Family Leguminose Juss. (Fabaceae)
Сем. LXII. БОБОВИ — LEGUMINOSAE JUSS.¹
Family Leguminose Juss. (Fabaceae)
Сем. LXII. БОБОВИ — LEGUMINOSAE JUSS.¹
Fam: Leguminose Juss. (Fabaceae)
English Name: Legume, Pea, or Bean familyDescription: Trees, shrubs, vines or herbaceous plants, often climbing or creeping. Root is usually with small tubers, containing Nitrogen-Fixing bacteria. Leaves consistent, rarely opposite, simple or double pinnate, rarely simple or palmate or stunted leaves, modified or completely absent; textured branches in kladodii; usually with stipule, last free or fused with petiole. Flowers in cluster, rarely in sunshade-similar, thyroid, or wheatear-like of head inflorescences or flowers single, correct or zigomorf, polygamous or hermaphroditic, rarely modified or completely missing. The calyx is usually from 5 rarely than 4, more or less fused leaflets, often bilabial, rarely sepals are free. Corolla many or 5 more or less identical leaflets or 5 unequal, differentiated in the flag, keel and wings, free or partially fused petals, corolla rarely reduced to a piece of leaflet. Stamens many - 5 (4), usually twice the petals, free or all fused into a tube, or 9 fused into a tube and a free. The carpophor of one leaflet of fruit with upper, single station go across sometimes, very rarely longitudinal bulkhead ovary, with one to many ovules attached to the abdominal seam. Fruit dry, break out into blossom two or more single seam, less less unhatched is, or beans disintegrating into smaller members. The seed usually without endosperm or the last minor, with or without appendage. Pollination by insects generally, the nectar was collected at the base of stamens, less often. kleystogam. Fruits usually endo- or ekzozoohor, rarely entire saplodie anemohor, seeds or distracted at bursting or the last positive gravitropism, burying in the ground where it ripens (geokarpiya). Reproduction mainly by seed.
Note. Within the family observed characteristic evolution in terms of adaptive mechanisms to disperse the fetus. In primitive generas beans with many seeds, simply arranged without devices zoohoriya and usually beans with many seeds, and splits still on the plant (Chamaecytisus, Corothamnus, Genista, Spartium, Robinla, Cercis, etc.), Releasing the seeds appearing in the case of unit variance. At senior (Medicago, Onobrychis, etc.) the bean has a variety of devices ekzozoohoriya and is usually with few seeds and no cracks - appears to be one of dissipation. Peak of this line with our representatives we see in the genus Trifolium. Beans is irrelevance to monocotyledon, with thin pericarp and usually covered by the calyx, but in some species and continued dry corolla. In the anemohoriya or ekzozoohoriya unit of variance is the whole flower or calyx, including beans as the main role in the play ekzozoohoriya teeth. In some cases, a unit of variance is the whole inflorescence (anemohorno) or extreme degree of specialization - avtohorno by geokarpiya (T. subterraneum etc.). In the latter case, some flowers are sterile and the teeth of the calyx play a major role in the only burial in soil (see. Van der Riil, 1969).
In this Family have been identified in the fossil state these taxa: Cassia ambigua Neeg (Chukurovo Miocene), C. hyperborea Ung. (Chukurovo Miocene), C. lignitum Ung. (Bobov Dol, Oligocene); Cassiophyllum berenices (Ung.) Krausel (Borovets, Oligocene); Podogonium knorri Neeg (Chukurovo Miocene D. Dobrevo Oligocene; Tsastrogor, Oligocene; Tselovo and Stavertsi, Miocene, Bobov Dol, Oligocene); Dalbergia sp. (Pelova, Miocene); Ceratonia emarginata A. Br (Brezhani, Oligocene); Gymnocladus cf. meoreatarica Kol. (Garmen Pliocene); Cesalpinia haidingeri Ett. (Brezhani, Oligocene); Sophora europaea Ung. (Brezhani, Oligocene).
Table setting of wild and cultivated generations
1 At least some of the double pinnate leaves ............................................................................................................................................................ 2
1* Leaves simply pinnate, triple, digitate, simple or leaves opadlivi, undeveloped or modified ………………………………………........................... 5
2 Trees ................................................................................................................................................................................................................... 3
2* Shrub 1.0 - 2.0 m tall ....................................................................................................................................................................... Caesalpinia L.
3 Flowers white or greenish white ................................................................................................................................................ Gymnocladus Lam.
3* Flowers yellow-green, yellow, pink or rose red ..................................................................................................................................................... 4
4 Stamens significantly exceed corolla .................................................................................................................................................. Albizia Durazz.
4* Stamens not exceed corolla ................................................................................................................................................................... Gleditsia L.
5 Corolla from a leaflet ........................................................................................................................................................................... Amorpha L.
5* Corolla from a 5 leaflets ......................................................................................................................... ............................................................... 6
6 Lleaves modified into tendril, with large leaf-shaped stipule .............................................................................................................37. - Lathyrus L.
