Nordic Journal of Botany 29: 518533, 2011 doi: 10.1111/j.1756-1051.2011.01104.x, # 2011 The Authors. Nordic Journal of Botany # 2011 Nordic Society Oikos Subject Editor: Arne Strid. Accepted 22 February 2011
Astragalus raphaelis (Fabaceae), a critical species from Sicily and taxonomic remarks on A. sect. Sesamei Cristian Brullo, Salvatore Brullo, Gianpietro Giusso del Galdo, Pietro Minissale and Saverio Sciandrello C. Brullo, S. Brullo (
[email protected]), G. Giusso del Galdo, P. Minissale and S. Sciandrello, Dipartimento di Botanica, Univ. degli studi di Catania, Via A. Longo 19, IT-95125 Catania, Italy.
The taxonomic position of Astragalus raphaelis, a rare and misappreciated species endemic to Sicily, has been examined. In addition, its geographical distribution, ecology and conservation status, diagnostic morphological characters, pod and seed micro-morphology and relationship with A. stella are provided. Phenetic and phylogenetic trees (based on morphological data) for A. sect Sesamei shows that A. raphaelis is a taxonomically quite isolated species, with some relationships to A. sesamoides.
The aim of this study is to taxonomically investigate Astragalus raphaelis G. Ferro, a critical species that is endemic to Sicily. It was described by Ferro (1978) from a population in Butera (south Sicily), that grow on steep and heavily eroded clayey badlands. Later, new records of this therophyte from other Sicilian localities were reported by Romano et al. (1994), Raimondo et al. (1994, 1995, 2004) and Minissale et al. (2009). Astragalus raphaelis belongs to A. sect. Sesamei DC., in which several annual species usually found in xeric environments are included, such as A. sesameus L., A. sinaicus Boiss., A. stella L., etc. (Gazer 1993). Ferro (1978) considered A. raphaelis to be closely related to A. sinaicus, from which it chiefly differs in having long pedunculate inflorescence, and different shape and arrangement of the legumes. Neither Pignatti (1982) nor Gazer (1993) reported A. raphaelis in their lists, while it was included by Conti et al. (2005) and Giardina et al. (2007). Recently, Podlech (2008) treated A. raphaelis as a synonym of A. stella, probably because of its long pedunculate inflorescence. In order to clarify the taxonomic position and phylogenetic relationships of A. raphaelis within A. sect. Sesamei, we did an in-depth survey focused on the macro- and micromorphological characters of this species. In particular, we compared A. raphaelis with closely allied species belonging to this section, and especially with A. stella.
Material and methods The morphological study is based on literature, herbarium and field investigations. Several herbarium specimens of 518
A. raphaelis and allied species, preserved in B, CAT, FI, G, M and PA, have been examined. The micro-morphology of seed testa and pod indumentum of A. raphaelis and A. stella was studied on dried material with the aid of a scanning electron microscope (SEM) Zeiss EVO LS10 (Huttunen and Laine 1983). A comparative scheme of the main morphological characters of the species belonging to A. sect. Sesamei is provided in Table 1. It is based on observations of herbarium specimens and literature data (Eig 1955, Chamberlain and Matthews 1970, Zohary 1972, Jafri 1980, Gazer 1993, Boulos 1999, Podlech 1999a, 1999b, 2008). The phenetic analysis was performed by using 31 morphological characters (Table 2). The selection was made considering the diacritic value of each character, mainly due to the occurrence of discontinuities among them. Eighteen characters used for the phenetic analysis were coded as binary, while 13 were coded as multi-state. The data matrix of the characters used for the phenetic analysis is shown in Table 3, in which all the species belonging to A. sect. Sesamei were examined with the exception of A. biovulatus Bunge, as it was not possible to find detailed information on its morphology. The morphological dataset was processed by the NTSYSpc package (Rohlf 2000). The same morphological dataset was also processed using parsimony as the optimality criterion with the programme PAUP* (Swofford 2003), thus highlighting the phylogenetic relationships among the species of this section. Character evolution was examined using the programme MacClade 4.05 PPC (Maddison and Maddison 1992). All characters were unordered and equally weighted (Table 4). The data matrix of the morphological characters used for the phylogenetic analysis is shown in Table 5. In addition to the 22 species surveyed in the present study, one species
Table 1. Main differential characters of the species of Astragalus sect. Sesamei. Asc. ascending, prost. prostrate. A. stella Stem habit Stem length (cm) Stem hairs (patent or appressed) Stem hairs length
A. raphaelis
A. triradiatus
A. tribuloides
A. transjordanicus
A. tadmorensis
A. sinaicus
A. sesamoides
A. sesameus
A. scorpioides
A. schimperi
prost.-asc. 436(50) appr.
asc.-erect 520 appr.
prost.-asc. 620 pat.
prost.-asc. 540 appr.
asc.-erect 1030 pat.
prost.-asc. 715 appr.
asc.-erect 525 pat.
prost.-asc. 736 appr.
prost.-asc. 736 appr.
prost.-asc. 440 pat.
prost.-asc. 1030 appr.
0.31.9
0.31
0.51.2
0.21.5
0.30.7
0.30.9
0.31.2
0.31
0.31.8
0.31.2
0.41.5
loose
dense
dense
loose
loose
loo or den
loose
loose
dense
loose
dense
adnate
adnate
connate
adnate
adnate
adnate
connate
adnate
adnate
adnate
adnate
green
membr.
green
membr.
membr.
green
green
membr.
green
membr.
green
Stipule length (mm) Stipule hairs (white or whitish-black) Stipule shape (triangular or lanceolate)
36(7) whitishblack triang.
23 whitishblack lanc.
45 whitishblack triang.
23 whitishblack triang.
12.5 white triang.
22.5 whitishblack triang.
45 whitishblack triang.
34 whitishblack triang.
36 whitishblack triang.
35 whitishblack triang.
24 whitishblack triang.
Stipule apex (acuminate or acute) Leaf (sessile or petiolate) Leaf length (cm) Leaf hairs (white or
acum.
acum.
acum.
acum.
acum.
acute
acum.
acum.
acum.
acum.
acum.
petiol.
petiol.
petiol.
petiol.
petiol.
sess.
petiol.
petiol.
petiol.
petiol.
petiol.
27.5(9) white
2.59 white
2.56 white
1.57(9) white
26.5 white
23.5 white
27 white
2.55.5 whitish-black
28 (95) whitish-black
3-8 white
1.54 white
whitish-black) Leaflet pairs Leaflet apex
710(11) obtuse
711 rounded
57(9) obtuse
49(10) acute
59 obtuse
58 rounded
810(12) obtuse
2.510(16) 1.53.5(5)
2.510 13.5
410 1.54
2.59(15) 0.82(4)
1.44 0.81.5
24.5 11.5
210 0.84
610(12) obtuse or retuse 310(14) 13.2(5.5)
69 rounded to retuse 513 14
(4)68 obtuse
Leaflet length (mm) Leaflet width (mm)
46 rounded to obtuse 415 25.5
Leaflet indumentum (upper side) Leaflet shape
hairy
hairy
hairy
hairy
hairy
hairy
hairy
hairy
hairy
glabrous
hairy
ellipt. to obl. pedun.
ellipt.
ellipt. to ov. ellipt.
ellipt.
ellipt.
ellipt.
ellipt. to obl.
obl. to obov.
pedun.
pedun.
pedun.
pedun.
pedun.
pedun.
pedun.
sess. or subsess.
ellipt. to ellipt. to obov. obl. sess. or sub- pedun. sess.
