
1913. Type: A. angustifolia Haw. The designa-
tion of A. angustifolia Haw. as type by Tre-
lease (1913: 9) is in accordance with ICN Art.
10.8. (Turland et al., 2018).
Gentry (1982), Continental North America
In the monumental treatment of the species of
Agave (s.s.) that occur in continental North Ame-
rica, and slightly beyond, Gentry (1982) presen-
ted his approximately 30 years of field work,
extensive institutional visits to study herbarium
material, and familiarity with the pertinent lite-
rature available at the time. Gentry subdivided
A. subg. Agave and A. subg. Littaea into informal
unranked infrageneric groups, eight in A. subg.
Littaea and 12 in A. subg. Agave (Gentry 1982: xii,
62, 269). He indicated that he followed Trelease
(1913) and Berger (1915) in his groupings but did
not hesitate to “redefine and make new groupings
whenever new evidence (specimens) recommen-
245
Bradleya 37/2019
Figure 8. Agave parviflora is the type of A. sect. Parviflorae [A. subg. Littaea]. Nogales, Arizona, USA.
2 August 2017. Photograph: Gideon F. Smith.

ded changes” (Gentry 1982: xi). In the diagnoses of
the individual infrageneric groupings, Gentry al-
ways designated these as “group” and refers to
“Group relationships”, designations which he used
throughout in the taxonomic accounts.
In a “Summary of Agave Taxa”, Gentry (1982:
xii) labels the groups as “Section[s]”, and he also
provided a “Sectional List of Species” for each sub-
genus, on page 62 for A. subg. Littaea and on page
269 for A. subg. Agave. However, Gentry (1982)
gave inadequate guidance on and was inconsis-
tent in how exactly he viewed the relationship bet-
ween his “Groups” and “Sections”. For example,
Gentry (1982: 61), the page before he gives a
sectional list for A. subg. Littaea, refers to these
same groupings as “Groups”. Based on the desig-
nation “sectional list” for both subgenera, Mot-
tram (2015), in his “Annotated checklist of the
infragenera of Agave L.”, treated all of Gentry’s
Groups as having been published at the rank of
section. We do not follow this view here, since all
infrageneric taxa are designated as “group” in the
diagnoses and by Gentry (1982: xi) himself where
he introduces his work, stating that “These groups
are tantamount to sections in more formal taxo-
nomy”, without unambiguously accepting them as
“sections”. In our view Gentry therefore stopped
short of treating his Groups as Sections. Under
ICN Article 37.1 (Turland et al., 2018) which re-
quires “...a clear indication of the rank of the
taxon concerned...” the group names that Gentry
(1982) proposed were not validly published. Fur-
thermore, where Gentry’s classification system
has been widely adopted by later agave taxono-
mists, they almost invariably use the designation
“group”, and not “section”, in taxonomic treat-
ments or when describing new taxa.
Agave subg. Littaea
Agave [infragen. unranked, “group”] Amolae
Gentry, Agaves Cont. N. Amer. 63. 1982 nom.
inval. Art 37.1. Type: A. attenuata Salm-Dyck
(Figure 22). Not validly published.
Agave [infragen. unranked, “group”]
Choritepalae Gentry, Agaves Cont. N. Amer.
89. 1982 nom. inval. Art 37.1. Type: A. elle-
meetiana K.Koch. Not validly published.
Agave [infragen. unranked, “group”] Filif-
erae Baker, Gard. Chron. ser. nov. 7: 171, 303.
1877. Type: A. filifera Salm-Dyck.
Gentry (1982: 101) did not refer to Baker
(1877) and treated this group as newly published
by himself. Baker (1877: 171) listed A. filifera and
A.schidigera Lem. as “Examples” (= syntypes),
and Gentry (1982) designated A. geminiflora as
type. The type is A. filifera according to ICN Art.
10.8. & Ex. 12 (Turland et al., 2018).
Agave [infragen. unranked, “group”] Mar-
ginatae Baker, Gard. Chron. ser. nov. 7: 171,
368. 1877. Type (lectotype): A. lophantha
Schiede ex Kunth (Figure9).
Gentry (1982: 124) did not refer to Baker
(1877) and treated this group as newly published
by himself. Baker (1877: 171) listed A. lophantha
and A. univittata Haw. as “Examples” (= syntyp-
es). The designation of A. lophantha as “Typical
species” by Gentry (1982) is viewed as a lectotyp-
ification.
Agave [infragen. unranked, “group”] Parvi-
florae Gentry, Agaves Cont. N. Amer. 195.
1982 nom. inval. Art 37.1. Type: A. parviflora
Torr. (Figure8). Not validly published.
Agave [infragen. unranked, “group”] Poly-
cephalae Gentry, Agaves Cont. N. Amer. 216.
1982 nom. inval. Art 37.1. Type: A. celsii Hook.
(accepted name: A. mitis Mart.). Not validly
published.
246 Bradleya 37/2019
Figure 9. Agave lophantha is the accepted name
of the type, A. heteracantha, of A. sect. Hetera-
canthae [A. subg. Littaea]. Cultivated, Wallace
Gardens, Phoenix, Arizona, USA, 10 July 2014.
Photograph: Gideon F. Smith.

Agave [infragen. unranked, “group”] Stri-
atae Baker, Gard. Chron. ser. nov. 7: 171, 8:
556. 1877. Type: A. striata Zucc. (Figure7).
Baker (1877) listed A. striata and A. dasyli-
rioides Jacobi & C.D.Bouché as “Examples” (=
syntypes), and Gentry (1982: 235), who referred
to Baker, designated A. striata as “Typical
species”; A. striata is the automatic type according
to ICN Art. 10.8. & Ex. 12 (Turland et al., 2018).
Agave [infragen. unranked, “group”] Urceo-
latae Gentry, Agaves Cont. N. Amer. 251.
1982 nom. inval. Art 37.1. Type: A. utahensis
Engelm. (Figure1). Not validly published.
Agave subg. Agave
Agave [infragen. unranked, “group”] Ameri-
canae Baker, Gard. Chron. ser. nov. 7: 171,
200. 1877 nom.inval. Art. 22.1. Type: A. amer-
icana L. (Figure 21).
Baker (1877) listed A. americana and A. pota-
torum Zucc. as “Examples” (= syntypes), and Gen-
try (1982: 270), who referred to Baker (with the
wrong citation “1888: 175”), designated A. ameri-
cana as “Typical species”. Agave americana is the
correct type according to ICN Art. 10.8. & Ex. 12
(Turland et al., 2018). Since this grouping includes
A. americana, the type of Agave, its valid name is
A. “group” Agave according to ICN Art. 22.1. (Tur-
land et al., 2018).
Agave [infragen. unranked, “group”] Cam-
paniflorae Trel., Annual Rep. Missouri Bot.
Gard. 22: 44. 1912 (dated 1911). Type (lecto-
type): A. aurea Brandegee (Figure10).
Trelease (1912) did not designate a type. Gen-
try (1982: 309), who referred to Trelease, designa-
ted A. aurea as “Typical species” which is viewed
as a lectotypification (Starr & Webb, 2015: 36).
Agave [infragen. unranked, “group"] Crena-
tae Gentry, Agaves Cont. N. Amer. 251. 1982
nom. illeg. Art. 53.3 & nom. inval. Art 37.1.
Type: A. inaequidens K.Koch.
Gentry (1982: 323) referred to A. “Unterreihe”
Crenatae A.Berger (1915: 194), but treats Crena-
tae as a “group” which is seen as independent pu-
blication of a new unranked infrageneric name
(cited above). As neither of these two names was
published with a clear indication of rank, neither
can be considered superfluous under Art. 52.1 of
Turland et al. (2018), as priority is rank-limited
(Art. 11.2), and for an unranked taxon there can-
not be any name that ought to have been adopted.
The designation of A. inaequidens K.Koch as
“Typical species” (= type) by Gentry (1982: 323) is
in accordance with ICN Art. 10.8. Consequently,
A.“group” Crenatae Gentry represents an illegiti-
mate homonym of A. “Unterreihe” Crenatae
A.Berger according to ICN Art. 53.3 (Turland et
al., 2018).
Agave [infragen. unranked, “group”] Deser-
ticolae Trel., Annual Rep. Missouri Bot. Gard.
22: 44–45. 1912 (dated 1911). Type (lectotype):
A. deserti Engelm. (Figure13).
Trelease (1911) did not designate a type. Gen-
try (1982: 354), who referred to Trelease, desig-
nated A. deserti as “Typical species” which is
viewed as a lectotypification (Starr & Webb, 2015:
36).
Agave [infragen. unranked, “group”]
Ditepalae Gentry, Agaves Cont. N. Amer.
416. 1982 nom. inval. Art 37.1. Type: A. shre-
vei Gentry (Figure12). Not validly published.
Agave [infragen. unranked, “group”] Hiemi-
florae Gentry, Agaves Cont. N. Amer. 465.
1982 nom. inval. Art 37.1. Type: A. hiemiflora
Gentry. Not validly published.
247
Bradleya 37/2019
Figure 10. Agave aurea subsp. aurea. Agave
aurea is the type of A. sect. Campaniflorae [A.
subg. Agave]. Cultivated, Arizona-Sonora Desert
Museum, near Tucson, Arizona, 13 April 2014.
Photograph: Gideon F. Smith.

