IAWA Bulletin n.s., Vol. I (1-2), 1980 59

VESTURED PITS IN THE NEW WORLD PITHECELLOBIUM (SENSU LATO)

by

Daniel L. Cassens* Agricultural Experiment Station, Purdue University, West Lafayette, Indiana 47907, U. S. A.

Summary Vestured pits in Pitheeellobium sensu lato Bailey's types of vestures, Cote and Day (1962), were observed with the SEM. In both the pit as well as Schmid (1965), reported that only chamber and on the pit aperture distinct types two types can be easily recognized. These are of vestures can be recognized. The quantity of branched and unbranched. Scurfield, et al. vestures in both locations also varied. However, (1970) have suggested classifying vestured pits due to inter- and intra-species variation, the into four groups - filamentous, bead-like, mas­ vesture types cannot be used to differentiate sive coralloid, and massive foliate. These au­ the many generic segregates of Pithecellobium. thors also noted that intra- and inter-species variation in the structure of vestures is such as Introduction to make any classification system of dubious Vestured pits are minute outgrowths from value. Other authors (Ishida & Ohtani, 1970; the free surface of the secondary wall into the Butterfield & Meylan, 1974; Meylan & Butter­ pit cavity of bordered and half bordered pit field, 1974; and Miller, 1977) report that dif­ pairs. The presence or absence of vestured pits ferent types of vesture morphology can be in various woody genera has been reported by identified, but due to excessive variation even numerous investigators. Record (1925) review­ within the same specimen a reliable classifica­ ed the early work. Bailey (1933) was the first tion system cannot be developed. More recent­ to do an extensive investigation of vestured pits ly, however, Van Vliet (1975) has indicated to determine their diagnostic and phylogenetic that three types of vestures within intervascular value. In his examination of 152 families, Bailey pits of the Crypteroniaceae are consistent. The observed vestured pits in only 24 families. Only three types of vestures, unlike other systems four of these did not have vestures throughout proposed, are based on the extent of vesturing every genus and in only Oleaceae did the occur­ on the vessel wall, in the pit chamber and on rence of vestured pits not follow any taxonom­ the pit aperture. Van Vliet (1978) also report­ ic subdivision. Thus, the diagnostic and phylo­ ed that two main types of vestures could be ob­ genetic significance of the occurrence of ves­ served in Combretaceae and that these types tured pits was established. Record (1936) and are in agreement with a subfamily classification Metcalfe and Chalk (1950) continued to inves­ system. He concludes, however, that each tigate vestured pits but significant new findings taxon must be examined separately because of were not reported. frequent intermediiltes and because of intra­ Bailey (1933) also noted variation in vesture generic variation. morphology. He alluded to vestures that ap­ An investigation of 87 specimens in the New peared as 'massive papillae', 'distinctly branch­ World Pithecellobium complex (Leguminosae­ ed or coralloid' and 'loose mats of branching Mimosoideae) was undertaken to characterize and anastomosing filaments'. The vestures may the types of vestures present. These different be attached to both inner and outer apertures vesture types might be used as an aid in the and spread into the lumens of the cells. Using classification of this difficult complex. Some the electron microscope, Cote and Day (1962) taxonomists prefer to recognize Pithecellobium were the first to substantiate different mor­ s.l. while others have proposed numerous segre­ phological categories of vestured pits. Because gate genera. These include Abarema, Arthrosa­ of considerable variation and overlapping in manea, Chloroleucon, Cojoba, Ebenopsis,

* The research for this paper was conducted at the U.S. Forest Products Laboratory, Madison, Wis­ consin. It is submitted as Journal Paper Number 7713, Agricultural Experiment Station, Purdue University, West Lafayette, Indiana 47907.

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Fig. I . Pseudosamanea guachapele (SJRw 27084). Cross section with vessel lumina (VL), pit canals (PC) and vestures (V). x 1530. -- Fig. 2. Zygia /ati/olia (SJRw 42739). Vessel lumen with few minute vestures (arrows), pit membrane (PM) and vestures in the pit chamber. x 1490. -- Fig. 3. Arthrosamanea pistaciae/olia (SJRw 1551). Filamentous vestured pits. x 4630. -- Fig. 4. Pithecel­ lobium (Marmaroxylon?) dinizii (SJRw 43205). Coralloid vestured pits. x 4170. -- Fig. 5. Pithe­ cellobium dulce (SJRw 16442). Bead-like to coralloid vestured pits showing pit rim (r). x 3870. -­ Fig. 6. Zygia longi/olia (SJRw 22402). Coralloid vestured pits. x 5400.

