Revision of the Genus PhyllolobiumFisch. (Leguminosae-Papilionoideae)

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  • Feddes Repertorium 117 (2006) 12 , 4164 DOI: 10.1002/fedr.200511090 Weinheim, Mai 2006

    2006 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim 0014-8962/06/1-205-0041

    Ludwig-Maximiliams-University Munich, Faculty of Biology, Systematic Botany and Mycology, Munich

    M.-L. ZHANG & D. PODLECH

    Revision of the Genus Phyllolobium FISCH. (Leguminosae-Papilionoideae)

    Summary According to the most recent molecular and pollen evidences, Astragalus L. subgenus Pogonophace is proposed to be extracted from Astragalus and erected as a new genus Phyllolobium FISCH. Based mainly on the specimens checking of major Chinese herbaria, a taxonomic revision regarding this genus is provided in the present paper. 1. Phyllolobium balfourianum (N.D.SIMPSON)

    M.L.ZHANG & PODLECH, comb. nov. 2. Phyllolobium camptodontum (FRANCH.)

    M.L.ZHANG & PODLECH, comb. nov. 3. Phyllolobium chapmanianum (WENNINGER)

    M.L.ZHANG & PODLECH, comb. nov. 4. Phyllolobium chinense FISCH., in SPRENGEL 5. Phyllolobium dolichochaete (DIELS) M.L.ZHANG

    & PODLECH, comb. nov. 6. Phyllolobium donianum (DC.) M.L.ZHANG &

    PODLECH, comb. nov. 7. Phyllolobium enneaphyllum (P.C.LI) M.L.ZHANG

    & PODLECH, comb. nov. 8. Phyllolobium eutrichus (HAND.-MAZZ.)

    M.L.ZHANG & PODLECH, comb. et stat. nov. 9. Phyllolobium flavovirens (K.T.FU) M.L.ZHANG

    & PODLECH, comb. nov. 10. Phyllolobium heydei (BAKER) M.L.ZHANG &

    PODLECH, comb. nov. 11. Phyllolobium lasaense (C.C.NI & P.C.LI)

    M.L.ZHANG & PODLECH, comb. nov. 12. Phyllolobium lineariaurifer (P.C.LI) M.L.ZHANG

    & PODLECH, comb. nov. 13. Phyllolobium milingense (C.C.NI & P.C.LI)

    M.L.ZHANG & PODLECH, comb. nov. 14. Phyllolobium pastorium (H.T.TSAI & T.T.YU)

    M.L.ZHANG & PODLECH, comb. nov. 15. Phyllolobium prodigiosum (K.T.FU) M.L.ZHANG

    & PODLECH, comb. nov. 16. Phyllolobium sanbilingense (H.T.TSAI &

    T.T.YU) M.L.ZHANG & PODLECH, comb. nov. 17. Phyllolobium siccaneum (P.C.LI) M.L.ZHANG &

    PODLECH, comb. nov. 18. Phyllolobium tingriense (C.C.NI & P.C.LI)

    M.L.ZHANG & PODLECH, comb. nov.

    Zusammenfassung

    Revision der Gattung Phyllobium FISCH. (Le-guminosae-Papilionoideae)

    Entsprechend moderner molekularer und pollenmor-phologischer Daten wird Astragalus L. subgenus Pogonophace aus der Gattung Astragalus ausgeglie-dert und in eine eigene Gattung Phyllolobium FISCH. berfhrt. Basierend auf dem reichen Material, vorwiegend in den grossen chinesischen Herbarien, wird eine Revision dieser Gattung vorgelegt.