6* Leaves developed with leaflets ............................................................................................................................................................................... 7
7 Leaflets pinnate (cloven-cirrus with 2, 4 or more leaflets, imparipinnate with 5, 7 or more leaflets) or digitate (with 5 - 7 (9 - 17) leaflets, attached at one point) ................................................................................................................................................................................................................... 8
7* Leaves simple or triple sometimes underdeveloped, reduced or modified, drop off ............................................................................................... 57
8 Leaves cloven-cirrus ending with tendril, awn or spinule .......................................................................................................................................... 9
8* Leaves imparipinnate, ending with a leaflet or digitate ............................................................................................................................................ 18
9 Trees or shrubs, rarely subshrubs ......................................................................................................................................................................... 10
9* Herbaceous plants ............................................................................................................................................................................................... 13
10 Flowers bright yellow or yellowish ....................................................................................................................................................................... 11
10* The flowers white, violet, purple or pink .............................................................................................................................................................. 12
11 Corolla 15.0 - 25.0 mm long, bright yellow. Flower stems articulated ............................................................................................ 12. Caragana L.
11* Corolla about 10.0 mm long, yellowish. Flower stems rigid .......................................................................................................... 13. Astragalus L.
12 Stipule converted into 2.0 - 6.0 cm long spines. Flowers 2 - 3 in bundles .................................................................................. Halimodendron DC.
12* Stipule not converted into spines. The flowers in clusters cluster …............................................................................................... 13. Astragalus L.
13 Peduncle after overblown extending, with a positive gravitropism, burying the fetus in the soil. Stipule fused with leaf axis. The upper lip of the calyx with 4, the lower with a 1-tooth ..................................................................................................................................................................... Arachis L.
13* Peduncle not burying the fetus. Stipule not accrete with leaf axis. Calyx regular or bilabial, but then the upper lip with 2 bottom 3 teeth .................. 14
14 Stems winged ................................................................................................................................................................................ 37. Lathyrus L.
14* Stems not winged ..................................................................................................................................................................................... ......... 15
15 The teeth of the calyx 2 - 4 (6) times longer than the tube, equal to or longer than corolla ....................................................................... 36. Lens L.
15* The teeth of the calyx shorter or equal, more rarely longer than the tube, shorter than the corolla .......................................................................... 16
16 Stipule (5,0-) 6.0 - 10.0 cm long, almost equal to the leaflets. The teeth of the calyx leaf-shaped teeth ............................................... 38. - Pisum L.
16* Stipule to 3.0 cm smaller than the leaflets. The teeth of the calyx are not leaf-shaped ............................................................................................ 17
17 The baluster fiber along its entire circumference, only the underside of naked stigma to the top extended Staminal tube finally severed right ................................................................................................................................................................................................................ 35. - Vicia L.
17* The baluster fiber only from the top, the stigma of top threadlike, staminal tube to end obliquely cut ............................................... 37. - Lathyrus L.
18 (8*) Climbing ligneous lianas to 20.0 meters long, Pale violet flowers ............................................................................................... - Wisteria Nutt.
18* Non climbing trees or shrubs. The flowers pink, yellow, white or dove to lilavobeli …...................................................................................….. 19
19 Trees ................................................................................................................................................................................................................. 20
19* Shrubs or herbaceous plants (at least in our conditions) ....................................................................................................................................... 23
20 Corolla pink or yellow. The flowers in loose panicle .............................................................................................................................. - Sophora L.
20* Corolla white, pink to purple. The flowers in drooping cluster inflorescences ………............................................................................................ 21
21 Flowers white .............................................................................................................................................................................. ..................... 22
21* The flowers pink to purple .................................................................................................................................................................. - Robiriia L.
22 Corolla 2.5 - 3.0 cm long. The bark of the trunk smooth ............................................................................................................... Cladrastis Rafin.
22* Corolla 1.5 - 2.0 cm long. The bark of the trunk cracked ..................................................................................................................... - Robinia L.
23 (19*) Leaves digitate or seemingly digitate. Leaflets attached more or less at one point .......................................................................................... 24
23* Leaves imparipinnate. Leaflets attached along the leaf axis …………................................................................................…………………....... 25
24 Leaflets 5 lower two leaflets attached to the stem, resembling stipule. Clusters of head. Keel dark red to black .......................... 20. - Dorychium L.
24* Leaflets 7 - 9 (11 - 17), lower not resemble stipule. Clusters cluster. The keel is not dark red to black .............................................. 3. - Lupinus L.
25 Shrubs or subshrubs ........................................................................................................................................................................................... 26
25* Perennial to annual herbaceous plants ……………………................................................................................………………………………. 30
26 A flag yellow ...................................................................................................................................................................................................... 27
26* A flag white, pink or purple ................................................................................................................................................................................ 28
27 Flowers in cluster inflorescences. Beans heavily swollen bubble, burst to the top ……………......................................................... 11. - Colutea L.
27* Flowers in sunshade-like, almost of head inflorescences. Beans linear composed of separate small members, which decays ........... 25. - Coronilla L.
28 Flowers in sunshade-similat, almost of head inflorescences. Beans composed of separate small members, which decays ................ 25. - Coronilla L.
28* Flowers single, two or many clusters in the cluster . The bean splits along ............................................................................................................ 29
29 All 10 stamens fused into a tube. Branches more or less fibrous glandular ........................................................................................... 29. - Ononis L.
