Phytotaxa 163 (3): 149–165
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Copyright © 2014 Magnolia Press
ISSN 1179-3155 (print edition)
Article
PHYTOTAXA
ISSN 1179-3163 (online edition)
http://dx.doi.org/10.11646/phytotaxa.163.3.2
Discovery of Hyptis pseudolantana in Jalisco and Michoacán, and description of
H. cualensis and H. macvaughii (Ocimeae, Lamiaceae), two new species from
western Mexico
JESÚS GUADALUPE GONZÁLEZ-GALLEGOS1, ARTURO CASTRO-CASTRO1, ALEJANDRA FLORESARGÜELLES1 & ARIOSTO RAFAEL ROMERO-GUZMÁN2
1
Herbario Luz María Villarreal de Puga (IBUG), Instituto de Botánica, Departamento de Botánica y Zoología, Universidad de
Guadalajara-CUCBA, km 15.5 carretera Guadalajara-Nogales, Las Agujas, Nextipac, Zapopan, CP 45110, Jalisco, México.
E-mails: xanergo@hotmail.com; arca68@hotmail.com
2
Desarrolladora de Proyectos y Edificaciones Ecoturísticas S.A. de C.V.
Abstract
The distribution of Hyptis pseudolantana is documented in the states of Jalisco and Michoacán, species previously
reported exclusively from the state of Guerrero, Mexico. An expanded description, distribution map and photographs are
provided for it. Hyptis cualensis and H. macvaughii are also described and illustrated by photographs, as two new species
from the Mexican states of Jalisco and Nayarit, respectively. Also, an identification key to species of tribe Hyptidinae in
western Mexico is provided.
Resumen
La distribución de Hyptis pseudolantana es documentada en los estados de Jalisco y Michoacán, especie registrada con
anterioridad únicamente del estado de Guerrero, México. Se proporciona una descripción ampliada, mapa de distribución
y fotografías de la misma. Hyptis cualensis e H. macvaughii son también descritas e ilustradas mediante fotografías,
como dos especies nuevas de los estados mexicanos de Jalisco y Nayarit, respectivamente. Además, se incluye una clave
para la identificación de las especies de la tribu Hyptidinae encontradas en el occidente de México.
Key words: Hyptis section Rhytidea, Hyptis subsection Mutabiles, Hyptis subsection Umbellatae, subtribe Hyptidinea
Introduction
The genus Hyptis Jacquin (1786: 101) is one of the richest Lamiaceae genera with almost 300 species. Hyptis
grows mainly in the New World with few species extending as weeds to the Old World (Epling 1936, Harley 1988,
Li & Hedge 1994, Harley et al. 2004, Pastore et al. 2011). The subtribe Hyptidinae (in tribe Ocimeae), which
includes Hyptis, can be recognized—excluding Asterohyptis Epling (1933a: 17)—because of the downwardly
directed stamens, and the thickened lower corolla lobe that acts as a hood, protecting them before anthesis, and
releasing them explosively when mature (Harley et al. 2004). However, the distinction of the genera embedded in
the subtribe is not entirely unambiguous (El-Gazzar & Rabei 2008). As stated in the keys provided by Harley
(1988) and Harley et al. (2004), the delimitation of the genera is achieved by means of a set of characters: structure
of the inflorescences, calyx bilabiate or not, appendiculate, spiny or simple calyx lobes, these reflexed or not,
stylopodium present or not, mericarp shape, amongst others. In Hyptis, the diagnostic character is the absence of a
stylopodium, even when the species with involucrate capitula might present it eventually.
Besides, phylogenetic analysis based on DNA sequences (Pastore et al. 2011) have brought to light Hyptis as a
polyphyletic genus since all other Hyptidinae genera appeared nested within Hyptis lineages in the cladograms.
Also, the polyphyly of the genus had been previously suggested by the phenetic analysis of morphological
characters (El-Gazzar & Rabei 2008). Given this complexity, two solutions may be taken, one is to reduce all other
Accepted by Zhi-Qiang Zhang: 10 Mar. 2014; published: 26 Mar. 2014
149
Hyptidinae genera under Hyptis, otherwise it should be recognized a much larger number of genera. In this context,
Pastore et al. (2011) claimed for the segregation of new genera to build a monophyletic delimitation of Hyptis as a
better solution rather than the submersion of all Hyptidinae within only one genus. Harley & Pastore (2012) did not
consider a good choice leaving a single genus with a broad range of morphological variation, purportedly
unmatched in any other genus of flowering plants. So they decided to integrate a new classification proposal for
Hyptidinae segregating new genera grounded mainly in the cladograms presented by Pastore et al. (2011).
Accordingly, they increased the number of genera within the subtribe from 8 (Harley et al. 2004) to 19, with the
coinage of six new generic names and status change of five infrageneric categories. Moreover, this approach
generates 142 new nomenclatural combinations.
However, little less than half of Hyptidinae species are represented in Pastore et al. (2011), and although the
clades are well supported in the topology shown, there are also several politomies in the cladograms that create
mistrust in the clade relationships in spite of their support. In addition, distinction of the genera they proposed is
ambiguous through the dichotomous key provided (Harley & Pastore 2012): characters are not precisely defined,
measurements are not always used in lieu of uncertain terms, the parallelism between two leads is inconsistent and
not clearly contrasting, several unqualified negative statements are done; these features are contrary to what is
recommended to construct an identification key (Judd et al. 2008). Moreover, morphological diversity is not
synonymous with taxonomic diversity, and a broad morphological variation is not unique of Hyptis s.l. For
example, in the large subtribe Euphorbiinae some authors decided to retain a single well-known and easily
identifiable genus, Euphorbia Linnaeus (1753: 450), instead of a vaguely circumscribed multitude of genera;
thereby Euphorbia becomes one of the most striking morphologically diversified genera of vascular plants, which
embraces perennial and annual herbs, erect and scandent shrubs, trees, geophytes and succulents, including
representatives of the three different kinds of carbon fixation C3, C4 and CAM, a high variability of ploidy levels,
but unified by the particular and conserved configuration of its inflorescences, the cyathia (Steinmann & Porter
2002, Horn et al. 2012, Yang et al. 2012, Dorsey et al. 2013, Riina et al. 2013). Another important consideration is
that a classification pursues conflicting goals, on one hand it tries to be a true picture of evolutionary history, and
on the other it is committed to be practical in order to be useful to general public and not being only an academic
whim; thus, an ethic position as a systematist should be to find a balance between both sides. The convenience of a
classification can be evaluated in terms that it implies easily diagnosable groups (i.e. well supported, preferably
with synapomorphic characters), and in second instance, few nomenclatural changes. Submerging all Hyptidinae
within Hyptis demands 32 nomenclatural changes almost 5 times less than the choice taken by Harley and Pastore
(2012). In summary, taking in consideration the uncertainty about phylogenetic results, the hardness in
morphologically delimiting Hyptidinae genera, we keep on recognizing provisionally the traditional Hyptis
definition instead of several new poorly defined genera. It is probable that the best solution in the near future will
be to submerge all Hyptidinae within Hyptis; though, more compelling evidence is needed to take a definitive
decision.
