JOURNAL
OF THE ARNOLD ARBORETUM HARVARD UNIVERSITY
ALFRED REHDER EDITOR
JOSEPH H. FAULL ann CLARENCE E. KOBUSKI ASSOCIATE EDITORS
VOLUME XVII
JAMAICA PLAIN, MASS. 1936
Reprinted with the permission of the Arnold Arboretum of Harvard University
KRAUS REPRINT CORPORATION New York
1968
DATES OF ISSUE
(pp. 1-51, pl. 166-178) issued January 24, 1936, (pp. 53-114, pl. 179-181) issued April 16, 1936. (pp. 115-240, pl. 182-194, map) issued July 30, 1936. (pp. 241-389, pl. 195-200) issued October 20, 1936.
Printed in U.S.A.
TABLE OF CONTENTS
STUDIES ON THE BIOLOGY OF GYMNOSPORANGIUM GLOBOSUM FAL. ‘Vith plates 166-175 and two text figures. By J.D. MacLachlan... 1
Tue Lire Histories or MILEs1A ScoLopeNpRII, M. Potypopi, M. VOGESIACA AND M. KRIEGERIANA. With plate 176. By Lillian M.
A CONVENIENT SAND-CULTURE ee With plate 177 and one text tipure. (By Mover! WW Ord eee oases odoin eas See eee CHROMOSOME COMPLEX AT PREMEIOTIC ANAPHASE AND MEIOTIC METAPHASE. With two text figures. By Haig Dermen ........ BURRETIODENDRON, A NEW GENUS OF TILIACEAE. With ous 178. BY Ee OER Ne 2055550 6 a oA SoU eae ee eae Two NEw SPECIES OF GYMNOSPORANGIUM FROM AsIA. With two RONG UUM Se OOM UPON 6 oss bsead eae dae eaneee caer
Notes ON THE LIGN PLANTS DESCRIBED BY LEVEILLE FROM EASTERN Asta. By “Alfred TCROCE «2 sisatecs Yip eae eee
CHROMOSOME NUMBERS AND PHYLOGENY IN THE GYMNOSPERMS. With two text figures. By Walter S. Flory ........ccccceeeeeee
APOSPORIC PARTHENOGENESIS IN A TRIPLOID since MALUS HUPE- HENSIS. With plates 179-181. By Haig Dermen ..............
FERTILIZATION IN THE BALDWIN APPLE, A TRIPLOID VARIETY. By Ha
ES PINON as 63 kh 4d ee nea SHR Oe 106 Two SpRUCE-INFECTING Rusts, CHRYSOMYXA PIPERIANA AND CHRYSOMYXA CHIOGENIS. By 7. POU a. Pee eae eee ae 109
MorPHOLOGY AND ONTOGENY OF THE SPERMOGONIA OF THE MELAMP- SORACEAE. With plates 182-188. By Lillian M. Hunter ........ 115 THE EXPERIMENTAL PRODUCTION OF PoLyPLompy. With plate 189. BE OR ats We ie oh aos oy + Wk e's Be GE STUDIES IN THE OAKS OF THE MOoUNTAINS OF NORTHEASTERN Mexico. By Cornelius H. Mueller .........ccccccccccseccceces 1
PHYTOGEOGRAPHIC STUDIES IN THE ATHABASKA—GREAT SLAVE LAKE Recion. With plates 190-194 and map. By Hugh M. Raup .... 180 PavroctocaaPsc STUDIES IN THE ATHABASKA—GREAT SLAVE LAKE REGION (contin.). With plates 195-200. By Hugh M. Raup .... 241 NoTES ON THE LiGNEOUS PLANTS DeEscRIBED BY LEVEILLE FROM EASTERN AsIA (contin.). By Alfred Rehder ......... 0.000005. 316 THE GeNuS CESTRUM IN GUATEMALA. By C. V. Morton ......... 341 A New SPECIES oF ACER FROM GUATEMALA. By Alfred Rehder .... 350
POLYPLoIDy AND GEOGRAPHIC DIsTRIBUTION IN SPIRAEA. By Karl Sax 352
Ai a A a eo Ae a a ee ee er ee Yer ee Ye Te a ee ee
iv TABLE OF CONTENTS
THe ARNOLD ARBORETUM DuRING THE FiscAL YEAR ENDED JUNE
ct ie |X, ee re rer eee eer errr errs yor rer ery 57 BIBLIOGRAPHY OF THE PUBLISHED WRITINGS OF THE STAFF AND
STUDENTS, JULY 1, 1935-JUNE 30, 1936 ....... cee cece eeceeeces 366 STAFF OF THE ARNOLD ARBORETUM, 1935-36 ....... 00sec eee ences 370 COMNORPINME ook bac ck pace ee cace P50 bd) wu RW e Rea ee areeTEes 371 > ee ee en ee ee ee eRe ee rr ree rr TT 373
JOURNAL
OF THE
ARNOLD ARBORETUM
VotuME XVII. JANUARY, 1936 NuMBER |
STUDIES ON THE BIOLOGY OF GYMNOSPORANGIUM GLOBOSUM FARL.
Toa. MaAcLACHLAN
With plates 166-175 and two text figures
TABLE OF CONTENTS
i TA TROUUCTION fe 64-24 6 0.05-05-556 8 E46 whe SRDS re es 1 II INGMENCEATOU RE op oiis Go ceed oi eee e teed e eee ene s 2 Ill RANGE AND EcoNnoMIc IMPORTANCE OF G. GLOBOSUM ........ 3 IV SYMPTOMATOLOGY OF THE DISEASES CAUSED BY G. GLOBOSUM.. 5 V Factors AFFECTING SPORE GERMINATION ......--eeeceeees 14 VI Tee Aree OF THE LELTAL SCORUS. .<sJ<0cns eae meee canes 16 Vil . INFECTION OF THE RED CEDAR 2.5 os cc bcss .citiew eee vet eee 18 APUEL PSUSMARY oo eee BNo 65k ea hs 5 45a SRR COO Oe 19 IX Pe) < (eh g ACL 6 1. 5 ¢ ee er err rere ark etree cera er era 20 x PLRIAOGE AT AY aoc se oe a Oe aes HE Ree awa 20 XI EXPLANATION OF LA DES crerettietcunts tite ceetaniechete sche aleus opeterere aye oe
I. INTRODUCTION
Stnce the pioneer work of Farlow (1880), who ascertained the alter- nate hosts of Gymnosporangium globosum Farl., many observations on various phases of the biology of this rust have been recorded; these observations, however, do not afford a complete survey of the biology of G. globosum, More than one hundred suscepts have been added, since 1880, to the host list, yet observations of the writer warrant the con- clusion that the number of suscepts, conservatively estimated, is more than six hundred; moreover, little information was previously available
2 JOURNAL OF THE ARNOLD ARBORETUM [voL. Xv
regarding their relative susceptibility. Many aspects of the progressive development of the symptoms and signs of the diseases caused by this rust have not been described in full; the incompleteness of this knowl- edge becomes more evident with the increase in the number of known hosts. No definite information has been available concerning the time when the aeciospores of G. globosum germinate or the method by which they produce infection on the Juniperus hosts; knowledge with respect to this question is a prerequisite to any attempt to control the rust on the red cedar. Aside from an academic consideration of the biology of G. globosum, this rust is of increasing economic importance and is causing great damage to both the ornamental and the orchard hosts in localized areas throughout the eastern part of the United States.
These desiderata have led the writer to investigate more fully the biology of G. globosum. A determination of the hosts and their rela- tive susceptibility has already been published (MacLachlan, 1935). The numerous inoculations and field observations that were conducted at that time afforded an opportunity to carry on extensive studies on other aspects of the biology of G. globosum; the results of these investi- gations have been incorporated with those of other writers and are now presented.
Il. NOMENCLATURE
The following names, chronologically arranged, have been given to the rust now known as Gymnosporangium globosum Farl. The stage in the life cycle of the rust to which the name refers precedes the name. III Gymnosporangium globosum Farl. in Bot. Gaz. 11: 236, 239
(Sept. 1886). I Aecidium _ bes Oxyacanthae Schw. Syn. Fung. Carol. 66 40), no. 432 (1822). I Caeoma Cvlindrites Lk. var. Crataegi-punctatae Schw. Syn. Fung. Am. Bor. (Trans. Amer. Phil. Soc. Il. 4: 294, no. 2899a. 32).
III Gymnosporangium fuscum DC. var. eal Farl. Gymno- sporan pl. 1, f. 7-11 (188 III Gymnosporangium Sabinae (Dicks. ) ie var. globosum Trel. rans. Wisc. Acad. 6: 133 (29) (July 1866). Ill es ee globosum Farl. in Bot. Gaz. 11: 236, 23 (Sept. 1886). I Roestelia lacerata y & zg Thaxt. ex Farl. in Bot. Gaz. 11: 240 (Sept. 1886). I Gy Wane heroin globosum I Thaxt. in Rept. Conn. Exper. Sta. 14 (20 891).
‘oO
I Roestelia globosa Shear, N. Y. Fungi Exsicc. no. 79 (1893). Corrected label.
1936] MacLACHLAN, GYMNOSPORANGIUM GLOBOSUM 3
III Puccinia globosa Kuntze, Rev. Gen. Pl. 37: 507 (1898). III] Roestelia globosa II] Kuntze in Bot. Centralbl. 77: 300 (15 Feb.
1899). III Tremella globosa Arth. in Proc. Ind. Acad. 1900:136 (June 1901)
I Aecidium globosum Farl. Bibl. Index N. Amer. Fungi, 49 (1905). III Aecidium globosum Arth. in Result. Sci. Congr. Bot. Wien, 1905: (1906).
The names Gymnosporangium globosum Farl. and Roestelia globosa Shear stand as the authentic names for the III and I stages, respectively. The name Gymnosporangium globosum Farl. is now accepted as refer- ring to either stage of the rust and is so used throughout this presentation.
III. RANGE AND ECONOMIC IMPORTANCE OF G. GLOBOSUM
A difficulty in determining the exact range and economic importance of G. globosum arises due to the many reports in the literature which refer collectively to the three rusts, G. globosum, G. Juniperi-virginianae, and G. clavipes. Nevertheless, sufficient evidence is extant to make possible accurate determinations of both range and economic importance of G. globosum.
Gymnosporangium globosum is confined in its range to the eastern and central parts of United States and to the southern parts of Ontario and Quebec. In Fig. 1 the distribution by states has been plotted; the circles and dots indicate the states in which the diseases caused by this rust have been reported on the Juniperus and pomaceous hosts, respectively.
A review of the literature indicates that the prevalence of this rust is steadily increasing; several factors may be involved in this phenomenon. Jones and Bartholomew (1915) offer the suggestion that the alternate hosts, the red cedar and the wild pomaceous hosts such as crab-apples and hawthorns, multiply more rapidly in open pastures and waste cut- over lands than they did in the original forests. The orchardist and ornamentalist, as well as bringing the alternate hosts closer together, have introduced on their estates many susceptible pomaceous trees that may serve as hosts. Whether or not the rust is increasing in virulence is open to question.. The rapid increase in scientific investigation during recent years, which brings to light new host species and through the discovery of these has extended the known range of the diseases caused by this rust, may give an appearance of increased prevalence and virulence.
Gymnosporangium globosum is causing more damage in some states
4 JOURNAL OF THE ARNOLD ARBORETUM [voL. Xvi
than in others; this, as well, may be due to the influence of man, who is constantly bringing into close range of each other, susceptible alter- nate hosts of the rust. In the eastern part of New York State, for example, this rust is almost on a par with G. Juniperi-virginianae with respect to the damage which it is causing. Stewart (1910) reports severe infection on Kieffer pears in an orchard at Long Island. According to W. D. Mills (Haskell, 1929) G. globosum was unusually prevalent in Duchess and Greene counties on the foliage of pears in 1928. Thomas and Mills (1929a) consider G. globosum as one of the three rusts (G.
Fic. 1. DistRIBUTION OF GYMNOSPORANGIUM GLOBOSUM BY STATES
globosum, G. Juntpert-virginianae and G. clavipes) destructive to apples in eastern New York State and report the occurrence of G. globosum on at least thirteen varieties of apples, with severe infection in some in- stances. In 1930 they (Thomas and Mills, 1930) list twenty-three varieties of apples on which the disease caused by this rust has been found. H. E. Thomas in a letter, dated Aug. 27, 1930, to the Plant Disease Reporter (Anonymous, 1930) writes that G. globosum was more common that year on the foliage of apples in Essex County than was G. Juniperi-virginianae. Another report (Anonymous, 1930) by W.S. Fields states that all the Crataegus plants of all kinds on an estate at Locust Valley, Long Island, were heavily infected. Other writers who have reported this rust in New York State include Grier (1925),
1936] MacLACHLAN, GYMNOSPORANGIUM GLOBOSUM 5
Martin (1925), Salisbury (1929), Crosby, Mills and Blauvelt (1929), Barrus, Boyd and Wood (1931) and Miller, Stevens and Wood (1933).
Severe local infections have been observed by the writer in eastern Massachusetts. Loci of infection may be found on estates where suscep- tible ornamental Crataegi are planted in vicinities that embrace the red cedar, and in open pastures and waste lands where wild Crataegus and Malus species are within close range of the cedar. Apparently no seri- ous outbreaks of the diseases caused by this rust have occurred, as yet, in commercial orchards in Massachusetts.
States in which severe to moderately severe local infections have been reported include Connecticut (Clinton, 1903), Florida (Martin, 1925), Illinois (verbal report from Morton Arboretum), Michigan (Martin, 1923), Minnesota (Martin, 1920) and Ohio (Martin, 1923). Bliss (1933) states that this rust is common on apple in Indiana.
Reports would indicate that the rust is not of great economical im- portance in Alabama (Underwood and Earle, 1897), Iowa (Bliss, 1933), Kansas (Bartholomew, 1899), Maine (Morse and Lewis, 1910) (Miller, Stevens and Wood, 1933), New Jersey (Cook, 1917), Ontario (Con- nors, 1934), Wisconsin (Jones and Bartholomew, 1915) and Vermont (Anonymous, 1928). Martin (1925) reports G. globosum as occurring in Alaska.
IV. SYMPTOMATOLOGY OF THE DISEASES CAUSED BY G. GLOBOSUM
A. Own Pomaceous Hosts
Observations were made concerning the progressive development of the symptoms and signs of the diseases caused by G. globosum as they appeared on the inoculations of the respective pomaceous hosts (Mac- Lachlan, 1935). For the sake of convenience the progressive develop- ment of the symptoms and signs will be considered from both the mor- phological and histological viewpoints as they appeared on the foliage of Crataegus; modifications of these symptoms and signs as they appeared on the foliage of other hosts and on the flowers, fruit and twigs, will follow; additional observations made by other writers will be included throughout the presentation.
(a) On the foliage of Crataegus.
The first morphological evidence of the disease caused by this rust is the exhibition of very small light-colored areas or flecks on the upper surface of the leaf. These appear within ten to twelve days after inocu- lation. In rare cases no further symptoms ever occur except that the
6 JOURNAL OF THE ARNOLD ARBORETUM [voL. Xvi
flecks may turn brown; normally, though, these flecks become more distinct and within approximately fifteen days after inoculation (see Table I) they show as bright yellow lesions varying from one to more than ten millimeters in diameter. Sections at this stage of development reveal intercellular hyphae among the palisade and mesophyll cells (Plate 168, Fig. 5) as well as densely intertwined masses of hyphae between the epidermis and palisade layer. These masses of hyphae are the spermogonia primordia which develop to form the mature spermo- gonia approximately twenty-three days after inoculation (see Table I). The mature spermogonia appear on the upper surface of the leaf, rarely on the lower, as small raised points in the center of the lesion. On rupturing the epidermis of the leaf the spermogonia exude a sticky fluid in which abundant spermatia may be found. In Plate 173, Fig. 1 may be seen a photograph of these spermatia (mag. « 545), obtained by placing a drop of the spermogonial fluid on a glass slide. Weimer (1917b) has illustrated the type of spermogonium in cross section.
TABLE I DATA ON THE TIME OF OCCURRENCE OF THE SYMPTOMS AND SIGNS OF THE DISEASE CAUSED BY G. GLOBOSUM ON THE FOLIAGE OF CRATAEGUS!
. days after inoculation No. Symptoms and signs minimum maximum — average species
Ist appearance of lesion 13 Zz 15.1 38 Ist exudation of spermogonial
uid 30 229 39
Spermogonia turned black 28 54 41.5 35
Ist appearance of swelling 47 59 530 at
Ist appearance of aecia vf 111 96.0 48
Exudation of the spermogonial fluid continues for approximately twenty days (see Table I), following which the spermogonia turn black and are quite conspicuous on the yellowish background (Plate 171, Fig. 1). Other than the color change of the spermogonia, little change in the lesion can be noted except for the formation, in many instances, of reddish borders around the lesions. These borders while manifested to a greater extent on some hawthorns than on others are not consistently formed even on a single host. Severe infections where more than fifty percent of the leaf area is diseased may cause yellowing of the leaf and defoliation at this stage in the development of the rust; scattered in-
1These data were obtained from typical representatives of the various groups of Crataegus inoculated with G. globoswm Farl. in May, 1932
1936] MacLACHLAN, GYMNOSPORANGIUM GLOBOSUM 7
fections, however, cause only minor injury to the leaf except for reduc- tion of photosynthetic area.
Swelling on the lower surface of the leaf, opposite the spermogonia may be observed approximately ten days after the spermogonia cease exudation (see Table I). Sections of the lesions reveal that this swell- ing is due to both hypertrophy and hyperplasia of the mesophyll tissue, resulting in long palisade-like cells closely packed together. Hyper- trophy of the leaf is confined to the area occupied by the lesion and may increase the thickness of the leaf to more than five times normal (Plate 168, Fig. 1). It is not until this stage in the development of the rust that severe leaf killing can be caused by relatively few lesions per leaf; this is especially true of vein infections (MacLachlan, 1935a).
Following the time during which hypertrophy of the lesion takes place, minute swellings may be seen on the lower surface, rarely on the upper surface, which, within two or three days time, break through the epi- dermis and appear as greyish acuminate cylinders about one millimeter in diameter. These are the peridia of the developing aecia and are first evident approximately ninety-six days after inoculation (see Table I). The peridia are not arranged in any definite pattern except when they occur on veins or petioles; in that case they may be arranged in two rows, one on each side of the vein or petiole (Plate 167, Fig. 4); they may develop to a length of more than six millimeters and may vary in number from one to more than fifty per lesion. Within each peridium are the aeciospores which are released by the splitting and shredding of the peridium a short distance from the distal end (Plate 171, Fig. 2). Unlike G. Juniperi-virginianae the peridial cells do not recurve with changes in humidity to release the aeciospores; the latter are shed through the shredded region of the peridium. High winds as well as the effect of one leaf rubbing on another have a marked influence in break- ing up the peridium to release the aeciospores.
The peridial cells, as described by Kern (1911), are “broadly lanceo- late in face view, 15-23 %* 60~90 uw, linear-rhomboid in side view, 13-19 u thick, outer wall about 1.5 p thick, smooth, inner and side walls 3—5 uw thick, rather densely rugose with ridge-like papillae of varying length.” As has been demonstrated by Thomas and Mills (1929b), the peridial cells are shorter and broader towards the distal end of the peridium and do not separate easily from each other.
The aeciospores are borne in chains from the base of the aecial cup and when mature are, as described by Kern (1911), “‘globoid or broadly ellipsoid, 15-19 & 18-25 wu, wall light chestnut brown, 1.5—2 w thick, finely verrucose.” In Plate 173, Fig. 2 may be seen a photograph of
8 JOURNAL OF THE ARNOLD ARBORETUM [voL. xv
aeciospores at the same magnification as the spermatia in Fig. 1. Two of the aeciospores (lighter color in the figure) were colorless; these colorless spores were found occasionally and proved to be viable.
Sections reveal that the aecium itself is long and narrow and deeply embedded in the hypertrophied tissue, the base of the aecium approxi- mating that of the palisade layer of the leaf (illustrated by Weimer, 1917b). In Plate 168, Figs. 1 and 2 may be seen photographs of a longitudinal section through several aecia; the non-median sections give the appearance that the aecia begin at various levels throughout the hypertrophied tissue.
The great majority of the aeciospores are distributed by the middle of September. Remnants of the peridia may remain on the lesion until the leaves normally drop, but usually the circular pits in the aecial cups are all that can be seen at this time. Sections through the lesions after the leaves have dropped reveal very few viable aeciospores; most of those that still remain in the base of the aecium are hollow and sterile.
Insects have been observed feeding on the aeciospores during August. The peridia containing the aeciospores are eaten out first, then the insects, after gaining entrance by way of the pits into the aecial cups, proceed to break down the whole interior of the lesion.
(b) On the foliage of pomaceous hosts other than Crataegus.
Pyrus: The lesions on the ornamental species are smaller than those found on Crataegus. They vary greatly. On some host species they are very minute, being less than a millimeter in diameter and producing a few spermogonia only (e.g. P. serrulata Rehd.); others may exhibit two or three aecia on each lesion (e.g. P. Balansae Decne.) ; still others may exhibit lesions which are two to three millimeters in diameter with several aecia each (e.g. P. betulaefolia Bge.) (Plate 166, Fig. 4). On P. betulaefolia the red borders around the lesions are conspicuous. Stewart (1910) describes the lesions on Kieffer pear as being bright yellow and pin-head in size on June first. He states, further, that by June fifteenth the lesions had turned brown to black with conspicuous red borders on the upper surface of the leaf but without the red borders on the lower surface. He adds that a few aecia are formed on the upper side of the leaf, but most of them are on the lower side, frequently on each side of the mid-rib. Hesler and Whetzel (1917) describe the lesions as being orange-colored in June, one quarter to one half inch in diameter with red borders showing on the upper side of the leaf. In the fall the lesions turned dark on the under side of the leaf and at this time of year showed no red border.
1936] MacLACHLAN, GYMNOSPORANGIUM GLOBOSUM 9
Sorbus: The lesions in all cases were very small, rarely measuring more than one to two millimeters in diameter, with an average of three to five aecial horns per lesion (Plate I, Fig. 3). As in Pyrus the lesions on certain species died and turned brown shortly after the spermogonia appeared.
Malus: Inoculation on ornamental apples resulted in small lesions that rarely measured more than one to two millimeters in diameter (Plate 166, Fig. 5). Certain species exhibited spermogonia only. Crosby, Mills and Blauvelt (1929), Sherbakoff (1932) and others have reported this small type of lesion on commercial apples and refer to it as one method of distinguishing G. Juniperi-virginianae from G. glo- bosum. Bliss (1931), by culture, obtained flecking only on nine vari- eties of commercial apples.
(c) On the flowers and fruit.
Infection of flowers and fruit of Crataegus is relatively rare when compared to the prevalence of foliar infection. All parts of the flower including pedicel, ovary, sepals, petals and even stamens may be attacked. Infections on the stamens and petals cannot persist but infec- tions of the ovary result in either dwarfing and killing of the fruit (Plate 167, Fig. 1) or premature ripening and dropping of the fruit. Abundant spermogonia and aecia may be produced (Plate 167, Fig. 2). A cross section of a diseased fruit reveals that only a portion of the fruit is affected. In Plate 168, Fig. 3 may be seen a photograph of such a section; the diseased area is represented by the darkened strip on one side of the section. On the opposite side of the section may be seen some of the diseased area of a second lesion. As in the leaf, hyper- trophied, palisade-like, densely packed host cells are evident. The fungal mycelium is intercellular and produces haustoria in the host cells. Some mycelium can be seen ramifying among the host tissue beyond the hypertrophied area.
Fruit infection of Pyrus was not observed in the Arnold Arboretum. Stewart (1910) reports the Kieffer pear as suffering infection from this rust at Long Island, New York. In particular he finds the diseased fruits are very small and misshapen, usually exhibiting circular black areas devoid of aecia, although a few show aecia. Hesler and Whetzel (1917) report somewhat similar symptoms on the fruit of pear.
Fruit infection of Malus has not been observed by the writer. Gar- man (1899), speaking collectively of G. Juniperi-virginianae and G. globosum, reported these rusts on the fruit of Malus; he found the in- fection to be most abundant on crab apples.
10 JOURNAL OF THE ARNOLD ARBORETUM [voL. XvII
(d) On the twigs.
Infection of the twigs of Crataegus is relatively rare and occurs on the current year’s growth only. Both spermogonia and aecia may be produced (Plate 167, Fig. 5). Similar infections of the thorns and axillary buds have also been observed (Plate 167, Fig. 3). A cross section of a diseased twig reveals that the infection is confined to the cortical region (Plate 168, Fig. 4). The same typical, densely packed, hypertrophied cells that occur in the leaves and fruit may be seen in the diseased area. The aecial cups, however, appear shorter and broader than those exhibited in diseased leaves. Several diseased twigs were tagged in the fall of 1932 to determine if the mycelium might be perennial in the twigs; such was not evident as the twigs were all dead the following spring.
B. On JuNripeRus Hosts
The advanced stage of the disease caused by G. globosum on the Juniperus host is expressed in the form of a woody gall attached firmly to a twig that may be more than two or three years old. This condition led early observers to conclude that infection originated in the stem. Farlow (1880) who named this fungus stated that the disease is first manifested by bursting through the stem at the point of attachment of the leaves. Kern (1911) described the gall as being caulicolous. Stewart (1915) gave an elaborate account of histological studies made, which he interpreted as showing that the gall originates from the axils of leaves and are evidently transformed axillary buds. Weimer (1917a), after examining Stewart's slides, showed that Stewart mistook an axillary bud for a gall. He (Weimer, 1917a) gave a full description of a normal juniper leaf and presented definite proof that the gall may originate in the leaf. Bliss (1933) has substantiated the results of Weimer and has illustrated stages in the progressive development of the gall on the leaf. Several brief descriptions of the mature gall as it occurs on the red cedar may be found in the literature; these descriptions are given, for the most part, as an aid in distinguishing the galls caused by G. globosum from those caused by G. Juniperi-virginianae.
To obtain a more complete picture of the progressive development of the various symptoms and signs of the disease caused by G. globosum on the red cedar, the writer has carried on further studies both morpho- logically and histologically. Material for study was obtained from a locus of infection that was isolated from other Gymnosporangium rusts. The symptoms and signs will be considered as they occur on the red cedar in eastern Massachusetts.
1936] MacLACHLAN, GYMNOSPORANGIUM GLOBOSUM 11
The initial manifestations of the disease caused by this rust may be found on one year old leaves of infected red cedars about the first. of August. The exhibition of a yellowish chlorotic zone or band on the leaf is closely followed by the formation of a slightly raised area on the inner (upper) surface of the leaf; the lifting of the epidermis over the point of infection is due to the development of a very young gall underneath.
Longitudinal splits in the epidermis immediately over the raised area allow the young gall to emerge. On long subulate leaves the gall may occur at any place throughout the length of the leaf; in Plate 169, Figs. 2 and 3, may be seen young galls developing near the tip, in the middle and near the base of the leaf. The usual type of leaf and young gall that is found is illustrated in Plate 169, Fig. 1; the leaves are short and the gall must of necessity be brought in close contact with the stem, a phenomenon which led early investigators to conclude that the gall originates in the stem at the base of the leaf.
A longitudinal section of a portion of a twig bearing two normal leaves as well as a diseased leaf, at approximately the same stage of gall de- velopment that is shown in Plate 169, Fig. 1, is illustrated in Plate 170, Fig. 1. A comparison of the two normal leaves with the diseased leaf reveals marked hypertrophy and hyperplasia of the mesophyll of the latter. Distortion of the vascular system may also be seen. Figs. 4 and 5 of Plate 170 exemplify further the distortion of the vascular elements. In Fig. 4 parenchymatous cells may be seen separating the vascular elements. In Figs. 2 and 3 a corky layer, several cells thick and com- pletely surrounding the portion of the gall not in contact with the twig, may be seen. This corky layer usually cuts off the tip of the leaf; it is of interest to note in Figs. 3 and 4 the sharp severing of the vascular strand. The tip of the leaf that is cut off together with the mycelium of the rust that may be found in it dies; remnants of this dead portion may be found on the gall for two or three years afterwards. Intercellular mycelium is abundant throughout the gall tissue but haustoria were not observed at this stage.
