Introduction
Apocynaceae and Asclepiadaceae constitute the most diverse lineage
of Gentianales and regarded as two closely related
families (Civeyrel et al. 1998). Endress et al.
(1996) and Sennblad and Bremer (2002)
reported that Apocynaceae sensu
lato (s.l.)
includes the former Asclepiadaceae, and this merge is based on both
morphological and molecular similarities. Melchior
(1964), Dahlgren (1983) and Jeffrey and
Cronquist (1984) separated Asclepiadaceae from Apocynaceae
and divided Apocynaceae s.l.
into two distinct families recognizing two subfamilies per each; Plumieroideae and Echitoideae
within Apocynaceae, while Cynanchoideae
and Periplocoideae within Asclepiadaceae.
It has long been known that
some taxa of Apocynaceae s.l.
are more morphlogically similar to Asclepiadaceae
than they are to their own family, a series of morphology-based papers focused on pollen have proved
to be the most important morphological character (Endress et al. 2014; Dabbub et al. 2020). The importance of stomatography
and leaf venation as a helpful taxonomic tool needs no emphasis, as these
characters have immense value in identifying species of Apocynaceae
s.l. (Chandra et
al 1968). Foliar epidermal characters
are considered as very important diagnostic tool to differentiate among the
species of the family (Ayaz et al. 2020).
Light microscopy (LM) and Scanning electron microscopy (SEM) on leaf of Apocynaceae
were believed to have a pivotal role in the classification and delimitation of
its species (Bashir et al. 2020).
A wide range of
molecular approaches has been used by numerous research groups to find the
natural relationship within various groups of Apocynaceae
(Potgieter and Albert 2001; Rapini et al. 2003; Rapini et al. 2007; Simões et al.
2007; Livshultz 2010). Fishbein (2001)
used chloroplast matk gene and confirmed the
monophyly of subfamilies and assumed a closer relationship of Periplocoideae to Apocynaceae s.s. than to the remainder of Asclepiadaceae. Sennblad and Bremer (1996) used the sequence data for rbcL gene of 24 species of Apocynaceae
and Asclepiadaceae to evaluate the existing familial and subfamial
classification, and concluded that Asclepiadaceae is nested into Apocynaceae. Judd et al. (1994) and Kunze (1996) reported that rbcL gene region did
not support the monophyly of Asclepiadaceae and Periplocaceae
as separate families, rather indicated a close relationship among Periplocaceae and Apocynaceae s.s.
A controversy over the taxonomic relationship between
the Apocynaceae and Asclepiadaceae persisted. Apocynaceae s.l. has been divided into five subfamilies viz.
Apocynoideae, Rauvolfioideae,
Asclepiadoideae, Secamonoideae
and Periplocoideae. The former two subfamilies belong
to Apocynaceae s.s.,
while the latter three belonging to Asclepiadaceae (Endress et al. 2000; Endress et al. 2007; Angiosperm Phylogeny Group
2016). The principle target of the current
investigation was to embrace whether Apocynaceae should be treated as distinct
entity or two separated taxonomic entities through the use of some diagnostic
morphological characters in phenetic analysis, as well as DNA barcoding based on the sequencing of rbcL gene that is used in phylogenetic
analysis.
Materials and Methods
Twenty species representing
four sub-families and 19 genera of Apocynaceae s.l. were
collected from natural habitats (wild species) and different botanical gardens
(cultivated species) in
Egypt and Libya (Table 1). The identification and Synonyms were authenticated
based on previous infornation (Bailey 1949; Tackholm 1974; Boulos 2000; IPNI 2012).
Macromorphological
characters investigation
The
macromorphological characters of the whole plant viz. habit, stem, leaf
and flower were examined, described from the fresh specimens and with the aid
of text books (Bailey 1949; Hutchinson and Dalziel 1954; Boulos 2000; Endress
and Bruyns 2000; Pandey and Pandey 2006). Huernia andreaeana is with rudimentary leaves, so the lamina vein
architecture is not dealt with. Lamina vein architecture is examined
from fresh leaves in Nerium
oleander, Cynanchum
acutum and Carissa carandas, while in other species, leaves
were cleaned and soaked in a strong household bleach
solution (sodium hypochloride less than 5%, sodium
hydroxide less than 5% and water) until they turn white. The leaves were
removed from bleach and rinsed with water then stained with 1% safranine (Johansen 1940).
Lamina
epidermal strips were prepared from fresh leaves of the studied species
(leaves rudimentary in H. andreaeana). The
epidermal strips were prepared by mechanically stripping (Johansen 1940) or by chemical methods (Pohl 1967) by taking fragment of 5˗10 mm2
from median portion of leaf, soaked in nitric acid and hydrogen peroxide
solution (1:1) for a period (2 h for two days) depending on the leaf thickness.
The epidermal strips were stained with 1% safranine, mounted on slides. Examination and photomicrographs were taken using Canon
power-shot A720, 8.0 mega pixels. For SEM investigation, small pieces (7 mm2)
of the fresh leaf material were fixed on SEM stubs with double-sided tape,
coated with gold in SPI-Module sputter coater. Only abaxial surface was
observed and photographed by Scanning Electron Microscope (JSM-5500 LV; JEOL
Ltd-Japan) by using high vacuum mode. Terminology of epidermal characteristics
was performed with the help of previous information (Metcalfe and Chalk 1950; Ash et
al. 1999; Prabhakar 2004; Stearn 2005).
