WOOD ANATOMY of CAESALPINIA S.S., COULTERIA, ERYTHROSTEMON, GUILANDINA, LIBIDIBIA, MEZONEURON, POINCIANELLA, POMARIA and TARA

IAWA Journal, Vol. 30 (3), 2009: 247–276

WOOD ANATOMY OF CAESALPINIA s.s., COULTERIA, ERYTHROSTEMON, GUILANDINA, LIBIDIBIA, MEZONEURON, POINCIANELLA, Pomaria AND TARA (LEGUMINOSAE, CAESALPINIOIDEAE, CAESALPINIEAE)

Peter Gasson, Kate Warner and Gwilym Lewis Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, United Kingdom

SUMMARY

Caesalpinia s.l. traditionally comprised c. 140 species in the New and Old World tropics, and contained a maximum of 25 generic synonyms. The genus in its broadest sense has been shown to be polyphyletic in molecular studies, and most species have now been assigned to reinstated segregate genera: Caesalpinia s.s. (c. 25 spp.), Coulteria (10 spp.), Erythrostemon (13 spp.), Guilandina (c. 7 spp.), Libidibia (8 spp.), Mezoneuron (c. 26 spp.), Poincianella (c. 35 spp.), Pomaria (16 spp.) and Tara (3 spp.). About 15 Asian taxa remain unassigned pending more data, especially DNA sequences. In this paper we describe the wood anatomy of these nine segregate genera, outlining the features that consistently help define some of them. We have examined the wood of 27 species representing all the woody segregate genera and found wood descriptions of three more species in the literature. Most species lack well defined growth rings, vessels are solitary and in radial multiples, intervessel pitting is alternate and vestured, fibres are mainly non-septate, axial parenchyma is aliform to confluent and irregularly storied, and the rays are mainly 1–2-seriate, mostly non-storied, and of varying height. Prismatic crystals are in chambered axial parenchyma cells in all except Erythrostemon gilliesii (Hook.) Link, and in ray cells in many species. Libidibia is well defined, with storied axial parenchyma, narrow short storied homocellular rays and lacking crystals in ray cells. Tara is also well defined with non-storied heterocellular rays and some ray cells containing crystals. The other genera are less consistent in wood characters. In Caesalpinia s.s. the rays are not storied, and most species lack crystals in ray cells. Coulteria has some species with storied rays and all have homocellular rays and crystals in ray cells. Poincianella is particularly poorly defined from a wood anatomical point of view, perhaps indicating that it can be further segregated. A few Poincianella species have septate fibres, which are otherwise seen only inLibidibia corymbosa. Mezoneuron has non-storied, heterocellular rays. The two species of Guilandina we examined have wide vessels and heterocellular rays containing crystals. Only two species of Erythrostemon were examined and E. gilliesii was unusual in having ring porous wood and very wide rays (but the sample was cultivated at Kew, and we do not know its porosity in its native range). Caesalpinia decapetala (Roth) Alston (originally described as Reichardia decapetala Roth) and Caesalpinia sappan L. from the Old World have not been reassigned to a segregate genus. Pomaria is mainly herbaceous and we have included some information on it. Key words: Caesalpinia s.s., Coulteria, Erythrostemon, Guilandina, Libidibia, Mezoneuron, Poincianella, Tara, Pomaria, wood anatomy.

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INTRODUCTION

Caesalpinia s.l. traditionally comprised c. 140 species in the New and Old World tropics, and contained 25 generic synonyms (Lewis 2005). The genus in its broadest sense has been shown to be polyphyletic in morphological and molecular studies (Lewis & Schrire 1995; Simpson & Miao 1997; Lewis 1998; Simpson 1998, 1999; Simpson & Lewis 2003, Simpson et al. 2003), and most species have now been assigned to reinstated segregate genera, whose names and/or status differ from the group names used in Lewis (1998) as follows: Caesalpinia s.s. (c. 25 spp.), Coulteria (10 spp., Brasilettia group in Lewis (1998)), Erythrostemon (13 spp., part of Erythrostemon/Poincianella group in Lewis (1998)), Guilandina (c. 7 spp., subgenus Guilandina), Libidibia (8 spp., Libidibia group), Mezoneuron (c. 26 spp, subgenus Mezoneuron), Poincianella (c. 35 spp., part of Erythrostemon/Poincianella group), Tara (3 spp., Russellodendron group in Lewis (1998)). About 15 Asian taxa remain unassigned pending further analysis, especially of molecular data. Pomaria was once largely included in Hoffmannseggia but is now considered more closely related to Caesalpinia s.l. based on molecular studies by Simpson et al. (2003). Some species of Caesalpinia s.l. (e.g. Poincianella pannosa and P. exostemma) have considerable morphological variation in their foliage, leaflet size, shape and indumentum, which has led to a proliferation of species names now included in synonymy. In this paper we describe the wood anatomy of the genus in its broadest sense, and outline the features (if any) that consistently define each segregate genus. Our major aim was to ascertain how closely the wood anatomy reflects the reclassification ofCaesalpinia s.l. into putatively monophyletic segregate genera. The wood anatomy of Caesalpinia s.l. is not simply of academic interest. Several species are well-known and of considerable economic importance. Caesalpinia echinata (currently tentatively placed in the Poincianella-Erythrostemon alliance, although proving to be molecularly distinct), locally known as pau-brasil (Brazil wood) and the wood after which the country Brazil was named, is an emblematic tree of the Atlantic forests of coastal Brazil and is the only wood used to make professional high-quality violin bows. There is a considerable literature on this species, which has recently been listed under Appendix 2 of CITES regulations as a timber not to be internationally traded. Another well-known species, Caesalpinia pulcherrima L. from Mexico and Guatemala, is widely cultivated in warm climates as an attractive ornamental and has various medicinal properties. Caesalpinia (Poincianella) pyramidalis Tul. is valued in the caatinga region of northeast Brazil as one of several species used for firewood and charcoal production, and a project is currently being carried out by Kew and the Associaçao Plantas do Nordeste (APNE) to ascertain the best management methods (coppicing or pollarding in the dry or wet season) to produce high quality fuel wood. It would be premature to publish all the new combinations necessary in the reinstated genera, mainly because there is still some doubt as to the correct generic allocation of some species, e.g. Caesalpinia echinata Lam. from Brazil, and 15 Old World taxa, mainly from Asia. These taxa cannot be placed with confidence until a comprehensive molecular phylogeny of Caesalpinia s.l. has been completed. Our observations on wood anatomy provide additional data testing species placement and generic relationships.

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In this paper we use binomials in the segregate genera where these already exist in the literature or Caesalpinia binomials with their associated segregate genus in brackets where new combinations are still required in the segregate. Gasson et al. (2003) found some variation in the wood anatomy of Caesalpinia s.l. in their survey of Caesalpinioideae. They identified several wood characters which can be used for distinguishing between species and genera in the subfamily: storeying of vessels, axial parenchyma and rays, ray size and cell composition, presence or absence of prismatic crystals in ray cells, vestured intervessel pits, intervessel pit size, axial parenchyma patterns, presence or absence of septate fibres, fibre wall thickness, presence or absence of silica bodies and axial canals. The last two characters, silica bodies and axial canals, which are useful diagnostic features in some genera of Caesal- pinioideae are not found in Caesalpinia s.l. This study further develops the analysis of wood characters in Caesalpinia s.l. and shows that some characters support generic segregation, some do not, and some are equivocal.

MATERIALS AND METHODS

Caesalpinia species covering nine recently reinstated segregate genera (Lewis 2005) and two unplaced Old World species were examined in detail, and a brief description of Pomaria is also included. We obtained data on c. 46 species by direct observation (see Table 1 and the Appendix for details of the specimens) and on three additional species from the literature (see Table 1). Specimens were sectioned in their transverse, tangential and radial planes at 20–40 µm thick using a Reichert sliding microtome and stained in 1% Alcian blue and 1% Safranin in 50% ethanol. Sections were then taken through a dehydration process using ethanol, cleared in histoclear and the three planes were mounted onto microscope slides using Euparal. Information on habit (tree, shrub, liana etc.), habitat and major uses is taken from Lewis (2005). The majority of specimens were from branches and therefore juvenile, and the Appendix indicates the radius of each sample and which ones we consider to be mature. The wood anatomical characters, vessel diameter and intervessel pit size classes fol- low the categories given in Wheeler et al. (1989), and Table 1 is in a similar format to that for the Caesalpinioideae in Gasson et al. (2003). The Appendix lists the specimens examined.

WOOD ANATOMY

A general description of the wood anatomy of Caesalpinia s.l. is not given because the genus, as traditionally circumscribed, is polyphyletic. However, a brief summary of similarities and differences amongst the generic segregates of Caesalpinia sensu lato is given below. Following the genus name are the number of species we examined and the total number of species in that genus (e.g. 5/25). Two unassigned Old World species are treated separately. Table 1 provides a summary of the characters we recorded for each genus and Table 2 condenses this information further for ease of reference.