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Havardia, Jupunba, Klugiodendron, Macrosa­ Pithecellobium collinum*: FDBG 3976 (type) rnanea, Marmaroxylon, Painteria, Pseudosama­ (SJRw 46465) - P. dinizii*: J. Cuatrecasas nea, Punjuba, Samanea, and Zygia. Cassens 17374 (SJRw 43205); B.A. Krukoff 1302 (1973) and Cassens and MilJer (publication in (MADw 25118) - Marmaroxylun racemosum: process) have reviewed the taxonomic history B.E. Dahlgren (SJRw 16780); A. Ducke 39** of Pithecellobium s.l. and investigated the wood (SJRw 20720); Capucho 274 (SJRw 21260); anatomy. The anatomical characteristics can be Capucho 377 (SJRw 22055); FDBG 3114 used to form groups that are about the same (SJRw 46469); G. Stahel 72 (SJRw 41145); as the segregate genera proposed by various Unknown (MADw 21013) - Pithecellobium taxonomists. umbriflorum*: B.A. Krukoff 6872 (SJRw 36937) - P. dulce: S.1. Record 43 (SJRw Materials and Methods 16442); Espina & Giacometto B8A (SJRw Samples representing 12 of the segregate 20990); A. Dugand 223; 60 (SJRw 22512); genera from Pithecellobium s.l. were taken A. Dugand 574 (SJRw 27104) - P. guadalu­ from specimens in the Madison (MADw) and pense: Stern & Brizicky 358 (SJRw 51174); Samuel J. Record (SJRw) wood collections; 87 Stern & Brizicky 362 (SJRw 51178); Stern & specimens were examined with the scanning Chambers 252 (SJRw 51464) - P. oblongum: electron microscope (SEM). Small samples S.J. Record & H. Kuylen G.120 (SJRw 10071); about 5 mm square, exposing a tangential sur­ W. Stern & K.L. Chambers 27 (SJRw 51544); face, were split from the original specimen, A. Dugand 847 (SJRw 29678) - P. unguis-cati: mounted on stubs and coated with gold for A.F. Wilson F-5 (MADw 15950); D.H. Caldwell examination. The splitting technique did less (SJRw 49290) - Pseudosamanea guachapele: damage to the structure than cutting with a Humberto Tasayco Tasayco 3 (MADw 22546); microtome knife or razor blade. During the H. Pittier 12184 (SJRw 9520); S.1. Record & examination of each specimen, no attempt was H. Kuylen G.126 (SJRw 10077); A. Dugand made to differentiate between inter-vessel, ves­ 540 (SJRw 27084); M. Acosta-Solis 11716 sel-ray, and vessel-parenchyma pits. (SJRw 45429); Navy Project 79 (SJRw 45518) - Punjuba racemiflora: Wm.R. Barbour 1014 (MADw 10284) - Samanea saman: Whitford & List of Pithecellobium (sensu lato) specimens Stadtmiller (MADw 10851); GilJ & Whitford examined (xylarium number in brackets): Ar­ 90 (SJRw 9101); L1. Williams 5495 (SJRw throsamanea pistaciaefolia: L1. Williams 12823 18758); Espina & Giacometto A202 (SJRw (MADw 25493); H.M. Curran 43 (SJRw 1551); 20977); Capucho 479 (SJRw 22770); M. Turner H.M. Curran 44 (SJRw 1552); H.M. Curran III (SJRw 45751; Felix Woytkowski 35145 300 (SJRw 1528); A. Dugand 138;483 (SJRw (SJRw 44592) - Zygia ampla: L1. Williams 23915); A. Dugand296 (SJRw 28504) - Chlo­ 14747 (SJRw 41738) - Z. cauliflora: B. Ma­ roleucon mangense: Navy Project 454 (SJRw guire 5573 (MADw 22954); A.C. Persaud 106 45679); Stern & Chambers 158 (SJRw 51650) (SJRw 9496); G. Stahel284 (SJRw 42513); L1. - Cojoba arborea: Britton & Kramer XV Williams 11424 (MADw 25120) - Z. cognata: (SJRw 3088); M.O. Hope II (SJRw 4797) - Ll. Williams 9208 (SJRw 34792) - Z. conzattii: Ebenopsis flexicaulis: Harold Nogle 99 (MADw L1. Williams 9438 (SJRw 34857) - Z. inaequa­ 13215); A.F. Wilson M-I (MADw 18344); lis: Espira & Giacometto An (SJRw 20847); MEXFw X-324 (MADw 25290); C.D. Gilbert A. Ducke 384 (SJRw 40412); Wurdack & Ad­ (SJRw 40706) - Havardia leiocalyx: V.M. Page derley 42674 (SJRw 54490) - Z. juruanum: 9615 (MADw 23863) -H pallens: A.F. Wilson B.A. Krukoff 4746 (MADw 18544) - Z. latifo­ M-7 (MADw 18342); Milton Scott (SJRw lia: B.A. Krukoff 5798 (MADw 25119); A.C. 44776) - H platyloba: Curran & Haman 400 Persaud 63 (SJRw 9471); B.A. Krukoff 6212 (SJRw 2775); Record & Don Jaca 51 (SJRw (SJRw 36517); J. Cuatracasas 14251 (SJRw 16450); A. Dugand 1158; 535 (SJRw 35287) 42739); FDBG 3618 (SJRw 46562) - Z. longi­ - Klugiodendron laetum: Ll. Williams 4190 folia: Gutierrez R.74 & R.93 (MADw 22402); (MADw 15942); B.A. Krukoff 6457 (SJRw Stern, Eyde & Ayensu 1897 (MADw 24320); 36693); R.M. King 6170 (MADw 25071) H.C. Kluge 23 (SJRw 7136); Cooper & Slater 54 (SJRw 10152); Record & Kuylen 14 (SJRw 16413) - Z. peckii: H.W. Winzerling (SJRw * The wood anatomy indicates these species 10 172) - Z. recordii: Whitford & Stadtmiller could be placed in the segregate genus Marmar­ 64 (SJRw 3724); Record & Kuylen G.5 (type) oxylon. (SJRw 8836); H.W. Winzerling (1-20) (SJRw **Published incorrectly as A. Ducke 30 in 9869) - Z. stevensonii: D. Stevenson II (type) Tropical Woods No. 63, p. 2. (SJRw 3338); LB. Kinlock 17 (SJRw 17141).

