IAWA Journal, Vol. 27 (1), 2006: 99–114

WOOD ANATOMY OF THE TRIBE CAESALPINIEAE (LEGUMINOSAE, ) IN

Narcisana Espinoza de Pernía and José Luis Melandri Laboratorio de Anatomía de Maderas, Facultad de Ciencias Forestales y Ambientales, Universidad de Los Andes, Mérida, Venezuela

SUMMARY We studied the microscopic wood anatomy of 8 genera and 30 species in the tribe Caesalpinieae, subfamily Caesalpinioideae, with a focus on the identification and comparative anatomy of these genera. Characters suitable for reliable identification include intervessel pit size, fibre wall thickness, septate fibres, storied structure, ray type, ray width, and silica bodies. A table of diagnostic characters, generic descriptions, and phot- omicrographs provide tools for identification and descriptive information for comparative and phylogenetic studies. Key words: Leguminosae, Caesalpinioideae, Caesalpinieae, wood anat- omy, Venezuela.

INTRODUCTION

The subfamily Caesalpinioideae comprises 160 genera and about 2175 species, most of which are tropical and subtropical trees and shrubs (Barneby et al. 1998; Herendeen et al. 2003). Recent studies divide it into four tribes: Caesalpinieae, Cassieae, Cercideae and Detarieae (GRIN 2001; ILDIS Legume Web 2002; Missouri Botanical Gardenʼs VAST 2002). Several authors have studied the wood anatomy of tribes, genera or species in the subfamily Caesalpinioideae. Cozzo (1951) studied the wood anatomy of the Argen- tinean Mimosoideae and Caesalpinioideae with many valuable observations. In 1955 Reinders-Gouwentak studied storied structure and taxonomic rank within the legumes. She showed that storied structure was an important feature and that many taxa have storied rays. Koeppen (1980) studied the arborescent Leguminosae with silica bodies in their wood. He found that only eight genera of subfamily Caesalpinioideae have silica bodies in their secondary xylem, including and Sclerolobium of the tribe Caesalpinieae. Loureiro and Silva (1981) described the wood of 7 species of Dimorphandra and Loureiro et al. (1983) described 5 species of Sclerolobium and 5 species of Tachigali. Barajas-Morales and León (1989) studied the wood anatomy of species from Mexico, two of which are spp. that also occur in Venezuela. Angarita (1991) described 7 species of Campsiandra from Venezuela. Later Espinoza de Pernía et al. (1998) described amazonicum and S. parahybum from

Associate Editor: Alex Wiedenhoeft

Downloaded from Brill.com10/04/2021 11:57:04AM via free access 100 IAWA Journal, Vol. 27 (1), 2006 Espinoza de Pernía & Melandri — Wood anatomy of Caesalpinieae 101

Venezuela. In 1981 Baretta-Kuipers published a comparative study of wood anatomy of the Leguminosae. She surveyed the entire family and distinguished two groups of Caesalpinioideae on the basis of ray features and parenchyma characteristics. Ranjani and Krishnamurthy (1988) showed a table of species of some Caesalpiniaceae with vestured pits. Wheeler and Baas (1992) studied the xylem evolution and ecological anatomy of legumes in fossil wood. In 1992 Nardi and Edlmann published a study of commercially important tropical woods used in Italy. They included a dichotomous key for diagnostic characters and described the macro and micro structure of 116 woods, 2 of which are Caesalpinieae from Venezuela, and also included photomicrographs. In 1999 Höhn reported on 23 West African genera of Caesalpinioideae and Mimosoideae. These were grouped into types according to wood structure, and although a key was not given, a synoptic table of characters was included. Miller and Détienne (2001) provided anatomical descriptions, photomicrographs and a dichotomous key of approximately 80 Guyanese timbers. Among these 20 legume genera were included. Herendeen (2000) reported the occurrence of morphological and anatomical characters with possible phylogenetic implications; some of these characters, such as the occurrence of vestures in vessel element pits may be phylogenetically informative in defining a large clade near the base of the family. In 1994 Gasson published on the wood anatomy of the tribe Sophoreae and related Caesalpinioideae and Papilionoideae, and in 2003 Gasson et al. gave a comprehensive account of the wood anatomy of the Caesalpinioideae. These papers reflect how wood anatomy can be used to derive diagnostic and phylogenetic information. The wood anatomy and identification of Venezuelan Caesalpinioideae is currently being studied by the present authors. Some genera of the tribe Caesalpinieae in Venezuela are trees with commercial tim- ber importance. particularly Caesalpinia coriaria, C. granadillo, excelsa, M. gong- grijpii (Mora 1974; JUNAC 1981; Arroyo 1983; INIA 1996; Aristeguieta 2003), Sclero- lobium (Aristeguieta 1973) and Tachigali paniculata (= Sclerolobium paniculatum) (Arroyo 1983). Others, such as Caesalpinia spp., Delonix regia (Aristeguieta 1973, 2003; Hoyos 1994) and (Hoyos 1994), are important as ornamental trees, in gardens, city avenues and parks. Caesalpinia coriaria is important in the tanning and dye industries (Aristeguieta 2003) and was exported to North America in earlier times (Hoyos 1994, 1985; Mabberley 1997). Caesalpinia granadillo is an endemic species in the dry zone of northern Venezuela (Hoyos 1985) and others such as Campsiandra curaara, C. emonensis, C. macrocarpa, C. nutans, C. pasibensis and C. velutina are endemic species in Venezuelan Guayana (Stergios 1996; Barneby et al. 1998). The present paper provides information on the microscopic structure of the wood, according to the proposed terminology in the List of Microscopic Features for Hard- wood Identification (IAWA Committee 1989), of 30 species of the tribe Caesalpinieae with their main distribution in Venezuela. The microscopic wood anatomy of the tribe Caesalpinieae was studied because of its great importance in the timber industry and the complexity of its anatomy. The anatomical descriptions, table of diagnostic features and photomicrographs provide tools for the identification of the genera and groups within the tribe Caesalpinieae.