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250 IAWA Journal, Vol. 30 (3), 2009

T = tree, S = shrub, C = climber = C shrub, = S tree, = T Habit:

T/S

S T/S S

RM = marginal ray cells, RP = procumbent ray cells, F = fi bres fi = F cells, ray procumbent = RP cells, ray marginal = RM

prismatic except D = druses, AP = axial parenchyma, axial = AP druses, = D except prismatic Crystals:

Rays homocellular / heterocellular / homocellular Rays

in cells in Ray height Ray

in cells in Ray width Ray

Y = storied, I = irregularly storied, N = not storied not = N storied, irregularly = I storied, = Y Ray storeying: Ray

Y = storied, I = irregularly storied, N = not storied not = N storied, irregularly = I storied, = Y

Axial parenchyma storeying parenchyma Axial

id = idioblasts occasionally, sc = sclerifi ed sclerifi = sc occasionally, idioblasts = id

, no. of cells per strand per cells of no. , Axial parenchyma Axial

Axial parenchyma banded parenchyma Axial

Axial parenchyma conﬂ uent conﬂ parenchyma Axial

V = vasicentric, A = aliform, sc = scanty = sc aliform, = A vasicentric, = V Axial parenchyma: Axial

Yes, No Yes, Septate ﬁ bres: ﬁ Septate

0 = thin, 1 = thick, 2 = very thick very = 2 thick, = 1 thin, = 0 Fibre walls: Fibre

m) (≥10 large = L m), (7–10 medium = M µ µ

m), (4–7 small = S m), (<4 minute = Mt Intervessel pit size: pit Intervessel µ µ

Caesalpinia s.l. reference literature present, = P

in radial multiples radial in Number of vessels of Number

m (IAWA Committee 1989) Committee (IAWA m 200 > = 43 m, 100–200 = 42 µ µ

m, m, 50–100 = 41 m, 50 < = 40 Vessel diameter: Vessel µ µ

R - ring, S - semi-ring, D = diffuse, T = tyloses = T diffuse, = D semi-ring, - S ring, - R Porosity:

Yes, Indistinct, No Indistinct, Yes, Growth rings: Growth – – 2-6 – – – V – – 2 – – 3-4(6) (7)20-25(60) Het R AP, Y Y D 41,42 2-4 M 0/1 N A & V (Y) N 1-4 Y Y 1-2 (3)8-12 Hom (Het) RM, RP) F, (AP, T N D 41,42 2-7 M 0/1 N (V) A sc N N 2-4 ? Y 1-2 (2)8-12 Hom (Het) (F) RM, RP, AP, T Y Y D ‘medium’ 2-5 – 0/1 – V sc – – – – Y 1-2(3) 10-15 Het R AP, I D 41,42 2-5 M 0/1 N A & V Y Y 1-2(3) I N (1-2)3-4 (4)10-25(41) Hom (Het) (RP) F, AP, T

N D 42 2-5 ML 0/1 N (V) A Y Y 2-5 I N 1-2(3) (2)8-40(45) Het RM, RP AP, C/S

Y Y D 41 2-16 SM 0/1 N V sc N N 2-3 ? N 1-2 (2)6-20(43) Het RM (AP), RP, S

N N D N D,T N 42 D 42 Y D 41,42 D,T 2-4 41 2-12 41,42 I 2-3 M – D 2-8 SM 2-5 0-1 1 ML 41 0/1 Mt N 0 N N (V) A 1 2-6 A A Y N N SM V sc (Y) (Y) Y A 1-4 0/1 N N Y I N 1-3 Y 2-4, id N V sc I N 2-4 I Y 1-2(3) N N 2-4 ? N 1 (1)6-25(33) N 1-2 I N ? (sc) Het 1-2 N I? (3)8-22(29) (2)6-10(15) 1 N (2)5-15(20) Hom Hom /(Het)AP RM, RP, 1(2) RM, RP) (AP, Het S/T (2)7-15 T/S (F) AP, (2)10-40 AP Hom Hom T F AP, (AP), RM, RP T Y Y D 41,42,43 2-9 SM 0/1 N A (Y) (Y) 1-3 I N (1)2-3 (3)15-50(70) Het RP) (AP, C/S Y Y R,T 41,42 2-4 S 0-1 N V & A (Y) (Y) 2 I N (1-3)4-6 (1)25-50(60) Het (D, in RP), (RP) S

(Hook.) Link S. Wats. Griseb. (Cozzo 1951) – (Mill.) Standl. Benth. (Britton & Rose) Standl. (Britton Benth. ex Hemsl. s.s. (5/25 spp.) (5/8 spp.) Caesalpinia bahamensis Lam. Willd. Caesalpinia cassioides Caesalpinia madagascariensis (R.Vig.) S. Senesse Caesalpinia nipensis Urb. Caesalpinia pulcherrima (L.) Sw. Caesalpinia gracilis Kunth (Babos & Cumana 1988) Coulteria mollis Caesalpinia platyloba Caesalpinia Coulteria Caesalpinia violacea Caesalpinia velutina Table 1. Wood Wood 1. anatomy Table of the segregate genera of Erythrostemon (3/13 spp.) Guilandina (2/18 spp.) Caesalpinia exilifolia Guilandina bonduc L. Guilandina major (DC.) Small Caesalpinia calycina gilliesii Erythrostemon

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T/C T T

(continued on the next page) (Het) RM, RP F, AP, T

Hom AP

Hom AP Hom RP AP, T D – 2-4 – – – – – – 2-4 Y – 2-4 15-22 – AP I D Y 41 D 41,42 2-3 N 2-7 N D SML D M 2 41,42 N 41,42 1/2 N 2-3 I? D A N 2-3 N? D M D 41,42 A & V SM 42 Y Y 0/1 42 2-3 2 Y Y Y N M 2-6 2-6 2-4 I D A & V N 2-6 Y SM Y D A Y 0/1 Y 41 SM D Y Y N 0-1 Y I 42 0-1? 1-3 Y 2-4 1-2 41,42 A N N D 2-6 N D Y A Y 2-3 2-4 (V) A (6)10-15 S (3)6-12 41,42 2-4 Y Y – Y SM S Y 1-2 Y Y 2-4 Y 1/2 Hom Y Hom 0/1 2-4 N D Y Y 1/2 SM N 1-2 (4) 2-4 (4)8-12 D 1-2(3) N 2-(4) I I A (F) AP, N 42 N 0/1 (F) AP, (V) A S I (2)10-15 41 (V) A I N D N N Y Hom 1-2(3) I Y (1)2-3 N Y D 2-4 1 (V) A T 41 Hom T Y 1-2(3) D 2-3 (2)6-30(40) N (3)10-35(40) Y Y Y SM 2-4 (F) 41 AP, Y (2)10-30(56) 2-4 1-4 Hom (Het) S 41 D Het Y 2-5 AP (A) V 0 Y AP Het 2-5 I I 2-4 Y 41 T S Y 2 Y 2-3 (F) AP, N I I 1-2 SM N A F? & V AP, (1)2-3(4) N 1-2(3) 0/1 SM 2-4 2-3 N (3)10-25(40) (Y) 0/1 A (2)10-20(30) (1)3-4(5) N (3)15-20(25) N C 1 Hom SM Y N (4)15-30(45) Hom C (V) A 2-4(6) Hom A & V Y Y Hom 1/2 N I Y (F) AP, RP, (1)2-3 Y A N AP Y N Y (2)15-25(30) Y (F) AP, T/SA & V 1-2 1-2 2-4 1-3 Y Hom Y Y I (3)6-30 Y (Y) I T I 1-2 (AP) 2 F, N N 1 1-2(3) I 1-3 Hom Y (2)8-25(30) T N (3)5-10(12) Y (2)15-25(100) 1-2(3) AP v.occ Hom (1)2-3(4) Het Hom (3)10-15(42) (3)5-35(41) T/S Hom RP) (F, AP, Het (F) (F) AP, AP, RM, RP, S T/S (F) AP, (AP) RM, RP, S S S Y Y D 41,42 P S 2 N A Y Y 2-4 Y Y 1-3 -15

N D 41,42 2-4 SM 2 N A Y Y 4 I Y D Y 1-2 41,42 2-7 4-15 SM 1 N A I D Y Y 41,42 4 2-4 I M N 0/1 1-3 N A (3)10-25(35) Hom (Y) N F (RP) AP, 1-4 T/S I N (1)2-3 (4)10-20(24) Het? AP RM, RP, F, T I D N – D 41 2-6 N D – 2-3 42 SM 2 2 2 Y N A SM A & V Y Y 2 Y Y N 2-4 – A Y Y Y Y 1-3(4) – Y 1(-2) Y (6)10-20(25) 2-4, sc I D Y Hom 8-12 N D Y 41,42 D 1-3 F AP, 41,42 2-3 40,41 (5)10-15(20) 2-4 S 2-3 Hom ML SM 0/1 0/1 AP N 2 Y V V N A & V Y Y Y Y Y Y 2-4 2-7 2-4 I I I N N N (1)2-3 (1)2-3 (1)3-4(5) (4)12-30(62) (2)8-20(30) (3)8-25(30) Hom Het Hom (RM, RP) AP, RM, RP F, RM, RP AP, F, AP, T T T/S )

C. granadillo ) C. paucijuga C.

Oliv.