Inflorescence (number of flowers) Inflorescence peduncle length (cm) Inflorescence hairs
411(15)
312
36
28
24
34
37
39
310
1(2)
25
211.5
1.54
15
02.5
12.5(3.5)
13
0.55
26
00.5
00.3
0.54.2
whitish-
white
whitish-
white
whitish-black
whitish-
whitish-black
white
white
white
whitish-
(white or whitish-black) Bract length (mm) Bract apex (acuminate or acute)
black 1.53 acum.
34.5 acute
1.52.5 acute
12(3) acute
11.5 acute
0.7 acute
12.5 acum.
23 acute
13 acute
45 acute
1.52(2.5) acum.
Bract shape (lanceolate, traingular, linear) Bract hairs (white or whitish-black) Flower (sessile or peduncolate)
lanc.
lanc.
triang.
triang.
linear
triang.
triang.
triang.
lanc.
triang.
triang.
whitishblack sess.
whitishwhitishblack black sess. or sub- sess. sess.
white
whitish-black
whitish-black
whitish-black
whitish-black
sess. or subsess.
sess.
whitishblack sess.
sess.
pedun.
sess.
whitishblack sess.
whitishblack sess.
Calyx length (mm) Calyx shape (tubular or campanulate) Calyx tooth shape (subulate, triangular or
4.57 tubul.
6.57.5 tubul.
56 camp.
45 tubul.
56 tubul.
34 camp.
4.36 tubul.
5.57 camp.
57 camp.
68 tubul.
57(8) tubul.
subul.
subul.
subul.
subul.
subul.
subul.
subul.
subul.
linear
subul.
subul.
1.23(3.8)
34.5
1.32.3
12
23
0.70.9
1.31.6
34
23
45
1.72.1
shorter
longer
shorter
shorter
subequal
shorter
shorter
longer
shorter
longer
shorter
whitishblack
whitishblack
whitishblack
white
whitishblack
whitishblack
whitishblack
whitishblack
whitishblack
whitishblack
whitishblack
(mm) Stem hairs (loose or dense) Stipules (adnate, connate or free) Stipules (green or membranaceous)
Inflorescence (sessile or peduncolate)
linear) Calyx tooth length (mm) Calyx teeth (longer/ shorter than tube) Calyx hairs (white or whitish-black)
black
black
310(12) 1.53
black
519
Table 1 (Continued)
A. saharae
A. pseudosinaicus
A. persepolitanus
A. longicaulis
A. kerkukiensis
prost.-asc.
Stem habit
asc.-erect
prost.-asc.
prost.-asc.
prost.-asc.
Stem length (cm) Stem hairs (patent or appressed) Stem hairs length (mm)
630 appr.
520 pat.
3.527 appr.
837 appr.
acaul. or prost.-asc. 05.5 appr.
0.21.3
0.41.3
0.10.3
0.31
Stem hairs (loose or dense) Stipules (adnate, connate or free) Stipules (green or membranaceous)
loose
unknown
dense
connate
connate
green
Stipule length (mm) Stipule hairs (white or whitish-black) Stipule shape (triangular or lanceolate) Stipule apex
A. gryphus
A. geniculatus
A. asterias
A. filicaulis
A. cornubovis
A. coronilla
prost.-asc.
asc.-erect.
prost.-asc.
prost.-asc.
715 pat.
acaul. or prost.-asc. 020 pat.
740 appr.
827 appr.
1526 appr.
515 pat.
0.20.7
0.31.5
0.52.3
0.21
0.31.1
0.50.7
0.20.5
dense
loose
loo or den
dense
dense
loose
loose
dense
adnate
adnate
adnate
adnate
adnate
adnate
adnate
adnate
adnate
membr.
membr.
green
membr.
green
green
green
green
green
membr.
23 whitishblack triang.
34 white
24.5 whitishblack triang.
1.52 white
45 white triang.
23.5 whitishblack triang.
2.54 whitishblack triang.
22.5 white
triang.
3.55 whitishblack triang.
12.5 white
triang.
12(2.5) whitishblack triang.
triang.
triang.
acum.
acum.
acute
acum.
acum.
acum.
acum.
acum.
acum.
acute
acum.
petiol.
sess.
petiol.
sess.
petiol.
petiol.
sess.
sess.
petiol.
petiol.
petiol.
1.54 white
2.54 white
27 white
26(7) white
312(15) white
2.54 white
25.5 white
24.5 white
58 white
2.55 white
35 (8.5) white
whitish-black) Leaflet pairs Leaflet apex
69 obtuse
36(7) retuse
610 retuse
(6)710 obtuse
(2) 34 retuse
35 (7) retuse
28.5 0.83 hairy
311 13 hairy
38 13 hairy
410 1.54 hairy
59 acute-obtuse 1.513 0.83.2 hairy
57 obtuse
29(11) 12.5(3) hairy
47 rounded to retuse 312(15) 1.54.5(6) glabrous
58 rounded
Leaflet length (mm) Leaflet width (mm) Leaflet indumentum
611 obtuse or retuse 58 24 hairy
29(13) 12.8 hairy
39.5 26 glabrous
312 35 hairy
ellipt. to obov.
obl. to obov.
oblong
obl. to obov.
ellipt.
ellipt. to obl. ellipt.
ovate
pedun.
pedun.
pedun.
(3)511
sess. or subsess. 24
pedun.
310
sess. or subsess. 36
26
28 (10)
39
sess. or subsess. 37
cun.to obov. pedun.
37
sess. or subsess. 36
ellipt. to obl. pedun.
15
01
0.57
15.5
00.6
00.3
1.55
1.54.5
1.56
00.5
0.55 (7)
whitishblack
white
whitish-black
whitishblack
white
whitishblack
whitishblack
whitishblack
whitishblack
white
white
12 acute
12 acum.
0.81.2 acute
12 acute
11.2 acute
35 acute
13 acute
12 acute
11.5 acute
1 acute
0.5 acum.
triang.
triang.
triang.
lanc.
triang.
triang.
triang.
triang.
lanc.
lanc.
triang.
whitishblack pedun.
white
whitish-black
white sess.
whitishblack sess.
whitishblack sess.
whitishblack pedun.
white
pedun.
whitishblack sess.
white
sess.
whitishblack sess.
sess.
pedun.
