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Phylogenetic Relationships in Artemisia spp. (Asteraceae) Based on Distribution of Foliar Trichomes


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The phylogenetic relationships within the genus Artemisia have been very controversial and need throughout investigations. In continuation to the previous paper, here we report the phylogenetic relationships in the genus Artemisia based on the foliar trichomes using light microscopy and scanning electron microscopy. The data from 24 taxa was analyzed for its phylogeny. In addition, eight new types of foliar trichomes in the genus were described. This study also revealed that foliar trichomes of the genus Artemisia are good taxonomic markers and can be utilized to resolve the taxonomic conflicts within the genus.
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To cite this paper: Hayat, M.Q., M. Ashraf M.A. Khan, G. Yasmin, N. Shaheen and S. Jabeen, 2009. Phylogenetic relationships in Artemisia spp.
(Asteraceae) based on distribution of foliar trichomes. Int. J. Agric. Biol., 11: 553–558
Phylogenetic Relationships in Artemisia spp. (Asteraceae) Based
on Distribution of Foliar Trichomes
Department of Plant Sciences, Quaid-i-Azam University, Islamabad, Pakistan
NUST Center of Virology and Immunology, National University of Science and Technology, Rawalpindi, Pakistan
National Center of Excellence in Geology, University of Peshawar, Peshawar, Pakistan
1Corresponding author’s e-mail:
The phylogenetic relationships within the genus Artemisia have been very controversial and need throughout investigations. In
continuation to the previous paper, here we report the phylogenetic relationships in the genus Artemisia based on the foliar
trichomes using light microscopy and scanning electron microscopy. The data from 24 taxa was analyzed for its phylogeny. In
addition, eight new types of foliar trichomes in the genus were described. This study also revealed that foliar trichomes of the
genus Artemisia are good taxonomic markers and can be utilized to resolve the taxonomic conflicts within the genus.
Key Words: Artemisia; Seriphidium; Anthemideae; Asteraceae; Anatomy; Trichomes; Phylogeny
Artemisia L. is one of the largest genera of the family
Asteraceae. Its members are wind pollinated and mainly
distributed in temperate areas of mid to high latitudes of the
northern hemisphere, colonize in arid and semiarid
environments, and only few representatives are found in
southern hemisphere (McArthur & Plummer, 1978; Valles
& McArthur, 2001). Many species of the genus have
economic value as medicines, food, forage, ornamentals or
soil stabilizers in disturb habitats; some taxa are toxic or
allergenic and some others are invasive weeds, which can
adversely affect harvests (Pareto, 1985; Tan et al., 1998;
Hayat et al., 2009a & b). Except few annuals or biannual,
most Artemisia species are perennial (Valles et al., 2003).
The genus Artemisia is also considered as indicator of
steppe climate (Erdtman, 1952) and moderate precipitation
(El-Moslimany, 1990).
Classically the genus has been divided into four
groups, which has been treated as sections or subgenera;
Absinthium (Tournefort) de Cand., Artemisia Tournefort
(=Abrotanum Besser), Dracunculus Besser, Seriphidium
Besser (Hooker, 1881; Martin et al., 2001 & 2003).
McArthur et al. (1981) proposed a new group Tridantatae
(Rydb.) McArthur, which is endemic to North America.
Ling (1982, 1991a & b, 1995a & b) separated Seriphidium
(Besser ex Hooker) as independent genus. Bremer and
Humphries (1993) and Bremer (1994) accepted this
separation. But Kornkven et al. (1998), Torrell et al. (1999)
and Watson et al. (2002) in their molecular studies again
merged Seriphidium with Artemisia. Nonetheless
classification of Artemisia and relationships among its
different taxa are still controversial.
Scotland et al. (2003) opined that rigorous and critical
anatomical studied of fewer morphological characters in the
context of molecular phylogenies is fruitful to integrating
the strengths of morphological data with those of sequence
data. Since the micromorphological characteristics of foliar
trichomes play an important role in plant taxonomy,
especially of particular groups at generic and specific levels,
attention of plant morphologists and systematists has been
mainly attracted to resolve the taxonomic conflicts (Hardin,
1979; Fang & Fan, 1993). Although many studies
conducted on the histochemistry of the secretory products of
the glandular trichomes of Artemisia (Smith & Kreitner,
1983; Slone & Kelsey, 1985; Ascensao & Pais, 1987; Duke
& Paul, 1993; Duke et al., 1994), little is known about the
systematic significance of trichomes in Artemisia. In the
present paper, we report the micromorphological
characteristics of foliar trichomes in Artemisia using light
microscopy (LM) and scanning electron microscopy (SEM).
