Legumes of the world [1st ed.] 9781900347808, 1900347806

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In nrtlrern Er-trope species f Gleclit,:sict, Lctlxrnttm ancl llc¡bitti,a have l>een wiclely g1'()wl1 as stl'eet, p:rrk ancl garclen trces ft>r centttries, 'lncl extics, sttcl-i as thc winter-flowering Ar-rstlalian acztci:is, are bccoming vcry popr-rlar. A hanclfil of species are solcl as ctlt flowers ('florist's mimsa' lteing, confr-rsingly, thc cmtnon name of a small nlttnbet' ol Australian Acctcict species).

Sweet peas (Lct.tbyrzl.s species), h,rpins (Lupirt'us species) ancl brf genera ancl species have been introch.rcecl intct hrticr.tltttre. Some of these, such as Acacict., Dicbrostc.tcbys, Lettcctentt, Mimr¡salncl Sesbanict., have becme weecly ancl invasive, having escapecl their natural pests ancl cliseases. Mtlltipr-rrpose trees ¿rncl shrults have long been selectecl ancl refinecl by local cc;mtnltnities fìrr shacle, ornAfilent, forage, foclcler, fltelwcl, bee fìrragc fol honey proch-tctin, ancl soil enrichment. Regional favottrites inch'rcle Butea, Dctlbergi,ct ancl Milleïtia in Inclia; C'ct'llictnclra, Gliricicli,ct., Inga ancl Lettc¿tettct in Central America (Pllrill, 1997) '¿ncI l;øiclberbía ancl Acctcict, in Africa. Some ale gr()wn as itnpenetrable spiny heclges, living fencelines ancl winclbleaks. In tl-ie 19il0s ancl 1990s a number r>f seecl banks targetecl mr-rltipr-llpost' leguminus trees for introclttctirestry systems ancl legtltnes

form :r large

percentage notnic notes which fonn part of the textr,rral blt>ck f each genr-rs in this

7.2.10

1.2.1t L2.12 r.2.13 7.2.14

72 73 .

Colophospennurn Ilarclwichia l)aniellia

/)

/('t

/6 77 77

Iìurypetalnrn.....

78

r.2.t5

Epen-ra Augc>uarclia

I,2.16

Stcrnonr>cctleus

r.2.17

Peltogyne Hyrnenaea

79 79 80

Guibr>urtia

u1

r.2.20 1.2.2t r.2.?2

Hylclenclron..,..

7.2.23

Tessrnannia

L2.2,i

Sìnclora

1.2.25 1.2.26

Sinckl'opsis Copaifela Detarir¡m

1.2.28

82 82 82 83

(ìilletioclenchon llaikiaea

r.2.29

Enclertia Lysiclice

1 2.30

Sarac¿t

1.2.i t

Leucstegane

1,2.3¿

Talbtiell¿r

1.2.33 7.2.3tt 1.2.3s 1.2.36

ScoroclophloeLls Cllrclia

r.2.37

7L)

Lcl¡rr-rnioclenclron .

83 83

.

Plagiosiphon..... Micklethwaitia...,

1.2.59 1.2.60

I.2,6I

.

Br()wn('()psis Macri>llc¡biurn . .

Dicymbe

Antlrr'evillerr

)iclelotia

I

r.2.7r

1 ) -7)

I'elleglinioclcnrl ton Gillrcltioclenrlt'on

t.2.73

Isobel'linil

1.2.74

Ocltl
t)

450

SESBANIEAE

3.21..1' Sesbania

3.22

453

LOTEAE

3.22.1 HipPocrePis .

.

3.22.2Scorpiurus.... 3.22.3 Securigera....

3.22.4 Coronilla..... 3.22.5 Podolotus

3.22.6 Anthyllis 3.22.7 Hymenocarpos 3.22.8 Pseudolotus... 3.22.9 Antopetitia . . . 3.22.1,0 Hosackia . . . .

439 440 440

3.24.1.1.

