LIBRARY 
 
 OF THE 
 
 UNIVERSITY OF CALIFORNIA. 
 
 GIFT OF 
 
 Class 3 
 
TERATOLOGY IN THE FLOWERS OF TWO 
 CALIFORNIAN WILLOWS 
 
 BY 
 
 WILLIAM WARNER MOTT 
 
 THESIS PRESENTED TO THE FACULTY OF THE COLLEGE OF NATURAL 
 SCIENCES IN PARTIAL FULFILLMENT OF THE REQUIRE- 
 MENTS FOR THE DEGREE OF MASTER OF SCIENCE. 
 
UNIVERSITY OF CALIFORNIA PUBLICATIONS 
 BOTANY 
 
 Vol. 2, No. 7, pp. 181-226, Pis. 19-20 December 16, 1905 
 
 TERATOLOGY IN THE FLOWERS OF TWO 
 CALIFORNIAN WILLOWS 
 
 BY 
 
 WILLIAM WARNER MOTT 
 
 CONTENTS. 
 
 PAGE 
 
 I. Introduction 181 
 
 II. A Teratological, mainly Staminate, Salix lasiolepis Benth 184 
 
 III. A Teratological, mainly Pistillate, Salix lasiandra Benth. X 
 
 ~babylonica L. and some related individuals 196 
 
 IV. Bearing of the Abnormalities described on some Moot Points 
 
 regarding the Salix Flower 203 
 
 V. Some Possible Causes of the Abnormalities described 200 
 
 VI. Summary 213 
 
 VII. Bibliography 216 
 
 VIII. Explanation of Plates 219 
 
 Introduction. 
 
 The normal flowers of the numerous species of willows are 
 unisexual, and the staminate and the pistillate flowers are born 
 on separate individuals (species dioecious). There is no peri- 
 anth, but the flowers are born in catkins, and the bracts of a 
 catkin closely subtend the sex-organs composing the flowers and 
 protect these organs in the young stage. The number of 
 stamens in a flower varies in different species, but two, three, 
 or five are common. The pistil is unicellular and bicarpellary 
 with two parietal placentae, and the fruit is a capsule. A small 
 honey-gland is borne at the base of the pistil and of the stamens, 
 on the side next to the axis of the catkin, and in some species a 
 second basal gland is borne on the bract opposite to the first. 
 
 172945 
 
182 University of California Publications. [BOTANY 
 
 Cases of monoecism (male and female flowers on one indi- 
 vidual), and of abnormal bisexual flowers with perfectly de- 
 veloped pistil and stamens, and of flowers with monstrous sex- 
 organs combining the characters of pistil and stamens in vari- 
 ous proportions have been frequently recorded in this genus 
 (Salix) and several writers have speculated at some length as 
 to the cause, and as to the significance, morphological, anatomical, 
 and physiological, of these abnormalities. A few of the articles 
 published have been accompanied by illustrations, notably a 
 paper by Camus (1899) and two by von Seemen (1886, 1895), 
 but none of the drawings show in detail a series of the monstrous 
 organs connecting stamens and pistil. The object of this paper 
 is mainly to present drawings of such a series with particular 
 note of certain facts well shown by the material at hand, some of 
 which have not been found described elsewhere. The material 
 was discovered by the writer on a predominantly male individual 
 of Salix lasiolepis Benth. (near Berkeley, in the spring of 
 1903). The male tree of this species has normally two stamens 
 to a flower, and this abnormal specimen showed* the two stamens 
 in various stages of transformation (so it appeared) into pistil. 
 The converse case of apparent transformation, namely, pistil 
 more or less changed into stamens, has also been found by the 
 writer on a predominantly pistillate willow-tree,f which is be- 
 lieved to be a hybrid of Salix lasiandra Benth. and Salix baby- 
 lonica L. ; and on this tree occurred also bisexual flowers with 
 perfectly developed pistil and stamens (the number of stamens 
 being commonly two), and also two extra, lateral floral glands. 
 Some notes of these abnormalities are made. 
 
 This pistillody of the stamens and staminody of the pistil, 
 and the occurrence of bisexual flowers with normally developed 
 sex-organs have not, it is believed, been hitherto reported in 
 Salix for this Coast, nor, therefore, for this species and this 
 hybrid. Also no mention has been found for this genus of the 
 intimate association, which is shown here in the Salix lasiolepis 
 
 * The tree was destroyed soon after it was found, by the opening of a 
 street. 
 
 t The tree is still growing, near Berkeley. It was found the same 
 season as the 8. lasiolepis first mentioned. 
 
VOL. 2] Mott. Teratology in Calif ornian Willows. 183 
 
 series, of the micro- and macrosporangial tissues, ovules pro- 
 truding through pollen-sacs, and causing malformations in their 
 walls.* Likewise there seems to be no record for Salix of the 
 occurrence of four floral glands in a flower. 
 
 The possible bearing of the irregularities described upoti 
 several matters of discussion regarding the genus Salix will be 
 briefly indicated, such questions, for example, as whether the 
 ancestral flower was perfect,f instead of unisexual, or whether 
 even complete ; t what, if the flower were perfect, was the num- 
 ber of stamens ; and whether the two carpels of the now normal 
 pistil stand median or transverse. || 
 
 The facts described seem of interest, too, in connection with 
 a wider biological question of what, in general, causes to be pro- 
 duced, from the apparently common undifferentiated tissue, in 
 one case, pistil, in another, stamen, in another, leaf, stem, or 
 root, the question, that is, of what are the determining factors 
 in the development of this embryonic basal tissue which seems 
 to possess the potentiality of producing all forms of organ-tissue 
 peculiar to a species. 
 
 The occurrence of bisexual flowers with distinct pistil and 
 stamens, on these willows, in place of normal unisexual flowers, 
 and the abnormal development of ovules, female bodies, in- 
 stead of pollen-sacs, male bodies, and also of both ovules and 
 pollen-sacs at once, from the embryonic tissue of the connective', 
 are matters of interest in connection with the general problems 
 of sex, not only in the plant world but in the animal world as 
 well. In the animal world, also, cases of casual and of normal 
 hermaphroditism, of different degrees of intimacy, occur. 
 
 The perfect flowers with distinct pistil and stamens suggest, 
 among animals, the case of one of the toads, the male of which 
 
 * Two cases somewhat similar, of polleniferous ovules in passion-vines 
 (Passiflora coerulea and P. palmata) have been described and illustrated 
 by Salter (Trans. Linn. Soc. XXIV, p. 143, tab. XXIV), and a case of the 
 same monstrosity in a rose (Kosa arvensis) has been reported by Masters 
 (Seemann's Journal of Botany, 1867, p. 319, tab. LXXII, figs. B 1-9). 
 An instance of ovuliferous anthers in a gourd (Cucurbita) is mentioned 
 by Berkeley (Gard. Chron. 1851, p. 499.) 
 
 t Perfect : having the essential organs, viz. pistil and stamens. 
 
 + Complete: having pistil, stamens, and also perianth. 
 
 Median: in the line of the bract and the axis of the ament. 
 
 !l Transverse: in a line at right angles to the median line. 
 
184 University of California Publications. [BOTANY 
 
 has frequently a rudimentary ovary in front of the testes, and 
 again the case of the leech, which has regularly both ovaries 
 and testes in one individual, the two sets of organs being quite 
 distinct from each other. Many fishes, such as the cod, and. a 
 little lower, the hagfish are often similarly hermaphrodite. 
 
 The case of the monstrous bisexual floral organs, where pollen- 
 sacs and ovules develop with apparent indifference from con- 
 nective tissue, is similar, in some measure, to the case of the 
 regularly hermaphrodite snail, where sperm and ova are derived 
 from identical cells originating in the so-called hermaphroditic 
 gland, the sexual differentiation of these reproductive cells de- 
 pending merely upon which of two glands they are conducted 
 into for their development. 
 
 In degree of anatomical intimacy, the hermaphroditism seen 
 in the monstrous floral organs, where the seats of pollen-sac- 
 and ovule-production are so very close together on one phyllome, 
 is analogous to that bisexualism occasionally found in the 
 common frog, where a testis may have an anterior ovarian por- 
 tion, or an ovary be anteriorly replaced by a testis. 
 
 These cases, then, of sexual abnormality among plants, like 
 the similar instances among animals, bear upon various general 
 questions of sex: is hermaphroditism the primitive condition 
 in all organisms ; and are male and female sex-elements, micro- 
 spores and macrospores, sperm and ova, universally devel- 
 oped by differentiation of sexless cells, or, better perhaps, of 
 cells potentially of either sex ; and so on. The facts here noted 
 may prove of interest in some such connection. 
 
 A Teratological, Mainly Staminate, Salix lasiolepis Benth. 
 
 PlSTILLODY OP THE STAMENS : DESCRIPTION OP A SERIES OP SEX- 
 UAL ORGANS CONNECTING THE TWO STAMENS (THE TYPICAL MALE 
 FLOWER OP THE SPECIES ) WITH A BICARPELLARY PISTIL (THE TYPI- 
 CAL FEMALE FLOWER OF THE SPECIES ). This series of forms 
 showing increasing amounts of pistillody of stamens is repre- 
 sented in the series of figures on plates 19 and 20 (figs. 1-15). 
 Only enough of the abnormal forms are shown to make evident 
 
VOL. 2] Mott. Teratology in Calif ornian Willows. 1.85 
 
 the connection between stamens and pistil; numberless other 
 phases might have been interpolated. Figures 1 and 15 are from 
 normal trees, as types of the male and female flowers, for com- 
 parison. The various forms that occur in this abnormal tree, 
 it is well perhaps to state clearly at once, are found composing 
 as many distinct flowers. That is, a given pair of sexual phyl- 
 lomes do not go through a transformation from a perfectly devel- 
 oped pair of stamens, to a perfectly developed pair of carpels 
 forming a pistil, but remain constantly of the general character 
 that marks them in the floral bud, * until they finally fall with 
 the catkin. However, for the sake of vividness in bringing out 
 certain points that are shown by the series of forms, the differ- 
 ent members of the series are described as though each repre- 
 sented a temporary stage in a progressive change from stamens 
 to pistil taking place in a single flower. To follow a description 
 of such a process of change it is well perhaps first to have more 
 clearly in mind the initial and final stages. 
 
 The normal staminate flower in this species (cf. pi. 19, fig. 1) 
 consists of the two stamens, standing transverse, with cylindrical 
 filaments more or less united. The amount of union of the fila- 
 ments is variable in the species. In the flower chosen to show 
 the normal type, the filaments are united about to the middle 
 as in the case of the most nearly normal staminate flowers on 
 our abnormal tree. The anthers are adnate, that is, attached 
 by their whole length to the ends of the filaments. Each anther 
 is composed of four pollen-sacs (or microsporangia), in two 
 pairs, the two pollen-sacs of a pair being closely united to each 
 other, and shedding pollen through a single slit. The anthers 
 in the young stage are back to back (extrorse) (cf. fig. IA, pi. 
 19), but by the time they are ready to shed their pollen, they 
 have turned, through torsion of the filaments, to face the honey- 
 gland. 
 
 The normal female flower (cf. figs. 14 and 15) consists of 
 the unicellular, bicarpellary pistil. The two carpels stand it 
 
 * Whether a pair of phyllomes which are finally, at least, abnormal 
 undergo such a change, more or less completely, in the very young stage in 
 the floral bud, or whether the pair are abnormal from their very inception 
 at the meristem these are points apparently still to be worked out. 
 
186 University of California Publications. [BOTANY 
 
 is commonly considered transverse, like the stamens. The stalk. 
 or stipe, is short. The two placentae are median, and on the 
 walls of the ovary (parietal), and only on the lower part of 
 the walls (see fig. 13). The number of ovules is from six to 
 eight. The upper barren portion of the ovary is tapering-, and 
 affords space for the tufts of hair arising from the ovule stalks, 
 or funiculi. The style is single and rather short, and the stigrma 
 is more or less bilobed transversely. Two longitudinal seams, 
 one in the anterior * wall of the ovary and one in the posterior f 
 wall, mark the sutures between the two carpels and, as well, the 
 position of the two placentae within. Two slight longitudinal 
 grooves, on the right and left sides of the ovary, follow the 
 midribs of the carpels. It is along these lateral grooves, and 
 not at the sutures, that dehiscence occurs (dehiscence loculicidal, 
 see fig. 12) ; hence the two valves of the capsule are median and 
 the seeds are found on the middle of the valves. 
 