G.P. Lewis G.P.

* (Rose) Britton & Rose Vatke (Benth.) Britton & Rose (DC.) Britton & Rose (Britton) Britton & Rose (Standl.) Britton & Rose (A. Gray) Britton & Rose (Brandegee) Britton & Rose (Greenman) Britton & Rose G.P. Lewis G.P. Kunth ( paipai ) Lam.

G.P. Lewis G.P. DC. var . peltophoroides (S. Wats.) Britton & Rose Wats.) (S. (Brandegee) Britton & Rose S. Sotoyo & G.P. Lewis S. Sotoyo & G.P. (Willd.) Britton (Willd.) Britton (as (Willd.) Britton (as (Willd.) (Jacq.) Schltdl.

(3/26 spp.) (19/35 spp.) (7/8 spp.) Libidibia Libidibia coriaria Libidibia corymbosa (Benth.) Britton & Killip (Latorre 1983) Mart. Caesalpinia ferrea Caesalpinia glabrata Lewis Libidibia paraguariensis (D. Parodi) G.P. Libidibia punctata (as L. ebano in Pernia & Melandri 2006) Libidibia punctata Libidibia punctata Britton & Rose Libidibia sclerocarpa Mezoneuron hildebrandtii Mezoneuron Voigt Arn. ex & sumatranum Wight Mezoneuron welwitschianum Mezoneuron Poincianella Poincianella caladenia Caesalpinia coccinea Caesalpinia echinata Poincianella eriostachys Poincianella exostemma subsp. exostemma Poincianella gaumeri Caesalpinia oyamae Caesalpinia hughesii Poincianella melanadenia Poincianella mexicana Poincianella myabensis Caesalpinia nicaraguensis Poincianella palmeri Poincianella pannosa Poincianella placida Caesalpinia pluviosa Lewis paraensis (Ducke) G.P. var. Caesalpinia pluviosa (Schmid 1915) –

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252 IAWA Journal, Vol. 30 (3), 2009

T = tree, S = shrub, C = climber = C shrub, = S tree, = T Habit:

S/C

T/S

RM = marginal ray cells, RP = procumbent ray cells, F = fi bres fi = F cells, ray procumbent = RP cells, ray marginal = RM

prismatic except D = druses, AP = axial parenchyma, axial = AP druses, = D except prismatic Crystals:

Rays homocellular / heterocellular / homocellular Rays

in cells in Ray height Ray

in cells in Ray width Ray

Y = storied, I = irregularly storied, N = not storied not = N storied, irregularly = I storied, = Y Ray storeying: Ray

Y = storied, I = irregularly storied, N = not storied not = N storied, irregularly = I storied, = Y

Axial parenchyma storeying parenchyma Axial

id = idioblasts occasionally, sc = sclerifi ed sclerifi = sc occasionally, idioblasts = id

, no. of cells per strand per cells of no. , Axial parenchyma Axial

Axial parenchyma banded parenchyma Axial

Axial parenchyma conﬂ uent conﬂ parenchyma Axial

V = vasicentric, A = aliform, sc = scanty = sc aliform, = A vasicentric, = V Axial parenchyma: Axial

Yes, No Yes, Septate ﬁ bres: ﬁ Septate

0 = thin, 1 = thick, 2 = very thick very = 2 thick, = 1 thin, = 0 Fibre walls: Fibre

m) (≥10 large = L m), (7–10 medium = M µ µ

m), (4–7 small = S m), (<4 minute = Mt Intervessel pit size: pit Intervessel µ µ

P = present, literature reference literature present, = P

in radial multiples radial in Number of vessels of Number

m (IAWA Committee 1989) Committee (IAWA m 200 > = 43 m, 100–200 = 42 µ µ

m, m, 50–100 = 41 m, 50 < = 40 Vessel diameter: Vessel µ µ

R - ring, S - semi-ring, D = diffuse, T = tyloses = T diffuse, = D semi-ring, - S ring, - R Porosity:

Yes, Indistinct, No Indistinct, Yes, Growth rings: Growth I N D D 41,42 41,42 2-4 Y 2-3 D SM SM 41,42,43 0 2-7 2 N SM N (A) V A 0/1 Y N N Y (A) V 2-4 Y N 2-4 I Y 2-4 N I (1)2-3 I N (2)6-25 1-2 N (1)2-4 (2)6-15(20) Het (5)15-50 Het (AP) RM, RP, Hom T/S RM, RP AP, F AP, T Y Y D N 41,42 D 2-9 41,42 M 2-5 0/1 SM N 0/1 A N A Y N (Y) (1)-4 (Y) I 1-4 N I 1-2 N (1)2(3) (2)15-50(55) (2)10-25(35) Hom Hom/(Het) (F) AP, AP (RM, RP), F, S/C T Y Y D 41 2-6 SM 2 N Y A D Y 42,43 Y 2-6 2 SML Y 0 Y N (1)2-3 V (3)6-10(43) N Hom N F AP, 2-4 I N 1-5(6) (2)8-30(40) Hom/(Het) (RP) AP, S/C N D 42 2-3 – 1/2 N A & V Y (Y) 4 I N 1-2(3) (2)8-25(30) Het F) (AP, RM, RP, T N D 41 2-4 SM 0/1 N A Y Y 1-2 (4) I N (1)2-3 (3)7-40(44) Hom (RP) AP, S

– D – 2-5 – – – V – – 2 – – 3-4(6) (7)20-25(60) Het R AP, N D 41 2-4 0/1 N A & V Y Y 2-4 I N (1)2-3 (3)10-45 Hom (F) AP, T/S ose R I D 42 2-3(4-7) SM 0/1 N A & V Y Y 2-5 I N (1)2-3(4) (3)9-30(48) Hom (RM, RP) F, AP, ST ritton &

) B in Poincianella may be incorrect. ) reenman (G subsp . chiapensis subsp . hondurensis I I D D 42 42 2-5 2-4(5) SM SM 0/1 0/1 (Y) (Y) A & V A & V Y Y Y Y 2-4 1-4 I I N N (1)3-4(5) (1)2-3(5) (3)10-30(47) (3)15-35(46) Hom Hom (RM, F) RP, AP, RM, (AP) RP, T T subsp. yucatanensis Tul. Tul. (= japonica (= sepiaria ) Alston (Roth) Britton & Rose L. Humb. & Bonpl.

Caesalpinia echinata

(Table 1 (Table continued) Caesalpinia vesicaria Caesalpinia yucatanensis Caesalpinia yucatanensis Caesalpinia yucatanensis Caesalpinia decapetala Caesalpinia sappan L. Caesalpinia decapetala Caesalpinia pyramidalis (3/3 Tara spp.) Caesalpinia cacalaco species unassigned World Old Pomaria (1/16) Tara spinosa (Molina) Britton & Rose Tara Pomaria rubicunda (Vogel) B.B. Simpson & Pomaria rubicunda (Vogel) Lewis (Cozzo 1951 as C. rubicunda (Vogel) G.P. Bentham) * Placement of Poincianella standleyi Poincianella yucatanensis Caesalpinia decapetala