Downloaded from Brill.com10/04/2021 11:57:04AM via free access 100 IAWA Journal, Vol. 27 (1), 2006 Espinoza de Pernía & Melandri — Wood anatomy of Caesalpinieae 101

MATERIALS AND METHODS

We examined 46 specimens representing 30 Venezuelan species from 8 of 11 genera in the tribe Caesalpinieae (Leguminosae, Caesalpinioideae): Caesalpinia (4/7), Camp- siandra (9/18), Delonix (1/1), Dimorphandra (4/8), Mora (2/2), Schizolobium (2/2), Sclerolobium (1/2), and Tachigali (7/19). No material was available to us of the genera Haematoxylon, Jacqueshuberia and Recordoxylon, which each have one species in Venezuela. The majority of the specimens were collected in Venezuela and are specimens from wood collections at the Laboratorio de Anatomía de Maderas de la Facultad de Ciencias Forestales y Ambientales de la Universidad de Los Andes, Mérida, Venezuela (MERw) and at the USDA Forest Service, Forest Products Laboratory, Madison, Wis- consin, USA (MADw and SJRw). Terminology and methodology followed the List of Microscopic Features for Hard- wood Identification (IAWA Committee 1989). For vessel diameters, vessel element lengths, fibre lengths and ray height 25 measurements were taken from each specimen and averaged. The values reported [e.g. 30 (50–110) 150 μm], are minimum value, range of averages, and maximum value. The measurements are accurate only to the 10 μm level, and are reported accordingly. For other quantitative values the most frequent range is reported. Generic descriptions follow in alphabetical order and features not listed in the generic descriptions are either absent or do not apply. Preparation of slides for the microscopic study was made following the methodology used at MERw (Corothie 1967). Photomicrographs were taken using a film camera with a light microscope. We followed the scientificnames in database ILDIS (http://www.ildis.org/Legume Web/) for all names except for Tachigali and Sclerolobium; for them we used the web site Grin Taxonomy (http://www.ars-grin.gov/cgi-bin/npgs/html/taxgenform.pl?language= sp) and W3Tropicos (http://mobot.mobot.org/W3T/Search/vast.html), because there is a discrepancy concerning synonymy between ILDIS and the other databases.

RESULTS Generic descriptions Caesalpinia L. — Fig. 1 & 2 Growth rings indistinct to distinct, marked by marginal parenchyma bands and/or thick-walled fibres in C. coriaria, C. ebano, C. granadillo; absent in C. sclerocarpa. Diffuse porous. — Vessels solitary and in radial multiples of 2–4, occasionally in clusters; 4–18 per mm2; 30 (50–110) 150 μm in diameter; 110 (130–170) 250 μm in element length. Simple perforation plates. Alternate intervessel pits; circular or oval, small to medium, 5–8 μm in diameter. Vessel-ray pits with distinct borders, similar to intervessel pits in size and shape; pits vestured. Dark brown gum-like deposits abundant in vessels. — Fibres non-septate; thick-walled to very thick-walled; 700 (850–1260) 1350 μm in length. — Axial parenchyma mostly in narrow bands or lines, up to three cells wide or bands more than three cells wide. Marginal banded parenchyma present in C. coriaria, C. ebano and C. granadillo. Paratracheal parenchyma vasicentric and aliform to confluent; aliform parenchyma winged, occasionally unilateral. Apotracheal parenchyma rarely diffuse-in-aggregates in C. ebano. Axial parenchyma 2–4 cells

Downloaded from Brill.com10/04/2021 11:57:04AM via free access 102 IAWA Journal, Vol. 27 (1), 2006 Espinoza de Pernía & Melandri — Wood anatomy of Caesalpinieae 103

Fig. 1–4. – 1: Caesalpinia coriaria, growth rings distinct (arrowheads); parenchyma in bands more than three cells wide and aliform. – 2: C. sclerocarpa, rays 1–3 cells wide and storied. – 3 & 4: Campsiandra curaara, parenchyma in bands more than three cells wide. – 4: C. emonensis, rays homocellular. — Scale bars = 250 μm.

Downloaded from Brill.com10/04/2021 11:57:04AM via free access 102 IAWA Journal, Vol. 27 (1), 2006 Espinoza de Pernía & Melandri — Wood anatomy of Caesalpinieae 103

per parenchyma strand. — Rays homocellular, cells typically procumbent; 10–16 per mm; 1 to 3 cells wide; 60 (130–200) 360 μm in height. — Storied structure in all the cell types, rays 4.5–6.5 tiers per axial mm (up to 7.5 in C. granadillo MERw 1162). — Prismatic crystals abundant in long chains in axial parenchyma cells, one crystal per cell or chamber. Material studied: 9 specimens, C. coriaria (Jacq.) Willd., MERw 1180, MADw 4851, MADw 4853, MADw 32283; C. ebano H. Karst., MADw 32287; C. granadillo (Pittier) Pittier, MERw 1162, MADw 20509, MADw 21236; C. sclerocarpa Standley, MERw 2502.

