IAWA Bulletin n.s., Vol. 4 (1),1983 53

WOOD ANATOMY OF LEANDRII H. HUMB. (PEDALIACEAE) AND ITS RELATION TO

by

R.W. den Outer and W.L.H. van Veenendaal Department of Cytology and Morphology, Agricultural University, Arboretumlaan 4, 6703 BD Wageningen, The Netherlands

Summary The secondary xylem of Uncarina leandrii H. tology and Morphology in the botanical gar­ Humb., endemic to Madagascar, is described. dens or greenhouses of the Agricultural Univer­ Comparisons have been made with woody re­ sity, Wageningen. - ST (Stahel) numbers col­ presentatives of c10sely related families, viz. lected by Stahel in Surinam (1944; herbarium Bignoniaceae, Scrophulariaceae and Acantha­ vouchers present in Uw, Utrecht). - V&O ceae. Only affinities with the Bignoniaceae are (Versteegh and Den Outer) numbers 1-749 apparent. collected by Versteegh and Den Outer in Ivory Coast (1969), 850-987 by Den Outer in Suri­ Introduction nam (1974), 988-1227 by Van Veenendaal The Pedaliaceae constitute a small family of and Den Outer in Madagascar (1978); herba­ herbs and shrubs or small trees, confined to the rium vouchers present in Lw (Leiden), Uw arid zones in Africa, Madagascar, Indomalaysia (Utrecht) and the Department of Plant Taxon­ and Australia. The family consists of 12 genera, omy and Geography (Wageningen). - WS (Wel­ with 60 (Humber, 1971) to 70 (Ihlenfeldt, vaartsfonds) number collected in Surinam (her­ 1967) species. The woody Uncarina (BaiI­ barium vouchers present in Uw). - ZW&R lon) Stapf of the Pedaliaceae is endemie in Ma­ (Zwart and Rood) numbers collected by the dagascar. As far as we know, its secondary Forest Research Institute, Bogor, Indonesia xylem has not been described before. Other (1925). - from WIBw (Department of Forest­ very brief wood anatomical descriptions of ry Techniques, Wageningen). - BW (Boswezen) some Pedaliaceae species are given by Metcalfe numbers collected by the former Dutch Forest­ and Chalk (1950) and for alatum by ry Service in West New Guinea, Indonesia, up Pawar and Kulkarni (1971). to 1962 (herbarium vouchers in Lw, Leiden). - CTFT (Centre Technique Forestier Tropical) Materials and Methods numbers collected by the Service Forestier in The collected Uncarina leandrii is a xero­ Madagasear (records are kept of the loeation of philous shrub of 2 m high, with a stern diame­ herbarium vouchers). ter of 6 cm. It was found in a vestigial forest, Transverse, radial and tangen tial sections of 20 km north-east of Sakaraha in south-wesJ the wood sampies were made with a sledge mi­ Madagascar. Stern sam pies were immediately erotome, varying in thiekness from 10-20 j..lm. fixed in FAA. They are stored at the Depart­ All sections were embedded in Kaiser's gelatin­ ment ofPlant Cytology and Morphology, the ac­ glycerin (1ohansen, 1940). Means and ranges of companying herbarium vouchers (V&O 1022) the number of wood rays per mm in tangential at the Department of Plant and direction, ray height and width, length of pa­ Plant Geography, both at Wageningen, the renchyma strands, radial vessel diameter and Netherlands. Other wood sampies from shrubs vessel member length are based on at least or trees were obtained from two institutional twenty-five measurements. The vessel member wood collections referred to according to Stern length was measured inc1uding the tails. Vessel (1978). If the presence of herbarium vouchers frequency was determined over an area of at is known, this is indieated. least 20 square mm, where possible these areas The following number indications are used: were not taken near the pith. For all quantita­ from WLw (Department of Plant Cytology and tive da ta mean values are given, preceded and Morphology, Wageningen, The Netherlands). - followed by extreme values between brackets. A (Arnoldo) numbers collected by Arnoldo­ We have used the definition of libriform fibres Broeders and De Jong in the Netherlands Antil­ given by Reinders (1935) and Janssonius (1940). Ies (1966). - PL (Plantkunde) numbers collect­ Wood ray types given in Table I are c1assified ed by the staff of the Department of Plant Cy- aecording to a modified system of Kribs (1959)