Once we have clarified our position in respect to the definition of Hyptis, we proceed to document the
occurrence of Hyptis pseudolantana Epling (1941: 555) in the states of Jalisco and Michoacán, and two new
species of Hyptis from western Mexico. We provide descriptions, pictures and distribution maps for the three taxa.
This is the result of recent field exploration and examination of specimens at several herbaria.
Hyptis pseudolantana Epling (1941: 555) (Figs. 1 and 2A–D)
Type:—MEXICO. Guerrero. Coyuca de Catalán: Aguazarca–Filo, District Mina, Guerrero, 9 November 1937 (fl, fr), G.B.
Hinton et al. 11266 (holotype UC, isotypes K!, MO!, US-00121889!, US-01014362!).
Perennial herb to subshrub, erect, 0.6–2(–2.5) m tall, arising from a small xylopodium, aromatic. Stems glabrous to
puberulent and pilose, with the hairs concentrated towards young branches. Leaves with petioles (1–)6–20 mm
long, pilose; blade ovate to ovate-elliptic, 1.5–7.1 × 0.9–4.1 cm, acute to acuminate at the apex, rounded to shortly
cuneate at the base, serrate margin, pilose in both surfaces. Inflorescences axillary or terminal, spiciform, 4–18 cm
long, with pectinate opposite cymes at each node, 6–18 flowers per cyme, peduncles 3–8.7 mm long (sometimes
hidden by the flowers); peduncle bracts conformed by reduced leaves 2.1–4.9 × (0.8–)1.5–2.2 mm, bracteoles
linear, 1.4–2.8 mm long; inflorescence axes, peduncles and bracteoles sparsely pilose and glandular-punctate.
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Flowers sessile or with pedicels less than 0.5 mm long. Calyx tube (1.7–)2.2–3.5(–5) mm long, and 1.6–1.8 mm
wide at the throat in flower (up to 4.4 × 2.2 mm in fruit), sparsely pilose and covered with some short glandularcapitate hairs and or sessile glandular dots, puberulent, internally with a ring of hairs attached 1–2 mm below the
line of the teeth or at the middle of calyx tube length; teeth triangular and attenuate towards the apex, 9–1.5 mm
long and straight in flower (up to 2 mm in fruit and backwardly oriented). Corolla pale violet, dull lavender to
white, sparsely pilose towards the lobes; tube (5.1–)6.5–7.3 mm long, (1.5–)2.3–2.8 mm wide at the throat, lobes
subequal, 1.7–3 mm long. Stamens 4, attached at 5.3–5.5 mm of corolla tube length, the posterior stamens attached
a little below than the anterior ones; filament (1.2–)1.5–1.6(–2.4) mm long, pilose; theca 0.2–0.4 mm long. Style
(3.3–)4.4–6.3 mm long, white and glabrous, without stylopodium; stigma bifurcate, the branches rounded at the
apex. Mericarp 4, ovoid to turbinate, 1–1.3 × 0.6–0.8 mm, slightly trigonous, light or dark brown, smooth to
muriculate, nitid, pulverulent, glabrous, with a markedly suture line in the external face.
Distribution, habitat and phenology:—Hyptis pseudolantana inhabits the western portion of the Sierra del
Halo, in the municipality of Tecalitlán, Jalisco, the eastern side of Sierra de Coalcomán, Michoacán, and
northeastern slopes of Sierra Madre del Sur in the municipality of Coyuca de Catalán, Guerrero (figure 1). It grows
in pine-oak and oak forests and in ecotones with montane cloud and tropical deciduous forests, at elevations from
(1400–)1650–2200 m. It shares habitat with Carpinus caroliniana Walter, Cosmos carvifolius Benth., Cunila
pycnantha B.L.Rob. & Greenm., Dendropanax arboreus (L.) Decne & Planch., Fuchsia arborescens Sims,
Galphimia glauca Cav., Inga micheliana Harms, Jungia pringlei Greenm, Pinus douglasiana Martínez, P.
hartwegii Lindl., Quercus scytophylla Liebm. It flowers and fructifies from August to May.
FIGURE 1. Distribution map of Hyptis pseudolantana (squares), H. cualensis (triangles), and H. macvaughii (dots) in Mexico.
Taxonomic relationships:—Hyptis pseudolantana was assigned by Epling (1941) to section Rhytidea Epling
(1933b: 80) due to the internal ring of hairs of the calyx placed at the middle of its length. However, this species is
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more similar to those from section Mesosphaeria Bentham (1833b: 122) subsection Pectinaria Epling (1933b: 97),
than to Hyptis rhytidea Bentham (1839: 21), the other member of section Rhytidea Epling (1933b: 80). It shares
with subsection Pectinaria the habit (herbs to subshrubs or rarely trees), petiolate leaves, ovate and rarely
lanceolate, pectinate or globose cymes with peduncles longer than the calyces and, linear floral bracts. It differs
from the species of section Rhytidea due to its membranaceous leaves (vs. coriaceous), cymes clearly pedunculated
(vs. subsessile), and linear floral bracts (vs. ovate). As stated in the introduction, the classification of the genus is
still unstable, so we have no elements to unequivocally support the inclusion of H. pseudolantana to one of the
infrageneric groups.
Hyptis pseudolantana is similar to H. pectinata (Linnaeus 1759: 1096) Poitier (1806: 474) and H. urticoides
Kunth (1817: 320) from subsection Pectinaria, but it can be easily distinguished from these because the calyx is
infundibuliform instead of tubular, with triangular teeth rather than linear, and calyx tube without an internal welldefined ring of hairs projecting between calyx teeth. It is also similar to Hyptis pinetorum from subsection
Mutabiles Epling (1933b: 102), but it can be differentiated by means of the glandular capitate hairs restricted to the
calyces and not spread throughout stems, leaves and inflorescences, usually shorter petioles [(2.5–)6–20 vs. (18–
)30–55 mm], flowers arranged in pectinate cymes instead of spherical cymes, shorter peduncles [3–7 vs. (8–)12–30
mm], shorter calyx tube (1.7–2.2 vs. 3–4.5 mm), and shorter style [(3.3–)4.4–6.3 vs. 6.2–7.7 mm] (table 1).
TABLE 1. Morphology, habitat and distribution comparison between Hyptis pseudolantana and similar species.
