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FOC | Family List | FOC Vol. 2-3

1. Pteridophytes (Lycophytes and Ferns)

蕨类植物 jue lei zhi wu

Authors: Youxing Lin, Li-Bing Zhang, Zhang Xianchun, He Zhaorong, Zhongren Wang, Shugang Lu, Sugong Wu, Prof. Fuwu Xing, Zhang Gangmin, Wenbo Liao, David S. Barrington, Xiang Jianying, Masahiro Kato, Wang Faguo, Kunio Iwatsuki, Qi Xinping, Michael G. Gilbert, Yan Yuehong, Ding Mingyan, Peter H. Hovenkamp, Liu Jiaxi, Hans P. Nooteboom, Shiyong Dong, Jefferson Prado, He Hai, Ronald Viane, Maarten J. M. Christenhusz, Zhang Qiaoyan, Shannjye Moore, George Yatskievych, Atsushi Ebihara, Zhaohong Wu, Li Zhongyang, Shunshuke Serizawa, Jin Xiaofeng, Barbara S. Parris, Bingyang Ding, Tom A. Ranker, Quanru Liu, Norio Sahashi, Elisabeth A. Hooper, Shi Lei, Julie Barcelona, Alexandr Shmakov, Harufumi Nishida, Sujuan Lin, Alan R. Smith, Michele Funston, Christopher Haufler, Nicholas J. Turland, Judith Garrison Hanks, John T. Mickel, Yoko Kadokawa, Kathleen M. Pryer, W. Carl Taylor, David M. Johnson, Edward R. Alverson, Jordan S. Metzgar & Shigeo Masuyama

Plants with a regular alternation between larger asexual sporophytes and mostly inconspicuous, sexual gametophytes, mostly free-living but retained within sporocarps of heterosporous ferns or developed mostly within spore walls of heterosporous lycophytes (Iseotaceae and Selaginellaceae). Sporophytes mostly with roots (absent in Psilotaceae), stems, and leaves, and with well-developed vascular strands. Stems mostly rhizomes, protostelic, siphonostelic, solenostelic, or dictyostelic, sometimes polystelic, some with limited secondary thickening, articulate in Equisetaceae. Leaves microphylls: scalelike or linear with a single vascular strand and a single axillary sporangium, or fronds (megaphylls): with branched vascular strands, lamina often divided, often compound, with many sporangia on abaxial surface, margin, or specialized sporophore, forked and subtending a 3-lobed sporangium in Psilotaceae. Sporangia thick- or thin-walled, homosporous or heterosporous, sessile or stalked, rarely enclosed within sporocarps. Spores trilete or monolete. Gametophytes filamentous or thalloid, autotrophic or mycotrophic. Male gametes (antherozoids) bi- or multiflagellate. Female gametophytes (egg cells) borne singly within flask-shaped archegonia (largely adapted from Kramer & Green in Kubitzki, Fam. Gen. Vasc. Pl. 1: 11. 1990).

Some 265-300 genera and 10,900-11,100 species recognized worldwide (numbers based largely on Smith et al., Taxon 55: 705-731. 2006): extant pteridophytes are cosmopolitan but are much better represented in the humid tropics, with only a few families (e.g., Dryopteridaceae) well represented in subtropical and temperate regions and rather few extending into alpine regions (e.g., Woodsiaceae) and more arid regions (most notably Pteridaceae subfam. Cheilanthoideae); 177 genera (three endemic, one introduced) and 2,129 species (842 endemic, four introduced) in China.

Pteridophytes are conventionally divided into four major groups, Psilotatae, Lycopodiatae (lycophytes or club mosses), Equisetatae (horse tails), and Filicatae (ferns) (Kramer & Green, loc. cit.), or five major groups when Iseotinae/Iseophytina is also recognized (e.g., Ching, Acta Phytotax. Sin. 16(3): 1-19. 1978). Molecular data shows that the lycophytes (Iseotaceae, Lycopodiaceae, and Selaginellaceae), characterized by microphylls and protostelic or polystelic vascular strands, are sister to all other vascular plants but Psilotatae and Equisetatae, along with the Ophioglossaceae and Marattiaceae, are better regarded as basal relatives of the true ferns (Osmundaceae onwards), forming a monophylletic group, the monilophytes, more closely allied to the spermatophytes, the seed-bearing gymnosperms and angiosperms than to the lycophytes (Pryer et al., Nature 409: 618-622. 2001; Smith et al., loc. cit.).

