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TRANSACTIONS
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WISCONSIN ACADEMY
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VOL. XX.
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NATURAE SPECIES RATIOGUE
MADISON, WISCONSIN
1921
i
The Fiftieth Anniversary volume of the Transactions of the
Wisconsin Academy of Sciences, Arts, and Letters is issued
under the editorial supervision of Charles E. Allen and the
Secretary.
Arthur Beatty,
Secretary.
TRANSACTIONS
OF THE
WISCONSIN ACADEMY
OF
SCIENCES, ARTS AND LETTERS
VOL. XX.
MADISON, WISCONSIN
1921
CONTENTS.
Page
Ordo Prophetarum. Karl Young . 1
Richard Wagner’s “Die Meistersinger von Niirnberg” and
its Literary Precursors. Edwin C. Roedder . 83
The Passing of a Historic Highway. F. E. Williams . 131
/ The Arsenical Solutionsj^No. 1. Liquor Potassii Arsenitis
(Fowler’s Solution). H. A. Langenhan . 141
\/ ''The Development of the Frenulum of the Wax Moth* Gal¬
leria mellonella Linn. (With Plate I.). William S.
Marshall . 199
/ The American Water Mites of the Genus Neumania.l\ (With
Plates II-IV.) Ruth Marshall . 205
\/ Syrphidae of Wisconsin.' (With Plates V-VI.) Charles
L. Fluke . 215
v Some Experiments with the Larva of the Bee Moth,' Galleria
mellonella L. Joy E. Andrews . 255
vThe Length of Life of the Larva of the Wax Moth, Galleria
mellonella L., in its different Stadia. (With one
Figure.) Ruth Chase . 263
v A Preliminary Study of the Digestive Secretions of Pickerel
and Perch. Caroline Walker Munro . 269
The Organization of the Nuclei in the Root Tips of Podo¬
phyllum Peltatum.\\ (With Plate VII.) James Bert¬
ram Overton . 275
v The Phytoplankton of the Muskoka Region, Ontario, (Canada.
(With Plates VIII-XIII.) Gilbert Morgan Smith. . 323
v Species of Lentinus in the Region of the Great Lakes.
(With Plates NIV-XXVIII.) Edward T. Harper.. 365
y Cytological Studies of the Lower Basidiomycetes ’ I.
Dacrymyces. (With Plate XXIX.) E. M. Gilbert
(One Figure) . 387
Notes on Parasitic Fungi in Wisconsin, VII. (With Plates
XXX-XXXII, and one Figure.) J. J. Davis . 389
Page
Notes on Parasitic Fungi in Wisconsin, VIII. (With three
Figures.) J. J. Davis. . . . 413
Experimental Work on the Relation of Soil Temperature to
Disease in Plants. (With Plates XXXIII-XXXVII,
and four Figures.) L. R. Jones . 433
Quantitative Studies of the Bottom Fauna in the Deeper
Water of Lake Mendota. (With one Figure.) Chan-
cey Juday . 461
A Survey of the Larger Aquatic Plants of Lake Mendota.
(With one Figure). R. H. Denniston . 495
A Quantitative Study of the Larger Aquatic Plants of Lake
Mendota. (With Plate XXXVIII.) H. W. Rickett 501
Notes on the Chemical Composition of Some of the Larger
Aquatic Plants of Lake Mendota. I. Cladophora and
Myriophyllum. Henry A. Schuette and Alice E.
Hoffman . 529
A Second Report on Limnological Apparatus. (With
Plates XXXIX-XL, and five Figures.) Edward A.
Birge . 533
The Rotifers of Wisconsin. (With Plates XLI-LXI, and
four Figures.) H. K. Harring and F. J. Myers . 553
Thure Kumlien (With Plates LXII-LXIV.) Publius V.
Lawson . 663
Proceedings of the Academy, 1919 and 1920 . 687
Programme of the Exercises in Commemoration of the
Fiftieth Anniversary of the Academy . 693
The Academy Medallion, 187CL1920 (With Plate LXV)
Edward A. Birge . 711
List of Officers and Members . 717
Charter of the Academy . 751
Constitution of the Academy . 754
General Index of Volumes I-XX . 759
ORDO PROPHETARUM
KARL YOUNG
As has often been observed by recent writers on the subject,1 the
liturgical plays of the Christmas season may be divided into the
following groups: (1) a play of the shepherds (Officium Pas-
torum), performed on Christmas Day;2 (2) a play of the Magi
(Officium Stellce), for Epiphany;3 (3) a play representing the
Slaughter of the Innocents (Or do Rachelis), for performance on
Epiphany or on Innocents Day (December 28) ;4 * 6 and (4) a play of
the prophets (Or do Prophet arum) , associated with Christmas Day,
or with the octave of Christmas.
In undertaking a special study of the Ordo Prophetarum 5 1 wish,
in both introduction and conclusion, to pay homage to M. Marius
Sepet, whose brilliant monograph Les Prophetes du Christ 6 not
1 See, for example, E. K. Chambers, The Mediaeval Stage , Vol. II, Oxford.
1903, pp. 41-56 ; or the present writer, Ordo Rachelis (University of Wisconsin
Studies in Language and Literature , No. 4, Madison, 1919), p. 3.
2 The most extensive treatment of this play, so far as I know, is a study by
the present writer entitled Officium Pastorum: A Study of the Dramatic De¬
velopments within the Liturgy of Christmas, in Transactions of the Wisconsin
Academy of Sciences, Arts, and Letters, Vol. XVII, Part I (1912), pp. 299-396.
3 The most thorough study of this play is by H. Anz, Die lateinischen Magier-
spiele, Leipzig, 1905.
4 This play has recently been treated by the present writer in a special study
entitled Ordo Rachelis, mentioned above.
6 For general purposes, I use the designation found at the head of the version
from Laon. See below, p. 40.
6 Sepet’s monograph appeared first in serial form in Bibliotheque de Vficole des
Chartes (Vols. XXVIII, 1867, pp. 1-27, 211-264 ; XXIX, 1868, pp. 105-139,
261-293; XXXVIII, 1877, pp. 397-443), and then as a separate volume, Les
Prophetes du Christ, Paris, 1878. I make my references throughout to the
separate volume. E. Mart&ne (Tractatus de Antiqua Ecclesioe Disciplina, Lyons,
1706, pp. 78, 106-107) was familiar with the phenomenon of the Ordo Prophet¬
arum, but he presents no full text of it, and does not trace its origin. E. Du-
M§ril (Les Origines Latines du TM&tre Moderne, Caen, 1849, p. 180), in editing
the Limoges version of the Ordo Prophetarum, bases upon a passage in Durandus
the following remark: “II [the Limoges version] avait aussi son origine dans
9
Wisconsin Academy of Sciences , Arts, and Letters.
only first disclosed the essential nature of this liturgical play, but
also described the early stages of its genesis in general terms that
may still be accepted. The discovery of new examples and new
facts, however, has rendered Sepet’s demonstration noticeably in¬
complete, and calls for a fresh survey of the phenomena. Upon
such a survey I now venture, calling attention to the fact that the
present study differs from that of Sepet in several particulars.
Whereas, in the first place, Sepet considered the prophet-play both
in its relations to the liturgy and in its later developments in the
vernaculars, I confine myself to the Ordo Prophetarum as a strictly
liturgical play. Secondly I re-edit from the manuscripts all the
dramatic texts known to me. Sepet presented none of these texts
in extenso, and with one important version he was not acquainted.7
Finally, from manuscripts and printed books, I offer materials that
are, I believe, brought into this consideration for the first time.
Within its limited field, then, I hope that my study may be found
relatively complete.
I
Among apocryphal works -of St. Augustine is found a sermon of
substantial length entitled Contra Judceos , Paganos, e’t Arianos
Sermo de Symbolo.* 1 Although the Augustinian authorship has
been generally discredited in modern times,2 the attribution to the
la liturgie catholique.” L. Petit de Julleville (Les. My sieves, Vol. I, Paris, 1880,
pp. 35-45) bases his account of the prophet-play frankly upon Sepet. P. Weber
( G-eistliches Schauspiel und kirchliche Kunst, Stuttgart, 1894, pp. 41-48) surveys
Sepet’s work with approval, and adds certain contributions of detail. W. Meyer
( Fragmenta Bur ana, Berlin, 1901, pp. 50-56) accepts Sepet’s demonstration of
the origin of the Ordo Prophetarum and adds important suggestions ; but upon
later parts of Sepet’s study he passes certain strictures, for which he is appro¬
priately rebuked by W. Creizenach ( Liter at urblatt fur germanische und roman-
ische Philologie, Vol. XXIII, 1902, col. 203.) Chambers (Vol. II, pp. 52-55) pays
explicit homage to Sepet, and follows him avowedly ; but he includes a consid¬
eration of the Laon play, which was unknown to the French savant, and adds
significant observations upon details. The one or two minor contributions of
Chasles (Les Prophetes du Christ, in La Vie et les Arts Liturgiques, 3e Annee,
No. 25, January, 1917, pp. 121-134) are mentioned in appropriate places below.
7 1 refer to the version from Laon. See below pp. 39—49.
1 This sermon is found in the Benedictine edition of St. Augustine’s works,
Sancti Aurelii Augustini . . . Opera, Vol. VIII, Paris, 1688, Appendix,
col. 11-20. It is reprinted by Migne, Pat. Lat., Vol. XLII, col. 1117-1130.
2 The Benedictine editors (loc. cit., col. 11-12) cite the dictionis barbaries and
the multa . . . delibata ex Augustino. See Weber, p. 41 ; Sepet, p. 8, note 2.
Young — Or do Prophet arum.
3
great bishop persisted throughout the middle ages.3 The sermon
was written manifestly during the period in which the Arian heresy
throve; hence, before the year 600.4 Since, moreover, the author
seems to have made use of a translation of the Old Testament older
than the Yulgate, we may place the date of composition before the
middle of the fifth century.5
Of the twenty-two chapters, or sections, into which the discourse
is divided in the Benedictine edition, the first six are addressed
to the Faithful by way of an exhortation to resist the devil and to
cling to their Christian faith, to be worthy of their baptism, and
to believe in the Trinity.6 In Sections seven to ten the preacher
arraigns the heretics and brings before them the relation of the Son
to the Father, and Christ’s birth from the Virgin.7 Section ten as¬
sails Herod for his cruel designs against Jesus, and passes into a
denunciation of the Jews as a whole. Sections eleven to eighteen,
addressed to the Jews, attempt primarily to convict them of error
through utterances of their own prophets and of certain Gentiles.
The preacher summons the prophets and Gentiles singly by
name, and delivers their successive testimonies.8 Section nineteen
inveighs once more against the Arians.9 The last three sections are
an appeal to the Faithful to prepare for the eternal life.10
3 For example, in Ms. Vatic. Regin. 125, a version of the passage beginning
Vos} inquam, convenio (fol. 74r-76v, written at Forcalquier in the thirteenth
century) bears the following heading: Sermo beati Augustini Episcopi de
Natale Domini. See also below, pp. 5, 18.
4 See Weber, p. 41.
5 See Meyer, p. 50. Chambers assumes (Vol. II, p. 52) that it was “probably
written in the sixth century.” In regard to date and provenience the Benedictine
editors (loc. cit., col. 11—12) write, “Iste quoque Sermo sub Arianorum dominatu
habitus est, eo tempore quo haereticis illis disputantibus, & catholicos, alios pol-
licitatione ac praemio, alios vi potentiaque deducere a fide nitentibus obsistere
nullus audebat.” Hauck (as reported by Weber, p. 41) infers as follows:
“Der Sermo gehort entweder nach Gallien unter die Westgotenherrschaft, Oder
nach Afrika unter die Vandalenherrschaft.”
6 The headings of the first six sections, given by the Benedictines in side-votes,
are as follows : Caput I, Christianse vigiliae ; Caput II, Diabolus qui sit ; Caput
III, Renuntiare satanae, quid ; Caput IV, Relapsus a baptismo ; Caput V, Triadis
Unitas ; Caput VI, Trinitas incomprehensibilis. These headings may be merely
editorial.
7 The headings are as follows : Caput VII, Contra Arianos ; Caput VIII,
^Equalitas personarum ; Caput IX, Christi nativitas ; Caput X, Crudelitas Herodis.
8 The headings are these : Caput XI, Contra Judseos ; Caput XII, Ex Daniele ;
Caput XIII, Testimonia ex Lege & Prophetis ; Caput XIV, Simeonis & Zacharies
testimonium ; Caput XV, Ex libris Ethnicorum ; Caput XVI, De Sibyllinis vati-
ciniis ; Caput XVII (no heading) ; Caput XVIII, Ex eventis.
9 Caput XIX, Contra Arianos.
10 Caput XX, Resurrectio ; Caput XXI, Differentia temporalis vitae & aeternse ,*
Caput XXII, De via ad patriam.
4 Wisconsin Academy of Sciences , Arts , and Letters .
The outstanding rhetorical characteristic of the sermon is its
lively dramatic quality.11 Aside from the part in which the proph¬
ets are summoned to confute the Jews, direct address is employed
effectively. Section seven, for example, begins as follows :12
Die mihi, HaBretice, qui audes dicere quod minor sit Dei sapientia: die
mihl, altitudo cseli quanta sit, latitudo terrae, profundum abyssi?
Section ten ends with the following passage :13
Die, Herodes, si te Christus ideo offendit, quia ejus Magis annuntiantibus
audisti regnum, quid te offenderunt bi qui tecum adversus Christum
sentiebant principes Judaeorum, quorum filios interficiens ipsos magis
atrocissima poena in suis filiis afflixisti? Nam Christum minime invenire
potuisti. Sed quid tecum diutius agam? Ipsos, ipsos conveniam Judaeos,
qui dum infantem Christum noluerunt agnoscere, filios suos cum illo
compulsi sunt amittere: quos quidem occidit mortique propinavit Herodes
vester amicus; sed his mortalitatem aeternamque vitam donavit Christus,
quern vestrum etiam nunc dicitis inimicum.
It appears, then, that at several points the sermon contains dra¬
matic elements and verges upon a dramatic presentation of its
theme.
II
For our present purpose, however, we may dismiss considerations
bearing upon the sermon as a whole and may center our attention
upon the sections addressed to the Jews (Cap. xi-xviii). Almost
the whole of this part is frequently found during the middle ages
as a separate liturgical lectio for use in Matins of Christmas or of
some other day of the Christmas season. This liturgical use is
seen in the following text found in a lectionary of the twelfth
century from Arles i* 1
11 On the literary qualities see Sepet, p. 9.
12 Benedictine edition, Vol. VIII, col. 14.
13 This passage serves as a transition to the part beginning Vos, inquam, con -
venio, O Judcei, to be studied below.
1 Paris, Bibl. Nat., Ms. latin 1018, Lectionarium Arelatense ssec. xii, fol.
129r— 132v. This is the text printed by Sepet (pp. 3-8). My text differs from
his only in a few details, all of which are recorded in my foot-notes. I have
seen no cogent reason for collating the texts in Paris, Bibl. Nat., Ms. latin 16819
(Lectionarium Compendiense ssec. xi, fol. 12v-14r), Oxford, Bodleian Library,
Ms. Canonici Liturg. 391 (Lectionarium ssec. xii, fol. llv-13v), and Rome, Bibl.
Vatic., Ms. Vat. Regin. 125 (Lectionarium Porcalqueriense ssec. xiii, fol. 74r-76v),
Young — Or do Prophet arum.
5
Sermo Beati Augwstini Episcopi de Natale DomiNi
Lectio vi
“Vos, inquam, conuenio, o Iudei, qui usqwe in hodiernum diem negatis
Filium Dei. Nonne uox uesfra est ilia quando eum uidebatis miracula
facientem atq ue temptantes dicebatis: Quousqwe animas nostras sus-
pendis? Si tu es Xpisiws, die nobis palam. Ille autem uos ad cowsidera-
tionem mittebat miraculorum, dicens: Opera que ego facio ipsa testi¬
monium perhibent de me; ut Xpisio testimonium dicerent now verba, s ecZ
facta. Vos autem no n agnoscentes Saluatorem qwi operabatwr salutem
in medio uesire terre, adicientes in malo aistis: Tu de te ipso testi¬
monium dicis; testimonium tuum now est uerum. Sed ad hec ille quid
uobis respowderit, aduertere noluistis. Nowne scriptum est in lege uestra
qwod duorwm hominum testimonium uerum sit? Preuaricatores legis,
intendite legem. Testimonium queritis de XpisZo: in lege uestra scriptum
est qwod duorwm hominum testimonium uerum sit. Procedant ex lege now
tantum duo sect eciam plures testes Xpisti, et cowuincant auditores legis,
now factores.* 2
Die, Ysaya,3 testimonium Xptsto. Ecce, inquit,4 uirgo in utero con-
cipiet et pariet filium, et uocabitwr nomew eiws Hemanuhel, qwod est
interpretatum : Nobiscum Dews.5
Accedat et alius testis.5
Die et tu, Iheremia,6 testimonium Xpisfo. Hie est, inqwit, Dews noster,
et now <fol. 129v> estimabitur alius absqwe illo qui inuenit omnem uiam
scientie et dedit earn Iacop puero suo et7 Israel dilecto suo. Post hec in
terris uisus est, et cum hominibws cowuersatws est.8 Ecce duo testes
idonei ex lege uestra ad quorwm testimonia non suwt compuncta corda
uestra. Sect alii atqwe alii ex lege testes Xpisti introducantur, ut frontes
durissime inimicorwm conterantwr.9
all of which are before me as I write. In my foot-notes I indicate the marginal
entries and unusual rubrications in Mss. Vat. Regin. 125 and Canonici Liturg.
391. The headings of the texts are as follows: in Ms. 391, Sermo Beati Augustini
Episcopi; in Ms. 125, Sermo Beati Augustini Episcopi de Natale Domini; in Ms.
16919, none. These three manuscripts are not mentioned by Sepet. The manu¬
scripts containing the sermo under discussion must be numerous.
2 Followed by a vertical line indicating separation or paragraph division
(Ms. 1018).
3 Ysaya] In the right margin, opposite the line in which the name occurs, is
written the rubric: Ysayas (Mss. 1018 and 391). All the marginal entries in
Ms. 391 seem to me to be of the 15th century. In the left margin, a later hand
has written: i prophetia (Ms. 125).
4 Ecce inquit . . . nobiscum Deus] Encircled by red lines, with marginal
numeral: i (Ms. 125).
5 Followed by a paragraph sign (Ms. 1018).
6 Iheremia] Opposite this name in the margin: Iheremias (Ms. 1018);
Ieremias (Ms. 391).
7 dedit earn Jacop puero suo et] Written in the margin by a later hand
(Ms. 1018).
8 Hie est inquit . . . conuersatus est] encircled by red lines, with marginal
numeral: ii (Ms. 125).
9 Followed by a paragraph sign (Ms. 1018).
6
Wisconsin Academy of Sciences, Arts , and Letters.
Veniat et ille Danihel10 sanctus, iuuenis quidem etate, senior uero
scientia ac mawsuetudine, et conuincat omwes falsos testes et sic ut conuicit
seniores impudicos, ita suo testimonio Xpisti conterat inimicos. Die,
sancte Danihel, die de Xpisto qwod nosti. Cum uenerit, inquit, Sanctus
Sanctorum, cessabit unetio.* 11 Quare, illo presente, cui insultantes dice-
batis: Tu de te testimonium dicis, testimonium tuum no n est uerum;
cessauit unetio uestral Nisi quia ipse est qwi venerat Sanctus Sanctorum.
Si enim, sicw£ uos dicitis, now dum uenit, s ed expectatur ut ueniat Sanctus
Sanctorum, demonstrate unctionem: si autem, quod uerum est, cessauit
uestra, unetio, agnoscite uenisse Sanctum Sanctorum. Ipse est enim et
lapis ille abscisus12 de monte sine manibws concidentium, id est Xpistus
natus de uirgine sine manibws complectentium, qui tantum creuit ut mons
magnus fleret, et impleret uniuersam faciem terre. De quo monte dicit
propheia: Venite, ascendamws in montem Domini; et de quo Dauid dicit:
Mons Dei, mons uber, ut quid suspicamini montes incaseatos, mowtem in
quo placuit Deo habitare in ipso. Cum enim ipse Dominws Xpistus dis-
cipwlos suos interrogaret quern dicerewt esse homines Filium hominis,
responderunt: Alii Helyam, alii Iheremiam aut unum ex prophets; et
ille, vt quid suspicamini montes incaseatos, montem in quo placuit Deo
habitare in eo, hunc cognouit Petrus dlcens: Tu es Xpistus, Filius Dei.
Agnouit montem et ascendit in montem; testimonium dixii ueritati, et
dilectus est a ueritate. Super petram fundatus est Petrus, ut montem
susciperet ilium amando quern ter negauerat timendo.13
Die et Moyses,14 legislator, dux popwli Israel, testimonium Xpisto.
Prophetam, inquit, uobis suscitabit Deus de fratribws uesJris; omwis awima
que now audierit prophetam ilium, exterminabitur de popwlo suo.15 Pro¬
phetam autem dictum Xpistum ipswm audi in Buangelio dicentem: Non
est,16 inquit, propheta sine honore, nisi in patria sua.17
Accedat autem Dauid18 sawetws, testis fidelis, ex cuiws semine processit
ipse <fol. 130r> cui lex et prophete testimonium dieuwt, dicat et ipse de
Xpisto. Adorabunt, inquit, eum omwes reges terre, omwes gentes seruient
illi.19 Cui seruiewt? die, cui seruient? Vis audire cui? Dixit Dominws
Domino meo: Sede ad dexteram meam, donee ponam inimicos tuos scabel-
hxm pedum tuorwm.20 Et expressiws atqwe nominatim: Quare, inquit,
10 Danihel] Opposite this name in the margin: Daniel (Ms. 391).
11 Cun uenerit . . . cessabit unetio] Encircled by red lines, with marginal
numeral: iii (Ms. 125).
12 abscisus] abcisus (Sepet).
13 Followed by a paragraph sign (Ms. 1018).
14 Moyses] Opposite this name in the margin: Moyses (Ms. 391).
15 Prophetiam inquit . . . de populo suo] Encircled by red lines, with il¬
legible marginal numeral (Ms. 125).
16 Non est . . . patria sua] Encirled by red lines, without marginal numeral
(Ms. 125).
17 Followed by a paragraph sign (Ms. 1018).
18 Dauid] opposite this name in the margin: Dauid (Ms. 391).
19 Adorabunt . . . seruient illi] Encircled by red lines, with marginal
numeral: v (Ms. (125).
20 Dixit Dominus . . . pedum tuorum] Encircled by red lines, without
marginal numeral (Ms. 125).
Young— Or do Prophet arum.
7
tumultuate sunt gentes, et popwli meditati sunt inania? Astiteruwt reges
terre et principes comienerunt in unum aduersus Dominwm et aduersws
Xpistum. ei us.21
Accedat et alius testis. Die et tu, Abacuch22 prophefa, testimonium de
Xpisto. Domine, inquit, audiui auditum tuum et timui; consideraui opera
tua et expaui.23 Que opera Dei iste mirato expauit? Numqsud fabricam
rnundi iste miratus expauit? Absi]t. S ed, audi, aliquid expauit. In
medio, inquit, duum awimalium cognosceris. Opera tua, Deus, Uerbmw,
caro factum est. In medio duum animalium cognosceris. Qui quousqwe
descendisti, expauescere me fecisti; Uerbum, per quod facta sunt omnia,
in presepe iacuisti. Agnouit bos possessorem suum et asinus presepe
Domini sui. In medio duum ammalium cognosceris. Quid est in medio
duum ammalium cognosceris? nisi aut in medio duorwm testamentor^m,
aut in medio duorwm latronum, aut in medio Moysi24 et Helye cum eo in
monte sermocinantium. Ambulauit, inquit, Uerbum et exiuit in campis.
Verbum caro factum est et habitauit in nobis. Hoc et Iheremias ait:
Post hec in terris uisus est, et cum hominibns conuersatus est.25 Ecce
quemadmodum sibi conueniunt testes Ueritatis, ecce quemadmodum conuin-
cunt filios falsitatis. SufRciunt uobis ista, o Iudei, an adhuc ad uesiram
confusionem ex lege et ex gente uesfra alios introducemus testes ut illi
testimonium perhibeant cui perdita mente insultantes dicebatis: Tu de
te ipso testimonium dicis, testimonium26 tuum non est verum? Quod si
uelim ex lege et ex prophetis omnia que de Xpisto dicta sunt colligere,
facilius me tempns quam copia deseret.27
Verumptamen senem23 ilium ex gente uesira natum, s ed in errore uestro
non relictum. Symeonem sanctum in medio introducam, qui meruit teneri
decrepitus in hac luce quousqne uideret lucem. Quern quidem iam etas
compellebat ire, s ed expectabat suscipere quern sciebat uenire; cum iste
senex admonitns esset a Spiriiu sancto quod non an te moreretur quam
videret Xpisfnm Dei natum, quern cognoscens perrexit ad templum.
<fol. 130v> Vbi uero eum portari matris manibns uidit et diuinam in-
fantiam pia senectus agnouit, tulit infantem in manibns suis. Ille qui¬
dem Xpis£nm <infantem>29 ferebat, s ed Xpistus senem regebat. Regebat
qui portabatur ne ille ante promissum a corpore solueretnr. Quid tamen
dixerit, quern tamen confessus fuerit aduertite inimici, non Xpisti, s ed
21 Followed by a paragraph sign (Ms. 1018).
22 Abacuch] Opposite this name in the margin: Abacuc (Ms. 391).
23 Domine . . . et expaui] Encircled by red lines, with marginal numeral:
vi (Ms. 125).
24 Moysi] Moyse (Sepet). The last letter of this word is blurred in the
manuscript. I print it as it appears to me.
25 Post hec . . . conuersatus est] Encircled by red lines, with marginal
numeral: vii (Ms. 125).
26 dicis testimonium] Supplied in the right margin by a later hand (Ms. 1018).
27 Followed by a paragraph sign (Ms. 1018).
28 senem] Opposite this word in the margin: Symeon (Ms. 391).
29 Following Sepet, I supply the word infantem from the Benedictine edition
of the entire sermon Contra Judceos, mentioned above. The word is not present
in Mss. 1018, 125, 16819, and 391.
8
Wisconsin Academy of Sciences, Arts , and Letters.
ues£ri. Benedicts Dominion exclamauit senex ille et dixit: Nunc dimit-
tis,30 Domine, seruum tuum in pace, quia uideruwt oc%li mei salutare
tuum.31
Illi eciam parentes Ioh<mnis, Zacharias et Elisabeth,32 iuuenes steriles,
in senecta fecuwdi, dicawt eciam ipsi testimonium Xpisto; dicant de Xpisto
quid sentiant et testem idoneum Xpisto nutriant. Aiunt enim suo paruulo
nato: Tu puer propheta Altissimi uocaberis, preibis enim ante faciem
Domini parare uiam eius.33 Ipsiqne matri et uirgini, Helisabeth ait:
Vnde mihi hoc ut ueniat mater Domini mei ad me? Ecce enim ut facta
est uox salutationis tue in auribns meis, exultauit in gaudio infans in
utero meo.31 Intelligens enim Iohannes35 matrem Domini sui uenisse ad
suam matrem inter ipsas angustias uteri adhuc po situs, motu salutauit
quern uoce non poterat. Qui postea ipse Iohannes precursor et amicns,
humiUimns et fidelissimns seruus, testis fidelis idoneus effectns, tanto
maior inter natos mulierum quanto existimabatur esse qnod non erat.
Xpistum enim eum esse Iudei credebant, s ed ille non se esse clamabat
dicens: Quern me suspicamini esse, non sum ego. Sed ecce uenit post
me cuins pedum non sum dignus soluere corrigiam calciamenti.33 O fidelis
testis et amice ueri sponsi, quanto te humiliauisses si ad corrigiam cal-
ciamenti eins soluendam dignum te esse, dixisses! Sed dum ad hoc non
te dignum dicis, ludeis falsis testibns contradicis. Et hec a te dicta sunt
antequam Xpistnm videres, qui cum ad te ipse uenit excelsus humilis,
implende dispensationis sue gratia, ut a te baptizaretur qui nullum habebat
omnino peccainm, quid responderis, quern cognoueris, qnod testimonium
protuleris audiant inimici qui audire nolunt. Ecce, inquit, agnus Dei,
ecce qni tollit peccaia mundi. Et adiecit: Tu ad me uenis baptizari.
Ego a te debeo baptizari.37 Agnouit seruus dominion, agnouit uincnlis
<fol. 131r> originalis peccofi obligates ab omni nexu peccaii liberum.8'
Agnouit preco iudicem, agnouit ereatura creatorem, agnouit paranimphus
sponsum. Nam et hec uox Ioh&nnis est: Qui habet sponsam sponsus est,
amicus autem sponsi qui stat et audit eum gaudio gaudet propter uocem
sponsi.39
30 Nunc dimittis . . • . salutare tuum] Encircled by red lines, with marginal
numeral: vlii, written twice (Ms. 125).
31 Followed by a paragraph sign (Ms. 1018).
32 Zacharias et Elisabeth] Opposite these names in the margin : Zacharias,
Helisabeth (Ms. 391).
83 Tu puer . . . uiam eius] Encircled by red lines, with marginal numeral :
ix (Ms. 125).
34 Vnde mihi ... in utero meo] Encircled by red lines, with marginal
numeral : x (Ms. 125).
35 Iohannes] Opposite this name in the margin: Iohannes (Ms. 391).
36 Quern me suspicamini . . . corrigiam calciamenti] Encircled by red
lines, with marginal numeral: xi (Ms. 125).
87 Ecce inquit agnus . . . debeo baptizari] Encircled by red lines, with¬
out marginal numeral (Ms. 125).
38 liberum] obligatum (Sepet).
39 Followed by a paragraph sign (Ms. 1018).
Young — Ordo Prophetarum.
9
Sufficiunt uobis ista, O Iudei, sufficiimt uubis tanti testes, tot testimonia
ex lege uesfra et ex gente uesfra? An adhuc impudentia nimia audebitis
dicere quod alterias gentis uel nationis homines Xpisto deberewt testi¬
monium perhibere? S ed si hoc dicitis, respondet quide?n ille uobis:
Non sum missus nisi ad oues que perierunt dom us Israel. S ed, sic ut uos
in Actibus aposfoloram increpat Paulus, uobis primum oportuerat an-
nuntiare Uerbum Dei, sed quia repulistis ilhtd nec uos dignos uite eterne
iudicastis: Ecce, inquit, cotiuertimzts nos ad gentes. Demonstremws
eciam nos ex gentibas testimonium Xpisto fuisse prolatum, quoniam
Ueritas no n tacuit clamando eciam per linguas inimicoram suorum.
Nonne quando ille poeta40 facutidissimi&s inter sua carmina:
lam nova progenies403 celo demittit^r alto,41
dicebat, Xpisto testimonium perhibebat? In dubium hoc ueniat nisi alios
ex gentibws idoneos testes pluraqwe dicentes in medio introducam.42
Ilium, ilium43 regem qui uesiram superbiam captiuando perdomuit,
Nabuchodonosor, regem scilicet Babilonis, no n pretermittam. Die, Na-
buchodonosor, quid in fornace uidisti quando tres uiros iustos iniuste
illuc miseras, die, die quid tibi fuerit reuelatum. Nonne, inquit, tres
uiros misinms in fornace ligatos? Et aiunt ei: Vere, rex. Ecce, inquit,
ego uideo quatuor uiros solutos deambulantes in medio ignis, et corruptio
nulla est in eis, et aspectus quarti simiUs est Filio Dei.44 Alienigena,
unde tibi hoc? Quis tibi annunciauit Filium Dei? Que lex? Quis pro-
phefa tibi annuntiauit Filium Dei? Nondum quidem mundo nascitur et
similitudo nascentis a te cognoscitur. Vnde tibi hoc? Quis tibi istwd
annuntiauit nisi quia sic te diuinus ignis i ntus illuminauit ut cum illic
apud te captiui tenerentur inimici Iudei, sic diceres testimonium Filio
Dei? S ed quia in ore duorttm uel trium testium stat omne Uerbum,
sic ut ipse Bominus uesiram contumaciam confutans: <fol. 131v> in
lege, inquit, uesira scriptum est quod duorum hominum testimonium
uerum sit; etiam ex gentibns45 tercius testis introducatur ut testimonium
ueritatis ex onmi parte roboretur.46
Quid Sibilla47 uaticinando etiam de Xpisto clamauerit in medium pro-
feramns ut ex uno lapide utrorztmqwe frontes percuciantur, Iudeornm
scilicet ntque Paganornm SL+que suo gladio, sic ut Golias, Xpisti omwes per¬
cuciantur inimici. Audite quod dixerit:
"poeta] Opposite this word in the margin: Virgilius (Ms. 391).
40a progenies] proienies (Ms. 1018).
41 lam noua . . . dimittitur alto] Encircled by red lines, without marginal
numeral (Ms. 125).
"Followed by a paragraph sign (Ms. 1018).
"ilium] Opposite this word in the margin: Nabuchodonosor (Ms. 391).
44 Ecce inquit ego . . . Filio Dei] Encircled by red lines, with marginal
numeral: xii (Ms. 125).
45 ex gentibus] exigentibus (Sepet).
46 Followed by a paragraph sign.
47 Sibilla] Opposite this word in the margin: Sibilla (Ms. 391).
10
Wisconsin Academy of Sciences , Arts, and Letters.
4SJvdicii sign um: tellus sudore madescet;
E celo rex adueniet per secla futurus,
<S>cilicet in carne presens ut iudicet orbem,
Ynde Deum cement incredulus atq ue fidelis
Celsum49 cum sanctis, euiB0 iam termino in ipso
Sic anime cum carne aderunt quas iudicat ipse,
Cum iacet incultus densis in uepribus orbis.
Reicient simulacra uiri cunctam quoq ue gazam,
Exuret terras ignis, pontumque polumque: <fol. 132r>
Inquirens tetri portas effringet Auerni;
Sanctorum sed enim cuncte lux libera carni,
Tradentur sontes, eterna61 flamma cremabit.
Occultos actus retegens, tunc quisque loquetur
Secreta, atq ue Deus reserabit pectora luci..63
Tunc erit et luctus, stridebunt dentibus omnes.
Eripitur solis iubar, et chorus interit astris,
Soluetur celum lunaris splendor obibit,
Deiciet colles, ualles extollet ab imo:
Non erit in rebus hominum sublime uel altum.
Turn equantur campis montes, et cerula ponti <fol. 132v>
Omnia cessabunt, tellus confracta peribit:
Sic pariter fontes torrentur fluminaq ue igni.
Et tuba tunc sonitum tristem demittet ab alto
Orbe, gemens facinus miserum, uariosque labores,
Tarthareumque chaos monstrabit terra de<h>i<s>cens
Et coram hie domino reges sistentur ad unum:
Decidet e celo ignisque et sulphuris amnis.
Hec de Xpisti natiuitate, passione, et resurrectione atq ue secuudo ei us
adueutu ita dicta sunt ut si quis in greco capita horum uersuum discer-
nere uoluerit inueniet: Ihesus Xpisfus, Yos Theu, Soter; quod in latino
ita interpretatur: Ihesus Xp istus, films Dei, Saluator; quod et in latinum
translatis eisdem uersibus, apparet, pret er hoc quod grecarum litterarum
proprietas nou adeo potuit obseruari. Credo iam uos, o inimici Iudei,
tantis testibus ita53 obrutos confutatosque esse ipsa ueritate ut nichil
ultra repugnare, nichil querere debeatis.64
In view of the importance of this lectio in the dramatic develop¬
ment under consideration, we must examine its content in detail.
48 The verses of the Sibyl (Judicii signum . . . sulphuris amnis) are fur¬
nished with musical notation in Ms. 1018, and the first two verses are thus
furnished in Ms. 125.
49 Celsum] Selsum (Ms. 1018).
60 eui] cui (Sepet) ; eui (Ms. 125); euis (Ms. 391); aeui (Ms. 16819).
61 eterna] eternaque (Sepet).
62 luci] lucis (Ms. 1018).
63 ita] Omitted (Sepet).
S4 Followed immediately by the rubric : Lectio Sancti Euangelii secundum
Lucam (Ms. 1018).
Young~Ordo Prophet arum.
11
But before we undertake this examination we may pause to observe
certain variations of content in the extant versions, and certain
facts concerning its position in the liturgy.
In some churches the lectio was substantially abbreviated. Thus
in a French breviary of the thirteenth century55 it appears as
Lectio vi of Christmas, but is reduced by almost one-half in length.
The lesson retains only the introduction, the prophecies of Isaiah,
Jeremiah, and Daniel, and the concluding prophecy of the Sibyl.
The shortest form that I have observed is the following from a
breviary of the fourteenth century from Saintes, used as the sixth
lectio of Christmas Matins :56
Lee tio via
Vos, inquam, conuenio, o Iudei, qui usqwe in hodiernum diem negastis
Filium Dei. Nowne uestra uox est ilia quando eum uidebatis miracula
facientem atqwe temptantes dicebatis: Quousqwe awimas nostras tollis?
Si tu es Xpistuc, die nobis palam. Ille autem ad consideracionem miracu-
lorwm uos mittebat dicews: Opera que ego facio ipsa testimonium peri-
bent de me, ut Xpisfo testimonium dicerewt non uerba set facta. Vos
astern now agnoscentes Saluatorem qui operabatur salutem in medio
uestre terre, adiciewfes maledixistis: Tu de te ipso testimowium peribes;
testimonium tuum now est uerum. Sed ad hec ille quod testimowium
uobis responderit auertere noluistis: Nowne scriptum est in lege quia
qwod duorwm hominum <tdstimonium> uerum sit? Preuaricatores legis,
intewdite legem. Testimonium queritis de Xpisto. In lege uestra scrip¬
tum est quod duorwm hominum testimonium uerum sit. Preuaricatores
legis intendite legem. Testimonium queritis de Xpisto: in lege uestra
scriptum est quo<d> duorwm hominum <testimonium> uerum sit.57
Procedant ex lege now tantum duo s ed eciam plures testes Xpisti cowuiw-
cant auditores legis et now faefores.
Die, Ysaya, testimonium de Xpisto. Ecce, inquit,58 uirgo in utero con-
cipiet et pariet filium, et uocabitur nomew eius Hemanuel, qwod est inter-
pretatum nobiscum Dews.
65 Paris, Bibliotheque Nationale, Ms. lat. 1255, Breviarium Gallicanum ssec.
xiii, fol 84r— 85r. Sepet (See p. 2, note 1) was familiar with this version. The
exact provenience of this manuscript I have not been able to determine. In
Catctlogus Codicum Manuscript orum Bibliothecae Regice, Vol III, Paris, 1744,
p. 99, the content of this manuscript is described as follows: Breviarium ad
usum monasterii cujusdam Gallicani, ordinis Sancti Benedicti. That this
breviary is not for monastic use, however, seems to be shown by the presence of
nine lectiones in Matins of Epiphany.
56 Paris, Bibl. Nat., Ms. lat. 16309, Breviarium Santonense saec. xiv, fol. 31r.
This version is now published for the first time. The same manuscript con¬
tains (fol. 40v-41v) another excerpt from the pseudo-Augustinian sermo , used
as the sixth lectio of Matins for the Circumcision. See below, note 77.
57 1 print the text as it stands in the manuscript, with its obvious repetition.
“inquit] inquid (Ms.).
12
Wisconsin Academy of Sciences, Arts, and Letters.
Accedat et alius testis. Die et tu, Iheremia, testimonium Xpisto. Hie
est <inquit> Deus noster, et non estimabitur alius absqne illo, qui inuenit
omnem uiam sciencie, et dedit earn Iacob puero suo et Israel dilecto suo.
Post hec in terris uisus est, et cum hominibns conuersatus est.
Ecce duo59 testes ex lege uestra ex quornm testimonio non snnt com-
puncta corda uestra.60
Since this version contains only the nsnal introduction and the
first two prophecies (Isaiah and Jeremiah), one might surmise that
other parts of the sermo supplied one or more of the succeeding
lectiones of Matins at Saintes ;61 hut such is not the case.
Somewhat different from the versions previously cited is the
following Lectio ix for Christmas Matins in a breviary of the four¬
teenth century from Carcassonne :62
Lectio ix
Inter pressuras atqne angustias presentis temporis in nostre seruitutis
officia eogimur, Dilectissimi, non tacere cum pocius expediat flere magis
quam aliquid dicere. Uerumptamen ne quid minus lucri arche dominice
accedens dicatur nobis: Serue neqnam et piger tu erogares peccuniam
meam et ego ueniens cum usuris exigerem.63
Procedant ex lege non tantum duo s ed etiam plures testes Xpisti et
conuinean£ uos Iudeos auditores legis non autem facJores.
Die, Ysaia, testimonium Xpisto.
Ecce, inquit, uirgo concipiet et pariet filium et uocabitur nomen eius
Emanuhel, qztod interpretatur : Nobiscum Dens.
Accedat et alius testis. Die, Iberemias, testimonium de Xpisto.
Hie est, inqnit, Deus nosier et non estimabiinr alius absqne illo qni
inuenit omnem scienfiam et dedit earn Iacob puero suo et Israel dilecto
suo.
Quid enim Sibilla uaticinando de Xpisto clamauerit in medium profer-
amus. Audite quid dixerit:
59 duo] duos (Ms.).
60 The sixth responsory follows immediately : Responsorium : Sancti et in-
maculata uirginitas.
61 For excerpts from the sermo serving in a series of lectiones see the use of
Sarum, cited below, note 74.
62 Paris, Bibl. Nat., ;Ms. lat. 1035, Breviarium Carcassonense ssec. xiv,
fol. 3 Or— 31r. This text is now published for the first time. In the manuscript
it is immediately preceded by the following henedictio: Initium et finis, Alius
Marie uirginis eruat nos a peccatis et uitiis.
63 This introductory paragraph (Inter pressuras . . . exigerem) consists
of the opening sentences of the complete Contra Judaeos, Paganos, et Arianos
Sermo de Symbolo. See above, pp. 2 ff.
Young — Or do Prophetarum.
Versus Sibille
Iudicii84 signum: tellus sudore madescet;
E celo rex adueniet per secla futurus
Scilicet in carne presens \it iudicet orbem.
Post: Iudicii.
Unde Deum cement incredulus atq ue fidelis
Celsum cum sanctis, euis iam termino in ipso.
Posi : Iudicii
Sic anime cum carne aderunt, quas iudicat ipse
Cum iacei incultus densis in uepribns orbis.
Posi: Iudicii.
Deicient simulacra uiri, cunctam quoqne gazam;
Exure£ terras ignis, ponthumqne polumqne.
Pos£: Iudicii.
Inquirens tetri portas effringei Auerni;
Sanctorum sed erit cuncte lux libera carni.
Pos£: Iudicii.
Tradentur sontes, eterna flamma cremabit.
Occultos actus retegens tunc quisqne looretur.
Post : Iudicii.
Secreta atqne Dens reserabit pectora luci.
Tunc erit et luctus, stridebunt dentibns omnes.
Post: Iudicii.
Eripitur solis iubar, et corpus interit astris;
Soluetnr celum, lunaris splendor obibit.
Post : Iudicii.
Deicient col<fol. 30v>les, ualles extoilet ab ymo;
Non erit in rebus hominum sublime uel altum.
Post : Iudicii.
Iam eqnuntur camp is montes, et cerula ponthi;
Omnia cessabunt, tellus confracta peribit.
Post: Iudicii.
Hie pariter fontes torrentnr flamineq ue igni,
Et tubarnm sonitum tristem dimittet ab alto.
Pos* : Iudicii.
Orbe gemens facinus miserum uariosqne labores
Tartareumqne chaos monstrabit terra deiscens.
Post : Iudicii.
Et coram hie Domino reges sistentnr ad unum:
Reddet e celo ignisqne et sulphuris amnis.
Post: Iudicii signum.
13
64 Iudicii] Iuditii (Ms).
14
Wisconsin Academy of Sciences, Arts, and Letters.
Lectio Nong^5
Hec de XpisZi natiuitate, passione, et resurrectione, atq ue secundo eius
aduentu iam dicta, siunt ut, si quis in greco capita horum nersuum dis-
cernere uoluerit, inuemet: Ihesits XpisZuc, Yos Theu, Sother; quod in
latinum <translatis> eisdem versibus apparet pret er quod litteranm
grecarum proprietas no n adeo potuit obseruari.
Credo uos, o inimici ludei, tantis testibws obrutos cowfutatosqwe <esse>
ipsa ueritate uZ nichiZ ultra repugware, nichiZ querere debeatis. T<u
autem, Domine, miserere nobis>.®8
The peculiarity of this version lies not so much in the elimination
of the majority of the prophecies from the interior of the dis¬
course, as in the substitution for the usual introduction (Vos,
inquam . . . verum sit ) of the opening words (Int &r press uras
. . . usuris exigerem) of the Contra Judceos, Paganos, et Ari-
anos Sermo de Symbolo ,67
But aside from these differences in content, the sermon Vos
inquam shows variations also in liturgical position.68 We have al¬
ready observed its use as the sixth69 or ninth70 lectio in Matins of
Christmas day. On the same day it sometimes served as the fifth71
or eighth72 lectio. But it is found in Matins of other days as well.
It served as the fourth73 lectio, or as the fourth, fifth, and sixth
65 For the repetition of this rubric I have no explanation.
68 Followed immediately by the responsory Descendit de celis Deus.
67 See above, pp. 2 ff.
68 See Chambers, Vol. II, pp. 52, note 4; 53, note 2.
69 See also Bibl. Nat., Mss. latins 1234, Ordinarium Utinense ssec. xiv, fol. 2v ;
1273, Breviarium Andegavense ssec. xiv, fol. 37r ; 1279, Breviarium Monasterii
Tutelensis ssec. xiv, fol. 27r; 752, Breviarium Arelatense ssec. xiv, fol. 70v— 71r ;
1028, Breviarium Senonense ssec. xiv, fol. 51r. See further U. Chevalier, Insti¬
tutions liturgiques de VCglise de Marseille ( Bibliotheque liturgique, XIV), Paris,
1910, p. 23.
70 See also Bibl. Nat., Ms. 1309, Breviarium Gerundense ssec. xv, fol. 31r-31v.
, 71 That this was the use of Paris is shown by the following breviaries from
churches of that place: Bibl. Nat., Ms. lat. 1025, ssec. xiv, fol. 126v, 128r— 128?;
Bibl. Nat., Ms. lat. 746, ssec. xiv, fol. 30v-31r; Bibl. Nat., Ms. lat. 746 A., ssec. xiv,
fol. 23v ; Bibl. Nat., Ms. lat. 1293, anni 1471, fol. 34r-34v; Bibl. Nat., Ms. lat.
1294, anni 1472, fol. 37r; Bibl. Mazarine, Ms. 342, ssec. xiv, fol. 109r; Bibl
Mazarine, Ms. 345, ssec. xv, fol. 160v. See also Bibl. Nat., Ms. lat. 747, Lec-
tionarium Parisiense ssec. xiii-xiv, fol. 9v-10r. A similar use at the Abbey of
St. Victor is seen in Bibl. Mazarine, Ms. 347, ssec. xiv, fol. 89v-90r. See also
Bibl. Nat., Ms. lat. 12035, ssec. xii, fol. 22v-23r, — a breviary of provenience un¬
known to me.
72 Vatican Ms. 7126, Breviarium Monasticum ssec. xiii, fol. 64r-64v.
73 Munich, Hofbibliothek, Cod. lat. 12635, Breviarium Ranshofenense ssec. xiii,
p. 7.
Young — Or do Prophet arum.
15
lectiones74 on the fourth Sunday of Advent; as the fourth lectio
on Christmas eve;75 as all three lectiones in ferial Matins on any
day of the week preceding Christmas ;76 and as the sixth lectio on
the day of the Circumcision.77 I can cite no evidence of its use
outside the season of Christmas.
But with these variations in content and in liturgical position
we are concerned only incidentally, for these matters have no im¬
portant bearing upon the dramatic development under considera¬
tion. Let us, then, return to an examination of the complete and
normal form of the lectio as printed above from the lectionary of
Arles (Ms. 1018). 78
The lectio opens with a direct arraignment of the Jews for their
perverse disbelief in Christ’s Messiahship. Since the Jews stub¬
bornly demand evidence, the preacher grimly proposes to fetch testi¬
mony from their own law. He first summons Isaiah, bidding him
testify concerning Christ. As if receiving a response directly from
Isaiah in person, the preacher reports the prophet’s utterance.
Similarly are summoned the prophets Jeremiah, Daniel, Moses,
David, and Habbakuk. After each summons, the preacher reports
the prophet’s utterance, and adds a few words of elucidation.
With the taunt that he might readily extend this succession of
prophets from the Old Testament, the preacher passes on to the
premonitions regarding Christ in the New Testament, quoting ut¬
terances from Simeon, Zacharias, Elisabeth, and John. With a
74 Breviarium ad usum insignis Ecclesice Sarum, edited by F. Procter and
C. Wordsworth, Cambridge, 1882, pp. cxxxv-cxliii.
76 See E. Mart§ne, Tractatus de Antiqua Ecclesice Disciplina, Lyons, 1706,
p. 78.
76 At Laon. See U. Chevalier, Ordinaires de I’figlise cathedrale de Laon, Paris,
1897, pp. 33-43.
77 Bibl. Nat., Ms. lat. 16309, Breviarium Santonense saec. xiv, fol. 40v-41v.
In this case the liturgical lectio contains only the following parts of the sermon
Vos inquam: the opening sentences of the introduction, the prophecies of Virgil,
Nebuchadnezzar, and the Sibyl, and the conclusion. The excerpt from the
sermon Vos inquam found in the same manuscript (fol. 31r) for use as the sixth
lectio on Christmas has been printed above.
78 From the present study I omit all investigation of the sources of the Latin
lectio before us. The probability that the sermon Vos, inquam, convenio is in¬
fluenced *by Greek sermons of dramatic content is discussed in the able mono¬
graph of Giorgio La Piana, Le Rappresentazioni Sacre nella Letteratura Bizan-
tina, Grottaferrata, 1912, pp. 283-302, 308-309. Once given the sermon Vos,
inquam, convenio, the Western development of this composition into drama
appears to me to be independent of Byzantine influence. The opposing opinion
of La Piana in the case of the Ordo Prophetarum from Limoges, I consider
below, pp. 86-37.
16 Wisconsin Academy of Sciences, Arts, and Letters.
tart reminder to the Jews that the testimony already adduced should
be ample, the preacher adds utterances from the Gentiles, Virgil79
and Nebuchadnezzar, and finally80 a prophetic passage in hexa¬
meters from the Erythraean Sibyl.81 After elucidating the Chris¬
tian acrostic of the Sibylline verses, the speaker darts at the hostile
Jews a concluding shaft of scorn.
Viewed as a literary production the lectio is particularly note¬
worthy for its energetic dramatic style. The direct denunciations
of the Jews are presented in effective, though artificial, antitheses;
and the verbal playfulness of some passages, however strained it
may appear, contributes at least a certain vivacity.82 The monot¬
ony immanent in the formula of repeated summonses and responses
is somewhat mitigated through variety in the elucidating narratives
accompanying the responses. The circumstance that the prophets
are not summoned in chronological order does not seriously impair
the organization of the composition.83
79 For information concerning the famous line from Virgil, see J. B. Mayor,
W. W. Fowler, and R. S. Conway, Virgil’s Messianic Eclogue , London, 1907.
80 It will be observed that the last seven witnesses ( Simeon, Zacharias, Eliza¬
beth, John, Virgil, Nebuchadnezzar, and Sibyl) are not summoned after the
formula (Die, Ysaia; Die et tu, Iheremia ; Die, sancte Danihel ; etc.) employed
for the six preceding. The actual presence of Simeon, Zacharias, and Elizabeth,
John, Nebuchadnezzar, and the Sibyl may be understood as suggested in various
utterances of the preacher, such as the following : Symeonem . . . intro -
ducamj Illi etiam parentes Ioannis, Zacharias et Elisabeth, . . . dicant ; O
fidelis testis ; In dubium hoc ueniat, nisi alios ex gentibus idoneos testes pluraque
dicentes in medio introducam. I find in the text nothing that could possibly
imply the actual presence of Virgil.
81 These prophetic verses. are quoted by St. Augustine in his Be Civitate Bei
(Lib. xviii, cap. 23. See Migne, Pat. Ldt., Vol. XLI, col. 579—581). Their ap¬
pearance throughout the middle ages is frequent. See F. Neri, Le Tradizioni
italiene della Sibilla, in Studi Medievali (ed. Novati and Renier), Vol. IV
(1912-13), pp. 220-221; Notices et Extraits des Manuscrits de la Bibliotheque
Nationale, Vol. XXXII, Part II, Paris, 1888, pp. 101-102 ; F. Kampers, Bie
Sibylle von Tibur und Vergil, in Historisches Jahrbuch, Vol. XXIX (1908),
pp. 3, 244; Chevalier, Repertorium Hymnologicum, No. 9876 ; La Piana,
pp. 308-309.
82 See Sepet, p. 9.
83 See Petit de Julleville, Vol. I, p. 35.
Young — Ordo Prophet arum.
17
III
Still more important than the matter of dramatic style and
content, however, is the question of dramatic delivery. There can
be no doubt that the sermon before us, like many other mediaeval
homilies, contains material suitable for dialogue and impersona¬
tion, and hence for drama. We may well ask, therefore, how far
the homily in the form before us actually proceeded in the direc¬
tion of dramatic rendition.
That the lectio as it stands was not delivered as a dialogue seems
perfectly clear from the use of the word inquit in the responses.1
The use of this word seems to prove conclusively that the utter¬
ances of the persons summoned are delivered not by separate
speakers, but by the lector himself.2 The most that can be assumed
is that the lector altered his voice in such a way as to distinguish
between summons and response.3
Although, then, the lectio in use at Arles could not have been re¬
cited as dialogue, there is no reason why there should not exist other
versions in which the several prophecies were spoken by separate
persons. Such a version is courageously conjectured by Sepet as
follows :
J’en conlus de plus qu’on a fini, a un moment donnd, par completer les
flexions de voix en leur donnant pour organes des lecteurs differents, et
qu’alors chaque ton, c’est-a-dire cliaque prophete, a eu son interprete, le
dialogue demeurant cependant toujours enchassd dans le recit. Cette con¬
clusion repose sinon sur des preuves pdremptoires, au moins sur des in¬
ductions raisonnables. C’est une conjecture, mais c’est, je crois, une con¬
jecture vraisemblable.4
Although Sepet could find no text with which to support this
conjecture, and although he constructed it upon feeble data,5 it has
1 For example: Ecce, inquit, uirgo, etc.; Hie est, inquit, Deus noster, etc.
2 Early in his argument Sepet seems to affirm the presence of dialogue in the
lectio (“la reponse suivant immediatement la question et par consequent con-
stituant un dialogue,” p. 9) ; but he subsequently corrects himself (“ce qui nous
indique bien qu’il n’y avait pas de dialogue proprement dit,” p. 23). Chasles
sees clearly (p. 123) that the presence of the word inquit precludes dialogue.
3 See Sepet, pp. 13, 23. The varying of a single lector’s (or cantor’s) voice
in the rendering of the passiones of Holy Week is discussed at length in my
article Observations on the Origin of the Mediaeval Passion-Play, in Publica¬
tions of the Modern Language Association, Vol. XXV (1910), pp. 309-333.
4 Sepet, p. 13. See also Sepet, pp. 42, 96-97.
6 Sepet bases (pp. 10, 13) his conjecture, in the first place, upon the analogy
of the Passiones of Holy Week. He asserts (p. 10) that as early as the thir-
2 — S. A. Li.
18
Wisconsin Academy of Sciences, Arts , and Letters.
been welcomed by competent critics as conforming to general prob¬
ability.6 I am particularly glad, therefore, to present a version of
the lectio that provides for Sepet ’s conjecture a genuine justifica¬
tion. In a service-book from the church of Salerno we find the
following :7
In Nattvitatis Nocte post primam , Miss am legitur Sermo Sancti Augus-
tini Episcqpi, more Salernitano.
Vos (inquam) conuenio, O Iudsei, qui vsqwe in hodiernum diem negatis
filinm Dei. Nonne vox vestra est ilia, quando eum videbatis miracula
facientem, atq ue tentantes dicebatis: Quousqae animas nostras suspendis?
Si tu es Christus, die nobis palam. Ille autem vos ad eonsiderationem
miraculorum mittebat, dicens: Opera que ego facio, ipsa testimonium
perhibent de me; vt Christo testimonium dicere^t, non verba, sed facta.
Vos autem, non cognoscentes Saluatorem, adijeientes in main dixistis :
Tu de te testimonium dicis; testimonium tuum non est verum. Sed ad
hsec ille quid vobis respondent, aduertere noluistis. Nonne scriptum est
(inquit) in lege vestra quod duorum hominum testimonium verum sit?
Prseuaricatores legis <p. 76> intendite legem. Testimonium quaeritis de
Christo; in lege vestra scriptum est quod duorum hominum < testi¬
monium > verum sit. Procedant de lege non tantum duo sed etiam plures
testes Christi, & conuincant auditores legis, nec factores. Die & tu,
Isaias propheta, testimonium Christo?
teenth century the Passio was sung by three persons, to whom respectively were
assigned the parts of Christ, the Jews, and the Narrator ; and he raises the
question as to whether some such assignment of parts may not have been prac¬
ticed in the delivery of the lesson Vos, inquam, convenio . Such slight force as
this analogy may be supposed to have virtually disappears before the evidence
that this practice of distributing the roles of the Passio among three persons
did not obtain before the fifteenth century. See my Observations on the Origin
of the Mediaeval Passion-Play, already cited, p. 331. Another analogy cited by
Sepet (pp. 11-12) is that of the Gradual of the Mass for Palm Sunday as
rubricated in a certain manuscript of the twelfth century (Bibl. Nat., Ms. lat.
9486, fol. 9r). The several sentences of the Gradual are here distributed with
somewhat unusual dramatic appropriateness between Duo cantores and Chorus,
and this text as a whole, like the pseudo-Augustinian lectio, may be viewed as
a mingling of monologue and dialogue. Although the analogy between this
musical piece and the recited lectio is remote, it may be allowed to stand oy
whatever strength it has. I regard it as negligible. Sepet perceives (p. 13) a
further hint toward a distribution of the r61es of the lectio in the manuscript
itself (Bibl. Nat., Ms. lat. 1018) : that is, in the paragraph-signs and in the
entry of the names of two prophets at appropriate points in the margin (I have
recorded these rubrications in my foot-notes to the text of the lectio above,
along with the more thorough-going rubrications from Ms. Vatic. Regin. 125
and Ms. Canonici Liturg. 391). I cannot accept Sepet’s eager interpretation of
these rubrications. Although they are undoubtedly significant as indicating
divisions in the content of the lectio, they in no way prove that the several
prophecies were assigned to separate speakers,
6 See Petit de Julleville, Vol. I, p. 35; Chambers, Vol. II, p. 53.
7 Officia Propria Festorum Salernitance Ecclesice, Naples, 1594, pp. 75—79. I
print from the copy in the British Museum.
Young — Or do Prophet arum.
19
Isaias:
Ecce virgo concipiet & pariet filium, & vocabitur nomen eius Emanuel,
quod est interpretatum: Nobiscum Deus.
Lector:
Accedat & alius testis. Die & tu, Hieremia, testimonium Christo?
Ieremias :
Hie est Deus noster, & no n sestimabitur alius absque eo qui inuenit
omnem viam scientiae, & dedit earn Iacob puero suo & Israel dilecto
suo. Post haec in terris visus est, & cum hominibus conuersatus est.
Lector:
Ecce duo testes idonei ex lege vestra, ex quorum testimonio non sunt
compuncta corda vestra. Sed alij atqwe alij ex lege Christi testes intro-
ducantur vt frontes durissimi inimicorum conterantur. Veniat & Daniel
sanctus, iuuenis quidem aetate, senior vero scientia ac mansuetudine,
conuincat omnes falsos testes, & sicut conuicit seniores impudicos, ita
suo testimonio Christi conterat inimicos. Die sancte Daniel, die de
Christo quod nosti?
Daniel:
Cum venerit Sanctus Sanctorum, cessabit vnetio vestra.
Lector :
Quare, illo praesente <cui> insultantes dicebatis: Tu de te ipso
testimonium dicis, testimonium tuum non est verum, cessauit vnetio
vestra nisi quia ipse est qui venerat Sanctus Sanctorum? Sed si, ut8 vos
dicitis, non dum venit, sed expectatur, vt veniat Sanctus Sanctorum,
demonstrate vnetionem. Si autem,. quod verum est, cessauit vnetio
vestra, cognoscite venisse Sanctum Sanctorum. Accedat & alius testis.
Die & tu, Moyses, legislator, dux populi Israel, testimonium Christo?
Moyses :
Prophetam vobis excitabit Deus de fratribus vestris; omnis anima
quae non audierit prophetam ilium, exterminabitur de populo suo.
Lector:
Prophetam dictum Christum ipsum Christum audi in Euangelio
<p. 77> dicentem: Non est propheta sine honore, nisi in patria sua.
Accedat enim Dauid sanctus testis fidelis, ex cuius semine processit
ipse cui lex & prophete testomonium dicunt; dicat & ipse de Christo.
Die & tu, Dauid, propheta, testimonium Christo?
Dauid:
Adorabunt eum omnes reges terrae, omnes gentes seruient ei.
Lector:
Cui seruient? Die cui seruient.
si, ut] sicut (Print).
20
Wisconsin Academy of Sciences , Arts , and Letters.
Dauid:
Vis audire cui?
Lector:
Volo.
Dauid:
Dixit Dominus Domino meo: Sede ad dexteram meam, donee ponam
inimicos tuos scabellum pedum tuorum.
Lector:
Et espressius atq ue nominatim: Quare (Quare, inquit) tumultuatse
sunt gentes, & populi meditati sunt inania? Astiterunt reges terrse, &
principes conuenerunt in vnum aduersus Dominum & aduersus Christum
eius. Accedat & alius testis. Die & tu Abachuc, propheta, testimonium
Christo?
Abachuc:
Domine, audiui auditum tuum & timui, consideraui opera tua &
expaui. In medio duorum animalium cognosceris. Opera tua, Dews, &
verbum caro factum est.
Lector :
Ecce quemadmodum comieniunt testes veritatis; ecce quemadmodum
comieniunt testes falsitatis. Sufficiuwt vobis ista, O Iudei? An adhuc
ad vestram cowfusionem & ex lege & ex gente vestra alios introducemus
testes, & illi testimonium praebeant cui perdita mewte insultantes dice-
batis: Tu de te ipso testimonium perhibes; testimonium tuum non est
verum? Accedat & alius testis. Benedicens Dominum Simeon, ex-
clamauit senex ille et dixit. Die & tu, Simeon, propheta, testimonium
Christo?
Simeon:
Nunc dimictis, Domine, seruum tuum secundum verbum tuum in pace,
quia viderunt oculi mei salutare tuum.
Lector:
Ille etiam parentes Ioannis, Zaccharias & Elizabeth, iuuenes steriles &
in senectute fecundi, dicant etiam ipsi testimonium Christo; dicawt
de Christo quod sentiuwt & testem idoneum Christo nutriant. Aiunt
enim suo paruulo nato. Die & tu, Zacharias, propheta, testimonium
Christo?
Zaccharias :
Tu puer, pro<p. 78>pheta Altissimi vocaberis, prseibis enim ante
faciem domini parare vias eius.
Lector:
Ipsique matri & virgini Elizabeth ait. Die & tu, Elizabeth, testi¬
monium Christo?
Young — Or do Prophetarum.
21
Elizabet:
Vnde hoc mihi vt veniat mater Domini mei ad me? Ecce enim vt
facta est vox salutationis in auribus meis, exultauit in gaudio infans in
vtero meo.
Lector:
Intelligens enim Ioa/mes Matrem Domini sui venisse ad suam matrem,
inter ipsas angustias vteri adhuc positus motu salutauit, quern voce no?i
poterat. Die & tu, Ioannes, testimonium Christo?
Ioannes :
Ecce Agnus Dei, ecce qui tollit peccata mundi.
Lector:
Et adiecit: Tu ad me venis baptizari; ego a te debeo baptizari.
Agnouit seruus Dominum, cognouit vinculis originalis peccati obligatus
omni nexu peccati liberum. Agnouit peccator iudicem, agnouit creatura
creatorem, cognouit paranymphus sponsum. Nam & hsec vox Ioannis
est: Qui habet sponsam, sponsus est. Nonhe quando ille facundissimus
poeta inter sua carmina dicit. Die & tu, Virgili, testimonium Christo?
Virgilius :
lam noua progenies ccelo dimittitur alto.
Lector:
Dicebat Christo testimonium propheta? In dubium hoc veniat nisi
alios ex gentibus idoneos testes pluraqwe dicentes in medium introducat.
Ilium regem qui vestram superbiam captiuando perdomuit, Nebuch-
donosor regem scilicet Babylonis, non prsetermittam. Die & tu Nabuch-
donosor testimonium Christo?
Nabuchdonosor :
Nonne tres viros misimus in fornacem ligatos?
Lector:
Et aiunt ei.
Famulus Regis:
Vere, Rex.
Nabuchdonosor:
Ecce video quatUor viros solutos deambulantes in medio ignis, & cor¬
rupts nulla est in eis, & aspectus quarti similis est filio Dei.
Lector:
0 alienigena, vnde tibi hoc? Quis tibi annunciauit filium Dei? Non-
dum quidem mundo nascitur, & similitudo nascerctis a te cognoscitur.
Quid Sibilla vaticinando etiam de Christo clamauerit, in medium profer-
amus, vt ex uno la<p. 79>tere vtrorumqwe frontes percutiantur, Iudse-
orum scilicet atq ue Paganorum atqwe suo gladio, sicut Golias, ChristI
omnes percutiantur inimici. Audite quid dixerit. Die & tu, Sybilla,
testimonium Christo?
22
Wisconsin Academy of Sciences , Arts , and Letters.
Sibilla Erytheaea:
Ivdicii signum: tellus sudore madescet;
E ccelo Rex adueniet, per ssecla futurus,
Scilicet in carne9 presens vt iudicet Orbem.
Vnde Deum cernewt incredulus atq ue fidelis
Celsum cum sanctis seui iam termino in ipso.
Sic animse cum carne aderunt, quas iudicet ipse
Cum iacet incultus densis in vepribus Orbis.
Deiiciewt simulacbra viri, cunctam quoqwe gazam.
Exuret terras ignis, pontumq^e polumq^e;
Inquirens tetri portas effringet Auerni.
Sanctorum sed enim cunctae lux libera carni
Tradetur. Sontes seterna flamma cremabit.
Occultos actus retegens tunc quisq^e loquetur
Secreta atqwe Deus reserabit pectora luci.
Tunc erit & luctus, stridebunt dentibus omnes.
Eripitur solis iubar, et chorus interit astris.
Soluetur caelum, lunaris splendor obibit.
Deiiciet colles, valles extoilet ab imo.
Non erit in rebus hominum sublime vel altum.
Iam aequantur campis montes et caerula powti.
Omnia cessabunt, tellus confracta peribit.
Sic pariter fontes torrentur, fluminaqzte igni;
Et tuba tunc sonitum tristem demittet ab alto.
Orbe gemews facinus miserum varTosqwe labores,
Tartareumq^e chaos mowstrabit terra dehiscews,
Et coram hie Domino reges sistewtur ad vnum.
Decidet e ccelo ignis & sulphuris amnis.
Lector:
Haec de Christi Natiuitate, Passione, et Resurrectione, atq ue secundo
eius Aduentu ita dicta sunt vt si quis in graeco capita eorum discere
voluerit, inueniet: IISVS, CHRISTOS YOS THEV, SOTIR. Quod est
in Latino: IESVS, CHRISTVS, FILIVS DEI SALVATOR. Quod & in
Latinum translatis eisdem versibus apparet praeter quod Graecarum lit-
erarum proprietas non adeo potuit obseruari. Tu<autem, Domine, mis-
ere nobis>.10
Although our information concerning the ceremonial connected
with this liturgical piece is slight,* 11 we can infer the essentials from
the text itself. The unusual liturgical position of the lectio is
made clear in the introductory rubric: In Nativifatis Node post
primam Missam. The reading occurs, then, at the conclusion of
9 carne] carnem (Print).
10 Followed immediately by the rubric : De officio Beati Matthaei in Feria
Quinta.
11 1 am not acquainted with any ordinaria from Salerno.
Young — Or do Prophet arum.
23
the first of the three Christmas Masses, and this first Mass is sung
immediately after the Te Deum of Christmas Matins. This posi¬
tion may be shown schematically as follows :12
Lectio ix (the last lectio of Matins)
Responsorium13
Te Deum
Missa de Nocte (First Mass)
Sermo Sancti Augustini
Laudes
Missa in Aurora (Second Mass)
Prima
Tertia
Missa in Die (Third Mass)
Clearly, then, the lectio is no longer connected with Matins,14
but is an independent liturgical piece for delivery more Salernitano.
The mos Salernitanus for the delivery of the sermo in its new
position is not difficult to discern. The Lector reads the introduc¬
tory address to the Jews, and then summons, in turn, the thirteen
witnesses. That each testimony is read by a separate person repre¬
senting the appropriate witness, is shown by the rubrics giving
the names of the speakers and by the absence of the word inquit
in the testimonies.15
It will be observed that with the text of the sermo the Lector
takes certain liberties : his own utterances are substantially reduced
in length, and the summonses are given a new uniformity.16 But
most noticeable is the expansion into dialogue of the roles of David
and Nabuchadnezzar.17 The expansion of the latter’s dialogue in¬
cludes the introduction of an additional personage to speak the
12 See, for example, U. Chevalier, Ordinaires de Vfiglise cathedrale de Laon,
Paris, 1897, pp. 47-48.
13 According to the mediaeval use, this last responsory of Christmas Matins
was frequently followed by a reading of the genealogy of Christ (Liber Genera-
tionis) from the Gospel of Matthew (i, 1—16). See Chevalier, Ordinaires de
Vftglise cathedrale de Laon, p. 47 ; and the present writer, in Transactions of
the Wisconsin Academy of Sciences, Arts , and Letters, Vol. XVII, Part I (1912),
pp. 313, 385-386, 391.
14 The closing formula Tu <^autem, Domine, miserere nobis^> may, or may
not, be a survival from the use of the sermo as a regular lectio in Matins.
15 The retention of inquit in connection with testimonies that are read by the
Lector himself is illustrated in the Lector’s utterance immediately after the
last speech of David.
16 Note, for example, Die & tu, Simeon and Die & tu, Ioannes.
17 This expansion of the role of Nebuchadnezzar anticipates the dramatic
elaboration surrounding this personage in the more highly developed versions
of the Ordo Prophetarum. See below, pp. 61-62, 65-66.
24 Wisconsin Academy of Sciences, Arts, and Letters.
words of Famulus Regis. There is, of course, nothing to show that
any of the personages named in the rubrics are, or are not, actually
impersonated. From the laconic legitur of the introductory rubric
one more naturally infers that there is no impersonation.
It cannot he asserted positively that the text before us represents
a chronological stage between the use of the sermo as an ordinary,
undialogued lectio in Matins and the use of it unmistakable drama.
Since the lectio of Salerno is known to us only in a printed version
of the late sixteenth century, one might contend that it represents
not a pre-dramatie stage of the development, but a later stage in
which the influence of the dramatic Ordo Prophetarum merely
lingers. To me, however, the very presence of a dialogued lectio in
a printed service-book of the year 1594 seems to indicate a long
tradition; and while awaiting further knowledge concerning the
mediaeval usages of the church of Salerno, I shall regard the text
before us as representing a stage between the normal liturgical
lectio and the fully dramatized Ordo Prophetarum.
IV
However dramatic the version from Salerno may seem, it still
retains the essential outline and language of the Augustinian lectio:
the prophecies are still merely incidental in an expository address, —
an address which has, to be sure, been reduced, but which still
forms the backbone of the piece. We pass naturally, then, to a
version from the monastery of St. Martial at Limoges, in which the
prophecies are virtually free from enclosing exposition, are given
a special literary form, and are set to music i1
1 Paris, Bibl. Nat., Ms. latin 1139, Troparium Martialense ssec. xii, fol. 55v-58r.
The manuscript is described by L.-J.-N. Monmerque and P. Michel, Theatre
frangais au Moyen Age, Paris, 1842, pp. 1-3, by E. de Coussemaker, Histoire de
VHarmonie au Moyen Age, Paris, 1852, pp. 126-127, and by the same author in
his Drames liturgiques du Moyen Age, Rennes, 1860, pp. 311-319. See also
W. Meyer, Fragmenta Burana, Berlin, 1901, pp. 50-51. Since this manuscript
contains the famous Sponsus, numerous notes upon it have been written by
students of romantic dialects (See W. Foerster and E. Koschwitz, Altfran-
zosiches Uehungshuch, Leipzig, 1902, col. 91-94). The text printed below has
been previously edited numerous times. I collate the editions of E. DuMeril
(Les Origines latines du Theatre moderne, Caen, 1849, pp. 179-187. — D. ), Mag-
nin (Journal des Savants, 1846, pp. 88—93. — M. ), Coussemaker (Drames
liturgiques du Moyen Age, pp. 16—20. — C. ), and Monmerqug and P. Michel
(Theatre frangais au Moyen Age, pp. 6-9. — N.). Although Sepet does not edit
the text as a whole, I collate the parts that he transcribes (pp. 15-21 passim)
from the manuscript (S). I do not consider the text of T. Wright (Early Mys-
Young — Or do Prophetarum.
25
<ORDO PROPHETARUM* 2>
<Cantor> :3
Omnes gentes
Congaudentes
Dent cantum leticie.
Dens homo
Fit de domo
Dauid4 natus hodie.5 * <fol. 56r>
O Iudei,
Uerbum Dei
Qui negatis hominem,
Vestre legis
Teste<s>® regis
Audite per ordinem.7
Et uos gentes
Non eredentes
Peperisse uirginem
Vestre gentis
Documentis
Pellite caliginem.
teries and Other Latin Poems of the Twelfth and Thirteenth Centuries, London,
1838, pp. 60-62), derived from P. Michel (See Wright, p. xiv). I omit from
consideration also the text of E. de Coussemaker in his Histoire de VHarmonie
au Moyen Age, — a text dependent upon Magnin and DuM§ril (See Histoire,
p. 133). A complete, but faulty, facsimile of the part of the manuscript with
which we are concerned is found in Coussemaker’s Histoire, Plates xviii-xxiii.
Contrary to the assumption of some persons, the Ordo Prophetarum from this
manuscript is not found in Raynouard, Choix des Poesies originates des Trouba¬
dours, Vol. II, Paris, 1817, pp. 139-143. The text as printed below is imme¬
diately preceded in the manuscript by the closing rubric of the Sponsus : Modo
accipiant eas demones et precipitentur in infernum, which is written in and
above the last four centimeters of the third line from the bottom of fol. 55v.
The first four words of the Ordo Prophetarum, — Omnes gentes congaudentes
dent, — occupy the first two-thirds of the next line. The last four centimeters
of this line are blank. This blank space appears to have been left for an in¬
troductory rubric that was never written. Except for one brief passage, indi¬
cated in my foot-notes, the entire text is furnished with musical notation.
2 1 supply this heading without reference to the several vernacular inventions
of previous editors.
3<Cantor>.] <Praecentor> (D. C. ) ; <Dicat Sacerdos> (M) ; omitted
(N). I enter the conjecture cantor at appropriate places throughout the text
merely for general intelligibility. Since the text is sung, the word is not
inappropriate ; and its lack of explicitness accords with our ignorance of the
precise facts. I myself should raise no objection to cantores or chorus . Du-
Meril’s Praecentor and Magnin’s Dicat Sacerdos, however, seem to imply undue
editorial certainty.
4 Dauid] Dauit (Ms. N. S.).
5 hodie] After this word D inserts the rubric: <Ad Judaeos>.
®Teste<s>] teste <m>- (D) ; Testem (M. C.) ; Teste (N).
7 ordinem] After this word D inserts the rubric: <Ad Gentes >.
26
Wisconsin Academy of Sciences, Arts, and Letters.
IsitaeL
<Cantor> :3
Israel, uir lenis,9 inque
De Xpisto <quae>10 nosti firme.
Responsum z* 11
Dux de Iuda non tolletur12
Donee adsit qui mit<t>etur13
Salutare Del uerbum
Expectabunt14 gentes mecum.
Moynes .
<Cantor> :15
Legislator, hue propinqwa
Et de Xpisto prome digna.
Responsum:16
Dabit Deus uobis uatem;
Huic ut17 m ilii aurem date.
Qui non audit hunc dicentem13
Expellitur19 sua gente.
ISAIAS
< Cantor > :20
Isayas, uerum qui scis,
Ueritatem cur <fol. 56v> non dicis?
Responsum:21
Est necesse
Uirga<m>22 Iesse
8 Israel <Cantor>] <Ad Israelem> (D) ; <Accedat> Israel <et dicat
Sacerdos^> (M) ; Israel (C. N. ).
9 lenis] D suggests fortis.
10<quae>] <quid> (D. C.) ; quae (M) ; quid (S) ; omitted (N).
11 Responsum] Israel (D). It will be observed that DuMeril ignores the rubric
Responsum throughout, and consistently misplaces the names of the witnesses.
12 tolletur] tollitur (Ms. M. C. N.).
13 mit<t>etur] notetur (D. C. N. ) ; vocetur (M. Magnin observes, however,
that the Ms. may read notetur) ; mittetur (S).
14 Expectabunt] expecta<b>unt (D).
15 Moyses < Cantor >] <Praecentor ad Moysem> (D) ; <Accedat> Moyses
<et dicat Sacerdos> (M) ; Moyses (C. N.).
16 Responsum] Moyses (D).
17 ut] et (M).
18dicentem] audientem (Ms. C. N.) ; audientem <loquentem?;> (D. S.).
19 Expellitur] expelletur (D. S. DuM6ril, however, records the Ms. reading
expellitur) .
20 Isaias < Cantor >] <Praecentor ad Isaiam> (D) ; <Accedat> Isaias <^et
dicat Sacerdos> (M) ; Isaias (C. N.).
21 Responsum] Isaias (D).
22 Uirga<m>] Virga (M. N.) ; Virgam (C).
Young — Ordo Propketarum.
27
De radice proue<h>i23
Flos deinde
Surge* inde
Qui est spiritus Dei.
Ieremias
< Cantor >
Hue accede, Ieremias;
Die de Xpisto propbetias.
Responsum:23
Sic est,
Hie est
Deus noster,
Sine quo non erit alter.
Daniel
<Cantor>:2S
Daniel, indica
Uoce prophetica
FaCta dominica.
Responsum:™
Sanctus sanctorum ueniet,
Et unctio deficie*.
<Abacuc
Cantor> :28
Abacuc,29 regis celestis
Nunc ostende quod 20 sis testis.
Responsum :31
Et32 expectaui,
Mox expaui
Metu mirabilium
Opus tuum
Inter duum
Corpus animalium.
23proue<h>i] provehi (M) ; provei (N).
24 Ieremias <Cantor>] <Praecentor ad Ieremiam]> (D) ; <Accedat]> Jere-
mias <et dicat Sacerdos> (M) ; Jeremias (C. N. ).
^Responsum] Jeremias (D).
26 Daniel <Cantor^>] <^Praecentor ad Danielem^> (D) ; <Accedat> Daniel
<et dicat Sacerdos> (M) ; Daniel (C. N. ).
27 Responsum] Daniel (D) ; preceded by a superfluous scribal capital S (Ms).
28 < Abacuc Cantor >] <;Praecentor ad Habacuc> (D) ; <Accedat Abacuc,
et dicat Sacerdos> (M) ; Abacuc (C) ; < Abacuc > (N).
29 Abacuc] <H>abacuc (D).
30 quod] quid (M. N).
31 Responsum] Habacuc (D).
32 Et] D observes that this word is superfluous.
28
Wisconsin Academy of Sciences , Arts, and Letters.
David
< Cantor > :33
Die tu, Dauid,34 de nepote
Causas que35 sunt tibi note.
R esponsumr*5
Vniuersus
Grex conuersus
Adorabat37 Dominurn,
Cui futurum
Seruiturum
Omne genus ho<fol. 57r>minum.
Dixit Dominus Domino meo:
Sede a38 dextris meis.
Simeon
< Cantor > :39
Nunc Symeon adueniat,
Qui responsum acceperat
Qui40 non <Ii>abere£41 terminum
Donee uidere* Dominum,
'Responsum :42
Nunc me dimittas, Domine,
Finire uitam in pace;
Quia mei modo cernunt ocwli
Quern misisti
Hunc mundum43 pro salute popwli.44
83 Dauid <;Cantor>] <Praecentor ad David> (D) ; <;Accedat> David <^et
dicat Sacerdos> (M) ; David (C. N.).
34 Dauid] Dauit (Ms. N.).
35 que] qui (M).
36 Responsum] David (D).
37 Adorabat] adorabit (D. C. DuMeril and Coussemaker, however, record the
Ms. reading adorabat) ; adorabat < adorabit ?> (S).
38 a] ad (Ms. N) ; a<d> (D).
39 Simeon <Cantor>] <Praecentor ad Simeonem> (D) ; <Accedat> Simeon
<|et dicat Sacerdos> (M) ; Simeon (C. N.).
40 Qui] Quod (D. M. Magnin, however, records the Ms. reading as qui).
41 <h>aberet] haberet (D. M. C.) ; aberet (N).
^Responsum] Simeon (D).
43 Hunc mundum] By printing these words in square brackets D appears to
regard them as superfluous.
44 For the entire part of Simeon (Nunc Symeon . . . salute populi) the
musical notation is omitted.
Young— Or do Prophetarum.
29
Elis abet
<Cantor>:45
Illud,46 Helisabet, in medium
De Domino profer47 eloquium.
R esponsum ;48
Quid est rei
Qttod me mei
Mater <h>eri49 uisitat?
Nam ex eo
TJentre meo
Letus infans palpitat.
< Johannes Baptista
Cantor> : 50
Die,51 Babtista,
Uentris cista clausus
Qua52 dedisti causa
Xpisto <fol. 57v> plausus.
Cui dedisti gaudium
Profer53 et testimonium.
R esponsum:5*
Venit talis
Sotularis85
Cuius non sum etiam
Tam benignus
Ut sim ausus™
iSoluere corrigiam.
46 Elisabet <Cantor>] <]Praecentor ad Elisabeth > (D) ; <Accedat> Elisa¬
beth <et dicat Sacerdos> (M) ; Elisabeth (C. N.).
40 Illud] Illuc (M. Magnin, however, records the Ms. reading as Illud).
47 profer] profert (Ms. N.) ; profer <t> (D).
48 Responsum] Elisabeth (D).
49 <h>eri] heri (D. M. C.) ; Eri (S. N.).
50 Johannes Baptista Cantor^] <Praecentor ad Johannem Baptistam>
(D) ; <[Accedat Johannes Baptista et dicat Sacerdos> (M) ; <Joannes Bap-
tista> (C. N.).
51 Die] De (Ms. N‘.).
52 Qua] Quod (Ms. N.).
63 Profer] Profert (Ms. N.) ; profer <t> (D).
54 Responsura] Johannes Baptista (D).
55 Sotularis] Solea nobis (M. Magnin, however, records the Ms. reading sotu¬
laris) ; salutaris (C).
50 ausus] dignus (D. DuMeril, however, records the Ms. reading ausus) ; ausus
<dignus?> (S).
30
Wisconsin Academy of Sciences , Arts , and Letters.
VlRGILI us
<Cantor>0T
Vates, Maro,68 gentilium
Da50 Xpisto testimonium.
R esponsum:™
Ecce polo demissa solo noua progenies est.
Nabucodo < no > SOR
<Cantor> :61
Age, fare, os laguene,63
Que de Xpisto nosti uere.63
Nabucodonosor, propheti<z>a,64
Auctorem omnium auctoriza.
Responsum .,65
Cum reuisi
Tres quo<s>66 misi
Uiros in incendium
Vidi iustis
Inconbustis
Mixtum Dei filium,
Viros tres in ignem misi
Quartum cerno67 prolem Dei.
Sibilla <fol. 58r>
<Cantor> :68
Vera pande iam, Sibilla,
Que de Xpisto pre<s>cis69 signa.
67Uirgilius <Cantor>] <Praecentor ad Virgilium> (D) ; <Accedat> Vir-
gilius <et dicat Sacerdos> (M) ; Virgilius (C. N. ).
58 Maro] Moro (Ms. N. ).
59 Da] dea (Ms. N.).
60 Responsum] Virgilius (D).
61 Nabucodo < no >sor <Cantor>] <Praecentor ad Nabuchodonosor ;> (D) ;
<Accedat> Nabucodonosor <^et dicat Sacerdos> (M) ; Nabuchodonosor
(C. N.).
63 os laguene] os laguene <nos loquere?> (D) ; os lagenae (M. Magnin, how¬
ever, records the Ms. reading laguene) ; os lagene (C).
63 uere] uere Responsum (Ms. N.) ; uere <; Alias > (M). Upon the scribal
insertion of the rubric Responsum, Magnin comments plausibly as follows :
“Codex : responsum. C’est une erreur du copiste. Ces deux vers ne peuvent
etre qu’une variante du distique precedent qui aura paru trop grossier.” See
Sepet, p. 33.
64 propheti< z> a] prophetia (M. N.) ; prophetiza (C).
65 Responsum] Nabuchodonosor (D).
66quo<s>] quos (C) ; quo (N).
67 cerno] cerna (Ms. N. ).
68 Sibilla < Cantor >] <Praecentor ad Sibyllam> (D) ; <Accedat> Sibylla
<et dicat Sacerdos> (M) ; Sibilla (C. N.).
69Pre<s>cis] Praescis (M. C.) ; precis (N).
Young — Or do Propbetarum.
31
R esponsum :70
Iudicii71 signum: Tellus sudore madescet;
E celo rex adueniet per secla futurus,
Scilicet in carne presens ut iudicet orbem.
<Cantor>:72
ludea incredula,
Cur manes73 adbuc inuerecunda?
<H>IC INCOANT BeNEDICAMWS.74
A mere glance suggests that this dramatic Ordo Prophetarum is
developed from the pseudo- Augustinian lectio. The two are identi-
70 Responsum] Sibylla (D).
71Iudicii] Iuditii (Ms. N. ).
72 <Cantor>] <Praecentor ad Judaeos> (D) ; omitted (M. C. N.).
73 manes] manens (Ms. D. N. )
74 <H>ic incoant Benedicamus] omitted (D. C.) ; Incoant Benedicamus (M) ;
Incohant benedicamus (N). Immediately after the rubric <^H^>ic incoant
Benedicamus is found the following metrical composition (printed without sub¬
stantial difference by Sepet, p. 26) :
Letabundi iubilemus,
Accurate celebremus
Xpisti natalicia
Summa leticia ;
Cum gratia
Produxit,
Gratanter mentibus
Fidelibus
Inluxit.
Eructauit Pater Uerbum,
Perdit hostis ius acerbum.
Quod in nobis habuit,
Quod diu latuit,
Tunc patuit <fol. 58v>
Arcanum.
Qui contra gariunt
Insaniunt
In uanum.
O re <s>- digna predicari.
Cui non ualent conparari
Quantauis miracula.
Ferit uirguncula
Per secula
Rectorem
Conceptum edidit
Nec perdidit
Pudorem.
Now follows the rubric Alium Benedicamus, and a fresh trope beginning:
Prima mundi seducta sobole, printed by Sepet, p. 84.
32 Wisconsin Academy of Sciences, Arts, and Letters.
cal in general subject, — the refutation of the Jews; and the similar¬
ity between the two lists of witnesses appears in the following :75
The list of the Limoges version differs only slightly from that of
the lectio. Israel has been added, and Zacharias has been dropped.
These changes are not weighty, however, since Israel is mentioned
in the original lectio,76 and Zacharias is sufficiently represented by
his wife Elisabeth.77 Although neither list is thoroughly chrono¬
logical, the Limoges version shows certain improvements in this
respect.78 The first six names are in approximately chronological
order, with Israel appropriately at the head. David, as progenitor,
appears appropriately enough after the first six witnesses and imme¬
diately before the contemporaries of Christ. The pagan witnesses
are left in their original position, and in chronological disorder
among themselves.
The detailed resemblances between the testimonies in the lectio
and those in the dramatic text of Limoges may be conveniently seen
in the following comparison:
75 In each column I present the names in the order of their appearance in the
text itself.
76 In the prophecy of Jeremiah. See above, p. 5.
77 See Sepet, p. 16.
78 The chronological disorder still remaining in the Limoges version is ob¬
served by Sepet, pp. 84-85.
Young— Or do Prophet arum.
33
Lectio
Isaiah
Ecce virgo in utero concipiet et
pariet filium, et vocabitur nomen
ejus Hemanuhel, quod est interpre-
tatum: Nobiscum Deus.79
Jeremiah
Hie est Deus noster et non sesti-
mabitur alius absque illo qui in-
venit omnem viam scientise, et dedit
earn Jacob puero suo et Israel di-
lecto suo. Post hsec in terris visus
est et cum hominibus conversatus
est.81
Daniel
Cum venerit Sanctus Sanctorum,
cessabit unctio.82
Moses
Prophetam vobis suscitabit Deus
de fratribus vestris; omnis anima
quas non audierit prophetam ilium
exterminabitur de populo suo.83
Ordo of Limoges
Isaiah
Est necesse
Virga<m> Jesse
De radice prove <h>i
Flos deinde
Surget inde
Qui est spiritus Dei.80
Jeremiah
Sic est
Hie est
Deus noster
Sine quo non erit alter.
Daniel
Sanctus sanctorum veniet,
Et unctio deficiet.
Moses
Dabit Deus vobis vatem
Huic ut mihi aurem date
Qui non audit hunc dicentem
Expellitur sua gente.
79 Ecce virgo concipiet, et pariet filium, et vocabitur nomen ejus Emmanuel
(Isa. vii, 14).
80 Et egredietur virga de radice Jesse, et flos de radice ejus ascendet. Et
requiescet super eum spiritus Domini (Isa. xi, 1-2). Cf. Isa. xi, 10 ; Rom. xv,
12-13 ; Num. xvii, 5-6.
81 Hie est Deus noster, et non sestimabitur alius adversus eum. Hie adinvenit
omnem viam disciplinae, et tradidit illam Jacob, puero suo, et Israel, dilecto suo.
Post hsec in terris visus est, et cum hominibus conversatus est. (Bar, iii, 36—38).
82 Septuaginta hebdomades abreviatse sunt super populum tuum et super urbem
sanctam tuam, ut consummetur praevaricatio, et finem accipiat peccatum, et
deleatur iniquitas, et adducatur justitia sempiterna, et impleatur visio, et pro-
phetia, et ungatur sanctus sanctorum (Dan. ix, 24).
83 Prophetam de gente tua et de fratribus tuis sicut me, suscitabit tibi Dominus
Deus tuus ; ipsum audies . . . Prophetam suscitabo eis de medio fratrum
suorum similem tui, et ponam verba mea in ore ejus, loqueturque ad eos omnia
quae praecepero illi. Qui autem verba ejus, quae loquetur in nomine meo, audire
noluerit, ego ultor existam (Deut. xviii, 15, 18-19).
3— S. A. L.
34
Wisconsin Academy of Sciences , Arts, and Letters.
David
Adorabunt eum omnes reges ter-
rae; omnes gentes servient illi.
Dixit Dominus Domino meo: Sede
ad dexteram meam donee ponam
inimicos tuos scabellum pedum
tuorum.84
Habakkuk
Domine, audivi auditum tuum et
timui; consideravi opera tua et ex-
pavi. In medio duum animalium
cognosceris.85
Simeon
Nunc dimittis, Domine, servum
tuum in pace, quia viderunt oculi
mei sa^utare tuum.86
David
Universus
Grex conversus
Adorabat Dominum
Cui futurum
Serviturum
Omne genus bominum.
Dixit Dominus Domino meo:
Sede a dextris meis.
Habakkuk
Et expectavi
Mox expavi
Metu mirabilium
Opus tuum
Inter duum
Corpus animalium.
Simeon
Nunc me dimittas, Domine,
Finire vitam in pace;
Quia mei modo cernunt oculi
Quern misisti
Hunc mundum pro salute populi.
Zacharias Zacharias
Tu, puer, propheta Altissimi vo-
caberis, preibis enim ante faciem (Wanting)
Domini parare viam ejus.87
84 Reminiscentur et convertentur ad Dominum universi fines terrae ; et adorabunt
in conspectu ejus universse familiae gentium ; quoniam Domini est regnum, et
ipse dominabitur gentium et semen meum serviet ipsi (Ps. xxi, 28-29,
31); Dixit Dominus Domino meo: Sede a dextris meis, donee ponam inimico3
tuos scabellum pedum tuorum (Ps. cix, 1). Cf. Matt, xxii, 44.
85 Domine, audivi auditionem tuam, et timui. Domine, opus tuum, in medio
annorum vivifica illud ; in medio annorum notum facies ; cum iratus fueris,
misericordise recordaberis (Hab. iii, 2).
86 Nunc dimittis servum tuum, Domine, secundum verbum tuum in pace ; quia
viderunt oculi mei salutare tuum, quod parasti ante faciem omnium populorum
(Luc. ii, 29-31).
87 Et tu, puer, propheta Altissimi vocaberis ; praeibis enim ante faciem
Domini parare vias ejus (Luc. i, 76).
Young — Or do Prophet arum.
35
Elisabeth
Unde mihi hoc ut veniat mater
Domini mei ad me? Ecce enim ut
facta est vox salutationis tuae in
auribus meis, exultavit in gaudio in¬
fans in utero meo.88
Johannes Baptista
Quern me suspicamini esse, non
sum ego. Sed ecce venit post me
cujus pedum non sum dignus solvere
corrigiam calciamenti.89
Virgilius
Jam nova progenies coelo demit-
titur alto.90
Nebuchadnezzar
Nonne tres viros misimus in for-
nace ligatos? Ecce ego video qua-
tuor viros solutos deambulantes in
medio ignis et corruptio nulla est in
eis, et aspectus quart! similis est
Filio Dei.91
Sibilla
Judicii signum: tellus sudore
madescet;
E coelo rex, etc.
Elisabeth
Quid est rei
Quod me mei
Mater <h>eri visitat?
Nam ex eo
Yqntre meo
Letus infans palpitat.
Johannes Baptista
Venit tails
Sotularis
Cuius non sum etiam
Tam benignus
Ut sim ausus
Solvere corrigiam.
'Virgilius
Ecce polo demissa solo nova pro¬
genies est.
Nebuchadnezzar
Cum revisi
Tres quo<s> misi
Viros in incendium
Vidi justis
Incombustis
Mixtum Dei filium.
Viros tres in ignem misi
Quartern cerno prolem Dei.
Sibilla
Judicii signum: tellus sudore
madescet;
E coelo, etc.
88 Et unde hoc mihi ut veniat mater Domini mei ad me? Ecce enim ut facta
est vox salutationis tuae in auribus meis, exsultavit in gaudio infans in utero
meo (Luc. i, 43-44).
89 Respondit Joannes dicens omnibus : Ego quidem aqua baptizo vos ; veniet
autem fortior me, cujus non sum dignus solvere corrigiam calceamentorum ejus
(Luc. iii, 16) ; Respondit eis Joannes dicens: Ego baptizo in aqua; medius autem
vestrum stetit, quern vos nescitis. Ipse est, qui post me venturus est, qui ante
me factus est ; cujus ego non sum dignus ut solvam ejus corrigiam calceamenti
(Jo. i, 26-27 >. Cf. Marc, i, 7.
"Iam nova progenies caelo demittitur alto (Virgil, Eel. iv, 6).
91 Tunc Nabuchodonosor rex obstupuit ; et surrexit propere, et ait optimatibus
suis: Nonne tres viros misimus in medium ignis compeditos? Qui respondentes
regi dixerunt : Vere, rex. Respondit et ait : Ecce ego video quatuor viros
solutos, et ambulantes in medio ignis, et nihil corruptionis in eis est, et species
quarti similis Fillio Dei (Dan. iii, 91-92).
36
Wisconsin Academy of Sciences, Arts, and Letters.
Israel Israel
Dux de Juda non tolletur
Donee adsit qui mit<t>etur
(Wanting) Salutare Dei verbum
Expectabunt gentes mecum.92
Aside from the presence of Israel in the Limoges version, and
the absence of Zacharias, the most conspicuous divergence from the
lectio is seen in the prophecy of Isaiah. For this prophecy the
Limoges author offers a stanza clearly based upon a prophetic
passage from the book of Isaiah unconnected with that quoted in
the lectio.93 In general, however, the testimonies in the Limoges
text may be characterized as versifications of the parallel utter¬
ances in the lectio.94: Particularly noticeable is the prophecy of
Daniel in the two versions, in that the Limoges text manifestly de¬
rives from the lectio rather than from the Vulgate.95
If one wished to add further details to the demonstration one
might cite such resemblances as the following, in the summonses:96
(1) Lectio: Die, et Moyses, legislator . . .
Limoges: Legislator , hue propinqua . . .
(2) Lectio: Symeonem sanctum in medio introducor . . . Cum
iste senex admonitus esset a Spiritu Sancto quod non
ante moreretur quam videret Christum Dei natum
Limoges: Nunc Symeon adveniat
Qui responsum acceperat
Qui non haberet terminum
Donee videret Dominum.
Assuming, then, that the lineage of the Limoges version is es¬
tablished,97 we may examine it in detail as an example of the
92 Non auferetur sceptrum de Juda, et dux de femore ejus, donee veniat qui
mittendus est ; et ipse erit expectatio gentium . . . Salutare tuum expectabo,
Domine (Gen. xlix, 10, 18).
93 This stanza is found also in the cantio (or trope) Gloriosi et famosi, dis¬
cussed below, pp. 77-80.
94 See Sepet, pp. 16-21.
95 One should observe also that in the prophecy of David both the lectio and
the Limoges version unite unconnected verses from separate Psalms.
9(5 Concerning these resemblances, and others, see Sepet, pp. 21-24.