The specific objectives were to: (a) identify and compare the
micromorphological characteristics of foliar trichomes in
different species of this genus and (b) reconstruct the
phylogenetic relationships among different taxa of
Artemisia based on characteristics of foliar trichomes.
All details about the origin and collection of leaf
material, processing of the tissues, basic terminology and
studies of the foliar trichomes were essentially the same as
HAYAT et al. / Int. J. Agric. Biol., Vol. 11, No. 5, 2009
we reported in the previous paper (Hayat et al., 2009a).
Source and collection information of remaining taxa is
given in Table I. Using modified methodology of Shaheen
et al. (2009) and Yasmin et al. (2009) foliar trichomes were
at first examined by OLYMPUS/BX-51 light microscope.
Pieces of leaves were dipped in 30% Nitric acid and boiled
along with 1.5 g of Potassium chloride in a test tube for 2-3
min. Then these leaf pieces were cleaned with distal water.
Epidermis was removed and kept in 60% Potassium
hydroxide solution for 2 h. Finally, these leaf sections were
suspended in Lactic acid and transferred on glass slides for
LM observations. For SEM investigations, the dried leaves
were mounted on aluminum stubs with the help double
adhesive tape, sputter coated with gold by SPI-Module
Sputter Coate and studied with a Jeol-JSM 5910 scanning
electron microscope. Basic terminology used for trichomes
categorization and elaboration was that recommended by
Ramayya (1972), Payne (1978), Bento et al. (2008), Popa
and Sipos (2009) and Shaheen et al. (2009). However,
straightforward self illustrative terms are incorporated to
recognize the particular type of trichome.
On the bases of presence or absence, all 16 foliar
trichomes types, noted here, were selected as a character
states for phylogenetic analysis of Artemisia (Table II). The
plesiomorphic or apomorphic state of each trichome was
determined using the criteria established by Boudreaux
(1979) and further explained by Crisci and Stuessy (1980).
In this study an imaginary outgroup was used for
comparison that contained all the ancestral characters. An
original binary data matrix was produced (Table III) for
phylogenetic analysis using the outgroup comparison
method (Watrous & Wheeler, 1981). The phylogenetic
analysis was done by PHYLIP computer program version
3.67 (Felsenstein, 2007). Most parsimonious trees (MPTs)
based on the binary matrix were constructed with MIX
program of PHYLIP using Wagner parsimony method
(Farris, 1970). A strict consensus phylogenetic tree of the
MPTs was generated using the CONSENSE program of
PHYLIP (Sokal & Rohlf, 1981). Finally, using
strict consensus phylogenetic tree of Artemisia were
Types of foliar trichomes. Based on observations of LM
(Fig. 1) and SEM (Fig. 2), the foliar trichomes in 24 taxa
from Artemisia were classified into 16 main types. Of these
a-h were described in Hayat et al. (2009a), i-p are
illustrated below:
(i). Vase-shape trichomes. These are glandular
trichomes. They are cylindrical in shape with constricted
apex and are characteristic of A. persica (Fig. 1A & Fig.
(j). Aduncate curly trichomes. These are also non-
glandular trichomes. They are long cylindrical in shape.
They are characteristic feature of A. roxburghiana (Fig. 1B).
(k). Dolabrate trichomes. They have shape like the head
of pick, with two divaricated or opposed terminal branches.
These non-glandular trichomes are found in A. vulgaris
(Fig. 1C).
(l). Unicellular peltate trichomes. These glandular
trichome are embedded in epidermis and are the
characteristic feature of A. vulgaris (Fig. 1D), A.
roxburghiana and A. japonica.
(m). Rope rolled trichomes. These are nonglandular
trichomes. They are exclusive characteristic feature of A.
moorcroftiana (Fig. 2B).
Fig. 1. Types of glandular and nonglandular foliar
trichomes in Artemisia by means of LM: A, A. persica;
B, A. roxburghiana; C-D, A. vulgaris (Scale bar = 50
Fig. 2. Types of glandular and nonglandular foliar
trichomes in Artemisia by means of SEM: A, A. persica
(Scale bar = 10 µm); B, A. moorcroftiana (Scale bar =
10 µm); C, S. kurramense (Scale bar = 10 µm); D, A.
stricta (Scale bar = 1 µm)
PHYLOGENY OF FOLIAR TRICHOMES IN GENUS Artemisia / Int. J. Agric. Biol., Vol. 11, No. 5, 2009
(n). Cylindrical trichomes. They are cylindrical in shape.
These non-glandular trichomes are the unique feature of S.
kurramense (2C).