Agave [infragen. unranked, “group”] Mar-
moratae Gentry, Agaves Cont. N. Amer. 507.
1982 nom. inval. Art 37.1. Type: A. marmorata
Roezl. Not validly published.
Gentry (1982: 507) referred to A.“Reihe” Mar-
moratae A.Berger (1915: 269), but treats Mar-
moratae as a “group”, which is interpreted as the
publication of a new, unranked infrageneric name
(cited above). Agave “Reihe” Marmoratae A.Ber-
ger and A.“group” Marmoratae Gentry are based
on the same type, A.marmorata, according to ICN
Art. 10.8. & Ex. 12. Agave “group” Marmoratae
and A.“Reihe” Marmoratae therefore do not re-
present homonyms according to ICN Art. 53.3.
Since both A. “Reihe” Marmoratae A.Berger and
A.“group” Marmoratae Gentry are typified by the
same element (i.e. A.marmorata), Gentry’s cita-
tion was merely the later use of Berger’s name.
Agave [infragen. unranked, “group”] Par-
ryanae Gentry, Agaves Cont. N. Amer. 520.
1982 nom. inval. Art 37.1. Type: A. parryi En-
gelm. (Figure15). Not validly published.
Agave [infragen. unranked, “group”] Rigidae
Baker, Gard. Chron. ser. nov. 7: 171, 8: 264.
1877. Type: A. rigida Mill. (accepted name:
A.angustifolia Haw.).
Gentry (1982) referred to A.“Reihe” Rigidae
A.Berger (1915: 226), omitting that his “group” Ri-
gidae was published earlier by Baker as cited
above. Baker (1877: 171) listed A. rigida and
A.xalapensis Roezl ex Jacobi as “Examples” (=
syntypes), but later (1877: 264) designated A.lu-
rida Aiton and A.ixtli Karw. ex Salm-Dyck “as
the types” (= syntypes). Starr & Webb (2015: 37)
designated A. ixtli as lectotype from these two.
Gentry (1982: 101) designated A. angustifolia
Haw. as “Typical species”. However, the type is
A.rigida Mill. according to ICN Art. 10.8. & Ex.
12 (Turland et al., 2018).
Agave [infragen. unranked, “group”] Salmi-
anae Gentry, Agaves Cont. N. Amer. 594.
1982 nom. inval. Art 37.1. Type: A. salmiana
Otto ex Salm-Dyck.
Gentry (1982: 594) referred to A. “Reihe”
Salmianae A.Berger (1915: 128), but treats Salmi-
anae as a “group” which is seen as publication of a
new unranked infrageneric name (cited above).
Agave “Reihe” Salmianae A.Berger and A.“group”
Salmianae Gentry are based on the same type,
A.salmiana, and according to ICN Art. 10.8. & Ex.
12 therefore do not represent homonyms, based on
ICN Art. 53.3. (Turland et al., 2018).
248 Bradleya 37/2019
Figure 11. Agave shawii subsp. shawii. Agave shawii is the type of A. sect. Umbelliflorae [A. subg. Agave].
Cultivated, Desert Botanical Garden, Phoenix, Arizona, 9 April 2014. Photograph: Gideon F. Smith.

Agave [infragen. unranked, “group”] Sisala-
nae Trel., Mem. Natl. Acad. Sci. 11: 8–9, 16,
47. 1913. Type: A. angustifolia Haw.
Gentry (1982: 619) referred to Trelease and de-
signated A. sisalana Perrine as “Typical species”,
but the type is A. angustifolia as designated by
Trelease (see chapter “Trelease (1913), Caribbean
region”).
Agave [infragen. unranked, “group”] Umbel-
liflorae Trel., Annual Rep. Missouri Bot.
Gard. 22: 44. 1912 (dated 1911). Type (lecto-
type): A. shawii Engelm. (Figure11).
Trelease (1912) did not designate a type. Gen-
try (1982: 635), who referred to Trelease, desig-
nated A. shawii as “Typical species” which is
viewed as a lectotypification (Starr & Webb, 2015:
38).
2) Nomenclatural assessment of infragene-
ric taxa published in Agave at sectional rank
by Salm-Dyck, Jacobi, Engelmann, Berger,
Ullrich, Webb & Starr, and Starr & Webb
Salm-Dyck (1861)
Salm-Dyck (1859), who is arguably better
known for his work on cacti (Salm-Dyck, 1850)
and aloes and mesembs (Salm-Dyck, 1836–1863),
published an informal infrageneric classification
of Agave (s.l.) and established five “Unterabthei-
lungen” (§. I.–§. V.) based on leaf characters, with
contextual Latin diagnoses provided for each Un-
terabtheilung. He divided 47 of the species known
to him among these Unterabtheilungen; he regar-
ded five species as requiring further investigation
to facilitate their correct placement (“Species du-
biosae, amplius inquirendae”).
Two years later, Salm-Dyck (1861) amended
his earlier classification system and recognised
the same five groupings at sectional rank and pu-
blished one further section “Juncineae” (§. I.–§.
VI), all with contextual Latin diagnoses, thus pro-
viding the first formal infrageneric classification
of Agave (s.l.) at sectional rank. Sectional rank for
all six groupings is clearly indicated by his conti-
nuous use of the denomination “Section” for them.
Salm-Dyck did not indicate types for any of the
sectional names that he published; lectotypes are
designated here where necessary.
Agave sect. Macracanthae Salm-Dyck, Wo-
chenschr. Vereines Beförd. Gartenbaues Kö-
nigl. Preuss. Staaten 4(23): 181. [6 Jun.] 1861
nom.inval. Art. 22.1. & Ex. 2. Type: A. ameri-
cana L. (Figure 21).
Salm-Dyck included 29 binomials in A. sect.
Macracanthae. Since A. sect. Macracanthae in-
cludes A. americana, the type of the genus Agave,
it is invalid under ICN Art. 22.1. & Ex. 2 (Turland
et al., 2018).
Agave sect. Heteracanthae Salm-Dyck, Wo-
chenschr. Vereines Beförd. Gartenbaues Ko-
nigl. Preuss. Staaten 4(23): 182. [6 Jun.] 1861.
Type: A. heteracantha Zucc. (accepted name:
A. lophantha Schiede ex Kunth) (Figure9).
Salm-Dyck included seven binomials that all
place in A. “group” Marginatae of A. subg. Littaea.
From these, A. heteracantha is the type according
to ICN Art. 10.8. & Ex. 12 (Turland et al., 2018).
At sectional rank, A. sect. Heteracanthae corres-
ponds with, and must be adopted for what Baker
(1877 [7]: 171, 368) later treated as A. “group”
Marginatae Baker.
Agave sect. Micracanthae Salm-Dyck, Wo-
chenschr. Vereines Beförd. Gartenbaues Kö-
nigl. Preuss. Staaten 4(23): 182. [6 Jun.] 1861.
Type: A. micracantha Salm-Dyck (accepted
name: A.mitis Mart.).
Salm-Dyck (1859) included eleven binomials
that fall in A.“group” Rigidae (three species) and
Caribaeae (one species) of A. subg. Agave,
A.“group” Polycephalae (five species) and Amolae
(one species) of A. subg. Littaea., and A. subg.
Manfreda (one species). The type is A. micra-
cantha Salm-Dyck according to ICN Art. 10.8. &
Ex. 12 (Turland et al., 2018), so that A. sect. Mi-
cracanthae is the valid name at sectional rank for
the group that corresponds with what Gentry
(1982: 216) treated as A. “group” Polycephalae
Gentry.
Agave sect. Inermes Salm-Dyck, Wochenschr.
Vereines Beförd. Gartenbaues Königl. Preuss.
Staaten 4(23): 182. [6 Jun.] 1861. Type (lecto-
type, designated here): A.attenuata Salm-
Dyck (Figure 22).
Salm-Dyck (1859) included three binomials
that fall in A.“group” Amolae (one species) and Fi-
liferae (two species) of A. subg. Littaea. From
these, A.attenuata of A.“group” Amolae is desig-
nated here as lectotype, so that A. sect. Inermes is
the valid name at sectional rank that corresponds
to and is homotypic with A.“group” Amolae Gen-
try (1982: 63).
Agave sect. Juncineae Salm-Dyck, Wochenschr.
Vereines Beförd. Gartenbaues Königl. Preuss.
Staaten 4(23): 182. [6 Jun.] 1861. Type (lecto-
type, designated here): A.striata Zucc. (Fig-
ure7).
Salm-Dyck (1861) included three binomials
that he earlier (Salm-Dyck 1859: 87) classified in
A. “Unterabtheilung” Inermes (see above); these
place in A. “group” Filiferae (one species) and
Striatae (two species) of A. subg. Littaea. From
these, A.striata of A.“group” Striatae is designa-
249
Bradleya 37/2019