  • 42 Feddes Repert., Weinheim 117 (2006) 12

    2006 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim www.fedrep.de

    19. Phyllolobium tribulifolium (BUNGE) M.L.ZHANG & PODLECH, comb. nov.

    20. Phyllolobium turgidocarpum (K.T.FU) M.L.ZHANG & PODLECH, comb. nov.

    Astragalus L. subgenus Pogonophace was established by BUNGE (1867) mainly based on a distinct character, the penicillately hairy stigma. Molecular analysis based on the ITS sequence (KANG et al. 2003) has shown that the subgenus is not a monophyletic taxon. Mean-while, the same character was observed from species among several other sections of the genus Astragalus, these sections are not mem-bers of subgenus Pogonophace, i.e., annual: Hispiduli, Annulares (PODLECH 1994); per-ennial: Erionotus (PODLECH 1988). BAKER (1876) described a new subgenus in the genus Astragalus, subgenus Trichostylus BAKER, which was based on the character of the style with hairs on the dorsal side below the stigma. In this subgenus, the species A. heydei BAKER has been thought of having the charac-ter of penicillately hairy stigma. But the posi-tion of these hairs at the style is not constant and even missing in many flowers of this spe-cies. Furthermore, this character can be found commonly in other sections of Astragalus, such as sections Caprini, Astragalus, Aegacantha and Pelta. Therefore, penicillately hairy stigma should not be thought to be a synapomorphic, but a parapomorphic or polymorphic character among the groups mentioned above, which should not be used as diagnostic character in the higher rank among these taxa. As a single character regarding the penicil-lately hairy stigma, it should be considered as having heterogenous origin. In some groups within subgenus Pogonophace we can find in-teresting pollen brush hairs (KANG & ZHANG 2004), those groups include the sections Bibrac-teolati, Phyllolobium and Trichostylus. Com-bined with the characters of non-interlocking keel and wing petals (for the absence of bosses on the keel and matching sockets on the wings), these sections form a very natural group and are similar to the genera which are clearly belonging to subtribe Coluteinae BENTH. & HOOK. f. instead to Astragalus (belonging to subtribe Astragalinae BENTH. & HOOK.f.).

    BARNEBY (1964) was the first, who found the strong similarities between Astragalus complanatus and the genera Sphaerophysa and Swainsonia of subtribe Coluteinae. However, those species only with hairy stigma but with-out the pollen brush, such as Astragalus ho-antchy FRANCH. belonging to former Pogono-phace, have interlocking Keel and wing petals, therefore they belong clearly to the genus As-tragalus. KANG & ZHANG (2004) proposed that those species with pollen brush hairs should be clas-sified as a new genus Phyllolobium, which was created by FISCHER in 1818 with the type spe-cies later named as Astragalus complanatus. During PODLECHs stay in China this year, the important characters of all species of sect. Bi-bracteolati and the related ones have been investigated once more. Combined with our previous studies (ZHANG 2000, 2002; KANG & ZHANG 2004), it is quite clear that all species of the sections Phyllolobium, Bibracteolati and Trichostylus form a natural group, namely the genus Phyllolobium FISCH. These species occur mainly in the southwestern high mountains of China (ZHANG 2003). A taxonomic revision of all species of this genus is given as follows.

    Phyllolobium FISCH. in SPRENGEL, Novi Provent.: 33. 1818

    Astragalus sect. Phyllolobium (FISCH.) BUNGE, Mm. Acad. Imp. Sci. Saint Ptersbourg 11(16): 3. 1868 A. sect. Chlorostachys subsect. Phyl-lolobium (FISCH.) PODLECH, in Fl. Iranica 174: 79. 1999 A. sect. Oliganthi N.D.SIMPSON, Notes Roy. Bot. Gard. Edinburgh 8: 251. 1915.

    Type: Phyllolobium chinense FISCH. (Syn.: Astragalus complanatus BUNGE). = Astragalus subgen. Trichostylus BAKER, in Fl.

    Brit. India 2: 118. 1876 A. sect. Trichostylus (BAKER) TAUBERT, in ENGLER & PRANTL, Natrl. Pflanzenfam. III, 3: 286. 1894. Type: A. heydei BAKER.

    = Astragalus sect. Bibracteolati N.D.SIMPSON, Notes Roy. Bot. Gard. Edinb. 8: 252. 1915.