(o). Floriform peltate trichomes. These glandular
trichomes have flower like appearance and are present in A.
stricta (Fig. 2D).
(p). Cup-shape peltate trichomes. They are unicellular
cup like glandular structures with broad apical opening.
They are found in A. tangutica (Fig 3).
The quantitative dimensions of all the trichomes types
studied are given in Table IV and V.
Phylogenetic analysis. Hundred MPTs were generated
using Wagner parsimony method (Farris, 1970) and then a
strict consensus phylogenetic tree of these MPTs was
obtained (Fig. 4 & Fig. 5). In the resulting phylogenetic tree,
Artemisia and Seriphidium is shown to be a monophyletic
Features of trichomes are generally considered as
valuable for ascertaining the taxonomic relations within the
genus Artemisia (Hall & Clements, 1923). Different forms
are reported to occur in different taxa of Artemisia (Ferreira
& Janick, 1995). Different authors reported only two types
of trichomes in the genus Artemisia: a) capitate glandular
trichomes (Smith & Kreitner, 1983 in A. ludoviciana Nutt.;
Kelsey, 1984 in A. nova Nelson; Slone & Kelsey, 1985 in A.
tridentata Nutt.; Ascensao & Pais, 1987 in A. compestris L.;
Lodari et al., 1989 in A. princeps Pamp.; Ferreira & Janick,
1995 in A. annua) and b) T-shape nonglandular trichomes
(Ascensao & Pais, 1987 in A. compestris ; Lodari et al.,
1989 in A. princeps, A. absinthium, A. capillaris & A.
japonica; Ferreira & Janick, 1995 in A. annua). In this and
our previous paper (Hayat et al., 2009a), we reported not
only 16 new types of glandular and non-glandular foliar
trichomes in Artemisia but also un-veiled the under lying
variations in capitate and T-shape trichomes of the genus.
In addition to capitate and T-shape trichomes and
those described in Hayat et al. (2009a), here we also
observed eight new types of trichomes of taxonomic
importance. Of these, unicellular peltate trichomes of A.
vulgaris (Fig. 1D), A. roxburghiana and A. japonica, vase-
shape trichomes (Fig. 1A & 2A) of A. persica; aduncate
curly trichomes (Fig. 1B) of A. roxburghiana; dolabrate
trichomes (1C) of A. vulgaris; rope rolled trichomes (Fig.
Table I*. List of taxa studied for foliar anatomy and their herbarium vouchers. ISL: Herbarium, Quaid-i-Azam
University, Islamabad. PUP, Herbarium, University of Peshawar, Peshawar
Taxon Collection data Herbarium Voucher
Section Artemisia Tournefort
A. rutifolia Spreng. Gilgit: Nattar valley. A. Rashid, 1988. PUP, 244 (1105)
A. santolinifolia Turcz. ex Krasch. Gilgit: Nattar valley. A. Rashid, 1986. PUP, 239 (1108)
A. vestita Wall. ex DC. Jahlum: Soon vally, Sakasar. M. Farooq, 2005. ISL, 20093
Section Seriphidium (Besser) Besser
A. maritima L. Skardu: M. Q. Hayat, 2007. PUP, PH003 (ART003)
S. brevifolium (Wall. ex DC.) Ling & YR Ling Mansehra: Ujtar, Naran to Lalusar lake track. M. Q. Hayat, 2007. PUP, PH007 (ART007)
S. stenocephalum (Krasch. ex Poljakov) Poljakov Gilgit: Chalas. M. Q. Hayat, 2008. PUP, PH010 (ART010)
S. turanicum (Krasch.) Poljakov Gilgit: Nattar vally. M. Q. Hayat, 2008. PUP, PH009 (ART009)
Section Dracunculus Besser
A. scoparia Waldst. et Kit. Islamabad: Quaid-i-Azam university campus. M. Q. Hayat, 2008. ISL, 32313
A. stricta Edgew. Kashmir: Muzafrabad. T. Malik, 1972. ISL, 25650
*Partly adapted from Table I, Hayat et al. (2009a)
Table II. Character and character states of foliar
trichomes for the phylogenetic analysis of Artemisia
(the number in brackets represents the codes of
character states; plesiomorphic, 0; apomorphic, 1)
Charecter Character states
a. Capitate Present (0), Absent (1)
b. Pluricellular Absent (0), Present (1)
c. Multicellular peltate Absent (0), Present (1)
d. Thin neck Absent (0), Present (1)
e. T-shape Present (0), Absent (1)
f. Macroform Absent (0), Present (1)
g. Unicellular tector Absent (0), Present (1)
h. Clavate Absent (0), Present (1)
i. Vase-shape Absent (0), Present (1)
j. Aduncate curly Absent (0), Present (1)
k. Dolabrate Absent (0), Present (1)
l. Unicellular peltate Absent (0), Present (1)
m. Rope rolled Absent (0), Present (1)
n. Cylindrical Absent (0), Present (1)
o Floriform peltate Absent (0), Present (1)
p. Cup-shape peltate Absent (0), Present (1)
Fig. 3. Types of glandular foliar trichomes in Artemisia
by means of LM: A. tangutica (Scale bar = 50µm)
HAYAT et al. / Int. J. Agric. Biol., Vol. 11, No. 5, 2009
2B) of A. moorcroftiana; cylindrical trichomes (Fig. 2C) of
S. kurramense; floriform trichomes (Fig. 2D) of A. stricta
and cup-shape peltate trichomes (Fig. 3) of A. tangutica, are
important one.
