Systematic Implications of Pollen Morphology in Subfamilies Lamioideae and Pogostemonoideae (Labiatae)

34 Pages·1994·12.1 MB·

by  Mones S Abu-Asab

SYSTEMATIC IMPLICATIONS
Mones Abu-Asab 2 and Philip D. Cantino
S.
MORPHOLOGY
POLLEN
OF IN
LAMIOIDEAE
SUBFAMILIES
POGOSTEMONOIDEAE
AND
(LABIATAE)
1
Abstract
Pollen morphology was surveyed in 57 of the 60 genera of Lamioideae and Pogostemonoideae sensu Cantino et
al. and two genera of uncertain subfamilial affinities: Anisomeles and Eurysolen. Pollen in all Pogostemonoideae,
most Lamioideae, Anisomeles, and Eurysolen tricolpate with suprareticulate sculpturing. Variants found in a few
is
genera of Lamioideae include tetracolpate and hexapantocolpate pollen and psilate, granulate, and rugulate sculpturing.
Most species in which pollen was examined in section have simple columellae, but seemingly branched columellae
were observed in Gomphostemma Chelonopsis, and one species of Stenogyne. With regard to the genera of uncertain
y
affinities, the findings presented here support a relationship between Anisomeles and Pogostemon; the position of
Lamiophlo-
Eurysolen remains unclear. Within subfamily Lamioideae, pollen morphology supports the segregation of
mis, Lagopsis, and Phlomidoschema from Phlomis, Marrubium, and Stachys, respectively. Hexapantocolpate pollen
is hypothesized to be a synapomorphy of Sideritis section Empedoclea. Similarities in pollen sculpturing were noted
between Brazoria, Macbridea, and Physostegia, and between Galeopsis and Synandra.
A (Cantino 1992),
pollen survey of subfamily Lamioideae sensu alternative classification et al.,
Erdtman (1945) was undertaken with the hope that which is followed here. Unless otherwise indicated,
might further our understanding of systematic references herein to subfamily Lamioideae imply
it all
relationships. A brief overview of the study as a the circumscription of Cantino et al. (1992).
& 55
whole has been published (Abu-Asab Cantino, Subfamily Lamioideae comprises genera, the
Subfam-
them Eurasian (Table
1992), with elaboration elsewhere on two small vast majority of 1 ).
subgroups Pogostemonoideae an Asian group of five
that appear not to be closely related to ily is
& Based on morphological and molecular ev-
the remainder of the study group (Abu-Asab genera.
Cantino, 1993a, b). It is our intent here to doc- idence, the two subfamilies are closely related (Can-
&
ument 1992a; Downie Palmer, 1992), and Eu-
in detail the pollen morphology of the bulk tino,
—
of Erdtman's Lamioideae those genera with rysolen resembles genera in both. (See Table 1 for
i.e.,
names and Appendix authors
a gynobasic authors of genus for
style.
A Anisomeles has been variably
we names.)
taxonomic needed because of species
clarification
is
have circumscribed subfamily Lamioideae differ- assigned to both subfamilies (see Discussion).
members
ently here than some our works Most pollen studies that included of
in of earlier (e.g.,
& Pogostemonoideae have centered
Cantino Sanders, 1986; Abu-Asab, 1990; Can- Lamioideae and
& Huynh, 1972; Bassett
genera
tino, 1990, 1992a; Abu-Asab Cantino, 1992). on particular (e.g.,
&
Although Munro, 1986) or geographical regions (e.g..
the pollen survey was initiated as a study
&
Verma, 1968;
of the Lamioideae sensu Erdtman 945), and our Waterman, 1960; Varghese
1
(
1956) and Pozhidaev(1989)
earlier papers adopted Erdtman's circumscription, Huang, 1972). Risch(
recent evidence (Abu- conducted broader surveys of the Labiatae, but
that taxon polyphyletic
this
is
Asab & microscopy, and his descrip-
Cantino, 1992; Cantino, 1992b) led to an Risch used only light
' This paper is based in large part on the doctoral dissertation of the first author, submitted to Ohio University in
WO.
Smithsonian Institution;
The Penological Laboratory at the
final portions of the study were completed in the
We permission
*e are most grateful for Joan Nowicke's advice and encouragement. thank the following institutions for
was
study
to remove BHO, BM, CH, K, MO, US. Financial support for this
pollen from herbarium specimens: A,
Fund
BSR-9006209 and Ohio Univer^.y Baker
Provided by BSR-8306878 and
National Science Foundation grants
Award
88-10
1 Department P.O. Box 14, Birzeit, West Bank, via Israel. Reprint
of Biology and Biochemistry, Birzeit University,
requests
should be sent to the second author. ..-.
1 Department of Environmental and Plant Biology, Ohio University, Athens, Ohio 45701-297 ^ U.>.A.
653-686. 1994
Card. 81:
Ann. Missouri Bot.
—
654 Annals of the
Garden
Missouri Botanical
Table Genera of Lamioideae and Pogostemonoide- Table Continued.
1 1.
.
Numbers number
ae. in parentheses: of species in the
—
genus/number of species studied (Appendix). Numbers of Paraphlomis Prain (8/3) Eastern Asia, Malesia
—
species were taken from a variety of sources, geographic Phlomidoschema (Benth.) Vved. (1/1) Iran
to
distributions mostly from Mabberley (1987) and Airy Shaw western Himalayas
—
(1973). Phlomis 100/7) Mediterranean
L. (ca. region to
China
Subfamily Lamioideae (28/5)—
Phyllostegia Benth. Hawaii, Tahiti
—
—
mum
Achyrospei Blume (10/6) Paleotropics Physostegia Benth. (12/2) North America
—
—
&
Acrotome Benth. ex Endl. (8/4) Tropical south- Prasium L. (1/1) Mediterranean region
—
ern Africa Pseuderemostachys Popov (1/1) Central Asia
—
Ajugoidcs Makino (1/1) Japan Pseudomarrubium Popov (1/0)— Central Asia
—
Alajja Ikonn. (1/1) Himalayas Roylea Wallich ex Benth. (1/1)— Himalayas
—
130/19)—
Ballota L. (35/7) Europe, Africa, western Asia Sideritis L. (ca. Mediterranean region to
—
Bostrychanthera Benth. (1/0) China southwestern Asia, Macaronesia
&
Brazoria Engelm. & Gray (4/4)— Oklahoma Stachyopsis Popov Vved. (3/1)— Central Asia to
A. Texas,
—
Chaiturus Temperate western China
Willd. (1/1) Eurasia
&
& 300/2)—
Chamaesphacos Schrenk ex Fischer C Meyer (1/ Stachys L. (ca. Northern southern tem-
—
& &
mountains
1) Central southwestern Asia perate subtropical; tropical
—
—
Chelonopsis Miq. (13/2) Himalayas, China, Japan Stenogyne Benth. (20/3) Hawaii
—
&
Colquhounia Wallich (5/2) Himalayas Sulaimania Hedge Rech. (1/1)— Pakistan
f.