3.23

ROBINIEAE

Tripodion Hammatolobium Cytisopsis

3.23.t

Hebestigma....

3.23.2

Lennea

Gliricidia

44t

.

44r 441.

3.23.s

Poitea

.

Olneya Robinia Poissonia Coursetia

3.4.9 Peteria 3.23.10 3.23.1

1

Genistidium.... Sphinctospermum

.

486 486

4ó/ 487

49r 491.

49r

Eversmannia

Hedysarum . . . Corethrodendron

3.25.9

Sulla Taverniera....

3.25.1.0

Onobrychis . .

.

40t

3.26.1 Cicer...

465 46>

3.27

472 472 472

485

4ót 486

.

Calophaca .... Caragana ..... Halimodendron Alhagi

3.26

47r

.

HEDYSAREAE

3.25 3.25.1.

462 462 463 464 464

468 468 468 469 469 470 470

482 482 483 483 483

484 484

3.24.23 Streblorrhiza 3.24.24 Galega

461.

Tetragonolobus

.

....

3.24.2I Clianthus . . . . 3.24.22 Carmichaelia .

461 462

....

482

....

3.24.79 3.24.20 Montigena

3.22.1,3 Kebirita 3.22.1.4 Omleya 3.22.1.5 Acmispon

Syrmarium Lorns . Dorycnium

481.

.

Swainsona....

3.25.7 3.25.8

3.22.16 3.22.17 3.22.18 3.22.19 3.22.20 3.22.27 3.22.22

479 479 480 480 480

3.24.1,5 Eremosparton 3.24.1.6 Sphaerophysa 3.24.17 Lessertia 3.24.1"8 Sutherlandia . .

461.

.

4/A

Barnebyella . .,

3.24.14 Smirnowia

Ornithopus 3.22.12 Dorycnopsis

.

478 478

3.24.1.2 Colutea 3.24.1.3 Oreophysa . . .

3.25.6

.

GALEGEAE

3.24.10 Ophiocarpus

3.25.2 3.25.3 3.25.4 3.25.5

.

442 442 442 443

4>8 458

3.24

3.24.1 Glycyrrhiza 3.24.2 Chesneya.... 3.24.3 Spongiocarpella 3.24.4 Gueldenstaedria 3.24.5 Tibetia 3.24.6 Erophaca.... 3.24.7 Oxytropis 3.24.8 Biserrula.... 3.24.9 Astragalus...

459 459 460 460 460

3.4.3 3.4.4 3.4.5 3.4.6 3.4.7

.

4>/ +>/

4>A

3.22.1,1,

Campylotropis

471

.

424

3.19.7 3.L9.2 3.L9.3 3.19.4 3.79.5 3.19.6 3.19.7 3.19.8 3.19.9

3.1.8.39 Rhynchosia

Chrysoscias . . .

....

DESMODIEAE

409 409 409 410

Bolusafra Carrissoa

3.78.74 Dolichos

3,19

408 408 408

.

3.18.7 3 Austrodolichos

423 423

....

i.tg.zs Melliniella.' '

3.20

422 423

.

.. . ..

3.18.83 Phaseolus 3.I8.84 Ramirezella 3,18.85 Strophostyles 3.1s.86 Dolichopsis 3.18.87 Macroptilium 3.18.88 Mysanthus . .

406 407

3.1.8.30 Vandasina

Sphenostylis . 3.18.7I Nesphostylis

ß

421.

422 422

.

443 444 444

26 AlYsicarPus á,g.27 Desmodiastrum q

i.tg.lo

42t

.

. t1c\l.- )5- Christia ,,t,

420 420 420 421,

\ù/ajira

404

406 .

Phylacir.rm . . Neocollettia .

404

4U>

4t9

.....

Glycine

3.t8.82 Oxyrhynchus

404

.

4r9

3.\8.64 Amphicarpaea 3.I8.65 Teramnus

3.1.8.72 Alistilus

402

.

41.9

399 400 400

.

