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STATUS of an ASIATIC MEMBER of the JUGLANDACEAE REGARDED AS a 'LIVING FOSSIL'l Richard A

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STATUS of an ASIATIC MEMBER of the JUGLANDACEAE REGARDED AS a 'LIVING FOSSIL'l Richard A 666 AMERICAN JOURNAL OF BOTANY [Vol. 40 FRENCH, R. C. 1953. The effect of growth regulators and stimulating substances and related chemicals. Amer, allied compounds on growth and respiration of maize Jour. Bot. 36: 368-378. tissues. Thesis. Purdue Univ. Lafayette, Indiana. NANCE, J. F., AND L. W. CUNNINGHAM. 19.50. Acetalde­ LARDY, H. A., AND H. WELLMAN. 1952. Oxidative phos­ hyde accumulation in excised wheat roots induced by phorylations: Role of inorganic phosphate and ac­ plant growth substances. Science 112: 170-172. ceptor systems in control of metabolic rates. Jour. BioI. Chern. 19.5: 21.5-224. RHODES, A., AND R. DE B. ASHWOIlTH. 19.52. Mode of action ---, ---, AND C. FELDOTT. 19.52. Mechanism of ac­ of growth regulators in plants. Nature 169: 76-77. tion of dinitrophenol and thyroxine. Abstracts, Spring YAMAKI, T. 1949. Einfluss der Wuchstoffe auf den At­ Meeting, Div. BioI. Chem., Amer. Chern. Soc. mungsvorgang der Koleoptile von Avena sativa. II. MITCHELL, J. E., R. H. BURRIS, AND A. J. RIKER. 1949. Mitteilung iiber Pflanzliche Wirkstoffe. Acta Phyto­ Inhibition of respiration in plant tissues by callus chim. 1.5: 1.51-167. STATUS OF AN ASIATIC MEMBER OF THE JUGLANDACEAE REGARDED AS A 'LIVING FOSSIL'l Richard A. Scott ~ THE DESCRIPTION of a living representative of a sis, Chevalier also inexplicably referred to it as genus known previously only from the fossil record Juglans indochinensis in the same paper. Later in is a rare event of considerable interest to both neo­ the same year Kuang (1941) independently pro­ and paleobotanists. The extent of this interest may posed another new genus, Rhamphocarya, whose be measured in part by the large number of papers single species, R. integrifoliolata, is also based upon concerned with Metasequoia glyptostroboides since material of the plant under discussion. the discovery of this 'living fossil' was reported in Hjelmqvist (1948), overlooking the earlier de­ 1948. Recently Dr. Hsen·Hsu Hu (1952), who had scriptions by both Dodeand Chevalier, transferred a part in the discovery of the modern Metasequoia, the species from Rhamphocarya to Carya, creating has concluded that a member of the walnut family the new combination C. integrifoliolata (Kuang) now found in China and Indo-China is a living Hjelmqvist. W. Y. Chun had previously referred species of Juglandicarya, a genus previously known the species to Carya as C. tsiangii, but this combina­ only from fossil fruits occurring in the Eocene tion was never published (Manning and Hjelm. London Clay formation of England. The writer, qvist, 1951). who recently examined the material of Juglandi­ Leroy (1950) concluded that the species consti­ carya in the collections of the British Museum [Nat­ tutes a distinct genus and, with full knowledge of ural History), believes that this assignment is in the pertinent literature, proposed the new com­ error. The following discussion reviews the avail­ bination Annamocarya sinensis (Dode) Leroy as able information on the living and fossil species as its name. Leroy's reason for taking the species out a basis for the contention that the species are not of Carya. was based chiefly upon the vascular struc­ congeneric. ture of the fruit (1951a, b). He reported that the Unlike Metasequoia glyptostroboides, this Recent vascular strands extend from the base to the apex species has been known to botanists for a number within the inner wall of the fruit rather than with­ of years. However, a lack of reference to the perti­ in the primary partition as is the case for both nent literature, in part unavoidable, by authors Carya. and luglans. dealing with this species has led to much confusion Manning and Hjelmqvist (1951), after examin­ with regard to its systematic position and nomen­ ing all available herbarium material but without clature. knowledge of Leroy's work, reaffirmed Hjelmqvist's Fruits of the species were described as Carya earlier opinion that this Asiatic species belongs in sinensis by Dode (1912), but his description was Carya; As Leroy had done earlier, they linked overlooked until 1950. Meanwhile the species was Dode's description of Carya sinensis to the material again described from other material by Chevalier at hand. After rejecting the possibility that the spe­ (1941), who placed it in a new genus, Annamo­ cies is closely related to Juglandicarya, they con­ carya. Although naming this plant A. indochinen- c~uded that its name should remain as Carya. sinen­ SIS. 1 Received for publication April 2.5, 19.53. The first intimation that this Recent species might The author wishes to thank Mr. W. N. Edwards of the British Museum (Natural History) for his courtesy in per­ be a 'living fossil' was made by Chevalier when mitting study of the material of Juglandicarya in the Mu­ he described it as Annamocarya indochinensis. He seum's collections. This paper was written while the au­ considered that its fruits showed features suggest­ thor was a Rackham postdoctoral fellow at the University ing both Juglans and Carya, and that the species of Michigan. 