The delimitation of families of extant pteridophytes had been very controversial in the past but a consensus has been emerging on overall relationships, based largely on molecular data from the chloroplast genome. This has shown that traditional characters, particularly those of venation, sori, and indusia, show many parallelisms and convergences such that related genera were placed in different, polyphyletic or paraphyletic, families. This had already been recognized by some botanists who identified many such anomalous genera and placed them within smaller, more homogenous families. The new molecular data showed that some of these families were nested within other families, rendering some families paraphyletic and thus untenable to some modern systematists. Thus, the decision was taken for the Flora of China to follow the most recent overall account of the pteridophytes at family level, that of Christenhusz et al. (Phytotaxa 19: 7-54. 2011), which is largely based on Smith et al. (loc. cit.). Christenhusz et al. proposed the recognition of 48 families, 38 of which occur within China. At generic level, various genera are recognized for Flora of China based on molecular and/or morphological evidence.

Pteridophytes were dominant land plants during the Carboniferous era and a major source of today’s coal and oil. Extant pteridophytes are cosmopolitan but are much better represented in the humid tropics, with only a few families (e.g., Dryopteridaceae) well represented in subtropical and temperate regions and rather few extending into alpine regions (e.g., Woodsiaceae) and more arid regions (most notably Pteridaceae subfam. Cheilanthoideae).

In contrast to the 177 genera and 2,129 species recorded from China, the Flora of North America, covering a similar area, has only 96 genera and 554 species. This illustrates the size and importance of the pteridophyte flora of China, which is much richer than that of other comparable temperate areas and is probably the most species-rich country in the world.

Detailed citations for the corresponding volumes of Flora Reipublicae Popularis Sinicae (FRPS), volumes 2 (1959), 3(1) (1990), 3(2) (1999), 4(1) (1999), 4(2) (1999), 5(1) (2000), 5(2) (2001), 6(1) (1999), 6(2) (2000), and 6(3) (2004), are provided under each family in this volume.