Calyx length (mm) Calyx shape (tubular or campanulate) Calyx tooth shape (subulate, triangular or linear)
56 camp.
45 camp.
2.53 camp.
45 camp.
2.53 camp.
45 camp.
6 camp.
34 camp.
34 camp.
3 camp.
2.53 camp.
subul.
subul.
subul.
subul.
subul.
subul.
subul.
subul.
subul.
triang.
subul.
Calyx tooth length (mm) Calyx teeth (longer/ shorter than tube) Calyx hairs (white or whitish-black)
1.51.8
1.52
0.81
1.52
1.51.7
22.5
22.5
11.3
1.82
0.80.9
0.51
shorter
shorter
shorter
shorter
longer
subequal
shorter
shorter
subequal
shorter
shorter
whitishblack
white
whitishblack
whitishblack
white
whitishblack
whitishblack
whitishblack
whitishblack
white
white
(acuminate or acute) Leaf (sessile or petiolate) Leaf length (cm) Leaf hairs (white or
(upper side) Leaflet shape Inflorescence (sessile or peduncolate) Inflorescence (number of flowers) Inflorescence peduncle length (cm) Inflorescence hairs (white or whitish-black) Bract length (mm) Bract apex (acuminate or acute) Bract shape (lanceolate, traingular, linear) Bract hairs (white or whitish-black) Flower (sessile or peduncolate)
520
ellipt. to obl. pedun.
26
Table 1 (Continued)
A. stella
A. raphaelis
A. triradiatus
A. tribuloides
A. transjordanicus
A. tadmorensis
A. sinaicus
A. sesamoides
A. sesameus
A. scorpioides
A. schimperi
Calyx hairs length (mm) Corolla colour
0.51.5
0.41.2
0.51.2
0.51.5
0.51
0.30.5
0.51.4
0.41
0.51.8
0.31.2
0.41.5
yellow to violet
pale blue
violet
violet
violet
yellow to violet
violet
yellow to lilac
yellow to blue
white to pale violet
Corolla (longer/
longer
shorter
longer
white to pinkishwhite longer
longer
longer
longer
shorter
longer
longer
longer
shorter than calyx) Standard shape
ellipt.
obov.-spat.
ellipt.
oblong
obl-rhomb
obov
ellipt.
ellipt.
ellipt.
ellipt.
Standard length (mm)
710(11)
5.56.5
811
410
912
89
68.2
5.58
79
911
rhom to obov 810(11)
Standard width (mm) Standard apex Wing length (mm) Wing limb shape Wing limb apex (emarginate or rounded)
24 retuse 68 oblong rounded
2.52.7 retuse 5.56 oblong rounded
33.5 retuse 67.5 oblong rounded
1.22.5 retuse 46 oblong retuse
34 retuse 8 oblong rounded
33.5 retuse 67 oblong rounded
23 retuse 57 oblong obtuse
22.5 retuse 45(5.5) oblong obtuse
23 retuse-apic. 6.27 oblong retuse
3 retuse 810 oblong retuse
2.53 retuse 78(8.5) oblong retuse
Wing limb length (mm) Wing claw length (mm) Keel length (mm) Keel limb shape
34
3.53.7
2.54.5
1.52.8
4
2.83
2.83.2
22.5
3.24
45
33.5(4)
34
22.3
33.5
23
34
2.83
2.33
22.5
33.5
4
3.54
56 triang.-
4.55 obov.-spat.
4.55 triang.-
35 triang.-
6 triang.-obov.
4 obov.
45 triang.-obov.
44.5 triang.-obov.
55.3 triang.-obov.
77.5 ellipt.
5.56.2 triang.-
obov. acute 2.52.7
obtuse 22.2
obov. acute 1.52
obov. obtuse 12.5
obtuse 3
obtuse 2
obtuse 1.51.8
acute 1.51.8
acute 2.5
acute 3.54
obov. obtuse 22.8
2.53.5 cut lanceolate
2.23 cut oblong
3 curved ovate
22.5 cut obl.-triang.
3 curved lin. to lanc.
2 23 cut cut lin. to ellipt. obl. to obov.
2.22.7 curved lin.-obl.
2.52.8 cut linear
3.54 cut linear
34.2 cut oblong
1015 2.53 patent long curved double
912 34 erect long curved double
1523 44.5 erect long curved double
49(12) 1.54 patent short acute double
1525 56 erect long curved single
912 33.5 patent short acute double
1018 2.54 erect short acute double
1215 3.54 erect long curved single
917 4.55 erect short curved double
3050 33.5 erect long curved double
1522 34 patent long curved double
appr. to pat.
pat.
pat.
appr.
pat.
appr.
appr. to pat.
appr.
appr. to pat.
appr.
appr.
47
35
7
410(12)
67
45
57
46
46(7)
many
1011(14)
0.050.2
0.10.2
0.10.4
0.050.15
0.10.2
0.10.2
0.10.2
0.050.3
0.10.3
0.52
0.51.3
1.32
0.51.5
1.52.3
0.51.4
0.51.2
0.41.3
11.6
0.50.7
0.52.4
not
yes
yes
yes
yes
yes
yes
not
not
not
yes
bigibbous
rounded
rounded
bigibbous
rounded
bigibbous
bigibbous
rounded
rounded
rounded
rounded
reptangular 11.5 1.21.5 brown
reniform 1.52 1.62 olivaceous,
reptangular 23 1.52 brown
reniform 12 11.5 pale brown
reniform 33.5 22.5 brown
reptangular 2 1.5 brown
Quadrangular quadrangular 1.11.5 1.51.8 0.91.2 1.51.8 Brown yellow
quadrangular 1.22 11.8 yellow
reptangular 3-3.5 22.5 brown
globose 11.4 11.4 brown
smooth
smooth
smooth
rugose
Keel limb apex Keel limb length (mm) Keel claw length mm Staminal tube Legume shape Legume Legume Legume Legume Legume Legume
length (mm) width (mm) arrangement apex size apex shape
indumentum Legume indument (patent or appressed) Number of seeds per locule Legume short hair length (mm) Legume long hair length (mm) Legume hair partly sitting on little tubercles Legume bigibbous or rounded at base Seed shape Seed length (mm) Seed width (mm) Seed colour Seed smooth surface
smooth
dotted rugose
smooth
smooth
Smooth
smooth
smooth
521
Table 1 (Continued)
A. saharae
A. pseudosinaicus
A. persepolitanus
A. longicaulis
A. kerkukiensis
A. gryphus
A. geniculatus
A. asterias
A. filicaulis
A. cornubovis
A. coronilla
Calyx hairs length (mm) Corolla colour
0.21.3
0.41.3
0.050.6
0.40.9
0.10.3
0.51.5
0.61.2
0.20.6
0.31
0.5
0.20.5
yellow
yellow
white to pale violet
violet
white
yellow
white to pinkishwhite
mauvecream
violet
w
yellowpink
Corolla (longer/shorter than calyx) Standard shape
longer
longer
longer
longer
longer
longer
longer
longer
longer
longer
longer
obl to ov
ellipt.