97 La Piana (op. cit., pp. 302-308) undertakes to show that the Limoges ver¬
sion borrows somewhat from a Greek homily of Hesy chius of Jerusalem
(printed by Migne, Patrologia Graeca, xciii, 1453-1460). That the Limoges
author is chiefly indebted to the pseudo -Augustinian lectio rather than to the
Greek homily is obvious, for example, from the fact that seven of the prophets
common to the lectio and the Limoges text are wanting to the Greek homily.
Young — Ordo Prophetarum .
37
dramatic Ordo Prophetarum. In the absence of adequate rubrics,
we are left in considerable uncertainty as to the manner in which
it was produced. We are at least sure that the piece was delivered
as a dialogue, or as a succession of dialogues. This is clear both
from the nature of the utterances themselves and from the use of
the rubric responsum. One may reasonably suppose that the in¬
troductory exhortations and the summonses are all sung by the
same cleric, or group of clerics, — we cannot be more specific.98
We may infer also that at each summons one of the prophets ap¬
peared in person and responded with his prophecy. Of impersona¬
tion, upon the part either of the summoner or of the prophets, we
cannot be sure. The fact that the name of each prophet is written
as a rubric immediately before the summons seems to indicate that
a separate speaker appeared at this point, and it suggests that he
impersonated the prophet concerned.99 As to the part of the church
in which the performance occurred, and as to mise en scene , we
have no knowledge.
In the absence of more precise information we may accept some
such conjectural description of the performance as the following
from Magnin:100
Moreover the lectio and the Limoges version agree in differing from the Greek
in the prophecy of Daniel. La Piana acknowledges these facts. The one sub¬
stantial suggestion of the Limoges author’s borrowing from the Greek is the
summons and prophecy of Israel (Jacob), which is wanting in the Latin lectio,
but which is found in the other two texts in the following forms :
Ordo of Limoges
Israel, vir lenis, inque
De Xpisto <qu8e> nosti firme.
Responsum :
Dux de Iuda non tolletur
Donee adsit qui mittetur
Salutare Dei verbum
Expectabunt gentes mecum.
Hesychius
II apdffTTjdL tw ’Ia/ccu/3 /cat rl Xeyei
/jLera^i) t&v evXoyi&v Kar&fiaOe.
Ovk etcXeixf/et apxwv tt- ’lovda, /cat
riyovfievos £k tCov /xrjpcov avrov, ecus ov
eX07f os diroKeiTai /cat avros TrposdoKia
ktiv&v.
(Migne, Pat. Or., xciii, 1460)
This parallel may, or may not, show that for the prophecy of Israel the
Limoges author resorted to Hesychius. If he borrowed at all, he borrowed little.
98 As I have said above (p. 25), DuMeril’s (and Coussemaker’s) con¬
jectured Praecentor and Magnin’s conjectured Sacerdos imply undue certainty
upon the part of these editors. Petit de Julie ville (Les Mysteres, Vol. I, p. 36)
adopts Magnin’s conjecture, and Chasles (p. 123), that of DuM§ril and Cousse-
maker. The term Appellatores of the Laon version (See below, pp. 41-45) and'
the words Chorus and Vocatores of the Rouen Festum Asinorum (See below,
pp. 50-63) suggest that the introductory verses and the summonses of the-
Limoges verses were not sung by a single person.
99 Sepet (p. 25) considers impersonation not improbable.
100 Journal des Savants, 1846, p. 88. A somewhat less venturesome conjecture-
is advanced by Sepet, pp. 25-26. See also Sepet, pp. 40-41. Both Magnin and;
Sepet acknowledge the conjectural nature of their descriptions.
38
Wisconsin Academy of Sciences , Arts , and Letters.
D’abord, les trois premieres strophes, qui sont comme le prologue ou
1’exposition du mystere, devaient etre dites ou chantees par un eccldsi-
astique elevd en dignite. Ensuite ce personnage appelait a haute voix
chacun des acteurs du drame, lesquels s’avangaient et prenaient succes-
sivement la parole. Ce principal interlocuteur etait, comme nous dirions
pour un spectacle profane, le meneur ou le directeur du jeu. II se tenait
probablement debout sur les degres de 1’ambon ou au milieu du jub£,
entoure des musiciens. Les autres personnages, pretres ou moines, vetus
du costume de leurs roles, etaient assis dans les stalles, attendant le
moment de se lever et de venir au milieu du choeur psalmodier ou chanter
leur verset.
We are left in uncertainty also as to the day on which the
Limoges Ordo was performed and as to its position in the liturgy.
One might infer that the words natus hodie of the opening verses
were sufficient for identifying the Ordo with Christmas day;101
but the same verses are found also in the Rouen Festum Asinorum ,
which was undeniably presented on the day of the Circumcision.102
As to the precise liturgical attachment of the Limoges Ordo our
only possible shred of information is the rubric at the end : <H>fc
incoant Benedicamus. If this rubric is to be regarded as belong¬
ing to the preceding dramatic text, it must, of course, be inter¬
preted as meaning, “Here let them begin the Benedicamus.” Since
the Benedicamus was sung at the end of each of the canonical
“ horse ”, this interpretation, upheld by Sepet,103 would locate the
dramatic performance at the end of one of these services. But the
rubric may be interpreted also as meaning, “Here begin the Bene-
dicamuses.” Since the rubric is followed by a series of metrical
pieces evidently composed as tropes of the Benedicamus , this second
interpretation, advanced by Magnin,104 is by no means inept. Be¬
tween the two interpretations I am unable to decide positively. It
may fairly be said that the plain facts of the manuscript, in which
tropes of the Benedicamus follow the rubric, give some support
101DuMeril (p. 180) chooses to assign the Limoges Ordo to the day before
Christmas; Sepet (pp. 14-15) is unconditionally for Christmas day.
102 See below, pp. 66—68.
103 Sepet (pp. 25-26, 84) regards as part of the dramatic text not only the
rubric under discussion but also some score or more of verses of text following
it. Petit de Julleville (Vol. I, p. 36) appears to accept Sepet’s view, as, ac¬
cording to Monmerque and Michel (p. 1), does Lebeuf. Monmerque and Michel
themselves do not (p. 3) regard the Benedicamus trope as part of the dramatic
text.
104 Magnin (p. 92) prints the rubric Inchoant Benedicamus, and translates it
as follows : “Ici commencent les Benedicamus.” Monmerque and Michel give
(p. 9) precisely the same translation.
Young— Ordo Prophet arum.
39
to the view of Magnin. On the other hand it may be said that a
liturgical play may normally be expected to end with a rubric of
some kind. For the present one may well withhold judgment.105
It is clear, in summary, that however uncertain we may be as to
external matters of production and liturgical relationship, we must
certainly recognize in the Limoges version a notable advance in the
literary treatment of the Ordo Prophetarum. Most noteworthy
are the elimination of the narrative and expository element of the
lectio, and the versifying of both summonses and responses. In
addition one must commend the force and aptness of the opening
stanzas, seen here for the first time.106
V
Although the advance shown by the Limoges version in literary
form is marked, the approach of this production toward acted
drama is, as we have seen, a matter of uncertainty, because of the
absence of rubrics indicating impersonation. We may proceed,
then, to a play of the thirteenth century from the Cathedral of
Laon, in which impersonation and specific dramatic action are un¬
mistakable i1
105 Sepet (pp. 24-26, 84) undoubtedly exaggerates the significance of the
rubric Hie incoant Benedicamus. He not only considers this rubric and the
subsequent lines as organically related to the preceding dramatic text, but he
also regards the dramatic text itself as a trope of the Benedicamus. As a
matter of fact, we cannot be sure that the dramatic text is to be connected
with the Benedicamus; and even if the dramatic text was performed imme¬
diately before the Benedicamus , only by exaggeration can it be called "un
trope du Benedicamus” (p. 25). L». Gautier (Les Tropes, Paris, 1886, p. 2) has
ably defined a trope as an extra-liturgical addition to a traditional liturgical
piece, by way of preface, interpolation, or conclusion. By an exaggerated
application of this definition every liturgical play may, of course, be regarded
as a trope, since every such play must take a place before or after some
liturgical element. But the exaggeration in calling Hilarius’ Daniel, for ex¬
ample, a trope of the Te Deum or of the Magnificat is obvious (See DuMeril,
pp. 241-254). When a dramatic piece has sufficient complexity and complete¬
ness for maintaining itself as an independent production, one may safely cease
regarding it as a trope of the liturgical element that happens to precede or
follow it.
108 The opening stanzas ( Omnes gentes congaudentes . . . pellite cal-
liginem) are found also as part of a longer composition Gloriosi et famosi to be
considered below, pp. 41, 78.
1 Laon, BibliothSque de la Ville, Ms. 263, Troparium-Hymnarium Laudunense
ssec. xiii, fol. 147v-149r. The manuscript is briefly described in Catalogue
40
Wisconsin Academy of Sciences , Arts , and Letters.
ORDO PROPHETARUM
YSAIAS, BARBATUS, DALMATICA INDUTUS, STOLA RUBEA per MEDIUm uerticis
ANTE et RETRO DEPENDENTE.2
IHEREMIAS SIMILITer, ABSQtte STOLA.
Daniel adolescens, ueste splendida indutus.
MOYSES, CUM DALMATICA, BARBATUS, TABULAS LEGIS FERENS.
Dauid, REGIO HABITU.
AbACUC BARBATUS, CURUUS, GIBOSUS.
Elisabeth, femineo habitu, pregnans.
lOHANNES BAPTISTA, PILOSA UESTE et LONGIS CAPILLIS, BARBATUS, PALMAM
TENENS.
VlRGILIUS, CUM CORNU et C ALAMO, EDERA CORONATUS, SCRIPTORIUm TENENS.
NABUGODONOSOR, REGIO HABITU, SUPerBO INCESSU.
SlBILLA, UESTE FEMINEA, DECAPILLATA, EDERA CORONATA, INSANIENTI SIMIL-
LIMA.
SYMEON, BARBATUS, CAPA SERICA IN <fol. 148r> DUTUS, PALMAM TENENS.
Balaam, super ASiNAm, curuus, barbatus, palmam tenens, calcaribrs
IJRGENS.
Gloriosi
Et famosi
Regis festum
Celebrantes
Gaudeamus.
Cuius ortum,
Uite portum,
Nobis datum,
Predicantes
Habeamus.
Ecce regem
Nouam legem
Dantem orbis
Circuitu
Predicamus.
general des Manuscrits des Bibliotheques publiques des Departements, Vol. I,
Paris, 1849, p. 155. The Ordo Prophetarum from this manuscript has been pre¬
viously published by U. Chevalier, Ordinaires de Vfiglise cathedrale de Laon
( Bibliotheque Liturgique, Vol. VI), Paris, 1897, pp. 385-389; and Chevalier’s
text is reprinted by Chasles in La Vie et les Arts liturgiques , Jan., 1917, pp. 129-
134. My own edition of the text from the manuscript differs in no essential
way from that of Chevalier. The opening- rubric Ordo Prophetarum is imme¬
diately preceded by the following unrelated incipit : De Sancto Nicholao : Con-
gaudentes. For the text of the Ordo Prophetarum no music is provided.
2 dependente] dependens (Ms).
Young— -Or do Prophet arum.
41
Duo Cantores:
Oirmes gentes
Congaudentes
Dent cantus letitie;
Deus homo
Fit de domo
Dauid, natus hodie.
Ad Iudeos :
O Iudei,
Uerbum Dei
Qui negastis hominem,
Vestre3 legis
Testes regis
Audite per ordinem.
Ad Paganos:
Et uos gentes
Non credentes
Peperisse uirginem
Vestre legis
Documentis
Pellite caliginem.
Appellatoees :
Isaias, uerum qui scis,
Ueritatem cur non dicis?
Isaias:
Est necesse
Uirgam Iesse
De radice prouehi
Flos deinde
Surget inde
Qui est filius Dei.
Appellatoees :
Iste cetus
Psallat letus;
Error uetus
Condempnetwr.
Omnis Chorus:
Quod Iudea
Perit rea
Hec chorea
Gratulatwr.
Appellatoees :
Hue accede, Iheremias;
Die de Xpis£o prophetias.
3 Vestre] Veste (Ms), — with an r faintly inserted by a later hand.
42
Wisconsin Academy of Sciences , Arts, and Letters.
Hieremias :
Sic est
Iste cetus.
iTErn chokws:
Qwod Iudea.
Qwod Iudea.
Ap<p>Eixa£ ores :
Die tu, Moyses, legislator,
Qwod Iudea.
Ap<p>ELLa fores ;
Die tu, Dauid, de nepote
Qwod Iudea.
AppEixafores :
Abacuc, regis celestis
Nunc ostende quod sis testis.
4 deficiet] defitiet (Ms. Chevalier).
Young — Or do Prophet arum.
43
Quod Iudea.
Ap<p>ell atores :
Illud, Elisabeth, in medium
De Domino profer eloquimn,.
Elisabeth :
44
Wisconsin Academy of Sciences , Arts , and Letters.
Mako:
Ecce polo dimissa sola noua progenies est.
Duo:
QttOd Iudea.
APPEULA TORES REDUCUNT DANIELEM, et DICIWT AD REGEm .*
Fuerum cum pueris,
Nabugodonosor,
Cum in igne uideris,
Quid dixisti?
Nabugodonosor :
Tres in igne positi pueri
Qttod Iudea.
Ap<p>ELLa£ores :
Tu, Sibilla,
Qwod Iudea.
Appellators :
Symeon, inter propbetas
Symeon accipiens PuERum dicit:
Tuum sub pacis tegmine
Seruum dimittis, Domine.
Young — Or do Prophet arum.
45
Ap<p>ELLGtfores :
Die, Balaam, ex Iudaica
Oriturum Dominion prosapia.
Balaam :
Exibit de Iacob rutilans noua stella
Et confringet ducum agmina
Regionis Moab maxima potentia.
Hie UENIAT ANGeL^S CUM GLADIO. BALAAm TANGIT ASINAM,
et ILLA Non PrOCEDENTE, DICIT IRATUS I
Quid moraris, asina,
Obstinata bestia?
Iam seindent calcaria
Costas et precordia.
PUER SUB ASINA RESPONDED
Angelas cum gladio,
Quern adstare uideo,
Probibet ne transeam;
Timeo ne peream.5
Here, for the first time, impersonation is manifest. In suggestive
details of costume and property the text leaves little to be desired.
Daniel’s youthful appearance, Moses’ tables of the Law, Elizabeth’s
pregnancy, John the Baptist’s hairy shirt and branch of palm,
Virgil’s writing materials and crown of ivy, the Sibyl’s expression
of mad inspiration, Balaam’s ass, — these are some of the details ex¬
plicitly provided for.
The performance itself begins with the singing of the Gloriosi et
famosi, presumably by the Prophetce themselves.6 Two cantors
now deliver three stanzas, the second and third of which are ad¬
dressed specifically to the Jews an Gentiles respectively, calling
upon them to prepare for enlightenment. Two7 special summoners
( Appellator es) now call forth the prophets, one by one, and after
each prophecy deliver the stanza I site ccetus, to which the chorus re¬
sponds with the stanza Quod Judcea.8
The close kinship of the Laon play to the Limoges version and to
the original lectio is obvious. Each of these three versions pre¬
sents thirteen witnesses, and twelve of these witnesses are common
5 Followed immediately by the rubric Or do Stelle, introducing a version of
the Epiphany play.
6 This presumption finds justification in the usage recorded in the Einsiedeln
fragment printed below, pp. 72-74.
7 Several of the later rubrics specify duo.
8 1 have not found these two stanzas apart from the prophet-plays of Eaon
and Rouen.
46 Wisconsin Academy of Sciences, Arts , and Letters.
to all three : namely, Isaiah, J eremiah, Daniel, Moses, David, Habak-
knk, Elisabeth, John the Baptist, Virgil, Nebuchadnezzar, Sibyl,
and Simeon. Balaam is peculiar to the Laon version, as Israel and
Zacharias are to the Limoges version and the lectio respectively.
The similarity of the three versions is further apparent in the
textual content of the prophecies. In eight of these prophecies9
the Limoges and Laon versions agree in presenting metrical ver¬
sions of the utterances provided by the lectio. In the utterance of
Isaiah the Limoges and Laon versions agree as against the lectio.
In the prophecies of Moses and Nebuchadnezzar the two dramatic
versions differ from each other in expression, but both derive from
the lectio. The utterance of Simeon in the Laon play is developed
somewhat beyond what is found either in the Limoges text or in the
lectio.
In general arrangement the Laon play differs considerably from
the two preceding versions. The non-chronological order in which
the first six prophets appear reproduces what we have observed in
the lectio. The presence of Simeon and Balaam at the end, after
Sibyl, suggests that those two prophets were attached to the series
in a relatively mechanical way. Possibly the copy from which
the present text was made was in disorder. This impression is in¬
deed strengthened by the inconclusiveness with which the play
ends.10
In considering the delivery of the prophecies from the dramatic
point of view, our attention centers in three : those of Nebuchadnez¬
zar, Simeon, and Balaam. The first of these may be dismissed
briefly, for although the rubric Appelatores reducent Danielem
suggests special dramatic action of some sort, the nature of this
action is not specified. The dramatic action accompanying the
role of Simeon can be visualized somewhat more clearly. It ap¬
pears that Simeon barbatus is represented as standing in the temple
in the presence of Joseph, Mary, and Jesus, and that after Simeon’s
first utterance has been heralded by the cantores and choir, he
takes the Child in his arms and speaks again, in the manner of a
Nunc dimittis.* 11
9 Jeremiah, Daniel, David, Habakkuk, Elisabeth, John the Baptist, Virgil, and
Sibyl.
10 Concerning the mechanical attachment of the part of Balaam to the end
of the play see Chambers, Vol. II, pp. 54, 56-57 ; Chasles, p. 127, note 2.
11 See Luc. ii, 22-32.
Young — Ordo Prophetarum.
47
The part of Balaam deserves special comment for its dramatic
liveliness. Balaam makes his appearance mounted upon an ass.* 1 2 3 * * * * * * * * 12
After he has uttered his prophecy an angel appears in his path, so
halting the progress of the animal that Balaam impatiently strikes
her and demands the reason of the delay. In defense, the puer sub
asina explains that an angel stands threateningly in the path ; and
thus the play abruptly ends. The action of this little dramatic
episode centering in Balaam is, of course, derived from the Biblical
narrative (Num. xxi, 21-31) ; but the speeches show no dependence
upon the Vulgate text. The speech of the Appellator es seems
clearly to be adapted from the following passage in a well-known
sequence of Epiphany : Quam Balaam ex Iudaica orituram dixerat
prosapia;13 and the response of Balaam consists in a passage from
another Epiphany sequence; Exibit ex Jacob rutilans, inquit ,
Stella , Quce confringet ducum agmina regionis Moab maxima po¬
tential The second speech of Balaam and the response of the
asina seem to be original compositions for the purpose at hand.15
It is clear that the parts of Simeon and of Balaam stand apart
from the rest of the performance as small dramatic units. In ac¬
tion and wise en scene they show an advance beyond the general
uniformity of the elements examined hitherto. These two bits of
composition seem to have been ambitiously added at the end. The
question as to whether Balaam ’s ass is introduced through a comic
13 See the following rubric in the list of dramatis persona? : Balaam, super
asinam, curvus, barbatus, palmam tenens, calcaribus urgens.
13 See the sequence Quem non prevalent, printed by J. Kehrein, Lateinische
Sequenzen des Mittelatters, Mainz, 1873, pp. 41-42. See Meyer, p. 52.
14 See the sequence Epiphaniam Domini canamus , printed by Kehrein, pp. 40-
41. See Meyer, p. 52.
15 In connection with the dramatic development of the episode of Balaam and
his ass, Sepet (p. 35) suggests three stages of the Ordo Prophetarum:
(1) Le drame primitif, immediatement sorti du sermon, et d’oti Balaam est
absent.
( 2 ) Le drame de seconde formation, oil Balaam apparait et se borne
reciter sa prophetie.
( 3 ) Le drame de troisieme formation, oh la prophetie de Balaam est devenue
un petit drame dans le grand.
Sepet’s “drame primitif” is exemplified, presumably, in the ordo of Limoges,
and his “drame de troisiSme formation,” in the Festum Asinorum of Bouen, to
be considered below. In so far as Sepet’s own evidence is concerned, the
“drame de seconde formation” must be considered a conjecture. Chasles now
suggests (p. 127) that the Ordo of Laon, — unknown to Sepet,— exemplifies the
drame de seconde formation.” I cannot accept Chasles’ suggestion, for the
Balaam of the Laon Ordo does not “confine himself to the reciting of his
prophecy.” The Balaam episode of Laon is not essentially different from that
of Rouen. The Rouen episode is merely longer.
48 Wisconsin Academy of Sciences , Arts , and Letters.
impulse, I do not undertake to answer. The evidence is not posi¬
tive.16
Since the text of the Laon play, like that from Limoges, provides
no evidence as to its position in the liturgy, we may seek indirect
evidence elsewhere. With the Laon text we may conveniently as¬
sociate the following description of the Ordo Prophetarum at
Tours :17
Ordo Prophetarum
Ad Matutinas cantant duo & duo versus Invitatorii & antiphonas
similiter, & Hymnos sicut ad Vesperas. Antequam legatur prima Lectio
accenditur primus circulus turris David, & duo corruptores cantant: Sal-
vetur noster circulus. Post dicuntur versiculi & prosse resp. in medio
chori in rota, & cantant responsoria duo et duo, cum Gloria in choro, ex-
ceptis tertio, sexto, & nono, quos cantent juvenes in pulpito candelis ac-
censis, & omnes a capite repetuntur. Dicto versiculo tertii Nocturni, ac¬
cenditur totum luminare, & veniunt Prophetse in capitulo revestiti, & post
cantant insimul Lumen Patris, & clericus solus dicit In g audio; & post
legitur septima Lectio. Post nonam Lectionem ducunt Prophetas de
Capitulo ad portam Thesaurarii cantilenas cantando, et post in chorum,
ubi dicunt cantori prophetias, et duo clericuli in pulpito cantando eos ap¬
pellant. Post dicitur nonum18 in pulpito.
In Laudibus incipiunt antiphonas duo & duo . . .
Ad ,Missam sunt in cappis sericis, & cantant juvenes duo & duo, &
Epistolam in pulpito, & alia dicunt sicut in liibro reperies.
Post prandium debent chores ducere in claustro in supelliciis donee
Ecclesia aperiatur, & totum luminare accendatur. Quo facto, incipiunt
Nonam & Vesperas; faciunt totum sicut ad primas,19 & dictis psalmis &
antiphonis, ducunt ad portam Thesaurarii Prophetas sicut ad Matutinum
& reducunt in chorum similiter, & habent clerici virgas plenas candelis
ardentibus. Vocant eos clerici duo sicut ad Vesperas. Post dicitur
Stirps Jesse, & cantant juvenes versum in pulpito cum Gloria , & post in
choro repetitur, & hunt pneumse. Post dicitur Regens gubernansque, &
hunt pneumse. Post levat se cantor super formam, cappam habens super
humeros, & incipit antiphonam, & dicit ter Deposuit, baculum tenens, &
si baculus capitur, Te Deum laudamus incipietur, & post versus p salmi
Esurientes cum antiphona & oratione. Post vadunt ad pedes cantando
r<esponsorium> Videbant, & antiphonam Tu principatum, post de B. Mar¬
tino antiphonam Confessor Domini cum Magnificat & oratione; & redeundo
Jubilemus sicut ad primas Vesperas.
16 Meyer (pp. 52-53) declares unhesitatingly for a comic intention.
17 E. Martene, Tractatus de Antiqua Ecclesice Disciplina , Lyons, 1706, pp. 106-
107. Mart&ne describes the manuscript as follows: “Turonensis S. Martini
ecclesias Rituale vetus ante annos 400 scriptum. Ex ms. ejusdem ecclesise.”
18 1, e. nonum < responsorium > .
19 1, e. ad primas <Vesperas)>.
Young— -Or do Prophet arum.
49
At Tours the Ordo took place in Matins of the Circumcision
(Jan. 1), after the reading of the ninth lectio and before the sing¬
ing of the ninth responsory.20 The prophetce entered the choir dur¬
ing the singing of certain cantilence. The summonses to the indi¬
vidual prophetce were delivered by duo clerici in pulpito, and the
several prophecies seem to have been addressed to a sort of presid¬
ing cantor. The repetition of the ordo at Vespers is mentioned,
but not described in detail. Although the actual utterances of the
participants are not known, the nature of the performance must
have been similar that of the Laon play. We may safely infer that
the prophetce were specifically impersonated, and that they spoke in
the formal sequence already familiar to us. It may be that at
Matins one or more of the preceding lectiones were supplied by the
pseudo-Augustinian sermon, and that the prophetce were introduced
after the final lectio by way of didactic illustration. But this is the
merest conjecture.
If, then, the Ordo Prophetarum at Tours resembles the Laon ver¬
sion in general procedure, one may reasonably conjecture a similar¬
ity in liturgical association. It may well be that the Laon Ordo
Prophetarum , — and the Limoges version also, — is to be associated
with the liturgy of the Circumcision. This conjecture, indeed,
receives further support from the circumstances of the Ordo
Prophetarum which we are to consider next.
VI
The two versions of the Ordo Propheta)rum already examined
closely resemble the pseudo-Augustinian lectio in the number and
names of the prophets presented. To the list of the lectio the
Limoges version adds only Israel, and the Laon version, only
Balaam. The possibilities in the way of addition are fully recog¬
nized only in such a play as the following, from the cathedral of
Rouen d
20 See the liturgical arrangement outlined above, p. 23.
XI base my text upon Rouen, Bibliotheque de la Ville, Ms. 384 (olirn Y. 110),
Ordinarium Rothomagense ssec. xiv, fol. 33r-35r, with variants from ibid.,
Ms. 382 (olim Y. 108), Ordinarium Rothomagense ssec. xv, fol. 31v-33r, and
from Paris, Bibl. Nat., Ms. lat. 1232, fol. 26r— 27r (manu saec. xvii). These are
the only manuscript sources known to me. I know of no previous edition in
which all three manuscripts are used. Sepet (p. 28) and Petit de Julleville
(Les My stores, I, 37) erroneously report that we possess no extant manuscript
sources at all. DuMeril mentions (p. 181, note) all three manuscripts as con-
4— S. A. L.
50
Wisconsin Academy of Sciences , Arts , and Letters.
<ORDO PROPHET ARUM >
Nqta, Cantor: Si Festum AsiNORim fiat, processio2 ordinetur post
Terciam; si non fiat Festum, tutic fiat procEssto ut prenotatur.3
Ordo PROCESsionis AsiNOR^m secunmiM RoTHOMAGensem Vsum.4
Tercia cantata, paratis ProPHeiis iu xta suum ordiwem, fornace in
MEDIO NAUIS ECCLesIE LINTHEO et STUPIS CONSTITUTA, PROCESSIO MOUEAT DE
CLAUSTRO, et DUO CuemCI DE iia SEDE IN CAPPIS PROCESSIONEm RECANT, HOS
versus canentes:5
Gloriosi
Et famosi
< Regis festum
Celebrantes
Gaudeamus>.
Chorus :
Gloriosi
<Et famosi
Regis festum
Celebrantes
Gaudeamus>.
'Versus:
Cuius ortum
<Vite portum
Nobis datum
Predicantes
Habeamus>.
taining the Ordo Prophetarum, but he does not edit the text. Meyer speaks
(p. 51) of the l;ext as found “in mehreren Handschriften des 14. und 15. Jahr-
hunderts in Rouen.” A general bibliography of the manuscripts and prints
bearing upon the liturgical plays of Rouen is given by the present writer in
Modern Philology, Vol. VI (1908), pp. 224-227. There can be little doubt that
Ms. 1232 is a copy of Ms. 384 ; and the copy seems to be surprisingly accurate
(See my variants). If, as Gast§ supposes (p. 2), Ms. 382 is a copy of Ms. 384,
the copyist departed freely from his model (See my variants, and Modern Philol¬
ogy, Vol. VI, pp. 224-225). Gaste provides (pp. 1-4) important notes on Mss.
384 and 382. None of the three manuscripts mentioned above provides music.
The Rouen Ordo Prophetarum (“Festum Asinorum”) has been previously
edited twice: (1) by Du Cange, Glossarium Medics et Infimce Latinitatis, Vol.
Ill, Niort, 1884, pp. 460-461 (and in earlier editions of this work), from an
unidentified “Ordinarius MS” of Rouen (See Modern Philology, Vol. VI, p. 226) ;
and (2) by A. Gaste, Les Drames liturgiques de la Cathedrale de Rouen, Evreux,
1893, pp. 4-20 (Concerning earlier forms of Gaste’s monograph see Modern
Philology, VI, 201), from Rouen Ms. 384, with some of the variants from Rouen
Ms. 382. I collate Gaste’s text throughout (G), taking into account his “Er¬
rata et emendanda” (p. 81).
2 processio] processio et (Ms. 382).
3 prenotatur] nunc prenotatur (G). As to the reference in the words ut
prenotatur see below, pp. 66—68.
4 Rothomagensem usum] usum Rothomagensem (Ms. 382).
5 hos versus canentes] cantantes hos versus (Ms. 382).
Young — Or do Prophet arum.
Chorus:
Versus :
Chorws :
Gloriosi.
Quem futurum
<Regnaturum
Prophetico
Admonitu
Nuntiamus>.
Gloriosi.
Versus:
Impior um
Iudeorum
<Corda negant
Regnaturo
Sua lege>.
Chorus :
Gloriosi
Versus:
Et gentiles
<Prius viles
Convertuntur
Majestate
^Etherea>.
Chorus :
Gloriosi.
Versus:
Sed Iudei
<Facti rei
Condemnantes
Sacrum regem
Damnabuntur>.
Chords :
Gloriosi.
Versus:
Israeli6
Infideli7
<Jam Maria
Natus scitur
Hie adesse>.
Chorus :
Gloriosi.
Israeli] Israel (G).
Infideli] Infideles (Ms. 382).
52 Wisconsin Academy of Sciences, Arts, and Letters.
Versus:
Gentes unde.8
Tunc processio in medio ecclcsie stet. Et vi Iudei sint ibi
PARATI, ET EX ALTerA <fOl. 33v> PARTE SEX GENTILES, ET
OMnes Gentes uocent ita Uocatores:
Onmesi gentes
<Congaudentes
Dent cantus laetitise>;
Deus homo
Fit9 <de domo
David natus hodie>.
HlC UERTANT SE UOCATORES AD lUDEOS DICENTES :
O Iudei
Uerbum Dei
<Qui negastis hominem>.
Versus:10
Vestre legis
Testes* 11 <regis
Audite per ordinem>.
Iudei respondeant:
Nos mandatum uobis.
Vocatores ad Gentiles dicant:
Et uos, gentes,
Non credentes
<Peperisse Virginem,
Versus:12
Vestrse legis13
Documentis
Pellite caliginem>.
Gentiles respondeant:
Deum uerum
Regem rerum.14
Vocatores prius Moysen,15 ita dicentes:
Tu, Moyses,
Legislator,
<Huc propinqua
Et de Christo
Prome digna>.
8 unde] Iudae (Ms. 1232). I can make no suggestion for the completion of
this stanza. The expansions suggested for various other initia, above and be¬
low, are made from the several versions of the Ordo Prophetarum printed in
the present study.
9 Deus homo fit] Dominus homo sit (Mss. 384, 382, and 1232).
10 Versus] omitted (G).
11 testes] testem (G). Through a typographical error Gast6 gives the im¬
pression that the words Vestre legis testes are not in Ms. 384.
12 Versus] omitted from the restoration (G).
13 legis] gentis (G).
14 Deum uerum regem rerum] Deum regem verum (Ms. 382).
15 Moysen] omitted (Ms. 382).
Young — Or do Prophet arum,
53
Tunc Moyses, tenews tabulas legis APerrAS, indutms alba et
CAPPA, et CORNUTA FACIE, BARBATUS, TENENS UIRGAM IN
manu, dicat:
Vir post me veniet exortus.
Hoc DICTO VOCATORES EUm DUCANT ULTRA FORNACEWl DICENTES :
Iste cetus
Psallat letus
<Error vetus
Condemnetur>.
Chorus :
Quod Iudea
<Perit rea
Hec chorea
Gratulatur>.
VOCATORES DICANT AD AMOS :
Amos mentis.
Tunc Amos, senex barbatus, spicam tenens, dicat:
Ecce dies uenient.16
Vocatores :
<Iste cetus>.
Chorus :
Qwod Iudea.
Vocatores dicant Ysaye:
Ysaias, verbum qui scis,17
<Veritatem cur non dicis>?
Ysaias, barbatus, alba indutus, per MEDiAm18 frontem rubea
STOLA10 DISTINCTWS,20 DICAT:
Est necesse
Uirga<m> Yesse
<De radice provehi:
Flos deinde
Surget inde
Qui est Filius Dei>.
Vocatores :
Iste cetus.
Chorus :
Quod Iudea.
Vocatores ad Aaron:
Aaron, doce populum.
16 See Amos, viii, 11 : Ecce dies veniunt, dicit Dominus, et mittam famem in
terram ; non famem panis, neque sitim aquae, sed audiendi verbum Domini.
17 scis] scit (Mss. 384, 382, and 1232, G).
18 mediam] medium (Ms. 382). 1
19 Stola] Stolla (Ms. 382).
20 distinctus] discrinitus (Mss. 384, 382, and 1232, G).
54
Wisconsin Academy of Sciences , Arts , and Letters.
Aaron, ornatus pontificalibws indumentis et mitra,21 bar-
BATUS, TENENS FLOREM DICAT *.
Virga Iesse florida.22
Vocatores:
Iste cetus.
Chorus:
Quod Iudea.
Vocacio IfterEMiE*.
Qui vocaris, Ifteremias,23
<Dic de Christo prophetias>.
I7ierEMiAS SAcerDOTALi ho-Situ ornatus et barbatus24 tenens
ROTULUM DICAT:
Sic est,
Hie est,
Dews noster;
<Sine quo non erit alter>.
VocATores:
Iste cetus.
Chorus :
Qwod Iudea.
VocAcio Danielis:
Daniel, indica
Uoce prophetica
<F'acta dominica>.
Daniel, indutus uiridi tunica, iuuenilem uultum Ha&eNS,
tenens spiCAm, dicat:
Sanctus sanctorum ueniet,
<Et unctio deficiet.>
VocATores ;
Iste cetus.
Chorus:
Quod Iudea.
Vocacio Abacuc : 25
Abacuc,25 regis celestis
<Nunc ostende quod sis testis>.
21 mitra] mittra (Ms. 384).
22 See Num. xviii, 5-8 : Quern ex his elegero, germinabit virga ejus . . .
fueruntque virgse duodecim absque virga Aaron . . . Sequenti die regressus
invenit germinasse virgam Aaron in domo Levi ; et turgentibus gemmis eruperant
(lores, qui, foliis dilatais, in amygdalas deformati sunt.
23 Iheremias] Omitted (Ms. 382).
24 ornatus et barbatus] indutus et ornatus barbatus (Ms. 382).
25 Abacuc] Abacuth (Ms. 382); Habacuc (Ms. 1232).
Young— Or do Prophet arum.
55
AbACUC,25 SENEX CLAUDUS, DALMATICA INDUTUS,26 HafteNS IN
PERAM RADICES, et LONGAS PALMAS HG&CNS UNDE GENTES
per<fol. 34r>cuciAT, comedews, dicat:
Opus tuum
Inter duum27
<Latus animalium
Ut cognovi
Mox expavi
Metu mirabilium>.
Vocatores :
Iste cetus.
Chorus :
Quod Iudea.
Duo missi a rege Balac28 dicant:
Balaam,29 veni et fac.
Tunc Balaam,29 ornatus, seders super asinam, uabeNs cajl-
CARIA RETINEAT LORA et CALCARIBWS PerCUCIAT ASINAM. ET
QUIDAm IUUENIS HafteNS alas,30 tenens gladium, obstet
ASINE. QuiDAm SUB ASINA DICAT:
Cur me cum calcaribws miseram sic leditis?
Hoc DICTO, ANGeLWS EI dicat :
Desine regis Balac preceptum perficere.
VocAcio Balaam:31
Balaam,31 esto uaticinans.
Tunc Balaam31 respondeat:
Exibit ex Jacob rutilans32 <nova stella,
Et confringet ducum agmina
Regionis Moab maxima potentia>.
VocATores :33
Iste cet us.
Chorus :
Quod Iudea.
VocAcio Samuelis:
Accede, Samuel.
Samuel, religiose indutws, dicat : 34
In Israel faciet rex verbum.35
26 indutus] Omitted (Ms. 382).
27 Inter duum] in triduum (Ms. 382). The whole speech of Habakkuk Gaste
prints as follows : <Expectavi, mox expavi, metu mirabilium^> Opus tuum
inter duum corpus animalium.
28 Balac] ballat (Ms. 382).
29 Balaam] Ballaam (Ms. 382).
30 alas] alias (Ms. 382).
31 Balaam] Ballaam (Ms. 382).
32 rutilans] rutillans (Ms. 382). Gaste does not expand this initium.
33 Vocatores] Vocant (Ms. 382).
34 dicat] doceat (Ms. 382).
35 See 1 Reg. iii, 11 : Et dixit Dominus ad Samuelem : Ecce ego facio verbum
in Israel, quod quicumque audierit, tinnient ambse aures ejus.
56
Wisconsin Academy of Sciences, Arts, and Letters.
Vocat ores :36
Iste cetus.
Chorus :
Quod Iudea.
VocACio Danin:
Die tu, Dauid, de nepote
Causas <que sunt tibi note>.
Dauid, ornatus reg alibis oreamewtis, dicat :
XJniuersus
Grex cowuersus
Adorabit Dominion,
<Cui futurum
Serviturum
Omne genus hominum>.
Vocat ores :
Iste cetus.
Chords :
Quod Iudea.
Vocacio Osee37
Auffer,38 Osee,37
Plebi Hebree
Cecitatem.
Osee,37 barbatus, dicat:
Deus nuweiauit de filio Dauid in presenti.39
Yocatores : 40
Iste cetus.
Chorus :
Quod Iudea.
VocACio Iohelis:41
Iobel,42 leua uocem cum ceteris.
lOHEL,42 DIuerSUM HUbeNS CULTUM, et BARBATUS DICAT I
Effur-dam de spiriZu meo, dicit dominus.43
Vocatores : 44
Iste cetus.
Chords :
Quod Iudea.
36 Vocatores] Vocant (Ms. 382).
37 Osee] Ozee (Ms. 382).
38 Auffer] aufer (Ms. 382).
39 See Os. iii, 5 : Et post hsec revertentur fllii Israel, et quserent Dominum
Deum suum, et David regem suum ; et pavebunt ad Dominum, et ad bonum
ejus, in novissimo dierum.
^Vocatores] Vocant (Ms. 382).
41 Iohelis] Ioelis (Ms. 382).
42 Iohel] Ioel (Ms. 382).
43 See Joel ii, 28: Et erit post hsec: Effundam spiritum meum super omnem
carnem ; et prophetabunt filii vestri et filise vestrae ; senes vestri somnia som-
niabunt, et juvenes vestri visiones videbunt.
44 Vocatores] Vocant (Ms. 382).
Young— Or do Prophet arum.
57
Vocacio Abdie:
Fac, Abdia, preconia uenturi iSaluatoris.
Abdias, DiuersuM Habeas cultum, barbatus, dicat:
Et in monte Syon saluacio erit.45
YocATores:46
Iste cetus.
Chorus :
Quod Iudea.
VocAcio Ione:
De persona
Xp£s£i, Iona,47
Que sunt in te mistica?
IONAS, CALUUS, AEBA INDUTUS, DICAT:
O Iudei,
Huius rei
Signum, genus fatuum.48
VocATores :
Iste cetus.
Chords:
Qwod Iudea.
Vocacio Michee:
Effice, Michea, qwod credat plebs.
Micheas, Diuersum cui/rum Ha&eNs, barbatus, dicat:
Descendet Domimts
Cui no n est terminus.49
VocATores ;
Iste cetus.
Chords :
Qwod Iudea.
VocAcio Naum:50
Naum,51 plebi Iudaice die.
45 See Abdias, 17 : Et in monte Sion erit salvatio, et erit sanctus ; et possidebit
domus Jacob eos qui se possederant.
^Vocatores] Vocant (Ms. 382).
47 Iona] Omitted (Ms. 382).
48 See Matt, xii, 39-40: Qui respondens ait illis : Generatio mala et adultera
signum quserit et signum non dabitur ei, nisi signum Jonae prophetae. Sicut
enim fuit Jonas in ventre ceti tribus diebus et tribus noctibus, sic erit Filius
hominis in corde terrae tribus diebus et tribus noctibus.
49 See Michceus i, 3 : Quia ecce Dominus, egredietur de loco suo ; et de¬
scendet, et cal cab it super excelsa terrae ; Mich, v, 2 : Et tu, Bethlehem Ephrata,
parvulus es in millibus Juda ; ex te mihi egredietur qui sit dominator in Israel,
et egressus ejus ab initio, a diebus aeternitatis ; Mich, v, 4 : Et convertentur,
quia nunc magnificabitur usque ad terminos terrae.
50 Naum] Nain (Ms. 382) ; Nahum (Ms. 1232).
61 Naum] Omitted (Ms. 382); Nahum (Ms. 1232).
58
Wisconsin Academy of Sciences , Arts , and Letters.
Nau m50 <fol. 34v>_senex RESjPONDeat:
Super montes62 euucmgelizantis.63
Vocatoees:
Iste cetus.
Chorus:
Quod Iudea.
VocAcio Sophonie:
Esto nobis, Sophonia.
SOPHONIAS,54 BABBATUS, DICAT I
In medio tui Syon55 rex regnabit.05
VocATores :
Iste cetus.
Choeus :
Qwod Iudea.
Vocacio Aggei:
Audiamus os Aggei ut exponat.
Aggeus, senilem57 uultum gebens, dicat:
Veniet cunctis gentibws rex desideratissimus.68
VocATores ;
Iste cetus.
Chorus:
Quod Iudea.
VocAcio Zachaeie:
Veni, Zacbaria,59 fili Barachie.
ZACHARIAS, BARBATUS, DICATI
En rex tuus uenit tibi60 iustus, filia Syon.61
VocATOres;
Iste cetus.
Chorus :
Qwod Iudea.
Vocacio Ezechielis:
Profer nobis, Ezechiel, aduentttm.
52 montes] montem (G).
63 euuangelizantis] euuangelisantis (Ms. 382). See Nahum i, 15: Eqce super
montes pedes evangelizantis, et annuntiantis pacem.
54 Sophonias] Sophonia (G).
53 Syon] Sion (Ms. 382).
56 See Sophon. iii, 14 : Lauda, filia Sion ; jubila, Israel ; lsetare, et exsulta in
omni corde, filia Jerusalem.
57 senilem] senillem (Ms. 382).
58 See Agg: ii, 8 : Et movebo omnes gentes, et veniet desideratus cunctis
gentibus ; et implebo domum istam gloria, dicit Dominus exercituum.
59 Zacharia] Zdcharie (G).
60 tibi] omitted (G).
61 See Zach. ix, 9 : Exsulta satis, filia Sion ; jubila, filia Jerusalem ; ecce rex
tuus veniet tibi justus, et salvator ; ipse pauper, et ascendens super asinam
et super pullum filium asinse.
Young — Or do Prophet arum.
59
RespONSIO EIUSDEM I
Per clausam ianuam rex intrabit solus.62
VocATores ;
Iste cetus.
Chorus:
Qt^od ludea.
VocAoio Malachie:
Palam nobis refer, Malachia.63
RespoNSio EiwsDem :
Scimus bee04 dieentem Deum.65
VocATores ;
Iste cetus.
Chorus :
Quod ludea.
VocAcio Zacharie patais S ancti IoHUfmis:
Zacbaria, os aperi.
Ipse, ornatus quasi Iudeus, dicat:
Per uiscera dulcifflue66
Dei mi sericordie.61
Vocat ores :
Iste cetus.
Chorws:
Qwod ludea.
VocAoio Elyzabeth:68
Illud,69 Elyzabeth,68 in medium
<De Domino profer eloquium>.
In persoNA alba, quasi pregnans, dicat:
Quid est rei
Q%od me70 mei
62 See Ezech. xliv, 1—3 : Et convertit me ad viam portae sanctuarii exterioris,
quae respiciebat ad orientem ; et erat clausa. Et dixit Dominus ad me : Porta
haec clausa erit ; non aperietur, et vir non transibit per earn, quoniam Dominus,
Deus Israel, ingressus est per earn ; eritque clausa principi.
63 nobis refer Malachia] refer nobis Mathia (Ms. 382).
64 hec] hoc (G).
65 See Malach. iii, 1 : Ecce ego m'itto angelum meum, et praeparabit viam ante
faciem meam ; et statim veniet ad templum suum Dominator quern vos quaeritis,
et angelus testamenti quern vos vultis. Ecce venit, dicit Dominus exercituum.
66 dulcifflue] dulciflue (Ms. 382) ; dulcifluae (Ms. 1232).
67 See Luc. i, 7 8 : Per viscera misericordiae Dei nostri, in quibus visitavit nos,
oriens ex alto. It will be observed that the prophecy assigned to Zacharias in
the Rouen version is not the same as that (Luc. i, 76) assigned to him in the
pseudo-Augustinian lectio. See above, p. 8.
68 Elyzabeth] Helizabeth (Ms. 382) ; Elisabeth (Ms. 1232).
69 Illud] Illud et (Ms. 1232).
70 me] Omitted (Ms. 382).
60
Wisconsin Academy of Sciences, Arts , and Letters.
< Mater heri visitat?71
Nam ex eo
Ventre meo
Laetus infans palpitat>.
VocATores :
Iste cetns.
Chords :
Quod Iudea.
VocAcio S ancti lonanms Bapt iste;
Da, Baptista,
Uentris72 cista
Clausus.
<Quos73 dedisti
Causa Christi
Plausus.
Cui dedisti gaudium,
Profer et testimonium >.
Ipse, nudus pedes, tenens textuw, dicat:
Venit talis
Sotularis
Cuius non sum <etiam74
Tam benignus
Ut sim dignus
Solvere corrigiam>.
Vocatores:
Iste cet us.
Chords :
Qwod Iudea.
VocACio Symeonis :
Quid dixisti, Symeon, cum in tua?
Symeon, senex, respondeat:
Dei nostri, Saluatoris, conspexerunt75 ocwli.
Vocat ores:
Iste cetus.
Chorus:
Qnod Iudea.
VocAcio Uirgilii :
Maro, Maro,7S uates gentilium,
Da Xpisfo < testimonium >.
71 Mater heri visitat] Gaste prints these words as if they were present in
Ms. 384.
72 Uentris] Ventrix (Ms. 382).
73 Quos] Quod (G).
74 etiam] Printed as if present in Ms. 384 (G).
75 conspexerunt] surrexerunt (Ms. 382). The entire speech of Simeon Gast§
prints as follows : <Nunc me dimittas, Domine, finire vitam in pace, quia>
Dei nostri salvatoris conspexerunt oculi Quern misisti (in hunc mundum) pro
salute populi>.
76 Maro Maro] Mars Mars (Ms. 382) ; Maro (G).
Young — Ordo Propketarum.
61
VlRGIUUS, IN IUUENILI Ha&ITU R6N6 ORNATWS, RESPOND eat:
Ecce polo demissa solo <nova progenies est>.
VocATores :
Iste cetws.
Chorws:
Qwod Iudea.
Interim Najbugodonosor,77 quasi Rex paratws, <fol. 35r> os-
iCNDENS YMAGINEW, DUOBWS ArMATIS DICAT :
Hue venite, uos Armati.
Tunc Armati ostendant ymaginew prius78 dicentes:
Regi gratum
Famulatum.
Interim ostendant79 ymaginem tribes Pueros dicentes:
Huic sacro simulacro.
Tunc Pueri ymaginem respuentes dicant:
Deo soli
Digno coli.
Hoc audito Armati Pueros Regi ducant,80 dicentes :
Quia ritum
Stabilitum
Non timetis?
Tunc ostendantS1 Regi Pueros,82 dicentes:
“Rex, tua saluentur.
Tunc Rex iratus dicat:
Ergo tales assumawtwr.
Tunc Armati ducentes Pueros ad fornacem dicentes:83
Reos digne84
lam in igne.
Tunc mittantur Pueri in fornace<m>, et accendatur. At
illi, Facti liberi, dicant:
Benedictus es, Domine Dews, et cetera.
Rex, hoc audiens, admirans dicat:85
En quid cantant86 illi tres?
Armati87 dicant:
Deum laudant.88
77 Nabugodonosor] Nabugodenosor (Ms. 382); Nabuchodonosor (Ms. 1232).
78 prius] pueris (Mss. 384 and 382, G). The reading prius (adopted from
Ms. 1232) results in the avoiding of a repetition of the summons to the Pueri , —
a repetition which may, or may not, be objectionable. See Gaste, p. 17, note.
79 ostendant] ostendat (Ms. 382).
80 regi ducant] ducant regi (G).
81 ostendant] respondeant (Ms. 382).
82 Regi Pueros] Pueros Regi (G).
83.83 omitted from Ms. 382.
84 Reos digne] Digne reos (Ms. 382).
85 dicat] hoc dicat (G).
85 cantant] cantent (Ms. 382).
87 tres Armati] Tunc Armati (Ms. 1232).
88 laudant] laudent (Ms. 382).
62
Wisconsin Academy of Sciences, Arts, and Letters.
Tunc Uocatores dicant Regi:
Puerum cum pueris,
Nabugodonosor,89
<Cum in igne videris,
Quid dixisti>?
Rex FORNACEm ostendens90 dicat:
Tres in igne positi pueri89
< Quarto gaudent comite liberi>.
Vocat ores :91
Iste cetus.99
Chords :
Quod Iudea.
Vocacio Sibille:
Tu, Sibilla,93
Uates ilia,89
<Dic adventum judicis.
Die signum judicii>.
Sibilla,94 coronata et muliebri hcj&itu ornata, dicat:
Iudicii signum: Tellus sudore <madescet>.95
VocATores :91
Iste cetus.
Chorus :
Qitod Iudea.
Quo finito omms ProPHetE et Ministri in pulpito cantent98
hos versus:
Ortum97 predestinacio
Poruo Sabbaii spacio.
Hoc finito Cantor incipiat ad introitum chori r esponsorium:
Confirmatum est cor Uirgims.98
Prophete et iMinistri regentes cHORum secunouM. suum ordi-
?IEM INCIPIANT AD MlSSAM OFFidUM
Puer natus" <est nobis >. 100
89 Gaste does not expand these initia.
90 fornacem ostendens] ostendens fornacem (Ms. 382).
91Vocatores] Vocant (Ms. 382).
92 cetus] Omitted (Ms. 382).
93 Vocacio Sibille Tu Sibilla] Vocatis Sibila En Sibila (Ms. 382).
94 Sibilla] Sibila (Ms. 382).
95 The entire speech of Sibilla Gaste prints as follows : Judicii signum : tellus
sudore madescet <E coelo rex adveniet per secla futura, Scilicet in carne
praesens ut judicet orbem>.
"cantent] cantant (Ms. 382).
97 Ortum] (H)ortum (G).
98 cor Uirginis] Omitted, (Ms. 382).
"natus] natus est et cetera (Ms. 382) ; natus est (G).
100 Puer natus est nobis are the opening words of the Introit for the Mass of
the Circumcision (Jan. 1), as well as for the third Mass of Christmas.
Young — Or do Prophet arum.
63
Kyrie et GLoriA festiue.101
Onatio: Deus, qui nobis nati salua<toris>.102
Nulla MEMoria fiat.103
Epistona: Multipharie.
Tees de secunvA sede r esponsum: Viderunt omwes
fi<nes>.104
Tees de maioribws : Alleluia,.
Versus: Multipharie.
SvQueistia: Letabuwdus.
EuaNGmwM: Postquam consummati sunt.
Credo in vNum.105
Off ertorium: Tui sunt celi.106
Secreta: Presta, quesumus, Domine.
Pref atio: Quia <fol. 35^> per incarnati.107
Sanctus et Agnus festiue.108
Com munio: Viderunt onmes.
Postcom munio: Presta,, quesumus, Domine, ut q%od109 sal-
uatom.110
The action of the Rouen Festum Asinorum runs, then, as follows :
Immediately after Terce the procession of prophetce, led by two
clerics, leaves the cloister and advances toward the nave, two
clerics singing antiphonally with the choir the Gloriosi et famosi.
In the middle of the church, — presumably at the junction of nave
and transepts, — the procession makes a station between six Jews
on one side and six Gentiles on the other. After exhorting both
groups to rejoice in Christ’s birth (Omnes gentes), the Vocatores 111
address in turn the recalcitrant Jews (0 Judcei) and the unbeliev¬
ing Gentiles (Et vos, Gentes), receiving a response from each.112
101 Gaste’s text and the text in Ms. 1232 end here. Ms. 1232 adds the follow¬
ing note : Haec supra scripta deprompta fuerunt ex antiquo codice Rituali
S. Rotom. Ecclesise in Festo Circumcisionis Domini. There can be little doubt
that the manuscript referred to is Rouen Ms. 384.
102 nobis nati salua<toris> ] nobi nati (Ms. 382).
103 fiat] Omitted (Ms. 382).
104 responsum : Viderunt omnes fi<nes>] Graduale : Viderunt. Versus:
Notum fecit (Ms. 382).
105 in vnum] Omitted (Ms. 382).
108 celi] Omitted (Ms. 382).
107 Quia per incarnati] Et communicantes, de Natiuitate (Ms. 382).
108 Sanctus et Agnus festiue] Omitted (Ms. 382).
109 quod] qui (Ms. 382).
110 Followed immediately by the rubric : Ad vi.
111 The number of Vocatores is uncertain.
112 I am unable to complete the initia of these responses.
64 Wisconsin Academy of Sciences, Arts, and Letters.
The Vocatores now call upon Moses to testify concerning Christ,
and after he has delivered his testimony, they conduct him to a
position beyond the furnace that has been constructed in the middle
of the nave. As they escort Moses to his new position they sing
Is'te coetus, to which the choir responds Quod Judcea. Next in order
Amos is appropriately summoned, utters his prophecy, and is con¬
ducted to a position beside Moses. Similarly are treated Isaiah,
Aaron, Jeremiah, Daniel, Habakkuk, Balaam, Samuel, David,
Hosea, Joel, Obadiah, Jonah, Micah, Nahum, Zephaniah, Haggai,
Zechariah, Ezekiel, Malachi, Zacharias, Elizabeth, Saint John the
Baptist, Simeon, Virgil, Nebuchadnezzar, and the Sibyl. After the
Sibyl has been escorted to her place among the prophets, the whole
company of prophets and ministers113 unite in singing in pulpito 114
the prose Hortum prmdestinatio . The procession now advances
into the choir for Mass, the prophets and ministers beginning the
Introit ( Officium ) and ruling the choir.
The most obvious difference between the Rouen play and the
versions previously considered is, of course, the notable increase in
the number of the prophets.115 Against the thirteen personages
summoned in the lectio, and in the Limoges and Laon versions, the
Rouen text provides twenty-eight, in non-chronological order, in¬
cluding all of the major and minor prophets of the Vulgate. Of
these twenty-eight, fourteen116 appear in none of the versions al¬
ready considered, whereas, excepting Israel,117 all the prophets of
the shorter versions are present in the Rouen play. This increase
in the number of prophets need occasion no surprise, for such an
accretion is natural to any literary development, the names of the
added personages lay ready to hand in the Vulgate, and the sermon-
izer of the pseudo-Augustinian lectio himself suggests the process:
Quod si velim ex lege et ex prophetis omnia quse de Christo dicta sunt
colligere, facilius me tempus quam copia deseret.118
113 1 infer that these ministri are the persons who assist the prophetce in
performing the play.
114 Pulpitum may mean the rood-loft over the choir-screen. See Gast§,
pp. 19-20.
115 Sepet (pp. 29-83), having no knowledge of the Laon play, compares the
text of Rouen only with that of Limoges.
118 Amos, Aaron, Samuel, Hosea, Joel, Obadiah, Jonah, Micah, Nahum,
Zephaniah, Haggai, Zechariah, Ezekiel, and Malachi.
117 Israel is present only in the Limoges version. See above, pp. 26, 32.
118 See above, p. 7.
Young — Or do Prophetarum.
65
Of the fourteen prophets that the Rouen version has in common
with the shorter versions, some nine119 use utterances that we have
already seen in both the Limoges and the Laon texts. The other
five present special cases. Balaam, found only in Laon and Rouen,
uses the same prophecies in the two versions. The Moses and
Simeon of Rouen, — if we may judge from their brief initia, — de¬
liver speeches unlike those of Limoges and Laon in text, but like
them in ultimate derivation from the lectio. Zacharias appears
only in. the lectio and in the Rouen play, and in the two versions
his utterances are not the same. The Rouen Nebuchadnezzar utters
the prophecy that we have already seen in the Laon version.
The peculiar dramatic interest of the Rouen play lies, however,
not in such details as these, but in two special episodes, centering
respectively in Balaam and Nebuchadnezzar.120
The dramatization of the story of Balaam is brief, but notably
more comprehensive than what we have seen in the Laon play.
Tn the spoken text of the Rouen play, to be sure, less attention is
given to the asina; but on the other hand specific notice is taken
of the relation of Balaam to King Balak. It is, indeed, two emis¬
saries of this king, rather than the official Vocato'res, who first sum¬
mon Balaam forth, and bid him come to Balak ’s court.121 Balaam
now rides forth upon an ass, vigorously plying his spurs, until the
person concealed under the animal cries out in protest (Cur me
. . . leditis ?).122 Meanwhile a youth costumed as an angel,
with wings and sword, stands in the path and commands Balaam to
cease serving Balak.123 The official Vocatores now call upon
Balaam for his prophecy, and he delivers the utterance that we
have already observed in the Laon play.124
For vivifying the prophecy of Nebuchadnezzar a still more
elaborate action is devised. The mise en scene consists in a furnace
119 Isaiah, Jeremiah, Daniel, Habakkuk, David, Elizabeth, John the Baptist,
Virgil, and Sibyl. See Sepet, pp. 30-32. Sepet (pp. 29-38) makes a detailed
comparison of the Limoges and Rouen versions, coming to the conclusion the
Rouen text is based upon that of Limoges, or upon something similar. It should
be remembered that Sepet did not know the Laon version.
120 See Sepet, pp. 33-36.
121 This summons, of which we have only the opening words (Balaam, veni et
fac), probably included a command that Balaam curse the Israelites. See Num.
xxii, 5-21.
122 Cur percutis me? (Num. xxii, 28).
123 See Num. xxii, 31-33.
124 For the source of the whole action see Num. xxii, 1-35. It will be ob¬
served that the dramatist does not use the Vulgate phraseology.
5 — S. A. L».
66
Wisconsin Academy of Sciences , Arts , and Letters.
constructed of cloth and oakum in the middle of the nave,125 and in
some sort of figure to serve as the golden idol of the Biblical ac¬
count.126 Pointing to the idol, Nebuchadnezzar begins the action,
promptly by ordering two of his soldiers to command the Pueri
(Shadrach, Meshach, and Abednego) to fall down in worship.127
After they have spurned the idol, the three young men are con¬
ducted into the presence of Nebuchadnezzar. Hearing of their
contumacy, the king angrily orders that they be cast into the fur¬
nace. After the order has been executed and the furnace lighted,
the Pueri sing the Benedictus. Aroused by the sound of their
voices, Nebuchadnezzar asks his soldiers what the young men are
singing, and is told that they are praising God. The Vocatores
now intrude into the action, asking the king what he said at this
juncture. Nebuchadnezzar closes the dramatic episode by deliver¬
ing the testimony already known to us from the Laon version.
As to the day on which the Rouen processus was performed
scholars have not agreed. Du Cange,128 Sepet,129 and Chambers130
have assigned the performance to Christmas Day; Gaste131 asso¬
ciates it with the Vigil of Christmas (Dec. 24) ; and DuMeril132
and Chasles133 speak positively for the Feast of the Circumcision
125 Fornace in medio navis ecclesie lintheo et stupis constituta. See above, p. 50.
128 Ostendens ymaginemj ostendant ymaginem.
127 It may be that the Armati deliver the command to the Pueri twice (See
above, p. 61). The general action of this whole dramatic episode is provided by
Dan. iii, 13-92. Except in the Benedictus sung by the Pueri, however, the
dramatist does not use the Vulgate phraseology.
128 Du Cange (G-lossarium medice et infimce Latinitatis, Vol. Ill, Niort, 1884,
p. 460) introduces his text as follows: “Festum Asinorum, cujus Officium, quod
die Christi Natalitio celebratur, ut et nomenclature rationem, accipe ex Ordi-
nario Ecclesiae Rotomagensis Ms. : Nota, Cantor, si ...” I assume that
a similar annotation occurs in the earlier editions of Du Cange, momentarily
inaccessible to me.
129 Sepet (p. 42) writes as follows: Le texte nous indique positivement que la
Procession de Vane faisait partie de l’office du jour de Noel. What textual evi¬
dence Sepet refers to I do not know.
130 Chambers (Vol. II, p. 55) assigns the Ordo to Christmas, rather than to
the preceding day, because “the eeIntroit with which the text concludes is
Puer natus est, which belongs to the Magna missa of the feast-day, and not to
the eve.” Chambers overlooks the fact that Puer natus est is the Introit also
of the Mass of January first. As Villetard has observed (op. cit. infra, p. 47,
note), the entire liturgy of the Circumcision is essentially a compilation from
other feasts, chiefly Christmas.
151 Gaste (p. 4) assigns the dramatic performance to “La veille de Noel”
without reference to evidence of any kind.
132 See DuMeril, p. 181, note. DuM6ril cites no evidence.
133 Chasles writes (p. 126, note 3) as follows: “La procession des Prophgtes
du Christ avait lieu le jour de la Circoncision. Jusqu’S, present on avait pens£,
a la suite de M. Sepet et de M. Gaste, que cette procession se faisait le jour de
Young — Or do Prophetarum.
67
(Jan. 1). There can be no doubt that this last date is the correct
one, and that the other dates have been arrived at only through
neglect of the plain facts of the manuscripts. In Rouen Ms. 384,
for example, which is our chief source for the text of the Festum,
Asinorum, the dramatic text is found in the very midst of the ordo
for the service of January first,134 and is attached directly to the
Mass for that day.135 This attachment is made clear by the rubrics
preceding and following the Festum Asinorum itself. The intro¬
ductory rubric is as follows :
Processio fiat ut in die Natiuitatis Domini excepto quod ad introitum
chori dicatur responsorium: Confirmatum est.
Nota Cantor: Si Festum Asinorum fiat, processio ordinetur post Ter-
ciam; si non fiat Festum, tunc fiat processio ut prenotatur.136
From this rubric it is clear, in the first place, that the Festum
Asinorum might or might not be performed in any particular
year.137 If the Festum Asinorum were omitted, the usual liturgical
processio occurred immediately after Terce and before Mass, the
processio being identical with that of Christmas except that as the
chant at the entrance to the choir (Ad introitum chori) the re-
Noel. On s’appuyait principalement sur l’introit Puer natus est nobis entonnS
par les prophetes. Cela ne prouve rien, puisque cet introit est aussi celui de la
Circoncision . . . Enfin, nous avons trouve dans un ordinaire de Bayeux
(Ms. 121, publie par U. Chevalier, Bib. liturg., t. VIII, p. 75) le r6pons : Con¬
firmatum est, a, la procession de la Circoncision. Or c’est justement ce meme
repons qu’entonne le chantre quand les prophetes sont rentres au choeur. On le
chante, d’ailleurs, encore aujourd’hui, au 2e nocturne des matines de la Circon¬
cision. Les faits semblent bien imposer notre conclusion.”
134 The Cursus and Mass for St. Sylvester (Dec. 31) are found on fol. 31v-32v,
followed immediately by the Cursus and Mass for the Circumcision (Jan. 1) on
fol. 32v-35v. The Festum Asinorum is found on fol. 33r— 35r.
135 The ordo for this Mass is printed above, pp. 62-63. This Mass can be identi¬
fied as that for Jan. 1 by comparison of its basic elements with the same ele¬
ments in the Mass of the Circumcision for the Feast of Fools at Sens (Villetard,
op. cit. infra., pp. 109—117). Further evidence that the Rouen Festum Asinorum
belongs on Jan. 1 is provided by the following note written at the end (fol. 27r)
of the seventeenth-century copy of the dramatic text found in Bibliotheque Na-
tionale Ms. lat. 1232 (see above p. 63) : “Hsec supra scripta deprompta
fuerunt ex antiquo codice Rituali S. Rotom. Ecclesise, in Festo Circumcisionis
Domini.”
138 Rouen Ms. 384 (Y. 110), fol. 33r. The second paragraph is printed at the
beginning of the Festum Asinorum above, p. 50.
137 As to the reason for this option any person may, in the absence of facts,
guess to his heart’s content. Sepet conjectures (pp. 38-40) that the giving of
the performance may have been contingent upon the decorous conduct of the
spectators during the performance of the previous year. Of this conjecture
Chasles (p. 128) very justly observes, “Cette idee nous semble d§nu£e de tout
fondement.”
68 Wisconsin Academy of Sciences , Arts , and Letters.
sponsory Confirmatum est supplanted the antiphon Hodie Christies
natus.138 In other words, when the Festum Asinorum was per¬
formed it merely supplanted the greater part of the normal pro -
cessio, the responsory Confirmatum est being retained ad introitum
chori ,139 From the liturgical point of view, therefore, the dramatic
performance may be regarded as a sort of introduction to the
Mass ; and it will be observed that at Mass the prophetce and their
assistants begin the Introit and rule the choir.140
The name of this dramatic performance, its attachment to the
liturgy of the Feast of Circumcision, and the presence of the
asina, — all these circumstances combine to raise the question of the
relation of the dramatic performance at Rouen to the ecclesiastical
ludi , or revels, of the Christmas season, particularly to the famous
Feast of Fools.141 This festival, variously known as festa asinaria,
or as festum stultorum, fatuorum, follorum, subdiaconorum, or
baculi, was usually celebrated on the Circumcision. The general
nature of the observance may be conveniently seen in the famous
manuscript inappropriately known as Missel des Fous, or Missel
de l’ cine, containing words and music for the Circumcision, written,
or compiled, by Pierre de Corbeil for use at Sens.142 This docu¬
ment is simply a choir-book containing troped, or embellished,
forms of the musical propria for the complete Canonical Office and
Mass of the Circumcision.143 The simple heading of the text as a
whole is Circumcisio Domini . The text itself begins, under the
rubric In januis ecclesie, with four lines of extra-liturgical verse,
138 In Rouen Ms. 384 (Y. 110) the Christmas processio is found (fol. 23v) as
follows : Ad Processionem cantor incipiat responsorium : Descendit de celis.
Prosa : Felix Maria. Versus : Tamquam sponsus. Prosa : Familiam custodi.
Et exiuit. Prosa : Te laudant alme. Conditor. In statione responsorium :
Verbum caro factum. Tres de maioribus in pulpito versum : In principio.
Plenum gratie. Ad introitum chori antiphona : Hodie Xpistus natus. Sequatur
Benedictio Archiepiscopi si presens fuerit. Hoc finito incipiatur Missa a can-
tore cappato.
139 See above, pp. 50, 67.
140 See above, p. 62.
141 The most convenient and comprehensive account of the Feast of Fools is
that by Chambers, Vol. I, pp. 274-335. The other ludi we need not, for our
present purpose, consider. Chambers considers the whole matter in masterly
fashion (Vol. I, pp. 274-371).
142 Ms. 46, of the thirteenth century, now kept in the municipal museum at
Sens. From this manuscript the so-called “Office des Fons” has been most re¬
cently edited by H. Villetard, Office de Pierre de Corheil (Bibliotheque Musi-
cologique, IV), Paris, 1907.
143 One may consult Villetard’s complete text, or the general descriptions
given by him (pp. 39-51) and by Chambers (Vol. I, pp. 279-284).
Young — Or do Prophet arum.
69
followed, under the rubric Conductus 144 ad tabulam, by the famous
“Prose of the Ass,” beginning,
Orientibus partibus
Aduentauit asinus.
Then follow the troped chants; for the complete Canonical Office
and Mass.
The rubrics accompanying the text are brief and apply, in gen¬
eral, strictly to the liturgical ceremonial. There is no suggestion
of burlesque.145 At Beauvais and elsewhere, however, the cere¬
monial included acts of revelry such as a drinking bout and a
censing with pudding and sausage.146 For our present purpose we
may omit a survey of these revels as a whole, and need mention
merely the fact that at Beauvais, during the singing of the “Prose
of the Ass, ’ ’ an asinus was actually brought into the church.147
What, then, is the relation of the Laon Ordo Prophetarum and
the Rouen Festum Asinorum to the Feast of Fools or similar eccle¬
siastical ludif For an answer, which I cannot improve, I adopt
the able words of Chambers :148
144 “Conductus design e un morceau de musique ou plutOt de chant qu’on exe-
cutait en marchant” (Villetard, p. 75).
145 Villetard (pp. 49-51, 73—82) considers and interprets the rubrics in detail.
One’s attention falls questioningly upon the rubric Conductus ad ludos, which
introduces the charming Noel (Natus est) just before the Te Deum of Matins;
but Villetard (p. 50) interprets this rubric as referring to the departure of the
congregation from the church “pour alle se reorder quelques instants.” The
serious tone of the observance at Sens is observed by Chambers (Vol. II, p. 281)
and by Villetard, the latter expressing himself (p. 51) as follows: “Rien, en
effet, k relever, dans cette vaste composition, qui soit de nature k choquer le
godt le plus exigeant. L’ouvrage, dans son ensemble, est incompatible avec
l’idee du plus leger desordre.” Chambers reminds us (Vol. I, p. 288), however,
that the more extravagant ceremonies would not be likely to be recorded in a
formal service-book.
148 See Chambers, Vol. I, pp. 286-287. I know of no complete edition of the
Circumcision office of Beauvais. Villetard (pp. 219-232) gives a complete
analysis of its content, and Chambers (Vol. I, pp. 284-287) gives a general de¬
scription. Chambers (Vol. I, pp. 279-281, 287-288) points out the fact that the
Circumcision office written for Sens by Pierre de Corbeil has undergone a
reform from which the Beauvais office escaped.
147 See Chambers, Vol. I, pp. 286, 331; Villetard pp. 49, 232. Concerning the
use of the ass in ecclesiastical ludi in general, see Chambers, Vol. II, pp. 330-
332. There is no proof that Pierre de Corbeil allowed an ass to appear in his
ceremony ; but his use of the “Prose of the Ass” indicates that at one period or
another the ass may have been used.
143 Chambers, Vol. II, pp. 56—57. The foot-notes are annotations of the present
writer.
70
Wisconsin Academy of Sciences , Arts , and Letters.
I think it must he taken for granted that the plays149 are the older in¬
stitution of the two. They seem all to have taken shape by the eleventh
century,150 before there is any clear sign that the Kalends151 had made their
way into the churches and become the Feast of Fools. The plays may
even have been encouraged as a counter-attraction, for the congregation,
to the Kalends outside. On the other hand, I do not hold, as some writers
do, that the riotous Feasts of Asses were derived from the pious and
instructive ceremony so called at Rouen.152 On the contrary, Balaam and
his ass are an interpolation in the Prophetce both at Rouen and, more ob¬
viously, at Laon. Balaam, alone of the Laon performers, is not from
the pseudo-Augustinian sermon.153 Is he not, therefore, to be regarded as
a reaction of the Feast of Fools upon the Prophetce, 15i as an attempt to
turn the established presence of the ass in the church to purposes of edi¬
fication, rather than of ribaldry? I think the explanation is the more
plausible one.
We may, then, with reasonable certainty account for the mis¬
nomer Festum Asinorum attached to the prophet-play in the Rouen
manuscripts: The Feast of Fools contributed to the prophet -play
the figure of the ass, and the conspicuous presence of the ass forced
upon the prophet-play the alien name Festum Asinorum. The
question remains as to whether the ass in the Rouen play, — and in
the Laon play as well, — brought with it the comic associations of
the Feast of Fools. In the absence of documentary evidence, one
can only surmise that however solemn the intention of the writers
of the prophet-plays may have been,155 the spectators must have
149 In this part of his statement Chambers is referring to the several liturgical
plays of the Christmas season : OJficium Pastorum, Officium Stellas, and Ordo
Prophetarum. See Chambers, Vol. II, pp. 41-55.
iso ;por an application of this date to the Ordo Prophetarum see Chambers,
Vol. II, p. 53, note 4.
151 The word Kalends here refers, of course, to the festivities of the folk at the
beginning of the year.
152 Chambers notes that Gast§ (pp. 20-24) holds this view. See also Chambers,
Vol. I, p. 332, note 1. Chasles (p. 126, note 3) seems not to discriminate at all
between the Ordo Prophetarum and the Feast of Fools.
153 See above, p. 46.
154 Elsewhere (Vol. I, p. 332) Chambers presents this view more positively as
follows : “It has been pointed out, and will, in the next volume, be pointed out
again, that the ecclesiastical authorities attempted to sanctify the spirit of play
at the Feast of Fools and similar festivities by diverting the energies of the
revellers to ludi of the miracle-play order. In such ludi they found a place for
the ass. He appears for instance as Balaam’s ass in the later versions from
Laon and Rouen of the Prophetce, and at Rouen he gave to the whole of this
performance the name of the festum or processio asinorum.
155 Sepet (p. 28) undertakes to assure us that the conduct of the ass itself
was altogether decorous.
Young — Ordo Prophet arum.
71
experienced, at least surreptitiously, some of the mirth openly pro¬
claimed in the festa asinaria .156
We have still to notice the special chants with which the dramatic
performance of Rouen begins and ends. Like the Laon play, the
version before us begins with a processional use of the Gloriosi et
famosi. It will be observed, however, that the Rouen play dis¬
closes157 several stanzas that we have not seen hitherto. The ap¬
propriateness of this chant to the fundamental theme of the Ordo
Prophetarum is so complete as to suggest that the Gloriosi et
famosi was composed specifically for use in the dramatic procession
of the Prophetce .15S
Unfortunately a like appropriateness cannot be claimed for the
prose II or turn prcedestinatio which the Prophetce et Ministri in pul -
pito sing after the concluding prophecy of the Sibyl. The com¬
plete text of this prose will be examined below.159 Suffice it to say
here that in content this composition is related, not to the Ordo
Prophetarum , but to the theme of Easter. I can offer no plausible
conjecture to account for its presence in the Rouen version before
us.160
Glancing back over our survey, we observe that of the extant
examples of the Ordo Prophetarum in its use as a strictly liturgical
play, the Rouen version shows the most extended development.
The importance of this development consists not so much in the
mere addition of new prophets as in the use of mise en scene and
dialogue for the roles of Balaam and Nebuchadnezzar. Whereas all
the other prophetce are provided merely with a conventionally ade¬
quate impersonation, these two personages are the centers of dia¬
logues that approach the status of independent dramatic episodes.
158 As to whether the Feast of Fools was cultivated at Rouen itself, alongside
the prophet-play, I have no information. Chambers (Vol. I, pp. 303-304)
traces the presence of a Feast of Fools at Laon from “about 1280” on into the
sixteenth century.
157 The Rouen text, here and elsewhere, provides only the initia.
158 For its use elsewhere, see below, pp. 72-80.
159 See pp. 75-77.
160 Nor can Sepet (p. 47) or Gaste (p. 20). I consider this matter in detail
below, pp. 76-77.
72
Wisconsin Academy of Sciences , Arts , and Letters.
VII
I have reserved for separate consideration two texts which are
directly related to versions examined above, but which raise special
questions.
The first of these texts is associated with Einsiedeln i* 1
AD PROPHETAS
Prophete venientes admonent:2 3 4 <p. 54 >
Gloriosi
Et famosl
Regis festum
Celebrantes
Gaudeamus;
Cuius ortum
Uite portum
Nobis datum
Predicantes
Habeamus.
1 Einsiedeln Ms. 366 (olim 179), Fragmenta liturgica ssec. xi-xii, pp. 53-54.
The manuscript is described by P. G. Meier, Catalogus Codicum manu scriptorum
qui in Bibliotheca Monasterii Einsidlensis O. S. B. servantur, Vol. I, Einsiedeln —
Leipzig, 1899, pp. 331-332. The part of the manuscript containing liturgico-
dramatic pieces (pp. 53-56) is more carefully described by W. Meyer, Frag¬
menta Burana, Berlin, 1901, pp. 51—52. I base my own text and observations
upon the manuscript itself. The liturgico-dramatic pieces are found as follows :
(1) p. 53 (beginning on the first line and ending on the next to the last line) :
a fragment of the Officium Stellce. For bibliography see a note of the present
writer in University of Wisconsin Studies in Language and Literature, No. 4,
Madison, 1919, p. 7, note 19. In this note I neglected to mention the fact that
this fragment is printed also by A. Schubiger, in Musikalische Spicilegien, Jahr-
gang IV, Lieferung II, Berlin, 1876, pp. 44-46.
(2) pp. 53 (last line) -5 4 (last line) : a fragment of the Or do Prophet arum ,
published by F. J. Mone, Schauspiele des Mittelalters, Vol. I, Karlsruhe, 1846,
pp. 10-12, and by Schubiger, op. cit., pp. 46-47 ; and republished herewith, with
collation of the texts of Mone and Schubiger.
(3) p. 55 (first three lines) : a fragment (latter part) of the prose Hortum
prcedestinatio, published by Mone, p. 12, and republished below, p. 74.
(4) pp. 55 (line 3) -5 6 (line 5) : a version the Visitatio Sepulchri, headed by
the rubric In Resurrectione ; published by Mone, pp. 12—13. See below, p. 74.
Since there is obviously no continuity between the last words on p. 54 and
the first words on p. 55 (See below, note 12) it is clear that one or more leaves
have been lost between (2) and (3). The lost part must have contained a
continuation of the Ordo Prophetarum and the beginning of the Hortum prce¬
destinatio. For further discussion of this lacuna in the manuscript see below,
pp. 76—77, and Meyer, pp. 51—52.
2 Ad Prophetas. Prophete venientes admonent] Omitted (Mone).
Young — Or do Prophet arum.
73
4 circuitu] circuitum (Schubiger; Mone, — who remarks, “Vor diesem Wort ist
per zu verstehen.” ) .
5Ammonitu] ammonitum (Mone).
6 Et famosi] etc. (Mone).
7Etfamosi] etc. (Mone).
74 Wisconsin Academy of Sciences , Arts f and Letters.
Chorus :
Gloriosi
Et famosi .8
Prophete:
Deum uerum
Regem regum
Confitentes
Per lauacrum
Saluabuntur.
Chords :
Prophete:
Sed Iudei
Facti rei
Condemnantes
Sacrum regem
Damnabuntur.
Gloriosi.9
Floruisse
Et dedisse
Nouum fructum
Dinoscitur
Radix Iesse.
Chorus :
Israheli
Infideli
lam Maria
Natus scitur
<Hic>10 adesse.* 11
Gloriosi.12
8 Et famosi] etc. (Mone).
9 Gloriosi] Gloriosi etc. (Mone) ; Gloriosi et fa. (Schubiger).
10 1 adopt Mone’s conjecture. Schubiger supplies nunc.
11 adesse] For this word and the word Gloriosi following, the music is omitted.
12Thus ends p. 54 of the manuscript. At the top of p. 55 begins the following:
centurio
Florem Marie proprio
Sepeliuit in tumulo.
Flos autem die tercio
Qui floret ab initio,
Refloruit e tumulo
Summo mane diluculo.
In Resurrectione
Angelus dicit :
Quern queritis.
Here follows a version of the Visitatio Sepulchri.
Young— Or do Prophetarum.
75
This fragment is obviously the processional beginning of a ver¬
sion of the Ordo Prophetarum. Although we must regret the ab¬
sence of the prophecies themselves, we are fortunate in having a
relatively complete form of the processional Gloriosi et famosi.13
From this text it appears that at Einsiedeln the Prophetce them¬
selves sang the stanzas of this cantio , a practice that was probably
followed at Laon, but certainly not at Rouen.
The most puzzling aspect of the text before us is the uncertainty
as to the relation of the fragmentary Ordo Prophetarum to the
fragment of the prose Hortum Prcedestinatio that follows it in the
manuscript. A complete version of this prose may be seen in the
following text:14
R esponsorium: Et ualde mane una sabbatorum ueniunt ad monu-
mentum, orto iam sole, alleluya. Versus: Et respicientes uiderunt
reuolutum lapidem ab hostio monumenti. Orto. Gloria Patri et Filio et
Spiritui Sancto. <Verstjs>: Et <respicientes>.
Prosa
<H>ortum predestinatio
Paruo sabbaifi spatio
Prouiderat in proximo
Ciuitatis proastio,
13 From this text we are able to complete one or two of the initia of the
Rouen text.
14 Paris, Bibliotheque Nationale, Ms. latin 2028, Breviarium Senonense ssec. xiv,
fol. 12 Or. So far as I know this text is now published for the first time. It
appears that texts of the Hortum prcedestinatio are relatively few. Chevalier,
Repertorium Hymnologicum, No. 8045, mentions only one text, in a breviary
of the year 1548. In the appendix to Analecta Bollandiana, Vol. XXXIII,
Fasc. 1, Jan., 1914, Chevalier’s information is supplemented by the following
note: “Ms. Douai, 170 (X. Is.), 72a. Rev. cath. Rouen, II, 868.” I have texts
also from Paris, Bibliotheque Mazarine, Ms. 349, Breviarium Farsemonasterii
ssec. xiii, fol. 82r— 82v, and Bibl. Mazarine, Ms. 346, Breviarium Sancti Maglorii
ssec. xv in., fol. 103v. From an unidentified Sens manuscript, the Hortum Prce¬
destinatio and accompanying Visitatio Sepulchri are published in Melanges de la
Societe des Bibliophiles , 1834, pp. 165—167, and this text is reprinted by Du-
Meril, pp. 98-100. It will be observed that the text now published from Paris
Ms. 1028 is from the use of Sens, and that, like the text reprinted by DuMeril, it
presents the Hortum prcedestinatio as associated with the Visitatio Sepulchri of
Easter. Although the Hortum prcedestinatio contains one or two rather awk¬
ward conceptions, I should not call it a “prose barbare”, as do the editors of
Melanges (p. 166, note 1).
15 proastio] proactio (Ms.). This reading explains the unintelligible pro fascio
of the Sens text reprinted by DuMeril, pp. 98-99. The reading proastio is found
in Bibl. Mazarine Ms. 346, fol. 103v, and prophastio, in Bibl. Mazarine Ms. 349,
fol. 82v. Proastium (“suburb”) may serve as the emendation desired by Meyer
(P. 51).
76
Wisconsin Academy of Sciences , Arts , and Letters.
<H>ortura pomorum uario
Non msignem edulio
Quantum uirtutis spatio18
Coeqnalem Elisio.
In hoc magnns decurio
Ac nobilis centurio
Florem Marie proprio
Sepeliuit in tumulo.
Flos autem die tercio,
Qni floret ab initio,
Refloruit e tumulo
Summo mane dilucnlo.
Orto <iam sole>.
Post representationem Mariarnm17 sequltur:
Te Deum laudamns.18
Here the prose Hortum prcedestinatio is used as a trope of the
third responsory of Easter Matins, and is followed, according to
the rubric, by a version of the dramatic Visit atio Sepulchri.1* It
appears, then, that both through its content and through its tradi¬
tional associations the Hortum prcedestinatio belongs to the liturgy
of Easter,20 and was commonly used as a trope, or appendage, of
the third responsory of Easter Matins.21 It will be remembered,
furthermore, that it is immediately after this third responsory
that the dramatic Visit atio Sepulchri is commonly found.
These facts ought to provide a sufficient basis for determining
the relations of the fragmentary Hortum prcedestinatio in the Ein-
siedeln manuscript. This fragment of a prose is, to be sure, pre¬
ceded, — after a lacuna of unknown length,22 — by the beginning of
an Ordo Prophetarum; but the prose is immediately, and appropri-
18 spatio] precio (Mazarine Mss. 346 and 349), with obviously better sense.
17 The Representatio Mariarum is, of course, the well-known Visitatio Sepulchri
traditionally performed immediately before the Te Deum of Easter Matins.
18 The rubric In Laudibus follows immediately.
19 The actual text of the Visitatio Sepulchri happens not to be given in this
manuscript. The text referred to was probably similar to that reprinted by
DuMeril, pp. 99-100.
20 Chevalier, Repertorium Hymnologicum, No. 8045, assigns it to Easter.
21 This prose is used as a trope of the third responsory of Easter Matins not
only in the text printed above from Bibl. Nat. Ms. 1028, but also in my unpub¬
lished texts from Mazarine Mss. 346 and 349, mentioned above, note 14.
22 See above, note 1.
Young — Or do Prophetarum.
77
ately, followed by a version of the Easter play, Visit atio Sepul -
chri ,23 We ought, therefore, to feel assured that the Hortum prce-
destinatio in the Einsiedeln text does not belong to the preceding
Ordo Prophetarum , but is to be related to the succeeding Visitatio
Sepul chri.
But our assurance is not quite complete ; for although in thought
the Hortum prcedestinatio is decisively alien to the Ordo Prophet-
arum and to the Christmas season, the Rouen Festum Asinorum
does actually conclude with the singing of this prose;24 and the
Einsiedeln dramatist may have allowed a similar incongruity. It
must be admitted, moreover, that in the Einsiedeln manuscript it¬
self there is nothing to disprove absolutely the inclusion of the
prose in the preceding prophet-play, now largely lost. In the
present state of our information, the precise dramatic relations of
the fragmentary Hortum prcedestinatio in the Einsiedeln manu¬
script must be left undecided.25
The second text for special consideration is the following, of
uncertain provenience :26
lx Epiphania Domini
Lectio Isaiae prophetae: Surge, illuminare, Ierusalem, quia venit
lumen tuum, et gloria Domini super te orta est.
Gloriosi
Et famosi
Regis festum
Celebrantes
Gaudeamus ;
Cuius ortum
Vitae portum
Nobis datum
Praedicantes
Aveamus.
23 See above, note 12.
24 See above, pp. 62, 71.
23 Since the Hortum prcedestinatio fragment is immediately followed (not in the
margin, as Meyer says [p. 51], but as the ending of line 3) by the rubric In
Resurrectione , and since the opening words (Quern queritis) are written in
uncials, Meyer holds (p. 51) that the Hortum prcedestinatio fragment is effect¬
ually separated from the Visitatio Sepulchri, and has “Nichts mit dem Oster-
spiel zu thun.” I myself do not regard the rubric and the uncials as decisive in
effecting the separation. The general situation under discussion may be
grasped from note 1 above. Even though the Hortum prcedestinatio fragment
be regarded as effectually separated from the succeeding Visitatio, the associa¬
tion of the' fragment with the Ordo Prophetarum is a matter of uncertainty.
28 1 take this text from Analecta Hymnica Medii 2Evi, Vol. XLIX, Leipzig,
1906, pp. 182-183. Concerning his source, the editor, Clemens Blume, writes
(p. 183) : “Cod. ms. saec. 14.— -Bannister, dem ich die Abschrift verdanke,
konnte leider den Fundort und die Provenienz der Handschrift nicht mehr
ermitteln.”
78
Wisconsin Academy of Sciences , Arts , and Letters.
Quia, ecce, tenebrae operient terrain et caligo populos.
Ecce, regem
Novam legem
Dantem orbis
Circuitu
Praedicamus.
Quern futurum
Regnaturum
Prophetico
Admonitu
Nuntiamus.
Super te autem orietur Dominus, et gloria eius in te videbitur.
Sunt impleta
Quae propheta
Quisquis dixit
De future
Summo rege.
Impiorum
Iudaeorum
Corda negant
Regnaturo
Sua lege.
Et ambulabunt gentes in lumme tuo et reges in splendor e ortus tui.
Dilatata Et gentiles
lam privata Prius viles
Fit regali Convertuntur
Potestate
Plebs ludaea.
Leva in circuitu oculos tuos et vide .
tibi.
Deum verum
Regem regum
Confitentes
Per lavacrum
Salvabuntur.
Maiestate
Aetherea.
omnes isti congregati sunt, venerunt
Sed ludaei
Facti rei
Non fatentes
Regem sacrum
Damnabuntur.
Filii tui de longe venient, et filiae tuae de latere surgent.
Omnes gentes
Congaudentes
De cantu laetitiae,
Quia homo
Sit de domo
Natus David hodie.
Tunc videbis et afflues, et mirabitur et dilatabitur cor tuum, quando
conversa fuerit ad te multitudo maris, fortitudo gentium venerit tibi.
Isaias, die, de Christo quid prophetizas?
Est necesse Flos deinde
Virgam Iesse Surget inde
De radice provehi; Qui est filius Dei.
Inundatio camelorum operiet te, dromedarii Madian et Epha.
0 ludaei
Verbum Dei
Cur negastis hominem?
Yestrae legis
Vestri regis
Audite nunc ordinem.
Young — Ordo Prophetarum.
79
Omnes de Saba venient aurum et tus deferentes, et laudem Domino
annuntiantes.
Confunduntur
Convertuntur
iMaiestatis aethere.
Omnes gentes
Non credentes
Peperisse virginem
Since for our present purpose the interest of this liturgical piece
is somewhat subsidiary, I deal with it only briefly. It is a troped
form of the Epistle for the Mass of Epiphany: among the sentences
of the Vulgate (Isa. lx, 1-6) are distributed metrical compositions
in the nature of embellishments of the orthodox liturgical text.27
It will be observed that the trope begins with the Gloriosi et famosi,
and consists of stanzas virtually every word of which we have al¬
ready encountered in one version or another of the Ordo Prophet-
arum. That this trope is in some way related to the prophet-play
is further apparent in the intrusive summons Isaias , die , de Christo
quid prophetizas ?
The question arises, then, as to whether this trope is one of the
sources of the Ordo Prophetarum , or is itself under the influence of
the dramatic tradition. That the latter relation is the valid one
is suggested, in the first place, by the uniqueness and late date of
the text before us. In order to influence the development of the
Ordo Prophetarum it would have been necessary that this Epistle-
trope be in existence and be well-known as early as the eleventh
century, the date of the Limoges Ordo Prophetarum, for this
dramatic text contains substantial passages that are identical with
passages in the trope.28 That fact that the editors of Analecta
Hymnica have discovered only one text of the trope, and that this
text is relatively late (fourteenth century), seems to show that the
trope can scarcely have had sufficient age or repute for serving
as a source of the prophet-play of Limoges.29
Further indication that the trope is not a source of the prophet-
27 1 follow Blume in printing the liturgical Epistle itself in italics.
28 See above, pp. 25-31. In the trope are found also the following passages of
the Limoges text : Omnes gentes . . . per ordinem ; Est necesse . . .
est spiritus (filius) Dei; Et vos (Omnes) gentes . . . peperisse virginem.
29 The Beverend H. M. Bannister, who contributed the trope under considera¬
tion, was complete master of the trope manuscripts of Western Europe, as one
may infer, for example, from his list of troparia in Analecta Hymnica , Vol.
XL VII, pp. 22-25.
80 Wisconsin Academy of Sciences, Arts , and Letters.
play, but rather an emanation from it, appears from such a stanza
as the following:
O Iudaei,
Verbum Dei,
Cur negastis horninem?
Whereas such an address to the Jews30 finds no special inspiration
in the liturgical Epistle that is being embellished, it accords per¬
fectly with the tenor of the pseudo-Augustinian lectio and of the
central motive of the Ordo Prophetarum. It would appear that
such a passage originated not from the liturgical Epistle of Epiph¬
any but from the dramatic tradition.
The text before us, then, may be regarded as an isolated liturgical
piece in which certain stanzas from the Ordo Prophetarum have
been used, with considerable appropriateness, as a trope of an
Epistle.31
VIII
In view of the detailed attention given above to the several ver¬
sions of the Ordo Prophetarum, our summary of the development
of this liturgical play may be brief.
There can be, in the first place, no doubt as to the origin of this
dramatic phenomenon. A re-examination of the matter has im¬
paired none of the essentials of Sepet’s original demonstration.1
The Ordo Prophetarum certainly arises from that part of the
pseudo-Augustinian sermon (Contra Judaeos, Paganos, et Arianos)
which was widely used as a lectio for Matins during the Christmas
season, this particular part addressing itself to the Jews and be-
30 See also the stanza beginning Sed Iudcei , and the stanza ending Plebs Iudcea.
31 1 can see no reason for reprinting here the Cantio de mulieribus beginning,
Recedite, recedite,
Ne mulieri credite !
Die tu, Adam, primus homo, etc.
In form this satirical composition may possibly show the influence of the Ordo
Prophetarum. A new version of the Cantio, with information concerning ver¬
sions previously published, is given by L. Suttina, Una Cantilena medievale
contro le donne, in Studi Medievali (ed. F. Novati and R. Renier), Vol. II
(1906-07), pp. 457-460. In the same volume (pp. 538-550), F. Novati, fetching
an analogy from the Ordo Prophetarum, undertakes to show that the Cantio
is “un dramma liturgico” for Ash Wednesday. I cannot accept the demon¬
stration.
1 See above, pp. 1-2.
Young — Ordo Prophetarum.
81
ginning with the words Vos , inquam, convenio, O Judaei. The
fresh documents brought forward in the present study merely ex¬
tend, elucidate, and confirm Sepet ’s conception of this liturgico-
dramatic development.
From the extant texts it appears that even as a procession of
prophets confined within the bounds of the liturgy the Ordo
Prophetarum attained a conspicuous development in the direction
of genuine and independent drama. It must be admitted, to be
sure, that in so far as the mere formula of the Ordo is concerned,
this dramatic invention tends toward a mediocre regularity. The
inevitable uniformity in the summoning of a succession of wit¬
nesses, and in the delivering of their testimonies, suggests
monotony.2 But it is also clear that the liturgical dramatists ac¬
complished a good deal in the way of variety. The Laon and
Rouen versions show great care in impersonation.3 The costumes
are at once brilliant and discriminating, and details of personality
are often penetratingly disclosed. More striking still is the provi¬
sion, for certain witnesses, of conspicuous properties or mise en
scene. The action centering in the ass of Balaam and in the fur¬
nace of Nebuchadnezzar, in the Rouen Ordo, develops into dramatic
episodes of independent interest; and since one of these episodes
is found near the beginning, and the other near the end, of a long
defile of prophets, the dramatist may have intended explicitly to
enliven a solemn formula with moments of comic suggestion.
Let it be understood, finally, that the present study does not
undertake to survey the career or the influence of the Ordo
Prophetarum beyond the confines of liturgical drama. Sepet pur¬
sues the matter much farther, and through demonstration and con¬
jecture, outlines a development of the Ordo through several later
and more comprehensive stages. He conjectures, in the first place,
a form of the dramatic procession in which new witnesses appear,
and in which prophets besides Balaam and Nebuchadnezzar were
the centers of dramatic expansion; and he holds that certain of
these special episodes may have been more extended than anything
that we have seen in the plays of Laon and Rouen.4 He then rea¬
sons that some of these special episodes detached themselves from
2 Upon this monotony of Meyer (p. 52) is particularly insistent.
3 It may be, of course, that the Limoges version also availed itself of im¬
personation, even though the rubrics of the extant text are silent. See above,
p. 37.
4 See Sepet, pp. 48-147.
6— S. A. L.
82 Wiscdnsin Academy of Sciences , Arts , and Letters.
the procession, and persisted as independent plays.5 Finally Sepet
assumes that these independent plays reunited in the form of Old
Testament dramatic cycles, such as we find in several of the ver¬
naculars of Western Europe.6
As to the validity of these comprehensive conjectures and infer¬
ences scholars have disagreed,7 and it may be that the extant texts
are insufficient for demonstrating with precision the part of the
Or do Prophetarum in the forming of the vernacular cycles upon
subjects from the Old Testament. In any case I mention the
matter here not for the purpose of passing judgment, but rather
in order to discriminate between the part of Sepet ’s study that
is demonstrably correct and the part into which enters tenuous
conjecture. However doubtful his more ambitious conjectures may
be, his derivation of the dramatic Or do Prophetarum from the
pseudo-Augustinian lectio is sound. Such additional evidences
and such corrections of detail as may be advanced in the present
study all combine in support of Sepet ?s important discovery.
5 See Sepet, pp. 49-80.
6 See Sepet, pp. 165-179.
7 See Meyer, pp. 53-56 ; W. Creizenach, in Literaturhlatt fur germanische und
romanische Philologie, Vol. XXIII (1902), col. 203 ; H. Craig, The Origin of the
Old Testament Cycles, in Modern Philology, Vol. X (1913), pp. 473—487 ;
Adeline M. Jenney, A Further Word as to the Origin of the Old Testament
Plays, in Modern Philology, Vol. XIII (1915), pp. 59-64.
To
ERNST VOSS
On His Sixtieth Birthday
October Thirteenth
Nineteen Hundred and Twenty
RICHARD WAGNER’S
“DIE MEISTERSINGER VON NURNBERG”
AND ITS LITERARY PRECURSORS
EDWIN C. ROEDDER.
The first performance of Richard Wagner’s musical comedy-
drama Die Meistersinger von Numb erg took place in the Royal
Opera House in Munich, June 21, 1868. As to its genesis we are
accurately and fully informed by the author himself, in passages
of his autobiographical writings as well as in his letters. Over a
score of years had elapsed between the first performance — the
most memorable one that any opera had ever had up to that
time, as an artistic achievement equalled only by the presentation,
in 1876, of T)er Ring des Nibelungen, and surpassed only by Par¬
sifal, in 1882 — and the first conception of the play which Wagner,
then musical director of the Royal Opera at Dresden, jotted down,
in 1845, during a stay at Marienbad in Bohemia, where he had
gone for the recuperation of his health. In his posthumous auto¬
biography My Life 1 he says :
“Owing to some comments I had read in Gervinus’s History of
German Literature* 2 , both the Mastersingers of Nuremberg and
Hans Sachs had acquired quite a vital charm for me. The very
name of the Marker, and the part he takes in the Mastersinging,
were particularly pleasing to me, and on one of my lonely walks,
without knowing anything particular about Hans Sachs and his
poetic contemporaries, I thought out a humorous scene in which
the cobbler — as a popular artisan-poet — with the hammer on his
last, gives the marker, who is forced to sing, a practical lesson,
thereby taking revenge on him for his pedantic misdeeds. To me
the force of the whole scene was concentrated in the two follow¬
ing points: on the one hand the marker, with his slate covered
*1 quote from the anonymous English translation, New York, 1911, vol. I,
page 366, taking the liberty of correcting some curious mistakes of the trans¬
lator.