Key to Families

1 Sporangia solitary in axils of simple or once-forked leaves   (2)
+ Sporangia several to very many together borne directly on surface of fronds or frond axes or on specialized sporophores borne on frond or in achlorophyllous strobili, sometimes enclosed within sporocarp or indusium   (5)
2 (2) Plants stemless, leaves all fertile, linear, in dense rosettes from subterranean corm, each with sporangium imbedded in base.   Isoetaceae
+ Plants with creeping or ascending stems, sometimes scandent, erect, or plants epiphytic and pendent; leaves small, often overlapping, sporangia superficial, absent on lower leaves   (3)
3 (3) Sporangia 3-lobed, subtended by forked sporophyll; stems with widely spaced, alternate, scalelike leaves.   Psilotaceae
+ Sporangia not lobed, subtended by simple undivided leaves; stems with closely spaced, ± overlapping, subulate to ovate leaves   (4)
4 (4) Spores of two kinds: large megaspores and much smaller microspores; vegetative shoots often dorsiventral with leaves in 4 ranks, 2 median, with smaller leaves, and 2 lateral, less often uniform and spirally arranged.   Selaginellaceae
+ Spores of one kind, always very small; vegetative shoots usually with leaves uniform and spirally arranged, rarely lateral branches obviously flattened but then leaves not in 2 ranks, sometimes fertile leaves reduced.   Lycopodiaceae
5 (5) Sporangia borne on hexagonal peltate sporophores organized into very distinct terminal achlorophyllous strobili; aerial stems usually hollow, longitudinally ridged, articulate, with base of internode surrounded by tubular sheath, branches absent or whorled, rarely irregular at node.   Equisetaceae
+ Sporangia borne on fronds, these sometimes modified into sporocarps enclosing sporangia; stems never hollow [rarely with irregular ant-infested chambers outside Flora area], not articulate, branches when present never whorled   (6)
6 (5) Sporangia enclosed within sporocarps; small ferns of very wet situations, often floating in water, pinnae 1-25 mm   (7)
+ Sporangia borne on surface of frond or on specialized sporophore arising from frond; mostly ferns of well-drained situations, if growing in water then fronds much larger, pinnately (or palmately- pedately) divided   (8)
7 (7) Plants rooted in mud; fronds long stipitate, with 4 palmate pinnules [fewer or frond linear outside Flora area].   Marsileaceae
+ Plants free floating (sometimes stranded on mud); fronds sessile, usually floating on water surface.   Salviniaceae
8 (8) Fronds 3-dimensional, divided near base (or middle) into a fertile terminal "panicle" or "spike" and a sterile segment (simple, pinnatifid, or ternate), usually fleshy, vernation usually nodding; caudex subterranean, short, usually erect (horizontal in Helminthostachys); apex surrounded by a sheath.   Ophioglossaceae
+ Fronds and caudex not as above, vernation circinate, rarely hooked in some Pteridaceae; apex not sheathed   (9)
9 (9) Fronds vinelike with a twining rachis.   Lygodiaceae
+ Fronds not vinelike, sometimes scrambling but never with a twining rachis   (10)
10 (10) Fronds membranous, 1 cell thick, or rarely with 2-4 cell layers without intercellular spaces and stomata; sporangia borne on an extended veinlet (receptacle); indusia tubular or 2-lipped, borne on tips or upper margins of segments.   Hymenophyllaceae
+ Fronds herbaceous to leathery, several cells thick with intercellular spaces and stomata; sporangia not borne on extended veinlets   (11)
11 (10) Ferns treelike with an erect trunklike rhizome, with large compound fronds in a crown at apex   (12)
+ Ferns with rhizome short, creeping or climbing, sometimes massive but then prostrate or hardly longer than wide, never treelike   (16)
12 (12) Fertile pinnae without visible lamina; sporangia in dense clusters directly on rachis and rachillae, not covered by indusium or modified lamina.   Osmundaceae
+ Fertile pinnae with distinct lamina; sporangia in orbicular or linear sori or along veins and eventually ± acrostichoid   (13)
13 (12) Stem usually over 8 cm in diam.; sori orbicular or linear along veins and eventually ± acrostichoid   (14)
+ Stem usually less than 8 cm in diam.; sori linear   (15)
14 (14) Sori discrete, orbicular, medial, with cuplike or scalelike indusia or exindusiate.   Cyatheaceae
+ Sori linear along veins and eventually ± acrostichoid.   Blechnaceae (Brainea)