29* A stamen separated from the rest 9 which are fused with each other. Branches without glandular hairs ........................................ 13. - Astragalus L.
30 (25 *) The main veins of the leaflets reach the edge, often ending in tooth ............................................................................................................... 31
30* The main veins of the leaflets anastomosed not reach the edge ............................................................................................................................. 32
31 All leaves feathery. Leaflets 7 - 17 number .......................................................................................................................................... 34 - Cicer L.
31* Only the lower leaves feathery and leaflets fewer than 7 ................................................................................................................... 29. - Ononis L.
32 Leaflets 5, lower two attached to the stem, resembling stipule. Keel dark red or black ........................................................... 20. - Dorycnium Mill.
32* Leaves feathery, the lower two petals not resemble prilistnitsi. The keel not dark red or black .............................................................................. 33
33 Inflorescence with stem, at the base with one or more in inflorescences leaves ...................................................................................................... 34
33* Inflorescence sessile or with a stem, but in this case basis without inflorescence leaves …..................................................................................… 35
34 Clusters of two or more in inflorescences leaves. Beans around the edge, shorter than the calyx ...................................................... 17. - Anthyllis L.
34* Clusters in the bosom of the leaves. Beans serrated edge, much longer than the calyx...................................................... 18. - Hymenocarpus Savi
35 Plants at least partly glandular hairs. Beans reddish brown, shiny ............................................................................................. 16. - Glycyrrhiza L.
35* Plants without glandular hairs. The bean is not reddish brown and shiny ............................................................................................................... 36
36 Flowers in sunshade or bundles. Flower stems attached at one point ................................................................................................................... 37
36* Flowers in cluster inflorescences or thyroid inflorescences or single ...................................................................................................................... 43
37 The keel on the front end curved and tapering in the form of beak ........................................................................................................................ 39
37* The keel on the front end blunt or pointed, but it is not curved as beak …………...................................................................................……….. 38
38 Flowers at 1 - 3 in the inflorescence, shorter than 5.0 mm. Beans disintegrated into separate small member. Annuals plants ........ 24. - Ornithopus L.
38* More than 3 flowers in the inflorescence, longer than 5.0 mm. Beans splits are, non-fissile of small members. Perennials .............. 13. - Astragalus L.
39 Flowers pink or white pink. Beans disintegrated into separate small members .................... ........................................................... 25. - Coronilla L.
39* Flowers yellow or white. Beans no cracks, does not disintegrate into separate small members ....................................………………………...... 40
40 Keel dark red or black ......................................................................................................................................................... 20. - Dorycnium Mill.
40* Keel is not dark red or black .............................................................................................................................................................................. 41
41 The leaflets 5, lower two approximated to the base of the leaf axis resemble stipule ............................................................................. 21. - Lotus L.
41* The leaflets more than 5, lower two not resemble stipule ..................................................................................................................................... 42
42 Beans articulated divided, on top without beak, splits are; the small members like a horseshoe, or with rounded sinus ............ 26. - Hippocrepis L.
42* Beans oblong, on top with 1.5 - 3.0 cm long by beak, it is not a member of, the seams bold, no cracks or later burst is ............. 19. - Securigera L.
43 (36 *) Leaflets with clear parallel veined ........................................................................................................................................ 37. - Lathyrus L.
43* Leaflets with feather-like veined or lateral veins unclear ..................................................................................................................................... 44
44 The keel there on the front end curved and tapering in the form of a beak or with 0.5 - 1.5 mm long tooth ...................................................... ..... 45
44* The keel on its front end blunt or pointed, not curved in the form of a beak and no tooth ...................................................................................... 47
45 The leaflets 5; lower two approximated to the base of the leaf axis, resembling stipule .......................................................................... 21 - Lotus L.
45* The leaflets more than 5; lower two not resemble stipule ..................................................................................................................................... 46
46 Inflorescences are sunshade similar heads or single flowers. Beans articulated inflorescences racemose …................................ 26. - Hippocrepis L.
46* Inflorescences racemose. The bean is not articulated inflorescences racemose ……………....................................……………..14. - Oxytropis L.
47 Leaves cloven-cirrus .......................................................................................................................................................................................... 48
47* Leaves imparipinnate .......................................................................................................................................................................................... 49
48 Spiny plants ............................................................................................................................................................................ 13. - Astragalus L.
48* Not prickly plants ............................................................................................................................................................................... 35 - Vicia L.
49 The keel at the top tapering ................................................................................................................................................................................ 50
49* The keel to the top blunt ..................................................................................................................................................................................... 52
50 All 10 stamens fused into a tube ...................................................................................................................................................... 10. - Galega L.
50* A stamen separated from the rest 9 which are fused with each other in the tube …....................................................................................……… 51
51 Corolla (12.0) 14.0 - 40.0 mm in length. Beans more or less flattened, always smooth, with clear small members, when ripe disintegrating on 8 segments …………….......................................................................................................................................................……..... 27. - Hedysarum L.