Craniotome Rchb. (1/1)— Himalayas Suzukia Kudo (2/1)— Taiwan, Ryukyus
Eremostachys Bunge ex Ledeb. sensu (60/4) Synandra Nutt. (1/1)— Eastern United States
lato
&
Europe to central Asia Thuspeinanta T. Durand (2/1)— Central south-
—
Eriophyton Benth. (1/1) Himalayas western Asia
—
&
(2/l)-South-
Meyer
Galeopsis L. (10/6) Temperate Eurasia JVeidemannia Fischer C.
Gomphostemma
Wallich ex Benth. (30/7)— South- western Asia
eastern Asia, Malesia
Subfamily Pogostemonoideae
Haplostachys (A. Gray) Hillebrand (5/2)— Hawaii —
Colebrookea Smith (1/1) India
Hypogomphia &
Bunge (1/1)—
Central southwest-
Comanthosphace Moore (7/3)— Eastern Asia
S.
ern Asia
— Leucosceptrum Smith (1/1)— Himalayas, China
Isoleucas Schwartz (1/0) Arabia
(71/10)— Malesia
Pogostemon Desf. China, India,
Lagochilus Bunge ex Benth. (35/3)— &
Central
Rostrinucula Kudo (2/1)— Eastern Asia
southwestern Asia
Lagopsis Bunge (4/2)— Western Genera
Siberia to Japan of uncertain affinities
Lamiophlomis Kudo (1/1)—
Himalayas (6/4)—
Anisomeles R. Br. Paleotropics
Lamium
(16/7)— Temperate Indoma-
L. Eurasia, northern Eurysolen Prain (1/1)— Southeastern Asia,
Africa
lesia
Leonotis (Pers.) R. Br. (15/3)— Tropical & southern
Africa, pantropical
1
Leonurus (24/2)—
L. Temperate Eurasia
morphology size,
tions emphasized gross pollen (i.e.,
Leucas R. Br. (100/17)— Pantropical, southern
Afri- used
survey
shape, Pozhidaev's (1989)
color).
ca, China
(SEM), but tne
scanning electron microscopy
Loxocalyx Hemsley (3/2)—
China, Japan
exine
on variation in
Macbridea suiting publication focused
ex (2/2)—
Elliott Nutt. Southeastern
mor-
pollen
United States structure in the family as a whole; the
Marrubium systematically
L. (30/5)— Europe, northern Africa, phology of individual genera was not
&
central southwestern Asia were provided for
described, and micrographs
Mrlittis L. (1/1)— Europe We are aware
than a third of the genera studied.
Metastachvdium Wu &
Airy Shaw ex C. Y. SEM only -
Li (1/ micrographs for
— of published pollen
1) Central Asia
Pogostemo-
of the 60 genera of Lamioideae and
Microtocna
Prain (25/4)—
Himalayas, China Annan
1978;
noideae 1976; Roca Salinas,
(Nabli,
Moluccclla (2/2)—
L. Southern Europe,
southwest- &
& Galan.
Ubera
1982;
Moore, 1982; Cantino,
ern Asia
& Sand-
&
Cantino
1983; Munro, 1986;
hotochaetc Benth. (1/1)— Himalayas Bassett
&
Abu-Asab
Otostegm Benth. (20/5)— ers, 1986; Pozhidaev, 1989, 1992;
Northeastern
tropical Afri-
Trude
&
1992;
ca to central Asia Cantino, 1992; Demissew Harley,
Panzerina Sojak (2/1)— Western & Few have been studies
Siberia Mongo- Morton, 1992). species
to
sur
lia in
variation
so the degree of intrageneric
Parakunium Dunn (1/1)— ge
Southwestern China unknown most of these
sculpturing for
is
Number
Volume 81 4 Abu-Asab & Cantino 655
,
1994 Pollen Morphology Lamioideae and
in
Pogostemonoideae
exceptions are Eremostachys, Paraphlomis,
lead citrate, and examined and photographed with
Phlomis, Physostegia, and Stachys. Transmission a Hitachi HS-8 transmission electron microscope.
(TEM)
electron microscope photographs have been
published for only three genera (Nabli, 1976; Abu-
Results
&
Asab Cantino, 992).
1
The
The objectives of this paper are to document results are summarized in Table 2, and
the pollen morphology of subfamilies Lamioideae representative pollen grains are illustrated in Fig-
ures 1-247. The between
and Pogostemonoideae, distinction foveolate,
including interspecific vari-
ation within most of the larger genera, and to microreticulate, punctate, reticulate, and scrobic-
evaluate the systematic ulate sculpturing follows Vezey et al. (1992), a
significance of the resulting
data. Because the resulting compilation of SEM more precise system of terminology than used in
our previous papers (Abu-Asab, 1990; Abu-Asab
photographs
includes nearly every genus these
in
&
Cantino, 1989, 1992, 1993a, Abu-Asab
subfamilies, should serve as a useful reference b; et
it
we
for those studying the systematics or palynology al., 1993). Pollen that previously referred to
as tectate-perforate described here as scrobic-
of the Labiatae. is
ulate or punctate.