Hyptis pseudolantana
H. pinetorum
H. pectinata
H. urticoides
restricted to calyces
on the stems, leaves
(mainly on lower
surface), and
inflorescences (floral
axis and bracts, pedicels
and calyces)
absent
restricted to calyces
Petiole length
(mm)
(1–)6–20
(18–)30–55
(7.5–)10–30(–50)
5–20(–39)
Blade shape
ovate to ovate-elliptic
ovate to triangular-ovate
ovate to ovate-deltoid
lanceolate to ovatelanceolate
Blade size (cm)
(1.5–)1.6–7.1
× 0.9–4.1
(5–)6–10.5
× (3.2–)4–6.5(–9.8)
1.7–3.5(–8) × (1.3–)2–4.5
3–8 × (1.4–)2.5–5
flower arranged in
pectinate cymes, each
one with 3–8.7 mm
long peduncles
flowers arranged in
spherical cymes, each
one with (8–)12–30 mm
long peduncles
flowers arranged in
pectinate cymes, each one
with 1–2.5(–3.7) mm long
peduncles
flowers arranged in
pectinate to semispherical (fewflowered) cymes, each
one with 10–25 mm
long peduncles
Shape
linear
linear-lanceolate to
linear-elliptic
linear
linear
Length (mm)
1.4–2.8
1.9–4.3
1.9–3.1(–4)
1.8–3.2
0–0.5
0–0.5
< 0.1
0–0.1
Tube size (mm)
(1.7–)2.2–3.5(–5) ×
1.6–1.8
3–4.5 × 1.2–1.9
0.9–1.2 × 0.5–0.7
1.4–2.6 × 1.2–1.6
Teeth length (mm)
0.9–1.5
1–1.4
0.5–0.7
1.1–1.5
Distribution of
glandularcapitate hairs
Leaves
Inflorescence
Arrangement
Floral bract
Pedicel
Length (mm)
Calyx
...... continued on the next page
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TABLE 1. (Continued)
Hyptis pseudolantana
H. pinetorum
H. pectinata
H. urticoides
Teeth shape
triangular
triangular
linear
linear
Well-defined ring
of hairs projecting
between calyx
teeth
absent
absent
present
present
(5.1–)6.5–7.3
× (1.5–)2.3–2.8
5–6.5(–7.6) × 1.8–2.1
1.4–1.5 × 0.2–0.3
3.2–4.3 × 0.5–1
(1.2–)1.5–1.6(–2.4)
1.2–1.4
0.2–0.4
0.5–1
(3.3–)4.4–6.3
6.2–7.7
1.1–1.7
2.4–4.4
Size (mm)
1–1.3 × 0.6–0.8
1–1.2 × 0.5–0.7
0.6–0.9 × 0.2–0.3
1.1–1.4 × 0.6–0.8
Habitat
pine-oak and oak
forests, and ecotones
with tropical
deciduous forest
pine-oak and montane
cloud forests
tropical deciduous forests
and secondary vegetation of
pine-oak forests
tropical deciduous and
subdeciduous, oak
forests, secondary
vegetation of pine-oak
and montane cloud
forests
Altitudinal range
(m)
(1400–)1650 –2200
900–2000
(10–)800–1700
(50–)600–2100
Distribution
Mexico. Jalisco and
Michoacán
Mexico. Jalisco.
Widely distributed in
America from the United
States to Peru, Brazil and
the Antilles; introduced in
tropical Africa and Asia. In
Mexico almost in all the
states except for those from
Baja California Peninsula.
Widely distributed
from Mexico to
Panama. In Mexico
almost in all the states
except for the northern
ones and those from the
Yucatán Peninsula.
Corolla
Tube size (mm)
Androecium
Filament length
(mm)
Gynoecium
Style length (mm)
Mericarp
Hyptis pseudolantana was formerly known only from the type collection series from Guerrero, but here,
several specimens from Jalisco and one from Michoacán are assigned to this species (figure 1). Hyptis
pseudolantana is not reported for the latter two states in most recent floristic lists, although the specimen McVaugh
22775 (ENCB) was included as an undetermined Hyptis for Michoacán (Rodríguez-Jiménez & Espinosa-Garduño
1996, Ramírez-Delgadillo et al. 2010). Our collections and other herbaria material reviewed differ in some
characters from the original description of the species. Nonetheless, these differences are not so abrupt and
consistent as to demand the need of naming the new discovered populations as a different taxon. The
morphological discrepancies are: petioles 1–2 mm long in the original description vs. 6–20 mm long in the new
discovered populations, axillary inflorescences vs. most of them terminal, calyces internally ornate with a ring of
hairs attached at the middle of its length vs. attached 1–2 mm below calyx teeth line, and white corollas vs. pale
violet to dull lavender. It is likely that the relatively low number of specimens that were surveyed to elaborate the
original description by Epling (1941) promoted these discrepancies. Moreover, the distribution of the species is
composed of three disjunct populations around Balsas Depression; it can be expected that more extensive
exploration in mountains between these points and with adequate elevations for the species establishment will
reveal new populations. This ultimately could help to better understand the morphological variation of H.
pseudolantana.
If the classification proposal of Harley & Pastore (2012) is followed, H. pseudolantana would fit better within
their definition of the genus Mesosphaerum Browne (1756: 257).
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Conservation assessment:—Hyptis pseudolantana has a wide geographical distribution and their populations
consist of numerous individuals, so it appears not to be in danger of extinction.
Additional specimen examined:—MEXICO. Jalisco. Tecalitlán: Sierra del Halo, near a lumber road leaving
the Colima highway, 7 miles SSW of Tecalitlán and extending southeasterly toward San Isidro, 1400 m, 13 August
1957 (fl, fr), R. McVaugh 16149 (MICH!); Sierra del Halo, near a lumber road leaving the Colima highway, 7 miles
SSE of Tecalitlán and extending SE toward San Isidro, 2000–2200 m, 28 November 1959 (fl), W.N. Koelz 1171 &
R. McVaugh (MICH!); 46 km carr. Ciudad Guzmán-Pihuamo, por la brecha Llanitos-Canutillo, a 16 km, 1650 m,
14 May 1988 (fl, fr), Pichardo 47 (MEXU!); 14 km al E de Llanitos, brecha a Canutillo, 1750 m, 18 March 1990
(fl, fr), Villa et al. 673 (IBUG!, IEB!); brecha a Mexicanillo, Sierra del Halo, 2060 m, 13 March 1996 (fl, fr),
Ramírez et al. 3599 (IBUG!); Sierra del Halo, cañada La Jabalina, predio Las Palomas, 3.5 km en línea recta al O
de Alotitlán, 19º15’35”N, 103º14’58”W, 1700–1750 m, 23 February 2012 (fl, fr), A. Castro-Castro, E.A. SuarezMuro & A. Frías-Castro 2619a (IBUG!); Sierra del Halo, predio Los Fresnos, Puerto de Ortiz, 2.5 km en línea
recta al O de Alotitlán, 19º15’3”N, 103º14’8”W, 1700 m, 23 February 2012 (fl, fr), A Castro-Castro, E.A. SuarezMuro & A. Frías-Castro 2747a (IBUG!); Sierra del Halo, cerca del río Canutillo en el predio Las Palomas,
19º15’35”N, 103º14’58”W, 1700–1750 m, 8 March 2012 (fl, fr), A. Castro-Castro 2680a, E.A. Suarez-Muro & A.