51* Corolla to 12.0 - 13.0) mm long. Beans elliptical or ovoid, no cracks, often with spines, prickles or warts ................................ 28. - Onobrychis L.
52 All 10 stamens fused into a tube ...................................................................................................................................................... 10. - Galega L.
52* One stamen separated from the remaining 9, which are fused into a tube .............................................................................................................. 53
53 The leaflets primarily in the upper half fine cartilage serrsted ................................................................................................................. 35 - Vicia L.
53* The leaflets entire .............................................................................................................................................................................................. 54
54 The bean with hard and prickly spines or highly prominent network of veins, no cracks ….................................................................................. 55
54* The beans without spikes or highly prominent network of veins, splits .................................................................................................................. 56
55 Flowers blue or yellow with a blue peak. Long beans, cut jagged edge .......................................................................................... 15. - Bisserula L.
55* The flowers white, pink or purple. The bean more or less roundish, usually around the edge serrated, convex on the sides with a network of veins or wells and veins often jagged ..................... ............................................................................................................................ 28 - Onobrychis L.
56 The flowers without bracts; fiber underneath the bar or on the entire round …....................................................................................... 35 - Vicia L.
56* The flowers with bracts; the bar naked ...................................................................................................................................... 13. - Astragalus L.
57 (7*) Leaves simple, sometimes reduced, modified, drop off or missing, replaced by photosynthetic twigs or greatly increased stipule and leaf petiole resembling simple leaves .................. ......................................................................................................................................................................... 58
57* Leaves triple, always well developed ................................................................................................................................................................... 67
58 Wood with large semicircular at the base deeply heart-shaped leaves .................................................................................................. 1. - Cercis L.
58* Annual or perennial herbaceous plants ................................................................................................................................................................. 59
59 The flowers yellow, orange or yellowish .............................................................................................................................................................. 60
59* The flowers pink, red or purple to whitish ............................................................................................................................................................ 61
60 The leaves linear lance. Beans longer than the calyx ....................................................................................................................... 37 - Lathyrus L.
60* Leaves narrowly to broadly elliptical or ovoid. Beans shorter than the calyx ................ .................................................................. 17. - Anthyllis L.
61 Annual herbaceous plants. Beans spirally curved on external ribs with spines ………………… .................................................. 23 - Scorpiurus L.
61* Perennial herbaceous plants or shrubs. Beans untwisted, spineless ....................................................................................................................... 62
62 Young stems with two wide to 2.0 (3.0 - 5.0) mm wings, indented in knots ............................................................... 5. - Chamaespartium Adans.
62* Young stems not winged or wings narrow to 1.0 mm wide, indented in the nodes ................................................................................................. 63
63 Perennial herbaceous plant. Calyx after overblown more or less bubble gummy. Beans shorter than the calyx ............................... 17. - Anthyllis L.
63* Shrubs or subshrubs. Calyx after overblown not bubble gummy. Beans longer than the calyx ................................................................................ 64
64 Calyx with a lip, on the top with 5 very short teeth. Corolla 20.0 - 30.0 mm long ………… ............................................................... - Spartium L.
64* Calyx bilabial, upper lip with 2, bottom with 3 teeth. Corolla (8.0) 10.0 - 15.0 (16.0 - 20.0) mm long ............... ................................................. 65
65 The upper lip of the calyx shallow, up to 1/3 two-pronged, lower 1/5 - 1/3 tridentate ......................................................................................... 66
65* The upper lip of the calyx up to the base bilabial, lower to middle or base tridentate ....................................................... .................. 4. - Genista L.
66 Small shrubs, 10.0 - 20.0 (40.0) cm high. Beans more or less evenly thick fibrous ............................................. 6. - Corothamnus (Koch) C. Presl
66* Shrub, 30.0 - 150.0 (250.0) cm high. Beans on the sides goal or almost goal, the edges with long hairs ................................. Sarothamnus Wimm.
67 (57*) The main side veined of leaflets finishing the edge; the latter often serrated ............................................................. ...................................... 68
67* The main side veined of leaflets anastomosed and do not reach the edge, sometimes unclear; edge entire .............................................................. 79
68 Plants fibrous glandular at least in the upper part ...................................................................................................................................... ........... 69
68* Plants without glandular hairs .............................................................................................................................................................................. 70
69 All 10 stamens fused into a tube. The keel with beak ....................................................................................................................... 29. - Ononis L.
69* One of the stamens clear separated from the rest 9, which are fused with each other. The keels without beak ............................... 31. - Medicago L.
70 The corolla at beans durable; at least one of the petals accreted to staminal tube ............................................................................. 33. - Trifolium L.
70* Corolla after after overblown fallen; the petals are not fused with Staminal tube ……......................................................................................….. 71
71 Beans shorter or equal to calyx. The stems of the stamens at least of 5 stamens atop extended ...................................................... 33. - Trifolium L.
71* Beans usually longer than the calyx. All stems of the stamens threadlike ……….....................................................................................………... 72
72 Beans spirally curved in one or more rounds ............................................................................................................................... 31. - Medicago L.