Materials and Methods
LAMIOIDEAE
The
study collection (Table comprised 183
1)
species representing genera of Pogoste- Pollen of subfamily Lamioideae suboblate to
all five is
&
monoideae, 52 55 Walker
of the genera of Lamioideae, euprolate (shape classification follows
and two genera of uncertain subfamilial affinities Doyle, 1975, based on P/E ratios in Table 2),
^m
15-59
(Anisomeles and Eurysolen). In most of the larger with the polar axis and the equatorial
genera, an effort was made to sample a broad axis 13-52 /im. The grains are inoperculate and
spectrum of species by reference to the infrageneric usually tricolpate; tetracolpate and hexapantocol-
classifications of Briquet (1895-1897) and avail- pate grains were found in a few genera (Table 2).
able monographs and Eremostachys, The exine most species scrobiculate (some-
revisions. in is
rhlomis, Physostegia, and Stachys were only su- times bordering on microreticulate or punctate).
perficially sampled because they have already re- The exine is microreticulate in seven species, re-
&
ceived considerable study (Azizian Moore, 1982; ticulate in one, and partially foveolate in one (Table
& &
Cantino, 1982; Ubera Galan, 1983; Bassett The sculpturing suprareticulate in most spe-
2). is
Munro, 1986; Demissew & Harley, 1992). The cies (referred to as bi-reticulate by some authors;
&
monotypic genera Bostrychanthera, Pseudomar- Demissew Harley, 1992; Harley et al., 1992),
rubium, and holeucas were not examined for lack but four other forms of sculpturing were found:
of material, but micrographs former two granulate, rugulate, and suprareticulate-
of the psilate,
were
provided by Pozhidaev 989); thus only the rugulate. The columellae are usually simple but
1
(
pollen of holeucas remains unknown. sparsely branched in a few species.
totally
Pollen obtained from mature buds (vouchers The pollen features in a few genera warrant
list-
ed in Appendix) was prepared SEM, TEM, and a more detailed description than is available in
for
"ght microscopy (LM) using the prolonged aceto- Table 2:
& Gomphostemma 62-76; Pozhidaev,
•ysis procedure Abu-Asab (Figs.
of Cantino (1989),
modified from Erdtman 960). For LM, including 1989): sculpturing exceptionally variable: psilate
1
(
size measurements, the pollen was mounted in gly- or with suprareticulate, rugulate, or granulate
cerine jelly and sealed with paraffin. In most cases, sculpturing; columellae branched.
seven
to ten grains were measured per species. The Lagopsis (Fig. 77): exine rugulate, psilate to-
ratio of polar to equatorial axis length (P/E) was ward the poles, apparently nonperforate.
oetermined mean 182-196; Huynh, 1972; Nabli,
for each measured grain, and the Sideritis (Figs.
P/E
ratio was calculated from these individual 1976; Roca Salinas, 1978; Pozhidaev, 1992): tet-
values. For SEM, pollen was coated with gold- racolpate (the amb circular to more or less square),
palladium and examined and photographed with a 6-pantocolpate (the amb circular), or rarely tri-
Hitachi HHS-2R amb exine psilate (ap-
scanning electron microscope. For colpate (the triangular);
'EM,
pollen was treated with osmium tetroxide, proaching a suprareticulate condition in section
'ained with uranyl acetate, and embedded in resin. Empedoclea (Raf.) Benth.; Figs. 192, 193), usu-
«nin sections were collected on carbon-stabilized ally scrobiculate (in S. montana, the polar regions
ormvar, mesocolpia foveolate, the
stained with aqueous uranyl acetate and and centers of the three
0)
Table 2. Summary of pollen morp hological data for subfamilies Lamioideae and Pogostemonoideae and genera of uncertain affinities. Colpi: 3, tricolpate; 4, tetracolpate; 6,
CJ1
hexapantocolpate. Columellae: simple vs. branched. Sculpturing: Upper layer: Gr, granulate; Ps, psilate; Ru, rugulate; Sr, suprareticulate-rugulate; Su suprareticulate. Lower layer
., ,
\
(or only layer if psilate): Fo, foveolate; Mi, microreticulate; Np, nonperforate; Pu, punctate; Re, reticulate; Sc, scrobiculate; slash indicates borderline designations.
P/E
Sculpturing
Polar axis itim) Equatori al axis (jrni) ratio
Mean Range Mean Range Mean Range Colpi Columellae Upper Lower Figures
Species
Subfamily Lamioideae
Achyrospermum aethiopicum 21.3 (20-22) 18.5 (18-20) 1.16 (1.0-1.3) 3 Su Sc 1,2
Achyrospermum carvalhl 26.9 (22-31) 23.9 (20-29) 1.13 (0.8-1.3) 3 Su Sc/Mi 3
Achyrospermum cryptanthum 24.0 (22-31) 19.1 (18-22) 1.25 (1.1-1.4) 3 Su Sc 4
Achyrospermum densijiorum 24.5 (22-26) 18.4 (15-22) 1.34 (1.0-1.7) 3 Su Sc 5
Achyrospermum parvijiorum 23.3 (22-24) 18.9 (18-20) 1.24 (1.1-1.3) 3 Su Sc 6,7
Achyrospermum schimperi 28.0 (22-33) 20.9 (20-22) 1.34 (1.1-1.5) 3 Su Sc 8,9
Acrotome 20.2 (20-22) 18.0 (17-20) 1.13 (1.0-1.3) 3 Su Sc 10, 11
angustifolia
? ?
Acrotome 20.7 (20-22) 22.0 (20-24) 0.94 (0.8-1.1) 3
fleckii
• •
Acrotome hispida 20.0 (-) 21.6 (20-22) 0.92 (0.9-1.0) 3 ?
* •
Acrotome 22.9 (22-24) 22.9 (22-24) 1.00 (0.9-1.1) 3 ?
inflata
•
Ajugoides humilis 31.9 (31-35) 23.5 (22-26) 1.36 (1.2-1.6) 3 Su Sc/Mi 12, 13
Alajja rhomboidea 30.1 (29-33) 28.4 (26-31) 1.06 (0.9-1.3) 3 Su Sc 14, 15
Ballota africana 22.7 (22-24) 20.7 (20-22) 1.10 (1.0-1.2) 3 Su Sc 16
Ballota andreuzziana 23.5 (22-24) 20.9 (20-22) 1.13 (1.0-1.2) 3 Su Sc/Mi 3
C/>
Ballota hirsuta 22.0 (-) 18.0 (17-20) 1.22 (1.1-1.3) 3 Su Sc 17, 18 o &
Ballota integrifolia 21.6 (20-24) 21.3 (20-24) 1.01 (0.9-1.1) 3 Su Mi 19 o
3.