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Similarities and differences in Caesalpinia s.l. All species have simple perforation plates, vestured alternate intervascular and vessel-ray pitting and paratracheal parenchyma, as do most Caesalpinioideae and legumes in general. Growth rings and their definition are variable. The wood is usually diffuse porous with the exception of Erythrostemon gilliesii which is ring porous, but this was a cultivated specimen and we cannot be sure its porosity reflects the situation in its natural range (Fig. 17). Most species have narrow to medium-sized vessels, in a few species just exceeding 200 µm in tangential diameter, e.g. Guilandina major (DC.) Small (Fig. 19), Guilandina bonduc L. (Fig. 21), Caesalpinia decapetala (Roth) Alston (Fig. 68) and C. sappan L. (Fig. 73). Vessels are sometimes in radial multiples and occasionally in clusters, and are frequently filled with a gum-like substance which stains red with safranin. Most species have non-septate fibres that are thin- to thick-walled. A few species of Poincianella have some septate fibres (see Table 1). They have also been reported in Libidibia corymbosa (Benth.) Britton & Killip by Latorre (1983). Prismatic crystals sometimes appear to occur in chambered fibres, but are not as common as crystals in chambered axial parenchyma cells and ray cells, and it is likely that the “chambered fibres” are actually axial parenchyma in a matrix of fibres. All species have paratracheal axial parenchyma, which is in a vasicentric, aliform and/or confluent pattern. A few species have indistinct paratracheal parenchyma where there is not much difference between the wall thickness of the parenchyma and adjacent fibres, but most have confluent parenchyma in narrow or wide bands. Most species have irregularly or non-storied parenchyma, apart from all Libidibia and some species of Coulteria and Poincianella which have storied parenchyma. Prismatic crystals are found in chambered axial parenchyma cells of all species except Erythrostemon gilliesii (Hook.) Link. Enlarged parenchyma cells (idioblasts) resembling oil cells are found in Caesalpinia madagascariensis (R.Vig.) S. Senesse (currently considered as a member of Caesalpinia s.s.), but their content is unknown. Rays vary from uniseriate, as in C. madagascariensis (Fig. 4), to up to six cells wide in Erythrostemon gilliesii (Fig. 18). Ray height also varies and can be up to 100 cells high (Poincianella pannosa (Brandegee) Britton & Rose, Fig. 56). The shortest rays are usually storied. Most species have non-storied rays, but in all Libidibia and some species of Coulteria and Poincianella the rays are storied. Rays are either homocellular, as in all species of Libidibia and Coulteria, or heterocellular as in all Mezoneuron, Tara and Erythrostemon species. Ray cellular composition varies within the remaining genera – Poincianella, Caesalpinia s.s. and Guilandina. Prismatic crystals in non-chambered ray cells are found in many species, and some species also have a few prismatic crystals in chambered ray cells.

Caesalpinia s.s. (5/25 species, Fig. 1–7) Caribbean, Mexico, Colombia, Ecuador, Peru, Paraguay, Argentina, Madagascar, Africa. Trees and shrubs from seasonally dry tropical woodland, wooded grassland, coastal thicket, bushland and thorn scrub, dry plains and riparian woodland, many species on limestone or sandstone. Caesalpinia sensu stricto is essentially confined

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Figures 1–7. Caesalpinia s.s. – 1 & 2. Caesalpinia cassioides. – 1. TS. Vessels in long radial multiples, axial parenchyma scanty paratracheal. – 2. TLS. Tyloses in vessels, rays not storied and crystals present in axial parenchyma. – 3–5. C. madagascariensis. – 3. TS. Sparsely distributed vessels, axial parenchyma in narrow bands. – 4. TLS. Rays short and narrow, not storied, some idioblasts. – 5. RLS. Homocellular rays and idioblasts. – 6 & 7. C. pulcherrima. – 6. TS. Distinct growth ring and mainly indistinct scanty paratracheal parenchyma. – 7. TLS. Rays narrow and not storied.

Downloaded from Brill.com10/11/2021 11:10:06AM via free access Gasson, Warner & Lewis — Wood anatomy of Caesalpinia sensu lato 255 to the succulent biome of Schrire et al. (2005) but it is possible that the taxon is not monophyletic. Recent unpublished molecular phylogenies of Sotoyo place the species of Caesalpinia s.s. (sensu Lewis) in at least three unrelated clades. The best known species in Caesalpinia s.s. is the widely cultivated ornamental C. pulcherrima which has various medicinal properties (Lewis 2005). The five species of Caesalpinia s.s. we examined were Caesalpinia bahamensis Lam., C. cassioides Willd. (Fig. 1 & 2), C. madagascariensis (R. Vig.) S. Senesse (Fig. 3–5), C. nipensis Urb. and C. pulcherrima (L.) Sw. (Fig. 6 & 7). The limited literature on wood anatomy includes Cozzo (1951) and Luo (1989). We have poor coverage of Caesalpinia s.s. All the species examined have non- storied rays and axial parenchyma. Caesalpinia cassioides has very long radial vessel multiples (Fig. 1) and an indistinct paratracheal axial parenchyma pattern (similar to C. (Erythrostemon) calycina, Fig. 14). Caesalpinia madagascariensis differs from all other Caesalpinia s.l. species in having idioblasts in the wood which look very like the

Figures 8–13. Coulteria. Caesalpinia (Coulteria) velutina. – 8. TS. Axial parenchyma scanty paratracheal and indistinctly aliform and confluent. – 9. TLS. Rays storied.Caesalpinia (Coul- teria) platyloba. – 10. TS. Axial parenchyma in wide bands with one indistinct narrow marginal band. – 11 & 12. TLS. – 11. Rays not storied. – 12. Crystals in chambered cells among fibres, probably in axial parenchyma. – 13. RLS. Heterocellular ray with some upright ray cells containing crystals.

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Coulteria (5/8 species, Fig. 8–13) Mexico and Central America, Cuba, Jamaica, Curaçao, Venezuela and Colombia. Trees and shrubs in seasonally dry tropical forest, deciduous woodland and dry thorn scrub, some species on limestone. We examined four species of Coulteria: Caesalpinia (Coulteria) gracilis Benth. ex Hemsl., Caesalpinia (Coulteria) velutina (Britton & Rose) Standl. (Fig. 8–10), Caesalpinia (Coulteria) violacea (Mill.) Standl. and Caesalpinia (Coulteria) platyloba S. Wats. (Fig. 11–13). Babos and Cumana (1988) described the wood of the type species, Coulteria mollis Kunth. Coulteria is locally used as a preferred firewood, and some species are grown as ornamentals in living fence lines (Lewis 2005).

Figures 14–18. Erythrostemon. Caesalpinia (Erythrostemon) calycina. – 14. TS. Axial parenchyma scanty paratracheal and indistinctly vasicentric, most vessels in radial multiples. – 15. TLS. Axial parenchyma and rays not storied. – 16. RLS. Crystals present in heterocellular rays and occasionally in axial parenchyma. – 17 & 18. Erythrostemon gilliesii. – 17. TS. Ring porous. – 18. TLS. Rays very wide and quite tall.

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The wood anatomy of Coulteria is variable. The only consistent features were the presence of prismatic crystals in ray cells and crystals in chambered axial parenchyma in all the species we examined. Some species have storied rays (Caesalpinia (Coulteria) velutina, Fig. 9, Caesalpinia (Coulteria) violacea and Coulteria mollis) while others do not (Caesalpinia (Coulteria) gracilis and Caesalpinia (Coulteria) platyloba, Fig. 11). All the species in Coulteria, except for Caesalpinia (Coulteria) gracilis, show a tendency towards heterocellular rays. Chambered cells containing crystals are present in Caesalpinia (Coulteria) platyloba (Fig. 12), but it is unclear whether they are fibres or diffuse axial parenchyma cells.

Erythrostemon (3/13 species, Fig. 14–18) South America, southern USA and Mexico. Shrubs and woody-based perennial herbs in seasonally dry tropical and subtropical semi-arid thorn scrub (including caatinga), spiny cactus scrub, woodland and grassland. Erythrostemon gilliesii is a widely cultivated garden ornamental and is used in revegetation projects (Lewis 2005). This description is based on only three species, and therefore may not fully reflect the range of wood features in the genus. We examined Caesalpinia (Erythrostemon) calycina Benth. (Fig. 14–16) and E. gilliesii (Hook.) Link (Fig. 17 & 18). Cozzo (1951) described the wood anatomy of Caesalpinia (Erythrostemon) exilifolia Griseb. and E. gilliesii. Caesalpinia (Erythrostemon) calycina has vessels in long radial multiples and unlike E. gilliesii and most species in Caesalpinia s.l., has ill-defined paratracheal parenchyma (Fig. 14). This specimen has a very small diameter and consists entirely of juvenile wood. All three species have heterocellular rays containing crystals. Caesalpinia (E.) calycina has narrow rays (Fig. 15) and, like E. gilliesii which has wide rays (Fig. 18), the rays are non-storied. Cozzo (1951) did not mention storeying in C. (E.) exilifolia. Cozzo (1951) also reported fibre tracheids and vascular tracheids in E. gilliesii, and vascular and vasicentric tracheids in C. (E.) exilifolia. Erythrostemon gilliesii was the only species in Caesalpinia s.l. we examined with ring porous wood (Fig. 17). It is also the only species in Caesalpinia s.l. where we could not find any crystals in the axial parenchyma. Druses were found in ray cells in the bark but not in the adjacent wood, and were not recorded in any other Caesalpinia s.l. species. Erythrostemon is essentially a west South American genus, with one species, C. (E.) calycina from eastern Brazil. Better sampling of the genus is needed, and as currently defined it might prove not to be monophyletic.