DNA extraction
and PCR amplification with rbcL primers
The fresh leaves of the studied species were collected
and subjected to molecular analysis, Solenostemma argel material was
practically unavailable and loaded from gene bank (ID HG530567). Leaves (100
mg) of the available material were ground to a powder using liquid nitrogen in
Eppendorf tube then DNA were isolated using CTAB (Cetyl-trimethyl ammonium bromide) protocol of (Doyle and Doyle 1987). The purified DNA
was used in PCR amplification of rbcL region
using universal primers as the following:
Forward
primer: 5´-ATG TCA ACA CAA ACA GAG ACT AAA GC-3´
Reverse primer:
5´-GAA ACG GTC TAT CCA ACG CAT-3´
The
amplification reactions were performed in 25 μL as follow; 5x GoTaq® Flexi
buffer 5μL, MgCl2 (25 mM) 2.5 μL,
dNTPs (10 mM each) 0.5 μL, forward primer (10 μM) 1.2 μL, reverse
primer (10 μM) 1.2 μL,
Go Taq™ (5 U/μL) 5 μL,
DNA Stock 2 μL, H2O 7.6 μL
up to make 25 μL total volume. The reaction conditions were
as the following:
Initial denaturation at 95°C for 5 min., 40 cycles at 94°C for 30 s., 58°C
for 30 s., 72°C for 45 s. and 72°C for 10 min. All positive PCR amplicons were prepared for the cleanup step
(purification) from other undesired substances as dimers, RNA, free nucleotides
and unamplified DNA fragments by using PCR product purification Kit (Thermo PCR
purification kit, USA). It is an essential step before automated
DNA-sequencing. The purified DNA was submitted for sequencing to Macrogen, Korea; 6F, 172, Dolma-ro,
Bundang-gu, Seongnam-si,
Gyeonggi-do (Jeongja-dong, Seoul National University Bundang Hospital Healthcare Innovation Park).
Phenetic
analysis
The
morphological data of 20 species were subjected to phenetic analysis based on
176 morphological characters states. Prior to analysis the presence for
character states were indicated
as numerical values in order to make comparison during similarity estimation
feasible. Thus, a uniform convention was used for all characters; “1” used for
present and “0” for absent.
All computations were carried out using NTSYS-PC version 2.02 software program (Rohlf 2000).
Phylogenetic
analysis
Crucianella angustifolia (family Rubiaceae) was designed as an out group for phylogenetic
analysis and loaded also from gene bank (ID X81094). The sequences were prepared for alignment to get
the best trees. The chromatograms in sequences were compiled using BIOEDIT V3
program (Hall 1999). Sequence service
chromatogram (fasta file) of sample was imported into
a new alignment file created in BIOEDIT. The MEGA7 a phylogenetic program was
used for assembling data and constructing the tree. Alignment fasta file was entered into the program for constructing
the phylogenetic tree. Tree was constructed according to Neighbor Joining
model, which is based on the number of distances between sequences. The number
of bootstrap tests was performed using 1000 replicates.
Results
Phenetic analysis
A summary of morphological characters states and its codes as revealed
by LM & SEM is presented
in Table 2 and some of the most specific structures were illustrated in Fig. 1.
The obtained phenogram (Fig. 2) resulting from the
cluster analysis of 176 morphological characters states of whole plant, lamina
architecture, stomatography (LM and SEM) showed that H.
andreaeana basally segregated with unresolved
relationship due to the lack of many character states (102 leaf character
states). The remaining 19 species were separated into two series; I and II.
Series I comprised Acokanthera
oblongifolia, Alstonia
scholaris, Adenium obesum,
C. carandas, C. macrocarpa, Cerbera odollam, Tabernaemontana divaricate, Wrightia coccinea, Kopsia arborea, Plumeria obtusa, Cascabela thevetia, N. oleander, Asclepias
curassavica, Cryptostegia
grandiflora, Calotropis procera, S. argel, Gomphocarpus sinaicus and
C. acutum, at a taxonomic distance 0.38 based on sharing of simple leaf, paracytic or anomocytic stomata,
smooth surface anticlinal wall, pinnate primary vein category and regular
polygonal reticulate quaternary vein category. Series II included Catharanthus
roseus as a single entity at a taxonomic distance 0.38. Owing to annual
herb, amphistomatic leaf type, eucamptodromus secondary veins, opposite percurrent
tertiary veins with straight course, wrinkled anticlinal wall of epidermal
cells.
Series I was divided into two sub-series; (A and B) at a
taxonomic distance 0.45. Sub-series A was divided into two clusters; C1 and C2
at a taxonomic distance 0.47. Cluster 1 was divided into two groups; G1 and G2
at a taxonomic distance 0.51. Group 1 included A. oblongifolia
and A. scholaris as one group due to presence fissured layers wax with ill-defned sculpture of
lamina abaxial epidermal cells and elliptic
leaf. Group 2 included nine studied species sharing the presence of narrow, depressed anticlinal walls with smooth
surface, and raised periclinal walls.