Frías-Castro (IBUG!); Sierra del Halo, 12 km en línea recta al E de Pihuamo, entre los predios Las Palomas y La
Esperanza, 19º15’35”N, 103º14’58”W, 1980 m, 29 March 2012, A. Castro-Castro, E.A. Suarez-Muro & A. FríasCastro 2842a (IBUG!). Michoacán. Aguililla: near the pass ca. 15 km S of Aserradero Dos Aguas (NW of
Aguililla), 1650–1700 m, 4 March 1965 (fl), R. McVaugh 22775 (ENCB!, MICH!).
Hyptis cualensis J.G.Gonzále & Art.Castro sp. nov. (Figs. 1, 2E–H and 3)
H. rhytidea affinis sed bracteis floralibus semper linearibus (vs. foliis reductis constitutus), plus floribus per verticillastum,
bracteolis brevioribus, calycum tubo in florem et in fructum brevioribus, trichomatum annulo interno affixo ad 1–2 mm
infra dentium lineam, calycum dentibus in florem et in fructum brevioribus et distinctis (vs. calycum dentibus superis ad
basem connatis), et nuculis plus minusve brevioribus differt.
Type:—MEXICO. Jalisco. Puerto Vallarta: Ojo de Agua, 20º30’43.5”N 105º12’20.5”W, 1227 m, 1 May 2013 (fl, fr), A. FloresArgüelles & A.R. Romero-Guzmán 662 (holotype IBUG!, isotypes IEB!, MEXU!).
Shrub up to 3 m tall, stems puberulent and covered with some sessile amber glandular dots, the hairs more
abundant in young branches and toward the apex, very aromatic. Leaves with petioles 5.6–9.6(–11.5) mm long,
pilose, puberulent with sessile amber glandular dots; blade lanceolate to narrow lanceolate, 5.7–9.9 × 0.8–2 cm,
decreasing in size toward the inflorescences, coriaceous, acute to acuminate at the apex, shortly rounded at the
base, margin obscurely serrate, upper surface lustrous, pilose in the main vein, puberulent with sessile amber
glandular dots, pilose in the main vein, tomentose beneath and with secondary and tertiary veins conspicuous.
Inflorescences terminal, spiciform and dense (6.5–)10.5–21 × 1.4–2.1 cm, with pectinate opposite, subsessile or
pedunculate cymes at each node, 7–10 flowerered, peduncles 1.2–4.6 mm long, peduncle bracts compose of
reduced leaves at the base of the inflorescence and progressively differentiated into linear structures (2.5–)4–12
mm long, bracteoles linear, 1.3–2.2 mm long; inflorescence axes, peduncles and floral bracts pilose, puberulent and
with some sessile glandular dots. Flowers sessile. Calyx tube 1.6–2.5 mm long, and 1.6–2 mm wide at the throat in
flower (up to 3.1–3.5 × 2.5–2.7 mm in fruit), purple, incanous, puberulent and covered with sessile glandular dots,
internally ornate with a ring of hairs attached from teeth line to 1–2 mm below; calyx teeth triangular, 0.8–1.3 mm
long in flower (up to 1.5 mm in fruit). Corolla pink, externally and internally tomentose; tube 5.6–6.2 mm long,
1.3–2.4 mm wide at the throat, lobes subequal, 1.7–2.1 mm long. Stamens 4, attached at 3.6–4.1 mm of corolla tube
length; filament 1.7–2.7 mm long, pilose; thecae 0.5–0.8 mm long. Style 5–7.8 mm long, brown and glabrous,
without stylopodium; stigma bifurcate at the apex, the branches subulate. Mericarp 4, oblong, 1.6–1.8(–1.9) × 0.8–
1 mm, triquetrous, bronze brown, nitid, muriculate, smooth to finely reticulate, and glabrous.
Distribution, habitat and phenology:—Hyptis cualensis grows in the Pacific slopes of Sierra de El Cuale,
and it is only known from the municipality of Puerto Vallarta, Jalisco (figure 1). It inhabits in open pine-oak forest
with savannoid elements, from 1100–1227 m elevation. It shares habitat with Bejaria mexicana Benth., Byrsonima
crassifolia (L.) Kunth, Nolina sp., Pinus maximinoi H.E.Moore, Quercus sp., and Salvia ramirezii J.G.González, in
higher elevations, and also with Quercus aristata Hook. & Arn. in lower elevations. It flowers and fructifies from
February to May.
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FIGURE 2. Hyptis pseudolantana terminal spiciform inflorescences (A), cyme, calyx and corolla detail (B, D), and habit (C). Hyptis
cualensis terminal spiciform inflorescences (E), pectinate opposite cymes, calyx and corolla detail (F, G), and infructescence (H).
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FIGURE 3. Hyptis cualensis holotype.
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Etymology:—The name of Hyptis cualensis honors the mountain range to which it belongs: Sierra de El
Cuale, which is a relevant spot of plant diversity in western Mexico, several novelties and species have been
described from there in the last recent decades (González-Gallegos & Castro-Castro 2012).
TABLE 2. Morphology, habitat and distribution comparison between Hyptis cualensis and Hyptis rhytidea.