ted here as lectotype, so that A. sect. Juncineae is
the valid name at sectional rank that corresponds
to and is homotypic with A. “group” Striatae
Baker (1877 [7]: 171, [8]: 556).
Agave sect. Herbaceae Salm-Dyck, Wochenschr.
Vereines Beförd. Gartenbaues Königl. Preuss.
Staaten 4(23): 182. [6 Jun.] 1861. Type (lecto-
type, designated here): A. brachystachys
Cav. (accepted name: A.scabra Ortega).
Salm-Dyck (1859) included six binomials that
fall in A.“group” Manfreda Thiede (Thiede, 2001:
11) of A. subg. Manfreda (five species of the for-
mer genus Manfreda), and in A. “group” Amolae
(one species) of A. subg. Littaea. From these,
A.brachystachys is designated here as lectotype,
so that A. sect. Herbaceae is the valid name at
sectional rank that corresponds to A. “group”
Manfreda.
Jacobi (1864, 1868)
Shortly after Salm-Dyck (1859, 1861) publis-
hed his classification for Agave, Jacobi (1864) pro-
posed a more elaborate classification for the genus
in which he separated Agave (s.l.) into four
“Hauptgruppen” (p. 465, = “main groups”), viz.,
Keratacanthae, Chondracanthae, Inermes, and
Herbaceae. However, he did not recognise these at
a formal rank, and they are therefore unranked
groups. In the Keratacanthae, Jacobi recognised
eight lower ranked, but still unranked groupings;
seven of these he published as new, whereas the
“Juncineae” were adopted from Salm-Dyck (1861).
Jacobi uses the informal denomination “Abthei-
lung” (p. 504) for his groupings and “Unterabthei-
lung” (p. 464, 507) for his subgroupings. A rank
therefore cannot be asigned to them, especially
also because his classification is essentially based
on that of Salm-Dyck (1859), where “Unterab-
theilungen”, not sections, were used. Further, the
unranked “Marginatae” represent a mere rena-
ming of Salm-Dyck’s A. sect. Heteracanthae
(Salm-Dyck, 1861) with a more descriptive name,
as he (Jacobi, 1864: 505–506) clearly admits. Four
years later, Jacobi (1868) slightly amended his
classification system by renaming A. “Abtheilung”
Subcarinatae as A. “Abtheilung” Aloideae.
Engelmann (1875)
Engelmann (1875: 296–297, 300–301, 304,
309) provided an infrageneric classification of
Agave in which he recognised three sections based
on inflorescence types. These sections correspond
to the three subgenera A. subg. Agave, A. subg.
Littaea, and A. subg. Manfreda. Engelmann did
not indicate types so that lectotypes are designa-
ted here where necessary.
Agave sect. Singuliflorae Engelm., Trans.
Acad. Sci. St. Louis 3: 296, 300–301. 1875.
Type (lectotype, designated here): A.virgi-
nica L. (Figure18).
Engelmann included three binomials that
place in A. “group” Manfreda of A. subg. Man-
freda. From these, A. virginica, the type of the
genus Manfreda, is designated here as lectotype,
so that A. sect. Singuliflorae becomes homotypic
with Manfreda and a heterotypic synonym of the
earlier A. sect. Herbaceae.
Agave sect. Geminiflorae Engelm., Trans.
Acad. Sci. St. Louis 3: 296–297, 304. 1875.
Type (lectotype, designated here): A. heter-
acantha Zucc. (accepted name: A.lophantha
Schiede ex Kunth) (Figure9).
At sectional rank, A. sect. Geminiflorae cor-
responds to A. subg. Littaea. Engelmann (1875)
included five binomials that place in A.“group”
Marginatae (one species), Parviflorae (two spe-
cies), Striatae (one species), and Urceolatae (one
species) of A. subg. Littaea. From these, A.hete-
racantha is designated here as lectotype. This re-
sults in A. sect. Geminiflorae becoming a later
homotypic synonym of A. sect. Heteracanthae, and
rids Agave nomenclature of a meaningless name
at the rank of section. Mottram (2015: 16) consi-
ders Littaea geminiflora Tagl. (≡ A.geminiflora
(Tagl.) Ker Gawl.) as “Autotype (Art. 7.4)” because
of the epithet chosen, “Geminiflorae”. Since
A. sect. Geminiflorae is a new name at sectional
rank, and since Engelmann (1875) did not include
A. geminiflora and mentioned Littaea in syno-
nymy we do not follow Mottram (2015).
Agave sect. Paniculatae Engelm., Trans. Acad.
Sci. St. Louis 3: 297, 309. 1875 nom.inval. Art.
22.1. & Ex. 2. Type: A.americana L. (Figure
21).
Engelmann (1875) included eight binomials,
and also mentioned A. americana. Since A. sect.
Paniculatae includes A. ameri cana, the type of
Agave, it is not valid under ICN Art. 22.1. & Ex. 2
(Turland et al., 2018).
Bentham (1883)
Bentham (in Bentham & Hooker 1883: 733),
in his general plant classification, combined Lit-
taea and Manfreda at sectional rank in Agave.
Agave sect. Littaea (Tagl.) Benth., Gen. Pl.
[Benth. & Hook.f.] 3(2): 733. 1883. Type: Lit-
taea geminiflora Tagl. (type of Littaea; = A.ge-
miniflora (Tagl.) Ker Gawl.) (Figure6).
Agave sect. Littaea is a new combination based
on the genus Littaea Tagl. and is the valid name
at sectional rank corresponding to A.“group” Fil-
iferae Gentry.
250 Bradleya 37/2019

Agave sect. Manfreda (Salisb.) Benth., Gen. Pl.
[Benth. & Hook.f.] 3(2): 733. 1883. Type: Agave
virginica L. (Figure18).
Agave sect. Manfreda is a new combination
based on the genus Manfreda Salisb. and is a he-
terotypic synonym of A. sect. Herbaceae.
Berger (1915)
In his monograph of Agave (s.l.), Berger (1915)
recognized the three subgenera A. subg. Man-
freda, A. subg. Littaea, and A. subg. Agave (then
called A. subg. Euagave nom. inval.). Whereas
A. subg. Littaea is subdivided into seven formal
sections, all published as new (partly with sub-
sections), A. subg. Agave is treated differently as
it is classified into 18 informal “Reihen” (not secti-
ons), partly with informal “Unterreihen”. Berger
did not indicate types for his new sectional names,
so that lectotypes are designated here where ne-
cessary.
Mottram (2015) treats the informal “Reihen”
in A. subg. Agave as published at sectional rank,
in analogy to A. subg. Littaea. Since Berger (1915)
used the designation “section” only for groupings
in A. subg. Littaea, but not in A. subg. Agave
where he used the designation “Reihe” throug-
hout, we do not here follow the view of Mottram
(2015).
Alwin Berger produced several enormously in-
fluential taxonomic works on a range of succulent
plant taxa, and his views on their classification
were adopted by a number of researchers who fol-
lowed after him (see for example Klopper et al.,
2013 on the genus Aloe).
Agave sect. Anacamptagave A.Berger, Agaven
38–39. 1915. Type (lectotype, designated
here): A.mitis Mart.
Berger (1915) included 27 binomials that place
in A. “group” Amolae (two species) and Polyce-
phalae (21 species) of A.subg. Littaea; two enti-
ties are hybrids, and two species are unplaced.
From the 27 binomials, A.mitis of A.“group” Po-
lycephalae is designated here as lectotype, so that
A.sect. Anacamptagave becomes a synonym of the
earlier A.sect. Micracanthae.
Agave sect. Xysmagave A.Berger, Agaven 68.
1915. Type (lectotype, designated here):
A.geminiflora (Tagl.) Ker Gawl. (Figure6).
Berger (1915) included 11 binomials that place
in A.“group” Filiferae (six species) and Parviflorae
(three species) of A.subg. Littaea; two entities are
hybrids. From these, A.geminiflora is designated
here as lectotype, so that A.sect. Xysmagave be-
comes a synonym of the earlier A.sect. Littaea.
Agave sect. Schoenagave A.Berger, Agaven 78.
1915. Type (lectotype, designated here):
A.striata Zucc. (Figure7).
Berger (1915) included three binomials that all
place in A.“group” Striatae of A.subg. Littaea.
From these, A.striata is designated here as lecto-
type, so that A.sect. Schoenagave becomes homo-
typic with, and a synonym of, the earlier A.sect.
Juncineae.
Agave sect. Chonanthagave A.Berger, Agaven
83. 1915. Type (lectotype, designated here):
A.dasylirioides Jacobi & C.D.Bouché.
Agave sect. Chonanthagave corresponds to A.
“Abtheilung” Loriformes (Jacobi, 1864); Berger
(1915) included the same two binomials that both
place in A.“group” Striatae of A.subg. Littaea.
From these, A.dasylirioides is designated here as
lectotype, so that A.sect. Chonanthagave becomes
homotypic with A. "Abtheilung" Loriformes, and a
heterotypic synonym of the earlier A.sect. Junci-
neae.
Agave sect. Pericamptagave A.Berger, Agaven
85. 1915. Type (lectotype, designated here):
A.lophantha Schiede ex Kunth (Figure9).
Berger (1915) included 27 binomials that place
in A.“group” Marginatae (24 species) and Urceo-
latae (one species) of A.subg. Littaea; one entity is
a hybrid, and one species is unplaced. From the
27 binomials, A.lophantha of A.“group” Margi-
natae is designated here as lectotype, so that
A.sect. Pericamptagave becomes a heterotypic sy-
nonym of the earlier A.sect. Heteracanthae.
Agave sect. Brachysolenagave A.Berger, Aga-
ven 116. 1915. Type (lectotype, designated
here): A.ghiesbreghtii Verschaff.
Berger (1915) included six binomials that
place in A. “group” Marginatae (two species) of
A.subg. Littaea; one entity is a hybrid, and three
species are unplaced. From the six binomials,
A.ghiesbreghtii of A.“group” Marginatae is de-
signated here as lectotype, so that A.sect. Bra-
chysolenagave becomes a heterotypic synonym of
the earlier A.sect. Heteracanthae.
Agave sect. Anoplagave A.Berger, Agaven 121.
1915. Type (lectotype, designated here):
A.attenuata Salm-Dyck (Figure 22).
Berger (1915) included six binomials that
place in A. “group” Amolae (four species) and
Choritepalae (two species) of A. subg. Littaea.
From these, A.attenuata of A.“group” Amolae is
designated here as lectotype, so that A.sect. Ano-
plagave becomes a homotypic synonym of the ear-
lier A.sect. Inermes.
Ullrich (1991–1995)
Ullrich (1991b, 1992, 1993, 1995) published
three new combinations at sectional rank for older
names, and one new name at sectional rank.
251
Bradleya 37/2019