Campsiandra Benth. — Fig. 3–6 Growth rings absent. Diffuse porous. — Vessels solitary and in radial multiples of 2–4, occasionally up to 7; 2–7 per mm2; 70 (120–200) 280 μm in diameter; 130 (260– 450) 610 μm in element length. Simple perforation plates. Alternate intervessel pits; circular or oval, small to medium, 6–10 μm in diameter. Vessel-ray pits with distinct borders, similar to intervessel pits in size and shape; pits vestured. Dark brown gum-like deposits in vessels, but not observed in C. macrocarpa. — Fibres non-septate; very thick-walled; 1000 (1150–1510) 1720 μm in length. — Axial parenchyma mostly ali- form to confluent, in bands more than three cells wide; aliform parenchyma lozenge type. Apotracheal parenchyma occasionally diffuse-in-aggregates in C. angustifolia. Axial parenchyma 3–8 cells per parenchyma strand. — Rays mostly homocellular, cells typically procumbent, occasionally marginal cells slightly enlarged, rays rarely hetero- cellular, one row of upright and/or square cells; 6–13 per mm; 1 to 3 cells wide, pre- dominantly biseriate except in C. guayanensis and C. pasibensis; 110 (200–410) 810 μm in height. — Prismatic crystals in long chains in axial parenchyma cells, one crystal per cell or chamber. Material studied: 13 specimens, C. angustifolia Spruce ex Benth., MERw 2644, MERw 5382; C. curaara Stergios, MERw 5101; C. emonensis Stergios, MERw 5378, MERw 5383; C. gomez-alvareziana Stergios, MERw 5015, MERw 5099; C. guaya- nensis Stergios, MERw 5384; C. macrocarpa, MERw 5096; C. nutans Stergios, MERw 5387, MERw 5385; C. pasibensis Stergios, MERw 5376; C. velutina Stergios, MERw 5372.

Delonix regia (Bojer ex Hook.) Raf. — Fig. 7 & 8 Growth rings distinct, marked by marginal parenchyma bands and thick-walled fibres. Diffuse porous. — Vessels solitary and in radial multiples of 2–4, sometimes up to 8, occasionally in clusters; 7 per mm2; 100 (140) 190 μm in diameter; 150 (210) 290 μm in element length. Simple perforation plates. Alternate intervessel pits; circular or oval, occasionally with coalescent apertures, medium, 8–10 μm in diameter. Vessel-ray pits with distinct borders, similar to intervessel pits in size and shape; pits vestured. Reddish brown gum-like deposits in vessels. — Fibres non-septate; thin- to thick-walled; 900 (1110) 1340 μm in length. — Axial parenchyma paratracheal mostly aliform to con- fluent; aliform parenchyma lozenge. Banded parenchyma occasionally in wide bands of more than three cells wide. Marginal banded parenchyma present. Axial parenchyma

Downloaded from Brill.com10/04/2021 11:57:04AM via free access 104 IAWA Journal, Vol. 27 (1), 2006 Espinoza de Pernía & Melandri — Wood anatomy of Caesalpinieae 105

Fig. 5–8. – 5 & 6: Campsiandra velutina. – 5: Parenchyma in bands more than three cells wide. – 6: Rays 1–3 cells wide predominantly biseriate. – 7 & 8: Delonix regia. 7: Growth rings distinct; parenchyma vasicentric, aliform and confluent, marginal band (arrowheads) in lower half of photo. – 8: Rays 1–3 cells wide and not storied; prismatic crystals in axial parenchyma cells. — Scale bars = 250 μm.

Downloaded from Brill.com10/04/2021 11:57:04AM via free access 104 IAWA Journal, Vol. 27 (1), 2006 Espinoza de Pernía & Melandri — Wood anatomy of Caesalpinieae 105

Fig. 9–12. – 9 & 10: Dimorphandra cuprea subsp. ferruginea. – 9: Parenchyma aliform to con- fluent, marginal band (arrowheads) in lower half of photo. – 10: Rays heterocellular. – 11: Mora excelsa, parenchyma aliform to confluent and unilateral. – 12: M. gonggrijpii, rays homocellular. — Scale bars = 250 μm.

Downloaded from Brill.com10/04/2021 11:57:04AM via free access 106 IAWA Journal, Vol. 27 (1), 2006 Espinoza de Pernía & Melandri — Wood anatomy of Caesalpinieae 107

2–4 cells per parenchyma strand. — Rays homocellular, cells typically procumbent; 5–8 per mm; 1–3 cells wide; 140 (210) 270 μm in height. — Prismatic crystals common in long chains in axial parenchyma cells, one crystal per cell or chamber. Material studied: 1 specimen of Delonix regia (Bojer ex Hook.) Raf., MERw 4995.

Dimorphandra Schott — Fig. 9 & 10 Growth rings distinct to indistinct, marked by marginal parenchyma bands in D. cuprea subsp. ferruginea and thick-walled fibres in D. macrostachya and D. pen- nigera; growth rings absent in D. davisii. Diffuse porous. — Vessels solitary and in radial multiples of 2–4, sometimes up to 7; 2–10 per mm2; 80 (150–210) 280 μm in diameter; 200 (310–370) 550 μm in element length. Simple perforation plates. Alternate intervessel pits; circular or oval, minute to small, 3–7 μm in diameter. Vessel-ray pits with distinct borders, similar to intervessel pits in size and shape; pits vestured. Red- brown gum-like deposits common. — Fibres non-septate; thin- to thick-walled in D. macrostachya, D. pennigera, and thick-walled to very thick-walled in D. davisii, D. cuprea subsp. ferruginea; 800 (920–1320) 1490 μm in length. — Axial parenchyma mostly aliform to confluent; aliform parenchyma lozenge type. Banded parenchyma in wide bands more than three cells wide in D. davisii. Marginal banded parenchyma present in D. cuprea subsp. ferruginea. Axial parenchyma 2–5 cells per parenchyma strand. — Rays homocellular with typically procumbent cells to heterocellular, with one row of upright and/or square cells; 4–10 per mm; 1 to 2, occasionally 3 cells wide; 100 (220–300) 440 μm in height. Rays sometimes irregularly storied or in echelon arrangement. — Prismatic crystals mostly common in long chains in axial parenchyma cells, but not common in D. pennigera, one crystal per cell or chamber. Material studied: 5 specimens, D. cuprea subsp. ferruginea Ducke, MADw 31798; D. davisii Sprague & Sandwith, MERw 193; D. macrostachya Benth., MERw 226, MERw 3339; D. pennigera Tul., MADw 31797. Note: Metcalfe & Chalk (1950) and Mainieri & Chimelo (1989) characterized the rays of Dimorphandra as homocellular, but Miller & Détienne (2001) described the rays as homocellular to heterocellular. In addition, Détienne et al. (1982) reported the rays as homocellular to sub-homocellular and Loureiro & Silva (1981) as heterocellular.