 The change from stamens to pistil is effected, in brief (a 
 detailed description follows later), by a median broadening and 
 also an apical development, and final union, of the two con- 
 nectives, to form the ovary of the pistil (1) the lower ovule- 
 bearing portion and (2) the upper barren portion and also 
 (3) the style and (4) the stigma, and further by a shortening, 
 and completion of the fusion, of the filaments, to form the stipe 
 of the pistil. The vegetative broadening of the connectives 
 occurs in such a way as, at an early stage in the series, to throw 
 the reproductive portions inward, so that the anthers, which are 
 produced from these portions, are, in the young stage, face to 
 face (introrse) (though normally extrorse) and the ovules, pro- 
 duced from the same reproductive regions, are, finally, enclosed 
 when the connectives unite. The change in the general form 
 of the phyllomes from that of a stamen to that of a carpel is 
 proportional (with some latitude) to the degeneration of the 
 pollen-sacs, and the replacement of them by ovules. That is, 
 all the male characters of the phyllome, being correlated, vary 
 approximately together, and the female characters the same. 
 
 * Anterior : toward the bract. 
 
 t Posterior : toward the axis of the catkin. 
 
VOL. 2] Mott Teratology in Calif ornian Willows. 187 
 
 The two phyllomes of a flower usually show the same amount 
 of change, but they may differ somewhat. Dehiscence of the 
 ovary, so long as it is monstrous to the extent of bearing anthers, 
 or anther-vestiges, is premature and at the sutures, so that the 
 transversely standing phyllomes remain intact ; after the anthers 
 have quite disappeared, the dehiscence is no longer premature, 
 and it occurs at the midribs (as in the flower on the normal tree) 
 so that the two valves of the capsule are anterior and posterior. 
 
 A detailed account of this change from stamens to pistil is 
 best given by describing in order the forms represented in the 
 series of drawings. Various facts that are exhibited by the 
 series as a whole, or by certain parts of it, will be touched upon 
 under those figures that are most suggestive of the particular 
 facts. In all the figures, it is the posterior side of the flower 
 which is shown, the side toward the axis of the ament. The 
 bract, honey-gland, and lower part of the united filaments are 
 in most cases not shown. Most of the figures are about ten times 
 natural size. 
 
 Fig. 1. This is a typical staminate flower from a normal 
 tree. It has already been described. The four pollen-sacs of 
 each anther are seen to be closely associated. Later on in tlie 
 series, these pollen-sacs, or microsporangia, degenerate, and the 
 vestiges often stand independent of each other, as their proto- 
 types in the Gymnosperms and Cryptogams quite commonly do. 
 Fig. IA shows a young diandrous flower (S. purpurea L.) with 
 extrorse anthers. 
 
 Fig. 2. This is the nearest approach to absolutely normal 
 flower that is found on the tree. It is a very common type. The 
 free ends of the two filaments (which we will call the "forks") 
 are apparently never perfectly cylindrical but are always slightly 
 grooved longitudinally on the sides that face each other. These 
 two grooves are the first indications of the formation of the 
 ovarian cavity of the pistil. As has been said, however, it is 
 the connectives which form the ovary, and later in the series, 
 these forks cease almost entirely to be developed, the pair of 
 filaments as a whole shortening to become the stipe of the pistil. 
 These two grooves at this stage arise as follows : 
 
188 University of California Publications. [BOTANY 
 
 While in the floral bud (and these first remarks are true 
 of stamens in general) the filaments attain about their full de- 
 velopment in thickness, but remain very short, until the anthers 
 have reached almost their full size and are nearly ready to dis- 
 charge their pollen. This keeps the anthers within the protec- 
 tion of the bract as long as possible. Just before dehiscence 
 of the pollen-sacs, the cells of the filaments increase greatly in 
 length, so that the anthers are exserted and the pollen is ren- 
 dered accessible to insects. Now while these abnormal stamens 
 are still in the bud, and the filament forks are short (remem- 
 bering that the anthers of the abnormal phyllomes, at least 
 while young, are introrse as explained later) there is a hollow 
 on the inner side of each fork, at the base of the anther. The 
 presence of this hollow is due to the fact that the connectives, 
 which bear the anthers directly, have commenced to become 
 grooved on the inner sides preparatory to the formation of the 
 concave ovary walls, and this grooving has extended downward 
 into the short filament forks. Hence as the cells of the forks 
 lengthen, these slight hollows develop as grooves along the whole 
 length of the forks. 
 
 The anthers here have twisted to face the honey-gland, and 
 the flower is otherwise normal. 
 
 Fig. 3. The forks and connectives here are broadened some- 
 what, medianly, and the grooving, or concaving, is more pro- 
 nounced. The anthers, which are still, from the bud. largely 
 facing each other, are seen to be introrse in their position on 
 the connectives; that is, the four reproductive portions of each 
 broadening connective, and hence the four pollen-sacs, have been 
 brought to face inward. This change is anticipatory of the 
 production of ovules which are to come from these same repro- 
 ductive portions for the ovules are finally to be enclosed by the 
 union of the broadened connectives.* 
 
 * The fact that the anthers in these abnormal forms are introrse, at 
 least in the young stage, instead of extrorse as in the normal flower of the 
 species, it is possible, it seems, to account for as follows: 
 
 In the young stamens and this is true of stamens in general of this 
 simple Salix type the four pollen sacs, or microsporangia, are produced 
 by a differentiation longitudinally of the tissue at the end of the stamen. 
 Two pollen sacs are formed in the anterior angles of the stamen, one right 
 
VOL. 2] Mott. Teratology in Calif ornian Willows. 189 
 
 The anthers are seen to have commenced to twist, mainly 
 through mere torsion of the forks, to face toward the honey- 
 gland. The tendency of the anthers to assume this position, 
 when they are mature, is present in the normal flower, and is 
 shown all through the series of abnormal forms as long as any 
 trace of the pollen-sacs remains. The advantage of the position 
 for insect pollination is obvious. 
 
 The posterior margins of the forks are ununited. Either the 
 twisting of the anthers to face the honey-gland has torn them 
 apart, or else the swelling of the still well-developed anthers 
 prevented them from ever uniting. This condition, namely, of 
 intact anterior suture and prematurely ruptured, or never 
 formed, posterior suture, is common in these monstrous forms 
 especially later in the series. In the earlier forms of the series, 
 while the anthers are still well developed, there is usually no 
 union of the forks at all. 
 
 Fig. 4. This figure shows the first appearance of ovules, 
 and the ovules are seen to be close to the basal end of the anther- 
 lobes. Only one of the stamens bears them. The two phyllomes 
 of a flower are not infrequently different in character to the 
 extent shown here. The right stamen is similar to both phyl- 
 lomes in fig. 2, though the groove is a little more evident. The 
 left fork has reached a little further developed stage. The 
 ovules are seen to point inward. There is only one ovule, as yet, 
 on each margin of the phyllome. The normal number on each 
 
 and one left, and two in the posterior angles, one right and one left, the 
 portion of the stamen between the two pairs being the "connective." The 
 position of the pollen-sacs in the mature state of a stamen is often dif- 
 ferent from this, owing to the later vegetative processes of growth in the 
 stamen-tissue, especially in the region of the connective. These are well- 
 established facts. Now in the normal flower of this diandrous willow, the 
 two stamens are close together, and it is apparently because the pollen-sacs 
 crowd against each other as they develop and swell with pollen, that they 
 come to face outward, or become extrorse. In the bud, the two extrorse 
 anthers, standing close together, appear as in fig. IA, pi. 19. In the 
 series of abnormal forms, where the two connectives gradually become 
 broadened medianly to form the concave ovary-wall, it is apparently be- 
 cause the vegetative increase necessary to effect this broadening is greatest 
 on that side of each connective which is farthest from the other connective, 
 that the four pollen-sacs are thrown to face inward, or become introrse. 
 
 The connectives in fig. 3 are not yet greatly broadened but there seems 
 to have been sufficient growth on their outer sides to throw the pollen-sacs 
 inward. 
 
190 University of California Publications. [BOTANY 
 
 margin of a carpel is three or four, since the normal numbor 
 on each placenta of the pistil is six or eight and the placenta 
 is formed by the union of two carpel margins. That these two 
 bodies are undoubtedly ovules is shown by the enlarged drawing, 
 fig. 16. This drawing represents an optical, longitudinal, me- 
 dian section of one such body through the raphe. A similar 
 section of an ovule from a pistil borne on a normal tree is iden- 
 tical with this. The characters are unmistakable: central nu- 
 cellus enclosed by two coats (integuments), basal micropyle 
 (ovule anatropus), and raphe, or prolongation of the stalk (funi- 
 culus), with its vascular bundle terminating at the (chalaza) 
 base of the two coats. 
 
 Fig. 5. Here the number of ovules has increased, and the 
 placental vascular strands have become evident as thickenings 
 on the margin of the forks. The vascular bundle in a normal 
 stamen is so little developed as scarcely to be detected, for only 
 enough nutritive material has to be conducted to the pollen-sacs 
 (microsporangia) to bring them to maturity, while to the ovules 
 (macrosporangia) sufficient food has to be conducted not only 
 to bring them to maturity as macrosporangia, but also to supply 
 their demands, after their fertilization, by the pollen, while they 
 are being developed to maturity as seeds. 
 
 There is an ovule at the base of each half -anther, or anther- 
 lobe, and the bodies a and c are ovules even protruding through 
 the anthers, so closely are the ovules associated with the pollen- 
 sacs in the production of both from the reproductive portions 
 of the phyllomes. In fact it is this close association, which is 
 seen commonly in the forms of this type, that shows that it is 
 a fairly definite part of a phyllome which is reproductive. In 
 other words, the connective region of the stamen and the pla- 
 cental regions of a carpel seem to be homologous. 
 
 The small bodies higher up, in the sinuses, or furrows, in 
 the face of the anthers, where two pollen-sacs unite to form a 
 half anther, bodies which look like ovules, are in reality lobes 
 in the walls of the pollen-sacs. There seems to be no question 
 that these lobes are caused by abortive ovules beneath, for though 
 in most cases there is nothing but pollen within such lobes, in 
 
VOL. 2] Mott. Teratology in Calif ornian Willoivs. 191 
 
 some instances there is a slight protuberance on the connective 
 beneath; and an ovule, in general, originates in this way. The 
 cases such as seen here (noted above) of ovules actually pro- 
 truding through pollen-sac tissue, are further evidence that the 
 lobes are caused by incipient ovules. The tissue &, for example, 
 around the base of the protruding ovules a is unmistakably 
 anther-tissue, and it would doubtless have constituted a lobe only 
 that the ovule succeeded in keeping from being overwhelmed, 
 as ovule and pollen-sac developed together; that is, the crowding 
 of the anther-tissue here was partially withstood by the devel- 
 oping ovule, while in other cases anther-tissue gained almost 
 complete supremacy, and yet lobes in the pollen-sac walls were 
 caused by the overwhelmed ovules.* 
 
 The pollen-sac lobe d looks particularly like an ovule ; it 
 stands out in the sinus almost independent of the main portion j 
 of the pollen-sac. The lobe e on the right half of the right 
 stamen, on the other hand, is merely a slight elevation in the 
 surface of the wall of its pollen-sac. Other lobes (also marked e) 
 are more or less intermediate in form. (The means of identify- 
 ing pollen-sac tissue are seen in figs. 17-19, and the matter of 
 this identification is further referred to under fig. 7.) 
 
 The right anther lobe has dehisced. 
 
 Fig. 6. The forks and connectives in this figure have become 
 considerably broader, and here, on the left phyllome at c, is seen 
 the first indication of apical development of the connective to 
 form the upper barren portion of the ovary, and the style and 
 stigma. The right connective is not yet so developed. (The 
 ovules are usually produced nearer to the anthers than shown 
 here. ) 
 
 Fig. 7. The right phyllome here is still predominantly stam- 
 ineal, the anther being well developed, and the fork elongated. 
 The left phyllome, however, is about equally divided in char- 
 acter between stamen and carpel. The anther has been reduced 
 to vestiges, and the fork accordingly is only partially developed, 
 
 * These effects of pollen-sacs and ovules upon each other, where they are 
 produced together, are such as one might expect, in consideration of the 
 processes by which pollen-sacs and ovules, in general, are produced by dif- 
 ferentiation of embryonic tissue of stamen and carpel respectively. 
 