3.18.2I Clitoriopsis

3.18.60 Eminia 3,18.61. Sinoclolichos 3.I8.62 Pseuderninia 3.18.63 Pseuclovigna

398 398 398 399 399

401,

3.18.20 Periandra

418

3.18.66 3.18.67 3.18.68 3,18.69 3.18.70

401.

3.1,8.79 Centrosema . . .

3.18.35 3,18.36 3.18.37 3.18.38

395 396 397 397 397

400

.

3.18.17 Clitoria 3.18.18 Barbieria

3.18.31" Spatholobus . .

395 395

3.18.58Ptterarta.". 3.18.59Nogra .. ..

Sartoria 3.25,r2 Ebenus 3.25.1.1"

.

492 492 493 493 493 494 494 495 495

CICEREAE 497

TRIFOLIEAE

3.27.1.

Parochetus

.

507

211

Trifolium .

.

501

''

3.27.3

Ononis

3.27.4

Melilotus .

3.27.5 3.27.6

Trigonella Medicago .

3.28

FABEAE

502 .

.

502 503 503

3.28.7

Vicia .

3.28.2

Lens

)00

3.28.3

3.28.4

Lathyrus Pisum

3.28.5

Vavilovia

507 508 508

.

506

INTRODUCTION 79

BiogeograPhY of the Leguminosae byß.D.Schrire,

G' P' Lewis and M' Lavin

Introduction 'moist equatorial megathermal' Morley (2000) implies a Ma in the Campanian ot 84-74 c' .r.lnin'of legumes, thus supporting Cretaceous, Upper the of Maãstrichtian

u

,Uø"r, Gonclwana

(or austrotropical) origin of the

& Polhill, 1981)' iamily (Raven & Axelrod, 1974;Raven

This long-held \flest Gondwana hypothesis for the origin of legumes requires the family diversification (i.el, that of the legume crown clacle; Fig. 3) to be at (Lavin et al., 2000; Davis et al., least 100 - 90 Ma in age South America were last in and Africa when ZO02b), Raven & Axelrod (1974) and although near contact (2000; that dispersal routes suggest 2003) Morley

existed over islands and ridges between these continents until c. 65 lvla. In addition, it is often stated

(e.g., Modey, 2000) that the tropical angiosperm fossil record is biased to Laurasian collection localities, and that if more fossils were available from South America and Africa, the stratigraphic record for many groups would be older.

Although

it is true that legumes are now highly

diverse in both Africa and South America, fossil data alone indicate that neither a moist megathermal origin, nor a Mesozoic age of legume diversification is likely. The clear message derived from fossil legume studies such as those of Herendeen et al. (1,992), Herendeen & Dilcher (7992), Herendeen (2001) andJacobs (2003)

is that all three subfamilies of legumes are well represented in the fossil record in North America, Europe, Africa, and Asia by at least fruits and leaves from recent times back to the palaeocene. putative earlier legume fossils inclucle only pollen ancl wood specimens that lack any specific legume

distribtrtion patterns. Four generalised areas of endemism predictive for legume distributions were identified at the biome level, including a newly

diversification of the family must have occurred soon after. By the middle Eocene (c. 50 Ma) nearly all of the major lineages have a fossil recorcl in North America, Africa, ancl Asia (e.g., Axelrod, 1))2; Lavin, ::::p", 1998; Herendeen et at., 7992; Herendeen, 2001). In a chronogram of the matKphylogeny (Lavin in press) shãws minimal resolution at the base tree between major crown clacle cliversifications, Ï.tlte the abrupr of all br.rt one major l1^o1:u,,1g ctade

1:dl.tion, et^a|.,

"-"rg".,." of legumes bärween 6o anct 50 Ma.