Mora Schomb. ex Benth. — Fig. 11 & 12 Growth rings distinct, marked by marginal parenchyma bands. Diffuse porous. — Vessels solitary and in radial multiples of 2–3, sometimes up to 5; 4–5 per mm2; 90 (140–170) 230 μm in diameter; 170 (300–420) 600 μm in element length. Simple perforation plates. Alternate intervessel pits; circular or oval and sometimes polygonal, minute to small, 2–5 μm in diameter. Vessel-ray pits with distinct borders, similar to intervessel pits in size and shape; pits vestured. Reddish brown gum-like deposits common. — Fibres non-septate; thick-walled to very thick-walled; 1050 (1290–1460) 1690 μm in length. — Axial parenchyma vasicentric, mostly aliform to confluent; ali- form parenchyma lozenge type, occasionally unilateral. Banded parenchyma occasion- ally in wide bands of more than three cells wide. Marginal banded parenchyma present.

Downloaded from Brill.com10/04/2021 11:57:04AM via free access 106 IAWA Journal, Vol. 27 (1), 2006 Espinoza de Pernía & Melandri — Wood anatomy of Caesalpinieae 107

Axial parenchyma 2–4 cells per parenchyma strand. — Rays homocellular, cells typically procumbent, marginal cells sometimes slightly enlarged; 5–9 per mm; 2 to 3 cells wide; 110 (250–290) 500 μm in height. — Prismatic crystals common in long chains in axial parenchyma cells, one crystal per cell or chamber. Material studied: 4 specimens, M. excelsa Benth., MERw 127; M. gonggrijpii (Kleinhoonte) Sandwith, MERw 1742, MERw 4563, MADw 24010. Note: Metcalfe & Chalk (1950) and Gasson et al. (2003) characterized the rays of Mora as heterocellular, JUNAC (1981) as homocellular to heterocellular, but Kribs (1968), Détienne et al. (1982), Nardi & Edlmann (1992) and Miller & Détienne (2001) described the rays as homocellular.

Schizolobium Vogel — Fig. 13 & 14 Growth rings distinct, marked by thick-walled fibres in S. parahyba, and absent in S. amazonicum. Diffuse porous. — Vessels solitary and in radial multiples of 2–3; 2–3 per mm2; 150 (200–230) 290 μm in diameter; 200 (310–380) 540 μm in element length. Simple perforation plates. Alternate intervessel pits; circular or oval, medium to large, 8–11 μm in diameter. Vessel-ray pits with distinct borders, similar to intervessel pits in size and shape; pits vestured. Deposits not observed. — Fibres septate; thin- to thick-walled; 940 (970–1240) 1620 μm in length. — Axial parenchyma scanty to thin vasicentric. Axial parenchyma 2–4 cells per parenchyma strand. — Rays mostly homocellular, cells typically procumbent, to occasionally heterocellular, with one row of upright and/or square cells; 4–7 per mm; 2–5 cells wide; 130 (320–350) 600 μm in height. — Prismatic crystals not common, when present crystals in short chains in axial parenchyma cells only in S. amazonicum, rarely present in upright and/or square ray cells of S. parahyba, one crystal per cell or chamber. Druses present in ray and axial parenchyma cells of S. amazonicum. Material studied: 3 specimens, S. amazonicum Huber ex Ducke, MERw 2548; S. parahyba (Vell.) S.F. Blake, MERw 3319, MERw 4886. Note: JUNAC (1981) reported storied fibres in S. parahyba, this feature was not observed in our specimens. Metcalfe & Chalk (1950), JUNAC (1981), Mainieri & Chimelo (1989) and Gasson et al. (2003) characterized the rays of Schizolobium as homocellular, but Espinoza de Pernía et al. (1998) described the rays as mostly homo- cellular to occasionally heterocellular.

Sclerolobium subbullatum Ducke — Fig. 15 & 16 Growth rings distinct, marked by thick-walled fibres. Diffuse porous. — Vessels solitary and in radial multiples of 2–3, sometimes up to 5; 2–3 per mm2; 140 (210) 260 μm in diameter; 280 (450) 610 μm in element length. Simple perforation plates. Alternate intervessel pits; circular or oval and polygonal, medium, 7–10 μm in diameter. Vessel-ray pits with distinct borders, similar to intervessel pits in size and shape; pits vestured. Deposits not observed. — Fibres non-septate; thin- to thick-walled; 1050 (1260) 1410 μm in length. — Axial parenchyma scanty to vasicentric, aliform paren- chyma lozenge type. Axial parenchyma 2–4 cells per parenchyma strand. — Rays homocellular, cells typically procumbent; 10–16 per mm; exclusively uniseriate;

Downloaded from Brill.com10/04/2021 11:57:04AM via free access 108 IAWA Journal, Vol. 27 (1), 2006 Espinoza de Pernía & Melandri — Wood anatomy of Caesalpinieae 109

Fig. 13–16. – 13: Schizolobium parahyba, growth rings distinct, marked by thick-walled fibres; parenchyma vasicentric. – 14: S. amazonicum, fibres septate; rays homocellular. – 15 & 16: Sclerolobium subbullatum. – 15: Growth rings distinct, marked by thick-walled fibres in latewood; parenchyma aliform. – 16: Rays exclusively uniseriate. — Scale bars = 250 μm.