Hyptis cualensis
Hyptis rhytidea
Petiole length (mm)
5.6–9.6(–11.5)
4–8(–14)
Blade shape
lanceolate to narrow lanceolate
elliptic-lanceolate to lanceolate
Blade size (cm)
5.7–9.9 × 0.8–2
5–11.5 × 1.6–3.1
(6.5–)10.5–21
15.5–24
Shape
foliaceus (ovate) at the base of the inflorescence,
and differentiated into linear structures toward
the apex
foliaceus (ovate-elliptic) throughout the floral
branch
Length (mm)
(2.5–)4–12
(2–)6.4–13.3(–21.2)
1.2–4.6
(2.6–)3.4–5.7
Flowers per cyme
7–10
3–7
Bracteole length (mm)
1.3–2.2
3.5–5.4(–7.4)
Calyx tube size (mm)
1.6–2.5 × 1.6–2; up to 3.1–3.5 × 2.5–2.7 in fruit
2.4–2.9 × 2.5–2.6(–3); up to 7–7.4 × 3.5–3.8 in
fruit
Teeth length (mm)
0.8–1.3; up to 1.5 in fruit
2.5–4.5; up to 7.3 in fruit
Corolla color
pink
lavender, violet, blue, purple or pink
Corolla tube size (mm)
5.6–6.2 × 1.3–2.4
5.5–6.9 × 1.7–2(–3.4)
Corolla lobes length (mm)
1.7–2.1
1.3–1.9(–2.6)
Filament length (mm)
1.7–2.7
1.8–2.6
Theca length (mm)
0.5–0.8
0.5–0.8
5–7.8
5.7–7
Size (mm)
1.6–1.8(–1.9) × 0.8–1
(1.9–)2.5–2.6 × (1–)1.4–1.6
Habitat
oak forests and ecotones with tropical deciduous
forests
pine-oak, oak and tropical deciduous forests
Altitudinal range
1100–1227
(400–)1600–2000(–2400)
Distribution
Mexico: Jalisco
Mexico: Durango, Jalisco, Nayarit, and Sinaloa
Leaves
Inflorescence
Inflorescence or floral
branch length (cm)
Floral bract
Peduncle
Peduncle length (mm)
Flowers
Calyx
Corolla
Androecium
Gynoecium
Style legth (mm)
Mericarp
THREE SPECIES OF HYPTIS FROM MEXICO
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Taxonomic relationships:—Hyptis cualensis is morphologically similar to Hyptis rhytidea due to its habit as
robust shrub, indumentum, coriaceous, lustrous leaves of similar shape and size, spiciform compact inflorescences
composed of pedunculated cymes of sessile flowers, and corollas similar in shape, size and color (table 2).
However, there are several differences between them that reveal H. cualensis as a distinct species. The floral bracts
of H. cualensis are always linear except for those of the base of the inflorescence (vs. reduced leaves as floral bracts
throughout the inflorescence), more flowers per verticillaster (7–10 vs. 3–7), shorter bracteoles [1.3–2.2 vs. 3.5–54(–7.4) mm], smaller calyx tube in flower [1.6–2.5 × 1.6–2 vs. 2.4–2.9 × 2.5–2.6(–3) mm] and in fruit (3.1–3.5 ×
2.5–2.7 vs. 7–7.4 × 3.5–3.8 mm), internal ring of hairs attached from teeth line to 1–2 mm below (vs. at the middle
of calyx tube length), shorter calyx teeth in flower (0.8–1.3 vs. 2.5–4.5 mm) and in fruit (up to 1.5 vs. 7.3 mm), and
all of them distinct till the base (vs. the three upper connate at its base, and the two lower distinct), and slightly
smaller mericarps [1.6–1.8(–1.9) × 0.8–1 vs. (1.9–)2.5–2.6 × (1–)1.4–1.6 mm]. Besides, H. rhytidea occupies a
much wider geographical and ecological distribution, since it grows from Michoacán to almost southern Sonora
(Ramamoorthy & Elliot 1998), dwelling in several different types of vegetation and elevation range (table 2). In
contrast, H. cualensis is known of one locality in the Pacific slope of Sierra de El Cuale in Jalisco, with a narrow
ecological distribution in terms of the types of vegetation and elevation range it occupies (table 2).
Hyptis rhytidea is one of two species assigned by Epling (1933b, 1941) to section Rhytidea. The similarity of
H. cualensis to the former suggests it should be placed in the same section according to Epling’s proposal. Harley
& Pastore (2012) treated section Rhytidea as unplaced in any of the genera they recognized, but stating that
probably belongs to Condea Adanson (1763: 504), since H. rhytidea was recovered as sister of Condea clade in the
cladograms shown by Pastore et al. (2011). However, H. rhytidea was not sampled in that research. Furthermore,
using the key to genera they provide leads to Mesosphaerum instead of Condea.
Conservation assessment:—Hyptis cualensis appears to be in danger of extinction, based on its restricted
geographic distributional range and population size. No subpopulation estimated to contain more than 250 mature
individuals, so it should be classified as endangered [EN, criteria C2a (i)] according to the IUCN (2008).
Additional specimen examined (paratypes):—MEXICO. Jalisco. Puerto Vallarta: Ojo de Agua, 100 m al SO
de donde nace el arroyo "Las Trancas", 20º31’00.3”N 105º12’8.2”, 1219 m, 19 February 2013 (fl), A. FloresArgüelles & A.R. Romero-Guzmán 644 (IBUG!); Ojo de Agua, 2 km al SO de donde nace el arroyo "Las Trancas",
20º31’30”N 105º11’14.6”W, 1139 m, 30 April 2013 (fr) A. Flores-Argüelles & A.R. Romero-Guzmán 660 (IBUG!);
Ojo de Agua, aprox. 400 m de donde nace el arroyo "Las Trancas", siguiendo el cauce, 20º30’55.9”N
105º11’54.6”W, 1104 m, 25 August 2013 (fr), A. Flores-Argüelles & A.R. Romero-Guzmán 798 (IBUG!).
Hyptis macvaughii J.G.González & Art.Castro sp. nov. (Figs. 1, 4 and 5)
Hyptis subtilis Epling affinis sed indumento trichomatum ramosis, petiolis plus minusve brevioribus, cymis sessilibus, plus
floribus per verticillastum, pedicellis longioribus, calycum dentibus in florem et in fructum plus minusve brevioribus,
corollarum tubo longiore, stylo longiore, plus nuculis fertilibus per florem et magnioribus differt.
Type:—MEXICO. Nayarit. Compostela: 3.5 km al O de Las Mesilla por la carretera 200D rumbo a Las Varas, 15–20 km antes
de llegar a esta población, 285 m, 21 November 2013 (fl, fr), J.G. González-Gallegos & A. Castro-Castro 1547 (holotype
IBUG, isotypes CIIDIR, ENCB, IEB, MEXU, ZEA).