Cluster 2 included N. oleander at a taxonomic
distance 0.47 owing to cladodromous secondary veins categoty, sunken stomata and curved anticlinal wall
adaxially, incomplete looped marginal veins, one-branched freely ending
ultimate veins, granulate
crystalloid epicuticular wax and colliculate sculpture of
lamina abaxially.
Sub-series B was divided into two clusters; C3 and C4 at
a taxonomic distance 0.55. C 3 was divided into two groups; G3 and G4 at a
taxonomic distance 0.60. Group 3 included A. curassavica
and C. grandiflora grouped together as one group. The latter group 4
included C. procera, G. sinaicus
and S. argel based on sharing morphological characters viz. ad medially ramified tertiary veins, amphistomatic leaf
type and sunken or leveled stomata elevation abaxially. The latter cluster 4 included C. acutum due to presence climbing
stem, cordate leaves with actinodromus primary vein category, ill-defned
sculpture of lamina abaxially with hight density of
non-entire platlets epicuticular.
Phylogenetic analysis
The obtained
sequences were uploaded to Genbank and gained
accession numbers as follow; BankIt2396104 A. oblongifolia MW208824, BankIt2396104 A. obesum
MW208825, BankIt2396104 A. scholaris MW208826, BankIt2396104 A. curassavica MW208827,
BankIt2396104 C. procera
MW208828, BankIt2396104 C. carandas
MW208829, BankIt2396104 C. macrocarpa
MW208830, BankIt2396104 C. thevetia MW208831, BankIt2396104
C. roseus MW208832, BankIt2396104 C.
odollam MW208833, BankIt2396104 C. grandiflora MW208834, BankIt2396104 C. acutum MW208835, BankIt2396104 G. sinaicus
MW208836, BankIt2396104 H. andreaeana MW208837, BankIt2396104 K. arborea MW208838, BankIt2396104 N. oleander MW208839, BankIt2396104 P. obtusa
MW208840, BankIt2396104 T. divaricata MW208841, BankIt2396104 W. coccinea MW208842.
Table
1: Data of
collection of the studied species
|
No. |
Taxa |
Locality/source |
|
1. |
Acokanthera oblongifolia Benth. & Hook.f. - Gen. Pl. [Bentham & Hooker f.] 2(2): 696.
1876 [May 1876]; nom. inval. (IK) = A. spectabilis
(Sond.) Hook.f. |
BG |
|
1.
2. |
Adenium obesum Roem. & Schult. - Syst. Veg., ed. 15 bis [Roemer & Schultes] 4: 411. 1819 (IK) = A. arabicum Balf.f |
OBG |
|
2.
3. |
Alstonia scholaris (L.) R.Br. - Mem. Wern.
Nat. Hist. Soc. 1: 75. 1811 [dt. 1809; issued in 1811] (IK) = Echites scholaris L. |
GZ |
|
3.
4. |
Asclepias curassavica L. - Sp. Pl. 1: 215. 1753 [1 May 1753] (IK) = A. margaritacea Hoffmanns. ex Schult. |
OBG |
|
4.
5. |
Calotropis procera W.T.Aiton
- Hort. Kew., ed. 2 [W.T. Aiton] 2: 78. 1811 (IK) = C. persica Gand. |
WML |
|
5.
6. |
Carissa carandas L. -- Mant. Pl. 52. 1767 [15-31
Oct 1767] (IK) = C. carandas var. congesta
(Wight) Bedd. |
BG |
|
6.
7. |
Carissa macrocarpa (Eckl.)
A. DC. - Prodr. [A. P.
de Candolle] 8: 336. 1844 [mid Mar 1844] (IK) = Arduina
grandiflora E.Mey. |
OBG |
|
7.
8. |
Cascabela thevetia (L.) Lippold. - Feddes Repert. 91: 52. 1980 (GCI)
= Thevetia peruviana (Pers.) K.Schum. |
BG |
|
8.
9. |
Catharanthus roseus (L.) G.Don.
- Gen. Hist. 4(1): 95. 1837 (IK) =Vinca rosea L. |
AUG |
|
9.
10 |
Cerbera odollam Gaertn. - Fruct.
Sem. Pl. 2: 193. 1791 (IK) = Odollamia malabarica Raf. |
AG |
|
10.
11. |
Cryptostegia grandiflora R.Br. - Bot. Reg. 5: t. 435. 1820 [1819 publ. 1820] (IK) = C. grandiflora Roxb. ex R.B. |
AMG |
|
11.
12. |
Cynanchum acutum L. - Sp. Pl. 1: 212. 1753 [1
May 1753] (IK) = C. excelsum Desf. |
AU |
|
12.
13. |
Gomphocarpus sinaicus Boiss.
- Diagn. Pl. Orient. ser. 1, 11: 80. 1849 [Mar-Apr 1849] (IK) = Asclepias sinaica (Boiss.) Muschl = Gomphocarpus schimperi C.Presl |
SK |
|
13.