192 University of California Publications. [BOTANY 
 
 though the united part of the two phyllomes is still of the normal 
 length in the stamen. The development of pollen-sacs and the 
 elongation of the cells of the filament are correlated male char- 
 acters of the phyllome, and the development and the elongation 
 vary in amount approximately together. The lengths of the 
 cells of the two forks in this case were found to be about in- 
 versely proportional to the lengths of the forks. On the left 
 phyllome, ovules have almost entirely supplanted the right lobe 
 of the anther, all that remains of the lobe being the vestige d. 
 At the same time, there has occurred a considerable apical devel- 
 opment of the connective toward the formation of the upper 
 pistil-parts (barren portion of the ovary, style, and stigma), 
 this being a female character of the phyllome correlated with 
 the production of ovules. The ovules here are all well devel- 
 oped. The beginning of the upper barren portion of the ovary 
 is seen at 7i', the style at li, and the stigma at i. The styles 
 and stigmas of the two phyllomes unite later on. The style here, 
 being still free, has shriveled and curled downward. 
 
 The body d on the left phyllome might be taken superficially 
 for an ovule, it being minute and otherwise similar to an ovule, 
 and being borne among ovules, but it proves under the higher 
 power of the microscope to be a pollen-sac vestige; just as the 
 still more ovule-like body d, pi. 19, fig. 5, proved to be of pollen- 
 sac tissue. Even under the lower power, the ovules appear 
 slightly fluted longitudinally and the oblong shape indicates the 
 polarity they possess; while the pollen-sac lobes and even the 
 least developed pollen-sacs have a smooth surface of small poly- 
 gonal cells such as normally developed pollen-sacs have. Pollen- 
 sac lobes and vestiges also often contain grains of pollen, normal 
 at least superficially (see pi. 20, fig. 18), (though these are often 
 lost in dissecting out the body for examination). Under the 
 higher power, d, for example, shows the cellular wall structure 
 which is characteristic of the normally developed pollen-sac. 
 This peculiar structure is illustrated by figs. 17 and 19. The 
 wall, is composed of an outer layer of thin-walled cells and an 
 inner layer of so-called fibrous cells. On approaching maturity, 
 the outer layer dries more rapidly than the inner, and contracts 
 
VOL. 2] Mott. Teratology in Calif ornian Willows. 193 
 
 more (as does the drying film of clay over coarser material on 
 the site of a recent pool), and the rupturing along the weakest 
 line, namely, the suture between two pollen-sacs, is dehiscence. 
 Hence the identity of ovules and of pollen-sac vestiges and lobes 
 is established. 
 
 Fig. 8. The connectives here are broadened almost to the 
 full amount ; the anterior margins are united ; a style and stigma 
 have developed from each connective, though they are mal- 
 formed and the styles and the stigmas are still ununited; the 
 old filament forks are both reduced, though the old united por- 
 tion of the filaments is still longer than a stipe. The anthers 
 at this stage are usually more degenerate. There is about a 
 normal number of ovules, on the anterior placenta. The ovarian 
 cavity is well developed, though the posterior margins of the 
 phyllomes (which were very likely united in a younger stage) 
 have been torn apart. This figure illustrates the fact that the 
 monstrous ovary dehisces prematurely and at the sutures, in- 
 stead of at the midribs. 
 
 Fig. 8. The connectives have now both reached full carpel- 
 width, both anterior and posterior margins have united below, 
 though the upper parts are still free. The tapering upper por- 
 tion of the ovary is not yet well developed, and the figure shows 
 (better than the last) a rather common persistence of abnorm- 
 ality of style and stigma. The anthers have become so nearly 
 eliminated that dehiscence is no longer premature, and yet seeds 
 are not matured, the organ falling early. As a whole, the two 
 phyllomes have come to suggest pistil rather than pair of stamens. 
 
 Fig. 10 shows close approximation to normal pistil. The 
 phyllomes are wholly united, the ovary is tapering above, and 
 style is single and there is clear demarcation of the style from 
 the ovary. The lobing of the stigma, however, is still right and 
 left, corresponding to the original stamens, and not yet to the 
 valves. The stipe is still abnormally long. (Even though, how- 
 ever, the lobes of the stigma in the normal pistil in this species 
 commonly correspond to the valves, dehiscence does not extend 
 through the style and stigma, but style and stigma either fall 
 intact from the ripe capsule or else break away from one valve 
 at the time of dehiscence, and remain attached to the other.) 
 
 
 ?x 
 
 ( UNIVERSITY ] 
 
 V OF / 
 
194 University of California Publications. [BOTANY 
 
 Fig. 11 shows the same specimen as fig. 10, but with the 
 nearer (posterior) half of the ovary removed, to show that the 
 pollen-sacs have completely disappeared, and that the ovules 
 are in their normal position: that is, they are median, for they 
 are borne on the margins of transverse phyllomes, as we have 
 seen in the preceding figures ; and they are on the lower part of 
 the ovary-wall, for the old filament forks below the reproduc- 
 tive portion of the phyllomes have been reduced and at the 
 same time the old connectives the reproductive portions them- 
 selves have been developed apically to form the upper barren 
 portion of the ovary-wall. 
 
 Fig. 12 shows the completion of the change. The stipe is of 
 normal length, the ovary has swollen with seeds and down, and 
 has matured as a normal capsule dehiscing at the carpel mid- 
 ribs so that the valves are anterior and posterior. Style and 
 stigma have fallen away. The valves separate as normally, only 
 half or two-third's of the way down. 
 
 Fig. 13 presents the anterior valve of the capsule in fig. 12, 
 pi. 20, with basal remnants of the other valve, to show the char- 
 acteristic position of the placenta, median and at the lower 
 half of the suture between the phyllomes. 
 
 Fig. 14, showing a pistil from a normal tree, has already 
 been described. It exhibits the typical characters which we 
 have seen developing through the series. The view of the pistil 
 is posterior, as in the preceding figures, for the sake of com- 
 parison. 
 
 Fig. 15 shows the same flower as in the last figure, but turned 
 somewhat to bring into view one of the slight grooves at the 
 carpel midribs along which normal dehiscence occurs, as con- 
 trasted with the abnormal dehiscence at the sutures in the mon- 
 strous forms. 
 
 There is thus a complete series of forms on this abnormal 
 Salix lasiolepis connecting, by slight gradations, a pair of 
 stamens and a bicarpellary pistil. 
 
 MANNER OF OCCURENCE OF THE ABNORMAL FLOWERS ON THE 
 CATKINS, AND EFFECT OF THEIR PRESENCE ON THE TIME OF FALLING 
 OF THE CATKINS. The flowers abnormal to the tree are, of 
 
VOL. 2] Mott. Teratology in Calif ornian Willows. 195 
 
 course, both those with any form of monstrous hermaphrodite 
 organ and also those with perfectly developed pistil, for the tree 
 as a whole was evidently predominantly staminate. The pre- 
 dominance of the male character of the tree as a whole was 
 shown by the fact that when the tree was in full bloom, anthers 
 were commonly enough present in the flowers to give almost as 
 much of a yellow cast to the tree as is characteristic of the 
 normal male individual. Many of the catkins were composed 
 wholly of practically normal male flowers, but the majority of 
 the catkins were made up of monstrous hermaphrodite flowers 
 which yet were more staminal in character than pistillary. 
 Quite a large number of catkins which were chiefly of monstrous 
 flowers showed more of a green cast than yellow, the majority 
 of the sex-organs here resembling pistils rather than pairs of 
 stamens. Again a considerable number of catkins were com- 
 posed wholly of perfectly developed pistillate flowers. Then 
 in addition to these four sorts of catkins named, composed 
 wholly or mainly of one type of flower, there were many catkins 
 showing combinations of these types of flowers in various pro- 
 portions; and the different types occuring on a given catkin 
 might be rather thoroughly intermingled, or might be in dif- 
 ferent parts of the catkin (as base, middle, and tip), but without 
 any regularity as to which type occurred on any part. 
 
 The presence of monstrous sex-organs on a catkin tended to 
 cause the catkin to drop early, earlier even than a wholly male 
 catkin drops. In general, of course, male catkins of the willow 
 fall earlier in the season than the female, for the function of 
 the staminate catkins in fulfilled earlier, that is, as soon as the 
 matured pollen is shed, while the pistillate catkins must per- 
 sist, after this escaped pollen has fallen upon the stigmas, until 
 the seeds have matured and scattered. These monstrous her- 
 maphrodite organs, however, unless very predominantly of one 
 sex or the other, soon shriveled after the opening of the flowers, 
 so that catkins wholly of monstrous flowers soon fell after 
 blooming. The majority of the catkins borne by the tree being 
 of the decidedly monstrous type, the tree as a whole had few 
 catkins hanging upon it at a time (March 26) when normal male 
 individuals near bv still had many catkins on them in bloom. 
 
196 University of California Publications. [BOTANY 
 
 A peculiar fact noticed was that a very few normally de- 
 veloped pistils on a catkin were sufficient to keep the catkin from 
 falling until the maturation of the seeds in these pistils, what- 
 ever the character of the remaining flowers, male, hermaphrodite, 
 or of both kinds. Hence, as catkins with just a few normal 
 pistils were not rare, toward the end of the season, when capsules 
 on trees generally were dehiscing, a considerable proportion of 
 what catkins still remained on this tree displayed only very 
 scattering capsules ; and search among the fallen catkins beneath 
 the tree revealed very few that had more than one or two un- 
 matured pistils. 
 
 Catkins throughout the season, whether blooming earlier or 
 later, showed in about the same proportion of cases the ab- 
 normalities, arid the diversity among the catkins was about the 
 same throughout the season. 
 
 A Teratological, Mainly Pistillate, Salix lasiandra Benth.X 
 babylonica L.* and Some Related Individuals. 
 
 ON THE TREE FIRST * NAMED : STAMINODY OF THE PISTIL ; 
 
 BISEXUAL FLOWERS WITH NORMALLY FORMED PISTIL AND STAMENS ; 
 EXTRA FLORAL GLANDS; OTHER NOTES; THIS TREE AND TWO SIMI- 
 LAR ONES ARE SCION-PROPAGATED. Staminody of the pistil was 
 one of the abnormalities shown by this mainly pistillate tree. 
 That is, in the normal position of the pistil there occurred 
 monstrous organs which possessed in various proportions the 
 characters both of pistil and of stamens. (See a pi. 20, fig 21.) 
 This is the converse of the apparent transformation of sex- 
 organs seen in the Salix lasiolepis: there the two stamens ap- 
 peared more or less changed to pistil, here the pistil is more or 
 less changed to two stamens. The monstrous pistils vary from 
 
 * There are (still growing) three trees of this general description, 
 standing near together (at the corner of San Pablo Avenue and McAvoy 
 Lane, near Berkeley), which were grown from cuttings planted some 
 fifteen years ago, and also about a dozen trees of similar description, but 
 of both sexes, standing not far from the first three in a small mixed grove 
 of willows (at the foot of Powder Works Hill), which were naturally 
 grown, probably from seed. The three cuttings came from this grove, and 
 were probably taken from one or more of the a'bnormal mainly pistillate, 
 naturally planted trees. 
 
OF THE 
 
 UNIVERSITY 
 
 OF 
 
 VOL. 2] Mott. Teratology in Calif ornian Willows. 197 
 
 the normal in a manner very much similar to that seen in the 
 various other cases of staminody of the pistil in Salix which 
 are described in the literature. The stipe is usually lengthened, 
 filament-like (or like two filaments fused) ; the ovary is short- 
 ened; the style and stigma are more or less nearly lost; the 
 posterior suture is prematurely ruptured; and two anthers, 
 either perfectly developed or more or less rudimentary, are dis- 
 closed, one on each side, borne on the inner wall of the ovary; 
 a reduced number of ovules are borne on the median, parietal 
 placentae; pollen is produced and discharged; the exposed 
 ovules soon shrivel and fall. Probably a complete series of 
 forms from pistil to stamens could be worked out as in the case 
 of the converse change. 
 
 The abnormal occurrence of bisexual flowers with normally 
 developed pistil and stamens (perfect flowers) is a striking 
 peculiarity of this tree (see pi. 20, fig. 20). The general appear- 
 ance of these flowers suggests (if one considers them with the 
 usually accepted theories of descent in mind), that their occur- 
 ence is due to reversion, and hence that the hermaphroditism 
 in them is of a different sort from the hermaphroditism seen in 
 the monstrous bisexual organs occurring either on this tree or 
 on the one first described. 
 