He et al. (2000) resolved the taxonomic conflicts in
the genus Actinidia Lindl. and related taxa by following the
same methodology. In this paper, the results obtained from
the phylogenetic tree (Fig. 4 & Fig. 5) based on the
micromorphological characteristics of foliar trichomes
support the view presented based on molecular studies
(Kornkven et al., 1998; Torrell et al., 1999; Watson et al.,
2002) that Artemisia and Seriphidium are monophyletic
From cladogram (Fig. 4) and phylogenetic tree (Fig. 5)
it is clear that foliar trichome evolution in Artemisia starts
form outgroup (assumed as ancestor of Artemisia). The first
evolved group (1) consist of those taxa, which have both T-
shape and capitate trichomes. The second evolve group (2)
has only capitate trichome and third group (3) is without
trichomes. A possible tendency for foliar trichome evolution
is marked in Fig. 4 and Fig. 5. Also we support the idea
about general degenerate evolutionary tedency of
Asteraceae (Gailing & Bachmann, 2003). we do not believe
on old classification of the genus, because it was based only
on floral characters and has many objections; for example
section Artemisia only differ from section Absinthium by a
Table III. Data matrix used in phylogenetic analysis of Artemisia (Character & character states are described in
Table II)
Taxon Abbreviation Trichomes*
a b c d e f g h i j k l m n o p
Outgroup Out 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A. amygdalina Amy 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A. biennis Bie 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A. dubia Dub 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0
A. moorcroftiana Moo 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0
A. roxburghiana Rox 1 1 0 0 0 0 0 1 0 1 0 1 0 0 0 0
A. rutifolia Rut 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A. santolinifolia San 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A. tournefortiana Tou 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0
A. vestita Ves 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A. vulgaris Vul 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0
A. absinthium Abs 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A. macrocephala Mac 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A. persica Per 0 0 1 1 1 1 0 0 1 0 0 0 0 0 0 0
A. siversiana Siv 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A. tangutica Tan 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1
A. maritima Mar 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
S. leucotrichum Leu 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
S. brevifolium Bre 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
S. kurramense Kur 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0
S. stenocephalum Ste 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0
S. turanicum Tur 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
A. japonica Jap 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0
A. scoparia Sco 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0
A. stricta Str 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0
*a, capitate; b, Pluricellular; c, multicellular peltate; d, thin neck; e, T-shape; f, macroform; g, unicellular tector; h, clavate; i, vase-shape; j, aduncate curly;
k, dolabrate; l, unicellular peltate; m, rope rolled; n, cylindrical; o, floriform peltate; p, cup-shape paltate
Table IV. Quantitative characteristics of glandular foliar trichomes of Artemisia
Taxa Vase-shape Unicellular peltate Floriform peltate Cup-shape peltate
Height x width (µm) Diameter (µm) Diameter (µm) Diameter (µm)
A. roxburghiana Absent 10-14 Absent Absent
A. vulgaris Absent 10-15 Absent Absent
A. persica 97-101x45-50 Absent Absent Absent
A. tangutica Absent Absent Absent 20-25
A. japonica Absent 9-13 Absent Absent
A. stricta Absent Absent 4-5 Absent
Table V. Quantitative characteristics of nonglandular foliar trichomes of Artemisia
Taxa Aduncate Dolabrate Rope rolled Cylindrical
Height × width (µm) Height × width (µm) Height × width (µm) Height × width (µm)
A. moorcroftiana Absent Absent 200-250x45-50 Absent
A. roxburghiana 700-750x15-20 Absent Absent Absent
A. vulgaris Absent 25-30x10-15 Absent Absent
S. kurramense Absent Absent Absent 35-40x8-10
PHYLOGENY OF FOLIAR TRICHOMES IN GENUS Artemisia / Int. J. Agric. Biol., Vol. 11, No. 5, 2009
single character i.e., receptacle naked (Artemisia) or
receptacle cover with long hairs (Absinthium) (Kaul &
Bakshi, 1984). However, we believe that more taxa of the
genus Artemisia, which includes sufficient taxa of each
classical section, need to be investigated in more detail and
especially focusing on the micromorphological characters of
foliar trichomes integrated with molecular approaches.