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given in the legends. A heterogeneous wood ray Discussion means a wood ray composed of procumbent Takhtajan (1969, 1980) placed the Pedalia­ cells together with upright and/or square cells; ceae in the Scrophulariales ne ar to the Scroph­ a homogeneous wood ray is a wood ray com­ ulariaceae and Bignoniaceae. These three fami­ posed of either entirely procumbent ceUs or lies are supposed to be closely rela ted, especial­ entirely upright and/or square cells. Iy the Scrophulariaceae to the Bignoniaceae. Also the Acanthaceae is placed in the same or­ der, even in the same suborder, viz. Scrophula­ Resuits riineae. Hutchinson (1973) emphasised only The secondary xylem of the investigated the connection of the Pedaliaceae with the Big­ Uncarina leandrii H. Humb. shows the follow­ noniaceae (both placed in the Bignoniales). In ing characters (see also Table land Figs 1-4). his opinion relationship between the Acantha­ Growth rings fairly distinct (marked by termi­ ceae and Scrophulariaceae (both placed in the nal parenchyma), wood diffuse-porous. Personales) on the one hand and Pedaliaceae Vessels solitary, in short radial multiples and and Bignoniaceae on the other is remote. in clusters; round to oval in cross section; aver­ Also Ihlenfeldt (1967) concluded that the age number 8 per mm2; radial diameter (40-) Pedaliaceae are closely related to the Bignonia­ 120(-160) JJ.m, thin-walled (2-3 JJ.m). Vessel ceae. The Scrophulariaceae and Acanthaceae member length (250-) 320(-400) JJ.m. Perfora­ are related to the Bignoniaceae, but only to a tions simple in oblique end walls. Inter-vessel lesser extent to the Pedaliaceae. pits alternate, 7-8 JJ.m, sometimes with coales­ Some general wood anatomical descriptions cent apertures. Vessel-ray and vessel-paren­ of the families Bignoniaceae and Acanthaceae chyma pits half-bordered to almost simple, 8- as given by Metcalfe and Chalk (1950) indica­ 25 JJ.m. Vessels usually in contact with axial ting a closer relationship of Uncarina leandrii parenchyma. Fibres libriform, thin-walled (2-3 with the Bignoniaceae than with the Acantha­ JJ.ill), non-septate, with mainly small bordered ceae, comprise the following. In the Bignonia­ pits with slit-like inner apertures, confined to ceae the vessels are medium-sized (100-200 the radial walls (often in two rows at the ends JJ.m), sometimes sm aller (50-100 JJ.m), solitary of the fibres); length (460-)590(-790) JJ.m. and in small multiples of 2 or 3 cells, sometimes Parenchyma moderately abundant, paratracheal in irregular clusters of small vessels, e.g. Catalpa; (a very small, not always complete sheath mostly 5-20 per square mm; sometimes ring­ around the vessels), apotracheal (2-3 cells porous in Bignonia, Catalpa, Chi/opsis and Fer­ wide tangential bands, about 3 per radial mm dinandoa, and spiral thickenings present in small including the terminal parenchyma) and irregu­ vessels ofCampsis, Catalpa andChilopsis. Paren­ lar diffuse. Strands of 3 cells, about 350 JJ.m chyma paratracheal, varying from narrowly va­ long. Rays uni- and multiseriate, composed of sicentric to abundant aliform and confluent upright and square cells, sometimes with a few types and sometimes very abundant. Rays typi­ procumbent cells; tails of multiseriate rays cally homogeneous composed entirely or main­ short, sometimes more than 4 marginal rows Iy of procumbent cells, but slightly heterogene­ present; average 2-3-seriate, uniseriate rays ous in some species; 4-11 per tangential mm. with or without small bi-seriate portions often Fibres only septate in climbers and a few other present; height (200-)550(-1100) JJ.ill; (5-)7 species; pits simple or with very small borders. (-9) per tangential mm. Crystals always pres­ In the Acanthaceae the vessels are typically ent in ray cells, rhomboidal or small styloids. very small (less than 50 JJ.ill), sometimes ex­ The results of the investigated species be­ tremely small (less than 25 JJ.m), e.g. in Whit­ longing to the Bignoniaceae, Acanthaceae and fieldia colorata; some multiples of 4 or more Scrophulariaceae are presented in Table I. cells and a tendency for these and the solitary Average values of the family as a wh oie are also vessels to be grouped in radial rows in Belo­ given. perone, Isoglossa, Pseuderanthemum, Aphelan-