ellipt.
ellipt.
ellipt.
ellipt.
ellipt.
nar.-ovate
Standard length (mm) 67.5
67
4.56(7)
610
45
7
8.512
narr.-ellip- elliptictic oblong 69 mm 68
6
nar.elliptic 57
Standard width (mm) Standard apex Wing length (mm) Wing limb shape
1.82.9 retuse 56.3 oblong
2.52.7 retuse 56 oblong
2 rounded 44.5 oblong
34 retuse 67.5 oblong
1.52 retuse 3.54 oblong
2.5 retuse 6 oblong
3.55 retuse 79 oblong
2.23.5 retuse 45 (6) oblong
22.5 retuse 4.55 oblong
Wing limb apex (emarginate or
retuse
retuse
obtuse
obtuse
obtuse
retuse
retuse
retuse
retuse
2 retuse 55.5 nar.oblong obtuse
2.5 retuse 4.56 nar.oblong retuse
23
23
2.22.5
2.83.5
22.3
3.5
35
23
3
33.3
34
22.5
2.53
1.82
2.53.5
1.31.7
2.5
33.5
22.5
1.82
1.52.2
12
45
4.5
33.8
45
2.83.3
5
6
(3) 44.5
3.54
5
44.5
triang.obov. obtuse 1.22.5
triang.-obov.
triang.-obov.
obov.
triang.-obl.
oblong
triang.-obov.
obov.
obov.
obov.
acute 1.51.8
obtuse 2.2
acute 1.51.8
rounded 1.8
acute 2.5
obtuse 3
triang.obov. acute 1.51.8
rounded 2
rounded 2.5
acute 22.5
22.5 cut
3 cut
1.82 cut
23.2 cut
1.31.5 curved
2.5 cut
3 cut
22.5 cut
1.52 cut
1.52.2 curved
2 cut
Legume shape
lin.-obl.
ov.-obl.
ov.-triang.
oblong
ovate
linear
oblong
1115(18) 33.5 erect long
1012 33.5 erect long
69 33.2 unknown long
915 2.53 patent long
6.57 33.5 erect short
1518 4 erect long
1216(19) 2.53 erect long
oblong to ovate 67(10) 3 patent short
lin.-oblonglanc. 1216 33.5 erect short
nar.-ovate
Legume Legume Legume Legume
lin.-oblong 1015 23 patent short
curved double
curved double
curved double
acute single
curved double
curved double
acute double
acute double
acute single
acuminate single
appr.
appr.
curved glabrous or single appr.
appr.
appr.
pat.
pat.
appr.
pat.
appr.
appr. to pat.
78
78
2
68
2
45
35
710
23
5
34
0.050.15
0.10.2
0.050.15
0.050.3
0.10.2
0.10.2
0.10.3
0.51.3
11.3
0.50.6
0.51.2
0.10.5
0.51.6
0.52.3
1
11.5
0.150.8
0.20.5
yes
yes
not
not
not
yes
not
yes
not
not
not
rounded
rounded
bigibbous
bigibbous
rounded
rounded
rounded
bigibbous
bigibbous
rounded
rounded
reptangular 1.82
reptangular 2
reptangular 1.22
reptangular 1.21.8
reptangular 22.5
reptangular 3.5
reptangular 23
reniform 22.5
reptangular 1.21.5
reniform 2.2
reptangular 33.2
1.21.4 unknown rugose
1.5 unknown smooth
11.8 yellowish unknown
11.5 yellowish smooth
1.82 yellowish smooth
3 brown smooth
2-2.5 brown smooth
12.2 yellow smooth
11.2 brown unknown
2 brown smooth
2 brown smooth
rounded) Wing limb length (mm) Wing claw length (mm) Keel length (mm) Keel limb shape Keel limb apex Keel limb length (mm) Keel claw length mm Staminal tube
length (mm) width (mm) arrangement apex size
Legume apex shape Legume indumentum Legume indument (patent or appressed) Number of seeds per locule Legume short hair length (mm) Legume long hair length (mm) Legume hair partly sitting on little tubercles Legume bigibbous or rounded at base Seed shape Seed length (mm) Seed width (mm) Seed colour Seed surface
522
1016 23 erect long
Table 2. Morphological characters and character state codes used for the phenetic analysis. No. Character
States
01
0 0.5; 0.5 1; 1 1
02 03 04 05 06 07 08 09 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31
Stem hairs max length (mm) Stipules Stipule texture Stipule hairs Stipule apex Leaf hairs Leaflet max length (mm) Leaflet indumentum (upper side) Inflorescence
0 adnate or free; 1 connate 0 green; 1 membranaceous 0 white; 1 white and black 0 acute; 1 acuminate 0 white; 1 white and black 0 6; 0.5 10 ; 1 10 0 glabrous; 1 hairy
0 sessile or subsessile; 1 peduncolate Inflorescence max number 0 1; 0.5 6; 1 6 of flowers Inflorescence hairs 0 white; 1 white and black Bract length (mm) 0 0-0.7; 0.5 1-3; 1 3-5 Bract apex 0 acute; 1 acuminate Bract hairs 0 white; 1 white and black Calyx length (mm) 0 2.5-3; 0.5 3-4; 1 4-8 Calyx teeth length (mm) 0 0.7-1; 0.5 1-3; 1 3-5 Calyx teeth/tube 0 short; 0.5 subequal; 1 longer Calyx hairs 0 white; 1 white and black Corolla (exserted from 0 slightly; 1 long calyx) Standard apex 0 retuse; 0.5 retuse-apiculate; 1 rounded Wing limb apex 0 retuse; 0.5 obtuse; 1 rounded Keel length (mm) 0 2.8-5; 1 5-7.5 Staminal tube 0 cut; 1 curved Legume length (mm) 0 4-9; 0.5 9-25; 1 30 Legume arrangement 0 erect; 1 patent Legume apex 0 acute; 1 curved Legume indumentum 0 simple; 1 double Number of seed per locule 0 1-3; 0.5 3-10; 1 10 Legume base 0 rounded; 1 bigibbous Seed shape 0 globose; 0.5 kidney; 1 rectangular Seed length (mm) 0 1-2; 0.5 2-3; 1 3-3.5
covered with white and black subpatent hairs. Leaves petiolate, 2.59.0 cm; rachis covered with white hairs. Leaflets green, 711-paired, 2.510.0 1.02.3 mm, elliptical, with appressed or sub-appressed hairs on both sides, rounded (or rarely obtuse) at apex. Inflorescences with a peduncle 1.54.0 cm long, hairy like the stem, with 312 flowers in a compact head. Bracts membranaceous, 3.04.5 mm long, lanceolate, acute, with subpatent white and black hairs, 0.81.2 mm long. Flowers sessile or sub-sessile. Calyx 6.57.5 mm long, tubular, covered with subpatent hairs, 0.41.2 mm long, white and black on the teeth, usually white in the tube; teeth subequal, subulate, 3.04.5 mm long, longer than tube. Corolla pale blue, slightly longer than calyx. Standard obovate-spathulate, 5.56.5 2.52.7 mm, retuse-apiculate at apex. Wings 5.56.0 mm long; limb oblong, 3.53.7 1.01.2 mm, rounded at apex, with auricle 0.8 mm long; claw 2.02.3 mm long. Keel 4.5 5.0 mm long; limb obovate, obtuse at apex, 2.02.2 1.2 1.4 mm; claw 2.23.0 mm long. Staminal tube straightly cut, 3 mm long, with filaments 0.40.9 mm long and anthers 0.150.20 mm long, rounded to obtuse at apex. Ovary short pedunculate, 3.03.5 mm long, with densely appressed hairs, 0.20.3 mm long; style glabrous, 1.01.2 mm long; stigma conic. Legumes erect, oblong, 912 34 mm, blackish, rounded at the base, ending in a patent straight beak, 1.01.2 mm long, deeply grooved dorsally and rounded ventrally, covered with two kinds of hairs: the short ones sub-appressed, 0.10.2 mm long, the long ones patent, 0.51.3 mm long, sitting on a little tubercle. Seeds compressed, 35 in each locule, kidney-shaped, sometimes sub-quadrangular, 1.52.0 1.62.0 mm, pale brown, minutely dotted, rugose. Iconography
Ferro (1978, Fig. 1). Distribution and ecology
(A. echinatus Murray) not belonging to the cycle of A. sect. Sesamei was included as an outgroup. The resulting most parsimonious trees were used to calculate a strict consensus tree. Branch support was evaluated using a multiple bootstrap analysis (Felsenstein 1985) with 1000 replicates, simple taxon addition and TBR branch swapping and collapse branch if minimum length is zero. Groups were retained with bootstrap percentage 50%.