 These perfect flowers are very numerous (though a pre- 
 ponderance of normal pistillate flowers on the tree as a whole 
 shows it to be predominantly female), and they are fairly uni- 
 form in type, consisting usually of a pistil between two stamens, 
 the latter standing right and left (transverse). The pistil and 
 stamens are in full bloom (in "anthesis") at different times 
 (flower "dichogamous"), as is very commonly the case in per- 
 fect flowers. And here the stamens are in anthesis last ; that 
 is, the anthers do not shed their pollen until the stigma is no 
 longer receptive (dichogamy " p rot ogy nous"). The figure (20) 
 shows that the filaments of the two stamens are still short, and 
 the pollen-sacs unopened, whereas the pistil is ready for pollina- 
 tion. This difference in time of anthesis of pistil and stamens 
 in the perfect flowers is rendered more striking by the fact that 
 there are other stamens on this tree, often on the same catkins 
 
198 University of California Publications. [BOTANY 
 
 with perfect flowers, yet usually occurring in purely male 
 flowers, which are in anthesis, with lengthened filaments, about 
 the same time that the pistils are in anthesis. Sometimes the 
 two stamens of a perfect flower, though normal in form, are 
 small and pale as though not fully restored. 
 
 Again there is suggestion of reversion to an ancestral form 
 of flower in the fact that there is a tendency on this tree, 
 especially in the perfect flowers, toward the establishment of 
 two extra glands, standing transverse, in addition to the usual 
 one or two median (fig. 20 does not show the extra glands). 
 These glands are referred to later as possible vestiges of an 
 ancestral perianth. (They are also described somew T hat more 
 completely later on.) 
 
 The male flowers mentioned as occurring on this hybrid 
 tree are rather common. There are usually two or three stamens 
 in a flower, though as high as five occur. The supposed parent 
 species are tri- and pentandrous respectively. 
 
 Numerous variations of the types of flowers mentioned 
 occur. Roughly classifying all the sorts observed on the tree 
 we have (1) normal pistil (a) alone, (&) between two short* 
 stamens; (2) monstrous pistil (a) alone, (b) between two short 
 stamens, (c) with one longf stamen; (3) one long stamen and 
 one short stamen; (4) one long stamen between two short 
 stamens; (5) one to five long stamens; (6) two long stamens 
 each with a distinctly double anther; (7) perfectly developed 
 ovary and anther on the end of one stalk. Doubtless other com- 
 binations and forms could be found. 
 
 The aments are very diverse, as in the case of the Salix lasio- 
 lepis. The majority are strictly pistillate, but yet very many 
 are almost wholly of perfect flowers, others are purely staminate, 
 and still others are made up chiefly of the flowers consisting of 
 the monstrous organs. Then there are catkins that combine in 
 various proportions all the types of flowers described. 
 
 * Short : that is, of the set which are late in developing to shed their 
 pollen. 
 
 t Long : that is, of the set which are early in developing to shed their 
 pollen. 
 
VOL. 2] Mott. Teratology in Calif ornian Willows. 199 
 
 Branchlets show the same diverseness, many being largely 
 female, some chiefly male, some mainly of perfect flowers, others 
 pronouncedly of monstrous organs. 
 
 An interesting feature of the teratology of this tree is that 
 in this third spring season the abnormality of the flowers is 
 more general over the tree than it was in the first season. (The 
 tree was not closely observed the second season). The first 
 spring, the lower part of the tree (the tree being about forty 
 feet tall, and of a spreading habit) was most affected, the upper 
 part being but little abnormal. This spring the lower part of 
 the tree is somewhat more affected than when first noticed, and 
 the upper part has become about as much abnormal as the lower. 
 Close to this tree are two other individuals, apparently of the 
 same cross, and both also mainly pistillate, the three having been 
 planted at the same time as cuttings; and these others have 
 shown something of the same seasonal increase of abnormality 
 that the first has shown. (To distinguish between the three 
 trees, the first described may be referred to as the north tree, 
 the other two as middle and south respectively.) The first 
 spring the middle and south trees were not noticeably abnormal 
 in any degree, though they were carefully examined. This 
 third spring, the middle tree shows abnormality in a small pro- 
 portion of the aments (the irregularities being of the same gen- 
 eral description as those shown by the north tree) and the south 
 tree has borne one bisexual flower with normal organs, a pistil 
 and one stamen. A thorough search over this south tree failed 
 to reveal any other abnormal flowers. 
 
 An irregularity in this one bisexual flower on the south tree, 
 which is rather striking if one is looking for evidence of rever- 
 sion, is the presence of two small lateral glands, since the normal 
 one or two median glands in Salix are quite commonly con- 
 sidered as vestiges of an ancestral perianth, and here apparently 
 is indication of two perianth phyllomes which are normally en- 
 tirely missing. The other flowers on the tree had but a pos- 
 terior gland. 
 
 It seems rather natural that restoration, in some degree, of 
 lost perianth phyllomes should accompany the restoration of lost 
 
200 University of California Publications. [BOTANY 
 
 sexual phyllomes. Many of the flowers also on the north tree, 
 especially the bisexual ones with normal organs, had, as noted 
 above, more or less evident lateral glands. 
 
 Some further notes as to the glands on these three hybrid 
 scion trees are of interest as bearing upon the question of what 
 are the species represented in the trees (the descent, however, 
 being largely judged from other facts). On the north tree 
 there are usually two median glands, one anterior and one pos- 
 terior, to each flower. (The glands are small yellow thallus- 
 like, or flattened, bodies at the base of the sexual organs, e and 
 /, pi. 20, fig. 20). Now, of the probable parents, Salix lasiandra, 
 in this locality, has commonly only the posterior gland in the 
 flowers of both sexes, and Salix babylonica has only the posterior 
 in the female flower, but both anterior and posterior in the male. 
 The north tree, then, which is quite largely bisexual, seems to 
 follow the male babylonica in having two median glands. It 
 should be mentioned here that a male Salix lasiandra found this 
 season, has quite uniformly four glands to the flower, though 
 the tree is otherwise apparently normal, and hence this hybrid 
 mother tree in showing a tendency to four glands, may be merely 
 exhibiting an occasional normal lasiandra character, and not be 
 giving evidence of reversion in the flower in this particular. (A 
 flower with the four glands from this Salix lasiandra is shown 
 in fig. 22, pi. 19. The five stamens and the bract have been cut 
 off). The middle hybrid tree, which has a majority of its 
 flowers pistillate, has commonly only the posterior gland, like 
 Salix lasiandra and the female babylonica; and the south tree, 
 which has all its flowers pistillate, excepting the one bisexual 
 flower with the two small lateral glands has aside from this one 
 flower only the posterior gland throughout. 
 
 TENDENCY TO ABNORMALITY PRESENT IN THE SCIONS HYBRID, 
 SEEMINGLY BY PERPETUATION FROM PARENT STOCK; PROBABLE 
 SOURCE OF THE SCIONS, AND EVIDENCE AS TO THE TWO ORIGINAL 
 PARENT SPECIES. The abnormalities in these three scion-propa- 
 gated trees being apparently not due to seasonal influences nor 
 to local conditions (which are normal), nor to such agencies as 
 
VOL. 2] Mott. Teratology in Calif ornian Willows. 201 
 
 insects or injuries (for the trees appear perfectly healthy and 
 sound), some effort was made this spring to find whether they 
 came from abnormal stock, and, as stated above in a foot-note, 
 their immediate origin was traced with considerable certainty 
 to some abnormal mainly pistillate trees in a naturally grown 
 mixed grove of willows near by.* With the latter are some 
 mainly staminate abnormal individuals of the same cross. 
 These naturally grown trees show the same general characters, 
 both normal and abnormal, that mark the scion-propagated 
 trees. But there are some abnormal features (some of which 
 are mentioned later), which are rather more common here than 
 on the scions, and there is some noticeable variation from the 
 scions in the relative prominence of the different abnormal 
 features.f For example, the tendency to four glands is con- 
 siderably more evident here. Various combinations of the four 
 glands occur in a flower, and they may be quite distinct, or more 
 or less united by their bases into a ring encircling the sex-organs, 
 like a miniature sympetalous corolla. 
 
 The hybrid origin of all these abnormal trees such descent 
 seeming quite certain is of interest as being very likely the 
 cause of the teratology which marks them, hybrids in general 
 being prone to show abnormalities, especially in the flowers. 
 The scion trees were first taken for lasiandras, since they re- 
 semble that species more than any of the other native species, 
 but later some planted exotic pistillate babylonicas, or weeping 
 willows, were found near one edge of the natural grove of wil- 
 lows, and these are probably the original maternal stock of all 
 the abnormal trees, while native lasiandras, which are common 
 in the neighborhood, are probably the paternal stock. The 
 babylonicas were planted some fifty years ago to mark some old 
 Spanish graves, and their flowers were probably pollinated at 
 some time from lasiandras and then seeds from them blew over 
 and gave rise to the abnormal trees in the grove. The grove 
 
 * Normal trees in the grove are : many Salix lasiolepis Benth., a few 
 S. lasiandra Benth., and a few S. laevigata Bebb. 
 
 t The matter of whether scions from these hybrid willows will per- 
 petuate the floral abnormalities, in some measure, as seems almost certainly 
 to have occurred, is being tested in the University gardens with some cut- 
 ting from one of the trees. There are no flowers this first season. 
 
202 University of California Publications. [BOTANY 
 
 came into notice only about twenty-five years ago, and the trees 
 forming it were still quite small when the three cuttings were 
 taken. 
 
 The probable origin of these abnormal trees by the crossing 
 of these particular species is indicated in the trees themselves 
 by their showing many characters intermediate between the 
 normal characters of 8. lasiandra and 8. babylonica, as to 
 habit, number of stamens, number of glands (as noted above), 
 and so on. It is true that the common presence in the perfect 
 flowers of two stamens might suggest that the male parent was 
 of some such native diandrous species as Salix lasiolepis Benth ; 
 for, of the two assumed parent species, Salix lasiandra has 5-4 
 stamens (for this region),* and Salix babylonica three, and, 
 furthermore the purely male flowers on the abnormal trees are 
 most frequently of two or three stamens; but yet the abnormal 
 trees have not uncommonly as high as five stamens in the purely 
 male flowers, like normal 8. lasiandra, and besides they have 
 many characters, aside from the number of stamens, which seem 
 to preclude the supposition of parentage on one side by any of 
 the native species other than 8. lasiandra.j 
 
 It may be noted in connection with this matter of parentage, 
 that Salix babylonica has been frequently reported as abnormal 
 in the flowers. 
 
 ON SOME NATURALLY GROWN INDIVIDUALS APPARENTLY RE- 
 LATED TO THE SCION TREES: SEPARATION, PARTIAL TO COMPLETE, 
 
 OF THE TWO CARPELS OF THE PISTIL WITHOUT BISEXUALISM, AND 
 
 ALSO MULTIPLICATION OF CARPELS. These two irregularities are 
 the most striking among those referred to above as occurring 
 commonly on some of the abnormal trees in the grove and not 
 to any extent on the scion trees. The abnormalities are most in 
 evidence on the mainly pistillate trees. Cases illustrative of 
 
 * Jepson, Flora of Western Middle California. 
 
 t A case of the occurrence of willows which were considered to be 
 Salix lasiandra X babylonica, from a cross between native staminate 
 lasiandras and planted exotic pistillate babylonicas was reported by Ander- 
 son (1890) from Santa Cruz, Calif. These trees were described on account 
 of being monoecious, staminate and pistillate flowers occurring together on 
 the catkins, but none of the other abnormalities that are present on these 
 Berkeley hybrids, are mentioned. 
 
VOL. 2] Mott. Teratology in Calif or nian Willows. 203 
 
 these two general manifestations of abnormality have been fre- 
 quently reported by writers,* so mere mention of some of the 
 forms in point found on our trees will be made. There are 
 found then here: (1) pistils showing various degrees of separa- 
 tion of the carpels; (2) two distinct unicarpellary pistils re- 
 sembling miniature pea-pods in one flower; (3) three such pistils 
 in one flower; (4) a normal bicarpellary and a unicarpellary 
 pistil in one flower; (5) two perfectly developed and distinct 
 bicarpellary pistils in one flower. Sometimes the unicarpellary 
 pistils were monstrous in being bisexual, like the bicarpellary 
 pistils described above, a more or less malformed anther bulging 
 out at the suture between the carpel margins. 
 