2 The passages in question are collected in the appendix to this paper.
86 Wisconsin Academy of Sciences, Arts, and Letters.
with chalk marks, and on the other Hans Sachs holding np the
shoes covered with the marks from his hammer blows, each inti¬
mating to the other that his singing had been a failure. To this
picture, by way of concluding the second act, I added a scene con¬
sisting of a narrow, crooked little street in Nuremberg, with the
people all running about in great excitement, and ultimately en¬
gaging in a street brawl. Thus suddenly the whole of my Meister-
singer comedy took shape so vividly before me that, inasmuch as
it was a particularly cheerful subject, and not in the least likely
to overexcite my nerves, I felt I must write it out in spite of the
doctor’s orders.”
In Wagner’s Mitteilung an meine Freunde 3, written six years
later, in 1851, we learn that his intention to carry out this new
plan was considerably strengthened by the advice of several well-
wishers, who wanted him to compose some opera of the lighter
kind, which would be likely to secure for his works admittance to
all German stages and thereby win for him the outward success
that he had so far missed and that was so necessary in his struggle
for existence. At the same time he mentions here another aspect
of the new plan : the Mastersinger drama was to be the humorous
converse of the tragic contest of the Minnesingers at the Wart-
burg, as treated in his Tannhduser, the score of which he had
completed in the month of April, and the first performance of
which took place on the 19th of October, 1845. Hans Sachs was
to personify the last appearance of the artistically productive
folk-spirit, and as such he was to be contrasted with the philistine
mastersingers, to whose comical pedantry, bound up entirely in
the rules of the “tablature,” Wagner expected to give a distinctly
personal expression in the figure of the marker.
The Marienbad sketch of the Meistersinger was jotted down
July 16, 1845. 4 The Mitteilung an meine Freunde contains a
synopsis of the plan from which the general reading public re¬
ceived its first knowledge of the subject. The final and defini¬
tive execution adheres quite closely to this first sketch, so far as
the general outward course of the action is concerned, but presents
vital departures in a number of points, especially in the charac¬
terization of Sachs and the youthful hero. The internal changes
are indeed so incisive that Wagner, when toward the close of the
3 Gesammelte Schriften, 4, 284 ff. (References to second and subsequent edi¬
tions.)
4 It is printed in full in Sdmtliche Schriften , 11, 344ft.
Boedder — “Die Meistersinger von Niirnberg 87
year 1861 he wrote out the present form of his work, could say
that the original plan offered him little or nothing.5
To gain an insight into the growth of Wagner’s spiritual life
as expressed in the play from its first inception to its completion,
and at the same time to obtain a satisfactory answer to the ques¬
tion as to how much Wagner is indebted to his “ sources” for
the plot, it will be necessary to give a fairly detailed synopsis of
the Marienbad sketch.6 In this the characters are not yet named,
excepting Hans Sachs, his apprentice David, and Magdalene,
David’s elderly sweetheart, the kinswoman and companion of the
unnamed daughter of the senior of the mastersingers ’ guild. The
later Walter von Stolzing appears merely as a junger Mann, Beck-
messer as the Merker, and Eva as the Preismadcken, For brev¬
ity’s sake we shall in the following insert the names used in the
final version.
Walter, a young knight, inspired by the reading of the Helden-
buch and the old Minnesingers, an ardent lover of poetry, descends
from his impoverished ancestral castle in Franconia, in order to
learn the art of the mastersingers in Nuremberg. To join the
guild, he presents himself at the house of its senior, Veit Pogner,
the goldsmith, and here meets Pogner ’s daughter, with whom he
quickly falls in love.7 Pogner has offered his daughter’s hand as
a prize to the master who shall defeat his competitors in a sing¬
ing contest to be held in the near future. The deciding vote in
the matter is to be left to the maiden herself. When at the meet¬
ing of the guild Walter presents himself for the master’s test,
which he must pass in order to participate in the coming competi¬
tion, Hans Sachs, who happens to be the warden of the law for
the time being, reads to him the rules of the guild, with a decided
admixture of irony, and Walter is rather nervous and intimidated.
Beckmesser, the all-important marker, — whose official business it
is to keep tally of the mistakes and infractions against the narrow
rules and laws of the mastersingers’ art, — himself aspires to the
fair maiden’s hand and her father’s gold. He scents a rival, and
naturally avails himself of this glorious chance to put him out
of the way. So he interrupts Walter’s singing before he is half
5 Richard Wagner an MatMlde Wesendonk, p. 293.
6 For a full synopsis of the drama in its present form see Henry Edward
Krehbiel, Studies in the Wagnerian Drama, New York, N. D. (1891), pages
72-77, or Henry T. Finck, Wagner and his Works. The Story of his Life with
Critical Comments, vol. II, New York, 1893, pages 217-223.
7 So it is not, as in the final version, his acquaintance with Eva and his love
for her that suggests to him the desire to become one of the mastersingers
himself, but in first visiting Pogner’s house he has no ulterior purpose beyond
his reception into the circle of the masters.
88 Wisconsin Academy of Sciences, Arts, and Letters.
through, and declares the young knight’s efforts to have been an
utter failure. Walter tries to defend himself, in his anguish begs
for another chance, and finally cries out in the greatest despair,
“Have mercy, masters!” and “ rushes away as though annihi¬
lated.”
When in the second act Walter meets Eva again, he gives vent
to this despondency and bitterness, and also to his disappointment
in the mastersingers ’ ways. In his desperate state of mind he
plans and attempts to abduct his lady-love, but Hans Sachs, who
has taken a fancy to him, frustrates this bold scheme, with the
best of intentions for the young couple. At the same time Sachs
utilizes the opportunity to vex and get even with the marker.
Beckmesser, intending to humiliate Sachs, has twitted him for not
finishing a pair of shoes that he had ordered from him. Now when
in the night prior to the great singing-contest the marker serenades
the girl with the song with which on the next day he hopes to
triumph over his rivals, Sachs, whose workshop is just opposite
Pogner’s house, likewise starts a loud song, since, as he explains
to the angry marker, he must sing in order to keep himself awake
when working so late at night; and no one knows better than
Beckmesser himself that the job must be done forthwith, for has
he not just a few hours since reminded Master Sachs most forcibly
that he is neglecting his duty as cobbler for the sake of his poetic
attempts. At last he promises the unfortunate marker that he
will stop singing, only the latter must permit him to indicate the
mistakes that he detects in Beckmesser ’s song in his own way, as
a shoemaker, by a hammer-stroke on the shoe over the last. Beck¬
messer sings, and Sachs uses his hammer very liberally. The
critical taps on the lapstone thoroughly infuriate and confuse the
marker, but all that he elicits from Sachs is the placid question
whether he is done. “Not nearly!” the marker cries, and Sachs
laughing holds up the shoes, the job having been finished merely
by the hammer markings. Beckmesser now bellows forth the re¬
mainder of his song without a pause, and fails most woefully in
the eyes of the female figure at the window. David has recognized
the latter as his beloved Magdalene and rushes out to fall upon
the singer, the noise calls out the other neighbors, and a battle
royal is the result. During this brawl Hans Sachs thwarts the at¬
tempt of Walter to abduct Eva, and takes him into his house.
On the next morning we listen to a long monolog of Sachs as
he sits in the bright sunlight, leaning back in his armchair, sur¬
rounded with books, a large volume on his knees, on which he
props up his arm while he muses on the state and condition of
poetry: “Is the fair art really coming to an end? Is it possible
that I, a shoemaker, should be the only one to breathe in the realm
of the great German past?” He continues philosophizing on the
decline of poetry, and sinks into a brooding mood, asking himself
whether his trade could dishonor him, — no indeed, for does it not
Roedder — “Die Meister singer von Numb erg .”
89
provide for him a better and more honorable livelihood than the
singing guild? Walter, who now enters, reveals to Sachs his keen
disappointment in his Nuremberg experiences: he had placed
high hopes on the mastersingers, from the repugnant present in
which he lived they were to lead him into a beautiful poetic life ;
here he had expected to find remnants of the ancient Thuringian
spirit, and now such disillusionment! He informs Sachs of the
poetry he has written hitherto, under the influence of the Helden-
buch and the great Wolfram, songs celebrating the great heroes
and emperors, and he submits to the older man his latest minne-
song, which makes Sachs exclaim, “You are a poet!” But, he
adds, “you can no longer thrive as such.” For he himself feels
keenly the pressure that weighs upon him: he was born to live
in an unpoetic age. “With a melancholy humor he depicts to Wal¬
ter the epoch in which they live, the imminent extinction of the
last mournful remnant of the old poetry, the mastersong! . . .
Believe me, for a long, long time poetry will be forgotten. Peo¬
ple will fight with other weapons than songs: with reason, with
philosophy, against stupidity and superstition ; aye, with the
sword they will defend these new weapons : you, who cherish such
fine, noble sentiments, are to join in such combat, and thus you
can achieve more than through the use of a gift that no one nowa¬
days appreciates. Sometime perhaps, after centuries have elapsed
and a new world has begun, they will turn back again and look
upon that which they once had: then they may possibly chance
upon Hans Sachs, and he will point the way farther back and
lead them to Walter, Wolfram, and the hero lays.” And when
the young knight asks him, “Advise me then, what am I to do?”
Sachs answers cheerfully, “Return to your castle, study what Ul¬
rich von Hutten and he of Wittenberg have written, and if it be
necessary defend what you have learned with the sword!” But
Walter’s thoughts run in another direction: “Very well, master!
But now I need a wife!”
Beckmesser, disconsolate over his failure in the previous night,
comes to Sachs, whom he holds responsible for his plight, and de¬
mands from him a new song for the competition. Sachs hands
him Walter’s song, pretending not to know where it came from,
and advising him to pay due attention to a suitable tune to sing
it to. The conceited marker deems himself invincible on this
score, and in the presence of the masters and the population of
Nuremberg who are to pronounce judgment, he sings the poem to
an utterly unsuited and disfiguring melody, so that he fails again,
and this time decisively. In his rage he charges Sachs with fraud,
in foisting on him a disgraceful poem. Sachs, however, asserts
that the poem is very good indeed, only it must be set to suitable
music. It is agreed that he who knows the right tune shall be de¬
clared victor. Walter naturally steps forward and with his sing¬
ing wins the bride, but disdains reception into the masters’ guild,
90 Wisconsin Academy of Sciences, Arts, and Letters.
which is now offered to him. Sachs then begins to sing the praise
of the mastersingers’ guild half ironically, half seriously, dwelling
on its good features and the excellent things that have been pre¬
served and fostered by it. Thereby he assuages the mastersingers
themselves and wins them over. He closes with the words:
‘E’en should the Holy Roman Empire perish,
We’d still have holy German art to cherish.’ ”
Fully sixteen years after the completion of the Marienbad draft,
on October 3,0, 1861, Wagner wrote to his publisher Franz Schott
informing him that he expected to resume an old plan for a comic
opera :
“The opera is entitled Die Meistersinger von Numb erg, and the
poetical and jovial chief hero is Hans Sachs. The subject matter
has in it very much that is genial and droll, and I am pleased to
think that I have hit upon something unexpected and ingenious
in this plan, which is entirely of my own invention.’ ’
And on November 20, he added:
“When I surveyed this year, which was in every respect lost
to me, I asked myself, what am I to do? All of a sudden my
whimsical Mastersingers rose up before me, and at one blow I felt
myself again master of my fate. Evidently my good star had
formerly suggested to me this unique and even jolly subject, in
order to help me with it at the most critical moment. ”
From these remarks it is manifest that Wagner, at least at the
time of the execution of his plan, regarded the subject as entirely
his own. Neither do we have any statement from him at the time
of its inception which would contradict his later assertion. And
so Professor Golther seems fully justified in saying:8
‘ ‘ So far as the Meistersinger is concerned, we can hardly speak
of sources, so irrelevant and insignificant are the few external
traits that emanate from literary sources. Werner and Welti
have collected the few pertinent data in Kursehners Wagner jahr-
buch, 1886. 9 As the historical coloring of the tenth century was
employed in Lohengrin in a novel and independent manner, so
Wagner introduced in the Meistersinger the cultural background
8 Richard Wagners dramatische Dichtungen im Verhaltnis zu ihren Quellen.
Biihne und Welt, I, 2 (1898-99), p, 579.
0This book was not at my disposal.
Boedder — “Die Meistersinger von Niirnberg.”
91
of the sixteenth century, especially from Wagcnseil’s well-known
mastersinger book. 1 no
In the Introduction to his edition of Wagner ’s works, p. 286,
Golther says,
“The original plan appears as a comic opera, which for the out¬
ward course of the action took a few things from Deinhardstein ’s
drama Hans Sachs (1827) and from Lortzing-Reger ’s opera, but
in the main was freely invented. ” * - . . ■ . ,
r
Over against this minimizing of foreign influences, we "have in
Miss Anna Maude Bowen’s Cornell dissertation (published at
Munich in 1897) a scrupulous, almost meticulous attempt to gather
every detail that Wagner might possibly have taken from others.* 11
Miss Bowen even deems it necessary to defend Wagner against a
possible charge of plagiarism for his adoption of motives. Wag¬
ner himself tells us in his autobiographical writings as a rule
quite freely and fully about the genesis of his poetic works — we
learn from his pages, e.g., that Das Liebesverbot was based on
Shakespeare’s Measure for Measure, and Die Feen on Gozzi’s La
Donna Serpent e — and informs us concerning the poetic process
of transforming the subject matter thus provided until it suits
his purposes. As he does not mention the plays named above,
and as he refers to his mastersinger comedy distinctly as an in¬
vention of his own, it is obvious that Golther and Miss Bowen con¬
nect very different ideas with the term sources. Curiously enough,
Miss Bowen overlooked Goethe’s Poem Hans Sachsens poetische
Sendung (1776), which, if anything, must be termed a source of
Wagner’s work if we turn our attention to the characterization
of Hans Sachs. That she should not have mentioned Gervinus ’s
Geschichte der deutschen Dichtung is not to be wondered at, since
at the time when the monograph was written Wagner’s autobiog¬
raphy Mein Leben had not yet appeared. Looking upon Wag¬
ner’s drama as a large stream the works that Miss Bowen cites
10 The reference is to Johann Christoph Wagenseil, Be Civitate Norimbergensi
Commentatio, with its appendix Buch von der Meistersinger holdseligen Kunst
Anfang, Fortubung , Nutzbarkeiten und Lehr-Satzen. Altorf 1697.
11 The subject was suggested to Miss Bowen by her erstwhile teacher, Pro¬
fessor Franz Muncker, who in his book Richard Wagner. Eine Skizze seines
Lebens und Wirkens (2nd ed., Bamberg 1909) , p. 99f., enumerates the works
quoted by Miss Bowen, without however attaching much importance to their
influence.
92 Wisconsin Academy of Sciences, Arts, and Letters .
do not appear to ns as its headwaters, but as little rivulets which
join it at some distance below and are carried along without for
a moment changing its original course. Dropping the figurative
speech we shall be content to speak of Wagner’s literary precur¬
sors rather than of his sources.
The most important of these is Johann Ludwig Franz Dein-
hardstein. Born in Vienna, June 21, 1794 — almost exactly three
hundred years after Hans Sachs — Deinhardstein became professor
of classical literature and aesthetics in the university of his native
city in 1827, and in the same year wrote his Hans Sachs, Schau-
spiel in vier Aufziigen.12 From 1832 to 1841 he was vice-director
of the Vienna court theater; from 1841 to his death, July 12,
1859, dramatic censor in the same institution. His plays, col¬
lected in seven volumes, have gone to well-merited oblivion, and
his stage versions of Shakespeare’s The Taming of the Shrew and
Twelfth Night have now been superseded by the originals on prac¬
tically all German stages. The only connection Deinhardstein
still has with the modern stage is through his Hans Sachs, and this
but indirectly, as the source of Lortzing’s comic opera. In its
day Deinhardstein ’s drama enjoyed an extraordinary success.
The first performance took place at the Royal Court Theater of
Berlin, February 13, 1828, preceded by a prolog which Goethe
had written for the occasion (“Da stehf ich in der Fremde ganz
allein”). How highly Goethe thought of Deinhardstein ’s work
is evidenced by the following lines :
“Und wie auch noch so lange getrennt
ein Freund den andern wiedererkennt,
hat auch ein Frommer neuerer Zeit
sich an des Vorfahren Tugend erfreut,
und hingeschrieben unit leichter Hand,
als stiind ■ es farbig an der Wiand,
und zwar mit Worten so verstdndig,
als wurde Gemaltes wieder lebendig
We no longer share Goethe’s high opinion of Deinhardstein ’s
drama. It probably was subconsciously due to the fact that Dein¬
hardstein here paid tribute to a figure that had meant so much to
the youthful Goethe. And it must be conceded that owing to its
unprecedented run on the stage it resuscitated and popularized the
12 Conveniently accessible in Reclams Universal-Bibliothek, No. 3215.
Boedder — “Die Meistersinger von Niirriberg.” 93
figure of Hans Sachs for the whole nation, just as Goethe’s Hans
Sacks ’s Poetic Mission had aroused the interest of the poets of
the Storm and Stress period, after the untutored artisan-poet had
for two centuries been despised as a wretched rhymester.
The rising curtain reveals to us Hans Sachs, a young man of
twenty-three years, seated beneath a blooming tree in his garden,
trying in vain to write a poem, more dreaming than composing.
One of his fellow mastersingers comes to return to him his poem
4 'The Goddess’s Nine Gifts;” with very adverse comment, since
Sachs has in it, as he thinks, grossly offended against the rules of
poetry and also failed to show the modesty becoming so young
a writer. A second mastersinger joins them and announces a
meeting of the guild, at which new privileges granted by Emperor
Maximilian are to be proclaimed. That these privileges are due
mostly to Sachs’s writings, which have found favor with the em¬
peror, the two mastersingers, stung with jealousy, carefully con¬
ceal from their young colleague. Nor is Sachs envied by his
brothers in Apollo alone: James the baker and Martin the grocer
likewise find fault with him, unconsciously feeling his superiority:
Jakob. Er ist fast jeglichem ein Dorn im Aug’,
der Meister Superklug.
Zweiter Meisters anger. • Er hat Talent,
das ist wohl wahr — allein —
Erster Meistersanger. Talent! — Talent! —
Wir brauchen kein Talent, Tabulaturam
soil er befolgen; die Aequivoca,
die Relativa und die blinden Worte
soil er vermeiden, keine Milben brauchen,
glatt singen soli er, das begehren wir ,
nicht aber dabei zucken, wie er’s tut ,
das macht den Dichter und nicht das Talent.
Talent kann jeder haben, aber nicht
das rechte Ohr und jene Sorgsamkeit,
so uns die Fehler klug vermeiden lassen,
und die sprech ’ ich ihm ab; er ist noch nicht
gesetzt genug, ihm macht die Phantasei
zu vielen Schaden noch.
Jakob (der erstaunt zugehort hat, zu Martin).
Das ist ein Mann
von anderm Schrote als der Meister Sachs;
wenn man den reden hort, da lernt man was.
I have quoted in full what I regard as the best passage in the
whole play. 0 si sic omnia! But unfortunately the spectator is
supposed to believe in the greatness of Deinhardstein’s hero more
94 Wisconsin Academy of Sciences , Arts , and Letters.
on the basis of the judgment of others, especially the rancorous
aspersions of his jealous associates, than on the basis of any posi¬
tive achievements of his own. This is the snag that any frail
craft sailing on the perilous waters of the Kiinstlerdrama is so
apt to strike, and which Wagner so skilfully avoided — in the Mei-
stersinger we are not merely invited but actually forced to believe
in the greatness of both Hans Sachs and Walter von Stolzing, be¬
cause we receive in both cases irrefragable proof of it, instead of
having to infer it by antithesis from Beckmesser ’s malicious sneers.
The scene changes, and the author takes us to a garden in front
of the house of Master Steffen, a rich goldsmith, the father of
Sachs’s lady-love, Kunigunde, who returns her lover’s affection
but is very anxious that he should give up his lowly calling, since
her purse-proud father would never consent to the marriage of his
only child with a cobbler. Sachs however takes a manly stand and
is not willing to humor her; on the contrary, he makes up his
mind to sue now openly for the girl’s hand. No sooner has Sachs
left than Steffen enters and announces to his daughter that he has
brought her a birthday gift, the Augsburg councillor Eoban Runge,
to whom he has promised his daughter in marriage. Eoban, to set
him off against Hans Sachs, is depicted as an intolerable coxcomb
and coward, and when Sachs shortly afterwards reappears on the
scene, a quarrel ensues in which Sachs easily shows himself the
superior of Kunigunde ’s unwelcome suitor.
The second act takes us to the square in front of Hans Sachs’s
house. We learn that at the meeting of the mastersingers ’ guild
he has been so mercilessly ridiculed on account of his latest poem
that he has left the assembly pale and trembling. He complains
of his humiliation in a monolog. In this mood he meets his rival
Eoban for the second time. Eoban has torn his shoe on the sharp
cobble-stones of the Nuremberg pavement, and seeing the shoe¬
maker’s sign, demands that the hole be mended forthwith. Learn¬
ing that his fellow suitor is a shoemaker, he hurries away to Kuni¬
gunde, to convey to her the supposed bit of news. Before he ar¬
rives, Kunigunde has almost coaxed her father into consenting to
her marriage with Sachs, provided that he can approve of her
lover’s calling, when Eoban rushes in, out of breath, to impart his
information. In her confusion she promises to wed the Augsburg
councillor if what he has stated be correct, hoping of course that
under the pressure of circumstances Sachs will now be willing to
give up his trade, which, being a wealthy man, he need not pursue
anyway. But Sachs, who comes to ask openly for Kunigunde ’s
hand, is now more determined than ever to remain true to his
work, and Kunigunde, exasperated beyond patience, bids him leave
her for good.
Boedder — “Die Meistersinger von Niimberg.”
95
Grieved and sad, Sachs has left Nuremberg in quest of a new
home elsewhere. In a large forest at some distance from his un¬
grateful native city, he meets a distinguished-looking stranger in
hunter’s garb, who is none other than the Emperor Maximilian,
traveling incognito, on his way to Nuremberg. The stranger wants
Sachs to show him the way, and on learning his guide’s name is
overjoyed, since he values him very highly as a poet, and the same
opinion he says is held by the monarch himself. The latter would
certainly be pleased to meet him, and if Sachs ever should have a
special request would do his best to comply with it. Thoroughly
appeased, Sachs returns to Nuremberg with his new friend, whom
he imagines to be a powerful count and high dignitary of the court.
Again the scene changes, and once more we are in Master Stef¬
fen’s garden. Eoban, decked out for the approaching wedding,
gets scant encouragement from the disconsolate Kunigunde, who
cannot forgive herself for her treatment of Sachs. He hopes to
compel her to accept him as her husband, and when a number of
citizens bring the announcement that Steffen has been elected
burgomaster of the city, Eoban is the first to convey this precious
piece of news to his prospective father-in-law, with the intimation
that he was chiefly instrumental in bringing about this decision.
Steffen promises Eoban that Kunigunde shall wed him on this very
day, and as he is about to force her to consent, Sachs, who with
the emperor’s retinue has just passed by outside, leaps over the
wall to protect the girl and threatens to invoke the law against the
father and would-be bridegroom. Steffen, who himself now repre¬
sents the majesty of the law, commands him to leave the premises
instantly. Kunigunde, awakening from a swoon, entreats Sachs
to forgive her and plights her troth to him anew. With the definite
promise that he will protect her, Sachs hurries away, confident of
success.
In the emperor’s ante-room, in Act Four, Sachs craves an im¬
mediate interview with the supposed count, and is directed to re¬
pair to the city hall square within an hour. Here, in the next and
final scene, the burghers are assembling for the festive installation
of the new mayor, and we learn that the council has decreed Sachs’s
banishment for having broken the peace in entering Steffen’s gar¬
den. Sachs is informed by Kunigunde of the impending edict,
and decides to leave the city at once. Kunigunde is resolved to
accompany him, though Sachs tries to dissuade her, and as she
clings to him, Steffen with the councillors and citizens steps forth
from the city hall and proclaims the decree against Sachs, on which
Eoban comments with malicious jeers. The emperor appears in
the nick of time. After eliciting Steffen’s legal opinion on a ficti¬
tious case in which a jewel was withheld from the man to whom it
rightfully belonged, he discloses his identity, and declares Kuni¬
gunde to be the jewel and Sachs the rightful owner according to
the burgomaster’s judicial wisdom. Steffen urges his debt of grati-
96 Wisconsin Academy of Sciences, Arts, and Letters.
tude to Eoban and the latter’s supposed assistance in clothing him
with his new dignity, but is quickly disillusioned, and as the Augs¬
burg councillor steals away, Kunigunde crowns Sachs with a laurel
wreath, and the whole assembly burst out in cheers for the emperor.
From Deinhardstein’s own prolog to his play, we are led to con¬
clude that Hans Sachs is not the real hero of the drama, or that at
least he shares this distinction with Emperor Max :
“Und so empfehlen deutsche Verse wir;
die schildern, wie ein grosser deutscher Fiirst
mit Macht und Weisheit GiLte auch vereint,
und eines deutschen Dichters Eigenheit
dem heissgelieMen deutschen Vaterlande .”
The characterization of Hans Sachs is the weakest part of the play.
The reader finds it difficult to conceive how the interest of the spec¬
tator in the hyper sentimental young man can possibly be sus¬
tained.13 About half a dozen times the stage directions prescribe
that his words and ways should be gutmutig • There is nothing
about Master Steffen to commend him to our sympathy, and Eoban
has no redeeming features whatever, not even as a comic figure.
The synopsis will have sufficed to show how much the play leaves
to be desired structurally.
When Philipp Reger (1804-1857) in 1839 wrote the book for
Lortzing’s opera Hans Sachs, jointly with his friends Philipp Jakob
Diiringer (1809-1870) and Gustav Albert Lortzing (1801-1851),
basing his text on Deinhardstein’s drama, almost all of the numer¬
ous changes he made were improvements on the original.14 Rene
Doumic says, “On ne ref ait pas une piece, on en fait une autre,”
and, naturally, either a better or a poorer one. To preface what
we have to say with what will be our final verdict, Lortzing-Reger ’s
opera ( Diiringer ’s share does not amount to very much) would to
this day enjoy well merited popularity had not Wagner ’s Meister-
singeir crowded it off the stage by its superior qualities. After see¬
ing Shakespeare’s masterpieces, only an antiquarian would desire
13 And yet the audiences at the time of the first appearance of the play seem
to have taken a great fancy to this character. In Hamburg the actor imper¬
sonating him received from the shoemakers’ guild a diploma promising to him,
his children, and his children’s children, free shoes for their lifetimes.
14 The text with a very illuminative introduction by G. R. Kruse, can be
had as No. 4488 of Reclams Universal-Bibliothek.
Boedder — “Die Meistersinger von NurnbergK” 97
to witness a performance of the plays of his predecessors, no mat¬
ter how good they might be in themselves.
The first scene takes place in Sachs’s shop. The journeymen are
busy working; Gorg, the apprentice, has fallen asleep over his job
and is unceremoniously, by a blow with the stirrup, awakened by
the others, who flout his pretensions to proficiency in poetry, his
master’s art. When the hour to stop work for the day has come,
Gorg, remaining behind alone, detects a manuscript of his master’s,
a poem written for the occasion of his sweetheart’s birthday, which
Gorg purloins, in order to read it to his beloved Kordula, the cou¬
sin of Sachs’s lady-love Kunigunde, daughter of Master Steffen,
the goldsmith. An unknown visitor, Emperor Maximilian, enters
to pay his respects to Sachs, whose poetry, he tells him, the monarch
and the whole court appreciate very highly. Sachs, who has en¬
tered from the garden, can at first scarcely believe such good news,
which indemnifies him for many hours of grief in his unapprecia¬
tive native city. He is now in the right mood to compose his
prize poem for the competition that is to take place on the next
day. Then, as he is about to leave for Master Steffen’s house,
Eoban Hesse15, an Augsburg councillor and also a mastersinger,
demands his professional services in repairing a hole in his shoe.
While Gorg attends to the job, with many facetious remarks at the
stranger’s expense, Sachs learns from the latter’s words his real
purpose in coming to Nuremberg : Master Steffen has promised him
the hand of his daughter, and Sachs’s happiness seems short-lived.
Kunigunde, whom we meet in the next scene in her father’s gar¬
den, is very much distressed at Eoban ’s wooing, but the energetic
Kordula encourages her, and Sachs, who with Gorg appears for
his visit, is confident of his success in next day’s competition,
which he thinks will win over Kunigunde ’s father. But the
haughty Master Steffen, unfortunately, is just now apprised of his
election as burgomaster, and Eoban is quick to enter complaint
against the shoemaker, from whose own mouth he has heard of his
love for Kunigunde. In the contest Sachs sings16 the praise of
15 Helius Eobanus Hesse — or Hessus, since according to the custom of the
scholars of his time he Latinized his name — was a noted German humanist
(1488-1540), who wrote Latin poetry of marvelous elegance of form and
since 1526 held the chair of poetics in the newly founded Nuremberg univer¬
sity. To his contemporaries he was best known through his Bohemian habits
and his stupendous feats in drinking. His portrait, preserved in a sixteenth
century woodcut, shows him as a handsome middle-aged man of serious mien.
Being a writer of Latin verses, he naturally would have laughed at the idea
of ever joining the mastersingers.
16 Or rather speaks, except the last line, which is sung. This is one of the
inexplicable oddities of the opera, and proves that the authors knew very little
about the rules of mastersong. To the mastersingers the tune was everything,
and the text was of very subordinate consideration. Cf. Gervinus, in the
Appendix.
7— S. A. Li.
98 Wisconsin Academy of Sciences , Arts, and Letters.
Ver Liebe Gluck, das Vaterland, and the assembled people are
unanimous for according to him the prize, but Steffen decides in
favor of Eoban, who recites an incredibly silly mastersong on Ab¬
salom’s rebellion against David. The jealous mastersingers side
with Steffen, and the latter publicly announces his daughter’s be¬
trothal to Eoban. During a public festival on a meadow outside
the city walls, Gorg reads to Kordula the poem which he has filched
from his master as his own, then loses the manuscript, which falls
into the hands of some crossbowmen of the emperor, and they
pocket it with the intention of handing it to Max, whose fondness
for poetry is universally known. Hans Sachs, deeply grieved by
his humiliation at the contest in the morning, has made up his
mind to leave Nuremberg, and comes to the festival only in order
to bid farewell to Kunigunde. Like her prototype in Deinhard-
stein’s play, she tries to coax Sachs into giving up his trade, but
is not so insistent and more than willing to ask his forgiveness
when she notices how she has hurt his feelings. As they embrace,
Steffen and Eoban step out of the burgomaster’s tent, and Steffen
after a brief conference with the councillors pronounces the decree
of banishment for Sachs. Sachs leaves, accompanied by the faith¬
ful Gorg, who is resolved to share his master’s exile. Steffen soon
repents of the council’s action: Emperor Maximilian, who has just
arrived in the city, wishes to meet the author of the verses that his
soldier has handed him — Sachs’s poem which Gorg had appropri¬
ated for his own use. With Maximilian, Sachs has returned to
Nuremberg, and during a clandestine visit at Steffen’s house over¬
hears an agreement between Steffen and Eoban according to which
Eoban is to pose as the author. In the last scene at the great ban¬
quet hall the emperor demands that Eoban prove his authorship,
and his attempt fails dismally. Sachs naturally recites his poem
by heart and is publicly acknowledged as the author. After this
recognition by the sovereign, the authorities of the city are of
course only too glad to receive Sachs back into their fold, Kuni¬
gunde is solemnly betrothed to Sachs, and the curtain drops amid
jubilant cheers for the monarch.
The few external features that Wagner’s work shows in com¬
mon with Reger and Lortzing’s adaptation of Deinhardstein’s
drama may be briefly summarized as follows: A young and en¬
thusiastic poet fails before the guild of the older men, as a result
of their incapacity and unwillingness to understand and appre¬
ciate his superiority. A poem composed by the hero is stolen by a
rival, and the latter is put to shame when asked to recite it. Gorg,
Hans Sachs’s apprentice in the earlier opera, is a votary at the
shrine of the Muses, as is David in the Meister singer. Gorg and
Kordula are the forebears of David and Magdalene, excepting
Boedder — “Die Meistersinger von Nurriberg.” 99
that in the latter ease the girl is about a dozen, or perhaps a score,
of years older than her swain.17 Eoban’s song of Absalom is as
silly as Beckmesser’s serenade. That is all. That there should be
choruses of journeymen, and cobbler songs, is almost imperative
in a musical work dealing with Hans Sachs. The invention of the
festival on the meadow in Lortzing’s opera and the blooming tree
in Hans Sachs’s garden in Deinhardstein’s play would not require
an exuberant poetic imagination and need not be accounted for on
the basis of “influence.” Both features are in Wagner’s play
much more organically connected with the action than in either
of the other two works.
Of internal relationship between Reger-Lortzing’s work on one
hand and Wagner’s on the other I am not able to detect any pro¬
nounced traces. Granting, for argument’s sake, that the poetic
idea of Wagner’s comedy is present in embryo in the earlier play,
it will be generally conceded that it has been very badly stunted,
if not shriveled and wizened in its growth. If, at the time of the
publication of Wagner’s text and the first performances of the
opera, any distinct similarity with Lortzing’s work had been felt,
Wagner’s numerous enemies, who attacked the Meistersinger as
viciously as his earlier creations, would most certainly have pointed
it out, to disparage the composer. But there is to my knowledge
not a trace of this in all the acrimonious criticism of the sixties,
which today strikes us as so irresistibly amusing.
If we point out similarities in Lortzing’s and Wagner’s works,
it is but fair to call attention to a radical difference. In the ear¬
lier opera, as in Deinhardstein’s drama, the emperor plays a
very significant part and toward the end overshadows the hero
completely. In fact, Emperor Max is a sort of deus ex machina,
without whose intervention in favor of the poet the latter’s fate
would remain very dubious. Neither does in Deinhardstein’s and
Reger-Lortzing ’s plays the principal figure fight for the recogni¬
tion of a new principle in poetry: here he is only, or is supposed
to be, the superior of his colleagues and thereby becomes the ob¬
ject of their open and secret hostilities. In Wagner’s play the
victory at the end is not bestowed as a present from above; it is
17 1 do not see the necessity of assuming that I^unigunde and Kordula have
been patterned after Agathe and Aennchen in Weber’s Freischiitz. They may
have been ; but cousins of about the same age and different temper and out¬
look on life do not appear to be so exceeding scarce in the pages of literature.
100 Wisconsin Academy of Science s, Arts, and Letters.
the natural and organic result of a battle royal that Walter von
Stolzing and, directly and indirectly with him, Hans Sachs have
been fighting. There is here no monarch, the weight of whose
personal opinion could have borne down with it the resistance of
the opponents. Indeed, it is characteristic that in Wagner’s Nu¬
remberg, the free imperial city, the emperor’s name should not
once be mentioned. May we not in this fact see an indication of
the democratic sentiments that filled Wagner’s soul in the years
prior to the Revolution of 1848, and that subsequently led to his
political activity in the interest of this movement as well as to
his flight from Germany and his long exile ? And is it not worthy
of recognition that Wagner in the final execution of the Meister-
singer did not introduce any features that might have detracted
from the glory of his Nurembergers’ civic pride and love of free¬
dom, although his intimate friendship with King Ludwig II. of
Bavaria might very easily have suggested such to him ?
A word will have to suffice concerning Wagner’s indebtedness
to E. T. A. Hoffmann’s narrative Meister Martin der Kiifner und
seine Gesellen,t to which Miss Bowen assigns the distinction of
having furnished the chief motive of the plot of the Meister singer,
viz., that Master Martin’s daughter Rosa is offered as a prize for
a masterwork (not, it must be noted, for a poetic or musical mas¬
terpiece), only on condition that she herself consent (1. c., p. 54) :
“ There can be no doubt that Wagner drew his principal motive
thus from this very tale.” While Hoffmann’s influence on Wag¬
ner must be admitted — Hoffmann’s Kampf der Sanger was one
of the works that inspired the Tannhduser — the motive in ques¬
tion here is so often used in the literature of the first half of the
last century that it seems indeed presuming to try to locate its
first appearance precisely. The frequency with which it occurs
is no doubt sufficiently accounted for by the fact that in the early
decades of the nineteenth century the paterna potestas disposing
of a daughter ’s hand was already greatly on the wane in Germany.
The other references to mastersinging in Hoffmann’s tale may
possibly have kindled the youthful Wagner’s interest in this fea¬
ture of the German past, but it is just as likely that in his very
intensive studies of German antiquities he had become acquainted
with it.
There remain three more works that must be mentioned in this
connection. One of these was unearthed from the archives of
j Roedder — “Die Meistersinger von Nurnberg.” 101
the Royal Opera at Dresden by Kurt Mey18 and bears the title
Hans Sachs. Im vorgeriickten Alter. Romantisch-komisches Sing-
spiel in zwei Akten. Musik von Adalbert Gyrowetz (1763—1850).
It seems that the opera was performed, or a performance contemp¬
lated, in 1834. The writer of the book is not named. The text
is very romantic indeed, but could not have offered Wagner any
suggestions. We need not enter here into its contents. The only
feature that should be recorded is the character of the hero, who
is depicted as a mature and experienced man, a roguish poet full
of humor, a citizen proud of his work, modest and yet superior to
all the rest. This I quote on the authority of Mey, and I have
to take it on the same authority that Wagner’s conception of the
character is infinitely deeper and his portrayal much more
poetic. It is questionable whether Wagner ever knew of the ex¬
istence of Gyrowetz’ opera. His position as musical director of
the Dresden opera may have given him a chance to become ac¬
quainted with it. It is however extremely unlikely that he ever
read it. We should be willing, even if there were documentary
proof of this fact, to accept in full Wagner’s word that Die Mei¬
stersinger is his own invention.19
A scene which in its broad humor and in its combination of
comic elements may be regarded as the archetype of the scene of
Beckmesser’s serenade and the attendant circumstances is found
in August von Kotzebue’s Die deutschen Kleinstadter (1802). 20
It is found in Act IY, scenes 3-5. While the lovelorn Sperling is
singing before the window of his chosen one, Sabine herself is in
the company of Herr Olmers, to whom she has plighted her troth,
and Sabine’s aunt, Frau Staar, is singing her evening hymn, in¬
terrupted by, and in turn disturbing, Sperling’s musical efforts.
To make confusion worse confounded, the nightwatchman appears
on the scene with his call and blowing his oxhorn, and Sabine and
Olmers are forced to hide behind a lamp post until the danger is
past. All the elements entering into the delightful scene in front
of Pogner’s house while Beckmesser sings his prize song, disturbed
18 Cf. his Der Meistergesang in G-eschichte und Kunst. Leipzig- o. J. (See-
mann) .
19 In this connection it is interesting to learn that Wagner at one time con¬
templated treating Hans Sachs’s second marriage — provided that a note of
Professor Golther’s in Biihne und Welt , I, 2, 579, is correct. I am unable to
verify it.
20 My attention was called to this by two of my colleagues independently,
Miss A. B. Ernst and Professor A. R. Hohlfeld.
102 Wisconsin Academy of Sciences, Arts, and Letters.
by Hans Sachs’s Schusterlied, and Walter and Eva are hiding
behind the bushes in Pogner’s garden, are here found together.
Wagner certainly was acquainted with Kotzebue’s play, which
was a great stage favorite when he was young, and the make-up
of the scene in question must have made its way into his comedy
unconsciously.
The last work that we have to consider is Ferdinand Raimund ’s
drama Die gefesselte Phantasie. Raimund (1790-1836) is known
to German literature as the master of the Vienna magic- and fairy-
play in the early decades of the last century. His <( Phantasy En¬
chained” was written in 1826 and represents his first attempt at
a serious drama that tries to attain its effects without resorting to
parody and travesty. It was not performed before 1828, January
28, a couple of weeks before the first performance of Deinhard-
stein’s Hans Sachs.21
Phantasy, the goddess to whom all poets owe their inspiration, is
captured by two direful sorceresses, the sisters Vipria and Arro-
gantia, at a time when her assistance to her protegees is most urg¬
ently needed; for Hermione, the youthful queen of the island of
Flora, has promised her hand and heart to the successful competi¬
tor in a poetic contest, confident that her beloved Amphio will be
the victor. This Amphio is a young prince, the son of the king
of Athunt, but in true fairy tale fashion has hitherto concealed his
identity and nevertheless won Hermione ’s love, tending her fa¬
vorite flock as a shepherd and writing enamored verses in praise
of his mistress. The contest is to take place in the evening of the
day on which the play opens. When the two magic sisters learn
that it is Phantasy on whom the poets depend for their inspiration,
they waylay her and then secure a most repugnant Vienna ale¬
house harpist by the name of Nachtigall, whom they plan to make
the husband of Hermione, on the terms proposed and sworn to by
the young queen when she announced the contest. Nachtigall is
introduced to the enchained goddess, and tries first by fair words
and finally by brute force to get her to dictate to him the neces-
21 1 believe that Raimund knew Deinhardstein’s drama when he wrote his
play. There seems to be an allusion to Hans Sachs in Act I, scene 2, where
Distichon, the court poet, says : Wir haben keinen Schnee, als wenn uns
Zephyr weisse Bliiten streut ; darum begeistert uns der ewige Blumenduft und
weihet uns zu Priestern des Apoll, so dass der Schuster selbst mit einer Hand
nur seinen Stiefel schafft, in der andern halt er hoch die goldne Leier:
“Sein kiihner Geist ist mit Apoll verwandt,
1st seine Leier gleich mit Schustergaim bespannt.>>
The quotation marks here, which are Raimund’s own, seem significant. In
another place Distichon says, “Ich werfe sie mit Knittelreimen tot.” The
doggerel verse is characteristic of Hans Sachs’s tales and farces.
Boedder — “Die Meistersinger von Nurnberg.”
103
sary lines for the prize poem. From the misunderstood sentences
that Phantasy in her distress utters he manufactures the veriest
nonsense, and finally runs away in despair, determined to sing a
popular street ballad and merely to substitute the queen’s name
for the heroine’s. The poets of the whole island are in the worst
possible plight — not one of them succeeds in finding a single rhyme,
and Hermione is horrified to see that even her beloved Amphio
has turned as stupid as all the rest. The fateful hour arrives—
Nachtigall is the only candidate to enter the lists, and Vipria and
Arrogantia, disguised as the priests of Apollo, declare that Her¬
mione must wed him if nobody produces a better poem, even
though all those assembled are incensed and furious at Nachti¬
gall ’s song. All seems lost when Phantasy, in the supreme moment
freed by Jupiter, appears on the scene and inspires Amphio to
sing Hermione ’s praise in a long poem at the close of which he
reveals his identity. Apollo appears in person to plunge the two
sorceresses into the deepest depths of Orcus, and Amphio and
Hermione are married, and, let us hope, live happily ever after.
It is to be regretted that Raimund’s poetic power did not suffice
to do justice to this idea, which would have been worthy of Shakes¬
peare’s pen. He lacked markedly what Coleridge calls ‘'the
shaping spirit of imagination.” Die gefesselte Phantasie is one
of his weakest plays, and appears on the modern stage only at
long intervals. Even Schubert’s music does not sustain it on the
boards.22 In the author’s own time this evidently was different:
22 The following extract from Eugen Kilian, Felix Mottl. Karlsruher Erin-
nerungen (in Siiddeutsche Monatshefte, 1911—12, I, 488—496 ; reprinted in Aus
der Praxis der modernen Dramaturgie. Der Dramaturgischen Blatter zweite
Reihe. Miinchen 1914, pp. 294-302), gives a good idea of the possibilities con¬
tained in Raimund’s play when properly staged : “Mottl was especially de¬
lighted with conducting the musical rehearsals of Raimund’s magic-play Die
gefesselte Phantasie, with Franz Schubert’s music arranged for this purpose.
This new musical adaptation of the old Vienna magic-play, the arrangement
of the music of Schubert’s long forgotten Magic Haw for Raimund’s work,
and its further embellishment with Schubert tunes and dances, was an ex¬
ceedingly felicitous and in its way ingenious idea of Mottl’s. This superb
score deserves first rank among all his works belonging under this head.
My co-operation with Mottl in the rehearsals of this play belongs among
the most beautiful reminiscences from the time of my Karlsruhe activity.
Mottl set his whole heart on, and watched over, this creation, on which the
double star of Raimund and Schubert beamed, with a truly touching tender¬
ness. The whole kindliness and warmheartedness of Old Vienna, the naive
and yet graceful, exquisitely humorous ingenuity of Raimund’s play was a
true heart’s delight to Mottl, the Viennese. It was blood of his blood, flesh
of his flesh that here he laid on the altar of art with patriotic pride. As he
was conducting the rehearsals at the piano, he would enjoy again and again
the charm and frolicsome humor of these Old Vienna figures, and could laugh
till he cried when Fritz Herz as Nachtigall in his duo with Phantasy chained
to the writing desk — one of the most superb gems of all humorous literature !
104 Wisconsin Academy of Sciences, Arts, and Letters.
he took the play along on his tours and appeared as Nachtigall in
Munich, Hamburg, and Berlin, with distinct success (1832). Wag¬
ner may have made the acquaintance of Die gefesselte Phantasie
while he was musical director at Madgeburg. Indeed he may have
had to direct a performance of it in this capacity. Again, he may
have witnessed a performance of the play during his prolonged
stay in Vienna in 1861-1862. It is at present impossible to verify
any such surmises. Wagner had a high opinion of Raimund’s
poetic genius. In his essay on Das Wiener Hofoperntheater (1863 ;
Gesammelte Schriften vii, 295ff.), he says:23
“What Vienna quite of itself can do for even Art, on the path
of a purely speculative, an un-subventioned commerce with an
imaginative, gay and genial public, is proved by two of the most
original and delightful products in all the realm of public art : the
Magic-dramas of Baymund and the Waltzes of Strauss. If you
don ’t wish for higher things, then be content with this : indeed its
intrinsic worth is nothing to be made light of. ’ ’
And again, in TJeber Schauspieler und Sanger (written in the
early seventies, Gesammelte Schriften, 9, 186) :
“Out of the Viennese popular farce, with its types still dis¬
tinctly akin to the Kasperl and Hanswurst of old, we see Ray-
mund’s magic dramas rise up to the realm of a truly ingenious
theatrical poetry.”
Rudolf Fiirst in the introduction to his edition of Ferdinand
Raimund’s works (Goldene Klassiker-Bibliothek) , page LXV, says:
“The possiblity that the scene of the singing contest, the tri¬
umphant lover Amphio, and the caviling critic Distiehon24, in
conjunction with NachtigalFs attempts to piece together a prize
song from misunderstood fragments, may have influenced the
poet of Die Meister singer von Numb erg, can hardly be set lightly
aside.”
• — unfolded his magnificent, heartfelt humor. And when Fritz Plank, the ever
memorable, as the Vienna tapster in the capital alehouse scene of the first
act set his heavy, stout limbs in droll motion to the catching tunes of Schu¬
bert’s German Dances, then rapturous contentment gleamed in the eyes of
Mottl the artist : that was home, those were images and visions whose radi¬
ance derived from the native soil of Mozart and Grillparzer !”
23 Translation by William Ashton Ellis, Richard Wagner’s Prose Works, vol.
Ill (London, 1907), page 386.
24 “Das Gedicht ist voller Fehler.” Which it is, indeed. Amphio’s prize
song is the weakest part of the play.
105
Eoedder — “Die Meistersinger von Nurnberg.”
In its guarded form this statement is more of an hypothesis
than of a thesis. There might be something in it provided that
it can be shown that Wagner knew Raimund’s play before he pen¬
ned the Marienbad sketch. For all the features mentioned occur
also, as we have seen, in Lortzing’s opera.
What if Wagner had consciously utilized the work of his pre¬
decessors? I hope to have shown that he did not. But if Mo-
liere, charged with literary piracy, had really said, “Je prends
mon bien oil je le trouve” — a repartee invented by one of his de¬
tractors — he would have done so with a perfect right. In all
art it matters little what materials the artist uses, or where he
gets them. It does matter what he makes of them. And Die
Meistersinger von Numb erg is one of the few truly great come¬
dies of all literature.
With this assertion we are not paying Wagner’s work tribute
above its meed. The year 1918 witnessed the semi-centenial of
its first performance. Half a century is time enough for both
fulsome panegyrics and unfair personal criticism against the
poet’s individuality to give way to soberminded judgment, and
half a century has rendered its verdict.
No other work of Wagner’s is so narrowly circumscribed as re¬
gards time and place, and yet scarcely another is more universal
in its human appeal. “As a picture of medieval life,” says
Finck, 1. c., page 224, “it is as realistic, accurate, and delightful
as the best of Scott’s novels.” It is a wonderfully correct por¬
trait of sixteenth century life in the free imperial city of Nurem¬
berg in its most flourishing state, with its industrious citizens, its
proud trade and craft guilds, its pedantic but worthy and honor¬
able mastersingers, and the towering figure of its Hans Sachs. In
this respect it is the most distinctly German work of the com¬
poser, and it is noteworthy that he wrote the text during his stay
in Paris in 1861-62. In Wollen wir Jioffenf (1879; Gesammelte
Werke, x, 119 f.) he says:
“In my execution and presentation of the Meistersinger, which
at first I hoped to bring out in Nuremberg itself, I was guided
by the intention of offering to the German public a picture of its
own true nature which heretofore had been presented to it only
very unsatisfactorily, and I , cherished the fond hope of winning
serious recognition from the heart of the nobler and superior ele¬
ments of the German middle classes. An excellent performance
in the Munich Royal Court Opera House met with the warmest
106 Wisconsin Academy of Sciences , Arts , and Letters.
reception ; but curiously enough it was some French guests present
on that occasion that with great animation recognized the popular
element of my work and welcomed it as such.”
Its broad human appeal may best be judged, aside from its en¬
thusiastic reception all over the globe, if in our minds we transfer
the battles that are fought in the Meister singer to other phases of
human activity, substituting for the mastersingers any other set
of more or less self-centered artists, or professional men, or in¬
deed men from any walk of life, in any country, at any time.
The comic elements will remain the same: there are the jealousies
and rivalries of the individuals making up the class among them¬
selves, the overt or covert animosities against the daring innova¬
tors that chafe against the narrow fetters of tradition, the con¬
certed fight against outsiders, the unconfessed but ever present
opposition of old age against the rising generation, and the event¬
ual triumph of youth. And there is the everlasting and irresist¬
ible comedy element, la lutte pour la femme, here aggravated
by the fact that the rivals are not only representatives of the
younger and the older generations, not only the impersonations
of romanticism and philistinism respectively, a dashing young
knight and a pompous old scribe, but additional zest is lent to
their fight because they are the representatives of young and old
in art. And these elements are treated with a humor
“essentially German and Wagnerian — a combination of play¬
fulness, exuberant animal spirits, practical jokes, puns, burlesque,
and withal an undercurrent of amiability, seriousness, passion, and
even sadness, as in all great humorous literature. Every form of
humor is represented, the lowest as well as the highest; from the
horse-play accompanying the riot scene, the pun on Vogelgesang’s
name, and the broad burlesque of Beckmesser’s serenade, to the
more subtle persiflage of Kothner’s address, the merry mockery
of the apprentices, the quaint spectacle of the watchman, the
chivalrous bluster of the knight, the rollicking cobbler songs, and
the subtle satire of Sachs. In this variety of humor, from the
lowest to the highest, Wagner resembles Shakespeare.”25
The chief charm of Wagner’s comedy-drama, however, lies not
in such comic incidents, no matter how delightful they may be in
themselves, but in the delineation of the characters. This is what
gives the play imperishable value.
25 Finck, l.c., page 225.
Roedder — “Die Meistersinger von Numberg.”
107
A word will have to suffice for Eva, this truly winsome and be¬
witching lineal descendant of our general mother in paradise, and
first cousin to Julia Capulet in her frankness and outspokenness
in love as well as her astounding daring in taking her fate in her
own hands, irrespective of the desire of her beloved father. She
is especially engaging in the manner in which she tries to draw out
her father and her old friend Hans Sachs concerning Walter von
Stolzing’s success in the singing test in St. Catherine’s Church,
in her playfully disguised apprehension that Sachs might enter
the fateful singing contest on St. John’s Day, in her pouting when
she misunderstands Sachs’s subtle irony as he refers to the other
mastersingers ’ feeling of inferiority towards Walter, and in her
complaint about all the trouble she has with the men folk when
Walter is about to rush upon Beckmesser in front of Pogner’s
house. She becomes a pathetic figure in the moment when the
realization of Sachs’s true feelings for her and his suppressed
grief overwhelms her, and yet how engagingly natural is her re¬
turn to the sun of her happiness in Walter’s presence immediately
thereafter. Eva is altogether one of the most fascinating youth¬
ful figures in all operatic literature.
On Magdalene, her confidante, David’s sweetheart, and on David
we need not dwell. Wagner avoided the pitfall of making these
two characters mere repetitions of types, and provided them with
sufficient individuality to impress them on the spectator’s memory
as beings of flesh and blood.
The mastersingers as a class are excellently described by Kreh-
biel, 1. c., p. 92f., where he sets forth the two melodies that char¬
acterize them throughout the comedy:
4 ‘Note that as the mastersingers belonged to the solid burghers
of old Nuremberg — a little vain, as was to be expected in the up¬
holders of an institution of great antiquity and glorious traditions ;
staid, dignified, and complacent, as became the free citizens of a
free imperial city, whose stout walls sheltered the best in art and
science that Germany could boast — so these two melodies are
strong, simple tunes; sequences of the intervals of the simple dia¬
tonic scale ; strongly and simply harmonized ; square-cut in rhythm ;
firm and dignified, if a trifle pompous, in their stride.” \
The musical characterization of the mastersingers alone would
suffice to convince any one that in the figure of Beckmesser Wag¬
ner did not mean to put on his feet the typical mastersinger, even
if we forgot for a moment that Hans Sachs is a mastersinger as
108 Wisconsin Academy of Sciences, Arts, and Letters.
well as Beckmesser. The latter simply is the personification of
mastersong carried to its last consequences, and as such he stands
out in the prelude to the work. In him Wagner chastises all that
is foolish in the mastersinger’s ways, with a smile, for all great
comedy ridendo castigat mores. For the subject of this chastise¬
ment Wagner
1 ‘ chooses only things which are temporary aberrations from the
good. What is strong and pure and wholesome in the art of the
mastersingers he permits to pass through his satirical fires un¬
scathed. Classicism in its original sense, as the conservator of
that which is highest and best in art, he leaves unharmed. ' ' (Kreh-
biel, p. 96).
Beckmesser is, then, the arch exponent of the rule-of-thumb pe¬
dantry and formalism; the worshipper of the ossified common¬
place ; a stubborn believer in the necessity of the unchanging main¬
tenance of a status which the progress of time has long since ren¬
dered untenable ; such an upholder of tradition that he would re¬
gard the best possible innovation as worthless in comparison to
the imitation of inferior models, provided that these be old; an
example of Robert Burns's statement, which is more emphatic than
is consistent with our latter day notions of good breeding, that
“a man may be an excellent judge of poetry by square and rule,
and after all be a d — — blockhead "-—except that Beckmesser has
ceased to be an excellent judge. But while he is pavilioned in the
glittering pride of his supposed accomplishments as a mastersinger,
he is by no means unaware of his lack of personal attractiveness
to the fair sex, and so he endeavors to assure himself of the sup¬
port of Eva's father, and does not approve of the stipulation that
Eva herself may reject the successful competitor in the singing
contest if he is not to her liking. As a further consequence, he
strives to remove his rival Walter von Stolzing by declaring him
versungen at the test ; and as his follies advance in geometrical pro¬
gression, he at last becomes a thief and steals what at the time he
regards as the song with which Hans Sachs would enter the lists
on St. John's Day. Many critics, sincere admirers of Wagner's
art among them, have taken exception to this character delineation.
Thus Krehbiel says, 1. e., page 80 :
“Beckmesser ought not to have been made the blundering idiot
and foolish knave that he appears to be in the stage versions, but
at the worst a shortsighted, narrowminded, and perhaps malicious
Boedder — “Die Meistersinger von Nurnberg.”
109
pedant. As he stands in the stage representations Beckmesser is
an ill-natured and wicked buffoon, a caricature of a peculiarly
gross kind, and only an infinitesimally tiny corrective idea lies in
the fact that a manly young knight who loves a pretty young wo¬
man should have saved her from falling into such a rival’s hands
by marrying her himself. He would have had the vote of the
public on his side if he had sung like a crow and Beckmesser like
Anacreon. ’ ’
This last statement, in order to clear one objection out of the
way, seems to me begging the question. First, the public had no
vote in the matter, Hans Sachs’s motion at the meeting in St.
Catherine ’s having been voted down ; and secondly, it was a mat¬
ter of outsinging the competitors and not of personal attractive¬
ness and similar advantages. But to revert to the charge, let us
quote Erich von Schrenck:26
“In Beckmesser there is so much grotesque exaggeration that
sober criticism should not be content with mere praise. One need
only think of the mutilated prize song, with which Beckmesser al¬
most ranges himself among the lunatics. It is not easy to see how
critics can pass over such matters as though everything were here
as it should be. ’ ’
On the other hand Finck, p. 228, has this to say :
“Another amusing actual feature in this comedy is that some
of the critics who feel more or less guilty of having once been
Beckmessers, still are a little sore on the subject and mercilessly
abuse actors who are intelligent enough to treat this part in a real
burlesque spirit. But Wagner shows by his whole treatment of
this role — the blackboard scene, the tuning and twanging of the
lute, the grotesque serenade, the antics (musical and mimic) in
Sachs’s room after the fight, and especially the laughable parody
of the prize song on the little stand on the meadow, that he in¬
tended this character to be essentially a burlesque, and not the
doleful, dignified duffer the critics referred to would have it. Wag¬
ner even rewrote the mock prize song and made it more extrava¬
gant than before. Beckmesser is naturally a silly fellow, and in
this case his pedantry, arrogance, and incompetence are aggravated
in such a manner by blinding jealousy that he cannot help mak¬
ing a fool of himself. If he did not make a fool of himself, why
should the people laugh at him loudly, and the Masters exclaim:
“What does this mean? Can he be crazy?” — A note to this page
adds: “Of course, the self -burlesque must be unconscious on
26 Richard Wagner als DicTiter, Munchen 1913, p. 167.
110 Wisconsin Academy of Sciences Arts , and Letters.
Beckmesser’s part. Wagner wrote to a tenor in 1872: ‘Be seri¬
ous throughout . . . Great pettiness and much gall. Take as
a model any captious critic. ’ ’ ’
Wagner’s mention of the “captious critic” brings up a matter
on which the dust of controversy has not yet settled, namely, the
amount of autobiographical material that is, or is supposed to be,
contained in his Meister singer.
“A word of caution,” says Krehbiel, p. 79f., “should be uttered
against the autobiographic stamp which some extremists have
wanted to impress upon it. The comedy is not rendered more in¬
teresting or its satire more admirable by thinking of Walter as
the prototype of Wagner himself, of Beckmesser as Wagner’s op¬
ponents, and of Hans Sachs as King Ludwig, embodying in him¬
self, furthermore, the symbol of enlightened public opinion, which
neither despises rules nor is willing to be ridden by them. Such
an exposition of its symbolism lies near enough in its broad lines,
but there is danger in carrying it through all the details of the
plot. When it is too far pushed, critics will ask in the future,
as they have asked in the past,' how this can be accepted as the
satirical motive of the comedy when the hero who triumphs over
the supposed evil principle in the drama does so, not to advance
the virtue which stands in opposition to that evil principle, but
simply to win a bride — a purpose that is purely selfish, however
amiable and commendable it may be. Walter does, indeed, dis¬
cover himself as the champion of spontaneous, vital art, and the
antagonist of the pedantry represented by the mastersingers ; but
this is not until after he has learned that he can only win the
young lady by himself becoming a member of the guild, and de¬
feating all comers at the tournament of song. Knowing none of
the rules, he boldly relies on the potency of the inspiration begot¬
ten by his love, and does his best under the circumstances ; that he
ultimately succeeds he owes to the help of Sachs, and the fact that
his rival defeats himself by resorting to foul means.”
The very fact that the naive spectator all over the world can
appreciate and enjoy Wagner’s Meistersinger without knowing
how much of the poet ’s own experience is contained therein, should
give us pause when we search for autobiographical material in this
play. This criterion furnishes the best of evidence that we are
dealing with a genuine work of art, one in which the gold has been
cleansed in the furnace of creative inspiration of all the dross of
individual experience which had clung to it. To be sure, Wag¬
ner’s works, like Goethe’s, and like all great works of art, are
‘ 1 fragments of a great confession, ’ ’ but the exact knowledge of the
Roedder — “Die Meistersinger von Niirnberg.,f 111
individual experiences that the artist had to undergo is not indis¬
pensable for the enjoyment of his creation; indeed, it may even
prove an obstacle. On the other hand it must be granted that it
may also augment its charm.
Now the last named feature is exactly what may be claimed for
Die Meistersinger. Of course, it would be narrowminded to state
in cut and dried fashion that Wagner identifies himself with Wal¬
ter, and the like. Wagner is identical with Walter, but he is also
identical with Hans Sachs, in both cases to a certain extent. We
know from his letters that in the delineation of Pogner, especially
m the musical part of this character, he meant to do homage to his
friend Otto Wesendonk, Mathilde’s husband. We know that for
some time he had intended to call the present Beckmesser by the
name of his bitterest critic, Hanslick — his name appears as Hans
Lick and as Hanslich in some of the drafts preceding the final ver¬
sion. On sober second thought he abandoned such purely personal
allusions, certainly not to the detriment of the finished work. The
figure of Beckmesser stands for the whole chorus of hostile critics
that greeted the appearance of every new work of Wagner’s with
a volley of vituperation. In writing out the final version he may
very well have had in mind the fate of his Tannhauser in Paris.
It may not be amiss to hear on this point the drastic words of El¬
bert Hubbard :27
“Mr. Henry I. Frick has compiled a list of over one hundred
names of musical critics who placed themselves on record in opposi¬
tion to Richard Wagner and his music. Only such men as proved
themselves past masters and adepts in abuse are given a place in
this Academy of Immortals. No writer, musician or artist who
ever lived brought down on his head an equal amount of contumely
and disparagement as Richard Wagner. Turner, Millet and Rodin
have been let off lightly compared with the fate that was Wagner’s ;
and even the shrill outcry that was raised in Boston at sight of
MacMonnies’ Bacchante was a passing zephyr to the storm that
broke over the head of Wagner in Paris when after one hundred
and sixteen rehearsals Tannhauser was produced. The derisive
laughter, catcalls, shouts, hisses and uproar that greeted the play
were only the shadow of the criticisms that filled the daily press,
done by writers who mistook their own anserine limitations for
inanity on the part of the composer. They scorned the melody
they could not appreciate, like men who deny the sounds they can¬
not hear, or those who might revile the colors they could not dis-
27 Richard Wagner, in Little Journeys to the Homes of Great Musicians ;
vol. VIII, No. 1 (January, 1901).
112 Wisconsin Academy of Sciences , Arts , and Letters.
tinguish. And worse than all this, the aristocratic hoodlums re¬
fused to allow any one else to enjoy, and would not tolerate the
thought that that which to them was 4 jumbling discord, seven times
confounded’ might be a succession of harmonies to one whose per¬
ceptions were more fully developed.”
Some of the criticisms that Beckmesser and a few of the other
masters hurl at Walter’s trial song sound as though Wagner had
taken them bodily from the paragraphs of the daily press dealing
with his works. But, it should be noted, there is, thanks to his
humor, not a trace of bitterness in Wagner’s treatment of these
matters, he has risen superior to it all.
The comedy character of the play demanded imperatively the
introduction of a figure of the type of Beckmesser. Wagner had
to show where mastersong, as the representative of formalism in
art, in its ultimate consequences would lead. And if we grant the
poet’s right of using his personal experiences, Beckmesser had to
resort to theft in order to show what would become of Wagner’s
art in the hands of those that were called but not elect. A cer¬
tain, if you will a large, amount of caricature and overdrawing is
present in this figure, but in Wagner’s case it would hardly be
appropriate to call this a concession to public taste — a good per¬
formance (the crucial test of every dramatic work) should also in
the case of Beckmesser not leave an unpleasant taste in one’s
mouth. One question which so far has not been raised by the
commentators is this, how does a man like Beckmesser attain to
the office of marker in the mastersingers ’ guild? It seems incon¬
ceivable, with such men as Hans Sachs, Pogner, and Nachtigall in
the society. Here, if anywhere, we have an unconscious concession
to comedy, which in motivation does not appear to make such rigo¬
rous demands as tragedy.
Why did Wagner choose the name of Beckmesser for this charac¬
ter? It is that of an historical figure, a mastersinger of Nurem¬
berg and contemporary of Hans Sachs, an honorable man who in
his life had done nothing to be held up to derision. Is it the sound
of the name which in itself strikes the ear as funny, reminding as
it does of the fussy bleating of a goat ? The etymological meaning
certainly has nothing to do with it.
Walter von Stolzing! There is music in the very name. And
indeed he seems a direct descendant of Walter von der Yogel-
weide, the greatest lyrical genius of the German Middle Ages. For
he is a poet, not merely a rhymester, but one who has something
Roedder-—“ Die Meistersinger von Nurriberg.” 113
new to say, thoughts that breathe, and is possessed of the gift to
say it beautifully, in words that burn. Even though his poetry
has about it something dreamlike, visionary, hazy, there is ‘ ‘ clearer
promise of refulgent day”, and Hans Sachs, who in clearness and
strength of vision and in luminous clarity of diction is Walter’s
superior, willingly yields homage to the genius of the younger man.
And if Beckmesser and his like are afflicted with arteriosclerosis,
Walter’s blood courses through his veins with leaps and bounds,
and it will take some time yet before it will flow in the even, warm
pulse-beat of Hans Sachs. In his make-up the elements of caution
and expediency are noticeably lacking — to this extent he is the
image of his creator— -and the sense of humor is entirely absent,
in which respect he differs widely from Wagner as well as Hans
Sachs. He presents a wonderfully impressive picture when “in
shape and gesture proudly eminent”, seeing his attempt at winning
the master’s title fruitless, he still continues to sing, hurling de¬
fiance at the masters’ code of rules, and yielding homage only to
the eternal laws of poetry — as he feels them. There is here a
marked contrast between the “young man” of the Marienbad
sketch and the Walter von Stolzing of the final version; and from
the former’s “Have mercy, masters!” to the latter’s “Nicht Mei-
ster, nein! Will ohne Meister selig sein!” seems indeed a far cry.
True, even in the original draft he finally disdains the distinction
conferred on him, but here it looks more like the petulant vexa¬
tion and self-sufficiency of the youth who exults in his victory over
old age, and not as the expression of a deep conviction. In refus¬
ing the masters’ badge in the final form, however, Walter is con¬
sistent, for he has from the beginning regarded his singing before
the masters in St. Catherine’s as merely instrumental in winning
the hand of Eva. It cannot be denied that the impression he cre¬
ates even in the final version by his blunt refusal is not pleasant
to the spectator, and is felt as indicating a lack of tact, which only
Hans Sachs’s grand speech “Veracktet mir die Meister nicht und
ekrt mir ikre Kunst” succeeds in effacing. Even Walter’s noble
discontent with his Nuremberg experiences and his burning feel¬
ing that “all is dross that is not Helena” should not permit his
passionate unrestraint to leap forth. For he forgets entirely that
what won him Eva’s hand and Nuremberg’s approval after all was
a mastersong, composed with the kindly assistance of his friend
Hans Sachs— even though he has enriched and ennobled the form
— and that he did not offer new wine in new vessels, but that it
8— S. A. L.
114 Wisconsin Academy of Sciences, Arts, and Letters .
sparkled in the old.28 Still the final effect after Hans Sachs’s ad¬
dress is one of perfect harmony, and we feel that now Walter has
indeed bowed to the older poet’s matnrer insight and judgment,
and is not simply submitting to the wish of Sachs that he refrain
from hurting the feelings of his elders. There is a beautiful sym¬
bolism of enduring value in the final scene, when Walter accepts
the mastersingers ’ chain and Eva places Walter’s crown on Sachs’s
head, and the orchestra bursts forth in magnificent strains glorify¬
ing the union of the new art and the old. The old art willingly
and happily passes on its laurel to the new, and the new is willing
and happy to recognize what there is of intrinsic worth in the old —
just as Goethe and Schiller in Rietschel’s monument instinctively
grasp the same wreath.
Hans Sachs’s great speech at the close of the play, in which the
influence of Gervinus’s vigorous portrait can be distinctly traced,
is, as we have seen, contained also in the first draft of Die Meister-
singer. But one who is familiar only with the final version can¬
not help being surprised by the passage quoted above, “Thereby
he assuages the mastersingers, and wins them over.” For in the
play as we have it today, Sachs’s leadership in the guild as well
as among the population of Nuremberg at large admits of no doubt.
Not so, however, the Hans Sachs of the Marienbad sketch. A man
who can advise an ardent lover of poetry like the young knight
there depicted to give up writing verses altogether, cannot feel so
sure of his ground as to become the recognized leader of his master-
singer associates. The sketch says, 4 ‘ They have their scruples about
him and doubt whether he means honorably by the guild. . . .
His conduct at times seems dubious to the masters.” Eva warns
Walter against him, “Do not trust him, he is a false man! . . .
Father has often told me.” This trait must appear passing
strange to any one who calls to mind even a few of the outward
signs of the charming relation between the aged master and fair
Eva, such as their playful chat in the second act (entirely lacking
If we wish to allow for the autobiographical element in this, the interpre¬
tation would be easy enough. Wagner says to his opponents, “Gentlemen, the
moment I make up my mind, I am master of the form you worship, and I can
handle it even better than you !” Was not the quintet in the first part of the
third act born of this spirit? Some interpreters call it a concesssion to the
old operatic form. No doubt Wagner here deviates from his musical theory.
But the beauty of this quintet, which has rightly been said to dwarf any simi¬
lar effort in all musical literature, is likely to reconcile even an extreme theorist
with this departure from the composer’s principles.
Roedder — -“Die Meistersinger von Nurnberg.” 115
in the original sketch), and the girl’s exclamation, “Ach, der hat
mich lieb!”
In view of the fact that the outward events in both the first and
final versions are virtually the same, Wagner’s statement that for
the final execution of the play the original draft offered little or
nothing would seem extraordinary. It becomes clear as soon as we
realize that in the final version Hans Sachs, heretofore merely an
ornament — even though owing to his historical character and his
intellectual caliber he rose considerably above his surroundings —
is now the vitalizing center of the whole play, and that everything
hinges on his love for Eva and his renunciation.29 The action is
now placed within his soul and finds expression in tones rather
than words. Only once, in the quintet of Act III, does Sachs
softly say to himself :
“For dem Einde lieblich hehr
mocht’ ich gem wohl singen;
doch des Herzens suss ’ Beschwer
gait es zu bezwingen.
’s war ein schoner Abendtraum:
dran zu deuten wag ’ ich haum”
For a long time he has loved Eva, and on St. John’s day, his
birthday, in the singing contest at the public festival, his heart’s
fondest wish is to be consummated : no longer young in years, but
buoyant in the undying youth of genius, he would outsing his com¬
petitors and carry off the fair prize. And everybody else has ex¬
pected this outcome: Eva herself, who up to this time has loved
the master with the fondness of a child ; her father ; David the ap¬
prentice; David’s friends; and the malicious Beckmesser. Now
Walter von Stolzing appears, and all is changed at once. Eva has
fallen in love with him at first sight: “Doch nun hat’s mich ge-
wdhlt zu nie gekannter Quad: und werd ’ ich heut ’ vermahlt, so
war’s ohn ’ alle Wahl! Das war ein Musseny war ein Zwang.”
But long before he finds out from Eva herself that she loves the
young knight, Sachs with quick intuition sees how matters stand:
Walter’s request to be received into the mastersinger ’s society, by
all the others treated as a surprising incident, opens his eyes, and
29 Cf. on this point especially Houston Stewart Chamberlain, Richard Wag¬
ner. Translated from the German by G. Ainslee Hight and revised by the
author. London and Philadelphia, 1900, page 281ff. Also the fine essay by
Julius Burghold, Wagners Meistersinger: Erlebnis und Dichtung } in Ludwig
Frankensteins Richard Wagner-Jahrbuch, I, 41f£.
116 Wisconsin Academy of Sciences , Arts, and Letters.
his resolve to leave the field to him is taken at once. To be sure,
his sly joke at Beckmesser’s expense, “Nickt dock, Herr Merher!
Aus jungrem Wachs als ich und Ihr muss der Freier sein, soil
Evchen ihm den Preis verleihn,” would seem to indicate that he
harbors no desire to enter the lists; still it is here nothing more
than a well-meant advice to Beckmesser not to make himself ridicu¬
lous by competing. But soon after, while Sachs continues “to
show to the world a serene and energetic face, ’ ’ a plaintive motive
from the orchestra reveals to us his real feelings ; it is repeated in
Act II, during his chat with Eva, at the words liJa Kind! eine
Freiung machte mir Not,” (the word Freiung being here taken in
a double sense), and soon after this in the third stanza of the merry
cobbler song a similar strain strikes our ears and hearts, and Eva
exclaims, “ Mich schmerzt das Lied, ich weiss nickt wie!” These
strong undercurrents are most pronounced in the prelude and the
first scene of the third act, picturing the last struggle in the aged
man’s soul, until in the “lofty mirth which heals our pains” he
achieves the triumph over his passion and “calmed and appeased
reaches the serenity of a mild and blissful resignation.”
This last quotation, from his Entwiirfe, Gedanken, Fragmente,
may be applied to Wagner’s own state of mind at the time when
he finished the drama. It is the culmination of one of the deepest
experiences in the poet’s own life, his love for Mathilde Wesen-
donk, his victory over himself, and his renunciation. “Now only
am I fully resigned,” he writes to Mathilde, as he is working on
the final draft, and “Look out for your heart when you meet Hans
Sachs! you will surely fall in love with him.” By giving it im¬
mortal expression in a creation of his genius, Wagner the artist
has risen above the painful experience of Wagner the man. Now
he has made himself truly a master — master of his art, and master
of himself, and the fact that from this time on Wagner loved to
call and sign himself der Meister gains a deepened significance.
Just as the historic Hans Sachs is one of the most interesting
and impressive figures of sixteenth century Germany, the Hans
Sachs of Wagner has become one of the most sympathetic charac¬
ters of the highest type of comedy, and it is difficult indeed not
to fall in love with him. What endears him to us is his truly
God-given humor — not simply an intellectual gift, to be defined as
the sense of proportion, but a quality of the heart that recognizes*
the ordained place for everything in the universe and is willing
to content itself with its place in the eternal scheme of things.
Roedder — “Die Meistersinger von Nurnberg.”
117
Its undercurrent of sadness makes it all the more lovable — indeed
this is the token of all true humor: “ There’s not a string at¬
tuned to mirth but has its chord in melancholy” sighs Thomas
Hood, and “ Chords that vibrate sweetest pleasure thrill the deep¬
est notes of woe” moans Robert Burns.
Such glorious humor is not the chief trait of the earlier Hans
Sachs, the one of the Marienbad sketch. He is far from conquer¬
ing the world; his humor is a rather pessimistic irony which at
best toys with, and generally scorns and derides, the world. The
changes in Hans Sachs’s philosophy of life incorporated in the
second version may easily enough be explained on general grounds.
When Wagner wrote the Marienbad sketch he had not yet reached
the prime of life. When he finished the play he was almost fifty
years old. His Lohengrin, the colossal Der Ring des Nihelungen,
his Tristan und Isolde ,, as well as most of the theoretical writings,
had in the meantime been completed. The happy period at Dres¬
den had been followed abruptly by his ten years’ exile in Switzer¬
land. New years of migration had succeeeded the latter. They
had brought numerous handicaps to his development and creative¬
ness. He had gone through the most serious domestic and finan¬
cial distress, and suffered the bitterest disappointments, such as
the open hostility of most of the contemporary critics to his work.
But all these factors combined could not make him lose faith in his
mission, and he felt now as sure of his ground and his place in the
history of art as Schiller did when to clarify his views he wrote
his treatise TJeber naive und sentiment alische Dichtung.
The victory that Walter von Stolzing with his art wins in the
Marienbad draft is at best half a victory — with Hans Sachs as
there depicted the game is scarce worth the candle. Considering
the superior greatness of Hans Sachs in the final execution, both
as a poet and as a man, Walter’s victory is complete. Nor is its
force lessened by the fact that in some respects Hans Sachs is
decidedly the younger man’s superior even as a poet. Such he is,
as Erich von Schrenck points out, 1. c., page 169ff., in the measure
in which the naive artist — he who is one with nature — is superior
to the sentimental one who seeks nature:
“Sachs’s poetic pictures are delineated with bold, firm strokes*
Walter is ever in danger of becoming verbose, sometimes even
diffuse. Sachs shines in sunny clarity, Walter has something
vague and romantic about him. . . . For Walter the poetry of
dreams and the prose of actual life are much farther apart than
118 Wisconsin Academy of Sciences , Arts, and Letters.
for Sachs. Sachs looks upon all life as poetic, Walter flees from
prosaic reality into the world of poetry. For Sachs everything, no
matter how trite and trivial it may seem, becomes an object of
poetic musing, Walter seeks his ‘poetic’ themes. Therefore Sachs
is more versatile than Walter. Sachs takes joy in all things, he
abandons himself to them. Walter selects, and always connects
with his subject matter a relation to his own soul. Sachs is an
objective poet with a subjective element, Walter is purely subjec¬
tive. . . . Sachs regards the hostile powers with the eyes of
the wise man: he laughs at them and outwits them; Walter looks
upon them with the eyes of a knight who places himself in posture
for battle. Therefore Sachs’s poetry, like its creator, is humor¬
ous; Walter’s, pathetic ... Of course, both types of poets
have their justification, their significance. This is beautifully
shown in Sachs’s conduct. For he, who certainly is the greater,
willingly bows before the young man who possesses one side of
art that he, the mature poet, lacks. The sweetness of dream poetry,
the perfume of romanticism have so captured Sachs’s susceptible
poetic soul that the fiery ‘poet-hero’ appears to him as the true,
heaven-gifted poet. His own workaday poetry seems to him irrele¬
vant in comparison . . . However, both types are needed : the
sunny poetry of daylight with its clear features and sharp con¬
tours, with its irradiation and transfiguration of sober reality, as
well as the vague moonlight poetry with its diffuse lines and myste¬
rious figures — naive and sentimental poetry. ’ ’
Naturally Wagner did not intend to exemplify in his work
Schiller’s famous distinction— this is a by-product of the composer’s
creation, and as such interesting in the extreme. Neither did he
attempt a portrait of the historical Hans Sachs ; at the time of the
origin of the first sketch he expressed in no uncertain terms his
disapproval of the biographical Goethe and Schiller dramas that
were popular in the forties and fifties. Wagner simply desired to
place among the circle of the mastersingers an eminent, real poet,
such as the historic Hans Sachs was. It is worth while to observe
that while the earlier Hans Sachs reminds us more of the Hans
Sachs of the thirties and forties as Gervinus portrays him— for, as
we have seen, Wagner designates Gervinus as one of his sources —
the Hans Sachs as we have him today resembles more closely the
Hans Sachs of the fifties in the same treatise. It may well be left
to the reader to trace out the corresponding features in the passages
quoted in the Appendix. When his general character was thus
altered, Wagner quite naturally dropped the allusion to the Refor¬
mation in the first draft. Such an allusion seemed a matter of
course in a drama the scene of which was laid in sixteenth century
Roedder — ‘‘Die Meistersinger von Niirriberg.”
119
Germany, but as it stands, the result of this great movement ap¬
pears destructive rather than constructive — the Reformation is not
treated as one of the great fructifying epochs of the race but as
subversive of the last remnants of poetry and artistic endeavor.30
By infusing into Hans Sachs’s veins drops of his own blood,
Wagner has endowed his hero with a broader human appeal than
the genial cobbler-poet of the sixteenth century could ever have
for our own time. His innermost life, as we have seen, is too deep
for words — a linguistic expression of the depth of his love, the in¬
tensity of his grief, and the greatness of his self-conquest would
have destroyed the glorious virility of this character. But twin-
born with the words, the expression of the masculine conceptual
thought of the poet, is music, in which the feminine lyrism of the
emotion finds its voice. It constitutes the lasting triumph of Wag¬
ner’s genius that in his dramas we have “ music not written for
the text nor text written for the music, but music and text created
at the same time, the melody mirroring forth the soul of the
words.” One need not himself be a poet, full of limitless feeling
struggling with the limited utterance of words, to fathom that a
new art has here been given birth, fulfilling the age-long yearning
of sage and poet. To the word-tone-poet the lines of Oliver Wendell
Holmes ‘ ‘ The flowering moments of our mind lose half their petals
in our speech” do not apply. Music imparts to these our flower¬
ing moments immortal beauty of color and imperishable fragrance.
30 The praise of practical political activity in connection with the reference
to the Reformation, so alien to Wagner’s ways of thinking in his later years,
Julius Burghold, l.c., attributes to the interest and sympathy with which the
coming Revolution of 1848 was even then filling Wagner’s soul. It seems to
me more likely that Gervinus had at least a large share in it — all the more
so since, as Golther points out, l.c., page 297, this praise of practical political
life is really a superfluous departure even in the original version, as is evi¬
dent from the young knight’s answer to Hans Sachs’s admonitions, “Very
well, master ! But now I need a wife.” Together with the allusion to tne
Reformation, all the extraneous elements of the original scholarly apparatus
were excised, i. e., the references to the Heldenbuch , to Wolfram and his Par-
zival, to the Nibelungenlied with Siegfried and Kriemhild, of whom Walter
was to sing before the masters in St. Catherine’s. Only the reference to Wal¬
ter von der Vogelweide remained — naturally enough, for the mastersingers
counted him as one of the founders of their art, and Walter’s baptismal name,
for the earlier Konrad, was given him with a definite purpose.
120 Wisconsin Academy of Sciences, Arts , and Letters.
APPENDIX
Extracts from G. G. Gervinus, Geschichte der deutschen Dichtung ,
2. Band, 4. Auflage, Leipzig 1853.#
Chapter VI. Decline of Knightly Poetry and Transition to
Popular Poetry. 10. Mastersong.
(Page 245.) But it may be wrong to look at these songs at all
from the viewpoint of poetry. For historical purposes it suffices
to have shown that their poetic texts mark the uttermost decline
of the old lyric poetry. These songs were not meant for publicity,
let alone immortality, they remained consequently unprinted in
their obscurity, and history had better let them rest there. Even
in the case of minnesong we disdained a too searching analysis,
and here the analytic method would be still less proper. A more
accurate characterization of mastersong would incontestably be the
task of the historian of music, if indeed its music were preserved.
The mastersingers, in the period when they had established real
singing schools, let themselves be heard only as singers. With
them, as well as with the French and Dutch rhetoricians, the high¬
est achievement was the invention of a new tune; in their tunes,
however, melody was the essential thing. Little depended on the
text ; it was permissible to render the same text in different tunes ;
only in melody were they inventive; and this must not encroach
on the tunes of other mastersingers to the extent of four syllables,
but melody and fioriture were to be entirely original. And so in
the examination of such new tunes great emphasis was laid on the
musical rendition. . . . If it was found that the melody did
not encroach on any other tune to the extent of four syllables, it
was registered, its parent had it christened and invited sponsors
to the ceremony.
(Page 246). We deemed the relation of minnesong to the moral
status of the people far more significant than its aesthetic side,
and we do this also in the case of the mastersong. There we had
* Sincere thanks are due to Professor B. Q. Morgan for generous help in the
preparation of this translation as well as for the revision of the entire manu¬
script.
Boedder — “Die Meistersinger von Niirriberg.” 121
found that the soulful minnesong exercised wonderful power in
helping to check the brutality and violence of the knights. In the
fifteenth and sixteenth centuries all the lower classes were in a
state of ferment ; a rush and race for gain, distinction and author¬
ity seized even the lowly, and . . . mutual jealousy, hostility,
and persecution between the various ranks of society and the vari¬
ous crafts disfigured the middle classes of the time. Considering
all the intensity of professional jealousy and the downright char¬
acter of this sort of people, what a solidity and soundness of nature
must we presuppose to account for the fact that men united and
segregated themselves in the collective guild of the singers, threw
a common bond about the citizens, and educated them to a touching
devotion to a purpose that was not infected by any selfishness, that
kept aloof from baseness and vulgarity, and could only found
friendship and mutual confidence. Even though the great ma¬
jority of artisans and craftsmen after their day’s work would re¬
pair to the ale-house, it was all the more, at a time when the physi¬
cal vices were so monstrously rampant, wholesome that at least a
number of stalwart masters applied their hours and days of rest
from labor to something worthier, in taking down the old art of
the courts into their circle and essaying to arouse and keep alive
an interest in it ; for had not Hans Sachs so popularized the gra¬
cious art that around him there were two hundred and fifty mas¬
ters in Nuremberg? These would sit down after the hard work
of the day and compose their songs, think out new tunes and prac¬
tice the old ones, copy everything in big books, and take joy in pre¬
serving for posterity what with love and gratitude they had in¬
herited from their predecessors. The dignity of the life and the
unselfishness of these masters compensates us for their labored art.
Heretofore poetry had begged for bread at the courts and even in
its most flourishing period had not cast off the parasitic note to¬
ward Maecenases and patrons, but mastersong is the foundation of
our modern independent poetry in this respect also, that it taught
people how in the heartfelt practice of a fine pursuit, even with
indifferent success, there is a felicity that needs no further reward.
With what self-denial did the good masters devote themselves with¬
out any compensation to instructing their apprentices and pupils
in the difficult tunes, depriving themselves of rest and sleep in
order to have leisure to recruit and educate new votaries to their
art, since the day was filled with their professional labors. And
with what love did the pupil then look up to his master !
122 Wisconsin Academy of Sciences , Arts , and Letters.
(Page 251). The mastersinger tablatures do to be sure repre¬
sent most defective poesy, the weakness of which as compared with
ancient classical poetry was at once recognized, and the reason why
the mastersingers affected so much secrecy as to their rules was
that they suspected that every one versed in the new poetry and
music who inquired about their rules did so only to ridicule them,
and because the new learned versifiers really did view the master-
singers’ art as far beneath them.
Chapter VII. Reception of Popular Poetry. 7. Hans Sachs.
(Page 409). Hans Sachs (of Nuremberg, 1494-1576) was the
first one to feel vaguely that all poetry had sunk to a low level in
which it could not possibly persist. As of everything truly na¬
tional that we possess in medieval poetry, so also of this man, as
a purely German phenomenon, we must say : we must appreciate
him historically to establish his merit and determine his worth
accordingly. He occupies as it were a middle ground between the
old and the new art, and his works both point to older creations
of the nation and also lay the foundation for later ones yet to
come ; he embraces the poetic past of the people, and treats in many
ways all the forms and subjects that had become popular since the
rise of middle-class poetry; he seizes on all the actualities of his
time, and participates in the whole course of its religio-politieal
poetry; then he is the first one to withdraw from this, removes
poetry from its trend toward actual life and devotes himself to
the dramatic form, which since his time has remained the chief
form of all modern poetry. He draws all history and the full ex¬
tent of all knowledge and action into poetry, tears down the bound¬
aries of nationality, and thus gives warning of what was to be
henceforth the most characteristic feature of German literature.
He is in a certain sense a reformer of poetry as Luther is of re¬
ligion, as Hutten in politics; more fortunate than Hutten, less
fortunate than Luther, of far more unconscious talent than both
of them, like them indefatigably at work, little recognized in his
true value, indeed for a long time reviled as the representative of
mastersong, from which he strove to break away, for which he
composed only in private, in which he esteemed only morally what
from the viewpoint of poetry he considered unworthy of printing.
Only in recent times did Goethe, uncovering the poetic germs in
his forms and language, bring him again to notice and recognition,
Boedder — “Die Meistersinger von Number g.”
123
so that one may now name the venerable old master as one of the
chief figures of the epoch of the Reformation, so prolific in great
minds and characters.
(Page 411). As much as the lack of knowledge of human na¬
ture had impaired medieval court poetry, just as much did exces¬
sive association with men of a low class injure the middle class
poets. As much as the lack of important domestic affairs had left
the poetry of chivalry impoverished and lifeless, just as much did
the momentous domestic events of the epoch of the Reformation
blight contemporary poetry. The vortex of these events had un¬
balanced so many, the coarse tone of the progressive party of the
period had marred the language and everything most needful to
poetry. Hans Sachs’s life falls in the middle of this cultural
period, is involved in the fortunate and unfortunate phases of the
new doctrine, and his early youth coincides with the first move¬
ments. If like so many others he had let himself be carried along,
it would not be cause for wonder, considering his zeal and his
talent; if after winning his first applause he had joined in the
general chorus, he might plead the example of such great men!
What a deep nature does it indicate that this man could with such
minute and penetrating versatility follow the aspects of his times
and his nation, and fathom and depict, praise and blame them,
without faltering in his sobermindedness, without sinking down
from the height from which he viewed things. The entire range
of contemporary life, the vast movement of his time are unfolded
for us in the countless writings of the honest cobbler, animate and
eloquent, but not passionate; vivid and penetrating, but without
unrest, effortless and undesigning. He takes us into the plebeian
crowds, but one sees at once that he is one of those finer natures
who had forsworn the vulgar mob in the interest of social refine¬
ment. He shows to us the whole world in its whirling motion and
haste, imperturbed himself from his quiet cell in which nothing
escapes him, nothing leaves him indifferent, but also nothing robs
him of his equanimity. He examines the empire’s manifold de¬
fects, but he does not wish to reform them. Only one sees that he
is the inhabitant of a city which then enjoyed an enviable pros¬
perity of public and domestic finances, as well as of culture ; whose
good fortune has been lauded by every poet since Rosenblut’s time,
described by every writer from Aeneas Sylvius on, whose constitu¬
tion was the envy of every enlightened person, which not only gave
124 Wisconsin Academy of Sciences, Arts, and Letters.
birth to, and held fast, great talents, but knew how to attract for¬
eign talents, something that scarcely any republic has been capable
of at one and the same time; which was prominent in commerce
and industry, in mechanics and inventions, the center and citadel
of mastersong; which through more than a hundred years, from
Rosenblut and Folz down to Hans Sachs, remained the cradle of
the German drama; and which sheltered in its walls the greatest
men in all branches, Regiomontanus, Celtes, Yischer, Diirer, Pirk-
heimer, Hans Sachs ; which showed such a fecundity in artists and
scholars that no other German city, and indeed not many German
states, can display such a line of artists and scholars, which is sur¬
passed in part only by the great Italian republics. In this refuge,
full of incitement and without excitement, it was easier for him
to observe, easy to rise above and master what he had observed;
he surveyed from a distance and did not get confused by prox¬
imity.
(Page 413). As his poetic share in the pamphlet (Eyn wunder-
liche W eyssagwpig von dem babstumb, 1527) was very harmless, so
also his further writings for Protestantism were indeed forceful
and definite, but always moderate and calm and entirely devoid of
all excesses in form and content. . . . He combatted the coarse
tone of life and art, not by imitating this rudeness, like Murner,
but by trying to elevate his language and presentation, and to keep
above common reality.
How he did this shows what an innate poetic talent was his. And
that attracted Goethe so much to him (who himself knew how diffi¬
cult it is to remain superior to irruptive epochal events), to see
with what playful ease the honorable master treated the world
and life, how securely and placidly he roamed about, how the really
creative power of the poet wrought in him, not passion (414) and
personal interest and excitement; how his poetry is not a stupid
copy of life, but a free reproduction of it. It is true, we can in
his case speak only of a rough draft, not of perfect execution;
only of vigor and expressiveness, and of the great humorous power
of his language, which under Goethe ’s finishing touches ingratiated
itself so much with us, while in his own case the monotony and
heedlessness with which he throws off his rhymes annoys and de¬
ters us. It is true again that there is in his works a good deal of
idle chatter, of awkwardness in treatment, of unconcerned seizing
upon the first theme that offers, and later of soulless versified
Roedder—“Die Meistersinger von Nilrnberg
125
scribbling from mere force of habit. However, one may be chari¬
tably inclined even toward such artless versifying, where it is meant
for an artless type of people, unpretentious and amusing, if only
in its inner core it is entirely sound, serene, conciliating, and en¬
couraging. ... It is admirable to remain truly human in
great events of public life with so much interest and wholehearted¬
ness; more admirable than his attempt to make a completely van¬
ished poetry to bloom afresh and produce new seed for further
plantings. It was a time when so many unbidden meddled with
things that did not concern them, when so many lost or misjudged
their place in life. But how Hans Sachs, after the Muses had in
his twentieth year called him to his poetic work, quickened him
with their gifts, inspired him to praise virtue and to relieve sad¬
ness, and he, chained to his modest trade, had at first followed
their call with less inclination, how from that time on, even when
the loud applause of Germany was already honoring him, he kept
to his limits in the same even temper, with modesty and self-knowl¬
edge, and (415) ever remained the artisan-poet and the poet-arti¬
san, how in his life he kept up the same tone that his poems evince,
— all that is easier to observe than to comprehend. He might have
argued with Hutten which one of them knew human nature bet¬
ter, watched conditions in Germany more keenly, felt more ardently
for the fate of his fatherland and its culture and improvement,
but yet his poems on the conditions of the times in comparison to
Hutten ’s form a perfect contrast of placidity to nervous agitation,
of self-restraint to bold self-confidence, of moderation to portentous
passion, and, so far as poetic treatment is concerned, of superior
mastery of the subject to being mastered by the subject. . . .
He did not suffer himself to be carried away by the coarse tone in
the writings of his times; in the greatest passion and indignation
he does not use invectives such as Luther and even the rulers of
the period did; his style is vigorous and rich almost beside that
of any other contemporary; it is innocent, animated, and lumi¬
nous beside Murner’s, much more poetic, plastic, impressive, and
noble than Hutten ’s; full of health and pure humor in compari¬
son to Fisehart’s, and next to Luther’s his language is by far the
most remarkable of the century; it is a treasure-house for every
future humorist and satirist.
(Page 416). In the youthful products of his muse he con¬
centrates entirely on the question of chaste love, which usually
126 Wisconsin Academy of Sciences , Arts, and Letters.
brings to every moral man Ms first battle with life. He shows
himself as a man with plain middle-class sentiments, and praises
conjugal love above the adventurous Mnd, as every rightminded
man of his time considers it his duty to do. At an early period
(1517), in his Venus’s Retinue, he manifests how little he would
be skilled to interpret love and its nature in any other fashion.
Early and late, in his youthful poem on Chastity Exiled (in which
the stringent commands that he imposes on himself do honor to
his fine character), as well as in the theme of Tristan, which he
treats late in life, he gives the admonition to reserve love for matri¬
mony; and the sanctity of this union is also in his serious and
comic works the constant pivot about which his moralizing poetry
almost prefers to circle. In musing meditatively with himself, or
when his private genius listens at the window-shutters, in looking
into the heart of family life, or in escorting Ulysses to the abodes
of Calypso and Circe, he makes it an occasion to praise wedded life,
to scourge the common infidelity, to ridicule and curse the evil
domestic life in cities and villages.
(Page 417). As early as 1523 he wrote his Wittenberg Night¬
ingale, and saluted the new doctrine with so much determination
that nothing but a glance at this poem is needed to recognize his
attitude toward the Reformation and at the same time to see how
Luther’s teaching struck and roused long dormant thoughts of the
honorable middle classes of Germany, how the upright intellect of
this class, with the guiding testament in hand, now secured for it¬
self light in all directions, how the honest citizens hailed the day
with the singing nightingale, how they let themselves be called
back by it from devious wanderings, from desert and darkness
whither the crafty lion had lured them, how they withstood all the
yelping of his aiding fiends.
(Page 418). The attentive study of religious conditions in Ger¬
many led Hans Sachs of itself to the German Empire and its state,
especially at the time of the Smalkaldic War. Hence in the fifth
decade of his life the master’s poetry is particularly occupied with
it. He scourges what Hutten and with him every unselfish man of
the period scourged, but he does it in his own peculiar manner.
He remains true to the conviction, which Hutten abandoned, that
public spirit and concord alone would be Germany’s salvation.
. . . (421) Most of those allegorical and other poems that with
a satiric whip persecute the aberrations of the time as vices, while
Boedder — “Die Meister singer von Niimberg.”
127
later, more mild and forbearing, lie but laughs at them, arise in
this sturdy period, during which he was also more active in public
life (fourth and fifth decades). The felicitous and unfaltering
observation of the world and ol men, which was natural to our
master’s genius, found plentiful food in the tendency of the sages
of antiquity to consider man’s inner nature; and their judicious
moderation fortified him in the placid calmness with which he views
unconfused the antlike medley of human beings, and confronts the
people with the mirror of his truthful paintings; their contempla¬
tive wisdom fostered his plastic sense. . . . (422) From ancient
history he emphasized for his contemporaries that which in school
we present in the same way to the minds of children, and in the
most direct manner he conducted the purest water of the spring he
had found even into the lowest classes of people. What two or
three centuries had already done would have been wholly lost if
at this time of the first printed books, when the common people
were really eager to learn and read, a man who had mastered the
tone of the common folk had not taken the whole mass of what
Thomasin, the Renner, and all the didactic poems and collections
of moral examples had long been spreading broadcast, and modern¬
ized it in a new language and suitable diction. Let us never fail
to give Hans Sachs credit for this service. He became a humanist
teacher of the common people, as the scholars became teachers of
the young. He was the first to popularize the ancients from their
purely moral side in our country, just as in recent times Wieland
introduced his Cicero, Lucian, and Horace from the side of the
philosopher of life and man of the world.