Results Astragalus raphaelis G. Ferro (1978, p. 45) (Fig. 1, Fig. 2, Fig. 3) Type: Sicilia. S. Nicola (Comune di Butera), 210 m a.s.l., campi abbandonati su substrato argilloso, 7 Apr 1974, Ferro 3183/8 (holotype: CAT). Annual, branched at the base, hairy. Stems leafy, 520 cm tall, with the main stem erect and numerous ascending basal stems, densely covered with appressed to sub-patent white hairs, 0.31.0 mm long. Stipules membranaceous, lanceolate, acuminate, 23 mm long, adnate to the petiole,
Based on literature, herbarium and field investigations, A. raphaelis is currently known from only a few localities in centralsouthern Sicily (Fig. 4). The species was first described from Butera (Ferro 1978), but was later found by various authors at other Sicilian localities, such as Alimena, Serradifalco and Petralia Sottana (Romano et al. 1994, Raimondo et al. 2004, Minissale et al. 2009). Recently, we found additional populations of A. raphaelis near Diga Comunelli (Gela), and Cozzo Niglio near Agira. It occurs between 80 and 230 m of altitude, within the thermo-mediterranean drysub-humid bioclimatic belt. It is usually a member of thermoxerophilous annual plant communities on clayey substrata, represented by budlands, chiefly colonized by Lygeum spartum L. grasslands. In these habitats many other therophytes, ecologically very specialized, are also present, such as Brassica souliei (Batt.) Batt. subsp. amplexicaulis (Desf.) Greuter & Burdet, Parapholis pycnantha (Hack.) C. E. Hubbard, Catananche lutea L., Astragalus echinatus, Podospermum canum C. A. Mey., Anthemis muricata (DC.) Guss., etc. From the phytosociological viewpoint, this community can be ascribed to the GaudinioPodospermion cani, an alliance of Saginetea maritimae, a class grouping annual haloxerophilous communities (Brullo and Giusso del Galdo 2003). 523
A. raphaelis
A. triradiatus
A. tribuloides
A. transjordanicus
A. tadmorensis
A. sinaicus
A. sesamoides
A. sesameus
A. scorpioides
A. schimperi
A. saharae
A. pseudosinaicus
A. persepolitanus
A. longicaulis
A. kerkukiensis
A. gryphus
A. geniculatus
A. asterias
A. filicaulis
A. cornubovis
A. coronilla
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31
A. stella
524
Table 3. Data matrix for the 31 selected characters used for the phenetic analysis of the investigated taxa of Astragalus sect. Sesamei. The character codes are explained in Table 2.
1 0 0 1 1 0 0.5 1 1 1 1 0.5 1 1 1 0.5 0 1 1 0 1 1 0 0.5 1 1 1 0.5 1 1 0
0.5 0 1 1 1 0 0.5 1 1 1 0 1 0 1 1 1 1 1 0 0.5 1 0 0 0.5 0 1 1 0.5 0 0.5 0
1 1 0 1 1 0 0.5 1 1 0.5 1 0.5 0 1 1 0.5 0 1 1 0 1 0 1 0.5 0 1 1 0.5 0 1 0.5
1 0 1 1 1 0 0.5 1 1 1 0 0.5 0 0 1 0.5 0 0 1 0 0 0 0 0 1 0 1 0.5 1 0.5 0
0.5 0 1 0 1 0 0 1 1 0.5 1 0.5 0 1 1 0.5 0.5 1 1 0 1 1 1 0.5 0 1 0 0.5 0 0.5 1
0.5 0 0 1 0 0 0 1 1 0.5 1 0 0 1 0.5 0 0 1 1 0 1 0 0 0.5 1 0 1 0.5 1 1 0
1 0 1 1 0 0.5 1 1 1 1 0.5 1 1 1 0.5 0 1 1 0 0.5 0 0 0.5 0 0 1 0.5 1 1 0
0.5 0 1 1 1 1 1 1 1 1 0 0.5 0 1 1 1 1 1 0 0 0.5 0 1 0.5 0 1 0 0.5 0 1 0
1 0 0 1 1 1 0.5 1 0 1 0 0.5 0 1 1 0.5 0 1 1 0.5 0 1 0 0.5 0 1 1 0.5 0 1 0
1 0 1 1 1 0 1 0 0 0 0 1 0 1 1 1 1 1 1 0 0 1 0 1 0 1 1 1 0 1 1
1 0 0 1 1 0 0.5 1 1 0.5 1 0.5 1 1 1 0.5 0 1 1 0 0 1 0 0.5 1 1 1 1 0 0 0
1 1 0 1 1 0 0.5 1 1 1 1 0.5 0 1 1 0.5 0 1 1 0 0 0 0 0.5 0 1 1 0.5 0 1 0
1 1 1 0 1 0 0.5 1 0 0.5 0 0.5 1 0 1 0.5 0 0 1 0 0 0 0 0.5 0 1 1 0.5 0 1 0
0 0 1 1 0 0 0.5 1 1 1 1 0.5 0 1 0 0 0 1 1 1 0.5 0 0 0 1 1 0 0 1 1 0
0.5 0 0 1 1 0 1 1 1 1 1 0.5 0 1 1 0.5 0 1 1 0 0.5 0 0 0.5 1 1 1 0.5 1 1 0
0.5 0 1 0 1 0 1 0 0 0.5 0 0.5 0 0 0 0.5 1 0 1 0 0.5 0 1 0 0 0 0 0 0 1 0.5
1 0 0 0 1 0 0.5 1 0 0.5 1 1 0 1 1 0.5 0.5 1 1 0 0 0 0 0.5 0 1 1 0.5 0 1 1
1 0 0 1 1 0 0.5 1 1 0.5 1 0.5 0 1 1 0.5 0 1 1 0 0 1 0 0.5 0 1 1 0.5 0 1 0.5
0.5 0 0 1 1 0 1 1 1 1 1 0.5 0 1 0.5 0.5 0 1 1 0 0 0 0 0.5 1 0 1 0.5 1 0.5 0.5
1 0 0 1 1 0 0.5 1 1 1 1 0.5 0 1 0.5 0.5 0.5 1 1 0 0 0 0 0 1 0 1 0 1 1 0
0.5 0 0 0 0 0 0.5 0 0 1 0 0.5 0 0 0 0 0 0 1 0 0.5 0 1 0.5 0 0 0 0.5 0 0.5 0.5
0 0 1 0 1 0 1 1 1 0.5 0 0 1 0 0 0 0 0 1 0 0 0 0 0.5 0 0 0 0.5 0 1 1
Table 4. Morphological characters and character state code used for the phylogenetic analysis. No.