 Bearing of the Abnormalities Described on Some Moot 
 Points Regarding the Salix Flower. 
 
 The notes following here are based mainly on the usually 
 accepted general theories of evolution. Most comment of 
 writers, suggested by similar abnormalities in Salix, has been so 
 made, though the conclusions to which different writers have 
 inclined have differed. A more recent fundamental theory, 
 under which such abnormalities may be interpreted, is referred 
 to later. 
 
 WAS THE ANCESTRAL FLOWER BISEXUAL OR UNISEXUAL? This 
 
 question is, perhaps, the one most commonly discussed, and the 
 answer to it seems to depend mainly upon the interpretation 
 one makes of the present sporadic instances of hermaphroditism 
 in the flower. In fact, it is generally the encountering of cases 
 of hermaphroditism that raises this question about the ancestral 
 flower. 
 
 On our trees there seem to be two distinct sorts of hermaphro- 
 ditism. On all the trees there are (1) the flowers with the 
 monstrous bisexual phy Homes, and (2), especially on the hybrid 
 tree first described, the flowers with the normally developed pistil 
 and stamen. On this latter tree, the two kinds of androgyny 
 
 * Von Seemen (1886 and 1895), in particular, has pictured and de- 
 scribed many abnormal forms coming under these two heads. 
 
204 University of California Publications, [BOTANY 
 
 are so associated as apparently to show clearly that they are 
 distinct phenomena ; and the perfect flowers with normal organs, 
 as noted in describing them, certainly show many of the gen- 
 erally accepted signs of reversion, entirely aside from any con- 
 sideration of the monstrous bisexual organs, and hence they 
 seem to indicate that the ancestral flower was bisexual. 
 
 Some writers (Camus, 1889, and Von Seemen, 1886 and 
 1895) have inclined to the belief that the perfect flowers often 
 seen have come about by sexual change of part of the phyl- 
 lomes of an always unisexual flower, and point to such cases 
 of complete substitution of one sexual organ for the other as 
 seen in our Salix lasiolepis series, to show the possibility of the 
 origin of bisexual flowers with normal organs in such a way;* 
 and on any polyandrous male individual such origin does seem 
 possible, for there are sufficient phyllomes there in one flower 
 for both stamens and pistil. But on a pistillate individual, 
 with its two phyllomes to a flower, such as our hybrid, we 
 should have to conceive of first a multiplication of phyllomes and 
 then a development of them as stamens instead of as carpels. 
 Multiplication of phyllomes is a common phenomenon, it is 
 true; the polyandry in some species of willowsf and the more 
 extreme cases of polyandry in the other genus of the family, 
 the poplars,| doubtless arose by reduplication, and the extra 
 unicarpellary and bicarpellary pistils on some of our mainly 
 pistillate hybrids are probably to be accounted for in this way ; 
 but it seems far-fetched, in the case of the perfect flowers on 
 our predominantly pistillate hybrid, which flowers consist rather 
 uniformly of pistil between two stamens, to assume that these 
 two lateral stamens are present through reduplication of phyl- 
 
 * Von Seemen and others, interpreting in this way the apparent mutual 
 convertibility of pistil and stamens, have held that pistils and stamens in 
 Salix are morphologically identical. But if, however, there are sufficient 
 facts which can be interpreted as indicating that the ancestral flower was 
 bisexual, there would seem to be no more reason for considering "the male 
 and female sex-organs morphologically equivalent" here than in flowers in 
 general. 
 
 f As high as twenty stamens occurring in some species, according to Pax 
 (in Engler and Prantl's Pflanzenfamilien). 
 
 \ As many as thirty stamens being sometimes present here, according to 
 Pax; and even seventy-one according to Jepson, Flora of Western Middle 
 California, 2nd ed., MS. 
 
VOL. 2] Mott Teratology in Calif ornian Willows. 205 
 
 lomes, and then that they are stamens by conversion. Although 
 monstrous bisexual organs occur on this tree, even in the same 
 catkin with the perfect flowers, these two lateral stamens are 
 not found connected by any intermediate forms, in the same 
 position in the flower, with carpels. They look like restored 
 organs, original microsporophylls. The bisexual bodies that do 
 occur, on the contrary, seem always to be more or less changed 
 fo'carpellary pistils, standing in the normal position of the pistil. 
 They may even occur in the place of the pistil in otherwise per- 
 fect flowers. 
 
 Further facts indicating reversion here, and hence indicat- 
 ing once perfect flowers, are, that the perfect flowers are pro- 
 togynous, that their stamens are sometimes small and pale, as 
 though incompletely restored, that the perfect flowers are numer- 
 ous and of a fairly uniform type, that there is a tendency to a 
 concomitant restoration of glands, which seem to be, with the 
 normal glands, perianth vestiges, and that there are only two 
 stamens in these perfect flowers, as though by restoration of an 
 original number (as opposed to a later tendency to multipli- 
 cation of stamens in the family). 
 
 Miiller believed the ancestor of the^present unisexual flower 
 was bisexual, and was led to this belief by consideration of the 
 same evidence of possible change of organs (as seen in our Salix 
 lasiolepis series) which has led others to believe the present occa- 
 sional bisexual flower with normal organs is derived from an 
 ancestral unisexual flower. Since stamens can thus change com- 
 pletely to pistils, and vice versa, he concluded that the modern 
 unisexual flower developed one way or the other from a bisexual 
 flower. But if that be true, why have the normal pistillate flowers 
 and the diandrous male flowers only two phyllomes each? Evi- 
 dently we must assume suppression there of the phyllomes of 
 the lost sex (suppression being in fact a common occurrence, 
 as noted later), and hence why not assume that there has oc- 
 curred uniformly a suppression of the phyllomes of one sex or 
 the other in an ancestrally bisexual flower, to produce the mod- 
 ern flower, and then that the increased number of phyllomes 
 in polyandrous species has occurred uniformly by reduplication ? 
 
206 University of California Publications. [BOTANY 
 
 This monstrosity of bisexualism occurring in one organ seems 
 capable of a double interpretation, and apparently is one of 
 those numerous teratological phenomena that must be consid- 
 ered as of no morphological significance. As noted above, this 
 hermaphroditism shown in the monstrous organs seems to be of 
 a sort distinct from that exhibited in the perfect flowers with 
 normal organs. 
 
 Aside from the evidence that has been noted, which it seems 
 may be accepted as showing that the ancestral flower was bisex- 
 ual although it is teratological evidence, there is the additional 
 strong support of the theory of original bisexualism in the 
 great similarity in most species practical identity aside from 
 the sexual organs of the male and the female aments, in all 
 characters, shape, size, coloring, position on the branches, char- 
 acter of the individual bracts, degree of hairiness, etc. In the 
 other amentiferous families with unisexual flowers, such as the 
 oaks, walnuts, hazels, and even birches and alders, there is 
 a very great difference between the floral clusters of the two 
 sexes. In such cases it might be difficult to imagine how a uni- 
 sexual flower could be the result of suppression of one of the 
 sexes in a perfect flower. But in Salix, unisexualism by such a 
 process is easy to conceive. 
 
 That unisexual flowers may arise by abortion of one sex, an- 
 droecium or gynaeceum, is shown, it is commonly considered, 
 beyond doubt, in such a case as that of the polygamous genus, 
 Acer, the maples. Here there are sometimes bisexual flowers 
 to be found with organs of both sexes normal ; in other flowers, 
 the organs of one sex have become vestigial, though they are 
 still recognizable as being remnants of such organs, and in still 
 others, the organs of one sex have suffered complete obliteration. 
 
 That suppression is common in the willow family, Pax * 
 thinks is shown in the reduction of the broad disk of Populus 
 to the usual one or two small glands in Salix. Evidently he con- 
 siders both disk and gland as reduced perianth. 
 
 On the whole, there seem good reasons for believing that the 
 ancestral Salix flower was bisexual. 
 
 * In Engler and Prantl's Pflanzenfamilien. 
 
VOL. 2] Mott. Teratology in Calif ornian Willows, 207 
 
 ARE THE FLORAL GLANDS VESTIGES OF AN ANCESTRAL PERIANTH ? 
 
 The glands of Salix have been held by several writers to be 
 vestigial perianth of a once bisexual flower. Velanovsky (1904), 
 especially, has recently worked out a ground plan for a com- 
 plete flower, basing his contentions as to perianth on some tera- 
 tological material of Salix aurita L. The flowers were bisexual, 
 and the posterior gland throughout, according to his interpre- 
 tation, was divided into two somewhat leaf-like bodies, which, 
 further, had been moved around so as to stand transverse. He 
 then restores a lost posterior gland to match the anterior one, 
 and has an ideal dimerous perianth, as in Myrica and Juglans, 
 which some consider as standing next in relationship to Salix. 
 
 The occurrence of four glands together as noted above, a 
 fact which apparently has not been recorded before in Salix. 
 would seem to make possible a much simpler restoration of the 
 same perianth. The four glands might be vestiges of four peri- 
 anth phyllomes, in the original positions of those phyllomes. 
 Then to get the modern normal flower, with its posterior gland 
 and sometimes also anterior gland, we have only to conceive 
 that the other glands, or phyllomes, have been entirely sup- 
 pressed. In the reversion to hermaphrodite flower in our hybrid, 
 there seems to be a tendency to restore these lost glands. 
 
 SUPPOSING THE ANCESTRAL FLOWER WERE BISEXUAL, WAS THE 
 NUMBER OF STAMENS TWO OR FOUR? ARE THE TWO CARPELS OF 
 
 THE PISTIL NOW MEDIAN OR TRANSVERSE. Velanovsky arguing 
 for a once perfect flower, concludes that it had four stamens. For 
 since the two carpels of the pistil stand transverse in Salix. and 
 since the pair of stamens also in the perfect flowers of his mate- 
 rial stand transverse (as in ours), to obtain a dimerous flower 
 with regular alternation of parts, he restores a median pair of 
 stamens and thus has four. However, since the actually occur- 
 ring perfect flowers on our hybrids give many indications of 
 reversion to the original type, and these flowers have two stamens 
 each, perhaps we could as well assume, to satisfy alternation, 
 that the two stamens were once median and were crowded around, 
 to stand transverse, by the closely appressed subtending bracts 
 
208 University of California Publications. [BOTANY 
 
 of the ament. Such displacement in general is not uncommon. 
 Yet, again, is it necessary that alternation be satisfied? It is 
 frequently lacking in these ancient types. Regularity in number 
 and in alternation of parts is a development of the modern 
 flower. It would seem that we can as well hold that the original 
 number of stamens was two. 
 
 Pax,* arguing for a once perfect flower, gives a ground plan 
 based on the case of abnormality reported by Heinricher (1883), 
 and shows in it only two stamens. However, he represents the 
 carpels as standing median, and thus there is alternation with 
 only the two stamens. 
 
 Yet to consider now whether the carpels are median or 
 transverse this median position given by Heinricher seems 
 wrong. He had but a single flower, and does not support his 
 statement as to the position of the carpels by any evidence. The 
 stipe of his pistil, too, is shown to be abnormally long, and in 
 such cases it is difficult to make sure of the position of the car- 
 pels. The formation of a perfectly normal pistil from two trans- 
 verse stamens, as seen in our 8. lasiolepis and, conversely, the 
 median rupturing of a pistil to form two transverse stamens, 
 seem safely to be taken as evidence (as noted by Wichura as 
 long ago as 1887), that the carpels of the pistil stand transverse. 
 This is the position accepted for the normal female flower by 
 Pax, by Sargent, and by others, in descriptions of the genus 
 (though Brendel [1880] appears to think the median position 
 generally held). 
 
 Hegelmaierf is led to believe in the transverse position of 
 the carpels by a study of the development of the pistil in the 
 bud. On the other hand, an examination of the winter flower 
 buds of Salix lasiolepis seems not to bear out Hegelmaier's view, 
 however much other evidence does. The pistil appears as a 
 small medianly compressed, urn-shaped protuberance, bilabiate 
 at the apex, with the lips anterior and posterior seeming to cor- 
 respond to carpel apices. There are even two slight lateral 
 
 * In Engler and Prantl's Pfianzenfamilien, III, 1, p. 31. 
 
 f tiber Bliitenentwickelungen bei den Salicineen. Jahreshef te des Ve- 
 reins fur Vaterlandischer Naturkunde in Wiirttemberg. Stuttgart, 1880. 
 XXXVI, pp. 204-244, Tafn. Ill, u. IV. 
 
