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WOOD ANATOMY of EXOSTYLES VENUSTA (SWARTZIEAE, PAPILIONOIDEAE, LEGUMINOSAE) by Peter Gasson & Polly Webley

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WOOD ANATOMY of EXOSTYLES VENUSTA (SWARTZIEAE, PAPILIONOIDEAE, LEGUMINOSAE) by Peter Gasson & Polly Webley IAWA Journal, Vol. 20 (1), 1999: 59-66 WOOD ANATOMY OF EXOSTYLES VENUSTA (SWARTZIEAE, PAPILIONOIDEAE, LEGUMINOSAE) by Peter Gasson & Polly Webley Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, United Kingdom SUMMARY The genus Exostyles consists of two to three species from Brazil. This paper describes the wood of E. venusta Schott, completing generic cov­ erage of the wood anatomy of the tribe Swartzieae as defined by Cowan (1981). Small twigs from herbarium sheets of E. glabra and E. ama­ zonica (labelled E. venusta) were also examined. Exostyles wood anat­ omy is compared with that of the other genera in the tribes Swartzieae (Gasson 1996) and Sophoreae (Gasson 1994; Fujii et al. 1994), and is very similar to three closely related genera of Swartzieae, Zollernia, Harleyodendron and Lecointea. Key words: Exostyles, Swartzieae, Sophoreae. INTRODUCTION Exostyles Schott is a poorly known and rarely collected genus, consisting of two species from Southeast Brazil (E. venusta Schott and E. glabra Vogel) and possibly a third (E. amazonica Yakovlev) from Amazon Brazil. Wood specimens of Exostyles were unavailable for a previous study of Swartzieae (Gasson 1996), and there is ap­ parently no previously published information on the wood anatomy of this genus. We have since discovered two wood samples of E. venusta, which enables us to complete generic descriptions for the tribe Swartzieae as defined by Cowan (1981) and forms the basis for a general discussion of wood anatomy in relation to the systematics of the closely related legume tribes Swartzieae and Sophoreae. Since there are differ­ ences in pollen structure between E. venusta and E. glabra (Ferguson & Skvarla 1988), we also examined small twigs from herbarium specimens of E. glabra and E. ama­ zonica (the latter originally labelled E. glabra, then E. venusta !). MATERIALS AND METHODS The description below is based on two samples of Exostyles venusta obtained from the wood collection at IPT, Sao Paulo: BCTw 13301, Jardim Botanico do Rio de Janeiro, placa no. 4640, A. Mattos Filho; BCTw 13328, Pernambuco, DardanoAndrade Lima. For full details of the IPT wood collection see Stem (1988). Two very small twigs from herbarium specimens were also examined: (1) E. glabra Vogel, colI. M. Glaziou 9410, near Rio de Janeiro (1878), twig approx. 4.5 mm in diameter; (2) E. Downloaded from Brill.com09/24/2021 04:00:51AM via free access 60 lAWA Journal, Vol. 20 (1), 1999 amazonica, colI. M.J. Pires & N.T. Silva 2054 (1968), Monte Dourado, Para, twig approx. 3 mm in diameter. This specimen comes from the type locality of E. amazonica (Yakovlev 1979), and was originally labelled E. glabra, then later E. venusta. It is probably E. amazonica, and is referred to as such in this paper. No anatomical meas­ urements of vessels, fibres or rays are given, because the original positions of the samples in the tree (i. e. whether from branch or trunk) and their cambial age are un­ known. WOOD DESCRIPTION (Fig. 1-9) Exostyles venusta: Wood diffuse-porous without growth rings. Vessels solitary, in pairs, radial mUltiples of 3 or 4 and occasional clusters of up to 6; some vessels con­ taining dark-staining deposits. Vessel elements storied, relatively short, with fine, al­ ternate, polygonal, vestured pitting and simple perforation plates. Vessel-ray pitting similar to intervascular pitting in size and shape. Fibres thick-walled with lumina narrow or absent, often gelatinous, non-septate, with slit-like apparently simple pits. Axial parenchyma predominantly paratracheal, but variable in distribution; mainly in tangential bands 2-4 cells wide in BCTw 13328, more scanty paratracheal and winged aliform in BCTw 13301, and confluent in both. Axial parenchyma storied, in strands of 2-4 cells. Rays mainly storied, short rays about the same height as adjacent axial parenchyma strands, axially fused in places to form rays 2, 3 or 4 stories high. Rays: in BCTw 13328 mainly 2 cells wide (but also some uniseriate and in a few places triseriate) and c. 10 cells high; in BCTw 13301 often 3 and occasionally 4 cells wide and c. 15 cells high. Rays heterocellular, with 1-3 rows of square or upright marginal cells. Crystals: prismatic crystals present in both axial parenchyma and ray cells, uncommon in non-chambered and chambered axial parenchyma cells, and common in non-chambered ray cells (usually square and upright), often in radial lines, a single crystal in each ray cell. Note: twig samples of E. glabra and E. amazonica were similar to the E. venusta samples described above, although no crystals were found in E. amazonica, and nei­ ther twig had storied rays or axial parenchyma. The absence of storeying raises the question that this could be partly dependent on cambial age as well as taxonomic affinity. There are several examples in legumes of variation in storeying within a genus, e.g. in Ateleia (Gasson 1994, and Dalbergia pers. obs.), which may be a func­ tion of cambial age, and this needs further