In conclusion, diversity in glandular and non-glandular
foliar trichomes is a valuable taxonomic tool. Non-etheless,
there is a need to develop better terminology and detail
comparative study of these micromorphological features to
resolve taxonomic conflicts in the genus. Their careful
anatomical and ontological studies along with the
integration of molecular data one can portray the complete
picture about the infrageneric classification of Artemisia.
Acknowledgement. We are grateful to Dr. Riaz and Mr.
Abdullah Jan, Department of Physics and Dr. M. Tahir
Shah, NCE in Geology, University of Peshawar, Pakistan
for their facilitation. We also acknowledge HEC Pakistan
for their financial assistance.
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Fig. 4. The strict consensus cladogram of Artemisia
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trichomes, Letters represent the abbreviation of taxa
(see Table III)
Fig. 5. The strict consensus phylogenetic tree of
Artemisia based on the micromorphological characters
of foliar trichomes, Letters represent the abbreviation
of taxa (see Table III)
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(Received20 May 2009; Accepted 29 May 2009)
... Different types of trichomes (glandular and non-glandular) have been reported in different Artemisia species [146]. The most common types in Artemisia are capitate type trichomes reported by Kelsey [147], Slone and Kelsey [143], Ferreira and Janick [146], Ascensao and Pais [148], Lodari et al. [149], Hayat et al. [150], Hussain et al. [131] and references therein. Ascensao and Pais [148] observed capitate trichomes in A. compestris, Smith and Kreitner [151] in A. ludoviciana, Ferreira and Janick [146] in A. annua, Kelsey [147] in A. nova, Slone and Kelsey [143] in A. tridentatae and Lodari et al. [149] in A. princeps. ...
... Another attempt validated the foliar epidermal structure of A. annua and its biochemical profile [132]. Based on foliar trichomes micromorphological data, Hayat et al. [150] described relationships and evolutionary patterns within Artemisia. They analyzed 24 species and reported eight trichomes types in Artemisia species. ...
... The morphology of plants is very important to recognize their taxonomic position with cladidtics approach [161]. Previous inquiries on Artemisia have recommended that the morphology of Artemisia is very challenging to address due to similarities in species which causes problems in their identification and delimitation [150]. This complexity is manifested in A. vulgaris complex [177]. ...
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Artemisia is the largest genus in the tribe Anthemideae having ecologically, morphologically, and chemically diverse species. These species are found mostly in the Northern hemisphere, with the distribution of fewer taxa in the Southern hemisphere of the world. Species of this genus have great therapeutic potential and constitute a remarkable natural asset and are still utilized as folk remedies against different health-related problems all over the world. The taxonomy of Artemisia has long been problematic essentially due to the morphological complexities within its species. The recent classification divides the genus into six major groups like Absinthium DC., Artemisia L., Dracunculus Besser, Pacifica, Seriphidium Besser, and Tridantatae (Rydb.) McArthur. However, its infrageneric classification is still indistinguishable. The current review comprehensively enlightens the ethnomedicinal significance and recent advancements in the taxonomy based on foliar anatomy, pollen features, morphology, and molecular phylogeny of Artemisia to understand the classification of this economically significant genus.
... Peeling of epidermis was performed and 60% potassium hydroxide solution was used to keep the peel for 2 h. Finally, these peels were transferred to lactic acid and glass slides were prepared for LM investigation (Hayat et al., 2009b). ...
... The majority of nonglandular trichomes observed in this study were not previously been reported in Artemisia species except for the aduncate and unicellular tector non-glandular trichomes. However, we found aduncate curly trichomes in A. indica and A. dubia whereas other studies showed these types of trichomes in A. roxbhurgiana (Hayat et al., 2009b). Moreover, this study found unicellular tector tichomes in A. herba alba whereas, other studies disclosed these types of trichomes in A. dubia (Hayat et al., 2009a). ...