Downloaded from Brill.com10/04/2021 11:57:04AM via free access 108 IAWA Journal, Vol. 27 (1), 2006 Espinoza de Pernía & Melandri — Wood anatomy of Caesalpinieae 109

110 (260) 360 μm in height. — Prismatic crystals not observed. Silica bodies frequent in the ray cells. Material studied: 1 specimen of S. subbullatum Ducke, MERw 2355. Note: Metcalfe & Chalk (1950), Kribs (1968), JUNAC (1981), Détienne et al. (1982), Mainieri & Chimelo (1989) and Miller & Détienne (2001) characterized the rays of Sclerolobium as homocellular, Gasson et al. (2003) reported the rays as homocellular and occasionally heterocellular. Loureiro et al. (1983) described the rays as heterocellular in S. subbullatum.

Tachigali Aublet — Fig. 17 & 18 Growth rings distinct, marked by thick-walled fibres present in T. chrysophylla, T. melinonii, T. paniculata var. alba, T. polyphylla, T. rigida, T. setifera, and absent in T. micrantha. Diffuse porous. — Vessels solitary and in radial multiples of 2–3; some- times up to 8, occasionally in clusters; 2–7 per mm2 in T. chrysophylla, T. melinonii, T. micrantha, T. paniculata var. alba, T. polyphylla, T. setifera and 7–15 per mm2 in T. rigida; 80 (110–230) 350 μm in diameter; 140 (300–500) 630 μm in element length. Simple perforation plates. Alternate intervessel pits; circular or oval and polygonal; small to medium, 4–8 μm in diameter. Vessel-ray pits with distinct borders, similar to intervessel pits in size and shape; pits vestured. Reddish brown gum-like deposits in T. micrantha and T. rigida; deposits not observed in T. chrysophylla, T. melinonii, T. paniculata var. alba, T. polyphylla and T. setifera. — Fibres non-septate; thin- to thick-

Fig. 17–18. Tachigali melinonii. – 17: Growth rings distinct, marked by thick-walled fibres at latewood boundary. – 18: Rays homocellular; silica bodies in the ray cells (arrow). — Scale bars = 250 μm.

Downloaded from Brill.com10/04/2021 11:57:04AM via free access

110 IAWA Journal, Vol. 27 (1), 2006 Espinoza de Pernía & Melandri — Wood anatomy of Caesalpinieae 111 Silica bodies Silica

• • • •

Druses

•

axial parenchyma cells parenchyma axial

Prismatic crystals in in crystals Prismatic ) )

• • • • • • • • • • • • • • • • • • • • • • • •

( ( Storied structure Storied • • • •

Ray width Ray 1 1 1 1 1 1 1–3 1–3 1–3 1–3 1–3 1–3 1–3 1–3 2–4 2–5 1–2 1–2

1–3 +2 1–3 +2 1–3 +2 1–3 +2 1–3 +2 1–3 +2 1–3 +2 1–2 (3) 1–2 (3) 1–2 (3) 1–2 (3) 1–2 (3) Rays heterocellular Rays ) ) ) ) ) ) ) ) )

• • • • • • • • • • • • • • • ( ( ( ( ( ( ( ( (

cumbent marginal cells marginal cumbent

slightly enlarged pro- enlarged slightly

Rays homocellular with homocellular Rays * • • • • • • • • • • • • • • • •

Rays homocellular Rays + + + + + + + + + • • • • • • • • • • • • • • • • • • • • • • • • • • • • • •

ber of cells per strand per cells of ber Axial parenchyma num- parenchyma Axial 2–4 2–4 2–4 2–4 3–5 3–5 4–8 3–5 3–6 3–6 3–7 3–5 3–5 2–4 3–4 3–4 2–4 3–4 2–4 3–4 2–4 2–4 2–4 2–4 2–4

2–4 (5) 2–4 (8) 3–4 (7) 3–4 (6) 3–4 (6)

Marginal parenchyma Marginal

• • • • • • • wide parenchyma wide

) ) ) ) ) )

Bands more than 3 cells 3 than more Bands + *

• • • • • • • • • • • • • • • • • • • ( ( ( ( ( (

cells wide parenchyma wide cells

Narrow bands up to 3 to up bands Narrow + + • •

Unilaterally parenchyma Unilaterally ) ) ) ) ) ) • • • • • • ( ( ( ( ( (

Confluent parenchyma Confluent )

+ * • • • • • • • • • • • • • • • • • • • • • •

• ( Aliform parenchyma Aliform

)

+ + + * • • • • • • • • • • • • • • • • • • • • • • • • • (

Vasicentric parenchyma Vasicentric ) + +

• • • • • • • • • • • •

• • (

Scanty parenchyma Scanty

• very thick-walled very

Fibres usually thick- to thick- usually Fibres * • • • • • • • • • • • • • • • • thick-walled

Fibres usually thin- to thin- usually Fibres * • • • • • • • • • • • • Septate fibres Septate • •

Intervessel pits size pits Intervessel 5–8 5–7 6–8 6–9 6–9 6–8 6–8 6–9 6–8 6–8 6–8 7–9 4–5 3–4 4–7 4–7 2–3 3–5 7–8 7–8 6–8 5–6 5–8 4–7 5–6 8–11 6–10 8–10 8–10 7–10

Vessels per mm2 per Vessels 4–5 2–5 2–3 2–6 2–6 2–7 4–5 2–5 3–4 4–5 5–7 4–5 2–5 6–8 4–5 4–5 2–3 2–3 2–3 2–5 2–3 3–5 3–6 4–5 4–7

5–10 7–15

16–18 15–17 14–18 Growth rings Growth

• • • • • • • • • • • • • • • • •

ferruginea

subsp.

alba

var. var.