15 (15) Sori parallel to lateral veinlets or costules, indusia often double.   Athyriaceae (Diplazium)
+ Sori parallel to costa, indusia not double.   Blechnaceae (Diploblechnum)
16 (16) Fronds erect, linear or dichotomously divided into linear lobes with sporangia borne on terminal tufts of linear lobes ("sorophores"); sporangia flask-shaped with subapical annulus.   Schizaeaceae
+ Fronds various, if with linear lobes then sporangia borne on surface of lamina; sporangia not flask-shaped, annulus vertical or oblique   (17)
17 (17) Fronds pseudodichotomously branched, with a dormant bud in axils of regularly dichotomous forks; ultimate branches pinnate or bipinnatifid; sori orbicular, exindusiate, sporangia few.   Gleicheniaceae
+ Fronds simple, pinnate, palmate, or pedate, never with buds in axils of branch forks (ignore budlike bulbils along costa or rachis, not associated with branching)   (18)
18 (17) Fertile fronds or pinnae ?without visible lamina; sporangia in dense clusters directly on rachis and rachillae, not covered by indusium or modified lamina   (19)
+ All pinnae with lamina, sporangia borne on abaxial surface or at margin, sometimes pinnae very narrow with lamina inrolled to cover sporangia   (23)
19 (19) Fronds simple, fertile lamina ?reduced to single costa.   Polypodiaceae (Leptochilus)
+ Fronds compound   (20)
20 (20) Climbing, later epiphytic, ferns with long rhizomes and widely spaced fronds.   Dryopteridaceae (Lomagramma, Teratophyllum)
+ Terrestrial ferns with stout erect rhizomes and clustered fronds   (21)
21 (21) Stipe with enlarged base; rachis with cushionlike or long and hornlike aerophores at bases of pinnae.   Plagiogyriaceae
+ Stipe without enlarged base; rachis without aerophores   (22)
22 (22) Sporangia opening by an apical slit, annulus lateral; spores green.   Osmundaceae
+ Sporangia opening by a lateral tear, annulus vertical; spores not green.   Dryopteridaceae (Bolbitis)
23 (23) Fronds with brown leathery stipules at base of swollen stipe, and a pulvinus at base of each pinna; sporangia in synangia or ?free, without annulus.   Marattiaceae
+ Fronds with neither stipules nor pulvini (Plagiogyriaceae with swollen aerophores at base of pinna); sporangia not fused into synangia, with obvious annulus of thick-walled cells   (24)
24 (23) Sporangia acrostichoid, uniformly covering abaxial side of lamina   (25)
+ Sporangia distributed along veins, discrete sori or coenosori on abaxial side of lamina or along margin (not acrostichoid)   (34)
25 (24) Lamina with stellate hairs or scales on one or both surfaces   (26)
+ Lamina with simple hairs or glabrous   (27)
26 (26) Fronds lobed to deeply divided, sterile fronds sessile, base deeply asymmetrically auriculate, strongly adpressed to substrate and hiding rhizome, fertile fronds ?stipitate, dichotomously lobed, not adpressed.   Polypodiaceae (Platycerium)
+ Fronds entire or 1-pinnate, sterile and fertile fronds stipitate, or if sessile then base attenuate to cuneate, not adpressed to substrate, rhizome not hidden; fertile fronds or pinnae similar to sterile fronds but longer and narrower.   Dryopteridaceae (Subfam. Elaphoglossoideae)
27 (27) Sterile fronds without a distinct costa, lamina simple, bifid, or palmatifid with 3-5 main longitudinal veins.   Dipteridaceae (Cheiropleuria)
+ Sterile fronds or pinnae each with a distinct costa   (28)
28 (28) Stipe with enlarged base; rachis with cushionlike or long and hornlike aerophores at bases of pinnae; scales absent.   Plagiogyriaceae
+ Stipe without enlarged base; rachis without aerophores; scales present   (29)
29 (29) Rhizome scales clathrate; lamina simple; fronds remote   Polypodiaceae (Leptochilus)
+ Rhizome scales not clathrate; lamina pinnate to pinnatifid or pinnatilobed; fronds often clustered   (30)
30 (29) Plants epiphytic or epilithic, with creeping or climbing rhizomes; pinnae articulate at base   (31)
+ Plants terrestrial or epilithic in forests, with erect or creeping rhizomes; pinnae not articulate   (32)
31 (31) Veins free.   Lomariopsidaceae (Lomariopsis)
+ Veins anastomosing.   Dryopteridaceae (Lomagramma)
32 (32) Fronds clearly dimorphic (except Tectaria coadunata) Tectariaceae (Tectaria s.l.)   (32)
+ Fronds ± monomorphic, fertile pinnae only slightly smaller   (33)
33 (33) Pinna margin entire, crenate, or lobed, with or without teeth or spines, rachis usually with bulbils; growing in forests, often on rocks near streams, often over 100 m.   Dryopteridaceae (Bolbitis)