From his sixth decade on another taste begins to predominate in
Hans Sachs’s poetry. He centers his attention more on rhymed
tales and carnival farces, he loves to attach the didactic to exam¬
ples, the ethical character of his poems becomes more plastic, his
German style of painting becomes more Dutch, his allegory is ex¬
changed for the fable, direct references to the present become
rarer, and he takes us from public into private life. He then sees
the classes and ranks of men less in their relation to their civic and
social obligations than to human nature and the rational world in
general ; he pictures the droll doings of men more humorously, and
laughs at them instead of scourging them as heretofore. His im¬
pressively severe teaching rather disappears beside his goodhu-
mored description, his censure becomes ironic delineation; his po¬
etry, which formerly was more bent on enjoining virtue, is now
128 Wisconsin Academy of Sciences , Arts , and Letters.
more directed toward the alleviation of sadness; the severity of
manhood wears off giving way to the benignity of old age. At all
times of his life the master wrote rhymed stories and tales, from
the fifties on, however, they become both more frequent and better.
The whole monotony and mechanical fertility of his (423) produc¬
tion shows itself in this species, but it is at the same time his high¬
est triumph. He had inherited this type of instructive tale, seri¬
ous or comic, from older periods, whose novelettes, folk tales, and
farces he modernized in countless numbers and multiplied by new
ones, but he also left them to posterity. No older story teller is
his peer in moral earnestness, few later ones in skill of treatment
and genuine humor. . . . The animation and accuracy of de¬
scription, the motley throng of subjects, and the uniform precision
and reliability of his brush impart an uncommon charm to these
pieces, and it was these qualities that attracted our Goethe, who in
Hans Sachs’s Poetic Mission erected the most reverent monument
to the old master. The figures move and stir before our eyes, and
if Hans Sachs praises the painter because he can so place every¬
thing before our eyes that one could not narrate it more clearly,
he himself narrates and describes everything so that one could not
paint it more plainly. The most playful and frolicsome humor
tints the pictures of the magic chest that he throws open to us,
. . . (424) and all this is to be compared only with the bur¬
lesques of Dutch painting. . . . Everything that characterizes
the good German middle class, craftsmanship, self-respecting trade-
unionism, common sense, honesty (425) and integrity, pious sim¬
plicity, a sound moral core, and practical insight into life, speaks
amiably from every tone and every line of these pieces, poor as
many of them are in content and stale jests.
In the last decades of Hans Sachs’s poetry, a distinct change
takes place. He himself complains repeatedly of the decline of art
in general. Formerly, he says, it flourished, all nooks and corners
were full of scholars, there were ingenious craftsmen and artists
enough and books in abundance, now the arts are common and
despised, few devotees remain, looked at ascance as visionaries;
the world is madly bent on lust and gold, the Muses are leaving
the country. His eulogy of good morals had brought him envy and
hatred; often he meant to stop writing, also because at last his
reason told him that his poetic power was on the wane. But yet
he means courageously to make the most of his talent; after forty-
four years of poetic activity he will not now desist from preaching
Roedder — “Die Meistersinger von Nurriberg.”
129
virtue and relieving sadness, and no defamer shall disturb him in
his sacred calling. . . . (426) The present furnished no more
themes for poetic treatment ; folk poetry, and the occasional poem
ran out of material, in addition the writers were weary of the
coarse tone, withdrew from the realities of life and sought after
another, nobler element for poetry. Before ancient art or its
Italian and Spanish imitations found reception, one pointed back
once more to the old romantic period and reproduced it in old and
new forms. A foolish idea. One might now, like Puschmann,
gather in prose form the rules of the art of yore, but it was no
longer to be kept alive; the Heldenbuch theorized on giants and
dwarfs, heroes and men. . . . (427) Nevertheless it remains
remarkable enough that Hans Sachs, just as in his treatment he had
always striven away from the vulgar and decadent manner of his
contemporaries, aspired toward the end of his life in the choice of
his subjects after something nobler, although in the absolutely ig¬
noble treatment of these heroic themes he showed how little the
period was suited to a revival of these things.
9— S. A. L.
g
THE PASSING OF AN HISTORIC WATERWAY
BY P. E. WILLIAMS.
The early development of any country depends primarily upon
the avenues of entrance to its interior. The opening of the in¬
terior of the United States was largely balked by the eastern moun¬
tains, which served to divert the entrance to northern routes. It
was natural that waterways should be sought, because of the greater
ease of travel in the canoe than on foot. With the practically
continuous water route of the St. Lawrence, Ottawa and the Great
Lakes, at their disposal, explorers soon found their way into Wis¬
consin. As early as 1634, Nicolet, following the north shore of
Lakes Huron and Michigan to Green Bay, thence the Lower Fox,
Lake Winnebago, and the Upper Fox reached a point in the vicinity
of Berlin, Wisconsin.1 In 1655 Badisson and Groseilliers2 crossed
from Lake Michigan to the Mississippi River following the Fox-
Wisconsin waterway which was for many years one of the most
used highways in the northwest. Probably some of the coureurs de
hois and voyageurs had earlier passed this way as we know it was
a path much used by the Indians. It soon became a regular route
for the French fur traders and Green Bay became one of the most
noted fur depots of the northwest. Along it came the great mot¬
ley procession of adventurers, explorers, priests, traders, trappers,
and soldiers who paddled their canoes and rowed or poled their
batteaux or, in later years, their Durham boats. “Throughout
127 years of French control, down through the 25 year period of
English control, and on to the days of the American railroad in the
’60’s, this nature-made highway of travel between the Great Lakes
and the Mississippi was a link in a great continental waterway — a
part of a Pathway of Empire.”*
1 Thwaites, Reuben Gold, Wisconsin, The Americanization of a French Settle¬
ment, New York 1908, pp. 20-32.
a Ibid, p. 40.
s Whitbeck, Ray Hughes, The Geography of the Fox-Winnebago Valley, BulL
XLII, Wis. Geol. & Nat. Hist. Survey, Madison, 1915. p. 27.
131
132 Wisconsin Academy of Sciences , Arts , and Letters .
With the knowledge of the success attained by the completion of
the Erie Canal in 1825 came an agitation for the construction of
canals in several parts of the country, especially in those states
which touch the Great Lakes. The introduction and improvement
of the steamboats was a further incentive to waterway improvement
as many of the streams used by the pioneer were too shallow or re¬
quired portages, both disadvantages in their use for profitable
steamboating. Such a highway was the Fox-Wisconsin. In 1829,
just four years after the completion of the Erie canal, an agitation
was begun for the improvement of the Fox-Wisconsin highway in
order that steamboats might be used.4 This agitation came not
only from the merchants and the usual boomers of real estate in
the small towns along the way, from the farmers and boatmen, but
also from representatives in Congress who took an active part in
bringing the proposed improvement to the attention of the govern¬
ment and the public. Great was the optimism in regard to the ad¬
vantages to be gained by those along the highway and to the whole
northwest. Cities situated along this water course were sure to in¬
crease rapidly in size and become the most important in the state.
Along this valley would move the pioneers to the region farther
west. In their wake would follow supplies and implements, and in
return would go the furs, the lead of southwestern Wisconsin, and
the products of the constantly increasing number of farms.
Since the Fox-Wisconsin route had been the one most travelled
between the Great Lakes and the Mississippi River in the early ex¬
plorations, in the fur trade, and subsequent communication, it was
but natural to believe that it would be the one most easily converted
into a waterway for boats of deeper draught and heavier cargo.
There was a great difference of opinion as to the navigability of
this waterway, depending probably on the season of the year in
which the several trips were made and consequent differences in
volume of water. All accounts agree in a general way as regards
the conditions on the Fox River, and when later surveys were made
the engineers agreed as to the manner of improvement and quite
closely as to the estimated cost. The Wisconsin, on the other hand,
had so many changes in volume, channels, and general behavior
that even able surveyors and engineers arrived at different con¬
clusions. Marquette says in his journal that “It is very broad,
with a sandy bottom, forming many shallows, which render navi-
Whitbeck, p. 29.
Williams — The Passing of An Historic Waterway. 133
gation very difficult.”* 5 Another writer says “that the general
depth of the river is, at the ordinary height of the water, from four
to five feet, but the sand-bars often extend entirely across the river
and have not more than eight or ten inches of water; the sands,
however, are quick and oppose but little resistance.”6 The other
extreme, high water stage, is shown by the fact that in 1828 the
fifth regiment of U. S. Infantry came in barges from St. Louis up
the Mississippi and Wisconsin Rivers and down the Fox to Fort
Howard without unloading.7
As time went on and the success of other canals became apparent,
the agitation for improvement increased. The government early
inaugurated a movement to improve the Fox River for navigation
purposes. In 1836 the war department had A. J. Center make a
survey from Fort Howard, at Green Bay, to Tail Point, a distance
of six miles.8 In 1837 a hurried survey was made of a consider
able portion of the river.9 In 1838 the improvement of the route
was recommended by the Secretary of War for the purpose of facil¬
itating troop and munition transportation.10 In 1839 a preliminary
survey of the Fox River was made by Captain Cram under the di¬
rection of the Secretary of War.11 In 1846, Morgan L. Martin,
territorial delegate from Green Bay, introduced a bill in Congress
for the improvement of the Fox and Wisconsin rivers. This bill
which was favored by President Polk, passed in that year, and con¬
tained a provision that a large amount of public land in the Terri¬
tory of Wisconsin be granted by Congress and the funds from the
sale of this land to be used for the purpose of making a water way
navigable from the mouth of the Wisconsin River to Green Bay, as
soon as Wisconsin became a state.12 Land equal to one-half of three
sections in width on each side of the Fox River and of the lakes
through which it passes was to be sold, the proceeds to pay for the
6 Thwaites, Reuben Gold, Down Historic Waterways , 2nd edition Chicago 1907,
p. 239, quoting from Marquette’s Journal.
6 Warren, Major G. K., Report on the Transportation Route along the Wis¬
consin and Fox Rivers , Executive Document. No. 28, 44th Cong., 1st session.
Government Printing Office, 1876.
7 Durrie, Daniel S., Early Outposts of Wisconsin, Green Bay for ZOO years ;
1639-1839. A paper read before the Wisconsin Historical Society, 1872, p. 10.
8 Warren, p. 25.
9 Ibid, p. 25.
10 Art Publishing Co. (no author) The Valley of the Lower Fox: Historical,
Descriptive, Picturesque , 1887.
11 Ibid.
12 McLenegan, Annie Susan, Pioneer Life in the Fox River Valley, Proc. Wis.
Hist. Soc. 1905, p. 279.
134 Wisconsin Academy of Sciences, Arts, and Letters.
improvement. The work was to begin within three years after Wis¬
consin’s admission as a state and was to be completed within twenty
years. An act of the Wisconsin legislature in 1848 provided that
5/6 of the sales should go to improve the Fox River and 1/6 of
the sales to improve the Wiseonsin.lsThis act shows how little and
how carelessly the facts about the Wisconsin River had been con¬
sidered. The legislature evidently believed that the Wisconsin
River with slight improvement, could easily be made navigable. In
1849, or about twenty years after the agitation began, the state com¬
menced work upon the improvement. By 1850 the lock at De Pere
rapids was completed. The canal connecting the Fox and Wis¬
consin Rivers was dug and a little dredging of shallower portions
of the river was accomplished.14 Land sales were much slower than
had been anticipated, money ran out, interest was accruing on the
first debt, and work was nearly stopped. An issue of state bonds
was proposed but this was held unconstitutional by the state legis¬
lature. So after spending over $400,000, the state resolved to sur¬
render the whole improvement, the remainder of the public lands
unsold, and the hydraulic privileges to a private company. With
a number of provisions regarding toll and free use by the United
States, the whole project was turned over to the “Fox and Wis¬
consin Improvement Company.’’15 In 1855 Congress passed an
act giving to the state an additional two sections of land making
in all five sections per mile for the whole length of the Fox River
and of the lakes through which it runs, a distance of about 216
miles.16 By 1856 the Lower Fox could be traversed by vessels draw¬
ing three feet of water and the Upper Fox by boats of slightly less
draught,17 Much rejoicing was caused by the arrival at Green Bay
of the Aquila, a small steamer that had come from Pittsburgh down
the Ohio and up the Mississippi and Wisconsin rivers and down the
Fox. A public celebration was held and many believed that the
long dreamed of waterway which would be used as one of the na¬
tion’s most important highways had come into its own.18 In 1856
the Wisconsin legislature passed another act providing that the
capacity of the Lower Fox be increased to a draught of 4 feet ; the
13 Warren, p. 27.
14 Whitbeck, p. 80.
15 Whitbeck, p. 80.
16 Warren, p. 39.
17 Whitbeck, p. 31.
18 McLenegan, p. 285.
Williams — The Passing of An Historic Waterway. 135
Upper Fox to 3 y2 feet; and providing for locks 160 feet long, by 35
feet wide. The work was begun and carried on vigorously by the
company until the panic of 1857. From this time on the com¬
pany encountered many difficulties and in the summer of 1866, be¬
cause it had failed to fully perform its agreement with the state,
the trustees sold the improvements, lands, and franchises at public
sale. The purchasing company was the “ Green Bay and Missis¬
sippi Canal Company.”19 A short time before this the national
government took a renewed interest in the waterway and during
the third session of the 37th Congress a resolution was passed
that a “committee on naval affairs be appointed to inquire into
and report upon the practicability and the probable cost and
time required to improve the Wisconsin and Fox Rivers, so as to
give an uninterrupted navigation from the Mississippi River to
Lake Michigan for vessels of war 200 feet in length, 34 feet beam,
and drawing not less than 6 feet of water. ’ J2° The committee which
reported in 1863 estimated the cost at $2,387,000 and suggested
that the Lower Fox be improved to a 12 foot draught with the
further suggestion that Lake Winnebago might become a naval
station provided this could be done under the “treaty of 1817.”
The full committee decided against the naval station proposal,
however, and closed the report with the following: “The true
ground as the committee think, upon which to place the propriety
of yielding assistance to this Wisconsin enterprise, is its great
natural importance in making cheaper and easier the intercourse
between the grain-regions of the Northwest and the manufacturing
and commercial states of the East. The expenditure of twenty
millions in the completion of this work and that of Illinois with a
corresponding enlargement of the means of conveyance in the East,
would be many times repaid in the increased general prosperity
which would result from it. Whenever some systematic and well-
matured plan shall be laid before Congress, which shall compass
this result, it is to be hoped that it may be adopted.”21
In 1870 the waterway was taken over by the United States Gov¬
ernment, but it did not take over the land grants or water power
franchises of the Green Bay and Mississippi Canal Company. This
company is still in existence and maintains the right of control over
19 Warren, p. 47.
^Warren, p. 46.
21 Ibid, p. 47.
136 Wisconsin Academy of Sciences , Arts, and Letters.
all the surplus water beyond the needs for navigation purposes.22
Agitation continued for years in the attempt to induce Congress to
complete the work already begun. The governors of four states
called a convention which was held at Prairie du Chien in 1868. The
delegates sent a memorial to Congress declaring that the ‘ ‘ immedi¬
ate opening of said channel is demanded by the interests of the
people of the entire country ; the work is one of national import¬
ance, required as a channel of commerce, as a ligament to bind to¬
gether the states, in securing national unity and as a defense in
case of war.”23 Between 1866 and 1875 a most thorough investi¬
gation of the problem was made by Major G. K. Warren for the
government. His report, issued in 1876, is very full and con-,
elusive. It contains a short outline of recommendations, made be¬
fore, and the work accomplished up to that time, as well as the
results of his own work and his conclusions. Major Warren spent
much of his time in a detailed study of the Wisconsin River for
he realized that the work previously done was more superficial here
than along the Fox River. It is true that other engineers working
either for the state, the two private companies, or for the govern¬
ment had made preliminary or reconnaisance surveys. Commis¬
sioners Croswell and Richardson as well as Mr. J enne believed that
the Wisconsin River would be improved by the construction of
wing-dams located so as to contract the shallow portions of the chan¬
nel. Seven of these dams had been built in 1852, but most of them
soon disappeared, — “A fate which attends all wing-dams not con¬
stantly cared for.”24 It was also proposed to keep boats running
at points where bars tended to form and thus keep the sand moving.
Earlier authorities believed that all that was necessary to keep a
three foot channel was to remove the snags and overhanging trees.25
The chief reason for these differences of opinion was the inaccurate
data of slope and current available to the early engineers. Major
Warren concluded that the use of the Wisconsin, especially for a six
foot channel as desired, was impossible because of the constant for¬
mation of sand bars. There are two particularly important require¬
ments for a rectified river. It must be narrow enough to give a
required depth at low water, and broad enough to carry off the
excess water in times of floods. Furthermore, such arrangements
22 Whitbeck, p. 32.
23 Whitbeck, p. 33.
24 Warren, pp. 32, 34.
25 Ibid, p. 30.
Williams — The Passing of An Historic Waterway. 137
must be made as to insure the return of the main stream to the nar¬
row channel.26 Inasmuch as the Wisconsin was broad and shallow
at low water, and as the wing-dams designed to narrow it were
either quickly buried or destroyed, Major Warren advocated the
building of a canal along the Wisconsin River, using the river itself
for short distances. This would necessitate locking in and out of
the river. He estimated the cost of the improvement of the Wis¬
consin alone at $4,000,000 and an additional $50,000 yearly for
maintenance.27 This large initial cost in conjunction with the de¬
crease in the use of canals in general caused the improvement of the
Wisconsin River to be shelved. The government, however, did en¬
large and improve the locks on the Fox river, and now maintains
navigation about 8 months of the year without toll charge.
Up to this time about $4,000,000 has been expended on the
waterway of which only $590,000 has been spent on the Wisconsin
River. The average yearly cost of operation and care has been
about $56,000, and since 1902 the yearly expense for operation has,
with few exceptions, exceeded this average.
After the opening of the canal, the major part of the traffic was
on the Lower Fox from Green Bay to and on Lake Winnebago.
Few records are found of boats of any size having made the con¬
tinuous passage from Green Bay to the Mississippi River. Men¬
tion has been made of the small steamer Aquila which made the
trip in 1856 at high water. In the early 50 ’s, there were regular
steamer routes for short distances along the Fox and six steamers
were running between Green Bay and Kaukauna, one from Fond du
Lac to Oshkosh and then up the Fox and Wolf Rivers to New Lon¬
don. During the sixties and the seventies, the decades in which
the waterway was of most importance, steamboat building was ac¬
tive on the Fox and Wolf rivers. Over 60 boats were built between
1844 and the present time, about 50 of which were constructed be¬
fore 1880. With the exception of coal barges and pleasure boats
however the Fox River is of little use at this time. In recent years
there have been approximately 800,000 tons of coal carried annu¬
ally, principally on the Lower Fox and Lake Winnebago.28
No regular lines of steamboats now run on the Lower Fox but two
or three small steamers or power boats make more or less regular
2« Warren, p. 92.
27 Ibid, pp. 106, 114.
28 Whitbeck, p. 35.
138 Wisconsin Academy of Sciences , Arts, and Letters.
trips on the Upper Fox and Wolf Rivers. The number of lockages
has increased since 1900 but this is due for the most part to pleasure
boats, especially small gasoline launches. Although the Fox River
has not measured up to anticipations as a highway for steamboats,
it was important in opening up that part of the state more rapidly
than the surrounding regions. The water power developed at the
dams, which were necessary for locks, has been of great advantage
in building up there a concentrated group of industrial cities not
equaled anywhere else in Wisconsin except along the shores of Lake
Michigan. However many of the plants in this section, and four
of the most modern, are now using steam power only.
Steamboat navigation on the Wisconsin has always been almost
negligible. In the 50 ’s following the improvement of the: Fox River,
two steamers were partially successful in the attempt to make regu¬
lar trips to carry lead from Galena to Fort Winnebago (near Port¬
age).29 In 1859 a steamer made regular trips on the Wisconsin
River between Sauk City and Portage.30 Many are the records of
failures in the attempt to navigate the Wisconsin except at high
water stages. In 1868 the Wisconsin, a side wheeler drawing only
two feet, made the attempt and met with little difficulty in the
Upper Fox but found it almost impossible to get down the Wiscon¬
sin at all. Later that same year a dredge boat tried to get up the
Wisconsin from the Mississippi but was unable to get into the Wis¬
consin. The boat drew 32 inches but only 24 inches of water could
be found on the bars of the Wisconsin for six miles up from its
mouth. In 1869 some work was done on the river between Portage
and Sauk City and two small side wheeled steamers were enabled
to make trips but with difficulty.31 In 1887 the bridge tender at
Helena on the Wisconsin River said that it had been four years
since he had swung the draw for a river craft. He told of an at¬
tempt of a small steamboat to make the passage on what was then
considered a good stage of water from Portage to Lone Rock. She
had been two weeks making the 21 miles from Arena to Lone Rock
and finally was abandoned on a sand bar.32 These are but a few
of the many attempts that might be cited. In 1916 I asked the
superintendent of the power plant at Prairie du Sac how many
times he had opened the gates of the dam during the year. He
29 Whitbeck, p. 32.
80 Warren, p. 42.
31 Ibid, p. 59.
82 Thwaites, Down Historic Waterways, p. 262.
Williams - — The Passing of An Historic Waterway. 139
answered: “Three; twice for small launches from Sauk whose
occupants were going up into the lake to hunt for ducks and once
for two new houseboats that were made in Oshkosh for use on the
Mississippi River.”33
Although the causes for the decline of this water way are many,
a few may be mentioned: 1. The shallow depth of most of the
channel. This applies to the Upper Fox and Wisconsin Rivers.
2. The crookedness of the Upper Fox. The devious, winding,
obscure course of this part of the river discouraged and perplexed
even the early canoeist.
3. The closing of the waterway by ice from three to five months
each year. This required the development of some other means of
transportation which could he used the whole year unless there is a
very great advantage in water transportation during the summer
months.
4. The direction or trend of the waterway is against it for
present day commerce. It extends northeast — southwest while the
principal trend of trade for all territory north and west of Chicago
is northwest — southeast. During the present difficulties of railway
transportation it has been proposed that a line of barges be estab¬
lished on the Fox River. This has met with but small encourage¬
ment, largely because of the reason just given.
5. The slowness of the waterway — its use for passengers especi¬
ally was possible only before other means of transportation were
developed.
6. The extremely poor natural facilities of the Wisconsin River
for navigation. This has been touched upon several times in this
article but deserves emphasis. Few writers point out the detri¬
mental effects which the Wisconsin River had on the attempt at
Fox-Wisconsin improvement. True, there have been other influ¬
ences that have been more potent in taking away the trade of this
highway, but if the Wisconsin River had been navigable in the
’50 ’s and ’60 ’s when the agitation and interest were highest, a
traffic would have been established which would have continued
for some time in spite of outside competition. It is a long step
from one of the most perfect natural canoe routes to an even
passable steamboat channel.
7. The chief cause of decline was the competition of other means
of transportation, to a small extent roads, but principally rail-
33 Greene, Kenneth, Personal Communication.
140 Wisconsin Academy of Sciences , Arts , and Letters.
roads. In the early ?50’s stage lines and other plank roads were
developed connecting tl principal places and diverted considerable
of the river passenger traffic.34 In 1857, the Milwaukee and
Prairie du Chien Railroad, crossing the Wisconsin River three times,
was completed to the Mississippi River. In 1862 the Chicago and
North Western Railroad reached Green Bay,35 so before the wa¬
terway had reached its highest stage of improvement, powerful
competitors entered the field and drew the trade into new channels
before the route had been fully tried out. The railroads were able
to accomplish this in spite of the fact that for the greater part of
the route, or from Spring Green to Oshkosh, no railway parallels
the waterway. In 1887, R. G. Thwaites made an excursion through
the waterway and has painted a vivid word picture of fallen
grandeur and prosperity. He says, 4 ‘The canal, like the most of
the Fox-Wisconsin improvement, is fast relapsing into a costly
relic. The timbered sides are shaky and worm-eaten, and several
moss covered barges and a stranded old ruin of a steamboat turned
out to grass tell a sad story of official abandonment.”36
Today the dream of a commercial waterway is past, but the
scheme has added a very pretty pleasure route to the beauties of the
state. There can be no doubt that this project, though a failure
in itself, was of great advantage to the state, in that it attracted
attention to the importance of the region. The route did serve as
an avenue of early access to the settlers and the agitation and partial
improvement hastened the coming of statehood. To the villages and
cities along the route, the failure of the Fox-Wisconsin to become
a great commercial highway was a bitter disappointment, but the
results of such delusions must inevitably be shared by all those
whose hopes for commercial greatness are tied to a small inland
waterway.
34 Warren, p. 40.
35 Whitbeck, p. 32.
36 Thwaites, Down Historic Waterways, p. 144.
THE ARSENICAL SOLUTIONS
No. 1. Liquor Potassii Arsenitis
(Fowler’s Solution)
H. A. LAN GENH AN.
INTRODUCTORY STATEMENT
The history of arsenic therapy is of considerable interest in the
evolution of the materia pharmaceutica. Natural compounds of
arsenic, such as realgar and auripigment were used during anti¬
quity,1 but primarily for external purposes. Because of the change
which they produced in the color of copper, arsenical compounds
have played an important role during the alchemistic period of
chemical history.2 As a result, new derivatives of arsenic were
discovered,3 some of which at least were destined to play a role in
the battles between the iatrochemists and the representatives of the
old or Galenical School of Medicine.4 A larger increase to the
number of arsenic compounds, however, did not come until the
close of the eighteenth and the beginning of the nineteenth cen-
1 For a concise statement of the knowledge of the ancients, see Kopp,
Geschichte der Chemie, vol. 4, p. 89.
Berendes, Des Pedanios Dioskurides Arzneimittellehre, p. 531 (Chap. 120 :
Arsenikon ; and Chap. 121: Sandaracha.)
Pliny, Natural History (Bohn’s Classical Library, vol. 6, p. 104 (Orpiment)
and p. 220 (Sandarach and Arrhenicum).
Scribonius Largus (Felix Rinne, Das Receptbuch des Scribonius Largus, vol.
5 of R. Robert, Historische Studien aus dem Pharmakologischen Institute der k.
Universitat Dorpat, p. 79 ( Auripigmentum) and p. 84 (Sandaraca.)
For a more detailed account see, “Historical Notes on Arsenic and its prin¬
cipal combinations: by G. F. Babcock, New Remedies, vol. 12, (1883) pp. 98,
131 and 170.
2 Comp. E. v. Meyer, History of Chemistry (3rd ed.) p. 58: “On account
of this behaviour, the pseudo-Geber calls arsenic Medicina Venerem dealbans”.
See also Kopp, Geschichte der Chemie, vol. 4, p. 94.
3 The arsenicum album (As2 03) of the alchemists; the arsenicum fixum of
Paracelsus (arsenic acid or its potassium salt). Kopp, Geschichte der Chemie,
vol. 4, p. 95.
4 Meyer, History of Chemistry (2d ed.) p. 94.
141
142 Wisconsin Academy of Sciences, Arts, and Letters.
tury.5 Still later followed the organic derivatives of arsenic that
formed the basis of the classical researches by Bunsen.6 7 Com¬
paratively few of these have found application in medicine, one of
the latest being the much discussed specific “606” or salvarsam.1
If the chemotherapy of arsenic includes but few of its com¬
pounds, pharmacopoeial preparations of the derivatives of this ele¬
ment are even less numerous. Those of the U. S. P. and N. F. are
but seven in number and can be grouped as follows :
1. ) Solutions of arsenous acid and its salts:
1. ) Liquor Acidi Arsenosi, U. S. P.
2. ) Liquor Potassii Arsenitis , U. S. P.
2. ) Solutions of arsenates:
3. ) Liquor Sodii Arsenatis , U. S. P.
4. ) Liquor Sodii Arsenatis, Pearson, N. F.
5. ) Liquor Arsenicalis, Clemen’s, N. F.
3. ) Solutions of arsenic trihalides (or rather “double salts”) :
6. ) Liquor Arseni et Hydrargyri lodidi, U. S. P.
7. ) Liquor Auri et Arseni Bromidi, N. F.
I. Solutions of Arsenous Acid and Its Salts.
The socalled “arsenious acid” of the older chemists, our arsenous
acid anhydride, or arsenic trioxide, is but sparingly soluble in wa¬
ter, yet sufficiently soluble to form solutions with a 1 p. c. As203
content. Nevertheless such a solution is not official in any of the
latest editions of the pharmacopoeias.8 9 Neither does the simple
solution appear to be prescribed at present by physicians, although
arsenic trioxide is frequently prescribed in solid form as pill,®
°Kopp Geschichte der Chemie, 4, p. 96.
6 The reprint of these has been edited by A. v. Baeyer in Ostwald’s Klassiker
der exakten Wissenschaften. vol. 27.
7 This epoch making discovery is discussed by Schweizer “Ehrlich’s Chemo¬
therapy— a new science” in Science, vol. 32, p. 809. (Dec. 9, 1910.)
8 In the French Codex of 1884 a Solute Arsenical ou Mineral (Boudin), one-
tenth as strong as the corresponding U. S. P. arsenical solution, was official,
but has been dropped in the latest edition. See also Dorvault, L’Offfcine, 15th
edition (1910), p. 1289, which also gives a formula for a Solute Arsenical (Is-
nard), which is only one-tenth of the strength of Boudin’s solution.
9 Arsenic trioxide enters the Pilulae Ferri, Quininae, Strychninae et Arseni
fortiores of the National Formulary 4th revision ; also Pilulae F. Q. S. et As.
mites , N. F. IV with less arsenic trioxide.
The French Codex of 1866 and 1884, but not the latest edition, contains.
Granules d’ Acide Arsenieux ; also Pilules Arsenicales (Pilules Asiatiques) ; see
also, L’Officine 15th edition (1910), p. 1061; Binz, Lectures on Pharmacology,
vol. 2, (1897), p. 88, translation by Latham.
Langenhan — Tine Arsenical Solutions.
143
powder,10 or tablet. The slow solubility of arsenic trioxide in wa¬
ter may be accelerated either by hydrochoric acid or by potassium
carbonate. The question may well arise, are these adjuncts mere
pharmaceutical conveniences or do the resulting reaction mixtures
differ sufficiently chemically to make the products different thera¬
peutic agents'? If essentially alike as therapeutic agents, another
pharmaceutical question may be asked, viz., are the two solutions
required in medical practice to avoid chemical incompatibilities
when prescribed with other substances? The first question is an¬
swered in the negative by modern pharmacologists.* 11 The socalled
neutral solution being unstable,12 the acid and alkaline solutions
afford pharmaceutical advantages only. Both are official so that
incompatibilities may be avoided by using either the one or the
other.13
In a general way, the chemistry of these two solutions has cer¬
tain aspects in common, hence may be considered together if for
no other reasons than to avoid unnecessary repetition. The reac¬
tions involved may be reviewed from the following points of view :
A. ) The hydration of arsenic trioxide.
B. ) The action of an alkali, such as potassium hydroxide or
potassium carbonate, on the hydration products.
C. ) The action of hydrogen chloride on the hydration products.
A. Hydration of arsenic trioxide.
When arsenic trioxide, arsenous acid anhydride, is dissolved in
water, its hydration to metarsenous and other arsenous acids may
be assumed. This hydration finds a ready expression in the fol¬
lowing structural formulas of these compounds :14
As^°
\ II
O + OH
AS\\
O
— - —
Arsenous acid
anhydride.
10 In the French Codex of 1866 there was official a Poudre d’ Acide Arsenieux.
11 See letters by Bernard Fantus, A. S. Loevenhart, and Torald Sollmann.
“See letter by Dr. Withering, in Fowler’s Report, p. 124.
m See letters by Dr. Fantus and Dr. Sollmann.
14 'Similar formulas are used by A. Stavenhagen in his article on “Beitraege
zur Kentniss der Arsenite”. Joum. pr. Chem. (1895) 159, p, 1.
As— OH
As— OH
Metarsenous acid.
144 Wisconsin Academy of Sciences, Arts , and Letters.
H
As + OH
\()H
Metarsenous acid
/OH
As— OH
\OH
Ortho-arsenous acid.
For a better understanding of the chemical possibilities of prod¬
ucts obtained upon the evaporation of the solution of the arsenites,
the following condensation products of ortho-arsenous acid with
itself may be referred to.
/OH
As— OH
\OH
— h2o
/OH
As-OH
\()H
Ortho-arsenous acid
w? r
/OH
As— OH
^O
As— OH
\OH
Di-arsenous acid (tetra basic)
Still another dehydration reaction may be indicated by the fol¬
lowing formulas :
HO
OH
As
O
-HsO
HO — As
OH
OH
As
As
OH
O
Uiarsenous acid (dibasic)
A trioxide of arsenic resulting upon complete dehydration of
this dibasic diarsenous acid which, however, is different structur¬
ally, hence in its properties, from the compound with which we
started out, readily follows:
O
.As\
OH '
OH
'As^'
O
H20
O
As
<To
As
Diarsenous acid (dibasic)
Diarsenous acid anhydride.
The possibilities of other hydration products of arsenic trioxide
need not here be considered.
15 Salts of this acid have been prepared by Stavenhagen, 1. c.
Langenhan — The Arsenical Solutions.
145
B. Action of Alkalies on Arsenous Acid.
Leaving out of consideration the diarsenous acid,16 as well as
other possible polyarsenous acids, it is quite apparent
1.) That metarsenous acid is monobasic and with metals may be
supposed to form the simple metarsenites :
As
\0— IT
Metarsenous acid
As
\(J_M
Metarsenites
2.) That orthoarsenous acid is tribasic and may be expected to
form three types of orthoarsenites, viz. :
/OH
As— OH
\OH
/OM'
As— OH
\OH
/OM'
As— OM'
\OH
/OM'
As— OM'
NOM'
o — Arsenous acid Mono-metal
dihydrogen
orthoarsenite
Di-metal Tri-metal
monohydrogen orthoarsenite
orthoarsenite
This leaves out of consideration, for the time at least, the possi¬
bility of the re-arrangement of the atoms of the orthoarsenous acid
molecule made possible by a change of valence, such as that of the
phosphorus, from three to five and the salts of the new ic acid.
As— H
\OH
\()H
ic Isomer of o — arsenous acid
Which of the above salts are formed depends, in part at least,
on the ratio of acid and alkali. Presumably a mixture, not a single
compound results. As to which may dominate in the equilibrium
resulting from a given formula for preparation need not be con¬
sidered here, but will receive such attention as seems desirable,
under the study of the pharmaeopoeial formulas in use.
18 Pyroarsenites have also been prepared by Stavenhagen, 1. c.
/OH
As— OH
\OH
o— Arsenous acid
10— S. A. L.
146 Wisconsin Academy of Sciences, Arts, and Letters.
C. Action of Hydrogen Chloride on Arsenous Acid.
For the present purposes the diarsenous acids and other more
or less complicated possibilities may be disregarded. Taking into
consideration only the simple meta and orthoarsenous acids, the
possibilities are indicated by the following structural formulas :
/OH /OH /OH ^C1
As— OH As— OH As— Cl As— Cl
\OH \C1 \C1 \C1
^O 5^0
As — OH As— Cl
Again it becomes apparent that when arsenic trioxide is dis¬
solved in water with the aid of hydrogen chloride, a number of
products, rather than a single one, are likely to result and that the
equilibrium, that finds its expression in a given pharmacopoeial
preparation, will depend on a number of factors.
LIQUOR POTASSII ARSENITIS (FOWLER’S SOLUTION).
History : — The arsenicum album17 of the alcemists, the arsenous
acid of the antiphlogistic nomenclature, our arsenic trioxide, ap¬
pears first to be mentioned in the Latin alchemistic manuscripts
formerly attributed to Geber.18 Under the ban of the phlogistic
theory, it was regarded as the calx of the regulinic arsenic, a half
metal (Brandt, 1733). 19 This conception paved the way to the
recognition of the “white arsenic” as an oxide of the element.20
The differentation of the two oxides as acide arsenieux and acide
arsenique in accordance with the new nomenclature of Morveau,
Lavoisier, Berethollet and Fourcroy (1787) naturally followed.
The recognition of the several arsenous acids: ortho, meta and
pyro and their several salts also of the isomeric trioxides came
later.
17 So called to contrast it with arsenicum citrinum (auripigment) and arseni-
cum rubrum (realgar), Kopp, Geschichte der Chemie, vol. 4, p. 90.
18 Kopp, Geschichte der Chemie, vol. 4, p. 90 ; also E. v. Meyer, History of
Chemistry (3rd ed.) p. 58.
1#Kopp, Geschichte der Chemie, vol. 4, p. 92.
“Kopp, Geschichte der Chemie, vol. 4, p. 99.
Lang enhan— The Arsenical Solutions.
147
As already pointed out, the medicinal use of arsenic compounds
was restricted largely to external purposes (depilatory, etc.)-
Whether arsenic, or more correctly, arsenous acid or its salts, or
possibly organic derivatives of arsenic, played a role in the famous
Aqua tofana of about 1700 may possibly never be definitely cleared
up.21 That white arsenic combines with aqueous alkalies was shown
by Maequer22 in 1746 and 1748, who called the products resulting
fois d’ arsenic , or arsenic liver, reminding of the times when ar¬
senic was supposed to be similar to sulphur.23
It was this property that Hughes, apothecary of the infirmary
at Stafford, made use of when requested by Fowler to analyze and
duplicate a quack remedy, known as “Tasteless ague or fever
drops” which the latter had found efficacious in the “cure” of
“agues, remitting fevers and periodic headaches”.24 This was in
1786. “It is universally recognized that Fowler introduced the
modern medicinal employment of the arsenic, but it should in
fairness be remembered that he was guided to his discovery by a
quack remedy, as he himself fully acknowledged”.25
Fowler’s solution quickly found its way into the pharmaco¬
poeias: first of all into the London Pharmacopoeia of 1809. 23 From
this it quickly passed into the first edition of the United States
Pharmacopoeia which was published in 1822. On the European
continent, it made its first appearance in the Pharmacopoea Han-
overana of 1819. Its rapid admission to the other state pharma¬
copoeias is indicated by the following chronological list:27
1822 Pharmacopoea Bawatrica
1823 Pharmacopoea Belgica
1826 Codex Medicament arius
21 Comp. Peters, Aug pharmazeutischer Vorzeit, neue Folge, sweite Aufl.,
(1899), p. 124.
22 See Kopp, Geschichte der Chemie, vol. 4, p. 95.
Maequer, Dictionary of Chemistry, pp„ 648 and 666 ; Maequer however, does
not differentiate between the potassium arsenite obtained by dissolving arsenic
trioxide in potassium carbonate solution, and potassium arsenate, obtained by
dissolving arsenic trioxide in aqueous potassium nitrate solution.
23 A similarity that is even today reflected in the classification of minerals
as illustrated by the isomorphous series of pyrites and arsenical pyrites.
34 Wootton, Chronicles of pharmacy, vol. 2, p. 134; See also Wilson’s patent
in appendix.
26 Wootton, p. 135.
28 Powell’s translation of the Pharmacopoeia Londinensis, 1809.
27 Whether in all cases the date given is that of the first appearance has not
been definitely settled because of the inability to consult complete sets of edi¬
tions in all cases.
148 Wisconsin Academy of Sciences, Arts, and Letters.
1827 Pharmacopoea Borussica
1831 Pharmacopoea Slesvico-Holsatica
1855 Pharmacopoea Austy*iaca
1868 Pharmacopoea Danica
While Macquer,28 who was professor of Medicine and Chemistry
at the University of Paris, points out that only quacks used arsenic
for internal administration, the regular medical practitioners of
the infirmary of Stafford, England, used the “Tasteless ague and
fever drops,” a secret preparation,29 during the years 1781, 1782,
and 1783 as a remedy against intermittent fevers. This adoption
of a “patent medicine”30 by the physicians of the infirmary,
caused the apothecary Hughes to analyse30 and imitate31 this secret
preparation, basing his experiments on the “pharmaceutic his¬
tory” of arsenic found in Lewis’s Materia Medica.32 Even of the
milder sulphide minerals, Lewis makes the following statement:
“The native minerals have been used as medicines in the eastern
countries, and by some imprudently recommended in our own”.
(Materia Medica, p. 104.)
According to the account of Dr. Withering of Birmingham, this
physician used a simple solution of arsemcum album, one grain to
each ounce of distilled water, the solution being “facilitated by
boiling for a minute or two in a Florence Flask, or other glass
vessel”. “Long continued boiling” he adds “disposes it to pre¬
cipitate again ’ \33 The same formula was employed by Dr. Arnold
of Leicester34 who directs the solution to be filtered. While this
solution was satisfactory in hospital practice, where the quantity
28 Diet, of Chemistry, p. 103: “Arsenic is a very violent corrosive poison:
it produces always the most painful symptoms and fatal effects, whether it be
taken internally or applied externally. It ought never to be employed medi¬
cinally, although some people not very intelligent, give small doses of it in
obstinate intermittent fevers, which it effectually cures, but is always attended
with bad consequences to the patients, such as phthisis, and other no less
troublesome diseases.”
29 Fowler, Medical Reports, Preface, p. vii.
30 Ibidem, Preface, p. vi ; See also Wootton, On patent medicines in Appendix.
Ibidem, p. 124, Withering, letter to Fowler.
31 Ibidem, Preface, p. vii.
32 An Experimental History of the Materia Medica of the Natural and Arti¬
ficial Substances Made Use of in Medicine. Containing A Compendious View
of their Natural History, An Account of their Pharmaceutic Properties, And
an Estimate of their Medicinal Powers, so far as they can be ascertained by
Experience, or by rational Induction from their sensible Qualities. By William
Lewis , London. 1761.
83 Fowler, Medical Reports, p. 124.
34 Ibidem, p. 117.
Langenhan — The Arsenical Solutions.
149
prepared was used up quickly, it did not suffice for private prac¬
tice because precipitation took place upon standing. In order to
overcome this, Mr. Baley, the apothecary of Birmingham, added
“nitre” to the solution “by the direction of Mr. Hughes”.35
Fowler, in his report states that he used “fixed vegetable alkali”.36
At the same time he points out that the “fixed vegetable alkali” is
not “perfectly pure; for that which is of the Druggists is fre¬
quently so impure, a double Proportion of purified nitre may be
substituted” instead of going to the trouble of purifying the al¬
kali.37
We have, therefore, at the very beginning, when arsenical ther¬
apy was transferred from the secret practices of nostrum venders
to the above board practice of the regular physician, three differ¬
ent pharmaceutical preparations:
1. ) The simple solution of arsenic trioxide in water;
2. ) The solution of arsenic trioxide in water with the aid of
potassium carbonate, a chemical solution socalled with the
formation presumably of a potassium arsenite;
3. ) The solution of arsenic trioxide in water with the aid of
potassium nitrate whereby presumably a potassium arsen¬
ate is formed.
All three solutions have found their way into pharmacopoeias
or similar standards. Thus the simple solution of arsenic trioxide
was official in the French Codex. That of the ratio of 1 :1000 as
Solute Arsenical ou Mineral (Boudin) in 1884. A Solute Arseni¬
cal Isnard, ratio 1 :10000, is likewise mentioned by Dorvault in his
L’Officine (1910), p. 1289. The solution of arsenic trioxide in
aqueous potassium carbonate is the one that became official in most
pharmacopoeias, with “Fowler’s Solution” either as title or syn¬
onym. The third solution has become official as Liquor Arsenicalis
Clemens, N. F., IV, though the oxidation is effected not by the
nitric acid set free from the nitre, but by bromine. Similar sodium
compounds are official as Liquor Sodii Arsenatis , U. S. P. and
Liquor Sodii Arsenatis, Pearson, N. F., IV.
Names and Synonyms. The compound resulting upon the union
of “white arsenic” (As203) and alkali in aqueous solution was
named fois d’ arsenic, liver of arsenic, by Macquer in 1776 because
of the similarity of arsenic to sulphur which produced the socalled
35 Ibidem, p. 125.
36 Ibidem, Preface p. viii ; also p. 79.
37 Ibidem, pp. 82 and 83.
150 Wisconsin Academy of Sciences, Arts, and Letters.
livers (liver of sulphur) under similar conditions. When Fowler
prepared his arsenical solution in 1786, the new nomenclature of
acids, bases and salts of Lavoisier and his committee of four (1787)
had not been worked out. Moreover, he desired particularly to
omit any reference to arsenic, hence he called his preparation
(t Mineral Solution”.38 Somewhat later the solution was merely
known as Liquor (solwHo) arsenicalis Fowleri and as such became
official in the London Pharmacopoeia of 1809.
In the U. S. P. of 1820 the English title is still “ Arsenical solu¬
tion”, but in the Latin title an attempt was made to make it con¬
form with the new nomenclature of the antiphlogistic system of
chemistry, hence Liquor Potassae Arseniatis.39 Whether this
should read Arsenitis and whether it is to be assumed that the ad¬
ditional “a” is due to a typographical error, it may be too late to
ascertain. The 1830 Philadelphia edition makes the correction :
Liquor Potassae Arsenitis, whereas the New York edition uses
Liquor Arsenicalis as the official Latin title and Solution of Arsen-
ite of Potass, as well as Fowler’ s solution as English synonyms for
Arsenical Solution.
The change from the oxygen acid to the hydrogen acid nomen¬
clature, viz. to Liquor Potassii Arsenitis ,, is made in 1870 and con¬
tinues throughout after that date. Fowler’s name appears only
in connection with the synonyms for the New York 1830 edition
and in the revisions of 1880, 1900 (index only) and 1910.
A list of recent pharmacopoeia! titles follows with references to
sources.40
38 In his report, preface page x, Fowler, makes the following statement :
“ as the idea of poison seems to be so strongly connected with that of ar¬
senic, it will be found to be very difficult to separate them in the mind, when¬
ever that term is named ; and therefore to correct as much as possible such
disagreeable ideas, in the practice of the healing art, the medicine now about
to be introduced to the notice of the public, will be distinguished by the name
Mineral Solution.”
39 Up to the present date the British Pharmacopoeia refuses to apply a chem¬
ical designation to this solution, but calls it Liquor Arsenicalis. In comment¬
ing on the British usage the editors of the U. S. Dispensatory, 13th ed. in 1872,
make the following statement: “The name by which this preparation is desig¬
nated in the U. S. Pharmacopoeia is the most correct. It has however, been
denied that the carbonate of potassa is decomposed by the arsenious acid,
which is supposed to be merely held in solution ; and, in this view of the nature
of the preparation the British name Arsenical Solution would be appropriate.”
40 For non pharmacopoeial names and synonyms, likewise for older pharma¬
copoeia! nomenclature see the glossary.
Langenhan — The Arsenical Solutions.
151
Arsenikdropper, Ph. Svenska (1908).
Arsenicalis Liquor Fowler H, Ph. Belg. (1906).
Arsenical Solution , Ph. B. (1914).
Fowler’s Arsenikdraaper, Ph. Norv. (1913).
Fowler sche Loesung , P. G. (1910).
Fowler’s Solution , U. S. P. IX.
Kalii arsenicosi liquor, Ph. Belg. (1906).
Kalium Arsenicosum solutum, Ph. Helv. (1907).
Licor Arsenical de Fowler , Ph. Hisp. (1905).
Liqueur Arsenicale de Fowler, Ph. Belg. (1906).
Liqueur de Fowler , Fr. Cod. (1908).
Liquor Arsenicalis , Ph. B. (1914).
Liquor Arsenicalis Fowleri,, Ph. Norv. (1913).
Liquor Arsenicalis ex Fowler, Ph. Hisp. (1905).
Liquor Arsenicale del Fowler (F. I.), Ph. Ital. (1909).
Liquor Arsenitis Kalici, Ph. Svensky, (1908).
Liquor Arseniitis Kalici, Ph. Norv .(1913).
Liquor Fowleri, Ph. Norv. (1913).
Liquor Kalii Arsenicosi , Ph. G. (1910).
Liquor Potassae arseniatis , U. S. P. (1820).
Liquor Potassii Arsenitis, U. S. P. (1916).
Potassium arsenit licuid, Ph. Roman. (1893).
Solucion de Arseniio Potasico, Ph. Hisp. (1905).
Solute d’ Arsenit e de Potasse, Fr. Cod. (1908).
Solutio Arsenitis Kalii composita, Nedrl. Ph. (1898).
Solutio Arsenicalis Fowleri, Ph. Russ. (1910).
Solutio Kalii arsenicosi, Ph. G. (1910).
Soluto le arsenito de Potassa, Ph. Port. (1876).
Solution of Potassiu\m Arsenit e, U. S. P. IX.
Solutum Arsenitis Kalici, Ph. Port. (1876).
Solutum arsenitis potassici, Ph. Hisp. (1905).
Soluzione del Fowler, Ph. Helv. (1907).
Soluzione alcalina di metarsenito di potassio, Ph. Ital. (1909).
The London and British Pharmacopoeias afford an interesting
example of apparent chemical misnomenclature. The Latin title
of the 1809 London Pharmacopoeia, the first to make Fowler ’s solu¬
tion official, is Liquor Arsenicalis. This title was used in the Lon¬
don Pharmacopoeia of 1824, but that of 1851 used the dualistic
title Liquor Potassae ArsenMis. The British Pharmacopoeia re-
i
152 Wisconsin Academy of Sciences , Arts , and Letters.
turned to the title Liquor Arsenicalis 41 and has maintained it
throughout its revisions ; the title of the London Pharmacopoeia of
1851, viz. Liquor Potassae Arsenitis being used as a synonym.
If the official title is to be justified because we really know little
about the exact chemistry of the solution, this excuse or line of
reasoning will not hold when we study the English practice as to
the designation of the principal ingredient of the solution. In the
1809 London Pharmacopoeia it was Arsenici Oxidum Sublimatum,
hence in accordance with the dualistie nomenclature. Yet in 1824
it was changed back to Arsenicum album sublimatum. In 1851 it
became Acidum Arseniosum, again in accordance with the dualistie
nomenclature. But whereas the designation of 1809, viz. Arsenici
oxtidum, is indicative of a base, that of 1851, Acidum Arseniosum ,
is plainly indicative of an acid. It is possibly because of this that
the phrase ( Acidum metallicum sublimation praeparatum) is added
in parenthesis. Whereas the element is still pronounced metallic
in character, its oxide is now, not a base, but an acid.
A peculiar incongruity in spelling is found in the words arseni-
ous and arsenite. The ending ous is that of the acid with lower
oxygen content, and becomes ite for the salts thereof. The ending
ic is that of acids of higher oxygen content and becomes ate for its
salts. Hence arsenic should yield :
A. ) arsenous acid, and its salts should be arsenites; also
B. ) arsenic acid, and its salts should be arsenates.
However the former acid is commonly spelled arsenious acid
(P. L. 1851 ; B. P. 1864, 1885, 1898, 1914.). Hence the salt should
be arseniite, but almost invariably it is spelled without the second
i i j, > > 42
The English pharmacopoeias are not the only offenders in this
direction. Koscoe and Schorlemmer, in their “Treatise on Chem¬
istry”, vol. II, p. 1246 (1911) still use arsenious acid and arsenites.
Our own Pharmacopoeias, after having clung to arsenious acid
(Latin, Acidum Arseniosum) from 1820 to 1880 inclusive, in 1890
changed its spelling to Arsenous acid (Latin, Acidum Arsenosum),
only to be changed in the subsequent editions to Arsenic Trioxide
(Latin, Arseni Trioxidum) .
41 “This appelation is certainly very objectionable, as it conveys an erroneous
idea of the preparation, even admitting that the term arsenic may be used to
designate the white oxide : it should have been Liquor Arsenialis Potassae or
Liquor alcalinus Oxidi Arsenici ”, A. T. Thomson, The London Dispensatory,
4th edition, 1826.
42 The single exception observed thus far is found in the Norwegian Pharma¬
copoeia of 1913: Liquor Arseniitis Kalici.
Langenhan — The Arsenical Solutions.
153
Pharmacopoeial. Text. In as much as Fowler’s solution was
made official in the first edition of the U. S. Pharmacopoeia, was
included in both the New York and Philadelphia editions of 1830,
and has remained official up to the present time, there are eleven
written texts to be considered. Moreover, the 1820 edition and
both of the 1830 editions contain the text in both Latin and Eng¬
lish. The detailed comments on the text, whether based on the
study of the literature of the subject or on laboratory experiments
or observations, are recorded in connection with those parts of the
text to which they have reference. A better idea of this part may
be had by a mere glance at the list of text subjects commented
upon:
1. ) Titles and Synonyms.
2. ) Definition.
3. ) Preservation.
4. ) Arsenic trioxide as an ingredient.
5. ) The form in which it is used.
6. ) Potassium carbonate or bicarbonate as an ingredient.
7. ) Ratio of ingredients.
8. ) Water.
9. ) Amount of water to effect solution.
10. ) Solution of arsenic trioxide.
11. ) Alcohol and compound tincture of lavender.
12. ) Volume of finished product.
13. ) Appearance of finished product.
14. ) Qualitative tests.
15. ) Assay.
16. ) Dose.
Note. For convenient reference, the numbers representing the sub¬
jects commented upon are incorporated in the text proper.
154 Wisconsin Academy of Sciences , Arts, and Letters.
LIQUOR POTASSAE ARSENIATIS1 U. S. P. 1820
Ingredients.
R. Acidi arseniosi,4 in pulverem
snbtilem redacti;5
Potassae subcarbonatis,6 utri-
usque grana sexaginta qua-
tuor.7
Aquae distillatae8 oetantem
unum.9
Arsenical
Ingredients.
Take of Arsenious acid,4 in fine
powder ;5
Subcarbonate of potass,6 each
sixty-four grains.7
Distilled water,8 one pint.9
Directions.
In vase vitreo concoque donee
acidum arseniosum omne resolva-
tor.10 Liquorem frigefaetum in
mensuram octariam infunde ; et
adde
Alcoholis fluidr'aehmas quatuor,11
Et aquae distillatae quantum
suffecerit ad mensuram complen-
dam.12
Solution1
Directions. ,
Boil them together in a glass
vessel until the arsenic is entirely
dissolved.10 When the solution is
cold, pour it into a pint measure,
and add thereto Alcohol,11 four
fluidrachms, and so much distilled
water, as shall fill up the measure.12
LIQUOR ARSENICALIS1 U. S. P. 1830 N. Y.
Ingredients.
R. Acidi arsenici4 in pulverem
subtilem redacti,5
Potassae subcarbonatis,6 utri-
usque grana sexaginta qua¬
tuor,7
Aquae distillatae8 oetantem
unum,9
Alcoholis fluidrachmas qua¬
tuor j11
Directions.
In case vitreo concoque donee
acidum arseniosum omne resolva-
tur.10
Liquorem frigefaetum in men¬
suram octariam infunde; et adde
alcohol, et aquae distillatae quant¬
um suffecerit ad mensuram comp-
lendam.
Langenkan — The Arsenical Solutions.
155
Arsenical Solution
Solution of Arsenite of Potass. Fowler’s Solution1
Ingredients.
Take of Arsenious acid,4 in fine
powder,5
Subcarbonate of potass,6 each
sixty-four grains,7
Distilled water8 one pint,9
Alcohol, four fluidrachms,11
Directions.
Boil them together in a glass
vessel, until the arsenic is entirely
dissolved.10. When the solution is
cold, pour it into a pint measure,
and add to it the alcohol and so
much distilled water, as shall fill
up the measure.12
LIQUOR POTASSAE ARSENITIS1 U. S. P. 1830 Phila.
Ingredients.
R. Acidi arseniosi4 in pulverem
subtilissimum triti,5
Potassae Carbonatis Puris-
simi,6 singulorum, grana
sexaginta quatuor;7
Aquae Destillatae8 quantum
satis sit;
Spiritus Lavandulae Compo-
siti fluidrachmas quatuor.11
Directions.
Acidum Arseniosum et Potassae
Carbonatem cum Aquae Destil¬
latae octario9 coque, in vase vitreo.
donee Acidum omnino liquetur.10
Liquori frigefacto ad j ice Spiri¬
tual Lavandulae, et postea Aquae
Destillatae quantum satis sit ut
mensuram octarii accurate im-
pleat.12
Solution of Arsenite of Potassa1
Ingredients.
Take of Arsenious Acid,4 in very
fine powder,5
Purest Carbonate of Potassa,
each,6 sixty-four grains,7
Distilled Water a sufficient
quantity ;
Compound Spirit of Lavender
four fluidrachms.11
Directions.
Boil the Arsenious Acid and
Carbonate of Potassa with a pint9
of Distilled Water, till the Acid is
entirely dissolved.10 To the solu¬
tion, when cold, add the Spirit of
Lavender, and afterwards suffi¬
cient Distilled Water to make it fill
up exactly the measure of a pint.12
156 Wisconsin Academy of Sciences , Arts , and Letters .
LIQUOR POTASSAE ARSENITIS U. S. P. 1840
Solution of Arsenite of Potassa1
Ingredients.
Take of Arsenious Acid,4 in
small fragments,5
Pure Carbonate of Potassa,
each,6 sixty-four grains;7
Distilled Water8 a sufficient
quantity ;
Compound Spirit of Lavender
half a fluidounce.11
Directions.
Boil the Arsenious Acid and
Carbonate of Potassa with twelve
fluidounces9 of Distilled Water, in
a glass vessel, until the Acid is en¬
tirely dissolved.10 To the solution,
when cold, add the Spirit of Lav¬
ender, and afterwards sufficient
Distilled Water to make it fill
exactly the measure of a pint.12
LIQUOR POTASSAE ARSENITIS U. S. P. 1850
Solution of Arsenite of Potassa1
Ingredients.
Take of Arsenious Acid,4 in
small fragments,5
Pure Carbonate of Potassa,6
each, sixty-four grains ;7
Distilled Water8 a sufficient
quantity ;
Compound Spirit of Lavender
half a fluidounce.11
Directions.
Boil the Arsenious Acid and
Carbonate of Potassa, in a glass
vessel, with twelve fluidounces of
Distilled Water, till the Acid is
entirely dissolved.10 To the solu¬
tion, when cold, add the Spirit of
Lavender, and afterwards suf¬
ficient Distilled Water to make it
fill exactly the measure of a pint.12
LIQUOR POTASSAE ARSENITIS U, S. P. 1860
Solution of Arsenite of Potassa1
Ingredients.
Take of Arsenious Acid,4 in
small pieces,5
Bicarbonate of Potassa,6 each,
sixty-four grains;7
Compound Spirit of Lavender
half a fluidounce;11
Distilled Water8 a sufficient
quantity.
Directions.
Boil the Arsenious Acid and Bi¬
carbonate of Potassa, in a glass
vessel, with twelve fluidounces9 of
Distilled Water till the Acid is
entirely dissolved.10 To the solu¬
tion, when cold, add the Com¬
pound Spirit of Lavender, and
afterwards sufficient Distilled Wa¬
ter to make it measure a pint.12
Lang enhan— -The Arsenical Solutions.
157
LIQUOR POTASSII ARSENITIS U. S. P. 1870
Solution of Arsenite of Potassium1
Ingredients.
Take of Arsenions Acid,4 in
small pieces,5
Bicarbonate of Potassium,6 each,
sixty-four grains;7
Compound Spirit of Lavender
half a fluidounce ;lx
Distilled Water8 a sufficient
quantity.
Directions.
Boil the Arsenious Acid and Bi¬
carbonate of Potassium, in a glass
vessel, with half a fluidounce9 of
Distilled Water, until the Acid is
entirely dissolved,10 and add twelve
fluidounces of Distilled Water.
Then add the Compound Spirit of
Lavender, and afterwards enough
Distilled Water to make it meas¬
ure a pint.12
LIQUOR POTASSII ARSENITIS U. S. P. 1880
Solution of Arsenite of Potassium
[Fowler’s Solution.]1
Arsenious Acid,4 in small pieces,5 one part 7 . . . 1
Bicarbonate of Potassium,6 one part 7 . . . . 1
Compound Tincture of Lavender,11 three parts . . 3
Distilled Water* a sufficient quantity,
To make one hundred parts 12 . 100
Boil the Arsenious Acid and Bicarbonate of Potassium in a
glass vessel with ten (10) parts 9 of Distilled Water, until the Acid
is completely dissovled.10 Then add the Compound Tincture of
Lavender, and enough Distilled Water to make the product weigh
one hundred (100) parts. Lastly, set the mixture aside for eight
days and then filter through paper.3
If 24.7 Cm. of the Solution are boiled with 2 Gm. of bicarbonate of
sodium, the liquid, when cold, diluted with 100 C.c. of water, and some
gelatinized starch added, should require from 48.5 to 50 C.c. of the vol¬
umetric solution of iodine, before the blue color ceases to disappear on
stirring (corresponding to 1 per cent, of arsenious acid of the required
purity).15
158 Wisconsin Academy of Sciences, Arts, and Letters.
LIQUOR POTASSI ARSENITIS U. S. P. 1890
Solution of Potassium Arsenite
[Fowler’s Solution.]1
Arsenous Acid,4 in fine powder,5 ten grammes 7 . 10 Gm.
Potassium Bicarbonate,6 twenty grammes T . 20 Gm.
Compound Tincture of Lavender,11 thirty cubic centimeters - 30 Cc.
Distilled Water,8 a sufficient quantity.
To make one thousand cubic centimeters12..- . 1000 Cc.
Boil the Arsenous Acid and Potassium1 Bicarbonate with one
hundred (100) cubic centimeters9 of Distilled Water, until solu¬
tion has been effected.10 Then add enough Distilled Water to make
the solution, when cold, measure nine hundred and seventy (970)
cubic pentimeters, and, lastly, add the Compound Tincture of Lav¬
ender. Filter through paper.
If 24.7 Cc. of the Solution be boiled for a few minutes with 2 Gm. of
sodium bicarbonate, and the liquor, when cold, diluted with water to 100
Cc., and mixed with a little starch T.S., it should require from 49.4 to 50
Cc. of decinormal iodine V.S. to produce the blue tint of iodide of starch
(corresponding to 1 Gm. of arsenous acid in 100 Cc. of the Solution).15
LIQUOR POTASSII ARSENITIS U. S. P. 1900
Solution of Potassium Arsenite1
An aqueous solution, which should contain Potassium Arsenite
corresponding in amount to 1 percent, of arsenic trioxide.2
Arsenic Trioxide,4 in fine powder,5 ten grammes'1 . 10 Gm.
Potassium Bicarbonate,6 twenty grammes 7 . . . 20 Gm.
Compound Tincture of Lavender,11 thirty grammes....- . 30 Gm.
Distilled Water,8 a sufficient quantity,
To make one thousand grammes 12 . 1000 Gm.
Boil the Arsenic Trioxide and Potassium Bicarbonate, in a
tared dish, with one hundred grammes9 of Distilled Water, until
solution has been effected.10 Then add enough Distilled Water to
make the solution weigh nine hundred and seventy grammes, and,
lastly, add the Compound Tincture of Lavender. Filter through
paper.
If 24.6 Gm. of Solution of Potassium Arsenite be diluted with water to
100 Cc., the mixture very slightly acidified with diluted hydrochloric acid,
and then made alkaline with 2 Gm. of sodium bicarbonate, it should re¬
quire not less than 50 Cc. of tenth-normal iodine V.S. to produce a perma¬
nent yellow tint (corresponding to 1 Gm. of arsenic trioxide in 100 Gm.
of the Solution.)15
Average dose. — 0.2 Cc. (3 minims).16
Langenhan — The Arsenical Solutions.
159
LIQUOR POTASSII ARSENITIS U. S. P. 1910
Solution of Potassium Arsenite
Jjiq. Pot. Arsen.— -Fowler’s Solution Liquor arsenicalis Fowleri P.I.
An aqueous solution containing potassium arsenite, correspond¬
ing in amount to not less than 0.975 per cent, nor more than 1.025
per cent, of As203 (197. 92). 2 Preserve the Solution in amber-col¬
ored bottles.3
Arsenic Trioxide,4 in fine powder,5 ten grammes 7 . . 10 Gm.
Potassium Bicarbonate,6 twenty grammes T . _.... 20 Gm.
Compound Tincture of Lavender,11 thirty grammes . _.. 30 Gm.
Distilled Water,8 a sufficient quantity ,9
To make one thousand grammes 12 . 1000 Gm.
Boil the arsenic trioxide and potassium bicarbonate in a tared
flask with one hundred grammes of distilled water, until solution
has been effected,10 and then add enough distilled water to make the
solution weigh nine hundred and seventy grammes; then add the
compound tincture of lavender and filter through paper.
Solution of Potassium Arsenite is a clear or faintly opalescent liquid
with a pink color and a slight odor of lavender.13 It is alkaline to litmus.14
Acidulate strongly 10 mils of the Solution with hydrochloric acid and
add an equal volume of hydrogen sulphide T.S.; a yellow precipitate is
produced which dissolves in ammonium carbonate T,S.14
Acidulate 4 mils of Solution of Potassium Arsenite with diluted nitric
acid, add 1 mil of silver nitrate T.S. and superimpose a layer of ammonia
water upon this liquid; no red or reddish-brown color is observed at the
line of contact (arsenate) . 14
Assay — Dilute about 20 mils of Solution of Fotassium Arsenite, accur¬
ately weighed, with 75 mils of distilled water, acidify the mixture very
slightly with diluted hydrochloric acid, then dissolve in the solution 2 Gm.
of sodium bicarbonate and titrate the resulting liquid with tenth-normal
iodine V.S., starch T.S. being used as indicator. It shows not less than
0:975 per cent, nor more than 1.025 per cent, of As203.15
Each mil of tenth-normal iodine V.S. used corresponds to 0.004948 Gm.
of As^Og. Each gramme of Solution of Potassium Arsenite corresponds
to not less than 1.97 nor more than 2.07 mils of tenth-normal iodine V.S.15
Average dose— Metric, 0.2 mil — Apothecaries, 3 minims.16
160 Wisconsin Academy of Sciences , Arts , and Letters.
Comments on U. S. P. Text
1. ) Titles and Synonyms. See Names and Synonyms.
2. ) Definition. A definition is not given previous to 1900. The
modification in the definition for 1910 was necessitated by the Pure
Food and Drugs Act of 1906 which demanded reasonable limits
rather than an absolutely fixed standard.
3. ) Preservation. The 1910 revision is the only one that directs
that the solution be kept in amber colored bottles to protect it
against light.
Fowler ’s Solution upon standing deposits a flocculent precipi¬
tate. This precipitate has apparently introduced a problem for
comments and criticisms as to the cause, the composition and the
ways and means of eliminating this objectionable factor. Unfor¬
tunately much of the work contributed on this topic is merely theo¬
retical and not the result of experimental labor, at least no data
of an experimental type are given in the majority of papers pub¬
lished.
The causes for this precipitate as given, may be considered un¬
der three groupings :
1st. The action of certain organisms in the water, resulting in
an algaceous growth.
2nd. The action of the alkali on the glass container.
3rd. The action of the alkali on one or more of the ingredients
of the alcoholic tincture or spirit used.
Those supporting the first theory are: J. Mueller43 who sug¬
gests the addition of 0.4 p. c. of borax, as a preservative ; Perschne44
who suggests the use of glycerin for the same purpose ; Kollo45 and
Benysek46 who claim that the alcohol present acts as a preserva¬
tive;47 while Feuer48 consideres its preservative action negligible;
and Dannenberg49 who speaks of the precipitate as an algaceous
growth. Brautigam50 apparently disagrees with the above, in his
claim that the precipitate is not an algaceous growth but results
from the action of the alkali on the glass container. This is sup¬
ported in part by Neinhaus51 who calls attention also, to the U. S. P.
43 Am. Jour. Pharm., 51, p. 249; from Pharm. Ztg. (1879.)
44 Am. Journ. Pharm., 55, p. 456; from Pharm. Ztschr. f. Russland (1887.)
46 Pharm. Post. 29, p. 233 (1896.)
48 Pharm. Post., p. 157 (1896.), see also ibid. 32, p. 49.
47 Presumably the preservative action of the alcohol tends to prevent the
growth of organisms that result in the formation of a flocculent mass.
48 Pharm. Post., 25, p. 381 (1895.)
49 Pharm. Centralh., 22, p. 319 (1881.)
50 Pharm. Centralh., 33, p. 324 (1892.)
61 Pharm. Ztg., 37, p. 331 (1892.)
Langenhan — The Arsenical Solutions.
161
of 1880 which directed that the solution be filtered after standing
eight days, to remove the precipitate formed. The editor of the
Chemiker Zeitung52 calls attention to a previous custom of neutral¬
izing the excess alkali with acetic acid53 to prevent this same reac¬
tion. Evidently in a parallel line of thought Garraud54 suggests
using a standard alkali55 to avoid excess alkalinity in the finished
product.
Seemingly most writers support the third theory, viz. the action
of the alkali on the ingredient or ingredients of the alcoholic tinc¬
ture or spirit used. M. Goeldner56 suggests the use of phenol-
phthalein as coloring agent in place of such a pigment as that of
red saunders of the compound tincture of lavender, evidently as¬
suming that the alkali and the pigment from the red saunders
cause the precipitate. J. Luettke57 attributes the “ dirty color”
appearing in the P. G. solution to the action of the alkali and com¬
pound spirit of melissa58 and suggests avoiding this by neutraliz¬
ing the solution. Nienhaus51 claims that the alkali and aromatic
tinctures and even distilled spirits used in the various pharma¬
copoeias are the main causes of the precipitate. He -substitutes
melissa water for the alcoholic distillate in the P. G. solution, and
does not obtain a precipitate. The substitution of compound
spirit of melissa by melissa water is also recommended by Pascal59
and Menhaus,60 while Mueller60a suggests using the simple spirit of
melissa. An unsigned article61 states that the precipitate is due
to the action of the excess alkali on the red saunders. The “ query”
editor of the Druggists Circular 62 suggests all of the already men-
62 Chem. Zt g., 16, p. 174 (1892.)
53 To what extent the solution is neutralized is not made clear. As the alka¬
linity of the solution is supposed to be, in part, due to the hydrolysis of the
formed arsenite (see qualitative tests No. 13) a complete change to potassium
acetate might take place if sufficient acid were added, before the solution
would respond to a “neutral test.”
64 Pharm. Ztg., 39, p. 753; from Bull. Soc. Pharm. de Bordeux (1894.)
65 Here again the alkalinity due to any potassium arsenite present has been
overlooked, apparently, or it. has been assumed that the alkalinity is negligible.
KBer. d. Pharm. Gesellch., 2, p. 127 (1892.), see also p. 302.
57 Pharm. Centralh., 559 (1890.)
68 Compound Spirit of Melissa P. G. is prepared by macerating a mixture of
melissa leaves, lemon peel, nutmeg, cinnamon and cassia buds with hydro¬
alcohol and subsequently distilling until a definite amount of distillate has been
obtained. This spirit apparently contains only volatile matter as compared
with the Compound Tincture of Lavender U. S. P. which is prepared by macera¬
tion and filtration.
69 Pharm. Ztg., 50, p. 1009 (1905.), from Bull. Commercial.
60 Nat. Drug., 21, p. 22 (1893.)
6°a Ber. d. pharm. Ges., 2, p. 302 (1892).
81 Proc. A. Ph. A., 46, p. 694 (1896.) from Merck’s Report
82 Drug. Cir., 52, p. 223 (1908.)
11— S. A. L.
162 Wisconsin Academy of Sciences, Arts, and Letters.
tioned reasons and adds the possible insoluble impurities in the ar¬
senic trioxide used. The impurities in the arsenic trioxide used
are also emphasized by Glueeksman.63 Chumaceiro64 refers to the
“International Formula”65 for preparing a clear permanent solu¬
tion. This evidently eliminates the alkalinity. Moreover this
formula directs that spirit of lavender be used in place of the tinc¬
ture. However the finished product in this case is not a solution
of an arsenite,65
The strength of Fowler ’s Solution is based on the arsenic trioxide
content. That a gradual oxidation of the trivalent arsenic to the
pentavalent arsenic takes place is generally accepted. The cause
of this oxidation however, has called forth some considerable in¬
vestigation and comment. Apparently all investigators agree that
the introduction of atmospheric oxygen, in various ways, is the
direct cause, but the conditions which accelerate or retard such
oxidation seem to offer a source of disagreement. G. Kassner66
attributes the oxidation to the introduction of atmospheric oxygen,
but does not mention any conditions influencing the rate. C.
Feuer67 claims that the oxidation due to atmospheric oxygen is
slight within a half year’s time, and that the alkalinity of the solu¬
tion accelerates this change. Fosenthaler68 merely points out that
oxidation of arsenous acid takes place. Lyons69 claims that under
like conditions the greatest amount of oxidation takes place in an
alkaline solution, the least in an acid solution, while in a solution
containing the (HC03) radicle the rate of oxidation lies between
the two. In a later publication Lyons70 claims that no oxidation
takes place in an acid solution. L. A. Brown71 suggests keeping
the solution in small well filled and closed containers to prevent
63 Pharm. Post., 29, p. 353 (1896.)
64 Ibid. 52, p. 323 (1908.)
65 According to the “International Formula” as given in this article, “spirit
of lavender (colorless distillate)” is used in place of the Compound Tincture of
Lavender U. S. P. Also some diluted hydrochloric acid is added, 3.5 grams
(13.67 p. c.) to 1 gram of potassium carbonate. This represents 0.458 grams
HC1, which requires 1.72 grams of potassium carbonate for neutralization. As
only 1 gram of potassium carbonate is used evidently the finished product is an
acid solution.
«8 Pharm. Ztg., 34, p. 419 (1889.)
« Pharm. Post., 28, p. 381 (1895.)
68 Pharm. Centralh., 47, p. 850 (1906.)
«9Proc. A. Ph. A., 56, p. 901 (1908.)
70 Proc. A. Ph. A., 57, p. 904 (1909.)
71 Ky. Agr. Exp. Sta. Bull. No. 150, p. 153 (1910.)
Langenhan — The Arsenical Solutions. 163
oxidation. A. N. Cook72 attributes the oxidation to the atmospheric
oxygen which is introduced every time the container is opened to
remove some of the liquid. E’we and Vanderkleed73 state that,
4 ‘ One sample of liquor potassium arsenite U. S. P. suffered an oxi¬
dation of 1.9 p. c. of its As203 as arsenite to arsenate in two years
and three months. Another sample had 1.8 p. c. of As203 as ar¬
senite oxidized to arsenate in one year and five months”. Engel-
hardt and Winters,74 after determining the amounts of arsenate
and arsenite respectively during a period of 14 months conclude
that only a slight amount of oxidation takes place in the solution.
4. ) Arsenic Trioxide as an Ingredient. The original formula of
Fowler calls for arsenicum album or “ white arsenic”, in which
manner the arsenic trioxide was distinguished from the yellow and
red sulphides. The London Pharmacopoeia of 1808 calls for Ar-
senici oxidum sublimatum. The U. S. P. of 1820 adopted the desig¬
nation arsenious acid in accordance with the new antiphlogistic-
nomenclature. For other changes, see Names and Synonyms.
5. ) The Form in Which it is Used. Fowler directs the “ white
arsenic” to be used “in pulverum subtilissimum triti, i. e. “tritu¬
rated to a very fine powder”. The U. S. P. of 1820 directs that
the “arsenious acid” be used in form of a “fine powder”; the
Philadelphia edition of 1830 directs a “very fine powder”.
“Small fragments” are suggested by the New York edition of
1830, also by the editions of 1840, 1850, 1860 and 1870. From 1880
on a “fine powder” is again directed.
The explanation for this is found in the following statement made
by the editors of the U. S. Dispensatory, 13th ed. (1872), p. 1267:
“In making this preparation, care should be taken that the arseni¬
ous acid is pure. This object is best secured by using the acid in
small pieces instead of in powder. Sulphate of lime is a common
impurity in the powdered acid, and if present will remain undis¬
solved, and cause the solution to be weaker than it should be”.
Although the U. S. P. of 1870 directs a qualitative test (com¬
plete volatilization) to be made, that would bar such an impurity
as sulphate of lime, the U. S. P. of 1880 is the first to direct the
assay of “arsenious acid”. It is, no doubt, due to this difference
in the requirements of the 1880 edition and its precursors from
72 Rep. So. Dak. P. & D. Com., 1912, p. 75 (1912.)
73 Proc. Pa. Ph. Assn., 37, p. 277 '(1914.)
74 Jour. A. Ph. A., 6, p. 134 (1917.)
164 Wisconsin Academy of Sciences , Arts , and Letters.
1840 on that the “fine powder” is again directed to be used in
1880.
6. ) Potassium Bica/rbonate or Carbonate as an Ingredient.
Fowler calls it Sal alkalinum fixum vegetabilis or “salt of fixed
vegetable alkali” to differentiate it from the fixed mineral alkali
or sodium carbonate and the volatile alkali or ammonium carbon¬
ate. The designation of the U. S. P. 1820 is in accordance with
the antiphlogistic nomenclature. The alkali used to facilitate the
solution of the “arsenious acid” varies. According to the U. S. P.
1820 and the New York edition of 1830 it is Potassae subcaxbonas
or “Subcarbonate of potass.”, which according to the 1820 edition
is the “Impure subcarbonate of potass.” or “Pearl ash”. The
Philadelphia edition of 1830 designates that “Purest Carbonate of
Potassa” or Potassae Carbonas Purissimup be used, which is to be
prepared from potassium bitartrate by ignition and lixiviation. (p.
172, not that in the Materia Medica list p. 20.) The editions of
1840, 1850 and 1860 direct that a “pure” carbonate be used. Ac¬
cording to the 1840 edition this is prepared from the bitartrate,
according to the 1850 and 1860 editions, by heating the bicarbonate.
From 1870 the bicarbonate is directed to be used in the formula.
Upon heating, as directed, the pure bicarbonate is changed to the
pure carbonate.
7. ) Ratio of Ingredients. Not only the ratio, but the exact quan¬
tities, viz. 64 grains of both the acid and alkaline ingredients, are
the same in all editions of the U. S. P. up to 1870 inclusive, as they
are in the original formula of Fowler. This would seem quite
rational but for two considerations, viz. :
1. The purity of the potassium carbonate originally employed,
no doubt, varied greatly. (For details of this see No. 6.)
2. In the 1870 edition, bicarbonate of potassium was substituted
for carbonate, but the amounts (64 grains of each) remained the
same.
Hence the ratio of acid (anhydride) to alkali (carbonate or bi¬
carbonate) fluctuated greatly in the several revisions from 1820 to
1870 inclusive. Although the U. S. P. 1880 is a relatively modern
treatise, as compared with its immediate precursors, the ratio was
not changed in 1880, but the change was made in 1890, so that
since that date the alkalinity of Fowler’s Solution was once more
what it may be supposed to have been originally, provided, how¬
ever, that the “alkaline vegetable salt” of Fowler’s formula was
Langenhan — The Arsenical Solutions.
165
pure potassium salt, which, in all probability it was not. In as
much as it is impossible to establish the purity of the potassium
carbonate used in the original formula, it would be vain to attempt
to duplicate, in this respect, the original formula.
It might be argued that the “white arsenic” of Fowler’s days
was not as pure as the present pharmacopoeial product is supposed
to be, and that the impurities in both ingredients compensated
each other. However, it is scarcely to be assumed that the “white
arsenic” or arsenic trioxide was ever as impure as the potassium
carbonate made from wood ashes, unless it was wilfully adulter¬
ated. (For table of ratios see Number 12.)
8. ) Water . According to his original formula, Fowler de¬
manded “distilled well water”. This is probably the reason why
distilled water has always been demanded by the U. S. P.75 A per¬
manently hard water might be considered as interfering with the
desired alkalinity of the finished product.76
9. ) Amount of Water Used to Effect Solution. According to
Fowler’s original formula only “a half pound” of water, neces¬
sary for the finished preparation, was used to effect the solution.
The London Pharmacopoeia of 1809, also the U. S. P. of 1820 di¬
rected “one pint” to be used, the loss resulting from evaporation
to be made up. So did both editions of 1830. The edition of 1840
reduced the amount to “twelve fluid-ounces”, as did the editions
of 1850, 1860 and 1870. In the 1880 edition, the amount was re¬
duced much more, namely to one- tenth of the volume of the finished
product. Practically the same ratio obtained in the editions of
1890, 1900 and 1910. The London Pharmacopoeia throughout
made no reduction. However, the first British Pharmacopoeia in
1864 reduced the amount, prescribing 10 ounces to be used, like¬
wise the edition of 1885. The 1898 edition directed either 10
ounces or 500 cc., depending upon whether the metric formula or
the apothecaries formula was used as both were given in this edi¬
tion. The 1914 edition prescribes 500 cc. In all B. P. formulas
the amount of water directed to be used to effect solution is equal
to one-half of the volume of the finished product.
75 The several editions of the London and British pharmacopoeias, also the
recent editions of all other pharmacopoeias consulted demand distilled water.
76 Such a water, which, however, no pharmacist would think of using, is the
public spring water of Sheboygan, Wis. It contains per liter 0.2394 Mg. equiv.
1.358 K^COg, and 1.0832 Ca. equiv. 3.448 K^COg, or a total of 4.806 K2C03. Hence
of the 13.804 KgCOg = 20.000 KHC03 used, about 9.000 K2C03 remain for the
solution of the 10 gm. As2Os and the desired surplus alkalinity.
166 Wisconsin Academy of Sciences , Arts , and Letters .
It has been commonly assumed that the original amount of water
used was reduced for the purpose of accelerating the reaction be¬
tween the acid and the alkali carbonate and thus increasing the
rate of solution. Thus E. Martin77 modified the U. S. P. process
by dissolving 64 grains each of arsenic trioxide and potassium bi¬
carbonate in two or three drams of water with the aid of heat.
When solution was effected he added the remainder of the water.
This procedure he claimed decreased the time ordinarily required
to make Fowler’s solution. Gr. Hamel78 placed the arsenic trioxide
and potassium carbonate, (3 grams each) in a test tube moistened
the mixture with a few drops of water and applied heat until a
liquid mass resulted. This he then diluted with a few mils of
water and heated until solution took place, after which more water
was added, up to the required volume. P. Maysse79 modified the
‘ ‘ Codex ’ ’ method by heating one gram each of arsenic trioxide and
potassium carbonate with five drops of water, in a test tube, until
the solution took place and then diluting up to 100 cc.
In order to test the correctness of this explanation, experiments
on the rate of solution in varying concentrations were performed.
A.) One gram of arsenic trioxide and two grams of postassium
bicarbonate (the ratio of As203 and KHC03 used in the U. S. P.),
were mixed with 25 mils, 50 mils, 100 mils, and 200 mils respec¬
tively, of boiling water, and the time required to obtain a clear
solution noted. The effervescence, caused by the decomposition of
the bicarbonate, had a tendency to carry the arsenic trioxide up
the sides of the container, hence to interfere with the action of the
solvent, even though a stirrer was used and vigorously at that.
To eliminate this objectionable feature the following modification
was adopted. The potassium bicarbonate was dissolved in water,
(20 gm. in sufficient water to make 100 mils.) and the solution
boiled until no more carbon dioxide was apparently given off. This
required about 30 minutes. An aliquot portion of this solution,
representing the amount of potassium acid carbonate desired, was
diluted with water up to a definite volume and the mixture heated
to boiling. The required weight of arsenic trioxide was introduced
into a test tube of suitable capacity, the boiling solution of the
potassium salt poured on to this and the tube immediately im-
77 Am. Jour. Pharm., 44, p. 202 (1872.)
78 Pharm. Ztg., 61, p. 402 (1906.)
79 Bull, des sc. pharmacol. 13, p. 106 (1906); through Digest of Comments,
1906.
Langenhan — The Arsenical Solutions.
167
mersed into a bath of boiling water. The mixture was constantly
agitated and the time required to obtain a clear solution was re¬
corded. Three separate determinations were made for each dilu¬
tion. The results are recorded in the following table:
Table I.
A glance at the table reveals the fact that the differences for the
several dilutions are no greater than those obtained from the same
dilution, hence fall within the limits of experimental error.
B.) In the next series an attempt was made to secure more
satisfactory results by increasing the amounts of the chemical
agents acting upon each other. Ten grams of arsenic trioxide and
the equivalent 20 grams of potassium bicarbonate (as carbonate)
with varying amounts of water were used. In this series one ex¬
periment only was performed with each volume of water. The
results are herewith tabulated.
proper conditions, the rate of solution is hastened to any appre¬
ciable extent by greater concentration.
C.) A third series was made in an attempt to note the effect of
the effervescence (mentioned in the preliminary experiments),
when larger quantities of reagents were used. Ten grams of ar¬
senic trioxide and 20 grams of potassium bicarbonate were mixed
with 25 mils, 50 mils and 100 mils of boiling water, and the time
of solution determined as before. The results are again tabulated.
168 Wisconsin Academy of Sciences , Arts , and Letters.
The slight increase in time is, no doubt, due principally to the
removal of the arsenic trioxide from the solution by the escaping
carbon dioxide.
D.) In a fourth series of tests ten grams of arsenic trioxide and
11.2 grams of potassium hydroxide, the equivalent of 20 grams of
potassium acid carbonate, the latter dissolved in 25 mils, 50 mils
and 100 mils respectively, of boiling water, were mixed, and the
time required for solution noted as above. In each case the time
was not more than one minute. Apparently the arsenic trioxide
dissolved as fast as it came in contact with the solvent quite ir¬
respectively of the degree of concentration of the alkaline solvent.
10.) Solution of Arsenic Trioxide. For the chemistry of the
solution see Hydration of Arsenic trioxide.
The formula for potassium arsenite given in the U. S. P. IX is
KAs02, potassium metarsenite. Whether it is assumed that the
salt exists as the metarsenite in the official solution is not made
clear. Apparently writers differ as to the composition of the
arsenite in the solution. L. Bullock80 speaks of the “chemically
indefinite liquor arsenicalis, ” M. H. Buignet81 suggests that only
part of the arsenous acid is combined with the alkali, the remainder
being in simple solution. H. K. B amber8 2 concludes that a reac¬
tion takes place between potassium carbonate and arsenous acid in
preparing the solution, basing his conclusions on the fact that
carbon dioxide was given off during the process. This drew forth
a number of comments by Redwood, Garrod, Attfield and others as
to its correctness, the main criticism being that the concentrations
of Bamber’s experiments were not comparable to the pharma¬
copoeia! conditions. A. F. Haselden83 refers to the controversy
relating to the correctness of the title “arsenite of potash” or
80 Pharm. Jour., 10, p. 357 (1850.)
81 Jour, de Pharm. et de Chim., 59, p. 439 (1856.)
82 Pharm. Jour., 17, p. 481 (1858.)
83 Pharm. Jour., 16, p. 541 (1857.)
Lange.nhan — The Arsenical Solutions.
169
“ Liquor Arsenicalis. ’ ’ 0. Olberg84 offers a formula for the solu¬
tion in which he substitutes potassium hydroxide for the potassium
bicarbonate. In this solution he claims that the arsenite exists as
the di-potassium-mono-hydrogen-ortho-arsenite, K2HAs03. After
determining the amount of potassium arsenite formed when 5
grams each of arsenous acid and potassium carbonate were heated
in the presence of water, for varying intervals, C. Lonnes85 con¬
cludes that the title “ Liquor Kali Arsenicosi” is a misnomer, and
that “Liquor arsenicalis” is more correct. L. H. Bernegau86
states that the metarsenite is formed when arsenic trioxide and po¬
tassium bicarbonate are heated gently in the presence of water,
(presumably as in the U. S. P. process), but that the dry salt has
the formula, (according to Schmidt), of KAs02, HAs02 + H20.
L. H. Bernegau87 also reports that the dry salt always contains
some uncombined potassium carbonate. Stocker88 in his discussion
on the P. Gr. solution points out that the amount of potassium
carbonate prescribed is insufficient to saturate all of the arsenic
trioxide. J. Zieghr89 disagrees with Stocker on this point as does
Schenk.90 The Comment or zum D. A.-B. V .91 states that the
alkalinity of Fowler’s solution is due in part to the excess of potas¬
sium carbonate present. This would also tend to agree with Zieghr
towards invalidating Stocker’s claim.
In an attempt to determine whether a reaction takes place,
towards the formation of an arsenite in Fowler’s solution, the fol¬
lowing experiments were performed.
I.) The solutions resulting from the rate of solution experiments
were reserved for further observations. Twenty-four hours after
their preparation a deposit of crystals was noticed in the solution
representing the greatest concentration. These crystals were sep¬
arated from the mother liquor by means of a force filter and freed
from adhering liquid as much as possible by pressing between
bibulous paper. Both crystals and mother liquor were assayed for
84 The Apothecary, 2, p. 1 (1892.)
85 Pharm. Ztg., 39, p. 90 (1894.)
88 Am. Jour. Phar., 79, p. 553 (1907.)
87 Proc. Pa. Ph. Ass’n., 33, p. 145 (1910.)
88 Apoth. Ztg., 26, p. 335 (1911.)
89 Apoth. Ztg., 26, p. 40 (1911.)
80 Apoth. Ztg., 26, p. 379 (1911.)
91 Kommentar zum Deutchen Arzneibuch, 5 Ausgabe, 1910, p. 64 by O. Ansel-
mino and Ernst Gilg.
170 Wisconsin Academy of Sciences, Arts, and Letters.
potassium acid carbonate and arsenic trioxide, using N/l acid and
N/10 iodine respectively. The following results were obtained: —
Wt.
Crystals 7.6 gm. 67. p. c. KHC03 4.5 p. c. As203
Liquid 24.0 gm. 39.4 p. c. K2C 03 35.5 p. C. As2Q3
II.) The experiment was repeated using 200 grams of potassium
acid carbonate dissolved in 250 mils of water, and boiled for about
thirty minutes. While boiling, this solution was added to 100
grams of arsenic trioxide contained in a flask and the whole im¬
mersed in a bath of boiling water until a clear solution resulted.
The solution was allowed to stand over night when the crystals
were collected and assayed as before. A duplicate experiment was
made. The following results were obtained: —
Crystals
Liquid
Total
Crystals
Liquid
Total
81. gm. 96. p. c. KHC03 1.7 p. c. As203
359. gm.
j 3.6 p. c. KHC 03
l 22.1 p. c. K2C03
20. p. c. As203
440. gm.
100. gm. 96.09 p. c. KHC03 2.13 p. c. As203
oofi j 2.0 p. c. KHC03 26.6 p. c. As808
330. gm. | 23a £ c KjCOj*
430. gm.
III.) The results obtained so far lead to a desire for more in¬
formation and the experiment was repeated on a still larger scale.
In this case four times the original quantities were used. Instead
of allowing the solution to stand over night it was cooled and the
crystals formed at definite temperatures were removed and exam¬
ined. The first crop of crystals was collected at 30° C., the second
crop at 20° C. Then the solution was placed out doors and al¬
lowed to stand over night. The next morning the liquid in the
flask registered — 10° C. A separation between the crystals and
the liquid was effected as quickly as possible. At room temperature
the separated crystals were apparently decomposing hence were
placed in a funnel and allowed to stand twenty-four hours. By
that time the crystals had lost approximately one-half of their bulk
with the formation of a liquid. Each lot of crystals, likewise the
Langenhan — The Arsenical Solutions.
171
liquid resulting from the decomposition of the last lot was assayed.
The following results were obtained: —
105 gm. 96.09 p. c. KHC 03 1,73 p. c. Ass03
65 gm. 94.09 p. c. KHC O 3 1 .24 p. c. AssO 3
. 96.09 p. c. KHCO3 0.07 p. c. As203
j 2.00 p. c. KHCO3 12.3 p. c. As203
. I 15.2 p. c. KsC03
After the removal of the crystals at — 10° C. the mother liquor
was again placed out doors. Twenty-four hours later a second lot
of crystals was removed, the temperature of the liquid being the
same. These like the former decomposed at room temperature.
The assay results are as follows: —
— 10°C Crystals 97.09 p. c. KHC03 1.62 p. c. As203
IV.) One gram of arsenic trioxide may be expected to react with
1.01, 2.02 and 3.04 grams respectively of potassium acid carbonate
to form arsenites. The ratio of arsenic trioxide to potassium
bicarbonate in the experiments I,, II, and III, was 1 to 2. Accord¬
ing to this there would be an excess of bicarbonate in the first com¬
bination, and this excess might be represented by the crystals that
had been obtained. Hence the equivalent amount of pure92 potas¬
sium carbonate was substituted for the bicarbonate and the experi¬
ment repeated on the quarter kilo basis. The solution was allowed
to stand over night, the crystals removed and assayed in the usual
The mother liqour after standing in an open beaker for several
days deposited a second lot of crystals93 which assayed : —
Crystals 10 gm. 89.1 p. c. KHC 03 2.9 p. c. As203
92 Pure potassium bicarbonate was heated over a Bunsen flame to a constant
weight. The resulting residue assayed 99.8 p. c. K2C03.
93 In this lot the crystals were much larger and more of a cubical form.
When powdered before assaying, a very moist powder resulted.
30° crystals
20° crystals
— 10° crystals
liquid
172 Wisconsin Academy of Sciences , Arts , and Letters.
The presence of the potassium bicarbonate crystals would seem to
indicate that a reaction had taken place between the arsenic tri¬
oxide and the potassium carbonate, and that the liberated carbon
dioxide combined r ifh some of the carbonate to form the bi¬
carbonate.
V.) One hundred and twelve grams of potassium hydroxide, the
equivalent of 200 grams of potassium bicarbonate, were dissolved
in 100 mils of water and to the boiling solution 100 grams of
arsenic trioxide were added. Then the mixture was immersed in a
bath of boiling water until a clear solution resulted. After stand¬
ing several days a solid mass94 separated out. This was assayed
with the following results: —
Weight 105.0, gm. 36.0 p. c. KOH 43 p. c. Asa03
The mother liquor was allowed to stand at room temperature in
an open dish and several portions of a similar mass were collected
at various intervals. These were assayed with the following
results : —
Second mass 36.5 p. c, KOH 43.0 p. c. As20 3.
Third mass 19.6 p. e. KOH 21.67 p. c. As203
Fourth mass 19.07 p. c. KOH 19.9 p. c. As203
VI.) Fowler’s solution is alkaline to litmus95 due to the hy¬
drolysis of the potassium arsenite.96 In order to ascertain the
alkalinity of the finished product three solutions were prepared of
different composition. The first was prepared by dissolving 10
grams of arsenic trioxide and 20 grams of potassium bicarbonate
in sufficient water to make 1000 grams; the second, 10 grams of
potassium bicarbonate in 1000 grams of solution and the third by
substituting an equivalent amount of potassium hydroxide for the
bicarbonate of the first solution. These solutions were assayed for
As203 and KHC03 with the following results: —
94 The mass could not be removed from the beaker without breaking the dish
so it was dissolved in water. Upon adding the water to this mass a lowering
of temperature was noticed while solution was taking place, differing decidedly
from the action of potassium hydroxide.
05 United States Pharmacopoeia.
99 Kommentar zum Deutschen Arzneibuch , 5. Ausgabe 1910.
Langenhan — The Arsenical Solutions.
173
As the results obtained agree with the amounts used, within the
limit of error,97 one of two conclusions may be offered : viz. that no
reaction takes place between the two compounds; or that the hy¬
drolysis goes on during the titration until all of the arsenite is de¬
composed into acid and hydroxide.
VII.) The results obtained from the assay of Fowler’s solution
lead to the examination of some potassium arsenite of the market.
A sample of Merck’s pure potassium arsenite was assayed with the
following results : —
Found Computed for K AsOa + R.6As03
As203 65. p. c. equiv. 49. p. c. As. 72.75 p. c. 55. p. c. As.
KOH 45. p. c. eqniv. 31. p. c. K. 20.58 p. c. 14.3 p. c. K.
HsO . 6.62 p. c.
According to Merck’s Report this salt has the composition
KAs02 + H3As03. Structurally it may be represented by : —
/OH
As —OH
/
As \OH
VOK
VIII.) For a better understanding of the analytical data, the
details of the method employed may be desirable : —
Ten grams of crystals or liquid were dissolved in water to make
100 mils. The alkalinity of these solutions, in ten mil titrations,
was determined, first with phenolphthalein, and secondly with
methyl orange. From the number of mils of N/l acid required
with each indicator the amounts of K2C03 and KHCOs were com-
97 Ten mils instead of ten grams were assayed. Hence the results are not
exactly on a percentage basis.
174 Wisconsin Academy of Sciences , Arts , and Letters.
puted. To the titrated portions some sodium bicarbonate was
added and the amount of As203 determined by titrating with N/10
iodine, using starch T. S. as an indicator. The computations were
based on the following reactions : —
I. K8C03 4- HOH = KOH + KHC03 )
>- Phenolphthalein.
II. 2 KOH + H2S04 = K8S04 + HaO )
III. 2 KHCO3 + H8S04 = K3S04 -f 2 H20 + 2 C02 Methyl orange.
Titrating with phenolphthalein as an indicator gives one half of
the carbonate. Then adding methyl orange, the remaining half of
the carbonate (as bicarbonate) and any bicarbonate originally
present is determined. Hence the quantity of acid consumed by
the original bicarbonate is represented by the total used with
methyl orange as the indicator (after neutrality has been reached
with phenolphthalein), minus that used with the first indicator.
The following table reveals the computed percentage of the sev¬
eral components of possible arsenites.
\OK
Langenhan — -The Arsenical Solutions.
175
A$-
OH
OK
O
-OK
.OK
K3HAs205 43.05
348.228
/OK
As— OK
>
As — OK
\OK
K4 As2 O 5
386.32
38.80
/OH KH2AsO
As— OH - -
<OK 164.06
45.68
33.68 0.29 22.94
40.58 .......... 20.70
23.83 1.22 29.25
KAsO 2
As=0 — - 51.32 26.77 .......... 21.90
x-OK 146.06
/-OH K2HAsO 3
As— OK -
\OK 218.168
99.96
100.08
99.98
99.99
34.35
35.84
0.46
29.33
99.98
176
Wisconsin Academy of Sciences , Arts , and Letters.
X.) For the sake of a more ready comparison with the theoretical
computations of the previous tables, the results of the experimehts
from I to V are herewith tabulated : —
N/I H2SO4 with:-
phenolphthalein
mils
methyl orange
mils
N /10 iodine
starch T. S.
I. crystals,
liquor.. .
II. crystals,
liquor... .
crystals. .
liquor . . .
III. 1st lot
30° C. crystals. ......
20° C. crystals .
crystals,
liquid . . .
2nd ot
crystals..
liquid . . .
-10° C. |
ot
( cr
C. ]
( lie
IV. crystals _
liquor .
2nd. crystals _
3rd. crystals...
4th. crystals....