The gall as seen in late autumn and winter is smooth, shiny and mahogany red in color, rarely exceeding a diameter of half a centimeter. This is in striking contrast to the larger, greenish and more or less con- voluted gall of G. Juniperi-virginianae.
About the last’of March of the following spring light orange colored markings may be seen over the surface of the gall. These markings represent early stages in the rupturing of the corky epidermis and are caused by the underlying masses of hyphae—the telia primordia—from
12 JOURNAL OF THE ARNOLD ARBORETUM [voL. XVII
which the teliospores arise. Progressive stages in the maturation of the teliospores from this dense hyphal layer are shown in Plate 171, Figs. 5, 6 and 7. Dodge (1918) has published an extensive cytological study of the development of the teliospores and has illustrated progressive stages of their maturation. The teliospores mature progressively from the centre towards the periphery of the layer of telia primordia. The corky epidermis is ruptured over the maturing teliospores and the latter emerge as a cCinnamon-brown, pulvinate mass (Plate 171, Fig. 4). The irregu- lar outline of this rupture readily distinguishes the galls caused by this rust from those caused by G. Juniperi-virginianae since the telial sori of the latter emerge through circular openings. The teliospores are borne on pedicels and the extension of these pedicels, as the teliospores pro- gressively mature from the centre towards the periphery of the basal layer of mycelium, enlarges the sorus to form a tongue or wedge-shaped structure, the centre of which may be hollow. This cavity is due to the developing teliospores around the peripheral regions of the sorus; these push up the mature teliospores in the centre and as a result the pedicels of the latter teliospores are broken off.
A mature telial sorus, if kept dry, is a tongue-like or wedge-shaped flange 1-3 mm. broad by 2—5 mm. long at the base and 6—12 mm. high. A section through such a sorus reveals a dense cinnamon-brown layer of teliospores, several spores thick, over the surface. Under the spore layer is a dense white mass composed entirely of long pedicels, one for each teliospore. The peripheral pedicels still remain attached to the layer of mycelium from which they arose; while the pedicels in the central region, which have been broken off from the layer of mycelium from which they arose, are suspended to form a conical amphitheatre-like cavity in the centre.
The teliospores are typically two-celled and are, as described by Kern (1911), “ellipsoid, 16-21 % 37-48 u somewhat narrowed above and below, slightly constricted at the septum, wall pale cinnamon-brown, 1-2 uw thick; pores 2 in each cell, near the septum.” Pammel (1905) has described one-celled teliospores. No single-celled teliospores have been observed by the writer; three- and four-celled teliospores, how- ever, as well as four-celled teliospores with one cell abortive have been found (Plate 173, Fig. 3).
Studies with respect to the nature of the pedicels of the teliospores and the effect of wetting will be considered later (page 16).
Wetting of the telial sori by rains in May causes them to expand to many times their original size. Progressive stages in the expansion of sori on galls as well as on the small infections that take place on the long
1936] MacLACHLAN, GYMNOSPORANGIUM GLOBOSUM 13
subulate leaves are illustrated in Plate 172. The hygroscopical nature of the pedicels (see page 16) is responsible for this expansion. If only a portion of the teliospores are mature at the time of wetting, the telial mass may, on drying, shrink to somewhat of its original shape and size. If, however, all the teliospores are mature at the time of original wetting and the humidity remains high for a sufficient length of time to fully expand the telial mass, the latter, on drying, will shrink to an amorphous sheet or strand and later fall away from the gall. Under favorable weather conditions practically all the teliospores are mature within three to five weeks from the time of their initial appearance.
Germination of the teliospores results in the production of basidio- spores. Progressive stages in the production of the latter are shown in Plate 174, Fig. 1. The basidiospores may be seen as a powdery, some- what velvety coating over the surface of the expanded telial sorus and are evident about five or six hours after the expansion takes place. These basidiospores will drop from the gall of their own accord but are usually carried away by the air currents. The basidiospores are ovate, flattened on one side, slightly tapering towards the end of attachment to the sterigmata; thin-walled (less than 1 u thick); protoplasm light orange- yellow in color containing numerous oil droplets which tend to aggregate (Plate 173, Fig. 4); size, 12-16 (average, 13.7 1) XX 7-11 (average, 8.9 1). Under favorable conditions they germinate immediately after their formation to form relatively long germ tubes (Plate 174, Fig. 4). If the spores are allowed to dry shortly after the germ tubes begin to develop, small secondary spores may be formed (Fig. 5).
As a rule, the majority of the teliospores have germinated by May 25. The dried up remnants of pedicels and empty teliospores blow away leaving a smooth orange colored scar bordered by the lacerated edges of the broken corky epidermis. This scar soon turns greyish brown and may persist on the gall surface during the remainder of the life of the gall (Plate 171, Fig. 4). Cross sections through the gall reveal that the tissue immediately underlying this scar dies and a corky layer develops, segregating this dead area from the remaining living tissue of the gall (Plate 170, Fig. 5).
The gall continues growth during the summer causing distortion of the neighboring twigs (Plate 169, Fig. 5) and in some cases killing of the twig beyond the gall (Plate 169, Figs. 4 and 6). The original single vascular strand of the leaf now shows as a many-branched structure (Plate 170, Fig. 5). New growth beyond the gall as well as the natural dropping of the older leaves and the formation of a corky epidermis on the twig surrounding the gall give the appearance that the gall originated
14 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XVI
in the stem rather than in the leaf. This illusion is further substantiated by the tendency of the gall to become woody in appearance as it grows older (Plate 169, Fig. 7).
The gall is commonly referred to as being perennial in nature. For how many years it will live and produce teliospores is not known; in Plate 169, Fig. 7 is shown a gall that produced teliospores for three con- secutive years. The telial sori are formed each year between the scars where teliospores were produced the previous years (Plate 171, Fig. 4). It is doubtful that many of the galls persist for more than three or four years, because, by that time the surface of a gall would be completely covered with dead tissue resulting from the formation of the telial sori. Long subulate leaves on which infection takes place usually drop after the first crop of spores; these infections, then, can persist for one year only.
V. FACTORS AFFECTING SPORE GERMINATION A. Tue Time Factor
The teliospores exhibit a low and inconsistent percentage of germina- tion when tested immediately after their appearance on the gall. If twigs bearing galls on which teliospores are just emerging are placed in water in the greenhouse, abundant germination can be obtained, regu- larly, in a week to ten days time. The maturation period of the telio- spores is longer, depending on the weather conditions, when the galls bearing them are left in the field. Normally, the teliospores will all have germinated within six to eight weeks after their formation. Under low temperature conditions, however, they have the potential ability to remain viable for a much longer time; in Plate 175, Fig. 3 may be seen severe infection obtained on leaves of Crataegus Jonesae Sarg. using, as inoculum, teliospores that had been kept in the refrigerator for one year.
The basidiospores have the ability to germinate immediately after their formation. The length of time that they will live is still a question; basidiospores of G. Juniperi-virginianae will live for many days above a humidity of 25% and below a temperature of 25° C. (MacLachlan, 1935b
Aeciospores will not germinate to the extent of more than 2 or 3% at the time of their formation. Nevertheless, if hawthorn leaves bear- ing aecia are collected in August before the aeciospores are shed and placed in a refrigerator at 0° C., abundant germination can be obtained within a month’s time and by the first week in October more than 80% germination can be obtained consistently. By keeping the infected leaves at this temperature, the same high degree of germination can be
1936] MacLACHLAN, GYMNOSPORANGIUM GLOBOSUM 15
obtained until the first of March of the following year; after this date, however, the percentage of germination that can be obtained falls off rapidly. Whether the infected leaves are kept in a dry container or a moist chamber makes no difference in the percentage germination obtained. It may be noted that on one occasion more than 75% germi- nation was obtained in August, 1932, using aeciospores taken from un- broken peridia of an infected leaf of Crataegus shirleyensis Sarg.
B. Tue Humipity Factor
Tests in the laboratory revealed that none of the three spore forms would germinate, even in saturated humidity (spores placed on cover glasses over water ), unless in direct contact with a drop of water. Excess water, however, causes irregularity in the percentage germination; basidiospores, especially, show a very low percentage of germination when immersed in large drops of water; under the same conditions aeciospores are not consistent in the percentage of germination that may take place; teliospores germinate in excess water but instead of produc- ing basidiospores the promycelia grow to great lengths, presumably to come in contact with the air, and may exhibit long side tubes (Plate 174, Fig. 3), or break up to form elongated spore-like bodies (Plate 174, Fig. 2). Farlow (1880) reported a type of spore, similar to that shown in Fig. 2, which he observed when the telial masses were quickly dried after moistening. He also found that, on remoistening, these spore-like bodies would send out germ-tubes similar to those of normal basidio- spores.
In the laboratory, optimum humidity conditions for germination of any of the three spore forms was obtained by placing the spores on a cover glass and inverting the latter on a Van Tieghem cell in a petri-plate lined with wet filter paper; enough water of condensation would accu- mulate on the lower (spore) surface of the cover glass to give optimum humidity conditions for germination. In the field, very satisfactory humidity conditions for infecting the pomaceous host were obtained by painting the leaves with an aqueous suspension of the teliospores and enclosing the inoculated twig in a celluloid cylinder, the ends of which were plugged with moist sphagnum.
C. Tue TEMPERATURE FACTOR Temperature has a marked effect on the percentage of germination that may be obtained. Miller (1932) found 24° C. to be the optimum temperature for the germination of teliospores and aeciospores, and 16° C, the optimum for basidiospores. In Table II may be found data
16 JOURNAL OF THE ARNOLD ARBORETUM [VoL. XVI
TABLE II
DATA ON THE PERCENTAGE GERMINATION OF THE SPORES OF G. GLOBOSUM WHEN SUBJECTED TO DIFFERENT TEMPERATURES!
Temp. Percentage germination of Ke Ge teliospores basidiospores aeciospores 2 0.0 —_ 0.0 5 0.3 0.0 4 7 0.7 7.2 8.2 10 80.4 68.7 11.3 15 90.9 79.2 70.2 20 96.3 83.9 88.7 25 98.6 63.1 60.7 30 59.8 14.5 35.3 35 0.9 0.9 0.6 38 0.0 0.0 0.0
on the percentage germination of the three spore forms when sub- jected to various temperatures; these data have been plotted in Fig. 2. Examination of Fig. 2 reveals that: (1) the teliospores have a very wide range of temperature within which more than 80% germination may take place, (2) the corresponding curve for basidiospore germination has the same general contour, except at a lower level, (3) the aecio- spores have a narrower range of temperature within which a high per- centage of germination may take place, the optimum being around 20° C., (4) below a temperature of 10° C. and above a temperature of 30° C., the percentage germination of all three spore forms drops off to almost zero.
VI. THE NATURE OF THE TELIAL SORUS
Reference was made under SyMp ocy (page 12) to the marked expansion of the telial excrescences on the galls during wet weather. In Plate 172 may be seen photographs of progressive stages in the expansion of these telia. Further studies were made with respect to this phenomenon.
It was found that the pedicels of the teliospores, when wetted, are responsible for this expansion process. No estimation has been made with respect to the number of teliospores and the corresponding number of pedicels that may exist in a single telium, yet, they must number in the millions. Crowell (1934) has given estimations which indicate that four to five million teliospores may exist in a telium of G. Juniperi-
11500 spores were counted in each sample. Counts were made 24 hours after the cultures were set up.
1936] MacLACHLAN, GYMNOSPORANGIUM GLOBOSUM 17
virginianae ; the number that may exist in a telium of G. globosum would be as many or more as the telia are characteristically larger in the latter rust. The pedicels, as well as being numerous, are relatively long; some idea of their length may be obtained in Plate 175, Fig. 5. Under strong light and on a dark background they appear white and opaque; they do not stain readily in aqueous methylene blue. In Plate 175, Fig. 4 may be seen a photograph of a pedicel that was immersed in 25% alcoholic
@-—TELIOSPORES 100 Q@-— BASIDIOSPORES O- AECIOSPORES r o 80] ra < z 2 60, uJ rT) 0 < 40) = z uJ ) tl a 20) ce) ll 7 Tt v ¥ ¥ as ) 5 10 15 20 25 30 35 40
alae A | DEGREESC.
Fic. 2. THe INFLUENCE OF TEMPERATURE ON THE acai GERMI- NATION THAT MAY BE OBTAINED FOR THE THREE SPORE FoR
methylene blue for fifteen hours; the pedicel itself did not stain to any extent but the stain was absorbed in the centre of the pedicel indicating the presence of a lumen. The hygroscopic nature of the pedicel is illus- trated in Plate 175, Figs. 1 and 2. The pedicels represented in Fig. 1 were subjected to 95% alcohol for twelve hours while those represented in Fig. 2 were placed in water for the same length of time. The two pictures are at different magnifications but when one compares the diameter of the pedicel with the size of the teliospore which it subtends, it is quite evident that the pedicels are capable of considerable expan- sion when wet; Fig. 1 illustrates the pedicel as it would appear when the telium is dry while Fig. 2 illustrates the pedicel as it would appear during
18 JOURNAL OF THE ARNOLD ARBORETUM [VoL. XvII
rainy weather in May. In Fig. 2 may also be seen small air bubbles existing in the region that was once the lumen but is now filled by the expanded wall of the pedicel. On the other hand, as may be seen in Fig. 1, the lumen is relatively wide with respect to the diameter of the desic- cated pedicel. Taking into consideration, then, the number of pedicels present in a telium as well as their length and the diameter they may attain when wet, one can readily see how a telium may expand to the size that is so common on infected cedars during rainy periods in May.
VII. INFECTION OF THE RED CEDAR A. TIME oF INFECTION
The actual time of year that infection of the red cedar by the aecio- spores takes place is still conjectural. About 2% of the aeciospores will germinate at the time of their dispersal in late August and when one realizes the large number of aeciospores that may be released from infected pomaceous hosts it can be presumed that possibly some infec- tion may take place immediately. Nevertheless, the fact that more than 80% germination can be obtained after the fresh aeciospores have been subjected to 0° C. for about six weeks time (cf. page 14) would indicate that both time and low temperature are involved in the matura- tion of the aeciospore. This would indicate that infection takes place in late fall after frosts have occurred. The fact that a high percentage of germination can be obtained in March of the following spring, using aeciospores that have been kept at 0° C. during the interim, would not necessarily mean that infection takes place in the spring; if the aecio- spores are mature in late fall, in all probability they will have all germi- nated long before March.
B. INFECTION PROCESS
Having access to an abundance of aeciospores which would germinate readily, attempts were made to trace the method by which infection of the red cedar by this rust takes place.
Leaves of the current year’s growth were removed from potted red cedars, washed carefully and dusted on the stomatal (inner and upper) surface with aeciospores. The inoculated leaves were then inverted and placed on moist filter-paper in petri-plates. Examination 24 hours later revealed abundant germination of the aeciospores; the germ tubes were long and slender and sometimes spiralled; the protoplasm had all mi- grated to the end of the germ tubes. Within 12 hours more, the ends of the germ tubes had enlarged to form irregular and sometimes convoluted
1936 ] MacLACHLAN, GYMNOSPORANGIUM GLOBOSUM 19
haustoria-like formations. On one occasion the germ tube was observed to bend at a sharp right angle to form the haustoria-like structure over a stoma; however, no consistent orientation of the germ tubes with respect to the location of stomata could be observed. The haustoria- like structures that happened to form over stomata were smaller than those that formed on the inter-stomatal regions of the leaf epidermis. Two samples of about fifty leaves each were removed from the culture and killed for embedding thirty-six and fifty-two hours, respectively, from the time of inoculation. These were sectioned (both transversely and longitudinally), stained with safranin and light green and exam- ined under the high power of the microscope in hopes of observing the method of penetration.
The length of the germ tubes made it impossible to obtain even a series of sections that showed an aeciospore connected to its germ tube throughout its entire length. This led to a difficulty in the identification of germ tubes that were observed entering the leaf and consequently the results obtained can only be considered as indications and not conclu- sive proof. The characteristic intercellular mycelium of G. globosum was found around the mesophyll cells in the immediate vicinity of the stomata. Hyphae of the same diameter as the germ tubes were found passing through the stomata; in some instances three or four passed through the same stoma. Some of the haustoria-like structures were found to be still clinging to the surface of the leaf but no penetration from them was observed.
Whether the germ tubes that were observed passing through the stomata were those of G. globosum or of some other fungus that hap- pened to be mixed with the aeciospores cannot be said for certain. It is quite safe to say, however, that the rust gains entrance through the upper epidermis of the leaf on which the stomata exist, as the character- istic intercellular hyphae of this rust were found around the mesophyll cells immediately under the stomata. Further studies on this problem will be carried out by the writer.
VIII. SUMMARY
At least thirteen names have been given to the rust now known as Gymnosporangium globosum Farl. Of these the authentic names for the III and O & I stages of the rust stand as Gymnosporangium globo- sum Farl. and Roestelia globosa Shear, respectively. The name Gymno- Sporangium globosum Farl. is now accepted as referring to either stages of the rust.
20 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XVII
Gymnosporangium globosum is confined in its range to the eastern and central parts of the United States and to the southern parts of Ontario and Quebec. The rust is increasing in prevalence and in local- ized areas is causing great damage to both the ornamental and orchard trees; this is especially true in eastern New York State.
The diseases caused by G. globosum occur on at least ten genera of the Pomoideae and on at least three species of Juniperus. The symp- toms and signs of the diseases caused by this rust may be found on the foliage and to a lesser extent on the flowers, fruit and twigs of the poma- ceous hosts and on the foliage and twigs of the Juniperus hosts. The progressive development of these symptoms and signs as they occur on the aforementioned organs of the respective hosts have been described and illustrated.
The factors of time, humidity and temperature have been considered with respect to the percentage germination of the teliospores, basidio- spores and aeciospores that may be obtained. The age of the three spore forms as well as the temperature to which they are subjected have a marked effect on the percentage germination that can be obtained. The amount of water present also modifies the percentage germination that can be obtained as well as the type of germ tubes that may be exhibited.
All evidences indicate that infection of the red cedar by G. globosum occurs, primarily, in late autumn. From studies of the infection process it would appear that infection of this host takes place through the upper and stomatal surfaces of the leaves.
IX. ACKNOWLEDGMENTS
To Professor J. H. Faull, who afforded the writer the opportunity to study this problem and for guidance and supervision during the pursu- ance of its investigation, sincere appreciation is expressed.
The writer is also grateful to Dr. Ivan H. Crowell for many helpful suggestions and to Mr. Francis M. Bryant for technical assistance.
X. BIBLfLOGRAPHY ee (1928). Apple rust (Gymnosporangium spp.). (Plant Dis. ep. 7:
(1930). Apple rust diseases in New York. (Plant Dis. Rep. 14: i nee Barrus, M. O. oyp and J. I. Woop (1931). Plant diseases in the United eee “geen Dis. Rep. Suppl. 81: 72.) BARTHOLOMEW, FE, (1899). The Kansas Uredineae. (Trans. Kan. Acad. : J
Sci. 16: 184
1936] MacLACHLAN, GYMNOSPORANGIUM GLOBOSUM 21
Buss, D. E. (1931). Physiologic specialization in Gymnosporangium Casual ee ee 4 Te BB ye
oo The pathogenicity and seasonal development of Gymno- een in Iowa. (Iowa State Coll. Agric. and Mech. Arts Res.
CLINTON, Ge (1903). Diseases of plants cultivated in Connecticut. (Ann. Rep. Conn. Agric. Exper. Sta. Part 4: Connors, I. L. 19 4). VI. Diseases of ornamental plants. (Ann. Rep.
Coox, M. T. (1917). Common diseases of apples, pears and quinces. . J. Agric. Exper. Sta. Circ. 44: 14.
Crossy, C. R., W. D. Mitts and W. E. BiavuveEtt (1929). Protecting orchard crops from diseases and insects. (Cornell Univ. Agric. Exper Sta. Bull. 498: 36.)
CroweE Lt, I. H. (1934). The hosts, life history and control of the cedar- apple rust fungus a ala Juniperi-virginianae Schw. (Jour. Arn. Arb. 15: 165-232
Dopce, B. O. (1918). Studies ¢ on the genus Gymnosporangium—III. The origin of the teleutospore. ipcole da: 10: 182-189. ) Fartow, W. G. (1880). The Gymnosporangia or cedar apples of the
United States. (Anniv. Mem. Bost. Soc. Nat. Hist. pp. 9, 18, 34-35.) GarMAN, H. (1899). Some pests likely to be disseminated from nurseries. ieols)
Grier, N. M. (1925). The native flora of the vicinity of Cold Spring Harbor, L. I., New York. (Amer. Midl. Nat. 9: 94.
HaskeELt, R. J. (1929). Diseases of fruit and nut crops in the United Sets in 1928. (Plant Dis. Rep. Suppl. 70: 210.
Hester, L. R. and H. H. Wuerzer (1917). Manual of fruit diseases. (MacMillan aad Co., New York, pp. 341- aay
Jones, L. R. and E. T. BARTHOLOMEW (1915). pie rust and its control in Wisconsin. (Wis. Agric. Exper. Sta. Bull. ar
Kern, F. D. (1911). A biologie and taxonomic ae of the genus Gym- nosporangium. (N. Y. Bot. Gard. Bull. 7: 468-469.
MacLacutan, J. D. (1935a). The hosts of Gymnosporangium globosum Farl. and their relative susceptibility. (Jour. Arn. Arb. 16: 98-142.)
35b). The ne eal of viable basidiospores of Gymno- sporangium rusts. our. Arn. Arb. 16: a ee.
MartTIN, G. H., Jr. (1920). mie of cotton, sugar cane, forest trees and miscellaneous plants in the United ieee in 1919. (Plant Dis. Survey Supp
19
VAN Diseases of forest and shade trees, ornamental and miscellaneous plants in the United oe in 1922. (Plant Dis. Survey Suppl. 418. — (1925). Diseases of forest and shade trees, ornamental and miscellaneous plants in the United States in 1923. (Plant Dis. Rep
Miter, P. R. (1932). Pathogenicity of three red-cedar rusts that occur on apple. ae 22: 728, 73.
Mitter, P. R E. STEVENS and J. I. Woop (1933). Diseases of plants in the naa: States i in 1931. (Plant Dis. Rep. Suppl. 84: 50. Morse, W. J. and C. E. Lewis (1910). Maine apple diseases. (Maine
PAMMEL, L. H. (1905). The ‘cedar apple fungi and apple rust in Iowa. (Iowa Exper. Sta. Bull. 84: 5-16.)
22 JOURNAL OF THE ARNOLD ARBORETUM [voL. xvu
SatisBury, W. S. (1929). Cedar ae (Plant Dis. Rep. 13: 7.) SHERBAKOFF, C. D. (1932). e more important diseases of apples in Tennessee. (Univ. of Tenn. pen Exper. Sta. Bull. 145: 34-37.) Stewart, A. (1915). Anatomical study of Gymnosporangium galls. (Amer. Jour. Bot. 2: 402-417.)
Stewart, F. C. (1910). Notes on New York plant diseases, I. (N. Y. — Exper. Sta. Bull. 328: 376.
Tuomas, H. E. and W. D. Miris (1929a). Rust diseases of the apple. ie 19: 87.)
—— (1929b). Three rust diseases of the apple. (Cornell Univ. Agric. Exper. Sta. Memoir, 123.
— (1930). Rust diseases of the apple. (Plant Dis. Rep. 14: 214-215.)
Unperwoop, L. M. and F. S. Earte (1897). A preliminary list of the Alabama fungi. (Ala. Agric. Exper. Sta. Bull. 80: 212.
Weimer, J. L. (1917a). The a and development of the galls pro- uced by two cedar rust fun (Amer. Jour. Bot. 4: 241-2
(1917a). hree ae rust fungi, their life histories and the
diseases they produce. (Cornell Univ. Agric. Exper. Sta. Bull. 390: 527-538. )
XI. EXPLANATION OF PLATES
PLATE 166 Aecial stage of G. globosum on the foliage of pomaceous hosts. Unless signified inoculations were artificia
Fig. 1. On Crataegomes pilus andehien Bean. Note leaf killing. Fig. 2. On Crataegus pruntfolia (Marsh.) Persoon.
Fig. 3. On Sorbus americana Marsh. The lesions are very small. Fig. 4. On Pyrus betulaefolia Bge.
Fig. 5. On Malus sp. (Natural infection. )
PLATE 167
Aecial stage of G. globosum on various parts of Crataegus hosts. (Arti- ficial inoculations. ) Fig. 1. Dwarfing and killing of fruit. Fig. 2. Abundant aecia produced on basal portion of a fruit Fig. 3. Ona terminal bud that was forming in the axil of a leaf. Fig. 4. On the petiole of a leaf. Fig. 5. On twigs of the current year’s growth.
PLATE 168 Histological sections of Crataegus showing diseased areas. Fig. 1. Cross section through a diseased Crataegus leaf showing the amount of hypertrophy that may occur in the lesion. Longitudi-
The section also uae: a near-median section of an aecium.
Fig. 3. Section through a diseased Crataegus fruit. Hypertrophied area of the lesion shows as a dark strip in the photograph. This area is composed of densely packed, eee like, cells similar to those which occur in the foliar lesions
1936] MacLACHLAN, GYMNOSPORANGIUM GLOBOSUM 23
Fig. 4. Cross section through a lesion on a Crataegus twig. ee tite and intercellular mycelium are confined to cortical regio The aecial cups are eet stierily shorter and broader aan those found on the foliage. Big so, Intercellular mycelium in the mesophyll of a diseased a eaf under a spermogonium before hypertrophy has s
PLATE 169 s of G. Pa eat in various stages on leaves and twigs of Juniperus ie Redes -cedar ). ig. 1. The eee type of young galls that may be seen aia through
Fig. 4. A one-year-old cal that has caused the death of the twig alate it.
Fig. 5. A two-year-old gall that is causing distortion of the tw
Fig. 6. A two-year-old gall that has killed the twig beyond the aon of hae ction. The dead portion of the twig has fallen away. (See
g. 4.) Fig. 7. aL old gall (3 or 4 years) that gives the appearance of having originated in the stem. The infection that resulted in this gall took place when the twig bearing it was young and covered with green leaves.
PLATE 170 Histological sections of Juniperus infected with G. paatieie ig. 1. Longitudinal section through a twig bearing two mal leaves
n infected leaf. Note hypertrophy of the a Peo leaf as
well as the distortion of the single strand of vascular elements. Fig. 2. A later stage than that shown in Fig. 1. A corky layer several cells thick has set in that surrounds the gall and cuts off the tip of the leaf. The tip of the leaf that is cut off together with the sail ralasehes that roa e found in it dies but may persist in the
or three
Fig. 3. ae portion of aii similar to that shown in Fig. 2. Note
Fig. 4. Enlarged portion of section shown in Fig. 3. ote par renchym a- like cells that are forming within the vascular strand splitting up e latter.
Fig. 5. A section through a two-year-old gall. The splitting up of the single vascular strand of the original leaf has now resulted in a
spring; a corky layer had set in, segregating the dead portion from the remainder of the gall.
PLaTE 171
Fruiting structure of G. globosum on Crataegus (Figs. 1 and 2) and
Juniperus (Figs. 3-7).
Fig. 1. Spermogonia in the center of a foliar lesion; they have turned and are quite conspicuous on the yellow background of the
lesion. Fig. 2. Typical aecia on the foliage. Note method of shredding of the peridial cells for the release of the aeciospores.
24 JOURNAL OF THE ARNOLD ARBORETUM [voL. xvi
ture the epi idermis readily Pike ge ier the galls caused by G. globosum from those caused by G. Juniperi-virginianae in fuo circular pit-like openings are formed. Note also the scars be- tween the sori; within the area occupied by these scars iat sori occurred in previous years Figs. 5, 6 and 7. Progressive stages in the development of the layer of Fie primordia that form under the corky epidermis of a gall in : corky layer is ruptured about the time that the Elie are formed.
PiaTE 172
Gelatinization of the telial excrescences. Figs. 1 and 2. Before and after wetting of two infections on a long ead leaf. No gall formation was evident on this el
Figs. 3, 4 and 5. a eal stages in gelatinization of three telial sori on ngle one-yea gall. Figs. 6 ae oe Before on after wetting of a single ecies on a long leaf. Note the formation of a gall on this Fig. 8. Gelatinization of the usual type of gall that is rae y
PLATE 173 Spore forms of G. globosum (Magnification * 545). Fig. 1. Spermatia.