VOL. 2] Mott Teratology in Calif ornian Willows. 209 
 
 grooves, appearing to indicate lateral sutures, and here are only 
 very slight indentations in the middle of each lip. This sug- 
 gests strongly that the carpels are anterior and posterior (me- 
 dian), and that the valves of the capsule, which are median, 
 correspond, therefore, to the carpels. However, such a series 
 of forms connecting a pair of transverse stamens and a normal 
 pistil, as shown in the drawings here of Salix lasiolepis, and 
 also the converse phenomenon, as indicated in the figure from 
 the hybrid, would seem, even though the evidence is terato- 
 logical, to make the contention for the transverse position of the 
 carpels prevail. 
 
 This view is strengthened by the median position of the two 
 placentae, since ovules are commonly, though by no means 
 always, marginal on the carpels. On the other hand, the trans- 
 verse dehiscence of the capsule in Salix might, by itself, lead 
 one to conclude that the sutures are transverse and hence the 
 carpels median for dehiscence of unilocular capsules is per- 
 haps more commonly at the sutures than at the midribs; but 
 in some such capsules, as in the tricarpellary fruit of Viola,* 
 the splitting does occur at the midribs, and in septate capsules, 
 dehiscence is quite commonly loculicidal. 
 
 The balance of evidence seems to show that the carpels in 
 Salix stand transverse ; and, yet, at the same time, the necessity 
 for assuming more than two stamens in the ancestral flower seems 
 questionable. 
 
 Some Possible Causes of the Abnormalities Described. 
 
 Any attempt to discover the ultimate cause of such abnorm- 
 alities as found in the two willows here described would doubt- 
 less entail an explanation of all the manifold teratological phe- 
 nomena that occur throughout the plant kingdom, and involve 
 one, as noted at the outset, in still wider biological problems. 
 The proximate cause in the case of the Salix lasiandra X baby- 
 lonica, at least (if we confine ourselves to consideration of the 
 fairly well established causes of teratology), seems to be the 
 
 * Sachs, Text Book of Botany. 
 
210 University of California Publications. [BOTANY 
 
 crossing of species. Considerable evidence points to this. Wil- 
 lows in general are very prone to this intermingling ("Wichura 
 by repeated crossing obtaining a hybrid combining no less than 
 six species). A large proportion of the abnormal cases reported 
 for Salix are of hybrids. In general, " malformations and odd 
 forms are apt to appear in hybrids, especially in the flowers;"* 
 "monstrous or abnormal forms of floral organs are much more 
 common in hybrids than in individuals of pure descent, ' 'f Mil- 
 lardet, in extended experiments with strawberries, found that 
 many hybrid descendants bore perfect flowers instead of uni- 
 sexual with the male and female on different plants. 
 
 It seems quite possible that many of the cases of abnormal 
 willows reported under a simple specific name would prove, on 
 careful investigation, to be hybrids, as happened with our 8. 
 lasiandra X babylonica. This might be true of our 8. lasiolepis, 
 though it appeared pure. In general, it is true, hybrids are 
 more or less intermediate in characters between the parents, yet 
 it is not uncommon 1 1 for the offspring to resemble one parent 
 exclusively. Or, in all outward characters, as of habit, foliage, 
 and fruit, offspring may resemble one parent, while in hardi- 
 ness, vigor, resistance to disease, fertility, etc., they may follow 
 the other. In such cases there would still be a mingling of blood, 
 as it were, to cause disturbance internally in the adjustment of 
 vital forces that had been reached in each species, and such dis- 
 turbance might easily lead to manifestation in morphological 
 abnormalities. 
 
 Sometimes teratological phenomena seem to be due to change 
 in nutrition, rich or unusually abundant food, cultivation, or 
 other change of normal conditions which occasions such a physio- 
 
 * Bailey, Plant Breeding, p. 231. 
 
 t Focke, quoted by Swingle and Webber, Hybrids and their Utilization 
 in Plant Breeding. Yearbook of the U. S. Dept. of Agric. for 1897, p. 410. 
 
 ; L. c., p. 398. 
 
 The native willows of the region, aside from Salix lasiolepis, which 
 might possibly have given rise to this tree by crossing are S. lasiandra 
 Benth., S. laevigata Bebb, and S. Nuttalli Sarg. var. brachystachys. Sarg. 
 A few introduced species, like S. babylonica, have been planted to some 
 extent in the locality. In general appearance, however, this abnormal tree 
 was identical with many individuals of S. lasiolepis with which it was grow- 
 ing. All these trees were naturally planted. 
 
 II L. 6:, p. 396. 
 
VOL. 2] Mott. Teratology in Calif ornian Willows. 211 
 
 logical disturbance as suggested above. In these two instances 
 of the willow, however, there is no indication of any such excit- 
 ing cause, for trees growing near by, under identical conditions, 
 are perfectly normal. 
 
 The author of one of the articles on Salix, writing also of 
 Carex (Hampe, 1840), thought that his observations showed 
 that dry conditions tend to produce change of male organs to 
 female, and moist conditions the reserve. But, as shown (by 
 Von Schlectendal) in a discussion which follows the article, this 
 explanation does not always hold. Our 8. lasiolepis, in fact 
 though changing to female, was on a small creek and none of 
 the other adjacent trees, in the same conditions, showed the 
 peculiarity. 
 
 Insects are doubtless responsible for some malformations, as, 
 for example, the willow-roses and galls. Their sting, or punctur- 
 ing of a growing shoot to deposit an egg, introduces foreign 
 chemicals, either directly with the egg, or in the course of devel- 
 opment of the larva, into the tissues of the plant; and this 
 causes a disturbance of the normal stimuli of growth. Para- 
 sitic fungi, too, give rise to disorder by chemical means and 
 they have been known to cause the production of bisexual flowers 
 where unisexual are normal.* 
 
 In the case of our S.'lasiandra X babylonica, however, there 
 was no notable evidence of insect action ; and in the case of the 
 8. lasiolepis, though there was considerable breeding of insects 
 on the tree, still adjacent trees were quite as much affected, and 
 they were normal. 
 
 Hybridizing here seems to be the most likely explanation of 
 both abnormalities, the monstrous organs and the perfect flowers. 
 
 There may be some elucidation, if not explanation, of these 
 and other teratological phenomena, in Sachs ' ' ' Stoff und Form ' ' 
 theory, or, perhaps better, hypothesis, first advanced in 1875, 
 and further developed and explained in 1892. f According to 
 his idea, the plant produces a special substance for each organ's 
 development out of the common organic compounds which go 
 
 * Strasburger : Biologisches Centralblatt, 20: 657. 1900. 
 t Sachs ' Gesammelte Abhandhmgen iiber Pflanzen-Physiologie. II, pp. 
 1159-1231. 
 
212 University of California Publications. [BOTANY 
 
 to form the mass of all the organs. For each organ there is a 
 different substance. The amount of this special substance may 
 be very minute in proportion to the total amount of tissue going 
 to form the organ. Under normal conditions each "special- 
 baustoff" is conducted to a particular part of a growing point 
 to produce the organ normal there, stem, root, leaf, stamen, 
 pistil, etc. If any unusual stimuli are introduced (as by cross- 
 ing, insects, change in nutrition), the attunement of forces in 
 the plant is disturbed, and there may result a misdirection of 
 these special substances a "misleitung der Specialbaustoffe "- 
 and pistil-substance, for example, will go to developing tissue 
 which would normally have become stamens, and will cause the 
 production of a pistil there; or stamen-substance will be sent 
 astray, and will invade the embryonic tissue which could nor- 
 mally have developed into pistil, and cause the production of 
 stamens there, and so on. Or there may be a combination, in 
 various proportions, of two of these special substances, and the 
 result will be, for example, a dual organ partaking of the char- 
 acters of both pistil and stamen, the preponderance of the one 
 set of characters or of the other depending upon the proportion 
 of the special substances entering in. 
 
 Flowers are most likely to be the region of such disturbances 
 owing to the close association there of several organs of very 
 different nature. A very minute deviation of a special sub- 
 stance on its way to produce one of these organs is sufficient to 
 throw it into a position which is normally the sphere of action 
 of one of the other special substances. 
 
 If there is overproduction, for any reason, of any special 
 substance, then, under the theory, there results a reduplication, 
 or multiplication, of the appropriate organ. And thus, for ex- 
 ample, there come to be, in many species of Salix, a greater 
 number of stamens than the original number, which we have 
 thought to be two. The theory could further be used, it would 
 seem, to account for the present occasional occurrence of perfect 
 flowers in Salix. Supposing, for the reason outlined above, the 
 flowers were once perfect, the plant then possessed the power, 
 and it was active to produce both the special stamen-substance 
 
ySv 
 
 \ 
 
 I UNIVERSITY | 
 
 VOL. 2] Mott. Teratology in Calif ornian Willows. 213 
 
 and the special pistil-substance. Then, owing to the rise of cer- 
 tain conditions, the power to produce either the one or the other 
 of these special substances became dormant, and the flowers be- 
 came unisexual, with male and female flowers on separate trees. 
 Finally, later conditions, as noted above, such as crossing, re- 
 awaken in certain individuals this dormant power, and there 
 occurs reversion to the bisexual form of flower. 
 
 The preceding discussion has been, of course, on the basis of 
 the long widely-accepted theories of descent and of reversion. It 
 is well, perhaps, finally, to recognize at least the more recent 
 theory of the origin of characters which is coming to be favor- 
 ably considered by many. This theory says, that if the willow, 
 which certainly has not been a complex organism since all time, 
 could once develop, for example, the power to produce perianth 
 and perfect flowers, it could lose that power completely, not 
 merely suffer its becoming dormant, and then develop it anew, 
 as at first, and the phenomenon could not be called reversion: 
 or it could be, that our willow has now developed these charac- 
 ters for the first time, and nothing is to be concluded as to 
 whether or not it ever possessed them before. And similarly 
 other abnormalities herein noted would be explained. 
 
 Summary. ,. 
 
 On a tree mainly staminate, of the diandrous Salix lasiolepis 
 Benth., abnormal flowers were produced which show a complete 
 series of forms in change of a normal pair of stamens into a 
 normal bicarpellary pistil (pis. 19, 20, figs. 2-13). 
 
 X given phyllome is constant in character. A pair usually 
 show -equal change (e.g., pi. 19, figs. 5 and 9), but may differ 
 somewhat (pi. 19, figs. 4 and 7). Through the series the form of 
 the phyllome varies from stamen to carpel in proportion (with 
 some latitude) as its pollen-sacs degenerate and ovules supersede 
 them. Pollen-sacs and ovules are produced from the same part 
 of the phyllome (pi. 19, figs. 4 and 8) ; connective and placental 
 regions seem homologous. While phyllomes are immature, an- 
 thers, when present are introrse (pi. 19, fig. 3) (though normally 
 in this species extrorse, pi. 19, fig. 1 A); for the phyllomes stand 
 
214 University of California Publications. [BOTANY 
 
 with their reproductive faces together as looking to the enclosing 
 of the ovules. Dehiscence of the imperfect, bisexual ovary is 
 sutural (pi. 19, figs. 7 and 8) ; phyllomes remain intact (standing 
 transverse). When pollen-sacs have disappeared, dehiscence (as 
 normally in Salix) is at the midribs, loculicidal (valves median) 
 (pi. 20, fig. 12). In the former case dehiscence is premature, 
 owing to swelling of the pollen-sacs and their twisting to face 
 (as normally) the honey-gland; in the latter case, it is at the 
 normal time. Exposed ovules do not become mature seeds. 
 
 A rudimentary ovule often causes an ovule-appearing lobe to 
 form in the wall of a pollen-sac developing around and over it 
 (d and e, pi. 19, fig. 5) ; or, developing normally, the ovule may 
 stand protruded through the pollen-sac (a, pi. 19, fig. 5). Ves- 
 tigial pollen-sacs often stand independent of each other, showing 
 the individuality of the pollen-sac as a microsporangium (d, pi. 
 19, fig. 7). Pollen-sac vestiges and lobes are distinguished from 
 ovules by their normal structural peculiarities, and often by 
 pollen-grains (pi. 20, figs. 17-19), and ovules are, in the same 
 way, identified by structural peculiarities (pi. 20, fig. 16). 
 