  • M.-L. ZHANG & D. PODLECH: Revision of the genus Phyllolobium FISCH. 43

    2006 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim www.fedrep.de

    Lec to type : (PODLECH 1990): A. camptodontus FRANCH.

    Plants perennial with mostly well developed stems, with basifixed hairs. Stipules free from the petiole and from each other, very rarely shortly connate behind the stem. Racemes several, with a distinct peduncle. Bracts persis-

    tent. Bracteoles present or more rarely absent. Calyx campanulate. Keel and wing petals not interlocking for the absence of bosses on the Keel and matching sockets on the wings. Style with a brush of straight, rigid, short hairs just below the glabrous stigma. Legumes unilocular or incompletely to completely bilocular.

    1 Plants with slender stems up to 90 cm long, at least in lower half with long, horizontally spreading side-branches; hairs slender, 0.10.4 mm long . . . . . . . . . . . . . 8. Ph. eutrichus

    Stems mostly shorter, without horizontally spreading side-branches, if sometimes as long, than with other kind of hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

    2 Plants in all parts covered with bladdery, scale-like hairs up to 0.1 mm long . 4. Ph. chinense Plants at least at the calyx with slender, longer, not bladdery hairs. . . . . . . . . . . . . . . . 3 3 Petals yellow, rarely with violet tipped keel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Petals blue or violet, rarely yellow at the base only . . . . . . . . . . . . . . . . . . . . . . . . 6 4 Plants 3075 cm tall; calyx 712 mm long, teeth 49 mm long; keel as long as the wings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. Ph. dolichochaete Plants 1050 cm tall; calyx up to 10 mm long, teeth up to 6 mm long; keel longer than the

    wings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 5 Leaflets in 713 pairs, 516 14 mm, distinctly more than 2 times longer than wide;

    peduncles 311 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9. Ph. flavovirens Leaflets in 510 pairs, 512 2.55 mm, ca. 2 times longer than wide; peduncles 1.5

    4 cm long; petals yellow with violet-tipped keel. . . . . . . . . . . . . . . . 2. Ph. camptodontus 6 Plants at least in part with spreading hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Plants with appressed to ascending hairs only . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 7 Calyx 810 mm long, white hairy, teeth 4.56 mm long; standard 1519 mm long; leg-

    umes with a stipe ca. 5 mm long, 2530 mm long, bilocular . . . . . . . 20. Ph. turgidocarpum Calyx shorter, white and black hairy, teeth mostly shorter; standard 713 mm long; leg-

    umes with a stipe up to 3 mm long or subsessile, up to 20 mm long, unilocular . . . . . . . 8 8 Leaflets 820 mm long; bracts 810 mm long; calyx 78 mm long, with teeth 45 mm

    long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12. Ph. lineariaurifer Leaflets distinctly smaller; bracts at most up to 4 mm long; calyx up to 6 mm long with

    shorter teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 9 Stipules 2.54.5 mm long; leaflets 315 mm long; bracts 1.54 mm long; bracteoles

    0.52 mm long; calyx teeth 1.54 mm long; legumes 56.5 mm high and wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19. Ph. tribulifolium

    Stipules up to 2.5 mm long; leaflets 26 mm long; bracts up to 2 mm long; bracteoles mostly absent, if present, than ca. 0.3 mm long (in Ph. milingense); calyx teeth up to 2.5 mm long; legumes either much higher than wide or much wider than high . . . . . . . . 10

    10 Stems with appressed hairs; leaflets loosely to rather densely hairy on upper side; bracts 1.52 mm long; legumes 1520 mm long, 812 mm high, strongly compressed laterally, without beak . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10. A. heydei

    Stems with partly spreading hairs; leafltes glabrous to subglabrous on upper side; bracts 0.61 mm long; legumes 1015 mm long, 2.53 mm high and 57 mm wide, with a short beak . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13. Ph. milingense