= distinct feature = occasionally = predominantly

C. ebano C. granadillo C. sclerocarpa C. curaara C. emonensis C. gomez-alvareziana C. guayanensis C. macrocarpa C. nutans C. pasibensis C. velutina D. davisii D. macrostachya D. pennigera M. gonggrijpii S. parahyba melinonii T. micrantha T. paniculata T. polyphylla T. rigida T. setifera T. • + Table 1. Synoptic table of Table anatomical features ( ) Species studied Caesalpinia coriaria Campsiandra angustifolia Delonix regia Dimorphandra cuprea Mora excelsa Sclerolobium subbullatum chrysophylla Tachigali

Downloaded from Brill.com10/04/2021 11:57:04AM via free access 110 IAWA Journal, Vol. 27 (1), 2006 Espinoza de Pernía & Melandri — Wood anatomy of Caesalpinieae 111

walled; 800 (940–1290) 1320 μm in length. Axial parenchyma scanty to vasicentric, aliform to confluent, aliform parenchyma lozenge type, occasionally unilateral. —Axial parenchyma 2–4 cells, occasionally up to 8 cells per parenchyma strand. — Rays homocellular, cells typically procumbent (marginal cells sometimes slightly enlarged); 7–14 per mm; exclusively or mostly uniseriate, occasionally biseriate in T. melinonii, T. micrantha and T. setifera; 90 (110–340) 450 μm in height. Prismatic crystals in axial parenchyma cells in T. paniculata var. alba, T. rigida and T. setifera; not observed in T. melinonii, T. micrantha, T. chrysophylla and T. polyphylla; one crystal per cell or chamber. Silica bodies frequent in the ray cells in T. chrysophylla, T. melinonii and T. polyphylla (rarely present in axial parenchyma cells and fibres); not observed in T. micrantha, T. paniculata var. alba, T. rigida and T. setifera. Material studied: 10 specimens, T. chrysophylla (Poepp.) Zarucchi & Herend., MERw 3346; T. melinonii (Harms) Zarucchi & Herend., MERw 2575, MERw 4287; T. micrantha (L.O. Williams) Zarucchi & Herend., MERw 2543; T. paniculata var. alba (Ducke) Dwyer, MERw 779, MERw 2090; T. polyphylla (Ducke) Poeppig., MERw 2439; T. rigida Ducke, MADw 23532, MADw 31298; T. setifera (Ducke) Zarucchi & Herend., MERw 2546. Note: Metcalfe & Chalk (1950), Détienne et al. (1982) and Richter & Dallwitz (2000) characterized the rays of Tachigali as homocellular, Gasson et al. (2003) reported the rays as homocellular and occasionally heterocellular, but Loureiro et al. (1983) described the rays as heterocellular in T. alba and T. plumbea. The feature silica bodies was not observed by Koeppen (1980) in T. melinonii (= Sclerolobium melinonii).

DISCUSSION

Like most genera and species in the family Leguminosae, the species we described have simple perforation plates, vestured pits, vessel-ray pits similar to the intervessel pits in size and shape, inconspicuous fibre pits, and axial parenchyma with mostly 2–4 cells per strand. Features that are variable in Leguminosae including growth rings, poros- ity, helical thickenings, septate fibres, axial parenchyma patterns, ray type, width and height, are also somewhat variable in the tribe Caesalpinieae. Table 1 provides tabulated anatomical information to help in the identification of the species and genera of the Venezuelan Caesalpinieae, but some separations are not so easy and other macroscopic features and explanations are necessary. Features within the tribe Caesalpinieae that are diagnostic include: septate fibres, fibre wall thickness, ray composition, ray width, storied structure and silica bodies. Quantitative features also vary (see Table 1), but most vary too much to be useful in identifications or comparisons. The exceptions are intervessel pit size and ray width. The size of the intervessel pits is a good diagnostic character. In most Caesalpinieae the pits are commonly medium to large; however, in Dimorphandra and Mora they are minute to small. Another good diagnostic character is the presence of septate fibres, but only Schizolobium (Fig. 14) has septate fibres in Venezuelan Caesalpinieae. Fibre wall thickness is typically not a good diagnostic character and was treated as variable