72* Beans crescent to the right, straight, ovoid or reniform ......................................................................................................................................... 73
73 Flowers white ..................................................................................................................................................................................................... 74
73* The flowers yellow or blue .................................................................................................................................................................................. 75
74 The flowers in racemose, loose, elongated inflorescences ......................................................................................................... 32. - Melilotus Mill.
74* The flowers single, two or compact cumulus or ovoid clusters .................................................................................................... 30. - Trigonella L.
75 Perennial plants. Long beans, crescent or …………..............................................................................……………................. 31. - Medicago L.
75* Annual plant- or biennial plant ............................................................................................................................................................................. 76
76 The flowers blue ........................................................................................................................................................................ 30. - Trigonella L.
76* The flowers yellow .................................................................................................................. ........................................................................... 77
77 The flowers in the racemose, elongated, loose, cluster .............................................................................................................. 32. - Melilotus Mill.
77* The flowers in cylindrical, ovoid back, short, cluster clusters or 1 - 2 in the axils of the leaves ................................................................................ 78
78 Beans linear to ovate with a long beak ........................................................................................................................................ 30. - Trigonella L.
78* Beans reniform without beak ...................................................................................................................................................... 31. - Medicago L.
79 (67*) Leaflets at the base with stipule .................................................................................................................................................................... 80
79* Leaflets at the base without stipule ...................................................................................................................................................................... 84
80 Shrub to 2.5 meters high. The flowers yellow ................................................................................................................................ Petteria C. Presl
80* Grassy plants. The flowers white, yellow, pink ..................................................................................................................................................... 81
81 Corolla longer than 9.0 mm. Plant bare or with white hairs ................................................................................................................................... 82
81* Corolla to 7.0 mm long. Plants with reddish hairs ………………….................................................................................………........ - Glycine L.
82 The keel there on top of a spiral twisted ........................................................................................................................................... - Phaseolus L.
82* The keel straight or curved, twisted not spiral ...................................................................................................................................................... 83
83 Stigma Mining, scar more than half the circumference of the seed ........................................................................................................ Lablab Savi
83* Stigma obliquely. Scar two times shorter than half the circumference of the seed .................................................................................. - Vigna Savi
84 Trees and shrubs ................................................................................................................................................................................................ 85
84* Herbaceous plants .............................................................................................................................................................................................. 90
85 Corolla red or pink violet ............................................................................................................................................................. Lespedeza Mich.
85* Corolla yellow ................................................................................................................................................................................................... 86
86 Clusters leafless drooping clusters ............................................................................................................................................ - Laburnum Medic.
86* Clusters like brooms, of head or cluster, but then upright ..................................................................................................................................... 87
87 The upper lip of the calyx to the base two-pronged .......................................................................................................................... 4. - Genista L.
87* The upper lip of the calyx at most middle two-pronged ........................................................................................................................................ 88
88 Calyx more or fewer pipe-like, upper lip of the calyx of 1/3 - 1/2 two-pronged ……………............................................ 8. - Chamaecytisus Link
88* Calyx bell-shaped, upper lip of the calyx completely shallow two-pronged ........................................................................................................... 89
89 Clusters outerwear, leafless. The seed without appendage .......................................................................... . 7. Lembotropis - Lembotropis Griseb.
89* Flowers single or two in the bosom of the leaves. Seed with appendage …….......................... ............................................ Sarothamnus Wimm.
90 (84*) Leaves with clear punctate glands ............................................................................................................................................ 9 - Psoralea L.
90* Leaves no clear punctate glands .......................................................................................................................................................................... 91
91 The upper lip of the calyx with 4 teeth, the bottom with a narrow tooth styliform, all longer than the tube of the calyx ............................................................................................................................................................................... 2. - Lotononis (DC.) Eckl. et Zeyh.
91* The upper lip of the calyx with 2, bottom with 3 tooth (or teeth obscure), all shorter than tube .............................................................................. 92
92 The flowers single or two in the bosom of the leaves ..................................................................................................... 22. Tetragonolobus Scop.
92* The flowers more than 3, gathered in clusters of head or sunshade-similar ........................................................................................................... 93
94 The leaves without stipule. All 10 stamens fused into a tube ......................................................................................................... 17 . - Anthyllis L.