Guilandina (2/18 species, Fig. 19–22) Pantropical from as far north as Japan and south to South Africa, 3 in the Caribbean, 1 in China, India, Myanmar, Indo-China, Hong Kong and Taiwan, 1 endemic to Mada- gascar, 1 in Australia and 2 widespread across the Old and New World tropics. There are possibly only 7 species in total, with many published binomials being synonyms (Lewis 2005). Scrambling prickly vines or scandent shrubs, coastal sands and thicket, secondary forest, lowland rainforest, some on limestone. Lewis (2005) gives various uses for the seeds and leaves of Guilandina, but none for the wood.

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Figures 19–22. Guilandina. – 19 & 20. Guilandina major. – 19. TS. Vessels of two distinct diameters. Abundant scanty paratracheal to aliform and diffuse-in-aggregates parenchyma. – 20. TLS. Rays tall and non-storied. – 21 & 22. Guilandina bonduc. – 21. TS. Indistinct growth ring boundary marked by difference in vessel diameter, vessels of two distinct diameters. Axial parenchyma much less frequent than in Fig. 19. – 22. RLS. Rays heterocellular, some ray cells are chambered and contain crystals.

Coverage of this segregate genus was very poor with only two species examined, Guilandina major (DC.) Small (Fig. 19 & 20) and Guilandina bonduc L. (Fig. 21 & 22). We found no additional information on the wood of this genus in the literature. Both species have wide vessels in radial multiples with much narrower ones in the same multiple. The rays and axial parenchyma are not storied. The heterocellular rays contain prismatic crystals in procumbent, square and upright cells, often in radial align- ment, and in G. bonduc some of the upright ray cells containing crystals are chambered (Fig. 22). The rays are 1–3 cells wide in both species (Fig. 20), vary considerably in height, but tend to be much taller in G. bonduc (see Table 1).

Libidibia (7/8 species, Fig. 23–30) Neotropical trees or shrubs in seasonally dry forest and scrub, thorn forest (including caatinga) and savanna woodland.

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Figures 23–30. Libidibia. L. paraguariensis. – 23. TS. Axial parenchyma scanty paratracheal to partially banded. – 24. TLS. Axial parenchyma and rays storied. – 25. RLS. Rays homocellular, crystals in chambered axial parenchyma cells. Caesalpinia (Libidibia) glabrata. – 26. TS. Axial parenchyma scanty paratracheal to banded, one indistinct growth ring, some vessels filled with gum or resin. – 27. TLS. Rays and axial parenchyma storied, many rays axially fused. – 28. RLS. Rays homocellular. – 29 & 30. Caesalpinia (Libidibia) ferrea. TS. – 29. Indistinct growth ring and aliform and confluent parenchyma forming incomplete narrow bands (slide 19792). – 30. Growth rings not seen and aliform and confluent parenchyma forming wider bands than in Fig. 29 (slide from Lewis 1619).

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We examined seven species of Libidibia: L. coriaria (Jacq.) Schltdl., Caesalpinia (Libidibia) ferrea Mart. (Fig. 29 & 30), Caesalpinia (Libidibia) glabrata Kunth (sy- nonym Caesalpinia paipai Ruiz & Pavon) (Fig. 26–28), Libidibia granadillo (Pittier) Pittier (probably = Libidibia punctata (Willd.) Britton), Libidibia paraguariensis (D. Parodi) G.P. Lewis (Fig. 23–25), Libidibia punctata (as Caesalpinia (Libidibia) paucijuga Benth. ex Hook.) and Libidibia sclerocarpa Britton & Rose (as Caesalpinia sclerocarpa Standl.). The wood of the eighth species, L. corymbosa (Benth.) Britton & Killip (probably = Caesalpinia (L.) glabrata Kunth), was described by Latorre (1983). The wood of Libidibia has several uses (Lewis 2005): it is prized in turnery and for parts of guitars and violins; Caesalpinia (Libidibia) glabrata and Libidibia paraguariensis (both known as partridge wood) are used in decorative inlay and cabinet work, some species are used in heavy construction (railway sleepers, beams, bridge sup- ports), for tool handles and firewood. SomeLibidibia species are also planted as ornamentals. There are wood descriptions and/or photomicrographs of the wood of Libidibia, usually as Caesalpinia species, in Schmid (1915), Record and Mell (1924), Williams (1939), Cozzo and Cristiani (1950), Cozzo (1951), Tortorelli (1956), Kribs (1959), Paula (1980, 1981), Mainieri et al. (1983), Luo (1989), Ilic (1991), Angyalossy et al. (2005), Espinoza de Pernía and Melandri (2006), and InsideWood (2004–onwards). Libidibia is well defined on the basis of wood anatomy. All species have short storied rays, storied axial parenchyma, and homocellular rays. Prismatic crystals were not found in any ray cells. Most species lack growth rings apart from Caesalpinia (Libidibia) ferrea (Fig. 29), which has indistinct growth rings and Libidibia paraguariensis, which has marginal lines of parenchyma. The sample labelled Libidibia granadillo (probably = L. punctata) was the only species of Caesalpinia s.l. examined with sclerified axial parenchyma.

Mezoneuron (3 / 26 species, Fig. 31–39) Mainly Asia, also Australia, Polynesia, Madagascar and Africa, with fossil records in Europe and North America where the genus no longer occurs. Prickly climbers, scandent shrubs and small trees, tropical and subtropical riverine forest, lowland rain forest, swamp forest, seasonally dry forest, thicket, vine forest and wooded grassland, especially along forest and river margins. Few uses of the wood have been reported, but Mezoneuron kauaiense Hildebr. from Hawaii was once used locally for spears and in house construction (Lewis 2005). The three species of Mezoneuron we examined were M. welwitschianum Oliv. (Fig. 31–33), M. sumatranum Wight & Arn. ex Voigt (Fig. 34–36) and M. hildebrandtii Vatke (Fig. 37–39). No literature was found on wood of this genus. These three species lack storeying and crystals are absent in ray cells. Both M. welwitschianum (Fig. 33) and M. sumatranum (Fig. 36) have heterocellular rays and M. hildebrandtii shows a tendency towards heterocellular rays (Fig. 39). Mezo- neuron sumatranum also differs slightly in having aliform, confluent and banded axial parenchyma (Fig. 34) whereas in the other two species the bands tend to be wider (Fig. 31 & 37).

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Figures 31–39. Mezoneuron. – 31–33. M. welwitschianum. – 34–36. M. sumatranum. – 37–39. M. hildebrandtii. – 31, 34, 37. TS. Axial parenchyma aliform, confluent and banded, bands becoming wider towards the bark in Fig. 34. – 32, 35, 38. TLS. Rays not storied. – 33, 36, 39. RLS. Heterocellular rays.

Poincianella (19/35 species, Fig. 40–57) Neotropics, North America and Caribbean south to northern Argentina. Trees and shrubs from seasonally dry tropical and subtropical woodland, arid thorn and cactus

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Figures 40–43. Caesalpinia (?Poincianella) echinata. – Fig. 40 & 41. TS. Axial parenchyma scanty paratracheal to confluent. The amount of parenchyma varies between specimens. Fig. 40 (slide 19784) from a violin bow has more parenchyma than Fig. 41 (slide 19786). – 42. TLS. Rays storied (left side) and irregularly storied (right side). – 43. RLS. Rays homocellular containing crystals. The number of crystals varies between specimens. scrub (including caatinga), beach thicket (including arboreal restinga) and coastal dunes, sandy washes and plains, sandy soils or limestones, less frequent in wooded grassland (cerrado), gallery forest and moist coastal forest, one species in mixed pine-oak forest, many on highly degraded sites. We examined 19 species of Poincianella sensu Lewis (2005). These were P. caladenia (Standl.) Britton & Rose, Caesalpinia (Poincianella) coccinea G.P. Lewis, C. (?Poincianella) echinata Lam. (Fig. 40–43), P. eriostachys (Benth.) Britton & Rose (Fig. 47–50), P. exostemma (DC.) Britton & Rose (subsp. exostemma), P. gaumeri (Greenman) Britton & Rose, C. (Poincianella ) oyamae S. Sotoyo & G.P. Lewis, C. (P.) hughesii G.P. Lewis, P. melanadenia (Rose) Britton & Rose, P. mexicana (A. Gray) Britton & Rose, P. myabensis (Britton) Britton & Rose, C. (P.) nicaraguensis G.P. Lewis, P. palmeri (S. Wats.) Britton & Rose, P. pannosa (Brandegee) Britton & Rose (Fig. 54–57), P. placida (Brandegee) Britton & Rose, C. (?Poincianella) pluviosa DC. var. paraensis (Ducke) G.P. Lewis (Fig. 44–46), C. (P.) pyramidalis Tul. (Fig. 51–53),

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Figures 44–50. Poincianella. – 44–46. Caesalpinia (Poincianella) pluviosa var. paraensis – 44. TS. Axial parenchyma confluent, in wide wavy paratracheal bands. – 45. TLS. Rays and axial parenchyma storied. – 46. RLS. Rays homocellular, crystals in chambered axial parenchyma cells. – 47–50. Poincianella eriostachys. – 47. TS. Axial parenchyma aliform, confluent, vasicentric and in wide wavy bands. – 48 & 49. TLS. Rays and axial parenchyma storied, some rays axially fused. Crystals in chambered axial parenchyma and fibres more abundant in some areas of the specimen (Fig. 48) than others (Fig. 49). – 50. RLS. Rays homocellular.