Ballota nigra 23.8 (22-26) 20.9 (18-24) 1.14 (1.0-1.3) 3 Su Sc 20
CD
~Z
O
ZJ
Su Sc/Mi 21,22
Ballota pseudodictamnus 29.3 (26-31) 20.5 (18-22) 1.43 (1.3-1.6) 3
Ballota rupestris 21.2 (20-22) 23.4 (22-24) 0.90 (-) 3 Su Sc
o
Brazoria arenaria 35.2 (33-40) 32.3 (31-33) 1.09 (1.0-1.2) 3 Su Sc 23
O
Brazoria pulcherrima 37.0 (35-42) 31.7 (29-33) 1.17 (1.1-1.5) 3 Su Sc 24
Brazoria scutellarioides 32.5 (29-35) 33.0 (29-35) 0.99 (0.8-1.2) 3 Su 9 25
m
Brazoria truncata 39.2 (37-42) 32.6 (29-35) 1.21 (1.1-1.5) 3 Su Sc 26,27 arden
Chaiturus marrubiastrum 21.2 (20-23) 20.5 (18-22) 1.04 (0.9-1.2) 3 Su Sc 28,29
Chamaesphacos 30.4 (24-33) 24.0 (20-26) 1.27 (1.1-1.4) 3 Su Sc 30
ilicifolius
Chelonopsis lichiangensis 28.4 (24-31) 27.3 (24-31) 1.04 (0.9-1.2) 3 Branched? Su Sc/Mi 31,32
Chelonopsis odontochila 28.0 (26-29) 26.4 (24-29) 1.06 (1.0-1.2) 3 Su Sc/Mi 33,34
Colquhounia coccinea 27.7 (26-31) 26.6 (26-29) 1.04 (0.9-1.2) 3 Su Sc
Colquhounia 23.3 (22-26) 21.6 (20-24) 1.09 (0.9-1.3) 3 Su Sc 35, 36
seguinii
37-39
Craniotonir furrnta 16.6 (15-18) 14.9 (13-15) 1.12 (1.0-1.3) 3 Su Sc
Kremostuchys 33.4 (31-37) 29.5 (24-33) 1.14 (0.9-1.6) 3 Su Sc
fetissovii
Krrmontachys UierwU 31.7 (.31-33) 27.1 (24-29) 1.17 (1.1-1.3) 3 Su Sc 40, 41
11
Table 2. Continued.
o
CO
~-
CD
P/E
Polar axis (/mi) Equatorial axis (/mi) ratio Sculpturing ^cl
09
Species Mean Range Mean Range Mean Range Colpi Columellae Upper Lower Figures
Z
Eremostachys rotata 34.5 (29-37) 32.3 (29-35) 1.07 (0.8-1.2) 3 Su Sc 42 c
3
Eremostachys speciosa 35.9 (33-37) 29.5 (26-33) 1.23 (1.1-1.4) 3 Su Sc 43
Eriophyton wallichianum 28.6 (26-31) 23.5 (22-26) 1.22 (1.0-1.4) 3 Su Sc 44,45 %
Galeopsis bifida 38.7 (33-46) 32.1 (26-37) 1.22 (0.9-1.4) 3 Su Sc 46,47 *»
Galeopsis ladanum 34.9 (31-40) 26.7 (24-31) 1.31 (1.2-1.5) 3 Su Sc
Galeopsis pubescens 32.1 (26-35) 26.0 (22-31) 1.24 (1.0-1.4) 3 Su Sc 48
Galeopsis segetum 35.2 (31-40) 26.8 (24-31) 1.32 (1.0-1.6) 3 Su ?
1
Galeopsis speciosa 37.8 (35-40) 27.3 (26-29) 1.39 (1.2-1.5) 3 Su Sc 49
Galeopsis tetrahit 38.5 (35-42) 29.0 (26-31) 1.33 (1.1-1.5) 3 Su Sc 50
Gomphoslemma
chinense 33.7 (31-35) 33.4 (33-35) 1.01 (0.9-1.1) 3 Gr Sc/Mi 62,63
Gomphostemma intermedium 32.1 (31-35) 32.6 (31-33) 0.99 (0.9-1.0) 3 Branched Ps Mi 64-66
Gomphoslemma javanicum 29.7 (26-31) 30.8 (29-33) 0.97 (0.9-1.0) 3 Ru Sc 69,70
Gomphostemma
leptodon 33.2 (29-40) 35.4 (29-42) 0.94 (0.9-1.2) 3 Ps Sc 67,68
Gomphostemma lucidum 32.3 (29-37) 32.6 (31-33) 0.99 (0.8-1.1) 3 Branched Ps Sc 71,72
Gomphostemma
parviflorum 26.6 (24-29) 25.7 (24-29) 1.04 (0.9-1.1) 3 Gr Sc 73, 74
Gomphostemma
wallichii 29.9 (29-31) 31.7 (31-33) 0.95 (0.9-1.0) 3 Su Sc 75, 76 >
~0
TJ
fiE?
Haplostachys haplostachya 51.0 (44-57) 48.4 (46-51) 1.06 (0.9-1.2) 3,4 Su Sc 51,52
Haplostachys linearifolia 49.7 (46-55) 46.2 (44-48) 1.08 (1.0-1.2) 3,4 Su Sc 53,54 8 S >
Hypogomphia turkestana 33.4 (29-40) 27.4 (24-29) 1.22 (1.0-1.5) 3 Simple Su Sc 55-57 ^
CD
o)
3 o
o-
Lagochilus aucheri 25.5 (24-26) 24.2 (22-26) 1.06 (1.0-1.2) 3 Su 58,59
•
3 0°
T3
Lagochilus diacanthophyllus 28.2 (26-29) 22.7 (20-24) 1.25 (1.1-1.4) 3 Su Sc 60,61
O
o
9.