Character
01 02 03 04 05 06 07 08 09 10 11 12 13 14 15 16 17 18 19 20
Stem hairs max length (mm) Stipules Stipule texture Stipule hairs Stipule apex Leaf hairs Leaflet max length (mm) Leaflet indumentum (upper side) Inflorescence Inflorescence max number of flowers Inflorescence hairs Bract length (mm) Bract apex Bract hairs Calyx length (mm) Calyx teeth length (mm) Calyx teeth/tube Calyx hairs Corolla (exserted from calyx) Standard apex
21 22 23 24 25 26 27 28 29 30 31
Wing limb apex Keel length (mm) Staminal tube Legume length (mm) Legume arrangement Legume apex Legume indumentum Number of seed per locule Legume base Seed shape Seed length (mm)
Conservation status
The populations in the surveyed sites are generally represented by few individuals (less than 300), with a scattered distribution. Despite their conspicuous fruit production, which is likely due to autogamy as frequently observed in the genus Astragalus, the number of individuals of A. raphaelis remains constantly low. Luckily, no relevant demographic decline was observed in the surveyed populations, but recent field investigations carried out in the ‘locus classicus’ (Butera) showed that this population has definitely disappeared, probably due to intensive agricultural exploitation of this area. Nevertheless, for its rarity and low number of populations, this species should be classified as ‘Critically Endangered’ (CR) (IUCN 2001, 2006). Additional specimens examined
Sicily: Diga Comunelli (Butera), 13 Apr 2007, Brullo and Sciandrello (CAT); Calanchi presso Diga Comunelli (Butera), 28 May 2009, Brullo, Giusso and Sciandrello (CAT); Calanchi presso Agira (Enna), 24 Apr 2009, Minissale and Sciandrello (CAT); Calanchi presso Cozzo Niglio (Agira), 9 Apr 2009, Minissale and Sciandrello (CAT); Cozzo Niglio (Agira), 19 May 2009, Minissale and Sorrentino (CAT).
Astragalus stella L. (1767, p. 734) (Fig. 5) Lectotype: Europae australis potissimum maritimis, D. Gouan (LINN 926.23!), designated by Podlech (cf. Turland and Jarvis 1997).
States 0 0.5; 1 1; 2 1 0 adnate or free; 1 connate 0 green; 1 membranaceous 0 white; 1 white and black 0 acute; 1 acuminate 0 white; 1 white and black 0 6; 1 10 ; 2 10 0 glabrous; 1 hairy 0 sessile or subsessile; 1 peduncolate 0 1; 1 6; 2 6 0 white; 1 white and black 0 0-0.7; 1 1-3; 2 3-5 0 acute; 1 acuminate 0 white; 1 white and black 0 2.5-3; 1 3-4; 2 4-8 0 0.7-1; 1 1-3; 2 3-5 0 short; 1 subequal; 2 longer 0 white; 1 white and black 0 slightly; 1 long 0 retuse; 1 retuse-apiculate; 2 rounded 0 retuse; 1 obtuse; 2 rounded 0 2.8-5; 1 5-7.5 0 cut; 1 curved 0 4-9; 1 9-25; 2 30 0 erect; 1 patent 0 acute; 1 curved 0 simple; 1 double 0 1-3; 1 3-10; 2 10 0 rounded; 1 bigibbous 0 globose; 1 kidney; 2 rectangular 0 1-2; 1 2-3; 2 3-3.5
In order to compare the morphological affinity with A. raphaelis, a detailed description of A. stella is given, as well as information on its ecology, chorology and nomenclature. In particular, for clarification of the taxonomic position of the two species in relation to the taxonomic treatment proposed by Podlech (2008). Annual, unbranched or branched from the base, hairy. Stems leafy, 435(50) cm tall, prostrate to ascending, loosely covered with appressed to subpatent white hairs, often more densely toward the nodes, 0.31.9 mm long. Stipules green, triangular-ovate, acuminate, (2.5)3.06.0(7.0) mm long, adnate to the petiole, covered with white and black patent hairs. Leaves petiolate, 2.59.0 cm; rachis covered with white hairs. Leaflets green, 711-paired, 2.510.0 (16.0) 1.5 3.5(5.0) mm, narrowly elliptic, with appressed hairs on both sides, obtuse to slightly retuse. Inflorescences with a 2.0 11.5 cm long peduncle, hairy like the stem but more densely and with more black hairs toward the raceme, with 411 (15) flowers in a compact head. Bracts membranaceous, 1.53.0(3.5) mm long, ovate-lanceolate, acuminate, with subpatent white and black hairs, 0.41.3 mm long. Flowers sub-sessile. Calyx 4.57.0 mm long, tubular, covered with subpatent white and black hairs, 0.51.5 mm long; teeth subequal, subulate, 1.23.0(3.8) mm long, shorter than the tube. Corolla pinklilac to violet, much longer than the calyx. Standard obovate-spathulate, 710(11) 2.54.0 mm, retuse at apex. Wings 68 mm long; limb oblong, 3 4 1.21.5 mm, rounded at apex, with an 0.50.6 mm long auricle; claw (2.5)3.04.0 mm long. Keel 56 mm long; limb obovate-triangular, acute at apex, 2.52.7 1.01.3 mm; claw 2.53.5 mm long. Staminal tube straightly cut, 525
526 A. rapahelis
A. triradiatus
A. tribuloides
A. transjordanicus
A. tadmorensis
A. sianicus
A. sesamoides
A. sesameus
A. scorpioides
A. schimperi
A. saharae
A. pseudosinaicus
A. persepolitanus
A. longicaulis
A. kerkukiensis
A. gryphus
A. geniculatus
A. asterias
A. filicaulis
A. cornubovis
A. coronilla
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31
A. stella
Table 5. Data matrix for the 31 selected characters used for the phylogenetic analysis of the investigated taxa of Astragalus sect. Sesamei.
