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Wood Anatomy of Goodeniaceae and the Problem of Insular Woodiness

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Wood Anatomy of Goodeniaceae and the Problem of Insular Woodiness WOOD ANATOMY OF GOODENIACEAE AND THE PROBLEM OF INSULAR WOODINESS SHERWIN CARLQuIsT ANN. MISSOURI B0T. GARD. 6: 358—390. 1969. WOOD ANATOMY OF GOODENIACEAE AND THE PROBLEM OF INSULAR WOODINESS’ SHERWIN CARLQuIsT’ ABSTRACT Goodeniaceae, a basically herbaceous family with a clear center of origin and diversffica tion in Australia, has become woody to a limited extent in several genera and genuinely arborescent in some species of Scaevola. The most woody of these are montane species of Pacific islands. The probable evolutionary patterns of these and correlations between wood anatomy and ecological conditions are reviewed. Quantitative and qualitative data are pre sented in tabular form for 78 collections of 43 taxa of the family. Xermorphic Goodeniaceae tend to have short, narrow vessel elements with helical sculpturing, whereas the reverse is true in mesic species. Absence of axial parenchyma and abundance of crystals also charac terizes xeric species. The predominance of erect ray cells or raylessness, as well as occurrence in a few species of scalariform or elliptical pits on lateral walls of vessels are probably in dicators of juvenilism (paedomorphosis). Goodeniaceae do not have libriform fibers, but rather tracheids or fiber-tracheids, with a tendency toward the latter in arborescent species. Multiseriate rays are tall. Correlations between wood anatomy and habit are summarized under the headings: Australian short-lived perennials and short-lived shrubs; montane Pacific species of Scaevola; and maritime species of Scaevola. INTRODUCTION Wood anatomy of Goodeniaceae has been little studied. This no doubt derives from the fact that no species in the family provides any economically used wood. Although a few species of Scaevola become small trees, most of the family can be described as herbs. However, a predominantly herbaceous family that becomes woody during its evolution provides interesting patterns of wood anatomy and ought to be an especially valuable subject for that reason. Goodeniaceae is an outstanding example of this mode of evolution. The center of origin and diversification of the family is obviously Australian. If we take into account Brunonia and the recently named Neogoodenia (Gardner & George, 1963) and Coopernookia (Carolin, 1967), ten of the 16 genera are endemic to Australia. In four genera, only one species extends beyond Australia, and that species occurs in Australia also: Selliera radicans (southern and south eastern Australia, Tasmania, New Zealand, Chile); Velleia spathulata (eastern Australia, southeastern New Guinea); Calogyne pilosa (northern Australia, Mo luccas, southern New Guinea, southern China); and Leschenaultia fihiformis (northern Australia, Moluccas, New Guinea). The remainder of these genera are exclusively Australian species. Goodenia koningsbergii occurs only in Indo nesia and southeastern Asia, but this is a single species of a large genus. That none of these genera (except Goodenia) has preponderantly primitive features for 1 This study has been supported by grants from the National Science Foundation, NSFG—23396, GB—4977X, and GB—14o92. Preliminary comments on this topic were pre sented at the 10th Pacific Science Congress. 2 Claremont Graduate School, Rancho Santa Ana Botanic Garden, Claremont Califor nia 91711. ANN. MISSOURI B0T. GARD. 56: 358—390. 1969. 19691 CARLQUIST—GOODENIACEAE 359 the family suggests strongly that the origin and diversificaion of Goodeniaceae has taken place within Australia. The only other genus with non-Australian species is Scaevola. This genus, too, has been regarded as specialized within the family (Krause, 1912; Carolin, 1959, 1966). It is the only genus of the family which becomes markedly shrubby or arborescent. The most arborescent species are S. gau dichaudiana and S. glabra of the Hawaiian Islands; these can form trees five or six meters tall. On the basis of circumstantial evidence alone, the family has an herbaceous origin and has become woodier on islands. Floral morphology and geographical distribution seem to support this view. Of the genera other than Scaevoltz, few show any appreciable woodiness. This seems to be related to adaptation to the dry temperate Australian environment. Many Australian Goodeniaceae are annuals or short-lived perennials. The latter are often branched from the base, so that little wood accumulates, and a wood sample, if one can be collected at all, has to be taken from the short rootstock. Wood in this structure is probably not really comparable to wood of a true shrub, and the grain is often so contorted that sections are difficult to prepare and of limited value. Growth forms such as these are related to