2 Present address: The Biological Laboratories, Harvard might represent a form ancestral to these two gen­ University. era. Merrill (1948), using the name Rhampho- November, 1953] SCOTT-ASIATIC MEMBER OF JUGLANDACEAE 667 carya, regarded the plant asa living representative TABLE 1. Summary of opinions regarding the possible af­ of the fossil genus Caryojuglans Kirchheimer. The finities of the species of [uglandicarya Reid and monotypic Caryojuglans, described from fruits Chandler. found in European brown-coal beds, is regarded by Kirchheimer (1938) as an intermediate between Nearest Modern Relative luglansand Coryo; Miidler (1939) believes that its Reid and Manning and Juglandicarya Chandler Kirchheimer Hjelmqvist single species should be assigned to luglans, and Species (1933) (1951) (1951) Leroy (1952) assigns the species to Carya. The J. lubbocki [uglans fruits of luglans and Carya have several features Engelhardtia Pterocarya, in common and, particularly in the fossil state, are J. cantia Juglans Engelhardtia, sometimes difficult to distinguish. Their character­ J. depressa Pterocarya, Pterocarya Juglans istics have been discussed by Miidler and by Kirch­ Juglans rupestris heimer (1951). Both Kirchheimer (1951) and J. crassa Specimens disintegrated in storage Manning and Hjelmqvist (1951), after examining fruits of the Recent 'Rhamphocarya', have rejected the possibility that it isa species of Caryo juglans. able features of the species of luglandicarya are Kirchheimer has also expressed the opinion that the for the most part general ones typical for the wal­ Asiatic species does not belong to luglandicarya. nut family and not necessarily limited to anyone Hu's (1952) transfer of the problematic species modern genus. Reid and Chandler's diagnosis of to the fossil genus luglandicarya is the most recent­ I. cantia is representative (1933, p. 142) : ly published opinion regarding its affinities. Of the "Endocarp globular, smooth, and without ex­ papers cited above, Hu referred only to Kuang's ternal nodulations, dehiscing into equal valves, one­ work. It should be noted, however, that because of loculed, one-seeded; walls thick, without cavities. the length of the period during which Hu's paper Seed erect, orthotropous, conforming to the shape was in press, he could not have seen any of the of the locule, simple above, two-lobed below, each papers published during 1951. Hu considered the lobe being slightly emarginate at the base. Diam­ earliest previous name for the species to be Rham­ eter of endocarp about 12 mm." phocarya integrijoliolata and created the new com­ The similarities upon which Hu bases the sup­ bination luglandicarya integriloliolata (Kuang) Hu posed congeneric relationship between 'Rhampho­ to designate it. carya' and luglandicarya are stated in this quota­ The net result of this involved sequence is that tion (1952, p. 264) : since it was first described this one juglandaceous 'Rhamphocarya has a smooth, globular to ellip­ species has been associated with six generic names: soid, one-loculedand one-seeded endocarp with Carya, luglans, Annamocarya, Rhamphocarya; thick wall without cavities. Its seed is erect, ortho­ Caryojuglans, and luglandicarya. There are cur­ tropous, conforming to the shape of the locule, sim­ rently three independent proposals regarding its ple or emarginate above, deeply two-lobed below, designation in the literature: Annamocarya sinen­ the lobes again being shallowly two-lobed by a sec­ sis (Dode) Leroy, Carya sinensis Dode, and the ondary septum. Its contours are smooth. These one to be discussed here, luglandicarya integri­ characteristics are similar to those of luglandi­ joliolata (Kuang) Hu. carya; only the size of the endocarp is much The genus luglandicarya was founded by Reid larger." and Chandler (1933, p. 140) to contain "Fruits This description of the fruit of 'Rhamphocarya' which, although clearly referable to the Juglanda­ is obviously similar to the diagnosis of luglandi­ ceae, are of doubtful generic relationship both to carya cantia and is also like that of I. lubbocki. Of living genera and to one another." It includes four the species of luglandicarya, Hu found these two to published species, one of which, I. crassa (Bower­ agree most closely with the modern species. The bank) Reid and Chandler, was based on material correspondence which he points out is not, how­ which had disintegrated in storage before the lat­ ever, adequate evidence for congeneric relation­ ter authors transferred it to luglandicarya. The ship. Hu's characterization of 'Rhamphocarya' is original description is not adequate for compari­ a generalized one which is also applicable in most sons. Opinions regarding the possible affinities of respects to fruits of other genera in the walnut fam­ the other three species are shown in table 1. These ily, for example, luglans and Carya. The only fea­ diverse opinions bear out the conclusion of Reid ture in which the agreement might indicate close and Chandler that the species assigned to lug­ relationship to luglandicarya is the stated lack of landiearya do not constitute a single natural genus.
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