Agave sect. Nizandensae (B.Ullrich) B.Ullrich,
Succulenta (NL) 70(6): 144. 1991 [1991b].
Type: A.nizandensis Cutak.
First published at the rank of series (Ullrich
1991a: 91). Its sole species was placed in
A.“group” Amolae of A.subg. Littaea by Gentry
(1982: 62). Agave sect. Nizandensae is placed in
the synonymy of A. sect. Inermes.
Agave sect. Guatemalenses (Trel.) B.Ullrich,
Kakt. and. Sukk. 43(9): 215. 1992. Type (lecto-
type): A.hurteri Trel. (Figures16 & 17).
First published as A.“group” Guatemalenses
Trelease (1915: 134) of A.subg. Agave, without des-
ignation of a type. Agave guatemalensis A.Berger
(1915: 201) was not mentioned by Trelease (1915)
who may not have been aware of Berger’s mono-
graph at the time. Ullrich (1992) designated
A.hurteri as “Typische Art”, which is interpreted
as a lectotypification. Agave sect. Guatemalenses,
which corresponds to A.“group” Hiemiflorae Gen-
try, must be adopted at sectional rank.
Agave sect. Crenatae (A.Berger) B.Ullrich, Pi-
ante Grasse 13(2): 53. 1993. Type: A. crenata
Jacobi (accepted name: A. inaequidens K.Koch).
First published as A.“Unterreihe” Crenatae of
A.subg. Agave. Neither Berger nor Ullrich desig-
nated a type; the type is A.crenata according to
ICN Art. 10.8. & Ex. 12 (Turland et al., 2018).
Agave sect. Crenatae is the name that must be
adopted at sectional rank for A.“group” Crenatae.
Agave sect. Yuccifoliae (A.Terracc.) B.Ullrich,
Piante Grasse 15(4): 29. 1995. Type: A.yucci-
folia F.Delaroche (sole element included) (ac-
cepted name: A.spicata Cav.).
First published as A. ser. Yuccifoliae (as ‘Yuc-
caefoliae’) of A. subg. Littaea (as A. subg. Apla-
gave) by Terracciano (1885: 14). Ullrich (1995)
wrongly used the rank “subseries” for the ba-
sionym; the rank “series” is clearly indicated by
Terracciano (1885: 14). The same combination
Agave sect. Yuccifoliae was not validly published
earlier by Breitung (1960: 181). This monotypic
section comprises only the taxonomically anom-
alous A. yuccifolia, which was placed in A. “group”
Amolae by Gentry (1982: 62). Agave sect. Yuccifo-
liae is a heterotypic synonym of A. sect. Inermes.
Hochstätter (2015)
Hochstätter (2015), in his illustrated synopsis
of Agave (s.s.), published many new infrageneric
taxa at the rank of section, series, and subseries.
Here, only the 10 new names and new combinati-
ons at sectional rank are considered.
Agave sect. Bechtoldiae Hochstätter, Agave I:
21. 2015. Type: A.victoriae-reginae T.Moore.
This new section in A.subg. Littaea is split off
from Gentry’s A.“group” Marginatae for A.victo-
riae-reginae and related species. It is placed here
in the synonymy of A. sect. Heteracanthae, the
valid name at sectional rank corresponding to
A.“group” Marginatae.
Agave sect. Choritepalae Hochstätter, Agave I:
14. 2015. Type: A.ellemeetiana K.Koch.
This new section in A.subg. Littaea provides a
valid name for Gentry’s A. “group” Choritepalae
at sectional rank with the same type. Hochstätter
(2015) uses the author designation “Gentry ex
Hochstätter” for this as well as his further new
names at the rank of section that correspond to
Gentry’s groups, but this is not followed here since
Gentry did not publish his “group” names at
sectional rank (see above).
Agave sect. Ditepalae Hochstätter, Agave I: 26.
2015. Type: A.shrevei Gentry (Figure12).
This new section in A.subg. Agave provides a
valid name for Gentry’s A. “group” Ditepalae at
sectional rank with the same type.
Agave sect. Hibernicae Hochstätter, Agave I:
31. 2015. Type: A.gentryi B.Ullrich.
This new section in A.subg. Agave comprises
the two species, A.gentryi and A.montana Villar-
real, which were both formerly placed in
A. macroculmis sensu Gentry (1982: 598), in
A.“group” Salmianae. Alsemgeest & Roosbroeck
(c. 2012) suggested this grouping earlier as “Par-
rasana groep” and also included A. parrasana
A.Berger.
Agave sect. Hiemiflorae Hochstätter, Agave I:
34. 2015. Type: A.hiemiflora Gentry.
This new section in A.subg. Agave provides a
252 Bradleya 37/2019
Figure 12. Agave shrevei subsp. shrevei. Agave
shrevei is the type of A. sect. Ditepalae [A. subg.
Agave]. Cultivated, Arizona-Sonora Desert Mu-
seum near Tucson, Arizona, USA, 11 April 1992.
Photograph: Joachim Thiede.

valid name for Gentry’s A.“group” Hiemiflorae at
sectional rank with the same type and is treated
here as a synonym of the earlier A.sect. Guate-
malenses.
Agave sect. Hodgsoniae Hochstätter, Agave I:
30. 2015. Type: A.murpheyi F.Gibson.
This new section in A.subg. Agave was split
off from Gentry’s A. “group” Ditepalae for
A.murpheyi and further cultigen species mainly
from pre-Columbian sites of habitation of native
Americans in Arizona. It is here treated in the
synonymy of A.sect. Ditepalae.
Agave sect. Inaguenses (Trel.) Hochstätter,
Agave I: 40. 2015. Type: A. nashii Trel.
This new section in A.subg. Agave is a new
combination at sectional rank based on A.“group”
Inaguenses Trelease (1913: 45).
Agave sect. Parryanae Hochstätter, Agave I: 29.
2015. Type: A.parryi Engelm. (Figure15).
This new section in A.subg. Agave provides a
valid name for A. “group” Parryanae Gentry
(1982: 520) at sectional rank with the same type.
Agave sect. Parviflorae Hochstätter, Agave I:
16. 2015. Type: A.parviflora Torr. (Figure8).
This new section in A.subg. Littaea provides a
valid name for A. “group” Parviflorae Gentry
(1982: 195) at sectional rank with the same type.
Agave sect. Urceolatae Hochstätter, Agave I:
18. 2015. Type: A. utahensis Engelm. (Fig-
ure1).
This new monotypic section in A.subg. Littaea
provides a valid name for A. “group” Urceolatae
Gentry (1982: 251) at sectional rank.
253
Bradleya 37/2019
Figure 13.
Agave deserti subsp. simplex. Agave de-
serti is the type of A. sect. Deserticolae [A. subg.
Agave]. Pima county, west of Tucson, Arizona, USA,
13 April 2014. Photograph: Gideon F. Smith.
Figure 14. Agave avellanidens is the type of A.
sect. Conicae [A. subg. Agave]. Cultivated, Ari-
zona-Sonora Desert Museum, Tucson, Arizona, 13
April 2014. Photograph: Gideon F. Smith.