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This study for the first time examined foliar epidermal anatomical characteristics (Epidermal cells, types of stomata and trichomes) of thirteen Artemisia species (Including rare ones) with Light microscopy (LM) and Scanning electron microscopy (SEM) from the Northeast (Gilgit-Baltistan) region of Pakistan. The epidermal cells varied from polygonal to irregular and elongate in shape, while wavy to smooth in margins. This investigation revealed four different types of stomata viz; Anomocytic, diacytic, anomotetracytic and anisocytic, which were unequally scattered on both the adaxial and abaxial surfaces of studied Artemisia species. Ten different types of trichomes were observed in the studied species. Four types of glandular trichomes including peltate, pluricellular, capitate and thin necked were observed, whereas six types of non-glandular trichomes including, Aduncate, unicellular calavate, conical type, stinging hair type, unicellular tector and unicellurar filiform were observed. This study authenticated that the morphology of these foliar anatomical characteristics in amalgamation with other features are helpful for the species delimitation within the genus Artemisia.
... All these 38 species are recorded from the arid and semi-arid areas of Baluchistan, Khyber Pakhtunkhwa, North Punjab and the temperate areas of Gilgit-Baltistan and Kashmir territory (Ghafoor 2002). Within Pakistan, the centre of diversity for the genus is the western Himalayan region (Hayat et al. 2009). Hayat (2011) initiated the phylogenetic study of Pakistani Artemisia using ITS and ETS sequences of nrDNA and found support for uniting the two genera. ...
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Background-Gilgit-Baltistan, the Northeast region of Pakistan, is well known for its floristic diversity, including members of the genus Artemisia. Artemisia is a large, taxonomically complex genus including ~500 species of both herbs and shrubs. This study was conducted to determine the phylogenetic position of ten undescribed Artemisia taxa from northern Pakistan, using nrDNA internal transcribed spacer (ITS), external transcribed spacer (ETS) and cpDNA intergenic spacer (psbA-trnH) regions. Methods-The phylogenetic relationships of 28 taxa of Artemisia using separate and combined data sets of sequences of three markers (ITS, ETS and psbA-trnH) were analysed with maximum parsimony, maximum likelihood, and Bayesian approaches. Key results-The results resolve northeastern Pakistani Artemisia, which represent five morphologically defined subgenera, into ten major clades. Subgenera Artemisia and Absinthium are shown to be polyphyletic, while Dracunculus, Pacifica and Tridentatae appear monophyletic. All species of subgenus Seriphidium are retrieved in a single clade that also includes annual species from subgenus Artemisia. In the Flora of Pakistan, Seriphidium is described as a separate genus but in this study, Seriphidium fell within the genus Artemisia. In addition, on the basis of phylogenetic analysis, we present evidence that ten as-yet undescribed taxa are present in northeastern Pakistan based on newly recognized three groups (Groups I, II and III) of taxa within the genus Artemisia. One undescribed taxon from group I was placed within the subgenus Dracunculus clade and the remaining nine taxa from groups II and III were placed in the subgenus Absinthium clade. Morphological studies coupled with modern molecular techniques may lead to a new infrageneric classification of the genus Artemisia. It will also clarify and characterize the undescribed taxa reported in this study.
... Ferreira and Janick in 1995 reported the presence of T-shape trichomes in A. annua. Hayat (2009b;2009d plus references there in) reported 16 main types of the trichomes and range of the capitate glands, differences in T-shaped hairs and existence of supplementary types of foliar trichomes in Artemisia. Artemisinin content in A. annua varies between 0.01 to 0.4 % depending on the genotype, variety, season, cultivation condition, and plant developmental stage (Lommen et al. 2007;Delabays et al. 2001;Yang et al. 2009;Davies et al. 2009) and a few clones have shown production [1 % (Delabays et al. 2001). ...
Artemisia annua L. (sweet sagewort) belongs to genus Artemisia L. of family Asteraceae and tribe Anthemideae. Artemisia L. approximately comprises of about 500 taxa and considered as largest genus of Asteraceae. A. annua, which are now widely and naturally distributed in various regions of the world (temperate, subtropical zones, Asia), have originated from China and are introduced and naturalized in Canada and USA. later on. On the basis of fossil records and phytogeographic data, it has been discovered that the Quaternary was the main time period for the variegation and dispersal of the taxa worldwide. Globally, many countries are currently cultivating A. annua on a large industrial scale, as it is a key source of a potent antimalarial drug, artemisinin. In this chapter, we will set out its origin, geological distribution, and phytography emphasizing on important characters, for instance pollen as pollen morphology is considered as worthy systematic marker for genus Artemisia. Furthermore, special protuberant structures viz. glandular trichomes and their role in artemisinin and various other phytotoxic secondary metabolites will be addressed. The second part of the chapter will draw attention to the possibilities and methods used to solve mysteries of Artemisia systematics on subtribal, generic, and subgeneric level over time. To date, for systematic studies at all taxonomic levels, none of the techniques alone is considered sufficient. In conclusion, the perspectives and issues for the further study of the plant and genus will be raised. © 2014 Springer-Verlag Berlin Heidelberg. All rights are reserved.