Fig. 3. rae yas The usual type are two- celled. Occasionally three- ace celled spores may be found. Note the four-celled telio- ith one cell abortive. Fig. 4. Bee Note the conspicuous yellow oil droplets.
PLATE 174 Germinating spores of G. globosum.
Fig. 1. Progressive stages in the germination of a teliospore to the dase of basidiospores. (A portion of the promycelium of er cell was out of focus and does not show in the last
ic Fig. 2. pictur promycelia of a teliospore that have broken up to form spore-like bodies ae of the normal production of basidio-
WwW
nating. They occur frequently if the teliospore is in too m water.
Fig. 3. A promycelium of a teliospore that has produced long projections instead of uipeun aie This as well occurs frequently if the teliospore is in too m ,
Fig. 4. Germinating basidio
Fig. 5. A germinated ay a that has produced a secondary spor Secondary spores are frequently found when ce ‘nies | ae es are allowed to dry shortly after germination begin
Fig. 6. A germinated pre that has produced a nee branch,
Fig. 7. Germinating aeciospor
Jour. Arnotp- Ars. VoL. XVII PLATE 166
(GYMNOSPORANGIUM GLOBOSUM Farl.
Jour. Arnotp Ars. VoL. XVII PLATE 167
GYMNOSPORANGIUM GLOBOSUM Farl.
Jour. ArNoLp Ars. VoL. XVII PLATE 168
GYMNOSPORANGIUM GLOBOSUM Farl.
Jour. ArNoLp Ars. VoL, XVII PLATE 169
¥
ay sop hed i crs Sa
GYMNOSPORANGIUM GLOBOSUM Farl.
Jour. ArNotp Ars. VoL. XVII PLATE 170
Pe Set — ' BS 5 rhe ri
EAN }' ~ , uy
ee
GYMNOSPORANGIUM GLOBOSUM Farl.
Jour. Arnotp Ars. VoL. XVIT
PuaTeE 171
1 Ais
a Hy)
4 By 77, ae? | f tf, FA hi 4i4 i de ULI es
‘J Opt, ee Ss saeahy apt
® &e® 8 ‘v, &
&
GY MNOSPORANGIUM GLOBOSUM Farl.
Jour. Arnotp Ars. VoL. XVII Piate 172
(8 ab
‘
GYMNOSPORANGIUM GLOBOsUM Farl.
Jour. Arnotp Ars. VoL. XVII PLaTE 173
GYMNOSPORANGIUM GLOBOSUM Earl.
Jour. AkNo_p Ars. VoL. XVII Prater 174
GYMNOSPORANGIUM GLOBOSUM Farl.
Jour. ARNotp Ars. VoL. XVII PLATE 175
GYMNOSPORANGIUM GLOBOSUM Farl.
1936]
MacLACHLAN, GYMNOSPORANGIUM GLOBOSUM 25
PuatTeE 175
Teliospores of G. globosum.
Fig.
Fig.
Ls
w&w
aN
wm
edicels and teliospores subjected to ou alcohol for twelve hours. Note the slender thin-walled p Pedicels and teliospores subjected to ae for twelve hours. The photograph is at a larger magnification than that of Fig. 1 (com- pare sizes of teliospores) but marked swelling of the pedicels has occurred; it is the swelling of these pedicels that causes the telial sori to swell during rainy periods in May. ote the air bubbles in the pedicel subtending the spore; these are in what was once the lumen but the latter has now lost its identity due to the swell- ing of the pedicel wall.
. Severe infection obtained on the foliage of Crataegus Jonesae
Sarg., the result of inoculation using Me Ms buise that had been kept in a refrigerator at 0° or on
. A teliospore and its pedicel that were sibieceal to 25% alcoholic
methylene blue for fifteen hours. rae all of the pedicel did not stain but the stain is evident in the
Portions of pedicels of teliospores. aoe idea of their length may be obtained from this photograph.
LABORATORY OF PLANT PATHOLOGY, ARNOLD ARBORETUM, Harvarp UNIVERSITY.
26 JOURNAL OF THE ARNOLD ARBORETUM [VoL. XVII
THE LIFE HISTORIES OF MILESIA SCOLOPENDRII, M. POLYPODII, M. VOGESIACA AND M. KRIEGERIANA
LittiAN M. HuNTER With plate 176
Heretorore the life histories of ten species of Milesia in all have been determined more or less completely by Klebahn (12), Faull (3), Kamel (9, 10, 11) and Mayor (14). Four more are now added.
During the year 1933-4 it was my privilege to make studies on the genus Milesia in England. Through personal collecting and the help of others telial material of Milesia Kriegeriana (Magn.) Arth. on Dry- opteris spinulosa (O. F. Miiller) Kuntze and on Dryopteris spinulosa dilatata (Hoffm.) Underw., M. Polypodii White on Polypodium vul- gare L., M. Scolopendrii (Fuckel) Arth, on Scolopendrium vulgare Smith and M. vogesiaca (Syd.) Faull on Polystichum angulare Pres] was obtained from Ireland, and the following from England—M. Blechni (Syd.) Arth. on Blechnum Spicant (L.) With., M. carpatica (Wrob.) Faull on Dryopteris Filix mas (L.) Schott and D. spinulosa, M. Poly- podii on Polypodium vulgare, M. Scolopendrii on Scolopendrium vulgare and M. Whitei Faull on Polystichum angulare. My experiments were restricted to M. Scolopendrii, M. Polypodii, M. vogesiaca and M. Kriegeriana. All were obtained from overwintered fronds in the spring months of 1934. Teliospore formation began in late March and con- tinued through April and May. It is of interest to record that search was made from October 1933 to early January 1934 for the teliospores of M. Scolopendrii, M. Polypodii, M. Kriegeriana and M. Whitei; but uredospores only were found.
The fern fronds bearing teliospores were sent in from the field and placed in a frigidaire, where they were kept at a temperature of approxi- mately 11° C. It was found that the range of temperature over which germination of the teliospores took place was between 16° and 20° C. Material was prepared for germination in the usual way. Pieces of rust- affected fronds, spread out on wet absorbent paper in petri dishes, were tested for germination in constant temperature chambers set at various temperatures ranging between 15° and 30° C. Material set up in a similar way was left in the laboratory and a thermograph record was kept of the room temperature. Records show that teliospore germina- tion was not secured when the temperature remained constantly above
1936] HUNTER, LIFE HISTORIES OF MILESIA 27
20°; but if it dropped to 20° for a part of the time some germination was secured.
After the rusted fronds had been stored in the frigidaire they dried out noticeably; moistening before setting the teliospores to germinate was necessary. To do that the leaves were placed in tap water for a few minutes to an hour and then surplus water was removed by placing them between layers of absorbent paper before arranging them in petri dishes. Excellent germination was secured when the correct temperature and moisture conditions were maintained and basidia appeared within 14 hours to slightly over 3 days. Germination of teliospores was also obtained in Van Tieghem cells set up in the way described by Ashworth (1). The basidiospores germinated equally well on the cover glass in a thin film of moisture or on a plain agar film. With bean agar the results were not so satisfactory; normal germination did not take place and grotesque bodies formed on germination of the spores.
Basidiospores of the species of Milesia described in this paper are slightly attenuated at the point of detachment from the sterigma of the basidium. A very conspicuous oil drop is usually present in the basidiospore. Basidiospores of M. Polypodii (Plate 176, Fig. 1) tend to be more spherical than those of M. Scolopendrii (Fig. 2) and M. Kriegeriana (Fig. 3). A table of measurements made from living basidiospores is given below:
Species Shape of
of rust basidiospore Breadth Length Milesia Polypodii Spherical 8-12 u 9-12 u Milesia Scolopendrii Elongated 5-7 wu 10-12 u Milesia Kriegeriana Elongated 5-8 wu 10-12 u
In normal germination of basidiospores a germ tube is usually sent out laterally from the spore. This tube may be only a few microns long, ending in a short, bulbous, conidial-like body; or it may grow to a length of more than 90 u (Fig. 4). Occasionally the contents of the basidiospore pass into the conidial-like body which increases in size as the protoplasm enters it, and the basidiospore wall collapses. A photo- micrograph of a stained preparation showing basidiospores and basidio- spore germination of M. Polypodii is shown in Fig. 5. Short germ tubes were put out two days after basidiospores were set to germinate and in from three to four days the tubes had attained a length of more than 90 u. No further development was observed to occur when experiments were left much longer than four days. Occasionally, in addition to the usual long germ tube extending from one side of the basidiospore, a
TABLE I.
RESULTS OF SUCCESSFUL INFECTION
No. of No. of needles
First First Peridermia Date of appearance appearance of rst Date needles with peri- inoculation Host of lesions spermogonia ruptured harvested infected dermia
g 27. 1V.34 Abies alba 26 days 26 days 75 days 94 days rd
us| 27.1V.34 Abies alba 25 days 25 days 67 days 70 days 14 22 2.V.34 Abies alba 21 days 21 days 89 days 101 days 5 mee, 30. V 1.34 Abies alba 23 days 23 days — — Tree wilted before 235 peridermia were fully
a 3 develo
Es.| 4.V1.34 Abies concolor 29 days 29 days 75 days 75 days Peridermia
8 insect eaten.
< 2.V.34 Abies alba 26 days 26 days 89 days 101 days s=2] 28.V.34 Abies concolor 21 days 25 days — 82 days Peridermia Sv insect eaten S$s mostly immature.
g&] 1.V1.34 Abies concolor 26 days 26 days
gs
8 588 30.1V.34 Abies alba 23 days 23 days — 62 days 4 Tree wilted before aS peridermia developed. S's = 2.V.34 Abies alba 21 days 21 days 99 days 108 days 28
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HUNTER, LIFE HISTORIES OF MILESIA
1936]
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30 JOURNAL OF THE ARNOLD ARBORETUM [VoOL. XVII
bulbous body was put out from the other side of the spore (Fig. 4). Normal germination of basidiospores was as high as 90 per cent.
The basidiospores stuck so fast to the surface on which they fell, regardless of whether they alighted on agar or on the cover glass in a thin layer of moisture, that they remained on the slides without requir- ing the use of egg albumen or other adhesive to secure them during the process of staining in iron alum haematoxylin. A study was made of stained germinated basidiospores and it was found that from two to four nuclei were in the germ tubes (Fig. 6).
Abies alba Mill., A. concolor Lindl. & Gord., A. grandis Lindl., A. Nordmanniana (Stev.) Spach, A. Veitchii Lindl., A. firma Sieb. & Zucc. and Picea pungens Engelm. were used for inoculation purposes. ‘Trees, three and four years old, were secured through the Forestry Commis- sion, London, and through reliable nurseries. Some of these were planted in pots and placed in greenhouses at the Chelsea Physic Garden, London, and some in the open at the Biological Station, Imperial Col- lege, Slough, Bucks. Hubert’s tube method as re-described by Faull (3) was used in making inoculations. In some cases the trees were so small that it was impossible to flex the short branches without breaking them, so in connection with these trees the teliospore inoculum was suspended from above so that the basidiospores fell directly upon the upper rather than the lower surface of the leaves. Experiments carried on in both ways gave successful results. Large celluloid tubes stuffed with wet sphagnum at either end to ensure a moist atmosphere were used to entirely enclose the smaller trees. Of all the species used, A. alba, A. concolor and A. grandis only gave positive results. The in- fected needles of A. concolor were slightly distorted but those of A. alba and A. grandis were not (cf. Faull, 3).
Successful infection results were obtained by the writer (7) for the ‘following species:—(1) Milesia Scolopendriti on Abies alba and A. con- color; (2) M. Polypodii on A, alba and A. concolor; (3) M. vogesiaca on A, alba; (4a) M. Kriegeriana (from Dryopteris spinulasa) on A. alba, A. concolor and A. grandis; (4b) M. Kriegeriana (from Dryop- teris Filix mas) on A, alba, A. concolor and A. grandis. Data on the experiments are assembled in Table I. One fir tree, simultaneously and under what were essentially identical conditions, bore four successful infections—each from a different rust species. The variations shown with regard to the periods of time that elapsed between inoculation and the first appearance of spermogonia and peridermia were similar to those recorded in the tables for the respective species. Many more inoculations than those recorded were later set up between June 6 and July 6 but no visible infection was observed.
1936] HUNTER, LIFE HISTORIES OF MILESIA 31
TABLE II. Milesia Scolopendrii from Abies to Various Ferns
Time from inoculation Fern From Date of Date to appearance inoculated Abies inoculation harvested of uredinia Results
Scolopendrium A. alba 7.VI1.34 30.VIII.34 54 days Uredinia vulgare
s ri 7.V1I.34 30.VIII.34 54 days Uredinia
7 _ “ 24.VI1.34 15.1X%.34 53 days Uredinia
24 : “ “ 24.VIL.34 = 30. VIII.34 37 days Uredinia
“ a “ VLVITIL.34 =18.1X.34 38 days Uredinia
a ag “ TLVIIL.34 ) =18.1X.34 38 days Uredinia
Blechnum « 24.VI1.34 = 19.X.34 — No infection Spicant
Dryopteris spinulosa eo "11.VIL.34 = :19.X.34 ~ No infection dilatata
Polystichum a 7.VII.34 = 19.X.34 — No infection angulare
a . « -11.VIIT.34 = 19.X%.34 _- No infection
TABLE III. Milesia Polypodii from Abies to Various Ferns
fob me from oculation Fern Fr Date of Date my appearance inoculated Abies inoculation harvested of uredinia Results
Polypodium A.alba 30.VI1.34 15.1X.34 47 days Uredinia are
ea « « — 30.VIL.34 15.1X.34 47 days eons
«4 “ “11, VIIL.34 18.X.34 68 days. Uredinia ue “ —‘11.VIIL.34 19.X.34 69 days No uredinia
a“ a ~ a
Polystichum 11.VIII.34 19.X.34 — No infection angulare
n a ~ ~
Scolopendrium 30.VII.34 19.X.34 _ No infection
vulgare
JOURNAL OF THE ARNOLD ARBORETUM [voL. XVII
TABLE IV. Milesia Kriegeriana (from Dryopteris Filix mas)
from Abies to Various Ferns
Time from a Fern From Date of Date to appear inoculated Abies inoculation harvested of iredinis Results Dryopteris A. con- 30.VI1.34 = 15.1X.34 47 days Uredinia Filix-mas color = « « 30.VII.34 = 15.1X.34 47 days erie nsect eaten = « 18.VIT1.34 15.1X.34 28 days Uredinia Drvyopteris « « 30.VI1.34 ©9330. VIII.34 31 days Uredinia spinulosa “ «© -18.VII1.34 = 19.%.34 — No infection TABLE V. Milesia Kriegeriana (from Dryopteris) from Abies to Various Ferns inoculation Fern From Date of Date to appearance inoculated Abies inoculation harvested of uredinia Results A. From D. spinulosa and D. spinulosa dilatata Dryopteris A. gran- 30.V1.34 31.VIII.34 64 days Uredinia Filix-mas dis Dryopteris A. gran- 30.V1.34 31.VIII.34 64 days Uredinia, spinulosa dis very slight infection B. From D. spinulosa dilatata Dryopterts A. alba 27.V1.34 31.VIII.34 — No infection spinulosa dilatata . . A. con- 30.VII.34 31.VIII.34 32 days Uredinia color “ “ A. con- 30.VII.34 19.X.34 — No infection col Dryopteris A. con- 30.VII.34 31.VIII.34 32 days Uredinia Filix-mas color Dryopteris A.con- 30.VII.34 15.1X.34 47 days Uredinia spinulosa color
intermedia
1936] HUNTER, LIFE HISTORIES OF MILESIA 33
Ferns for inoculation experiments were procured from Perry’s Hardy Plant Farm and from the field in Devon. Those from Devon were obtained between April 1 and 23. An endeavor was made to select plants that were not already rusted. All the overwintered fronds were cut away fairly close to the underground stem but the young unfolded fronds attached to the stem were left untouched. They were planted in pots at the Biological Station, Slough, in a shaded part of the garden well removed from firs. No rust was observed on any of the ferns as late as August 19, so it is thought that if the ferns from Devon bore natural infection there should have been evidence of it before that date. No naturally occurring ferns bearing rusts were found at the Biological Station.
Aéciospores from culture experiments were used in inoculating vari-
ous ferns with the result that uredospores were obtained for the follow- ing species—(1) Milesia Scolopendrii on Scolopendrium vulgare; (2) M. Polypodii on Polypodium vulgare; (3a) M. Kriegeriana (from Dryopteris spinulosa) on Dryopteris Filix-mas, D. spinulosa, D. spinu- losa dilatata and D. spinulosa intermedia (Muhl.) Underw.; (3b) M. Kriegeriana (from Dryopteris Filix-mas) on Dryopteris Filix-mas and D. spinulosa. Data on the experiments are assembled in Tables II to V. The successful culturing of Milesia Scolopendrii, M. Polypodii, M. vogesiaca and M. Kriegeriana on their alternate hosts now makes pos- sible a completion of the description of these rusts. The uredinial (2, 4-6) and telial (2) phases have already been adequately described. Heretofore the spermogonial and the aecial phases of M. Scolopendrii, M. Polypodii and M. vogesiaca have remained wholly unknown; and the same phases of M. Kriegeriana have been described by Mayor (14) from field collections only, a part of the material of which was used in making successful cultures on the fern host—material not free from the uncertainty of intermixtures. The spermogonia and the aecia of these four species are described below from the writer’s culture material on Abies, that is, from what is known to be authentic, representative material.
(1) Milesia Scolopendrii
O. Spermogonia on needles of current season, epiphyllous and hypo- phyllous, immersed, abundant, inconspicuous, colorless, plane, hemi- spherical to slightly flask-shaped in sectional view, subcuticular, 120- 228 uw broad by 100-188 u high; spermatia hyaline, narrowly elliptical, 1.5-2.0 K 4-5 u.
34 JOURNAL OF THE ARNOLD ARBORETUM [vVOL, XVII
I. Aecia hypophyllous on needles of current season, in two irregular rows, one on each side of midrib, white, cylindrical, 0.4—0.5 mm. in diameter by 0.7-1.5 mm. high; ese tlcire colorless, delicate, rupturing at the apex; peridial cells polygonal, elongated vertically, overlapping, in a single layer, 20-36 28-56 ut, with outer walls smooth, and inner walls finely and densely warted, the warts arranged in elevated, short lines; aeciospores ellipsoid, ovoid or globose, mostly elongated, white, 22-44 28-48 u, very densely and rather coarsely verrucose with warts irregular in outline, tapering to a blunt point and somewhat decidu- ous; walls hyaline, thin, about 1 p thick.
(2) Milesia Polypodii
O. Spermogonia on needles of current season, epiphyllous and hypo- phyllous, immersed, abundant, inconspicuous, colorless, plane, hemi- spherical to slightly flask-shaped in sectional view, subcuticular, 120— 228 uw broad by 105-194 uw high, usually broader than high; spermatia hyaline, narrowly elliptical, 1.5-2.0 & 4-5 u.
I. Aecia hypophyllous on needles of current season, in two irregular rows, one on each side of midrib, on slightly yellowish discolored por- tions of affected needles, white, cylindrical, 0.5-0.7 mm. in diameter by 1.0-1.5 mm. high; peridium colorless, delicate, rupturing at the apex; peridial cells polygonal, elongated vertically, overlapping, in a single layer, 22-42 >< 28-60 u, with outer walls smooth and inner walls with elevated, coarse, short, irregularly oriented ridges; aeciospores ellipsoid, ovoid or globose, mostly elongated, white, 20-36 28-54 u, densely and rather coarsely warted, warts irregular in outline, tapering to a very blunt point and somewhat deciduous; walls hyaline, thin, about 1 u thick.
(3) Milesia vogesiaca
O. Spermogonia on needles of current season, epiphyllous and hypo- phyllous, mostly epiphyllous, very abundant, immersed, inconspicuous, colorless, plane, 154-241 uw broad by 168-214 wu iat: spherical to slightly flask-shaped in sectional view, subcuticular; spermatia hyaline, narrowly elliptical, 1.5-2.0 « 4-5 wu.
I. Aecia hypophyllous on needles of current season, in two irregular rows, one on each side of midrib on slightly yellowish discolored portions of affected needles, white, cylindrical, 0.5-0.7 mm. in diameter by 0.6-1.0 mm. high; peridium colorless, delicate, rupturing at the apex; peridial cells polygonal, elongated vertically, overlapping, in a single layer, 20-32 & 32-48 u, with outer walls smooth and inner walls warted or with elevated, coarse, short, irregularly oriented ridges; aeciospores
1936] HUNTER, LIFE HISTORIES OF MILESIA 35
ellipsoid, ovoid or globose, mostly elongated, white, 24-30 32-46 u, densely warted, warts very irregular in outline and tapering to a very blunt point, somewhat deciduous; walls hyaline, thin, about 1 p thick. (4) Milesia Kriegeriana
O. Spermogonia on needles of current season, epiphyllous and hypo- phyllous, mostly epiphyllous, numerous, irregularly scattered, incon- spicuous, colorless, plane, abundant, immersed, hemispherical in sec- tional view, subcuticular, 98-168 u broad by 94-168 u high; spermatia narrowly elliptical, 1.5—-2.0 « 3.5—5.0 wu.
I. Aecia hypophyllous on needles of current season, in two irregular rows on slightly -yellowish discolored portions of affected needles, sub- circular, ellipsoid or compressed laterally in transverse section, cylindri- cal, 0.3-0.8 mm. in diameter by 0.5-1.3 mm. high; peridium colorless, delicate, rupturing at the apex; peridial cells polygonal, elongated verti- cally, overlapping, in a single layer, 16-52 32-68 u, with outer walls smooth and inner walls with fine elevated ridges, irregularly oriented ; aeciospores ellipsoid, ovoid or globose, mostly elongated, white, 20-36 22-48 u, finely warted, warts tapering to a blunt point, somewhat decidu- ous; walls hyaline, thin, about 1 p thick.
Mayor (14) refers to the late appearance of the aecia from natural infection in the field and believes that the younger needles of Abzes are not susceptible to Milesia Kriegeriana. It would seem from the writer’s experience that the late appearance of the aecia of M. Kriegeriana is because of the long period of development of the fungus from the time of inoculation until the peridermia make their appearance and that young leaves are more susceptible to the rust than older ones.
A comparison of the foregoing descriptions shows that these rusts can- not be separated on morphological differences in their peridermia but, on the other hand, that the spermogonia do afford some help. The spermo- gonia of Milesia Scolopendrii and M. Polypodii are hemispherical to slightly flask-shaped while those of M. vogesiaca are spherical to slightly flask-shaped, and are on the average much larger. The spermogonia of Milesia Kriegeriana are hemispherical in shape and are on the average consistently smaller than those of the other three species. The sub- cuticular areas directly overlying the spermogonia of M. Kriegeriana, M. Polypodii and M. Scolopendrii are relatively greater than that in M. vogesiaca. Indeed in M. vogesiaca the area is so small that in lateral vertical sections the spermogonium appears to be subepidermal. The spermogonia of M. Scolopendrii and M. Polypodii are much alike in form and size, although those of the latter are on the average a little larger. The detailed data are recorded in a paper to be published soon.
36 JOURNAL OF THE ARNOLD ARBORETUM [voL. Xvi
SUMMARY 1, By means of controlled cultures the O I stages of Milesia Scolo- pendriu, M. Polypodti, M. vogesiaca and M. Kriegeriana have been obtained for the first time.
O I hosts II III hosts (1) Milesia Scolopendrii Abies alba, Scolopendrium vulgare A. concolor (2) Milesia Polypodi Abies alba, Polypodium vulgare A. concolor (3) Milesia vogesiaca Abies alba Polystichum angulare (4a) Milesia Kriegeriana Abies alba, Dryopteris spinulosa, (from Dryopteris A. concolor, D. spinulosa dilatata, Spinulosa) A. grandis D. spinulosa intermedia,
D. Filix-mas (4b) Milesia Kriegeriana Abies concolor Drvyopteris spinulosa, (from Dryopteris D. Filix-mas Filix-mas )
2. The spermogonia and the aecia of M. Scolopendrii, M. Polypodii and M. vogesiaca are described for the first time. The spermogonia and the aecia of M. Kriegeriana are described for the first time from culture material,
3. The spermogonia of M. Kriegeriana and M. vogesiaca are dis- tinguishable by their form and size from one another and from those of M. Scolopendrit and M. Polypodii. The spermogonia of M. Scolopen- drii and M. Polypodii are similar in form and size.
ACKNOWLEDGMENTS
Studies recorded in this paper form part of an investigation carried out by the writer under the direction of Professor J. H. Faull of the Arnold Arboretum, Harvard University. This particular phase of the work was made possible by a fellowship granted by the Canadian Fed- eration of University Women and was mostly done in England under the sponsorship of Professor William Brown of the Imperial College of Science and Technology, London University. Sincere thanks are ten- dered to Professors Brown and Faull. The author also gratefully acknowledges various services rendered by Mrs. L. Allen, Mr. D. Blockey, Dr. G. D. Darker, Dr. E. Milton, Mr. H. B. S. Montgomery, Dr. Muriel P. Hall, Dr. P.O’Connor, Mr. A. H. G. Alston, Mr. W. Hales, Mr. W. Williams, Mr. J. Ramsbottom, Mr. A. D. Cotton, Miss E. M. Wakefield, Dr. E. J. Butler, Mr. E. W. Swanton, Mr. J. Ogilvie, the Torquay Museum of Natural History and the Forestry Commission of Great Britain.
Jour. ARNoLD Ars. VoL. XVII PLATE 176
Mites1a Potypopit, M. ScoLopENpDRII, M. KRIEGERIANA AND M. VOGESIACA
FULL-TONE— MERIDEN
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1936] HUNTER, LIFE HISTORIES OF MILESIA 37
Ne
oO. © NW np w bo
LITERATURE CITED AsHuwortH, D. Puccinia Malvacearum in monosporidial culture. (Trans. Br. Myc. Soc. 16: 177-202. 1931. FauL., J. H. Taxonomy and geographical distribution of the genus Milesia (Contrib. Arnold Arb. 2: 1-138. The biology of Milesian rusts. (Jour. Arnold Arb. 15: 50-86. 1934. Grove, W. B. The British rust fungi. (Cambridge, 1913.) — —— The British species of Milesina. (Jour. Bot. 59: 109. 1921.) ——— Mycological notes VI. (Jour. Bot. 59: 311-313. 1921.) Hunter, L. M. preliminary note on life history studies of Euro- pean species of Milesia. (Jour. Arnold Arb. 16: 143. 1935. ———— The genus ie in Great Britain and Ireland. (Trans.
. Kamel, S. Note Sve na vogesiaca Sydow on Polystichum
ana Miyabe et Kudo, A. firma Sieb. et Zucc. and A. sachalinensis ast. yas, Sapporo Nat. His. Soc. 11: 141-147. 1930. ———. A new species of Milesina Sig yee on ae vulgare L Cras Sapporo Nat. His. Soc. 12: 27-33. On new species of heteroecious yee rusts. teoas Sapporo Nat. a Soc. 12: 161-174. 1932.
. KieBpaAun, H. Kulturversuche mit Rostpilzen. (Zeitsch. Pflanzenkr.
26: 357.277, 1916. Maanus. P. Weitere ee uber die auf Farnkrautern auftre- tenden Uredineen. (Ber. Deutsch. Bot. Ges. 19: 578-584. 1901.)
. Mayor, Euc. Notes be Le VIII. (Bull. Soc. Neuchat. Sci.
Nat. 58: 23-26. 1933.) HITE UCHANAN. Note on the zoology and botany of Glen Tilt. (Scottish Naturalist, 4: 160-163. 1877-8. )
EXPLANATION OF PLATE 176
ig. 1. Basidiospores of Milesia Polypodii B. White. Note variation in
form and germination.
x 495. ioc: Basidiospores of Milesia apts (Fuckel) Arth. Note
variation in form and germinati
3. Basidiospores of Milesia Kriegeriana (Magn. ) Arth. from Dry- opteris Filix- ions ia Schott. Note variation in form and
ermination.
4. Germinating Sed cere of Milesia Polypodii. Note the vari- ous stages in germination and the formation of a secondary spore. 5.
. 5. Germinating basidiospores of Milesia Polypodii. X 200.