Lagochilus hirtus 25.5 (22-29) 21.1 (20-24) 1.22 (0.9-1.3) 3 Su Sc
^ D
CD
Ru+Ps
Lagopsis marrubiastrum 21.2 (20-23) 19.6 (18-21) 1.08 (0.9-1.2) 3 Np? 77 r
Lagopsis supina 21.1 (20-22) 21.8 (20-24) 0.97 (0.9-1.1) 3 ? ? o
5"
•
Lamiophlomis rotata 24.2 (22-26) 20.0 (18-22) 1.21 (1.1-1.3) 3 Ps Sc 78,79
^K »
Lamium album 25.1 (22-29) 23.8 (22-26) 1.06 (0.8-1.3) 3 Gr Sc 80,81
3
Lamium o
fiexuosum 23.5 (22-26) 21.8 (18-24) 1.09 (0.9-1.3) 3 Simple Su Sc 82,83
Lamium 5.
galeobdolon 30.8 (29-33) 22.9 (20-26) 1.36 (1.1-1.6) 3 Su Sc 86,87
CD
Lamium garganicum 28.6 (26-33) 26.8 (24-29) 1.07 (1.0-1.2) 3 Simple Su Mi 84,85 0)
CD
Lamium
moluccellifolium 34.1 (33-35) 30.1 (26-33) 1.14 (1.0-1.3) 3 Su 9
0>
•
—J
Lamium moschatum 21.2 (20-22) 20.6 (20-22) 1.03 (0.9-1.1) 3 Gr Sc 88,89 Q.
Lamium purpureum
3 Su Sc 90,91
Leonotis bequaertii 30.7 (29-33) 29.9 (28-33) 1.02 (0.8-1.2) 3 Su Sc 92,93
Leonotis leonitis 28.6 (26-31) 24.9 (24-26) 1.15 (1.1-1.3) 3 Simple Su Sc 94-96
III
Leonotis mollissima 31.9 (29-35) 28.6 (22-33) 1.13 (0.9-1.5) 3 Simple Su Sc 97 ^4
Table Continued.
2.
658
P/E Sculpturing
Polar axis (/mi) Equatorial axis (/mi) ratio
Species Mean Range Mean Range Mean Range Colpi Columellae Upper Lower Figures
Su Sc 98,99
Leonurus cardiaca 22.7 (22-24) 21.1 (20-24) 1.08 (1.0-1.1) 3
Leonurus 25.5 (22-29) 21.3 (20-24) 1.20 (1.0-1.4) 3 Su Sc 100
sibiricus
Leucas abyssinica 20.5 (18-22) 19.1 (18-20) 1.07 (1.0-1.1) 3 Su Sc 107, 108
Leucas alluaudii 26.0 (22-31) 22.9 (20-26) 1.14 (1.0-1.4) 3 Su Sc 109
Leucas aspera 24.0 (22-26) 19.6 (18-22) 1.23 (1.1-1.4) 3 Su Sc/Mi 110, 111
Leucas 29.3 (26-31) 23.1 (22-26) 1.27 (1.1-1.4) 3 Su Sc
biflora
Leucas calostachys 25.3 (24-26) 22.4 (20-24) 1.14 (1.0-1.3) 3 Su Sc 112, 113
Leucas 23.3 (22-24) 22.7 (22-24) 1.03 (0.9-1.1) 3 Su Sc
ciliata
Su Mi 114
Leucas eriostoma 20.9 (20-22) 17.6 (15-20) 1.20 (1.1-1.4) 3
Leucas hirta 20.5 (20-22) 16.7 (15-20) 1.24 (1.0-1.4) 3 Su Sc 115
Leucas 20.0 (-) 18.1 (17-20) 1.10 (1.0-1.1) 3 Su? ? 116
inflata
i
Leucas javanica 26.4 (22-31) 24.2 (22-26) 1.09 (0.9-1.3) 3 Su Sc/Mi 117
Leucas lanata 27.3 (26-29) 20.5 (18-22) 1.34 (1.2-1.5) 3 Su Sc
Leucas lavandulifolia 26.4 (24-29) 22.7 (20-24) 1.17 (1.1-1.3) 3 Su Sc
Leucas marrubioides 22.9 (20-24) 17.8 (15-20) 1.29 (1.0-1.4) 3 9 ?
1
Leucas martinicensis 22.6 (22-24) 20.1 (19-22) 1.13 (1.0-1.2) 3 Su Sc
Leucas mollissima 29.6 (29-31) 23.0 (22-24) 1.29 (1.2-1.4) 3 ? ?
• * 3
at
Leucas rosmarinifolia 22.2 (20-24) 18.5 (18-20) 1.21 (1.1-1.4) 3 Simple Su 9 118
8
• i.
Leucas stelligera 21.1 (20-24) 18.9 (18-20) 1.12 (1.0-1.3) 3 Su Sc
Loxocalyx ambiguus 23.8 (22-26) 21.3 (20-22) 1.12 (1.0-1.3) 3 Su Sc/Mi 101
~
03
O
Loxocalyx 24.0 (22-26) 20.9 (20-24) 1.16 (0.9-1.3) 3 Su Sc 102, 103 17
urticifolius
IT?" ft)
Mac
bridea alba 47.3 (42-53) 34.5 (31-37) 1.37 (1.2-1.5) 3 Su Sc 104 3
o'
Macbridea caroliniana 38.1 (35-42) 40.5 (40-42) 0.94 (0.9-1.1) 3 Su Sc 105, 106
9L
Marrubium anisodon 27.5 (24-31) 29.0 (26-31) 0.95 (0.8-1.1) 4 Ps Sc 122, 123 O
if
Marrubium cuneatum 24.9 (24-26) 26.8 (24-29) 0.93 (0.8-1.1) 3 Ps Sc 0)
s.
Marrubium heterodon 26.4 (24-29) 27.7 (26-29) 0.95 (0.8-1.1) 3 Ps Sc/Pu 124, 125
CD
3
Marrubium incanum 52.3 (51-55) 46.9 (44-51) 1.12 (1.1-1.2) 3 Ps Sc/Pu
Marrubium supinum 24.0 (-) 25.9 (24-26) 0.93 (0.9-1.0) 3 Simple Ps Sc 126-128
Melittis melissophyllum 45.8 (40-51) 35.6 (31-40) 1.29 (1.2-1.5) 3 Su Sc 119, 120
Metastachydium sagittatum 22.2 (22-24) 23.1 (22-24) 0.96 (0.9-1.1) 3 Su Sc 121
Microtoena delavayi 3 Simple Su Sc/Mi 129-131
Microtoena insuavis 3 Su Mi 132
Microtoena robusta 32.8 (29-37) 28.8 (24-33) 1.15 (0.9-1.4) 3 Su Mi 133, 134
Microtoena 24.6 (22-26) 28.2 (24-31) 0.88 (0.8-0.9) 3 Su Mi 135, 136
urticifblia
Su Sc/Mi 137
34.8 (33-40) 28.2 (26-31) 1.24 (1.1-1.5) 3
iYfofui-i t'f/u laei
i
i J
994
olume
Table
Continued. oo
2.