Downloaded from Brill.com10/04/2021 11:57:04AM via free access 112 IAWA Journal, Vol. 27 (1), 2006 Espinoza de Pernía & Melandri — Wood anatomy of Caesalpinieae 113 or inconsistent by Metcalfe & Chalk (1950), Mainieri & Chimelo (1989), Miller & Détienne (2001) and Gasson et al. (2003). However, we observed very thick-walled fibres inCaesalpinia, Campsiandra, Dimorphandra cuprea subsp. ferruginea, D. davisii and Mora. Axial parenchyma is typically a very good diagnostic feature for the Leguminosae, but in the Venezuelan Caesalpinieae it is not particularly useful (see Table 1). Generally the parenchyma is paratracheal in most species: vasicentric, aliform to confluent, bands up to three cells wide in Caesalpinia coriaria and C. ebano; more than three cells wide commonly in Caesalpinia (Fig. 1), Campsiandra (Fig. 3 & 5), Dimorphandra cuprea subsp. ferruginea, D. davisii and Mora gonggrijpii. Marginal in Caesalpinia (except in C. sclerocarpa), Delonix regia (Fig. 7), Dimorphandra cuprea subsp. ferruginea (Fig. 9) and Mora. Ray composition is not consistent in the literature (see notes in the generic descrip- tions), but it is still a useful feature if applied correctly. Within Caesalpinieae there are three groups that can be separated using ray composition or type. One group has ex- clusively homocellular rays: Caesalpinia, Delonix regia, Mora (Fig. 12), Sclerolobium subbullatum, Tachigali (Fig. 18). Another group has homocellular and heterocellular rays: Campsiandra (Fig. 4), Dimorphandra (Fig. 10) and Schizolobium (Fig. 14). There is also an intermediate group that has homocellular rays with slightly enlarged procumbent marginal cells: Campsiandra, Mora and Tachigali. However, Metcalfe and Chalk (1950) and Gasson et al. (2003) reported the rays as hetercellular in Mora. Ray width is a very good diagnostic character in Caesalpinieae. Rays are exclusively or predominantly uniseriate in Sclerolobium subbullatum (Fig. 16) and Tachigali, 4 or more cells wide in Schizolobium, and mostly 1–3 cells wide in the other genera (Fig. 2 & 6). Storied structure is not common in Caesalpinieae but it is present in Caesalpinia studied in this paper (Fig. 2) and occasionally irregularly storied rays (rays in echelon) occur in Dimorphandra. Prismatic crystals are typically present in long chains in the axial parenchyma, one crystal per chamber of most species, but their presence in the ray cells of Schizolobium parahyba is diagnostic. Druses occasionally occur in the ray cells and axial parenchyma of Schizolobium amazonicum (Espinoza de Pernía et al. 1998) and are quite diagnostic when present. The absence of prismatic crystals and the presence of silica bodies in the ray cells of Sclerolobium subbullatum, Tachigali chrysophylla and T. melinonii is also diagnostic.

CONCLUSIONS

Many of the Venezuelan species in the tribe Caesalpinieae can be separated using a number of diagnostic features in combination. The most important features are: rays exclusively or predominantly uniseriate, ray type, silica bodies, septate fibres, fibre wall thickness, storied structure, intervessel pit size and marginal parenchyma bands. The interpretation of homocellular versus heterocellular rays in some genera is not consistent in the literature and might need to be re-examined in other legume woods and possibly re-defined and re-interpreted for use in keys and descriptions.

Downloaded from Brill.com10/04/2021 11:57:04AM via free access 112 IAWA Journal, Vol. 27 (1), 2006 Espinoza de Pernía & Melandri — Wood anatomy of Caesalpinieae 113

ACKNOWLEDGEMENTS

We wish to thank Dr. Regis B. Miller, USDA Forest Service, Forest Products Laboratory, Madison, Wisconsin, USA for helpful suggestions and also for providing the sectioning blocks of some material studied. For the financial help we wish to thank the C.D.C.H.T of the Universidad de Los Andes (Project: FO-392-96-01-B), Mérida, Venezuela.

REFERENCES Angarita, de T. Rita, N. 1991. Estudio anatómico del leño de siete especies del género Campsian- dra. Trabajo de ascenso, Universidad de Los Andes, Facultad de Ciencias Forestales, Mérida, Venezuela. Aristeguieta, L. 1973. Familias y géneros de los árboles de Venezuela. Instituto Botánico, Dirección de Recursos Naturales Renovables, Ministerio de Agricultura y Cría, Caracas, Venezuela. Aristeguieta, L. 2003. Estudio dendrológico de la Flora Venezolana. Vol. 38. Academia de Ciencias Físicas, Matemáticas y Naturales, Caracas, Venezuela. Arroyo, J. 1983. Propiedades tecnológicas y posibles usos de 131 maderas Venezolanas. Universidad de Los Andes, Facultad de Ciencias Forestales, Consejo de Publicaciones, Talleres Gráficos Universitarios, Mérida, Venezuela. Barajas-Morales, J. & C. León. 1989. Anatomía de maderas de México: especies de una selva baja caducifolia. Universidad Nacional Autónoma de México, Publicaciones Especiales del Instituto de Biología 1, México D.F. Baretta-Kuipers, T. 1981. Wood anatomy of Leguminosae: its relevance to taxonomy. In: R.M. Polhill & P.H. Raven (eds.), Advances in Legume Systematics, Part 2: 677–705. Royal Botanic Gardens, Kew. Barneby, R., B. Stergios, R. Cowan, P. Berry, J. Zarucchi, R. Wunderlin, D. Kearns, M.F da Silva, A. Sprada, D. Velaázquez, N. Xena & G. Aymard. 1998. Caesalpiniaceae. In: P. Berry, B. Holst & K. Yatskievych (eds.), Flora of the Venezuelan Guayana. Vol. 4: 1–121. Missouri Botanical Garden Press, St. Louis, USA. Corothie, H. 1967. Estructura anatómica de 47 maderas de la Guayana Venezolana. Laboratorio Nacional de Productos Forestales, Universidad de los Andes, Ministerio de Agricultura y Cría, Mérida, Venezuela. Cozzo, D. 1951. Anatomía del leño secundario de las Leguminosas Mimosoideas y Cesalpinioideas Argentinas. Rev. Inst. Nac. Invest. Ci. Nat. II-2. Détienne, P., P. Jacquet & A. Mariaux. 1982. Manuel dʼ identification des bois tropicaux. 3. Française. Centre Technique Forestier Tropical, Nogent-sur-Marne, France. Espinoza de Pernía, N., O. Araque & W.J. León. 1998. Cristales en la madera de Schizolobium ama- zonicum y S. parahybum (Caesalpinioideae, Leguminosae). Rev. Forest. Venez. 42: 9–13. Gasson, P. 1994. Wood anatomy of the tribe Sophoreae and related Caesalpinioideae and Papilion- oideae. In: I.K. Ferguson & S. Tucker (eds.), Advances in Legume Systematics, Part 6: 165–203. Royal Botanic Gardens, Kew. Gasson P., C. Trafford & B. Matthews. 2003. Wood anatomy of Caesalpinioideae. In: B.B. Klit- gaard & A. Bruneau (eds.), Advances in Legume Systematics, Part 10: 63–93. Royal Botanic Gardens, Kew. GRIN Taxonomy 2001. The Germplasm Resources Information Network. http://www.ars-grin.gov/ npgs/tax/taxgenform.html [2004, April 15]. Herendeen, P.S. 2000. Structural evolution in Caesalpinioideae (Leguminosae). In: P.S. Heren- deen & A. Bruneau (eds.), Advances in Legume Systematics, Part 9: 45–64. Royal Botanic Gardens, Kew. Herendeen, P.S., A. Bruneau & G.P. Lewis. 2003. Phylogenetic relationships in Caesalpinioid Legumes: A preliminary analysis based on morphological and molecular data. In: B.B. Klit- gaard & A. Bruneau (eds.), Advances in Legume Systematics, Part 10: 37–62. Royal Botanic Gardens, Kew.