. 6. Germinating basidiospores of Milesia Polypodii. Note nuclei in germ tubes. he
. 7. Young tree of Abies alba Mill. infected with four species of
Milesia, namely,—(1) M. Polypodii on upper left branch (inocu- lation experiment No. 7); (2) M. vogesiaca (Syd.) Faull on up- per right branch (inoculation experiment No. 8) ; (3) M. Krieg- ertana on lower left branch (inoculation experiment No. 9); (4) M. ehatia’ on lower right branch (inoculation experiment No. 10). xX %.
LABORATORY OF PLANT PATHOLOGY,
ARNOLD ARBORETUM, HARVARD Lwiv ERSITY.
38 JOURNAL OF THE ARNOLD ARBORETUM [vVoOL. Xvi
A CONVENIENT SAND-CULTURE APPARATUS Rosert W. Warp With plate 177 and one text figure
One of the essential requirements in nutritional studies on higher plants is a culture apparatus in which replications of an extended series may be studied simultaneously. It is also always desirable, if not actu- ally necessary, that the time expended in the routine operations of water- ing and feeding the plants be reduced to a minimum. The most im- portant consideration, however, from the physiological point of view is that the component parts of the apparatus with which the plants, medium, or nutrient come in contact be constructed of materials which are chemically resistant, as well as non-porous, and capable of affording leak-proof connections.
An apparatus which seems to satisfy these requirements better than any heretofore described has been designed and put into use by the writer in a study on physiological disorders of apple trees.
The apparatus consists primarily of an inverted, bottomless, one- gallon jug connected to an inverted, standard, one-half-gallon jug by means of rubber- and glass-tubing (Fig. 14). The bottomless jug B, in which the plants are grown in white silica sand, is of standard dimensions but lacks both bottom and handle and has the neck grooved so that the rubber stopper may be wired in position. It is glazed externally accord- ing to standard requirements with a white glaze and internally with a special acid-resistant glaze. The sand found satisfactory for apple cul- ture is a mixture of 5 parts of Columbia No. 4, a coarse, angular, silica sand, and 6 parts of Ottawa No. 20, a silica sand having almost spherical grains. The lower part of the jug, as shown in the illustration, is pro- vided with the Columbia No. 4 sand to ensure good drainage. The sand is prevented from entering the glass tubing by the use of a small circle of Monel metal screening, No. 32 mesh, laid against the rubber stopper. The nutrient solution is placed in the one-half-gallon jug A, from which it is flooded into the sand by elevating this jug to the proper height and into which it is later drained by lowering again. The connecting tubings H are of heavy-walled gum rubber and standard Pyrex glass. As much of the latter is used as is conveniently possible (Plate 177). The tubing J, which facilitates movement of air in the nutrient jug, may also serve as a suction tube to which a very weak suction pump may be attached
1936] WARD, SAND-CULTURE APPARATUS 39
if more aération of the sand culture medium is desired than can be obtained by normal drainage of the nutrient solution. As many of these pieces of apparatus as are necessary or can be used conveniently are assembled on a wooden stand, as shown in Fig. 1, in which the culture jugs B...B are placed in twin beds C, C and the nutrient jugs A...A in a movable rack E. The writer was thus enabled to set up twenty-four cultures in triplicate, seventy-two cultures in all, on one greenhouse bench.
XX
C
x» Go PPT Figure 1. SANp CULTURE ApparRATus. 1, 2, 3. Working drawings of stand and assembled jugs. 4. Vertical section of jugs in flooding position.
The wooden stand which holds the jugs has been designed to fit on an ordinary greenhouse bench (Fig. 11:3; Plate 177). It is composed of twin beds C, C resting on horses D...D, placed about four feet apart, and a movable median rack E, which, in the raised or flooding position, rests on removable horses of a predetermined height, placed about eight feet apart (Plate 177) and which later, in the lowered or draining posi- tion, comes to rest on two stout spiral-spring blocks (Fig. 11?-F), The beds are fastened firmly to their respective horses and the ensemble is made immovable by use of the upright structures G...G (Fig. 13; Plate 177), which also serve as guides for the rack E. The lumber used
40 JOURNAL OF THE ARNOLD ARBORETUM (VOL. Xvi
in the construction of the stand is ‘‘western fir,” 2 inches by 4 inches, and white pine, 1 inch by 4 inches. All exposed parts are painted with a good white waterproof paint.
Although details of construction of the stand are obvious in the accompanying working drawings (Fig. 1) there are several important features to be noted. First, the scantlings supporting the inverted bottomless jugs B...B in the beds C, C are mitered instead of being inserted at right angles or parallel to the sides of the bed. This makes it possible to adjust the beds to the proper width so that the ensemble of beds and rack may be accommodated on the greenhouse bench. e mitered angle of the scantlings depends naturally on the width of bed desired. The scantlings are also spaced accurately, both in the beds and rack, so that the inverted jugs may rest snugly on their shoulders between adjacent pieces. Secondly, an intermediate portion of the upright part of the beds and the entire bottoms are covered with gal- vanized wire netting of a coarse mesh. This permits the bottomless jugs in the beds to be packed in wet sphagnum moss, an arrangement which tends to keep the cultures at a uniform temperature due to the regulatory properties of evaporating water. Thirdly, the horses D...D supporting the beds must be of sufficient height to allow proper and adequate drainage of the cultures. Finally, it is imperative that at least two sets of heavy spiral springs F...F, of the shock-absorber type be placed in such a position under the rack E as to give the latter maxi- mum support when fully loaded. This precaution is necessary to pre- vent shock and probable damage to the glass tubing when the rack with its heavy load of nutrient jugs and attached glass- and rubber-tubings are lowered to the draining position. —
The operations of raising and lowering the rack and nutrient jugs are accomplished by means of two quadruple, self-locking block and tackle sets. These are attached to the rack at the proper supporting points and to a solid superstructure directly above the rack, the slings in each case being of Manila rope or some equally reliable uecanl, The single operation of raising or lowering the rack can be accomplished by one operator and occupies about three minutes.
The height to which the rack should be raised for proper flooding of the cultures is determined in the following manner. Two liters of water are placed in the nutrient jug A, which is then set in the rack and con- nected to its respective culture jug B by means of the rubber- and glass- tubing H described above. The rack is then raised until the water level reaches exactly to the surface of the sand (Fig. 14). This height is carefully recorded and light-weight wooden horses are constructed to support the rack when raised to this position (Plate 177).
Jour. ArNoLp Ars. VoL. XVII PLATE 177
SAND CULTURE APPARATUS WITH APPLE TREES IN CULTURE. (Photos by A. B. Hatch)
a oe oe a oe
Bn
re
a 7 a —— 7 a 7
— — oe
a _ _ -
7
1936] WARD, SAND-CULTURE APPARATUS 41
At this time tests are made with several of the units to determine the average volume of water retained in the sand after drainage. Later, when the culture solutions are made up, this volume is taken into consideration.
The routine daily care of the cultures resolves itself briefly into the following operations. The rack is raised to the flooding position and, after a lapse of a few minutes to allow adjustment of the liquid level, distilled water is added directiy to the sand to compensate for loss by evaporation and transpiration, until the original level at the surface of the sand is reached. Thus the original volume of the culture solution is retained throughout the course of the experiment. When the cultures have been flooded for a sufficient length of time the rack is lowered, allowing the sand to be drained and aérated naturally at the same time.
For purposes of convenience, the writer placed a carboy of distilled water on a high stand at the rear of the beds. Leading from this, along either of the outer sides of the beds, were two lines of 12 mm. glass-tubing provided with several outlets (Plate 177). A short length of rubber- tubing was attached successively to these outlets and by this means the complement of distilled water was added to the cultures directly from the carboy.
The frequency with which the culture solutions should be renewed depends upon the technique of the operator and the plants in culture. Since all of the nutrients can be accounted for, either in the plants or in the solutions, it is possible to make periodic analyses of the latter and adjust the time of renewal according to the interpretation of the results obtained.
This apparatus was designed and constructed in the laboratory of Professor J. H. Faull of the Arnold Arboretum, Harvard University, in connection with investigations carried on under his direction. This research was made possible through scholarships from the Harvard Forest and the Arnold Arboretum. The writer wishes to express his sincere thanks to Professor Faull for the facilities afforded and his con- tinued interest. Much credit is due Professor P. R. Gast of the Harvard Forest, who contributed the basic principle of the apparatus devised from his own researches and assisted in many ways with time and equipment. Grateful appreciation is expressed to my former labora- tory associates, Drs. A. B. Hatch and J. D. MacLachlan for their many constructive criticisms.
LABORATORY OF PLANT PATHOLOGY,
RNOLD ARBORETUM, HARVARD UNIVERSITY AND Division OF HorTICULTURE,
ENTRAL EXPERIMENTAL FARM, OTTAWA, CANADA.
42 JOURNAL OF THE ARNOLD ARBORETUM [voL. XvII
CHROMOSOME COMPLEX AT PREMEIOTIC ANAPHASE AND MEIOTIC METAPHASE
Haic DERMEN With two text figures
A NUMBER of workers have reported in the case of plants that the individual chromosomes at anaphase are at least double. In this labora- tory we have seen this duality in the root-tips (Dermen 1933), during all stages of microsporogenesis in Rhoeo, and in the telophase stage of embryo sac tetrad nuclei in Lilium regale. In the present paper we are presenting conclusive evidence indicating that this duality also exists in
Fic. 1.
Ficure 1. Preme1otic ANAPHASE FROM RHOEO DISCOLOR.—FIGURE 2. Metotic First METAPHASE FROM A Hysrip TRADESCANTIA (T. REFLEXA x T. PALUDOSA).
the last premeiotic anaphase chromosomes which are destined to become meiotic leptotene threads. In addition to this the meiotic metaphase chromosomes were studied and were found to be structurally eight-parted.
The anthers of a young bud which is the third in series from that in the reduction stage in Rhoeo discolor were used for the study of pre- meiotic anaphase (Fig. 1). The cells of these anthers are for the most part in early prophase of meiosis but a few may be found in somatic metaphase or anaphase stage. The material showing the structure of meiotic first metaphase chromosomes (Fig. 2) was from a_ hybrid Tradescantia, T. reflexa * T. paludosa.
1936] DERMEN, CHROMOSOME COMPLEX 43
MATERIAL AND TECHNIQUE
The anthers in either case were smeared and pretreated for 5 to 10 seconds in 30% alcohol, 40 cc. in a staining jar, to which was added approximately 4 to 6 drops of ammonia water (26%), and the smear was then flooded with aceto-carmine 2 or 3 times for about 30 seconds. The slide was drained, any residual anther material that remained on the slide was removed, and a drop of aceto-carmine was added and a cover glass placed on the smear. The material was first located and studied, and if the preparation was successful the slide was gently heated over an alcohol lamp until small bubbles began to form. The cover glass was then pressed using a piece of filter paper for the purpose. To make these temtporary preparations permanent either McClintock’s acetic-alcohol or Metz’s acetic-alcohol-xylol method may be used.
For a more permanent smear using the crystal violet-iodine stain, the following method is prescribed: Smear and pretreat as described above and place the slide, smear side down, in a small shallow dish containing a few drops of % to 34 strength Flemming’s modified solution for %4 to 1 minute. Rinse in water in a staining jar for one to two minutes, then stain in crystal violet for 15 seconds or longer depending upon the source of the material. Some plant smears may require over one minute of staining. If material so stained is thoroughly washed with xylol future destaining is prevented and the stain is as permanent as haemotoxylin preparations.
DESCRIPTION
A photomicrograph of an anaphase genom at last premeiotic stage from Rhoeo discolor is shown in figure 1. It is one of the two anaphase groups seen from a slightly oblique polar view. It shows on the lower right the short arm of a chromosome pointing down. From this posi- tion the structure of this arm could be clearly made out indicating unmistakably that it is split. The halves seem to be independently coiled and are not intimately associated. Where these chromosomes are twisted they appear as if constricted.
A meiotic first metaphase from a hybrid Tradescantia (T. reflexa T. paludosa) is shown in figure 2, also a photomicrograph. A bivalent chromosome is seen in a horizontal plane. This pair of chromosomes at the connection point shows two kinds of split, one major and one minor. This type of opening was not confined to the rod-bivalents alone. In other cells in the ring-bivalent chromosomes at both ends of their connections, these double, large and small, diamond shaped open- ings could be observed, but for the purpose of illustration the rod- bivalent type was found best, for it is easily flattened out by pressure on the cover glass to show this feature.
44 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XVII
DISCUSSION
The duality in the premeiotic chromosomes has been shown by Kauf- mann (1926) in plant material. In animals, McClung (1928) and Robertson (1931) have demonstrated this duality of chromosomes at a corresponding stage. Koshy (1934) has illustrated the duality of pre- meiotic anaphase chromosomes and leptotene threads, though it appears from his drawings that he may have confused leptotene with pachytene stage.
Nebel (1932 and 1933), Stebbins (1935) and Goodspeed, Uber and Avery (1935) are in agreement that telophase chromosomes are four- parted. It is indicated that the leptotene threads, although optically single, are actually four-parted. Goodspeed et al. believe further, by estimation, that the meiotic metaphase chromosomes are sixteen-parted, thus homologizing with Drosophila salivary chromosomes which are suggested to be sixteen-parted.
Huskins (1932) and Huskins and Smith (1934 and 1935) have ad- mitted the duality of anaphase chromosomes elsewhere but deny it in the premeiotic anaphase. They also have reported eight-parted meiotic metaphase chromosomes. Darlington (1935) on the other hand has denied the duality of anaphase chromosomes in all regions. He believes that anaphase chromosomes are sing!e in both somatic and premeiotic phases because of the fact that leptotene threads appear to be single.
Our own evidence (Figs. 1 and 2) shows conclusively that premeiotic anaphase chromosomes are at least two-parted and eight-parted at mei- otic metaphase. The indication is that mitotic and meiotic phases are essentially of the same fundamental nature, with this difference, that instead of each chromosome remaining as a unit and perpetually dividing as in mitosis, in meiosis similar chromosomes (homologues) pair to- gether (barring the crossing-over phenomenon from the discussion) and during the first metaphase move to opposite poles. In mitosis homo- logues do not form intimate association while in meiosis their associa- tion is quite intimate and complex. The mitotic chromosomes are four- parted at metaphase, in meiosis they are eight-parted in the bivalents, and correspondingly there are four in each half of the bivalents.
I tried to analyze the same situation of duality in Lilium ovules where as in any other plant there is usually only one cell that becomes meiotic. The preparations I made (sectioned material) were of a somewhat advanced stage.- However, there is one obvious thing that should be considered here, that this meiotic cell is a product of a somatic cell division. Here one cell only becomes meiotic while its sister cell con- tinues the-somatic cycle. It is hard to imagine if the anaphase chromo-
1936] DERMEN, CHROMOSOME COMPLEX 45
somes in the somatic cell are split, as Huskins admits (1932), how the anaphase chromosomes of one such group become normally split while those in the opposite group remain unsplit to justify Huskins’ assump- tion that leptotene threads are single by origin. In both sister cells chromosomes are either split or unsplit since these halves are supposedly mirror images of each other. If the chromosomes were single in this stage there should be two cells alike and both meiotic. However, there is normally only one such cell. The obvious conclusion is that if chromosomes are split in the somatic cells at anaphase elsewhere, they must be split here too and that undoubtedly late premeiotic anaphase chromosomes of the egg-mother-cell are of a dual nature.
SUMMARY
Conclusive evidence is presented showing that last premeiotic ana- phase chromosomes in microsporogenesis are split and that metaphase chromosomes in pollen-mother-cells are optically eight-parted. It is argued that the same is true in the ovule.
A method which enabled us to bring out the coiled structure in meiotic chromosomes is described in some detail. This method involves pre- treatment of smear preparations with ammonia in alcohol.
LITERATURE CITED
DarLINGTON, C. D. (1935). The old Sige reer Ae and the new analysis
of mae aaa ee r. (Ann. 49: 579-586. )
DeRMEN, H. (19 Origin and behavi or of the nucleolus in plants. (Jour. Arnold res 14: 282-323. )
GoopspEED, T. H., F. M. User, & P. Avery (1935). Application of the Altmann freezing- drying technique to plant cytology. III. Chromosome structure in Lilium longiflorum. (Univ. Calif. Publ. Bot. 18: 33-44.)
Husxins, C. L. (1932). Observations ‘bearing o on the mechanism of meiosis and crossing over. (Proc. Sixth Int. Congress Genetics, 2: 95-96.
—— & S. G. SmitH (1934). Chromosome division and pairing in Fritillaria Meleagris: The mechanism of meiosis. (Jour. Genetics, 28: 6.)
————— ————. (1935). Meiotic chromosome structure in Trillium erectum L. (Ann. Bot. 49: 119-150.) KAuFMANN, B. P. (1926). Chromosome structure and its relation to the chromosome cycle. I. Somatic mitosis in Tradescantia pilosa. (Amer. 0
Kosuy, T. K. (1934). Chromosome studies in Allium. II. The meiotic
McCiune, C. E. (1928). ’ Differential chromosomes of Mecostethus gracilis. (Zeit. Zellf. Microsk. Anat. 7: 756-778).
46 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XVII
Neset, B. R. (1932). Chromosome structure in Tradescantiae. I. Method and morphology. (Zeit. Zellf. Microsk. Anat. 16: 251-284. —— (1933). Chromosome structure in Tradescantiae. IV. The
Rogertson, W. R. B. (1931). Chromosome studies. II. Synapsis in the Tettigidae with special reference to the presynapsis split. (Jour. ree i -_ Physiol. 51: 119-146.
STEBBINS, G. (193 ). Chromosome structure and the mechanism of eo hi in ae (Am. Nat. 69: 81.)
CyTOLoGIcAL LABORATORY, ARNOLD ARBORETUM, HARVARD UNIVERSITY.
1936] REHDER, BURRETIODENDRON 47
BURRETIODENDRON, A NEW GENUS OF TILIACEAE ALFRED REHDER With plate 178
Burretiodendron, gen. nov.
Flores dioeci, 5-meri, in cymis axillaribus paucifloris bracteatis pedunculatis petiolo multo brevioribus dense stellato-pilosis interdum in apice ramulorum brevium in paniculam brevem congestis; flores masculi breviter pedicellati, basi bracteolis 2 late ovalibus vel 3 ellip- ticis obtusiusculis concavis quam sepala fere dimidio brevioribus extus dense stellato-pilosis glandulis interspersis, intus glabris ante anthesin alabastrum arcte includentibus instructi; sepala 5, libera, anguste elliptica, acuta, extus dense stellato-pilosis, intus glabra basi area oblonga elevata dense glanduligera trientem sepali aequante instructa; petala 5, libera, alba aestivatione contorta, sepalis paulo longiora, unguiculata, lamina late obovata, apice fere truncata et plus minusve erosula et cilio- lata basi in unguem tenuem fere dimidiam laminam aequantem attenu- ata, margine ciliolata excepta glabra; stamina circiter 30, petalis triente breviora, filamentis filiformibus infra medium in phalanges 5 et basi in tubum connatis, antheris lineari-oblongis bilocularibus basifixis, loculis rima longitudinali dehiscentibus; rudimentum ovarii parvum tubo staminali inclusum ovoideum rostratum. Flores feminei non _ visi. Capsula oblonga, stipitata, 5-alata, septicide 5-cocca, alis capsulae papyraceis bilamellatim partitis, coccis verticaliter bialatis indehiscenti- bus monospermis; semen oblongo- obovatum, basin versus attenuatum, apice obtusum et chalaza ovali notata, parce albuminosum; pee ae rectus, cotyledonibus foliaceis latis tenuibus plicatis sacreule recta.— Arbor ramulis hornotinis dense pilis stellatis vestitis, annotinis glabris. Folia alterna, decidua, ovata vel late ovata, acuminata, basi cordata, palminervia nervis basalibus 5—7 et insuper utrinsecus nervis 4—5 angulo angusto a costa media divergentibus et ascendentibus, nervis lateralibus a costa media et a nervis basalibus divergentibus ad marginem ceterum integram in mucronem exeuntibus, utrinque pilis stellatis satis dense, in costa et venis densius vestita, costa nervisque supra fere planis subtus distincte elevatis et trabeculis reteque venularum prominulis conjunctis; petioli satis graciles, dense stellato-pilosi; stipulae caducae, vestigia tantum adsunt.
Species unica Chinae occidentalis incola.
48 JOURNAL OF THE ARNOLD ARBORETUM [voL. Xvi
Burretiodendron Esquirolii (Lévl.), comb. nov.
Pentace Esquirolii Léveillé in Fedde, Rep. Spec. Nov. 10: 147 (1911) ; Fl. Kouy-Tchéou, 419 (1915).— Burret in Notizbl. Bot. Gart. Mus. Berlin, 9: 620 (1926).
Eriolaena Esquirolii Léveillé, Fl. Kouy-Tchéou, 405 (1915).
Character generis.
Petiolus 2.5—5 cm. vel interdum ad 11 cm. longus; folii lamina 6-12 cm. longa et 4-8 cm. lata vel interdum ad 28 20 cm. magna. Inflo- rescentia pedunculo brevi incluso circiter 3 cm. longo; bracteae 7 mm. longae; flores circiter 2 cm. diam.; sepala circiter 10 mm. longa et 4 mm. lata; petala ungue 3 mm. longo incluso circiter 11-12 mm. longa, lamina circiter 7-8 mm. lata; stamina 7 mm. longa antheris 2 mm. longis; ovarii rudimentum 1 mm. longum. Capsulae 3—3.5 cm. longa et circiter 1.5 cm. lata, alis circiter 6 mm. latis; semina 8 mm. longa et 3-4 mm. lata.
DISTRIBUTION: Southeastern Yunnan and Kweichou.
Cuina. Kweichou:. ouest de Lo-fou (Kouai-kou), J, Cava- leri, no. 2648, Nov. 1905, “arbre moyen, moucilagineux” (holotype of Pentace Esquirolii; photo. in A. A.); same locality, J. Esquirol, no. 817, and Yang-ly, J. Esquirol, no. 2717, both cited in Fl. Kouy-Tchéou under P. Esquirolii; Yang-ly, J. Esquirol, no. 2717, Aug. 1911, “arbre 8-10 métres, fleur blanche” (holotype of Eriolaena Esquirolii; photo. in A. A.); Lohu, 170 m. in lightly shaded ravine, Y. Tsiang, no. 7290, Oct. 3, 1930, “tree 10 m. high, diam. of trunk (breast high) 20 cm., leaves dull green above, light green below, fruit yellowish green” (New York Bot. Gard.) Yunnan: Red River, A. Henry, no. 9572 (New York Bot. Gard.); Red River, near Manhao, A. Henry, no. 9573, “tree 25 ft., flowers all fallen off, young green fruit” (New York Bot. Gard.).
This new genus does not seem to be closely related to any of the genera of the Tiliaceae or Sterculiaceae, but is best perhaps placed in the Tiliaceae near Luehea; from the Sterculiaceae it differs in the dry winged fruit dividing septicidally into one-seeded carpels broadly winged all around, and in the androecium which in the latter family has the episepalous stamens sterile or lacking, while in Burretiodendron they are all fertile. With Luehea it agrees in the lack of an androgynophore and in pentadelphous stamens, but differs markedly in the unisexual flowers, in the slender-clawed petals without glandular spot at the base and in the fruit separating into 5 one-seeded carpels. In the presence of a nectary at the base of the sepals Burretiodendron differs from all Tiliaceae. In the fruit separating into one-seeded winged carpels, it resembles Colona, but is easily distinguished by the unisexual flowers in axillary short cymes, the lack of an androgynophore, the clawed petals
Jour. ARNoLp Ars. VoL, XVII PLATE 178
2B oP be roe Sogee nok (eer? J J Chol aor. FP care G 6 OU.
Det Alfred Rehder ARXHLD Antounrow SY, war
BURRETIODENDRON Esgurrotit (Lévl.) Rehder.
7
-
- OO Be BS
1936] REHDER, BURRETIODENDRON 49
without nectary and the pentadelphous stamens. Eriolaena in which Lé veillé placed the flowering specimen, differs among other characters chiefly in its ligneous loculicidally dehiscent capsule with many winged seeds. Pentace, to which he referred the fruiting specimen, is easily distinguished by the rather small bisexual flowers in large terminal panicles and the much smaller indehiscent one-seeded fruit.
The specimens collected by Henry near Manhao, about 25 kilometers north of the border of Tonkin, agree exactly with Cavalerie’s and Esquirol’s specimens, except that the former have the leaves generally more deeply cordate like those of Tsiang’s specimen. The genus may possibly extend into northern Tonkin and northwestern Kwangsi.
I take pleasure in naming this new and very distinct genus in honor of Dr. M. Burret who has made important contributions to the knowl- edge of the Tiliaceae.
EXPLANATION OF PLATE 178
1. Fruiting specimen, Y. Tsiang, no. 7290. x 2/5. . 2. Single carpel of fruit from the ventral and dorsal side. x 2/5. Fig. 3. Axillary cyme with one open flower, from Henry, no. 9572. Natural size. 4. Terminal panicle of flower buds, from Henry, no. 9572. Natural size.
HERBARIUM, ARNOLD ARBORETUM, HARVARD UNIVERSITY.
50 JOURNAL OF THE ARNOLD ARBORETUM [voL. Xvu
TWO NEW SPECIES OF GYMNOSPORANGIUM FROM ASIA Ivan H. Crowe. With two text figures
Gymnosporangium magnum Crowell, sp. nov. (Fig. 1).
Spermogonia foliicolae, subepidermalia. Aecidia hypophylla, cupuli- formia, tumoribus subglobosis insidentia; cellulae peridii rhomboideae, hyalinae, pariete interiore marginibusque subspinulosae, 22.5-30.6 50.0-64.4 ut, plus minusve 24.3 59.2 4; aecidiosporae globosae, pallide brunneae, verruculosae, 24.2-29.0 > 30-32 u, plus minusve 26.1 29.8 w.
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pur
Gees Kio — Fic. 2.
IGURE 1. GYMNOSPORANGIUM MAGNUM a —Ficure 2. GyYMm- NOSPORANGIUM LEVE Crowell.—AECIOSPORE ND PERIDIAL CELLS.
Hab. in foliis Crataegi sp. in China centrali. Ex Herb. G. Lagerheim, Botaniska Andelning Riksmuseet, Stockholm, Sweden (Smith, no. 917)
The specific name is suggested by the unusual size of the aeciospores, a feature in which they are surpassed by G. juniperinum (L.) Mart. only.
Gymnosporangium leve Crowell, sp. nov. (Fig. 2).
Spermogonia foliicolae, sibepideuans, Aecidia hypophylla, roes- telioidea, tumoribus tumidis insidentia, 1-2 mm. alta, 0.25—0.5 mm. lata; cellulae peridii elongatae, hyalinae, pariete interiore spinulosae, mar-
1936] CROWELL, NEW SPECIES OF GYMNOSPORANGIUM 51
ginibus et pariete exteriore gradatim plus breviter spinulosae, 12.8-18.2 X 80-120 u, plus minusve 15.2 102 uw; aecidiosporae globosae, pallide brunneae, leves, 19.2-22.4 20.8-24.0 u, plus minusve 20.8 xX 22.6 wu.
Hab. in foliis Mali Sieboldii (Reg.) Rehd. in China. Ex Herb. G. Lagerheim, Botaniska Andelning Riksmuseet, Stockholm, Sweden (Smith, no, 4221).
The material from which this species is described was collected by Smith at Karlong, Szechuan, China, altitude 3200 m., Aug. 27, 1922. The specific name is suggested by the smooth surface of the aeciospore wall, a character of rare occurrence in this genus of rusts.
Comparatively few species of Gymnosporangium have so far been described from Asia. One feature common to them, generally speaking, is the occurrence of spinulose embossments on the peridial cells of the aecia. Among the American species this feature is found in G. exiguum Kern only.
The specimens which now become the types of Gymnosporangium magnum and G. leve were loaned from the Herbarium of the Natur- historika Riksmuseet, Stockholm, Sweden, through the courtesy of Professor Dr. Gunnar Samuelsson.
LABORATORY OF PLANT PATHOLOGY, ARNOLD ARBORETUM, HARVARD UNIVERSITY.