14. |
Huernia andreaeana (Rauh)
L.C.Leach - J. S. African
Bot. 40(1): 21. 1974 (IK) = H. appendiculata
Berger. |
WML |
|
14.
15. |
Kopsia arborea
Blume. - Cat. Gew. Buitenzorg (Blume) 13. 1823; et Bijdr. Fl. Ned. Ind. 16: 1030 [Oct 1826-Nov 1827].
(IK) = K. jasminiflora Pit. |
BG |
|
15.
16. |
Nerium oleander L. - Sp. Pl. 1: 209. 1753 [1 May 1753] (IK) = N. carneum Dum.Cours. |
AUG |
|
16.
17. |
Plumeria obtusa L. - Sp. Pl. 1: 210. 1753(IK) = P. apiculata Urb. |
AUG |
|
17.
18. |
Solenostemma argel Hayne - Getreue
Darstell. Gew. ix. t.38.
1825. (IK) = Cynanchum arghel
Delile. |
A |
|
18.
19. |
Tabernaemontana divaricata (L.)
R.Br. ex Roem. & Schult. - Syst.
Veg., ed. 15 bis [Roemer & Schultes] 4: 427. 1819
(IK) = Taberna discolor (Sw.) Miers |
BG |
|
19.
20. |
Wrightia coccinea Sims - Bot. Mag. 53: t. 2696.
1826 (IK) = Nerium coccineum Roxb. ex Hornem. = N. coccineum Lodd. |
AG |
BG: Botanical Garden, Botany Department, Faculty of
Science, Ain Shams University. AU: Al-Azhar University Garden. A: Asswan. OBG:
Orman Botanical Garden, Giza. AUG: Al-Zawia
University Garden in Libya. WML: Western Mountain in Libya. AG: Al-Zohriya Garden, Gizzira. SK:
Saint Katherine, South Sinai. AMG: Agriculture Museum Garden. GZ: Giza Zoo
The generated cladogram (Fig. 3),
which was rooted by C.
angustifolia as an outgroup, supported the
monophylly of the taxa under investigation, which can be differentiated into
four lineages. The first one
included P. obtusa and K. arborea as
sister taxa due to sharing
of intermarginal secondary veins, well developed areolation, two or more branched freely ending ultimate veins, straight anticlinal wall of ab/ adaxial epidermal cells and leveled stomata (synapomorphic characters). The second lineage included A. oblongifolia, C. carandas,
C. macrocarpa, A. scholaris, C. odollam and C. thevetia. All taxa are nested
together at 0.771 bootstrap value after separation of C. roseus and T.
divaricata at earlier basal level. A. oblongifolia, C. carandas and C.
macrocarpa are nested together at 0.787 bootstrap value based on synapomorphic characters viz. shrub habitat, salver
corolla tube and union of carpels (ovary, style and stigma) throughout. In the third lineage, C.
grandiflora and W. coccinea were nested together at a high bootstrap value (0.792) due to
the presence of opposite leaves, weak
brochidodromous secondary veins and paracytic lens-shaped stomata (synapomorphic characters). In the last lineage, eight studied species were nested at 0.772
bootstrap value. A. obesum and N. oleander were
separated consequently at basal level, while A. curassavica,
G. sinaicus, C. procera, C.
acutum, S. argel and H. andreaeana were
nested together at 0.993 bootstrap value. The latter six species shared many synapomrphic characters such as campanulate corolla tube,
presence of gynostegium, two carpels (free below and
united above) with true pollinia. These species were separated consequently
from H. andreaeana at basal level due to autapomorphic characters viz. succulent stem, redumintary leaves features in addition to winged pollinium. C. acutum was separated based on
autapomorphic characters viz. climbing stem, cordate leaf and erect
orientation of pollinium. C. acutum was delimited from all the studied
species of Asclepiadeae on the basis of rhizome stem
and presence of opposite petiolated leaves, reaching G.
sinaicus and A. curassavica
that forming sister species based on synapomorphic
characters viz. globose head stigma, oblong-ovate shape of pollinium sac
and yellow colour pollinium. S. argel is considered a sister group to
C. procera, A. curassavica
and G. sinaicus.
Discussion
The generated phenogram supports the
treatment of Apocynaceae s.l.
as two distinct families, Apocynaceae s.s. and Asclepiadaceae where there is a clear separation of
series I into two subseries; one of them belongs to Apocynaceae
and the other with the remaining taxa belongs to Asclepiadaceae (Brown 1810; Bessey 1915; Hutchinson 1959; Takhtajan
1980; Goldberg 1986; Dabbub et al.
2020). Within Apocynaceae A.
oblongifolia and A. scholaris
were nested in one group and this is in accordance with (Byng et al. 2016) and (Endress and Bruyns 2000) where they included
them within subfamily Rauvolfioideae. Group 2
contained two species from Apocynoideae (A. obesum and W. coccinea) and seven
species from Rauvolfioideae according to (Byng et al. 2016) and (Endress and Bruyns 2000). Sub-series B was divided into two clusters; C3 and C4. C
3 was divided into two groups; G3 and G4. Group 3 included A. curassavica and C. grandiflora grouped kept
together as one group. The latter group 4 included C. procera,
G. sinaicus and S. argel. This group is
comparable with tribe Asclepiadeae according to (Melchior 1964; Endress et al.