0.0
0.0
1.6
0.0
0.0
0.0
1.1
0.0
0.0
2.03
0.0
0.0
1.42
9.6
1.9
9.4
1.8
9.6
9.4
9.6
1.3
9.7
9.6
0.0
8.9
9.77
4.57
Langenhan — The Arsenical Solutions .
177
The results obtained from experiment V, viz. KOH + As203
are herewith tabulated. It must be remembered that no definite
crystals were obtained in this experiment but a noncrystalline mass.
N/l Acid. N/10 Iodine.
11.) Compound Tincture of Lavender. Fowler’s original for¬
mula contains “a half ounce”98 of Compound Spirit of Lavender.
Of this addition he says99 “The small Proportion of Compound
Spirits of Lavender is added, merely for the sake of giving a
medicinal appearance, least, from its being colorless and tasteless,
those patients who may happen to be intrusted to drop it for them¬
selves, should be tempted to use it with too great freedom ; the
consequences of which might frequently prove troublesome, if not
sometimes dangerous.” In this connection two things are note¬
worthy: firstly, that the Compound Spirit of Lavender contained
the alcoholic extractive of cinnamon, nutmeg and red saunders100
in addition to the aroma of the lavender, and secondly, that the
patented preparation of Wilson,101 after which Fowler’s prepara¬
tion was modeled,, contained red saunders as coloring matter though
it contained no aromatic.
The London Pharmacopoeia of 1809, which apparently was the
first to adopt Fowler’s solution, likewise prescribed the use of
Compound Spirit of Lavender. The authors of the U. S. P. of
1820, however, apparently regarded this addition or disguise as
unnecessary or even undesirable. Thus Bigelow102 states that
98 Fowler’s Report, p. 79 (See Appendix).
99 Ibidem, p. 82.
100 Pharmacopoea Collegii Regalis Medicorum Londinensis M. D. CCC. IX Edito
altera. Lipsiae, 1816.
101 See description of patent in appendix.
103 A treatise on Materia Medica intended as a sequel to the Pharmacopoeia of
the U. S. (1882), p. 39.
12— S. A. L.
178 Wisconsin Academy of Sciences , Arts, and Letters.
“Pure alcohol is substituted instead of Compound Tincture of
Lavender, which, from the variety of its ingredients may produce
chemical changes. ” This is also true of the New York edition of
1830. Presumably, the alcohol addition was regarded as a pre¬
servative. The Philadelphia edition of 1830, however, came back to
the original Compound Spirit of Lavender. This remained, until
in 1880 when it was replaced by Compound Tincture of Lavender.
This change is one of title only as the composition of the two was
practically the same. The editors of the United States Dispensa¬
tory103 state that “The spirit of lavender is added to give it taste,
and prevent its being mistaken for water. ’ ’
With the alkali of the arsenical solution the pigment of the red
saunders possibly combines. (See No. 3 Preservation. )
The French Codex of 1818 uses spirit of melissa as flavoring
agent, and adds no coloring matter, a precedent apparently fol¬
lowed by the German Pharmacopoeia up to this day.
12.) The volume of the finished product. According to Fowler’s
formula, the finished product was diluted to exactly one pint or, by
weight, “fifteen ounces and a half.” The U. S. P. in all its edi¬
tions from 1820 to 1870 inclusive directs dilution to one pint. In
the subsequent editions the ratio of arsenic trioxide to finished
product is that of one part by weight to 100 parts by volume, what¬
ever the amounts of the formula. The corresponding ratio of the
earlier editions is 0.84 to 100.
The amounts and ratios of the London and British pharma¬
copoeias respectively are herewith tabulated : —
103 13th edition, 1872, p. 1267.
Langenhan — The Arsenical Solutions.
179
It becomes apparent that the London Pharmacopoeia followed
Fowler’s formula and that when, in 1851, the Imperial pint of
20 fid. oz. was substituted for the wine pint of 16 lid. oz. the amounts
of arsenic trioxide and potassium carbonate were increased ac¬
cordingly leaving the percentage of arsenic trioxide the same. In
1885, however, the strength of the solution was made approximately
that of one per cent, as had been done by the U. S. Pharmacopoeia
of 1880 published in 1883. The more accurate approximation was
made in 1898. With the abandonment of the old systems in 1914,
the exact ratio of 1 to 100 was the simplest and hence the only
natural one.
This does not imply, however, that the finished product contains
1 p. c. of arsenic trioxide or its chemical equivalent, for the Arseni
Trioxidum of the U. S. P. and the Arsenous Anhydride of the B. P.
are not necessarily 100 p. c. As203. For further details on the
percentage strength of the solution see No. 14.
The “fifteen ounces and a half,” the weight equivalent of Fowl¬
er’s pint can be explained by calling attention to his statement in
his original report (p. 80) in which he points out that the weights
are expressed in Troy ounces and wt. avoirdupois “is always im¬
plied in the medical profession. ’ ’ According to the London Phar¬
macopoeia of 1836 (Phillips translation) the “wine” pint is equiva¬
lent to 7290 grains, and the Troy ounce contains 480 grains. Hence
7290 grains (the equivalent of the wine pint) are equal to 15.1
Troy ounces and not 15.5 ounces as stated by Fowler.
In 1837, Salles points out that whereas in those pharmacopoeias,
in which the same system of weights and measures is used (Apothe¬
caries weights and Wine measure) viz. “American of 1820, Ant¬
werp of 1812, Batavia (Dutch) of 1805, Belgian of 1825, Edin¬
burgh of (?),, Hanover of 1823,” etc. the strength of the finished
product is the same, in the French Codex of 1818 it is different
because the formula of the London Pharmacopoeia has been used
“more or less literally” without taking into consideration the dif¬
ferences in weights and measures. Whereas the ‘ ‘ oxide of arsenic ’ ’
content of the London Pharmacopoeia is one-one hundred and
twentieth, that of the .French Codex is one-one hundredth. In
order to correct this discrepancy, he suggests a formula, based on
that of the London Pharmacopoeia (1 : 120) expressed in grammes
or “parts by weight” so that it may be translated in any language
and system without alteration of the strength of the finished
product.
180 Wisconsin Academy of Sciences , Arts , and Letters.
13. ) Appearance of the finished product. The 1880 edition is
the first edition of the U. S. P. that directs the finished product to
be filtered after the reaction mixture has been allowed to stand
eight days. Neinhaus104 points out that this procedure was adopted
to allow the formation of the precipitate that appears in Fowler’s
Solution and the subsequent removal by filtration.
14. ) Qualitative tests. The last edition of the U. S. P., viz. that
of 1910 is the only one that gives qualitative tests. Apparently
these were regarded as necessary because of the enforcement of the
Pure Food and Drugs Act of 1906. The first paragraph of these
tests is descriptive. The faint opalescence of the solution is due
to the oils105 in the compound tincture of lavender; the pink color
is that of the Santalum Rubrum also in the tincture. The alka¬
linity is due to the excess of potassium bicarbonate105 used in the
preparation, also to the hydrolysis of the potassium arsenite.106
The second paragraph is an identity test for arsenic. Upon
acidulating the solution with hydrochloric acid and saturating with
hydrogen sulphide a yellow precipitate of arsenous sulphide is
formed. This precipitate is soluble in Ammonium Carbonate T. S.
The reaction with ammonium carbonate has been indicated as
follows :105
2AsgS3 F 2(NH4)2C Os = 2C 02 +NH4As02 + 3NH4AsS2
Structurally, the formulas involved may be represented in the
following manner :
As — S
/ONH4
F c=0 IFF c F As F As
As=S \()NH4 FF) \()NI14 \SNH4
With Silver Nitrate T. S. the acidulated solution forms a yellow
precipitate of silver arsenite. This has been expressed by the fol¬
lowing equation :105
KAsO 2 F HgO F 3AgNOs = 2HNO^F KNOs F As(OAg)3
As /TI /OA g /OH /OK /OA g
\OK F o F N=0 IFF N=0 F N=0 F As— OAg
\H \() \0 Ag
104 Pharm. Z tg., 37, p. 331 (1892.)
105 Commentar zur Pharmacopoeia Helvatica (1896), p. 196.
1116 Kommentar zum Deutschen Arzneibuch, vol. 2, p. 64. (1910.)
Langenhan — The Arsenical Solutions.
181
The third paragraph is a test for the absence or presence of
arsenate. In the presence of hydrochloric acid the precipitate of
silver arsenate does not form due to its solubility in the acid solu¬
tion. If to the acidulated solution then, some Silver nitrate T. S.
is added and a layer of ammonia water superimposed upon it a
reddish brown color at the line of contact indicates the presence of
silver arsenate. The reaction has been expressed in the following
manner :105
/OK /-OAg-
0= As —OK + 3 AgN03 =B 3KN03 + 0= As — O Ag
\OK \OAg
Supplement to Tests. According to the U. S. P. “An aqueous
solution of Arsenic Trioxide is faintly acid to litmus. ’ ’ Whether
this same acidity is shown towards other indicators is not stated.
Hence a number of indicators were tested according to the following
procedure :■ —
Aqueous solutions of arsenic trioxide and potassium hydroxide
were prepared, their strength being represented by the following
formulas :
No. 1, As(0H)3; No. 2, KOH; No. 3, 2K0H; No. 4, 3K0H;
5 5 5 5
respectively. Six test tubes of the same size were used for each
series. Into these test tubes were placed the following solutions : —
It will be noted that each solution as contained in the tubes was
N/10 in strength, comparable in strength to the arsenical solutions.
182 Wisconsin Academy of Sciences, Arts, and Letters.
The indicators used were of U. S. P. strength. Those not offi¬
cial were prepared according to a table given by Clark and Lub.107
The following table represents the results obtained.108
A survey of the table reveals that only six out of twenty indica¬
tors show an acid reaction towards an aqueous solution of arsenic
trioxide. Three of these viz. litmus, turmeric and hematoxylin are
plant pigments, the composition of which is not well understood.
The other three phenol red and cresol red are phenol-sulphone-
phthalein, and ortho-cresol-sulphone-phthalein respectively, and
phenolphthalein.
A peculiar change was noticed in the two series of solutions con-
107 Journal of Bacteriology, vol. 2, p. 1 (1917.)
108 Tube No. 6 (H20) was taken as a standard, i. e. it was considered neutral,
and any other tube showing same color was considered neutral.
109 Phenolphthalein naturally does not reveal the acid reaction of the As2Os
solution.
Langenhai i — -The Arsenical Solutions.
183
taining iodeosin and eosin, the former being tetra-iodo-fluorescein,
and the latter a tetra-brom-fluorescein. Tubes 2, 3, and 4 of these
series developed an intense yellow fluorescene when viewed by re¬
flected light, which was not evident in the remaining tubes.
A third series worth mentioning is the one that contains alkali
blue. In this series tubes 3 and 5 have the same color which dif¬
fers from that of any of the remaining four. These tubes viz. 3
and 5, contained
As(OH)3 + 2KOH , and KOH
5 5 5
respectively. Whether one may assume that one molecule of potas¬
sium hydroxide is uncombined in each tube, leaving one molecule
combined with the As (OH) 3 in the first tube is possibly open to
question. Yet the sameness of color of these two tubes was so
striking that the fact seemed worth mentioning.
15.) Assay. From 1880 on the U. S. P. gives directions for the
assay of Fowler’s Solution. The quantitative determination of
trivalent arsenic as As203 is not only a check on the amount of
arsenic trioxide employed, but also on the possible deterioration
of the solution, brought about by the oxidation of the arsenite to
arsenate, in the presence of water, as expressed by the following
reactions : —
H20 + I2 =2HI+ O
As303 + 20 = As20 5 , or
AsO.OK + O = AsO aOK , or
As(OH)2OK + O = AsO(OH)2OK, hence
1 4 equiv. 02 equiv. Ass03 ,
126.92 equiv. 197.92 Qr
0.012692 equiv. 0.004948
The amount of As203 as determined by the prescribed volumetric
assay varies to a slight degree. The U. S. P. of 1880, the first to
introduce an assay method, states that 24.7 gm. of the solution
“should require from 48.5 to 50 cc. of volumetric solution of
iodine” — (“corresponding to 1 per cent, of arsenious acid of the
required purity”.) The As203 equivalents of 48.5 and 50 cc. re¬
spectively of N/10 I. Y. S. are 0.9709 gm. and 1.001 gm. As the
purity rubric for “arsenious acid” was “at least 97 per cent.
184 Wisconsin Academy of Sciences , Arts , and Letters.
pure”, ten grams, the amount directed to be used to make 1000 cc.
of the Liquor, would represent 0.9700 gm. As203 to 100 cc. of the
solution. Hence, unless the “arsenious acid” used assayed slightly
higher than the minimum U. S. P. requirement, the finished prod¬
uct would fall short by a very slight amount (within the limits of
experimental error) of the given standard.
The U. S. P. of 1890 states that 24.7 cc. of the solution “ should
require from 49.4 to 50. cc. of decinormal iodine Y. S.” — “ (cor¬
responding to 1 gm. of arsenous acid in 100 cc. of the solution) ”.
The As203 equivalents of 49.4 cc. and 50. cc. N/10 I. Y. S. ac¬
cording to this revision represent 0.9883 gm. and 1.004 gm. respec¬
tively of As203 in 100 cc. of the solution. As the purity rubric
for “ arsenous acid” in this revision is “at least 98.8 percent of
arsenic trioxide”, ten grams, the amount prescribed to make 1000
cc. of the Liquor, would represent 0.988 gm. to 100 cc. of the solu¬
tion. In this case the minimum requirement would be met with.
The U. S. P. of 1900 states that 24.6 gm. of the solution “should
require not less than 50 cc. of tenth-normal iodine Y. S. — ” “ (cor¬
responding to 1 gm. of arsenic trioxide in 100 gm. of the solution) ”.
The As203 equivalent of 50 cc. N/100 I. Y. S. according to this re¬
vision represents 0.9981 gm. of As203 in 100 gm. of the solution.
The new purity rubric for arsenic trioxide is “not less than 99.8
per cent, of pure Arsenic TrioxideY Hence ten grams of arsenic
trioxide, as directed to be used for the preparation of 1000 grams
of the Liquor, would represent only 0.998 gm. of As203 to 100 gm.
of the solution. In this instance the conditions meet the require¬
ment of 10 gm. of arsenic trioxide, which represents 9.98 gm. of
As203.
The U. S. P. of 1910 merely demands a definite percentage
strength, viz. “not less than 0.975 nor more than 1.025 gm. As203
in 100 gm. of the solution, ’ ’ and makes no mention of the amounts
of N/10 I. Y. S. to be used. The purity rubric for arsenic trioxide
for this revision is “not less than 99.8 per cent, of As203.” It is
quite evident however, that the maximum limit of As203 content
of the solution will probably not result when the prescribed ten
grams of arsenic trioxide are used, as this represents only 0.998 gm.
of As203 to 100 gm. of the solution. The lower limit of the rubric
viz. 0.975 gm. to 100 gm. of solution, is undoubtedly to allow for a
slight loss of arsenic trioxide, as determined by the assay process,
due to the oxidation of the trivalent arsenic to the pentavalent ar¬
senic.
Langenhan — The Arsenical Solutions. 185
Two factors influence the variations mentioned above: firstly,
the variation in the As203 equivalent of N/10 I. V. S. due to
changes in atomic weights, and secondly, the purity rubric of the
arsenic trioxide. The Cc. equivalent for 1880 was 0.004945; for
1890, 0.004942 ; for 1900, 0.004911 ; and for 1910 it is 0.004948. The
purity rubric for arsenic trioxide for 1880 was 97 p. c. As203; for
1890, 98.8 p. c. As203 ; for 1900, 99.8 p. c. As203 and for 1910 it is
99.8 p. c. As203.
In as much as only one gram of arsenic trioxide is used to pre¬
pare 100 grams of the solution, it becomes quite apparent that any
solution assaying 1 p. c. As203, (the generally accepted U. S. P.
requirement) would result only from carelessness in weighing, un¬
less it be that the U. S. P. formula were ignord and the equivalent
of 10 gm. of 100 p. c. pure arsenic trioxide were used, or that a 100
p. c. arsenic trioxide were used.
16.) Bose. The 1900 revision of the U. S. P. is the first one to
include the dose. This is given as “ average dose — 0.2 C. (3
minims)”. The 1910 revision likewise contains the dose given as
“ average dose — Metric, 0.2 mils — Apothecaries, 3 minims”.
“ Doses were introduced into the Pharmacopoeia to meet a popu¬
lar demand — mainly to add to the value of the book to the young
physician, but also to serve as an authoritative statement for the
benefit of the dispenser, of the ordinary practice of the physicians
in prescribing different remedies”.110 The subject of the introduc¬
tion of doses into the Pharmacopoeia evidently called forth some
little discussion, as to whether a maximum, a minimum, or an aver¬
age dose should be designated. The objection to establishing a
definite dose was the possibility of legal difficulties if this dose were
overprescribed.* 111 In reviewing the method in vogue for desig¬
nating doses in foreign pharmacopoeias, the question of overpre¬
scribing and thereby ignoring the standard has apparently been
overlooked, or considered unimportant. Or the revisors of the
pharmacopoeias had not unlimited confidence in the prescribers
knowledge of drugs and medicines, as to place no restrictions what¬
soever on the quantity he may prescribe, for out of ten of the more
recent revisions of foreign pharmacopoeias consulted, seven give a
maximum dose. Three give no dose, but state that the solution
must be dispensed with caution (or its equivalent). Not only is
110 Lyons, Cir. of the Comm, of Revision of U. S. P. VIII, Cir. 447, p. 1934.
111 Remington, Ibidem, Cir. 447, p. 1821.
186 Wisconsin Academy of Sciences, Arts , and Letters.
the maximum single dose given, but the maximum amount to be
given “per diem” is prescribed. “Overprescribing the maximum
amount in most of the European countries is indicated by having
the physician place an exclamation mark ( ! ) after such a pre¬
scribed dose. This reveals to the pharmacist that the prescriber
wants the effect of something more than the ‘ average ’ maximum
dose; this cautions him in his dispensing, and relieves him of the
responsibility of ‘too large a dose’.112”
The following table gives the dose of the European pharmaco¬
poeias consulted :
Bose
1910 German . max, single, 0.5 g. max. daily, 1.5 g.
1909 Italian. ............... . max. single, 0.5 max. daily, 1.5
1905 Holland. ............... max. single, 500 mg. max. daily, 1.5 gm.
1907 Switzerland. .......... . max. single, 0.5 max. daily, 1.5
1913 Norwegian. ... - - .... max. single, 0.5 gm. max. daily, 2.0gm.
1905 Spanish. . . max. single, 0.6
1914 British . . max. single, 0.5 centimils.
1910 U. S. . . . average 0.2 mils
1906 Belgian. ............... no dose given,
1908 French ................ no dose given,
1908 Swedish.. . . no dose given,
112 Kremers, Ibidem, Cir. 500, p. 2048.
Langenhan — The Arsenical Solutions.
187
APPENDIX
English Patents. (Patent Medicines.) “In the early days of
English commerce, monopolies were granted by the sovereigns at
their own pleasure, and often for their personal profit. Queen
Elizabeth so largely abused her power in this direction that to¬
wards the end of her reign the discontent of her subjects compelled
her to promise she would offend no more: and her successor, James
I, gave a similar undertaking. The abuse, however, was continued
until the Statute of Monopolies, passed in 1624, regulated all such
grants, placing the power in the hands of Parliament, and limit¬
ing the period of privilege to fourteen years.
For the first century or thereabouts of the administration of this
Act, specifications of processes or formulas were not a condition
of the patent. The idea was the introduction into the country of
new industries, and it was supposed that artificers who would have
to be employed in any such industries would certainly acquire such
necessary skill and knowledge about any new manufacture as would
prevent any perpetuation of the monopoly. It was during the
reign of Queen Anne that the law officers began to require that
specifications should be filed before letters patent were issued. But
the condition was not by any means uniformly or intelligently in¬
sisted upon, as will be seen immediately in the case of certain pat¬
ent medicines.
The term ‘patent medicines’ as now popularly used, means gen¬
erally secret medicines, and the meaning is therefore in exact con¬
tradiction to the expression. Truthfully to declare the composi¬
tion of many of these proprietary compounds would ruin their
sale. Not that the ingredients are often improper or injurious;
this rarely occurs; but because the success of these remedies in
most instances is rather on the mystery with which the maker can
surround them than on their exceptional merit.
But some old medicines which became popular, including a few
the reputation of which lives today, were actually patented. The
first compound medicine for which a patent was granted under the
Act of 1624 was No. 888, and was dated October 22, 1711. It was
granted to Timothy Byfield for his sal oleosum volatile, ‘which
188 Wisconsin Academy of Sciences , Arts, and Letters.
by abundant experience hath been found very helpful and beneficial
as well in uses medicinal as others,’ No particulars of the in¬
gredients or method of manufacture are given.” (Wootton,
Chronicles of Pharmacy, Yol. II, pp. 161-2.)
Medical Compound.
Wilson’s Specification.
To all to Whom these presents shall come, I, Thomas Wilson, of
Snow Hill, in the Parish of Saint Sepulchre’s, in the City of
London, Chymist, send greeting.
Whereas His most Excellent Majesty King George the Third,
by his Letters Patent under the Great Seal of Great Britain, bear¬
ing date at Westminister, the Sixteenth day of February, in the
twenty-first year of his reign, did give and grant unto me, the said
Thomas Wilson, His especial licence that I, the said Thomas Wilson,
during the term of years therein expressed, should and lawfully
might use, exercise, and vend, within England, Wales, and Town of
Berwick-upon-Tweed, my invention of “A Medicinal Composition,
which after much Experience hath beert found to be an Infallible
Remedy for Agues and Intermitting Fevers, even in the most ob¬
stinate Cases where the Bark and every other Medicine hath proven
ineffectual;” in which said Letters Patent there is contained a
provisoe obliging me, the said Thomas Wilson, under my hand and
seal, to cause a particular description of the nature of the said In¬
vention to be inrolled in His Majesty’s High Court of Chancery
within four calendar months after date of the said recited Letters
Patent, as in and by the same (relation being thereunto had) may
more fully and at large appear.
Now Knowe Ye, that in compliance with the said provisoe, I,
the said Thomas Wilson, do hereby declare that my said Invention
of a Medical Composition, which, after much Experience hath been
found to be an Infallible Remedy for Agues and Intermitting
Fevers, is described in the Manner following, (that is to say) : —
Take of centarium minus, or common centaury, any quantity,
burn it to ashes; take of these ashes, boil them in water for three
hours, evaporate the liquid to dryness; take this mass, calcine it
for four or five hours, keeping it constantly stirring. Take cob-
Langenhan — The Arsenical Solutions .
189
bait,113 powder it fine, put it in a crucible ; sublime the flowers ; take
these flowers, add to the above, melt them together and boil them
for two hours in water ; then take of the santalum rubrum, or red
sanders, boil it in water for four hours ; mix all together.
In witness whereof, I, the said Thomas Wilson, have hereunto
set my hand and seal, this Eleventh day of June,. 1781.
Thomas Wilson. (L. S.)
And Be it Remembered, that on the same Eleventh day of June, in
the year above mentioned, the aforesaid Thomas Wilson came be¬
fore our said Lord the King in His Chancery and acknowledged
the Specification aforesaid, and all and every thing therein con¬
tained and specified, in form above written. And also the Specifica¬
tion aforesaid was stampt according to the tenor of the Statute
made in the sixth year of the reign of the Late King and Queen,
William and Mary of England, and so forth.
Inrolled the same day and year above mentioned.
Medical Reports of the Effects of Arsenic in the Cure of
Agues, Remitting Fevers, and Periodic Headaches.
by
Thomas Fowler M. D.
Physician to the General Infirmary of the County of Stafford.
Together with a Letter from Dr. Arnold, of Leicester, and Another from
Dr. Withering, describing their Experience of the Effects
of Arsenic in the Cure of Intermittents.
London, MDCCL XXXVI.
(Title page of report.)
Preface, p. ii.
113 “This kind of pyrites miners call cobaltum , if it is allowed to me to use our
German name. The Greek call it cadmia. The juices,, however, out of which
pyrites and silver are formed, appear to slifify into one body, and thus is pro¬
duced what they call cobaltum . There are some who consider this the same
as pyrites, because it is almost the same. There are some who distinguish it
as species, which please me, for it has the distinctive property of being ex¬
tremely corrosive, so that it consumes the hands and feet of the workmen, un¬
less they are well protected, which I do not believe that pyrites can do.”
(Bermannus, p. 459.) Hoover’s Engl, transl. of Agricola, De re metallica,
p. 113. Hoover adds: “It is desirable to bear in mind that the mines familiar to
Agricola abounded in complex mixtures of cobalt, nickel, arsenic, bismuth, zinc,
and antimony. Agricola frequently mentions the garlic odour from cadmia
metallica which, together with the corrosive qualities mentioned below (above)
would obviously be due to arsenic.”
190 Wisconsin Academy of Sciences , Arts , and Letters.
The Medical Reports,, which the Author lately published, relative
to the Effects of Tobacco, in the Cure of Dropsies and Dysuries,
having been favourably received by the Public; and the general
diuretic Property of that Medicine, confirmed by furthur Experi¬
ence; he is encouraged to pursue the same Plan, with Regard to
another powerful Article of the Materia Medica.
Arsenic is a Mineral which has long been reputed one of the
most violent Poisons hitherto known; and accordingly has been
reprobated in the strongest Terms by almost every medical Writer,
that has ever deigned to notice it ; and yet there is a good Reason
to believe it (p. Ill) bids fair to hold a Place, among the best and
most valuable Medicines ; and to rank with peruvian Bark in the
Cure of Agues, remitting Fevers, and periodic Headachs.
Efficacious Medicines are certainly of the utmost Importance in
Practice, the Investigation of which, has been frequently recom¬
mended by many celebrated Philosophers and Physicians, wTho
have done Honour to Science amongst the Ancients; and by some
Authors of the highest Reputation amongst the Moderns, particu¬
larly Bacon, Boyle, Baglivi, and Boerhaave. The present Materia
Medica, however, notwithstanding the many Volumes that have been
written upon the Subject, makes but a very humble Appearance
with Regard to Medicines of real Efficiency and Importance; and
therefore, if by an experimental Enquiry, another efficacious Medi¬
cine is likely to be added to the Number of the few valuable Ones
already established, it is to be hoped, that such an Attempt will
not be deemed unworthy of Attention.
(p. IV.) Perhaps by some it may be alleged, that the Ague is a
very common and insignificant Disease, because frequently cured
both by Art and Nature, and that consequently there is no Occa¬
sion for the Investigation of a new Medicine on that Account.
But certainly it is a Disease of much Importance; for whenever
it continues long, Coldness, Paleness, and Debility are its constant
Concomitants, and evince it to be the fertile Parent of Cachexy;
and when it occurs in old Age, or in Constitutions broken down
by Intemperance, or some previous Disease, how often do we find
it accelerate the Accession of Jaundice, Consumption, Dropsy, or
some other Mischief of fatal Consequence.
(p. V.) Fatal however as the Disease has been in its Conse¬
quences, it is well known to practitioners that there are many Cases
wherein not only the minor Remedies have proved ineffectual, but
Langenkan — The Arsenical Solutions.
191
even the Bark itself has failed, or otherwise its Administration not
been admissible; and therefore a powerful vacarious Remedy for
so frequent a Distemper, becomes an important Object.
(p. VI) It may also be objected, that to introduce a Poison to
the Public as a general Medicine, is to put a two-edged Sword into
the Hands of the Ignorant. But the same Argument will equally
apply against several of the most efficacious Medicines in daily Use,
in particular Opium, and some of the more active preparations of
Mercury and Antimony.
Or if for a Moment, the Force of this Argument be admitted, yet
when it is considered that a Patent Medicine has, for some years
past, been largely circulated through the Kingdom, as a specific for
Agues, under the Title of Tasteless Ague and Fever Drops; that
from its Effects in Practice there is the strongest Reason to believe it
to be an arsenical preparation ; and that on this very Ground, it is
at present imitated in different Parts of the Kingdom, the Objec¬
tion becomes nugatory.
Therefore,, if any new and important Medicine, either from its
active Nature, or an imprudent Administration thereof, is found to
be sometimes connected with distressing and deleterious Effects, it
becomes highly requisite (p. VII) that the Public should speedily
be made acquainted with such Effects, as far as they are known ; to¬
gether with such Precepts, Cautions, and Restrictions, as may tend
to unite the greatest Degree of Safety with its Efficacy. Nothing
however, of this Sort, has been done; which Consideration, it is
presumed, will be an Apology for an early Publication of so inter¬
esting a Subject, and for such Imperfections as may have taken
Place from that Circumstance.
When the Patent Ague Drops began to acquire some Reputation
in the Country, they were occasionally adopted in the Hospital
Practice of this Place, in 1781, 1782, and 1783, and were found
efficacious.
In the beginning of October 1783, Mr. Hughes, the Apothecary to
the Infirmary, (whose Industry, Attention and Abilities in his pro¬
fessional Line, justly merit Applause) informed the Author that
he had tried to imitate the Ague Drops, and, from a number of
Experiments, had so far succeeded (his Medicine having produced
similar Effects) that he was convinced they were a Preparation of
Arsenic.
(p. VIII) In Consequence of this general Intimation the Author
192 Wisconsin Academy of Sciences, Arts, and Letters.
of these Sheets consulted Lewis’s Materia Medica upon arsenic,114
and agreeable to his pharmaceutic History of that Article, he dis¬
solved a small Portion of the white Sort in a solution of fixed vege¬
table Alkali.115 On trying and comparing the Effects of this Solu¬
tion, with those of the Patent Ague Drops, he found the Medicine
to be similar, but the former was too strong; he therefore diluted
the Solution, by doubling the Proportion of Water, and then com¬
paring their Effects, found the Medicines nearly of an equal
Strength.
Having thus become acquainted with what he esteemed a powerful
Medicine, he was strongly impressed with an Idea of its Importance ;
and therefore has kept a regular and connected History of the Cases
in which he has prescribed it. Hence the following Reports con¬
tain an Account of the curative Effects of the Medicine, described
in a just Proportion of Examples of Successes and Failure, in the
Course of its Administration. They also contain a particular
(p. IX) Account of its operative Effects, which being sometimes
troublesome, both in Degree and Number, will be found to merit a
serious Attention. They are likewise divided into Sections, in order
that Cases of similar Nature, Treatment or Event; and also the
Observations more immediately appertaining to each particular
Part of the Subject, might appear in a more clear and practical
Point of View.
The cases drawn out at large are but few, and those which are
abstracted are not so numerous as Some might think necessary,
considering the Number which the Writer has collected. But cer¬
tainly it is sufficient for an Author, to deliver, in as concise a man¬
ner as possible, such a View of the Result of his Experiments, with
the Mode if conducting them,, as may be clearly understood and
readily applied to Practice. Beyond this Point, a Multiplicity of
Cases and Arguments is of no Consequence ; because every Publica¬
tion which appeals to the Reality of Facts for its Importance, will
soon have its Propositions either confirmed or rejected, by the
daily Experience of the Public.
(p. X) The Author professes himself a strong Advocate for an
effective and powerful Materia Medica ; on which Account, the Sub¬
jects of his late and present Reports have engaged a Share of his
114 For complete title see p. 148.
115 Of “white arsenic” Lewis (Mat. Med., p. 101) states that it is “soluble
plentifully in alkaline lixivia.” See also statement in Wilson’s patent on p. 188.
Langenhan — The Arsenical Solutions.
193
most serious Attention. The Properties however of mild and
simple Medicines, have occasionally been the Objects of his In¬
vestigation; and his Enquiries in that Line have convinced him,
that among the many trifling and insignificant Articles, which are
still retained in the Materia Medica, there are some valuable Medi¬
cines, that are not in the Reputation they are justly entitled to ;
a few Examples of which, it will give him Pleasure to communicate
in some future Publication. In the meantime, as the Idea of a
Poison seems to be so strongly connected with that of Arsenic, it
will be found very difficult to separate them in the Mind, whenever
that Term is named; and therefore to avoid as much as possible,
such a disagreeable Association of Ideas, in the Practice of the
healing Art, the Medicine now about to be introduced to the Notice
of the Public,, will be distinguished by the Name of Mineral
Solution.
Advertisement, (p. XI)
All Cases beginning with the Words In-Patient, and Out-Patient,
belong to the Infirmary ; the Rest were chiefly such as the Author
gave Advice to at his own House.
When the Mineral Solution is ordered thrice a Day, six o’clock
in the Morning, two in the Afternoon, and ten o’clock in the
Evening ; and when twice a Day, ten in the Morning, and ten in the
Evening, are to be considered as the medicinal Hours, or Periods
for its Administration.
Unless the Solution be ordered in some particular Vehicle it is
always to be understood, that Adults are to take in a Tea cupful of
Water, and the Doses for Children are to be given in the same
Vehicle, proportionately diminished.
Every Medicine prescribed by a general title, without a Refer¬
ence, is to be found in the London Dispensatory.
Contents, (p. 1.)
Sect. I.
(pp. 1-40)
Sect. II.
(pp. 41-61)
Sect. III.
(pp. 62-64)
Cases of Agues, cured by the Mineral Solution,
with Observations.
Cases of Agues, cured by the Assistance of the
Peruvian Bark, after Failure from the Mineral
Solution, with observations.
Cases of Agues, suspended, releived and not re-
leived, by the Mineral Solution, from the irregular
Attendance of Patients, with Observations.
13— S. A. L.
194 Wisconsin Academy of Sciences , Arts , and Letters.
Sect. IY.
(pp. 65-74)
Sect. Y.
(pp. 75-78)
Sect. YI.
(pp. 79-83)
Sect. VII.
(pp. 84-95)
Sect. VIII.
(pp. 96-102)
Sect. IX.
(pp. 103-108)
Sect. X.
(pp. 108-116)
Cases of Remitting Fevers, treated by the Mineral
Solutions.
Cases of Periodic Headaches,, treated by the Min¬
eral Solution, with Observations.
Formulae Medicamentorum, with Observations.
Observations on the Doses and Mode of administer¬
ing the Mineral Solution, in the Cure of Agues,
interspersed with Practical Rules and Cautions.
Observations on the operative Effects of the Min¬
eral Solution.
Observations on the curative Effects of the Mineral
Solution.
General Observations, and practical Conclusions.
Sect. YI p. 79.
Formulae Medicamentorum :
with Observations.
Solutio Mineralis. (No. 1.)
Recipe Arsenici albi in Pulverem substillissimum triti;
Salis alkalini fixi vegetabilis purificati; singulorum Grana
Sexaginto quatuor,
Aquae fontanae distillatae Libram dimidiam.
Immittantur in Ampullam florentinum, qua in Balneo Arenae
posita, Aqua lente ebulliat, donee Arsenicum perfeete solutum
fuerit. Deinde Solutioni frigidae adde
Spiritus Lavendulae compositi
Unciam dimidiam,
Aquae fontanae distillatae Libram dimidiam, plus velminus, adeo
(p. 80) ut Solutionis mensura Libra una accurate, fit, vel potius
Pondere# quindecim cum dimidia.
# I have said, vel potius Pondere, because weight is a much more
accurate Method of ascertaining Proportions than Measure, in pre¬
paring Medicines of great Activity, of which Sort the Mineral Solu¬
tion is a strong Example. It is perhaps superfluous to remind the
Reader, that Troy Weight (which differs materially from Avoirdu¬
pois) is always implied in medicinal Professions.
Langenhan — The Arsenical Solutions.
195
Observations p. 81.
The first medicine is a chemical Preparation as simple as conld
be wished for, and if proper Care be taken that the Solution of the
Mineral is complete, it will be found of a certain and uniform
Degree of Strength. Its being also (p. 82) in a liquid Form,
renders it convenient, for administration by Drops, and thereby
nicely varying the Doses, according to the Exigencies of the Case ;
a matter of some Importance, where the Activity of the Medicine
is so great.
The small Proportion of Compound Spirits of Lavender is added,
merely for the Sake of giving a medicinal Appearance, lest, from
its being colourless and tasteless, those Patients who may happen
to be intrusted to drop it for themselves, should be tempted to use
it with too great Freedom; the Consequences of which might fre¬
quently prove troublesome, if not sometimes dangerous.
To a Pint of the Solution, sixty-four Grains were added, for the
purpose of a more ready Calculation; therefore one Ounce of the
Solution contains exactly four Grains of the Mineral and conse¬
quently each Dram, just half a Grain; I have found by repeated
Trials of dropping the Solution, from a two Ounce Vial (with a
broad Margin) little more than half full, that each Dram, by a two
Ounce graduated Glass Measure, contains about eighty Drops.
We ought to be well assured, that the fixed vegetable Alkali we
make Use of is perfectly pure; for that which is bought of the
Druggists is frequently so impure, as to be inadequate for the
Purpose of producing a perfect Solution of the Mineral; a Cir¬
cumstance which would occasion great Confusion and Uncertainty,
in the Doses (p. 83) of the Medicine, if therefore any One desirous
of preparing the Mineral Solution, should not be provided with the
pure alkali ; and would not choose to be at the Trouble of purifying
it, a double Proportion of purified Nitre (which is a preparation of
some certainty, and supposed to be always at Hand), may be sub¬
stituted; for the Arsenic having the Property of discharging the
nitrous acid, will unite with the Alkali.
The Formula which I have given with the Alkali, contains my
own preparation, alluded to in the Preface, and is that on which
my experience is chiefly founded.
Mr. Hughes has likewise used it prepared with Nitre, and on
trial found it equally efficacious. I have also used it a Number of
196 Wisconsin Academy of Sciences , Arts, and Letters.
Times, and found it successful, and believe the two Solutions to be
the same Medicine, provided they are accurately prepared with
either of the purified Salts.
A Letter from Dr. Arnold, Physician to the General Infirmary , and to the
Dispensary, \ at Leicester, (p. 117).
Dear Sir:
I was pleased to learn, by your last, that your intended publication rela¬
tive to the cure of intermittents by means of an arsenical Preparation,
is in such forwardness; and shall willingly contribute my assistance, to
establish the credit of a remedy of which I am led by experience to form
a very favourable opinion. With this in view you at length receive a
short sketch of my observations, on the use of Arsenic in the cure of
intermittents; which I flatter myself will be found to correspond in a
great measure with your own, and those of our friend, Doctor Wither¬
ing ; who first communicated to me the uses of this new Medicine, and
the method of preparing it.
I would most readily have sent you a complete summary of my prac¬
tice in this respect; in which I would have left it to facts to speak for
themselves; but am obliged to decline an undertaking, which would re¬
quire more time than is compatible with your haste, and my present
leisure.
(p. 118) The medicine I have made use of is a solution of the common
arsenicum album of the shops; of which the following is the formula.
Recipe Arsenici albi pulveris subtilissimi grana octo,
Aquae distillatae uncies octo.
Digire in blaneo mariae donee penitus solvatur arsenicum. Caveas
autem ne aquae vel guttula evaporetur; quae non statim, solution! Anita,
reponatur. Per chartam deinde cola.
Dosis a guttis quindecim ad quadraginta, ter in die, inter febrium in-
termittentium paroxysmos, in menthae vulgaris infusi uncies duabus.
(Note) The remainder of the letter refers to the dosage, symptoms and
diagnosis, of cases observed).
(Signed) (p. 123) Your faithful fried and fervent;
Thomas Arnold.
West-Cotes
March 27th, 1786.
A Letter from W. Withering. M. D. F. R. 8. (p. 12i3)
Birmingham, May 21st. 1786.
Dear Sir:
The Arsenical Solution was first used here in the Autumn of the year
1783, at which time Intermittents of various denominations were very
prevalent both in this and in adjourning Counties. The general use of
Tasteless Ague and Fever Drops of this period made me solicitous to know
the composition of that medicine, and I was informed it was made from
the Ore of Coblat; but as it did. not answer the usual tests for that Semi-
Lcmgenhan — The Arsenical Solutions.
197
metal, I thought it was probably Arsenic, which is known to abound
very generally in coblat Ores.* Whilst I was intending to submit the
drops to a more (p. 124) effectual examination, my good friend and your
excellent Apothecary Mr. Hughes informed me, that he had made an
analysis of the drops, and found them to be a solution of Arsenic.
At first we gave the medicine cautiously,.... (This part of the letter
deals with the dose and treatment of patients) . In the Autumn of
1784 it was almost constantly prescribed, and has ever since maintained
its credit with us under a very great number of trials.
At first we used a simple solution of Arsenicum album, one grain to
each Ounce of distilled Water. The solution is facilitated by boiling for
a minute to two in a florence Flask, or other glass Vessel, but long con¬
tinued boiling disposes it to precipitate again. After some time Mr.
Baley (p. 125) added Nitre to the solution, by the direction of Mr.
Hughes, to prevent the precipitation, which in the rapid consumption in
the Hospital had hardly been observed to take place, but in the smaller
demands of private practice seldom fails to happen. I should also men¬
tion that an ounce of alcohol has lately been added to twenty Ounces of
the Solution, but I am doubtful of the propriety of this addition.
(The. remainder of the letter deals with the dose and method of admin¬
istration)
(Signed) (p. 127) Your obliged and faithful friend
W. Withering.
* See footnote to Thomas Wilson’s patent, p. 189 for chemical explanation.
THE DEVELOPMENT OF THE FRENULUM OF THE WAX
MOTH, GALLERIA MELLONELLA LINN.
WILLIAM S. MARSHALL.
The following quotation is taken from Comstock’s (6) recent
book on the wings of insects :
“In most moths there is a strong spine-like organ or bunch of
bristles borne by the hind wing at the humeral angle: this is the
frenulum or little bridle. As a rule the frenulum of the female con¬
sists of several bristles; that of the male, of a single strong, spine-like
organ. In the males of certain moths, where the frenulum is highly
developed, there is a membranous fold on the fore wing for receiving
the end of the frenulum thus more securely fastening the two wings to¬
gether: this is the frenulum hook.”
Griffith (9) has given an account of the frenulum in the Lepi-
doptera, and the following historical remarks are in part ex¬
tracted from his paper. De Geer (7) in 1752 was the first to
describe the frenulum; he records finding it in all the species of
noctural Lepidoptera he examined. He failed to notice its pres¬
ence in the females, and, although he recognized its ability to
hold the fore and hind wings together, he was uncertain as to
its use. Harris (10) in 1767 described and figured the frenulum;
he observed its presence in the males only, but he found that the
female moths had four small bristles instead. Giorna (8) listed
seventy species of moths in which the males possess a frenulum;
he called attention to a corresponding structure present in the
females of many species of the moths he examined. References
to the frenulum are found in Burmeister (2), Westwood (15),
Poey (13), etc., etc. The last mentioned entomologist found that
the frenulum is simple in the male specimens but multiple in the
female. Chenu (3) held that the frenulum is used to hold the
wings in a horizontal position during repose.
199
200 Wisconsin Academy of Sciences , Arts , and Letters.
The following quotations are relative to the frenulum. Kirby
and Spence (11), Letter XXIII:
“The wings of many butterflies, hawk-moths, and moths are dis¬
tinguished by a remarkable apparatus, noticed by de Geer, and since
by many other naturalists, for keeping them steady and underanged
in their flight.”
These authors also noticed the difference in the frenulum be¬
tween the two sexes :
“The females, which seldom fly far, often have the bristles, but
never the hook.”
Comstock (4) has the following:
“Except in the Microlepidoptera the frenulum of the male consists
of a single strong spine, that of the female of two or more bristles.”
From another account by Comstock (5) ;
“If one of the bristles of the compound frenulum of the female be
examined it is found to be hollow, containing a single cavity. But
when the frenulum of a male is examined it is found to contain sev¬
eral small cavities. Evidently the frenulum of the male is composed
of several bristles as is that of the female, but these bristles are
grown together, forming a single spine.”
One of the latest accounts of the frenulum is by Tillyard (14).
In this work the wing-coupling apparatus of the Lepidoptera and
of other orders of insects is discussed.
The frenulum of the wax moth appears as a slightly curved
bristle on the costal margin of each hind wing near the humeral
angle. It is borne on the humeral lobe. In the male moth a
single bristle is present on each wing; in the female there are
generally three. There is a variation in this number in the fe¬
male (this will be discussed later), but at present we will assume
that three bristles, the most regular number, are present.
In studying the development of the frenulum, one is early im¬
pressed with the fact that it is similar to that of all other hairs,
etc., on the wings and on the bodies of insects; such an out¬
growth is the product of a single cell, a trichogen. Very early
in the development it is apparent that each frenular bristle of
the female arises from a single trichogen and that it remains at
all times separate from the others ; in the male the large frenu-
Marshall — Development of the Frenulum of the Wax Moth. 201
lum is formed from at least a dozen trichogens, in a group adja¬
cent to each other, the bristles from which, at first separate, soon
unite into a single piece.
A section through a hind wing soon after it has emerged from
the peripodial cavity will show that over the entire surface there
is a regular unicellular layer of hypodermal cells even on that
part where the frenulum later develops. As the wing becomes
slightly larger, one finds in this same area that certain of' these
hypodermal cells are larger than the surrounding ones and that
they soon sink below the surface, a position they continue to oc¬
cupy. At a slightly later stage it is noticed that the wings ex¬
amined can be divided into two groups: in one group there are
but three of these enlarged trichogens in each wing, in the other
there are a dozen or more of them grouped closely together. It
is also apparent that where there are but three of these tricho¬
gens present they are individually larger than in those wings in
which there is a greater number. This difference in the size and
number of these trichogens makes it possible to distinguish the
female from the male moths. As this difference in size, as well
as in the final development, is quite noticeable in the two sexes,
each one will be treated separately. Figures 2 and 4 (Plate I)
are drawn at about the same magnification from a male and a
female specimen and show, at about the same age, the relative
size of the trichogens in the two sexes.
Male moth. When the wing has reached about one fourth to one
fifth its normal size, there can be seen along the costal margin
near the humeral angle a group of enlarged hypodermal cells.
Sections show that these cells and their nuclei have, from their
enlarged size, become very distinct and that they have pushed in
from the surface of the wing. As yet the surface at this place
shows only a slight elevation above the surrounding area. As
development goes on, these enlarged cells (trichogens), become
more clearly defined and form a group of cells each one of which
lies close to the others. Early in their development there ap¬
pears from the free end of each trichogen a small, thin, papilla¬
like outgrowth; these at first protrude at right angles to the
margin of the wing, but they soon bend to point towards its
apex, thus taking up the position the frenulum occupies in the
mature moth. At first each small outgrowth remains separate
from all the others; soon their distal ends come in contact with
202 Wisconsin Academy of Sciences , Arts , and Letters.
each other and thus become permanently attached; this attach¬
ment, noticeable at first at the free ends only, soon extends to
the base of the bristles, which thus become united throughout
their entire length, forming the large compound frenulum of the
male moth. As far as can be noticed, the secretion which forms
the bristles is also used to cement them together.
An examination of the frenulum of a mature moth under the
microscope will at once disclose its compound structure, the
wall and lumen of each bristle being clearly seen. The number
of these bristles is not easily determined. Sections were made
both of the frenulum of the mature moth and of that of the moth
just before leaving the pupal case and before the cuticular cov¬
ering had hardened. Even with these sections it was difficult
to determine the exact number of bristles, but we are led to
believe that the frenulum of the male wax moth contains from
twelve to sixteen separate bristles.
Female moth. The frenulum of the female begins its develop¬
ment at a corresponding stage to that at which it appears in the
male moth. Instead of several enlarged hypodermal cells there
are only three present; they and their nuclei soon become much
larger than the trichogens of the male. One can thus readily
determine the sex of a pupal moth by an examination of the
frenular area of the hind wing. The frenulum-forming tricho¬
gens in the female have each to produce a much greater amount
of secretion, and they and their nuclei are correspondingly
larger.
Here, as in the male, the trichogens sink below the surface of
the wing but become separated from each other (fig. 5). The
bristles are at first at right angles to the margin, but soon bend
towards the apex of the wing and at all times remain entirely
distinct from each other. The bristles are borne on the outer
margin of the frenular lobe, and the base of each becomes envel¬
oped by a layer of hypodermal cells (fig. 6).
It was earlier stated that the number of the bristles forming
the frenulum in the female wax moth varies but that three to
each hind wing is the commonest number. Having found many
mature moths with but two bristles on one of the wings, it was
decided to examine a number of mature female moths to ascer¬
tain the relative number of those bearing three or some other
number of bristles. The following table shows that there is a
Marshall — Development of the Frenulum of the Wax Moth. 203
very close division into one group containing three bristles on
each hind wing, which we can call the normal number, and an¬
other group in which the number varies from this.
Right wing with 3 bristles, left wing with 3_. _ 456
Right wing with 2 bristles, left wing with 2. _ 121
Right wing with 2 bristles, left wing with 3 - 183
Right wing with 3 bristles, left wing with 2 _ 136
898
From the table it will be seen that 456 female moths had the
normal number and that 442 were abnormal.
Besides these recorded specimens there was one moth which
had but a single bristle on each wing.
The pupae of the wax moth when removed from their cocoon
are found to be covered with a cuticula which is quite hard and
thick over certain parts of the body. It was almost impossible
to remove this covering from living specimens and leave the body
intact ; it was found that the pupae must be hardened before the
wings from the young ones could be removed in a perfect con¬
dition. After trying several of the commoner preserving fluids,
Bouin’s was found to be the most satisfactory and was used in
the following way: with a hypodermic syringe the fluid was in¬
jected into the body, care being taken not to make the pressure
such that it would burst the pupae. The specimens were then
placed in a dish of Bouin’s fluid, where they were allowed to re¬
main for twenty-four hours or longer and then transferred to
seventy per cent alcohol. At the time of transfer many of the
specimens were injected with alcohol. In a few days the cuti¬
cula could be removed from over the wings, these cut out, and
then mounted entire or prepared for sectioning.
The University of Wisconsin
Bibliography
1. Boisduval, J. A. Species generale des Ldpidopteres. Paris, 1836.
2. Burmeister, H. Handbuch der Entomologie. Berlin, 1832-1844.
3. Chenu, J. G. Encyclopedic d’histoire naturelle. Papillons noc¬
turnes (avec E. Desmarest). Paris, 1858.
4. Comstock, J. R. Manual for the study of insects. 1895.
5. Comstock, J. H. Evolution and taxonomy. Reprinted from The
Wilder Quarter-Century Book, 1893.
6. Comstock, J. H. The wings of insects. Ithaca, 1918.
204 Wisconsin Academy of Sciences, Arts, and Letters.
7. de Geer, C. Memoires pour servir a l’histoire des insectes.
Stockholm, 1752.
8. Giarrm, E. Account of a singular conformation in the wings of
sc^e species of moths. Trans. Linn. Soc. Lond. 1: 135.
1791.
9. Griffith, G. C. On the frenulum of the Lepidoptera. Trans.
Entom. Soc. Lond. 1897, p. 121.
10. Harris, M. Tendons and membranes of the wings of butterflies.
London, 1767.
11. Kirby, W., and Spence, W. An introduction to entomology.
London, 1815.
12. Meyrick, E. Handbook of British Lepidoptera. London, 1895.
13. Poey, P. Ann. Soc. Entom. France. 1: 1832.
14. Tillyard, R. J. The wing-coupling apparatus, with special refer¬
ence to the Lepidoptera. Proc. Linn. Soc. New South Wales.
43: 286. 1918.
15. Westwood, J. O. Introduction to the modern classification of
insects. London, 1839-1840.
Explanation of Plate I
Fig. l.
Fig. 2.
Fig. 3.
Fig. 4.
Fig. 5.
Fig. 6.
All figures drawn with a camera lucida.
View of the frenulum-bearing area of a wing, young speci¬
men of male pupa, showing the enlargement of several of
the hypodermal cells. Near the free surface can be seen
a number of normal cells of the hypodermis; these are in
reality between the enlarged cells. X 18 6.
View of the same area from an older male pupa. The frenu-
lar lobe has not yet formed, and part of the trichogens are
not seen. The prolongation of the cells beyond the sur¬
face is shown; as yet they are separated from each other.
X 146.
Drawing of the frenular lobe of a still older male pupa. The
prolongations of the cells are not drawn throughout their
entire length, but they have in part fused to form the
single large bristle characteristic of the male moth. Cellu¬
lar divisions and nuclei of hypodermal cells not drawn.
X 186.
The frenular area of a young female moth of about the same
age as that represented in figure 2. The three enlarged
hypodermal cells are clearly seen. X 18 6.
Section through the frenular area of the pupa of a female
moth. The three large trichogens are shown and the
normal hypodermal cells. The bristles developing from
the trichogens have not been drawn. Cuticula shown.
X 146.
A single trichogen from the pupa of a female moth; the
bristle is cut near its base. X 186.
TRAMS. WIS. ACAD., TOIL. XX
PLATE 1
MARSHALL— —WAX MOTH
NEW AMERICAN WATER MITES OF THE GENUS
NEUMANIA
RUTH MARSHALL
The genus Neumania was established by H. Lebert in 1879 and
named in honor of the Swedish hydrochnologist, C. J. Neuman.
It belongs to the family Hygrobatidae, as given by Wolcott.
This group of water mites has long been known, the type species,
N. spinipes, having been described by 0. F. Muller in 1776, al¬
though it was then placed in the genus Hydrachna. Cochleo-
phorus of Piersig (1894) is a synonym for Neumania.
Twenty-one species are recognized up to this time. Eleven
species have been described for Europe, two of which are re¬
ported also from east Africa. Five others are described from
Africa, four from Java and Ceyloji ; while but one, described from
material from Central America, has been reported so far from
the Western Hemisphere.
The characteristics of the genus have been reviewed in recent
years by Piersig, Koenike and Wolcott. The body is oval, of
moderate size for these animals, the length of the body averag¬
ing about one millimeter. The integument is soft, with some
tendency to the development of small plates or points of chitin.
The epimera are relatively large, covering the greater part of
the ventral surface; they are in three groups separated by nar¬
row spaces. The fourth pair are largest and are nearly rec¬
tangular. From the .first group there extends back on each side
a long triangular piece, like a brace, underneath the third and
fourth pairs. The genital area is large and close to the epimera.
In the male the plates of either side are united around the slit
and form an oval area at the end of the body; in the female the
slit is longer and the plates are rounded and quite separated.
The outlines of the large eggs are sometimes seen in the body
(PI. II, fig. 1). In both sexes the genital plates have conspicu¬
ous acetabula (“Napfen”). Rosette-shaped glands underneath
205
206 Wisconsin Academy of Sciences , Arts , and Letters.
the outer ends of the plates show conspicuously in prepared
slides (fig. 4).
The palpi are small and more slender than the first pair of
legs. The third joint has stout bristles, and the flexor side of
the fourth joint has two small, hair-bearing papillae. The fifth
palpal joint ends in three or four little finger-like processes.
The first leg is the most characteristic one for the genus. It
is stout and bears many dagger-like spines set in chimney-like
papillae on the flexor surfaces of the middle joints; it usually
appears strongly flexed in the proximal third of its length. The
second pair of legs are much like the first but are somewhat
shorter. The third and fourth legs are more slender than the
first two and bear long, slender bristles, as well as many short,
stout ones, the latter sometimes feathered. All the legs end in
delicate double claws. The basal joints are stout; the successive
joints become longer and slimmer.
The Neumanis are beautiful little creatures when viewed alive
under the low power of the miscroscope. The delicate integu¬
ment is semi-transparent, and the internal organs appear as
brown areas with pink, yellow or white patterns. The legs and
plates are often blue-tinged. The legs are long, and these mites
are active swimmers. They are fairly abundant in the shallow
waters of ponds and lakes where there is some current.
Eleven species are described in this paper, all of them new.
Material from four states has been examined, which yielded
seventy- two individuals. The author’s material was collected
in Wisconsin, Illinois and Michigan. It was supplemented by
valuable collections taken from the lakes at Madison, Wisconsin,
by Dr. R. A. Muttkowski, and by other material from Louisiana,
collected by Dr. E. A. Birge, and very kindly turned over to the
author.
Neumania ovata nov. spec.
Plate II, figure 1.
This is a large mite, the female measuring 1.2 mm. It resem¬
bles N. limosa (C. L. Koch) and N. triangularis Piersig, two widely
distributed European species, but differs from them in the form
of the epimera and genital areas. The epimeral region is small ;
the plates are bluish and widely separated, the space between
the posterior ones gradually widening. The braces from the an-
Marshall — American Water Mites of the Genus Neumania. 207
terior groups are short. The genital plates of the female are
likewise small, rounded, with the acetabula crowded, small, and
not very distinct. The genital cleft is long. The entire genital
area is removed from the posterior end of the body. The legs
are deep blue, stout and well provided with bristles. The fourth
and fifth joints of the fourth legs have long coarse hairs and a
row of stout feathered bristles; a few of the latter are to be
found also on the third joint.
The male is unknown. Three females were found in Wiscon¬
sin lakes: in Lake Wingra, at Madison, and in Lake Spooner,
near Spooner.
Neumania semicircularis nov. spec.
Plate II, figures 2, 3.
This is the largest mite in the present collection, the female
having a length of 1.4 mm. The body is light in color, oblong,
narrowed dorso-ventrally at the anterior end. The epimeral re¬
gion is not extensive, and the spaces between the plates are large.
The genital area is removed from the epimera and the posterior
end of the body; in the female the plates are broadly semicircu¬
lar in outline, and the acetabula are small. Feathered bristles
are conspicuous on the third and fourth legs.
The male is unknown. Six females were found in Wisconsin
material: from Lake Spooner, near Spooner, and from Mirror
Lake, near Delton, by the author; and from the Madison lakes
and the Yahara River, by Dr. R. A. Muttkowski.
Neumania tenuipalpis nov. spec.
Plate II, figures 4-6.
The body is ovate, rather elongated. The epimeral region is
small, the spaces between the anterior and posterior groups
large. The braces from the first group of plates are close to¬
gether. The genital plates of the male are not extensive. The
heavy bristles and their sockets are not as conspicuous as in
some forms. The fourth and fifth joints of the fourth legs have
each a cluster of long hairs and a row of short, stout bristles;
both legs have a few feathered bristles. The palpi are unusually
small, and the second joint is short. The male measures 1.00 mm;
the female is unknown.
208 Wisconsin Academy of Sciences, Arts, and Letters.
Three males were found: one was in material from WinchelTs
Pond, near Kilbourn, Wisconsin; one in material collected by
Dr. E. A. Birge in Audubon Park, New Orleans; and one was
found by Dr. R. A. Muttkowski in material from the lakes at
Madison, Wisconsin.
Neumania fragilis nov. spec.
Plate II, figures 7-9
This is a small and delicate mite. The body is rounded, the
anterior end narrowed as seen laterally. The braces from the
first pair of epimera are broad and extend to the middle of the
fourth pair. The fourth pair of plates are relatively narrow.
The genital area is faintly outlined, and the plates are narrow
in the female, with few acetabula. The legs are slender and
bear feathered bristles; the latter are quite conspicuous on the
fourth pair. The body length in the female is 0.90 mm. The
male is unknown.
Four individuals were examined : one came from a pond at Ep-
worth Heights, near Ludington, Michigan; and the others were
found in the lakes at Madison, Wisconsin, by Dr. R. A. Mutt¬
kowski.
Neumania muttkowskii nov. spec.
Plate II, figure 10.
This new species is large and dark. The epimeral area is mod¬
erately large, the anterior group is separated from the posterior
by a wide space, and the braces underlying the groups are nar¬
row. The genital area of the male is distinct, rounded, and small
with the acetabula somewhat remote from each other; it is re¬
moved from the leg plates and from the posterior end of the
body. The legs are stout, and the palpi are small and slender.
Three males were found; the average body length is 1.00 mm.
One female was found, but not in good condition for study. The
species is named in honor of Dr. Richard A. Muttkowski who
found the material in the lakes at Madison, Wisconsin, in May,
1915.
TRANS. AVIS. ACAD., VOL. XX
PLATE II
MARSHALL — WATER MITES
TRANS. WIS. ACAD., VOL. XX
PLATE III
MARSHALL— -WATER MITES
Marshall — American Water Mites of the Genus N eumania. 209
Neumania armata nov. spec.
Plate III, figures 11-13.
The members of this species are large, stout, dark mites with
heavy plates. The posterior groups of plates are nearer to each
other than are the anterior group to them. The fourth leg plates
are broad, and their inner posterior corners end in conspicuous
rounded points. The genital areas are small, with small aceta-
bula. The legs are bluish, rather slender, with a moderate de¬
velopment of bristles and coarse hairs. The males measure about
0.90 mm., the females about 0.95 mm.
The individuals of this new species were found by the author
in the lakes at Delton, and at Madison, Wisconsin; and also by
Dr. R. A. Muttkowski in the Madison lakes. A total of eight
males and six females was examined.
Neumania extendens nov. spec.
Plate III, figures 14-16.
The epimeral area is extensive, the fourth plates are large,
and the spaces between the groups are small. The genital area
in the male is at the extreme end of the body, heavily outlined,
broad, with slight extensions on the outer borders. The legs are
rather slender with a moderate number of bristles, some of which
are feathered. They are blue-tinged in mature individuals.
The largest males measure 0.90 mm. ; one small individual was
found which measured but 0.50 mm. The female is unknown.
Four individuals were examined. They were found by the
author in collections from Epworth Heights, near Ludington,
Michigan, and in two Wisconsin lakes, Mirror Lake, near Del-
ton, and Lake Spooner, near Spooner; and one individual was
found by Dr. R. A. Muttkowski in collections from Madison,
Wisconsin.
Neumania papillator nov. spec.
Plate III, figures 17-20 ; Plate IV, figure 21.
This new Neumania resembles N. spinipes (Miill.), the type
species of the genus, widely distributed throughout Europe and
reported also by Koenike from Africa. From this the American
14— S. A. L.
210 Wisconsin Academy of Sciences , Arts , and Letters.
species differs in several details, most conspicuously in the form
of the genital plates. N. papillator also resembles N. fragilis and
N. extendens, differing from them in the shape of the genital areas
and in the size of the fourth leg plates. The groups of epimera
are large. The genital plates are a little removed from the
epimera and are placed at the extreme end of the body. The
papilla which is always found near the outer margin of the geni¬
tal plate of each side is very conspicuous in the male of this
species and is set into a concavity of the outer surface of each
plate. In the female the plates are small, pear-shaped, remote
from the cleft, and have the large papillae near them also. The
acetabula are not very distinct. The epimera and legs are tinged
with blue. The first three pairs of legs are well provided with
heavy spines set into deep sockets. The last joint of the first
leg is long, slim, and slightly curved; the fourth leg has long,
stout hairs, while the short spines of the middle joints are
slightly feathered. The palpi are large for the genus, especially
in the second joint.
This is a small mite, the male measuring 0.60 mm., the female,
O. 78 mm. Eight individuals were found: by the author in the
Lauderdale Lakes, southern Wisconsin ; by Dr. R. A. Muttkowski,
in the lakes at Madison, Wisconsin; and by Dr. E. A. Birge at
Slidell, Louisiana.
Neumania distincta nov. spec.
Plate IY, figures 22, 23.
This large member of the genus is readily identified by the un¬
usual form of the male genital plates, which are narrow, con¬
stricted near the center, and extend far out from the genital
cleft. The acetabula are small and scattered. The epimera are
heavy, large, and close together; the fourth pair have conspicu¬
ous posterior corners. The brace from the first group of plates
is short and shaped like a curved blade. The legs are stout and
bluish. The first and second pairs have the usual dagger-like
bristles set in large sockets; the third legs have many stout
bristles, moderately long; the last three joints of the fourth legs
have each a row of short, stout bristles with a bunch of longer
bristles on the distal ends of the fourth and fifth joints. The
palpi are rather stout; the fourth joint is broad at the distal end,
and the hair papillae are large, while the fifth joint is very small
TRANS. WIS. ACAD., VOL. XX
PRATE
MARSHALL— W ATER MITES
Marshall — American Water Mites of the Genus Neumania. 211
and curved. The body is slightly narrowed dorso-ventrally at
the anterior end. The male measures 1.20 mm. in length. The
female is unknown.
Four individuals were found in the lakes at Delton, Wisconsin,
in August, 1910.
Neumania brevibrachiata nov. spec.
Plate IY, figures 24, 25.
This mite has a stout body and the epimeral plates cover nearly
the entire ventral surface. The groups of plates are closely ap¬
proximated. The underlying braces from the first group are
very short, barely reaching to the beginning of the fourth pair,
and they are very narrow. The genital area is crowded to the
extreme end of the body and touches the fourth pair of epimera ;
the plates in the male join at the anterior border, and the cleft
is short. The legs are stout, the bristles moderately long; the
last two joints of the fourth legs are unusually long. The length
of the body in the male is 0.90 mm. The female is unknown.
Two individuals of the species were found; one came from
Lake Mason, Briggsville, Wisconsin, and the other from a small
pond at Epworth Heights, near Ludington, Michigan.
Neumania punctata nov. spec.
Plate IV, figures 26-32.
The epimeral region is very large, and very narrow spaces
separate the three groups. The underlying braces from the first
group extend back through half of the width of the fourth pair
of plates. These last epimera are broad, with the posterior bor¬
ders concave and the corners sharply marked off. The genital
plates are large and lie close to the fourth epimera, at the ex¬
treme end of the body; the clefts are large. In the male, the
papillae at the outer ends of the plates are conspicuous. The
first pair of legs are stout; the middle joints are well provided
with long and heavy dagger-like bristles which arise from con¬
spicuous sockets. The fourth legs are thickly beset with bristles
and coarse hairs of different lengths. In the palpi, the second
joint is somewhat shorter and more bent then is usual; the hair
papillae on the elongated fourth joint are small and bear fine
hairs, while other fine hairs are found on the outer margin. The
212 Wisconsin Academy of Sciences , Arts , and Letters.
integument shows many small rounded pieces of chitin, while
the appendages have many small areas of tiny points scattered
irregularly over their surfaces. This feature, although not pe¬
culiar to this species, is unusually pronounced here and has sug¬
gested the specific name. This is a small mite ; the body length
is variable, but averages about 0.75 mm.
The species is described from the study of thirteen males and
seven females, all from Wisconsin lakes. The collections came
from lakes near Delton, Big Spring, Briggsville, Spooner,
Neshota and Madison, the last mentioned material added by Dr.
R. A. Muttkowski.
Bibliography
Croneberg, A. 18 9 9. Beitrag zur Hydrachnidfauna der Umgegend
von Moskau. Bull. Soc. Imp. Nat. Moscou, 1899, no. 1: 69-71.
PI. IV, figs. 4, 5.
Daday, E. von. 18 98. Microskopische Siisswasserthiere aus Ceylon.
Termesz. Fiiz. 21: 85-9 0. figs. 41-43.
1910. Unter Siissw.-Mik. Deut.-Ost. Zool. 59: 250.
Koenike, E. 1893. Die von Herrn Dr. F. Stuhlmann in Ostafrika
gesammelten Hydrachniden des Hamburger naturhistorischen Mu¬
seums. Jahr. wiss. Anst. Hamburg, 10: 25-29. PI. II, figs. 19,
20.
— - — . 1895a. Ueber bekannte und neue Wassermilben. Zool. Anz.
18: 387-389, figs. 16, 17.
— — 1895 b. Die Hydrachniden Ost-Afrikas. Deut.-Ost-Afrika
4: 13-15, figs. 1, 2. Geog. Verlags. Diet. Reimer, Berlin.
— — . 1905. Vier neue Wassermilben. Zool. Anz. 29: 548-550,
fig. 1.
— 1 . 19 06. Hydrachniden aus Java. Jahr. Hamburg. Wissen.
23: 111-122, Pis. I, II, figs. 6-15.
- . 1908. Ein neues Hydrachniden-Genus und eine Neumania-
Species. Zool. Anz. 33: 706-7 07, figs. 3, 4.
- — — . 1909. Die Susswasserfauna Deutschlands 12: 99-104, figs.
151-158.
Neuman, C. 1880. Om Sveriges Hydrachnider. Konigl. Svenska
Vet.-Akad. Handlgr. 173: 25-32, PI. Ill, figs. 3, 4.
Piersig, R. 18 97. Deutschlands Hydrachniden. Bibliotheca Zoolog-
ica 22: 62-73, 468, PI. II, fig. 4; PI. IV, fig. 7; PI. V, figs. 8, 11,
12.
1901. Hydrachniden (und Halacaridae) . Das Tierreich 13: 223-
227, figs. 60-61.
Stoll, O. 1887. Hydrachnidae. Biol. Centr.-Amer. 59: 9, 10, Pis.
7-9, fig. 1.
Thon, K. 1900. Hydrachniden Durchforschung von Bohmen. Rozp.
Ceske Ak., 92 nr. 15: 135-139, figs. 1, 2.
Marshall — American Water Mites of the Genus Neumania. 213
Viets, K. 1913a. Diagnosen neuer Hydracarinen. Abh. Nat. Ver.
Bremen 222: 227-227.
1913&. Hydracarinen-Fauna yon Kamerun. Arch. Hydro, und
Plank., 0: 40-53, PI. IV, figs. 18-21; PI. V, figs. 22-24.
Wolcott, R. H. 1905. A review of the genera of the water mites.
Trans. Amer. Micr. Soc., 26: 210, PL XXV, fig. 75. (Reprint
Studies from the Zool. Lab., Univ. of Nebr., no. 66.)
EXPLANATION OF PLATES
Plate II
Fig. 1.
Fig. 2.
Fig. 3.
Fig. 4.
Fig. 5.
Fig. 6.
Fig. 7.
Fig. 8.
Fig. 9.
Fig. 10.
Neumania ovata, female, ventral view.
Neumania semicircular is, female, ventral view.
Neumania semicircular is, last segments of third and fourth legs,
right side, ventral view.
Neumania tenuipalpus, male, ventral view.
Neumania tenuipalpus, right palpus and base of first leg, ventral
view.
Neumania tenuipalpus, left first leg, ventral view.
Neumania fragilis, female, ventral view.
Neumania frag ilis , left palpus, ventral view.
Neumania jragilis, last segments of fourth leg.
Neumania muttkowskii, male, ventral view.
Fig. 11.
Fig. 12.
Fig. 13.
Fig. 14.
Fig. 15.
Fig. 16.
Fig. 17.
Fig. 18.
Fig. 19.
Fig. 20.
Fig. 21.
Fig. 22.
Fig. 23.
Fig. 24.
Fig. 25.
Fig. 26.
Fig. 27.
Fig. 28.
Fig. 29.
Fig. 30.
Fig. 31.
Fig. ,32.
Plate III
Neumania armata, female, ventral view.
Neumania armata, genital area of male.
Neumania armata, left palpus.
Neumania extendens, male, ventral view.
Neumania extendens, right fourth leg, ventral view,
Neumania extendens, palpi.
Neumania papillator , male, ventral view.
Neumania papillator, genital area of female.
Neumania papillator, first three legs, left side, ventral view.
Neumania papillator, right palpus.
Plate IV
Neumania papillator, left fourth leg, ventral view.
Neumania distincta, male, ventral view.
Neumania distincta, left palpus.
Neumania Krevibrachiata, male, ventral side.
Neumania 'brevi'brac'hiata, right fourth leg, ventral view.
Neumania punctata, dorsal view.
Neumania punctata, male, ventral view.
Neumania punctata, chitin patches in the integument.
Neumania punctata, genital area of female.
Neumania punctata, left palpus.
Neumania punctata, left first leg, ventral view.
Neumania punctata, left fourth leg, dorsal view.
SYRPHIDAE OF WISCONSIN
CHARLES L. FLUKE, JR.
The predaceous forms of Syrphidae have seldom received the
credit which is due them for their share in the suppression of the
endless increase of individuals in colonies of plant lice. Hereto¬
fore whenever the enemies of aphids were mentioned the lady
beetles usually took first place, and oftentimes the syrphus flies
were not even mentioned.
Today the entomologist and even many others are beginning
to realize the true value of Syrphidae as plant-lice predators.
Instead of being placed at the bottom of the list, they are now
treated as the most important factors in the checking of unusual
outbreaks of plant lice. Most aphidologists are now recognizing
this fact. A large part of this is due to education along this
line; and the life-history studies of these forms as carried on by
such men as Metcalf have been a great factor in the enlighten¬
ment received regarding these most interesting creatures.
The European grain louse has never been a destructive pest
to apple trees in Wisconsin. This spring (1920), as in many
others past, the young lice hatched in countless numbers and
soon began to multiply. From all appearances spraying would
be necessary to reduce their numbers. A careful examination
of the opening buds showed, almost without exception, one to
several syrphid eggs placed among the plant lice on each grow¬
ing tip. Repeated observations soon made evident the reason
of the non-destructiveness of the aphids. The syrphid larvae
preyed upon the plant lice with such voraciousness that few sur¬
vived to migrate to their summer hosts.
For several seasons biological notes of many of the common
species occurring in Wisconsin have been gathered. It has been
decided to give only a classification of the adults at this time,
with the intention of publishing later the life-history observa¬
tions.
The Syrphidae have been intensively studied but there is still
215
216 Wisconsin Academy of Sciences , Arts , and Letters.
plenty to learn, specially in snch difficult groups as Syrphus,
Platychirus, and Helophilus. I have tried to make the keys sim¬
ple enough for the average entomological student to identify the
species in his collection. In 1910 Dr. Graenicher published a
list of 92 species for the state; the present list contains 137,
which indicates the richness of Wisconsin’s fauna.
I am indebted to the members of the Department of Economic
Entomology of the University of Wisconsin for numerous speci¬
mens ; to Professor H. P. Wilson for his encouragement and for
many helps in securing literature ; to Dr. Graenicher and Profes¬
sor T. E. B. Pope for permission to study the specimens in the
Milwaukee Public Museum, and to Dr. W. S. Marshall for his
kindness in showing me many references and permitting me to
study his collection. To all these I am most heartily grateful.
Characteristics of the Family
“ Small to rather large flies. Head hemispherical, often elon¬
gated or produced in the lower part ; as broad or a little broader
than the thorax. Face moderately broad, bare or clothed with
dust or short pile; excavated in profile under the antennae and
projecting below, or with a distinct convexity near the middle
part, never with longitudinal furrows or lateral ridges, usually
convex transversely, sometimes with a median ridge. Oral open¬
ing large; proboscis rarely much elongated. Front never exca¬
vated. Antennae usually porrect and approximated at their
base, three-jointed, usually with a dorsal arista. Eyes large,
bare or pilose; in the male usually contiguous above. Ocelli al¬
ways present. Thorax comparatively large and robust, moder¬
ately arched above. Squamae of moderate size. Abdomen com¬
posed of five or six visible segments, rarely with only four.
Hypopygium usually not prominent. Legs usually of moderate
strength, never long. Bristles rarely present in any part of the
body, never on the head; the body generally thinly pilose or
bare, but sometimes clothed with thick pile. Wings compara¬
tively large ; third longitudinal vein never forked ; marginal cell
open or closed; the fourth vein terminates in the third at or
before its tip ; three posterior cells ; basal cells large ; anal cell
always closed before the border of the wing; between the third
and fourth longitudinal vein and nearly parallel with them, a
false or spurious vein, nearly always present and characteristic of
the family. ’ Williston.
Fluke — Syrphidae of Wisconsin.
217
Table of Genera (after Williston)
(Genera marked with an asterisk are not known to occur
in Wisconsin.)
1. Antennae with a terminal style .
Antennae with a dorsal (rarely subterminal) arista .
2. Antennae cylindrical, the first two joints elongated; first pos¬
terior cell with a stump of a vein (Geria) . .
. SPHYXIMORPHA (p. 249)
First two joints of the antennae short .
3. Eyes bare; small species . *Pelecocera
Eyes pilose; larger species . *Callicera
4. Marginal cell of the wings closed and petiolate .
Marginal cell open . . .
5. Anterior cross-vein of the wings distinctly before the middle of
the discal cell, almost always rectangular .
Anterior cross-vein near or beyond the middle of the discal cell,
usually oblique .
6. Antennae elongate (if arista plumose, see 56) .
Antennae short . . .
7. Mesonotum with yellow lateral stripes; large species, the abdo¬
men always with distinct yellow bands . chrysotoxum ( p. 223 )
Mesonotum not with distinct yellow lateral stripes or margins. .
8. Face rounded, not tuberculate, pilose; oral margin not project¬
ing .
Face not evenly arched; tuberculate or the oral margin not pro¬
jecting .
9. Moderately large to large species; scutellum flattened, often
with spines or tubercules on its border; a stump of a vein
in the first posterior cell from the third longitudinal vein. .
Small species; scutellum without spines; no stump of vein in
first posterior cell . Pipiza (p. 224)
10. Abdomen much narrowed at the base . . .
Abdomen not or but little narrowed at the base.MiCRODON (p. 222)
11. Face swollen and prominent below . *Rhopalosyrphus
Face not swollen and prominent below . *Mixogaster
12. Body clothed with sparse tomentum; all the femora thickened
and with spinous bristles below (Lepromyia) .*Lepidostola
Body not clothed with flattened tomentum ; the femora not thick¬
ened with spinous bristles below .
13. Face partly or wholly yellow, tuberculate below, the epistoma not
projecting; face and front not wrinkled . Paragus (p. 225)
Front and face black in ground color .
14. Face black in ground color .
Face more or less yellow in ground color . . .
2
4
3
48
5
6
55
7
14
8
9
12
10
11
13
19
15
26
218 Wisconsin Academy of Sciences, Arts, and Letters.
15. Abdomen with only four visible segments, very convex, the
venter excavated; first two joints of the antennae very
short, the third large, subquadrate, with a short subtermi¬
nal arista . *iNausigaster
Abdomen with more than four visible segments .
16. iHind femora distinctly thickened .
Hind femora but little or not at all thickened .
17. Scutellum unusually large, nearly square; males dichoptic _
. *Chalcomyia
Scutellum not unusually holoptic, with facial tubercule .
. . . . *Myiolepta
18. Face rounded, not tuberculate, the oral margin not pro¬
jecting . . . Fipiza (p. 224)
Face tuberculate or the oral margin projecting .
19. Epistoma projecting; small, black species . . .
Face tuberculate, the oral margin not projecting .
20. Front in the female and usually the face in both sexes with
transverse wrinkles; spurious vein obsolete; antennae
short or long . . Chrysogaster (p. 223)
Front and face not wrinkled; face pilose . *Psilota
21. Metallic green, metallic green and black, or black species; facial
orbits separated by a slender parallel grove .
. . . Chilosia (p. 226)
Black with more or less metallic green or blue, with yellow,
yellowish, or metallic cross-bands on the abdomen; face not
with orbital grooves . .
22. Face with transverse grooves in the middle; antennae elon¬
gate . *iRhysops
Face not with transverse grooves or wrinkles. . .
23. Wings not longer than the abdomen; ocellar tubercle large;
abdomen depressed, long elliptical, somewhat narrowed to¬
ward its base, the markings ferruginous or orange yel¬
low . . Pyrophaena (p. 228)
Wings longer than the abdomen, abdomen with yellow or
greenish-yellow, or shining metallic cross-bands; usually
elongate species .
24. Front tibiae distally and the tarsi of the male dilated, those of
the female slightly widened . platycheirtjs (p. 229)
Front tibiae and tarsi slender in both sexes . .
25. Rather large, blackish species, with a large flat, elliptical ab¬
domen . *Xanthandrus Verrall
More elongate and slender species; abdomen not elliptical in
outline . Melanostoma (p. 231)
26. Abdomen narrowed toward the base, distinctly club-shaped or
spatulate in outline . . . .
Abdomen oval or slender, not spatulate or club-shaped in out¬
line . . .
27. Third longitudinal vein bent deeply into the first posterior
cell . *Salpingogaster
Third longitudinal vein straight or gently curved .
16
17
18
19
20
21
22
23
24
25
27
30
28
Fluke — Syrphidae of Wisconsin.
219
31
38
32
33
42
34
28. Hind femora slender, front of female long, narrowed above; the
cheeks very narrow below the eyes; abdomen often very
slender . . . Baccha (p. 227)
Hind femora thickened; front not unusually long in female. ... 29
29. Epistoma produced anteriorly, the face in profile deeply concave
from antennae to tip; third joint of antennae rounded -
. ...Sphegina (p. 238)
Epistoma produced more downward, in profile gently concave;
third joint of antennae not rounded - Neoascia (p. 239)
30. Front long, much narrowed above in the female; cheeks very
narrow, the eyes approaching each other at the lower third
of the head, wings usually with dark picture; abdomen
more or less elongate . . .Ocyptamus (p. 228)
Flies not having the above described assemblage of characters
31. Mesonotum with distinct yellow lateral margins .
Mesonotum not with yellow lateral margins . . .
32. Abdomen with definite yellow cross-bands .
Abdomen not with definite yellow cross-bands .
33. Hind femora extraordinarily thickened . .Syritta (p. 246)
Hind femora slender . . .
34. iSixth abdominal segment in the male as long as the two preced¬
ing together, cylindrical; fifth segment of the female one-
half as long as the preceding . Eupeodes (p. 232)
Sixth abdominal segment of the male not peculiar; the fifth
segment of the female one-third or one-fourth as long as the
preceding . . . . . .
35. Front very convex; eyes of male with an area of enlarged facets
above (Lasiopticus) . *Scaeva
Front not remarkably convex . . . . .
36. Third longitudinal vein with distinct curvature into the first
posterier cell; third joint of antennae elongate oval....
. Didea (p. 232)
Third longitudinal vein straight or gently curved; epistoma
not produced (if produced snout-like, Rhingia) .
37. Males holoptic . ,Syrphus (p. 232)
Males broadly dichoptic; arista thickened .... *Chamaesyrphus
38. Mesonotum with median cinerous line; ocelli usually remote
from the vertex . . .
Mesonotum not with a median cinereous linear stripe .
39. Hind femora in the male thickened and arcuate, the tibiae
dilated at the tip . . Toxomerus (p. 237)
Hind femora simple and straight; tibae not dilated at the
tip . . . . . . . Mesogramma (p. 237)
40. Eyes of male with an area of enlarged facets above; fourth
segment of abdomen with two median yellow stripes and
oblique side spots . Allograpta (p. 235)
Eyes of male not with an area of enlarged facets above; fourth
abdominal segment not so marked. . . . 41
35
36
37
39
40
220 Wisconsin Academy of Sciences , Arts , and Letters.
41. Face projecting below; slender species, the hypopygium often
large . . Shapaerophoria (p. 238)
Face receding; abdomen oval _ Xanthogeamma (p. 235)
42. Thickly pilose species; the abdomen black, the basal part light
yellow . . . *iLeucozona
Thinly pilose species, abdomen not so marked .
43. Hind femora thickened . . .
Hind femora slender .
44. iSpecies wholly or chiefly reddish or lutescent .
Black species, sometimes with luteous spots on face, humeri,
and basal angles of abdomen .
45. Scutellum unusually large, nearly square in outline; males
dichoptic . *Chalcomyia
Scutellum oval; males holoptic . *Myiolepta
46. Face carinate; abdomen oval . Brachyopa (p. 239)
Face tuberculate; abdomen more elongate. . *Hammerschmidtia
47. Epistoma produced into a long porrect snout . Rhingia (p. 239)
Epistoma not produced (compare Paragus sp. if small) ; facial
orbits limited by a slender groove . Chilosia (p. 226)
48. Third vein bent deeply into the first posterior cell .
Third vein not bent deeply into the first posterior cell; third
antennal joint elongate . . .
49. Arista very densely plumose, appearing as a solid mass, the
arista itself also thickened . *Copestyium
Arista feathery-plumose .
50. Hairs of arista retrorse; males dichoptic . ,Megametapon
Hairs of arista not retrorse; males holoptic. Volucella (p. 239)
51. Hind femora with a sharp tooth-like projection below neai dis¬
tal end; sixth vein beyond anal cell strongly curved’
large species . . . .Milesia (p. 248)
Hind femora without such tooth .
52. Frontal triangle of male strongly protuberant, rather large dark-
colored species . . . Eristalis (p. 240)
Frontal triangle not protuberant.... .
53. Epistoma produced into a long porrect snout. . *Eicastrirhyncha
Epistoma not produced .
54. Thorax with distinct yellow markings; hind femora thickened;
hypopygium enlarged . . . *Meromacrus
Thorax not with distinct yellow markings, sometimes white-
fasciate; femora sometimes thickened; hypopygium not
conspicuously prominent . Eristalis (p. 240)
55. Arista plumose .
Arista bare or pubescent .
56. Antennae elongate, the third joint more than twice as long as
wide (Phalacromyia, Glaurotricha) - Volucella (p. 239)
Antennae short, the third joint not more than twice as long
as wide; third vein nearly straight, or gently or consider¬
ably curved into first posterior cell . . .
57.
58.
59.
60.
61.
62.
63.
64.
65.
66.
67.
68.
69.
70.
71.
72.
73.
Fluke — Syrphidae of Wisconsin. 221
Thinly pilose; abdomen with yellow bands .
Thickly pilose, abdomen not with yellow bands .
Third vein straight or moderately curved .. Sericomyia (p. 239)
Third vein considerably curved . Condidea (p. 240)
Eyes pubescent . *Pyritis
Eyes bare . ,Arctophila
Third longitudinal vein deeply curved into 1st posterior cell. . . .
Third vein only gently curved .
Hind femora thickened . 1 .
Hind femora slender .
Abdomen elongate, narrowed at base, spatulate in outline
(11) . *Salningogaster
Abdomen oval, with yellow, interrupted bands; mesonotum with
yellow margins; antennae elongate . Chrysotoxum (p. 223)
Face carinate or subcarinate ; hind femora with an angular pro¬
tuberance or spur below at outer end: .
Face protuberant in profile . Tropidia (p. 241)
Face concave in profile, subcarinate; spur of femora bifid .
. *Senogaster
Face tuberculate or arched, not keeled .
Abdomen much narrowed at base, club-shaped . *Ceriogaster
Abdomen not at all pedunculate or basally narrowed .
Antennae elongate; eyes pubescent; wings colored anteriorly..
. *“Platynochaetus ” niger female
Antennae not elongate .
Third joint of antennae broad; mesonotum not vittate .
Third joint of antennae oval:
Males holoptic; femora with protuberance below . .
. *Merodon
Males dichoptic; femora not with protuberance .
Ocelli remote from each other; mesonotum not vittate .
. *Asemosyrphus
Ocelli not remote; mesonotum vittate .... Helophelus (p. 242)
Thickly pilose species . Malotta (p. 245)
Thinly pilose .
Hind tibiae of male with an internal spur.TEUCHOcxEMis (p. 41)
Hind tibiae without such spur (Triodonta) . *Pterallastes
Thorax with distinct yellow markings other than on the humeri ;
wasp-like flies . .
Thorax not with distinct yellow markings of the ground color
other than rarely on the humeri .
Hind femora swollen and with a protuberance or spur below
distally; face carinate or subcarinate .
Hind femora not with such protuberance or spur .
Face concave in profile, femoral spur bifid . *Senogaster
Face protuberant in profile; femoral protuberance not spur-like.
. Tropidia (p. 241)
Abdomen narrowed basally; slender species (see 27) .
Abdomen in no wise club-shaped .
58
59
61
70
63
62
64
65
66
68
67
69
81
71
72
73
74
222 Wisconsin Academy of Sciences, Arts, and Letters.
74. Face transversely arched, not produced, not tuberculate; abdo¬
men more or less elongate and nearly bare . 75
More or less thickly pilose species, often large . 76
75. Hind femora extraordinarily thickened; anterior cross-vein rect¬
angular, and before the middle of discal cell (see 33) _ _ _
Hind femora distinctly thickened, but the cross-vein distinctly
oblique, and near or beyond middle of discal cell .
. . . . . . . Xylota (p. 246)
1 6. Scutellum, margin of thorax, and pleurae with distinct bristles,
femora slender (Chrysochlamys) . *Ferdinandea
No bristles anywhere on body . 77
77. Face short, not produced, concave from antennae to oral mar¬
gin, not tuberculate; hind femora thickened . 78
Face produced, long . 79
78. Abdomen elongate . Brachypalptts (p. 247)
Abdomen very broad; thorax densely pilose; middle femora of
male sometimes (Hadromyia) with a stout, basal, inferior
spur . . . . . *Pocota
79. Face produced forwards, pointed, concave from antennae to tip,
not tuberculate; hind femora thickened . Crioprora (p. 247)
Face not evenly concave in profile, but tuberculate or convex .... 80
80. Third joint of antennae produced above into an anteriorly di¬
rected, conical process, terminating in the thickened
arista . *Merapioidus
Third joint of antennae obliquely oval; hind femora rarely
thickened . Criorhina (p. 247)
81. Hind femora with a conical, tooth-like protuberance below near
distal end; antennae more or less elongated; sixth vein di¬
rected obliquely outward beyond anal cell.SpiLOMYiA (p. 248)
Hind femora without much protuberance; sixth vein beyond
anal cell not unusual . 82
82. Antennae inserted low down, near middle of head in profile,
the face not longer than front . Temnostoma (p. 248)
Antennae inserted on a conical process; front short, the face
much produced downward; antennae long or short. . .
. . . *Sphecomyia
Microdon Meigen
Hind metatarsi slightly or not at all thickened; third segment of
antennae nearly as long as the first; scutellum with apex emar-
ginate and armed on each side with a minute tooth of the same
color; dark-colored species . tristis
Hind metatarsi of male greatly dilated; third joint of antennae
shorter than the first; scutellum large, trapezoidal in shape,
wholly without spines or tubercles . . . ( gloUosus)
1. M. tristis Loew. One female Dane County (Marshall), Mil¬
waukee County (Graenicher). Figures 1, 2.
(M. globosus) Fabr. Mich., N. J., Fla., Car., Tex. Figure 3.
Fluke — Syrphidae of Wisconsin.
223
Chrysotoxum Meigen
Lateral margin of the third abdominal segment entirely yellow - *
. . . . . . . laterale
Lateral margin of the third segment never entirely yellow, usually
black . . pubescens
2. C. laterale Loew. One female from Price County which has
the second joint of the antennae shorter than the first and
third. Sides of the third segment of the abdomen nar¬
rowly yellow ; all bands touch the side margins. Bases of
front and middle femora black.
3. C. pubescens Loew. One female from Madison, Dane County,
August 13, 1918. Figure 4. This specimen indicates a
linking between laterale and pubescens , as Johnson in
Psyche (1907, p. 78) has already noted. The specimen I
have labeled pubescens is 13 mm. in length, and has the
lateral margins of the third abdominal segment yellow in¬
terrupted. The bases of the front and middle femora are
somewhat tinged with black, especially toward the outside.
The specimen is very close indeed to the female in the
Milwaukee Public Museum called laterale.
Chrysogaster Meigen
1. Base and tip of tibiae, or at least first two joints of tarsi,
yellow or yellowish red; last section of fourth longitudinal
vein rectangular; antennae elongate . 2
Legs wholly black; last section of fourth vein curved or bent,
third joint of antennae nearly rounded, front of female
strongly rugose . nigripes
2. Eyes with linear markings, last section of fourth vein joins the
third beyond the tip of the second vein, rectangular with a
stump of a vein in the middle; second joint of the antennae
nearly as long as the third . . . . . nitida
Eyes unicolorous, last section of fourth vein joins third oppo¬
site or before the tip of the second . 3
3. Cross-veins of the wings clouded, antennae distinctly longer than
from base of antennae to upper mouth edge, i. e., longer
than the face . pictipennis
Cross veins of wings not clouded, antennae scarcely as long as
from base of antennae to upper mouth edge, i. e., not longer
than the face . pulchella
4. C. nigripes Loew. Milwaukee and Washington Counties.
Figure 5. Also recorded from N. Y., N. J., Conn., N. C.,
Nebr., and Que.
224
5.
6.
1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
Wisconsin Academy of Sciences 9 Arts , and Letters.
C. nitida Wied. Vernon, Grant, Milwaukee, Marathon, Ju¬
neau, and Buffalo Counties. Late summer and fall. Fig¬
ures 6, 7.
C. pictipennis Loew. Milwaukee, Marathon, Winnebago, Dane,
Jefferson, and Sauk Counties. Late summer. Figures 8,
9. Females often caught while sweeping along sewer
drain ditches or swamp cat-tails.
C. pulchella Will. Buffalo, Vilas, Milwaukee, Racine, Dane,
Washington, and Douglas Counties. July to late Septem¬
ber. Figures 10, 11.
Pipiza Fallen
Abdomen with basal interrupted yellow fascia
Abdomen uniformly dark . .
Wings with a brown spot in the middle . f estiva
Wings without a brown spot in the middle . femoralis
Third joint of antennae at least twice as long as wide .
Third joint scarcely twice as long as wide, or rounded .
Small species — 4 or 5 mm. long — pile rather long, dilutely lutes-
cent, that of face white . pubescens
Larger species — at least 6 or 7 mm. long — pile short, light or
dark .
Wings nearly hyaline, pile of face white, body chiefly white
pilose . radicum
Wings distinctly smoky; pile chiefly dusky or black. .. .salax?
Wings with a brownish cloud . femoralis, form alMpilosa
Wings without such cloud .
Hind coxae of male armed with a slender process, dilated and
compressed toward the end . calcarata
Hind coaxe without such process . (pisticoides)
2
3
4
6
5
7
P. pubescens Loew. Wisconsin (Loew).
P. calcarata Loew. One male Madison, Dane County, June
14, 1918. Figure 12. This species is easily told by the
peculiar slender process on the hind coxae and also in a
smaller form on the middle coxae. It was described from
New York; I also have specimens from Colorado and Cali¬
fornia.
P. salax Loew. Two males taken at Madison, one June 29,
1917, and the other August 18, 1918. Figure 13. I re¬
fer these specimens to salax with considerable doubt, as
will be noticed by the following notes :
Fluke — Syrphidae of Wisconsin .
225
Male 6.5 to 7.5 mm. long. Pile chiefly dusky or black ;
pile of vertex and frontal triangle long black, of the face
slightly brownish. Face and frontal triangle blue-black,
shining; face with a distinct line of white pollen along
each eye-margin. Pile of eyes brownish, lighter below.
Antennae brownish black, reddish beneath the third joint,
arista reddish. Occiput with white pollen and pile.
Thorax shining bronze-black with brownish erect pile,
lighter on the dorsum; tegulae black, brownish ciliate.
Abdomen shining bronze-black on basal half to two thirds,
blue-black on apical third; pile short, black, except on
basal corners, where it is longer and brownish. Legs
black with black or brownish pile, the following parts
luteous : extreme tip of all the femora, basal two thirds of
the front and middle tibiae, and basal joints of front and
middle tarsi; hind tarsi slightly incrassate. Wings dusky,
more hyaline toward the base. Hypopygium rather long
and prominent.
11. P. femoralis Loew. Milwaukee and Dane Counties.
“Johnson (Psyche 14: 75) says femoralis and albipilosa are
different forms of the same species. Four specimens collected
at Milwaukee point in the same direction. One male and two
females have wings hyaline, slightly infuscated on the outer
half. In the other female the wings are conspicuously
clouded and insect is altogether darker than the typical
femoralis , having black antennae and darker legs. In one of
the females the interrupted yellow fascia across the second
abdominal segment is twice as broad as in the others. ’ ’ —
G-raenicher.
12. P. f estiva Meig. One specimen each from Dane and Pierce
Counties.
13. P. radicum Walsh and Riley. Pierce, Grant, Milwaukee,
Dane, and Burnett Counties. June to September. Fig¬
ure 14.
(P. pisticoides Will.) Recorded from N. Hamp., Alaska, Me.,
N. Y., Colo., N. Mex., Nebr., Ohio, Br. Col. Figure 15.
Paragus Latreille
1. Eyes uniformly pilose without a vertical glabrous stripe, face
of both sexes with a black median stripe, seutellum without
a yellow border . . . . . tibialis
Eyes pilose with a distinct vertical glabrous stripe, face of male
without a black stripe, seutellum usually with a yellow
border . . . . . . . 2
15— S. A. L.
226 Wisconsin Academy of Sciences, Arts, and Letters .
2. Abdomen with more or less red on the dorsum, scutellum us¬
ually with a distinct yellow border . Mcolor
Abdomen wholly greenish black, scutellum testaceous only at
the tip . . . . anyustifrons
14. P. angustifrons Loew. Several specimens from Dane County.
Mid-May to Mid-September. I have females of this doubt¬
ful species taken in copulo with males distinctly bicolor.
15. P. bicolor Fabr. Numerous specimens from Vernon, Grant,
Pierce, Milwaukee, Washington, Burnett, Winnebago,
Fond du Lac, and Dane Counties. Middle May to last of
September. Figures 16, 17.
16. P. tibialis Fallen. A common species, specimens taken in
Vernon, Grant, Manitowoc, St. Croix, Pierce, Dane, Mil¬
waukee, Washington, Marathon, Burnett, and Taylor
Counties. May to September. Figures 18, 19.
Chilosia Meigen
1. Eyes thinly pilose; antennae yellow . hoodiana
Eyes bare . . .
2. Arista plumose . . .
Arista bare or pubescent .
3. Scutellum with distinct marginal bristles . species?
Scutellum without distinct marginal bristles .
4. (Legs almost entirely yellow, face not pilose . prima
Legs mostly black .
5. Abdomen wholly shining, metallic green . comosa
Abdomen with second segment and an hour-glass spot on the
third opaque . sororcula
6. Shining metallic blue, wings nearly twice the length of the
abdomen . cyanescens
Shining metallic greenish . f ^allipes
1 tmstis
17. C. hoodiana Bigot. Several females from Douglas, Burnett,
and Manitowoc Counties.
18. C. prima Hunter. Four males and one female from Milwau¬
kee County.
19. C. sororcula Will. Specimens from Burnett, Sheboygan,
Douglas, and Manitowoc Counties.
Fluke — Syrphidae of Wisconsin .
227
Male: Eyes bare, arista bare, pile dusky or whitish,
no bristles on thorax or abdomen. Thorax shining metal¬
lic blue. Abdomen shining, except second segment and
an hour-glass spot on the third segment. Face black-
shining, covered with whitish pollen. Antennae reddish,
arista darker. Tegulae whitish with light brownish hairs.
Wings tinged with brown. Legs black except knees, base
of tibiae, slightly at tip of tibiae, which are yellowish
brown; tarsi brown.
20. G. comosa Loew. One male, Madison, May 10, 1917 ; one male
and three females, Madison, May 11, 1919. Figure 20.