 Monstrous bisexual organs tend to make a catkin fall early ; 
 a very few normal pistils keep a catkin otherwise male or of 
 monstrous organs from falling until the seeds are ripe. 
 
 The hermaphroditism here seems to arise by sexual substitu- 
 tion in the stamens, either at the time of origin of the phyllomes, 
 or soon after. 
 
 A mainly pistillate, scion-propagated, hybrid willow (S. 
 lasiandra Benth. X babylonica L.) by staminody of the pistil, 
 also shows this sort of hermaphroditism (a, pi. 20, fig. 21). 
 This tree has, besides, many bisexual flowers with normal organs, 
 mostly pistil between two stamens (pi. 20, fig. 20). This her- 
 maphroditism seems to arise by reversion. 
 
 This tree has also purely male flowers, commonly of two or 
 three, but up to five, stamens, the parents being pent- and tri- 
 androus. Rarely a filament with double anther occurs, also 
 normal ovary and anther on one stalk. Stamens are in two 
 sets: One, mostly in male flowers, in anthesis with the pistils; 
 the other, common in perfect flowers, in anthesis later (pro- 
 
VOL. 2] Mott. Teratology in Calif ornian Willows. 215 
 
 togyny) (pi. 20, fig. 20). Numerous variations occur of the 
 types of flowers described, by different combinations, in one 
 flower, of the different normal and abnormal sex-organs. Cat- 
 kins are varied ; so also are branchlets. 
 
 Especially in the perfect flowers, there is a tendency towards 
 two transverse glands in addition to the normal one or two 
 median. 
 
 This tree has borne abnormal flowers three seasons; most in 
 the third. A neighboring also pistillate scion tree of the same 
 cross was apparently normal the first season, somewhat abnormal 
 the third. A third tree, apparently normal the first season, bore 
 one perfect flower the third. Some similarly abnormal, mainly 
 pistillate, naturally planted hybrids near by (with also trees 
 mainly staminate) seem to be the parent stock. Apparently 
 scions perpetuate the abnormalities. Some of the mainly pistil- 
 late naturally grown trees show also apparent reduplication of 
 phyllomes, e.g., in one flower, three unicarpellary pistils (some- 
 times hermaphrodite, sometimes not) , or two normal pistils. The 
 same trees bear also pistils in various stages of division into two 
 unicarpellary pistils. 
 
 A 8. lasiandra, otherwise normal, shows the tendency to four 
 glands (pi. 19, fig. 22). 
 
 Viewed speculatively, the abnormalities seem to indicate that 
 the ancestral Salix flower consisted of pistil and two stamens 
 with a four-part perianth, and that it has become unisexual by 
 suppression of the organs of one sex or the other; the series of 
 monstrous sex-organs apparently shows that the two carpels of 
 the normal pistil stand transverse, etc. \ 
 
 Of the usually recognized possible causes of teratological 
 phenomena, hybridization seems, here, to offer the most likely 
 explanation. 
 
216 University of California Publications. [BOTANY 
 
 BIBLIOGRAPHY. 
 
 Penzig's Pflanzen-Teratologie, Bd. II, contains an approxi- 
 mately complete list of references to the literature on teratology 
 in Salix, up to the time of publication of this work, in 1894, Of 
 the articles therein mentioned, those especially were searched for 
 in the original, and are here noted, which are accompanied by 
 illustrations. For the period since 1894, up to March, 1905, 
 this bibliography pretends to be practically complete for all arti- 
 cles, whether with or without figures. 
 
 * Anderson, C. L. 
 
 1890. A Monoecious Willow. Zoe, I, p. 41. 
 fAubert, S. 
 
 1894. In Arch. sc. phys. nat., Geneve 3, per., t. 31, p. 307. 
 Bail. 
 
 1877. Neuere Beobachtungen der Androgynie bei Salicineen. Schr. d. 
 Phys.-okon. Gesellscli. zu Konigsberg, 18 Jahrg., 1 Abth., pp. 
 94-5. 
 Brendel, F. 
 
 1880. Transformation of anthers into ovaries. American Naturalist, 
 XIV, p. 442. 
 
 Burkill, I. H. 
 
 1898. Changes in the Sex of Willows. Ann. of Bot., XII, p. 557. 
 Camus, E. G. 
 
 1899. Fleurs faussement hermaphrodites et anomalies florales dans le 
 
 genre Salix. Bull. Soc. Bot. de Fr., XLIV, pp. 185-91, pis. 
 
 IV-VI. 
 Eichelbaum. 
 
 1888. Note. Botan. CentralU., XXXV, p. 114. 
 Hampe. 
 
 1840. In Linnaea. XIV, pp. 367-8, followed by six pages by publisher, 
 
 Von Schlechtendal. 
 Haring, J. 
 
 1894. Abnorme Katzchenbildungen bei Salix caprea L. und bei Salix 
 cinerea L. (Ester. Bot. Zeitsch., Wien, XLIV, pp. 386-7, 
 415-18. 
 Hartman. 
 
 1841. Note. Flora, XXIV, p. 199. 
 
 * Not referred to by Penzig. 
 
 t Have not seen the original article. 
 
VOL. 2] Mott. Teratology in Calif ornian Willows. 217 
 
 Hegelmaier, F. 
 
 1887. Abnormitaten einiger einheimischen diclinen Pflanzen. Jahresb. d. 
 Ver. f. Naturk., Wiirttemb., XLIII, pp. 307-15, Taf. III. 
 
 Heinricher, E. 
 
 1883. Eine Zwitterbliithe von Salix Caprea L. Sitz-ber. d. K. ATcad d. 
 Wissensch., Wien ; 1 Abth., Bd. LXXXVII, pp. 129-31, Taf. 
 II, Fig. 10. 
 
 Kirschleger. 
 
 1841. Note. Flora, XXIV, Literaturberichte, Nro. 1, p. 59. 
 1845. Note. Flora, XXVIII, p. 402. 
 
 Leefe, J. E. 
 
 1841. Observations on some curious metamorphoses of the pistil in 
 Salix Capraea. Transact, of the Botan. Soc. of Edinburgh, 
 I, 2, pp. 113-114, pi. VI, figs. 16-21. 
 
 Lowe, J. 
 
 1856. On an abnormality in the flowers of Salix Andersoniana. Ann. 
 and Magaz. of Nat. Hist., pp. 254-5, 9 figs. 
 
 Masters, M. T. 
 
 1869. Vegetable Teratology. 
 
 MUller, H. 
 
 1868. In Botan. Zeitg., pp. 843-4. Tab. XIV, B. 
 
 Panek, J. 
 
 1891. Weiden und Weidenbastarde aus der Umgebung von Hohenstadt 
 in Mahren. (Ester. Bot. Zeitsch., Wien, XLIV, 381-5. 
 
 Sauter. 
 
 1837. Note. Flora, XX, Beibldtter ler, Bd., p. 40. 
 
 Schimper. 
 
 1829. In Flora, XII, pp. 422-3. 
 
 Schnitzlein. 
 
 1850. Note. Bot. Zeitg. VIII, p. 746, Taf. VIII, Fig. 2-7. 
 
 Shimek, B. 
 
 1895. Perfect flowers of Salix amygdaloides Ands. Proc. Iowa Ac. Sc., 
 Ill, pp. 89-90. 2 figs. 
 
 Tausch. 
 
 1833. Note. Flora, XVI, p. 230. 
 
 Velanovsky, J. 
 
 1904. Vergleichende Studien iiber die Salix-Bliite. Beihefte zum Bot. 
 Centralbl., XVII, pp. 123-8. Taf. 2. 
 
218 University of California Publications. [BOTANY 
 
 Von Schlechtendal. (See Hampe.) 
 
 Von Seemen, O. 
 
 1886. Einiges iiber abnorme Bliitenbildungen bei den Weiden. Ver- 
 
 handlungen des Botan. Ver. der Prov. Brandenburg, XXVIII, 
 pp. 1-14, Taf. I. (48 small figs.). 
 
 1895. Abnorme Bliitenbildungen bei einer Salix fragilis L. (Ester. Bot. 
 Zeitsch., Wien, XLV, pp. 254-7, 289-95. Tafn. XII, u. 
 XIII. 
 
 Wigand, A. 
 
 1887. Beitrage zur Pflanzen-Teratologie. Botanische Hefte, II, Mar- 
 
 burg, p. 106 and pp. 121-22. 
 
VOL. 2] Mott. Teratology in Calif ornian Willows. 219 
 
 EXPLANATION OF PLATES 19 AND 20. 
 
 All figures (except figs. 16-19) show the posterior side of the flower, 
 i.e., the side which faces the axis of the ament. Bract, honey-gland, and 
 lower part of united filaments have been removed in most cases. 
 
 All figures (excepts figs. 1 A and 16-19) circa 10. Figs. 7 and 22 
 somewhat larger. 
 
 FIG. 1-19 SALIX LASIOLEPIS BENTH. 
 
 Figs. 1-15 show a series of pairs of sexual phyllomes varying in form 
 from two stamens to a bicarpellary pistil. Each pair is constant in form, 
 from the bud, but the series are described as though each member repre- 
 sented a temporary stage in a progressive change from stamens to pistil 
 taking place in a single flower. Fig. 1 from normal male individual; 
 figs. 14 and 15 from normal female; figs. 2 and 13 from abnormal male. 
 Fig. 1. Normal male flower (from specimen No. 5049 in the Herbarium 
 of the University, collected at Willow Camp, Marin Co., Calif.) 
 
 a, anther-lobe, composed of two pollen-sacs (microsporangia) 
 which dehisce by a common slit above the fused wall be- 
 tween the pollen-sacs. 
 
 &, slit for escape of pollen. 
 
 c, cylindrical filaments united to about the middle. 
 
 d, upper limit of normal union of filaments. 
 
 e, hairy bract of ament, subtending the flower. 
 
 f, honey-gland at base of filament. 
 
 Fig. IA. Young diandrous flower of Salix purpurea L., with filaments 
 united and anthers extrorse. 
 a.a.j anthers. 
 Z>, united filaments. 
 
 c, hairy bract. 
 
 d, honey gland. 
 
 Fig. 2. Beginning of transformation. 
 
 a, grooves in filament-forks, preparatory to formation of 
 ovarian cavity. Forks still free and of normal length, 
 and entire flower otherwise normal. The barren por- 
 tions of the forks are finally reduced, the connectives 
 developing to form the ovary while the filaments shorten 
 and complete their union to form the stipe. 
 
220 University of California Publications. [BOTANY 
 
 Fig. 3. Beginning of union of grooved filament forks. 
 a, anterior margins, united. 
 6, fc, posterior margins, free. 
 
 c, c, filaments, widened slightly medianly and with more evi- 
 
 dent grooves than in fig. 2. 
 
 d, beginning of ovarian cavity; no ovules yet. 
 
 e, connective, almost normal. 
 
 f, f, anthers, normal, but introrse and twisting to face honey- 
 
 gland as they swell in maturing. 
 
 Fig. 4. First appearance of ovules. (Normal number on a placenta is 
 six or eight, hence normal number on a phyllome margin is three or 
 four, since two margins unite to form a placenta). 
 
 a, a, ovules, marginal and close to base of anther-lobes. That 
 these bodies are ovules beyond doubt is shown in the 
 enlarged drawing, fig. 16. 
 &, b, developing marginal placenta. 
 
 c, anther, still normally developed. 
 
 d, stamen, normal except for groove; no ovules. 
 
 Fig. 5. Ovules (some well developed, some abortive) more numerous, and 
 on both phyllomes. The ovules are so closely associated with the 
 pollen-sacs that some of the ovules cause malformations in the pollen- 
 sacs, and two ovules even protrude through the pollen-sac tissue. 
 
 a, ovule, borne on a connective, and protruding through the 
 wall-tissue of pollen-sac (*), and carrying some of the 
 adjacent tissue outward with it as it develops. 
 6, wall-tissue of pollen-sac (i) adhering to base of ovule (a). 
 
 c, ovule protruding through pollen-sac tissue similarly to 
 
 ovule (a). 
 
 d, lobe in wall-tissue of pollen-sac (j), resembling an ovule 
 
 in form and probably caused by rudimentary ovule on 
 connective beneath. (Ovule fails to develop). Iden- 
 tity of tissue of lobe determined by its possession of the 
 same peculiar characters which mark the normal pollen- 
 sac. (Fig. 17). 
 
 e, e, e, lobes in pollen-sac wall similar to (d), but less promi- 
 
 nent. 
 
 f, f, slits between pollen-sacs at time of dehiscence. 
 
 g, g, ovules, borne free from anthers. 
 
 h, h, h, h, vascular strands of the developing placentae now 
 evident as thickenings on the margins of the phyllomes. 
 Margins still ununited. 
 