2014; Byng et al. 2016; El-gazzar et al. 2018).
Phylogentically, P. obtusa
and K. arborea showed a sister relationship both belonging to Rauvolfioideae, this is comparable with Melchior (1964) and Sennblad and Bremer
(1996) who gathered them under
subfamily Plumeroideae (Apocynacese s.s.), while Endress and Bruyns (2000) and Byng et al.
(2016) put them in subfamily Rauvolfioideae (Apocynacese s.l.). A. oblongifolia, C. carandas,
C. macrocarpa, A. scholaris, C. odollam and C. thevetia were nested together (Rauvolfioideae). While C. roseus and T. divaricata were separated at earlier basal level; the pattern of
separation clarified that Rauvolfioideae is not
monophyletic group (paraphyletic). (Simões et
al. 2007) reported six monophyletic tribes within Rauvolfioideae viz.
Table 2:
Morphological
character, states (176) and its codes of the studied species used for phenetic analysis. (0 = Absent; 1 = Present)
|
Organ |
Character |
Species Character states |
Acokanthera oblongifolia |
Adenium obesum |
Alstonia scholaris |
Asclepias curassavica |
Calotropis procera |
Carissa carandas |
Carissa macrocarpa |
Cascabela thevetia |
Catharanthus roseus |
Cerbera odollam |
Cryptostegia grandiflora |
Cynanchum acutum |
Gomphocarpus sinaicus |
Huernia andreaeana |
*Kopsia arborea |
Nerium oleander |
Plumeria obtusa |
Solenostemma argel |
Tabernaemont-ana ivaricata |
Wrightia coccinea |
||
|
Whole plant |
Duration |
Perennial |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
||
|
Annual |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
||||
|
Habit |
Herb |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
|||
|
Shrub |
1 |
1 |
0 |
0 |
1 |
1 |
1 |
1 |
0 |
0 |
1 |
0 |
1 |
0 |
1 |
1 |
1 |
0 |
1 |
0 |
||||
|
Succulent herb |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
||||
|
Tree |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
||||
|
Stem |
Stem latex |
Milky |
1 |
0 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
1 |
1 |
1 |
1 |
0 |
1 |
0 |
1 |
1 |
1 |
1 |
||
|
Watery |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
||||
|
Stem strength |
Erect |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
1 |
0 |
1 |
1 |
1 |
1 |
1 |
1 |
|||
|
Weak |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
||||
|
Stem texture |
Glabrous |
1 |
1 |
1 |
1 |
1 |
0 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
1 |
1 |
1 |
1 |
1 |
1 |
|||
|
Spiny |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
||||
|
Leaf |
Occurrence |
Rudimentary |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
||
|
Foliage |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
1 |
1 |
1 |
1 |
1 |
1 |
||||
|
Arrangement |
Alternate |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
|||
|
Opposite |
1 |
0 |
0 |
1 |
1 |
1 |
1 |
0 |
1 |
0 |
1 |
1 |
1 |
0 |
1 |
0 |
0 |
1 |
1 |
1 |
||||
|
Whorled |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
||||
|
Leaf petiole |
Sessile |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
|||
|
Petiolate |
1 |
0 |
1 |
1 |
0 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
1 |
1 |
1 |
1 |
1 |
1 |
||||
|
Lamina shape |
Ovate |
0 |
0 |
0 |
0 |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
1 |
|||
|
Ob-ovate |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
||||
|
Oblong-ovate |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
||||
|
Elliptic |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
||||
|
Lanceolate |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
1 |
1 |
0 |
0 |
||||
|
Cordate |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
||||
|
Lamina
apex |
Acute |
1 |
0 |
0 |
1 |
1 |
0 |
0 |
1 |
0 |
0 |
1 |
1 |
1 |
0 |
1 |
1 |
0 |
1 |
0 |
0 |
|||
|
Obtuse |
0 |
1 |
1 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
||||
|
Acuminate |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
1 |
1 |
||||
|
Flower |
Inflorescence |
Cymose |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
1 |
1 |
0 |
1 |
1 |
1 |
1 |
0 |
1 |
1 |
||
|
Racemose |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
||||
|
Sepal texture |
Hairy |
0 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
0 |
0 |
1 |
1 |
1 |
0 |
0 |
1 |
0 |
1 |
0 |
0 |
|||
|
Glabrous |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
1 |
1 |
0 |
0 |
0 |
1 |
1 |
0 |
1 |
0 |
1 |
1 |
||||
|
Corolla tube shape |
Salver-like |
1 |
1 |
1 |
0 |
0 |
1 |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
|||
|
Campanulate |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
1 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
||||
|
Funnel |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
1 |
1 |
0 |
1 |
1 |
||||
|
Rotate |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
||||
|
Petal color |
White |
1 |
0 |
1 |
0 |
0 |
1 |
1 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
1 |
0 |
1 |
1 |
0 |
|||
|
Violetish-white |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
||||
|
Pink |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
||||
|
Red |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
1 |
||||
|
Yellow |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
||||
|
Pinkish- white |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
||||
|
Petal
apex |
Acut |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
|||
|
Acuminate |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
||||
|
Obtuse |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
1 |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
||||
|
Petal texture |
Hairy |
1 |
0 |
1 |
0 |
0 |
1 |
1 |
1 |
1 |
0 |
1 |
0 |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
|||
|
Glabrous |
0 |
1 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
1 |
0 |
1 |
0 |
1 |
1 |
1 |
1 |
1 |
||||
|
Androecium |
Stamens adhesion |
Epipetalous |
1 |
1 |
1 |
0 |
0 |
1 |
1 |
1 |
1 |
1 |
0 |
0 |
0 |
0 |
1 |
1 |
1 |
0 |
1 |
1 |
||
|
Gynostegium |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
1 |
1 |