Fig. 1-4. Secondary xylem of Uncarina leandrii H. Humb. (Pedaliaceae). - I: Transverse section showing long tangential parenchyma bands of different width within a ground tissue of libriform fibres. - 2: Transverse section, enlargement of apart of Fig. 1. In the middle a 2-seriate ray. Note the bi-bordered pit-pairs in the radial walls of the libriform fibres. - 3: Radial section showing a ray composed ofsquare and upright cells. Note vessels \\ ith simple perforation plates and many bordered pits in the ends of the libriform fibres. - 4: Tangential section showing uniseriate and multiseriate rays with usually short tails. Note the parenchyma strands at the right hand side.

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Tablc I. Secondary xylem characters of the investigated species.

spccimcns number diam. at growth vessels breast rings height rd. diam. in pm av. nb. perf. inter· average in cm per vessel member mm2 pils length in pm in pm

Pedaliaceae Unearina leandrii H. Humb. V&O 1022 + (40-) 120(-160) 8 8 320 Acanthaceae Aphelandra deppeana SchI. et Cham. V&O 970 6 ± (15-) 30(- 50) 150 3-4 390 BarIeria her R. Benuist V&O 1056 5 + (15-) 30(- 60) 170 390 Crossandra poissunii R. Benoist V&O 1101 3 (25-) 85(-130) 16 7-8 145 Forsythiopsis vincoides R. Benoist V&O 1102 3 + (10-) 30(- 50) 350 3-4 270 Forsythiopsis vincoides R. Benoist V&O 1153 3 + (15-) 25(- 55) 150 3-4 300 cf. Hypocstes V&O 1041 2 + (10-) 20(- 30) 360 3 310 Strobilanthes spec. V&O 1023 4 + (10-) 30(- 50) 270 3-4 260 Trichanthera gigantea Steud. ST 175a >10 ± (60-)120(-170) 4 6 690 Average value [ar Ihe [amily (20-) 45(- 75) 180 4-5 345 Bignoniaceae Catalpa bignonioides Wal!. PL 2823 >10 + (30-)110(-240) 20 7 210 Deplanchea tetraphylla F.v.M. BW 11612 >10 (40-)130(-220) 3-4 7 590 Dolichandrone spathacea K. Schum. BW 3247 >10 + (30-) 90( -170) 7 s; ±rt 5 360 Jacaranda copaia (AubI.) D. Don WS 1061 >10 (40-)240(-360) 3 8 540 Jaearanda rhombifolia G.F.W. Meyer V&O 903 10 + (30-) 60(- 95) 14 6 320 Kigelia africana (Larn.) Benth. V&O 601 10 ± (30-) 80(-130) 6 4-5 250 Markhamia cf. ubtusifolia Sprague V&O 321 + (30-) 120( -220) 10 s;±rt 3-4 190 Newbouldia laevis Seem. ex Bureau V&O 545 6 + (30-) 75(-120) 12 5 230 Phyllarthron ilicifolium H. Perr. V&O 1154 4 + (30-) 70(-100) 37 3-4 260 Phyllarthron i1icifolium H. Perr. V&O 1139 6 + (30-) 60(-100) 19 3 260 Radernlachera gigantea (BI.) Miq. ZW&R36 >10 + (50-)140(-220) 12 4-5 340 Radermachera glandulosa (BI.) Miq. ZW&R 142 >10 + (30-) 95(-160) 27 s;±rt 3-4 270 Rhigozum madagascariense Drake V&O 1099 8 + (15-) 40(- 60) 200 s;±rt 3-4 160 Spathudes campanulata P. Beauv. V&0650 >10 + (40-) 150(-240) 6 s; ±rt 4-5 160 Stereospermum acuminatissimum K.Sch. V&O 587 10 + (30-) 105( -170) 4 4-5 290 Stereospermum arcuatum H. Perr. CTFT 137 >10 + (40-)130(-320) 6 5 260 Stereuspermum euphorioides De. V&O 1005 >10 + (40-) 120( -180) 40 3 220 Tabebuia capitata Sandw. ST 114 >10 + (30-)120(-160) 16 9 260 Tabebuia ehrysantha Nichols. A3541 >10 + (40-) 60(-100) 44 12 220 Tabebuia insignis Sandw. ST 174 >10 + (40-) 130( -210) 4 7 380 Tabebuia serratifolia Nichols ST 101 >10 + (40-) 80(-130) 9 II 325 Tecoma stans (L.) H.B.K. V&0391 3 + (30-) 55(-100) 48 2-3 260 Average value [ar the [amily (35-)100(-170) 25 5-6 290 Scrophulariaceae Paulownia tomentosa (Thunb.) Steud. PL 2478 >10 + (25-) I OO( -175) 12 8 210