study. SYSTEMATIC DISCUSSION The systematics of the closely allied tribes Swartzieae and Sophoreae are currently in a state of flux (Polhill 1981, 1994; Herendeen 1995); they include both South Amer­ ican taxa (e.g., Aldina, Holocalyx, Exostyles, Zollernia, Harleyodendron, Lecointea and Acosmium) and African taxa (e.g., Cordyla and Mildbraediodendron). Fossil leg­ ume pollen and wood have been reported from the Upper Cretaceous (Herendeen etal. 1992; Wheeler & Baas 1992), by which time (90-80 million years before present) Africa and South America were well separated (Fig. 13.12 & 13.15 in Howarth 1981), Downloaded from Brill.com09/24/2021 04:00:51AM via free access Gasson & Webley - Wood anatomy of Exostyles 61 Fig. 1-4. Exostyles venusta. TS. - 1 & 2: Slightly undulating bands of aliform and confluent parenchyma, many gelatinous fibres (BCTw 13301). - 3 & 4: much straighter bands of axial parenchyma than 1 & 2, also showing some aliform and confluent (BCTw 13328). Scale line for 1 & 3 (on 3) is 200~, for 2 & 4 (on 4) is 100 ~. Downloaded from Brill.com09/24/2021 04:00:51AM via free access 62 IAWA Journal, Vol. 20 (1), 1999 Fig. 5-9. Exostyles venusta. 5-7: TLS. 8 & 9: RLS. - 5: Storied axial parenchyma strands and irregularly storied rays, often extending axially alongside two stories of parenchyma strands (BCTw 13301). - 6-9: BCTw 13328. 6 & 7: Storied narrow vessel elements, axial parenchyma and rays. Prismatic crystals present in some ray cells in 7. - 8 & 9: Rays slightly heterocellu­ lar, with a single row of square cells at ray margins and within body of ray, some ray cells con­ taining a single prismatic crystal. Scale for 5 & 8 (on 5) is 200 J.lII1, for 6, 7 & 9 (on 9) is 100 J.lII1. Downloaded from Brill.com09/24/2021 04:00:51AM via free access Gasson & Webley - Wood anatomy of Exostyles 63 Caesalpinieae (Sclerolobium, Diptychandra) Acosmium (Sophoreae) o rphanodendron (Caesalpinieae) ALDINA (Swartzieae) Sophoreae (6 genera: Baphia, Baphiastrum, Leucomphalos, Airyantha, Bowringia, Dalhousiea) Swartzieae (6 genera: SWAR1ZIA, ++BOCOA, CANDOLLEODENDRO~CORDYLA, MILDBRAEDIODENDRON, BAPHIOPSIS) Holocalyx (Sophoreae) LECOINTEA (Swartzieae) HARLEYODENDRON (Swartzieae) EXOSTYLES (Swartzieae) ZOLLERNIA (Swartzieae) Sophoreae (11 genera: Myroxylon, *Amburana, *Ateleia, *Cyathostegia, Alexa, Castanospermum, Luetzelburgia, Bowdichia, Clathrotropis, Dussia, Ormosia) Fig.l0. Diagram summarising strict consensus tree in Herendeen (1995). Genera in upper case italics are Swartzieae. Asterisked genera were included in Sophoreae by Polhill (1981), but in Swartzieae by Polhill (1994). - ++Bocoa was treated as two taxa by Herendeen and they did not group together, suggesting that the genus is polyphyletic. In the unweighted analysis one Bocoa taxon grouped at the base of the Baphia-c1ade and the other with the Swartzia-clade. so the taxa on the two continents have had ample time to have diverged. Herendeen (1995) used both unweighted and successively weighted cladistic analyses of flower, fruit, pollen and leaf characters of genera of Swartzieae and some Sophoreae. His results indicated that Swartzieae and Sophoreae are para- or polyphyletic, and that Exostyles belongs in a clade (the' Lecointea-clade') consisting of five genera: Exostyles, Harleyodendron, Lecointea, Zollernia and Holocalyx (Fig. 10), which is united by four characters: toothed leaflet margins, basifixed anthers, stomata with heavily cuti­ nized flanges flanking the guard cells and ovoid leathery fruits. The wood anatomy of Exostyles was compared here with that of all the genera described by Gasson (1994) and Fujii et al. (1994) for Sophoreae, Gasson (1996) for Swartzieae, Bedell in Cowan (1979) for Harleyodendron and Baretta-Kuipers (1981) for both tribes and many other legume woods. Several Swartzieae and a few Sophoreae are similar in wood anatomy to Exostyles, the most similar genera being Zollernia, Downloaded from Brill.com09/24/2021 04:00:51AM via free access 64 lAWA Journal, Vol. 20 (1),1999 Table 1. Ray characters in Exostyles and six of the most similar genera in Swartzieae and Sophoreae. Key: I = irregularly storied. Crystals in Heterocellular Uniseriate Storied rays ray cells rays rays Exostyles Harleyodendron 1). Lecointea 1-2 Zollernia 1). Holocalyx Acosmium 1). Aldina Harleyodendron and Lecointea (all Swartzieae) and to a lesser extent, Holocalyx (Sophoreae according to Polhill1981 and Herendeen 1995, Swartzieae in Polhilll994) (Table 1). There is strong similarity in wood anatomy between the five genera of the Lecointea-clade. The presence of crystals in ray cells is relatively uncommon in wood of Papilion­ oideae, although prismatic crystals in chambered axial parenchyma cells are almost ubiquitous. This is therefore a potentially useful taxonomic character at the genus or species level. In the Lecointea-clade, Exostyles, Zollernia, Harleyodendron and Le­ cointea (but not Holocalyx) all have crystals in ray cells (Table 1), although these were inconspicuous and initially overlooked in the two samples of Lecointea amazonica examined by Gasson (1996). On the other hand, Acosmium (Sophoreae), which some­ times has crystals in ray cells, was relatively distant to the Lecointea-clade in Heren­ deen's (1995) analysis (Fig.
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