1 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
||||
|
Stamens Union |
Adhere
to stigma |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
|||
|
Adhere
to style& stigma |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
1 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
||||
|
Not
adhere to style nor stigma |
1 |
1 |
1 |
0 |
0 |
1 |
1 |
1 |
1 |
1 |
0 |
0 |
0 |
0 |
1 |
1 |
1 |
0 |
1 |
1 |
||||
|
Gynoecium |
Ovary |
Carpels |
Free
below and united above |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
1 |
1 |
1 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
|
|
United throughout |
1 |
1 |
1 |
0 |
0 |
1 |
1 |
1 |
1 |
1 |
0 |
0 |
0 |
0 |
1 |
1 |
1 |
0 |
1 |
1 |
||||
|
Texture |
Hairy |
0 |
1 |
1 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
|||
|
Glabrous |
1 |
0 |
0 |
1 |
1 |
0 |
1 |
1 |
0 |
1 |
1 |
1 |
1 |
0 |
1 |
0 |
1 |
1 |
1 |
1 |
||||
|
Style |
Number |
One |
1 |
1 |
1 |
0 |
0 |
1 |
1 |
1 |
1 |
1 |
0 |
0 |
0 |
0 |
1 |
1 |
1 |
0 |
1 |
1 |
||
|
Two |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
1 |
1 |
1 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
||||
|
Stigma |
Shape/ head |
Globose head |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
||
|
Pentagonal head |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
||||
|
Armed |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
||||
|
Cylindrical |
0 |
0 |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
||||
|
Bilobed |
1 |
1 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
1 |
1 |
||||
|
Nectary disc |
Present |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
|||
|
Absent |
1 |
1 |
0 |
1 |
1 |
1 |
1 |
0 |
0 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
1 |
1 |
1 |
1 |
||||
Table 2 Continued
%201111-1119_files/image001.png)
Table 2: Continued
|
Lamina
architecture |
1°
vein
category |
Pinnate |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
1 |
0 |
1 |
1 |
1 |
1 |
1 |
1 |
|
Basal
actinodromous |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
||
|
2°vein category |
Weak brochidodromou |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
1 |
1 |
|
|
Festooned
brochidodromous |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
||
|
Brochidodromous |
0 |
1 |
0 |
1 |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
||
|
Eucamptodromous |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
||
|
Intramarginal
vein |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
1 |
0 |
0 |
0 |
||
|
Cladodromous |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
||
|
2° vein spacing |
Decreasing
toward base |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
|
|
Increasing
toward base |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
||
|
Irregular |
0 |
0 |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
||
|
Uniform |
0 |
1 |
1 |
1 |
1 |
0 |
0 |
1 |
1 |
1 |
0 |
0 |
1 |
0 |
1 |
1 |
1 |
0 |
0 |
1 |
||
|
2° vein angle |
Uniform |
0 |
1 |
1 |
0 |
1 |
0 |
0 |
1 |
1 |
0 |
0 |
1 |
0 |
0 |
1 |
1 |
1 |
1 |
0 |
1 |
|
|
Smoothly
decrease toward base |
1 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
||
|
Smoothly
increase toward base |
0 |
0 |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
||
|
Inter-2° vein |
Absent |
0 |
0 |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
1 |
|
|
Weak |
0 |
0 |
0 |
1 |
0 |
1 |
0 |
1 |
0 |
0 |
1 |
1 |
1 |
0 |
1 |
1 |
0 |
1 |
0 |
0 |
||
|
Strong |
1 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
||
|
3°vein category |
Regular
polygonal reticulate |
0 |
1 |
0 |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
|
|
Random reticulate |
1 |
0 |
0 |
0 |
0 |
0 |
1 |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
||
|
Alternate
percurrent |
0 |
0 |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
1 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
||
|
Opposite
percurrent |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
||
|
Mixed
opp/alt |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
1 |
1 |
||
|
3°vein
course |
Exmedially |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
|
|
Admedially ramified |
0 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
1 |
0 |
1 |
1 |
0 |
1 |
1 |
1 |
1 |
1 |
0 |
||
|
Straight |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
||
|
Sinuous |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
||
|
4° vein category |
Ill-developed |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
|
|
Regular
polygonal reticulate |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
1 |
1 |
1 |
1 |
0 |
1 |
1 |
1 |
0 |
1 |
1 |
||
|
5°
vein category |
Ill-developed |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
1 |
0 |
0 |
|
|
Regular polygonal |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
1 |
0 |
||
|
Dichotomizing |
0 |
1 |
1 |
1 |
0 |
1 |
0 |
0 |
0 |
1 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
||
|
Aereolation |
Lacking |
1 |
1 |
1 |
1 |
1 |
0 |
0 |
0 |
1 |
1 |
1 |
1 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
|
|
Well-developed |
0 |
0 |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
1 |
1 |
0 |
0 |
1 |
||
|
Poorly
developed |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
||
|
Free
ending ultimate vein |
Absent |
1 |
0 |
0 |
0 |
1 |
0 |
1 |
1 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
|
|
1-
branched |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
||
|
2-
or more branched |
0 |
1 |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
1 |
0 |
0 |
1 |
0 |
1 |
0 |
1 |
1 |
||
|
Marginal
ultimate venation |
Looped |
1 |
1 |
0 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
0 |
1 |
0 |
1 |
1 |
1 |
1 |
|
|
Incomplete
loop |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
||
|
Lamina epidermal
characters (LM) |
Leaf
type |
Ill-defined |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
|
Hypostomatic |
1 |
1 |
1 |
1 |
0 |
1 |
1 |
1 |
0 |
1 |
1 |
1 |
0 |
0 |
1 |
0 |
1 |
0 |
1 |
1 |
||
|