2 0 0 1 1 0 1 1 1 2 1 1 1 1 2 1 0 1 1 0 2 1 0 1 1 1 1 1 1 2 0
1 0 1 1 1 0 1 1 1 2 0 2 0 1 2 2 2 1 0 1 2 0 0 1 0 1 1 1 0 1 0
2 1 0 1 1 0 1 1 1 1 1 1 0 1 2 1 0 1 1 0 2 0 1 1 0 1 1 1 0 2 1
2 0 1 1 1 0 1 1 1 2 0 1 0 0 2 1 0 0 1 0 0 0 0 0 1 0 1 1 1 1 0
1 0 1 0 1 0 0 1 1 1 1 1 0 1 2 1 1 1 1 0 2 1 1 1 0 1 0 1 0 1 2
1 0 0 1 0 0 0 1 1 1 1 0 0 1 1 0 0 1 1 0 2 0 0 1 1 0 1 1 1 2 0
2 1 0 1 1 0 1 1 1 2 1 1 1 1 2 1 0 1 1 0 1 0 0 1 0 0 1 1 1 2 0
1 0 1 1 1 1 2 1 1 2 0 1 0 1 2 2 2 1 0 0 1 0 1 1 0 1 0 1 0 2 0
2 0 0 1 1 1 1 1 0 2 0 1 0 1 2 1 0 1 1 1 0 1 0 1 0 1 1 1 0 2 0
2 0 1 1 1 0 2 0 0 0 0 2 0 1 2 2 2 1 1 0 0 1 0 2 0 1 1 2 0 2 2
2 0 0 1 1 0 1 1 1 1 1 1 1 1 2 1 0 1 1 0 0 1 0 1 1 1 1 2 0 0 0
2 1 0 1 1 0 1 1 1 2 1 1 0 1 2 1 0 1 1 0 0 0 0 1 0 1 1 1 0 2 0
2 1 1 0 1 0 1 1 0 1 0 1 1 0 2 1 0 0 1 0 0 0 0 1 0 1 1 1 0 2 0
0 0 1 1 0 0 1 1 1 2 1 1 0 1 0 0 0 1 1 2 1 0 0 0 1 1 0 0 1 2 0
1 0 0 1 1 0 2 1 1 2 1 1 0 1 2 1 0 1 1 0 1 0 0 1 1 1 1 1 1 2 0
1 0 1 0 1 0 2 0 0 1 0 1 0 0 0 1 2 0 1 0 1 0 1 0 0 0 0 0 0 2 1
2 0 0 0 1 0 1 1 0 1 1 2 0 1 2 1 1 1 1 0 0 0 0 1 0 1 1 1 0 2 2
2 0 0 1 1 0 1 1 1 1 1 1 0 1 2 1 0 1 1 0 0 1 0 1 0 1 1 1 0 2 1
1 0 0 1 1 0 2 1 1 2 1 1 0 1 1 1 0 1 1 0 0 0 0 1 1 0 1 1 1 1 1
2 0 0 1 1 0 1 1 1 2 1 1 0 1 1 1 1 1 1 0 0 0 0 0 1 0 1 0 1 2 0
1 0 0 0 0 0 1 0 0 2 0 1 0 0 0 0 0 0 1 0 1 0 1 1 0 0 0 1 0 1 1
0 0 1 0 1 0 2 1 1 1 0 0 1 0 0 0 0 0 1 0 0 0 0 1 0 0 0 1 0 2 2
Figure 1. Astragalus raphaelis. (A) flowers with bract, (B) open calyx, (C) bracts, (D) standard, (E) wings, (F) keel, (G) stamens, (H) pistil, (I) stigma, (J) pods, lateral view, (K) pods, ventral view, (L), pods, dorsal view, (M) fructiferous inflorescence, (N) seeds, (O) leaflets.
3.23.6 mm long, with filaments 0.71.0 mm long and anthers 0.250.30 mm long, apiculate at apex. Ovary sessile or sub-sessile, 2.73.2 mm long with densely subpatent hairs, 0.51.5 mm long; style glabrous, 1.01.3 mm long; stigma transversally flat. Legumes star-like spreading, linear-lanceolate, greyish, 1015 2.53.0 mm, bigibbous at base, ending in an erect or nearly hooked beak, 0.71.2 (1.5) mm long, deeply and narrowly grooved dorsally, keeled ventrally, covered with two kinds of hairs: the shorter ones appressed, 0.050.20 mm long, the longer ones sub-appressed or subpatent, 0.51.9 mm long, sitting on a small tubercle. Seeds
not compressed, 47 in each locule, quadrangular, 1.01.5 1.21.5 mm, brown, not dotted, rugose. Iconography
Jafri (1980, Fig. 17DN), Bonnier (1914, Pl. 144, Fig. 781), Dominguez (1987, Fig. p. 185), Chamberlain and Matthews (1970, Fig. 1), Coste (1937, Fig. 941, p. 364). Distribution and ecology
According to Gazer (1993), A. stella is widely distributed in the Mediterranean region. It chiefly occurs in northern 527
Figure 2. Inflorescence of Astragalus raphaelis.
Africa (Morocco to Egypt), Canary Islands, Iberian Peninsula and southern France. The species has its optimum in very xeric habitats, also in desert areas, where it is a member of therophytic communities belonging to the StipoTrachynietea distachyae, Poetea bulbosae and Schismo-Crythopsietea (Brullo and Furnari 1978, Brullo 1985). It does not seem to be connected to any particular soil type. Additional specimens examined
Figure 3. Fructified inflorescence of Astragalus raphaelis.
Libya, Cyrenaica: El Carruba, 1 Apr 1975, Brullo and Furnari (CAT); Algeria, Hauts Plateaux Hukallis (Oran), 26 Apr 1896, Chabert (FI); France, Pass de Lanc¸iers, B. du Rh., terrains en friche pres de la gare, 3 May 1876, Autheman (FI); Dep. Vancluse, collines dominat la Chartreuse, 22 May 1993, Girerd 16143 (FI).
Figure 4. Geographical distribution of Astragalus raphaelis. Locus classicus (star) and other localities (dot).
528
Figure 5. Astragalus stella. (A) flowers with bract, (B) open calyx, (C) bracts, (D) standard, (E) wings, (F) keel, (G) stamens, (H) pistil, (I) stigma, (J) pods, lateral view, (K) pods, ventral view, (L) pods, dorsal view, (M) fructiferous inflorescence, (N) seeds, (O) leaflets.