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Artemisia chinensis L. referred as Crossostephium chinensis (L.) Makino in the Flora of China is a rare and conceivably threatened plant species with an unclear origin in Asia. The species has been acknowledged so far from some islands of Taiwan. However, as it is extensively cultivated for ornamental and medicinal purposes in Japan, China, and the Philippines, it is still challenging to delimit its native range. This study confirms the presence of A. chinensis from Northern (Gilgit-Baltistan) Pakistan using molecular phylogenetic analysis and by assessing its distribution. The species were found in one site in the Skardu District of GB Pakistan and phylogenetic analysis indicated a close resemblance of the collected A. chinensis from the Skardu region with species of subgenus Pacifica of the genus Artemisia reported globally. According to the outcomes of the present study, it is proposed that broader field surveys should be conducted to acknowledge the distribution of A. chinensis plant from other districts of GB and cities of Pakistan as well. It is proposed that A. chinensis plant is present in North Pakistan and this plant should be mentioned and retained as rare species in the flora of Pakistan.
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Artemisia is a polymorphic and widespread genus from the Asteraceae family. This genus includes more than 500 species out of which 37 species have been reported from Iran. Widely distributed in the desert area of Iran that makes this genus as one of the main elements of the Irano-Turanian region. In this study, eight species of Artemisia from different parts of Isfahan province were analyzed using scanning electron microscopy (SEM) and light microscopy (LM) techniques based on both qualitative and quantitative features of foliar epidermal cells. Type of epidermal cells varied from polygonal to irregular and elongated in shape with wavy to smooth in margins. This study, defined three different types of stomata: anemocytic, anisocytic, and paracytic on both the adaxial and the abaxial sides of Artemisia leaves species. Six different types of both glandular and non-glandular trichomes including capitate, peltate, aduncate, tectorial, conical type, and stinging hair types were observed in these species.
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Monardella is a western North America genus ranging from the Rocky Mountains to the Pacific Ocean and from southern Canada to the southern tip of Baja California Sur. The levels of unrecognized and cryptic taxa within Monardella are similar to other taxonomically complicated western North American genera, such as Eriogonum (Polygonaceae), Penstemon (Plantaginaceae), and Oreocarya (Boraginaceae). We present variation in trichome morphology as a taxonomic tool within Monardella, being comparable in importance to trichome variation in other plant families and genera (e.g., Brassicaceae, Draba; Loasaceae, Mentzelia; Solanaceae, Solanum). Trichome morphology within Monardella is a useful character to differentiate taxa and resolve taxonomic ambiguities. We present nomenclatural novelties and revisions for Monardella occurring in the Hells Canyon region along the lower Snake River of west-central Idaho and northeastern Oregon. We introduce the following taxonomic and nomenclatural changes. Monardella odoratissima Benth. var. neglecta Cronquist is recognized as a later homonym; a new name and rank are presented for it: M. perplexans Elvin, R. B. Kelley, & B. T. Drew. Two new species are described: M. kruckebergii Elvin, R. B. Kelley, & B. T. Drew and M. walwaamaxsia Elvin, R. B. Kelley, & B. T. Drew.
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Investigations on the anatomical features of the leaf and petiole of three species of Stachytarpheta (S.angustifolia, S.cayannensis and S.jamaicensis) that were common in Awka, South East Nigeria was made with the aid of a light microscope. Anatomical significant features were found in the leaf and petiole. The leaves and petioles were dorsiventral. The epidermis was conspicuous but one cell thick. The epidermal cells in the leaves were not of uniform size in both surfaces. Chlorenchyma was present as a very narrow portion of the leaves and petiole tissue while the palisade layer was one to three cells thick. These features however, could not proffer any taxonomic relevance to the delimitation of any of the species in this genus.
The foliar epidermal surfaces, leaf venation and trichome studies of different plant parts of the three species of Stachytarpheta Vahl. family Verbenaceae found in Nigeria were studied and compared. Observations were carried out by light microscope. Across the genus, leaves were amphistomatic and amphitrichomic. Qualitative foliar anatomical characters found to be most useful in species delimitation were stomata shape, trichome types, anticlinal cell wall pattern on adaxial surface and veinlet termination number. Quantitative characters of taxonomic importance, as revealed by analysis of variance (ANOVA) and duncan multiple range test (DMRT) were length and width of epidermal cells, stomatal index and stomatal size. Two trichome types, glandular and non-glandular were observed in the genus. The non-glandular trichomes were of higher numerical density on adaxial surface than on abaxial surface. The implication of this in terms of protection against excessive radiation, chemical control strategies and higher temperatures was mentioned. On the abaxial surface however, the glandular secreting trichomes were more numerically than the non-glandular. The importance of the preponderance of these trichomes on the abaxial surface in relation to the role they play in the protection of essential oils earlier reported in the family and genus, was discussed. Stomatal abnormalities observed in the genus were, contiguous stomata arrangement in S. cayennensis and aborted guard cells in S. indica. Leaf venation type in the genus is pinnate craspedodromous simple with recurved secondary veins. Trichome distribution survey in the different plant parts revealed specific combination types that would be useful in species delimitation, especially in fragmentary parts identification.