Amphistomatic |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
||
|
Abaxial
cell shape |
Polygonal |
1 |
1 |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
1 |
0 |
1 |
0 |
1 |
1 |
0 |
0 |
|
|
Irregular |
0 |
0 |
0 |
1 |
0 |
1 |
1 |
1 |
1 |
1 |
0 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
1 |
||
|
Abaxial
anticlinal walls |
Straight |
0 |
1 |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
1 |
0 |
1 |
0 |
1 |
1 |
0 |
0 |
|
|
Curved |
1 |
0 |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
||
|
Undulate |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
1 |
1 |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
1 |
||
|
Adaxial
cell shape |
Polygonal |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
1 |
0 |
1 |
1 |
1 |
0 |
1 |
0 |
1 |
1 |
1 |
0 |
|
|
Irregular |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
1 |
||
|
Adaxial
anticlinal walls |
Straight |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
1 |
1 |
1 |
1 |
1 |
0 |
1 |
0 |
1 |
1 |
1 |
0 |
|
|
Curved |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
||
|
Undulate |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
||
|
Abaxial stomatal type |
Wanting |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
|
|
Paracytic |
1 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
1 |
||
|
Anomocytic |
0 |
0 |
0 |
1 |
1 |
0 |
1 |
0 |
1 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
||
|
Anisocytic |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
||
|
Sunkun |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
||
|
Paracytic& Anisocytic |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
||
|
Adaxial
stomatal type |
Wanting |
1 |
1 |
1 |
1 |
0 |
1 |
1 |
1 |
0 |
1 |
1 |
1 |
0 |
0 |
1 |
1 |
1 |
0 |
1 |
1 |
|
|
Anomocytic |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
||
|
Anisocytic |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
||
|
Trichomes |
Wanting |
1 |
0 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
1 |
1 |
1 |
0 |
0 |
1 |
0 |
1 |
0 |
1 |
1 |
|
|
Eglandular |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
1 |
0 |
0 |
||
|
Glandular (clavate) |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
||
|
Lamina epidermal characters (SEM) |
Epicuticular
wax |
Striated |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
|
Fissured
layers |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
||
|
Smooth film |
0 |
0 |
0 |
0 |
0 |
1 |
1 |
1 |
0 |
1 |
0 |
0 |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
||
|
Granulate
crystalloid |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
||
|
Crystalloid
threads |
0 |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
1 |
1 |
||
|
Non entire
platelets |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
||
|
Sculpture |
Rugose |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
|
|
Ruminate |
0 |
1 |
0 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
1 |
1 |
||
|
Ill-defined |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
||
|
Reticulate-foveate |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
||
|
Colliculate |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
||
|
Abaxial anticlinal
walls width |
Not obvious |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
|
|
Narrow |
0 |
1 |
0 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
1 |
0 |
1 |
1 |
1 |
0 |
1 |
1 |
||
|
Broad |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
||
|
Abaxial
anticlinal walls elevation |
Not obvious |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
|
|
Raised |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
||
|
Depressed |
0 |
0 |
0 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
1 |
0 |
1 |
1 |
1 |
1 |
1 |
1 |
Table 2: Continued
Table 2: Continued
|
|
Abaxial
anticlinal wall surface |
Ill-defined |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
|
|
Smooth |
0 |
1 |
0 |
1 |
1 |
1 |
1 |
1 |
0 |
1 |
1 |
0 |
1 |
0 |
1 |
1 |
1 |
1 |
1 |
1 |
|||
|
Wrinkled |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
|||
|
Abaxial
periclinal wall elevation |
Ill-defined |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
||
|
Raised |
0 |
0 |
0 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
1 |
0 |
1 |
1 |
1 |
1 |
1 |
1 |
|||
|
Depressed |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
|||
|
Abaxial
periclinal wall surface |
Ill-defined |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
||
|
Smooth |
0 |
1 |
0 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
0 |
1 |
0 |
1 |
0 |
0 |
0 |
1 |
1 |
|||
|
Wrinkled |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
|||
|
Granulated |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
|||
|
Stomata |
Shape |
Circular |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
|
|
Lens-shaped |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
1 |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
|||
|
Elliptic |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
|||
|
Ovate |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
|||
|
Semi-circular |
0 |
0 |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
|||
|
Sunken in
groove |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
|||
|
Elevation |
Leveled |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
1 |
1 |
1 |
1 |
||
|
Sunken |
0 |
0 |
0 |
0 |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
1 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
|||
|
Raised |
0 |
1 |
0 |
1 |
0 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
|||
Alyxieae, Carisseae, Hunterieae, Plumerieae, Tabernaemontaneae and
Willughbeieae. On the other hand, phylogenetic
studies reported the non-monophyletic groups in Rauvolfioideae
(Potgieter and Albert 2001). The
subfamily Plumerioideae of the Apocynaceae
has often been interpreted as a paraphyletic group, from which the other
subfamilies of the Apocynaceae have evolved (Fallen 1986; Endress et al. 2014). A. oblongifolia, C. carandas and C.