Webb & Starr (2015) and Starr & Webb (2015)
In creating a framework for a revision of Agave
on Baja California, Webb & Starr (2015) publis-
hed two new sections (A.sect. Conicae and A.sect.
Intermediae), and four new combinations at
sectional rank for informal, rankless group names
introduced by Trelease (1912). At the rank of
section these names were at first not validly pu-
blished in Webb & Starr (2015) and later valida-
ted by Starr & Webb (2015).
Agave sect. Conicae R.H.Webb & G.D.Starr,
Haseltonia 20: 71. 2015. Type: A.avellanidens
Trel. (Figure14).
This new section in A.subg. Agave includes
three species formerly placed in Gentry’s A.
“group” Deserticolae (A.avellanidens, A. gigan-
tensis Gentry, A.moranii Gentry), as well as the
recently described A. turneri R.H.Webb &
Salazar-Ceseña.
Agave sect. Intermediae R.H.Webb &
G.D.Starr, Haseltonia 20: 87. 2015. Type:
A.vizcainoensis Gentry.
This new section in A.subg. Agave includes
A.vizcainoensis that was formerly placed in Gen-
try’s A. “group” Deserticolae, as well as the re-
cently described A.azurea R.H.Webb & G.D.Starr.
Agave sect. Campaniflorae (Trel.) R.H.Webb &
G.D.Starr, Cact. Succ. J. (US) 87(2): 36. 2015.
Type (lectotype): A. aurea Brandegee (Fig-
ure10).
This new section in A.subg. Agave provides a
valid name for A.“group” Campaniflorae Trelease
(1912: 44) at sectional rank.
Agave sect. Deserticolae (Trel.) R.H.Webb &
G.D.Starr, Cact. Succ. J. (US) 87(2): 36. 2015.
Type (lectotype): A. deserti Engelm. (Fig-
ure13).
This new section in A.subg. Agave provides a
valid name for A.“group” Deserticolae Trelease
(1912: 44–45) at sectional rank.
Agave sect. Rigidae (Baker) R.H.Webb &
G.D.Starr, Cact. Succ. J. (US) 87(2): 37. 2015.
Type: A.rigida Mill. (accepted name: A.an-
gustifolia Haw.).
This new section in A.subg. Agave provides a
valid name for A.“group” Rigidae Baker (1877 [7]:
171, [8]: 264) at sectional rank.
Agave sect. Umbelliflorae (Trel.) R.H.Webb &
G.D.Starr, Cact. Succ. J. (US) 87(2): 38. 2015.
Type (lectotype): Agave shawii Engelm. (Fig-
ure11).
This new section in A.subg. Agave provides a
valid name for A.“group” Umbelliflorae Trelease
(1912: 44) at sectional rank.
3) Updated infrageneric classification of
Agave (s.l.)
Numbers and letters given in square brackets
(e.g., “[1]” and “[1a]”, etc.) are the identifiers for
the individual taxa, as used by Thiede (2001), and
revised and updated in Thiede (2019). Accepted
names are in bold font.
Agave L., Spec. Pl. [ed. 1], 323. 1753. Type:
A. americana L. (Figure 21) (lectotype, desig-
nated by Britton & P. Wilson, Sci. Survey
Puerto Rico, 5: 156. 1923 (fide Index Nominum
Genericorum [ING])).
= Polianthes L., Sp. Pl.: 316. 1753.
= Pothos Adanson, Fam. Pl. 2: 57. 1763 nom. illeg.,
Art. 53.1.
= Tuberosa Heist. ex Fabr., Enum.: 2. 1759 nom.
illeg., Art. 52.1.
= Bonapartea Willd., Enum. Pl. Suppl. [Willde-
now] 18. 1814 nom. illeg., Art. 53.1.
= Littaea Tagl., Bibliot. Ital. (Milan) 1: 106. 1816.
= Bravoa Lex. in P.de La Llave & J.M.de Lexarza,
Nov. Veg. Descr. 1: 6. 1824.
= Coetocapnia Link & Otto, Icon. Pl. Rar.: 35.
1828.
= Robynsia Drapiez, Hort. Universel 2: 127. 1841
nomen rejiciendum, Art. 56.1.
= Ghiesbreghtia Roezl, Gartenflora 10: 122. 1861
nom. illeg., Art. 53.1.
= Manfreda Salisb., Gen. Pl.: 78. 1866.
= Allibertia Marion, Rev. Hort. Bouch. Rhone,
Nov. 1882. nom. illeg., Art. 53.1.
= Prochnyanthes S.Watson, Proc. Amer. Acad.
Arts 33: 457. 1887.
= Leichtlinia H.Ross, Index Seminum (PAL,
Panormitani) 1893: 48. 1893.
= Delpinoa H.Ross, Boll. Reale Orto Bot. Palermo
1: 116. 1897.
= Pseudobravoa Rose, Contr. U.S. Natl. Herb. 5:
155. 1899.
= Runyonia Rose, Addisonia 7: 39. 1922.
= ×Mangave D.Klein, Kakt. and. Sukk. 61: 212.
2010.
= ×Polifreda W.Ritchie & J.T.Lindstr., J. Envi-
ronm. Hort. 32: 52. 2014.
[1] Agave subg. Littaea (Tagl.) Baker, Handb.
Amaryll. 164. 1888. Type: Littaea geminiflora
Tagl. (≡ A.geminiflora (Tagl.) Ker Gawl.) (Fig-
ure6).
[1a] Agave sect. Juncineae Salm-Dyck, Wo-
chenschr. Vereines Beförd. Gartenbaues Kö-
nigl. Preuss. Staaten 4(23): 182. [6. Jun.] 1861.
Type (lectotype): A.striata Zucc. (Figure7).
≡ A.[infragen. unranked, “group”] Striatae Baker,
Gard. Chron. ser. nov. 7: 171, 8: 556. 1877
Type: A. striata Zucc. (Figure 7) ≡ A. subsect.
254 Bradleya 37/2019

Filiferae (Baker) A.Terracc., Prim. Contr.
Monogr. Agave: 6–9, 15. 1885.
≡ A.sect. Schoenagave A.Berger, Agaven 78. 1915.
Type (lectotype): A.striata Zucc. (Figure7).
= A. [infragen. unranked, “Abtheilung”] Lorifor-
mes Jacobi, Hamburger Garten- Blumenzei-
tung 20: 502, 510–511. 1864 Type (not
indicated, lectotype, designated here):
A.dasylirioides Jacobi & C.D.Bouché.
= A.sect. Chonanthagave A.Berger, Agaven 83.
1915. Type (lectotype): A.dasylirioides Jacobi
& C.D.Bouché.
[1b] Agave sect. Inermes Salm-Dyck, Wochen-
schr. Vereines Beförd. Gartenbaues Königl.
Preuss. Staaten 4(23): 182. [6 Jun.] 1861. Type
(lectotype): A. attenuata Salm-Dyck (Figure
22).
≡A.sect. Anoplagave A.Berger, Agaven 121. 1915.
Type (lectotype): A.attenuata Salm-Dyck (Fig-
ure 22).
≡A.[infragen. unranked, “group”] Amolae Gen-
try, Agaves Cont. N. Amer.: 63. 1982 nom.
inval. Art 37.1. Type: A.attenuata Salm-Dyck
(Figure 22).
=A.[infragen. unranked, “Abtheilung”] Margine
integerrimae Jacobi, Hamburger Garten- Blu-
menzeitung 20: 501–502, 509–510. 1864 nom.
inval. Art. 21.2. Ex. 2 & 32.1.c. Type: A.glau-
cescens Hooker nom.illeg. Art. 53.1, sole ele-
ment included (accepted name: A.attenuata
Salm-Dyck).
=A.sect. Yuccifoliae (A.Terracc.) B.Ullrich, Pi-
ante Grasse 15(4): 29. 1995. Type: A.yuccifolia
F.Delaroche (accepted name: A.spicata Cav.) =
A. ser. Yuccifoliae A.Terracc., Prim. Contr.
Monogr. Agave: 14. 1885.
= A. [infragen. unranked, “group”] Serrulatae
Baker, Gard. Chron. ser. nov. 8: 748, 808.
1877. Type: A.pruinosa Lem. ex Jacobi, sole
element included (accepted name: A.attenuata
ssp. dentata (Roezl) B.Ullrich).
255
Bradleya 37/2019
Figure 15.
Agave parryi [subsp. parryi] var. parryi. Agave parryi is the type of A. sect. Parryanae [A. subg.
Agave]. Mexico, Chihuahua, Rosales Mpio., c. 19.5 miles (c. 31km) SE Chihuahua towards Delicias, 29 March
1992. Photograph: Joachim Thiede.