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7
JOURNAL
OF THE
ARNOLD ARBORETUM
VoLtuME XVII APRIL, 1936 NUMBER 2
NOTES ON THE LIGNEOUS PLANTS DESCRIBED BY H. LEVEILLE FROM EASTERN ASIA’
ALFRED REHDER
IN THE COURSE of my revision of the ligneous plants described by Léveillé from Eastern Asia, it became apparent that after once going over the ground in systematic order, I would have to publish a supplement to include those plants which for various reasons, had been omitted in the first account.
When I started this work in 1929, after a short sojourn in Edinburgh in the summer of the preceding year, I based it chiefly on notes and photographs I had taken and on duplicates of types generously given to me whenever feasible, by Sir William Wright Smith, Regius Keeper of the Royal Botanic Garden. Later in the course of my work, Sir William kindly offered to send me on loan all the types I had not examined while in Edinburgh; this enabled me to treat the later families more fully. Another reason for the omission of certain species in their proper places was the fact that in many instances, species referred by Léveillé to cer- tain genera, did not belong there but to families previously treated, e.g. Clerodendron Esquirolii which turned out to be a Tacca. There remains also to be cited important additional literature dealing with Léveillé’s plants and not yet mentioned in my previous publications.
After having finished my first attempt of an enumeration of Léveillé’s ligneous plants in systematic order from the Taxaceae to the Compositae, I journeyed last year once more to Edinburgh to go over the Léveillé herbarium, and again Sir William facilitated my work in every possible way, so that I was able to accomplish much in the limited time at my disposal.
It is to be expected that with the following supplement, the critical enumeration of the ligneous species described by Léveillé from eastern Asia will be fairly complete with the exception, perhaps, of a few species which could not be located in Léveillé’s herbarium.
1Continued from Vol. 16: 311-340; for preceding parts see Vols. 10: ag 132, 164- 196; 12: 275-281; 13: 299-332; 14: 223-252; 15: 1-27, 89-117, 267-3
54 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XVII
TAXACEAEF!
Podocarpus neriifolius D. Donn. — Rehder in Jour. Arnold Arb. 10: 108 (1929).
Alyxia Schlechteri Léveillé, Fl. Kouy-Tchéou, 30 (1914), quoad Cavalerie, no. 3463; not Léveillé (1911).
Cutna. Kweichou: Lo-fou, riviére de louest, J. Cavalerie, no. 3463, March 1909, “petit arbre, 4—5 m.” (sub Alyxia Schlechteri in herb. Lévl.; duplicate in A. A.)
Leaves of Cavalerie no. 3463 were sent in 1916 to the Arnold Arbore- tum by Léveillé in answer to our request for a fragment of Alyxia Schlechteri. For typical A. Schlechteri see Jour. Arnold Arb, 15: 316 (1934).
Of P. macrophylla var. Argyi Léveillé in Mem. R. Acad. Ci. Barce- lona, ser. 3, 12: 547 (1916), nom. nud., I have seen no description and no specimen; it is probably not different from P. macrophylla.
Taxus chinensis (Pilger) Rehder in Jour. Arnold Arb. 1: 51 (1919). Tsuga Mairei Lemée & Léveillé in Monde PI. ser. 2, Sit 20 (1914). — Léveillé, Cat. Pl. Yun-Nan, 58 (1916). — Synon CHINA. Yunnan: commun sur les collines be ae 700-800 m., E. E. Maire, May, 1912 (holotype of Tsuga Mairez; cola: and inane in A. A.). The type specimen is sterile. To 7. chinensis probably also belongs T. cuspidata in Léveillé, Fl]. Kouy-Tchéou, 112 (1914).
PINACEAE?
Pinus Massoniana Lambert. — Rehder in Jour. Arnold Arb. 10: 109 (1929).
Pinus Argyi Lemée & Léveillé in Fedde, Rep. Spec. Nov. 8:60 (1910). — Léveillé in Mem. R. Acad. Ci. Barcelona, ser. 3, 12: 547 (Cat. Pl. Kiang-Sou, 7) (1916
Pinus Argyi var. longevaginans Léveillé j in aes lc. and Mem. L.c.
Pinus Cavaleriei Lemée et Léveillé in Fedde,
Pinus tabulaeformis Carr. sensu Rehder in . “Arnold Arb. 10: 109 (1929) quoad syn. P. Argyi; non Hort. ex Carriére.
CHInA. Kiangsu: Che-sang and Zuosé, Ch. d’Argy (syntypes of P. Argyi,; photos. in A. A.); locality illegible, Ch. d’Argy (holotype of P. Argyvi var. longevaginans ; photo.in A. A.). Kweichou: Pin-fa, montagnes, J. Cavalerie, no. 1695, March 22, 1902 (holotype of P. Cavaleriei; photo. in A.
1See Vol. 10: 108. 2See Vol. 10: 108.
1936] REHDER, LIGNEOUS PLANTS DESCRIBED BY LEVEILLE 55
When I enumerated Léveillé’s species cited above in 1929, I had not seen any of the type specimens. I have since had the opportunity to examine the types, and find that they all belong to P. Massoniana and bore already on the sheets identification labels, two (P. Cavaleriei and P. Argyi var. longevaginans) by V. Komarov, and the other two signed by M. Y. O[rr].
Cunninghamia lanceolata (Lamb.) Hooker in Bot. Mag. 54: t. 2743 (1827).
Cunninghamia sinensis var. prolifera Lemée & Léveillé in Monde PI. ser. 2, 16: 20 (1914). — Léveillé, Cat. Pl. Yun-Nan, 56
CuinA. Yunnan: Tong-tchouan, pagode, alt. 2500 m., E. E. Maire, March [1910-13] ‘“‘rameaux secondaires pendants, naissant tous du centre d’un fruit” (holotype of C. sinensis var. prolifera; photo. in A. A.)
The abnormality that the axis of the cone continues growing into a leafy branch is found occasionally in this species, e.g. in Wilson’s no. 4076 from Szechuan. It can hardly be considered a variety; at best it is a monstrous form.
Cryptomeria japonica (L.f.) D. Don in Trans. Linn. Soc. 18: 166,
t. 13, fig. 1 (1841). — Léveillé, Fl. Kouy-Tchéou, 111 (1914); Cat. Pl. Yun-Nan, 56 (1916).
Cupressus Mairei Léveillé, Cat. Pl. Yun-Nan, 56 (1916). —Synon.
nov. ae er Duclouxiana Hickel sensu Rehder in Jour. Arnold Arb. 110 (1929), quoad syn. Cupressus Mairei; non Hickel.
a, Yunnan: plaine de Tong-tchouan, pagodes, alt. 2500 m., E. E. Maire, Jan. [1910-13], “grand et superbe resineux, petits rameaux decourbants.” (holotype of Cupressus Mairei; photo. and merotype in A. A.)
In 1910 when I referred Cupressus Mairei doubtfully to C. Du- clouxiana Hickel, I had not seen the type specimen.
Thuja orientalis Linnaeus, Sp. Pl. 1002 (1753). — Léveillé, FI.
Kouy-Tchéou, 112 (1914); Cat. Pl. Yun-Nan, 56 (1916). Thuja orientalis L. var. Argyi Léveillé & Lemée in Monde PI. ser. 2, 17:15 (1915).
CutInA. Kiangsu: Ch. d’Argy, [1848-66] (holotype of Th. orientalis var. Argyi; photo. and merotype in A. A.).
The specimen which bears staminate flowers differs slightly from the usual form in its more slender and distant branchlets, but the differences are hardly sufficient to distinguish it as a variety or form.
56 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XVII
Juniperus squamata Lambert, Descr. Gen. Pinus, 2: 17 (1824). — Rehder & Wilson in Jour. Arnold Arb. 9:17 (1928).— Handel- Mazzetti, Symb. Sin. 7: 8 (1929).
Juniperus ike ih Léveillé, Cat. Pl. Yun-Nan, 57 (1916), nom. seminud. — Komarov in a Syst. Herb. Hort. Bot. Reipub. Ros 5: 30 (1924), cum ds
CHINA. Yunnan: au an du col de Yentzchay, J. Delavay, no. 3112, July 19, 1887 “arbrisseau a tiges redressées” (ex Komarov; syntype in herb. Paris, photo. in A. A.).
Juniperus Franchetiana, of which the type could not be located in the Léveillé herbarium is known to me only from a photograph of a Delavay specimen in the Paris herbarium. Komarov distinguishes it from J. Fargesii and from J. squamata chiefly by its fruit and seed char- acters, while Handel-Mazzetti l.c. places it under J. squamata.
Juniperus Franchetiana is based by Léveillé on J. recurva Franchet, non Hamilton, which in Plantae Wilsonianae (2: 58. 1914) was cited as a synonym of J. squamata, Franchet (Pl. David. 1: 292. 1884) cites under J. recurva a specimen collected in Shensi without any further remarks; the type of J. recurva Franchet, non Hamilton, and conse- quently the type of J. Franchetiana, would therefore be the David specimen from Shensi and not Delavay’s specimen from Yunnan.
Juniperus squamata Lamb. var. Fargesii (Komar.) Rehder & Wilson in Sargent, Pl. Wilson. 2: 59 (1914).
Juniperus Fargesii Komarov in sched. Herb. Paris, Feb. 1911; in Not. Syst. Herb. Hort. Bot. Reipub. Ross. 5: 30 (1924), cum descript. Juniperus Lemeeana Léveillé & Blin in Léveillé, Fl. Kouy-Tchéou, (1914). — Handel-Mazzetti, Symb. Sin. 7:7 (1929).
Cutna. Kweichou: Gan-chouen, J. Cavalerie, no. 3922 (holotype of J. Lemeeana; photo. in A. A.)
By Komarov and by Handel-Mazzetti this juniper is considered a distinct species, but it seems closely connected with J. sguamata by inter- mediate forms, and I prefer to keep it, at least for the present, as a variety of the latter. If considered a distinct species, the correct name will be J. Lemeeana Lévl. & Blin which has priority.
GNETACEAE Gnetum Cavaleriei Léveillé, Fl. Kouy-Tchéou, 186 (1914).— Markgraf in Engler & Prantl, Nat. Pflanzenfam. ed. 2, 13: 440 (1926). Curna, Kweichou: Pin-fa, sur un arbre au sud a Pai-tchen, J. Cavalerie, no. 2336, March 30, 1905 (holotype; merotype in A. A.). The specimen is fragmentary; it consists of bare branches and broken,
1936] REHDER, LIGNEOUS PLANTS DESCRIBED BY LEVEILLE 57
immature, catkin-like inflorescences covered with a ferrugineous woolly tomentum. By Markgraf (in Bull. Jard. Buitenzorg, ser. 3, 10: 490. 1930) it was referred to Populus, but it certainly does not belong to that genus. From a certain resemblance of the branches and the pubescence to Lannea, I thought it might belong to the Anacardiaceae, and Dr. I. W. Bailey who kindly examined the wood structure, made the following statement: ‘The plant has five traces and five gaps at each node and the wood is clearly ring porous. The various anatomical characters, as a whole, would appear to place the plant in either the Anacardiaceae or the Burseraceae. I should be inclined to place it in the Anacardiaceae except for the fact that this family is stated to have trilacunar nodes. The Burseraceae have five traces, but I know of no species which is ring porous. The transition from three to five traces is an easy one and it may be that certain Anacardiaceae have five traces.”
To the Gnetaceae Léveillé (l.c.) doubtfully refers Cavalerie no. 2764 from Pin-fa, Li-tséou-gai, April 1906, “grand arbre a tronc écailleux, les rameaux coupés laissent couler une sorte de colle.” The specimen consists of branches with long pendulous staminate catkins which slightly resemble the spikes of Gnetum. It belongs to Carpinus and is probably referable to C. Fangiana Hu with which it agrees in the large winter-buds with strongly striate scales.
ARACEAE
Rhaphidophora Dunniana Léveillé in Fedde, Rep. Spec. Nov. 9: 325 (1911); Fl. Kouy-Tchéou, 39 (1914
CuinaA. Kweichou: Kuao-siang, lieux ombreux et frais, foréts profonds et ruisseaux, J. Esquirol, no. 246, Jan. 1905, “gros lierre qui monte aux arbres et rochers” (holotype; photo. in A. A.).
The specimen consists only of some bare branches, an immature in- florescence and detached leaves which measure 8-15 & 5-—9.5 cm. and are abruptly contracted into a short acumen about 5 mm. long. It is similar to Rh. Calophyllum Schott from Khasia and Sikkim, but may possibly belong to Scindapsus.
STEMONACEAE
Stemona japonica (Bl.) Miquel, Prol. Fl. Jap. 386 (1867). Helwingia Argyi Léveillé & Vaniot in Bull. Herb. Boissier, ser. 2, 6: 506 (1906). Stemona ae (Lévl. & Vant.) Léveillé in Fedde, Rep. Spec. Nov. 1 (1912); in Mem. Acad. Ci. Barcelona, ser. 3, 12: 560 (Cat. Pl. Kiang-sou, 20) (1916). — Synon. nov.
58 JOURNAL OF THE ARNOLD ARBORETUM [VoL. XVII
Cumna. Kiangsu:_ Sou-tchéou-fou, Ch. d’Argy [1848-66] (syntype of Helwingia Argyi; merotype in A. A.). Anhwei: Ning-koue-fou, Ch. d’Argy [1848-66] (syntype of H. Argyi; ex Léveillé).
Neither C. H. Wright (in Jour. Linn. Soc. Bot. 32: 493. 1896), nor Schlechter (in Notizbl. Bot. Gard. Mus. Berlin, 9: 193. 1924) credit S. japonica to China, but Hemsley (in Jour. Linn. Soc. Bot. 36: 95. 1903) cites a specimen from Kiangsu collected by Carles (no. 357) on the hills near Shanghai.
The two specimens in the Léveillé herbarium are in a cover which bears on the outside the localities “Sou-tchéou Ning-koue-fou,” but there is no indication from which locality each specimen came, nor any reference to Anhwei (Ngan-hoei).
LILIACEAE!
Smilax herbacea L. var. acuminata Wright in Jour. Linn. Soc. Bot. 36: 97 (1903). Smilax herbacea L. var. foetida Léveillé in Mem. Pontif. Accad. Nuov. Lincei, 23: 351 (Liliac. 23) (1905); Fl. Kouy-Tchéou, 256 14
914). Smilax aa L. var. heterophylla Léveillé l.c. (1905).
CuinaA. Kweichou: Pin-fa, bois, bords des torrents, lieux ombrageux, J. Cavalerie, no. 1314, May 25, 1902 (holotype of S. herbacea var. foetida; photo. in A. A.); same locality and collector, no. 1314, May 22, 1902 (holotype of S. herbacea var. heterophylla; photo. in A. A.).
Both of Léveillé’s varieties cited above are referable to var. acuminata Wright with which they agree in the gradually acuminate leaves, but they differ widely in the width of their leaves, var. heterophylla having on one branch leaves about 8 cm. wide, on the other 2 cm. in width, while var. foetida is intermediate with about 3—4 cm. width.
Smilax glabra Roxburgh, Fl. Ind. ed. 2, 3: 792 (1832). — Léveillé, Fl. Kouy-Tchéou, 256 (1914). — Rehder in Jour. Arnold Arb, 10: 110 (1929).— Wang & Tang in Sinensia, 5: 417 (1934).
Smilax glabra Roxb. var. maculata Bodinier ex Léveillé in Mem. Pontif. Accad. Nuov. Lincei, 23: 351 (Liliac. 23) (1905).
Smilax Dunniana Léveillé in Fedde, Rep. Spec. Nov. 9: 446 (1911).
Smilax Blinii Léveillé, Fl. Kouy-Tchéou, 256 (1914).
Cutna. Kweichou: Ma-jo, J. Cavalerie, no. 2973, July 1908 (holotype of S. Dunniana and S. Blinii; merotype in A. A.). Hong-
1See Vol. 10: 110.
1936] REHDER, LIGNEOUS PLANTS DESCRIBED BY LEVEILLE 59
kong: dans les rocailles d’un torrent 4 mi-céte du Pic Victoria, E. Bodinier, Feb. 24, 1893 (holotype of S. glabra var. maculata; photo. in
Smilax Dunniana is not cited in Flore du Kouy-Tchéou, but S. Blinii is described as a new species in that work, based on the same number. The name S. Dunniana does not appear on the type specimen in the Léveillé herbarium.
Smilax glabra var. maculata is hardly different enough to be main- tained as a variety; it is at best a slight form. Specimens of S. glabra with leaves variegated with irregular white spots occasionally occur, e.g. T. N. Hsiung 494 from Kiangsi, J. B. Norton 1252 from Fukien and U. Faurie 950 from Formosa.
Smilax microphylla Wright in Kew Bull. 1895: 117.— Wang & Tang in Sinensia, 5: 418 (1934).
Smilax Labordei Léveillé & Vaniot in Mem. Pontif. Accad. Nuov. Lincet, 23: 355 (Liliac. 27) (1905). — Léveillé, Fl. Kouy-Tchéou, 257 (1914).
Smilax gracillima Léveillé & Vaniot, 7 354 (Liliac. 26) (1905). — Léveillé, FI. Kouy-Tchéou, 256 (1914).
Smilax Darrisii Léveillé in Fedde, Rep. Spec. Nov. 12: 533 (1913); Fl. Kouy-Tchéou, 256 (1914).—Rehder in Jour. Arnold Arb. 10: 111 (1929).— Wang & Tang in Sinensia, 5: 419 (1934).— Synon. nov.
Smilax Mairei Léveillé in Bull. Géog. Bot. 25: 39 (1915); Cat. PI. Yun-Nan, 169 (1916).—Rehder in Jour. Arnold Arb. 10: 111 (1929).
Cuina. Kweichou: environs de Kouy-yang, mont. du Col- lége, rocailles, June 2, 1898 (flowers), environs de Tsin-gay, Dec. 9, 1897 (fruit), J. Laborde in herb. Bodinier, no. 2318 “petit buisson épineux” (holotype of S. Labordei; photo. in A. A.); Tsin-gay, bois, J. Cavalerie, no. 1156, July 15, 1903 (holotype of S. gracillima, photo. in A. A.); steppes et bois de Gan-chouen, J. Cavalerie, no. 3819, Dec. 1909, J. Esquirol, no. 3145, May, 1911 (syntypes of S. Darrisii; mero- type of Cavalerie’s and sitioks: of Esquirol’s specimen in A. A.). Yunnan: rochers a mi-mont de La-kou, 2400 m., E. E. Maire, June 1912 (holotype of S. Mairei; photo. in A. A.).
Smilax Darristi is enumerated as a distinct species by Wang & Tang (l.c.) as it was by me (l.c.), but I do not see now that it can be sepa- rated from S. microphylla. On the other hand, Wang & Tang refer S. castaneiflora Lévl. to S. microphylla, but that is undoubtedly a different species.
60 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XVII
Smilax scobinicaulis Wright in Kew Bull. 1895: 117.
Smilax brevipes Warburg in Bot. Jahrb. 28: 256 (1900).— Wang &
ang in Sinensia, 5: 419 (1934).
Smilax ocreata Léveillé & Vaniot apud Léveillé in Mem. Pontif. Accad. Nuov. Lincei, 23: 354 (Liliac. 26) (1905). — Léveillé, FI. Kouy-Tchéou, 257 (1914).— Non A. De Candolle (1878).
Smilax Martini Léveillé & Vaniot in Léveillé in Mem. Pontif. Accad. Nuov. Lincei, 23: 355 (Liliac. 26) (1905). — Léveillé, Fl. Kouy- Tchéou, 257 (1914).
Smilax Cavalerici Léveillé & Vaniot apud Léveillé in Mem. Pontif. Accad. Nuov. Lincei, 23: 355 (Liliac. 27) (1905). — Léveillé, Fl. Kouy-Tchéou, 257 (1914) ; Cat. Pl. Yun-Nan, 168 (1916).
Cutna. Kweichou:_ environs de Gan-pin, au Grandes Rocailles, trainant sur les rochers, L. Martin in herb. Bodinier, no. 1871, Sept. 17, 1897 (holotype of S. ocreata; photo. in A. A.); Pin-fa, bois, J. Cavalerie, no. 1315, May 3, 1902 (holotype of S. Martini; photo. in A. A.); Pin-fa, montagnes, J. Cavalerie, no. 1313, May 3, 1902 (holo- type of S. Cavaleriei; photo. in A. A.).
The three species of Léveillé’s had been referred by Wang & Tang to S. brevipes Warb. which can hardly be separated from S. scobinicaulis as a distinct species.
Smilax leucocarpa Léveillé & Vaniot apud Léveillé in Mem. Pontif. Accad. Nuov. Lincei 23: 354 (Liliac. 26) (1905); 24: 344 (Liliac. 10) (1906). — Wang & Tang in Sinensia, 5: 420 (1934).
Smilax Esquirolii Léveillé, Fl. Kouy-Tchéou, 256 (1914).
Curna. Kweichou: environs de Kouy-yang, dans les bois de Kien-lin-chan, E. Bodinier, no. 2124, Apr. 14 (flowers) and July 7 (fruit), 1898, ‘“arbuste épineux, a tiges dressées, non sarmenteuses”’ (holotype of S. leucocarpa; photo. in A. A.); plateau de San-tou, 1000 m., J. Esquirol no. 2109, May 1, 1910 (type of S. Esquirolii; photo. in A. A.); Gan-pin, J. Cavalerie, no. 2120 (op. cit. 24: 344 under S. leuco- carpa; photo, in A, A.)
This species resembles somewhat in its small leaves S. microphylla Wright, but the leaves are cuneate and obtuse and the fruits are yellow; from the related S. megalantha Wright and S. China L. it is at once dis- tinguished by the very small obovate to obovate-oblong obtuse leaves and smaller fruits on shorter peduncles.
In the Flore du Kouy-Tchéou Léveillé cites S. leucocarpa as a syno- nym of S. Esquirolii, a name which appears on the sheet of Esquirol 2109, but does not seem to have been published before 1914.
Smilax megalantha Wright in Kew Bull. 1895: 118.— Wang & Tang in Sinensia, 5: 420 (1934).
1936] REHDER, LIGNEOUS PLANTS DESCRIBED BY LEVEILLE 61
Smilax loupouensis Léveillé in Bull. Géog. Bot. 25: 38 (1915) ; Cat. Pl. Yun-Nan, 169 (1916).
Smilax ferox Wall. sensu Rehder in Jour. Arnold Arb. 10: 110 (1929), p-p., quoad synon. S. loupouensis; non Wall. ex DeCandolle.
Cuina. Yunnan: “brousse des mont. a Lou-pou, 3000 m., E. E. Maire, Sept. 1912, ‘“arbuste grimpant, un peu épineux, feuilles blanches au dessous, fleurs grises” (holotype of S. loupouensis ; mero- type in A. A.).
Smilax loupouensis resembles S. discotis Warb. in its stipules extend- ing the entire length of the short petiole, but that species has black fruits. From S. ferox, to which it seems closely related, it differs in the only slightly spiny branches, in the leaves being abruptly contracted at the apex and very glaucous beneath and in the stipules. It apparently agrees best with S. megalantha, but differs from the type in the shorter petioles with the stipules extending the whole length.
Smilax China Linnaeus, Spec. Pl. 1029 (1753).— Léveillé, FI. Kouy-Tchéou, 256 (1914), p.p. excl. syn. S. Bodinieri; Cat. Pl. Yun- Nan, 168 (1916). — Norton in Sargent, Pl. Wilson. 3: 4 (1916).
Smilax Taquetii Léveillé in Fedde, Rep. Spec. Nov. 10: 372 (1912). Smilax China L. f. obtusa Léveillé in Mem. Pontif. Accad. Nuov. Lincei, 24: 344 (Liliac. 10) (1906).
Korea. Quelpaert: in silvis Hogno et Hallisan, E. Taquet, nos. 3306, 3307, 3308, Jan., May, Aug. 1909 (syntypes of S. Taquetii; photo. of 3306 and isotypes of nos. 3306, 3307 and 3308 in A. A.).
Cuina. Kiangsu: Tou-gou-lin, Ch. d’Argy, [1848-66] (holo- type of S. China f. obtusa; photo. in A. A.).
Smilax Taquetii was first identified with S$. China by Norton (l.c.). Smilax China f{. obtusa does not seem to be different from typical S. China.
Smilax glauco-china Warburg in Bot. Jahrb. 29: 255 (1900).— Wang & Tang in Sinensia, 5: 420 (1934).
Smilax Bodinieri Léveillé & Vaniot apud Léveillé in Mem. Pontif. Accad. Nuov. Lincei, 23: 355 (Liliac. 27) (1905).
Smilax China “L.” sensu Léveillé, Fl. Kouy-Tchéou 256 (1914), quoad syn. S. Bodinieri; non Linnaeus.
Cutna. Kweichou: monts de Hin-y-fou et Hin-y-hien, dans les haies, E. Bodinier, no, 1581, Apr. 13, 1897 (holotype of S$. Bodiniert ; photo. in A. A.).
Smilax Bodinieri was cited as a synonym of S. China L. by Léveillé in his Flore du Kouy-Tchéou, but Wang & Tang referred it to S. glauco- china.
62 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XVII
Smilax menispermoidea A. De Candolle, Monog. Phan. 1: 108 (1878). — Wang & Tang in Sinensia, 5: 420 (1934). Smilax luteocaulis Léveillé in Fedde, Rep. Spec. Nov. 13: 339 (1914) ; Cat. Pl. Yun-Nan, 169 (1916). Cuina. Yunnan: brousse du mont To-chan, 3200 m., E. E. Maire, June, 1913 (holotype of S. /uteocaulis ; merotype in A. A.). The specimen agrees perfectly with typical S. menispermoidea.
Smilax stemonifolia Léveillé & Vaniot apud Léveillé in Mem. Pontif. Accad. Nuov. Lincei 23: 356 (Liliac, 28) (1905).— Wang in Bull. Fan Mem. Inst. Biol. 5: 117 (1934).
Cuina. Hongkong: bois le long de Bowenroad, trés rare, E. Bodinier, no. 804, Aug. 4, 1894 (holotype; photo. in A. A.).
The type specimen of this species is sterile, but Wang (l.c.) refers to it two other specimens, one from Hunan collected by H. F. Chow and P. Ou-yang, no. 46102, with young fruit, and one from Anhwei col- lected by F. T. Wang, no. 23860, sterile. Wang states that it is closely related to S. Bockii Warb. which differs in its much narrower leaves rounded at the base and in longer pedicels. It may also be near S. longipes Warb.
Smilax castaneiflora Léveillé in Bull. Acad. Géog. Bot. 25: 39 (1915); Cat. Pl. Yun-Nan, 168 (1916).— Rehder in Jour. Arnold Arb. 10: 111 (1929).
Smilax microphylla “Wright” sensu Wang & sea in a 5: 418 (1934) quoad synon. S. castanetflora; non Wri
CHINA. Yunnan: brousse des mont. a nitie ee, alt. 2700 m., E. E. Maire, June 1912, “arbuste grimpant, fleurs chocolat, fruits noirs” (holotype; merotype in A. A
In general appearance this species resembles S. Bockii Warb., but the branches are angled and quite spiny, the leaves are short-cordate, lanceolate, broad at the base and partly constricted below the middle into the narrow long-acuminate upper part. By Wang & Tang the species was referred to S. microphylla Wright, but it differs from that species in the long peduncle, the gradually acuminate cordate leaves, often somewhat restricted below the middle and green beneath. I have been unable to identify this species with any Chinese species I know. Handel-Mazzetti, no. 10379, from Kweichou may belong here, but it has broader leaves and is not spiny.
Smilax micropoda A. DC. var. reflexa Norton in Sargent, Pl. Wil- son. 3: 6 (1916). Smilax Lebrunii Léveillé, Fl. Kouy-Tchéou, 257 (1914). Pehies & Tang in Sinensia, 5: 421 (1934), “Lebranii.” — Synon
1936] REHDER, LIGNEOUS PLANTS DESCRIBED BY LEVEILLE 63
CHINA. Kweichou: bois sacré de Ban-gnien, 1100 m., J. Esquirol, no. 3518, March 19, 1912 (holotype of S. Lebrunii; photo. in
Wang & Tang keep S. Lebrunii as a distinct species and refer to it three specimens from Kwangsi, R. C. Ching 5846, 6992 and 7201 which I have not seen.
Smilax tortopetiolata Léveillé & Vaniot apud Léveillé in Mem. Pontif. Accad. Nuov. Lincei 23: 354 (Liliac. 26) (1905); Fl. Kouy- Tchéou, 258 (1914). — Wang & Tang in Sinensia, 5: 422 (1934), excl. synon.