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Fig. 7. Samanea saman (SJRw 45751). Coralloid vestures on pit apertures. x 3400. - - Fig. 8. Mar­ maroxylon racemosum (SJRw 21013). Coralloid vestures on coalesced pit apertures. x 4680. -­ Fig. 9. Pseudosamanea guachapele (SJRw 27084). Filamentous vestures on pit apertures. x 3830. -- Fig. 10. Ebenopsis flexicaulis (MADw 18344). Mostly coralloid to some bead-like vestures on coalesced pit apertures. x 4210. -- Fig. II. Pithecellobium dulce (SJRw 20990). Vestures absent from pit apertures. x 4170. -- Fig. 12. Zygia latifolia (SJRw 36517). Very few minute vestures (ar­ rows) on pit apertures. x 3490.

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Results and Discussion A transverse section with vessel lumina (VL), pit canals (PC) and vestures (V) in the pit chambers can be seen in Pseudosamanea guach­ apele (Fig. I). Zygia latifolia (Fig. 2) shows the inside of a vessel lumen (left) with few minute vestures on the pit apertures (arrows), the pit membrane (PM) and vestures in the pit cham­ ber. A portion of the secondary wall (pit bor­ der) arching over the pit cavity is shown at a. Three different types of vestures were ob­ served in the pit chambers of the specimens ex­ amined. Filamentous type vestures can be seen in Arthrosamanea pistaciaefolia (Fig. 3), coral­ loid in Pithecellobium (Marmaroxyion?)* dini­ zii (Fig. 4) and bead-like to coralloid in Pithe­ cellobium dulce (Fig. 5). These various vesture types are similar to those proposed by Bailey (1933). The filamentous type vesture forms a loose mat with continuously branching and anastomozing structure generally filling the pit chamber. Coralloid type vestures may be mas­ sive and trunk-like and dichotomize into thin­ ner branches. They generally do not complete­ ly fill the pit chamber but in some cases reach to the pit membrane. In Figure 5 the bead-like or papillary type appears to intergrade into a coralloid type rather than being completely rounded or bead-like throughout. Vestures can also be observed in and around the pit apertures. They may spread away from the pit aperture and cover, in part, the lumen side of the vessel. Two different types of ves­ tures were observed on the pit apertures. The coralloid type was observed in Samanea saman (Fig. 7) and on the coalesced apertures of Mar­ maroxylon racemosum (Fig. 8) while filamen­ tous vestures were observed in Pseudosamanea guachapele (Fig. 9). Figures 3 and 4 of the pit chambers and Fig­ ures 7, 8, and 9 of the pit apertures show dis­ tinct types of vestures which can be easily clas­ sified. However, not all vestures fit into one of these categories. Looking at the pit chambers, the vestures in Pithecellobium dulce (Fig. 5) for example appear mostly bead-like but some coralloid type structures can be seen (arrows). Intergrading of coralloid and bead-like vestures can also be observed on the pit apertures of Ebenopsis flexicaulis (Fig. 10). Between species the quantity of vestures in the pit chamber as well as on the pit aperture also varies. For example, the pit chambers are Fig. 13-15. Marmaroxylon racemosum. - 13: completely filled with vestures in Pseudosama- (SJRw 22055). Extensive coralloid vestures on pit apertures. x 3760. - 14: (SJRw 41145). Coralloid vestures on pit apertures. x 3910. - * The wood anatomy indicates these species 15: (SJRw 46469). Coralloid vestures on pit could be placed in the segregate genus Marmar­ apertures. x 3700. oxylon.