    11 Leaflets on both sides loosely to densely hairy . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 Leaflets on upper side glabrous or rarely sparsely hairy . . . . . . . . . . . . . . . . . . . . . . 13 12 Stipules 23 mm long; bracts 46 mm long, with spreading hairs; calyx 78 mm long,

    covered with spreading white and brown blackish hairs . . . . . . . . . . . 3. Ph. chapmanianus

  • 44 Feddes Repert., Weinheim 117 (2006) 12

    2006 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim www.fedrep.de

    Stipules 47 mm long; bracts 610 mm long, with appressed to ascending hairs; calyx 910 mm long, covered with appressed to ascending white and shorter black hairs . . . . .

    . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11. Ph. lasaense 13 Standard up to 10 mm long; bracts up to 2 mm long; bracteoles absent . . . . . . . . . . . . 14 Standard at least 12 mm long; bracts mostly distinctly longer . . . . . . . . . . . . . . . . . . 16 14 Plants up to 50 cm tall; calyx 78 mm long; legumes 2530 mm long . . . . . Ph. siccaneum Plants up to 30 cm tall; calyx up to 5.5 mm long; legumes up to 13 mm long . . . . . . . . . 15 15 Stem and peduncles loosely to rather densely hairy; leaflets narrowly elliptic, 311

    12 mm acute; keel as long as the wings . . . . . . . . . . . . . . . . . . . 15. Ph. prodigiosum Stem and peduncles glabrous to sparsely hairy; leaflets elliptic, 49 1.54, obtuse; keel

    distinctly longer than the wings . . . . . . . . . . . . . . . . . . . . . . . . 16. Ph. sanbilingense 16 Stem, rachides and peduncles glabrous or sparsely hairy; leaflets 1230 3.510 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14. Ph. pastorium Stem, rachides and peduncles loosely to rather densely, sometimes only sparsely hairy;

    leaflets up to 12 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 17 Stem, rachides and peduncles covered with hairs up to 0.15 mm long and also with much

    shorter, minute, nearly globular hairs; calyx teeth 2.53 mm long; legumes very shortly acuminate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7. Ph. enneaphyllum

    Stems without minute subglobular hairs; at least the longer calyx teeth 34.56 mm long; legumes with a beak 24 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

    18 Stipules 13 mm long, the lower ones completely embracing the stem but free from each other; keel shorter than wings . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6. Ph. donianum

    Stipules 35 mm long, the lower ones not or up to half embracing the stem; keel longer than wings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

    19 Plants 2040(60) cm tall; racemes 27(9)-flowered; petals violet or in lower part yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. Ph. balfourianum

    Plants 1025 cm tall; racemes 14-flowered; petals purple red, standard in the middle with a white, purple-striate spot . . . . . . . . . . . . . . . . . . . . . . . . . . . 18. Ph. tingriense

    1. Phyllolobium balfourianum (N.D.SIMPSON) M.L.ZHANG & PODLECH, comb. nov.

    Astragalus balfourianus N.D.SIMPSON, Notes Roy. Bot. Gard. Edinburgh 8: 123. 1913.

    Type: (WENNINGER 1991): [China] Yunnan, eastern flank of Lijiang (Lichiang) Range, 12000, ix. 1910, FORREST 6606 (lectotype: E!: foto MSB!; isotype: W!). = A. minutebracteolatus N.D.SIMPSON, Notes Roy.

    Bot. Gard. Edinburgh 8: 243. 1915. Type: W. China, 3300 m, viii. 1903, WILSON 3420 (K!: foto MSB!). = A. schneideri ULBR., Notizbl. Bot. Gart. Berlin-

    Dahlem 8: 85. 1921. Lec to type (WENNINGER 1991): [China] Yunnan, Lijiang (Lichiang), 3600 m, 01. 09. 1914, SCHNEIDER 2337 (K!: foto MSB!). Note : the original material at B is de-stroyed.

    = A. prattii N.D.SIMPSON, Notes Roy. Bot. Gard. Edinburgh 8: 2...

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