Polar axis (/mi) Equatorial axis (/mi) P/E
ratio Sculpt uring
'.
Species Mean Range Mean Range Mean dumber
Range
Colpi Colun Upper Lower
nellae Figures
*
Moluccella spinosa 29.5 (29-31) 30.4 (29-31) 0.97 (0.9-1.1) 3 Su Sc 138, 139
Notochaete hamosa
30.1 (26-33) 29.3 (26-31)
1.03 (0.8-1.2) 3 Ps Sc 140, 141
Utostegia
fruticosa 21.8 (20-22) 21 (20-22)
1.04 (1.0-1.1) 3 Su
Sc 142, 143
Utostegia
integrifolia 29.0 (26-33) 23.8 (22-26) 1.23 (1.0-1.4) 3 Su Sc/Mi 144
Utostegia minuccu
24.2 (22-26) 21.1 (20-22)
1.15 (1.0-1.2) 3 Su
Sc
Utostegia refxinda 20.9 (20-22) 17.2 (15-18)
1.22 (1.1-1.4) 3 Su
Sc 145
Utostegia tomentosa
21.1 (18-22) 18.3 (18-20) 1.16 (1.0-1.3) 3 Simple Su Sc 146, 147
!
anzenna
lanata
/ 23.3 (22-26) 19.4 (18-22)
1.21 (1.0-1.5) 3 Su Sc 148-150
aralammm
gracile
/ 16.3 (15-20) 15.0 (13-18) 1.10 (0.9-1.5) 3 Su
Sc 151
araphlomis javanica
/ 30.1 (29-33) 32.8 (29-35) 0.92 (0.8-1.0) 3
Sr Sc 152, 153
arapfuonus
/ lanceolala 26.8 (26-29) 26.0 (24-29) 1.04 (1.0-1.2) 3 Su Sc/Mi
154, 155
It
1 Iaraphlomifst rugosa 9
/ 30.0 (26-37) 31.2 (29-37) 0.96 (0.8-1.1) 3 Su
Sc 156
Phlomidoschema parvijlorum 24.2 (22-26) 21.6 (18-26) 1.14 (1.0-1.4) 3
Ps Sc 157, 158
V h 1
onus agraria
/ til 29.5 (29-31) 25.7 (24-29) 1.15 (1.0-1.3) 3 Su Sc 160 Pollen
159,
Pogost Abu-As
hlomis
/ crinita 33.9 (33-35) 25.1 (22-26) 1.36 (1.3-1.5) 3 Su ?
^
•
I'hlomis hcrfm-i enti 34.1 (31-37) 28.4 (26-31) 1.20 (1.1-1.4) 3 ? ? CD 0)
3 o
• * o-
I*hlomis lanata 40.3 (37-44) 29.0 (26-33) 1.39 (1.2-1.6) 3 Su
Sc 161, 162
/ hlomis maximoviczii 20.9 (20-22) 18.9 (18-20) 1.11 (1.0-1.3) 3 Gr o O
Sc 163 2.
/ hlomis tuberosa 27.3 (24-29) 25.5 (24-26) 1.07 (1.0-1.2) 3 Su logy
Sc 164
deae
hlomis umbrosa antin
/ 19.6 (18-22) 16.1 (15-18) 1.22 (1.1-1.3) 3 Gr Sc/Mi
165, 166
— ^ o
-^^ ^^- y -^^- -^h^- 5"
rhyllt'Strgia grandiflora 42.5 (40-44) 45.8 (44-48) 0.93 (0.8-1.0) 3 Ps Re
167, 168
Ph
yllostegia hirsuta 39.8 (37-44) 41.1 (35-46) 0.97 (0.9-1.1) 3 Su
Sc 170
169, to
3
I'hvllostrgia hispida 35 (33-37) 36.3 (35-37) 0.96 (0.9-1.1) 3 Su
Sc
171, 172
o
Ph
yllostegia lantanoides 36.3 (35-37) 38.1 (37-40) 0.95 (0.9-1.0) 3 Su
Sc 173
Pkyllostegia ra< emosa 33.2 (33-35) 36.1 (33-37) 0.92 (0.8-1.0) 3 Su CD
Sc 174
0)
Ph\ sostrgia longisepala 55.9 (53-59) 48.4 (46-52) 1.16 (1.1-1.3) 3 Su CD
Sc 175,*i1i76
» \^ v^
Ph • 0)
vsostcgia pnlt hclla 45.8 (43-48) 42.3 (36-46) 1.09 (1.0-1.3) 3 Su 3
Sc
Q.
Pro sium majus 35.9 (33-40) 30.6 (26-44) 1.19 (0.9-1.4) 3 Su
Sc 177
659
o
Table Continued.
2.