= A.sect. Nizandensae (B.Ullrich) B.Ullrich, Suc-
culenta (NL) 70(6): 144. 1991 [1991b]. Type:
A.nizandensis Cutak = A.ser. Nizandensae
B.Ullrich, Succulenta (NL) 70(4): 91. 1991
[1991a].
[1c] Agave sect. Choritepalae Hochstätter,
Agave I: 14. 2015. Type: A. ellemeetiana
K.Koch
≡ A.[infragen. unranked, “group”] Choritepalae
Gentry, Agaves Cont. N. Amer.: 89. 1982 nom.
inval. Art 37.1. Type: A.ellemeetiana K.Koch.
[1d] Agave sect. Littaea (Tagl.) Benth., Gen. Pl.
[Benth. & Hook.f.] 3(2): 733. 1883. Type: Lit-
taea geminiflora Tagl. (type of Littaea; ≡
A.geminiflora (Tagl.) Ker Gawl.) (Figure6).
≡ A.sect. Xysmagave A.Berger, Agaven 68. 1915.
Type (lectotype): A. geminiflora (Tagl.) Ker
Gawl.) (Figure6).
= A.[infragen. unranked, “group” Filiferae Baker,
Gard. Chron. ser. nov. 7: 171, 303. 1877. Type:
A. filifera Salm-Dyck ≡ A.subsect. Filiferae
(Baker) A.Terracc., Prim. Contr. Monogr.
Agave: 6–9, 17. 1885.
[1e] Agave sect. Parviflorae Hochstätter, Agave
I: 16. 2015. Type: A. parviflora Torr. (Fig-
ure8).
=A. [infragen. unranked, “group”] Parviflorae
Gentry, Agaves Cont. N. Amer.: 195. 1982
nom. inval. Art 37.1. Type: A.parviflora Torr.
(Figure8).
[1f] Agave sect. Micracanthae Salm-Dyck,
Wochenschr. Vereines Beförd. Gartenbaues
Konigl. Preuss. Staaten 4(23): 182. [6 Jun.]
1861. Type: A. micracantha Salm-Dyck (ac-
cepted name: A. mitis Mart.).
= A.[infragen. unranked, “Abtheilung”] Subcari-
natae Jacobi, Hamburger Garten- Blumenzei-
tung 20: 501, 509. 1864. Type (lectotype,
designated here): A.mitis Mart.
= A.[infragen. unranked, “Abtheilung”] Aloideae
Jacobi, Jahresber. Schles. Ges. Vaterl. Cult.
1867, 45: 71. 1868. Type (not indicated, lecto-
type, designated here): A. mitis Mart.
=A.sect. Anacamptagave A.Berger, Agaven 38–
39. 1915. Type (lectotype): A.mitis Mart.
= A.[infragen. unranked, “group”] Polycephalae
Gentry, Agaves Cont. N. Amer.: 216. 1982
nom. inval. Art 37.1. Type: A.celsii Hook. (ac-
cepted name: A.mitis Mart.).
[1g] Agave sect. Heteracanthae Salm-Dyck,
Wochenschr. Vereines Beförd. Gartenbaues
Königl. Preuss. Staaten 4(23): 182. [6 Jun.]
1861. Type: A. heteracantha Zucc. (accepted
name: A.lophantha Schiede ex Kunth) (Fig-
ure9).
= A. [infragen. unranked, “Abtheilung”] Mar-
ginatae Jacobi, Hamburger Garten- Blumen-
zeitung 20: 499, 504–506. 1864. Type
(lectotype, designated here): A.lophantha
Schiede ex Kunth (Figure9).
≡A.sect. Geminiflorae Engelm., Trans. Acad. Sci.
St. Louis 3: 296–297, 304. 1875. Type (lecto-
type): A.heteracantha Zucc. (accepted name:
A.lophantha Schiede ex Kunth) (Figure9).
= A. [infragen. unranked, “group”] Marginatae
Baker, Gard. Chron. ser. nov. 7: 171, 368.
1877. Type (lectotype): A.lophantha Schiede
ex Kunth (Figure9) ≡ A.subsect. Marginatae
(Baker) A.Terracc., Prim. Contr. Monogr.
Agave: 6–9, 30. 1885.
=A.sect. Pericamptagave A.Berger, Agaven 85.
1915. Type (lectotype): A.lophantha Schiede
ex Kunth (Figure9).
= A. sect. Brachysolenagave A.Berger, Agaven
116. 1915. Type (lectotype): A.ghiesbreghtii
Verschaff.
= A. sect. Bechtoldiae Hochstätter, Agave: I: 21.
2015. Type: A.victoriae-reginae T.Moore.
[1h] Agave sect. Urceolatae Hochstätter, Agave:
I: 18. 2015. Type: A.utahensis Engelm. (Fig-
ure1).
≡ A. [infragen. unranked, “group”] Urceolatae
Gentry, Agaves Cont. N. Amer.: 251. 1982
nom. inval. Art 37.1. Type: A.utahensis En-
gelm. (Figure1).
[2] Agave subg. Agave Type: A. americana L.
(Figure 21)
Continental North America
[2a] Agave sect. Agave. Type: A.americana L.
≡A.sect. Macracanthae Salm-Dyck, Wochenschr.
Vereines Beförd. Gartenbaues Königl. Preuss.
Staaten 4(23): 186. [6 Jun.] 1861 nom.inval.,
Art. 22.1. & Ex. 2. Type: A.americana L. (Fig-
ure 21).
≡A.[infragen. unranked, “Abtheilung”] Carnosae
Jacobi, Hamburger Garten- Blumenzeitung
20: 499, 506. 1864 nom.inval. Art. 22.1. & Ex.
2. Type: A.americana L. (Figure 21).
≡A.sect. Paniculatae Engelm., Trans. Acad. Sci.
St. Louis 3: 297, 309. 1875 nom.inval. Art.
22.1. & Ex. 2. Type: A.americana L. (Figure
21).
≡A. [infragen. unranked, “group”] Americanae
Baker, Gard. Chron. ser. nov. 7: 171, 200. 1877
nom.inval., Art. 22.1. & Ex. 2.Type: A.amer-
icana L. (Figure 21) ≡ A.subsect. Americanae
(Baker) A.Terracc., Prim. Contr. Monogr.
Agave: 6–9, 36. 1885 nom.inval., Art. 22.1. &
Ex. 2.
256 Bradleya 37/2019

≡A. [infragen. unranked, “Reihe”] Americanae
A.Berger, Agaven 148. 1915 nom.inval., Art.
22.1. & Ex. 2. Type: A.americana L. (Figure
21).
=A.[infragen. unranked, “Abtheilung”] Subco-
riaceae Jacobi, Hamburger Garten- Blumen-
zeitung 20: 501, 508–509. 1864. Type
(lectotype, designated here): A.vera-cruz
Mill. (accepted name: A.lurida Aiton).
[2b] Agave sect. Salmianae (A.Berger) Verloove
& Thiede, in Verloove et al., submitted. Type:
A.salmiana Otto ex Salm-Dyck ≡ A.[infragen.
unranked, “Reihe”] Salmianae A.Berger,
Agaven 128. 1915.
≡A. [infragen. unranked, “group”] Salmianae
Gentry, Agaves Cont. N. Amer.: 594. 1982
nom. inval. Art 37.1. Type: A. salmiana Otto
ex Salm-Dyck.
[2c] Agave sect. Sisalanae Thiede & Gideon
F.Sm. nom. nov. pro A. [infragen. unranked,
“Unterreihe” Sisalanae] A.Berger, Agaven
230. 1915 nom. illeg. Art. 53.3. (replaced syn-
onym). Type: A. sisalana Perrine (not indi-
cated, according to Art. 10.8. & Ex. 12 (Tur-
land et al., 2018)).
[2d] Agave sect. Crenatae (A.Berger) B.Ull-
rich, Piante Grasse 13(2): 53. 1993. Type:
A. crenata Jacobi (accepted name: A. i nae-
quidens K.Koch) ≡ A. [infragen. unranked,
“Unterreihe”] Crenatae A.Berger, Agaven
194. 1915.
=A.[infragen. unranked, “group”] Crenatae Gen-
try, Agaves Cont. N. Amer.: 251. 1982 nom.
illeg. Art. 53.3 & nom. inval. Art 37.1. Type:
A.inaequidens K.Koch.
[2e] Agave sect. Campaniflorae (Trel.)
R.H.Webb & G.D.Starr, Cact. Succ. J. (US)
87(2): 36. 2015. Type (lectotype): A. aurea
Brandegee (Figure10) ≡ A. [infragen. unran-
ked, “group”] Campaniflorae Trel., Annual
Rep. Missouri Bot. Gard. 22: 44. 1912 (dated
1911) ≡ A.sect. Campaniflorae (Trel.) R.H.
Webb & G.D. Starr, Haseltonia 20: 68. 2015
nom.inval., Art. 41.1.
257
Bradleya 37/2019
Figure 16. Agave hurteri is the type of A. sect. Guatemalenses [A. subg. Agave]. Cultivated, north-cen-
tral South Africa, 2 April 2015. Photograph: Gideon F. Smith.

≡ A.[infragen. unranked, “Reihe”] Campaniflorae
A.Berger, Agaven 168. 1915. Type (not indica-
ted, lectotype, designated here): A.aurea
Brandegee (Figure10).
[2f] Agave sect. Umbelliflorae (Trel.) R.H.Webb
& G.D.Starr, Cact. Succ. J. (US) 87(2): 86.
2015. Type (lectotype): A.shawii Engelm. (Fig-
ure11) ≡ A.[infragen. unranked, “group”] Um-
belliflorae Trel., Annual Rep. Missouri Bot.
Gard. 22: 44. 1912 (dated 1911) ≡ A.sect. Um-
belliflorae (Trel.) R.H.Webb & G.D.Starr,
Haseltonia 20: 68. 2015 nom.inval., Art. 41.1.
≡A.[infragen. unranked, “Reihe”] Umbelliflorae
A.Berger, Agaven 170. 1915. Type (not indica-
ted, lectotype, designated here): A.shawii
Engelm. (Figure11).
[2g] Agave sect. Rigidae (Baker) R.H.Webb &
G.D.Starr, Cact. Succ. J. (US) 87(2): 85. 2015.
Type: A.rigida Mill. (accepted name: A.an-
gustifolia Haw.) ≡ A. [infragen. unranked,
“group”] Rigidae Baker, Gard. Chron. ser. nov.
7: 171, 8: 264. 1877 ≡ A. subsect. Rigidae
(Baker) A.Terracc., Prim. Contr. Monogr.
Agave: 6–9, 42. 1885 ≡ A.sect. Rigidae (Baker)
R.H.Webb & G.D.Starr, Haseltonia 20: 90.
2015 nom.inval., Art. 41.1.
= A.[infragen. unranked, “Abtheilung”] Canalic-
ulatae Jacobi, Hamburger Garten- Blumen-
zeitung 20: 502, 510. 1864. Type (not
indicated, lectotype, designated here):
A. rubescens Salm-Dyck (accepted name:
A. angustifolia var. rubescens (Salm-Dyck)
Gentry).
= A. [infragen. unranked, “group”] Sisalanae Trel.,
Mem. Natl. Acad. Sci. 11: 8, 9, 16, 47. 1913.
Type: A. angustifolia Haw.
≡ A. [infragen. unranked, “Reihe”] Rigidae
A.Berger, Agaven 226. 1915. Type: Not indica-
ted, A.rigida Mill. according to Art. 10.8. &
Ex. 12 (Turland et al., 2018) (accepted name:
A.angustifolia Haw.).
= A. [infragen. unranked, “group”] Tequilanae
Trel., Trans. Acad. Sci. St. Louis 23(3): 134.
1915. Type: Not indicated, A. tequilana
F.A.C.Weber according to Art. 10.8. & Ex. 12
(Turland et al., 2018).
[2h] Agave sect. Ditepalae Hochstätter, Agave
I: 26. 2015. Type: A. shrevei Gentry
(Figure12).
≡ A.[infragen. unranked, “group”] Ditepalae Gen-
try, Agaves Cont. N. Amer.: 416. 1982 nom.
inval. Art 37.1. Type: A.shrevei Gentry (Fig-
ure12).
= A. [infragen. unranked, “group”] Applanatae
Trel., Annual Rep. Missouri Bot. Gard. 22: 85–
86. 1912 (dated 1911). Type: Not indicated,
A.applanata Hort. Tonel ex K.Koch according
to Art. 10.8. & Ex. 12.
= A.sect. Hodgsoniae Hochstätter, Agave I: 30.
2015. Type: A.murpheyi F.Gibson.
[2i] Agave sect. Deserticolae (Trel.) R.H.Webb
& G.D.Starr, Cact. Succ. J. (US) 87(2): 84.
2015. Type (lectotype): A.deserti Engelm. (Fig-
ure13) ≡ A.[infragen. unranked, “group”] De-
serticolae Trel., Annual Rep. Missouri Bot.
Gard. 22: 44–45. 1912 (dated 1911). ≡ A.sect.
Deserticolae (Trel.) R.H.Webb & G.D.Starr,
Haseltonia 20: 76. 2015 nom.inval., Art. 41.1.
≡A. [infragen. unranked, “Reihe”] Deserticolae
A.Berger, Agaven 260–261. 1915. Type (not in-
dicated, lectotype, designated here): A.de-
serti Engelm. (Figure13).
[2j] Agave sect. Conicae R.H.Webb & G.D.Starr,
Haseltonia 20: 71. 2015. Type: A.avellanidens
Trel. (Figure14).
[2k] Agave sect. Intermediae R.H.Webb &
G.D.Starr, Haseltonia 20: 87. 2015. Type:
A.vizcainoensis Gentry.
258 Bradleya 37/2019
Figure 17. Inflorescence of Agave hurteri.
Photograph: Gideon F. Smith.