+ Pinna margin entire; bulbils absent; growing in coastal areas, often in mangrove forests, below 100 m.   Pteridaceae (Acrostichum)
34 (34) Aquatic ferns; fronds 2- or 3-pinnatifid; sori marginal, covered by reflexed lamina margin.   Pteridaceae (Ceratopteris)
+ Terrestrial, epiphytic, or epilithic ferns; fronds and sori often not as above   (35)
35 (34) Sori exindusiate, superficial, or sometimes sunken or borne in grooves, not covered by a reflexed lamina margin   (36)
+ Sori with a true indusia, or covered by ?modified reflexed lamina margin (false indusium)   (50)
36 (36) Lamina narrowly linear, grasslike, erect or pendent; sporangia in coenosori, borne in strictly marginal grooves, in 2 submarginal lines, or a single line along costa.   Pteridaceae (Subfam. Vittarioideae p.p.)
+ Lamina not grasslike, sori not as above   (37)
37 (36) Sporangia in indefinite sori, scattered along veins, not parallel to costa   (38)
+ Sporangia in definite sori, or coenosori   (42)
38 (38) Epiphytic ferns; fronds simple, elliptic or oblanceolate, with linear or clavate paraphyses interspersed with sporangia   Pteridaceae (Antrophyum)
+ Terrestrial ferns; fronds pinnatifid to pinnately compound   (39)
39 (39) Fronds dimorphic, fertile fronds with much-reduced lamina.   Tectariaceae (Tectaria s.l.)
+ Fronds ± monomorphic, fertile frond sometimes with lamina slightly reduced but not conspicuously so   (40)
40 (40) Fronds 2-4-pinnate, sometimes also simple or pinnatifid, abaxially glabrous, farinose, densely covered with a mass of brown hairs, or sparsely covered with scales; typically ferns of exposed and/or dry situations.   Pteridaceae (Subfam. Cheilanthoideae, Pityrogramma)
+ Fronds 1- or 2-pinnate or pinnatifid, or simple, abaxially hairy or glabrous; typically ferns of forest understory, often along streams   (41)
41 (41) Lamina rough, with dense hooked thick hairs on both surfaces; veinlets reticulate, areoles in 3 or 4 rows.   Thelypteridaceae (Dictyocline)
+ Lamina glabrous or with hairs on one or both surfaces; veins free or rarely anastomosing near midrib, then free.   Pteridaceae (Coniogramme)
42 (42) Fronds fan-shaped, deeply cleft into 2 halves, each half dichotomously divided into linear lobes; sori many, small, orbicular.   Dipteridaceae
+ Fronds not as above   (43)
43 (43) Fronds simple, pinnatifid, or 1-pinnate, rarely pedately lobed.   Polypodiaceae
+ Fronds bipinnatifid to decompound   (44)
44 (44) Rhizome, stipe, and lamina without scales or ordinary hairs; lamina delicate, with blunt, yellow, glandular hairs.   Dennstaedtiaceae (Monachosorum)
+ Rhizome, stipe, and/or lamina with scales and/or hairs; lamina hairs when present not all blunt, yellow, and glandular   (45)
45 (44) Fronds hairy or scaly, especially on abaxial side of costae   (46)
+ Fronds glabrous or with sparse hyaline or pale yellow glands   (48)
46 (46) Plants epiphytic or epilithic, often in moss; lamina 8-25 cm Polypodiaceae   (46)
+ Plants terrestrial; lamina (25-)50-100+ cm   (47)
47 (47) Fronds with scales.   Cyatheaceae
+ Fronds with hairs.   Thelypteridaceae
48 (48) Pinnae not articulate; fleshy hornlike processes borne in grooves at base of costae and costules.   Athyriaceae (Cornopteris)
+ Pinnae articulate to rachis; fleshy hornlike processes absent   (49)
49 (49) Plants terrestrial; lateral veins simple or occasionally forked, terminating at margin.   Cystopteridaceae (Gymnocarpium)
+ Plants epiphytic; veins in ultimate lobes simple, not reaching margin.   Polypodiaceae (Gymnogrammitis)
50 (50) Indusia 2-lipped, borne along margins of segments, near their bases; fronds large, tripinnate; rhizome and stipes covered with long golden brown hairs.   Cibotiaceae
+ Indusia not 2-lipped; rhizome and stipe base without long brown hairs   (51)
51 (50) Sori marginal or submarginal   (52)
+ Sori between costae and margin, occasionally also with a few borne near lamina margin   (58)
52 (52) Rhizome and stipe with unicellular or multicellular hairs or rarely bristles.   Dennstaedtiaceae
+ Rhizome and stipe scaly, at least at base, scales sometimes very narrow   (53)
53 (52) Sori protected by true indusia opening toward margin   (54)
+ Sori protected by revolute lamina margin   (56)
54 (54) Indusia orbicular-reniform; fronds pinnate with pinnae articulate to rachis.   Nephrolepidaceae
+ Indusia linear, oblong, cup-shaped, or tubular   (55)