94* The leaves well developed webbed or grassy, more or less fused with the stem of the petiole. A stamen separated from the other 9, which are fused between themselves .................................................................................................................................................................. 33. - Trifolium L.¹ characteristic of family and tables to determine the genera developed by B. Kouzmanov
From „Флора на НР България”, том VI, БАН, София, (1976)
* * *
The Fabaceae, Leguminosae or Papilionaceae,[6] commonly known as the legume, pea, or bean family, are a large and economically important family of flowering plants. It includes trees, shrubs, and perennial or annual herbaceous plants, which are easily recognized by their fruit (legume) and their compound, stipulated leaves. The family is widely distributed, and is the third-largest land plant family in terms of number of species, behind only the Orchidaceae and Asteraceae, with about 751 genera and some 19,000 known species[7][8][9] . The five largest of the genera are Astragalus (over 3,000 species), Acacia (over 1000 species), Indigofera (around 700 species), Crotalaria (around 700 species) and Mimosa (around 500 species), which constitute about a quarter of all legume species. The ca. 19,000 known legume species amount to about 7% of flowering plant species.[8][10] Fabaceae is the most common family found in tropical rainforests and in dry forests in the Americas and Africa.[11]
Recent molecular and morphological evidence supports the fact that the Fabaceae is a single monophyletic family.[12] This point of view has been supported not only by the degree of interrelation shown by different groups within the family compared with that found among the Leguminosae and their closest relations, but also by all the recent phylogenetic studies based on DNA sequences.[13][14][15] These studies confirm that the Fabaceae are a monophyletic group that is closely related to the Polygalaceae, Surianaceae and Quillajaceae families and that they belong to the order Fabales.[16]
Along with the cereals, some fruits and tropical roots a number of Leguminosae have been a staple human food for millennia and their use is closely related to human evolution.[17]
A number are important agricultural and food plants, including Glycine max (soybean), Phaseolus (beans), Pisum sativum (pea), Cicer arietinum (chickpeas), Medicago sativa (alfalfa), Arachis hypogaea (peanut), Lathyrus odoratus (sweet pea), Ceratonia siliqua (carob), and Glycyrrhiza glabra (liquorice). A number of species are also weedy pests in different parts of the world, including: Cytisus scoparius (broom), Robinia pseudoacacia (black locust), Ulex europaeus (gorse), Pueraria lobata (kudzu), and a number of Lupinus species.Etymology
The name 'Fabaceae' comes from the defunct genus Faba, now included in Vicia. The term "faba" comes from Latin, and appears to simply mean "bean". Leguminosae is an older name still considered valid,[6] and refers to the fruit of these plants, which are called legumes.
Description
Fabaceae range in habit from giant trees (like Koompassia excelsa) to small annual herbs, with the majority being herbaceous perennials. Plants have indeterminate inflorescences, which are sometimes reduced to a single flower. The flowers have a short hypanthium and a single carpel with a short gynophore, and after fertilization produce fruits that are legumes.
Growth habit
The Leguminosae have a wide variety of growth forms including trees, shrubs or herbaceous plants or even vines or lianas. The herbaceous plants can be annuals, biennials or perennials, without basal or terminal leaf aggregations. They are upright plants, epiphytes or vines. The latter support themselves by means of shoots that twist around a support or through cauline or foliar tendrils. Plants can be heliophytes, mesophytes or xerophytes.[3][8]
Leaves
The leaves are usually alternate and compound. Most often they are even- or odd-pinnately compound (e.g. Caragana and Robinia respectively), often trifoliate (e.g. Trifolium, Medicago) and rarely palmately compound (e.g. Lupinus), in the Mimosoideae and the Caesalpinioideae commonly bipinnate (e.g. Acacia, Mimosa). They always have stipules, which can be leaf-like (e.g. Pisum), thorn-like (e.g. Robinia) or be rather inconspicuous. Leaf margins are entire or, occasionally, serrate. Both the leaves and the leaflets often have wrinkled pulvini to permit nastic movements. In some species, leaflets have evolved into tendrils (e.g. Vicia).[3][8][17]
Many species have leaves with structures that attract ants that protect the plant from herbivore insects (a form of mutualism). Extrafloral nectaries are common among the Mimosoideae and the Caesalpinioideae, and are also found in some Faboideae (e.g. Vicia sativa). In some Acacia, the modified hollow stipules are inhabited by ants and are known as domatia.Roots
Main article: Root nodule
Many Fabaceae host bacteria in their roots within structures called root nodules. These bacteria, known as rhizobia, have the ability to take nitrogen gas (N2) out of the air and convert it to a form of nitrogen that is usable to the host plant ( NO3− or NH3 ). This process is called nitrogen fixation. The legume, acting as a host, and rhizobia, acting as a provider of usable nitrate, form a symbiotic relationship.Flowers
The flowers often have five generally fused sepals and five free petals. They are generally hermaphrodite, and have a short hypanthium, usually cup shaped. There are normally ten stamens and one elongated superior ovary, with a curved style. They are usually arranged in indeterminate inflorescences. Fabaceae are typically entomophilous plants (i.e. they are pollinated by insects), and the flowers are usually showy to attract pollinators.
In the Caesalpinioideae, the flowers are often zygomorphic, as in Cercis, or nearly symmetrical with five equal petals in Bauhinia. The upper petal is the innermost one, unlike in the Faboideae. Some species, like some in the genus Senna, have asymmetric flowers, with one of the lower petals larger than the opposing one, and the style bent to one side. The calyx, corolla, or stamens can be showy in this group.
In the Mimosoideae, the flowers are actinomorphic and arranged in globose inflorescences. The petals are small and the stamens, which can be more than just 10, have long, coloured filaments, which are the showiest part of the flower. All of the flowers in an inflorescence open at once.