Polar axis (jim) Equatorial axis 0*m) P/E ratio Sculpturing
Mean Range Mean Range Mean Range Colpi Columellae Up,>er Lower Figures
Species
179
Su Sc 178,
Pseuderemostachys sewertzowii 37.0 (31-42) 29.0 (26-31) 1.28 (1.0-1.5) 3
3 Su Sc/Mi 180, 181
Roylea cinerea 22.2 (20-24) 22.0 (20-24) 1.02 (0.8-1.2)
183
31.7 (31-33) 35.0 (33-37) 0.91 (0.8-1.0) 4 Ps Sc 182,
canariensis
Sideritis
184
4 Ps Sc
44.0 (42-46) 40.3 (35-44) 1.10 (1.0-1.3)
candicans
Sideritis
192
(-) 6 Ps Sc 191,
30.4 (29-33) 30.4 (29-33) 1.00
Sideritis chlorostegia
185
4 Ps Sc
26.6 (24-29) 22.2 (20-24) 1.20 (1.1-1.3)
curvidens
Sideritis
193
(-) 6 Ps Sc
euboea 31.7 (29-33) 31.7 (29-33) 1.00
Sideritis
4 Ps Sc
gomerae 36.3 (35-40) 33.9 (31-37) 1.08 (0.9-1.2)
Sideritis
4 Ps Sc
31.9 (31-33) 29.7 (26-33) 1.08 (1.0-1.2)
Sideritis hirsuta
194
(-) 6 Ps Sc
Sideritis hololeuca 31.5 (29-33) 31.5 (29-33) 1.00
27.5 (24-31) 25.5 (22-29) 1.09 (0.9-1.4) 4 Ps Sc 186
Sideritis hyssopifolia
Sc/Pu 187
4 Ps
32.8 (31-35) 24.9 (22-29) 1.33 (1.2-1.5)
Sideritis folia
ilici
incana 26.2 (24-29) 24.6 (24-26) 1.06 (1.0-1.2) 4 Ps Sc
Sideritis
188
lagascana 31.2 (29-33) 24.0 (22-26) 1.31 (1.1-1.4) 4 Simple Ps Sc
Sideritis
lanata 29.7 (29-33) 28 (26-31) 1.07 (0.9-1.6) 4 Ps Sc
Sideritis o
=.
28.6 (26-31) 28.6 (26-31) 1.00 (-) 6 Ps Sc
Sideritis libanotica
O
marshchalliana 30.5 (29-33) 30.5 (29-33) 1.00 (-) 6 Ps Sc
Sideritis
CD
0)
montana 26.4 (24-29) 25.7 (20-29) 1.03 (0.9-1.3) 3 Ps Sc+Fo 195, 196 3
Sideritis
O
6 Ps Sc
pullulans 35.2 (31-40) 35.2 (33-37) 1.00 (0.9-1.1)
Sideritis
189
romana 26.0 (24-29) 21.3 (20-24) 1.22 (1.1-1.3) 4 Ps Sc
Sideritis
30.7 (29-33) 28.4 (26-31) 1.07 (0.9-1.2) 4 Ps Sc 190
Sideritis villosa
2.
197-199
Stachyopsis oblongata 23.3 (22-24) 21.6 (20-24) 1.09 (0.9-1.2) 3 Simple Su Sc
CD
3
3 Su Sc 200, 201
Stachys
riddellii
202-204
Stachys sylvatica 28.2 (26-31) 24.4 (24-26) 1.15 (1.1-1.3) 3 Simple Su Sc
206
Stenogyne haliakalae 39.6 (35-44) 36.5 (33-42) 1.09 (1.0-1.2) 3 Branched? Su Sc 205,
208
Stenosyne kamehamehae 45.8 (44-48) 41.4 (40-44) 1.11 (1.0-1.2) 3 Su Sc 207,
Su Sc 209, 210
Stenogyne purpurea 38.5 (33-44) 31.7 (28-35) 1.22 (1.0-1.4) 3 Simple
21.6 (20-22) 20.0 (19-22) 1.08 (0.9-1.1) 3 Su Sc 211
Sulaimarua otostcgioides
»
2 c
3
Tahlk
Continued. CD
2.
00
Polar axis (jim) Equatorial axis (/*m) P/E ratio
Sculpturing
Species Mean Range Mean 3
Range Mean
Range
Colp Columellae Upper Lower
i Figures
Suzukm
luchuensU
27.1 (24-29) 24.2 (22-26) 1.12 (1.0 -1.3) 3 Su
Sc 212,213
Synandra
hispidula
40.3 (39-42) 30.8 (29-33)
1.31 (1.2 -1.4) 3 Simple Su Sc 214,215
Thuspananta
persica 39.6 (33-42) 29.7 (24-35)
1.35 (1.0 -1.6) 3 Su Sc 216,217
m
a
iede a
11 itni orientttiis 25.1 (24-29) 18.0 (15-20) 1.40 (1.2 -1.7) 3 Su Sc 218,219
Subfamily Pogostemonoideae
CoUbrookta
oppositifblia 18.7 (18- 22) 18.9 (15 20) 0.99
(0.9 1.3) 3 Simple Su Sc 240, 241
Comanthosphace
japonica
23.5 (22- 26) 19.4 (18- 22) 1.22
(1.0 1-4) 3 Su Sc 242-244
Comanthosphace
stcllipila 24.2 (20- 26) 20.0 (18 22) 1.23
(0.9 1.5) 3 Su Sc
Comanthosphact
subianceoiata 22.0
(20- 24) 18.5 (15 22) 1.20
(1.0 1.4) 3 Su Sc
Leucosccptrum canum
27.1 (24- 29) 24.9 (24 26) 1.09
(1.0 1.2) 3 Simple Su Sc 246
245,
Pogostcmon
brachystach vus 26.0
(24 29) 23.1 (22 24) 1.12 (1.1 1.3) 3 Su Sc 228
227,
Pogostcmon
vablin 26.0 (- ") 24.2 (22 26) 1.09
(1.0 1.2) 3 Su Sc 229, 230
>
Pogostcmon
CTUCJatUS 24.9
(24 26) 26.4 (24 29) 0.95
(0.9 1.0) 3 Su Mi
231 O
& cr
Pogostcmon c
giaber 24.9 (20 29) 19.4 (18 22) 1.30
(0.9 1.5) 3 Su Mi 232, 233 o
5
>
Pogostcmon hevneanus
27.3 (26 29) 20.2 (18 22) 1.35 (1.2 1.6) 3 Su Sc
m c/>
Pogostemon
vosuroulcs 23.8 (20 24) 17.4 (15 20) 1.37 (1.1 1.6) 3 Su Mi 234, 235 C3D o
a-
Pogostcmon
plcctrantlioulcs 27.1 (24 29) 23.5 (20 26) 1.16 9-3
(1.1 1.2) 3 Su Mi 237
236,
Pogostcmon sampsonii 24.0 (- -) 22.0 (- -) 1.10 (- 3 Su O
?