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Figures 51–57. Poincianella. – 51–53. Caesalpinia (Poincianella) pyramidalis. – 51. TS. Axial parenchyma aliform and confluent, forming wavy bands. Vessels commonly in radial multiples. – 52. TLS. Rays and axial parenchyma storied and crystals in axial parenchyma. – 53. RLS. Rays homocellular and some vessels containing gum. – 54–57. Poincianella pannosa. – 54 & 55. TS. Growth rings wide and vessels sparsely distributed in Fig. 54, much narrower and vessels densely distributed in Fig. 55. Axial parenchyma mainly scanty paratracheal. – 56. TLS. Rays tall (up to 100 cells high), commonly 1–2 cells wide, but in places much wider. – 57. RLS. Rays heterocellular, some ray cells containing crystals.

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Figures 58–67. Tara. – 58–61. Caesalpinia (Tara) vesicaria. – 58. TS. Axial parenchyma scanty paratracheal to confluent near the pith. – 59. TS. Axial parenchyma abundant, confluent and banded, further from the pith than in Fig. 58 (same scale). – 60. TLS. Rays not storied. – 61. RLS. Ray heterocellular with abundant crystals in ray cells. – 62–64. Caesalpinia (Tara) cacalaco. – 62. TS. Growth ring boundary distinct, vasicentric to confluent parenchyma. – 63. TLS. Rays not storied. – 64. RLS. Heterocellular ray. – 65–67. Tara spinosa. – 65. TS. Axial parenchyma pattern is less distinct than in the other two members of Tara. – 66. TLS. Rays not storied. – 67. RLS. Heterocellular ray.

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P. standleyi Britton & Rose, P. yucatanensis (Greenman) Britton & Rose, P. yucatanensis (subsp. chiapensis), P. yucatanensis (subsp. hondurensis) , P. yucatanensis (subsp. yucatanensis). Some species of Poincianella have local medicinal uses. Caesalpinia echinata was once the source of a red dye and is now the only wood used for high quality violin bows, and C. (Poincianella) pyramidalis from the caatinga of northeast Brazil is used for fuelwood and charcoal (Figueirôa et al. 2006). The literature includes Schmid (1915, on C. peltophoroides Benth. (= C. (Poincia- nella) pluviosa DC. var. peltophoroides (Benth.) G.P. Lewis), Record and Mell (1924, on C. (P.) echinata), Paula (1980, on C. (P.) pyramidalis) and Mainieri et al. (1983, on C. (P.) echinata). Richter (1988) gives the history, timber properties and a wood description of C. echinata (as Guilandina echinata). More recently, De Lima, Lewis and Bueno (2002) provide a history of C. echinata and its uses in Brazil, and its wood anatomy and use in musical instruments are discussed in Angyalossy et al. (2005) and Amano (2007). Poincianella is the largest and most diverse wood anatomically of the nine segregate genera. Most Poincianella species have non-storied rays and irregularly storied axial parenchyma, apart from P. gaumeri, P. eriostachys (Fig. 48–49), C. (Poincianella) pluviosa var. paraensis (Fig. 45), P. myabensis and C. (P.) pyramidalis (Fig. 52). The placement of P. eriostachys (Fig. 47–50) and C. (Poincianella) pluviosa (Fig. 44–46) in Poincianella needs to be reassessed in light of recent molecular data which place them as sister species, but not clustered with other Poincianella species. Their wood anatomy is very similar. In the C. (?Poincianella) echinata specimens we examined the rays were either storied, irregularly storied or both (see Fig. 42), an observation also made by Angyalossy et al. (2005) and Amano (2007). Caesalpinia echinata is most probably misplaced in Poincianella. Most Poincianella species have crystals in ray cells and homocellular rays. A number of Poincianella species have septate fibres (see Table 1), an anatomical feature otherwise apparently found within the Caesalpinia s.l. complex only in Caesalpinia (Coulteria) platyloba. Prismatic crystals are found in chambered axial parenchyma in all species, as they are in most other legumes.

Tara (3/3 species, Fig. 58–67) Mexico, Caribbean and South America, widely cultivated including Canary Islands, Malta, India and Africa. Trees and shrubs in seasonally dry tropical forest to semi-arid thorn scrub. Tara species are often planted as living fence lines, and Tara spinosa is used for firewood and gums, tanning leather, dyeing and ink production in Peru (Lewis 2005). We examined the wood of all three species of Tara. These were Caesalpinia (Tara) vesicaria L. (Fig. 58–61), C. (T.) cacalaco Humb. & Bonpl. (Fig. 62–64), and Tara spinosa (Molina) Britton & Rose (Fig. 65–67). There is a Tara spinosa description in Détienne & Jacquet (1983). Tara, like Libidibia, is well defined by its wood anatomy. All three species have non- storied, mainly heterocellular rays and axial parenchyma and have indistinct growth rings. Tara spinosa differs from the other two species by having a less well defined axial parenchyma pattern and thin walled fibres (Fig. 65).

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Figures 68–75. Old World unassigned species. – 68–70. Caesalpinia decapetala ( Hort. Kew 1952). – 68. TS. Semi-ring porous. – 69. TLS. Rays 1–5 cells wide and tall. – 70. RLS. Rays tending towards heterocellular. – 71 & 72. Caesalpinia decapetala (J. Prior 287, Swaziland). – 71. TLS. Rays 1–2 cells wide (narrower than in Fig. 69) and very tall. – 72. RLS. Rays homocellular. – 73–75. Caesalpinia sappan. – 73. TS. Distinct growth ring boundary, semi-ring porous, axial parenchyma marginal and confluent. – 74. TLS. Rays and parenchyma irregularly storied. – 75. RLS. Ray homocellular containing crystals.

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Unassigned Old World species (Caesalpinia decapetala (Roth) Alston and C. sappan L., Fig. 68 –75) Approximately 15 Asian taxa have not been reassigned to segregate genera. The two species we examined were Caesalpinia decapetala (Fig. 68–72) and C. sappan (Fig. 73–75). Kanehira (1921), Ramesh Rao and Purkayastha (1972) and Tang (1973) have described the wood of C. sappan. Caesalpinia decapetala is included in the FFPRI key with images and IAWA codes for descriptions (http://f030091.ffpri. affrc.go.jp/index-E3.html). Lemmens and Wulijarni-Soetjipto (1991) outline the use of C. sappan wood for production of a red dye, but only describe some macroscopic features of the wood. There are also descriptions of the perforation plates and spiral thickenings in C. japonica Siebold & Zucc. (= C. decapetala) by Ohtani and Ishida (1977 & 1978).

Caesalpinia decapetala Specimen HK 1952 of C. decapetala (labelled C. japonica) is similar to Erythro- stemon (Fig. 17 & 18), especially on the basis of ray size. It resembles Erythrostemon gilliesii (Fig. 17) in having a tendency towards vessels of two distinct sizes (Fig. 68) and non-storied, wide and tall rays (Fig. 69) which are heterocellular (Fig. 70). How- ever, the other two specimens of C. decapetala have homocellular rays (Fig. 72), which are narrower and taller (Fig. 71) than E. gilliesii and specimen HK 1952.

Caesalpinia sappan Caesalpinia sappan has non-storied rays and irregularly storied parenchyma (Fig. 74), an absence of crystals in ray cells and small to medium-sized vessels (Fig. 73). Caesalpinia sappan does not clearly belong to any of the currently recognised segregate genera of Caesalpinia s.l. Caesalpinia sappan was historically used for dye, in much the same way as Caes- alpinia (?Poincianella) echinata, but apparently no timber uses have been reported.

Pomaria (2/16). Not illustrated Pomaria species are shrubs and perennial herbs and barely woody, mainly from subtropical dry areas of grassland and wooded grassland and in degraded sites, many on limestone. We examined some TS and RLS sections from a 3 mm diameter, 3 year old stem of Pomaria austrotexana B.B. Simpson. Cozzo (1951) gave a brief wood description of P. rubicunda (Vogel) B.B. Simpson & G.P. Lewis (as Caesalpinia rubicunda (Vogel) Benth.), which had heterocellular rays and calcium oxalate crystals in the axial parenchyma. In P. austrotexana we found well-defined growth rings, vessels in radial multiples and clusters, scanty paratracheal and initial parenchyma, very thick- walled fibres, many gelatinous, rays 1–4 cells wide and tall in RLS so presumably not storied, and some ray cells containing a single prismatic crystal. The limited juvenile material of Pomaria available and the brevity of the literature do not allow us to make sensible comparisons with the other genera.