macrocarpa are nested together (tribe Carissae) at 0.787
bootstrap value, which is inaccordance with the
previous reports (Pichon et al. 1950; Endress and Bruyns 2000; Sennblad and Bremer 2002;
Endress et al. 2014; Byng et al. 2016). Likwise, C. grandiflora (Priplocoideae) and W. coccinea (Apocynoideae) were nested
together contrary of Melchior (1964), Endress et al. (2014) and El-gazzar et al. (2018) whom separated
them; W. coccinea in Apocynoideae and C.
grandiflora in Priplocoideae.
A. obesum and N. oleander (Apocynoideae)
were
separated at basal level, while A. curassavica, G.
sinaicus, C. procera, C.
acutum, S. argel and H. andreaeana (Asclepiadoideae) were clustered together. This clade is corresponding to
tribe Stapeliae passing with C. acutum as a separate clade (Chase et al.
2016), C. acutum is delimited from all the studied species of Asclepiadeae, reaching G. sinaicus
and A. curassavica that forming sister species
(Sennblad and Bremer 1996). S. argel (Glossonematinae)
is considered a sister group to C. procera, A.
curassavica and G. sinaicus
(Asclepiadinae) (Melchior
1964). From the pattern of the fourth lineage, there is a
strong support to the monophylly of tribe Asclepiadoideae
(Chase et al. 2016; El-gazzar et al. 2018).
Melchior (1964) divided Apocynaceae s. l. into two distinct families and two
sub-families for each; Apocynaceae (Plumieroideae and Echitoideae)
and Asclepiadaceae (Cynanchoideae and Periplocoideae). Apocynaceae and Asclepiadaceae have been treated as one
family and are recognized five sub-families; three of Asclepiadaceae (Periplocoideae, Asclepiadoideae
and Secamonoideae) and two of Apocynaceae
s.s. (Rauvolfiodeae
and Apocynoideae) (Endress 2004; Endress et al. 2007; Endress et al.
2014; Chase et al. 2016). The obtained cladogram nested studied species of Asclepiadoideae in a separate lineage and the remaining
studied species of Periplocoideae, Apocynoideae and Rauvolfioideae
into three lineages. The phylogenetic analysis in the present study supports
the treatment of Apocynaceae and Asclepiadaceae as
one family (Apocynaceae s.l.)
with four subfamilies, which is accirding to previous
reports (Endress 2004; Endress
et al. 2014; Chase et al. 2016).
Conclusion
The phenetic analysis in the present study supports
the treatment of Apocynaceae s.l. as two distinct families Apocynaceae s.s. and
Asclepiadaceae contrary of the phylogenetic one that supports the
treatment of Apocynaceae s.s. and Asclepiadaceae
as one family (Apocynaceae s.l.)
with four subfamilies viz. Apocynoideae, Rauvolfiodeae, Asclepiadoideae
and Periplocoideae.
Author Contributions
MS drafted the manuscript and statistically analyzed the
data, AM analyzed and interpreted the results, MT planned the study conception
and design and made critical revision, HD acquainted the data and UA made
illustrations, made the write up and responded to journal reviewers.
Conflict of Interest
The authors declare that they have no known competing
financial interests or personal relationships that could have appeared to
influence the work reported in this paper.
Data
Availability Declaration
Authors declare that all data are available upon request
from corresponding author
Conformation
to Ethical Guidelines
All material that were used in
the current research do not need to ethically approved, human or animal materials
not included
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%201111-1119_files/image003.jpg)
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%201111-1119_files/image005.jpg)
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%201111-1119_files/image007.jpg)
Fig. 3: Neighbor joining phylogenetic tree of the studied taxa of Apocynacaeae s.s. based
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