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Wood Anatomy of the Altingiaceae and Hamamelidaceae

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Wood Anatomy of the Altingiaceae and Hamamelidaceae IAWA Journal, Vol. 31 (4), 2010: 399–423 WOOD ANATOMY OF THE ALTINGIACEAE AND HAMAMELIDACEAE Elisabeth A. Wheeler1, Sung Jae Lee2 and Pieter Baas3 SUMMARY Wood anatomical data for all three extant genera of the Altingiaceae and 23 of the 27 extant genera of the Hamamelidaceae were compiled in an effort to find features distinctive to genera, tribes, or subfamilies within these families. All genera studied have diffuse porous wood (except Corylopsis which tends to be semi-ring porous), vessels are predominantly solitary and narrow (<100 µm, usually < 50 µm) and angular in outline, vessel elements are long (>800 µm) with scalariform perforation plates with average bar numbers of 9–44, inter- vessel pits are mainly scalariform to opposite, vessel-ray parenchyma pits are scalariform with slightly reduced borders and usually are in the square to upright marginal ray parenchyma cells, rays are heterocellular and narrow, usually 1–3-seriate. Although the wood anatomy of both families is relatively homogeneous, it is possible to key out many genera using a combination of qualitative (presence/absence and location of helical thickenings in vessel elements and fibers, crystal occurrence, axial parenchyma abundance, degree of ray heterogeneity) and quantitative features (number of bars per perforation plate and ray width). Helical thickenings are present throughout the vessel elements in three genera (Loropetalum, Altingia, Semiliquidambar) and are restricted to the vessel element tails in two genera (Corylopsis, Liquidambar). Loropetalum has helical thickenings in ground tissue fibers as well. Axial parenchyma abundance varies from scarce to relatively abundant diffuse to diffuse-in-aggregates. One clade of the tribe Fothergilleae (Distylium, Dis- tyliopsis, Sycopsis, Shaniodendron, Parrotia, Parrotiopsis) has more abun- dant axial parenchyma and is characterized by narrow, usually interrupted bands of apotracheal parenchyma. Nearly exclusively uniseriate rays occur in some species of Hamamelis and in Exbucklandia, Chunia, Dicoryphe, and Fothergilla. These data on extant Altingiaceae and Hamamelidaceae not only provide information relevant for systematic, phylogenetic and ecological wood anatomy and wood identification, but also give context for reviewing the fossil woods assigned to them. A new combination is proposed for the Miocene Liquidambar hisauchii (Watari) Suzuki & Watari from Japan: Altingia hisauchii (Watari) Wheeler, Baas & Lee. Key words: Altingiaceae, Hamamelidaceae, systematic wood anatomy, Eocene, fossil wood, Hamamelidoxylon. 1) Department of Forest Biomaterials, North Carolina State University, Raleigh, NC 27695-8005, U. S. A. [E-mail: [email protected]]. 2) Gangwon Forest Development Research Institute, 200-140 Chunchon, South Korea. 3) National Centre for Biodiversity Naturalis, National Herbarium of the Netherlands, P.O. Box 9514, 2300 RA Leiden, The Netherlands. Associate Editor: Frederic Lens Downloaded from Brill.com09/29/2021 12:32:35AM via free access 400 IAWA Journal, Vol. 31 (4), 2010 INTRODUCTION The Hamamelidaceae constitute a family of trees and shrubs, with a tropical to temperate distribution, being especially diverse in Eastern Asia. The family is placed in the Saxi- fragales, with the Altingiaceae, Cercidiphyllaceae, and Daphniphyllaceae recognized as the most closely related families within the order (APG III 2009; see also Stevens 2001-onward). Genera of the Altingiaceae (Altingia, Semiliquidambar, Liquidambar) formerly were considered members of the Hamamelidaceae. The Angiosperm Phylog- eny Website lists three subfamilies for the Hamamelidaceae: 1) Exbucklandoideae with the genera Chunia, Exbucklandia, Mytilaria, and Rhodoleia (composition according to Endress 1989), occurring in East Asia, including Assam, East Malesia to Sumatra, 2) Disanthoideae comprised of the monotypic Disanthus cercidifolius of Japan, and the largest subfamily 3) Hamamelidoideae. Li and Bogle (2001) considered Hamamelidoideae monophyletic, and their analysis of morphological and molecular features recovered three monophyletic groups within the Hamamelidoideae. They proposed recognition of two tribes within each group (see Fig. 1). There are differ- ent estimates of the number of species per genus, but only three genera (Corylopsis, Dicoryphe and Distylium) are estimated to have more than 10 species. A remarkably high proportion of the genera, 11 of 27, are monotypic. Corylopsis (Asia) Loropetalum (Asia) Tetrathyrium (Asia) Matudaea (S. America, Mexico) Embolanthera (Asia) Maingaya (Asia) Dicoryphe (Madagascar) Trichocladus (Africa) Neostrearia (Australia) Noahdendron (Australia) Ostrearia (Australia) Eustigma (Asia) Fortunearia (Asia) Sinowilsonia (Asia) Molinadendron (Central America) Distylium (Asia) ⎫ ⎪ Distyliopsis (Asia) ⎪ ⎪ Sycopsis (Asia) ⎬Banded parenchyma ⎪ Shaniodendron (Asia) ⎪ Parrotia (Asia) ⎪ ⎪ Parrotiopsis (Asia) ⎭ Fothergilla (N. America, 1s rays only) Hamamelis (Asia: 1–3s rays; N. America: 1s rays only) Figure 1. Phylogenetic relationships within the Hamamelidoideae. Redrawn from Li & Bogle (2001) and Radtke et al. (2005). Downloaded from Brill.com09/29/2021 12:32:35AM via free access Wheeler, Lee & Baas — Altingiaceae and Hamamelidaceae 401 The Hamamelidaceae and Altingiaceae have a fossil record dating back to the Late Cretaceous, including an inflorescence interpreted as a basal “altingioid” with pollen characters found in the Hamamelidaceae (Zhou et al. 2001) and a Late Cretaceous (Campanian) flower attributed to the tribe Hamamelideae (Magallon-Pueblaet al. 1996). Some genera now endemic to Asia (Corylopsis, Disanthus, Fortunearia) occurred in North America and Europe during the Tertiary (Manchester et al. 2009). Hamamelis and Liquidambar (Altingiaceae) are classic examples of genera with a present-day disjunct distribution (e.g., Manchester 1999). Tracing the history of the family from fossil seeds is made difficult because “seeds [of Hamameloideae] have converged on very similar morphology … some of the genera might be difficult or impossible to dis- tinguish based on seed morphology alone …” (Manchester et al. 2009). Consequently, it is of interest to determine if any genera, tribes, or subfamilies of Hamamelidaceae can be distinguished by their wood anatomy so that fossil wood might be used to better understand the history and diversification of the Hamamelidaceae. The Kew Micromorphology database and Gregory (1994) list publications describing wood anatomy of the Hamamelidaceae and Altingiaceae. An especially useful reference, although it lacks figures, is Tang’s paper (1943), which tabulates data and includes a key to 19 genera (Altingiaceae and Hamamelidaceae). This paper apparently was a major source for the family description by Metcalfe and Chalk (1950). Subsequent relatively comprehensive treatments, with figures, are by Skvortsova (1975, 14 genera, 20 species studied) and Huang (1986, 9 Chinese genera studied). Materials AND METHODS Wood samples were obtained from various institutional wood collections. Unfortunately, some of the wood samples lack information on simultaneously collected herbarium material (see Barker 2008 for risks entailed in erroneous identifications). Following, in alphabetical order by genus within family, is a list of specimens examined and the information available on their provenance. The numbers of species per genus are from the Flora of China (FOC) (Zhang et al. 2003) and Mabberley (2008). Slides are deposited in the National Centre for Biodiversity Naturalis - National Herbarium of the Netherlands. Additional images are available on the InsideWood web site (InsideWood 2004-onwards). Images of additional samples and species on InsideWood, provided by FFPRI, Tsukuba, Japan (TWTw and courtesy of S. Noshiro), CSIRO (FPAw and courtesy of J. Ilic), National Herbarium of the Netherlands (Uw, UN) and Imogen Poole (slides at Kew, Kw) are listed below in brackets. Altingiaceae (3 of 3 genera examined) Altingia Noronha (Evergreen trees, Indomalesia, 8–10 species, 4 examined): A. chi- nensis (Champ. ex Benth.) Oliv. ex Hance, SJRw 21920, Coll. C.L.Tso, Kwangtung, China, Fan Memorial Institute Biology — A. excelsa Noronha, TWTw 3123 (= BZFw N4472), Forest Research Institute, Bogor, Indonesia; [FPAw 10616] — A. gracilipes Hemsl., CAFw 11260, mature tree, China — A. obovata Merr. & Chun, SJRw 29562, Ngai Yuen, Hainan, China. Downloaded from Brill.com09/29/2021 12:32:35AM via free access 402 IAWA Journal, Vol. 31 (4), 2010 Liquidambar L. (Deciduous trees, east Asia, east Mediterranean, southeast North America to northern Central America, 3–5 species, 3 examined): L. formosana Hance, CAFw 18047, mature tree, China; [TWTw 17405] — L. orientalis Miller, MADw 14115, cultivated, Riverside Co., CA, USA — L. styracifluaL., TWTw 6459, cultivated Osaka City, Japan; TWTw 15982, cultivated Forest Tree Breeding Institute, Kasahara-cho, Mito City, Ibaraki Pref., Japan; BWCw 8148; [FPAw u.45, Kw Liqstyr, Tw 52940, Uw 6759, RBHw 1447, Hw 17232]. Semiliquidambar H.T. Chang (Evergreen-deciduous trees, eastern China, 3 species, 1 examined): S. cathayensis H.T. Chang, Liu Peng s.n., Kwangtung, China, 23° N, 113° E. Hamamelidaceae (23 of 27 genera examined) Chunia H.T. Chang (Evergreen tree, China, 1 species): C. bucklandioides H.T. Chang, Tw 41881, coll. Forest Research Institute, Guangdong, China; Liu Peng s.n., Kwangtung, China, 23° N, 113° E. Corylopsis Siebold & Zucc. (Shrub to small tree, evergreen to semi-evergreen, China, Japan, India, 7–29 species, 3 examined): C. gotoana Makino, TWTw
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