Shrub 1–2.5 m tall, stems puberulent, with branched hairs, young branches reddish, scarcely aromatic. Leaves with
petioles 1.7–3.1(–4) cm long, puberulent, densely covered with branched hairs; blade lanceolate, 6.2–14.5(–17) ×
(1.5–)2.5–5.6 cm, acute to attenuate at the apex, rounded at the base and slightly oblique, margin serrate, both
surfaces covered with branched hairs, mainly in the veins, and beneath. Inflorescence terminal, spiciform, lax to
dense flowered, 18–40 cm long (sometimes the spiciform inflorescences branched and forming a raceme), cymes
sessile (with no apparent peduncle), each node composed by a verticillaster with 30–40 flowers, floral bracts linear,
0.7–2.2(–4) mm long, bracteoles linear 0.8–1.2 mm long; inflorescence axes, floral bracts and pedicels covered
with appressed hairs and sometimes with branch hairs as those in the stems and leaves; inflorescences become
pendulous and flowers resupinate as the axis is overcome by the weight. Flowers with pedicels (6.7–)9–15 mm
long. Calyx tube 2–3.2 mm long and 1.8–2.1 mm wide at the throat in flower (up to 5–5.5 × 2.5–3 mm in fruit),
calyx teeth 0.2–0.4 mm long in flower (up to 0.5 mm long in fruit), densely covered with pale purple branched
hairs. Corolla dull violet to lavender, sparsely pilose at the apex including the lobes; tube (4.6–)5.2–6.4 mm long,
1.5–2 mm wide at the throat, lobes subequal, 2–2.5(–2.9) mm long. Stamens 4, attached at 3.6–5.5 mm of corolla
tube length; filament 1.7–2.5 mm, pilose; thecae 0.2–0.7 mm long. Style (8.7–)10–11.6 mm long, long exserted,
158 •
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white and glabrous, without stylopodium, bifurcate at the apex, the branches acute. Mericarp 4, ovoid, triquetrous,
1.2–1.4 × 0.8–0.9 mm, black, tiny verrucose and glabrous.
FIGURE 4. Hyptis macvaughii holotype.
THREE SPECIES OF HYPTIS FROM MEXICO
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FIGURE 5. Hyptis macvaughii spiciform and racemose terminal inflorescences (A, C, E), stem (B), verticillaster, calyx and corolla
detail (D), infructescence (F, G).
Distribution, habitat and phenology:—Hyptis macvaughii inhabits in western Jalisco, in the municipality of
Puerto Vallarta, and southern Nayarit, in the municipality of Compostela (figure 1). It grows in savannoid
vegetation and ecotones of this with tropical subdeciduous forests, from 200–430(–1204) m elevation. It shares
habitat with Alvaradoa amorphoides Liebm., Barleria oenotheroides Dum. Cours., Bejaria mexicana, Brosimum
alicastrum Sw., Bursera fagaroides (Kunth) Engl. var. elongata McVaugh & Rzed., Byrsonima crassifolia (L.),
Kunth, Chamaedora pochutlensis Liebm., Clethra rosei Britton, Cochlospermum vitifolium (Willd.) Spreng.,
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GONZÁLEZ-GALLEGOS ET AL.
Cryosophila nana (Kunth) Blume ex Salomon, Curatella americana, Dioscorea sp., Erythroxylum havanense
Jacq., Glossostipula blepharophylla (Standl.) Lorence, Hyptis suaveolens (L.) Poit., Mandevilla subsagittata (Ruiz
& Pav.) Woodson, Maranta arundinacea L., Miconia saxicola Brandegee, Pinus maximinoi, Quercus acutifolia
Née, Q. aristata Hook. & Arn., Tabebuia sp., Vitex mollis Kunth, V. pyramidata B.L.Rob., Waltheria lundelliana
J.G.Saunders, Zamia sp. and several species of grasses. It flowers and fructifies from September to December and
probably extending until March.
Etymology:—Hyptis macvaughii is dedicated in recognition to the efforts of Rogers McVaugh (1909–2009),
an outstanding botanist who contributed greatly to the exploration and knowledge of western Mexican flora. This
new name is one of many more coined to honor his contributions (Carvajal & González-Villarreal 2012).
Taxonomic relationships:—Hyptis macvaughii can be easily distinguished from the members of section
Umbellatae Epling (1949: 197) by means of its flowers directed attached to thicken portion of main floral axis
instead of forming an umbel or panicle through the insertion of pedicels to a secondary floral axis (peduncle),
together with its inconspicuous linear floral bracts and indumenta composed of branched hairs throughout stems,
leaves, inflorescence axes, pedicels and calyces (figures 4 and 5). Hyptis subtilis Epling (1933b: 79) is its
morphologically most similar species, but H. macvaughii can be separated from this because of the peculiarities
already mentioned about the inflorescence without a peduncle and plant indumenta composed of branched hairs;
the slightly shorter petioles [1.7–3.1(–4) vs. 2.7–6.1 cm], more flowers per floral node (30–40 vs. 15–21), longer
pedicels (6.7–15 vs. 4–5.8 mm), slightly shorter calyx teeth in flower (0.2–0.4 vs. 0.5–0.6 mm in flower) and in
fruit (up to 0.5 vs. 0.8 mm), longer corolla tube [(4.6–)5.2–6.4 vs. 2–2.7 mm), longer style [(8.7–)10–11.6 vs. 4.3–
4.9 mm), production per flower of 4 fertile mericarps instead of only 1 and these smaller (1.2–1.4 × 0.8–0.9 vs. 1.5–
2.1 × 1.4–1.6 mm), are other characters that supports the recognition of the former. The elevation range and type of
vegetation both species occupy are not substantially different, both grow in tropical environments, though the
distribution of H. macvaughii is much more restricted (figure 1, table 3).
TABLE 3. Morphology, habitat and distribution comparison between Hyptis macvaughii and Hyptis subtilis.
Hyptis macvaughii
Hyptis subtilis
Petiole length (cm)
1.7–3.1(–4)
2.7–6.1
Blade shape
lanceolate to narrow lanceolate
ovate-lanceolate to lanceolate
Blade size (cm)
6.2–14.5(–17) × (1.5–)2.5–4(–5.6)
8.5–15 × 3.3–8.8
Inflorescence or floral
branch length (cm)
18–40
20–30
Peduncle length (mm)
0
5–11
Shape
linear
linear
Length (mm)
0.7–2.2(–4)
3.7–5.7
Flowers per node
30–40
15–21
Bracteole length (mm)
0.8–1.2
0.6–2.5
Pedicel length (mm)
(6.7–)9–15
4–5.8
Calyx tube size (mm)
2–3.2 × 1.8–2.1; up to 5–5.5 × 2.5–3 in fruit
2.6–3.6 × 1.3–1.9; up to 5–5.4 × 2.3–2.5 in fruit
Teeth length (mm)
0.2–0.4; up to 0.5 in fruit
0.5–0.6; up to 0.8 in fruit
Leaves
Inflorescence
Floral bract
Flowers
Calyx
..... continued on the next page
THREE SPECIES OF HYPTIS FROM MEXICO
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• 161
TABLE 3. (Continued)
Hyptis macvaughii
Hyptis subtilis
Color
dull violet to lavender
dull violet to lavender
Tube size (mm)
(4.6–)5.2–6.4 × 1.5–2
2–2.7 × 1.6–2
Corolla lobes length (mm)
2–2.5(–2.9)
2–2.6
Filament length (mm)
1.7–2.5
1.8–2.2
Theca length (mm)
0.2–0.7
0.5–0.7
(8.7–)10–11.6
4.3–4.9
Size (mm)
1.2–1.4 × 0.8–0.9
1.5–2.1 × 1.4–1.6
Habitat
savannoid vegetation and ecotones of this with
tropical subdeciduous forests
tropical subdeciduous forests and rarely into oak
forests
Elevation range
200–430(–1204)
(60–)200–600(–1000)
Distribution
Mexico: Jalisco and Nayarit
Mexico: Oaxaca, Guerrero, Estado de México,
Michoacán, Colima, Jalisco and Nayarit
Corolla
Androecium
Gynoecium
Style length (mm)
Mericarp
Hyptis iodantha Epling (1939: 16) is the third representative of section Umbellatae in Mexico. The three
species altogether constitute an interesting case of disjunction since the other species of this section grow in South
America (Argentina, Bolivia, Brazil, Colombia, Paraguay, Peru, and Uruguay) without any continuous connection
between both areas of distribution (Fernández-Alonso 2010). Finally, H. macvaughii would belong to the genus
Condea according to the proposal of Harley & Pastore (2012).