55 (55) Stipe not articulate to rhizome; indusia linear or oblong; rhizome scales very narrow.   Lindsaeaceae
+ Stipe articulate to rhizome; indusia tubular, cuplike, or scalelike; rhizome scales broad.   Davalliaceae
56 (56) Fronds usually monomorphic, if dimorphic then pinna margin not inrolled to costa.   Pteridaceae
+ Fronds strongly dimorphic, pinna margin inrolled nearly to costa   (57)
57 (57) Fertile fronds green; sori orbicular or elliptic, confluent when mature; false indusium broad, continuous, covering abaxial surface making fertile segment appear podlike.   Pteridaceae (Cryptogramma)
+ Fertile fronds often becoming purplish brown; sori orbicular, with raised receptacles and indusiate, or confluent into linear coenosori.   Onocleaceae
58 (51) Sori oblong to linear, straight or curved   (59)
+ Sori orbicular or rarely ?elliptic   (63)
59 (59) Sori parallel to costae and/or costules.   Blechnaceae
+ Sori parallel to lateral veins, at angle to costa; stipe base with 2 vascular bundles   (60)
60 (60) Veins anastomosing to form 2-4 rows of areoles; indusia sometimes adhering at their margin and rupturing irregularly.   Diplaziopsidaceae
+ Veins usually free and not forming rows of areoles (anastomosing in some species of Asplenium); indusia not adhering at margin and not rupturing   (61)
61 (61) Acroscopic base of pinna and pinnule much larger than basiscopic base.   Rhachidosoraceae
+ Bases of pinna and pinnule equilateral or sometimes inequilateral, or lamina imparipinnate   (62)
62 (62) Scales dull, not finely clathrate; two vascular strands at base of stipe uniting in upper stipe to form a single U-shaped strand; indusia curved, J-shaped, or reniform and crossing a veinlet.   Athyriaceae
+ Basal stipe scales clathrate; two vascular strands at base of stipe uniting in upper stipe to form a single X-shaped strand; indusia straight.   Aspleniaceae
63 (63) Sori long stalked, one per ultimate segment, indusia dark brown to black.   Dryopteridaceae (Dryopteris sect. Peranema)
+ Sori sessile, often more than one per segment, indusia paler in color   (64)
64 (64) Indusia completely surrounding receptacle and composed of filaments or scalelike segments forming a cup around sorus or membranous and completely enclosing sorus; costae abaxially without scales.   Woodsiaceae
+ Indusia attached centrally or laterally, not completely surrounding receptacle; costae abaxially with or without scales   (65)
65 (64) Fronds with stipes articulate to phyllopodia, or fronds 1-pinnate with pinnae articulate to rachis   (66)
+ Fronds with stipes and pinnae not articulate   (70)
66 (66) Fronds simple.   Oleandraceae
+ Fronds pinnate   (67)
67 (66) Individual pinnae articulate   (68)
+ Frond articulate at base of stipe, pinnae not articulate   (69)
68 (68) Rhizome without stolons; sori in several rows between midrib and margin.   Lomariopsidaceae (Cyclopeltis)
+ Rhizome forming stolons; sori in a single row between midrib and margin.   Nephrolepidaceae
69 (69) Fronds 3- or 4-pinnate; phyllopodia short and indistinct.   Hypodematiaceae (Leucostegia)
+ Fronds 1-pinnate; phyllopodia long and stipelike.   Tectariaceae (Arthropteris)
70 (65) Rachis with an adaxial groove confluent with grooves of rachillae   (71)
+ Rachis without an adaxial groove, or if grooved then groove not confluent with grooves of rachillae   (73)
71 (71) Base of stipe with several vascular bundles.   Dryopteridaceae
+ Base of stipe with 2 vascular bundles   (72)
72 (72) Veins free, reaching segment margin; indusia when present basal, a minute hoodlike scale, arching over sorus, frequently deciduous.   Cystopteridaceae
+ Veins anastomosing or free, usually ending before segment margin; indusia lateral, vaulted or essentially flat, opening along lateral margin, usually persistent.   Athyriaceae (Anisocampium)
73 (70) Veins anastomosing   (74)
+ Veins free   (75)
74 (74) Indusium reniform.   Tectariaceae
+ Indusium peltate.   Dryopteridaceae (Cyrtomium)
75 (75) Fronds 3- or 4-pinnate.   Hypodematiaceae
+ Fronds 1- or 2-pinnate   (76)
76 (76) Stipe, rachis, costae, and veins with multicellular scalelike or moniliform hairs and/or scales, rarely glabrous and then lamina simple or pinnatilobate.   Athyriaceae (Deparia)
+ Costae glabrous or sometimes with sparse short terete hairs adaxially; lamina 2-pinnatifid.   Tectariaceae (Pteridrys)

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