Symbols and abbreviations used: +: present; -: absent; ± scarcely present; av.: average; diam.: diameter; ob.: number; rd.: radial; rt: reliculate perforation; s: simple perforation.

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(Table 1 continued, see also next page) fibres rays axial parenchyma

type av. height av. width type pred. nb. per rare or diff. nb. lung tg. para· av. length inJ..ll11 in cells cell tg.mm absent mult. bands tracheal uf strands (maximum) type per rd. mm type in J..II11 (nh. (rd. width of cells) in cells)

In 550(-1100) 2-3 Ho (He 11) s; u 7 ± 2-3 C± 3) ±v 350 (3)

In 310(- 750) 2-3 He 11 13 + In 420(-1600) I (2) Ho III s; u 14 + In;±ls >10000 12 He 11 1-2 5 (2-5) ISO (3) Is 470( - 920) 2 He/Ho 11 s; u 17 .±v 240 (3) Is 310(- 900) He 11 p 13 + In 130(- 850) 1-2 He 11 II + Is 440(- 940) 2 He I/li s; u 18 + Is 1200(-3900) 3 He 11 560 (3) 1660( -2480) 3-4 12 310 (3)

In 175( - 440) 2 He 11 p 4 + 260 (4) In 51O( - 870) 2 He 11 p 6 + 2 (2-5) 740 (4) In 160(- 360) I (2) Ho III P 9 + 1-2(2-3) v;±a 450(4) In 380(-1600) 2-3 Ho 11 P + I (2-3) v; a;±c 770 (6) In 200(- 650) I (2) Ho Il P 13 + 4 (2-3) v; a 390(5) In 190(- 440) 2 Ho [] P 4 ± 1(2-5) v; a; c 300 (2) In 240(- 640) 3-4 He [] p 6 ± 3-4 (3-8) v;±a 285 (4) In;±ls 240(- 480) 3 He 11 p + 0-1 (3-5) v; a; c 280 (3) In 170( - 320) I (2) Ho III P v; ±a; .tc 260 (2) In 190(- 360) I (2) Ho III P v; ±a; ±c 330 (3) In 240(- 450) 2-3 Ho 11 p ± 1 (5) v; a; c 450 (4) In 260(- 400) 2- 3 Ho II P 4 ± 1(3-5) v; ±a; ±c 340 (3) In 200(- 480) I (2) Ho [] P 12 ± v:±a 160 (2) In 240(- 480) 3-4 He 11 p 6 3 (2-30) v; a;±c 240 (2) In;±tr 220(- 440) 3 Ho [] P 4 2(3-7) v; a; c 320 (2) In 300(- 720) 3-4 Ho [] P 4 2 (3-8) v; a; c 290 (4) In 230(- 480) 3-4 Ho 11 P 5 3 (8-20) v; a; c 215 (3) In 180(- 210) 2 Ho [] P 9 ± v:±a 290 (3) In 140(- 160) 1-2 Ho III P + 3 (2-3) v; a;±c 190 (2) In 280(- 560) 1-2 Ho III P 7 ± v; ta 480 (4) In 170(- 200) 2 Ho [] P 9 .± v: a; c 300 (2) Is 250(-1250) 2 He 1/11 11 ±v 280 (3) 235(- 545) 2 6-7 350 (3)

In 130(- 670) 2-3 Ho 11 P 4 I (5-20) v: a; c 280 (3)