21. C. species. Two males caught at Madison, June 14, 1918, I
am unable to place. They have the eyes bare, bristles on
the margin of the scutellum, arista nearly bare, antennae
rather dark, and tibiae largely yellow.
22. C. cyanescens Loew. Milwaukee and Dane Counties. Early
June. Figure 21. Males show the opaque black cross¬
band on the second abdominal segment; one male meas¬
ures 7 mm. in length.
23. C. pallipes Loew. Two males, Jefferson County, September
27, 1917 ; taken on flowers of goldenrod near a wooded
area.
24. C. tristis Loew. One female, Jefferson County, September 27,
1917. I have referred this specimen to tristis rather than
to pallipes as the humeri are without any light coloration.
Baccha Fabricius
X. Third joint of antennae elongate-oval, obtusely pointed; face
very prominent . . . . . clavata
Third joint short-oval, obtusely rounded, face not prominent. . 2
2. Wings subcinerescent; all the veins, except the last segment
of the fourth, broadly clouded with brownish, confluent
near the costa . . . . lugens
Wings hyaline with only a broad median cross-band reaching
to the fourth posterior cell .......... . . . . . 8
8. Scutellum and posterior part of the thoracic dorsum shining
golden; abdominal bands yellowish red, broadly inter¬
rupted. .................................... fascipennis
Scutellum translucent, yellowish brown, with a metallic reflec¬
tion; abdominal bands blood-red and only subinter¬
rupted . . . . . . . . . lemur
228 Wisconsin Academy of Sciences , Arts, and Letters.
25. B. clavata Fabr. Three males and three females, Milwaukee.
26. B. lemur 0. S. One female from Tenderfoot Lake, Vilas
County. Figure 22.
27. B. fascipennis Wied. Taken in Dane, Milwaukee, Washing¬
ton, and Polk Counties. Early Summer. Figure 23.
28. B. lugens Loew. “N. Wisconsin ” — Loew.
Ocyptamus Macquart
29. 0. fuscipennis Say. Dane and Milwaukee Counties only.
Pyrophaena Schiner
Abdomen largely reddish orange, front and middle tarsi of the male
flattened . granditarsis
Abdomen black with front corners of third and fourth segments
with reddish yellow or metallic spots, tarsi simple - rosarum
30. P. granditarsis Forster. Burnett and Dane Counties, 22
males and 13 females. May 22 to September 21. Figures
24, 25. A common species in the marshes.
31. P. rosarum Fabr. Vilas, Sheboygan, and Dane Counties, 12
males and 4 females. Figure 26. July 7 to September 10.
Variety rosarum Fabr. In the European specimens the male
is black with the basal corners of only the third abdominal
segment reddish yellow, the fourth segment being entirely
black. In the female the fourth as well as the third seg¬
ment have reddish yellow or metallic like spots, but no
markings on the second segment. One female of this
variety taken in Dane County.
Variety duplicata n. var. In this variety the male has distinct
spots on the basal corners of the fourth segment as well as
the third segment. In the female there is not so much
diffrence, there being in addition only an indistinct U-
shaped marking on the second segment. I also have one
female from Vilas County which lacks the markings on
segment four.
Fluke — Syrphidae of Wisconsin. 229
Platychirus St. Fargeau and Serville
(Table of males, adapted from Yerrall)
1. Front tibiae abruptly dilated at the extreme tip, the basal joint
of front tarsi exceedingly larger than the other joints,
front femora with a remarkably coarse ciliation behind on
almost the whole length, antennae pale beneath the third
joint . . . peltatus
Front tibiae gradually dilated or dilated soon after the base,
antennae nearly all black . 2
2. Front femora with a row of isolated long black hairs behind. . 3
Front femora without a row of isolated long black hairs behind 4
3. Front femora with a peculiar long white hair at the base, and
with about six isolated black hairs behind; front tibiae
with no kink above; longitudinal and black cross^bands
of abdomen very narrow . immarginatus
Front femora without a peculiar long white hair at the base,
and with about five isolated black hairs behind; front
tibiae with a kink above; black cross-bands and longi¬
tudinal stripe of abdomen broader . scambus
4. Abdomen with four pairs of abdominal spots . 5
Abdomen with three pairs of abdominal spots . 7
5. Front tibiae only slightly dilated, and with the outer angle
below sharp and very little produced downward; front
femora without a peculiar long white hair at the base. . . .
. . species?
Front tibiae considerably dilated, and with the outer angle
below more rounded and produced downward, front femora
with a peculiar long white hair at the base . 6
6. Middle femora at the base with a bunch of long dense pile, ab¬
domen with a distinct median longitudinal black stripe. .
. quadratics
Middle femora at the base without such bunch of long dense
pile; abdomen with scarcely any black, the median longi¬
tudinal stripe being nearly obsolete . perpallidus
7. Front femora with a few weak black hairs behind; basal ab¬
dominal spots small, nearly circular; wings brownish;
small rather dark species . .hyperboreus
Hairs behind on front femora mostly yellow, basal abdominal
spots usually more nearly the size of the second pair; wings
nearly hyaline; small, narrow, more yellowish spe¬
cies . angustatus
230 Wisconsin Academy of Sciences , Arts, and Letters.
Platychirus St. Fargeau and Serville
(Table of females, after Verrall)
1. Antennae absolutely without yellowish markings beneath the
base of the third joint . 2
Antennae partially yellow, at least beneath the base of the
third joint . 7
2. Hind legs with at least a distinct dark band on the femora and
tibiae . 3
Hind legs entirely orange, without the faintest trace of any
dark markings on at least the hind femora . 6
3. Paler species, with usually only a slight dark ring on the hind
femora and tibiae . 4
Darker species, with usually conspicuous blackish markings on
the hind femora and tibiae, but this character not a very
safe one . . . 5
4. :Larger species; frontal side dust spots golden and well de¬
fined (I have been unable to make use of this character) . .
. scambus
(Smaller species; frontal side dust spots grey and ill defined..
. immarginatus
5. Larger species; side dust spots on the frons moderately
large . Tiyperboreus
Smaller species; side dust spots on the frons small , angustatus
6. Abdominal spots distinctly reddish and separated by a broader
black median line and black cross-bands . quadratus
Abdominal spots distinctly yellowish and separated by nar¬
rower black lines . perpallidus
7. Hind legs entirely yellow, at least on the femora and tibiae. . . .
. quadratus
Hind legs black or brown . peltatus
32. P. angustatus Zett. Two males at Madison, May and August.
33. P. Tiyperboreus Staeger. Sheboygan, Manitowoc, Yilas, Pierce,
Dane, Portage, Milwaukee, Washington, Burnett, and Polk
Counties. May to October. Very closely related to, if
not the same as, clypeatus Meig.
34. P. immarginatus Zett. Fond du Lac and Dane Counties. 5
males. Figure 27. June to September.
35. P. peltatus Meig. Price, Vilas, Milwaukee, Sheboygan, Wash¬
ington, Dane, and Pierce Counties. Mostly late summer.
Figure 28.
36. P. perpallidus Verrall. Barron, Dane, and Fond du Lac
Counties. May to September.
Fluke — Syrphidae of Wisconsin.
231
37. P. quadratus Say. Numerous specimens from Dane, Yernon,
St. Croix, Pierce, Buffalo, Barron, Monroe, Jackson, Mil¬
waukee, Washington, and Bacine Counties. May to Octo¬
ber. Figure 30. Very common in the swamps.
38. P. scambus Staeger. Two males from Vilas County. I be¬
lieve chaetopodus Will, is synonymous with scambus
Staeg.
39. P. species? Sauk County, July 29, 1918. One male among
the reeds in Wisconsin River bottoms. I am unable to
place this specimen with any described species, but until
more material can be secured I prefer not to describe it
as new, especially in this difficult genus. Length 9 mm.
Front tibiae dilated very little, the outer angle below
sharp and very little produced downward; front femora
without a peculiar white hair at the base, with only a few
weak black hairs and a small amount of light-colored pile
intermixed. Figures 32, 33.
Melanostoma Schiner
1. Darker species, abdominal bands distinctly metallic, not red¬
dish yellow, pollen of face somewhat striate on the
sides . . obscurum
More yellowish species; abdomen of male with three pairs of
quadrate yellow spots, female with four pairs of triangular
or drop-like spots . 2
2. Larger species; yellow of fifth segment entire, not interrupted,
all the spots larger . angustatum
Smaller species; yellow of fifth segment separated into spots,
abdominal spots much smaller . mellinum
40. M. obscurum Say. Manitowoc, Vilas, Juneau, Milwaukee,
Washington, Jefferson, and Dane Counties. Common in
early spring. Figure 34.
41. M. mellinum Linn. Specimens from Dane, Mnnitowoc, St.
Croix, Pierce, Buffalo, Monroe, Milwaukee, Washington,
Douglas, Burnett, and Polk Counties. Early spring to
late fall. Figure 36.
42. M. angustatum Will. Numerous specimens, Madison, Dane
County. Late summer and fall. Figure 35. A common
species in the marsh.
232 Wisconsin Academy of Sciences , Arts , and Letters.
Eupeodes Osten Sacken
43. E. volucris 0. S. St. Croix and Marathon Counties.
Didea Macquart
44. J). fasciata Macq. Milwaukee and Vilas Counties.
Syrphus Fabricius
1. All the abdominal bands interrupted or greatly incised - ...
Bands on third and fourth segments, at least, entire. . . .
2. Face with a black shining median line, at least on lower part. .
Face entirely yellowish above upper mouth opening .
3. Basal pair of abdominal spots very small, widely separated,
all the spots triangular; abdomen narrow . oronoensis
Basal pair of spots larger, not so widely separated, other spots
nearly square . . . disjectus
4. Abdomen narrow, basal pair of spots large, whitish .
Abdomen oval, basal spots not larger than others, usually more
yellowish . . . .
5. Smaller species, 8 mm., basal spots large and nearly square,
very narrowly separated, the other spots narrowly linear,
pile of pleurae white . .fisherii
•Larger species, 10 mm., basal spots are more triangular; pile
of pleurae yellowish . (umbellatarum)
6. Spots on third segment joined by a very narrow line, those
on the fourth more broadly connected — the yellow spots
or bands occupying the greater part of the segments,
thorax metallic green . species no. 1
Spots of abdomen distinctly separated, thorax deep metallic
blue .
7. Third longitudinal vein distinctly curved into first posterior
cell pile of pleurae yellow, a single biarcuate black band
above the antennae . arcuatus
Third vein nearly straight, pile of pleaurae white, two rounded
separated spots above antennae . . .per plexus
8. Second and third abdominal bands reach side margins .
Bands separated from side margins .
9. Bands more or less attenuated before reaching side margins . .
Bands going over the side margins in practically their full
width . . .
10. Hind femora yellowish or reddish, at least on basal half .
Hind femora black on basal half .
11. Eyes hairy, faintly so in the female . torvus
Eyes distinctly glabrous . . . . . . .
2
8
4
3
5
6
7
9
20
10
16
13
11
12
Fluke — Syrphidae of Wisconsin.
233
12.
13.
14.
extreme base, thorax shining.
{
15.
16.
17.
18.
19.
20.
Cross-bands rather broad and undulating, reaching side mar¬
gins very narrowly on their upper sides only, thorax
r nitens
brightly shining . . . j _ _ . ,
1 abbreviatus
Bands not so undulating and reaching side margins more
broadly . . . male ribesii
Third joint of antennae at least twice as long as wide; cheeks
black, face with a median stripe . species no. 3
Third joint usually less than twice as long as wide, cheeks us¬
ually yellow . . .
Bands undulating and reaching the side margins by their
upper corners only, front and middle femora black at the
nitens
abbreviatus
Bands not so undulating and reaching side margins more
broadly, thorax with indistinct light grey lines .
Hind femora of both sexes reddish or yellowish at the base. . .
. . - . protritus
Hind femora of male black at the base . . ribesii
Hind femora black at the base . . . grossulariae
Hind femora yellow at the base . . . . .
Lateral margins of the thorax with yellow stripes in ground
color, antennae pointed . Tcnabi
Lateral margins without yellow stripes in ground color, an¬
tennae rounded .
Smaller species, 6 to 8 mm . . . var. ribesii
Larger species, 10 to 12 mm . . . .
Scutellum yellow-haired, bands broad . xanthostomus
Scutellum black-haired, bands narrower . protritus
Antennae reddish or yellowish, seldom blackish, third joint,
nearly as wide as long, pile of pleurae yellowish, bands
f nitens
^ abbreviatus
of abdomen undulating
21.
22.
Antennae blackish or deep brown, at least on the upper side
of the third joint, which is usually nearly twice as long
as wide, abdominal bands more straight .
Abdomen rather narrow with nearly parallel sides in the male;
smaller species; bands broadly biconvex posteriorly, nar¬
row in the female . species no. 2
Abdomen more oval; larger species; bands more straight .
Hind femora black at the base, the middle femora on the out¬
side with a row of rather heavy black pile with a few
pale hairs intermixed; antennae somewhat larger .
. species no. 3
Hind femora of female yellow at the base, middle femora with
a row of mostly yellowish pile, very seldom any black hairs,
present; antennae smaller . americanus
14
15
17
18
19
21
22
234 Wisconsin Academy of Sciences , Arts , and Letters.
45. S. oronoensis Metcalf. One larva collected by L. G-. Gentner
at Eau Claire, June 4, 1917, later pupated and within
three weeks a perfect male adult appeared. All three
stages of this species agreed with Metcalf’s full descrip¬
tion.
46. S. fisherii Walton. Two males, numerous females. July to
September. Dane and Milwaukee Counties. Figures 37,
38. The Wisconsin specimens have the antennae and the
facial stripe deep black, not brown; pile of thorax very
short, longer on the sides, whitish, that of the scutellum
short and mostly black; black of second segment mostly
opaque.
(S. umbellatarum) . The specimens labeled as this species in
the Milwaukee Public Museum are fisherii Walton.
47. S. disjectus Will. One specimen, Dane County, September 3,
1917. Specimens in the Milwaukee Public Museum labeled
disjectus belong to Xantho gramma fragila n. sp., loc. cit.
48. S. species no. 1. Two specimens, Dane County. Figure 39.
49. S. arcuatus Fallen. Vilas, Milwaukee, Dane, Burnett, and St.
Croix Counties. Early spring. Figure 40.
50. S. perplexus Osburn. Numerous specimens from Dane County,
taken in early spring only. Figure 43.
51. S. species no. 2. Two males and two females, all taken at
Madison, May to September. Figure 42. These speci¬
mens are very closely related to diversipes.
52. S. nitens Zett. Dane County. Early May only. Figure 43.
This species has been recorded from Maine. I have been
unable to separate this species from abbreviatus Zett.
Can they be the same species ?
53. S. abbreviatus Zett. Numerous specimens collected from the
marshes, late summer and fall. These seem to have the
bands of the abdomen broader and less undulating than
nitens , otherwise I can see no differences between the two
species.
54. S. ribesii Linn. Pierce, Buffalo, Dane, Manitowoc, Vilas,
Price, Grant, Milwaukee, Washington, Marathon, Douglas,
and Burnett Counties. Throughout the season. I have a
complete series ranging from 6 to 14 mm. in length. The
smallest have the bands going over the sides of the abdo¬
men entirely, face all yellow. About half of the larger
Fluke — Syrphidae of Wisconsin.
235
specimens have a very decided brown stripe on the face;
all of the large specimens and even the males of the
smaller ones have the bands distinctly attenuated.
55. 8. torvus 0. S. Dane and Milwaukee Counties. Early spring
and summer. May easily be told from all other species
of the genus Syrphus in Wisconsin by the pilosity of the
eyes.
56. 8. protritus 0. S. Seven females and two males from Dane
and Dunn Counties. July to October.
57. 8. americanus Wied. Common throughout the State from
May to October.
58. S. species no. 3. Dane and Manitowoc Counties. May to
October.
59. 8. kndbi Shannon. Dane County. June to August. Figure
44. Several males and two females, mostly bred speci¬
mens.
60. 8. xanthostomus Will. One female, Dane County, May 26,
1918. Figure 45.
61. S. grossulariae Meig. Milwaukee, Dane, Jefferson, and Vilas
Counties.
Allograpta Osten Sacken
62. A. obliqua Say. Common in almost every part of the State.
Xanthogramma Schiner
1. Dorsum of thorax before the scutellum with two coalescent
light yellow spots; wings with a brown anterior margin
. . . aequalis
Dorsum of thorax without yellow spots on the hind margin;
wings not brown in front . . . 2
2. Hind femora pure yellow without any dark ring about the
middle, pleurae with distinct bright yellow spots of
ground color . flavipes
Hind femora nearly always dark or with a brown ring near
the middle, pleurae with indistinct bright yellow spots of
ground color . 3
3. Yellow band on second abdominal segment separated from the
lateral margins by a distinct black interval . . . 4
Yellow band on second segment reaches the side margins . 6
236 Wisconsin Academy of Sciences, Arts, and Letters.
4. Smaller species, 6.5 to 8.5 mm. long, with slender, narrow ab¬
domen, sides of mouth-edge black; bands on abdomen of
female all interrupted; on the male the third is entire. . . .
. fragila
Larger species, 9 to 11 mm. long, abdomen more oval, cheeks
and mouth-edge yellow .
5. Abdominal bands broadly interrupted. . .divisa
Bands entire or only slightly interrupted . felix
6. Yellow band on third segment of abdomen goes over side mar¬
gins in nearly its full width . .
Yellow band on third segment usually separated from the lat¬
eral margins, or at least greatly attenuated; bands either
entire or interrupted . . emarginata
7. Small, narrow species, 7 mm. long, with abdominal bands of
female interrupted; male unknown . tenuis
Larger, 10 to 11 mm. long, abdomen oval with bands on third
and fourth segments entire or only sub-interrupted, male
unknown . aenea
63. X. aenea Jones. Figure 46. Two females from Dane County,
August and September, I place here although they differ
somewhat, as the following notes will show :
Black stripe of front extends up to the antennae and
expands. Third joint of antennae almost quadrate in out¬
line, rather large and opaque black, there being no indica¬
tion of yellow on the lower side. Lateral thoracic stripes
rather distinct. Scutellum black at extreme basal corners.
Band on fourth segment of abdomen slightly interrupted.
Hind legs mostly black, yellow at base of femora and at
knees. Wings slightly smoky.
64. X. aequalis Loew. One female, July 6, 1905, Door County.
65. X. divisa Will. Milwaukee, Bacine, and Burnett Counties.
Also one female taken in Dane County, September 19,
1918.
66. X. felix 0. S. One female from Vilas County, August, 1913.
67. X. emarginata Say. Milwaukee, Douglas, Burnett, Polk,
Dane, and Dunn Counties. June to September. Figure
47. I have one male specimen in which the frontal tri¬
angle is entirely yellow, with scarcely any indication of
black ; the antennae are also reddish.
68. X. flavipes Loew. Numerous specimens from Dane and Mil¬
waukee Counties. May to August.
69. X. tennis Osburn. One female from Milwaukee County, July
4, 1907.
Fluke — Syrphidae of Wisconsin.
237
70. X. fragila n. sp. One male and one female from Dane County,
also specimens from Burnett and Douglas Counties. Type
locality, Dane County. Type male in author’s collection.
July 29, 1917. Figures 48, 49. Specimens were taken in
a wood lot among thick low growing vegetation.
Male : Length 7.5 mm. A small, frail looking species
with a narrow abdomen. Face yellow with whitish pollen
and pile, lower mouth-edge and part of cheeks reddish
yellow and connected with the yellow of the face with a
dark area. Frontal triangle yellow with an obscure dark
spot above each antenna, pile black; antennae mostly red¬
dish yellow, darker on upper side of third joint. Verti¬
cal triangle black with black pile; pile behind and along
the occiput yellowish. Thorax shining metallic greenish
black, side stripes somewhat indistinct, pile yellow ; scutel-
lum yellow with basal corners black, pile black and white
mixed. Abdomen semi-opaque black, with yellow as fol¬
lows: first segment broadly on the sides, which slightly
encroaches on the anterior corners of the second segment ;
second segment with a pair of spots, broadly separated
and not reaching side margins ; third segment with a band
deeply emarginate behind but well separated from the
side margins; fourth segment with a pair of spots; fifth
with lateral edges only yellow; the posterior margins of
segments four and five and the anterior corners of the
third and fourth also yellow. Legs yellow, front and
middle femora brown at base ; hind legs with broad brown
rings on the femora and tibiae. Wings hyaline.
Female: Similar with the following exceptions:
Larger, 8.5 mm. long. Front shining greenish black,
dusted whitish on sides below. Antennae larger and much
darker. Pile of pleurae and lower part of occiput whit¬
ish. Abdomen with yellow bands broadly interrupted and
well separated from the margins. Hind legs darker, the
tibiae and tarsi nearly all dark brown.
Mesogramma Say (Toxomerus Macq.)
1. Hind femora blackish or at least with a distinct brown ring
in the female; thickened and arcuated in the male; scu-
tellum dark with a distinct entire yellow border .
. . . . geminata
Legs almost entirely yellowish, hind femora of male simple;
scutellum nearly always yellowish, seldom dark with a
yellow border . 2
238 Wisconsin Academy of Sciences, Arts, and Letters.
2. Front of female only a little narrowed above; smaller species,
5 or 6 mm. long, abdominal markings variable . . marginata
Front of female considerably narrowed above; larger species, 8
or 9 mm. long; abdominal markings more regular, rather
“blocked out” . polita
71. M. geminata Say. Numerous everywhere. Early spring to
late fall. Figures 50, 51, 52.
72. M. marginata Say. More common than geminata; throughout
the season. Figures 53, 54, 55.
73. M. polita Say. Milwaukee, Dane, Fond du Lac, and Winne¬
bago Counties. Midsummer to late fall.
Sphaerophoria St. Fargeau and Serville
(After C. W. Johnson)
1. Lateral thoracic stripe distinctly ending at the suture . 2
Lateral thoracic stripe continuous . 3
2. Face entirely yellow, abdominal bands beyond the second seg¬
ment present in the female, usually obsolete or wanting
in the male; length 8 mm . cylindrica
Face with a wide black stripe, abdominal bands similar in
both sexes ; length 6 mm . novaeangliae
3. Abdomen in the male with a series of yellow spots, in the fe¬
male with widely interrupted bands narrowly connected
with the lateral margin; length 8 mm . (strigata)
Abdomen with bands on the first four segments except the first,
the others irregularly marked with black; length 8 to
10 mm . . . scripta
74. S. cylindrica Say. Cosmopolitan, May to October. Figure
56.
75. S. novaeangliae Johnson. One female, Vilas County, and one
male, Dane County.
76. S. scripta Linn. Cosmopolitan. May to October. Figure
57. Often confused with cylindrica.
Sphegina Meigen
77. S. campanulata Kobertson. One female from Vilas County.
78. S. lob at a Loew. Vilas County.
Fluke— SyrpMdae of Wisconsin.
239
N^oascia Williston
Abdomen scarcely narrowed at the base, uniformly bright metallic,
antennae longer . . distincta
Abdomen distinctly narrowed toward the base, usually with yel¬
lowish or reddish spots or bands, antennae shorter, .globosa
79. N. globosa, Walk. Pierce, Milwaukee, Racine, Sheboygan,
Washington, and Dane Counties. Mostly early spring.
Especially numerous among grass sweepings near sewer
drains.
80. N. distincta Will. One specimen from Milwaukee County.
Rhingia Scopoli
81. R. nasica Say. Vilas, Price, Pierce, Milwaukee, Polk, Wash¬
ington, Marathon, Dane, and Dunn Counties. June and
September, mostly at the latter date. Figure 58.
Brachyopa Meigen
82. B. notata 0. S. One female from Dane County, June 6, 1918.
Volucella Geoffroy
Thorax heavily pilose; non-metallic species . evecta
Thorax nearly bare; brightly metallic-colored species _ vesiculosa
83. V. evecta Walk. Washington, Milwaukee, Marathon, and
Dane Counties. Figure 60.
84. V. vesiculosa Fabr. Milwaukee and Dane Counties. One
specimen taken from a wasp’s nest, a species of Crabro.-
Graenicher.
In the Milwaukee Public Museum there are specimens of
V. pusilla Macq. labeled as taken in Milwaukee County.
Graenicher tells me that without doubt the specimens are
mislabeled. This species occurs in Cuba, Florida, and
Porto Rico.
Sericomyia Meigen
Second abdominal segment with two yellow dots, or wholly black;
hind femora of male considerably thickened . . militaris
Second abdominal segment with an interrupted yellowish fascia,
the bands oblique . chrysotoxoides
240
85.
86.
87.
1.
2.
3.
4.
5.
6.
7.
8.
9.
Wisconsin Academy of Sciences, Arts, and Letters.
S. chrysotoxoides Macq. Specimens from Manitowoc, Vilas,
Milwaukee, Dunn, Winnebago, and Jefferson Counties.
Late summer and early fall. Five females and four males.
S. militaris Walk. Price, Vilas, Milwaukee, Washington,
Dunn, and Jefferson Counties. August and September.
Condidea Coquilett
C. lata Coq. Two specimens, Washington and Vilas Counties.
Eristalis Latreille
*
Third segment of abdomen wholly shining .
Third segment with opaque black markings .
Eyes with small round black dots, pile not confined to a ver¬
tical stripe; arista bare; abdomen entirely shining .
. aeneus
Eyes without such black dots .
Legs black, except hind tarsi, which are distinctly reddish; pile
of lower part of face black; large hairy species, bumble¬
bee-like . flavipes
Legs unusually dark, but hind tarsi not reddish, pile of face
quite heavy, whitish or light brownish .
Basal part of hind tibiae light colored, thorax brightly shining
. latifrons
Hind tibiae entirely dark brownish or blackish, thorax usually
somewhat dull; drone fly . tenax
Thorax with distinct one or more light dusted or dark cross¬
bands . . .
Thorax without such transverse bands .
Arista bare, pile of lower part of front and frontal triangle
wholly white . vinetorum
Arista pubescent at base, pile of front and frontal triangle
black . transversus
Thorax entirely shining, or if not entirely shining the third
segment of the abdomen has a rounded, isolated, anterior
dorsal opaque spot . . . . .
Thorax not entirely shining, no such isolated spot on the third
segment . . . . .
Arista bare, basal two-thirds of wings brown in front, pile
dusky, third joint of antennae large . . . . compactus
Arista pubescent at base, third joint of antennae not unusually
large .
Third segment of abdomen with an entire velvety cross-band,
wings with a distinct brown spot . saxorvt-m
Third segment without an entire velvety cross-band, wings
hyaline or nearly so . .
2
5
3
4
6
7
8
11
9
10
Fluke — Syrphidae of Wisconsin.
241
10. iScutellum with more or less black pile on the dorsum, thorax
usually with indistinct oqaque black stripes; fourth seg¬
ment of abdomen with a rounded opaque spot . . dimidiatus
Scutellum without black pile, entirely yellow, thorax entirely
shining; fourth segment of abdomen shining, no opaque
markings . latifrons
11. Thickly pilose species, thorax deep opaque black . bastardi
Thinly pilose species, thorax with brownish vittae . 12
12. Basal two-thirds of basal joint of middle tarsi light-colored. . . .
. arbustorum
Middle metatarsi wholly black . meigenii
88. E. tenax Linn. Grant, Pierce, Milwaukee, Washington,
Douglas, and Dane Counties. Mid-summer to November.
Figure 61.
89. E. aeneus Scopoli. Two specimens from Milwaukee and Dane
Counties.
90. E. dimidiatus Wied. Cosmopolitan throughout the season.
91. E. saxorum Wied. One female from Milwaukee County.
92. E. latifrons. Loew. Dane, Pierce, and Milwaukee Counties.
Late summer and fall. Scarce.
93. E. meigenii Wied. Cosmopolitan from early spring to late
fall. Figures 62, 63.
94. E. arbustorum Linn. Cosmopolitan in the southern part of
Wisconsin. Early spring to late fall. Figures 64, 65, 66.
95. E. bastardii Macq. Common from very early spring to late
fall, in nearly every part of the State.
96. E. flavipes Walk. Milwaukee, Washington, Burnett, Iron,
Douglas, Jefferson, Winnebago, and Dane Counties.
August and September only. Figure 67.
97. E. compact us Walk. Two males from Tomah, Wis., April 6
and 24, 1910.
98. E. transversus Wied. Cosmopolitan. June to mid-October.
99. E. vinetorum Fabr. Two specimens from Milwaukee County.
Tropidia Meigen
100. T. quadrat a Say. Vernon, Buffalo, Milwaukee, Washington,
Ashland, Manitowoc, Dane, and Jefferson Counties. June
to October. Figure 68.
16— S. A. L.
242
Wisconsin Academy of Sciences, Arts, and Letters.
Helophilus
Table of species
1. Face, between oral opening and base of antennae, with a bare
shining stripe, without pile or pollen . . .
Face without such shining stripe, entirely covered with pollen.
t. Abdominal bands on third and fourth segments distinctly in*
terrupted or entirely absent; facial stripe deep black-
shining .
Abdominal bands on third and fourth segments always present,
usually not interrupted; facial stripe light-colored, or at
most reddish, not entirely black. . . . .
3. Pile of scutellum entirely yellow, fourth abdominal segment
black, without yellowish spots or bands.... . . bruesi
Pile of scutellum black on disk, fourth segment with spots or
whitish cross-band . . . . . obscurus
4. First two joints of antennae black, third usually so; front of
female black pilose only near the ocelli, front of male
broad . latifrons
First two joints of antennae reddish, third reddish at least
on the under side; front of female black pilose, front of
male narrow above . . . similis
>5. Antennae entirely reddish or yellowish . . . . .
Antennae dark, at least first two joints blackish .
6. Abdomen oval, larger yellowish species, with distinct yellowish
cross-bands, femora mostly yellowish . laetus
Abdomen more slender, with nearly parallel sides; darker
species, cross-bands more whitish .
7. Hind coxae of male with a long, stout process below; only a
slight indication of this spur in female; abdominal cross¬
bands of female attain lateral margin . chrysostomus
Hind coxae without such protuberance .
8. Third and fourth segments of male mostly black-shining, spots
small or indistinct; cross-bands on third segment of female
do not attain lateral margin (if so only slightly), hind
femora of female with long black spots . bilinearis
Third and fourth segments of both sexes with interrupted lunate
cross-bands which distinctly attain side margins; hind
femora of female largely yellow . lunulatus
9. Face jutting forward into a sharp cone, first and second joints
of antennae black, third yellowish red. . . . conostomus
Face not jutting forward into a sharp cone, only moderately
produced . . . . .
10. Abdomen velvet-like black, each segment with a triangular
black-shining spot in the middle, and a hoary spot on each
side . porcus
Fluke — Syrphidae of Wisconsin. 243
Abdomen with definite yellowish or whitish cross-bands...... 11
11. Hind coxae of male with a stout process below; grey thoracic :
stripes of female narrower than intervening black ones..
. chrysostomus
Hind coxae without such process; grey thoracic stripes of fe¬
male broader than the black lines. ............ .distinctus
101. H. latifrons Loew. Walworth, Milwaukee, Washington,
Door, Winnebago, Manitowoc, and Dane Counties. Early
spring to late fall. Figures 69, 70.
102. H. similis Macq. Dane, Manitowoc, Vilas, Milwaukee, Door,
and Washington Counties. Figure 71.
103. H. obsciDrus Loew. Vilas and Dunn Counties. One male
and two females. July to September. Figure 72.
This species has been recorded from Colorado and Wy¬
oming. Loew’s description is rather short, but the speci¬
mens agree in almost every detail. The specimen from
Wyoming noted by Moodie “ agrees with the description
save there is no brown ring on the hind tibiae, the abdo¬
men is more shining and some of the cross-bands are not
interrupted”. Notes on the Wisconsin specimens are as
follows :
Male : Antennae dark reddish, almost blackish on up¬
per side of the third joint. Face with a black-shining
stripe, without pollen, extending from oral margin to an¬
tennae, going around the base of the antennae and ex¬
tending quickly to a point in the front, otherwise the face
covered with yellowish pollen and pile; vertex opaque
black with black pile. Cheeks shining black and con¬
nected by a narrow margin with the facial stripe. Front
a trifle wider than that of similis. Thorax opaque black
with yellowish pile and four yellow pollinose stripes; the
median ones are subinterrupted about midway by a cup-
like laterad depression, the stripes then nearly fade out
but suddenly broaden before reaching the scutellum, which
is translucent yellow. Abdomen opaque black (slightly
shining on the posterior margins of the segments) with
three distinctly interrupted yellow crossbands; first seg¬
ment opaque black, lateral margins yellow; second seg¬
ment with large yellow spots separated by about one half
their own width, reaching the lateral margins and joining
the yellow of the first segment; spots on third segment a
trifle smaller, touching the fore margins of the segment
hut reaching the lateral margins by the anterior corners
244 Wisconsin Academy of Sciences , Arts, and Letters.
only, the black separating the spots being broader an¬
teriorly and posteriorly than midway. The band on the
fifth segment smaller and not quite reaching the lateral
margins, distinctly interrupted; hypopygium yellow;
there is a small posterior, median, elongated, slightly red¬
dish spot on the second segment. Pile on scutellum black
on disk, yellow and longer on the margin ; that of abdomen
yellow or black according to ground color. Legs black
with the following parts yellowish or reddish: extreme
base and pre-apical spot of hind femora, base and a me¬
dian ring on the hind tibiae ; tips of the four anterior fe¬
mora, base of front tibiae, middle tibiae, and first two
joints of mid-tarsi. Pile underneath the hind tarsi dis¬
tinctly ferruginous. Sixth vein only slightly sinuate.
Female : Similar with the following exceptions : Front
with black of vertex connecting with the shining spot at
base of antennae; facial stripe scarcely reaching base of
antennae. Abdominal spots a trifle smaller, fifth segment
with spots similar to those on the fourth, no posterior spot
on second segment. In one female the first segment is
entirely black.
104. H. bruesi Graenicher. Vilas County. One female. Figure
73. Described in Bull. Wis. Nat. Hist. Soc. 8: 40. 1910.
105. H . laetus Loew. Dane, Milwaukee, Washington, Fond du
Lac, Eau Claire, Pierce, and Douglas Counties. Common
June to September.
106. H. chrysostomus Wied. Dane, Milwaukee, and Washington
Counties. Late June to early September. Figures 74, 75.
The male is easily told by the spur on the hind coxae,
the female has only a slight indication of this spur but it
is evident enough to distinguish the species after a little
practice. Female is easily separated from the female of
bilinearis by the pollinose bands on the third and fourth
segments which reach the side margins. The legs of
chrysostomus — female — are darker and the abdomen is
slightly constricted about the middle of the second seg¬
ment, making the species easily distinguishable from the
same sex of lunulatus. In fact, the constriction of the
abdomen is characteristic of this species and separates it
from other members of this genus occurring in Wisconsin.
107. H. bilinearis Will. Milwaukee and Dane Counties. Figures
76, 77. Caught only in May or June.
Fluke — Syrphidae of Wisconsin.
245
Female: Length 10:5 mm. Antennae yellowish,
slightly obscure above, face covered with yellow pollen
and pile, no median stripe, pile absent on median ridge ;
slightly concave below antennae and then convex to oral
margin which is slightly produced, lower part of face
shining black ; front black with greyish pollen below, pile
black. Four broad silvery pollinose stripes on the thorax
expanding so as to meet before reaching the scutellum, the
lateral intermediate black stripes narrower than the me¬
dian ones. Scutellum yellow-translucent, pile yellowish
or whitish. First segment of abdomen entirely silverly
pollinose, second segment opaque black, shining poster¬
iorly, with a median, interrupted pollinose cross-band
which expands laterally to reach the fore margin of the
segment. Third and fourth segments mostly shining with
whitish pollinose bands interrupted (only slightly on the
fourth) and not reaching the side margins, fifth entirely
white-pollinose. Fourth segment with a posterior reddish
cross-band, third has posterior angles also reddish. Legs
yellowish with the following dark markings: inner and
outer sides of all femora with irregular spots, more pro¬
nounced on hind femora; inner side of hind tibiae; pos¬
terior tarsi and last two joints of middle tarsi. Wings
hyaline.
108. H. lunulatus Meigen. Numerous specimens of this interest¬
ing species taken in Jefferson and Dane Counties. Early
spring only. Figures 78, 79.
109. H. conostomus Will. Dane and Milwaukee Counties. A
very common species in June and September. Figure 80.
110. H. porous Walker. One specimen from Douglas County,
July, 1909.
Mallota Meigen
Eyes pilose; more or less of the base of the abdomen with yellow
• • • . . . . . posticata
Eyes bare; wholly black pilose . cimMciformis
111. M. cimbiciformis Fallen. Manitowoc, Vilas, Milwaukee,
Dane, and Washington Counties. May to July.
112. M. posticata Fabr. Specimens from Dane, Milwaukee, Vilas,
and Washington Counties.
Triodonta Williston
113. T. curvipes Loew. Three specimens in the Milwaukee Public
Museum taken in Milwaukee County.
246 Wisconsin Academy of Sciences, Arts , and Letters .
Teuchocnemis Osten Sacken
114. T. lit ur at us Loew. Milwaukee and Dane Counties. May.
Scarce.
Syritta St. Fargeau and Serville
115. S. pipiens Linn. Cosmopolitan, early spring to late fall.
Figure 81.
Xylota Meigen
1. Legs wholly black; hind femora much thickened .
Legs not wholly black .
2. Abdomen chiefly red; the second segment with a basal black
triangle; wings nearly hyaline . pigra
Wholly black; wings black . chalybea
3. Front and middle legs and a large part of hind femora yellow;
large species . curvipes
Front legs in large part black .
4. Abdomen long, slender, with two yellow spots on second seg¬
ment in male, absent in female . angustiventris
Abdomen not unusually slender .
5. Abdomen with two pairs of yellowish spots .
Abdomen wholly black .
6. Arista wholly black . . . obscura
Arista luteous at the base . anthreas
7. Hind femora much thickened; smaller species . fraudulosa
Hind femora moderately thickened .
8. Spots on second and third segments of the abdomen distinctly
separated by a broad black line . ejuncida
Line separating the abdominal spots nearly obsolete. subfasciata
2
3
4
5
7
6
8
116. X. angustiventris Loew. Milwaukee and Dane Counties.
May to late summer. Figure 82. Common in woods.
117. X. anthreas Walker. One female from Douglas County.
118. X. chalybea Wied. Milwaukee and Dane Counties.
119. X. curvipes Loew. One male from Vilas County.
120. X. ejuncida Say. Numerous specimens from Manitowoc,
Vilas, Milwaukee, Marathon, Douglas, Burnett, Polk,
Door, St. Croix, Washington, Waushara, and Dane Coun¬
ties. Late summer.
121. X. fraudulosa Loew. St. Croix, Pierce, Dane, Milwaukee,
Washington, and Douglas Counties.
122. X. obscura Loew. One specimen, Sheboygan County.
123. X. pigra Fabr. One female, July, 1912, from Vilas County.
124. X. subfasciata Loew. One male, August 2, 1918, from Dane
County. Figure 83.
Fluke — Syrphidae of Wisconsin .
247
Male: Antennae slightly reddish beneath the base of
the third joint. Face and frontal triangle black with
white pollen. Thorax and scutellum shining bronze. First
segment of abdomen mostly shining black; second and
third segments mostly yellow, base of second with an
opaque black cross-band with a triangular projection into
the yellow portion, continued as a light brown median line
to the narrow brownish posterior band; third segment
with no anterior dark band but with an indistinct brown
posterior band covering about one third of the segment;
there is also a slight indication of a median dark line;
fourth and fifth segments shining black. Femora black
except at extreme tips; tibiae and first three joints of tarsi
light yellow in front legs; tibiae and first two joints of
middle legs light brownish; hind tibiae and tarsi mostly
dark. A dense patch of short white hair on hind coxae,
less on the other coxae ; distinct black small spines at tip
of middle tibiae. “ Coxal” spur quite prominent, tibial
spur rounded. Squamae and halteres light yellow.
Brachypalpus Macquart
125. B. frontosus Loew. Two males from Dane County. May.
Crioprora Osten Sacken
126. O. cyanogaster Loew. One specimen, Milwaukee County.
Criorhina Meigen
'Face produced downwards and forwards; males dichoptic; ground
color black, thickly pilose . . . sub-genus (Criorhina)
Face produced downwards, convex below, males holoptic, ground
color with yellow markings, moderately pilose . . .
. subgenus Cynorhina
Face produced downwards, convex below; eyes dichoptic, moder¬
ately pilose, in large part yellow . sub-genus Somula
Cynorhina Williston
Face with a median black stripe; second segment of the abdomen
on the sides, continued more narrowly on the third, yellow. . . .
. badia
Face yellow, without a black median stripe; abdomen black, except
a large part of the fourth and the whole of the fifth segments,
which are bright orange yellow . . . . analis
248 Wisconsin Academy of Sciences , Arts , and Letters.
127. C. analis Macq. Dane and Milwaukee Counties. Rare.
128. C. ~badia Walker. Milwaukee County.
129. Somula decora Macq. Several specimens from Milwaukee
and Dane Counties. Figure 84.
Milesia Latreille
130. M. virginiensis Drury. Milwaukee County only.
Spilomyia Meigen
1. Second joint of antennae much longer than the first; second
segment of the abdomen with a broad, arcuate yellow
band, the third and fourth with a slender median inter¬
rupted one . . . . quadrifasciata
Second joint of the antennae not much longer than the first;
abdomen not so marked . . . 2
2. Abdomen yellow; first segment, except the lateral angles, black,
each of the following segments with two narrow black
bands, the one on the anterior margin entire, the others
interrupted . hamifera
Abdomen black with yellow cross-bands . 3
3. First and second abdominal segments wholly black, third with
a posterior cross-band, fourth with two, the anterior one
interrupted; abdomen broadly oval . fusca
First segment with a yellow side spot, the following each with
two cross-bands . longicornis
S. fusca Loew. Specimens from Milwaukee, Ozaukee, Bur¬
nett, Washington, and Vilas Counties. Late summer.
Figure 85.
S. hamifera Loew. One specimen, Milwaukee County.
S. longicornis Loew. Milwaukee, Washington, Dane, and
Manitowoc Counties. Late summer and early fall.
S. quadrifasciata Say. Taken in Milwaukee, Washington,
Dane, and Dunn Counties. August and September. Fig¬
ure 86. I have specimens varying from 12 mm. to 16 and
17 mm. in length.
Temnostoma St. Fargeau and Serville
Abdomen with three or four pollinose cross-bands of nearly equal
width . . . bombylans
Abdomen broadly oval, with more than four cross-bands, of unequal
width; all, or at least the front, femora broadly black, .aequalis
131.
132.
133.
134.
Fluke — Syrphidae of Wisconsin.
249
135. T. aequalis Loew. Specimens taken in Milwaukee, Manito¬
woc, Douglas, and Sheboygan Counties.
136. T. bombylans Fabr. Milwaukee, Washington, Douglas, and
Manitowoc Counties.
Sphiximorpha Bondani
137. S. dbhreviata Loew. Only a few specimens taken in Dane,
Milwaukee, and Washington Counties.
Brief Bibliography
The following publications are given mainly to indicate those
that have been of the most help in the preparation of this paper.
Aldrich, J. M. 1905. A catalogue of North American Diptera.
Smithsonian Miscell. Coll. no. 1444.
Barth, G. P. 1907. The occurrence of the Syrphid fly Condidea lata
Coq. in Wisconsin. Bull. Wis. Nat. Hist. Soc. 5: 161.
Coquillett, D. W. 1900. Diptera of the Harriman Alaska Exposition.
Proc. Wash. Acad. Sci. 11: 389-464.
- . 1904. Notes on the Syrphid fly Pipiza radicum Walsh and
Riley. Proc. Entom. Soc. Wash. 6: 200.
— . 1907. New genera and species of Diptera. Canad. Entom.
39: 75.
Forbes, S. A. 1905. Corn feeding Syrphus fly. Rept. Entom. Ill.,
23: 162-163.
Graenicher, S. 1900. Syrphidae of Milwaukee County. Bull. Wis.
Nat. Hist. Soc., July, 167-177.
; - . 1910. A preliminary list of the flies of Wisconsin belonging
to the Families Bombyliidae, Syrphidae, and Conopidae. Bull.
Wis. Nat. Hist. Soc. 8: 3 2-43. Lists 92 species.
- - . 1910. Some rare Diptera. Canad. Entom. 42: 28-29.
Hunter, W. D. 1896. A contribution to the knowledge of North
American Syrphidae. I. Canad. Entom. 28: 87-101.
- . 1896. A summary of the genus Chilosia Meigen in North
America, with descriptions of new species. Canad. Entom. 28:
227-233.
— — . 1897. A contribution to the knowledge of North American
Syrphidae. II. Canad. Entom. 29: 121-144.
Jones, P. R. 1907. Notes on some little known North American
Syrphidae. Entom. News 18: 238—241.
- . 1907. A preliminary list of Nebraska Syrphidae with descrip¬
tions of new species. Jour. N. Y. Entom. Soc. 15: 87-100.
Johnson, C. W. 1907. Some North American Syrphidae. Psyche
14: 75-80.
250 Wisconsin Academy of Sciences, Arts, and Letters.
- . 1910. Some additions to Dipteran fauna of New England.
Psyche 17: 229-231.
— — . 1916. On the Criorhinja< intersistens Walker and an allied spe¬
cies. Entom. News 24: 293-5.
- . 1916. Some New England Syrphidae. Psyche 23: 75.
Metcalf, C. la. 1913. Syrphidae of Ohio. Ohio Biol. Surv. J: no. 1.
- . 1916. Syrphidae of Maine. Maine Agr. Exp. Sta. Bull. 253.
- . 1917. Syrphidae of Maine. Maine Agr. Exp. Sta. Bull 263.
Moodie, R. L. 1905. A new Milesia from Arizona with notes on
some Wyoming Syrphidae. Entom. News 16: 138-143.
Osbnm, R. C. 1910. Studies on Syrphidae. I. Syrphus arcuatus
Fall, and a related new species. Jour. N. Y. Entom. Soc. 18: 55.
- . 1915. Studies on Syrphidae. IV. Species of Eristalis new to
America with notes on others. Jour. N. Y. Entom. Soc. 23: 139—
145.
Robertson, C. 1901. Some new Diptera. Canad. Entom. 33: 284.
Verrall, G. H. 19 01. British flies. Vol. VIII. London. Indispens¬
able in a thorough study of the family.
Walton, W. R. 1911. Notes on Pennsylvanian Diptera, with two
new species of Syrphidae. Entom. News 22: 318—320.
Williston, S. W. 188 6. .Synopsis of the North American Syrphidae.
U. S. Nat. Mus. Bull. 31. This work is the basis of all North
American studies of Syrphidae. I am always using it; most of
the keys in this paper are based on those of Williston’s.
- . 1908. Manual of North American Diptera, 3rd edition. Syr¬
phidae on pages 246—260. I have quoted almost verbatim from
this work for the table of genera given above.
TRANS. WIS. ACAD., VOL. XX
PLATE V
FLUKE— SYRPHIDAE
Fluke — Syrphidae of Wisconsin.
251
Fig. 1.
Fig. 2.
Fig. 3.
Fig. 4.
Fig. 5.
Fig. .6.
Fig. 7.
Fig. 8.
Fig. 9.
Fig. 10.
Fig. 11.
Fig. 12.
Fig. 13.
Fig. 14.
Fig. 15.
Fig. 16.
Fig. 17.
Fig. 18.
Fig. 19.
Fig. 20.
Fig. j21.
Fig. 22.
Fig. 23.
Fig. 24.
Fig. 25.
Fig. 26.
Fig. 27.
Fig. 28.
Fig. 29.
Fig. 30.
Fig. 31.
Fig. 32.
Fig. 33.
Fig. 34.
Fig. 35.
Fig. 36.
'Fig. 37.
Fig. 38.
Fig. 39.
Fig. 40.
Fig. 41.
Fig. 42.
Fig. 43.
Fig. 44.
EXPLANATION OF PLATES
PLATE V
Microdon tristis : Scutellum (redrawn after Metcalf).
M. tristis: antenna (redrawn after Metcalf).
M. globosus: antenna (after Williston).
Chrysotoxum pubescens : head of female.
Chrysogaster nigripes: antenna.
G. nitida: wing.
G. nitida : antenna.
G. pictipennis : wing.
G. pictipeinnis : head of female.
C. pulchella: wing.
G. pulchella: head of female.
Pipiza calcar ata: hind leg, a. “coxal process.”
P. salax? : head of male.
P. radicum ; antenna.
P. pisticoides: antenna.
Paragus bicolor: wing.
P. bicolor: antenna.
P. tibialis : antenna.
P. tibialis: wing.
Ghilosia prima: face of male.
G , cyanescens : face of male.
Baccha lemur: side of face of male.
B. fascipennis Wied.: abdomen of female.
Pyrophaena granditarsis : abdomen of female.
P. granditarsis : front tibia and tarsi of male.
P. rosarum var. : abdomen of male.
Platychirus immarginatus : front leg of male.
P. peltatus: front leg of male.
P. perpallidus: front leg of male.
P. quadratus: middle leg of male.
P. scambus: front leg of male.
P. species?: front leg of male.
P. species?: middle leg of male.
Melanostoma obscurum: face of male.
M. angpistatum: abdomen of female.
M. mellinum: abdomen of female.
Syrphus fisherii: 1st and second segments of abdomen of female.
S. fisherii: 1st and second segments of abdomen of male.
5. sp. no. 1: profile of face of male.
6. arcuatus : wing.
S. perplexus : wing.
$. sp. no. 2 : abdomen of male.
S. nitens: antenna.
S. knabi : antenna.
252
Wisconsin Academy of Sciences , Arts, and Letters.
fig. 45.
Fig. 46.
Fig. 47.
Fig. 48.
Fig. 49.
Fig. 50.
iFig. 51.
Fig. 52.
Fig. 53.
Fig. 54.
Fig. 55.
Fig. 56.
Fig. 57.
Fig. 58.
Fig. 59.
Fig. .60.
Fig. 61.
Fig. 62.
Fig. 63.
Fig. 64.
Fig. 65.
Fig. 66.
Fig. 67.
Fig. 68.
Fig. 69.
Fig. 70.
Fig. 71.
FIG. 72.
Fig. 73.
Fig. 74.
Fig. 75.
Fig. 76.
Fig. 77.
Fig. 78.
Fig. 79.
Fig. 80.
Fig. 81.
Fig. 82.
Fig. 83.
Fig. 84.
Fig. 85.
Fig. 86.
PLATE VI
Syrphus xanthostomus : abdomen of female.
Xantho gramma aenea: antenna of female.
X. imarginata: antenna of female.
X. fragila n. sp.: profile of male.
X. fragila: abdomen of female.
Mesogramma geminata: hind leg of male.
M. geminata: hind leg of female
M. geminata: wing of male.
M. marginata : wing of female.
M. marginata: hind leg of male.
M. marginata: hind leg of female.
Sphaerophoria cylindrica: profile of head and thorax of male.
S. scripta: tip of abdomen of male from side.
Rhyngia nasica: head of female.
Brachyopa notata: head of female.
Volucella evecta: antenna.
Eristalis tenax: antenna.
E. meigenii: middle tibia and tarsus.
E. meigenii : wing.
E. arhustorum: abdomen of male, showing yellow areas (white
portion) opaque bands (solid black) and shining portions
(stippled area).
E. art)ustorum: female.
E. arbustorum: middle tibia and tarsus.
E. flavipes: antenna.
Tropidia quadrata: antenna.
Helophilus latifrons: antenna.
H. latifrons: head of male from above.
H. similis: head of male from above.
H. ot)seurus: head of male.
H. hruesi: head of female.
H. chrysostomus : hind leg of male.
H. chrysostomus: abdomen of female.
H. Mlinearis: abdomen of male.
H. bilinearis: abdomen of female.
H. lunulatus: head of male.
H. lunulatus: abdomen of male.
H. conostomus : head of male.
Syritta pipiens : hind leg.
Xylota angustiventris : face of female.
X. suhfasciata: abdomen of male.
Somula decora: face of male.
Spilomyia fusca: antenna.
S. quadrifasciata : antenna.
TRANS. WIS. ACAD,, VOL. XX
PLATE VI
FLUKE— -SYRPHIDJ3
Fluke — Syrphidae of Wisconsin.
253
APPENDIX
138: Microdon fuscipennis. Macq. Three specimens from Door County;
August, 1919 and August, 1920.
CMlosia comosa Loew. I am mistaken in the identification of this
specimen; it should be C. prima Hunter.
139: Syrphus venustus Meigen. Two females caught in the woods along
Lake Michigan shore line in Door County. June 20, 1921.
140: Helophilus distinctus Williston. Washington and Milwaukee Coun¬
ties.
141: Xylota vecors O. S. June 17, 1921, Door County.
142: Temnostoma alternans Loew. Two specimens, male and female,
Manitowoc County.
SOME EXPERIMENTS WITH THE LARVA OF THE BEE-
MOTH, GALLERIA MELLONELLA L
Joy E. Andrews
The following is an account of some experiments with the wax-
worm, or bee-moth. The effects of different degrees of tempera¬
ture on the length of the larval life, of light on both larvae and
imago, and of different kinds of food on the larvae were tried.
These experiments were carried out in the zoological laboratory of
the University of Wisconsin.
As a basis for the work, an account of the moth by F. B. Pad-
dock of the Texas Experiment Station1 was used. The life his¬
tory of the moth as given by him was made use of, but the habits
of the moths observed differed somewhat from those noted by
Paddock. He says that “moths emerge entirely at night”, name¬
ly, during the early part of the evening; but in my work it was
found that while this is usually true, yet emergence does occur
during the day. Cases of emergence have been noted both in the
morning and in the afternoon. Consequently a difference was found
in the mating of the moths, which Paddock says takes place at
night. Since mating occurs soon after emergence, I have ob¬
served moths to mate in the daytime as well as at night. These
differences may have resulted from the artificial conditions under
which fhe moths were reared. The constant-temperature rooms
in which the larvae were kept were heated by electric lights, with
the result that the rooms were light at all times, although the jars
containing the moths were shielded from the direct light by pieces
of paper.
Another point of difference was in the shape of the egg. Pad-
dock says that the eggs are elliptical, but I have found them to
be also spherical and pear-shaped. The shape of the eggs seemed
to depend largely upon the place where they were laid: they
1 Paddock, F. B. The life history and control of the bee-moth or wax-worm.
Investigations Pertaining to Texas Bee-Keeping. 1913.
255
256 Wisconsin Academy of Sciences , Arts, and Letters.
might be crowded into crevices, in the comb or in boards, and
flattened on one, two, or three sides. The eggs of one female
might show all these variations.
Effect of Temperature
In trying the effect of temperature on the length of the life
history of G. mellonella L., four different temperatures were
used: namely, room temperature (24°C.), 29°C., 37°C., and 45°C.
Boom temperature. Eggs laid from September 23 to September
28, 1918, hatched about October 23, and the larvae were placed
in glass finger bowls and small homeopathic vials containing brood
wax. The dishes were left in the laboratory, where the tempera¬
ture was about 24° C. The larvae grew very slowly and were
rather inactive during the winter months. On February 13, 1917,
they averaged 6.5 mm. in length; on June 4, 1917, the average
length was 10 mm., or about one-third the mature length. As a
result of their slow growth, only part of a generation, the egg
stage, could be used in comparing their life history with that of
the larvae kept at higher temperatures. In the experiment not
less than 20 larvae were used. From September 23, 1916, to June
4, 1917, only part of a life cycle was completed, the egg stage and
part of the larval life.
29° C. A temperature of 29 °C. was maintained in a constant-
temperature compartment by using a 16 candle power carbon elec¬
tric light globe.2 On March 10, 1917, half-grown larvae were placed
in several large crystallizing dishes in which was a brood comb for
food ; these dishes were placed in the constant temperature compart¬
ment. A large evaporating dish filled with water was placed in
the compartment to keep the atmosphere moist. The larvae grew
rapidly and were very active. Soon after pupation occurred,
moths of average size hatched, first males and then females. Mat¬
ing took place, and from the eggs laid, larvae hatched. The aver¬
age life of the moths was five days for females, seven days for
males. From this point on, the moths went through their life
cycle rapidly; from March 10 to May 29, 1917, when the last ob¬
servation was made, there had been three generations. The num¬
ber of larvae and moths in each generation was very large, being
not less than five hundred.
2 The temperatures of 29°, 37°, and 45° C. were obtained in the three com¬
partments respectively by experimenting with electric light globes of different
powers until those continuously giving the desired temperature were procured.
Andrews — Experiments with Larva of the Bee Moth. 257
37° C. In a second const ant-temperature compartment the tem¬
perature of 37° was obtained by using a 40- watt and a 60- watt
Mazda light. Newly hatched larvae were put in tall glass jars
containing brood comb and placed in this constant temperature
compartment on October 28, 1916. They seemed to be normal and
throve well. Pupation took place, the moths emerged, mating
occurred, and eggs were laid. About fifty per cent, of the eggs
did not hatch, but seemed dried up by the heat; the rest hatched
into normal larvae. The length of life of the moths was about four
days for females and six days for males. The females sometimes
emerged before the males, a condition which was never found to
obtain among the moths kept at 29 °C. From the time when the
larvae were put in the incubator until May 29, 1917, the life cycle
had been completed five times. At no time during the experiment
were there less than two hundred and fifty larvae in our dishes.
At each instance of egg-laying under this temperature, the same
difficulty of the eggs not hatching was found. The heat did not
seem to affect the young larvae after they had hatched.
45° G. In the third constant-temperature compartment the de¬
gree of temperature desired was produced by using two Mazda
globes of 60 watts each, and one of 40 watts. On March 10,
1917, nearly mature larvae were put in glass jars containing
brood comb, and the latter were placed in this compartment. A
dish of water was also put in the incubator. At the end of forty-
eight hours, the larvae appeared healthy and were spinning co¬
coons. A day later they seemed to be dying, and ninety-six hours
after they were first put in, the larvae were all dead. About
twenty-five were used for the experiment. It did not seem neces¬
sary to repeat it, because if the older larvae, which are hardier
than moths, young larvae, or eggs, could not withstand a tempe¬
rature of 45°C., the other forms surely could not.
Table 1 gives the result of the experiments.
17— S. A. L.
258 Wisconsin Academy of Sciences, Arts, and Letters.
Table 1. — The Effect of Temperature on the Length of Life History of
G. mellonella L.
(In each case only the average number of days is given.)
* Larvae at room temperature hatched October 19 to 23, 1916, had not com¬
pleted their larval life on June 2, 1917.
The effect of heat on the color of the moths was not especially
marked, and there was considerable variation between individual
moths in the same generation, but other things being equal (such
as amount of moisture and food), the moths kept at 37° tended
to be darker in color than those kept at 29 °C. The latter were
more often of a light tan color, while the former were dark gray
tinged with brown.
Effect of Light
The larvae of Galleria mellonella respond differently to light
stimuli at different stages in their development. When they are
newly hatched, they seem to be neither positively nor negatively
phototropic. For experimentation with these young larvae a 40-
watt Mazda electric light bulb was used. When the rays of the
bulb were directed upon the larvae which had been kept in the
dark, they continued their wandering, making no effort to seek the
dark. If they did happen to run out of the circle of light, they
did not turn back into it. Sunlight also was found neither to
repel nor to attract them. It is probable, however, that they are
irritated by light, because they seem to move faster and in a more
aimless fashion when exposed to light rays.
The larvae remain thus unresponsive to light stimuli until they
are 2 to 5 mm. in length, when they become negatively photo¬
tropic. On being put under a 40-watt Mazda globe they at once
Andrews — Experiments with Larva of the Bee Moth. 259
crawled from the light and tried to find a dark hiding place. This
was true in the case of stimuli of varying intensities, although
they are less sensitive to weaker stimuli. From the time they
were 5 mm. long the larvae were found to be affected more and
more by light rays. When taken out of the comb, placed on the
table, and the light of a 40-watt Mazda globe turned on them, the
older larvae invariably crawled into some dark crevice. Sunlight
affected them in the same manner. When comb containing lar¬
vae was put in the sunlight or under the 40-watt globe, the lar¬
vae which were on top scurried through their tunnels to the bot¬
tom of the comb.
Light stimuli, both artificial and sunlight, did not affect the
pupae at all. They always responded, however, to mechanical
stimuli, such as jarring or the touch of a dissecting needle, by
moving their abdomens.
Newly emerged moths were not found to be very sensitive to
light, but those several days old, especially females which were
ovipositing, responded to light and would fly up from their nest¬
ing places in the dishes on being put into sunlight or into the
light from an electric globe. Sunlight seemed to disturb them
more than artificial light.
Feeding Experiments
Problems in the feeding of Galleria mellonella were undertaken ;
bee-bread, dead moths, dead bees, and larvae being used for food
as well as brood comb.
PoUen. Since it has been held that the larvae of Galleria mel¬
lonella eat beeswax for the pollen it contains and derive no nour¬
ishment from the wax itself, the question was raised as to whether
the larvae would mature on bee-bread alone.
In most of the brood-comb used, a great many of the cells were
partly filled with tightly packed bee-bread, the lumps of which
were removed from the cells, and placed in Petri dishes with sev¬
eral larvae a day or two old; the dishes were placed in the incu¬
bator at 37 °C. All these larvae died. Then the effect of mash¬
ing up the bee-bread was tried, and other young larvae were put
in the dish with the same result as before. Finally it was dis¬
covered that part of the bee-bread was stale ; after that it was cut
into slices and only the freshest portion was used. Upon the bee-
bread prepared in this manner, three larvae out of the hundred
and one used in the experiment were raised. They acted nor-
260 Wisconsin Academy of Sciences , Arts, and Letters.
mally, built tunnels of pieces of bee-bread and their silk, and grew
rapidly. The average length of the life history from the newly
hatched larva to the moth was thirty-nine days. Two of the
moths were males and one a female.
Dead Bees. A similar experiment with dead bees was tried,
but none of the young larvae lived. Both whole and crushed
bees were placed in the dish, and in spite of the abundance of
food, all the larvae died.
Larvae ,. If larvae were kept in a crowded condition, it was
noticed that the nearly mature larvae would eat pupae in the
cocoons, as soon as pupation had taken place. It was once noted
that when a larva which had been accidentally injured was put
back into the dish it was immediately eaten by the others. Hence
the idea arose of ascertaining whether larvae were so cannibal¬
istic that they could be raised on animal food alone.
Seventeen larvae, ranging in size from 10 to 13 mm., were
placed in a Stender dish, and a lacerated larva was placed in the
same dish. The dish was then placed in the constant-tempera¬
ture room at 37 °C. In a short time the larvae began to eat the in¬
jured one, which, however attempted to defend itself. They finally
consumed all of the larva except the head and the brown chitinous
part of the first thoracic segment. Then the larvae wandered
aimlessly about the dish and after two or three hours were raven¬
ously hungry. At the end of twenty-four hours the larvae were
observed again, and only fourteen were left. The heads of the
devoured larvae were lying with the refuse in the bottom of the
dish. Two days later twelve larvae remained, the following day
but ten. At the end of six days only two larvae were to be
found; one of these was dead, while the other was in a small co¬
coon made of excrement, the heads and half-eaten bodies of lar¬
vae, and silk. The larva did not pupate but seemed to dry up
in the act of transforming into a pupa.
At another time thirty-six larvae were equally divided between
two Stender dishes and the latter were placed in the constant-
temperature room. The following day no decrease in number
was noticed in either dish, but the larvae were very restless and
showed a tendency to bite each other. A day later only etight re¬
mained in one dish and eleven in the other. After four days
more there were two larvae in one dish and one in the other.
These spun cocoons and pupated, but the pupae dried up and did
not hatch.
Andrews — Experiments with Larva of the Bee Moth. 261
Another attempt was made to raise larvae on animal food. Six¬
teen larvae, most of them 6 mm. long but a few 12 mm. in length,
were placed in a Stender dish which was put in the constant-
temperature compartment at 37° C. At the end of nine days it
was found that two of the largest larvae were left, one of these
was dead, and the other had pupated. Six days later a moth
emerged from the cocoon; it was a male, rather small and very
light in color. This male was functional, mating with an un¬
fertilized female taken from another lot of moths.
Dead Moths. One day in cleaning out a dish which contained
larvae about one fourth grown, several dead moths were found
having young larvae in them apparently living on the dead tissues
of the moths. The bodies of over one half of the other moths ex¬
amined were mere shells, all the interior portion having been eaten
out. To ascertain whether larvae could be raised on this food
alone, twelve young larvae, two or three days old, were placed in
a Petri dish with a dozen dead moths, and the dish placed in the
incubator at 37° C. All these larvae died within forty-eight hours.
The experiment was tried again, using fifteen larvae and the same
dish and moths, with the same result. Finally ten larvae were
placed in the dish with fresh moths; of these three larvae lived:
and seemed to thrive. They did not grow to be very large and at
the end of thirty days spun cocoons and pupated, being then about
J4 or % of an inch long. Unfortunately the moths did not
hatch; hence it was not known whether larvae fed on dead moths
would produce sexually mature moths or not. The experiment
has been tried a number of times since, but the larvae always
died within 48 hours.3
Zoological Laboratories,
University of Wisconsin
3 Professor W. S. Marshall tells me that he has allowed a number of moths
to remain in a dish until they were all dead. An examination of these dead
moths showed that egg's had been laid upon the body of many of them and
generally concealed between the wings and the body.
THE LENGTH OF LIFE OF THE LARVA OF THE WAX
MOTH, GALLERIA MELLONELLA L, IN ITS
DIFFERENT STADIA
Ruth W. Chase
The work carried on with the wax moth, Galleria mellonella,
was done entirely in the zoological laboratory of the University
of Wisconsin, beginning with the first of October, 1916, and con¬
tinuing into the following May. The regular food used for the
larvae was the brood-comb; some of it dry and pollen-filled, and
some of it moist with honey. An excess of honey in the comb
caused the death of several larvae, but on the other hand, young
larvae did not thrive in too dry a comb ; so a mixture was neces¬
sary. Great difficulties were experienced in keeping the young
larvae alive when separated for experimentation from their nor¬
mal habitat, either because their tender skins could not endure
handling, or because the food was too coarse for them at this age,
or because some conditions of their surroundings were unfavor¬
able. Therefore, many different plans had to be tried, for suc¬
cessful separation in the early stages was the chief factor in the
study of the life history. The length of the larval, pupal, and
adult life being known through Paddock’s work1, the object was
to obtain knowledge of ecdyses and, under these conditions, of
the length of each stadium.
Laboratory conditions were not suited to the life of the moth.
Experiment showed that temperatures of 20° to 22° C. in winter
did not result in normal life histories. The young larvae, while
feeding, grew but little, remaining extremely small for months,
and the time taken for the eggs to hatch was longer than that
given by Paddock; thus a change of conditions was found neces¬
sary. In December moths were therefore put into a constant-
temperature compartment at 35° C. (93° F.), which for at least
1 Paddock, P. B. Life history and control of the wax moth or bee moth.
Texas Agr. Exp. Sta., Bull. 155. 1913.
263
264 Wisconsin Academy of Sciences , Arts , and Letters.
an hour a day, and often longer, by opening tbe door, was cooled
to a considerably lower temperature. The length of the lives of
the moths was at once greatly shortened, and they came more
nearly to what Paddock considered normal growth and develop¬
ment. In this compartment the air was kept moist with stand¬
ing water, and the heat was regulated by a thermostat. Artificial
conditions of darkness were in part supplied by paper coverings
about the closed dishes in which the larvae were kept.
Food experiments were also made. At first in the work in the
laboratory, before temperature regulation was resorted to, bits
of dry comb were used. The larvae were, however, usually con¬
cealed by boring into it out of sight, and digging them out was
often fatal. Pieces of wax pressed hard and thin were used in¬
stead, but seemed to offer too great resistance to larval attacks.
Finally, for the second and successful attempt to raise the lar¬
vae in the constant temperature compartment, the wax of the
comb was chopped up, mixed with bee bread, soft dead moths,
and honey, and kneaded into a small loaf, which was allowed to
dry out a little and from which thin slices were cut to supply the
larvae with food. This proved the most successful method. When
about half grown, they were put on a diet of brood comb which
they then had no difficulty in eating. It was noticeable that the
newly hatched larvae, when allowed to remain together, first se¬
lected for food the dead bodies of the moths, later feeding on the
comb.
Many schemes for housing • the larvae were tried and found
ineffective. The most satisfactory quarters in which one to six
of them could live together proved to be thick glass slides with
a deep hollow center which could be closed with cover slips held
in place by a thin film of vaseline. The larvae kept in this way
were easily accessible and could be placed for observation under
the binocular microscope.
The problem of marking larvae without killing them was the
most difficult of all. While, after careful watching, it seems cer¬
tain that the larvae do not ordinarily eat their exuvia, which
were found at all stages of development after ecdyses, yet these
could not be relied on to show each molt. It was necessary to
mark the larvae with a substance which would dry quickly and
not easily rub off, with a brush which would not injure them. The
most successful brush was a bird’s feather cut to a fine point,
though single hairs were first tried. India ink was easily rubbed
Table 1.
Life histories at 35° C.
Chase— Length of Life of the Larva of the Wax Moth. 265
from partly grown larvae, and oil paint and colored balsam did
not dry quickly enough. Finally two satisfactory solutions were
found: the one, a wood-alcohol solution of shellac colored with
gentian violet, safranin, methyl green, or other aniline dye; the
other, a water solution of gum arabic colored with the same dyes.
The earliest larvae successfully to complete their life history were
marked with the first solution. On the smallest larvae, the gen¬
tian violet, turning to a maroon color when used, was found
most satisfactory, but for the older and darker larvae a bright
safranin was more conspicuous. Experimentation with a spray
or vaporizer, which would obviate the necessity of so much hand¬
ling of the larvae, was then tried. In this gum arabic was used.
After some practicing, it was found easy to mark the tiny newly
hatched larvae, transferring them with a soft camel’s hair brush
from their receptacles to a cover slip, spray them, and put them
into a slide. This method was used only for the first instar, the
rest having to be marked by brush, and it had about the same
percentage of success as the other.
It was found that eggs in masses gave a larger percentage of
larvae than if placed singly, and they were therefore allowed to
remain in masses and the larvae were separated. The most suc¬
cessful method of rearing them was to put two to six in the same
compartment. In the last experiments three or four individuals
from one brood of larvae were marked with different colors and
put together in a slide. The survivors, when old enough, were
then separated. From about a hundred marked in the last of
February, four lived to complete their life histories as shown in
table 1. A later attempt with the spray on another group was a
complete failure. The last trial was made with about fifty lar¬
vae, using slides which had been used before and were not cleaned
out, and then putting from one to six larvae, all marked alike
with vaporizer or brush, into one slide. This was the most suc¬
cessful. The larvae after the first ecdysis had, of course, to be
separated, and this resulted in some irregularities. Tables 1 and
2 show in tabulated form the results of all the life-history experi¬
ments.
Table 1 contains the records of the larval ecdyses. The ma¬
jority of the life histories show either eight or nine ecdyses, al¬
though two, as far as could be discovered, had only seven. The
number of ecdyses and the length of larval and pupal life seem
to bear no relation to the sex of the moth, for both sexes were
266 Wisconsin Academy of Sciences , Arts , and Letters.
represented in the long and in the short periods. A second table
was made, using the life histories in table 1 and material from a
number of incomplete records, to show the length of stadia in
days, and from it were plotted curves showing the variation in
stadia. It will be noticed that in the average life the first stadium
is nearly four days long. From this the stadia gradually decrease
in length until the sixth is reached. Then an increase in sta¬
dium duration begins and goes rapidly on until the last stadium,
which is always much the longest of any, because of the time
taken for cocoon spinning. The moths emerge in about seven days
from the date of pupation.
in
R
c>
4— >
I .2 3 + 5 6 7 8 ?
Ttu^nteT Of Ecdyses
Fig. 1. Curves showing variations in stadia of larval life.
_ Curve of larval life with eight ecdyses.
. Curve of larval life with nine ecdyses.
- Curve giving combination of both.
Table 2.
Giving in days the maximum, minimum, and average length of each period of larval and pupal life.
PERIODS
Chase — Length of Life of the Larva of the Wax Moth . 267
The larvae when they hatch are very small, under two milli¬
meters in length, colorless, and extremely active. They run about
rapidly, using only the true legs and not allowing the rest of the
body to touch the surface on which they move. When disturbed,
they can run backward nearly as readily as forward. As soon as
they are hatched, they spin a silken thread by which they drop
from one part of the comb to another. After the first ecdysis,
a change takes place in their appearance and movements. They
can still run rapidly, but are in the comb eating and tunneling
and in less danger. They are longer in proportion to their width
than before, and are beginning to use the prolegs as well as the
true legs in crawling. From this period on in succeeding ecdyses,
the larvae continue to change slowly. The body grows somewhat
gray in color and the chitin of head and tarsus becomes browner,
though immediately after each ecdysis the body is colorless. At
the time of the third or fourth ecdysis, fine hairs become visible
forming a protection for the body, the motion is slower, and the
dorsal body surface curves slightly upward, making the shape
of the mature larva differ from that of the first instars which are
flattened dorsoventrally. At the beginning of the last stadium,
whether eighth or ninth, the larva becomes very dark, almost black
in color. This condition continues for three or four days, and
then the larva becomes normally gray, remaining so until pupa¬
tion. This color change at one definite period in the life has not
yet been explained. The larva just before pupation is between
two and three centimeters in length and nearly round in cross
section. It wanders about until a suitable place is found, usually
outside the comb, and there spins its cocoon, taking from one to
three days to complete it. The cocoon of white silk with bits of
wax and excreta fastened to it, when finished, has at one end a
partial opening of two, three, or four Y-shaped flaps fastened to¬
gether with a few silk threads to make escape easy for the emerging
imago. The opening is probably cut by the larva after spinning
rather than spun in such form originally.
Zoological Laboratories,
University of Wisconsin
A PRELIMINARY STUDY OF THE DIGESTIVE SECRE¬
TIONS OF PICKEREL AND PERCH
CAROLINE WALKER MUNRO
Apparently very little experimental work on the digestive secre¬
tions of fish has been done up to this time. That the stomach of
a fish contains some other acid than hydrochloric (Stewart, 1905),
and that the secretion of definite enzymes is not limited to fixed
regions of the alimentary canal as in the mammals (Bridge,
1910), is known, but what the composition of the enzymes is, or
what foods the secretions of the different regions of the alimen¬
tary canal are adapted to digest, are questions that have appar¬
ently had little attention. The purpose of this paper is to state
the results of a few experiments performed with extracts of the
digestive juices of perch and pickerel, in an attempt to determine
which enzymes are adapted to digesting specific classes of foods.
The experiments were performed in accordance with suggestions
contained in Stewart’s (1905) Manual of Physiology, which out¬
lines methods of determining the digestive action of mammalian
enzymes, and a comparison was made of the relative strength of
prepared solutions of panereatin and pepsin with that of fish
enzymes. A number of preliminary experiments were carried out
on the digestive secretions of perch. After this two series of
parallel experiments were made on the enzymes obtained from
pickerel and perch, as compared with commercial solutions of the
corresponding mammalian enzymes.
Pickerel1
The extracts of the enzymes were obtained from a pickerel
weighing 10.3 pounds, which was found to contain a partially
digested perch about 6 inches long in its stomach. The mucous
membranee of the stomach was stripped off, triturated in a mor-
1 Esox lucius Linnaeus.
269
270 Wisconsin Academy of Sciences , Arts, and Letters.
tar with clean sand, and placed in a test tube with sufficient gly¬
cerine to cover it. Similar glycerine extracts of the intestine and
the pancreas were made, and the test tubes were allowed to stand
for two days in order that the glycerine might extract the en¬
zymes.
Peptic Digestion
Three test tubes were set up, containing, respectively: (1) 1
gram egg albumen, and 20 drops of glycerine extract of the
stomach; (2) 1 gram egg albumen and 20 drops of solution of
commercial pepsin; (3) 1 gram egg albumen. The three tubes
were filled with 10 cc. of a 2% solution of hydrochloric acid and
placed in a bath at a temperature of 35 to 40° C. for one hour.
In test tube (1) the albumen swelled up into large flakes which
soon disappeared, and a practically clear solution was obtained at
the end of the hour; in tube (2) the flakes of egg albumen dis¬
appeared less rapidly, and in tube (3) they disappeared still more
slowly. The contents of the test tubes were then filtered and neu¬
tralized with sodium hydroxide. In tubes (1) and (2) a heavy
flocculent precipitation was formed upon addition of the base ; in
tube (3) no such precipitate was formed. This precipitate prob¬
ably consisted of acid-albumens. An excess of sodium hydroxide
was then added to the contents of each test tube, and then a drop
of very dilute copper sulphate. A faint rose reaction in tubes
(1) and (2) showed the presence of proteoses and peptoses,
whereas the absence of such a reaction in tube (3) showed the
absence of these products of digestion.
The reaction of the contents of the stomach of a large pickerel
on litmus paper was tested and found to be decidedly acid. Eight
drops of a 20% solution of sodium hydroxide were required to
neutralize it.
Pancreatic and Intestinal Digestion
With proteins. Four test tubes were set up, containing, res¬
pectively: (1) 0.5 gram egg albumen and 20 drops glycerine ex¬
tract of intestine; (2) 0.5 gram egg albumen and 20 drops gly¬
cerine extract of pancreas; (3) 0.5 gram egg albumen and 20
drops of commercial pancreatin solution; (4) 0.5 gram egg al¬
bumen. Ten cc. of 1% solution of sodium carbonate were then
placed in each tube and the tubes were placed in water at a
temperature of 35-40° C. for one half hour, at the end of which
time the egg albumen had gone into solution completely in all the
Munro—Dig estiva Secretions of Pickerel and Perch. 271
tubes. A test with copper sulphate showed by the rose reaction
that in the tubes containing pancreatin and gycerine extract of
the intestine proteoses and peptoses were present, whereas in the
other two tubes there was no evidence of their presence.
With cane sugar. Four test tubes were set up, containing,
respectively: (1) 2 cc. cane sugar solution and 30 drops of
glycerine extract of the intestine; (2) 2 cc. cane sugar solution
and 30 drops of glycerine extract of the pancreas; (3) 2 cc. cane
sugar solution and 30 drops of commercial pancreatin solution ;
(4) 2 cc. cane sugar solution. Ten cc. of a 1% solution of sodium
carbonate were then placed in each tube and the tubes were
placed in a bath at a temperature of 30° C. for one hour. At the
end of this time the contents of each tube were treated with 4 cc.
of Fehling’s solution. The latter was soon completely reduced
by the contents of the tube in which commercial pancreatin had
been placed, whereas no reaction was obtained in the other three
tubes although they were allowed to stand for several days. This
showed that the cane sugar had been inverted to dextrose by the
action of the commercial pancreatin solution only.
Bile. The action of the bile in emulsifying oils was tested, and
it was found that one drop of the bile of the pickerel was suffic¬
ient to form a thick emulsion with 10 drops of olive oil.
Pergh2
Similar experiments to those described for the pickerel were
performed with glycerine extracts of the enzymes of the perch.
Since, however, no structure which can be definitely identified as
a pancreas had been found in the perch (Pratt, 1905), it was
possible to run these experiments with only two extracts, those of
the stomach and the intestine. The combined secretions of two
fishes were used on account of the small size of each fish.
Peptic Digestion
Three test tubes were set up, containing, respectively: (1) 23
drops glycerine extract of stomach and 0.2 gram egg albumen;
(2) 23 drops commercial pepsin solution and 0.2 gram egg al¬
bumen; (3) 0.2 gram egg albumen. All were filled with a 2%
solution of hydrochloric acid and placed in a bath at a tempera¬
ture of 28° C. for 40 minutes. The contents of the tube contain-
2 Perea flavescens Mitchill.
272 Wisconsin Academy of Sciences , Arts, and Letters.
ing commercial pepsin digested the fastest, that of the tube in
which no enzyme had been placed the slowest. The test with di¬
lute copper sulphate in an alkaline solution showed the presence of
proteoses and peptoses in tubes (1) and (2) but not in tube (3).
The contents of the stomach of the perch, while distinctly acid
to litums, was so small in quantity that it would require frac¬
tions of drops in order to titrate for the acidity. Such an ex¬
periment was not performed.
Intestinal Digestion
With proteins. Three test tubes were filled, respectively with
(1) 0.2 gram egg albumen and 20 drops of glycerine extract of
the intestine; (2) 0.2 gram egg albumen and 20 drops commercial
pancreatin solution; (3) 0.2 gram egg albumen. All three were
filled with a 1% solution of sodium carbonate, and placed in a
bath of 40° C. for 35 minutes. The commercial pancreatin di¬
gested the fastest and the contents of the tube in which no enzyme
was placed, the slowest. The test with dilute copper sulphate
showed the presence of proteoses and peptoses in tubes (1) and
(2) but not in tube (3).
With cane sugar. Three test tubes were set up, containing,
respectively: (1) 10 drops intestine extract and 2 cc. cane sugar
solution; (2) 10 drops commercial pancreatin solution and 2 cc.
cane sugar solution; (3) 2 cc. cane sugar solution. These were
half filled with a 1% solution of sodium carbonate, and placed
in a bath at 35° C. for 45 minutes; 2 cc. of Fehling’s solution
were then added to each, and all were allowed to stand for twen¬
ty-four hours. At the end of that time the Fehling’s solution
was completely reduced in the test tube containing commercial
pancreatin, showing that the cane sugar had been inverted. A
slight inversion had taken place in the tube containing glycerine
extract of the intestine, and none at all in the test tube in which
no enzyme had been placed.
Bile. The bile of the perch was very active in emulsifying
oils, one drop forming a thick emulsion with 20 drops of olive oil.
Discussion and Conclusions
The pickerel, whose diet in the adult stage consists almost
wholly of fish (Pearse, 1915), apparently has no enzyme adapted
to the inverting of sugars, an essential function of the digestive
Munro — Digestive Secretions of Pickerel and Perch. 273
juices of animals living on vegetable matter. The perch, on the
other hand, which is a versatile feeder (Pearse, 1915), is pro¬
vided with an enzyme which to a small extent inverts sugars. The
enzymes of the pickerel, however, are highly adapted to the thor¬
ough digestion of proteins, as evidenced by the fact that the ex¬
tracts of the stomach and intestine of the pickerel acted more
rapidly than a comparable enzyme from mammalian digestive
glands.
It appears, moreover, that the perch, in which a pancreas is
lacking, is provided with active enzymes in the intestine which
perform the functions ordinarily performed by the pancreas in
forms possessing that gland. The pancreas of the pickerel does
not appear from these experiments to be active in digestion, since
no action was obtained from the use of an extract of it.
It may be concluded, therefore, that the digestive secretions of
the fish studied are correlated with their food habits, and that the
lack of a digestive gland (the pancreas) found in higher verte¬
brates, or the non-functional state of that gland if present, are
compensated for by activity on the part of other digestive glands
present.
Zoological Laboratories,
University of Wisconsin
Bibliography
Bridge, T. W. 1910. Fishes. Cambridge Natural History 7: 141-
537. London.
Howell, W. H. 1912. A textbook of physiology. 1018 pp. Phila¬
delphia and London.
Pearse, A. S. 1915. The food of the shore fishes of certain Wiscon¬
sin lakes. !U. S. Bureau Fisheries, Bull. 25: 249-290. Wash¬
ington.
Pratt, H. S. 1905. A course in vertebrate zoology. 299 pp. Bos¬
ton.
Stewart, G. N. 1905. Manual of physiology with practical exercises.
911 pp. London.
18— S. A. L.
\
THE ORGANIZATION OF THE NUCLEI IN THE ROOT
TIPS OF PODOPHYLLUM PELTATUM
JAMES BERTRAM OVERTON
Introduction
Although the nuclear divisions of somatic cells have been fre¬
quently studied and the somatic chromosomes during certain stages
of division have received a great deal of attention, a thorough
study of their structure and behavior during rest has until re¬
cently been largely neglected. Thus far the structure and rela¬
tion of the chromosomes during rest has been determined in com¬
paratively few forms. A number of valuable papers have ap¬
peared on this phase of the subject, notably those of van Wisse-
lingh (’99), Gregoire (’03, ’06, ’08, ’12) and his students, Haecker
(’04, ’07), Sijpkens (’04), Strasburger (’05), Bonnevie (’08),
Nemec ’99, ’10), Digby (’10, ’19), Fraser and Snell (’ll), Lun-
degard (’10a, b, ’12 b, c.), von Schustow (’13), Sharp (’13, ’20a),
and others, so that a general interest in the subject has greatly
increased
The opinions held by some cytologists, especially those of the
English school, that in both diploid and haploid nuclei a separa¬
tion of the chromosomes into two equivalent portions on the spindle
is already foreshadowed by their fission in the preceding telo¬
phase, as well as the interpretation of Dehorne (’ll) and others
that the chromosomes are in all stages of division associated in
pairs, each chromosome having the value of a longitudinally split
chromosome, have increased interest in the subject of somatic
mitosis.
The observations described in the following paper are the out¬
come of several years of investigation and thought upon the
structure and organization of the nuclei in the root tips of Po¬
dophyllum pelt at um , the result of which have already been briefly
275
276 Wisconsin Academy of Sciences , Arts, and Letters.
reported (Overton, ’096, ’ll). In order to be certain that the
figures and conclusions drawn from them are correct, the prepar¬
ations from which the drawings were made and many other slides
have been many times re-examined during the last ten or twelve
years, especially in the light of the interpretations above referred
to. In the present account the aim will be to give a careful de¬
scription of the somatic chromosomes during the reconstruction
stages, to follow in detail their structure and arrangement in the
resting nucleus, and to determine how they are again reformed
preparatory to division.
Materials and Methods
The material was collected in early Spring before the roots had
attained great length by removing entire plants from the humus
in which they grew. The soil was carefully removed, and the
root tips were severed and dropped at once in the various fixing
fluids. In order to compare the effects of different fixing fluids
upon the nuclear and cytoplasmic structures, a large number of
fixatives were used, chiefly Flemming’s chromic-acetic mixtures in
various strengths and modifications, Carnoy’s alcohol-acetic and
alcohol-acetic-chloroform mixtures, Hermann’s platinic chloride-
acetic-osmie mixture, Merkel’s chromic-platinic chloride mixture,
Juel’s zinc chloride-acetic-alcohol mixture, Guignard’s chromic-
iron chloride-acetic mixture, Kaiser’s sublimate-acetic mixture,
picric-acid, mixtures, Worcester’s formalin-sublimate-acetic mix¬
ture, Gilson’s sublimate-nitric-acetic mixture, and Zenker’s subli¬
mate-potassium bichromate-acetic mixture, with Tellyesniczky’s
modification. Fairly good results were obtained with Flemming’s
stronger osmic fluid. As this is the mixture most generally used
for cytological work in botany, I have compared its fixation very
carefully with that obtained by the other fluids and find that it
apparently often causes artifacts, which will be discussed in the
description of my observations. Merkel’s fluid gave by far the
best results, preserving the nuclei and cytoplasmic structures
equally well, and preparations fixed in this fluid were found to
be far superior to all others. The results described in this paper
are, therefore, based mainly upon preparations fixed in this fluid.
Comparisons were always made, however, with osmic fixation,
which apparently shrinks the nuclear structures and may thus
easily lead to misinterpretation. Throughout this paper Merkel’s
fixation is considered the normal one.
Overton — On the Boot Tips of Podophyllum Peltatum. 277
Sections were cut from three to eight microns thick, and the
preparations were stained with Flemming’s triple combination or
by either Heidenhain’s or Benda’s haematoxylin methods. Care¬
ful comparisons were always made between preparations stained
by the various methods.
Observations
I shall describe the structure of the chromosomes during mi¬
tosis in the root tips of Podophyllum peltatum L. beginning with
the equatorial plate stage, and follow the changes which occur
during the reconstruction of the daughter nuclei, and also the
structure and organization of the resting nucleus in so far as I
have been able to determine and interpret the phenomena.
Metaphases and Anaphases
Figure 1 shows a section of an equatorial plate in lateral view
in which five entire chromosomes are present and a portion of a
sixth, whose remaining part has been cut away. In general the
chromosomes lie in a plane nearly perpendicular to the long axis
of the spindle. It can be seen that in some cases a bundle of
fibers is attached at about the middle of each chromosome while
in other cases the attachment is at or nearer one end. Although
the chromosomes lie usually with one end in or near the equato¬
rial plane, their free ends may often assume various other posi¬
tions with reference to the axis of the spindle. Examining the
chromosomes shown in figure 1 more in detail one can see that
they are not homogeneous in structure, as they have been de¬
scribed and figured by most observers. In each chromosome may
be seen lighter and darker portions. In the chromosomes in the
center of this figure a double row of bodies is distinctly visible,
making this chromosome appear longitudinally split. There ap¬
pear to be two stainable substances present in these chromosomes,
a less stainable supporting homogeneous trama in which granular
portions are imbedded. The trama appears between the indi¬
vidual granules and between the two rows of granulas as a less
deeply stained streak. The darkly staining bodies in each row
lie opposite each other. "With the triple combination the bodies
stain bluish red or purple like chromatin, while the substratum
stains an orange red like linin. These chromatic bodies are not
278 Wisconsin Academy of Sciences, Arts , and Letters.
homogeneous in structure but consist of numerous smaller granu¬
les. The linin on the other hand shows scarcely if any granular
structure.
Although it is evident that the chromosomes of the equatorial
plate are double structures, the halves always lie close together
and are never separated in the way as has been described for the
chromosomes of certain other plants. In no case have I observed
a transverse segmentation of the chromosomes at this or any other
stage.
As mentioned above, the spindle fibers may be attached at the
middle or at or near one end. In the case the attachment is near
one end, each half as it is drawn toward the pole assumes the form
of a J as is shown in figure 2 and in the chromosome at the right
of figure 3. When the attachment is at one end, more or less
straight rods pass to each pole. This condition is evident from
the anaphasic chromosome in the center of figure 3. Sometimes
the attachment of fibers is about the middle of the chromosomes,
in which case Y-shaped figures result. This condition is repre¬
sented at the left in figure 3. As the anaphasic chromosomes are
passing to the poles, they are quite separate from one another and
do not mingle very closely until the poles are reached and then
not so closely as described by some authors. It is noteworthy
that the anaphasic chromosomes, being in the forms of Y’s, J’s,
or I’s, are easily distinguishable from one another. I have ob¬
served that there is a tendency for the different forms habitually
to pass in pairs toward the poles. In figure 5 three pairs of
chromosomes may be seen at one pole, two Y-shaped chromosomes
at the left, two more or less straight rod-shaped ones at the right
and two J-shaped ones in the center.
Chromosomes, occurring in pairs on the spindle, have been re¬
ported by Strasburger, (’05, ’075, ’08, ’ll), and more recently
by Muller (’09, T2) and Gates (’12). That chromosomes occur
in pairs in resting somatic nuclei is also apparent in many plants
containing prochromosomes, as I have previously reported (Over-
ton, ’05, ’09a). Lundegard (’10c) denies the existence of such
a pairing in Allium and Yicia. Sharp (’13) fails to find such
pairing in Yicia, and von Schustow (T3) denies its existence in
Allium.
That the metaphasic and anaphasic chromosomes are not homo¬
geneous, but consist of two stainable substances, is evident from
figures 2 and 3. During the passage of the chromosomes from the
Overton — On the Root Tips of Podophyllum Peltatum. 279
equatorial plate to the poles, numerous more or less irregular
masses and transparent spots appear (fig. 2). This appearance
is probably due to the absorption of liquid material and its col¬
lection in droplets and masses of irregular shape. These droplets
may anastomose, giving the appearance shown in these figures.
This condition gradually increases until conspicuous anastomos¬
ing vacuoles appear irregularly distributed in each chromosome,
(figs. 2, 3, 5, 6, 7). I have, however, never found a line of vac¬
uoles arranged regularly along the chromosome axis, such as has
been described by Gregoire and Wygaerts (’03), Gregoire (’06),
and more recently by other observers. At their very earliest ap¬
pearance some of these vacuoles seem to anastomose, as is shown
in figures 2 and 3. In the late metaphasic and early anaphasic
chromosomes the more deeply staining substance or chromatin oc¬
curs in granules connected by finer threads, which stain less
densely and take up the orange in the triple combination like
linin. From these observations it might appear as though in
each chromosome the achromatic substance was being dissolved
or digested or removed by some means, leaving behind a granular
chromatic substance and a reticulate achromatic portion or linin.
However, I am inclined to believe that the appearance of vacuoles
is due to the absorption of liquid substance from without, prob¬
ably by osmotic action similar to that described by Lawson ( ’03,
’ll, ’12). I am also inclined to look upon the chromosomes as
osmotic systems, the less dense achromatic substance becoming
vacuolated and thus separating the denser granular chromatic
material, which it supports. In figure 3 the chromosomes are
represented as they appear when stained with the triple stain.
A less stainable reddish purple substratum is distinctly visible,
in which appears a more densely staining irregular, granular,
reticulate substance. By careful study of such preparations one
can see by focusing that alveoli or bubbles of sap-like material
also occur in the substratum and that they often anastomose.
The structure of these chromosomes apparently very strongly
supports the view that they are to be regarded as a colony of
granules or discrete particles imbedded in a substratum. From
the staining reactions the substratum appears to be linin of a
rather homogeneous or gelatinous consistency. The chromatic
bodies, which are present in the chromosomes of the equatorial
plate, and which are divided or separated as the chromosomes are
divided, are also granular in structure. Undoubtedly these larger
280 Wisconsin Academy of Sciences , Arts , and Letters.
granules or chromomeres are composite in structure, being com¬
posed of still smaller elements. By means of the progressive al-
veolation of the ground substance or linin of the chromosomes, in
which the granules are imbedded, these granules are separated.
Part of the linin may be dissolved or removed in this process, while
the remainder becomes reticulate by the anastomosing of vacuoles.
The chromatic granules of the chromosomes become simply spread
out or distributed by this process and are supported by the linin
reticulum. That the anaphasic chromosomes are granular reticu¬
late structures is shown by figures 3. The progressive alveolation
always occurs internally. I have never observed a chromosome
showing any appearance of erosion.
Although I have been able to stain the chromosomes so that one
may see through them, and even study their internal structure
and organization, I have not been able to discover in Podophyllum
any such internal structural changes as have been described by
Miss Bonnevie (’08) for Ascaris and Allium. She reports a cen¬
tral longitudinal axis in each chromosome, which divides longi¬
tudinally as the chromosome halves are separated. She regards
this central axial portion as being composed of newly formed ach¬
romatic substance. I have considered this point carefully, and in
some instances have found a central more darkly staining streak
but always in preparations which I regard as poorly fixed. The
same may be said concerning the massing of the chromatin in a
peripheral cylinder. When the chromosomes are not well fixed, as
I find in some preparations fixed in sublimate solutions, a denser
peripheral sheath may sometimes be differentiated, but I can place
no importance upon this appearance. Under no circumstances
have I been able to make out any quadripartite arrangement of
granules as described by Miss Merriman (’04) for Allium. Lun-
degard ( ’10a, ?12c) considers the axial vacuolation of the an¬
aphasic chromosomes of Allium and Vicia as a true split, while
Nemec ( ’10) places no such interpretation on the central vacuola¬
tion of the anaphasic chromosomes of Allium. Although Sakamura
( ’14) finds an anaphasic central vacuolation of the chromosomes of
Vicia, he does not regard it as a longitudinal splitting of these
chromosomes.
I have been unable to find any evidence of a longitudinal split¬
ting of the chromosomes during the anaphases as described by
Granier and Boule (’ll) in the root tips of Endymion nutans. I
cannot agree with the conclusions of Dehorne (’ll), based on his
Overton — On the Boot Tips of Podophyllum Peltatum. 281
studies of Salamandra and Allium that the anaphasic chromo¬
somes of Podophyllum consist of two lots of secondarily split
chromosomes. Dehorne’s conception has been criticized by Gre-
goire ( ’12).
Sharp (’13) explains in detail how the various appearances
above described may be brought about by the decolorizing of sec¬
tions, but holds that in Yicia no evidence is found for the real
vacuolation of the chromosomes during anaphase.
Telophases
Gregoire ( ’06) reports that the daughter chromosomes of Allium
become massed into a “tassement polaire”, which is quite dense
and from which the extremities of the chromosomes emerge. As
before mentioned, I have never found in normally fixed prepara¬
tions the chromosomes so densely massed at the poles of the spindle
as has been described for some other forms. Figure 4 represents
a cell, fixed in Flemming’s strong solution, which much resembles
the condition described by Gregoire and which corresponds to his
figure 1 for Allium cepa. In the cell represented by figure 4 the
chromosomes are densely crowded. That the cell is much shrunken
is evidenced by the vacuolar condition of the cytoplasm and by the
large space between the plasma membrane and the cell wall, which
is not here represented. The normal condition of the cytoplasm
is better shown in figures 1, 8, 11, 20, and 25. In figures 5, 6,
and 7, which are drawn from thin sections, the telophasic chromo¬
somes can be seen to be entirely separated from one another ex¬
cept laterally at certain points, their apparent connection being
due perhaps to their increased size.
I have been unable to observe the lateral anastomoses between
the telophasic chromosomes at this stage, either in the form of
pseudopodia as described by Boveri (’04), and similarly by Gates
(’12), Lundegard (’12 b), and von Schustow (’13), or as portions
formed by adhesion of the telophasic chromosomes as held by
Gregoire and his students. Beer (’13) finds that in the preme-
iotic divisions of Equisetum the substance of the telophasic chro¬
mosomes becomes distributed along numerous fine branches and
anastomosic connections which develop between them. He finds
no internal vacuolation of the chromosomes.
In my opinion the polar tassement , in which the chromosomes
are closely massed, is due entirely to the action of the fixing fluids.
282 Wisconsin Academy of Sciences, Arts, and Letters.
In preparations fixed in Merkel's fluid I have never observed this
massing of the chromosomes at the poles (see figs. 5, 6, 7). In
these figures no anastomoses are present. I have constantly ob¬
served the massing of the telophase chromosomes (fig. 4) and the
anastomoses in preparations fixed in Flemming’s osmic mixtures.