Smilax opaca (A. DC.) Norton in Sargent, Pl. Wilson. 3: 11 (1916). — Synon. nov.
CutInA. Kweichou: Pin-fa, bois, J. Cavalerie, no. 1312, March 18, 1902 (holotype of S. tortopetiolata; photo. in A. A.).
This species had not yet been reported from western China; it seems common in southeastern China and is represented in this herbarium by many specimens from Fukien, Kwangtung, Hainan and Kwangsi. I have been unable to find any specific difference between S. opaca and S. tortopetiolata which has to replace the former on account of priority.
Smilax cocculoides Warb. var. lanceolata Norton in Sargent, Pl. Wilson. 3: 11 (1916).
Smilax Pinfaensis Léveillé & Vaniot apud Léveillé in Mem. Pontif. Accad. Nuov. Lincei, 23: 355 (Liliac. 27) (1905). — Léveillé, Fl. Kouy-Tchéou, 257 (1914). — Synon. nov.
Smilax tortopetiolata “Lévl. & Vant.” sensu Wang & Tang in Sinensia, 5: 422 (1934), p.p., quoad synon. S. pinfaensis.
Cutna. Kweichou: Pin-fa, J. Cavalerie no. 1311, March 18, 1902, “fl. verte” (holotype of S. pinfaensis ; photo. in A. A.).
Smilax pinfaensis is apparently identical with S. cocculoides var. lanceolata Norton, the type of which is in fruit, and which I have before me. The flowers agree with those of typical S. cocculoides, as repre- sented by Wilson, no. 3252. Wang & Tang (l.c.) refer S. pinfaensis as a synonym to S. tortopetiolata, but it is easily distinguished from the latter by the narrow-lanceolate, long-acuminate leaves, the slender peduncles much longer than the petioles and the short stamens.
Smilax Lyi Léveillé in Fedde, Rep. Spec. Nov. 8: 171 (1910); Fl. Kouy-Tchéou, 257 (1914). — Wang & Tang in Sinensia, 5: 421 (1934).
Cuina. Kweichou: Pin-fa, montagnes, J. Cavalerie, no. 1406, Sept. 27, 1903 ‘fl. vert-jaune” (holotype; photo. in A. A.).
64 JOURNAL OF THE ARNOLD ARBORETUM [voL. xv
This species was kept distinct by Wang & Tang. The rather meagre specimen which consists of a branch with two leaves and an inflo- resence in bud, resembles S. stenophylla A. Gray from the Liukiu Islands and Formosa, but with the scant material at hand, I hesitate to consider these geographically widely separated species identical.
Smilax perfoliata Loureiro, Fl. Cochinch. 622 (1790).— Wang &
Tang in Sinensia, 5: 422 (1934). Smilax perulata Léveillé & Vaniot in Fedde, Rep. Spec. Nov. 9:78 (1910). — Léveillé, Fl. Kouy-Tchéou, 257 (1914).
Cuina. Kweichou: Lo-fou, J. Cavalerie, no. 3648, March 1909 (holotype of S. perulata; merotype in A. A.).
This species was identified first by Wang & Tang with S. perfoliata Lour. (S. ocreata A. DC.).
Heterosmilax Gaudichaudiana (Kunth) Maximowicz in Bull. Acad, St. Pétersb. 17: 176 (1872); in Mél. Biol. 8: 415 (1872).— A. DeCandolle, Monog. Phan. 1: 44 (1878). — Léveillé in Mem. Pontif. Accad. Nuov. Lincei, 23: 350 (Liliac. 22) (1905).
Heterosmilax Gaudichaudiana var. latifolia Bodinier ex Léveillé 1.c. 1905).
Cuina. Hongkong: bord de l’Aquéduc, prés Richmond Ter- race, FE. Bodinier, Aug. 22, 1894 (holotype of H. Gaudichaudiana var. latifolia; photo. in A. A.).
In the shape and measurements of the leaves Léveillé’s variety agrees with DeCandolle’s var. « Gaudichaudiana (1.c.), which represents the type of the species. It seems doubtful whether this and var. 3 hong- kongensis (Seem.) A. DC., (l.c.) are sufficiently distinct to be main- tained as varieties.
TACCACEAE
Tacca Esquirolii (Lévl.), comb. nov. Clerodendron Esquirolii Léveillé in 7 Rep. Spec. Nov. 11: 298 (1912) ; Fl. Kouy-Tchéou, 439 (1915). Tacca Paxiana Limpricht f. in Engler, Pflanzenreich, IV. 42: 16 (1928).— P’ei in Mem. Sci. Soc. China, 1, no. 3, p. 162 (1932). Cuina. Kweichou: bois de Ta-Thamn, trés ombreux et chaud, J. Esquirol, no. 2802, May 1912, “toute l’inflorescence rouge, lie de vin, feuilles radicales, fleur 1 m. de haut sur tige simple” (holotype of Clero- dendron Esquirolii; photo. in A. A.). Léveillé’s species was first identified by the writer with Tacca Paxtana (see P’ei, l.c.).
1936] REHDER, LIGNEOUS PLANTS DESCRIBED BY LEVEILLE 65
SALICACEAEF}
Populus adenopoda Maximowicz in Bull. Soc. Nat. Moscou, 54: 50 (1879). — Léveillé, Fl. Kouy-Tchéou, 380 (1915); Cat. Pl. Yun-Nan, 250 (1917). — Handel-Mazzetti, Symb. Sin. 7: 58 (1929).
Populus macranthela Léveillé & Vaniot in Bull. Soc. Bot. France, 142 (March 1905); in Monde Pl. 12:9 (1910); in Fedde, ep. Spec. Nov. 8: 446 (1910).— Rehder in Jour. Arnold Arb. 10: 111 (1929). Populus Duclouxiana Dode in Bull. Soc. Nat. Hist. Autun, 18: 190, t. 11, f. 329 (Extr. Monog. ined. Populus, 32) (1905). Populus alee aa Griff. var. Duclouxiana (Dode) Gombocz in Mat mesz. Kozl. 30: 130 (Monog. ae Populi) (1908). — Schneider i in Sargent, Pl. Wilson. 3: 25 (19 Populus rotundifolia Griff. « macranthela ae in Bot. Kozl. 10: 25, (7) (1910). — Léveillé, Fl. Kouy-Tchéou, 380 (1915).
Cuina. Kweichou: Pin-fa, J. Cavalerie, no. 974, April 1, 1903, “grande arbre” (holotype of P. macranthela; photo. in A. A.). Yunnan: _ Ko-kouy prés Tchao-tong, Pére Marc Mey in herb. F. Ducloux, no, 667, in 1906 (in hb. Léveillé and in hb. Univ. Calif. sub P. macranthela ; isotypes of P. Duclouxiana Dode; photos. in A. A.).
In my earlier account of P. macranthela, I had considered it a species distinct from P. adenopoda, chiefly on account of the absence of the glands at the base of the blade of the leaves, their glabrousness and coarser serration. In comparing a large series of specimens of P. adeno- poda one finds a great variability in these characters; though the glands are usually conspicuous, they may occasionally be small and partly lacking, as in Wilson’s no. 724 from Hupeh, and in Steward, Chiao and Cheo, no. 294, from Kweichou. One of the two mature leaves of Duclours specimen in the herbarium of the University of California has an almost suborbicular shape, but on the material from trees cultivated in the Arnold Arboretum similar leaves can be found. Both specimens I have seen of Ducloux’s no. 667 bear the name P. macranthela in Lé- veillé’s handwriting. Neither Dode nor Gombocz give an exact citation of the specimen upon which P. Duclouxiana is based, but I assume that the species is based on Ducloux’s no. 667.
Salix Cavaleriei Léveillé. — Rehder in Jour. Arnold Arb. 10: 113 (1929). — Handel-Mazzetti, Symb. Sin. 7: 61 (1929).
Salix erioclada Léveillé.— Rehder, l.c. 115 (1929). — Handel- Mazzetti, l.c. 64, fig. 1,1, 2 (1929).
Handel-Mazzetti gives a detailed description of the species and figures staminate and pistillate flowers.
1See Vol. 10: 111.
66 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XVI
Salix luctuosa Léveillé.— Rehder, lc. 115 (1929). — Handel- Mazzetti, I.c. 68 (1929).
Handel-Mazzetti gives a full description of the pistillate flowers and refers S. dyscrita Schneid. as a synonym to this species.
Salix Duclouxii Léveillé. — Rehder, l.c. (1929).— Handel-Maz- zetti, lc. 87 (1929).
Salix kouytchensis (Lévl.) Schneider in Sargent, Pl. Wilson. 3: 171 (1916). — Rehder, I.c. 117 (1929).
Handel-Mazzetti unites S. kouwytchensis (Lévl.) Schneid. with S. Duclouxii and discusses the close relation of the latter to S. Bocki Seemen and S. variegata Franch., but prefers to keep these species dis- tinct for the present.
MYRICACEAE!
Myrica esculenta Hamilton ex D. Don, Prodr. Fl. Nepal, 56 (1825). Litsea myricopsis Léveillé, Cat. Pl. Yun-Nan, 150 (1916).— Synon. nov.
Cutna. Kweichou: prés Tchang-pou, J. Cavalerie, no. 2697, Nov. 3, 1905, “petit arbre” (holotype of Litsea myricopsis ; merotype in A. A.).
Myrica esculenta has been collected in Kweichou also by Handel- Mazzetti (no. 10852).
Myrica adenophora Hance in Jour. Bot. 21: 357 (1883). — Lé-
veillé, Fl. Kouy-Tchéou, 281 (1914). Litsea Mairei Léveillé, Cat. Pl. Yun-Nan, 150 (1916).— Synon. nov.
Cutna. Yunnan: collines arides incultes de Tché-hay, alt. 2600 m., E. E. Maire, March, 1912 “arbousier, arbuste rameuse, fleur blanc-verdatre” (holotype of Litsea Mairei; merotype in A. A.).
Myrica adenophora does not seem to have been recorded from Yunnan before, nor from Kweichou, for I feel sure that Esquirol no. 3296 cited by Léveillé in his Flore du Kouy-Tchéou under that species (1.c.) does not belong here, but I have not seen it.
BETULACEAF? Alnus Fauriei Léveillé in Bull. Soc. Bot. France, 51: 423 (1904). — Rehder in Jour. Arnold Arb, 10: 118 (1929). APAN. Hondo: in monte Gurvassan, U. Faurie, no. 783, Sept. 28, 1897 (holotype; photo. in A. A.). The type specimen bears an identification label by Koidzumi with
1See Vol. 10: 118. 2See Vol. 10: 118.
1936] REHDER, LIGNEOUS PLANTS DESCRIBED BY LEVEILLE 67
the legend “Alnus cylindrostachya (Winkl.) Koidz.”” Though Wink- ler’s name, A. glutinosa var. cylindrostachya, was published June 17, 1904, while Léveillé’s name did not appear before November 1904, Léveillé’s binomial has precedence over Winkler’s varietal name.
Alnus nepalensis D. Don, Prodr. Fl. Nepal. 58 (1825). — Léveillé,
Fl. Kouy-Tchéou, 124 (1914); Cat. Pl. Yun-Nan, 66 (1916). Alnus Mairei Léveillé in Bull. Géog. Bot. 24: 283 (1914).
Cuina. Yunnan: vallées et montagnes de Tong-tchouan, 2500— 2700 m., E. E. Maire, Nov. 1913, “grand arbre, chatons jaunes” (holo- type of A. Mairei; photo. in A. A.).
Alnus Mairei was referred to A. nepalensis by Léveillé himself in 1916.
Carpinus Seemeniana Diels in Bot. Jahrb. 29: 279 (1900).— Schneider in Sargent, Pl. Wilson. 2: 430 (1916). Carpinus pinfaensis Léveillé & Vaniot in Bull. Soc. Bot. France, 52: 142 (1905). Carpinus pubescens Burkill sensu Winkler in Bot. Jahrb. 51: 501 (1914), p.p., quoad syn. C. pinfaensis et C. Seemeniana. — Léveillé, Fl. Kouy-Tchéou, 125 (1914).—Hu in Sunyatsenia, 1: 119 (1933). Cuina. Kweichou: Pin-fa, J. Cavalerie, no. 1011, May 28, 1903, “arbre” (holotype of C. pinfaensis; photo. in A. A Carpinus pinfaensis was first referred to C. pubescens (including C. Seemeniana) by Winkler in 1914, but in 1916 it was placed under C. Seemeniana by Schneider, where it certainly belongs if C. Seemeniana and C. pubescens are considered distinct. These two species and also C. austrosinensis Hu are closely related and perhaps best united under C. pubescens with one or two varieties, but at present I prefer to keep C. Seemeniana as a species, since the extreme forms are quite distinct. Carpinus austrosinensis is nearer to the latter.
Carpinus laxiflora Blume, Mus. Bot. Lugd.-Bat. 1: 309 (1850) :—
Schneider in Sargent, Pl. Wilson. 2: 438 (1916). ? Carpinus laxiflora Bl. var. chartacea Léveillé in Bull. Soc. Bot. France, 51: 424 (1904)
JAPAN: without locality and collector (holotype of var. chartacea ; ex Léveillé).
No specimen named C. laxifolia var. chartacea can be found in the Léveillé herbarium, but I have no doubt that it is but a slight variation of C. laxiflora under which it was placed by Schneider in 1916.
68 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XVI
FAGACEAF!
Castanea mollissima Blume in Mus. Bot. Lugd.-Bat. 1: 286 (1850). Castanopsis yunnanensis (Franch.) Léveillé, Cat. Pl. Yun-Nan, 66 (1916), Syn.: “(C. vulgaris L., p.p.)” — Camus, Chataigniers, 485
The combination Castanopsis yunnanensis was published by Léveillé without any citation of a specimen. It is based on Castanea vulgaris var. yunnanensis Franchet which is a synonym of C. mollissima; also the citation by Léveillé of C. vulgaris L., p.p., as a synonym seems to indicate that it is a true Castanea. It can, therefore, hardly be referred to any other species than C. mollissima which is common in Yunnan; probably to this species belongs also C. sativa Mill. cited in Cat. PI. Yun-Nan, 66.
Castanopsis cuspidata (Thbg.) Schottky in Bot. Jahrb. 47: 625
(1912). Castanea Fauriei Léveillé & Vaniot in Bull. Soc. Bot. shige 52: 142 (1905). — Camus, Chataigniers, 241 (1929). — Synon. nov.
JapAN. Kiushu: autour de Nagasaki, U. eRe no. 3681, June 1899 (holotype of C. Fauriei,; isotype in A. A.).
Miss Camus enumerated C. Fauriei under the doubtful species of Cas- tanea with a note that it is probably a Castanopszs.
Castanopsis hystrix (Hook. f. & Th.) A. DeCandolle in Jour. Bot. 1: 182 (1863); Prodr. 167: 111 (1864). — Léveillé, Cat. Pl. Yun-Nan, 66 (1916). — Rehder in Jour. Arnold Arb, 10: 118 (1929).
Castanea de Léveillé & Vaniot in Bull. Soc. Bot. France, 52: 142 (1905). — Léveillé, Cat. Pl. Yun-Nan, 66 (1916).— Camus, Chataigniers, 241 (1929).
Castanopsis Bodinieri (Lévl. & Vant.) Koidzumi in Tokyo Bot. Mag. 30: 100 (1916) ‘“Boodinieri.”
Quercus brunnea Léveillé in Fedde, Rep. Spec. Nov. 12: 364 (1913) ; Fl. Kouy-Tchéou, 127 (1914). — Camus, Chataigniers, 482 (1929) ; Atlas, pl. 28, fig. 9-11, 1x. f. 9-17 (1928).
Castanopsis brunnea (Lévl.) Camus, Chataigniers, Atlas, pl. 28 (1928).
Cu1na. Kweichou: Pin-fa, bois, J. Cavalerie, no. 2299, Apr. 13, 1905 (holotype of Q. brunnea; photo.in A.A.). Yunnan: en- virons de Yun-nan-sen, bois de la pagode de Kiang-tchou-se, E. Bo- dinier, Feb. 2, 1897, “grand arbuste ou petit arbre, fruits en long épis a glands sessiles serrés” (holotype of Castanea Bodinieri ; photo. in A. A.).
Castanea Bodinieri is enumerated by Miss Camus (l.c.) under in-
1See Vol. 10: 118.
1936] REHDER, LIGNEOUS PLANTS DESCRIBED BY LEVEILLE 69
completely known species of Castanea. Quercus brunnea she describes under doubtful species of Castanopsis and says “probablement var. ou sous-espece du C. Hystrix.”
Castanopsis Eyrei (Champ.) Tutcher in Jour. Linn. Soc. Bot. 37: 68 (1905). — Chun in Sunyatsenia, 1: 217, pl. 35 (1934), excl. syn. Q. Castanopsis.
Castanopsis caudata Franchet in Nouv. Arch. Mus. Paris, ser. 3, 7: 87 (Pl. David. 1:277) (1884).—Rehder in Jour. Arnold Arb. 10:119 (1929), excl. syn. Q. Castanopsis. — Handel-Mazzetti, Symb. Sin. 7: 28 (1929).
Quercus trinervis Léveillé in Fedde, Rep. Spec. Nov. 12: 364 (1913) ; Fl. Kouy-Tchéou, 128 (1914
Quercus cepifera Léveille, l.c. 364 (1913) ; lc. 127 (1914), pro parte, quoad specimina foliifera.
Castanopsis asymetrica Léveillé, Fl. Kouy-Tchéou, 125 (1914).— A. Camus, Chataigniers, 472 (1929) “asymmetrica”; Atlas, pl. 72, fig. 15, xxxiv, fig. 17-19 (1928).
Castanopsis neo-Cavaleriet A. Camus, ee ai 375- (1929): Atlas, pl. 72, fig. 1-12, xxx1v, fig. 8-15 (1928).
Castanopsts trinervis (Lévi.) A. Camus, op. cit. 400 (1929); Atlas, pl. 70, fig. 1-7, xxx111, fig. 1-7 (1928).
Castanopsis tribuloides var. echidnocarpa King sensu Rehder in Jour. Arnold Arb. 10: 119 (1929) ; non King apud Hooker f. (1888).
Cuina. Kweichou: J. Cavalerie, no. 3275, Aug. 20, 1908 (holotype of Q. trinervis ; photo. in A. A.) ; sud de Pin-fa, rare, J. Cava- lerie, no. 2341 (in part, except fruit), June 8, 1905 ‘fruits ci-joints trouvés par terre; apartiennent-ils 4 cet arbre” (holotype of Q. cepifera, except fruit; photo. in A. A.); Kouy-yang, monts du College, J. Chaffan- jon in herb. Bodinier, no. 2235, Apr. 15, 1898, ‘‘arbre, fleurs blanches”’ (syntype of Castanopsis asymetrica; merotype in A, A.); Kouy-yang, bois d’une pagode, J. Cavalerie, no. 2078 [cited as 2178 by Léveillé], “arbre” (syntype of C. asymetrica; photo. in A. A.); Kien-lin-chan, J. Esquirol, no. 17, May 2, 1904, “feuilles de camélia” (syntype of C. asymetrica; photo. in A. A.).
I agree with Chun in considering Castanopsis Eyrei and C. caudata identical. Miss Camus places C. asymmetrica, C. Eyrei and C. caudata among the incompletely known species and describes C. trimervis as a distinct species. Handel-Mazzetti cites Castanopsis asymetrica as a synonym of C. caudata Franch.
Castanopsis neo-Cavaleriei A. Camus in Bull. Bimens. Soc. Linn. Lyon, 8: 87 (1929); Chataigniers, 375 (1929) ; Atlas, pl. 72, fig. 1~12, XXIV, fig. 8-15 (1928).
70 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XVII
Quercus Cavaleriei Léveillé & Vaniot in oe Soc. Bot. France, 52: 142 (1905); Fl. Kouy-Tchéou, 127 (191
Quercus Castanopsis Léveillé in rt om Spec. Nov. 12: 363 (1913); Fl. Kouy-Tchéou, 127 (191
Castanopsis caudata Franchet sensu See in sbeakd aaa Arb. 10: 119 (1929), quoad syn. Q. Castanopsis; non Fra
Castanopsis tribuloides var. echidnocarpa King sensu | Rehder, lc. (1929) ; non King ex Hooker f.
Castanopsis eyrei (Champ.) Tutcher sensu Chun in Sunyatsenia, 1: 217 (1934), quoad syn. Q. Castanopsis; non (Champ.) Tutcher.
Cutna. Kweichou: Pin-fa, hautes montagnes, J. Cavalerie, no. 57 (in part; young fruit), July 15, 1902, “petit arbre” (holotype of Quercus Cavaleriei; photo. in A. A.); Pin-fa, Ma-jo, hautes montagnes, rare, J. Cavalerie, no. 57 (in part; flowers), May 1902 (syntype of Q. Castanopsis; merotype in A. A.); same locality, J. Cavalerie, no. 57, Aug. 1908 (syntype of Q. Castanopsis; merotype in A. A.); same lo- cality, J. Cavalerie, no. 1268, July 1908 (syntype of Q. Castanopsis ; ex Léveillé; ? merotype in A. A.).
This species is closely related to C. Eyrei, but differs in the thinly scurfy-tomentulose brownish under side of the leaves — quite glabrous in C. Eyrei — and in the larger, very oblique cupula with more distinct fascicles of spines nearly wanting on the ventral side. From C. Fargesii Franch, it differs in the broader leaves less densely tomentulose be- neath, — in C. Fargesii the dense brown tomentum obscures the vein- lets, — and in the larger apparently indehiscent cupula. Two fruiting spikes are figured by Miss Camus on pl. 72; fig. 3 apparently represents the young fruits of Q. Cavaleriei, while fig. 4-6 represent mature fruits of Q. Castanopsis collected in August.
The name C. neo-Cavaleriei for this species has to stand, since there exists already a Castanopsis Cavaleriei Lévl. of 1913 which, however, is not a Castanopsis, but represents Sloanea sinensis (Hance) Hu (see Jour. Arnold Arb. 15: 89. 1934), and the specific epithet of the second synonym cannot be transferred to Castanopsis, since this would result in a tautonym.
Castanopsis tibetana Hance. — Rehder in Jour. Arnold Arb. 10: 119 (1929),
Quercus Franchetiana Léveillé, Fl. Kouy-Tchéou, 128 (1915).— Camus, Chataigniers, 485 (1929).
Miss Camus states that C. Franchetiana is perhaps identical with C. tibetana ; in a note she says that the leaves of the former are quite differ- ent from those of Hance’s type in shape, are very lustrous above and more waxy and less pubescent beneath but that she has seen an inter- mediate form.
1936] REHDER, LIGNEOUS PLANTS DESCRIBED BY LEVEILLE 71
Castanopsis cryptoneuron (Lévl.) Rehder in Jour. Arnold Arb. 10: 119 (1929).— Camus, ChAataigniers, Atlas, pl. 54, fig. 1-4 (1928), nom, tantum; text, 418 (1929).
Miss Camus cites the combination as dating from her Atlas of 1928, but since the original binomial is not cited there, it cannot be considered as validly published.
Castanopsis spec. Rehder in Jour. Arnold Arb. 10: 120 (1929). Quercus Argyi Léveillé in Mem. Acad. Ci. Art. Barcelona, ser. 3, 12: 548 (Cat. Pl. Kiang-Sou, 8) (1916).
Cuina. Kiangsu: Sonosé, Ch. d’Argy, nos. 842, 856 [1846— 66]. In the cover of Q. Argyi in Léveillé’s herbarium, there are two speci- mens, no. 842 with pistillate and no. 856 with staminate flowers, both belonging apparently to the same species related to C. Fargesii Franch., but they have broader and more serrate leaves. Cavalerie’s no. 2078 (not 2178) of which I stated (l.c.) that it may belong here, belongs to C. Eyrei (Champ.) Tutch. together with the other specimens of Casta- nopsis asymetrica,
Castanopsis spec.
Myrica Cavaleriei Léveillé in Fedde, Rep. Spec. Nov. 12: Sa (1983); pro parte, quoad specim. Q
Cuina. Kweichou: haut plateau, route de Pin-fa a Kouy- yang, J. Cavalerie, no. 3184, June 1908 (holotype of Myrica Cavaleriet, in part; photo. in A. A.).
Cavalerie’s no. 3184 consists of two leafless branches, for which see Quercus spec. (p. 72), and a leafy branch with short axillary spikes of pistillate flowers and with lanceolate slender-petioled leaves 10-12 cm. long and about 2 cm. broad.
Quercus serrata Thunberg. — Rehder in Jour. Arnold Arb. 10: 120 (1929).
Since the name Q. serrata Thbg. which correctly belongs to the species named Q. glandulifera Bl. has been applied by almost all authors, fol- lowing Siebold and Zuccarini, to Q. acutissima Carruthers, it is advis- able to consider Q. serrata a nomen ambiguum and use Q. acutissima and Q. glandulifera for the two species involved.
Quercus guyavaefolia Léveillé in Fedde, Rep. Spec. Nov. 12: 363 (1913); Cat. Pl. Yun-Nan, 67 (1916) “guyavifolia.” — Rehder in Jour. Arnold Arb. 10: 121 (1929).— Camus, Chénes, Atlas, 1: t. 35 (1934).
72 JOURNAL OF THE ARNOLD ARBORETUM [VOL. Xvi
Quercus semecarpifolia Sm. var. longispica Handel-Mazzetti, Symb. Sin. 7: 39 (1929).
Handel-Mazzetti cites Q. guyavaefolia as a synonym of his Q. seme- carpifolia var. longispica, but does not enumerate the type of Léveillé’s species, and apparently bases his description on other specimens from Yunnan and also from Szechuan; Miss Camus’ illustrations also are partly based on other specimens. The type specimen of QO. guyavaefolia has all the leaves entire.
Quercus Prainiana Léveillé. — Rehder in Jour. Arnold Arb. 10: 121 (1929),
This species is figured by Léveillé in the manuscript work Cat. III. Pl. Seu-Tchouan on pl. 23 under the name Q. Prainii.
Quercus glauca Thunberg, Fl. Jap. 175 (1784).— Rehder in Jour. Arnold Arb. 10: 121 (1929). — Handel-Mazzetti, Symb. Sin. 7: 50 (1929).
Quercus Blakei Skan var. Vaniotii (Lévl.) Chun in Jour. Arnold Arb. 9: 153 (1928).
Chun’s new combination is based on Q. Vaniotii Lévl. and on Cava- lerie, no. 3274 (not 3264, as cited), which undoubtedly belongs to Q. glauca, he evidently placed too much faith in Léveillé’s statement, that Q. Vaniotii is very closely related to Q. Blakei Skan. The type speci- men consists only of sterile branches; the leaves show the closely appressed pubescence of Q. glauca, while Q. Blakei has the leaves gla- brous and green beneath, more gradually attenuate at base and with shorter appressed and fewer teeth. Handel-Mazzetti cites (l.c.) QO. Vaniotti as a synonym of Q. glauca.
Quercus spec. — Rehder in Jour. Arnold Arb. 10: 122 (1929). Myrica Cavaleriei Léveillé in Fedde, Rep. Spec. Nov. 12: 537 (1913), pro parte, quoad specim. ¢.
Curna. Kweichou: haut plateau, route de Pin-fa 4 Kouy- yang, J. Cavalerie, no. 3184 in part (4), April, 1908 (holotype of Myrica Cavaleriei, in part; photo. in A. A.).
To Quercus belong two leafless branches with immature staminate catkins; for the pistillate specimen see Castanopsis spec. (p. 71).
MORACEAE! Vanieria tricuspidata (Carr.) Hu in Jour. Arnold Arb. 5: 228 (1924). Cudrania_ tricuspidata Bureau in Lavallée, Arb. Segrez. 243 (1877). — Nakai, Fl. Sylv. Kor. 19: 114 (1932).
1See Vol. 10: 123.
1936] REHDER, LIGNEOUS PLANTS DESCRIBED BY LEVEILLE 73
Morus integrifolia Léveillé & Vaniot in Bull. Acad. Intern. Géog. Bot. 17, no. 210-11, p. iii (1907).— Schneider in Sargent, Pl. Wilson. 3: 301, in nota (1916)
Cuina. Hopei: Tchao-tchao, cimetiére, L. Chanet, no. 49, June 25, 1905. “fruit rouge a la maturité” (holotype of Morus integri- folia; photo. in A. A.)