20

LEGUMES OFTHEWORLD

P'

exhaustive sampling of various local subclades. A strict supertree (sensu Sanderson et al., 1998) was readily constn-rcted manually because of the high compatibility of all the component trees. Essentially, the large-scale

much before 60 Ma. Remarkably, the rapid

throughout the Cenozoic, and the abrupt absence of deciduous legume leaflets and pods prircr to the Late Palaeocene, the origin of legumes ii unlikely to be

Tribe Caesalpinieae Photograph by

Iargely stable, although the circumscriptions of terminal monophyletic groups will continue to be refined as more taxa ane sampled; it is, however, already requiring us to modify our traditional concepts of legume biogeography. Data from these large-scale molecular phylogenies have recently been compiled by Schrire et al. (2005), to identify the major subgroups of legumes and the biogeographical inter-relationships among these groups. The chloroplast matK phylogeny representing a comprehensive sampling of all major legume groups (\ùlojciechowski et a1.,2004) served as the backbone for the supertree (summarised with modifications in Fig. 3). In addition, major legume subclades,detected in the phylogenetic analyses of chloroplast trnL and rbcL sequences noted above, were used in supertree construction because they represented a more

of diverse legume macrofossils

spatial continuity

-

2003a; Forest (unpubl. data) and Mimosoideae (Luckow et al., 2000; 2003; Miller & Bayer (2003), as well as chloroplast trnL and rbcL region analyses of Papilionoideae (Doyle et al., 2000 Pennington et al., 2007; Kajita et al., 2001.). The emerging supertree is

matK phylogeny (\Øojciechowski e/ al., 2004) represented all major clades of legumes, and the other molecular phylogenetic studies identified the broader constituents of these monophyletic matK subclades. The local subclades were then scrutinised for global

synapomorphies, and even then such fossils go back only to the latest Cretaceous. Given the temporal and

Bolsamocarpon brevlfolium

The legume supertree (Fig. 1) is derived from a familywide phylogeny based on DNA sequences of the chloroplast rnatK region (rüØojciechowski et al., 2004) and complemented by chloroplast trnL region analyses of the Caesalpinioideae (e.g., Bruneau et al., 2000; 2001; Fougère-Danezan et al., 2003; Herendeen et al.,

recognised amphi-Atlantic Succulent Biome. Schrire et

al. (2005)

subjected the resulting taxon-biome

supertree (Figs.4-I4) to cladistic vicariance analyses, to detect a generalisecl pattern of biome relationships for legumes. Chronograms derived from rate-smoothed bayesian consensus trees of the matK phylogeny (Lavin et a1.,2004; in press) provided comparative clade ages within the family, based on thirteen time constraints derived from fossil evidence.

Boxter

BIOGEOGRAPHY OFTHE LEGUMINOSAE 2T

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LEGUMES OF TH

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biome (abbreviatecl here to Íoresr, thicket and btrshland 2) a fire-tolerant, sttcculent-poor and S-), or Succulent srass-rich, seasonally dry tropical forest, woodland and

Ër3F

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graclients. The generalised combination of these

in recognising the four biomes which are These include 1) a semi-arid, firebelow. ¿"scrib",l srtcculent-rich and grass-poot dry tropical

Êylo

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6

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(fire-histoly to no fire..-,.rerate), and disturbance

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llã"rc.ilred in general ecological terms involving (tropical to I..'.,irt,,r" (wet to dry), temperatllre

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È I å i ;Ê ii¡ i i* îãÈ ã íZzi ? 1i: 1 i i : * ã; iîi î: à : iå i I åç ?:?'=lllazË ååfr í g ïçÈ ããËiã +i|,*z î i! ; i ã; l;? iù : ä Fë ã 1 àËiÈiå iï ;i;: 3i I ã i¡ i: ËÉiEã+i{lã i ' ; È âå I lsi:i-ls iËi i ièåå ' ã¡ í i å ;* i âÈ i; ¡ i iåi i i i I* r rí.'?,i-+ I is;SFF€ iãË

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1 (e)

(60',)

Desm. 527

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IRLC (inct. IRLC

54 (4357)

Millettioids)

lM¡[.,37]

LEGUMI NOSAE

731 (79327)

7 (21) 16 (31)

289 (40)

173 (24) 4487 (23)

(3e)

28 (54)

(15)

(3t, (t4)

6524

7

7.2

42

(
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