Symbols and abbreviations used: +: present; -: absent; ±: scarcely present; I: uniseriate rays and multiseriate rays with lang uniseriate taHs; 11: uniseriate rays and multiseriate rays with short uniseriate taHs; IlI: only uniseriate rays present; a: aliform; aggr.: aggregates; av.: average; c: cont1uent; diff.: diffuse; He: heterogeneaus wuod rays; Ho: homogeneous wood rays (see modified definition under Materials and Methods); In: libriform fibres, non·septate; Is: libriform fibres, septate; muh.: multiseriate; nb.: number; p: procumbent wood.ray cell; pred.: predominant; rd.: radial; s: square wood· ray cell; tg.: tangential; tr: fibre·tracheids; u: upright wood·ray cell; v: vasicentric.

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(Table I continued)

specimens crystals further information (if present in rays)

Pedaliaceae Uncarina leandrii terminal parenchyma 2-4·seriate Acanthaceae Aphelandra deppeana In ± thick·walled Barleria her included phloem Crossandra poissonii average width rays 250 J.llTl Forsythiopsis vincoides s;ist cL Hypoestes terminal parenchyma uniseriate Strobilanthes spec. Trichanthera gigantea Is wilh simple pits in tangential walls Bignoniaceae Catalpa bignonioides ring-porous; tyloses; spiral thickenings Deplanchea tetraphylla Dolichandrone spathacea terminal parenchyma 1-3-seriate Jacaranda copaia In with bordered pits in radial walls: tyloses Jacaranda rhombifolia terminal parenchyma 2-4-seriate Kigelia af(icana ground tissue 30% parenchyma Markhamia cL obtusifolia ground tissue 30% parenchyma; semi ring-porous Newbouldia laevis terminal parenchyma uniseriate Phyllarthron ilicifolium In thick·walled Radermachera gigantea Radermachera glandulosa st terminal parenchyma 1-3·seriate Rhigozum madagascariense terminal parenchyma 1-4-seriate; ± storied structure Spathodes campanulata ground tissue ± 30% parenchyma; tyloses Stereospermum acuminatissimum ± terminal parenchym.; spiral thickenings Stereospermum arcuatum terminal parenchyma l-3-seriate; In thick-walled Stereospermum euphorioides ground tissue 90% parenchyma; semi ring-porous Tabebuia capitata In thick-walled; storied structure Tabebuia chrysantha In thick-walled; storied structure Tabebuia insignis In with bordered pils in radial walls; terminal parenchyma uniseriate Tabebuia serratifolia In thick-walled; storied structure Tecoma stans often > 3 stores in rays Scrophulariaceae Paulownia tomentosa ring-porous

Symbols and abbreviations used: In: Iibriform fibres, non·septate; Is: Iibriform fibres, septate; r: raphide crystal; s: simple crystal; st: styloid crystal.

dra andPachystachys; often over 100 per square eloser relationship of Uncarina leandrii with mm. Parenchyma scarce, paratracheal limited the Bignoniaeeae than with the Acanthaeeae. to narrow sheaths or a few cells around the ves­ This is not only obvious by vessel charaeteris­ sels, oeeasionally diffuse or terminal in Anisa­ ties like the number per square mm, diameter canthus. Rays often high, markedly heterogen­ and arrangement, especially if Rhigozum mada­ eous and often eomposed of square and upright gascariense is exeluded, but also by type and cens with a few procumbent eens, sheath eens number of rays per tangential mm, the amount sometimes present. Fibres septate exeept in and type ofaxial parenehyma and type and pit­ Aphelandra, Pachystachys andPseudoblepharis; ting of the libriform fibres. With the Aeantha­ pits smalI, simple. ceae, Uncarina leandrii only has in common the Dur data, as presented in Table I, like the predominanee of square and upright ray eells. general deseription given above, also reveal a Within the Aeanthaeeae, Crossandra poissonii