Conservation assessment:—Hyptis macvaughii appears to be in danger of extinction, based on its limited
geographic range. It should be classified as endangered [EN, criteria B1a] according to the IUCN (2008). Changes
in land use are the greatest threat to this species.
Additional specimen examined (paratypes):—MEXICO. Jalisco. Puerto Vallarta: Ojo de Agua, 8.3 km en
línea recta al SO de Llanitos, 1204 m, 28 November 2013, A. Flores-Argüelles & A.R. Romero 873 (IBUG); cresta
Palo María y Chupalodo, hacia el mirador de la cascada del río Mismaloya, 430 m, 30 November 2013, E. IñarrituMedina et al. s.n. (IBUG). Nayarit. Compostela: 5–6 miles W of Mazatán, 350 m, 17 September 1960 (fl), R.
McVaugh 19093 (MICH!); 8 km above (E of) Las Varas, new road to Compostela, 200 m, 20 December 1970 (fl),
R. McVaugh 25584 (MICH!); 11 km by road E of Las Varas toward Compostela, 200 m, 28 October 1971 (fl),
J.V.A. Dieterle 3933 (MICH!); along highway 200, W of Compostela, 2.5 miles W Mesillas, 280 m, 9 October
1978 (fl), T.F. Daniel 1070 (MICH!); 3.5 km al O de Las Mesillas por la carretera 200D rumbo a Las Varas, 15–20
km antes de llegar a esta población, 285 m, 15 August 2013 (fr), J.G. González-Gallegos & A. Castro-Castro 1536
(IBUG!, IEB!, MEXU!, ZEA!).
Key to species of tribe Hyptidinae in western Mexico
[Asterohyptis nayarana B.L.Turner (2011: 2) is not included in the key below because we consider this as synonym
of A. seemannii Epling (1933: 20), an issue that is being justified in an ongoing publication]
1.
–
Inflorescence lax capitate generally with less than 5 flowers; calyx lobes reflexed in fruit (backwardly oriented); mericarps
cymbiform .................................................................................................................................... Marsypianthes chamaedrys
Inflorescences variable, racemiform to spiciform, if capitate, these are compact and built by more than 10 flowers, except
in Hyptis suaveolens which has 5 or less flowers per cyme; calyx lobes straight in fruit (uprightly or laterally, but not backwardly, oriented); mericarps ovoid to flattened...................................................................................................................... 2
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2.
–
3.
–
4.
–
5.
–
6.
–
7.
–
8.
–
9.
–
10.
–
11.
–
12.
–
13.
–
14.
Plants gynodioecious; calyx lobes thin and resembling spines, laterally oriented, which give to the calyx a five-pointed star
appearance in sectional view; corollas white, middle lower corolla lip not thickened and not wrapping the stamens, which
are not explosively released ................................................................................................................................................... 3
Plants monoecious with bisexual flowers; calyx lobes variable in shape and generally uprightly oriented, without a fivepointed star appearance in sectional view; corollas white, blue, purple, violet, to pink, middle lower corolla lip thickened
and wrapping the stamens, which explosively releases them when these are mature and in response to vibration .............. 4
Floral nodes in blossom 5–7 mm diameter; calyx lobes 0.5–0.7 mm long. Plants from northern Jalisco, Nayarit, Durango,
Sinaloa, Sonora and Chihuahua........................................................................................................... Asterohyptis seemannii
Floral nodes in blossom 8–12 mm diameter; calyx lobes 1.5–3.5 mm long. Plants widely distributed from Sonora (Mexico)
to Central America ................................................................................................................................Asterohyptis stellulata
Plants with branched hairs in the stems, leaves and inflorescences (in Hyptis oblongifolia these hairs are more abundant in
the stems and leaves, but sometimes are difficult to be appreciated, and at times are absent) .............................................. 5
Plants without branched hairs ................................................................................................................................................ 7
Flowers sessile, arranged in opposite glomeruli throughout the floral axis, with peduncles 3–10 mm long. Plants from oak,
pine-oak and montane cloud forests ......................................................................................................... Hyptis oblongifolia
Flowers pedicellate, pedicels 2–14 mm long, directly inserted in floral axis, without a secondary peduncle. Plants typical
from roadsides, tropical deciduous forests, savannoid and secondary vegetation ................................................................. 6
Leaves ashy green above, white tomentose below; floral axes, pedicels and calyces white due to the pubescence; pedicels
2–3 mm long; calyx teeth 1.5–1.8 mm long. Plants from roadsides, tropical deciduous forests and secondary vegetation;
widespread throughout Mexico ...........................................................................................................................Hyptis albida
Leaves bright green on both sides, sparsely pubescent; floral axes, pedicels and calyces pale purple to green; pedicels (6.7–)9–
15 mm long; calyx teeth 0.2–0.4 mm long. Plants from savannoid vegetation and subtropical deciduous forests; endemic to
western Jalisco and southern Nayarit. ........................................................................................................ Hyptis macvaughii
Inflorescences in spherical glomeruli 1.3–3 cm in diam., covered at the base with an involucre of lanceolate bracts, each
glomerulus arranged to leaf axil with a peduncle (1.6–)3–12 cm long; flowers sessile or with pedicels up to 0.5 mm long .