In the Faboideae, the flowers are zygomorphic, and have a specialized structure. The upper petal, called the banner, is large and envelops the rest of the petals in bud, often reflexing when the flower blooms. The two adjacent petals, the wings, surround the two bottom petals. The two bottom petals are fused together at the apex (remaining free at the base), forming a boat-like structure called the keel. The stamens are always ten in number, and their filaments can be fused in various configurations, often in a group of nine stamens plus one separate stamen. Various genes in the CYCLOIDEA (CYC)/DICHOTOMA (DICH) family are expressed in the upper (also called dorsal or adaxial) petal; in some species, such as Cadia, these genes are expressed throughout the flower, producing a radially symmetrical flower.[18]Fruit
The ovary most typically develops into a legume. A legume is a simple dry fruit that usually dehisces (opens along a seam) on two sides. A common name for this type of fruit is a "pod", although that can also be applied to a few other fruit types. A few species have evolved samarae, loments, follicles, indehiscent legumes, achenes, drupes, and berries from the basic legume fruit.
Physiology and biochemistry
The Leguminosae are rarely cyanogenic, however, where they are, the cyanogenic compounds are derived from tyrosine, phenylalanine or leucine. They frequently contain alkaloids. Proanthocyanidins can be present either as cyanidin or delphinidine or both at the same time. Flavonoids such as kaempferol, quercitin and myricetin are often present. Ellagic acid has never been found in any of the genera or species analysed. Sugars are transported within the plants in the form of sucrose. C3 photosynthesis has been found in a wide variety of genera.[3] The family has also evolved a unique chemistry. Pterocarpans are a class of molecules (derivatives of isoflavonoids) found only in the Fabaceae.
Ecology
Distribution and habitat
The Fabaceae have an essentially worldwide distribution, being found everywhere except Antarctica and the high arctic.[9] The trees are often found in tropical regions, while the herbaceous plants and shrubs are predominant outside the tropics.[3]
Biological nitrogen fixation
Cross-section through a root nodule of Vicia observed through a microscope.
Biological nitrogen fixation (BNF, performed by the organisms called diazotrophs) is a very old process that probably originated in the Archean eon when the primitive atmosphere lacked oxygen. It is only carried out by Euryarchaeota and just 6 of the more than 50 phyla of bacteria. Some of these lineages co-evolved together with the flowering plants establishing the molecular basis of a mutually beneficial symbiotic relationship. BNF is carried out in nodules that are mainly located in the root cortex, although they are occasionally located in the stem as in Sesbania rostrata. The spermatophytes that co-evolved with actinorhizal diazotrophs (Frankia) or with rhizobia to establish their symbiotic relationship belong to 11 families contained within the Rosidae clade (as established by the gene molecular phylogeny of rbcL, a gene coding for part of the RuBisCO enzyme in the chloroplast). This grouping indicates that the predisposition for forming nodules probably only arose once in flowering plants and that it can be considered as an ancestral characteristic that has been conserved or lost in certain lineages. However, such a wide distribution of families and genera within this lineage indicates that nodulation had multiple origins. Of the 10 families within the Rosidae, 8 have nodules formed by actinomyces (Betulaceae, Casuarinaceae, Coriariaceae, Datiscaceae, Elaeagnaceae, Myricaceae, Rhamnaceae and Rosaceae), and the two remaining families, Ulmaceae and Fabaceae have nodules formed by rhizobia.[19][20]
The rhizobia and their hosts must be able to recognize each other for nodule formation to commence. Rhizobia are specific to particular host species although a rhizobia species may often infect more than one host species. This means that one plant species may be infected by more than one species of bacteria. For example, nodules in Acacia senegal can contain seven species of rhizobia belonging to three different genera. The most distinctive characteristics that allow rhizobia to be distinguished apart are the rapidity of their growth and the type of root nodule that they form with their host.[20] Root nodules can be classified as being either indeterminate, cylindrical and often branched, and determinate, spherical with prominent lenticels. Indeterminate nodules are characteristic of legumes from temperate climates, while determinate nodules are commonly found in species from tropical or subtropical climates.[20]
Nodule formation is common throughout the leguminosae, it is found in the majority of its members that only form an association with rhizobia, which in turn form an exclusive symbiosis with the leguminosae (with the exception of Parasponia, the only genus of the 18 Ulmaceae genera that is capable of forming nodules). Nodule formation is present in all the leguminosae sub-families, although it is less common in the Caesalpinioideae. All types of nodule formation are present in the sub-family Papilionoideae: indeterminate (with the meristem retained), determinate (without meristem) and the type included in Aeschynomene. The latter two are thought to be the most modern and specialised type of nodule as they are only present in some lines of the Papilionoideae sub-family. Even though nodule formation is common in the two monophyletic subfamilies Papilionoideae and Mimosoideae they also contain species that do not form nodules. The presence or absence of nodule-forming species within the three sub-families indicates that nodule formation has arisen several times during the evolution of the leguminosae and that this ability has been lost in some lineages. For example, within the genus Acacia, a member of the Mimosoideae, A. pentagona does not form nodules, while other species of the same genus readily form nodules, as is the case for Acacia senegal, which forms both rapidly and slow growing rhizobial nodules.Evolution, phylogeny and taxonomy