VOL. 2] Mott. Teratology in Calif ornian Willows. 221 
 
 Fig. 6. Broadening of the connectives to form ovary; and beginning of 
 style on one phyllome the broadening extends into the still elongated 
 forks. (The ovules are usually not distant from the anthers as 
 here). 
 
 a, a, broadened connectives and forks. 
 6, &, ovules. 
 
 c, beginning of apical development of connective to form 
 barren upper part of ovary and style and stigma. 
 Connective of the other stamen is not yet so developed. 
 
 Fig. 7. Increase of carpellary character of one phyllome, and degeneration 
 of its anther. Filament fork of this phyllome is now unelongated, 
 and the connective is shown to be the chief seat of reproductive 
 activity. 
 
 a, b, c, vestiges of the two pollen-sacs of the left anther-lobe. 
 Eight anther-lobe reduced to one small sac (d). Yet 
 (a), (b) and (c) all contain normal pollen, 
 a, left pollen-sac, still well developed, but largely independent 
 
 of the right. 
 &, right pollen-sac, not well developed. 
 
 c, lobe of (&), probably, similar to (d), fig. 5. 
 
 d, vestigial pollen-sac representing right anther-lobe. The 
 
 other pollen-sac has entirely failed to develop. (Figs. 
 17-19 show this body to be a pollen-sac and not an 
 ovule, though it is about as minute as an ovule, and is 
 easily mistaken for one). 
 
 e, e, ovules near base of anther-lobe, as before. 
 
 f, f, four ovules, occupying former position of pollen-sacs (this 
 
 seeming to indicate that connective and placental re- 
 gions are homologous). 
 
 g, filament-fork unelongated, owing to decadence in develop- 
 
 ment of pollen-sacs of this phyllome. 
 
 h, style, being apical development of connective (style is 
 curled downward; phyllome at this stage soon withers). 
 
 h 1 , upper barren portion of ovary-wall, this also being de- 
 veloped from the connective (corresponds to (&), 
 fig. 11). 
 
 i, stigma, first clear definition. 
 
 j, ruptured suture. 
 
 ~k, anther, still well developed. 
 
 I, three ovules. 
 
 m, filament fork, grooved, but still about normally elongated, 
 
 the male character of the phyllome predominating. 
 f n, honey-gland. 
 
 o, bract. 
 
222 University of California Publications. [BOTANY 
 
 Fig. 8. Ovarian cavity now well developed, though the swelling of the 
 enclosed pollen-sacs causes premature dehiscence, and dehiscence is 
 at the sutures instead of at the midribs. (The pollen-sacs are usually 
 more degenerate at this stage. The ovules should be shown more 
 elongated, as possessing polarity). 
 
 a, suture between phyllomes, remaining intact. 
 
 b, b, suture ruptured. 
 
 c, c, ovary-wall, from later and apical development of the 
 
 two connectives. 
 
 d, d, styles and stigmas, not yet united as in the normal 
 
 pistil; they are imperfectly developed. 
 
 e, old crotch of filament forks, corresponding to (d), fig. 1. 
 
 f, f, filament forks, both unelongated. 
 
 g, g, pollen-sacs, dehisced. 
 
 Fig. 9. A closed ovary, though still an imperfect one, has been evolved. 
 Anthers practically lost. 
 
 a, ovary, not yet tapering above. 
 
 b, b, styles, malformed and not yet united, and stigmas not yet 
 
 clearly differentiated. 
 
 c, stipe, still abnormally long. 
 
 d, d, phyllomes, formerly staminal, now predominantly car- 
 
 pellary. They still show clearly that they stood right 
 and left, transverse. 
 
 e, posterior suture between phyllomes (carpels). 
 
 f, honey-gland. 
 
 g, bract. 
 
 Fig. 10. Transition to normal pistil almost complete; ovary swollen below 
 and tapering above, style single, pollen-sacs have disappeared (fig. 
 11). Yet stipe is still abnormally long, and stigma has lobes right 
 and left, corresponding to phyllomes, or carpels. 
 
 a, swollen lower portion of ovary containing ovules. 
 
 b, tapering barren upper portion of ovary, to accommodate 
 
 seed-hairs. 
 
 c, single style, well defined. 
 
 d, demarcation of style from ovary as in fully normal pistil 
 
 (d, fig. 14). 
 
 e, e, stigma-lobes, being the ununited tips of the carpels. 
 
 f, posterior suture (placentiferous below) between carpels 
 
 (corresponds to (e), fig. 9). Same on anterior wall. 
 
 g, stipe, still abnormally long. 
 h, bract. 
 
 i, honey-gland. 
 
VOL. 2] Mott. Teratology in Calif ornian Willoivs. 223 
 
 Fig. 11. Same specimen as shown in fig. 10, but with posterior half of 
 ovary-wall removed to show total lack of development of pollen-sacs, 
 and typical position of ovules (median and on lower half of wall) ; 
 the number of ovules is normal. 
 
 a, swollen lower portion of ovary containing ovules. 
 &, tapering barren uppe-r portion of ovary, to accommodate 
 seed-hairs. 
 
 c, single style, well defined. 
 
 d, demarcation of style from ovary as in fully normal pistil 
 
 (d, fig. 14). 
 
 e, e, stigma-lobes, being the ununited tips of the carpels. 
 
 f, posterior suture (placentiferous below) between carpels 
 
 (corresponds to (e), fig. 9). Same on anterior wall. 
 
 g, stipe, still abnormally long. 
 h, bract. 
 
 i, honey-gland. 
 
 j, ovules on anterior placenta formed by union of phyllome 
 margins (ovules are marginal here as shown to be the 
 case in preceding figures, as fig. 7, (f). 
 
 Fig. 12. Dehisced capsule, showing completion of change of two stamens 
 into a bicarpellary pistil. (Style and stigmas fallen.) Dehiscence 
 is loculicidal (at the midribs) for valves are median while carpellary 
 phyllomes are transverse. In the monstrous forms, separation occurs 
 between the transverse phyllomes and each remains intact. The 
 series shows that the carpels in the normal flower in this genus stand 
 transverse. 
 
 a, normal stipe. 
 
 6, ovary, swollen with seeds and seed-hairs. 
 
 c, seed-hairs. 
 
 d, posterior placenta, at suture between carpels (corresponds 
 
 to /, fig. 10). 
 -* e, e, midribs of carpels, along upper part of which dehiscence 
 
 occurs (dehiscence loculicidal). 
 f,f, valves, standing median (anterior and posterior). 
 
 Fig. 13. Anterior view of capsule in fig. 12, to illustrate typical position 
 of placenta. 
 
 a, anterior valve. 
 
 ft, placenta (seeds removed), along middle of lower half of 
 valve. 
 
 c, lower limit of natural dehiscence. 
 
 d, line of fracture in tearing away posterior valve. 
 
 e, e, fragments of posterior valve. 
 
224 University of California Publications. [BOTANY 
 
 Fig. 14. Normal female flower from typical tree of Salix lasiolepis Benth, 
 (Strawberry Creek, Grounds of the University of California. Fall 
 catkin. September, 1904). Posterior view as in preceding figures. 
 a, swollen lower portion of ovary, containing ovules. 
 &, tapering barren upper portion of ovary to accommodate 
 seed-hairs. 
 
 c, single style, well defined. 
 
 d, demarcation of style from ovary. 
 
 e, stigma lobed transversely and only slightly lobed medianly. 
 /, posterior suture (placentiferous below) between the carpel 
 
 appearing as a seam in the wall. Same on anterior 
 
 wall. 
 
 g, stipe, short. 
 In, bract. 
 i, honey-gland. 
 
 Fig. 15. Partly side-view of pistil in fig. 14, to show slight groove at 
 midrib of carpel, along upper part of which dehiscence occurs, 
 a, groove. 
 &, transverse groove in stigma. 
 
 Fig. 16. Section of ovule (macrosporangium), optical longitudinal, median, 
 through raphe, to establish identity of ovules in figs. 4-7. (Oblong 
 form shows polarity. Vestigial, pollen-sacs and the pollen-sac lobes 
 are more likely to be isodiametric, though in general appearance and 
 size they are easily mistaken for ovules). 
 
 a, nucellus. 
 
 &, inner integument. 
 
 c, outer integument. 
 
 d, basal micropyle (ovule anatropus). 
 
 e, raphe, 
 
 /, vascular bundle in raphe. 
 g, chalaza. 
 h, funiculus. 
 t, placenta. 
 
 Fig. 17. Two cells, one from each layer of wall of pollen-sac in fig. 19, 
 enlarged, to show difference in character of layers which results in 
 dehiscence when wall dries. 
 
 a, thin-walled cell (shown in section) of outer layer, marked 
 
 (6) in fig. 19. 
 ~b, "fibrous-cell" of inner layer, marked (c) in fig. 19. 
 
VOL. 2] Mott. Teratology in Calif ornian Willows. 225 
 
 Fig. 18. Pollen-grain (microspore) enlarged from vestigial pollen-sac, 
 (d} fig. 7. 
 
 Fig. 19. Vestigial pollen-sac (microsporangium), (d) fig. 7, enlarged, to 
 show identifying characters. (View into the sac from side attached 
 to connective.) 
 
 a, line of rupture in tearing sac from connective. 
 Z>, layer of thin-walled cells forming outer part of sac-wall. 
 c, layer of so-called ' ' fibrous-cells ' ' forming inner part of 
 sac-wall. (Unequal contraction of (6) and (c) in dry- 
 ing causes dehiscence; (&) contracts the more.) 
 dj pollen-grains (microspores). Most of the pollen has fallen 
 out. 
 
 FIGS. 20, 21. SALIX LASIANDEA BENTH .X BABYLONICA L. 
 
 Fig. 20. Bisexual flower, parts normally formed; predominating type of 
 flower on the tree. Glands often four, the two not shown being 
 usually smaller, and transverse. 
 
 a, pistil of two transverse carpels. 
 
 6, suture between carpels, placentiferous below. 
 
 c, c, stamens, transverse; only two, though five normal for 
 
 S. lasiandra (in this locality) in staminate flower and 
 three normal for S. ~babylonica. 
 
 d, filament, not yet elongated (flower protogynous) . 
 
 e, posterior honey-gland. 
 
 f, anterior honey-gland. 
 
 g, bract, subtending flower. 
 
 Fig. 21. Bisexual flower with hermaphrodite organ in place of pistil ; 
 one of the numerous variations on this tree of the typical flower, 
 fig. 20. It shows one stage of change of pistil to two stamens; 
 and verifies evidence shown by S. lasiolepis series that carpels of 
 normal pistil in this genus stand transverse. 
 
 a, monstrous hermaphrodite organ. 
 
 b, 1), phyllomes, transverse. 
 
 c, c, anther-lobes of two pollen-sacs, each somewhat malformed 
 
 and misplaced yet producing and discharging pollen. 
 
 d, vestigial anther-lobe. 
 
 e, two ovules, on anterior sutural placenta. 
 
 f, posterior suture of monstrous ovary, prematurely ruptured 
 
 by swelling of abnormally enclosed anthers. 
 
 g, g, stamens, transverse, with filaments not yet elongated. 
 h, h, honey-glands, median. 
 
226 University of California Publications. [BOTANY 
 
 FIG. 22 (PL. 19). SALIX LASIANDEA BENTH. 
 
 Fig. 22. Staminate flower showing four glands, possibly vestiges of 
 4-part perianth. Only the posterior gland is commonly present in 
 this species; tree otherwise normal. (The five stamens and the bract 
 have been cut off.) 
 
 a, posterior gland. 
 
 &, anterior gland. 
 
 c, c, lateral glands. 
 
 d, d, bases of five stamens. 
 
 e, bract. 
 
 Of THE 
 
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UNIV. CALIF. PUB . B OT. VOL . 2 
 
 [MOTT] PLATE 19 
 
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UNIV. CALIF. PUB . B OT. VOL . 2 
 
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