Downloaded from Brill.com09/30/2021 01:43:54PM via free access IAWA Bulletin n.s., Vol. 4 (1),1983 59 and Trichanthera gigantea occupy a quite ex­ Acknowledgements ceptional position. Not only according to the We sincerely thank Mrs. M. H. van den Bergh number of vessels per square mm and the radial for preparing the slides used for this study. vessel diameter, but also concerning ray height (especially Crossandra poissonii), number of rays per tangential mm and amount ofaxial pa­ References renchyma. Wood anatomically Crossandra pois­ Campbell, D.H. 1930. The relationships of Paul­ sonii and Trichanthera gigantea possibly occu­ ownia. Bull. Torrey Bot. Club 57: 47-50. py an intermediate position between the Acan­ Humbert, H. 1971. Pedaliaeees. In: H. Humbert, thaceae and Uncarina leandrii. Flore de Madagasear et des Cornores, Farn. Uncarina leandrii exhibits more affinities 179: 1-46. Paris. with Paulownia tomentosa, a member of the Hutehinson, J. 1973. The families of flowering sole tree genus of the Scrophulariaceae, than . 3rd Ed. Clarendon Press, Oxford: with the Acanthaceae (including Crossandra 482-483. poissonii and Trichanthera gigantea). The posi­ Ihlenfeldt, H.-D. 1967. Über die Abgrenzung tion of Paulownia tomentosa within the Scro­ und die natürliche Gliederung der Pedalia­ phulariaceae is doubtful; it should be assigned ceae R. Br. Mitt. Staatsinst. Allg. Bot. to the Bignoniaceae according to many authors Hamburg 12: 43-128. like Campbell (1930), Westfall (1949). Wood Janssonius, H.H. 1940. Anatomische Bestim­ anatomically Uncarina leandrii has not much in mungstabelle für die javanischen Hölzer. common with other Scrophulariaceae species Brill, Leiden. described by e.g. Metcalfe and Chalk (1950). Johansen, D.A. 1940. Plant mierotechnique. Sterns of herbs and young shrubs show a con­ McGraw-Hill Book Co. Ine., New York, siderable range of structure when viewed in London. transverse seetion. This is mainly caused by va­ Kribs, D.A. 1959. Commercial foreign woods riations in the size and distribution of the ves­ on the American market. Edwards Broth., sels. Vessels are typically sm all (less than 100 Michigan. J.tm) and sometimes extremely small (less than Metealfe, C.R. & L.Chalk.1950. Anatomy ofthe 25 J.tm, e.g. in some species of Anastrabe, Cal­ Dieotyledons II. Clarendon Press, Oxford. ceolaria, Monttea, Penstemon and Veronica) Scrophulariaeeae: 978-988; Bignoniaceae: like in the Acanthaceae (see also Table I). Pa­ 1002-1013; Pedaliaeeae: 1013-1014; renchyma is usually very sparse or absent. Rays Acanthaceae: 10 14-1 023. when present are usually heterogeneous, com­ Pawar, J. S. & A. R. Kulkarni. 1971. Contribu­ posed of mixed procumbent and square to up­ tion to the morphology of Pedaliaceae. right cells; typical medullary rays are absent Part 6. Wood anatomy of Sesamum ala­ from herbaceous and from many woody spe­ turn. J. Shivaji Univ. 4: 117-120. eies. In this description for the family as a Reinders, E. 1935. Fibre-tracheids, libriform whole, Paulownia is always an exception. Wood wood fibres and systematics in wood anat­ anatomically Paulownia resembles much more omy. Trop. Woods 44: 30-36. the investigated Bignoniaceae species and Unca­ Stern, W.L. 1978. Index Xylariorum. Institu­ rina leandrii (Table I). tional Wood Collections of the World. 2. So, if one considers the wood anatomy of Un­ Taxon 27: 233-269. carina leandrii only, the position of the woody Takh tajan, A. 1969. Flowering plants, origin Pedaliaceae nearthe Bignoniaceae (nearly always and dispersal. Oliver & Boyd, Edinburgh. trees or shrubs) seems justified, as stated for -- 1980.0utline of the classification of flow­ the families as a whole by Takhtajan (1969, ering plants. The Botanical Review 46 (3): 1980), Hutchinson (1973) and Ihlenfeldt (1967). 225-359. A relation to the woody Acanthaceae is remote, Westfall, J.J. 1949. Cytological and embryolog­ whieh has also been mentioned by for instance ical evidences for the reclassification of Hutehinson (1973) and Ihlenfeldt (1967). Paulownia. Amer. J. Bot. 36: 80S.

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