[2l] Agave sect. Parryanae Hochstätter, Agave
I: 29. 2015. Type: A. parryi Engelm.
(Figure15).
≡ A. [infragen. unranked, “group”] Parryanae
Gentry, Agaves Cont. N. Amer.: 520. 1982
nom. inval. Art. 37.1. Type: A.parryi Engelm.
(Figure15).
[2m] Agave sect. Hibernicae Hochstätter, Agave
I: 31. 2015. Type: A.gentryi Ullrich.
= A. [infragen. unranked, “group”] Parrasanae
Alsemgeest & Roosbroeck, privately published
on the web. c. 2012? nom.inval., Art. 29.1.
Type: not indicated, A. parrasana A.Berger ac-
cording to Art. 10.8. & Ex. 12 (Turland et al.,
2018).
[2n] Agave sect. Marmoratae (A.Berger) Thiede
& Gideon F.Sm. comb. & stat. nov. Type:
A.marmorata Roezl ≡ A.[infragen. unranked,
“Reihe”] Marmoratae A.Berger, Agaven 269.
1915 (basionym).
≡A.[infragen. unranked, “group”] Marmoratae
Gentry, Agaves Cont. N. Amer.: 507. 1982
nom. inval. Art. 37.1. Type: A. marmorata
Roezl.
[2o] Agave sect. Guatemalenses (Trel.) B.Ull-
rich, Kakt. and. Sukk. 43(9): 215. 1992. Type
(lectotype): A.hurteri Trel. (Figures16 & 17) ≡
A. [infragen. unranked, “group”]
Guatemalenses Trel., Trans. Acad. Sci. St.
Louis 23(3): 134. 1915.
= A.[infragen. unranked, “Reihe”] Scolymoides
A.Berger, Agaven 181. 1915. Type (not indica-
ted, lectotype, designated here): A.potato-
rum Zucc.
259
Bradleya 37/2019
Figure 18. Agave virginica is the type of A. subg.
Manfreda and of A. ser. Virginicae [A. subg. Man-
freda]. From N.J. von Jacquin, Icones plantarum
rariorum vol. 2: t. 378. 1786–1793.
(Courtesy of The Biodiversity Heritage Library).
Figure 19. Agave amica is the accepted name of
the type, Polianthes tuberosa, of A. sect. Po-
lianthes and of A. ser. Polianthes [A. subg. Man-
freda]. From Botanical Register vol. 1: t. 63. 1815.
(Courtesy of The Biodiversity Heritage Library).

= A. [infragen. unranked, “group”] Hiemiflorae
Gentry, Agaves Cont. N. Amer.: 465. 1982
nom. inval. Art. 37.1.: Type: A. hiemiflora Gen-
try.
= A. sect. Hiemiflorae Hochstätter, Agave I: 34.
2015. Type: A. hiemiflora Gentry.
= A. [infragen. unranked, “group”] Atrovirentes
Trel., Trans. Acad. Sci. St. Louis 23(3): 134.
1915. Type: Not indicated, A.atrovirens Karw.
ex Salm-Dyck according to Art. 10.8. & Ex. 12.
Caribbean region
[2p] Agave sect. Antillanae (Trel.) Thiede &
Gideon F.Sm. comb. & stat. nov. Type: A.an-
tillarum Descourt. ≡ A.[infragen. unranked,
“group”] Antillanae Trel., Mem. Natl. Acad.
Sci. 11: 9, 16, 29. 1913 (basionym).
≡A. [infragen. unranked, “Reihe”] Antillanae
A.Berger, Agaven 205. 1915. Type (not indica-
ted, lectotype, designated here): A.antil-
larum Descourt.
[2q] Agave sect. Antillares (Trel.) Thiede &
Gideon F.Sm. comb. & stat. nov. Type:
A. willdingii Tod. ≡ A.[infragen. unranked,
“group”] Antillares Trel., Mem. Natl. Acad. Sci.
11: 9, 16, 42. 1913 (basionym).
≡ A. [infragen. unranked, “Reihe”] Antillares
A.Berger, Agaven 271–272. 1915. Type (not in-
dicated, lectotype, designated here):
A.willdingii Tod.
[2r] Agave sect. Bahamanae (Trel.) Thiede &
Gideon F.Sm. comb. & stat. nov. Type: A.ba-
hamana Trel. ≡ A. [infragen. unranked,
“group”] Bahamanae Trel., Mem. Natl. Acad.
Sci. 11: 9, 16, 38. 1913 (basionym).
≡ A. [infragen. unranked, “Reihe”] Bahamanae
A.Berger, Agaven 202. 1915. Type: Not indica-
ted. A.bahamana Trel. according to Art. 10.8.
& Ex. 12.
[2s] Agave sect. Caribaeae (Trel.) Thiede &
Gideon F.Sm. comb. & stat. nov. Type: A.
karatto Mill. ≡ A. [infragen. unranked,
“group”] Caribaeae Trel., Mem. Natl. Acad.
Sci. 11: 9, 16, 21. 1913 (basionym).
= A. [infragen. unranked, “Reihe”] Caribaeae
A.Berger, Agaven 213. 1915. Type: Not indi-
cated. A.caribaea Baker nom.illeg. Art. 53.1,
according to Art. 10.8. & Ex. 12.(accepted name:
A. karatto Mill.).
[2t] Agave sect. Inaguenses (Trel.) Hochstätter,
Agave I: 40. 2015. Type: A. nashii Trel. ≡ A.
[infragen. unranked, “group“] Inaguenses
Trel., Mem. Natl. Acad. Sci. 11: 9, 16, 45. 1913.
= A. [infragen. unranked, “Reihe”] Inaguenses
A.Berger, Agaven 268–269. 1915. Type: Not
indicated. A.inaguensis Trel. according to Art.
10.8. & Ex. 12. (Accepted name: A. karatto
Mill.).
[2u] A.sect. Viviparae (Baker) Verloove &
Thiede submitted. Type: Agave vivipara L. ≡
A. [infragen. unranked, “group”] Viviparae
Baker, Gard. Chron. ser. nov. 8: 780, 808. 1877
(basionym) ≡ A. subsect. Viviparae (Baker)
A.Terracc., Prim. Contr. Monogr. Agave: 6–9.
1885.
= A.[infragen. unranked, “group”] Vicinae Thiede
nom. inval. Art 37.1., Ill. Handb. Succ. Pl.,
Monocotyledons: 11. 2001. Type: A. vicina
Trel.
[2v] Agave sect. Columbianae (A.Berger)
Thiede & Gideon F.Sm. comb. & stat. nov.
Type (not indicated, lectotype, here desig-
260 Bradleya 37/2019
Figure 20. Agave coetocapnia is the accepted
name of the type, Bravoa geminiflora, of A. ser.
Bravoa [A. subg. Manfreda]. From Curtis’s Botan-
ical Magazine vol. 79 [ser. 3, vol. 9]: t. 4741. 1853.
(Courtesy of The Biodiversity Heritage Library).