Mediterranean climates. In the present study, Dampiera brownii and D. glabrescens represent caudex bearing perennials, branched from near the base. Short-lived shrubs, usually less than a meter high, are formed by Goodenia scapigera, G. ovata, Coopernookia polygakicea, C. strophiolata, and Verreauxia reinwardtii. Of these, only Goodenia ovata grows in a mesic situation, the eucalypt forests in which tree ferns may also be found. The other shrubby species listed grow in relatively dry sandy areas of Western Australia. Within the genus Scaevola some of the Australian species correspond to the above growth forms, but some are more genuinely woody. Scaevola sericophylla and S. fasciculata are short-lived shrubs of Western Australian scrub heath, whereas S. crassifolia, S. nitida, and S. porocarya are larger shrubs (to one meter or more), probably of relatively short duration (10 years or less). These three species occupy sites near the coast: S. crassifolia is typically a dune plant. Scaevola spinescens is a shrub, usually less than a meter in height, that grows near salt lakes in Western Autralia and other states of Australia. Scaevola tornentosa, al though a close relative of S. spinescens, is a prostrate shrub of coastal limestones in Western Australia. Scaevola holosericea and S. paludosa are Australian species, little more than perennial herbs, in which only a few older stems have sufficient wood for study. None of the Australian scaevolas could be called clearly mesic in habitat except S. oppositifolia, a scandent species of disturbed places in wet forests of Queens land. In localities north of Australia, including islands as far north as the Philip pines, S. oppositifolia has been given a multiplicity of segregate specific names. These have been justifiably designated as synonyms by Leenhouts (l9S7a). With respect to taxonomy, Krause (1912) has placed all of the Australian species studied here in section Xerocarpaea except S. spinescens, S. tomentosa, S. oppositifolia, and the wide-ranging S. taccada. Scaevola spinescens and S. tomentosa [VoL. 56 360 ANNALS OF THE MISSOURI BOTANICAL GARDEN FIGURES 1—4.—Wood sections of Coopernookia. — 1—2. Coopernookia polygalacea (Carl quist 3475).—1. Transection. Note narrowness of vessels.—2. Tangential section. Rays are numerous, narrow. — 3—4. Cciopernookza straphiolata (Cariquist 3393).—3. Transection. Note growth rings.—4. Tangential section. Rays are few, wide; light portions are the wider vessels from early wood. — Scale to the left of Fig. 1 is a stage micrometer photographed at the same scale as Fig. 1—41. Length shown is 1.8 mm; divisions 1Os each. 1969) CARLQUIST—GOODENIACEAE 361 are members of sect. Crossotoma; S. oppositifolia belongs to sect. Enantiophyllum, and S. taccada belongs to sect. Scaevola. Species of section Scaevola (formerly sect. Sarcocarpaea), sometimes termed the “fleshy-fruited scaevolas,” bear purple or white fruits with a fleshy exocarp attractive to birds and a woody endocarp in which seeds are embedded. Because of the excellence of this dispersal mechanism for long-distance dispersal (Carl quist, 1967), the fleshy-fruited scaevolas have spread around the world. This is undoubtedly a relatively recent, although pre-human, pattern. The widespread pan-Pacific beach Scaevola, S. taccada, occurs in Australia and might have orig inated there. Scaevola plumieri does not occur in Australia, but its distribution might be said to represent a radiation from the Australian region. The species of section Scaevola are otherwise non-Australian. However, perhaps the closely- related fleshy-fruited sections, Crossotoma and Enantiophyllum represent remnants of this stock on the Australian continent. Scaevola calendulacea, a beach species of eastern Australia, has fleshy white fruits, tinged purple, that float in seawater and remind one of fruits of S. taccada. It supposedly belongs to section Xerocarpaea, but if so (and that disposition needs to be reinvestigated), it shows rapid evolution of a beach species with fleshy fruits both attractive to birds and dispersible by flotation in seawater. Either interpretation of S. calendulacea as a derivative of dry-fruited Scaewola species or hypothesizing it as a relict of sect. Scaevola stresses the same result—origin of the fleshy-fruited stock termed section Seaevola in the Australian region. Two species of section Scaevola are exclusively maritime and distributed by oceanic drift of the fruits: S. taccada (also known as S. sericea, S. frutescens, S. koenigii, etc.) and S. plumieri (also known as S. lobelia). Guppy (1917) demon strated that S. taccada fruits can float for a year or more in seawater, while fruits of S. plumieri can withstand only four or five months of flotation. Scaevola taccada ranges across the Pacific, reaching all tropical Pacific coasts (except the west coasts of Mexico, Central and South
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