In figures 22 and 22 a, which are drawn from preparations fixed in
osmic mixtures, these structures are very distinct. Figure 22a;
represents an enlarged portion of the two chromosomes to the left
in the upper nucleus of figure 22. Figure 23 represents two
daughter cells also fixed in Flemming’s osmic mixture. Anasto¬
moses are also present. Figure 23 a is an enlarged drawing of a
portion of the upper nucleus of figure 23. The relationship of the
parts is carefully represented. In this figure the outlines of two
chromosomes are distinctly visible but cross anastomoses also ap¬
pear, which, from the appearance of the preparation when com¬
pared with Merkel preparations, are doubtless artifacts. These
figures represent the identical stage drawn by Gregoire and Wy-
gaerts (see their figs. 3 and 4). Their preparations were fixed in
Hermann’s fluid which may account for this appearance. I have
also observed similar structures in preparations fixed in this fluid.
In figure 8 drawn from material fixed in Merkel’s fluid, which
represents a more normal condition than figures 22 and 23, no
anastomoses are present between the adjacent chromosomes except
at their polar ends. This is better shown in the lower nucleus of
this figure. In this nucleus the chromosomes lie in the newly
forming nuclear vacuole perfectly distinct from one another. In
figures 6 and 7 similar conditions are shown less clearly. The
above mentioned authors describe the anastomoses as characteristic
in form, chromatic in nature and content, being under stress with
a thin portion in the middle and a cone-shaped portion where they
join the chromosomes. This is in my opinion apparently the case,
but I regard them as unnatural lateral portions due entirely to the
sticking together of the colloidal or gelatinous chromosomes, which
are brought into close contact by the fixing fluids and which again
become separated. The separation may perhaps be due to the
formation and increasing amount of nuclear sap. I cannot admit
that these anastomoses in Podophyllum take any important part
in the formation of the nuclear reticulum. The alveolation and
reticulation, which is beginning to take place in the anaphasic
chromosomes in the manner above described, continues in the telo-
phasic chromosomes. The chromatic and achromatic portions be-
Overton— On the Boot Tips of Podophyllum Peltatum. 283
come still more distinct. The vacuoles increase in size, number,
and distinctness (figs. 6, 7). Gregoire (’06) describes the con¬
ditions at this period thus:
“On voit le corps de chaque chromosome creuse de nombreuses cavites,
petites et de formes tres variables, contenant un liquide non miscible a
la substance chromosomique, non miscible non plus & l’enchyleme proto-
plasmique et qui constitituera plus tard l’enchyleme nucleaire”.
With this statement my observations, so far as they go, are in
accord. By means of this alveolar-reticulate process the chro¬
matic granules, imbedded in the linin, as I have described above,
become more widely separated. The formation of vacuoles and
their inosculation produces a spongy structure with threads and
plates in the linin substance, having denser and less dense por¬
tions, in which the chromatin remains suspended. This process
continues until a truly reticulate condition arises with the chro¬
matin mainly at the junctures of the threads. In figure 7 an¬
astomosing vacuoles may be seen as well as unreticulated portions,
which appear in the form of granules, platelets, and threads of
different forms and sizes, so that the substance of each chromo¬
some, as Gregoire states, soon appears
“en membranules, en lamelles, en filaments, de sorte que la vraie struc¬
ture de chaque chromosome est plutot celle d’une band spongieuse,\
but this process finally gives rise to the linin reticulum, in which
are imbedded the chromatic granules. Beginning on the left of fig¬
ure 7, this process may be followed by successively observing the
chromosomes to the right of this figure. The portion of the
chromosome on the right is nearly entirely distributed. I do not
find any marked tendency for the vacuoles at this stage to be ar¬
ranged along the chromosome axis, as claimed by Gregoire and
others, although this condition may sometimes occur as shown in
figure 6. I do not find a central undistributed axis, as seen by
Haecker (’04) in Siredon. The vacuoles are distributed irregu¬
larly as a rule, and the entire chromosome finally becomes reticu¬
late.
Gregoire (’06) states that the alveolation of the chromosomes is
most distinct during the telophases, which is true of Podophyllum,
but I think that in Podophyllum the process begins much earlier,
even during the metaphases. The compactness of the chromo¬
somes often causes the stain to cover up entirely any traces of
284 Wisconsin Academy of Sciences , Arts , and Letters.
alveolation in the earlier stages. Gregoire (’06) has criticized
Miss Merriman ’s ( ’04) interpretations, which do not need further
discussion at this point. Suffice it to say that I have not found
the quadripartite granules described by her for Allium in the
telophasic chromosomes of Podophyllum, nor do I find any indi¬
cation of a longitudinal splitting of these chromosomes at this
time or at any earlier stage on the spindle as has been claimed by
some observers. As pointed out by Sharp (’13), the fact that
vacuoles may occur in almost every conceivable position in these
chromosomes is of great importance as it bears upon the question
of the splitting of the chromosomes, which is held by many to
occur during the telophases.
Lundegard (’09) observed a median vacuolation of the chromo¬
somes of Trollius, which he did not at that time describe as a
longitudinal split, but in 1910 he often observed in Vicia and Al¬
lium what he regards as a true longitudinal split in the telophasic
chromosomes, both in fixed and in living material. He failed to
observe the spiral structure described by Bonnevie, but describes
the chromosomes as fragmenting and joining to form karyosomes.
More recently Lundegard (’12c) emphasizes the double nature of
the telophasic chromosomes. Nemec (’10) states that the telo¬
phasic chromosomes of Allium undergo a vacuolation without a
true splitting. Fraser and Snell (’ll) believe that they have
found that the telophasic chromosomes of Vicia undergo a true
longitudinal splitting by means of median vacuolation. The
chromosomes are described as being connected by anastomoses,
and these authors believe that, as the nucleus enlarges, the two
halves of the telophasic chromosomes are drawn apart along the
line of vacuoles. Digby (’19) holds that in all of the arches-
porial divisions in Osmunda the chromosomes undergo a longi¬
tudinal splitting in varying degree during the early telophases,
which split in agreement with Fraser and Snell is held to persist
throughout the resting stages until it is completed on the spindle
of the following division.
In my studies on Podophyllum I have been unable to observe
any of the above described phenomena, which in any way re¬
semble in the slightest degree a telophasic longitudinal splitting
of the chromosomes. Thus far in the consideration of the chromo¬
somes of Podophyllum I have been able to identify each chromo¬
some, which persists as an autonomous element although under¬
going a series of internal changes. Some authors maintain that
Overton— On the Boot Tips of Podophyllum Peltatum. 285
the daughter chromosomes unite end to end to form a daughter
spirem. Such a spirem has not been found by Gregoire or by any
of his co-workers, and has not been observed by Lagerberg (’09)
in Adoxa. In all the plants which possess prochromosomes, as
first described by Rosenberg (04) and since by several other in¬
vestigators, the absence of a continuous chromatic spirem has
been noted. In agreement with most recent work on other forms,
and as has been emphasized by Sharp (’13), I have been unable
to find a continuous spirem formed from the daughter chromo¬
somes of Podophyllum. Although during the reconstruction
stages of Podophyllum I have sometimes observed appearances of
a dispirem (figure 23), I have never been convinced of its uni¬
versal existence in this plant, and am inclined to the view held by
Gregoire that it does not obtain either in this plant or in Allium.
If there is anything comparable to a dispirem, it certainly is not
completely and continuously chromatic in character. In figures
5, 6, and 7 there seems to be no tendency for the chromosomes to
unite endwise into a spirem. Each chromosome is undergoing
the internal modifications above described. In figure 8, in the
nucleus of the lower cell, each chromosome is still free and not
united with any of its associates.
In the lower cell of figure 8 each chromosome may be seen to be
surrounded by a definite membrane much as has been described
for karyomeres or chromosome vacuoles in certain animal cells.
The cytoplasm in many places may also be seen to extend in be¬
tween these alveolated structures. By a constant progressive al-
veolation and reticulation of these chromosomes they come to lie
closer and closer together, so that the cytoplasm is shoved aside
and the chromosomes touch each other in places. This is evident
in the upper nucleus in figure 8. If the chromosomes first touch
at the polar ends, peculiar lobe-shaped structures arise giving a
lobed appearance to the nuclei, as shown in figure 8 and also in
figure 12. Gregoire (’06) in his figure 3 has represented a simi¬
larly lobed nucleus in Allium, and I have also often observed like
conditions in the same plant. Gregoire, however, shows these lobes
as distinctly anastomosed. Although the lobes in the upper nu¬
cleus in figure 8 appear in contact at the polar ends, I do not
believe that they are anastomosed or that they unite in any way
except by the inosculation of the vacuoles. This view is further
supported by a study of such nuclei as are shown in figures 11
and 12. These are resting nuclei in the interphase stage, in which
286 Wisconsin Academy of Sciences, Arts , and Letters.
some individual alveolated and reticulated chromosomes are still
visible as distinct independent entities. In figure 12 the lobed
condition of the nucleus is still apparent, although this nucleus
is in an entirely resting condition, as is shown by the cell itself.
The nuclear membrane may perhaps be only the limiting portion
of these alveolated chromosomes where they abut upon the cyto¬
plasm. Somewhat analogous formation of a nuclear membrane
occurs in some animals as described by Vejdovsky (’07), Reuter
(’09), and others. However, the vacuoles may interosculate with
each other and may perhaps communicate with the surrounding
cytoplasmic substance through the sap, although I have never
seen anything to indicate the existence of such a communication.
That a chromosome as it becomes reticulated may have its own
nucleolus is shown in figure 8. I have often counted several such
nucleoli in like position in the different chromosomes. As indi¬
cated, several nucleoli lie scattered in the nuclear reticulum (see-
figs. 8, 9, 11, 12, 13, 16, 17, 19, 24, 25). As seen from these
figures, the form and size of the nucleoles may vary, but they are
usually spherical and often vacuolate. No extranuclear nucleoli
were observed in the cytoplasm, as reported for certain Liliaceae
and also for Adoxa by Lagerberg (’09). In Podophyllum there
appears to be no direct connection between the nucleoli and the
chromosomes, such as reported by Wager ( ’04) for Phaseolus and
by Strasburger (’07a) for Marsilia, and which has also been de¬
scribed for certain animals. The origin of the nucleoli I have
not been able to determine. They apparently arise de novo dur¬
ing the early telophases, without any morphological connection
with the chromosomes. According to Mano (’04), in Phaseolus
and Solanum the nucleoli appear independently of the chromatin.
Fraser and Snell (’ll) hold that they appear as a droplet or
several droplets usually in some relation to the chromosomes;
Sharp (’13) finds the nucleoles appearing early among the loosely
packed chromosomes, and suggests that there may be an indirect
physiological connection between the nucleoli and the chromo¬
somes. I am inclined to the view of Strasburger (’95, ’00) that
they are reserve materials which are used up in the kinoplasmie
formations of the cell rather than that they are stores of chroma¬
tin, which may again be distributed upon the linin as Strasburger
more recently holds. This view, however, is supported by no di¬
rect evidence in Podophyllum.
Overton — On the Boot Tips of Podophyllum Peltatum. 287
Interphase
As Sharp (’13) points out, we are indebted to Lundegard
( ’125) for this term, which is applied to the period in the nuclear
history between two successive intervals of division in tissues
composed of rapidly dividing cells. I am in agreement with these
two workers that the transformation of the telophasic chromosomes
does not in general proceed as far in such nuclei as in older tis¬
sues, in which the divisions proceed more slowly. Figures 11 and
12 represent stages in which many nuclei are found to pass the
interphase. I have been unable in the root tips of Podophyllum
to find nuclei with a more uniform reticulum than is shown in
figures 9 and 10. In this stage one or two nucleoli are present,
as shown in figure 9. Eeticulation may proceed further in older
portions of the root, as is noted by Sharp ( ’13) . Although larger
chromatic masses may be present, I have been unable to distinguish
any distinct caryotin clumps or karyosomes as described by Lun¬
degard (’10 a, ’12 b) for Vicia, and have never found these karyo¬
somes split as described by him.
The Early prophases
The resting nucleus, as I have described it, consists of elemen¬
tary reticula, which in Podophyllum may often be recognized as
distinctly independent alveolar-reticulate bands or areas, while in
other cases these areas are not distinguishable (compare figs. 9 and
11) . During the prophases a reconstruction or condensation oc¬
curs to form the chromosomes of division. During the very early
prophases the first evidence of a preparation for division is notice¬
able in a somewhat greater affinity of these bands for stains, the
reticulum becoming more dense and chromatic in appearance es¬
pecially along the bands. Each band or chromosome becomes
more distinct, resembling those of the reconstruction stages (fig.
12) , The vacuoles become fewer and the chromatic portions
larger and closer together. The process continues until the con¬
densing alveolar-reticulate portions become quite distinct, with
clear areas between, as shown in figure 13. The chromosomes thus
condensing appear as rather definite bands, each band represent¬
ing a chromosome of the telophase.
As pointed out above, the resting reticulum may be a rather
uniform network (fig. 9), or the chromosomes may appear less
288 Wisconsin Academy of Sciences, Arts , and Letters.
distributed as shown in figures 11 and 12. Sharp (’13) points
out that in certain resting nuclei of Vicia heavier bands repre¬
sent the reticulate chromosomes of the telophase joined together
by fine anastomoses as a continuous net. My results in general
agree with those of Sharp, but I am of the opinion that, even
when the chromosomes become indistinguishable as such in the
resting reticulum, they again condense during the prophases.
Sharp does not represent the stages shown in my figures 11 and
12, which I regard as very early prophase stages that should come
before the prophase stage in Sharp’s figures 1 and 2, which latter
figures I regard as representing a rather advanced prophase
stage, and not a true resting condition.
These bands formed from the resting reticulum are at first
broad and very reticulate. In figure 13 two such bands may be
seen on the right of the nucleus. This represents about the same
stage as Gregoire (’06) figures for Allium, and I have often seen
similar figures in that plant. This author holds that the reti¬
culum is transformed into a series of alveolar-reticulate or spongy
bands, united by lateral anastomoses. I can agree with this in¬
terpretation, except that I do not find the lateral anastomoses
abundant in normally fixed preparations. As the chromosomes
do not anastomose during the reconstruction stages of the nuclei,
no laterial anastomoses appear, except in material improperly
fixed. In figures 24 and 25, which are drawn from preparations
fixed in Flemming’s fluid, such anastomoses are present, but I
do not regard this as a normal appearance. As Gregoire says,
the reticulum is thus resolved into a series of bands more or less
parallel, each of which possesses the same structure as the nuclear
reticulum.
“C’est comme si, dans ce reseau, on avait fait passer au rateau. Image
seulement, car se qui s’est reellement produit, c’est une. concentration.”
Sharp ( ’13, ’206) also represents the chromosome bands as
connected by lateral anastomoses, and points out that the inter¬
pretation placed upon the resting nucleus does not correspond
with that of Fraser and Snell ( ’ll) and of Lundegard ( ’09,
’10 a, ’12c). It may also be added that Digby ( ’19) agrees with
the opinion of Fraser and Snell that the resting reticulum con¬
sists of faint granules in which the individual chromosomes are
indistinguishable. Such preparations as that shown in figure 13
are the only evidence that I can find that the chromosomes, which
Overton— On the Boot Tips of Podophyllum Peltatum . 289
I have described as paired at the telophase, may again reform in
pairs during the prophases, but I regard this as rather strong
evidence. I can find no evidence from my studies to support the
hypothesis of Fraser and Snell (’ll) and of Digby (’19) that
each prophasic chromosome is formed of two units derived from
a single chromosome and that the units of each pair are separated
during the subsequent division.
Sharp (’13, 20b) holds that the first indication of prophasic
changes is the breaking down of the lateral anastomoses between
the heavier portions of the reticulum, so that each chromosome
stands out as an irregular reticulate, alveolar band, and points
out that such bands may probably be visible all through the period
of rest. With these statements my results accord, except, as
above stated, that I do not find evidence for lateral anastomoses
in all cases and that even during the period of comparative rest
their absence may be evident (fig. 11). The succeeding events
of chromosome condensation as found in Podophyllum harmonize
very well with those described by Sharp (’13) for Yicia. I find
exactly the stages represented by Sharp’s figures 3 to 17, except
that, as stated above, I am of the opinion that there exist both
chromatin and linin, while Sharp states that Vicia furnishes no
good evidence for the existence of more than one substance in the
chromatic structures. Mottier (’07) in his studies on the reduc¬
tion division in Podophyllum speaks of the nuclear reticulum as
being made up of both linin and chromatin.
In examining figures 11 and 12, the reticulum of the bands
can be seen to be made up of irregular granules imbedded in the
linin substratum. Bridges of linin appear to connect the granules
of a band together very regularly. On closer examination and
study, however, the structure will be seen to be very similar to
that shown in figures 9 and 10, only more condensed. As the
prophases proceed, the linin connections between any two or more
granules appear coarser and more chromatic in content. This
condition is especially observable in the two bands favorably placed
for study near the periphery on the right of the nucleus shown
in figure 13. These bands appear to be made up of irregular
granular chromatic platelets connected by linin strands. The
alveoli appear fewer and may open freely into neighboring al¬
veoli or into the nuclear sap. Out of these granular platelets by
some method, which I have not been able to make out to my en¬
tire satisfaction, and out of the linin connections a long, thin,
19— S. A. L.
290 Wisconsin Academy of Sciences, Arts, and Letters .
much coiled and twisted thread is finally evolved (figs. 13, 14, 15,
17). Sharp ( ’13) holds that the chromatic material condenses
along a very irregular region around the open spaces and cavities.
It appears from a most careful study of my preparations that the
granules, which are present in the prophasic reticulate bands, be¬
come condensed along certain strands or lamellae to form ir¬
regular platelets. These platelets are placed more or less, at right
angles to the long axis of the bands. On examining figure 13
again in detail, such condensations may be seen with many of the
collections or irregular platelets more or less at right angles to
the length of each band. This is especially observable at the
bottom of the figure. Linin threads or lamellae may still be seen
connecting them in various places.
The chromatin granules collect on the periphery of the alveoli
of the bands in such a manner as to form a more or less zigzag
arrangement across the bands. This would also account for the
arrangement of the platelets. By the dissolution or withdrawal
of the free linin portions, the platelets would thus be arranged
in a zigzag spiral-like line or thread. This final arrangement is
indicated in figure 14. In figure 15 the spiral arrangement of
the individual bands is very distinct. In the upper part of the
nucleus transformation has not so far progressed, the bands being
still quite reticulate. Figure 16 represents a nucleus in which
most of the bands have been cut transversely. Irregular chro¬
matin platelets are quite distinct at various places in the nucleus.
Figure 17 represents a nucleus in which all the bands have been
transformed into threads. The whole thread in its entire length
is long, slender, spirally coiled, and much eonvulated in the
nuclear cavity. From my observations I am convinced that there
are breaks in this thread and that no continuous spirem is formed.
Sharp (T3) describes similar zigzag bands in the prophases
of Yicia, points out that most authors have omitted these stages
from their accounts, and holds that these stages are of the greatest
importance in interpreting the splitting of the chromosomes.
Sharp (’205) also describes like stages for Tradescantia. Such
stages are exceedingly abundant in Podophyllum and are omitted
by Fraser and Snell (’ll), and as Sharp points out, are not de¬
scribed by Digby (’19) for Osmunda. Lundegard (’09, ’10a,
’12a, 5, e), also fails to describe such zigzag threads for Yicia.
Although I have searched carefully and have followed in detail
the formation of these zigzag bands, I have never found any evi-
Overton — On the Root Tips of Podophyllum Peltatum. 291
denee that they arise or that they are formed in lateral pairs
closely associated. In every case I am convinced that they are
formed from a single chromatic band as above described, and as
Sharp (’13) maintains for Vicia. In my opinion the zigzag stage
of the prophase in Podophyllum always precedes the formation
of the thicker coiled spirem, which is apparently the conclusion
of Sharp (’13) regarding Yicia.
Most authors place the spirally coiled spirem in the same place
in the series as I have done for Podophyllum. In examining
Strasburger’s (’82) early figures of corresponding prophase stages
of the endosperm nuclei of Fritillaria imperalis (his fig. 72, PI.
II ) , I find that he evidently saw the same sort of spirem, which he
places immediately after the resting stage of the nucleus. He
also showed a similar stage for Salamandra (see his fig. 182, PI.
III) , and Flemming (’82) also figured and described a somewhat
similar spirem for Salamandra (his fig. 31, PI. Ilia). He said:
“Die Anordnung des Fadenknauls zeigt in den Anfangsformen noch
vielfach scharfere winklige Knickungen, wie solche an den Balkchen des
ruhenden Kernnetzwerks die Regel sind. Je weiter aber der Process
bedeiht, desto mebr gleichen sich diese Knickungen zu welligen Biegungen
aus, die schliesslich durchweg vorkommen.”
The thin homogenous threads of the zigzag or coiled spirem split
longitudinally, as shown in figures 17 and 18. As pointed out
by Sharp (’13) for Yicia, not all portions of the thread split at
the same time, and all stages intermediate between vacuolation
and a complete split may be seen.
Strasburger (’05, ’075, ’ll) holds that the splitting of the
spirem is brought about by the division of chromatic units or
chromomeres, a view supported by Allen ( ’05) and others. Mul¬
ler ( ’12) holds that the achromatic material between the chromo¬
meres first splits, and that the splitting of the chromosomes then
follows. Sharp (’13) finds no evidence for the support of either
view from his studies on Yicia. Sakamura ( ’14) finds no evidence
of chromomeres in Vicia cracca.
In my former studies (Overton (’05, ’09a) on reduction divi¬
sions in certain dicotyledonous plants, I was unable to find the
regular arrangement of chromomeres above referred to. This
was especially the case in the heterotypic division of Podophyllum.
Mottier (’07), however, figures chromomeres in the heterotypic
divisions of this plant. In the present studies on the somatic
292 Wisconsin Academy of Sciences , Arts, and Letters.
divisions of this plant I have also found difficulty in destinguish-
ing chromatic bodies in the early spirem, which I could interpret
as chromomeres (see figs. 13, 14, 15, 17), although such collec¬
tions are undoubtedly present in later stages, especially after the
spirem is split. Perhaps these bodies are present in such young
stages as are represented in figure 17, but they may be obscured
by the uniformity of the staining quality at this stage. Even in
the very early split spirems no chromomeres are visible, as ap¬
pears from figure 19. Certain irregularly placed portions of this
spirem stain more densely, but there are no definite chromomeres
distinguishable. In figure 18, however, representing a nucleus
in which the spirem is completely split, a double row of chromatic
granules or chromomeres is apparently differentially stained as
distinct from the linin supporting strand. In such figures and in
the succeeding stages these granules, which stain distinct from the
linin, even in the chromosomes of the equatorial plate, are always
distinguishable. Whether or not they are always opposite each
other, and whether or not during the splitting of the spirem each
chromomere divides to form two, I have not been able to de¬
termine. To all appearances this is the case. I am led to this
conclusion from the number and arrangement of the chromomeres
in the two parts of the split spirem and especially from their po¬
sition in the chromosomes as they appear in division stages (see
figs. 18, 20, 21).
The Later Prophases
In the stages immediately following the longitudinal splitting of
the chromosomes, and while they are still long and much coiled
in the nuclear cavity, a thickening of the threads occurs as shown
in figures 18 and 19. At this stage the longitudinal split of the
threads is very evident, and my interpretation of the split agrees
with that of Muller (’12), namely, that the portions of the thread
between the chromomeres splits first and that this splitting is fol¬
lowed by, or brings about, a splitting of the chromomeres. Numer¬
ous such examples as are shown in figure 18 support this view,
rather than that of Strasburger (’07&, ’ll) that the splitting of
the thread is initiated by the division of the chromomeres. In
agreement with the description of Sharp (’13) for Vicia, I find
that the longitudinally split threads or chromosomes have a gen¬
eral longitudinal orientation in the nucleus, making them easily
Overton — On the Boot Tips of Podophyllum Peltatum. 293
studied, as shown in figure 19. Although the longitudinal halves
lie very close together, the split is distinct throughout the length
of each chromosome. I agree with Sharp (’13) that the appear¬
ance of free ends of the threads in thick sections may be taken
to indicate that no continuous spirem is formed. I do not find
the fine lateral anastomoses described by Sharp (‘13) for Vicia and
by him ( ’20b) for Tradescantia at this stage, except in cases which
I do not regard as normal.
The chromosomes continue to shorten and thicken and assume
various shapes and positions in the nuclear cavity as shown in
figure 20. In general the chromosomes lie about the periphery
of the nucleus as shown in this figure, but as they are compara¬
tively long, portions often extend into the center of the cavity.
The longitudinal split in each chromosome at this period is distinct,
each chromosome shows a double row of chromatic granules or
chromomeres. As the chromosomes shorten and thicken still
more, these chromomeres become still more distinct (fig. 21),
which distinction is maintained in the chromosomes of the equa¬
torial plate as shown in figure 1. Even though the achromatic por¬
tions of the chromosomes may unite more or less closely, it is
evident that the chromomeres remain distinct. Sharp (T3) be¬
lieves that although the halves of the chromosomes of Yicia may
lie very close together they do not fuse. My results in the main
support this view. It is evident from the two chromosomes ly¬
ing in the upper part of the nucleus shown in figure 20 that the
homologous chromosomes exist in pairs.
I have not studied the formation of the spindle in detail. The
phenomena observed in Podophyllum appear to be similar to
those described for spindle formation in somatic divisions. That
the formation of the spindle is preceded by the appearance of
nuclear caps as described by Densmore ( ’08 ) for Smilacina and
by Sharp (’13) for Vicia,, is evident from figure 20. I have ob¬
served a rapid contraction of the nucleus at this stage, as pointed
out for Smilacina and other plants by Lawson ( ’ll) and by Sharp
(’13) for Vicia. I agree with Sharp (’13) that the fact that the
fibres are differentiated in two opposite regions before the nucleus
begins to contract does not support Lawson’s view of the origin
of the spindle.
294 Wisconsin Academy of Sciences , Arts , and Letters.
General Considerations
As described above, the telophasic chromosomes of Podophyllum
become transformed into reticulate bands by progressive internal
alveolation. These bands are held to represent the individual
chromosomes, which again condense during the prophases into
irregularly coiled zigzag threads. These threads thicken and un¬
dergo a longitudinal split, which split persists until the daughter
chromosomes are separated at metaphase. As pointed out, I find
no evidence from my studies on this plant to support the view
that the telophasic chromosomes undergo a median vacuolation
to bring about a longitudinal split as held by von Schustow,
(’IB), I am also unable to accept the view held by Dehorne (’ll),
Granier and Boule (’ll), and Lundegard (’12a, c ) that a longi¬
tudinal splitting occurs in the chromosomes during the anaphases,
and cannot agree with Lundegard, who maintains that the ana-
phasic split is quite as evident and distinct as in the prophases,
and who holds with Fraser and Snell ( Tl) and with Miss Digby
(’10, 19) that each chromosome appears from the resting reti¬
culum already double. I agree with the interpretation of Gre-
goire and especially with that of Sharp (’13) that the duality of
the individual chromosomes is the result of the longitudinal
splitting of the prophasic threads, which are evolved by progres¬
sive condensation from the reticulate chromosomes of the resting
nucleus. My view as to the method of the vacuolation of the
chromosomes also accords in general with that of Lawson (’03,
’ll, ’12).
Sharp (T3) points out that the telophasic alveolation of the
chromosomes is too irregular to permit of its being regarded as
a longitudinal splitting. The telophasic figures of Fraser and
Snell (’ll) and of Lundegard (’12 a, c), have been criticized by
Sharp as lacking details of alveolation. Lundegard (’15) points
out that von Schustow ’s ( T3) results support his views as to the
duality of the caryotin and further calls attention to the fact that
Sharp ( T3) fails to take into account that Lundegard is the only
one who has studied the arrangement of the caryotin in the living
cell and maintains that fixed material does not represent the
facts.
The chromosome halves are always superposed at the point of
insertion, as emphasized by Gregoire (’12), and as stated by Lun-
Overton — On the Root Tips of Podophyllum Peltatum. 295
degard (’12), Sharp (’13), and von Schustow (’13) for certain
other plants. In Podophyllum the points of insertion of the various
chromosomes lie in a single plane, as has been pointed out by
von Schustow (’13) for Allium, by Sharp (’13) for Yicia, and
by Lundegard ( ’12c) for other forms. Sakamura ( ’15) states
that in Vida fab a the chromosomes do not lie in an equatorial
plane. In Podophyllum the spindle fibers may be attached at
any point to the chromosomes, this attachment resulting in var¬
iously formed figures as the halves of the chromosomes are pulled
apart ; this has been described by other authors and especially by
Lundegard (’125, c), Sharp (’13) von Schustow (’13), and
others. This condition seems to be the most common one in
plants.
In certain plants and animals the chromosomes have been de¬
scribed as segmenting transversely. In Vida fab a such pheno¬
mena have been mentioned by Lundegard (’125, T4), Fraser and
Snell (’ll), and Sharp (’13), and have been fully described by
Sakamura (’15), who holds that it is the usual occurrence in this
plant. Such a transverse segmentation of the chromosomes of
Podophyllum has not been observed by me.
Lundegard ( ’12c) states that on account of the changes which
the different fixing agents and stains cause it is difficult to de¬
termine the exact condition or constitution of the chromosomes.
He holds that the chromosomes of Allium do not contain chromo-
meres and describes the chromosomes in the living condition as
being cylindrical with smooth surfaces which become roughened
or wringled during fixation, a condition held by him to be an
artifact. This wrinkled condition may account, according to
Lundegard, for the conception of chromomeres, and he further
states that on destaining the stain is retained longer in the thicker
portions of the chromosomes.
Strasburger and others have held that the splitting of the chro¬
mosomes is initiated by the splitting of discrete units or chromo¬
meres arranged serially along the chromosomes, which conception
has been denied by Gregoire and his students. Bonnevie (’08)
and Stomps (’10, ’ll) find no chromomeres in Allium and Spina-
cia. Muller (’12) described definite chromomeres in Najas, but
holds that the linin thread splits first, which splitting is followed
by a splitting of the chromomeres. Sharp (’13, 205) fails to find
evidence of chromomeres in Vida faba and Tradescantia and be¬
lieves that their appearance is brought about by a removal of
296 Wisconsin Academy of Sciences } Arts , and Letters.
stains from denser portions of the chromosomes, a view some¬
what similar to that of Lundegard. Sharp holds that in the meta¬
phase the chromosomes show little differentiation, while Saka-
mnra (’14) finds chromomeres in Vida cracca.
That the prophasic and anaphasic chromosomes of Podophyllum
are not homogeneous but consist apparently of two stainable sub¬
stances has been stated above. I have expressed the opinion that
these two substances are linin and chromatin and that the chro¬
matin apparently exists in these chromosomes as distinct mor¬
phological units or chromomeres arranged in regular rows op¬
posite each other in the metaphasic chromosomes and are present
in a single row in the anaphasic chromosomes, which fact I have
considered as fairly good evidence that the appearance of chro¬
momeres is not due to fixation as held by Lundegard ( ’12c) . On
decolorizing such chromosomes I do not find such irregularly ar¬
ranged spots and masses as described by Sharp (’13) for Yicia.
Although I do not wish to make a positive assertion that chromo¬
meres are present in these chromosomes, nor that their appear¬
ance may not be due to fixation, I am inclined to the view of
Strasburger (’07b, ’ll) that there exist in the chromosomes defi¬
nite morphological units the halves of which are separated during
the separation of the daughter chromosomes.
In my former work on the reduction divisions in Podophyllum
I failed to find evidence of chromomeres, but Mottier (’07) figures
such bodies and describes the nuclear reticulum as being made up
of both chromatin and linin. Muller (’12) finds both chromatin
and linin bodies in Najas before the chromosomes split, and, as
stated, holds that the splitting is initiated in the linin intervals,
the chromomeres dividing later. Alhough Sharp (T3) holds there
is no trustworthy evidence to support the view that chromatic
morphological units exist in Vicia, he recognizes the 11 highly sug¬
gestive nature of the arrangement of the chromatin in a thin
thread and its accurate separation into two equal parts”.
During the anaphases several authors have described a longi¬
tudinal splitting of the chromosomes. Gregoire (’12) discusses
fully the metaphasic and anaphasic phenomena, especially in the
light of Dehorne’s (’ll) interpretation, and maintains that De¬
home’s scheme is contrary to the generally observed facts as Gre¬
goire describes them. Sakamura ( T4) holds that in Vicia cracca
neither during the metaphases nor anaphases is there any evi¬
dence of tetrads as held by Miss Merriman (’04), nor evidence
Overton — On the Root Tips of Podophyllum Peltatum. 297
of the longitudinal split of the next division. Yon Sclrus tow
(IS) has never observed a true longitudinal splitting of the ana-
phasic chromosomes in Vicia fab a as described by Dehome, nor
does von Schustrow find evidence to support Miss Merriman’s
view. Lundegard (’12c) states that the anaphasic longitudinal
splitting is quite as distinct as in the prophases, but von Schus-
tow interprets Lundegard ’s figures as telophasic ones. I agree
with Lundegard that his objection to using fixed material alone
as a basis of interpretation is a valid one. I have also compared
fixed material with that in the living condition not only in Podo¬
phyllum but in many other plants, and am convinced that in many
cases the fixed material presents the same configuration of the
nuclear structures as is seen in the living material. I can find
no evidence from a study of either living or properly fixed ma¬
terial that the telophasic chromosomes undergo a regular median
vacuolation such as would cause them to be longitudinally split.
I hope in the future to be able to present evidence from living
material on this point. Sharp (’13) holds that the telophasic
median vacuolation in Vicia is only apparent and does not really
divide the chromosomes, which observations the results of Saka-
mura ( ’14) on Vicia cracca as well as my own on Podophyllum
support, and I cannot agree with von Schustow (’13) that the
telophasic figures of Lundegard (’10a, ’12c), Dehorne (’ll),
Gregoire (’06), Muller (’09), Fraser and Snell (’ll), and his
own should be interpreted as indicating a splitting of these chro¬
mosomes.
The interpretation of Bonnevie (’08) that the cross sections of
the anaphasic chromosomes at first show a tetrad structure which
later becomes circular, the chromatic material being denser at the
periphery while the center is solid but less dense, is the reverse
of that described by von Schustow (’13), who observes a darker
cylindrical region and a lighter vacuolated inner region in the
chromosomes of Allium. In later anaphases von Schustow de¬
scribes the outer cylindrical region as becoming polygonal in cross
section with the chromatin on the periphery of the chromosomes.
Lundegard (’10a, ’12c), holds that the axis of the anaphasic
chromosomes is vacuolate and regards this vacuolation as bringing
about a true longitudinal splitting. Nemec (TO) also observed
what he regards as a true anaphasic vacuolation, often axial, but
does not describe an anaphasic splitting of the chromosomes.
Fraser and Snell (’ll) and Miss Digby (T9) find no vacuolation
298 Wisconsin Academy of Sciences , Arts, and Letters.
until telophase is reached. Sharp (’13), although finding an ir¬
regular anaphasic differentiation, holds that it does not represent
a true vacuolation as seen in the telophases. Gregoire and Wy-
gaerts (’03) show that vacuolation is evident as early as the meta¬
phase. There seems to be no evidence to support the view of
anaphasic splitting as described by Granier and Boule (’ll).
The structure of the telophasic chromosomes as described by
Bonnevie (’08), due to a disappearance of the achromatic sub¬
stance of these chromosomes and the inner appearance of a fine
spiral thread, finds no support in my observations on Podophyllum
and is discussed by von Schustow (’13) who holds that the con¬
traction of the chromosomes at this stage is so marked as not to
permit of a study of their internal organization, although he
maintains that in cross section the chromosomes are polygonal as
in the anaphases, which latter view also my results do not sup¬
port. Sharp (T3) holds that the figures of Bonnevie (’08-T1)
and of Dehorne (’ll) are apparently schematizations of the oc¬
casional spiral aspect of the alveolar bands.
As already stated, I find no evidence from my studies on Podo¬
phyllum to support the view held by Fraser and Snell (’12) and
Miss Digby (TO, T4, T9), and supported in principle by Lunde-
gard (’12c) and von Schustow (T3), that the anaphasic or telo¬
phasic split described by them in somatic nuclei persists during
the reconstruction stages, the period of rest, and the prophases,
until the component halves are separated at metaphase. I agree
with Sharp’s (’20 a) criticism of Miss Digby ’s statement that
workers on mitosis are in general agreement that the constitution
of the chromosomes is one of duality which persists throughout the
cycles of dissolution and reconstruction. I can find no evidence
in Podophyllum that the early prophasic reticulum possesses any
such duality or that it passes directly into the double spirem.
In certain cells, especially those fixed in Flemming’s fluids, as
shown in figures 22 to 25, aspects are presented which might be
interpreted as indicating a persistent duality of the chromosome
elements. Although there is such an apparent duality, I can not
place any significance on such appearances, for reasons stated in
the description of my observations.
During the polar tassement and at the time when the nuclear
vacuole begins to appear, Gregoire and Wygaerts (’03) believe
that anastomoses are formed between the chromosomes of the
telophase. The exact origin and nature of these anastomoses they
Overton— On the Boot Tips of Podophyllum Peltatum. 299
are unable to explain. They hold that they are the natural con¬
sequence of the close association and arrangement of the chromo¬
somes of the telophase at the poles. Lagerberg (’09), who finds
no true alveolar structure of the chromosomes of Adoxa until after
the nuclear membrane is formed, describes anastomoses between
the undistributed chromosomes. Nemec (’10) holds, in common
with Bonnevie ( ’08 ) , that there are lateral anastomoses between
the chromosomes but that these anastomoses are not entirely due
to the sticking together of the chromosomes of the telophase. Ac¬
cording to de Vries’ interpretation of Stomps’ (’10) results, and
as later described by Stomps (’ll) himself, the lateral anastomoses
are only appearances due to the presence of walls of numerous
small swelling vacuoles, which lie between the chromosomes of
the telophase. Digby (’10) holds that the lateral anastomoses are
present in all stages of nuclear division, even during the period
of rest. The absence of a true polar tassement and a consequent
absence of anastomoses has been described by Vejdovsky (’07) for
the chromosomes of certain animals. As described by Reuter
(09), the chromosomes of Pediculopsis graminum, which pass to
the poles of the spindle as distinct achromatic karyomeres, show
no traces of lateral anastomoses. Von Schustow (’13) holds that
no cross anastomoses exist during the telophases in Allium and
that these anastomoses arise later by an active dislocation of the
chromatic substance, not, as Gregoire (’06) and Sharp (’13) hold,
by the marginal portions of the telophasic chromosomes, which are
at first always in contact, adhering to one another as the chromo¬
somes separate. Von Schustow ’s view as to the origin of the anas¬
tomoses is somewhat similar to that of Boveri (’04), who holds
that the anastomoses are put out like pseudopodia from the chro¬
mosomes, a similar view being also held by Gates ( ’12) and Lun-
degard (’12c), while Strasburger (’05), Dehorne (’ll), and
Muller (’12) all believe that the anastomoses may arise by both
methods. Bonnevie (’08) describes the chromosomes of the telo¬
phases in Allium, which at first lie close together at the poles, as
separating so that anastomoses are formed between the chromatic
spirals. Stomps (’10, ’ll), studying the mechanism of division,
finds the chromosomes of Spinacia at first closely massed at the
poles, but soon separating by the swelling of numerous small
vacuoles which lie between them. The walls of the vacuoles, as
De Vries (’10) describes them, may be seen in the “shape of fine
lines of linin, giving the image of threads stretching from one
300 Wisconsin Academy of Sciences , Arts , and Letters.
chromosome to another”. Fraser and Snell (’ll) describe the
telophasic chromosomes of Yicia as becoming attached laterally
to one another. Beer (’12, ’13) holds that the chromosomes of
Crepis and Eq' isetum at the poles are joined by fine cross anas¬
tomoses. Sakamura (’14) holds that lateral anastomoses of the
telophasic chromosomes of Yicia play no part in their transfor¬
mation into a reticulum.
As stated, in my opinion the polar tassement, in which the
chromosomes are closely massed, is due entirely to fixation, as is
the appearance of cross anastomoses. I have been unable to ob¬
serve in Podophyllum lateral anastomoses between telophasic
chromosomes, either in the form of pseudopodia or as marginal
adhesions, in material which I regard as well fixed. In this re¬
spect my observations on Podophyllum agree with those of von
Schustow (’13) on Allium.
I have already emphasized the fact that I have found very
few stages in the study of Podophyllum which might lead to the
interpretation that the chromosomes of the telophase undergo a
longitudinal splitting by means of median vacuolation or other¬
wise. When the telophasic stages, which apparently show such
splitting, are studied more in detail, as has been done by Sharp
(’13) in Yicia and as I have done for Podophyllum (illustrated
in figs. 22 and 22a), it will be seen that the appearance of such
a longitudinal split is only apparent and not real.
As pointed out by Lundegard ( ’12c) , and as held by von
Schustow (’13) and Sharp (’13), certain nuclei, especially those
of the meristematic region, do not proceed as far in their telo¬
phasic transformations as do others. My results upon Podophyl¬
lum agree with these observations that the nuclei of the meriste¬
matic region show less chromatic distribution than those in older
regions of the root tip. The interpretation of nuclear transfor¬
mation as described by Gregoire has been confirmed in more or
less detail by Kowalski (’04), Berghs (’04- ’05), Mano (’04),
Lee (T2), De Smet (’13), and Sharp (T3, 205) and is supported
in principle by Haecker (’04), the Schreiners (’06), Lagerberg
(’09), Yamanouchi (’10), Nichols (TO), Stomps (TO, ’ll),
Lundegard (’10a, ’12c), Nemec (TO), Miss Digby (TO, T9),
Fraser and Snell (’ll), von Schustow (T3), Sakamura (T4),
and others.
The investigations of Mano (’04) on the root tips of Solanum
tuberosum and Phaseolus vulgaris , the nuclei of which have short
Overton— -On the Root Tips of Podophyllum Peltatum. 301
chromosomes, are of especial interest. In these plants the chro¬
mosomes undergo only a partial or incomplete vacuolation, the
portions not being used to form the anastomoses remainning as
rather large distinct chromatic masses. These masses undergo
very little change, their substance appearing to be more concen¬
trated and the anastomoses between them less distinct than in
the forms studied by Gregoire. During the prophases the anas¬
tomoses slowly disappear and the thicker masses enter into the
definite chromosomes.
Haecker (’04) observed in the epidermal cells of Siredon lar¬
vae that the chromosomes of the telophases become alveolized from
without inward, so that he was able to distinguish a peripheral
“ grosswabigen Alveolenmantel” and an axillary <(gehornelt er-
scheinenden Chromatinstrang” which later becomes further al¬
veolate.
The transformation of the chromosomes by means of a grad¬
ual alveolation into alveolar-reticulate bands or elementary reti¬
cula, as described by Gregoire (’06) and others, has already been
discussed. Sharp (’13) holds that the true alveolation of the
telophasic chromosomes begins about the time they begin to sep¬
arate from one another, while von Schustow (’13) thinks that the
vacuolation occurs much later. Sharp describes not only a median
and irregular internal, but also a peripheral, vacuolation, the in¬
ternal vacuoles occurring in almost every conceivable position, and
states that some of the internal vacuoles may rupture and break
the surface of the chromosomes, so that each chromosome finally
becomes a ragged chromatic band joined to its neighboring bands
by fine anastomoses. Sharp noted that in the process vacuoles
may finally be so placed as to give the chromatic regions of the
chromosomes a spiral aspect, while in other cases the vacuolation
may be more or less along the median line, which may lead to the
erroneous interpretation of a longitudinal split, as held by cer¬
tain authors whose work has already been mentioned. The phe¬
nomena of telophasic transformation as described above is also
supported by the observations of Muller (’12) on Najas and by
Sakamura (’14) on Vida cracca.
Lundegard (’10&) finds that in no case can he trace a direct
transformation of the chromosomes of the telophases to those of
the prophases in Allium and Yicia. In Allium parts of the vac¬
uolated chromosomes may persist as clumps, threads, or masses,
a condition not found in all resting nuclei. Stomps (’10, ’ll)
302 Wisconsin Academy of Sciences, Arts , and Letters.
describes in Spinacia the compact structure of the chromosomes
as being changed into a reticular one by means of numerous small
vacuoles, which slowly increase in size, thereby distending the
surrounding material, so that each chromosome is transformed
into the nuclear reticulum, which is a complex (< res eau de reseaux ”
as described by Gregoire. Nemec (’99) describes a peripheral
and internal vacuolation of the telophasic chromosomes of Allium
and the presence of an axial row of vacuoles and small alveoli
within the chromosomes, the chromatic substance becoming re¬
solved into numerous fine granules, which spread out along the
anastomoses of the linin reticulum. In a later work Nemec ( ’10)
supports Gregoire ’s view of the telophasic transformation, ob¬
serving in many telophasic chromosomes the same vacuolated ap¬
pearance, often showing a distinct axial row, or one large vacuole
running throughout the length of the chromosomes. Nemec notes
certain nodal points during the distribution of the chromosomes,
located at their periphery, and states that if we are to accept Bon-
nevie’s (’08) idea of spiral differentiation we must admit the
presence of several peripheral spiral bands. Nemec maintains
that these nodes or knots are not independent elements, but that
they are located where several lamellae or threads of chromatic
substance meet. Not only may the densely colored portions of
the differentiating and distributing chromosomes appear as spi¬
rals, but Nemec thinks they may show as rings, rods, etc., which
may account also for Miss Merriman’s (’04) conception of te¬
trads.
Miss Digby (’10) in contrast to Gregoire ’s observations, finds
no “such diagrammatic vacuolation of all the chromosomes” in
Galtonia, but observes that vacuolation may be seen in certain
regions and not in others. By transverse segmentation the chro¬
mosomes first break up into certain definite portions of various
sizes, which remain connected as they become distributed into a
more or less beaded reticulum. In her later work on Crepis and
Osmunda, Miss Digby (’14, ’19) supports in principle the view
stated above.
The phenomena of the telophasic transformation as described
by Muller (’12) for Najas agree essentially with the descriptions
of Gregoire.
Lawson ( ’03, ’ll, ’12) first studied the vacuolation of chromo¬
somes in Passiflora and Equisetum, but describes a like condition
for those of certain other plants; this process he thinks is a
Overton — On the Boot Tips of Podophyllum Peltatum. 303
constant normal phenomenon in the organization of daughter
nuclei. He would regard the vacuoles or lacunae, which are pres¬
ent within the chromosomes, as being due to osmotic phenomena.
The liquid in the vacuoles enters the chromosomes by osmosis,
and by means of this endosomosing sap the lacunae are formed.
In Allium Lawson observes an accumulation of the karyolymph
within the chromosomes, appearing as minute lacunae, which in¬
crease in size and flow together, the chromosomes thus appearing
vacuolated. As this karyolymph increases in amount the chromo¬
somes become finely divided, the chromatin finally appearing as
a delicate reticulum suspended in the large vacuole of nuclear
sap. According to this view, each chromosome has an enveloping
plasmic membrane, and each chromosome, therefore, is an os¬
motic system, there being as many osmotic systems as there are
chromosomes. Lawson’s view as to the origin of the chromosome
membrane has been discussed by Farmer (’12).
In Podophyllum I find the metaphasic and anaphasic chromo¬
somes apparently consisting of two distinctly stainable substances,
which I regard as fairly good evidence that both linin and chro¬
matin are present. During the passage of the chromosomes to
the poles numerous more or less irregular, transparent spots ap¬
pear, which I have suggested as probably due to absorption of
liquid material that collects in droplets or masses often irregular
in shape, the irregularities being due perhaps to the anastomosing
of neighboring vacuoles. As stated, I look upon the chromosomes as
osmotic systems as described by Lawson (’03, ’ll, ’12), except
that I do not agree with him as to the origin of the chromosome
membranes. Each chromosome absorbs liquid material from with¬
out, thus by a process of progressive vacuolation of the achro¬
matic portion of the chromosomes separating the denser chro¬
matic material which the chromatin supports. The chromatic
granules thus become distributed in the linin reticulum by this
process. I have never observed any indication of such an erosion
of the chromosomes from without as has been described by some
authors. The formation of vacuoles in the chromosomes thus pro¬
duces a spongy granular structure with numerous lacunae, finally
giving rise to the elementary reticulate bands found in the rest¬
ing nucleus.
I have described a peculiar lobed condition of the daughter
nuclei. Gregoire (’06) figures a like condition for Allium but
304 Wisconsin Academy of Sciences , Arts , and Letters.
represents the lobes as distinctly anasomosed. I do not find these
chromosome lobes always anastomosed and believe that the anas¬
tomosing is due largely to fixation. These chromosome lobes in
Podophyllum suggest that each chromosome becomes distributed
independently, each having its own osmotic membrane, and that
perhaps the nuclear membrane may be only the limiting portion
of the alveolated chromosomes. A somewhat analogous formation
of the nuclear membrane is described by Vejdovsky (’07) for
certain animals. He believes that the membranes of the kary-
omeres go to form the nuclear membrane, which is therefore,
nuclear in origin. It appears from my description that possibly
the nuclear membrane may be in some way closely associated
with, or related to, the outer limiting layer of the chromosomes.
Sharp (’13), in common with most authors, would look upon the
nuclear membrane as being formed by the reaction between the
cytoplasm and the chromosomes, the membrane being formed by
the outer boundaries of small vacuoles between the two. Sharp,
therefore, would regard the nuclear membrane as cytolasmic. My
observations on Podophyllum do not entirely support this view.
That alveolar-reticulate chromosomes much resemble karyomeres,
especially those described for some animals, is apparent. Kary¬
omeres are regarded by many as distributed chromosomes, their
origin being traceable to the chromosomes of the spindle. As
they are described by Reuter (’09), who has observed them to
divide mitotically in Pediculopsis graminum, each karyomere
maintains a complete individuality and continuity within its own
membrane throughout all the stages of division, even persisting
during the resting period. Richards (’07), who describes chro¬
mosome vesicles in Fundulus, and who holds that these vesicles
persist and still maintain their unity of structure during rest,
reviews the recent literature. Nemec (’10) holds that the kary¬
omeres described by Vejdovsky (’07), Bonnevie (’08- ’ll), and
others are formed by the vacuolation of the substance of the chro¬
mosomes; the karyomeres lie near each other and later fuse to
form one large nucleus. Nemec believes it is quite possible to
hinder or to retard this fusion by external conditions. In chlo-
ralized roots of Visum sativum or in those treated with benzol
vapor, Nemec observed an irregular diffuse distribution of the
chromosomes in the cytoplasm, each chromosome forming a small
nucleus. A wound stimulus also causes the chromosomes to be
unequally distributed in the cell space and each chromosome may
Overton— On the Root Tips of Podophyllum Peltatum. 305
form a small nucleus. That chromosomes may regularly be un¬
equally distributed and form small nuclei is found by Juel (’97)
and Schurhoff (T3) in the pollen mother cells of Hemerocallis
fvlvu, and I have observed a like condition in the pollen mother
cells of Paeonia mutan.
Typical karyomere formation in somatic cells of plants, as
Nemec (’10) pointed out, had been observed up to that time only
by Gregoire ( ’03), during the second division of the pollen mother
cells of Trillium cernuum. In this plant several nuclei arise each
of which is either mono- or polychromosomatic, but Gregoire holds
that normally sooner or later the single nucleus results from a
confluence of the chromosome vesicles. Nemec (’10) describes
similar chromosome vesicles in Chara fragilis , in which the chromo¬
somes do not closely mass after metakinesis, as is usually the case.
Nemec states that in small cells a single nucleus is constructed from
all the chromosomes, but in large cells several individual nuclei
are formed which soon after reconstruction fuse to form a single
nucleus. Sehiirhoff (T3) also reports that the karyomeres of
Hemerocallis fuse to form one nucleus. Lundegard (’12a) would
regard the formation of karyomeres as a special and not a usual
phenomenon in nuclear division, connected with the fact that the
chromosomes are in some cases not closely associated, which fact
Lundegard holds may explain the tendency of the young daugh¬
ter nuclei of Allium to show alveolated lobes, formed from the
separated ends of the daughter chromosomes which have become
surrounded by a membrane.
As described above, I have observed a constant though not a
universal lobing of the daughter nuclei similar to that described
and figured for Allium by Gregoire (’06) and by Lundegard
(’12c). I am of the opinion that these karyomeres are formed,
as Lundegard suggests, by the rather widely separated chromo¬
somes becoming alveolated. Instead of the chromosomes becom¬
ing surrounded by a membrane as held by Lawson (’ll, ’12), I
believe that the membrane about each chromosome is always pres¬
ent and that the karyomeres never fuse in the sense that they an¬
astomose, as held by Gregoire (’03), Nemec (TO), and Lunde¬
gard ( T2c) even at their polar ends, but that the only connec¬
tion between the chromosomes is by the inosculation of neighbor¬
ing vacuoles of two adjacent chromosomes. This view finds sup¬
port in the fact that in Podophyllum the chromosome bands are
20— -S. A. L.
306 Wisconsin Academy of Sciences , Arts , and Letters.
distinctly visible in many cases in the resting nucleus. The fact
that in this plant a chromosome may have its own nueleole seems
to support the persistence of karyomere-like structures in the
resting reticulum although the membranes of the karyomeres
may be broken down by the inosculation of neighboring vacuoles.
Sharp ( ’13) points out that in certain resting nuclei of Yieia
heavier bands are present which represent reticulate chromosomes,
but he observed them joined by fine anastomoses to form a con¬
tinuous net. It appears to me that these fine anastomoses repre¬
sent only the boundaries of inosculating vacuoles of the chromo¬
some bands.
Lundegard ( '12 c) fails to find in Allium any definite orienta¬
tion of the chromosome bands; Sharp ( ’13) holds that the reticu¬
late bands of Yieia represent the chromosomes of the telophases,
and figures such bands as more or less polarized and visible dur¬
ing rest; Fraser and Snell ('ll) and Lundegard ('09, '10a, '12c)
state that the chromosomes of Vicia become indistinguishable
during rest. Sakamura ('14) figures the chromosome bands of
the telophases in Vicia cracca, as they become alveolated to form
the resting nucleus, as remaining more or less polarized and states
that these alveolated chromosomes are distinct during rest. Digby
('10), Fraser and Snell ('ll), Beer (’12) and others have also
described a polarity in the arrangement of the chromosomes. Mul¬
ler (’12) states that in rapidly growing merismatic regions of
Najas, often the telophasic structures are carried over into the
next succeeding prophase. Additional evidence of the persistence
of the chromosomes during rest has been brought out by the ob¬
servations of Gregoire and his students, and the work of other ob¬
servers tends to show that the loss of chromosome identity dur¬
ing the reticulated condition is only apparent. The study of the
formation of a finely divided reticulum in the telophases and its
behavior in the prophases has led in many cases to the identifica¬
tion of the reticulated chromosomes during rest.
Many authors have recently come to the conclusion that a con¬
tinuous chromatic spirem does not exist either at prophase or at
telophase. Strasburger ('05) was unable to find such a spirem
in the somatic cells of Funkia and Galtonia. Gregoire and Wy-
gaerts (’03) do not figure a long, thin spirem in Trillium. Gre¬
goire (’06) figures a long slender spirem in Allium but holds that
it is not continuously chromatic. Gregoire and his students are
of the opinion that a continuously chromatic spirem does not ex-
Overtorir—On the Root Tips of Podophyllum Peltatum. 307
ist in somatic or germ nuclei. Although I ( ’04, ’05, ’09a) found
an apparently continuously chromatic spirem in the germ nuclei
of Podophyllum, Campanula, Helleborus, and Richardia, I was
unable to find one in Calycanthus and Thalictrum. I (’09a)
have already suggested that in plants like Allium, etc., in which
there is an apparently continuous chromatic spirem, there prob¬
ably exist individual segments or chromosomes very closely united
without any very extensive linin intervals. Evidence against a
continuous chromatic spirem at any stage is abundant, as is shown
by that presented, especially by Lundegard (’10a, ’12c), Stout
(’12), von Schustow (’13), and others. As stated, I do not find
evidence of a continuous spirem in the root tips of Podophyllum
which, contrary to the statement made in my preliminary reports
(Overton, ’09&, ’ll), in which it was stated that, while the spirem
is not continuously chromatic, the chromosomes appear to be con¬
nected by visible linin intervals into a definite spirem. That the
serial arrangement and association of chromosomes has a signifi¬
cant bearing on the problems as to the general organization of
the nucleus has been pointed out by Stout (’12) and Sharp (’13).
In my former studies on the organization of the nuclei of cer¬
tain plants (Overton, ,05, ’09a), and in common with Rosenberg
( ’04) and several more recent authors, I have presented consider¬
able evidence to show that the chromosomes persist as individual
structures in both somatic and germ cells of plants. In certain
plants I have shown that in the somatic nuclei the chromosomes
are represented by certain definite bodies, the prochromosomes,
arranged in parallel pairs with apparent double linin intervals,
and have also suggested that these heterogeneous spirems prob¬
ably remain distinct throughout the life history of the sporophyte.
I have also pointed out that in such plants possessing a hetero¬
geneous spirem during the first meiotic division, each of the two
parts composing each diakinetic chromosome represents a somatic
chromosome. Each univalent portion of each bivalent diakinetic
chromosome undergoes a longitudinal fission, forming apparent
tetrads.
That in certain plants the chromosomes in resting stages and
in very early prophases do not form a complete reticulum, but
can be observed as unit masses at all stages of nuclear develop¬
ment, is evidence supporting the doctrine of the individuality of
the chromosomes. The presence of prochromosomes or similar
bodies, as first reported by Rosenberg ( ’04) for Capsella, Zostera,
308 Wisconsin Academy of Sciences, Arts, and Letters.
and Calendula, and by Overton (’05, ’09a, ’ll) for several plants,
has been observed by several other investigators.1
The doctrine that the chromosomes are permanent and that the
parental elements remain separate throughout the life cycle of
the organism was first suggested by Haecker. The fact that
chromosomes of different sizes in certain plants are associated in
pairs, as decribed by Strasburger (’05) for Funkia and Galtonia,
has been substantiated by the subsequent observations of several
authors on other plants. The nucleus is, therefore, not only double
in the sense that it contains two sets of parental chromosomes,
but these chromosomes are arranged in homologous pairs.
It seems apparent, therefore, that the duality of somatic nuclei
is due to the association of the homologous chromosomes in pairs,
each element of which splits during the prophases, as above de¬
scribed, rather than that the duality is due to the splitting of the
telophasic chromosomes as held by Fraser and Snell (’ll), Lun¬
degard (’12a, c), Miss Digby (TO, T9), von Schustow (T3), and
others. I agree with Sharp (T3) that it is not necessary but
rather unsafe to rely upon the principle of telophasic splitting
as a premise for the conclusion that the approximation of thin
threads in the early heterotypic prophase represents the reassoci¬
ation of the halves of a single split chromosome, as is done by
the authors cited. The arrangement of entire chromosomes in
pairs and the presence of tetrads in diakinesis, as I have described
them during meiotic divisions of certain plants, argue against the
necessity of such an assumption.
Lundegard (’10a, b, ’12c, T5) has emphasized what he believes
to be a dual tendency of the caryotin during all stages of nuclear
activity, which tendency is expressed morphologically, according
to Lundegard by the appearance of double earyosomes or double
caryotin threads. Although Lundegard holds that morphological
duality is the same in both somatic and generative prophases, he
maintains that the duality signifies a qualitative difference in the
heterotypic prophases, and says:
“Es geht jetzt hervor, dass die heterotypische Doppelfaden aus einem
Paarungsvorgang hervorgegangen sein mussen, denn eine qualitative
Spaltung ware hochst unwahrscheinlich”.
According to this view the halves of the heterotypic chromo¬
somes are qualitatively different, while in the somatic divisions the
1 See lists given by Stout (’12), Digby (’14), and De Smet (’14).
Overton— On the Root Tips of Podophyllum Peltatum. 309
chromosome halves are quantitatively and qualitatively alike. Al¬
though von Schustow ( ’13) agrees with Lundegard as to the pres¬
ence of chromatic strands of double origin in somatic prophases,
he maintains that a like duality in the heterotypic prophases offers
no proof for the theory of a parallel conjugation during matura¬
tion.
Regarding the methods of reduction of chromosomes, Lunde¬
gard agrees with many others that the doubleness of the spirem
in the early heterotypic prophases, unlike that of the somatic
prophases, is due to a lateral conjugation of chromosomes. Sykes
(?08) interpreted the double structures, which she observed in
early somatic and heterotypic prophases, as a pairing of entire
chromosomes, and I, in common with others who have found pro¬
chromosomes, have furnished evidence for such a view. Even
those adherents of the parallel conjugation theory who have not
been able to follow the entire history of the chromosomes admit
that the lateral conjugation of threads in the early heterotypic
prophases represents the conjugation of entire chromosomes to
form the bivalents which are later separated during the first mei-
otic division.
According to the interpretation of Digby (’19), in the hetero¬
typic prophase a distinct parallelism arises just as in somatic
telophasic chromosomes and not by a conjugation of chromosomes.
Following Farmer’s view of meiosis, she holds that bivalents are
formed by a conjugation end to end of segments of this double
spirem, these segments separating in the first division and the
original split functioning in the second division, so that the
homoeotypic division is regarded as a continuation of the last pre-
meiotic division.
Sharp (’20a), in a review of Miss Digby ’s results on Osmunda,
points out that her view has certain advantages.
“It allows one interpretation to be placed upon the double spirem in
both somatic and heterotypic prophases, irrespective of the exact time at
which the split originates, and it also helps to explain the sudden appear¬
ance of the split for the second maturation mitosis in the anaphase of
the first.”
Sharp further says that this question must, however, be settled
by direct evidence and that its solution obviously depends upon
the exact manner in which the telophasic transformation of the
chromosomes and their derivation from the reticulum in the pro-
310 Wisconsin Academy of Sciences , Arts, and Letters.
phases are accomplished. Sharp points out that his investigations
on the somatic cells of Yicia show not only that the telophasic
alveolation of the chromosomes is too irregular to permit of its
being looked upon as a splitting, but also that the reticulate con¬
dition of the somatic prophases, instead of developing directly
into the definite split, gives rise to simple thin threads in which
a new split occurs. Sharp ( ’20b) in his work on Tradescontia
virginiana comes to a like conclusion.
Although I agree with Sharp that the interpretation of Miss
Digby has certain advantages, it appears to me that the best so¬
lution of the problem is to be found not so much by determining
the exact manner of telophasic transformation and the derivation
of the chromosomes from the prophasic reticulum, as by the fact
that the chromosomes remain distinct throughout all stages of
the heterotypic and homoeotypic divisions, and that in certain
cases their complete history both during rest and division can be
followed. As mentioned, I have found that in the root tips of
Calycanthus the prochromosomes in the resting nuclei are associ¬
ated in pairs and are quite as large as the ordinary somatic chro¬
mosomes and have the same shape. In these root tips I have
also found the definite chromosomes of division associated in pairs
just as Strasburger ( ’05) has described them for Funkia and
Galtonia and as they have since been described by several other
workers for other forms. I have already pointed out that in the
vegetative cells of Podophyllum the chromosomes are arranged in
pairs. The only indication of their doubleness in the early pro¬
phases in this plant is due undoubtedly as in other cases to the
associated chromosomes being in pairs. Of course, as both Sharp
and I have definitely shown, each chromosome undergoes an ulti¬
mate longitudinal split in the later prophases. Although Vicia and
Podophyllum are favorable subjects for study of chromosome vac-
uolation and of the prophasic evolution and splitting of the definite
chromosomes, they are not as well adapted to solve certain phases
of the problem as are plants with shorter, less definitely reticulated
chromosomes. In such forms as Funkia, in which Strasburger
(’05) found the chromosomes of different sizes in pairs not only
during division but also during the prophases, it should be pos¬
sible to find a satisfactory solution of the problem, especially in
plants in which not only the pairing is evident but the reticula¬
tion during rest is small.
Overton — On the Boot Tips of Podophyllum Peltatum. 311
Blackman (’98), Chamberlain (’99), and especially Miss Fer¬
guson (’04) have shown that in Pinus the parental elements re¬
main distinct during fertilization, observing that during the first
division of the fertilized egg the parental chromatin elements form
two separate spirems. The independent formation of maternal
and paternal chromosomes has also been described in Tsuga by
Murrill (’00), in Juniperus by Noren (’07), and in Abies by
Hutchinson (‘15), the later of whom describes the maternal and
paternal chromosomes as being formed independently as in most
conifers, but becoming paired side by side and twisting about
each other. The number of paired chromosomes is haploid, each
pair of which, Hutchinson holds, segments transversely to form
the diploid number. Chamberlain (’16) finds a similar pairing
of the chromosomes in Stangeria at fertilization. In the angio-
sperms few detailed accounts of the behavior of the chromosomes
during fertilization and the first division of the egg are at hand,
except that of Sax (’18) on Fritillaria and Triticum in which he
holds that there is no indication that the chromosomes of the male
and female gametes are in separate groups, or that they pair in
the first division of the zygote. It is apparent that the pairing of
the chromosomes may take place at the time of fertilization, as
Hutchinson and Chamberlain have described, or later as indi¬
cated by the results of Blackman, Chamberlain, and Ferguson on
Pinus and by those of other workers on other conifers. From my
own studies upon the vegetative nuclei of plants showing pro¬
chromosomes I believe that the parental elements remain distinct,
but I am convinneed that there is no permanent spatial separation
of the parental chromosomes as described for the first division of
the egg of Pinus. Miss Nothnagel (’18) reports for Lilium mar-
tag on and Trillium grandi folium the presence in the fertilized egg
of two distinct and separate spirems arising from the parental
chromatin elements, and two separate and distinct groups of chro¬
mosomes, which she regards as evidence that the chromosomes
maintain their individuality from one generation to the next.
Miss Weniger (’18) also finds that in Lilium there is no fusion
of the parental chromatic elements in the fertilized egg and that
each parental spirem segments into the chromosomes which come
together in pairs, in the manner described by Hutchinson, on the
equatorial plate. I am further convinced from my studies on
plants showing prochromosomes as well as from my present
312 Wisconsin Academy of Sciences , Arts , and Letters.
studies on Podophyllum that the doubleness of the early pro-
phasic somatic nuclear chromatic elements is due to the lateral
association of the parental elements rather than to a splitting of
the telophasic chromosomes.
As I have stated in my previous work (Overton, ’09a), the
nucleus is not only double in the sense that it contains two sets of
parental chromosomes, but these chromosomes are so placed that
there may be an interaction between homologous parallel pairs,
and the association of homologous chromosomes probably occurs
during fertilization or shortly thereafter, although the actual in¬
terchange or mutual influencing of parental parts occurs during
synapsis or related stages. It may further be added that the se¬
rial paired arrangement of homologous parental elements in so¬
matic cells suggests the probability of mutual interaction and in¬
fluence throughout the life history of the sporophyte. From my
present studies on Podophyllum and from a detailed study of the
literature I can find nothing to cause a change of view.
Since the above was written an important note by de Litardiere
(’21) has appeared, dealing with the structure and behavior of
the chromosomes in the root tips of Podophyllum peltatum. Cer¬
tain results as stated in this note do not agree with my (’09 b) in¬
terpretation. In my preliminary report it was stated that the
alveolation of the chromosomes occurs during the passage of the
anaphasic chromosomes toward the poles and that there was evi¬
dence that there were two substances in the chromosomes, the
chromatic granules being separated by a progressive vacuolation
of a linin ground substance. De Litardiere holds that the alveo¬
lation begins as early as the anaphasic dissociation, alveoli being
visible in only slightly stained sections. As stated in the present
paper, I have observed the beginnings of vacuolation as early as
the metaphase. I still maintain that the chromosomes are com¬
posed of two substances, as previously stated. De Litardiere de¬
scribes an axial row of vacuoles in each chromosome separated from
each other by thin partitions. As a general prenomenon, I have
never observed the appearance of such an axial arrangement of
vacuoles. One can often find individual chromosomes with such
a row of granules, but it is not the usual appearance in what I
regard as well fixed material. I have found the alveoli scattered
at all times so that the alveolar-reticulate bands have a spongy
appearance, as described in Gregoire.
At the poles and during the telophase stages, de Litardiere de-
Overton — On the Root Tips of Podophyllum Peltatum. 313
scribes the alveoli as increasing in number and dimensions and as
becoming distributed irregularly in the chromosomes, much as I
have described them. He also maintains that nothing leads to
the conclusion that there is a longitudinal splitting of the chromo¬
somes by a process of alveolation. There is, therefore, no ana-
phasic or telophasic splitting of the chromosomes, as held by some
authors.
De Litardiere describes the walls of the alveoli of the anaphasic
and telophasic chromosomes as of unequal thickness with certain
thin and thick spots, which might be interpreted as granules, but
he denies the existence of granules massed in a linin substratum.
These statements are not in accord with my observations that in
Podophyllum evidence exists that the chromosomes are made up
of two substances, linin and chromatin, and that the latter often
appears in the form of granules. I see no reason to change my
view in the light of de Litardiere ’s description.
De Litardiere describes anastomoses between the chromosomes
while at the poles, which he believes are formed as pseudopod-like
projections from neighboring chromosomes, much as described by
Boveri. He holds that these anastomoses are not stretched ad¬
hesions formed at a time the chromosomes are in contact. These
results are at variance with my observations, as I have never ob¬
served anastomoses formed by any method at any stage in well
fixed material.
In my preliminary report on Podophyllum (Overton, ’09&), I
stated that the telophasic chromosomes appear united into a con¬
tinuous spirem, but in the present paper I have shown that no con¬
tinuous spirem is ever present at any time in this plant. De
Litardiere finds no telophasic spirem.
De Litardiere also finds the extremities of some of the chromo¬
somes terminating at or on the nuclear membrane, much as I have
described, often forming marked projections, a fact which I have
cited as indicating the individual character of the chromosomes
in this plant. The appearance of zigzag bands is also described
in interphasic nuclei. His suggestion that, by observing cross
sections of chromosome bands, I have been led to conclude that
the chromosomes are formed by a condensation process is not well
taken. My conception of the condensation process is fully dis¬
cussed in the main body of this paper.
Although Mottier and I have found sixteen diploid chromo¬
somes in Podophyllum peltatum , de Litardiere finds only twelve
314 Wisconsin Academy of Sciences, Arts, and Letters.
and suggests that there may be several races of this species differ¬
ing in the number of chromosomes.
Resume
1. The resting nucleus does not in general contain a continuous
reticulum. Elementary alveolar-reticulate bands which represent
the chromosomes can be distinguished, often forming lobes in the
nucleus.
2. No karyosomes or prochromosomes are present in the rest¬
ing nucleus.
3. During prophases the alveolar-reticulate chromosomes con¬
dense to form zigzag threads which later thicken and shorten.
4. These thin threads split longitudinally, probably by means
of axial vacuolation, the split persisting until the chromosomes are
separated at metaphase.
5. No continuous spirem is present at any time.
6. Evidence exists that the chromosomes consist of two sub¬
stances, linin and chromatin, the latter often appearing in the
form of definite chromomeres.
7. The chromosomes are arranged on the spindle with their
halves superposed. Spindle fibers may be attached at any point
to the chromosomes.
8. The chromosomes show a tendency to be arranged in homol¬
ogous pairs on the spindle. A similar arrangement can not al¬
ways be seen during rest, but strong evidence is often found of
such pairing during the prophases.
9. There is no massing at the poles of the telophasic chromo¬
somes, each chromosome remaining separated from its neighbors.
10. The telophasic chromosomes are not normally connected
by cross anastomoses.
11. Each chromosome undergoes an irregular alveolation, the
beginnings of which become slightly evident as early as the meta¬
phases, but the alveolation continues to increase through the ana¬
phases and telophases until each chromosome by complete vacuo¬
lation forms an alveolar-reticulate structure of the resting reti¬
culum.
12. Each chromosome for a time is apparently surrounded by
its own membrane, thus resembling a karyomere. The chromo¬
some membrane apparently enters into the formation of the nu¬
clear membrane. During rest the chromosome membrane is broken
Overton — On the Boot Tips of Podophyllum Peltatum. 315
down between adjacent chromosomes by the interosculation of
neighboring vacuoles.
13. The anaphasic and telophasic vacuolation of the chromo¬
somes has no direct relation to the longitudinal splitting of the
chromosomes, this splitting being entirely prophasic.
14. Evidence exists that the chromosomes are individualized
structures persisting as such in different forms throughout all
stages of division and rest.
15. The heterotypic division separates whole chromosomes
which have previously been laterally paired (see Overton, ’05).
16. No evidence was found to support either Merriman’s or
Bonnevie’s theory of chromosome structure.
17. The results do not support the interpretation of Fraser and
Snell, Lundegard, Digby, and others as to the duality of nuclear
structures.
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Description of Figures in Plate VII
Podophyllum peltaum
Fig. 1. Equatorial plate stage, showing five entire chromosomes and por¬
tion of a sixth. Zenker’s fluid. Heidenhain’s iron-alum-
haematoxylin stain.
Fig. 2. Metaphasic chromosome, showing internal chromatic and achro¬
matic differentiation. Vacuolation already begun. Merkel’s
fluid. Triple stain.
Fig. 3. Anaphasic chromosomes, showing different chromosome forms
resulting from place of attachment of spindle fibers. Inter¬
nal vacuolation also shown. Merkel's fluid. Triple stain.
Fig. 4. Cell showing telophasic chromosomes densely crowded at poles.
Cell much shrunken. Cytoplasm appears poorly fixed.
Flemming’s strong solution. Triple stain.
Fig. 5. Telophasic chromosomes rather widely separated from one
another. No cross anastomoses present. Chromosomes shown
tend to be arranged in pairs. Merkel’s fluid. Iron-alum-
haematoxylin stain.
Fig. 6. Telophasic chromosomes, showing lack of cross anastomoses and
irregular internal vacuolation. Merkel’s fluid. Triple stain.
Fig. 7. Telophasic chromosomes, showing lack of cross anastomoses and
still further internal vacuolation. Anastomosing vacuoles and
unreticulate portions are shown. Merkel’s fluid. Triple
stain.
Fig. 8. Two daughter cells, showing nuclei in process of reconstruction.
Chromosomes undergoing internal vacuolation. In the upper
nucleus chromosomes are free from adjacent ones at polar
ends but form lobes at the free ends. In the lower nucleus
some chromosomes can be seen entirely free from one
another, but their substance is rather widely distributed.
Absence of cross anastomoses. Merkel’s fluid. Triple stain.
OVERTON.— OR
TRANS. WIS. ACAD., VOL. XX.
OVERTON. — ORGANIZATION OF NUCLEI.
Overton — On the Boot Tips of Podophyllum Peltatum. 321
Fig. 9. Two daughter cells, showing what appears to be complete distribu¬
tion of the substance of the chromosomes into a resting reti¬
culum. No evidence of chromosome individuality, absence
of prochromosomes, and no karyomeres. Flemming’s strong
solution. Triple stain.
Fig. 10. Enlarged portion of the upper nucleus shown in figure 0. In
this nuclear reticulum the linin portions are stained orange
and the enclosed chromatic substance violet with the triple
combination.
Fig. 11. Portion of a cell showing prohpasic nucleus with reticulate chro¬
mosomes distinct in the foreground. Merkel’s fluid. Triple
stain.
Fig. 12. Portion of a cell showing prophasic nucleaus with elementary
reticulate chromosomes or bands, one such band forming a
lobe on the left of the figure. Merkel’s fluid. Triple stain.
Fig. 13. Later prophasic stage, showing distinct, reticulate, condensing
chromosomes or bands on right of figure. Merkel’s fluid.
Triple stain.
Fig. 14. Portion of nucleus in prophase stage, showing condensation of
portions of chromosomes and spiral aspect of the thread at
the right of figure. Merkel’s fluid. Triple stain.
Fig. 15. Portion of a prophasic nucleus, showing condensation of the
chromosomes and evolution of spiral threads. Zenker’s fluid.
Triple stain.
Fig. 16. Prophasic stage of nucleus showing chromosome bands cut trans¬
versely and irregular arrangement of chromatic platelets of
each band. Merkel's fluid. Iron-alum-haematoxylin stain.
Fig. 17. Prophasic Stage showing simple zigzag arrangement of threads
with indication of longitudinal split. Zenker’s fluid. Triple
stain.
Fig. 18. Portion of zigzag thread showing longitudinal split. Merkel’s
fluid. Iron-alum-haematoxylin stain.
Fig. 19. Later prophasic stage showing disappearance of zigzag arrange¬
ment of thread and its complete longitudinal fission. Chro¬
mosomes show polar orientation in nuclear cavity. Zenker’s
fluid. Triple stain.
Fig. 20. Cell showing longitudinally split chromosomes in nuclear cavity.
Chromomeres evident in each half of split chromosomes.
Polar caps in cytoplasm. Zenker’s fluid. Iron-alum-haemat¬
oxylin stain.
21— S. A. L.
322 Wisconsin Academy of Sciences , Arts , and Letters.
Fig. 21. Two late prophasic longitudinally split chromosomes, showing
halves closely pressed together. Chromosome pair evident
at right of figure. Merkel’s fluid. Triple stain.
Fig. 22. Late telophase stage. Inuclear vacuole forming. Yacuolation of
chromosomes distinct in each, showing the appearance of
longitudinal split and cross anastomoses. Flemming’s
strong fluid. Triple stain.
Fig. 22 a. Detail of the two left-hand chromosomes of the upper nucleus in
figure 22. No central line of vacuoles appears in the left-
hand chromosome, although the vacuolation of the right-
hand chromosome might be so interpreted.
Fig. 23. Cell showing reconstruction stages of the two daughter nuclei and
apparent parallel arrangement of dispirem threads, but in
reality showing stage in alveolation of chromosomes into
irregular bands. Flemming’s strong fluid. Triple stain.
Fig. 23a. Detail portion of the upper nucleus of figure 23 to show that the
parallel aspect is only apparent. Figure 23 & represents a
similar condition.
Figs. 24, 25. Prohpasic nuclei showing conditions which might be inter¬
preted as a parallelism of threads but in reality represent¬
ing stages in condensation of chromosomes comparable to
that shown in figure 15. Flemming’s strong fluid. Triple
stain.
Fig. 25a. Detail of portion of nucleus shown in figure 25. Apparent parall¬
elism of chromatic portions of condensing chromosomes
shown.
THE PHYTOPLANKTON OF THE MUSKOKA REGION,
ONTARIO, CANADA1
Gilbert Morgan Smith
This report on a reconnaissance survey of the August and Sep¬
tember phytoplankton in the lakes of the Muskoka region in
southern Ontario represents an extension to another region of the
type of survey which I have been carrying on in the State of
Wisconsin for the past few seasons. The Muskoka region was
chosen because lakes are abundant and because of the geological
nature of the district. The expectation, on the basis of the Wests’
theory of geological correlation, that the region would prove rich
in desmids was fully realized, and many lakes were found with
a rich desmid flora.
The general nature of the plankton flora is much the same as
that of other lakes in areas of old igneous rocks: namely, small
bulk of plankton, scarcity of Myxophyceae, and few species of
Protococcales. In general, the same organisms are found in On¬
tario as in Wisconsin, but the relative abundance is not the same.
Thus, for example, Pediastrum Boryanum, Kirchneriella obesa,
and Coelastrum microporum are widely distributed in Wisconsin
but poorly represented in the plankton of Ontario. Conversely,
Pediastrum araneosum, Dimorpho coccus lunatus , and Dactylococ-
copsis rhaphidioides are much more abundant in Ontario. The
most striking feature of the desmid portion of the plankton in
Ontario is the frequent occurrence of Staurastrum limneticum var,
burmense, an alga that has been recorded from but a single lake
in Wisconsin. A comparison of the desmid floras of the two
regions will not be made at the present time since the account of
the Wisconsin region has not, as yet, been completed.
1 These investigations were made possible by grants for traveling expenses
from the Research Fund of the American Association for the Advancement of
Science and from the Research Fund of the University of Wisconsin.
323
324 Wisconsin Academy of Sciences , Arts, and Letters.
Table 1.
Showing the location and size of lakes from which plankton collections
were made. The first column refers to the sheet of the Canadian
Geographical Survey on which the lake is mapped, P. S. referring to
the Parry Sound sheet and T. to the Toronto sheet.
Smith — Phytoplankton of the Muskoka Region. 325
The method of presentation of the data is the same as is em¬
ployed in the account of the Wisconsin phytoplankton2, and the
same symbols are used to denote the frequency of occurrence of
the different species. Citation of the original publication of the
species has been omitted, however, in all cases where it has been
given in connection with Wisconsin algae. Citation of descrip¬
tions of the Desmidiaceae is likewise omitted since the ground is
amply covered by Nordstedt’s Index Desmidiacearum. The lakes
from which plankton samples were collected are shown in table 1.
Class MYXOPHYCEAE
Order Coccogoneales
CHROOCOCCUS Nageli 1849
Chroococcus dispersus var. minor G. M. Smith
Gull (2) (rr).
Chroococcus limneticus Lemm.
Basil (rr). Big Dudley (rrr), Black (2) (sss), Butterfly (r). Clear (1)
(rrr), Clear (2) (r), Corner (rrr), Hamer (rr), Harbon (rrr), Henshaw
(s), Joseph (s), (Lake of Bays (r), -Little Dudley (rrr), Long (1) (r),
Long (2) (r), McGowan (ss), Marion (rr), Muldrew (r), Muskoka (rr),
Otter (sss), Portage (rr), Rosseau (rr), Rowley (r), St. Mary (r), Silver
(1) (r), Silver (2) (r), Vernon (r).
Chroococcus turgidus (Ktz.) Nag.
Clear (ss),Gull (1) (rrr), Mill (rrr), Vernon (rrr).
MERISMOPEDIA Meyen 1839
Merismopedia elegans A. Br.
Basil (rr), Big Dudley (rrr), Clear (2) (rrr), Gull (1) (rrr), Marion
(rrr), Mill (rrr), Strange (rrr).
Merismopedia glauca (Ehr.) Nag.
Silver (1) (rrr).
Merismopedia tenuissima Lemm.
Butterfly (rrr), Black (1) (sss), Gull (2) (rrr), Henshaw (s), Muskoka
(rrr), Nine Mile (rrr), Rowley (sss), Vernon (ss).
2 Smith, G. M., Phytoplankton of the inland lakes of Wisconsin Part I. Wis.
Geol. and Nat. Hist. Survey Bull. 57, 1920.
326 Wisconsin Academy of Sciences, Arts, and Letters.
COELOSPHAERIUM Nageli 1849
COELOSPHAERIUM KUETZINGIANUM Nag.
Basil (rr), Big Dudley (rr), Black (1) (rrr), Burned Rock (rrr), Butter¬
fly (r), Henshaw (r), Joseph (rrr), Little Dudley (rrr), Long (1) (r),
Long (2) (rrr), McGowan (rr). Mill (rr), Muldrew (rr), Muskoka (rr),
Nelson (rrr), Otter (r), Round (2) (rr), Rowley (r), St. Mary (sss), Sil¬
ver (1) (rrr), Silver (2) (r), Vernon (rrr).
Coelosphaerium Naegelianum Unger
Basil (rrr). Big Dudley (rrr), Black (1) (rrr), Black (2) (rrr), Brush
(s), Burned Rock (rr), Butterfly (rrr), Clear (1) (rr), Clear (2) (rr),
Echo (rr), Gull (1) (sss), Gull (2) (sss), Harbon (rrr), Henshaw (sss),
Joseph (rrr), Little Dudley (rrr), Long (1) (rr), Long (2) (rr), Marion
(sss), Mill (r), Muldrew (rr), Muskoka (r), Nelson (r), Nine Mile (rrr).
Pine (rrr), Portage (rrr), Rowley (r), Rosseau (r), St. Mary (rr), Silver
(1) (r), Silver (2) (rrr), Strange (rr) , Thompson (rr), Vernon (r).
GOMPHOSPHAERIA Kiitzing 1836
Gomphosphaeria lacustris Chodat.
Basil (rr), Harbon (rr), Henshaw (r), Lake of Bays (r), Long (1) (rr).
Long (2) (rr),Mill (rr), Muskoka (rr), Rosseau (r), St. Mary (r), Strange
(rrr), Vernon (rr).
Gomphosphaeria aponina Ktz.
Long (1) (rrr).
MICROCYSTIS Kiitzing 1833
Microcystis aeruginosa Ktz.
Henshaw (ss).
var. major (Wittr.) G. M. Smith
Harbon (rr).
Microcystis flos- aquae (Wittr.) Kirchner
Big Dudley (rrr), Black (1) (rrr), Brush (rr), Butterfly (rrr), Clear
(2) (rrr), Echo (rrr), Gull (2) (rrr), Lake of Bays (ss), Little Dudley
(rrr). Loon (rrr), Marion (ss), Mill (rrr) , Muskoka (rrr), Rosseau (rr).
Round (2) (r), Thompson (rrr), Vernon (r).
APHANOCAPSA Nageli 1849
Aphanocapsa elachista var. conferta W. & G. S. West
Basil (r), Clear (2) (ss), Round (2) (sss), Rowley (sss), Thompson
(ss).
Smith — Phytoplankton of the Muskoka Region. 327
var. planctonica G. M. Smith
Black (1) (rr), Black (2) (cc), Burned Rock (rr), Butterfly (r), Gull
(1) (ss), Lake of Bays (ss), Leech (rr), Long (2) (ss), Muskoka (ss),
Marion (rrr), Strange (rrr), Vernon (sss).
Aphanocapsa Grevillei (Hass.) Kab.
Pine (rr), Otter (rrr).
APHANOTHECE Nageli 1849
Aphanothece clathrata W. & G. S. West
Vernon (rrr).
Aphanothece stagnina (Spreng.) A. Br.
Leech (rrr).
GLOEOTHECE Nageli 1849
Gloeothece linearis Nag.
Cells 13.5-20 fx long; 3. 5-4.5 [x broad.
Long (1) (rrr).
DACTYLOCOCCOPSIS Hansgirg 1888
Dactylococcopsis rhaphidioides Hansg.
Cells 2. 5-3. 5 /x broad; 7.5-21 fx long.
Basil (r), Clear (1) (rr), Clear (2) (rr), Hamer (cc) , Lake of Bays (rr),
Long (1) (rr), Muldrew (rrr), Muskoka (rrr), Nelson (rrr), Rowley
(rrr), St. Mary (rrr), Silver (2) (rrr), Vernon (rrr).
This seems to be a widely distributed alga in the Muskoka region.
The organism resembles Rhabdoderma linear e Schmidle & Lau-
terb. in general appearance, but differs in the cell ends being at¬
tenuated to a rounded point instead of having cylindrical cells that
are not attenuated. The two daughter cells resulting from the
division of any cell remain unchanged for some time so that the
two poles of a cell are different, as in Elaktothrix gelatinosa Wille.
The gelatinous envelope of the individual cells was evident in
many colonies, but in others it was completely confluent with the
colonial envelope. The distribution of the cells within the co¬
lonial envelope is not regular, although a majority of the ^ells
have their long axes parallel.
Dactylococcopsis acicularis Lemm.
Clear (2) (rrr).
328 Wisconsin Academy of Sciences , Arts , and Letters.
Order Hormogoneales
TRICHODESMIUM Ehrenberg 1830.
Trichodesmium lactjstre Klebahn.
Harbon (rrr), Loon (rrr).
ANABAENA Bory 1822
Anabaena limnetica G. M. Smith
Pine (rrr).
Anabaena Bornetiana Collins
Vegetative cells 1R-13 y broad; heterocysts 10-13.75 fx broad:
spores 12.5-16.5 y, broad, 46-111 fx long.
Black (1) (r), Butterfly (r), Gull (1) (rrr), Gull (2) (rrr), Long (1)
(rr), Loon (rr), Nelson (rr), Rowley (rr).
The spores of this species are always adjacent to and in the
great majority of cases on both sides of the heterocysts. The
dimensions given above are slightly different from those of Collins.
Anabaena spiroides Klebahn
Cells 7. 5-8. 5 fx broad; heterocysts 7.5-8.75 [x broad; spores 10
[x broad, 27-37.5 jx long.
Clear (2) (rrr), Lake of Bays (rr).
Anabaena Lemmermanni P. Richter
Black (2) (r), Burned Rock (rrr), Harbon (sss), Joseph ( rrr ), Lake of
Bays (r), Long (2) (rrr), Portage (rr).
Anabaena flos-aquae (Lyng.) Ralfs
Basil (rr), Big Dudley (rr), Black (1) (rr), Butterfly (r), Clear (1)
(aa), Henshaw (rr),Long (1) (rr), Marion (rr),Pine (rr),Rosseau (rrr),
Round (2) (rr), Rowley (rr), St. Mary (rr), Strange (rrr), Thompson
(rr).
APHANIZOMENON Morren 1838
Aphanizomenon flos-aqae (L.) Ralfs
Gull (1) (rrr), Lake of Bays (rr), Marion (rrr), Mill (rr), Rosseau
(rrr), St. Mary (rrr), Thompson (rrr).
Smith — Phytoplankton of the Muskoka Region,
329
GLOEOTRICHIA J. G. Agardh 1842
Gloeotrichia echinulata (Eng. Bot.) P. Richter
Clear (2) (r).
A form of this species was noted from Long Lake with ovoid
hetereocysts and longer and narrower spores.
Heterocysts 8.75-10 y broad, 11.5-13 /* long; spores 7.5-8.75 //.
broad, 41-60 //. long.
Class PHAEOPHYCEAE
Order Chromulinales
CHRYSOSPHAERELLA Lauterborn 1896
Chrysosphaerella longispina Lauterborn
Burned Rock (s), Butterfly (s), Gull (1) (c), Echo (rrr), Hamer (r),
Harbon (s), Henshaw (rr), Joseph (aa), Leech (r), Long (1) (rr), Long
(2) (r). Loon (sss), Muskoka (cc), Otter (ss), Pine (s), Portage (cc),
Rosseau (cc), Round (1) (rrr), Round (2) (ccc), Rowle'y (s), Silver (1)
(s), Strange (ss), Vernon (r).
Order Isochrysidales
SYNURA Ehrenberg 1838
Synura dvella Ehr.
Basil (s), Black (1) (rr), Brush (rr), Burned Rock (r), Butterfly (r),
Cornell (s), Echo (ccc), Gull (2) (a), Harbon (sss), Henshaw (s), Joseph
(sss), Lake of Bays (r), Leech (sss), Long (1) (r). Loon (ss), Little Dud¬
ley (ss), McGowan (cc), Marion (sss), Mill (ccc), Muskoka (cc), Nelson
(s), Nine Mile (cc), Otter (r), Pennsylvania (rr), Pine (sss). Round
(1) (rr), Rosseau (ss), Rowley (ss), Silver (1) (sss), Strange (c),
Vernon (r), White (r).
Order Ochromonadales
UROGLENOPSIS Lemmermann 1899
Uroglenopsis Americana (Calk.) Lemm.
Gull (1) (rrr), Henshaw (rrr), Joseph (r), Leech (rr), Loon (rr),
Marion (rr), Mill (rrr), Muskoka (r), Otter (rr), Rowley (rrr). Strange
(r).
330 Wisconsin Academy of Sciences, Arts, and Letters.
DINOBRYON Ehrenberg 1835
Dinobryon bavaricum Imhof
Gull (1) (rr), Leech (ss), Strange (r).
Dinobryon stipitatum Stein
Musk oka (ss).
Dinobryon setularia Ehr.
(Burned Rock (sss), Henshaw (aaa), Leech (ss), Marion (ss), Mus-
koka (sss), Strange (r).
Dinobryon divergens Imhof
Rowley (r), Strange (r).
Order Phaeocapsales
PHAEOCOCCUS Borzi 1892
Phaeococcus planctonicus var. ovalis var. nov. Pl. VIII, fig. 20.
Cells ovoid to ellipsoid, scattered in small groups throughout the
gelatinous envelope. Chloroplasts two, laminate, with smooth
edges.
Breadth cells 5-6 u, length 8-9 u.
Leech (ss).
Class HETEROKONTEAE
CHLOROBOTRYS Bohlin 1901
Chlorobotrys limneticus G. M. Smith
Black (1) (r), Long (1) (rrr), Nine Mile (rrr), Vernon (rr).
BOTRYOCOCCUS Kutzing 1849
Botryococcus Braunii Ktz.
Big Dudley (rr), Black (1) (rr), Black (2) (rrr), Butterfly (rrr),
Gull (1) (rr), , Hamer (r), Harbon (rr), Henshaw (rrr), Lake of Bays
(rrr), Leech (rr), Little Dudley (r), Long (2) (rr), Marion (rr), Nelson
(rrr), Pennsylvania (rrr), Round (2) (rrr), Silver (2 (rr), Thompson
(rrr), Vernon (rrr).
Botryococcus protuberans var. minor G. M. Smith
Brush (rr), Loon (rrr), Muldrew (rrr), Rosseau (rrr), Rowley (rr),
St. Mary (rr), Vernon (r).
Smith — Phytoplankton of the Muskoka Region .
331
OPHIOCYTIUM Nageli 1849
Ophiocytium capitatum Wolle
Big Dudley (rr), Black (1) (rr), Harbon (rr), Marion (rrr), Mill
(rrr), Otter (rr), Thompson (rrr), Vernon (rrr).
var. longispinum (Mobius) Lemm.
Brush (rrr), Burned Rock (rrr), Loon (rrr), Nine Mile (rr).
Class CHLOROPHYCEAE
Order Volvocales
GONIUM Muller 1773
Gonium pectorale Muller
Clear (2) (rrr), Long (2) (rrr).
PANDORINA Bory 1824
Pandorina morum Bory
St. Mary (rrr).
EUDORINA Ehrenberg 1832
Eudorina elegans Ehrenberg
Brush (rrr), Burned Rock (rr), Butterfly (rr), Echo (rrr), Gull (rrr),
Harbon (rrr), Henshaw (rrr), Lake of Bays (rrr), Long (2) (rrr), Mill
(rrr), Mud (r), Muldrew (rrr), Nelson (rrr). Nine Mile (rrr), Otter
(rr), Pennsylvania (rr), Pine (rrr), Strange (r), Vernon (rrr).
VOLVOX L. 1758
Volvox aureus Ehrenberg
Mud (rrr).
Order Protococcales
Family PALMELLACEAE
GLOEOCYSTIS Nageli 1849
Gloeocystis gigas (Ktz.) Lag.
Clear (2) (rr), Silver (2) (rrr).
332 Wisconsin Academy of Sciences , Arts, and Letters.
SPHAEROCYSTIS Chodat 1897
Sphaerocystis Schroeteri Chodat
Big Dudley (rr), Black (1) (rr), Black (2) (ss), Brush (r), Burned
Rock (rr), Butterfly (rr), Cornell (rr), Corner (r), Hamer (rr), Hen-
shaw (r), Joseph (rr), Bake of Bays (sss), Leech (rrr), Long (2) (r),
Loon (rr), McGowan (r), Marion (ss), Mill (r), Mud (rr), Muldrew (rr),
Muskoka (rr), Nelson (rr), Nine Mile (rr), Otter (rrr), Pennsylvania
(rr), Rosseau (r), Rowley (r), St. Mary (sss), Strange (rrr), Vernon
(r), White (r).
GLOEOCYSTOPSIS G. M. Smith 1916
Gloeocystopsis limneticus G. M. Smith
Black (1) (rrr), Butterfly (rrr). Otter (rrr), Silver (2) (rr), Vernon
(rr).
ASTEROCOCCUS ScherfM 1908
Asterococcus limneticus G. M. Smith
Black (1) (r), Harbon (rr), Long (1) (rrr), McGowan (r), Muldrew
(rr), Muskoka (rrr), Otter (sss), Rowley (rr), Silver (1) (rrr).
Family DICTYOSPHAERIACEAE
DICTYOSPHAERIUM Nageli 1849
Dictyosphaerium Ehrenbergianum Nag.
Gull (1) (rrr), Pine (rrr).
Dictyosphaerium pulchellum Wood
Big Dudley (rrr), Black (1) (r), Brush (s), Burned Rock (r). But¬
terfly (rr), Cornell (r), Clear (1) (rr), Clear (2) (rr), Echo (rrr),
Hamer (rrr), Harbon (rrr), Henshaw (rrr), Lake of Bays (rrr), Leech
(rrr), Little Dudley (rr), Long (2) (rrr), Loon (rrr), McGowan (rrr),
Mud (rrr), Muldrew (rrr), Muskoka (rrr), Nelson (rr), Nine Mile (rr),
Pennsylvania (rr), Rosseau (rrr), Rowley (rrr), St. Mary (rr), Silver
(1) (rr), Vernon (rrr), White (r).
DIMORPHOCOCCUS A. Braun 1855
Dimorphococcus lunatus A. Br.
Basil (r), Big Dudley (rrr), Black (1) (rrr), Burned Rock (rr), But¬
terfly (rrr), Gull (1) (sss), Leech (rrr), Long (1) (rrr), Loon (rr),
Marion (rrr), Nelson (rr), Pine (rrr), Silver (1) (r).
Smith— Phytoplankton of the Muskoka Region .
333
WE ST ELLA de Wildeman 1897
Westell a botryoides (W. West) de Wild.
Butterfly (rrr), Mud (rrr), Muskoka (rrr), Portage (rrr), Round (1)
(rrr), Vernon (rr).
Family AUTOSPORACEAE
OOCYSTIS Nageli 1845
Oocystis Borgei Snow
Butterfly (rrr), Otter (r), Rowley (ss).
Oocystis parya W. & G. S. West
Black (1) (rrr), Leech (rrr).
Oocystis lacustris Chodat
Loon (rrr).
TETRAEDRON Kiitzing 1845
Tetraedron trigonum (Nag.) Hansg.
Black (1) (rr), Burned Rock (rrr), Butterfly (rr), Cornell (rrr),
Harbon (rr), Marion (rrr), Otter (rr), Pennsylvania (rrr), Pine (rrr),
Rowley (rrr), Silver (1), (rrr), White (rrr).
var. gracile (Reinsch) de Toni
Butterfly (rrr).
Tetraedron minimum (A. Br.) Hansg.
Black (1) (rrr), Butterfly (rrr), Clear (2) (rrr), Harbon (rrr).
Tetraedron qnadrilobatmn sp. nov. PL VIII, figs. 14-18.
Cells small, flattened, four-cornered, generally with all four
corners in the same plane, sometimes with halves of cell cruciate.
Sides of cells deeply emarginate and giving the cells a cruciform
appearance. Angles broadly rounded, smooth, without spines.