Evolution
The order Fabales contains around 7.3% of eudicot species and the greatest part of this diversity is contained in just one of the four families that order contains: Fabaceae. This clade also includes the Polygalaceae, Surianaceae and Quillajaceae families and its origins date back 94 to 89 million years, although it started its diversification some 79 to 74 million years ago.[9] In fact, the Fabaceae have diversified during the early tertiary to become a ubiquitous part of the modern earth’s biota, along with many other families belonging to the flowering plants.[12][21]
The Fabaceae have an abundant and diverse fossil record, especially for the Tertiary period. Fossils of flowers, fruit, leaves, wood and pollen from this period have been found in numerous locations.[22][23][24][25][26] The earliest fossils that can be definitively assigned to the Fabaceae appeared in the late Palaeocene (approximately 56 million years ago).[27][28] Representatives of the 3 sub-families traditionally recognised as being members of the Fabaceae – Cesalpinioideae, Papilionoideae and Mimosoideae — as well as members of the large clades within these sub-families – such as the genistoides – have been found in periods a little later, starting between 55 and 50 million years ago.[21] In fact, a wide variety of taxa representing the main lineages in the Fabaceae have been found in the fossil record dating from the middle to the late Eocene, suggesting that the majority of the modern Fabaceae groups were already present and that a broad diversification occurred during this period.[21] Therefore, the Fabaceae started their diversification approximately 60 million years ago and the most important clades separated some 50 million years ago.[29] The age of the main Cesalpinioideae clades have been estimated as between 56 and 34 million years and the basal group of the Mimosoideae as 44 ± 2.6 million years.[30][31] The division between Mimosoideae and Faboideae is dated as occurring between 59 and 34 million years ago and the basal group of the Faboideae as 58.6 ± 0.2 million years ago.[32] It has been possible to date the divergence of some of the groups within the Faboideae, even though diversification within each genus was relatively recent. For instance, Astragalus separated from the Oxytropis some 16 to 12 million years ago. In addition, the separation of the aneuploid species of Neoastragalus started 4 million years ago. Inga, another genus of the Papilionoideae with approximately 350 species, seems to have diverged in the last 2 million years.[33][34][35][36]
It has been suggested, based on fossil and phylogenetic evidence, that legumes originally evolved in arid and/or semi-arid regions along the Tethys seaway during the Palaeogene Period.[5][37] However, others contend that Africa (or even the Americas) cannot yet be ruled out as the origin of the family.[38][39]
The current hypothesis about the evolution of the genes needed for nodulation is that they were recruited from other pathways after a polyploidy event.[40] Several different pathways have been implicated as donating duplicated genes to the pathways need for nodulation. The main donors to the pathway were the genes associated with the arbuscular mycorrhiza symbiosis genes, the pollen tube formation genes and the haemoglobin genes. One of the main genes shown to be shared between the arbuscular mycorrhiza pathway and the nodulation pathway is SYMRK and it is involved in the plant-bacterial recognition.[41] The pollen tube growth is similar to the infection thread development in that infection threads grow in a polar manner that is similar to a pollen tubes polar growth towards the ovules. Both pathways include the same type of enzymes, pectin-degrading cell wall enzymes.[42] The enzymes needed to reduce nitrogen, nitrogenases, require a substantial input of ATP but at the same time are sensitive to free oxygen. To meet the requirements of this paradoxical situation, the plants express a type of haemoglobin called leghaemoglobin that is believed to be recruited after a duplication event.[43] These three genetic pathways are believed to be part of a gene duplication event then recruited to work in nodulation.Phylogeny and taxonomy
Phylogeny
The phylogeny of the legumes has been the object of many studies by research groups from around the world. These studies have used morphology, DNA data (the chloroplast intron trnL, the chloroplast genes rbcL and matK, or the ribosomal spacers ITS) and cladistic analysis in order to investigate the relationships between the family’s different lineages. The studies have confirmed that the traditional sub-families Mimosoideae and Papilionoideae are each monophyletic but both are nested within the paraphyletic sub-family Caesalpinioideae. All the different approaches have yielded similar results regarding the relationships between the family's main clades, as shown in the cladogram below.[9][44][45][46][47][48][49][50][51]
From Wikipedia, the free encyclopedia* * *
References: „Флора на НР България”, том VI, БАН, София, (1976), Wikipedia, the free encyclopedia
Distribution in Bulgaria: (Conspectus of the Bulgarian Vascular Flora) = conspectus&gs_l= Zlc.
Distribution:GENERA:
Genus Anthyllis L. - Rosid dicot genus, Kidney-vetch Genus Astragalus L. - Milkvetch, Locoweed Genus Cercis L. - Redbuds Genus Chamaecytisus Link - x x x Genus Chamaespartium Adans. - x x x Genus Coronilla L. - False crown-vetch Genus Galega L. - Goat’s rue, Rosid dicot genus Genus Genista L. - Broon Genus Laburnum Medic - Golden chain Genus Lathyrus L - Vetch Genus Lotus L. - Bird's-foot trefoils, Bacon-and-eggs, Deervetches Genus Medicago L. - Burclover Genus melilotus Mill. - Melilot or Sweet-clover Genus Onobrychis - Sainfoins Genus Ononis L. - Restharrows Genus Robinia L. - Locust Genus Trifolium L. - Clover Genus Vicia L. - Vetches Genus Wisteria Nutt. - x x x
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