nated): A. cundinamarcensis A.Berger ≡
A.[infragen. unranked, “Reihe”] Columbianae
A.Berger, Agaven 222. 1915 (basionym).
[3] Agave subg. Manfreda (Salisbury) Baker,
Handb. Amaryll. 164. 1888. Type: A.virginica
L. (Figure18) ≡ Manfreda Salisb., Gen. Pl.: 78.
1866.
= Polianthes L., Sp. Pl.: 316. 1753.
= Bravoa Lex. in P.de La Llave & J.M.de Lexarza,
Nov. Veg. Descr. 1: 6. 1824.
= Allibertia Marion ex Baker, Gard. Chron. ser.
nov. 19: 176. 1883.
= Delpinoa H.Ross, Boll. Reale Orto Bot. Palermo
1: 116. 1898.
= Prochnyanthes S.Watson, Proc. Amer. Acad.
Arts 33: 457. 1887.
= Leichtlinia H.Ross, Index Seminum (PAL,
Panormitani) 1893: 48. 1893.
= Pseudobravoa Rose, Contr. U.S. Natl. Herb. 5:
155. 1899.
= Runyonia Rose, Addisonia 7: 39. 1922.
[3a] Agave sect. Herbaceae Salm-Dyck,
Wochenschr. Vereines Beförd. Gartenbaues
Königl. Preuss. Staaten 4(23): 182. [6 Jun.]
1861. Type (lectotype): A.brachystachys Cav.
(accepted name: A.scabra Ortega).
= A.sect. Singuliflorae Engelm., Trans. Acad. Sci.
St. Louis 3: 296, 300–301. 1875. Type (lecto-
type): A.virginica L. (Figure18).
≡ A. sect. Manfreda (Salisb.) Benth., Gen. Pl.
[Benth. & Hook.f.] 3(2): 733. 1883. Type: A.vir-
ginica L. (Figure18) ≡ A.[infragen. unranked,
“group”] Manfreda Thiede nom. inval. Art
261
Bradleya 37/2019
Figure 21. Agave americana is the type of the
genus Agave, the type of A. subg. Agave, and the
type of A. sect. Agave. Of all agaves, A. americana
subsp. americana is arguably the most widely nat-
uralised one. Here it grows in the Algarve in
southern Portugal at Ponta da Piedade, near
Lagos, 16 September 2013.
Photograph: Gideon F. Smith.
Figure 22. Although not widely naturalised,
Agave attenuata is arguably the most widely cul-
tivated Agave species in amenity and domestic
gardens globally. Here it grows in the Royal
Botanic Garden Sydney, New South Wales, Aus-
tralia, 2001. Photograph: Gideon F. Smith.

37.1., Ill. Handb. Succ. Pl., Monocotyledons:
11. 2001.
= Manfreda subg. Eumanfreda Rose, Contr. U.S.
Natl. Herb. 5: 153. 1899 nom.inval. Art. 21.3.
Type: A.virginica L. (Figure18).
= Manfreda subg. Pseudomanfreda Rose, Contr.
U.S. Natl. Herb. 5: 153. 1899. Type: Not indi-
cated.
[3a1] Agave ser. Brunneae (Hochstätter) Thiede
& Gideon F.Sm. comb. nov. Type: Manfreda
brunnea (S.Watson) Rose (≡ A. brunnea S.Wat-
son) ≡ Manfreda ser. Brunneae (as ‘Brun-
neaea’) Hochstätter, Manfreda Polianthes
Prochnyanthes: I: 9. 2016 (basionym) ≡ A. [in-
fragen. unranked, “Subgroup”] A. brunnea
Thiede nom. inval. Art 37.1., Handb. Succ. Pl.,
Monocotyledons: 11. 2001 ≡ Manfreda sect.
Brunneae (as ‘Brunneaea’) Hochstätter, Man-
freda Polianthes Prochnyanthes: I: 9. 2016.
[3a2] Agave ser. Scabraea (Hochstätter) Thiede
& Gideon F.Sm. comb. nov. Type: Agave
scabra Ortega ≡ Manfreda ser. Scabraea
Hochstätter, Manfreda Polianthes Prochnyan-
thes: I: 10. 2016 (basionym) ≡ A. [infragen. un-
ranked, “Subgroup”] A. scabra Thiede nom.
inval. Art 37.1., Ill. Handb. Succ. Pl., Mono-
cotyledons: 11. 2001 ≡ A. sect. Scabraea
Hochstätter, Manfreda Polianthes Prochnyan-
thes: I: 10. 2016.
[3a3] Agave ser. Guttatae (Hochstätter) Thiede
& Gideon F.Sm. comb. nov. Type: A. guttata
Jacobi & C.D.Bouché ≡ Manfreda ser. Guttatae
(as ‘Guttataea’) Hochstätter, Manfreda Po-
lianthes Prochnyanthes: I: 13. 2016 (ba-
sionym) ≡ A.[infragen. unranked, “Subgroup”]
A. guttata Thiede nom. inval. Art 37.1., Ill.
Handb. Succ. Pl., Monocotyledons: 13. 2001 ≡
Manfreda sect. Guttatae (as ‘Guttataea’)
Hochstätter, Manfreda Polianthes Prochnyan-
thes: I: 13. 2016.
[3a4] Agave ser. Virginicae (Hochstätter)
Thiede & Gideon F.Sm. comb. nov. Type: A.
virginica L. (Figure18) ≡ Manfreda ser. Vir-
ginicae (as ‘Virginicaea’) Hochstätter, Man-
freda Polianthes Prochnyanthes: I: 14. 2016
(basionym) ≡ A. [infragen. unranked, “Sub-
group”] A. virginica Thiede nom. inval. Art
37.1., Ill. Handb. Succ. Pl., Monocotyledons:
11. 2001. ≡ A.sect. Virginicae (as ‘Virginicaea’)
Hochstätter, Manfreda Polianthes Prochnyan-
thes: I: 14. 2016.
[3a5] Agave ser. Yucatanae Thiede & Gideon
F.Sm. ser. nov. Type: A.paniculata (L.Hern.,
R.A.Orellana & Carnevali) Thiede. Rosettes
30–100 cm in diameter; leaves with terminal
spine; inflorescences panicles or less often
racemes (A. petskinil (R.A.Orellana L.Hern. &
Carnevali) Thiede p.p.), with 1–5 branches;
flowers pedicellate, solitary, subtended by a
single bracteole. Distribution: Yucatán Penin-
sula (Mexico).
[3b] Agave sect. Polianthes (L.) Thiede &
Gideon F.Sm. comb. & stat. nov. Type: Po-
lianthes tuberosa L. (≡ A. amica (Medikus)
Thiede & Govaerts) (Figure19) ≡ Polianthes
L., Sp. Pl.: 316. 1753 (basionym).
[3b1] Agave ser. Polianthes (L.) Thiede Gideon
F.Sm. comb. & stat. nov. Type: Polianthes
tuberosa L. (≡ A. amica (Medikus) Thiede &
Govaerts) (Figure19) ≡ Polianthes L., Sp. Pl.:
316. 1753 (basionym) ≡ A. [infragen. un-
ranked, “group”] Polianthes Thiede nom. inval.
Art 37.1., Ill. Handb. Succ. Pl., Monocotyle-
dons: 11. 2001
[3b2] Agave ser. Bravoa (Lex.) Thiede & Gideon
F.Sm. comb. & stat. nov. Type (sole element
included): Bravoa geminiflora Lex. (≡ A. coeto-
capnia (M. Roemer) Govaerts & Thiede) (Fig-
ure 20) ≡ Bravoa Lex. in P.de La Llave &
J.M.de Lexarza, Nov. Veg. Descr. 1: 6. 1824 ≡
Polianthes subg. Bravoa (Lex.) M.Roemer,
Fam. Nat. Syn. Monogr. 4: 245. 1847.≡ A.[in-
fragen. unranked, “group”] Bravoa Thiede
nom. inval. Art 37.1., Ill. Handb. Succ. Pl.,
Monocotyledons: 11. 2001.
= Prochnyanthes S.Watson, Proc. Amer. Acad.
Arts 33: 457. 1887. Type: Prochnyanthes viri-
descens S. Watson.
Transfers from Manfreda and Polianthes
into Agave
Two species that were recently described in
Manfreda, and one that was described in Po-
lianthes, are here transferred to Agave with new
combinations, as follows:
Agave occidentalis (Art.Castro & Aarón Rodr.)
Thiede & Gideon F.Sm., comb. nov.
Basionym: Manfreda occidentalis Art. Castro
& Aarón Rodr., in Castro-Castro et al. 64(–65).
2017. Type: A. Rodríguez et al. 5924 (holotype
IBUG, isotypes CIIDIR, SLPM, IEB, MEXU).
Agave santanamichelii (Art.Castro, Aarón
Rodr. & P.Carrillo) Thiede & Gideon F.Sm.,
comb. nov.
Basionym: Manfreda santana-michelii
Art.Castro, Aarón Rodr. & P.Carrillo, in Castro-
Castro et al. 497(–502). 2018. Type: P. Carrillo-
Reyes & G.I. Tinoco-Villa 8160 (holotype IBUG,
isotypes CIIDIR, MEXU).
262 Bradleya 37/2019

Agave venustuliflora (E.Solano, García-Mend.
& R.Ríos-Gómez) Thiede & Gideon F.Sm.,
comb. nov.
Basionym: Polianthes venustuliflora E.Solano,
García-Mend. & R.Ríos-Gómez, in Solano et al.
[5]–([8]) 2019. Type: E. Solano y R. Ríos Gómez
1793 (holotype MEXU, isotypes CHAPA, FEZA,
IEB, MO, UAMIZ).
The page reference cited refers to the PDF doc-
ument of Acta Botanica Mexicana 126: e1441.
published online.
Acknowledgement
The authors are much indebted to Graham
Charles (UK) for his editorial work.
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