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33. Lappula Moench, Methodus. 416. 1794.

鹤虱属 he shi shu

Echinospermum Sweet ex Lehmann.

Plants annual or biennial, rarely perennial, pilose, strigose, rarely sericeous throughout, hairs tuberclelike or discoid at base. Leaves alternate. Inflorescence a drepanium, elongated after anthesis, bracteate. Calyx 5-parted nearly to base; lobes frequently enlarged in fruit. Corolla light blue, rarely white, campanulate or low salverform; tube short; throat appendages 5, trapeziform; limb 5-parted. Stamens included. Ovary globose, 4-parted. Style not exserted; stigma capitate. Gynobase subulate veined, ca. as long as or surpassing nutlets, rarely shorter. Nutlets 4, adherent to gynobase by entire adaxial rib or only by rib base, erect, homomorphic or heteromorphic; margin of abaxial surface usually with 1(-3) rows of glochids; bases of glochids separated or contiguous or confluent into a wing, rarely reduced to tubercles.

About 61 species: temperate regions of Africa, Asia, Europe, and North America; 36 species in China.


1 Gynophore free end (vertical part of gynobase) rising above nutlets as thick tetrahedral mucro with short style at apex.   (2)
+ Gynophore entirely hidden between nutlets (gynophore should not be confused with the thin style sometimes elevated above nutlets as in L.brachycentra and L.microcarpa).   (4)
       
2 (1) Nutlets 3-5 mm, lustrous, stonelike, tuberculate, with thick walls and without ribs or disc or anchor-like tip on tubercles   1 Lappula spinocarpos
+ Nutlets 2-3 mm, finely tuberculate, with very short glochids on margin of inconspicuous disc, without glochids on margin of well-developed disc.   (3)
       
3 (2) Calyx lobes exceeding fruit; pedicel always straight   3 Lappula occultata
+ Calyx lobes short, appressed to base of much longer fruit; pedicel declining, sometimes recurved   2 Lappula sinaica
       
4 (1) Developed nutlets 1, the other 3 aborting   31 Lappula monocarpa
+ Developed nutlets 4.   (5)
       
5 (4) Disc of nutlets narrow and nearly closed, marginal parts of nutlets involute on disc.   (6)
+ Disc of nutlets well developed, marginal parts of nutlets not involute on disc.   (7)
       
6 (5) Glochids on margin of disc ca. 0.5 mm   4 Lappula balchaschensis
+ Glochids on margin of disc ca. 1.5 mm   5 Lappula deserticola
       
7 (5) Disc margin of all or (1 or) 2 nutlets with well-developed mostly broad wing.   (8)
+ Disc margin of nutlets without well-developed wing, marginal prickles of disc sometimes widened and confluent into a very narrow winglike border.   (16)
       
8 (7) Perennial herbs.   (9)
+ Biennial or annual herbs (when uncertain, stems straight, tall).   (10)
       
9 (8) Herbs forming small tufts of leaf rosettes, with several or sometimes many lateral ascending stems; nutlets 5-7 mm across   33 Lappula transalaica
+ Cespitose herbs, with 1 or 2 straight stems; nutlets ca. 3 mm across   15 Lappula caespitosa
       
10 (8) Nutlet with entire wing, pubescent or not.   (11)
+ Nutlet with dentate wing, not pubescent.   (12)
       
11 (10) Nutlets tuberculate, densely pubescent, without prickles   32 Lappula lasiocarpa
+ Nutlets neither tuberculate nor pubescent, 2 nutlets with a single row of marginal glochids   30 Lappula alatavica
       
12 (10) Biennial or ?perennial herbs, with well-developed lateral rosette of leaves; stems (2 or) 3-5, few branched or not, straight   27 Lappula ferganensis
+ Annual herbs without rosette of leaves after anthesis; stems usually branched.   (13)
       
13 (12) All nutlets homomorphic, with narrow wings to 1 mm, sometimes 1 nutlet wingless, disc of nutlets granulose   19 Lappula granulata
+ Nutlets heteromorphic, 2 with leathery wings and 2 with 2 rows of glochids on disc margin.   (14)
       
14 (13) Corolla limb (5-)5.5-6 mm   34 Lappula lipskyi
+ Corolla limb 2.5-3.5(-4) mm.   (15)
       
15 (14) Marginal glochids in 2 rows, uniform in length   29 Lappula heteromorpha
+ Marginal glochids in 1 row, some short glochids forming a second row near base of nutlets   28 Lappula anocarpa
       
16 (7) Style projecting 1-1.5 mm above nutlets, filiform, persistent at fruiting.   (17)
+ Style hidden among apices of nutlets or projecting less than 0.5 mm above nutlet.   (18)
       
17 (16) Prickles along disc margin 0.5-0.9 mm, mostly developed in 1 row (sometimes a second row of prickles 0.1-0.2 mm, with very small tubercles and anchorlike tips)   10 Lappula microcarpa
+ Prickles along disc margin 0.1-0.2 mm or absent and replaced by tubercles   11 Lappula brachycentra
       