)
Pogostcmon n
stcHut us 18.7 (15 22) 16.5 (15 18) 1.15 (0.9 1.4) 3 Su ? 9- 0*)»
Pifg < Uetnon yatn hen n us 24.2 (22 26) 18.7 (15 22) 1.31 (1.0 1.7) 3 Su Mi 238, 239 "<
O
Rostrtnm dependms
uln
23.1 (22 26) 21.8 (20 22) 1.06
(1.0 1.2) 3 Su Sc 247
Genera
of uncertain affinities
3
Anisomeles h<\ neana 27.0 (24-29) o
25.1 (22 29) 1.09
(0.9 1.3) 3 Su Mi 220
a
Anisomeles indica 31.2 (29-35)
32.1 (29 35) 0.98
(0.8 1.2) 3 Su Mi 221,222
CD
Anisomeles main horn a 35.2 (33-40) 34.1 (31 37) 1.04 (0.9 1.3) 3 Su Mi 223 Q>
CD
Anisomeles suliu folia 28.2 (26-31) 25.5
(24 26) 1.11 (1.0 1.3) 3 Su Mi
224, 225
Eur\ tolen gracilis 23.8 (22-26)
18.9 (18 20) 1.26
(1.1 1.5) 3 Simple Su
Sc 226
CJ)
662 Annals of the
Garden
Missouri Botanical
rest of the grain scrobiculate); columellae simple reticulate sculpturing and simple columellae. All of
1976). the Pogostemonoideae, most Lamioideae, and
(Nabli, the
two genera of uncertain affinities exhibit these
ple-
POGOSTEMONOIDEAE siomorphic conditions, but the following
derived
occur some Lamioideae:
states in tetracolpate and
Pollen of subfamily Pogostemonoideae (Figs.
6-pantocolpate pollen, branched columellae, and
227-247)
oblate spheroidal to euprolate, with
is
four forms of sculpturing (psilate, granulate, ru-
15-29
and
the polar axis Jim the equatorial axis
and
gulate, suprareticulate-rugulate).
15-29 The and
/im. grains are tricolpate inoper-
The
culate. exine scrobiculate or microreticulate,
is
NUMBER OF APERTURES
with suprareticulate sculpturing (poorly developed
in Rostrinucula, approaching a psilate condition). Within the study group, hexapantocolpate pol-
In some species of Pogostemon (Figs. 231, 235; len (Figs. 191, 194) was found in Sideritis sect.
Pozhidaev, 1989), the lumina contain one or more Empedoclea. Elsewhere in the family, this feature
large central perforations surrounded by small ones. has been reported in only three species (tradition-
The columellae are unbranched. ally assigned to the Verbenaceae but treated here
Car
as Labiatae, following Cantino et 1992):
al.,
Discussion yopteris nepalensis Moldenke, Clerodendrum a-
&
trinum and Huxleya Ewart B.
Ridley, linifolia
Pollen morphology provides no
distinction be-
Rees (Abu-Asab 1993). These species are
et
al.,
tween subfamilies Lamioideae and Pogostemonoi-
Lamioideae
only distantly related to subfamily
deae, the variation within the former encompassing
(Cantino, 1992a). Because 6-pantocolpate pollen
that found in the latter. At a lower taxonomic level,
can be
1992a),
a derived condition (Cantino, it
is
similar pollen features suggest relationships within
synapomorphy of Sideritis
treated as a putative
and between certain genera. In some cases, these
congruence
Empedoclea pending a for
sect. test
similarities are thought to be derived, based on a
we examined only
Although
with other characters.
previous more
cladistic analysis of a inclusive study
many others
Huynh
(1972) studied
five species,
group (Cantino, 1992a; see below), and can there-
within the
and found this feature to be constant
fore be cited as putative synapomorphies (de Pinna,
genus.
section and absent elsewhere in the
1991) until they are tested for congruence with
were ob-
derived state)
Tetracolpate grains (a
other characters through a parsimony
analysis.
Marrubium anisodon,
served Haplostachys,
in
Other
features are not clearly ancestral or derived
Haplostachys,
many
and species of Sideritis. In
and
therefore contribute only to an assessment of
where-
the colpi are arranged in two pairs (Fig. 53),
phenetic
relationship.
(rigs.
spaced Sideritis
as the colpi are equally in
Based on outgroup comparison
to the Scroph- Mar
arrangement in
182, 186, 189); the colpal
we &
ulariales, argued (Abu-Asab
Cantino, 1992)
In
rubium anisodon unclear (Fig. 122).
is
that the following pollen character states are ple-
com-
a clade
tetracolpate pollen delimits
eritis,
siomorphic
in the Labiatae: three apertures, ab-
1972).
(Huynh,
prising four of the six sections
sence of operculum,
simple columellae,
psilate ex-
tricolpate
Haplostachys,
We both examined species of
ine. further hypothesized that suprareticulate
sam
the
and tetracolpate grains were found in
sculpturing a synapomorphy
is of a large clade
may be a relia
not
presence
flower, thus their
comprising
the gynobasic
-styled Labiatae (includ-
feature of the genus.
ing all taxa under study here), some of the genera
of Labiatae
that lack a gynobasic style, and a few
COLUMELLAR STRUCTURE
genera
traditionally assigned to the Verbenaceae.
foun^
and
This hypothesis was
corroborated by plesiomorphic
a cladistic Simple columellae are
85 (Abu-Asa
analysis of characters (Cantino, 1992a), five of in most gynobasic -styled Labiatae
^
which pertained to pollen. Within the clade marked Cantino, 992). The only apparent excePt,on^
1
^^e.
by
suprareticulate pollen, subfamilies Lamioideae are aware of within the Lamioideae or
and o
Pogostemonoideae occupy Gomphostemma
a relatively derived monoideae are (Figs, ,
Chelonop^
position. Therefore, although
suprareticulate sculp- Bostrychanthera (Pozhidaev, 1989),
Stea0&^
turing apomorphic
is in the Labiatae as a whole, and perhaps
lichiangensis (Fig. 32),
it is plesiomorphic within the groups of concern haliakalae Wawra 206). In the lalter ^
(Fig.
efS
here
(Cantino, 1992a). Thus, the plesiomorphic seem branch apically while ° ^
columellae
to
e
pollen type from
in subfamilies Lamioideae and unclear
Pogoste- appear to be simple. It is *
trUC.
monoideae branched s r
is tricolpate, inoperculate, with supra- available photographs whether the