.......................................................................................................................................................................... Hyptis capitata
Inflorescences spiciform and racemiform, if similar to a glomerulus, then less than 10 mm in diam., without an involucre
at the base, not arranged to leaf axil (but to a inflorescence axis) and peduncles shorter than 2 cm long; flowers mostly with
pedicels 1 mm long................................................................................................................................................................. 8
Leaves coriaceous, petioles 2–10(–11.5) mm long; inflorescences compact with sessile flowers, arranged in sessile pectinate opposite cymes or with peduncles up to 5 mm long ...................................................................................................... 9
Leaves thin, petioles regularly longer than 15 mm long; inflorescences lax with sessile to pedicellate flowers, arranged in
glomeruli with peduncles longer than 8 mm long ................................................................................................................ 10
Peduncle bracts composed by reduced leaves throughout the inflorescence; flowers per cyme 3–7; bracteoles 3.5–5.4(–
7.4) mm long; calyx bilabiate, upper lip composed by three connate teeth, lower lips composed by the other two teeth;
calyx tube 2.4–2.9 × 2.5–2.6(–3) mm in flower, and 7–7.4 × 3.5–3.8 mm in fruit; calyx teeth 2.5–4.5 mm long in flower
and 7.3 mm long in fruit; mericarp (1.9–)2.5–2.6 × (1–)1.4–1.6 mm. Plants widespread in Jalisco, Nayarit, and southern
Sinaloa and Durango, from (400–)1600–2000(–2200) m elevation ................................................................Hyptis rhytidea
Peduncle bracts composed by reduced leaves only at the base of the inflorescence and then differentiated into linear structures; flowers per cyme 7–10; bracteoles 1.3–2.2 mm long; calyx actinomorphic, without two sets of teeth; calyx tube 1.6–
2.5 × 1.6–2 mm in flower, and 3.1–3.5 × 2.5–2.7 mm in fruit; calyx teeth 0.8–1.3 mm long in flower, and up to 1.5 mm
long in fruit; mericarp 1.6–1.8(–1.9) × 0.8–1 mm. Plants exclusive of the Pacific slopes of Sierra de El Cuale, Jalisco,
growing from 1100–1227 m elevation ........................................................................................................... Hyptis cualensis
Inflorescences spiciform cylindrical, compact (floral axis surface is hidden by flowers and floral bracts); floral bracts linear (less than 0.5 mm wide) with rigid central vein and ciliated at margin; flowers sessile with calyces hidden by floral
bracts ............................................................................................................................................................. Hyptis spicigera
Inflorescences racemiform, very rarely with spiciform appearance; floral bracts ovate (more than 1 mm wide) to linear,
without rigid central vein and not ciliated at margin; most flowers with pedicels at least 1 mm long, calyces not completely
hidden by floral bracts ..........................................................................................................................................................11
Floral bracts ovate to ovate-lanceolate; calyces tubular ................................................................................ Hyptis mutabilis
Floral bracts linear; calyces infundibuliform ...................................................................................................................... 12
Inflorescences in purple lax racemes; pedicels 3–12 mm long; calyx lobes triangular, less than 0.5 mm long, which gives to
the calyx a truncate appearance at apex ........................................................................................................... Hyptis subtilis
Inflorescences in green to yellowish compact racemes; pedicels absent or shorter than 3 mm long; calyx lobes linear or
shortly triangular at base and then linear, longer than 1 mm, hence the calyx without a truncate appearance at apex ....... 13
Flowers up to 5 in each cyme; calyx tube 5–8 mm long and 6–7 mm wide at the throat, in fruit (there is a striking difference between flowering and fruiting calyx tube size in a same individual)................................................ Hyptis suaveolens
Flowers 10 or more in each cyme; calyx tube 2–4 mm long and less than 2.5 mm wide at the throat, in fruit (size difference
between flowering and fruiting calyx tubes in the same individual is not apparent) ........................................................... 14
Flowers in compact spherical glomeruli, densely white tomentose, the hairs hidding almost whole floral bracts and calyx
tubes, usually with branched hairs mainly in the leaves and stems, but which sometimes are difficult to be appreciated ....
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................................................................................................................................................................... Hyptis oblongifolia
Flowers in hemispherical glomerulus or pectinate cymes, not densely tomentose, the hairs not hidding floral bracts and
calyx tubes, never with branched hairs ................................................................................................................................ 15
15. Calyx lobes linear (triangular only in the base for 1/4 to 1/5 of its length), with a ring of hairs clearly defined inserted internally in the throat and projecting between the lobes ........................................................................................................... 16
– Calyx lobes triangular, with a ring of hairs diffusely or well-defined inserted internally in the throat but not projecting
between the lobes ................................................................................................................................................................. 17
16. Flowers arranged in pectinate cymes with peduncles 1–2 mm long; calyx tube surrounded with a thickened vein just below
its apical portion, being perpendicular to calyx lobes .................................................................................... Hyptis pectinata
– Flowers arranged in not pectinate cymes with peduncles 10–30 mm long; calyx tube without a thickened vein surrounding
it at its apical portion ..................................................................................................................................... Hyptis urticoides
17. Stems, petioles, and leaves without glandular-capitate hairs; blades 1.6–5.5 × 0.9–3.7 cm, rounded to short cuneate at the
base, irregularly serrate at the margin; flowers arranged in pectinate cymes, peduncles 3–7 mm long; calyx 2.6–3.7 mm
long in flower, with a diffuse ring of hairs internally inserted just below the throat. Plants only known from western sides
of Sierra del Halo, Jalisco, eastern Sierra de Coalcomán, and northeastern slopes of Sierra Madre del Sur in Guerrero.......
............................................................................................................................................................... Hyptis pseudolantana
– Stems, petioles, and lower blade surface covered with glandular-capitate hairs; blades 6–10 × 4–6 cm, cordate at the base,
serrate at the margin; flowers in spherical glomerulus, peduncles 8–20 mm long; calyx 4–5.9 mm long in flower, with a
well-defined ring of hairs internally inserted below the throat. Plants only known from the mountain range in San
Sebastián del Oeste and Sierra de El Cuale, Jalisco......................................................................................Hyptis pinetorum
–
Acknowledgments
We thank the curators and colleagues from the herbaria consulted (ENCB, IBUG, IEB, MEXU, MICH, ZEA), who
kindly provided us with all facilities needed. The online provision by several herbaria (K, MO) of type specimen
images is also much appreciated. Alfredo Frías Castro, Luis F. Bugarín Navarro and Esteban A. Suárez Muro
helped considerably with fieldwork in Sierra del Halo, Tecalitlán, Jalisco. Thanks to Marco Antonio Carrasco Ortiz
for generous assistance on the preparation of the distribution map of the species. Luz María González Villarreal and
María Ivonne Rodríguez Covarrubias helped with scanning of the holotypes. Financial support to ACC and JGGG
was partially provided by CONACYT (Comisión Nacional de Ciencia y Tecnología) and Universidad de
Guadalajara, Mexico. The fulfillment of this research was possible through a research stay by JGGG under the
guidance of Kenneth Sytsma (University of Wiscosnin-Madison), and scholarship granted by CONACYT for
research stays abroad. Finally, thanks to Idea Wild organization for the equipment given to ACC.
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