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The internal transcribed spacers (ITS-1 and ITS-2) of nuclear ribosomal DNA were sequenced for 33 populations of 31 species of Artemisia. The sequence data support the monophyly of the genus in its classical, broad sense, including Seriphidium and Oligosporus. Five main clades are defined, corresponding largely to the four main groups traditionally recognised in the genus: "Artemisia" (s.str.), "Absinthium", "Dracunculus", and "Seriphidium", plus the American "Tridentatae" segregated from the latter, except that "Artemisia", scattered over four different clades, is in need of a redefinition and restriction to the A. vulgaris clade. ITS data also support the hypothesis that the "Tridentatae" and "Seriphidium" are independent groups, and help differentiate the species in the A. herbaalba complex.
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The reconstruction of evolutionary trees by either conventional or cladistic methods depends upon the determination of evolutionary directionality (or polarity) of character states. Many criteria have been used to determine primitive states of characters, but these sometimes have duplicated the same concept and sometimes have not been based on evolutionary reasoning. Some apply only to specific taxa. We recognize two broad categories of criteria of primitiveness: first-level criteria, in which nothing is known about any primitive condition; and second-level criteria, in which at least one primitive condition already is known. We recognize nine criteria, the first six being first-level and the last three being second-level criteria: (1) fossil evidence; (2) common is primitive; (3) co-occurrence of primitive states; (4) earliest ontogenetic state; (5) minor abnormalities of organogenesis; (6) vestigial organs; (7) association; (8) correlation; and (9) group trends. The many specific criteria of primitiveness (e.g., hypogyny preceding epigyny in the angiosperms) are regarded as tendency statements of restricted utility. The only absolute criterion of primitiveness is that the oldest character state is primitive, which can never be determined with certainty because we lack the true phylogeny. The criteria, therefore, can only help approximate real phylogenies. Some criteria are more generally useful than others, but their efficacy depends to some extent on the nature of characters and character states within individual taxa.
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Plant volatiles are important cues for the orientation of herbivorous insects. It is possible that these compounds indicate whether the plant is suitable for feeding and larval development, or for mating aggregation. Vernonia condensata (Asteraceae) is known to attract species of leafhoppers, most of them important vectors of the citrus variegated chlorosis (CVC). In this study, we evaluated the role of volatiles of V. condensata on the orientation of Bucephalogonia xanthophis (Hemiptera: Cicadellidae). Four-arm olfactometer bioassays showed that only males were attracted to the volatiles of the host-plants Citrus sp. and V. condensata. Furthermore, fresh leaves of V. condensata induced a stronger response than volatiles from hexane-extracted leaves. This study opens the possibility to utilize V. condensata volatiles for pest management programs of B. xanthopis.
The visibility of leaf glandular trichomes with 10× magnification was examined for 7 sagebrush taxa. In 75% of the Artemisia nova plants the glands were easily observed protruding through a layer of nonglandular trichomes (hairs). On A. arbuscula and the 3 subspecies of A. tridentata the glands were covered by the hairs and were not visible. These structures can be observed in the field with a hand lens and provide a supportive characteristic for the identification of A. nova.
Nonglandular trichomes in Artemisia ludoviciana Nutt. are multicellular T-shaped structures which vary in density and hair cell length. Glandular trichomes are similar in structure to those of other Artemisia species, and their density is weakly correlated to nonglandular trichome density. The specialist feeder Hypochlora alba (Dodge) ingests these trichomes along with leaf tissue, but does not digest the nonglandular trichomes.
The out-group comparison method of character analysis. Syst. Zool., 30:1–11.—An operational rule for analyzing character polarity with out-group comparison is presented and a series of observations, including potential problems in applying the rule, are discussed. The “commonality principle” (=“frequency of occurrence,” “common equals primitive”) for determining character polarity is reviewed and dismissed as a reliable alternative to out-group comparison. Based on the rule and observations, a general method for character analysis is synthesized.