Downloaded from Brill.com09/27/2021 07:23:15AM via free access 64 lAW A Bulletin n.s., Vol. I (1-2), 1980 nea guachapele (Fig. I) and Arthrosamanea pis­ Cunn. (Proteaceae). IAWA Bull. 1974/1: taciaefolia (Fig. 3) whereas the vestures are lo­ 10-15. cated down and away from the pit membranes Cassens, D.L. 1973. Systematic wood anatomy in Pithecellobium (Marmaroxylon?) dinizii of the New World Pithecellobium (Legumi­ (Fig. 4) and Pithecellobium dulce (Fig. 5) thus nosae-Mimosoideae) complex. Doct. Thesis. exposing more of the pit rim (r). On the pit Univ. of Wisconsin, Madison, WI. aperture, vestures may be completely absent as -- & R.B. Miller. Wood anatomy of the New in Pithecellobium dulce (Fig. II). Note that World Pithecellobium (sensu lato). In prep­ vestures may be seen in the pit chambers (ar­ aration. row). Arrows show very few minute vestures Cote Jr., W.A. & A.C. Day. 1962. Vestured pits on the pit apertures of Zygia latifolia (Fig. 12) - fine structure and apparent relationship while abundant vestures may be seen in Pseu­ with warts. TAPPI 45: 906-910. dosamanea guachapele (Fig, 9) and Marmar­ Ishida, S. & J. Ohtani. 1970. Study of the pits oxylon racemosum (Fig. 13), for example. of wood cells using scanning electron micro­ Wide variation in the quantity of vestures loca­ scopy. I. An observation of the vestured ted on the pit aperture within Marmaroxylon pits in black locust, Robinia pseudo acacia racemosum was also observed (Fig. 13, 14, 15). Linn. Res. Bull. Coli. Exp. For. Hokkaido In the 87 specimens of Pithecellobium s.l. Univ. 27: 347-354. examined filamentous, coralloid and bead-like Metcalfe, C.R. & L. Chalk. 1950. Anatomy of to coralloid vestures were observed in the pit the Dicotyledons. Clarendon Press, Oxford. chambers while filamentous, coralloid and cor­ Meylan, B.A. & B.G. Butterfield. 1974. Occur­ alloid to some bead-like vestures were observed rence of vestured pits in the vessels and on the pit aperture. Variation in the quantity fibres of New Zealand woods. N.Z. J. Bot. and type of vesturing throughout the Pithecel­ 12: 3-18. lobium complex and even within some species Miller, R.B. 1977. Vestured pits in Boragina­ was observed. In some specimens, the different ceae. IAWA Bull. 1977/3: 43-48. types of vesture classifications intergrade. Record, SJ. 1925. Pits with cribriform mem­ Therefore, it seems futile to classify vestures branes. Trop. Woods 2: 10-13. for the Pithecellobium complex into any par­ - 1936. Classification of various anatomical ticular category. features of dicotyledonous woods. Trop. Woods 47: 12-27. Schmid, R. 1965. The fine structure of pits in Acknowledgements hardwoods, pp. 291-304. In: Cellular ultra­ The author is grateful to Dr. I.B. Sachs and structure of woody plants (ed. W.A. Cote Mr. R.E. Kinney of the U.S. Forest Products Jr.), Syracuse Univ. Press, Syracuse, NY. Laboratory for their assistance and use of the Scurfield, G., S.R. Silva & H.D. Ingle. 1970. SEM. Vessel wall structure: An investigation using scanning electron microscopy. Aust. References J. Bot. 18: 301-312. Bailey, I.W. 1933. The cambium and its deriva­ Vliet, G.1.C.M. van. 1975. Wood anatomy of tive tissues. VIII. Structure, distribution, Crypteroniaceae sensu lato. J. Microscopy and diagnostic significance of vestured pits 104: 65-82. in dicotyledons. 1. Am. Arb. 14: 259-273. - 1978. Vestured pits of Combretaceae and Butterfield, B.G. & B.A. Meylan. 1974. Vestured allied families. Acta Bot. Neerl. 27: 273- vessel and fiber pits in Persoonia toru A. 285.

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