Nakai (l.c.) seems to have first referred Morus integrifolia to this species. As Cudrania Trecul, though proposed as a nomen conservan- dum, did not receive the unanimous vote of the Committee, I retain here the oldest generic name Vanieria Lour.
As Chanet collected his specimen in a cemetery, it was probably from a planted tree the specimen has not been recorded as growing wild in Hopei, but we have specimens in the herbarium from the adjoining provinces — Shansi, Honan and Shantung.
Ficus gibbosa Bl. var. cuspidifera King. — Rehder in Jour. Arnold Arb. 10: 123 (1929). Ficus cuspidifera Miquel in London on Bot. 7: 434 (1848).— Handel-Mazzetti, Symb. Sin. 7:92 (19 Handel-Mazzetti cites F. rhomboidalis tél, & Vant. as a synonym of F. cuspidifera.
Ficus glaberrima Blume, Bijdr. 451 (1825).
Ficus suberosa Léveillé & Vaniot in Fedde, Rep. Spec. Nov. 8: 549 (1910). — Léveillé, Fl. Kouy-Tchéou, 433 Cee — Rehder in Jour. Arnold Arb. 10: 131 (1929).— Synon. n
Ficus Blinti Léveillé ba Vaniot, I.c. 550 (1910). wn Leveillé, lc. 429 (1915). — Synon. n
Ficus Feddei Léveillé & ‘Vanier op. cit. 9: 19 (1910). — Synon. n
Ficus Kingiana Léveillé, Fl. Kouy-Tchéou, 431 (1915), pro ae non Hemsley. — Synon. nov
Cuina. Kweichou: Lo-fou, J. Cavalerie, no. 3597, March 1909, (holotype of F. suberosa; fragments in A. A.); Pia-ouai-lo, alt. 600 m., J. Esquirol, no. 2197, Sept. 1910 (cited under F. suberosa in FI. Kouy-Tchéou; duplicate in A. A.); Lo-fou, J. Cavalerie, no. 3595, March 1909 (holotype of F. Blinii; merotype in A. A.) ; Lo-fou, J. Cava- lerie, no. 3634, Oct. 1908 (holotype of F. Feddei; photo. in A. A.) ; Hua- kiang, J. Cavalerie, no. 2172, June 3, 1904 (syntype of F. Kingiana; photo. in A. A.).
In his Flore du Kouy-Tchéou, Léveillé refers F. Fedde as a synonym to F. suberosa, but keeps F. Blinii distinct. Cavalerie’s no. 2172 is cited as a syntype of F. Kingiana by Léveillé, but the characters under which F. Kingiana appears in the key, exclude it, and therefore the other syntype which belongs to F. pyriformis Hook. & Arn. must be considered the real type of the species.
74 JOURNAL OF THE ARNOLD ARBORETUM [voL. xvi
Ficus retusa Linnaeus, Mant. 129 (1767). — Léveillé, Cat. Pl. Yun-
Nan, 275 (1917). — Handel-Mazzetti, Symb. Sin. 7: 100 (1929). Ficus retusiformis Léveillé in Fedde, Rep. Spec. Nov. 8: 549 (1910); Fl. Kouy-Tchéou, 433 (1915).
Cuina. Kweichou; Lou-fou, J. Cavalerie, no. 3601, March 1909 (holotype of F. retusiformis; merotype in A. A.).
Ficus retusiformis was first referred to F. retusa as a synonym by Handel-Mazzetti (l.c.).
Ficus lacor Hamilton.— Rehder in Jour. Arnold Arb. 10: 124
(1929 Ficus pili Miq. sensu Handel-Mazzetti, Symb. Sin. 7: 92 (1929), uo . F. Tent; vix Mig
ie pial unnan: * Kiao-kia, Siméon Ten, no. 733, Jan. 5, 1906, “grand arbre” (holotype of F. Tenii in hb. Lévl., isotype in hb. Ducloux in hb. Univ. Calif.; photos. in A. A.).
Ficus Tenu Lévl. was referred by Handel-Mazzetti (l.c.) to F. superba. Ficus pseudoreligiosa Lévl. has smaller longer-stalked recep- tacles and distinctly cuneate leaves.
Ficus obscura Blume, Bijdr. 474 (1825). — Gagnepain in Lecomte, Fl. Gén. Indo-Chine, 5: 792 (1928).
Ficus asymetrica Léveillé & Vaniot in Mem. Acad. Ci. Art. Barcelona, ser. 3, 6: 147 (Ficus Spec. Chin. 9) (1907); in Fedde, Rep. Spec. Nov. 4:82 (1907). — Léveillé, Fl. Kouy-Tchéou, 429 (1915).— Rehder in Jour. Arnold Arb. 10: 128 (1929). — Synon. nov
CuHInA. Kweichou: bord du Hoa-kiang, L. Martin in hb. Bodinier, no. 2577, Feb. 18, 1899 (holotype of F. asymetrica ; merotype in A. A.); Lo-fou, J. Cavalerie, no. 3596, March 1909 (cited in Fl. Kouy- Tchéou [as 3796]; duplicates in A. A.); bord du Ta-ras riviére, J. Esquirol, nos. 2692, July 20, 1911, and 3549 (cited in Fl. Kouy-Tchéou; duplicates in A. A.).
Ficus asymetrica agrees well with F. obscura which has been reported from Kweichou by Gagnepain (l.c.) apparently based on Cavalerie 3596 named in the Paris Herbarium F. obscura. It is a species of wide dis- tribution extending from the Malayan Archipelago to Tonkin, southern China and the eastern Himalayas.
Ficus longepedata Léveillé & Vaniot.— Rehder in Jour, Arnold Arb, 10: 127 (1929). — Handel-Mazzetti, Symb. Sin. 7: 93 (1929).
Handel-Mazzetti gives (l.c.) a complete description of the species, to which he refers F. trichopoda Léveillé and F. sordida Hand.-Mazz. as synonyms.
1936] REHDER, LIGNEOUS PLANTS DESCRIBED BY LEVEILLE 75
Ficus laevis Blume, Bijdr. 437 (1825). Ficus Jamini Léveillé & Vaniot in Fedde, Rep. Spec. Nov. 8: 550 (1910). — Léveillé, Fl. Kouy-Tchéou, 430 (1915).—Rehder in Jour. Arnold Arb. 10: 129 (1929). — Synon. nov. Cutna. Kweichou: Lo-fou, J. Cavalerie, no. 3601, March 1909 (holotype of F. Jamini; merotype in A. A.). The specimen cited represents the typical glabrous form to which be- longs also Henry’s no, 11823 from Yunnan.
Ficus scandens Roxburg, Fl. Ind. 3: 536 (1832).
Ficus heapicagenigt Bodinier ex Léveillé in Mem. Acad. Ci. Art. Bar- celona, ser. 3, 6: 148 (Ficus Spec. China, 10) etry: in Fedde, Rep. Spec. a 4: 83 (1907). — Synon. nov.
Cuina. Hongkong: introduit 4 Hongkong, ot il est main- tenant trés commun sur les murs, E. Bodinier, no, 1253, July 7, 1895 (holotype of F. cantoniensis ; merotype in A. A.).
Ficus scandens is not recorded in Hemsley, Index Florae Sinensis, nor in Dunn & Tutcher, Flora of Kwangtung and Hongkong, but it has been collected more recently in southern Yunnan and in Hainan.
Ficus foveolata Wall.— Rehder in Jour. Arnold Arb. 10: 124 (1929). — Handel-Mazzetti, Symb. Sin. 7: 95 (1929).
Handel-Mazzetti (l.c.) cites F. rufipes Lévl. as a synonym and points out that of the three syntypes cited only Cavalerie 340 belongs here, while Esquirol 75 and 76 represent Rubiaceae; the two last-named syn- types I have not yet seen.
Ficus foveolata var. Thunbergii (Maxim.) King.— Rehder in Jour. Arnold Arb. 10: 125 (1929).
Ficus stipulata Thunberg, es Ficus, 5, 8 (1786).— Nakai, FI. Sylv. Kor. 19: 124, t. 40 (1932).
Ficus hederera Léveillé. — Rehder, l.c. 129 (1929
Ficus Fauriei var. macrocarpa Léveillé in Fedde, Rep. Spec. Nov. 11:65 (1912).
Korea. Quelpaert, add: scandens in rupibus, E. Taquet, no. 5972, Dec. 1911 “feuilles et fruits doubles du F. Fauriez’ (holotype of F. Fauriei var. macrocarpa; photo. in A. A.); Hong-no, in declivibus altis, repens in rupibus et tapetem formans, E. Taquet, no. 4425, July 2, 1910 (holotype of F. hederifolia Lévl.; merotype in A. A.).
Nakai enumerates under F. stipulate. (l.c.) as synonyms F. hedert- folia Lévl. and F. Fauriei var. macrocarpa Lévl.
Ficus Martini Léveillé & Vaniot.— Rehder, l.c. 127 (1929).— Handel-Mazzetti, Symb. Sin. 7: 95 (1929).
76 JOURNAL OF THE ARNOLD ARBORETUM [VOL. xviI
Ficus botryoides Léveillé & Vaniot in Mem. Acad. Ci. Art. Barcelona, ser. 3, 6: 148 (Ficus Sp. Chin. 10) (1907); in Fedde, Rep. Spec. Nov. 4:83 (1907). — Léveillé, FI. ane Tchéou, 429 (1915), — Rehder in Jour. Arnold Arb. 10: 128 (1929).
Ficus lacrymans Léveillé, Fl. Kouy-Tchéou, 431 (1915).— Rehder in Jour. Arnold Arb. 10: 130 (1929).
Ficus kwangtungensis Merrill in Jour. Arnold Arb. 8:3 (1927).— Synon. nov.
Cuina. Kweichou, add: Ta-rin, alt. 900 m., J. Esquirol, no. 5373, April 12, 1913, “figuier pleureur” (holotype of F. lacrymans ; merotype in A. A.); environs de Tsin-gay, rochers au bord de la riviére, a Cha-teou-tchay, FE. Bodinier, no. 2653, June 27, 1899 (holotype of F. botryoides ; merotype in A. A.).
Handel-Mazzetti |.c. places F. lacrymans with F. Martini, and I now think that he is right; leaves of thinner texture with slight reticulation occur occasionally on specimens from various localities and the habit seems to be extremely variable; on many specimens the branchlets are very long and slender and probably hang down from scandent stems, while other specimens are described as upright shrubs, as is the type of F. kwangtungensis. Though I have not seen the type of the last-named species, I have seen specimens from Kwangtung and Hainan determined by its author.
Ficus tikoua Bureau in Jour. de Bot. 2: 213, pl. 7 (1888). — Lé- veillé, Cat. Pl. Yun-Nan, 275 (1917). Ficus Bonatit Léveillé in Fedde, Rep. Spec. Nov. 6: 112 (1908). — 8
ynon. nov. Ficus Bonatiana Léveillé, Cat. Pl. Yun-Nan, 274 (1917).
CuinaA. Yunnan: Yunnan-sen, ravins du mont Tchong-chan, F. Ducloux, no. 732, Aug. 18, 1905 (holotype of F. Bonati in herb. Bonati in Univ. Calif.; photo. in A. A.; isotype in herb, New York Bot. Gard.)
The specimen cited above differs somewhat from most of the material in this herbarium in the uniformly short petioles not exceeding 1 cm.
Ficus erecta Thunberg. — Rehder in Jour. Arnold Arb. 10: 125 (1929). — Handel-Mazzetti, Symb. Sin. 7: 100 (1929). — Nakai, FI. Sylv. Kor. 19: 121 (1932).
Nakai cites F. Taquetii as a synonym of F. erecta, and Handel- Mazzetti cites as synonyms besides F. pseudopiriformis and F. Taqueti also F. Mairei Lévl. which according to fragments from the type speci- men belongs to F. heteromorpha Hemsl. (see Rehder, op. cit. 126).
1936] REHDER, LIGNEOUS PLANTS DESCRIBED BY LEVEILLE 77
Ficus erecta Thunb. var. Sieboldi (Miq.) King in Ann. Bot. Gard. Calcutta, 1: 142 (1888).— Nakai, Fl. Sylv. Kor. 19: 123, t. 39 (1932). Ficus Bhi piriformis Léveillé & Vaniot in Fedde, Ree. Spec. Nov. 5: (1908).— Nakai in Jour. Coll. Sci. Tokyo, 31: 199 (FI. ah (1911). Ficus erecta Thunb. — Rehder in Jour. Arnold Arb. 10: 125 (1929), in Korea. Quelpaert: Hong-no, secus torrentes, U. Faurie, no. 1991, July 1907; in petrosis, U. Faurie, 2024, July 1907 (syntypes of F. pseudo-piriformis ; fragments of no. 1991 in A. A.; no. 2024 ex Lé- veillé) ; secus torrentes, alt. 400 m., E. Taquet, no. 4422, July 28, 1910 (in hb, Léveillé sub F. pseudo-piriformis; merotype in A. A.). According to Léveillé’s description Faurie no. 2024 belongs here, but an isotype of this number in the A. A. herbarium is typical F. erecta. On the other hand, a duplicate of Taquet no. 900, which is cited by Léveillé under F. Taqueti, belongs to var. Sieboldi. Ficus Taqueti, according to Léveillé’s description, represents typical F. erecta, while F. pseudo-piriformis agrees with var. Sieboldi. Nakai (l.c. 1932) cites F. pseudo-piriformis under F. erecta var. Sieboldi.
Ficus hirta Vahl. — Rehder, l].c. 126 (1929). — Handel-Mazzetti, Symb. Sin. 7: 99, 100 (1929).
Handel-Mazzetti (l.c. 100) identifies F. Porteri Lévl. & Vant. with F. hirta Vahl.
Ficus pandurata Hance in Ann. Sci. Nat. ser. 4, 18: 229 (1862). Ficus cuneata Léveillé & Vaniot in Mem. Acad. Ci. Art. Barcelona, ser. 3, 6: 149 (Ficus Sp. Chin. 11) (1907); in Fedde, Rep. Spec. Nov. 4:84 (1907). — Léveillé, Fl. Kouy-Tchéou, 429 (1915).— Rehder in Jour. Arnold Arb. 10: 128 (1929).— Pro parte, quoad
specim. Bodinier 2363. — Non Blume, nec Wallich, nec Miquel.
CHInA. Kweichou: environs de Kouy-yang, mont du Col- lege, gorge de Yan-pa, rocailles (May 16, 1898), environs de Tou-chan (May 31), E. Bodinier, no. 2363, “arbuste” (syntype of F. cuneata; photo. in A, A.).
On the label of Bodinier’s no. 2363, two localities are given, but there is only one specimen on the sheet; the name F. cuneata does not appear on the label.
Here belongs probably also F. comata Hand.-Mazz. as suggested by F. P. Metcalf in a note on one of the sheets of that species.
Ficus heteromorpha Hemsley. — Rehder in Jour. Arnold Arb. 10: 126 (1929). — Handel-Mazzetti, Symb. Sin. 7: 98, 100 (1929).
78 JOURNAL OF THE ARNOLD ARBORETUM [VoL. XVI
Ficus cuneata Léveillé & Vaniot in Mem. Acad. Ci. Art. Barcelona, ser. 3, 6: 149 (Ficus Sp. Chin. 11) (1907); in Fedde, Rep. Spec. Nov. 4:84 (1907). — Léveillé, Fl. Kouy-Tchéou, 429 (1915). — Rehder in Jour. Arnold Arb. 10: 128 (1929).— Pro parte, quoad specim. Cavalerie 1351. — Non Blume, nec Wallich, nec Miquel. — Synon. nov.
Cuina. Kweichou, add: Pin-fa, J. Cavalerie, no. 1351, Sept. 8, 1903, (syntype of F. cuneata; photo. in A. A.).
The other syntype of F. cuneata differing in the stalked receptacles seems referable to F. pandurata Hee.
Handel-Mazzetti (l.c. 100) refers F. pinfaensis to his F. comata, but the type of F. pinfaensis, Cavalerie 532, differs from F. comata in the sessile fruits, longer petioles and in the venation of the generally oblong and quite glabrous leaves rounded at base. To F. heteromorpha Handel- Mazzetti refers F. Stapfii Lévl., but Léveillé’s species, though apparently related to F. heteromorpha, differs in its leaves being rather densely hirsute on both sides and scarcely acuminate.
Ficus formosana Maximowicz. — Rehder in Jour. Arnold Arb. 10:
127 (1929). — Handel-Mazzetti, Symb. Sin. 7: 98, 100 (1929). Ficus ane Léveillé, FI. Kouy-Tchéou 433 (1915), p.p., quoad specim. Esquirol 3267. — Non Léveillé & Vaniot (1907).
Cuina. Kweichou: Teng-Tchéou, jardin du P. Marchand, Esquirol, no. 3267, June 22, 1902 (cited in Fl. Kouy-Tchéou; duplicate in A, A.).
The type of F. rhomboidalis belongs to F. gibbosa var. cuspidifera (Miq.) King.
Ficus lageniformis Lévl. & Vant. was referred by Handel-Mazzetti (l.c. 100) to F. formosana, as already identified by the writer (1.c.)
Ficus laceratifolia Léveillé & Vaniot.— Rehder in Jour. Arnold Arb. 10: 130 (1929).
This species has been referred tentatively by Handel-Mazzetti (Symb. Sin. 7: 100) to his F. comata as a form with lacerated leaves, but looks quite distinct. It is represented in this herbarium by a photograph of the type with fragments and by two specimens collected by W. P. Fang in Szechuan, no. 2415, from Mt. Omei and no. 5645 from Nanchuan Hsien.
Ficus Cavaleriei Léveillé & Vaniot.— Rehder in Jour. Arnold Arb. 10: 128 (1929).
This species seems related to F. pyriformis Hook. & Arn., but it is quite glabrous and the veins of the leaves diverge at nearly right angles;
1936] REHDER, LIGNEOUS PLANTS DESCRIBED BY LEVEILLE 79
it also resembles F. stenophylla Hemsley, but the leaves are broader and gradually narrowed from the middle to the ends.
Ficus pyriformis Hooker & Arnott, Bot. Beechey Voy. 216 (1841). — Léveillé, Cat. Pl. Yun-Nan, 275 (1917), “piriformis.” — Handel-Mazzetti, Symb. Sin. 7: 98, 100 (1929).
Ficus Nerium Léveillé & Vaniot in Fedde, Rep. Spec. Nov. 4: 66 (1907). — Léveillé, Fl. Kouy- oc 432 (1915).—Rehder in Jour. Arnold Arb. 10: 130 (1929
Ficus Kingiana Léveillé, FI. nee 431 (1915), pro parte; non Hemsley. — Synon. nov.
CuInA. Kweichou: without locality, J. Esquirol (holotype of F. Nerium,; fragments in A. A.); Tchen-fong, J. Esguirol, no. 914 (cited in Fl. Kouy-Tchéou; duplicate in A. A.); Ly-po, J. Cavalerie, Feb. 15, 1900 (syntype of F. Kingiana; merotype in A. A.)
Ficus Nerium was first referred to F. pyriformis by Handel-Mazzetti (l.c. 100). Ficus Kingiana appears in Fl. Kouy-Tchéou without de- scription, but as its characters can be partly ascertained from the key, it is not a nomen nudum; of the two syntypes, only the specimen from Ly-po belongs here and must be considered the type (lectotype) since it fits the characters given in the key, while the other specimen does not.
Ficus pyriformis Hook. & Arn. var. ischnopoda King in Ann. Bot. Gard. Calcutta, 1: 158, pl. 201c (1888).
Ficus macropodocarpa Léveillé & Vaniot in Fedde, Rep. Spec. Nov. 4:66 (1907). — Léveillé, Fl. Kouy-Tchéou, 431 ee — Rehder in Jour. Arnold Arb. 10: 130 (1929). — Synon
Cuina. Kweichou: without locality, a hae. no. 838 (holotype of F. macropodocarpa; merotype in A. A.); riviére de Lou- fou, J. Esquirol, no. 2216, Sept. 1910 (cited in Fl. Kouy-Tchéou; dupli- cate in A. A.).
Ficus macropodocarpa was first referred by Handel-Mazzetti to F. pyriformis (l.c. 100); on account of the long-peduncled receptacles, it seems referable to var. ischnopoda.
Ficus Esquirolii Léveillé & Vaniot in Mem. Acad. Ci. Art. Barce- lona, ser. 3, 6: 150 (Ficus Spec. Chin. 12) (1907); in Fedde, Rep. Spec. Nov. 4: 84 (1907). — Léveillé, Fl. Kouy-Tchéou, 430 (1915).
Ficus stenophylla Hemsley in Hooker’s Icon. 26: t. 2536 (1897), p-p., excl. parte typ. — Handel-Mazzetti, Symb. Sin. 7: 98 (1929).
Cut1na. Kweichou: préfecture de Hin-y-fou, J. Esquirol in hb. Bodinier, no. 2588, April 1899 (holotype of F. Esquirolii ; photo. in A. A.)
80 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XVII
This species seems to be closely related to F. pyriformis Hook. & Arn. and to F. stenophylla Hemsl.; from the former it differs in the narrower long-caudate leaves, subsessile subglobose receptacle covered with short appressed hairs, and from the latter by the thinner caudate leaves slightly hairy below and the subsessile subglobose and hairy receptacle.
Hemsley based his species mainly on Henry’s no. 8716 from Hainan which has to be considered the type of his species and is the plant figured on the plate except figure 7 representing a single receptacle which is from Faber’s no. 446 and apparently belongs to F. Esquirolii. The de- scription also applies mainly to the Hainan plant if under “Reptacula” the words “‘subsessilia vel . . . vel sphaeroidea . . . vel leviter puberula” are taken out.
Typical F. stenophylla is represented in this herbarium by plants from Hainan (W. T. Tsang no. 378, McClure, no. 9584), Kwangtung (C. L. Tso, no. 21048, Levine & Kwok, no. 875), Fukien (L. Y. Tai, no. 11673), Chekiang (R. C. Ching, no. 1692) and from Kweichou (Steward, Chiao & Cheo, no. 877); the last named specimen seems to approach F, Esquirolii in the shorter stalk of the receptacle. Ficus Esquirolii is represented besides by the photograph of the type also by Handel-Mazzetti, no. 10695, Y. Tsiang, no. 6053 and Steward, Chiao & Cheo, no. 147, all from Kweichou.
Under Ficus Esquiroliana Léveillé (in Bull. Geog. Bot. 24: 252) says of F. Esquirolii “extra genus probata est” but he still enumerates it the following year in his Flore du Kouy-Tchéou as F.. Esquirolii, while he changes F, Esquiroliana to F. laus-Esquirolit.
Ficus silhetensis Miquel in Ann. Mus. Bot. Lugd.-Bat. 3: 223, 291 (1867). — Léveillé, Cat. Pl. Yun-Nan, 275 (1917).
Ficus Cyanus Léveillé & Vaniot.—Rehder in Jour. Arnold Arb. 10: 129 (1929) — Synon. nov.
Ficus Cyanus var. viridescens Léveillé & Vaniot in Mem. Acad. Ci Art. Barcelona, ser. 3, 6: 149 (Ficus Spec. Chin. 11) (1907); in Fedde, Rep. Spec. Nov. 4: 84 (1907).
sh! i la Léveillé & Vaniot in Léveillé, Fl. Kouy-Tchéou, 429
15).— ae in Jour. Arnold Arb. 10:128 (1929).— Seay no
Cu1na. Kweichou:_ Si-liéou-gay, trous profonds, J. Cava- lerie, no. 169, July 3, 1902 (holotype of F. Cyanus;; merotype in A. A.) ; arrivée a Tong-kai (ruisseau), J. Esquirol, no. 3031, July 25, 1911 (cited in Fl. Kouy-Tchéou under F. Cyanus; photo, and duplicate in A. A.); Pin-fa, J. Cavalerie, no. 1099, June 23, 1903 (holotype of F. Cyanus var. viridescens; photo. in A. A.); Tché-chou, bois, J. Esquirol, no, 2506,
Sept. 1909 (holotype of F. congesta; merotype in A. A.). Ficus Cyanus differs slightly from F. silhetensis as figured and de-
1936] REHDER, LIGNEOUS PLANTS DESCRIBED BY LEVEILLE 81
scribed by King (in Ann. Bot. Gard. Calcutta, 1: 154, pl. 194, 1888) in the shorter petioles and cuneate base of the leaf, but otherwise agrees. The variety viridescens does not seem to differ from typical F. Cyanus, and in Flore du Kouy-Tchéou, Léveillé enumerates it under the species. Ficus congesta resembles closely Cavalerie’s no. 1099 (F. Cyanus var. viridescens) except that the leaves are somewhat smaller.
The species has also been collected in Kweichou by Y. Tsiang, no. 4404, determined as F. silhetensis Miq. by E. D. Merrill.
Ficus Stapfii Léveillé.— Rehder in Jour. Arnold Arb. 10: 131 (1929).
Ficus Stapfi is represented only by a sterile specimen (photo. and fragments in A. A.). It was referred by Handel-Mazzetti (Symb. Sin. 7: 100) tentatively to F. heteromorpha Hemsl., but differs in its rather dense pubescence; in this respect it resembles F. silhetensis Miq., but the leaves are oblong and obtusish to acutish, not acuminate at the apex.
Ficus Abelii Miquel in Ann. Mus. Bot. Lugd.-Bat. 3: 281 (1867). Ficus Schinzii Léveillé & Vaniot in Fedde, Rep. Spec. Nov. 8: 550 (1910). — Léveillé, Fl. Kouy-Tchéou, 433 (1915).—Rehder in Jour. Arnold Arb, 10: 131 (1929).— Non F. Schinziana Warb. — Synon. nov.
Cuina. Kweichou: Lo-fu, J. Cavalerie, no. 3592, Apr. 1909 (holotype of F. Schinzii; merotype in A. A.); Tsao-ly, bord de riviére, alt. 600 m., J. Esquirol, no. 3503, Feb. 5, 1912 (cited in Fl. Kouy- Tchéou; duplicate in A. A.).
By Handel-Mazzetti (l.c. 100) F. Schinzii was referred to F. pyri- formis and its allies. It certainly belongs to F. Abelii which has not yet been recorded from Kweichou, though it has been collected in Szechuan and Hunan.
Ficus Roxburghii Wallich. Num. List. 4508 (1832).— Handel- Mazzetti, Symb. Sin. 7: 100 (1929).
Ficus macrocarpa Léveillé & Vaniot in Mem. Acad. Ci. Art. Barcelona, ser. 3, 6: 152 (Ficus Spec. Chin. 14) (1907); in Fedde, Rep. Spec. Nov. 4:85 (1907). — Léveillé, Fl. Kouy-Tchéou, 431 (1915), sub F. macrocarpa Wight.—Rehder in Jour. Arnold Arb. 10: 130 (1929).— Non Wight.
Cuina. Kweichou: without precise locality, J. Cavalerie in 1899 (holotype of F. macrocarpa; ex Léveillé); ouest de Lo-fou, J. Cavalerie, no. 3593, March 1909, “bel arbre” (cited in Fl. Kouy-Tchéou; photo. in A. A.).
Handel-Mazzetti refers to F. Roxburghii besides F. macrocarpa also F, Letaqui Lévl. & Vant. which see below.
82 JOURNAL OF THE ARNOLD ARBORETUM [VoL. Xv
Ficus Roxburghii Wall. must be considered a valid publication of a new name, since he cites as a synonym “F. macrophylla Hb. Roxb. (haud Desf.)” a species described in Roxburgh, Fl. Ind. 3: 556 (1832).
Ficus Letaqui Léveillé & Vaniot in Fedde, Rep. Spec. Nov. 8: 550 (1910). — Léveillé, Fl. Kouy-Tchéou, 531 (1915).— Rehder in Jour. Arnold Arb. 10: 130 (1929).
This species was referred by Handel-Mazzetti (Symb. Sin. 7: 100) to F. Roxburghii, but it differs markedly from F. Roxburghii in the much smaller subglobose and subsessile receptacles, borne mostly in pairs on the nodes of long and slender leafless shoots; the leaves are similar in shape, but scabrid above and borne on longer and slenderer petioles.