Downloaded from Brill.com10/04/2021 11:57:04AM via free access 114 IAWA Journal, Vol. 27 (1), 2006

Höhn, A. 1999. Wood anatomy of selected West African species of Caesalpinioideae and Mimosoi- deae (Leguminosae): a comparative study. IAWA J. 20: 115–146. Hoyos, F. Jesús. 1985. Flora de la Isla de Margarita. Sociedad y Fundación de Ciencias Naturales La Salle. Monografía N° 34. Caracas, Venezuela. Hoyos, F. Jesús. 1994. Guía de árboles de Venezuela. Sociedad de Ciencias Naturales La Salle. Monografía N° 32. Caracas, Venezuela. IAWA Committee. 1989. IAWA list of microscopic features for hardwood identification. IAWA Bull. n.s. 10: 219–332. ILDIS Legume Web. 2002. International Legume Database & Information Service. http:// www.ildis.org/LegumeWeb/online.shtml [2003, May 03]. INIA. 1996. Manual de identificación de especies forestales de la subregión Andina. Instituto Nacio- nal de Investigaciones Agrarias, Talleres Gráficos de Asociación Editorial Stella, Lima, Perú. JUNAC. 1981. Descripción general y anatómica de 105 maderas del grupo Andino. PADT-REPORT, Junta del acuerdo de Cartagena (JUNAC), Lima, Perú. Koeppen, R.C. 1980. Silica bodies in wood of arborescent Leguminosae. IAWA Bull. n.s. 1: 180–184. Kribs, D.A. 1968. Commercial foreign woods on the American market. Dover Publications, New York. Loureiro, A.A. & M.F. da Silva. 1981. Estudo dendrológico e anatômico do lenho de 7 espécies e 3 subespécies de Dimorphandra (Leguminosae–Caesalpinioideae). Acta Amazonica 11: 561– 581. Loureiro, A.A., F.J. Vasconcellos & M.F. da Silva. 1983. Contribuicao ao estudo anatômico do lenho de 5 espécies de Sclerolobium e 5 espécies de Tachigalia (Leguminosae) da Amazónia. Acta Amazonica 13: 149–170. Mabberley, D.J. 1997. The -book. A portable dictionary of the vascular . Ed. 2. Cam- bridge University Press. Mainieri, C. & J.P. Chimelo. 1989. Fichas de características das madeiras brasileiras. Ed. 2. Instituto de Pesquisas Tecnológicas, Divisao de Madeiras, São Paulo, Brazil. Metcalfe, C.R. & L. Chalk. 1950. Anatomy of the . Vol. I. Clarendon Press, Oxford. Miller, R.B. & P. Détienne. 2001. Mayor timber trees of Guyana. Wood anatomy. Tropenbos Series 20, The Tropenbos Foundation, Wageningen, The Netherlands. Missouri Botanical Gardenʼs VAST. 2002. http://www.mobot.org/ [2003, June 20]. Mora, J. 1974. Características tecnológicas de 37 maderas Venezolanas. Universidad de Los Andes, Facultad de Ciencias Forestales, Escuela de Ingeniería Forestal, Mérida, Venezuela. Nardi Berti, R. & M. Edlmann Abbate. 1992. Legnami tropicali importati in Italia: anatomia e iden- tificazione. Vol. II. América Latina. C.N.R., Firenze. Ranjani, K. & K.V. Krishnamurthy. 1988. Nature of vestures in the vestured pits of some Caes- alpiniaceae. IAWA Bull. n.s. 9: 31–33. Reinders-Gouwentak, C.A. 1955. The storied-structure-features and the taxonomic rank of the leguminous taxa. Acta Bot. Neerl. 4: 460–470. Richter, H.G & M.J. Dallwitz. 2000. Commercial timbers: descriptions, illustrations, identification and information retrieval. [Online] In English, French, German and Spanish. Version: 18th October 2002. http://biodiversity.uno.edu/delta/ [24 May 2002]. Stergios, Basil. 1996. Contributions to South American Caesalpiniaceae. II. A taxonomic update of Campsiandra (Caesalpiniaceae). Novon 6: 434–459. Wheeler, E.A. & P. Baas. 1992. Fossil wood of the Leguminosae: a case study in xylem evolu- tion and ecological anatomy. In: P.S Herendeen & D.I. Dilcher (eds.), Advances in Legume Systematics, Part 4, The Fossil Record: 281–301, Royal Botanic Gardens, Kew.

Downloaded from Brill.com10/04/2021 11:57:04AM via free access