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DISCUSSION

All Caesalpinia s.l. woods have vestured pits, paratracheal parenchyma, simple perforation plates, and alternate intervascular and vessel-ray pitting. Prismatic crystals in chambered axial parenchyma cells are found in all species apart from Erythrostemon gilliesii. The only wood anatomical characters that help to separate the genera within Caesalpinia s.l. are the presence or absence of ray and axial parenchyma storeying, ray cell composition (i.e. homocellular versus heterocellular), and the presence or absence of prismatic crystals in ray cells (Table 2). This is a small number of characters on which to base taxonomic circumscription. All the species in Libidibia and Coulteria have homocellular rays and all the species in Tara, Guilandina and Mezoneuron have heterocellular rays. Poincianella and Caes-

Table 2. Summary of wood characters in the segregate genera of Caesalpinia s.l. Growth rings distinct Growth rings indistinct Axial parenchyma storied Rays storied Axial parenchyma irregularly storied Rays irregularly storied Rays homocellular Rays heterocellular Septate fibres Crystals in axial parenchyma Crystals in fibres Crystals in rays

Caesalpinia s.s. 1/5 4/5 N N 4/5 N 3/5 3/5* N Y 2/5 2/5 (5 /25 spp) Coulteria 2/5 2/5 1/5 3/5 2/5 N 4/5 (4)/5 1/5 Y 3/5 Y (5/8 spp) Erythrostemon 2/3 N? N N 1/3 N N Y N 2/3 N Y (3/13 spp) (1 RP) Guilandina 1/2 1/2 N None Y N N Y N Y N Y (2/18 spp) Libidibia 2/7 2/7 Y Y N N Y N 1?/7 Y 4/7 1?/7 (7/8 spp) Mezoneuron N 1/3 N N Y N 1/3 2/3 N Y 1/3 N (3/26 spp) Poincianella 6/19 8/19 5/19 4/19 14/19 2/19 15/19 5/19* 5/19 Y 14/19 12/19 (19/35 spp) Tara N 1/3 N N Y N N Y N Y 1/3 Y (3/3 spp) Old World 1/2 N N N Y N Y (Y)* N Y Y 1/2 (2 spp) Pomaria 1 ? ? ? ? ? ? 1 ? 1 ? 1 (2/16 spp)

Where numbers of species quoted in this table are lower than those in Table 1 and the Appendix, the character in question could not be assessed in all species.

* Some specimens from a given species showed this character while others did not. Y = yes (present in all), N = no (absent from all), ( ) = equivocal or poorly shown.

Downloaded from Brill.com10/11/2021 11:10:06AM via free access 270 IAWA Journal, Vol. 30 (3), 2009 alpinia s.s., as currently circumscribed, have both, but it is well known that hardwoods in many groups tend to have rays that become less heterocellular, i.e. more homocellular with maturity (increasing cambial age), and most of our samples are from relatively narrow stems and branches rather than mature wood (see Appendix). However, heterocellular rays occur in many more genera of Caesalpinioideae than in Papilionoideae, and are very uncommon in Mimosoideae (Baretta-Kuipers 1981; Gasson et al. 2003; Evans et al. 2006). Libidibia is the only genus in which all species have storied rays and storied axial parenchyma (Fig. 25–30). Erythrostemon, Caesalpinia s.s., Tara, Guilandina and Mezoneuron have non-storied rays and non-storied to irregularly storied parenchyma. Coulteria and Poincianella (sensu Lewis) have some species with storeying and some without. Caesalpinia (?Poincianella) echinata is particularly interesting, because it has both storied and irregularly storied rays in the same specimen (see Fig. 42). Amano (2007) and Angyalossy et al. (2005) describe and illustrate this variation well. Storey- ing is usually considered by wood anatomists to be fairly consistent within a species, but clearly can be quite variable even in a single TLS (another very good example in an unrelated family is Swietenia in Meliaceae, pers. obs.). Prismatic crystals in ray cells have considerable diagnostic value in Caesalpinia s.l., can be found in procumbent, square or upright ray cells, and are often in radial lines. They were found in all species of Coulteria, Erythrostemon, Guilandina and Tara; most species of Poincianella, two out of five species of Caesalpinia s.s. and one of the two unplaced Old World species (C. decapetala). Crystals were not found in ray cells of Libidibia and Mezoneuron. As in most legumes prismatic crystals were found in chambered axial parenchyma strands in all species except for Erythrostemon gilliesii, based on a cultivated specimen at Kew; the species is native to southern South America. However, the abundance of crystals can vary considerably in a given species (although their occurrence in a given cell type is very consistent), and this is probably related to the availability of calcium in the soil and climatic conditions. In African acacias (Leguminosae, subfamily Mimosoideae), Gourlay and Grime (1994) found crystals to be more abundant in latewood and in trees growing in drier areas. It is often difficult to tell whether chains of chambered crystals are in axial parenchyma or fibres (see Fig. 12 for an example inCoulteria ). Ranjani and Krishnamurthy (1991) discussed the possibility that in Caesalpiniaceae (= Caesalpinioideae) the crystals are not in the axial parenchyma but are in separate apotracheal chambered crystal strands (presumably fibres). They were, however, unable to find any crystal strands in the specimen of C. pulcherrima they examined, but these are present in our slides. Caesalpinia madagascariensis is unique in having enlarged idioblastic parenchyma cells in the wood (Fig. 3–5). The presence of such cells in a scattered fashion across the legumes is puzzling. Miller (1989) found similar enlarged cells in Obolinga (now part of Zygia of subfamily Mimosoideae) and Gasson (1997) found them in Inga alba and I. lateriflora (Mimosoideae). Several species of Caesalpinia s.l. have similar idioblasts in their floral parts (C. (Coulteria) velutina, C. bahamensis, C. (Tara) vesicaria, C. pulcherrima, C. nipensis, but C. madagascariensis has not been examined: Rudall et al. (1994)).

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Tara spinosa (Fig. 65–67) from dry Andean valleys has a less well defined axial parenchyma pattern than its two congeners, Caesalpinia (Tara) cacalaco (Fig. 62–64) and C. (Tara) vesicaria (Fig. 58–61) from the Yucatan Peninsula in Mexico. We have been careful to distinguish between differences in wood anatomy of each species that could be attributed to variation in stem diameter and hence cambial age rather than taxonomic position. We have observed variation in cell type proportions that may be due to differences between juvenile and mature wood. Poincianella pannosa, for example, showed an increase in axial parenchyma proportion with cambial age (Fig. 54 & 55), which is also a reflection of much slower radial growth in this case. The specimen of Caesalpinia (Erythrostemon) calycina we examined has a much lower axial parenchyma proportion, both a higher proportion of vessels in radial multiples and more vessels per radial multiple (Fig. 14) and a slightly narrower stem (see Appendix) than our specimen of Erythrostemon gilliesii (Fig. 17), although possibly these two species do not belong to the same genus. In this broad survey, we have not explored variation within a given species, largely because we do not have enough samples to do this. However, there are some studies on Caesalpinia s.l. that explore wood variation for particular reasons. Silva (2006) has examined several Caesalpinia (Poincianella) pyramidalis branch and trunk wood samples of varying age, and although there is variation in cell type proportions, the overall appearance of the wood varies little from pith to bark. His work was primarily aimed at assessing whether the variations in cell proportions affected the density and hence suitability of the wood for burning. Another species, Caesalpinia (?Poincianella) echinata, is world renowned as the most important source of wood for violin bows, and some research has been aimed at ascertaining why, for example by Angyalossy et al. (2005), Amano (2007) and Segala Alves et al. (2008).

CONCLUSIONS

Libidibia (storied axial parenchyma, short storied rays, homocellular rays, rays without crystals) and Tara (non-storied mainly heterocellular rays, crystals in ray cells) are well defined on the basis of wood anatomy, whereas the other genera as currently circumscribed, all vary with respect to storeying, ray composition and distribution of crystals. In Caesalpinia s.l. it appears that storeying either does not reflect currently understood generic limits or that it is of variable occurrence within most of the genera. Ray cell composition is also well known to vary between juvenile and mature wood in many woody species, usually being more heterocellular in juvenile wood. In view of the small diameter of many of the samples we examined, using the distinction between homocellular rays and heterocellular ones to support the definition of genera should be treated with some caution. A similar caveat is necessary with crystals in ray cells. Although the location of crystals in a certain cell type is often diagnostic, low numbers of crystals can be overlooked, and their occurrence depends at least to some extent on environmental conditions. Only when a fully comprehensive combined analysis of molecular, anatomical and morphological characters of all Caesalpinia s.l. has been undertaken will it become clear how significant wood anatomy is in helping to define the various genera.