Ficus Vanioti Léveillé, Fl. Kouy-Tchéou, 434 (1915), pro parte.
Cutna. Kweichou: bois sacré au Che-chen de Na-é, J. Esquirol, nos. 3077 and 3078, end of June and July 1 (syntype of F. Vanioti; photo. in A. A.).
The name F. Vanioti was published in Flore du Kouy-Tchéou with F. ouangliensis as a synonym and was intended apparently as a new name for that species which is based on Cavalerie no, 2568, and is not a Ficus, but an Aglaia. The two specimens cited above, however, repre- sent a species of Ficus, but the material is so fragmentary, consisting only of bare branches, broken leaves and a few broken receptacles, that exact identification is not possible; the material seems to belong in the affinity of F. formosana Maxim.
LORANTHACEAE
Tolypanthus Maclurei (Merr.) Danser in Bull. Jard. Bot. Buiten- zorg, ser. 3, 10: 355 (1929). nme Maclurei Merrill in Philipp. Jour. Sci. 21: 494 (1922). Loranthus Esquirolii Léveillé, Rev. Ann. Chine, 1916: 21 (MS).— Nomen non rite sabliontus,
Cuina. Kweichou: Pin-fa, Heou-tcheng, J. Esquirol, no. 52, “liane parasite sur le bibassier” (type of Loranthus Esquirolii; photo. in A. A.).
Léveillé’s name Loranthus Esquirolii appeared in a manuscript publi- cation, of which only a few copies were distributed and cannot be con- sdeced validly published; Merrill’s specific epithet therefore, remains vali The name Loranthus Esquirolii does not appear on the original label a the type specimen which was identified by Danser as Tolypan- thus Maclurei.
(To be continued)
HERBARIUM, ARNOLD ARBORETUM,
HARVARD UNIVERSITY.
1936] FLORY, CHROMOSOMES IN GYMNOSPERMS 83
CHROMOSOME NUMBERS AND PHYLOGENY IN THE GYMNOSPERMS
WALTER S. FLory With two text figures
CRITICAL sTuDIES of chromosome morphology in most groups of living gymnosperms have been published during the past few years. Geitler (1929) and Florin (1932) have described the chromosomes of Wel- wttschia and Ephedra of the order Gnetales. Sax and Beal (1934) inves- tigated the chromosome situation in all nine genera of the Cycadales. The most comprehensive chromosome survey in the gymnosperms is that of the Coniferales by Sax and Sax (1933). Fifty-three species from six- teen genera of the subfamilies Cupressineae, Taxodineae, Abietineae, and Taxineae were studied. The sole surviving member of the Gink- goales was included in the last mentioned analysis and has been the object of cytological studies by other workers as well. This leaves only the sub- families Araucarineae and Podocarpineae and the genus Gnetum with representatives unanalyzed in recent cytological papers. With regard to two of these groups Sax and Beal (1934) write, ‘“ .. . recent studies ae that Podocarpus has 20 pairs of chromosomes’; and, also . Gnetum has a large number of relatively small eheoiiosomnien in erie: accounts Burlingame, Ishikawa and Schurhoff (from Sax and Sax, 1933, p. 368) have found 12 chromosomes in several species of Podocarpus.
Only two genera are included in the subfamily Araucarineae. They are Araucaria and Agathis. An early morphological study (Lopriore, from Coulter and Chamberlain) on one species of Araucaria indicated the presence of 12 pairs of chromosomes here. No references have been found to critical chromosome work on either genus of this subfamily. Questions concerning the phylogenetic sequence of conifer subfamilies involve the Araucarineae at important points. One question concerns the comparative antiquity of the Araucarineae and Abietineae; another the relationship between the Araucarineae and the Podocarpineae. This investigation was made to see if a knowledge of chromosome number and morphology in the Araucarineae would aid in solving problems of the nature mentioned. To further that end, the chromosomes of several species of Podocarpus have been examined, and the results obtained will be briefly discussed.
84 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XVII
Material for this work has been secured from several sources. The greenhouses of the Bussey Institution, the Arnold Arboretum, and the Biological Institute of Harvard University, have been sources of some species of Agathis and Araucaria. Root tips of two species of Podocarpus and of two species of Araucaria were obtained at Wellesley College. Other material of Araucaria and Podocarpus was collected at the Brook- lyn Botanic Garden and at the New York Botanical Garden. We realize that the species names of such specimens may not be correct, but it was not possible to verify the classification of these immature plants.
Absolutely accurate chromosome counts of conifer species are difficult to secure in root tip cells, — endosperm material being much preferred for preparations of somatic chromosomes. However, root tips were the only material available for the present study. The chromosomes of all species examined were studied in aceto-carmine smears of root tip cells. Some recent advances in technique allow somewhat more critical prepa- rations to be made than were formerly possible. A modification of Warmke’s (1935) method of making temporary root tip smears was used. Root tips were fixed in aceto-alcohol overnight; treated with a hydrochloric acid and 95% alcohol mixture; hardened in aceto-alcohol ; then sliced and macerated. After the preparations had been allowed to stand for from 5 to 15 minutes, they were heated almost to boiling; then considerable pressure was exerted upon the cover glass, and this was fol- lowed by a second heating, after which the mounts were sealed. By this method metaphase plates were secured, in several species, in which the chromosomes were well scattered, enabling them to be readily counted.
Rehder’s (1927) classification has been followed with regard to divi- sion of the Coniferales, and elsewhere when possible. This has been sup- plemented by use of Pilger’s (1926) classification for the forms not gen- erally grown in the temperate United States.
DESCRIPTION
Agathis robusta, Araucaria brasiliana, and A, Cunninghamii each con- tains 13 pairs of chromosomes. Material from a plant of Agathis bearing a doubtful specific name also showed 26 somatic chromosomes. Numer- ous plates from Araucaria Bidwilli were also studied. The chromosome picture appeared to be identical with that found in the other two species of Araucaria studied. Counts of a number of metaphase figures seen in side view indicated the presence of 26 somatic chromosomes in this spe- cies. The chromosomes of species belonging to the same genus were very similar. The chromosomes of Agathis robusta (Fig. 1) and of Araucaria brasiliana (Fig. 2) were studied in more detail than those of the other
1936] FLORY, CHROMOSOMES IN GYMNOSPERMS 85
species. In each of these species there were 9 pairs of chromosomes in which the arms were about equal in length, or in which the longest arm was, at most, not twice the length of the shortest. In addition to these chromosomes with median or submedian attachments, both species apparently have 4 pairs with nearly terminal attachment points. The chromosomes of Agathis and Araucaria are similar not only in morphol- ogy but also in size. Chromosomes of Agathis, in aceto-carmine, vary in length from 13 u to 22 uw; those of Araucaria from 12.5 uto 20 u. In Araucaria the elements appear to be slightly greater in diameter than in Agathis, but the difference is too small to be significant.
TEXT FIGURE 1 Drawings of chromosomes of root tip cells. x 900. Fig. 1. Agathis robusta, 26 chromosomes. Fig. 2. Araucaria brasiliana, 26 chromosomes. Fig. 3. Podocarpus falcatus, 24 chromosomes. Fig. 4. Podocarpus macro- phyllus, 38 chromosomes. Fig. 5. Podocarpus neriifolius, 36 chromosomes.
The chromosomes of five forms of Podocarpus were examined. These were P. macrophyllus, P. falcatus, P. neriifolius, P. chinensis, and P. andinus. Pilger indicates that P. chinensis is merely a form of P. macro- phyllus, which is cultivated in botanic gardens, and is not a separate species. The first four forms listed belong in the generic section Eupodo- carpus. Podocarpus andinus is placed in Pilger’s section Stachycarpus.
In these five forms there are two different types of genoms. The first type of genom was found in a species growing in the greenhouse at Wel- lesley College and tentatively identified by Dr. Alice Ottley as P. fal-
86 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XVII
catus R. Br. Several plates quite clearly showed 24 chromosomes in root tip cells of this plant (Fig. 3). These plates, quite apparently, were almost complete. Each, however, had its elements quite scattered, owing to pressure; and it is possible that a few chromosomes were lost. Of the 24 chromosomes observed, 2 pairs were approximately isobrachial; the other 10 pairs were distinctly heterobrachial. Chromosome measure- ments were, for the longest, 15 ; for the shortest, 5.5 u; and the average length was approximately 11 u. In morphology the complement is similar to that found in many conifers. However, the chromosomes are smaller than those found in most species.
In P. macrophyllus many plates were found which showed all but a few chromosomes distinct and unentangled (Fig. 4), but no complete plate was beyond question at some point. The number is about 38, many plates seeming to have this count. In several cases there was a suggestion of 39 chromosomes. No plate seemed to have 40 but per- haps, for safety, the number should be placed as between 38 and 40. The best figures, those appearing to contain 38 elements, were analyzed. Ten pairs are heterobrachial, 3 of these approaching the isobrachial con- dition. Nine pairs appear to have attachment points that are terminal, or very nearly so. Chromosome measurements give, for the longest pair, 12.5 uw; for the shortest, 3.5 4; and the average length was about 7.5 uw.
The P. nertifolius complement has much the same appearance as that of the species just described. Approximately 38 chromosomes are pres- ent. However, the majority of these appear terminal or nearly so, the only apparent exceptions being one short isobrachial pair, and the longest pair, which have submedian attachment constrictions. The longest chromosomes are 12.5 ut, the shortest 3 u. Figure 5 shows 36 well-spaced chromosomes in a plate from which a few have been lost.
Podocar pus chinensis and P. andinus (belonging to a different section from the other podocarps mentioned here) were seen to have the same general type of complement as P. macrophyllus, having nearly 40 chro- mosomes, many with terminal or near terminal attachment points.
CHROMOSOME NUMBERS AND PHYLOGENY
Numerous morphological studies have been made on gymnosperms. These, with discussions of suggested evolutionary tendencies, are best summarized by Chamberlain (1935 a and b). A diagram (Text fig. 2) representing the phylogeny of gymnosperm orders and families, as rep- resented by the best available evidence, has been made. In this diagram ideas and material from. a number of sources (chiefly: Chamberlain, 1935b, diagram p. 184; Jeffrey, 1917, especially pp. 355-6; Sax and Sax,
1936 | FLORY, CHROMOSOMES IN GYMNOSPERMS 87
1. c.; Sax and Beal, 1.c.; Florin, 1. c.; Matsuura and Suto, 1935; and the present work) are incorporated in one figure. Since representatives of all major gymnosperm groups have now been subjects of critical cyto- logical examination, it seemed appropriate that some effort be made to see how the cytological facts fit in with those of grosser morphology.
A description of Text figure 2 follows. Beneath each major division are placed its genera from which representatives have been cytologically
Zami 8 i) Pinus 12 Cryptowmerial iy St 8 = 5 : : her ia a; = noi" Cedrus IL Taxodium [ut eratozama 6 3 Encephalartos i Un 8 3 "3 7F aes : ; ‘. phals g 4 2 v o|tl Picea " f]3 Boweni 5 0 ¥ 12 = Ts uga lo “ Dioon q wW F = Foo Abie 5 : ™Macreza icf 0 2 ; ; +2 a mua Oo w Oe Poevdotsu 0) A Cycas st S c z 7 al Yala Y ca £|'3 £ evdolarix]22 sla 9} 39 Microcycas 13 a ALC c ee ras "a c : | = v © ) q y 3 = E rie : £ a 2 v 3 ad “SE
‘
Cycadales 8B Coni fev ates
leo
TEXT FIGURE 2 Phylogenetic relationships of the living gymnosperms based on morpho- logical characters, and the basic chromosome numbers of the genera in the various families.
analyzed. The chromosome count is given in basic haploid numbers. Extinct orders are indicated by four small perpendicular lines, placed on the main evolutionary lines at about the geological level at which ex- tinction occurred. The most doubtful points in the suggested relation- ships are shown by dotted lines or by question marks.
There are apparently three basic groups of chromosome numbers in
88 JOURNAL OF THE ARNOLD ARBORETUM [voL. xvi
the living gymnosperms. The basic numbers are predominantly 8 or 9 for the Cycadales, 7 for the Gnetales, and 12 for the Ginkgoales and Coniferales. The deviations from these basic numbers in these groups can be attributed to elimination of one or several chromosomes following segmental interchange or polyploidy, and an increase in chromosome number resulting from fragmentation or chromosome duplication. Polyploidy is rare in the gymnosperms. The chromosomes of species of gymnosperms are much more uniform in number, size, and general morphology than those in the species of angiosperms.
The count of 38 somatic chromosomes in certain species of Podocarpus at first suggested a basic number of 19. This is the basic number in certain primitive angiosperms, and Anderson (1934) has suggested an allopolyploid origin of such angiosperms from primitive gymnosperms with basic chromosome numbers of 7 and 12. The presence of about 12 pairs of chromosomes in one species of Podocarpus suggests, however, that the forms with about 38 somatic chromosomes may be triploids with 13 chromosomes as the basic number, or that chromosome fragmenta- tion at the fiber attachment point has effected the change in these genoms.
The cytological analysis of the Coniferales seems to show some corre- lation with the morphological evidence regarding family relationships. The Cupressineae and Taxodineae have the same basic chromosome numbers. The basic chromosome number is 13 in the Araucarineae and is 12 or 13 for the Podocarpineae. The presence of 12 chromosomes in most genera of the Abietineae supports the theory that this may be the main line from which the other families have been derived. The basic number of 12 chromosomes in the Ginkgoales also supports the sugges- tion that 12 is the basic number from which both families, Taxaceae and Pinaceae, originated.
SUMMARY
The basic chromosome number is 13 for Agathis and Araucaria, and the genoms are similar in morphology. Chromosome numbers were obtained in five species of Podocarpus. In one species about 24 somatic chromosomes were counted, but in the other species there were about 38 chromosomes in the root tip cells. The basic chromosome number is 12 or 13, and the four species with about 38 chromosomes presumably are the result of polyploidy or chromosome fragmentation.
A survey of chromosome numbers in the gymnosperms indicates three basic numbers, — 8 or 9 for the Cycadales, 7 for the Gnetales, and 12 for the Ginkgoales and Coniferales. There is some correlation between
1936] FLORY, CHROMOSOMES IN GYMNOSPERMS 89
chromosome numbers and morphological relationships of the various taxonomic units.
This work was done at the Bussey Institution of Harvard University while the author was a National Research Council Fellow.
LITERATURE CITED ANDERSON, EpcGar (1934). Origin of the Angiosperms. (Nature, 133: 462.
CHAMBERLAIN, C. J. (1935). Gymnosperms, structure and evolution. Univ. of Chicago Press
a 5). The Gymnosperms. (Bot. Rev. 1: 183-209.)
Coutter, J. M. and C. J. CHAMBERLAIN (1910). Morphology of Gymno- sperms. one of Chicago Press.
FLorIn, R. (1932). Die ee ean ae bei Welwitschia und einigen Ephedra-Arten. (Svensk Bot. Tidskr. 26: 205-214.
GEITLER, L. (1929). Zur Zytologie von Ephedra. (Oester. Bot. Zeitschr. 8: 242 ff.
) JEFFREY, E. C. Ces The anatomy of woody plants. University of Chic
cago
Matsuura, H. and T. Suto or pas eget to the idiogram aired in Bh oe Heke plants. I. (Jour. Fac . Hokkaido Imp. Uni 5: 33-75.
Pricer, R. (1926). Gymmospermae (in Engler and Prantl’s “Die nattr-
en Pflanzenfamilien,” 2d ed.
REHDER, ALFRED (1927). Manual of cultivated trees and shrubs. The Macmillan Co., New York.
Sax, Kari and J. M. BEAL (1934). Chromosomes of the Cycadales. (Jour. Arnold Arb. 15: 255-258.
Sax, Kart and Hatty J. Sax (193 3). Chromosome number and morphol- ogy in fe conifers. (Jour. Arnold Arb. 14: 356-375.)
CYTOLOGICAL LABORATORY, ARNOLD ARBORETUM, HARVARD UNIVERSITY.
90 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XVII
APOSPORIC PARTHENOGENESIS IN A TRIPLOID APPLE, MALUS HUPEHENSIS
Haic DERMEN With plates 179-181
A NUMBER of years ago Dr. Karl Sax, in his attempt to make inter- generic crosses in Pomoideae, found that whenever Malus hupehensts was used as the female parent, seeds were always formed. Preliminary experiments were made to discover the basis of this peculiarity (Sax, 1931) and it was found that this species behaved as a typical partheno- genetic plant. I was designated to investigate the problem and the present report is a cytological analysis of this phenomenon.
Malus hupehensis (Pamp.) Rehd. (M. theifera Rehd.) (Rehder, 1933) is known to be a pure species, distributed widely in mountain regions of China extending south into Assam. It was introduced into this country in 1900. All plants of this species growing in the Arnold Arboretum, some of which have been propagated from seeds here and some brought in from outside sources, fruit heavily; the fruits con- taining the usual number of seeds. All the seeds appear to be deformed in outline in comparison with those of other species.
MATERIAL AND METHODS
In order to have a sounder basis for our conclusions concerning par- thenogenetic behavior in M. hupehcnsis it was thought advisable to make parallel observations on another species, the embryonic development of which was normal. Malus arnoldiana (Rehd.) Sarg. (M. baccata X floribunda), a hybrid, chosen as a control plant, produces an abundance of fruit, and normal seeds in contrast to M. hupehensis.
In the summer of 1931 a preliminary examination was made of the chromosome number from root-tips of a M. hupehensis seedling and it was found to be a triploid form. The analysis of the ovules was made beginning the first week of May and continuing through the summer during the years 1932, 1933 and 1934. The study of this problem dur- ing the first two years was exploratory in character and it was in 1934 that a more complete analysis of the ovules of both M. arnoldiana and M. hupehensis was made.
Specimens at different stages of ovular development were fixed sepa- rately. The fixing of material was begun during the first week of May
1936 ] DERMEN, PARTHENOGENESIS IN MALUS 91
when the buds began to turn pink and the development of the female meiotic phase was initiated. This phase is completed just before shed- ding of petals when the egg apparatus in the embryo sac (ES) is formed and fertilization has taken place (normally). From the pink stage to the time of shedding of petals six separate collections were made. Two collections were made between shedding of petals and remaining parts of perianth. After that the collections were continued at 5 to 7 day intervals in order to follow up the sequence of embryo development.
In fixing material for early stages, the fleshy part of the ovary is trimmed off with a sharp knife in order that the fixative may penetrate the ovules quickly. For later stages young seeds are removed from the fruit and are fixed directly. For quick penetration of fixative in older seeds the seed coat may be slightly slashed. For more mature seeds the whole testa may be removed.
The first year Flemming’s solution was used as fixative but was found unsuitable for this material. Navashin’s solution was found preferable for the meiotic and earlier stages of embryo sac formation. However, being doubtful of the effect of the latter fixative on later stages, Lewitz- ky’s solution (5% formalin and 0.5% chromic acid) was used for those, since I had found earlier (Dermen, 1933) that it facilitates at least the staining of nucleoli with either crystal violet-iodine or haematoxylin. In this way I hoped to have at least the nucleoli to go by as landmarks, so to speak, in determining the nuclear number and other features in the development of embryo sacs. The results eventually showed this assump- tion to be justified. To prevent severe distortion of embryo sacs in changing fixed material into alcohol for paraffin sectioning, the material was allowed to remain in the fixative for twelve hours or more, washed in water and run up in alcohol gradually beginning with 5%. The results later showed that there was some shrinkage of the embryo sac and endo- sperm layer away from the nucellar tissue, though not enough to make observations confused.
Paraffin sections of early stages to the beginning of ES formation were cut 20 u thick and from ES formation on up 35-40 u, thick enough to obtain whole embryo sacs and young embryos, in order that confusion might be avoided in studying the origin and sequence of development.
Meiotic stages were stained with crystal violet-iodine, and ES and later stages with haematoxylin. Root-tips were fixed in Lewitzky’s solution and stained with crystal violet-iodine; while for meiosis in the anther aceto-carmine technique was used.
Due to technical difficulties in illustrating various phases, it was necessary to use different scales. The scale used is indicated under each drawing.
92 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XVII
DESCRIPTION
The features illustrated which were considered important in the analysis of parthenogenetic behavior in M. hupehensis are presented in 38 figures in three plates. The drawings are made in diagrammatic form to bring out the points pertinent to this study and to give the most essential details of origin and development of both embryo sac and embryo. The comparative illustrations of the control plant, M. arnold- iana, and M. hupehensis run from Figs. 18 and 19 to Figs. 34 and 35 in- clusive. Other figures are from M. hupehensis alone.
The somatic chromosome number in M. hupehensis, studied from root-tips, was 3n = 51 (Fig. 1), a triploid number, the diploid being 2n — 34. The next figure (Fig. 2) represents the chromosome set from a tetraploid seedling of M. hupehensis with 4n = 68, the discussion of which will be found elsewhere in this paper. The size of these chromo- somes is about one micron in length.
The meiotic development in anthers took place about 10 days earlier than the corresponding phase in the ovules. In the anthers the homolo- gous chromosomes form trivalents and bivalents and a few remain uni- valents (Sax 1932). Figure 3 is II M from anthers with 25+ chromo- somes at each plate. No mature pollen grains are found in the anthers. Some cells begin to degenerate at I M and others at later stages, and most degenerate after they have reached the tetrad stage when the individual microspores fail to break apart although they are fully differ- entiated and each has its own wall.
A similar process of degeneration was found in the ovules. Here the chromosomes either remain univalents at I M or pair variously and divide at random, Figure 4 (a and b) represents I M in an egg mother cell (EMC). The chromosomes here are in univalent form. In the two sections, a and b, the total is exacly 51. It cannot be stated with cer- tainty when, if ever, in univalent form there was any division in the EMC. However, in a few instances observed where the chromosomes were unpaired the cell was in a stage of degeneration. Such a cell is shown in Fig. 11. An anaphase stage is given in Fig. 5 (a and b). The b group of chromosomes from the lower pole was in a following serial section. Here also the total number of chromosomes could be accurately counted.
Altogether in M. hupehensis over two hundred ovules in the early stages of meiosis were examined and only rarely were there found ovules lacking a meiotic cell, either in prophase, in division stages, or in a de- generative process. There were found some older ovules which lacked embryo sacs. One young ovule was found with two egg mother cells.
1936] DERMEN, PARTHENOGENESIS IN MALUS 93
There were observed many ovules divided into two nucellar regions, a twin ovule, each with a simple or a compound ES, enclosed together within a common integument.
An EMC is shown in Fig. 6 at pachytene stage. Usually around a meiotic cell there are one or more somatic cells each with a nucleolus as large or larger than that in the meiotic nucleus, which, as will be pointed out later, may develop into embryo sacs. Very rarely there were found ovules with two egg mother cells. Such a case is shown in Fig. 7. Around these meiotic cells are found a number of large nucleolated somatic cells. Figure 9 is a small scale drawing of the same cell to indicate the distance of a meiotic cell from the upper end of the nucellus. Here the EMC is 6 cell layers deep, while in other ovules the number of layers may vary. In Lilium regale (original observation) this layer was always one layer deep from an early stage to the time of fertilization (later stages were not observed). Figure 10 illustrates a typical young ovule with two layers of integumental tissue as it appears in longitudinal section. The cell in it is the same as in Figs. 8and 9. The tissue around this cell is the nucellus. This figure shows the approximate location of the EMC in the ovule taken as a whole.
As was mentioned above the EMC may either degenerate at I M or later as in the anther. A degenerating cell at I M is shown in Fig. 11. Here the cell was found compressed from the sides and the chromosome stain was blurry showing these bodies more or less clumped together. As a whole its degenerated feature could be easily distinguished when compared with other cells at the same stage. As well as could be made out the chromosomes in this cell were not paired, which fact, I believe, may be correlated with degeneration of the cells at this stage.
Figures 12 and 13 show a parallelism in division with EMC and in development of one of the resulting cells into, supposedly, an ES. Nor- mally an EMC should divide twice and from this division four cells should result. One of these, which is the furthest from the micropyle should be the megaspore destined to become an ES. However, as will be shown below, this expectation is not fulfilled in M. hupehensis. In Fig. 12 we find three cells. The one nearest to the micropyle has degen- erated. This cell corresponds to one in Fig. 13 which was in metaphase and in a stage of degeneration. In this latter figure the second division was found to be irregular; some of the chromosomes at opposite poles were at a distance from the main chromosome groups. If this cell divided it would result in two unequal sized cells, because these two groups of chromosomes are not found at equal distance from the ends of the cell. This is what is found in Fig. 12. Here one cell is degenerating.
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One of the sister cells from the second division is small, the other large, and in this large cell there is a small chromosome mass outside the main nucleus, indicating that this cell is not an ordinary somatic cell but the product of an irregular second division of an EMC.
Direct as well as indirect evidence shows that meiotic cells at some stage of their development degenerate while somatic cells in their imme- diate vicinity usually grow in size and develop into embryo sacs. Figure 14 illustrates this point clearly. Here is shown a rather large somatic cell with a large nucleolus, above which there is a meiotic cell in a stage of degeneration. In Fig. 15 is shown such a cell that has grown in size. The nucleus has divided and the cell is in a two-nucleate stage and beginning to develop into an ES. At the base of this are shown four cells in a row that have annular xylem thickenings, an unexpected mani- festation in an ovule, this type of cell being characteristic of vegetative tissues. Similar cells were found at later stages of ES development and in one case in a complex ES. Serial sections from either side of this were scrutinized and no trace of degenerating meiotic cells was found in the neighborhood of this binucleate ES. Therefore, it may be that these four annulated cells represent the tetrad resulting from two meiotic divi- sions. Their degeneration was expressed by transformation into xylem elements. Such cases were found only in 1932 material.
In the majority of cases the embryo sacs were structurally complex. These were found to have resulted from the fusion of two or more embryo sacs. A simple case is shown in Fig. 16 where two embryo sacs were found growing simultaneously. In other cases the number of these young embryo sacs growing side by side may reach six or more. In this figure one is four-nucleate and the other below it two-nucleate. In Fig. 23 two adjacent sacs are shown in mature egg apparatus stages. Figure 17 represents the middle section of five serial sections in which a total of 30 nuclei could be counted which may have resulted from fusion of four embryo sacs, boundaries of which could not be made out clearly.
Illustrations from Fig. 18 to Fig. 35 inclusive are diagrammatic rep- resentation of comparative development of embryo sacs and embryos in M. arnoldiana and M. hupehensis. Figures 18 and 19, the former from M. arnoldiana and the latter from M. Aupehensis, are from material fixed at the time of full bloom. Both represent the stage when the embryo sacs have completed the cycle of the development with three antipodal nuclei cut off, each having formed a separate small cell. The main portion of the ES now functions as a distinct body in which there is an egg cell with two synergid cells together forming the egg apparatus, below which the two polar nuclei have approached each other but com-
1936 ] DERMEN, PARTHENOGENESIS IN MALUS 95
plete fusion is delayed, perhaps till joined with a sperm. A similar situation was observed in Lilium regale (original observation) where first the upper polar nucleus is joined with a sperm and these two jointly meet half way with the lower polar nucleus. By the time of fertilization the egg cell is larger than the synergids (Fig. 20, M. arnoldiana). In these drawings the upper limits of the egg and synergid cells are not shown because they were vague in my preparations. In Fig. 19 (M. hupehensis) a dotted nucleolus of one of the polar nuclei and a dotted synergid cell are superimposed on the main portion from an adjacent section in a single drawing. The outer broken line represents the inner limit of the nucellus. A more mature ES is shown in Fig. 20 of M. arnoldiana from material fixed at the time of calyx shedding (the last remnant of the perianth with dried up styles). Here only nucleoli are shown. At this stage the antipodal cells had disappeared and the sac had grown longer but not much wider. At this time of the ES develop- ment fertilization has usually taken place, but perhaps due to climatic conditions some cells remain unfertilized. In the ES shown in Fig. 21 (M. arnoldiana) there is a clear indication that fertilization has taken place, judging by the amount of nucleolus in the egg nucleus in compari- son with the same in the synergids. In the other figures (Figs. 18-20, taking these individually) the size of the nucleoli in the egg and the synergids is remarkably similar. Fertilization in Fig. 21 is further indicated by the fact that there appears the beginning of endosperm nuclear division, due to fusion of the polar nuclei with a male gamete. The embryo sacs shown in Figs. 22, 23 and 24, all of M. hupehensis, are from material fixed at the