18 (16) Disc margin of nutlets with only 1 row of glochids (sometimes a medial row of glochids present).   (19)
+ Disc margin of nutlets with 2 or 3 rows of glochids.   (29)
       
19 (18) Perennial cespitose herbs, with several to many stems, branched only above   15 Lappula caespitosa
+ Annual or rarely biennial herbs.   (20)
       
20 (19) Corolla sky blue, limb 5-8 mm   35 Lappula macrantha
+ Corolla blue or white, limb to 3.5 mm.   (21)
       
21 (20) Corolla white; nutlets with marginal glochids to 0.5 mm, disc with 1 or 2 medial rows of glochids on tubercles   6 Lappula macra
+ Corolla blue or white-blue; nutlets with longer marginal glochids or disc finely tuberculate.   (22)
       
22 (21) Heteromorphic nutlets present, 2 with short glochids, 2 with longer glochids; homomorphic nutlets sometimes also present.   (23)
+ All nutlets homomorphic.   (24)
       
23 (22) All nutlets heteromorphic, disc and adaxial surface smooth   16 Lappula karelinii
+ Nutlets in lower part of inflorescences homomorphic, disc and adaxial surface tuberculate   13 Lappula duplicicarpa
       
24 (22) Marginal glochids slender, 0.5-0.9 mm   9 Lappula tenuis
+ Marginal glochids 1-2.5(-4.5) mm.   (25)
       
25 (24) Marginal glochids 1-2 mm; plants branched mainly on upper part of stems.   (26)
+ Marginal glochids 2-2.5(-4.5) mm; plants usually branched basally.   (27)
       
26 (25) Plants 25-40(-50) cm high; marginal spines 1-1.2 mm, disc not keeled   8 Lappula intermedia
+ Plants 15-20(-30) cm high; marginal spines ca. 1.5 mm, disc with protruding keel   7 Lappula stricta
       
27 (25) Marginal glochids hollow, terete   36 Lappula physacantha
+ Marginal glochids solid, or subglobose.   (28)
       
28 (27) Pedicel 1.5-4 mm, usually curved; nutlets 3.5-4 mm excluding prickles   12 Lappula semiglabra
+ Pedicel to 1 mm, straight; nutlets ca. 3 mm excluding prickles   14 Lappula patula
       
29 (18) Perennial herbs; stems several, 7-15 cm tall, ascending; basal leaves numerous, forming a rosette; glochids on disc 0.2-0.5 mm.   (30)
+ Biennial or annual herbs; stems few, usually 30-45 cm tall; basal leaves few, rarely forming a rosette; glochids on disc 1-3 mm, rarely smaller.   (31)
       
30 (29) Herbs silvery sericeous; basal leaves linear to filiform, to 4 cm × 1-2 mm; corolla ca. 5 mm; center line of disc of nutlet without glochids   20 Lappula sericata
+ Herbs green-gray pilose; basal leaves spatulate, to 3 cm × 2-4 mm; corolla 2-2.5 mm; center line of disc of nutlet with glochids   21 Lappula himalayensis
       
31 (29) Biennial herbs; corolla with spreading limb, 4-6(-8) mm across.   (32)
+ Annual herbs; corolla with campanulate limb, 2-3.5(-4) mm across.   (35)
       
32 (31) About 2/3 of adaxial angle of nutlets affixed to gynobase; gynobase longer than nutlets   25 Lappula ramulosa
+ Entire adaxial angle of nutlets affixed to gynobase; gynobase shorter than nutlets.   (33)
       
33 (32) Corolla sky blue, limb to 6-8 mm across, throat appendages blue or dark blue   22 Lappula tianschanica
+ Corolla blue, limb to 4-5 mm across, throat appendages white, yellowish in herbarium specimens.   (34)
       
34 (33) Glochids on disc numerous but short, glochids of main row on margin of disc 1-1.5 mm; herbs to 30(-35) cm tall   24 Lappula pratensis
+ Glochids on disc usually 3 along center line, to 1 mm, glochids of main row on margin of disc 1.5-2.5 mm; herbs to 40-45 cm tall   23 Lappula tadshikorum
       
35 (31) Calyx lobes stellate spreading in fruit, elongated to at least 7-8 mm; leaves light green or green, with sparse spreading hairs on large white tubercles; main marginal glochids with large anchorlike tip   8 Lappula intermedia
+ Calyx lobes spreading in fruit, not stellate, elongated to 4-5 mm; leaves usually densely appressed pubescent, grayish green; marginal glochids with small anchorlike tip.   (36)
       
36 (35) Upper part of nutlets separated by deep oval or elliptic pit; style protruding above nutlets (excluding glochids) 0.5-1 mm   26 Lappula consanguinea
+ Upper part of nutlets without pit; style protruding above nutlets to 0.2-0.3 mm.   (37)
       
37 (36) Main marginal prickles widened basally, confluent near base, 0.4-1.5(-2) mm   18 Lappula shanhsiensis
+ Main marginal prickles all separated, 1.5-2 mm (or more)   17 Lappula myosotis

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