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ACKNOWLEDGEMENTS We are very grateful to the following for supplying some of the wood samples: Regis Miller (Madison, USA), Jorgo Richter (Hamburg, Germany), Beryl Simpson (Texas, USA) and Jenny Sotoyo (Mexico). Kasia Zieminska helped produce the plates. The contribution of the third author was, in part, supported by National Science Foundation grant DEB-0316375.

REFERENCES

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Appendix: Specimen details

Slide Caesalpinia species ** Specimen origin Collector/s and field no. Specimen ref. no. radius (cm) *** Caesalpinia s.s. s.n. C. bahamensis Cuba BFH 6544 1 (M) 21532 C. cassioides Colombia R.B. White 105.1897, 21446 1.2 (M) 19799 C. madagascariensis Madagascar G.P. Lewis et al. 2158 2.7 19808 C. nipensis Cuba G.P. Lewis et al. 1814 1.5 19807 C. nipensis Cuba G.P. Lewis et al. 1838 2.5 21520 C. pulcherrima Mexico G.P. Lewis et al. s.n. 28.3.1989 1.4 19814 C. pulcherrima S.W. Tropics 0.5 19813 C. pulcherrima Central America U.S.P.I.G 1 (M)

Coulteria 19795 C. gracilis Mexico G.P. Lewis et al. 2066 1 21523 C. platyloba? Mexico L. Rico & S.V. Contreras 1017 1.7 19818 C. velutina Guatemala G.P. Lewis & C.E. Hughes 1713 2.2 25014 C. velutina Guatemala J.G. Salas 1463, MADw32262 1.2 (M) 19822 C. violacea Guatemala G.P. Lewis & C.E. Hughes 1755 3.2 25016 C. violacea MADw13738 1.4 (M)

Erythrostemon 21471 C. calycina Brazil G.P. Lewis et al. 1858 0.7 25017 E. gilliesii RBG Kew Hort Kew 1988-68 WLKR 1

Guilandina 25023 G. bonduc Jamaica 5970 3.1961, 21447 0.7 25022 G. bonduc India J.S. Gamble 0.4824, 21448 1.2 (M) 19800 G. major Cook Islands July 1991 410, 69870 1.3 (M)

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Libidibia 19778 L. coriaria Mexico G.P. Lewis & C.E. Hughes 1777 0.5 19779 L. coriaria El Salvador G.P. Lewis & C.E. Hughes 1745 2.5 21521 C. ferrea Brazil G.P. Lewis et al. 1619 0.6 19792 C. ferrea Brazil J.A. Araujo Filho s.n. 1.5 (M) 7187 C. glabrata Ecuador G.P. Lewis et al. 2328 1.3 (M) 19796 L. punctata (as C. granadillo) Venezuela Pitt 4166.3.1961 1 (M) 19801 L. paraguariensis (as C. melanocarpa) Argentina No. 27 1 (M) 19811 L. paraguariensis Argentina G.P. Lewis & B.B. Klitgaard s.n. 1994 1.3 (M) 19812 L. punctata (as C. paucijuga) West Indies U.S.P.I.G 1 (M) 25021 L. punctata (as C. paucijuga) Trinidad University of Poland D 999 2/1981 1 (M) 19815 L. sclerocarpa Mexico G.P. Lewis & C.E. Hughes 1778 2.5

Mezoneuron 20177 M. hildebrandtii Madagascar G.P. Lewis et al. 2137 1.7 48594 M. sumatranum Malaysia W. Mejer 122531, MADw 0.8 (M) 37668 M. welwitschianum Angola Dechamps, Murta & Da Silva 1609, MADw 0.8 (M) Poincianella 21524 P. caladenia Mexico J.L. Contreras J. 2728 1.4 21525 P. caladenia Mexico J.L. Contreras J. 2729 1.6 19776 P. caladenia Mexico G.P. Lewis et al. 2072 1.4 (M) 21477 C. coccinea Mexico G.P. Lewis et al. 1802 2.5 19787 C. echinata G.P. Lewis et al. 1626 1.3 (M) 19788 C. echinata Brazil 1920 1 (M) 19780 C. echinata “Pechtl” Brazil wood only 1 (M) 19781 C. echinata “Joâo” Brazil wood only 1 (M) 19782 C. echinata “Joâo” Brazil wood only 1.2 (M) 19783 C. echinata “Satori” Brazil wood only 1.1 (M) 19784 C. echinata Brazil G.P. Lewis, violin bow 1 (M) 19785 C. echinata Brazil G.P. Lewis, violin bow 1 (M) 19786 C. echinata Brazil G.P. Lewis s.n. 1.2 (M) 21528 P. eriostachys Mexico G.P. Lewis et al. 1719 1.7 21469 P. exostemma subsp. exostemma Honduras G.P. Lewis & C.E. Hughes 1708 1.5 21476 P. exostemma subsp. exostemma Nicaragua D.J. Macqueen et al. 5 2 19794 P. gaumeri Mexico G.P. Lewis & C.E. Hughes 1764 2 19797 P. oyamae Mexico D.J. Macqueen et al. 428 2.5 21463 C. hughesii Mexico G.P. Lewis et al. 1795 2 19804 P. melanadenia Mexico G.P. Lewis & C.E. Hughes 1784 1 19803 P. melanadenia Mexico G.P. Lewis & C.E. Hughes 1787 2 19802 P. melanadenia Mexico G.P. Lewis & C.E. Hughes 1792 3 19805 P. mexicana Mexico C.E. Hughes et al. 1606 1.3 (M) 19806 P. myabensis Cuba G.P. Lewis et al. 1845 2 21473 C. nicaraguensis Nicaragua C.E. Hughes 1406 2.2 19809 P. palmeri Mexico J.L. Contreras J. 2732 0.7 21522 P. pannosa Mexico J.L. Contreras J. 2726 0.6 21527 P. pannosa Mexico J.L. Contreras J. 2713 0.7 19810 P. pannosa Mexico J.L. Contreras J. 2715 0.7 19770 P. pannosa (as arenosa) Mexico J.L. Contreras J. 2727 0.6 21466 P. placida Mexico G.P. Lewis et al. 2031 0.6 21465 C. pluviosa var. paraensis Brazil G.P. Lewis et al. 1628 1.5 10929 C. pyramidalis Brazil BFH 1.2 (M) 21470 P. standleyi Mexico J.L. Contreras J. 2694 0.6 19825 P. yucatanensis 20.9.89 1.2 (M) 19823 P. yucatanensis Mexico G.P. Lewis & C.E. Hughes 1765 1 (M) 19824 P. yucatanensis Mexico G.P. Lewis & C.E. Hughes 1766 1.2 (M) 21467 C. yucatanensis subsp. chiapensis Mexico C.E. Hughes et al. 1684 1.7 21472 C. yucatanensis subsp. hondurensis Honduras C.E. Hughes 1448 2.2 21474 C. yucatanensis subsp. yucatanensis Mexico G.P. Lewis & C.E. Hughes 1765 2.1 21475 C. yucatanensis subsp. yucatanensis Mexico G.P. Lewis & C.E. Hughes 1765 1.7

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Tara 19774 C. cacalaco Mexico G.P. Lewis & C.E. Hughes 1782 1.7 19773 C. cacalaco Mexico G.P. Lewis & C.E. Hughes 1789 2.5 19775 C. cacalaco Mexico C.E. Hughes 1516 2.2 19817 T. spinosa Ecuador Smithsonian Institution 1088 1 (M) 19820 C. vesicaria Cuba G.P. Lewis et al. 1847 2.2 19819 C. vesicaria Mexico G.P. Lewis & C.E. Hughes 1767 2 19821 C. vesicaria Nicaragua C.E. Hughes et al. 1376 2

Old World miscellaneous 21531 C. decapetala Swaziland J. Prior 287, Kw75473 1.3 19798 C. decapetala (as japonica) Japan HK 1952 1.3 (M) 19816 C. decapetala (as sepiaria var. japonica) TI-4829 1.9 (M) 21530 C. sappan Burma India Museum, Kw 25906 1.3 (M)

** in this column, C. always means Caesalpinia. Kw = Kew slide collection. BFH = Bundesforschungsanstalt für Forst und Holzwirtschaft, Hamburg, Germany. MADw = Forest Products Laboratory, Madison, Wisconsin, USA. For Caesalpinia echinata, the names “Pechtl”, “Joâo” and “Satori” refer to morphological variants, especially of the foliage (leaflet size). *** radius from pith to bark is not in bold, more mature samples where pith is not included are in bold followed by an (M).

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