Pod and seed micro-morphology The micro-morphological characters of the outer coat and fruit hairs, as well as those of the seed coat surface, have remarkable diagnostic value and phylogenetic significance (Chuang and Heckard 1972, Barthlott 1981, Behnke and Barthlott 1983, Zarre 2003, Karamian and Ranjbar 2005, Vural et al. 2008). In fact, the surface structure of mature seeds is characteristic and constant for a given species or group of allied species. When comparing A. raphaelis and
A. stella, it is quite clear that both their pods and seeds are micro-morphologically well-differentiated (Fig. 6). The outer pod coat of A. raphaelis is characterized by a slightly rugose surface, with several sub-appressed, minutely and sparsely papillose short hairs, with a deep groove ending in a basal fovea, mixed to scattered simple hairs, obliquely inserted on a prominent tubercle surrounded by a rugose semi-lunar annulus (Fig. 6, A3). In contrast, A. stella has pods with a markedly reticulate-rugose surface, with several appressed simple short hairs, densely papillose, mixed to
529
Figure 6. SEM microphotographs of (A) Astragalus raphaelis and (B) A. stella. (1) seed, (2) testa, (3) pod surface.
scattered simple long hairs, obliquely inserted on a inconspicuous tubercle not surrounded by an annulus (Fig. 6, B3). In addition, both species are clearly different for the shape, size and surface of seeds. Astragalus raphaelis has kidneyshaped, flattened, pale-brown seeds with dark dots, slightly rugose, 1.52.0 mm long and 1.62.0 mm wide (Fig. 6, A1), while A. stella has quadrangular seeds, not compressed, brown without dots, markedly rugose, 1.01.5 mm long, and 1.21.5 mm wide (Fig. 6, B1). Relevant differences are also observed in the cellular arrangement of the testa. The seed epidermal surface of A. raphaelis is formed by isodiametric cells, 5.56.5 mm wide, slightly rugose-colliculate, bordered by minute, slightly branched rays (Fig. 6, A2), while A. stella has elongated stellate cells, 4.55.5 mm wide with long prominent irregular rays (Fig. 6, B2).
Discussion and conclusions Based on morphological surveys carried out on literature, living plants and herbarium specimens, A. raphaelis is welldifferentiated from the other known species of A. sect. Sesamei, chiefly by the occurrence of small flowers with corolla slightly exserted from calyx, pods blackish at maturity, and seeds flattened, kidney-shaped and dotted. Most of the species belonging to this section are characterized
530
by longer corolla, pods greyish to pale brown, and seeds rectangular, not compressed. The only other species having a corolla slightly exserted from the calyx is A. sesamoides Boiss., while a few species show kidney-shaped seeds, such as A. cornubovis Lipsky, A. asterias Hohen., A. tribuloides Delile and A. transjordanicus Sam. With regards to the similarities between A. raphaelis and A. stella, it is clear that the two species are morphologically well-differentiated, as long pedunculate inflorescences is the only character they share. In fact, A. raphaelis shows an erect stem with ascending basal branches, never exceeding 20 cm, leaves with membranaceous , lanceolate, 23 mm long stipules, an inflorescence with bracts 3.04.5 mm long, acute, a calyx with teeth 3.0 4.5 mm long, longer than the tube, a pale blue corolla with smaller petals (standard 5.56.5 mm, wings 5.56.0 mm, and keel 4.55.0 mm long), shortly exserted from the calyx, an ovary covered by appressed hairs, 0.20.3 mm long, conic stigma, erect pods, blackish, rounded at the base, 34 mm wide, and pale-brown, minutely dotted, kidney-shaped, seeds, 1.52.0 1.62.0 mm. Whereas, A. stella is characterized by prostrate to ascending stems, up to 35(50) cm tall, leaves with green, triangular-ovate, 36(7) mm long stipules, an inflorescence with bracts 1.53.0 mm long, acuminate, a calyx with teeth 1.23.0 mm long, shorter than the tube, a pinklilac to violet corolla with larger petals (standard 710 mm, wings 68 mm, and keel 56 mm long), long exserted from the calyx, an ovary covered by sub-patent
Figure 7. UPGMA phenogram of the species belonging to Astragalus sect. Sesamei based on morphological data (Table 2 and 3).
hairs, 0.51.5 mm long, transversally flattened stigma, pods that are star-like spreading, greyish, bigibbous at the base, 2.53.0 mm wide, and brown seeds without dots, not flattened, rectangular, 1.01.5 1.21.5 mm. Thus, based
on these relevant morphological differences, together with the micro-morphological ones, A. raphaelis is clearly differentiated from A. stella. The outcomes of the morphological analysis are also strongly supported by the phenetic tree
Figure 8. Strict consensus tree (maximum parsimony) of the species belonging to Astragalus sect. Sesamei based on morphological data (Table 2 and 3). Bootstrap values ( 50%) are indicated under each branch.
531
(Fig. 7). In fact, these species fall into different clades: clade A (A. stella) and clade B (A. raphaelis), while a closer morphological relationship is indicated between A. raphaelis and A. sesamoides, the latter confined to the IranoTuranian region (Gazer 1993, Podlech 1999b). We obtained the same results when performing a phylogenetic parsimony analysis (Fig. 8). A total of 139 maximum parsimonious trees of 201 steps in length with a consistency index (CI) of 0.367 and a retention index (RI) of 0.454 were obtained. All characters were parsimony informative. Within A. sect. Sesamei, the strict consensus tree also supports a clear separation between A. stella and A. rapahelis (the latter being closer related to A. sesamoides). In fact, both A. raphaelis and A. sesamoides have inflorescences long pedunculate, with several crowded flowers, calyx teeth that are longer than the tube, corolla slightly exserted from the calyx, erect legumes, rounded at base and ending into a reflexed beak, but some important differences can be observed between these species. In particular, A. sesamoides differs from A. raphaelis by having ascending stems up to 36 cm tall, stipules triangular, 34 mm long, leaflets 46-paired, 23 mm long bracts in the inflorescence, legumes 1215 mm long, and seeds yellow, not dotted, and smooth. Therefore, the hypothesis that A. raphaelis is closely related to A. stella, or merely a synonym of the latter, can definitely be discarded, while a possible relationship with A. sesamoides should be taken into consideration. On the basis of that, it can be emphasized that A. raphaelis is morphologically well-differentiated from the other species of sect. Sesamei, and show phylogenetic correlations with A. sesamoides only, from which it is geographically isolated (Gazer 1993). Genomic investigations on the species belonging to this section would better clarify the phylogeny and the taxonomical position of A. raphaelis and its allied species. Acknowledgements We are very grateful to the Prof. Dr D. Podlech (Mu¨nchen) for his useful suggestions and critical review of the manuscript. Furthermore, we are indebted to the curators of B, CAT, FI, G, M and PA Herbaria for the facilities during the examination of the collections.
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