Side view of flattened cells broadly ellipsoid. Chloroplast single,
parietal, without a pyrenoid.
Diameter of cells 7.5-17 thickness of cells 4-6 /a.
Clear (2) (rrr). Corner (rrr), Lake of Bays (rrr), Vernon (rr).
This species is about the size of the familiar T. minimum (A.
Br.) Hansg., but the sides are more deeply emarginate and the
334 Wisconsin Academy of Sciences , Arts , and Letters .
corners broadly rounded. In the emargination of the sides and
in the rounded cell angles it resembles T. trilobatum, but I feel
that it should not be considered a variety of this species since
T. trilobatum is always triangular.
Tetraedron regulare Ktz.
Pine (rrr).
var. torsum (Turner) Brunnth.
Black (1) (rrr), Burned Rock (r), Butterfly (r), Cornell (rr), Harbon
(sss), Loon (rrr), Pennsylvania (rrr), Silver (1) (rrr).
Tetraedron victorieae var. major G. M. Smith
Butterfly (rrr). Black (1) (rrr).
Tetraedron caudatum (Corda) Hansg.
Burned Rock (rrr), Clear (2) (rrr).
Tetraedron gracile (Reinsch) Hansg. PI. VIII, figs. 12-13.
Butterfly (r), Gull (1) (rrr), Harbon (rrr), Pine (rrr).
The branching of the processes in certain of the cells collected
from Harbon and Butterfly lakes was greatly reduced, but this
reduction in the branching does not seem sufficiently constant to
warrant varietal recognition.
Tetraedron limneticum Borge.
Butterfly (rr), Clear (1) (rrr), Harbon (rr), Long (1) (rrr).
Tetraedron spiniferum sp. nov. PL VIII, figs. 9-11.
Cells of medium size, tetragonal or pentagonal, flattened or py¬
ramidal. Processes at angles slightly attenuated, once or twice
branched. Each process ending in two or three (generally two)
long, slender, divergent spines that are either straight, slightly
incurved or recurved.
Diameter of cells 45-52 /x; length of spines 7.5-11 p.
Burned Rock (r).
The cell shape and branching of the processes is quite similar
to that of T. limneticum , but the terminal spines are quite differ¬
ent. These spines are true spines and not similar to the setae
found in Polyedriopsis .
Smith— Phytoplankton of the Muskoka Region.
335
Tetraedron planctonicum G. M. Smith
.Black (1) (rrr), Butterfly (rrr), Cornell (rr), Cull (1) (rr), Otter (rr),
Pine (rr), Silver (1) (rrr), Silver (2) (rrr).
Tetbaedron enorme (Balls) Hansg.
Cornell (rrr), Rowley (rrr), St. Mary (rrr).
MICRACTINIUM Fresenius 1858
Micractinium pusillum Fresen.
Round (1) (rrr).
LAGERHEIMIA Chodat 1895
Lagerheimia subsalsa Lemm.
Yernon (rrr).
SELENASTRUM Reinsch 1867
Selenastrum gragile Reinsch
Black (1) (rr), Brush (rrr), Cornell (rrr), Pine (rrr), Vernon (rrr),
White (rrr).
Selenastrum Bibraianijm Reinsch
Butterfly (rrr).
ANKISTRODESMUS Corda 1838
Ankistrodesmus Falcatus (Corda) Ralls
Big Dudley (sss), Black (1) (rr), Cornell (rrr), Long (2) (rrr), Nine
Mile (rrr), Thompson (rr).
var. mirabilis (W. & G. S. West) G. S. West
Butterfly (rrr), Silver (1) (rrr), Thompson (rrr).
Ankistrodesmus convolutus Corda
Almanach de Carlsbad 1838: 199. pi. 2. fig. 19. 1838.
Black (1) (sss).
The chloroplast ol the cells was frequently interrupted and with¬
out pyrenoids.
336 Wisconsin Academy of Sciences, Arts , and Letters .
Ankistrodesmus spiralis var. fasciculatus var. nov. PL VIII,
fi g. 19.
Cells curved or sigmoid, twisted around one another, united in
colonies of 50-200 cells with the median portion of the cells ap¬
posed and the apices free.
Cells 3.75-5 y broad, 55-70 /x long. Colonies 75-180 fx in
diameter.
Lake of Bays (rrr). Little Dudley (rr), Rosseau (rrr), Silver (1)
(rrr), Vernon (rrr).
The variety is distinguished by the large number of cells in the
colony, the cells being twisted around one another to form a
dense mass. The typical A. spiralis as I have found it has but
four or eight cells in the colony while in this variety the number
of cells is always large.
CLOSTERXOPSIS Lemmermann 1898
Closteriopsis longissimum var. tenuissimum var. nov. PL VIII,
fig. a.
Cells much narrower, arcuate or lunate.
Cells 2-3 jul broad, 175-210J p long.
Harbon (rrr).
This variety is much narrower than the variety tropicum or
the typical C. longissimum . The cells are much longer than An¬
kistrodesmus falcatus var. mirabilis , while the central row of py-
renoids at once excludes the alga from Ankistrodesmus . The alga
looks even more like a small Closterium species, but cannot be
considered a Closterium since there is but a single chloroplast in
the cell.
SCHROEDERIA Lemmermann 1898
Schroederia setigera (Schrjoder) Lemm.
Marion (ss).
Schroederia Judayi GL M. Smith
Big Dudley (ss), Black (1) (rr), Brush (rr), Corner (rr), Harbon
(rrr).
Smith — Phytoplankton of the Muskoka Region.
337
QUADRIGULA Printz 1915
Quadrigula Pfitzeri (Schrpder) G. M. Smith.
Basil (rr), Big Dudley (sss), Black (1) (rrr), Black (2) (rr), Brush
(rr), Burned Rock (rr), Butterfly (sss), Cornell (rrr), Corner (rr),
Gull (1) (rr), Henshaw (rrr), Lake of Bays (rr), Little Dudley (rrr),
Long (2) (rr), Marion (rrr), Mill (rr), Muldrew (rrr), Muskoka (rr),
Nine Mile (rrr), Otter (rrr), Round (2) (rr), Rowley (rrr), Silver (1)
(rrr), St. Mary (r), Thompson (rrr), Vernon (r), White (rr).
ELAKTOTHRIX Wille 1898
Elaktothrix gelatinosa Wille
Black (1) (rrr), Clear (1) (rrr), Hamer (rrr), Leech (rr), Long (1)
(rrr), Mill (rr) , Muskoka (rrr), Otter (rr), Strange (rr), Vernon (rrr).
KIRCHNERIELLA Sehmidle 1893
Kirchneriella lunaris (Kirchner) Mobius
Basil (r), Big Dudley (rrr), Brush (rrr), Burned Rock (rrr), Butterfly
(r), Echo (rrr), Gull (1) (rrr), Gull (2) (rrr), Hamer (rr), Harbon
(rrr), Henshaw (rr), Leech (rrr), Long (1) (rrr), Long (2) (rrr), Loon
(rr), Mill (r), Muldrew (rrr), Muskoka (rr), Nine Mile (rrr), Otter (r),
Pine (rrr), Rowley (rr), St. Mary (rr), Silver (1) (rr), Thompson (rrr),
Vernon (ss).
var. irregularis G. M. Smith
Silver (1) (rrr).
Kirchneriella obesa (W. West) Sehmidle
Basil (rrr), Black (1) (rrr), Brush (rrr), Vernon (rrr).
Kirchneriella arcuata sp. nov. PI. VIII, figs. 1-3.
Cells with apices somewhat attenuated to a rounded apex, ar¬
cuate, lying some distance from one another within the homo¬
geneous gelatinous colonial envelope. Colonies generally four-
celled, rarely with more than sixteen cells. Cellular arrangement
within the colony indefinite. Chloroplast on convex side of cells,
without a pyrenoid, hyaline spot on concave side of cell conspic¬
uous.
Cells 2.S-5.3 p broad, 8-17.5 p long.
Big Dudley (s), Muskoka (rrr). Silver (1) (rrr).
The species is distinguished by the cell shape and the shape
of the chloroplast as well as by the small number and arrange¬
ment of the cells within the colonial envelope.
22— S. A. L.
338 Wisconsin Academy of Sciences , Arts , and Letters.
Kirchneriella contorta (Schmidle) Bohlin
Black (1) (rrr) , Butterfly (rr), Corner (rrr), Marion (rrr), Otter
(rrr) , Silver (2) (rrr), Thompson (rr), Vernon (rrr).
Kirchneriella elongata G. M. Smith
Harbon (rrr), Muldrew (rrr), Nine Mile (rrr).
CRUCIGENIA Morren 1830
Crucigenia rectangularis (Nag.) Gay
Big Dudley (rrr), Lake of Bays (rrr), Long (1) (rrr), Muldrew (rrr).
Nine Mile (rrr),iRound (2) (rrr), Rowley (rrr), Vernon (rr).
Crucigenia truncata G. M. Smith
Butterfly (rr), Vernon (r).
Crucigenia apiculata (Lemm.) Schmidle
Big Dudley (rr), Black (1) (rrr), Harbon (rrr).
Crucigenia quadrata Morren
Black (1) (rr), Butterfly (r), Harbon (rrr), Marion (rrr), Muldrew
(rrr), Nine Mile (rrr), Rowley (rrr), St. Mary (rrr), Vernon (rr).
Crucigenia Tetrapedia (Kirchner) W. & G. S. West
Big Dudley (rrr), Black (1) (rrr), Burned Rock (rrr), Vernon (rr).
SCENEDElSMUS Meyen 1829
SCENEDESMUS OBLIQUUS (Turp.) Ktz.
Black (1) (rrr), Long (1) (rrr), Mud (rrr).
SCENEDESMUS DIMORPHUS (Turp.) Ktz.
Vernon (rrr).
Scenedesmus INCRASSATULUS var. mononae G. M. Smith PL VIII.
figs 4-7.
Trans. Wis. Acad. Sci., Arts, & Lett. 183: 440. PI. 29, figs. 81-83. 1916.
Cells 3.75-6 p broad, 12.5-16 p long (without apiculations).
Colonies 12.5-26 p broad, 17.5-26 p long.
Butterfly (r).
Smith — Phytoplankton of the Muskoka Region. 339
The cells of this variety as found in Butterfly Lake were slightly
longer than those from Lake. Monona in Wisconsin. A few cases
of eight-celled colonies with the two-ranked arrangement shown
in figure 4 were observed.
SCENEDESMUS BIJUGA (Turp.) Lag.
St. Mary (rrr), Vernon (rr).
Scenedesmus ARCUATUS Lemm.
Burned Rock (rrr), Butterfly (r), Vernon (rrr).
var. platydisca G. M. Smith
Butterfly (rrr). Gull (1) (rrr), Harbon (rrr), St. Mary (rrr), Vernon
(rrr).
Scenedesmus armatus (Chodat) Gr. M. Smith
Butterfly (rr), Harbor (rrr), Pine (rrr).
var. major G. M. Smith
Burned Rock (rrr).
Scenedesmus brasiliensis Bohlin
Basil (rrr), Big Dudley (rrr), Black (1) (rrr). Clear (2) (rrr), Cor¬
nell (rrr), Little Dudley (rr), Loon (rrr), Mud (rrr), Portage (rrr),
Silver (1) (rrr), Vernon (rrr), White (rrr).
Scenedesmus hystrix Lag.
Black (1) (rrr), Nine Mile (rrr).
Scenedesmus denticulatus Lag.
Big Dudley (rr), Black (1) (rrr), Cornell (rrr), Marion (rrr), Pine
(rrr), Round (1) (rrr), Vernon (rrr), White (rrr).
Scenedesmus abundans var. longicauda G. M. Smith
Big Dudley (rrr), Clear (2) (rrr).
Scenedesmus quadricauda (Turp.) Breb.
Big Dudley (rr), Clear (2) (rrr), Harbon (rrr), Rowley (rrr), Silver
(1) (rrr).
var. quadrispina (Chodat) G. M. Smith
Burned Rock (rrr).
var. longispina (Chodat) G. M. Smith
Black (1) (rrr), Brush (rrr), Butterfly (rrr), Harbon (rrr).
340 Wisconsin Academy of Sciences , Arts, and Letters .
var. Westii G. M. Smith
Butterfly (rr), Clear (2) (rrr), Little Dudley (rrr), Otter (rrr).
The spines of the specimens from Clear Lake were somewhat
longer than those of this variety from other stations and reached
a length of 22.5 /x.
var. maximum W. & G. S. West
Harbon (rrr), Marion (rrr), iStrange (rrr).
COELASTRUM Nageli 1849
COELASTRUM MICROPORUM Nag.
iBlack (1) (rrr), Brush (r), Burned Rock (rrr), Butterfly (rrr),
Harbon (rrr), Joseph (rrr), Rosseau (rrr), Silver (2) (rrr), St. Mary
(rrr), Vernon (rrr).
Coelastrum CAMBRicuM Archer
Basil (rr), Big Dudley (rr), Black (1) (r), Brush (rrr), Butterfly
(rrr), Cornell (rr), Echo (rrr), Gull (1) (rrr), Harbon (rrr), Leech
(rrr), Little Dudley (rr), Loon (rrr), Marion (rr), Nine Mile (rr),
Otter (rrr), Round (1) (rrr), Round (2) (rrr), Silver (1) (rrr), Thomp¬
son (rrr), Vernon (r), White (r).
var. intermedium (Bohlin) G. S. West
Jour. Linn. Soc. Bot. 88: 137. 197.
Coelastrum pulciirum var. intermedium Bohlin, iBih. Kgl. Svenska Vet-
Ak. Handl. 28, Afd. 3, No. 7; 85. PI. 2, figs. 16, 71. 1897.
Butterfly (rrr).
Coelastrum Morus W. & G. S. West
Jour. Bot. 84; 381. PI. 361. fig. Jf. 1896; Trans. Roy Soc. Edinb. 41:
607. PI. 6, fig. 12. 1905.
Diameter of cells 10-12 /x. Diameter of colonies 35-38 /x.
Silver (2) (rrr).
The record for this species is based upon two colonies. The
dimensions given above are smaller than the Wests give, and the
presumption is that the colonies observed in Silver Lake were
immature.
SORASTRUM Kiitzing 1845
SORASTRUM SPINULOSUM Nag.
Basil (rrr), Harbon (rrr).
Smith — Phytoplankton of the Muskoka Region .
341
Sorastrum americanum (Bohlin) Schmidle
Butterfly (rrr), Clear (2) (rrr), Harbon (rrr).
Family HYDRODICTYACEAE
PEDIASTRUM Meyen 1829
Pediastrum integrum Nag.
Marion (rrr).
Pediastrum araneosum Racib.
Bu-rned Rock (rrr), Clear (2) (rrr), Cornell (rrr), Cull (1) (rrr), Hen-
shaw (rrr), Lake of Bays (rrr), Little Dudley (r), Long (1) (rr), Long
(2), (rrr), Marion (rrr), Muskoka (rrr), Nine Mile (rr), Pennsylvania
(rrr), Rowley (rrr), St. Mary (rrr), Silver (1) (rrr), Strange (rrr),
Vernon (rrr), White (rrr).
Pediastrum Boryanum (Turp.) Menegh.
Big Dudley (rrr), Butterfly (rrr), Gull (1) (rrr), Joseph (rrr), Long
(2) (rrr), Mill (rrr), Nine Mile (rrr), Silver (1) (rrr).
var. longicorne Racib.
Clear (2) (rrr).
Pediastrum duplex Meyen
Gull (1) (rrr), Long (1) (rrr), Pine (rrr), Muskoka (rrr), St. Mary
(rrr), Silver (1) (rr).
var. clathratum (A. Br.) Lag.
Big Dudley (rr), Black (1) (r), Brush (sss), Burned Rock (r), Loon
(rr), Mill (rr), Rowley (rr), Vernon (rrr).
var. reticulatum Lag.
Cornell (rr),Echo, (rrr), Gull (2) (rrr), Pine (rrr), White (rrr).
var. gracillimum W. & G. S. West
Butterfly (r), Harbon (r).
var. cohaerens Bohlin
Butterfly (rrr), Cornell (rrr), Harbon (rr), Vernon (rrr).
Pediastrum tetras (Ehr.) Ralfs
Big Dudley (rrr), Black (1) (rrr), Brush (rrr), Cornell (rrr), Harbon
(rrr), Joseph (rrr), Leech (rrr), Long (1) (rrr), Marion (rrr), Round
(2) (rrr), Rowley (rrr), St. Mary (rrr), Silver (1) (rrr), White (rr)..
Vernon (rrr).
342 Wisconsin Academy of Sciences , Arts , and Letters.
Family PLANOSPORACEAE
CHARACIUM A. Braun 1849
Characium stipitatum (Bachm.) Wille
Basil (rrr), Black (1) (rr), Brush (sss), Butterfly (rrr), Echo (rrr),
Gull (1) (rr), Gull (2) (rr), Marion (r), Muldrew (rr), Nelson (r).
Nine Mile (rr), Rosseau (rrr), Silver (1) (r), Thompson (rrr), Vernon
(rr).
Characium curvatum G. M. Smith
Big Dudley (rrr), iBlack (1) (rr), Lake of Bays (rrr), Muskoka (rrr),
Rowley (rr), Silver (1) (rr), Vernon (rr).
Family DESMIDIACEAE
GONATOZYGON DeBary 1856
Gonatozygon aculeatum Hastings
Brush (rr).
NETRIUM Nageli 1849
Netrium digitus Itzighs. & Roth.
Pennsylvania (rrr).
PENIUM Brebisson 1844
Penium margaritaceum (Ehr.) Breb.
Gull (2) (rrr).
CLOSTERIUM Nitzsch 1817
Closterium costatum Corda
Long (1) (rrr).
Closterium Ehrenbergii Menegh.
Leech (rrr).
Closterium angustatum Ktz.
Brush (rr).
Closterium rostratum Ehr.
Marion (rrr).
Smith — Phytoplankton of the Muskoka Region.
343
PLEUROTAENIUM Nageli 1849
Pleurotaenium Subcoronulatum var. detum W. & G. S. West
Mill (rrr), Nelson (rrr), Nine Mile (rrr), Round (1) (rrr) .
Pleurotaenium maximum (Reinsch) Lundell
McGowan (rrr).
Pleurotaenium Ehrenbergii (Breb.) DeBary
Cornell (rrr).
TRIPLOCERAS Bailey 1851
Triploceras gracile Bailey
Leech (rrr), Long (1) (rrr), Nelson (rrr).
EUASTRUM Ehrenberg 1832
Euastrtjm didelta (Turp.) Ralfs
Brush (rrr).
Euastrum pulchellum Breb.
Silver (1) (rrr).
Euastrum verrucosum var. alatum Wolle
Mud (rrr).
var. reductum Nords.
Gull (1) (rrr).
MICRASTERIAS C. A. Agardh 1827
Micrasterias pinnatifida (Ktz.) Ralfs
Otter (rrr).
Micrasterias laticeps Nords.
Black (1) (rrr), Brush (rr), Pennsylvania (rr), Strange (rrr).
Micrasterias muricata (Bailey) Ralfs
Leech (rrr). Long (1) (rrr).
Micrasterias denticulata var. angulosa (Hantzsch) W. & G. S.
West
Little Dudley (rrr), Long (2) (rrr).
Micrasterias apiculata var. fimbriata (Ralfs) Nords.
Burned Rock (rrr), Clear (2) (rrr), Little Dudley (rrr).
344 Wisconsin Academy of Sciences , Arts , and Letters.
var. apiculata forma spinosa (Bisset) W. & G. S. West
Brush (rrr), Long (1) (rrr), Pennsylvania (rrr).
Ml CR AS TERI AS sol (Ehr.) Ktz.
Big Dudley (rrr), Burned Rock (rrr), Otter (rrr), Rowley (rrr).
var. ornata (Nords.) W. & G. S. West
Corner (rrr), Otter (rrr).
var. ornata forma elegantior G. S. West
Echo (rrr), Muskoka (rrr).
Micrasterias radi at a Hassall
Clear (2) (rrr). Little Dudley (rrr).
var. simplex (Wolle) comb. nov. PL IX, figs. 1-5.
Micrasterias furcata var. simplex Wolle. Bull. Torrey Bot. Club. 12:
128. PI. 51, figs. 6, 7. 1885.
Big Dudley (rr), Black (1) (r), Brush (rrr), Hamer (rr), Long
(1) (rr), Nelson (rrr), Otter (rrr), Round (2) (rr), White (rrr).
The great variation in the subdivision of the lateral lobes of
this species is well known to all students of the group. Johnson
has studied these variations (Bot. Gaz. 19: 58-60, PI. 6 , figs. 7-13
1894) and reached the conclusion that it is impossible to recognize
any of the varieties which have been based upon this reduction
in the number of lateral lobes. The Wests in their Monograph
of the British Desmidiacaeae (2: 114—115) arrive at the same con¬
clusion.
In the collections that I have made from Ontario lakes the re¬
duction in the branching of the lateral processes to a simple bifur¬
cation is the predominant type. Although there are variations
from this in that some of the semicells approach the typical M.
radiata. I feel that the variety with the reduced number of
lateral branches is sufficiently distinct to warrant recognition and
that Wolle was justified in establishing the variety simplex.
var. gracillima var. nov. IX, figs. 6-11.
Semicells with a single lateral process that is deeply incised to
form two divergent branchlets. Unbranched portion of lateral
lobes much narrower than in var. simplex and with sides sub-
Smith — Phytoplankton of the Muskoka Region. 345
parallel. Length of polar lobe nearly twice the greatest breadth
of lateral lobes. Polar lobe narrower, with parallel sides.
Length 135-165 /x ; breadth 130-150 /x ; breadth basal portion po¬
lar lobe 15-20 /x; greatest breadth lateral processes 13-15 /x;
breadth isthmus 15 /x.
Butterfly (rrr), Burned Rock (rrr), Gull (1), (c), Muldrew (r), Pine
(rrr) , Silver (1) (s).
This variety is remarkably constant in character, and all indi¬
viduals noted had but the single subdivision of the lateral lobe.
It is much more delicate than the variety simplex , the margin of
the lateral lobes never showing the marked convexity found in
that variety. The polar lobe is also much more prominent and
narrower than in the typical form.
Micrasterias Mahabuleshwarensis forma dichotoma forma nova.
PL IX, figs. 12-14; PL X, fig. 1.
Differing from the typical form in the development of rudi¬
mentary accessory processes on the inner side of the incision of
the lateral lobes.
Length 150-165 /x; breadth 150-160 /x; breadth polar lobe 20-25
/x ; breadth isthmus 20-31 /x.
Black (1) (rrr). Brush (rrr), Nelson (rrr), Nine Mile (rrr), White
(r).
The secondary lobelets on the lateral lobes of this form are
quite irregular and are not normally found on all eight lateral
branches of a cell but only on some of them. They may be re¬
duced to a small mammillate spine, but are always present in
one form or another. This form possesses a great resemblance to
M. americana (Ehr.) Ralfs, but is placed with M. Mahabulesh¬
warensis because both sides of the lateral lobes are denticulate and
the polar lobe has the shape of M. Mahabuleshwarensis with long
apical processes.
Micrasterias foliacea Bailey
Gull (1) (rrr), Rowley (rrr).
COSMARIUM Corda 1834
Cosmarium contractum var. papillatum forma minor forma nova.
PL X, figs. 2-5.
Length 30-32.5 n; breadth (without papillae) 23-25 /x; papillae
1.25 /x long; breadth isthmus 6.25-7.5 y; thickness 15-17.5 /x.
Thompson (aa).
346 Wisconsin Academy of Sciences , Arts , and Letters.
Cosmarium quadrum Lundell
Brush (rrr).
Cosmarium ornatum Ralfs
Black (1) (rrr).
XANTHIDIUM Ehrenberg 1834
Xanathidium armatum var. ceryicorne W. & G. S. West.
Leech (rrr), Long (1; (irr).
Xanthidium tetracentrotum var. hexagonum var. nov. Pl. X,
figs. 6-8.
Apex of semieells hexagonal, without spines, basal angles of
semicells with one or two long, outwardly divergent spines. Cen¬
tral area of semicells smooth, slightly thickened but without orna¬
mentation.
Length 35-40 /x; breadth (without spines) 37.5-40 u, (with
spines) 65-70 g.; thickness 22 /x ; breadth isthmus 10-12 /i.
Portage (rr).
The hexagonal upper half of the semicell is quite unlike that of
any other known variety of this species. The variety is also char¬
acterized by longer spines and by a central area without ornamen¬
tation.
Xanthidium subhastiferum var. Toweri (Cushman) comb. nov.
PI. X, figs. 9-13.
Xanthidium hastiferum var. Toweri Cushman; Bull. Torrey Bot. Club 31:
163. PI. 7, fig. 9. 1894.
Length (without spines) 35-38/x, (with spines) 50-60 /x; breadth
(without spines) 35-40 /x, (with spines) 65-75 /x; breadth isthmus
8.5-11.5 /x; length spines 12.5-22 /x.
Echo (rrr), Loon (rrr), Strange (rr).
This variety comes more appropriately under X. subhastiferum
than under X. hastiferum. The spines on both the upper and the
lower angles of the semicells are upwardly divergent in a grace¬
ful curve, while the spines of the typical X , subhastiferum are
straight and much shorter. At times there is a supplementary
pair of smaller spines, a single spine, or the rudiments of spines
Smith — Phytoplankton of the Muskoka Region.
347
on the apex of the semicells. The dimensions of the individuals
that I have found are smaller than those found by Cushman.
Xanadithium antilop aeum (Breb.) Ktz.
Leech (rrr).
var. limneticuxn var. nov. PI. X, figs. 14-16.
Semicells with spines on lower angles gracefully curved and
those on upper angles straight and divergent. Central area color¬
less or brownish, generally with an arc of 4-8 subapical pores,
more rarely with an irregular ring of pores.
Length (without spines) 51-56 y, (with spines) 81-90 y;
breadth (without spines) 47-50 y, (with spines) 87-100 y;
breadth isthmus 12-14 y.
Gull (1) (ss). Hamer (rrr), Muskoka (rrr), Pennsylvania (r), Sil¬
ver (1) (s).
The elegant curvature and length of the spines in this variety
distinguish it from the typical X. antilopaeum. The ornamenta¬
tion below the apex suggests the variety polymazum, but in this
variety the marking is due to pores while in the variety polymazum
the marking is due to small rounded granules.
var. polymazum Nords.
Black (1) (rrr), Burned Rock (rrr), Gull (1) (rr), Joseph (rrr).
Little Dudley (rrr), Long (2) (rrr), Marion (rrr), Otter (rrr), Penn¬
sylvania (rrr), Round (2) (rrr), Rowley (rrr), Silver (1) (rrr), White
(rrr).
var. minneapoliensis Wolle
White (rrr).
ARTHRODESMUS Ehrenberg 1836
Arthrodesmus incus (Breb.) Hass.
Otter (rrr).
var. ralfsii W. & G. S. West
Strange (rrr).
var. indentatus W. & G. S. West.
Clear (1) (rrr).
348 Wisconsin Academy of Sciences , Arts, and Letters.
Arthrodesmus constrictus sp. nov. PI. X, figs. 17, 18.
Cells of medium size, breadth (without spines) slightly greater
than the length, deeply constricted, sinus rectangular, with sides
outwardly divergent; semicells broadly cuneate with convex sides
and a slightly convex apex, lateral angles somewhat pointed and
furnished with strong, straight, somewhat divergent spines.
Isthmus cylindrical, elongate, with a slight constriction at the
point where cell division takes place. Vertical view elliptic, each
pole furnished with a single spine.
Length 22.5-25 /a; breadth (without spines) 28-30 /a, (with
spines) 58-68 /a; breadth of isthmus 4 /a; length of isthmus 5 /a;
spines 15-19 /a long.
Long (1) (rrr). Pine (rr).
The semicells of this species are much broader in proportion
to their length than those of any other species of the genus. The
isthmus is also peculiar in that the point of origin of the new
semicell is shown by an annular depression, whereas in other
species the point of union of the two semicells is not discernible.
Arthruuesmus triangularis Lag.
Hamer (r), Harbon (rr), Vernon (rr).
var. inflatus W. & G. S. West
Portage (rr).
var. subtriangularis W. & G. S. West
Black (rr), Burned Book (rr), Gull (2) (rrr), Joseph (r), Long (2)
(rrr), Muskoka (rr), Nelson (sss), Silver (1) (r), Thompson (r).
Arthrodesmus quiriferus var. subparallelus var. nov. PI. X,
figs. 19, 20.
Sides of cells straight, not curved, and with the long spines at
the angles only slightly divergent.
Length (without spines) 20-22.5 /a; breadth (without spines)
20-24 /a; breadth isthmus 6-7 /a; spines 28-31 /a long.
Corner (rrr), Henshaw (rrr), Loon (rr), Rowley (rr).
The dimensions of the cell in the variety are approximately the
same as in the typical form, while the spines are somewhat shorter
in the variety. This variety approaches biradiate forms of
Staurastrum jaculiferum W. & G. S. West ever more than the
Smith — Phytoplankton of the Muskoka Region. 349
typical A. quiriferus , but the retuse apices of the semicells are
not found in S. jaculiferum.
Arthrodesmus subulatus Ktz.
Big Dudley (rrr), Muskoka (rr), Vernon (rr).
STAURASTRUM Meyen 1829
Staurastrum muticum Breb. Pl. X, figs. 21, 22.
Length 27.5 n; breadth 23-26 breadth isthmus 6 y.
Butterfly (sss), Harbon (rrr).
Staurastrum grande Bulnh.
Rowley (rrr).
Staurastrum cuspid atum Breb.
Big Dudley (rrr), Black (rrr), Butterfly (rrr), Gull (1) (rrr), Joseph
(rrr), Lake of Bays (rrr), Otter (rr), Portage (rrr), Rowley (sss).
var. divergens Nords.
Brush (rrr).
var. canadense var. nov. PI. X, figs. 23, 24.
Isthmus of cells with a slight constriction at the point of origin
of new semicells,
Length 22-25 breadth (without spines) 22-24 y, (with spines)
45-58 fi ; breadth isthmus 5 y.
Clear (2) (rrr), Echo (rrr), Muskoka (rrr), Pine (r), Rosseau (rrr),
Round (1) (rrr), Strange (rrr).
Staurastrum dejectum Breb.
Burned Rock (rrr), Echo (rrr), Harbon (rrr), Henshaw (rrr), Loon
(r), Muskoka (rrr), Round (2) (rr), Rowley (r).
var. Tellami W. & G. S. West
Portage (rr).
Staurastrum curvatum W. West
Black (1) (rrr), Burned Rock (rrr), Butterfly (rr), Harbon (rrr),
Loon (rr), Muldrew (sss), Muskoka (r), Nelson (rrr), Pennsylvania
(rrr), Round (2) (rrr), Rowley (r), Silver (2) (rrr), Vernon (ss).
Staurastrum megacanthum Lund.
Gull (2) (rrr), Lake of Bays (rrr), Long (1) (rrr), Long (2) (rr),
Nelson (rrr), Strange (rrr).
350 Wisconsin Academy of Sciences , Arts, and Letters.
var. scoticum W. & G. S. West
Gull (1) (rr), Hamer (rrr), Silver (1) (rr).
Staurastrum Brebissonii var. paucispinum var. nov. PL X,
fig. 25; PL XI, figs. 1-5.
Cells small, with three or four large spines at the angles and
one or two small spines on the apex of the semicells just within
the angles.
Length 31-34 fx; breadth (without spines) 32-40 /*, (with
spines) 45-52 /x; breadth isthmus 8.5-10 /x.
Black (1) (ss), Burned Rock (rrr), Echo (rrr), Muldrew (rrr).
This variety is quite similar to the variety heterocanthum but
differs in the possession of only one or two smaller spines at the
cell angles and in a general occurrence of four large spines at the
angles.
Staurastrum setigerum var. brevispinum var. nov. Pl. XI,
figs. 6-8.
Semicells with several stout aculei at the corners and with two
rows of 6-8 short aculei across the median portion of the semi-
cell. Vertical view with one row of the smaller spines in profile
on the margin and an inner row of smaller spines following
the gentle concavity of cell margins. Large spines at the angles
as in the front view.
Length 35-40 fx; breadth (without spines) 37-^3 fx, (with
spines) 47-55 /x; breadth isthmus 12-14 fx.
Gull (1) (rrr), Gull (2) (rrr), Pennsylvania (rrr). Pine (rr). Rowlev
(rrr), Silver (rr), Vernon (rr).
Staurastrum minnesotense Wolle. Pl. XI, fig. 9.
Muldrew (rrr), Otter (rrr).
The Wests (Trans. Linn. Soc. 2 Ser. Bot. 5; 260. Pl. 17, fig.
15, 1896) infer that regular specimens with pairs of spines in each
semicell are quite rare. Practically all specimens collected in On¬
tario possessed the six stout and twelve delicate spines on each
semicell. The chief variation from the type is the lack of some
of the delicate spines on the sides of the semicells.
Staurastrum subnudibrachiatum W. & G. S. West
Brush (rr). Burned Rock (rrr), Gull (1) (rrr), Joseph (rrr), Loon
(r), Portage (r), Silver (2) (r).
Smith — Phytoplankton of the Muskoka Region. 351
Staurastrum tetracerum var. trigonum Lund.
Black (r), Butterfly (rrr), Harbon (rrr), Leech (rrr), Otter (rrr).
Staurastrum leptocladum Nords.
Basil (rr), Big Dudley (rr), Black (1) (ss), Burned Rock (sss), But¬
terfly (sss), Harbon (ss), Gull (1) (rrr), Gull (2) (rrr), Little Dudley
(sss), Long (1) (cc), Long (2) (c), Loon (s), Muldrew (ss), Muskoka
(ss), Nelson, (ss), Otter (rrr), Rosseau (rrr), Round (2) (rrr), St. Mary
(rrr), Thompson (rrr), Vernon (ss). .
W. & G. S. West (Jour. Linn. Soc. Bot. 33:317, 1898) have
reached the conclusion that Nordstedt overlooked the subapical
spines in his original description of the species, so that the variety
cornutum described by Wille cannot be recognized.
var. denticulatum var. nov. PI. XI, fig. 14.
Variety with a broad, slightly emarginate sub-apical tooth in¬
stead of a sharp spine.
Dimensions as in the typical form.
Corner (r), Echo (rrr), Nine Mile (ss), Pennsylvania (rr), Pine (sss),
Round (1) (rrr), Silver (1) (rr), Silver (2) (rrr).
var. sinuatum Wolle
Clear (ss), Gull (2) (rrr), Harbon (rrr), Mud (rrr), Round (2) (rrr),
White (rrr).
var. insigne W. & G. S. West.
Pennsylvania (rrr).
Staurastrum Johnsonii W. & G. S. West
Basil (sss), Big Dudley (ss), Black (1) (sss), Black (2) (rr), Brush
(c), Burned Rock (sss), Butterfly (rr), Clear (1) (rrr), Clear (2) (rr),
Echo (sss), Gull (1) (s), Gull (2) (r), Harbon (rr), Henshaw (sss),
Leech (c), Little Dudley (rr), Long (1) (r), Long (2) (s), Loon (rr),
Mill (rrr), McGowan (rrr), Muldrew (ss), Muskoka (rrr), Nelson (sss),
Nine Mile (sss), Otter (rr), Pine (rrr), Portage (rrr), Round (2) (sss),
Rowley (rr), St. Mary (rrr), Silver (1) (r), Silver (2) (rrr), Strange
(ss), Thompson (sss).
Staurastrum americanum (W. & G. S. West) comb. nov.
Staurastrum grallatorium var. americanum W. & G. S. West, Trans. Linn.
Soc. 2 Ser. Bot. 5: 265. PI. 17 , fig. 15. 1896.
This alga, which was described by the Wests as a variety of
S. grallatorium Nords., is sufficiently differentiated to warrant
352 Wisconsin Academy of Sciences, Arts, and Letters.
specific recognition. It differs from S. grallatorium in the smooth
convex apex of the semicells, the deeper constriction at the isthmus,
and the denticulation of the arms. I have not noted the typical
organism in Ontario although I have it from the plankton of Wis¬
consin lakes.
var. longiradiatum var. nov. PL XI, figs. 10, 11.
Semicells with longer, gracefully divergent processes ; wall some¬
what thicker at the apex of semicells than in other portions.
Length (without processes) 30 ft, (with processes) 80-95 /x ;
breadth (without processes) 25 ft, (with processes) 132-150 ft;
breadth isthmus 7. 5-8. 5 ft.
Gull (1) (rr), Silver (1) (rrr).
var. triradiatum var. nov. PL XI, figs. 12, 13.
Semicells triangular, not flattened, with the angles produced
into long, very slightly tapering processes with undulate sides.
Apex of semicells with walls slightly thickened.
Length (without processes) 25 ft, (with processes) 62-68 ft;
breadth (without processes) 17.5-20 ft, (with processes) 60-70 ft;
breadth isthmus 7.5-10 ft.
Butterfly (ss).
Staurastrum aspinosum var. verrucosum var. nov. Pl. XI,
figs. 17-20.
Semicells with two apical verrucae between each adjacent pair
of the three processes. Arms of semicells lying above one an¬
other and not alternately arranged.
Length (without processes ) 25-28 ft, (with processes) 50-64 ft;
breadth (without processes) 22 ft, (with processes) 65-75 ft;
breadth isthmus 10 ft.
©lack (1) (sss), Burned Rock (rr), Gull (1) (sss), Gull (2) (rr),
Hamer (sss), Joseph (rrr), Long (1) (rrr), Muldrew (rr), Portage (rrr),
Silver (2) (sss).
The processes of this variety possess the peculiar spines that
characterize the species but they are laxer than the type. The
apical verrucae found in this variety are not mentioned by Wolle.
Staurastrum paradoxum Meyen.
Brush (rr), Clear (1) (rrr), Thompson (rrr).
Smith — Phytoplankton of the Muskoka Region . 353
var. longipes Nords.
Basil (rr), Clear (2) (rrr), Corner (rrr), Echo (rr), Henshaw (rrr),
Long (1) (r), Long (2) (s), Loon (sss), Otter (r), Portage (ss), Rosseau
(rr), Rowley (rr), Silver (1) (rrr), Silver (2) (sss), Vernon (ss).
Staurastrum subiongipes n. sp. PI. XI, figs. 15, 16.
Cells of fairly large size, about twice as broad as long (with
processes), moderately constricted, sinus obtuse angled and widely
open; semicells cyathiform, with concave apices, ventral margin
smooth, apical margin with a single emarginate verruca, angles
prolonged into long gradually tapering processes that are smooth
on the lower surface and denticulate on the upper, processes
terminating in three small spines. Vertical view triangular, with
a single emarginate verruca on the body of the cell between the
angles ; processes with a double row of spines ; the angles of one
semicell alternating with those of the other.
Length (without processes) 37-40 /a, (with processes) 55-65 y.
breadth (without processes) 18-20 /a, (with processes) 80-105 y;
breadth isthmus 8.7 /a.
Burned Rock (r), Corner (rrr), Gull (2) (rrr), Joseph (rr), Lake of
Bays (ss), McGowan (rrr), Muskoka (rrr), Pine (r), Rosseau (ss).
Round (2) (rr), Rowley (c), iSt. Mary (r), Silver (2) (r).
The species has a considerable resemblance to S. paradoxum var.
longipes , but differs in the shape of the upper portion of the semi¬
cells and the apical verrucae.
Staurastrum cingulum (W. & G. S. West) comb. nov.
Staurastrum paradoxum var. cingulum W. & G. S. West, Jour. Linn. Soc.
Bot. 35: 548. PI. 18, figs. 6, 7. 1903.
Gull (1) (rr), Hamer (r), Leech (rr), Muskoka (sss), Otter (rrr),
Rosseau (rrr), Rowley (rr), Silver (1) (r), Vernon (rr).
Study of the plankton of the Ontario lakes shows that there
are a number of closely related forms that are characterized by
a ring of small spines at the base of the semicells. These seem to
form a natural group and the only named form, which has been
considered a variety of S. paradoxum , is taken as the type for the
species.
var. tortum var. nov. PI. XII, figs. 1, 2.
Semicells with the processes at the cell corners curved upwards
in the outer half only. Processes of one semicell alternating with
those of the other semicell.
23— S. A. L.
354 Wisconsin Academy of Sciences, Arts, and Letters.
Length (without processes) 25-28 /a, (with processes) 32-50 /a;
breadth (without processes) 16-20 /a, (with processes) 56-75
breadth isthmus 6-8 /a.
Big Dudley (rrr), Gull (1) (rr), Long (2) (rrr), Otter (sss), Round
(2) (rr), iRowley (s).
var. obesran var. nov. PL XII, figs. 3-5.
Semicells stouter and with the processes sub-parallel. Processes
slightly upturned at their apices.
Length (without processes) 35-40 /a, (with processes) 47-60 /a;
breadth (without processes) 25 /a, (with processes) 60-68 /a;
breadth isthmus 10-12 /a.
Burned Rock (r), Echo (rrr), Gull (1) (r), Miuldrew (rrr), Silver
(1) (sss).
Statjrastrum pseudosebaldi Wille
Clear (1) (rrr).
Statjrastrum vestitum Rails
Basil (rrr), Pennsylvania (rrr).
var. subanatinum W. & G. S. West
'Clear (2) (r), Long (2) (rrr).
Staurastrum anatinum Cooke & Wills
Butterfly (sss), Clear (2) (rr), Harbon (ss), Henshaw (rrr), Long (1)
(rrr), Otter (rrr), Thompson (rrr).
var. longibrachiatum W. & G. S. West
Echo (r), Gull (1) (rr), Leech (rrr), Loon (rrr), Muskoka (sss),
Nine Mile (r), Round (2) (rr), Rowley (sss), Silver (1) (sss), Vernon
(sss).
var. curtrnn var. nov. PI. XII, figs. 6, 7.
Body of cell smooth below the apex. Processes short, parallel,
not divergent.
Length 30 /a; breadth (without processes) 25 /a, (with processes)
68-75 n ; breadth isthmus 11 /a.
Big Dudley (rr), Brush (rr), Burned Rock (r).
This variety shows considerable resemblance to the variety
longibrachiatum, but differs in the nature of the processes which
are shorter and not divergent. The length of the cells is also con¬
siderably less than that of other varieties of the species.
Smith — Phytoplankton of the Muskoka Region.
355
Staurastrum protectum var. planctonicum var. nov. PL XII,
figs. 8-12.
Variety with simple verrucae irregularly distributed or in rings
on the processes. Body of semicells smooth between the processes
in front view. Vertical view with two emarginate verrucae be¬
tween the processes. Processes ending in two long divergent
spines.
Length (without processes) 22.5-27.5 /x, (with processes) 45-50
/ 1 ; breadth (with processes) 58-69 /x; breadth isthmus 6.25-8.50 n .
Basil (r), Black (1) (rrr), Burned Rock (rr), Echo (rr), Gull (1) (rr),
Long (1) (rrr), Long (2) (rrr), Loon (r), McGowan (r), Muldrew (rrr).
Nelson (rrr), Otter (rrr), Round (2) (sss), Rowley (rrr), Silver (1)
(rr), Strange (rrr), Vernon (rrr).
The verrucae on the processes are quite variable in distribution
and may be in horizontal rows or unevenly distributed. The
variety is also distinguished by the lack of ornamentation on the
smooth sides of the semicells in front view. An abnormal cell is
shown in figure 12 where instead of terminal spines there is a
branching of the processes that ends in two or three minute spines.
The alga should be compared with S. patens var. planctonicum G.
S. West which has a somewhat similar cell shape but a different
ornamentation.
Staurastrum lacustre sp. nov. PL XII, figs. 13-15.
Cells of medium size, a little longer than broad (with processes),
deeply constricted, sinus broadly open and apex acute; semicells
obversely triangular, apex slightly concave, with two emarginate
verrucae, ventral margin straight, angles prolonged into long
processes with straight, nearly parallel sides and terminating in
two large divergent spines. Processes with 3-4 rings of simple
verrucae, scattered verrucae, or 1-2 verrucae on the apical side
only. Body of semicell smooth, or with a few scattered verrucae
between the processes. Vertical view triangular, sides slightly
concave, with two emarginate verrucae on each side between the
angles, angles ending in processes with straight sides, ornamenta¬
tion of processes as in front view. Terminal spines of processes
lying above one another.
Length (without processes) 26-34 /x, (with processes) 68-85 /x;
breadth (without processes) 22-24 /x, (with processes) 75-100 /x;
breadth isthmus 8.5 /x; length terminal spines 9-12.5 /x.
Big Dudley (s), Gull (1) (rrr), Loon (rrr), Mud (rrr), Nelson (rrr).
Portage (rr), Silver (1) (rrr), Vernon (rr).
356 Wisconsin Academy of Sciences, Arts, and Letters.
This is a very distinctive species that I have found in all three
regions of North America where I have studied the plankton. The
long, smooth processes that terminate in a pair of spines are the
distinctive feature of the species. There is considerable variation
in the ornamentation, and the processes of some cells are prac¬
tically destitute of the small verrucae while other cells have 3-4
rings of verrucae on the processes. The bodies of the semicells
are usually without the simple verrucae on the sides of the cells
but usually have the emarginate verrucae on the apex of the cells.
The form shown in figure 15 with three spines at the ends of
the processes in one semicell is most unusual and has been ob¬
served but twice.
Staurastrum cerastes Lund.
Burned Rock (rrr), Hamer (rrr), Pennsylvania (rrr).
var. triradiatum var. nov. Pl. XII, figs. 16-18.
Cells larger, front view with processes not so sharply incurved
as in the typical form. Semicells with emarginate apical verrucae
and a row of trifid or quadrifid verrucae across the ventral mar¬
gin. Vertical view triangular, with emarginate verrucae on the
bases of and between the processes; central area containing a tri¬
angle of tridenticulate verrucae with three verrucae on each side.
Length 50 p ; breadth (without processes) 27.5 p, (with
processes) 70 p; breadth isthmus 10 p.
Gull (1) (rr), Silver (1) (rr).
\
This variety is distinguished by the laxer curving of the pro¬
cesses at the corners of the semicells and the consequent greater
breadth in proportion to the length. The cells are always tri-
radiate and the verrucae much more prominent than in the typical
form. The alga should also be compared with S. elegantissimum
Johnson.
Staurastrum pentacerum (Wolle) comb. nov. PL XII, figs. 19-22.
Staurastrum Ophiura var. pentacerum Wolle. Bull. Torrey Bot. Club
9: 28. PI. 13, fig. 5. 1882; Desmids of the United States 134. PI. 43, fig.
13; PI. U, fi9- 3. 1884.
Length 28-30 p ; breadth (without processes) 17.5-22.5 p, (with
processes) 81-100 p; breadth isthmus 7.5 p.
Big Dudley (s), Black (2) (r), Brush (rr) , Burned Rock (s), Corner
( rrr ) , Harbon (rr), Henshaw (s), Leech (c) , Little Dudley (s), Long (1)
Smith — Phytoplankton of the Muskoka Region.
357
(r) , Marion (ss), Muldrew (rr), Nine Mile (rrr), Pennsylvania (r),
Round (2) (rrr), Round (1) (r), Rowley (rr), Strange (sss), Thompson
(sss).
In 1882 Wolle described the varieties tetracerum and penta-
cerum of S. Ophiura. These have, as their name implies, four-
and five-radiate semicells which are shaped somewhat like the
semicells of S. Ophiura. In a vertical view of S. Ophiura there
is a coronet of quadrifid papillae with one papilla between each
two adjacent rays of the semicell. The arms have bifid projec¬
tions at the base while the terminal portion is serrate and with
a simple projection. The organism which I believe identical with
Wolle ’s S. ophiura var. pentacerum has an emarginate verruca
between each two adjacent rays while the sides of the rays are
denticulate and at times with a row of denticulations on the top.
I have observed five-rayed specimens of S. Ophiura from New
York, and in these individuals there is the typical marking of the
seven- and eight-rayed individuals. I think, therefore, that
Wolle ’s plant cannot be considered a variety of S. Ophiura but
must be regarded as a distinct species. The number of rays is
generally five, but may be six. The four-rayed specimens have
the same marking and are discussed below in connection with the
variety tetracerum. The specific name pentacerum has been chosen
because the name tetracerum has already been used in another
connection.
forma obesum forma nova. PI. XIII, figs. 1, 2.
Body of semicells stouter and apex more elevated.
Length 38^2 y; breadth (without processes) 25-27.5 y, (with
processes) 105-117 y; breadth isthmus 14 y.
Long (2) (rr), Rowley (c).
var. tetracerum (Wolle) comb. nov. PI. XIII, figs. 3-5.
Staurastrum Ophiura var. tetracerum Wolle. Bull, Torrey Bot. Club 9:
28. PI. IS, fig. Jf. 1882; Desmids of the United States 134. PI. M, figs. 1, 2.
1884. W. & G. S. West, Trans. Linn. Soc. 2 Ser. Bot. 5: 269. PI. 18, fig.
16. 1896.
Length 35-38 y; breadth (without processes) 15-18 y , (with
processes) 85-115 y; breadth isthmus 10-11.5 y.
Basil (r), Big Dudley (sss), Black (1) (sss), Brush (rrr), Butterfly
(s) , Burned Rock (r), Clear (1) (rrr), Clear (2) (rr), Gull (1) (rr),
Gull (2) (r), Hamer (ss), Long (1) (ss), Long (2) (s), Loon (r),
358 Wisconsin Academy of Sciences , Arts , and Letters.
McGowan (rr), Mill (rrr), Mud (rr), Muldrew (s), Muskoka (s), Nel¬
son (rrr), Rosseau (rrr), Round (2) (r), Rowley (s), Silver (1) (r),
Vernon (c), White (rrr).
In view of Wolle ’s rather diagrammatic figure of this variety,
the determination rests upon the figure given by the Wests. The
specimens that I have observed are not so long as the Wests state,
and the breadth of the semicells with the processes may also be
less. I have found the alga in quantity in several lakes without
finding a single five-rayed individual and think, therefore, that it
is a distinct variety. In some collections both four- and five-
rayed individuals are present, but I consider this a mingling of
the widespread type and variety and not a normal variation of
the type.
The twisting of the semicells 30 that the rays of one semicell
are equidistant between the rays of the other semicell is a constant
character and furnishes another distinction between this species
and S. Ophmra.
forma major forma nova.
Processes much longer than in the -foregoing, body of semicells
of the same size as in the variety.
Breadth (with processes) 145-155 /x.
Round (2) (r).
Staurastrum ankyroides Wolle PL XIII, figs. 13-15.
Bull. Torrey Bot. Club 11; 14. PI. U, fi9- 4- 1884; Desmids of the United
States 137. PI. 51, fig. h. 1884.
Length 60-68 /x; breadth (without processes) 20-24 /x, (with
processes) 92-132 /x; breadth isthmus 11-13.5 /x.
Brush (rr), (Burned Rock (r), Butterfly (ss), Echo (rrr), 'Gull (1)
(rrr), Nelson (ss), Nine Mile (rr), Little Dudley (r), Marion (rrr).
Silver (1) (rr).
This species has not been noted since Wolle ’s discovery of it in
1884. I have found an alga in the plankton of Ontario which I
believe is the organism that Wolle described. It is four-rayed,
with the rays of the two semicells alternating with one another. The
ornamentation of the rays is very similar to that of S. Oplniura,
but the ornamentation of the apex is quite different, there being
four heavy semicircular verrucae with four exterior lobes. The
Smith— Phytoplankton of the Muskoka Region.
359
body of the semicell is also narrower than in S. Ophiura. The
shape of the semicells and the sharp incurving of the rays are as
Wolle describes them for S. ankyroides, but I find the ventral
sides of the semicells straight instead of slightly undulate as he
figures. S. ankyroides should be compared with S. pentacerum
var. tetracerum (Wolle) ($. Ophiura var. tetracerum Wolle),
from which it differs in the cell shape, heavier apical verrucae,
lateral spines on rays and incurving of the rays.
Staurastrum Arachne Ralfs,
Big Dudley (rrr), Black (1) (rrr), Brush (sss), Harbon (rrr).
var. curvatum W. & G. S. West
Black (2) (r), Corner (rrr), Echo (sss), Henshaw (s), Otter (r) Pine
(rr), Silver (2) (sss).
Staurastrum Ophiura Lund.
Brush (rrr), Burned Rock (rr"). Clear (1) (rrr), Clear (2) (rrr),
Long (1) (rrr), Muldrew (rrr).
Staurastrum rotula Nords.
Henshaw (rr).
Staurastrum limneticum var. burmense W. & G. S West. PL
XIII, figs. 6-11.
Cells five- or six-radiate.
Zygospores spherical, furnished with long, slender, gradually
tapering spines that are bifurcate at the apex, the bifurcate apical
portion being recurved.
Length cells (without processes) 35-44 /x, (with processes) 47-
55 /a; breadth (without processes) 20-22.5 /x, (with processes)
75-87.5 fi; breadth isthmus 8.5 /x.
Diameter zygospore (without spines) 42-44 fi; (with spines)
85-92 /x; length spines 20-24 /x.
Black (1) (rrr), Black (2) (s), Brush (rrr), Burned Rock (r), Butterfly
(ss), Clear (1) (rrr), Echo (r), Harbon (rrr), Gull (2) (ss), Joseph (ss).
Loon (c), Muldrew (ss), Muskoka (ss), Nelson (s). Otter (r), Portage
(rrr), Rosseau (r), Rowley (r), Round (2) (r), Silver (1) (rrr), Strange
(rr), Vernon (ss).
S. limneticum var. burmense has been collected from but one
lake in Wisconsin, but is widely distributed in the portion of On¬
tario where these investigations were carried out. The Ontario
specimens differ slightly from those the Wests found in Burma
360 Wisconsin Academy of Sciences , Arts , and Letters.
since there is a ring of small conical verrncae on the apex of each
semicell, the number of verrucae corresponding to the number of
rays. The majority of the cells collected are five-armed. A cur¬
ious malformation of a semicell is shown in figure 11.
The alga was forming zygospores in the plankton of Lake
Joseph. This discovery of zygospores in the plankton is most
unusual, and only two cases are on record of zygospore formation
by a plankton Desmid.
forma tetragona forma nova. PI. XIII, fig. 12.
Semicells with four processes, spread of processes greater than
in the typical five- or six-rayed variety.
Breadth (without processes) 20-22.5 n; (with processes) 93-
105 /x.
St. Mary (rr).
Four-rayed cells were found in St. Mary’s Lake but not noted
among the thousands of cells collected from other lakes.
Staurastrum leptocanthum Nords.
Black (1) (aaa), Burned Hock (rrr), Butterfly (s), Clear (rrr), Echo
(rr), Gull (1) (c), Gull (2) (r), Joseph (rrr), Little Dudley (rrr), Long
(2) (r), McGowan (r), Muldrew (rrr), iNelson (s), Otter (rrr), Pennsyl¬
vania (rr), Pine (rrr), Rosseau (rrr), Rowley (rrr), Silver (1) (ss),
Strange (rr).
Staurastrum Arctiscon (Ehr.) Lund.
Basil (rrr), Big Dudley (rrr), Burned Rock (rr), Clear (2) (rrr), Lit¬
tle Dudley (sss), Long (1) (rrr), Loon (rrr), Nelson (rrr), Otter (rrr),
Strange (rrr).
var. GLABRUM W. & G. S. West
Rowley (rrr).
Staurastrum tohopekaligense Wolle
Loon (rrr).
var. trifurcata W. & G. S. West
Gull (1) (rr), Muskoka (rrr), Rowley (rr), Silver (1) (rr), Strange
(rrr).
COSMOCLADIUM Brebisson 1856
Cosmocladium saxonium DeBary
Marion (rrr), Rosseau (rrr), Rowley (rr).
Smith — Phytoplankton of the Muskoka Region.
361
SPHAEROZOSMA Corda 1834
Sphaerozosma granulatum Roy & Biss.
Basil (rr), Round (2) (rrr), Silver (1) (r).
Sphaerozosma excavatum Ralfs.
Mill (rrr), Strange (rrr).
ONYCHONEMA Wallieh 1860
Onychonema laeve var. micracanthum Nords.
Round (2) (rr).
A form with the spines about half the size that Nordstedt gives
was noted in Round Lake in Wood Township (PL XIII, fig. 16).
var. latum W. & G. S. West
Brush (rrr).
SPONDYLOSIUM Brebisson 1844
Spondylosium planum (Wolle) W. & G. S. West
Burned Rock (r), Joseph (rrr), Long (2) (rr), Muskoka (rr), St. Mary
(rrr).
Spondylosium pulchrum (Bailey) Arch.
Gull (1) (rr), Otter (rrr), Pennsylvania (rrr), Silver (1) (rrr).
HYALOTHECA Ehrenberg 1841
Hyalotheca mucosa (Dillwyn) Ehr.
Black (2) (rrr), Brush (rr), Cornell (rrr), Little Dudley (rr), Long
(1) (rrr), Loon (rrr), Marion (rrr), Mill (rr), Mud (rrr), Strange (rrr).
DESMIDIUM C. A. Agardh 1824
Desmidium aequale W. & G. S. West
Burned rock (rrr).
Desmidium cylindricum Grev.
Pennsylvania (rrr).
362 Wisconsin Academy of Sciences, Arts, and Letters.
Desmidium aptogonum Breb.
Hamer (rrr), Little Dudley (rrr).
Desmidium Swartzii C. A. Agardh
Clear (2) (rrr), White (rrr).
Desmidium Baileyi (Ralfs) Nords.
Black (2) (rrr), Marion (rrr), Mill (rrr), Otter (rrr).
GYMNOZYGA Ehrenberg 1841
Gymnozyga moniliformis Ehr.
Big Dudley (rrr), Black (1) (rr), Brush (r), Leech (rrr), Mill (rrr),
Otter (rr). Pine (rrr), Round (2) (ss), White (rrr).
var. gracilescens Nords. Pl. XIII, fig. 17.
Hamer (rr), Nelson (rrr).
SMITH — ALGAE
COCKAYNE BOSTON
TRANS, WIS. ACAD., VOL. XX
PLATE IX.
SMITH — ALGAE
COCKAYNE BO STOTT
TRANS. WIS. ACAD., VOL. XX
SMITH — ALGAE
TRANS. WIS. ACAD., VOL. XX
SMITH — ALGAE
COCKRTHSTE BO STOTT
Smith — Phytoplankton of the Muskoka Region.
363
Figs. 1-3.
Figs. 4-7.
Fig. 8.
Figs. 9-11.
Figs. 12-13.
Figs. 14-18.
Fig. 19.
Fig. 20.
Figs. 1-5.
Figs. 6-11.
Figs. 12-14.
Fig. 1.
Figs. 2-5.
Figs. 6-8.
Figs. 9-13.
Figs. 14-16.
Figs. 17, 18.
Figs. 19, 20.
Figs. 21, 22.
Figs. 23, 24.
Fig. 25.
Figs. 1-i5.
Figs. 6-8.
Fig. 9.
Figs. 10-11.
Figs. 12-13.
Fig. 14.
Figs. 15-16.
Figs. 17-20.
Explanation of Figures
plate VIII
Kirchneriella arcuata (x 530)
Scenedesmus incrassatus var .mononae (x 530).
Closteriopsis longissimum var. tenuissimum (x530).
Tetraedron spiniferum (x 530).
Tetraedron gracile formae (x 530).
Tetraedron quadrilobatum (x 530).
Ankistrodesmus spiralis var. fasiculatus (x 530).
PTiaeococcus planctonicus var. ovalis (x 666).
PLATE IX
Micrasterias radiata var. simplex (x 200).
Micrasterias radiata var. gracillima . (fig. 6 x 400; figs.
7-11 x 200).
Micrasterias Mahahuleshwarensis forma dichotoma (fig.
12 x 400; figs. 13, 14 x 200).
PLATE X
Micrasterias Mahahuleshwarensis forma dichtoma (x 200).
Cosmarium contractum var. papillatum forma minor (x 400).
Xanthidium tetracentrotum var. hexagonum (x 400).
Xanthidium subhastiferum var. Towerii (x 400).
Xanthidium antilopaeum var. limneticum (x 400).
Anthrodesmus constrictus (x 400).
Arthrodesmus quiriferus var. subparallelus (x 400).
Staurastrum muticum (x 400).
Staurastrum cuspidatum var. canadense (x 400).
Staurastrum BreMssonii var. paucispinum (x 400).
PLATE XI
Staurastrum BreMssonii var. paucispinum (x 400).
Staurastrum setigerum var. hrevispinum (x 400).
Staurastrum minnesotense (x 400).
Staurastrum americanum var. longiradiatum (x 400).
Staurastrum americanum var. triradiatum (x 400).
Staurastrum leptocladum var. denticulatum (x 400).
Staurastrum suhlongipes (x 400).
Staurastrum aspinosum var. verrucosum (x 400).
364
Wisconsin Academy of Sciences, Arts , and Letters.
Figs. 1,
Figs. 3-
Figs. 6-
Figs. 8-
Figs. 13-
Figs. 16-
Figs. 19-
Figs. 1,
Figs. 3-
Figs. 6-
Fig. 12.
Figs. 13-
Pig. 16.
Fig. 17.
PLATE XIX
2. Staurastrum cingulum var. tortum (x 400).
-6. Staurastrum cingulum var. obesum (x 400) .
-7. Staurastrum anatinum var. curtum (x 400).
-12. Staurastrum protectum var. planctonicum (x 400).
-15. Staurastrum lacustre (x 400).
-18. Staurastrum cerastes var. triradiatum (x 400).
-2,2. Staurastrum pentacerum (x 400).
PLATE XIIX
2. Staurastrum pentacerum forma obesum (x 400).
- 5. Staurastrum pentacerum var. tetracerum (x 400).
-11. Staurastrum limneticum var. bur mens e (x 400).
Staurastrum limneticum var. burmesse forma tetragenum
(x 400).
-15. Staurastrum ankyroides (x 400) .
Onychonema laeve var. micracanthum forma (x 400).
Gymnozyga moniliformis var. gracilescens (x 400).
TRANS. WIS. ACAD., VOL. XX
SMITH — ALGAE
COCKAYNE BO STOTT
TRANS. WIS. ACAD., VOL. XX
PLATE XIII.
SMITH — ALGAE
COCKAYNE BOSTON
SPECIES OF LENTINUS IN THE REGION OF THE GREAT
LAKES
Edward T. Harper
The species of Lentinus belong to the series of white-spored
agarics which are tough and leathery and which revive on the ap¬
plication of moisture. Their nearest relatives are species of Panus
and Pleurotus. The species of Lentinus have the edges of the
lamellae serrate while in Panus they are entire. The character
represents a difference in the tissue of the two genera. Typical
species of Lentinus have the gills split and lacerate on the edges
as shown in Plate XIV, C, not merely rough from an abundance
of cystidia as is the case in many of the higher agarics. The torn
edges of the gills are due to the fact that the hyphae run cross¬
wise, allowing the gills to split vertically (Plate XIV, A), while
in Panus the hyphae run more nearly parallel to the edges of the
gills and allow them to be torn lengthwise. The growth of the
pileus is more circinate in Panus than in Lentinus. In Panus,
moreover, the fibers of hyphae are looser and the tissue is more
homomorphic than in Lentinus. According to Fayod, this fact
places the species of Lentinus higher in the line of development
than those of typical Panus. In the genus Pleurotus the tissue is
still further developed and the plants do not revive on the appli¬
cation of moisture.
The natural home of the leathery species of agarics is in the
warm regions of the globe. The carpophores dry up and persist
during the drought and revive in the rainy season. Both Europe
and the United States are largely north of their zone of greatest
abundance. Hence the species of these genera in our region are
fewer and less variable than those of the genera whose natural
home is in temperate and cold regions such as Clitocybe, Cortin-
arius, etc.
365
366 Wisconsin Academy of Sciences , Arts , and Letters.
Stevenson reports 12 species of Lentinns from the British Isles,
Constantin and Dufonr recognize 14 species in France, Winter 17
in Germany, and Britzelmayr collected 13 species in Bavaria. The
species lepideus, tigrinus, adhaerens, cochleatus, flabelliformis, and
ur sinus or vulpinus are common to all these lists. TJr sinus and
vulpinus are very closely related, and flabelliformis appears to be¬
long to the cochleatus group. The species in the lists not found
in all the countries appear to be closely related to these principal
species, except perhaps suavissimus or odorus, reported from
France and Germany. There appear, therefore, to be six well
marked groups of species of Lentinus in Europe the types of which
are lepideus , tigrinus , adhaerens , suavissimus , cochleatus , and
ur sinus.
These are also the abundant and well known species in our re¬
gion. Here as in Europe, moreover, minor forms have been given
specific names in each group : spretus , obconicus, and maximus
in the lepideus group, umbilicatus % americanus , and piceinus in
the cochleatus group, etc. These minor forms have not been identi¬
fied with the minor forms in the groups in Europe, though the
close relationship of many, such as umbilicatus and omphalodes, has
been noted.
That the above mentioned are the common and well known spe¬
cies, and hence the proper species to give names to the groups in
Europe and America, is also shown by the number of times they
have been illustrated. According to Volume 21, of Saccardo’s
Sylloge Fungorum , 89 of the over 200 described species of Len¬
tinus have been illustrated. Lentinus lepideus has been illustrated
20 times, L. tigrinus 26 times, L. adhaerens 4 times, L. cochleatus
21 times, and L. ursinus or vulpinus 10 times. Of the other species,
L. flabelliformis , belonging to the cochleatus group, has been illus¬
trated 6 times, L. suffrutescens, a form of the lepideus group, 5
times, L. dunalii, a form of L. tigrinus , and L. degener each 4
times. 12 species have been illustrated 2 or 3 times apiece, and the
remaining 66 only once each, usually by the author of the species.
The only species commonly illustrated are lepideus , tigrinus , coch¬
leatus , and ursinus. Adhaerens, though usually reported in Eu¬
rope, has been figured only 4 times, and suavissimus or odorus ap¬
pears never to have been illustrated.
Peck’s work in New York State is by far the most careful ex¬
amination of a local flora that has been made in this country. In
his summary of New York species of Lentinus (Bulletin 131), he
Harper- — Species of Lentirms in Great Lakes Region. 367
reports nine species, of which lepideus , tigrinus, suavissimus, coch-
leatus , and ursinus are the type forms. He describes two minor
forms which he thinks sufficiently distinct to merit names : spretus,
a form of lepideus ,, which he says European mycologists have prob¬
ably overlooked, and umbilicatus, which he recognizes as closely
related to cochleatus, but which he considers not quite the same as
omphalodes of Europe. Peck also reports Inaematopus, the red¬
stemmed form of suavissimus , and L. sulcatus the same as L. ful-
vidus of Bresadola, a rare species found only once in the state.
Peck found no form of the L. adhaerens group.
No one of the seven or more species of Lentinus described by
Peck from material sent him by correspondents from different parts
of the country has group value. Murrill, in the North American
Flora, does not admit any of them as valid species. They should
not, however, be dumped in as synonyms. They represent forms
worth noting and which future collectors must reckon with. Their
peculiarities should be noted in their proper groups.
As has already appeared, the Lentinus floras of the Northeastern
United States and of Central Europe are very nearly alike. The
details of the variations are given more fully in the description of
the species, but a few general facts are noted here.
1. The species of the Lentinus tigrinus group are so universally
parasitized in this country that the plants appear to have been
stunted. Large, luxuriant forms like those illustrated in Cooke,
lUust., 1138, and Rolland, Atlas, 45, are not reported. The para¬
site has not been found in Europe.
2. The sticky, pulverulent, and resinous species of the Lentinus
adhaerens group are poorly represented in this region, but are com¬
mon in Europe, where leontopodius, pulverulentus, resinaceus, ad¬
haerens, and adhaesus are distinguished.
3. On the other hand, the Lentinus suavissimus group appears
better represented here than in Europe. The plants are quite com¬
mon in Northern Michigan, and besides the lateral stemmed forms
which appear typical of the group we have the larger central
stemmed plants represented by L. odorus or L. jugis. We have
also the red-stemmed form L. haematopus, which is much more
marked than the European L. anisatus.
4. The variation in the Lentinus cochleatus group appears very
similar in both countries. The small central-stemmed forms are
referred to L. omphalodes there and to L. umbilicatus here. The
368 Wisconsin Academy of Sciences, Arts, and Letters .
dimidiate imbricated form is L. flabelliformis there and L. Ameri¬
canos here.
5. The same is true also of the Lentinus lepideus group. An¬
nulate and exannulate forms are found in each country. Obconicus
here resembles suffrutescens there, spretus here may be closely
allied to homotinus there. Many monstrous forms are found in
the group in both countries.
6. In the Lentinus ursinus-vulpinus group, the corrugated
scrupose forms (L. vulpinus), the even tomentose forms (L. ur-
sinus ), and the smaller, smoother forms (L. castoreus) are found in
both countries.
This great similarity between the species of Lentinus in North¬
eastern North America and Central Europe was to be expected
from what is known of the relationship between the floras of the
two regions in general. The cool temperate zone in the northern
hemisphere is divided into an eastern and a western floral region
in both the old world and the new, middle Europe and the China-
Japan regions in the old world and the Great Lakes region and
the Northwestern states in this continent. The floras of Europe
and the Great Lakes region are very closely akin, the China -Japan
region comes next while the flora of the Northwestern states is
farthest removed. This is due to geological history even more
than to climatic conditions. The Lentinus flora of Western Amer¬
ica is not well known, but according to the North American Flora
the Lentinus lepideus group appears to be the only one of the
groups found in the region of the Great Lakes which has repre¬
sentatives on the Pacific coast. It is found also in Asia. L. ti-
grinus extends west to the Rocky Mountains, and the Lentinus
cochleatus group has a single known representative, L. piceinus in
Idaho. The other groups, so far as known, are confined to Europe
and the eastern half of the United States and Canada in the cool
temperate zone.
In the warm temperate zone and in the tropics, species of Len¬
tinus become very abundant, but none of our forms appear to be
found in those regions except tigrinus which extends as far south
as Cuba and is reported also from Australia. Our knowledge of
local floras is, however, too scanty to permit any definite statements
as to the character and distribution of the Lentinus flora of the
globe as a whole.
Fries’ scheme for the division of the genera of tough and woody
fungi into sections is based on the position of the stem or point of
PLATE XIV.
TRANS. WIS. ACAD.. VOL. XX.
HARPER — LENTINUS
COCKAYNE BOSTON
LENTINUS LEPJDEUS FR. A. B. C. D. ANNULATE FORM. E. EXANNULATE FORM.
Harper — Species of Lentinus in Great Lakes Region. 369
attachment of the pilens to the substratum. The divisions are
Mesopus, Pleuropus, Merisma, Apus, Reflexo-resupinati, and Re-
supinati, according as the stem or attachment is central, lateral,
branching, or dorsal. Lentinus and related genera are poorly
adapted to this classification. The tough and pliable substance
readily adapts itself to circumstances, and the position, on the top,
side, or underneath a log, erumpent from a crack or from under
bark, gives a great variety of form to the carpophores of the same
species. Forms belonging to more than one of Fries’ sections are
found in each group. In the Lentinus cochleatus group repre¬
sentatives of all the sections may be found. Murrill uses these
sections for the division of the genus into genera in the North
American Flora , which places plants belonging to the same group
in different genera. For example, L. cochleatus and umbilicatus
are scarcely more than habitat forms but are placed in widely
separated genera. Emphasis must be laid on the fruiting bodies,
the nature of the tissue, and other phylogenetic characters in de¬
termining the groups, and especially must there be large familiarity
with the plants as they grow in different localities and positions.
Synopsis of the Species
I. Stem central. Scales large, appressed, spot-like. The Lentinus
lepideus group.
1. Annulate form _ -Plate XIV, A-C
2. Exannulate form _ Plate XIV, D, E
3. Form with decurrent lamellae and long slender stem
(L. spretus Pk.) _ Plate XV, A
Related species. L. hornotinus Fr.
4. Form with an obconic pileus and long stem (L. obconi-
cus Pk.) _ _ _ _„__Plate XV, B
Related species. L. suffrutescens Fr.
5. Large plants growing on the ground (similar to L. mag-
nus Pk.) _ _ _ Plate XVI
Related species. L. maximus Johnson.
6. Form with finely furfuraceous stem _ Plate XVII, A
7. Form growing from a sclerotium _ _ _ Plate XVII, B
II. Stem central. Scales of the pileus small, fibrous, tufted. The
Lentinus tigrinus Group.
1. Form with the pileus thin, • irregular, splitting on the
edge _ _ _ _ _ _ Plate XVIII
24— S. A. L.
370 Wisconsin Academy of Sciences , Arts , and Letters.
2. Form with the pileus regular, orbicular, not splitting
on the edge _ __ _ _ _ _ _ —Plate XIX
Related species. L. dunalii Fr. L. ravenelii B. & C.
3. Form with the pileus irregular. Stems long and
flexuous _ _ _ _ _ _ _ -Plates XX, XXI, A
4. Form with a thick, short stem _ Plate XXI, B-D
III. Stem central. Pileus not scaly, gummy, resinous, or pul¬
verulent. The Lentinus adhaerens Group.
1. L. adhaesus Britz _ Plate XXI, E
Related species. L. adhaerens Fr. L. resinaceus Fr. L. pulverulentus Fr.
IV. Stem central or lateral, short, reddish brown mycelium at the
base. Odor pleasant. The Lentinus suavissimus
Group.
1. Stem white with little or no red mycelium at the base
(L. suavissimus Fr.) _ _ _ Plate XXII, A-C
2. Stem wholly reddish brown (L. haematopus Berk.)
_ _ _ Plate XXII, D-F
Related species. L. anisatus P. Henn. L. odorus vill. L. jugis Fr.
V. Plants typically irregular, clustered, with the stems concres-
cent. The Lentinus cochleatus Group.
1. Dense clusters of large plants _ —Plate XXIII
Related species. L. cochleatus var. occidentalis Bres.
2. Dimidiate imbricated forms. (L. flabellifo'rmis Fr.)
Plate XXIV
Related species. L. piceinus Pk.
3. Stem central grooved (L. omphalodes Fr.) -Plate XXV
Related species. L. curtisii S. & C. L. micheneri B. & C.
4. Stem central, smooth and even (L. umbilicatus Pk.)
_ _ _ Plate XXVI, A, B
Related species. L. americanus Pk.
5. Form with long, relatively thick, even stems, growing
among moss _ _ _ _ _ _ _ -Plate XXVI, C
VI. Large plants with dimidiate imbricated pilei the Lentinus
ur sinus— vulpinus Group.
1. Pileus ridged and reticulated on the margin, corrugated
behind (L. vulpinus Fr.)_, _ _ — Plate XXVII
2. Pileus even on the margin, coarsely pubescent behind
(L. ursinus Fr.) _ _ _ _ _ _ _ Plate XXVIII
Related species. L. pelliculosus Schw. L. castoreus Fr. L. pectinatus
Schw.
Note. L. sulcatus Berk., which appears to be the same as L. fulvidus
Bres. and L. pholiotoides Ell. & Anders., is a rare plant and it is not
certain to what group it belongs.
TRANS. WIS. ACAD., VOL. XX. PLATE XV.
A.
A. LENTJNUS SPRETUS PK.
B. LENTJNUS OBCONICUS PK.
HARPER— LENTJNUS
cockatoo: boston
Harper — Species of Lentinus in Great Lakes Region. 371
Description of the Groups
The Lentinus lepideus Group
Stem central, pilens with large appressed spot-like scales.
The plants of this group are of medium to large size with lacerate
serrate lamellae usually sinuate at the base. The pileus and stem
are coarsely scaly, and the spores are oblong, 3-6 x 8-12/x. The
sporophores are usually found on wood of coniferous trees. Rail¬
road ties and timbers in docks and bridges are especially liable to
attack. The mycelium is very destructive. The timbers in a dock
at Neebish, Michigan, were scattered all over with the fruiting
bodies showing that the wood was everywhere penetrated by the
mycelium. The plants are usually regular in shape, but many ab¬
normal and monstrous forms occur. ( See Stevenson 2 : 155, Sow.
t. 382; etc.)
Pileus 2-3 inches broad, fleshy, compact, tough, hard when dry,
convex or umbonate to depressed, sometimes irregular or eccentric,
solitary or caespitose, cuticle whitish, pale ochraceous or dark
brown, breaking up into adpressed, spot-like scales which sometimes
become blackish. Flesh tough, white. Odor pleasant. Lamellae
broad, sinuate-decurrent, transversely striate, torn and serrate on
the edges, white. Stem 1-2 inches long, l/2 inch thick, solid, woody,
often flat or pointed at the base where it emerges from the wood,
more or less scaly. Annulus near the apex of the stem, more or
less evident. Spores : ovoid to oblong, 6 x 10/*. Peck 4-6 x 10-
12y2/JL. Stevenson 5 x 11/*. Britzelmayr 3-4 x 20-12/*,.
On structural timber, usually of coniferous trees.
1. Plate XIV, A. B. This is the common form with a short stem
and well marked annulus. The annulus is broad and thick and
covers the gills in the young carpophore. It is sometimes very
broad and marked on the upper side with ragged ridges where the
partial veil was torn from the margins of the gills.
2. Plate XIY, Z), E. The exannulate form. The stem and
margin of the pileus are finely scaly, but there is no evident an¬
nulus. The plants photographed grew on pine timbers in a side¬
walk.
3. Plate XY, A. A form with slightly scaly pileus, more decur¬
rent lamellae, and long, slender stem destitute of an annulus.
Such forms were associated with others on timbers at Neebish,
Michigan. The form is Lentinus spretus Pk. (N. Y. State Mus.
372 Wisconsin Academy of Sciences, Arts , and Letters.
Bull. 105: 24, and 131: 43). The species is reported from Ohio by
Stover. Peck says :
“The species has probably been confused with Lentinus lepideus
from which it may be separated by its more slender habit, thinner
pileus, smaller scales, more decurrent lamellae without a sinus and
especially by its smaller spores. In our specimens there is no
evidence of a veil. ’ ’
In Bulletin 131 Peck adds that the dimensions of the spores of
Lentinus lepideus as given by European authors vary enough to
include this form. Peck gives the measurements as 4-5 x 7%-10/a.
In our collections of L. lepideus the spores are often no larger
than this. Murrill gives a wide range of size to the spores of L.
lepideus , 3-6 x 7-15/a. Peck’s description of L. spretus reads:
“Pileus thin, tough, convex, becoming nearly plane, obtuse or
umbonate, rimose-squamulose, grayish brown or pale alutaceous,
sometimes more highly colored in the center. Flesh white.
Lamellae close, rather narrow, decurrent, whitish, serrate-dentate
on the edges, sometimes lacerate. Stem equal or sometimes nar¬
rowed toward the base, sometimes thickened, solid, subsquamose,
often eccentric, whitish, often brownish toward the base. Spores
white, oblong, 4-5 x 7%-10/a.
“Pileus 2-5 inches broad, stem 1-3 inches long, 3-6 lines thick.
Decaying wood of pine and railroad ties. July to September.”
Note. L. spretus has much in common with L. hornotinus Fr. That
species is said to be villous-pulverulent rather than scaly. The spores
are 3 x 10-12/a. It is illustrated by Britzelmayr 522.3. It is quite, likely,
however, that L. hornotinus belongs to the Lentinus adhaerens group.
4. Plate XV, B. A form with obconic pileus, long decurrent
lamellae and slender, somewhat scaly stem. This is Lentinus ob-
conicus Pk. The photograph was made from plants of the type
collection loaned to me by Dr. Whetstone. They were found in a
lumber yard, but the exact position was not noted. They evi¬
dently grew from some crevice in the timber. The loose strands
of mycelium, composed of hyphae 4/a in diameter, which make up
the sporophore, the club-shaped basidia 4-6 x 20-30/a, and the
spores A- 5 x 8-10/a are all characteristic of L. lepideus . It is an ex¬
treme form of the nature of L. spretus. The species has been found
but once. Peck’s description (Bull. Torrey Bot. Club 33: 215)
reads :
TRAMS. WIS. ACAD., VOL. XX.
HARPER™ LENTINUS
COCKXTNE BOSTON
FORM NEAR LENTINUS MAGNUS PK.
Harper — Species of Lentinus in Great Lakes Region. 373
“Pileus obconic, fleshy, sometimes slightly depressed in the cen¬
ter, whitish with tawny brown squamules in the center. Flesh
white. Lamellae narrow, close, very decurrent, lacerate and den¬
tate on the edge. Stem long, flexuous, solid, whitish at the top,
tawny brown and squamose toward the base. Spores oblong,
4-5 x 8-10 ix.
“Pileus 2.5-6 cm. broad, stem 5-8 cm. long, 6-10 mm. thick.
Caespitose, decaying wood in a lumber yard, Minneapolis, Minn.
“The thick flesh and obconic shape of the pileus with the long,
decurrent lamellae are the prominent distinguishing features of
this species. The flesh of the dried specimens cuts easily. The
scales of the pileus are smaller than in L. lepidem Fr. It is closely
related to L. cyathiformis (Schaefl.) Bres.”
Note. Peck relates the species to the European Panus cyathiformis
which is considered a Lentinus by Bresadola. It is, however, a form of
Lentinus lepideus, and probably the same as the European L. suffrutescens
Fr. illustrated by Schaeffer (t. 248-249), which is considered by Murrill
(N. A. F. 9: 296) a form of L. lepideus.
5. Plate XVI. Large plants with irregular spot-like scales on
the pileus, decurrent lamellae and thick squarrose stems. Len¬
tinus magnus Pk.
The plants grew on the ground by rotten logs in woods at
Frankfort, Michigan. They were very large with large areolate
scales on the pileus. The stem was squarrose with thin, flake-like
scales. They agree with Peck’s L. magnus except that the scales
on the pileus are darker colored. All the characters of the spores,
basidia, and mycelium are those of L. lepideus. They grew on
ground full of humus. The peculiar surface of the pileus and the
thick lamellae emphasized by Peck can be seen in the photograph.
The spores averaged 3 x 7-8/a, a little narrower than the measure¬
ments given by Peck. The species is described (Bull. Torrey Bot.
Club 23: 413, 414) :
“ Pileus thick, hard, convex, slightly depressed in the center,
glabrous, dingy white, the surface cracking into broad areolae or
scales, margin involute. Flesh whitish. Lamellae broad, close,
thick, slightly decurrent, coarsely dentate or lacerate on the edge,
pallid. Stem stout, hard, solid, squamose, slightly thickened at
the base, colored like the pileus. Spores oblong-elliptical, 4 x 7^/*.
Pileus 6 inches or more broad, stem about 4 inches long, 1 inch or
374 Wisconsin Academy of Sciences , Arts , and Letters.
more thick. Gregarious on ground abounding in humus, Mount
San Antonio, California. August.
“This large species was found at an elevation of 1000 ft. It
is well marked by the peculiar areolate and scaly cracking of the
surface of the pileus. The scales of the stem are similar to those
of the pileus. The lamellae are thicker than those of Lentinup
lepideus and the spores are smaller. The scales are concolorous
not spot-like as in that species.”
Note. Lentinus maximus Johnson (Bull. Minn. Acad. Sci. 1878: 338)
has nothing according to the brief description to distinguish it from a
large form of L. lepideus.
6. Plate XVII, A. A form with an even, finely furfuraceous,
branching stem and small scales on the pileus.
The plants are peculiar in the character of the stem and in the
habitat on deciduous trees. The stem is covered with a brownish
furfuraceous coat like that on the pileus of Polyporus brumalis.
The pilei are regular, somewhat umbonate, yellowish tan color,
with small, darker colored, spot-like scales. The general appear¬
ance, spores, and substance were those of Lentinus lepideus. The
plants were collected at Geneseo, Illinois.
7. Plate XVII, B. Sporophore growing from a sclerotium.
The plants grew in sawdust by an old slab pile at Neebish, Michi¬
gan. They were typical Lentinus lepideus , except that the stem
was bulbous and black furfuraceous where it emerged from the
sclerotium. The sclerotium was tough, black, white and finely
fibrous within. It was dying at one end and had a growing point
at the other with some lateral buds. The formation of the sclero¬
tium was probably due to the growth of the plants in the loose
sawdust. Plants with a similar sclerotium but more irregular in
shape have been found growing in very rotten wood at Geneseo,
Illinois.
Species of Lentinus which form sclerotia have long been known.
Pries made a special section in the genus (Scleroma) for them
with L. tuber-regium as the type. These sclerotia vary from the
solid, black-coated masses of hyphae of L. woermanni, described by
Schroeter to simple conglomerations of earth and fungal hyphae.
T. Petch (Ann. Eoy. Bot. Gard. Peradeniya, August, 1915) has
described another form produced by L. similis and L. infundibuli-
formis ,, which consists of masses of rotten wood penetrated and
TRAMS. WIS. ACAD., VOL. XX,
PLATE XV ii
HARPER— LENTiNUS
COCKAYNE BOSTON
TRANS. WIS. ACAD., VOL, XX.
PLATE XVIII
LENTINUS TIGRINUS (BULL.) FR.
HARPER- LENTINUS
COCKrOTlE BOSTON.
Harper — Species of Lentinus in Great Lakes Region. 375
compacted by the persistent hyphae of the fungus. The sclerotium
I have illustrated resembles a rhizome with definite buds from
which the carpophores are produced. I have found the same kind
of sclerotium produced by Polyporus radicatus and will discuss
its nature more fully in an article on that species. Petch leaves
the question open whether different kinds of sclerotia are confined
to different species of fungi or whether the hyphae may assume
different forms under different conditions in the same species.
The Lentinus tigrinus Group
Plants smaller, scales finer and fibrous-tufted, not spot-like, and
spores smaller than in the Lentinus lepideus group. Usually found
on deciduous trees.
Pileus 1-3 inches broad, coriaceous, somewhat fleshy, thin, con¬
vex to plane, umbilicate, easily splitting on the margin, covered
with fibrous-tufted, innate, brown or blackish scales on a whitish
background. Flesh whitish, often turning reddish when bruised.
Lamellae narrow, very crowded, decurrent, finely serrate or erose
on the edges. Stipe 1-3 inches long, 1-5 lines thick, solid, flexuous
or curved, whitish, more or less scaly like the pileus. Spokes ob¬
long, obliquely apiculate 3-4 x 6— 9/x. Basidia club shaped, 4-5 x 20-
25/*.
On stumps of deciduous trees.
The large, luxuriant forms of tigrinus such as are illustrated by
Cooke (Illust. 1138), and Rolland (Atlas 45), are not found in our
region. The reason may be that the carpophores are so universally
parasitized that they have become stunted. There is much variety
in the group and I show four well marked forms in the photographs.
1. Plate XVIII. Rather small plants with the pileus thin and
easily splitting on the margin. White with small dark scales on the
pileus and stem. Stem short, equal, slender, flexuous or curved.
This is the form shown in Cooke’s Handbook, Volume 1, figure 56,
and copied by Winter in Kryptogamen Flora 1 : 488. It appears
to be the usual form in Europe, though the form shown in Plate
XIX is more common with us. The gills of the plant in figure E
are partially parasitized.
2. Plate XIX. Small plants with regular orbicular pileus which
is thick and incurved, not splitting on the margin. It resembles
somewhat L. dunalii as illustrated by Cooke (Illust. 1139 B), which
376 Wisconsin Academy of Sciences, Arts, and Letters.
is considered by Berkeley a form of L. tigrinus ; but that species is
said in the descriptions to have an irregular pileus. A small, thin
form, of this nature appears to be L. ravenelii B. & C., described
from specimens collected in South Carolina.
This is by far the most common form of L. tigrinus in this coun¬
try, and it is usually parasitized, showing the characters appearing
in the photographs in Plate XIX. Morgan based his Lentodium
squamulosum on this form (Jour. Cincinnati Soc. Nat. Hist. 1895
PI. I, fig. 3). Morgan appears never to have seen the form with
normal gills. Dr. Moffatt found the normal and parasitized forms
about equally common in one locality near Chicago. He says in a
letter to me:
(<Lentinus tigrinus was abundant a few years ago in a small wet
swale in the woods at Glen Ellyn, Illinois, 23 miles west of Chi¬
cago. The timber had been cut from this swale and the plants
grew on the decaying oak stumps. The normal form and the
Lentodium were about equally common. One stump might have
dozens of sporophores showing normal gills, while another a couple
of yards away would have only the distorted ones. Several sea¬
sons I looked for transitional forms but never found any, nor did
I ever find both forms on the same stump. When I came upon a
colony of these plants I would try to determine from an inspection
of the top and sides of the cap and stems whether or not gills
would be present, but never was able to do so. Both forms were
identical in size and in appearance.”
All of Moffatt ’s plants were like those illustrated in Plate XIX.
3. Plate XX. Larger plants with irregular pilei and long, flexu-
ous stems. The pileus and stem were coated with coarsely tufted
umber-brown scales. The gills were very narrow and close. They
grew on the sides of stumps at Geneseo and were very different in
appearance from the plants illustrated in Plate XVIII, yet the
whole character, including the spores, shows that they were a form
of the same species. The parasitized condition of this form is
common and is shown in Plate XXI, A. The parasitized forms
were often shapeless masses glued together in clumps. The plants
bore some resemblance to Cooke’s figure (Illust. 1139 A.)
4. Plate XXI, BD. More robust plants with very thick stems,
the pileus thick and not splitting on the margin, covered with
blackish scales. These plants resemble those illustrated by Britzel-
mayr (Lentinus 10). They were usually parasitized. The plant
TRANS. WIS. ACAD., VOL. XX.
PLATE XIX.
HARPER- LENTINUS
COCKAYNE BOSTON
FORM — LENTODIUM SQUAMULOSUM MORG.
TRAMS. WiS. ACAD.., VOL. XX.
PLATE XX
HARPER™ LEMTi NUS
COCKSTKE B0ST01T
LENTIMUS TIGRINUS (BULL.) FR.
Harper — Species of Lentinus in Great Lakes Region. 377
shown in Plate XXI, C has the gills partially parasitized and the
one in figure B wholly so. They grew on stumps of bass wood at
Geneseo, Illinois.
The plants of this group, like those of the Lentinus lepideus
group, show many monstrosities (see Nuov. Giorn. Bot. Ital. 1895,
p. 56). Lyman also in the cultures referred to below produced
many monstrous forms.
The nature of the form with abnormal gills has been much dis¬
cussed since Morgan claimed that it was autonomous and named it
Lent odium squamulosum. The most critical work on the subject
has been done by George R. Lyman in his Studies on Polymorphism
of Hymenomycetes (Proc. Boston Soc. Nat. Hist. 33: 181-193).
Lyman succeeded in growing Lentodium squamulosum from the
spores and produced carpophores with abnormal gills like those
from which the spores were taken. He agreed with Morgan as to
the autonomy of the form for two reasons: first, because it had
never been found growing with Lentinus tigrinu$, nor had inter¬
mediate forms been reported. Second, because of the permanence
of its essential peculiarities in cultures. The first reason does not
hold, for Dr. Moffatt and I have found abundance of normal forms
of several varieties and also forms with the gills partially covered
and partially free. As to the second reason, Lyman does not dis¬
cuss the possibility of the ‘ ‘ veil, ’ ’ as he calls the mass of mycelium
covering the gills, being the hyphae of a parasite. The conidial
spores which he finds on this veil may have been present in the
sowings. To make the evidence complete, cultures should have
been made from normal forms and then from mixtures of the
spores from both forms.
The connection of the normal and the abnormal forms as they
are found about Chicago and Geneseo strongly suggests a para¬
site, especially as several different forms of the fungus have their
gills affected in the same way.
The resemblance of the affected carpophores to species of Agarics
and of Boleti parasitized by Hypomyces is very striking. The
gills or pores are deformed and covered with a very similar cot¬
tony veil. The parasite appears to be a Hyphomycete of a genus
like Verticillium or Spicaria, constituting the conidial stage of a
Hypomyces the ascigerous stage of which does not develop or has
not been discovered. The spores of the plants with normal gills
are more even in size and as large as the larger spores in the
forms with abnormal gills, 3 x 7-9//,. Very many of the much more
378
Wisconsin Academy of Sciences, Arts , and Letters.
abundant spores of the abnormal forms are smaller, 2-3 x 4-6/*, and
not obliquely apiculate at the base. They may be the eonidial
spores of the parasite. As Lyman demonstrated, there are many
such eonidial spores on the hyphae of the veil.
The Lentinus Adhaerens Group
Plants not scaly. The pileus and stem pulverulent and more or
less gummy or resinous.
The group is represented in Europe by L. adhaerens Fr., L. pul-
verulentus Fr. with its variety L. resinaceus Fr., and L. adhaesus
Britz. The plants of these species are of about the same size, aver¬
aging 2 inches high and broad. Their distinguishing feature is
the smooth, more or less pulverulent gummy or resinous surface of
the pileus. In those species in which measurements are reported
the spores are a little smaller than in the Lentinus lepideus group,
3-5 x 6—10/*.
No members of the group have been reported from the United
States as far as I am aware. I have a single plant which is shown
in the photograph in Plate XXI, E. It grew on a log at Frank¬
fort, Michigan. It agrees very closely with L. adhaesus Britz.
Lentinus 8. The pileus is smooth and slightly sticky, bright red¬
dish yellow. The stem is reddish below and whitish above. The
gills are lacerate-serrate, pale reddish yellow, with spores 3-4 x 6-8/*.
Basidia club-shaped, 6 x 30-40/*. Britzelmayr ’s brief description
reads :
“Pileus convex, yellowish red, stipe yellowish above, reddish
brown below, subequal, rooting. Lamellae whitish or pale yellow¬
ish red. Spores 4-5 x 6-8/*. On stumps in autumn. ’ ’
The Lentinus suavissimus Group
The plants of this group are known by their fragrant odor, more
or less evident brown or red mycelium at the base of the short
stem, and the small spores, 2-3 x 5-6/*. The plants are small, whit¬
ish to yellowish in color, often with a yellow margin to the pileus.
They vary from umbilicate to infundibuliform with a short central
stem to dimidiate or with a short lateral stem.
1. Plate XXII, A-D. A form with the stem white. Lentinus
suavissimus Fr. The plants were not rare on sticks of willow or
birch at Neebish, Michigan, in August and September. The odor
was very noticeable.
TRANS. W!S. ACAD., VOL. XX.
PLATE XX f.
HARPER— LENTINUS
COCKATRE BOSTON
LENTINUS TIGRINUS (BULL.) FR.
TRANS. WIS. ACAD., VOL. XX.
PLATE XXII.
HARPER— LENT1NUS
COCKA.THE BOSTON
BERK
Harper — Species of Lentinus in Great Lakes Region. 379
Pileus 1-2 inches broad, thin, tough, convex or centrally de¬
pressed, smooth, even or striatulate on the margin, whitish or yel¬
lowish, often with a yellow margin. Flesh white. Odor strong,
pleasant. Lamellae denticulate, decurrent, wavy, sometimes
anastomosing at the base, colored like the pileus. Stem short, cen¬
tral or eccentric to lateral, whitish, 1-2 lines long and thick.
Spores 2-3 x 5—6/a.
Peck reports the species as rare in New York State, having been
collected only three times, always on willow. In our specimens
there are often traces of reddish mycelium at the base of the stem,
connecting it with the next species.
Note. The form with the central stem and the pileus bordered with
yellow appears to be the same as that called L. odorus Vill. or L. jugis Fr.
in Europe.
2. Plate XXII, C — F. Like the preceding, but with the stems
wholly reddish. Lentinus haematopus Berk. Our plants vary
from central- to lateral-stemmed. They are smaller, more regular,
and neater than L. suavissimus. They have the same fragrant
odor.
Pileus 1 inch or less broad, thin, tough, plane or umbilicate or
centrally depressed, smooth, ochraceous, more or less striate or
ridged on the margin, often lobed. Flesh whitish. Odor pleas¬
ant. Lamellae decurrent, serrate, whitish or coneolorous with the
pileus. Stem short, blood-red, with red or brown mycelium at the
base, smooth or laccate, 1-2 lines long and thick. Spores 2-3 x 5-6/a.
Murrill, 3-4x6-7/a.
On sticks of willow, birch, etc. The species is illustrated by
Boudier (PI. 82).
Note. L. anisatus P. Henn. does not differ except that it is said to be
sometimes subvillous, the stem pallid or reddish brown at the base, the
spores 3 y2 x 6-7/a.
The Lentinus cochleatus Group
The plants of this group grow on stumps, logs, and buried sticks,
and even on ground rich in humus. They are most variable in
size and shape, ranging from large, dense, irregular clusters with
long, grooved, branching, concrescent stems, to single plants with
regular, even, umbilicate pilei and central even stems in one direc-
380 Wisconsin Academy of Sciences, Arts , and Letters.
tion, and to dimidiate-umbricate forms with short lateral stems
or stemless in another direction. The pilei are smooth, tough, and
leathery, with tan colors often with rosy or pink tints. The gills
are serrate and ragged with sphaeroid spores 4-6/x in diameter. A
number of forms have been named as separate species.
1. Plate XXIII, Large caespitose plants with concrescent
branching stems. This is the typical form of well developed plants.
The clusters are found on stumps and logs in damp woods. Those
photographed were collected at Neebish, Michigan.
Pileus 1-3 inches in diameter, thin, tough, flaccid, irregular,
centrally depressed or infundibuliform, brownish tan with rosy or
flesh colored tints, paler when dry, margin even, often lobed.
Flesh whitish. Lamellae broad, deeurrent between the grooves
on the stem, edges lacerate-serrate, whitish or flesh colored. Stem
central or eccentric, caespitose, branching and grown together in
various ways, solid, glabrous, sulcate, concolorous with the pileus,
often darker below. Spores subglobose, 4-6 /a. Single plants are
shown by Liard (fig. 183). Clusters of branching forms with
mamT small plants intermixed are shown in the European illustra¬
tions. Cf. Stevenson, volume 2, page 153.
Note. Canthar ellus multiplex Underwood (Bull. Ton*. Bot. Club 26:
254) may be a form of this species with poorly developed lamellae.
There is in Europe what is considered a large caespitose form of Len-
tinus cochleatus, with entire gills. It is called var. occidentalis Bres.
(see Britzelmayr’s fig. 547.18). Murrill makes it the same as Panus
concavus Berk, from tropical America. The character of the lamellae
should determine whether it is a Lentinus or a Panus.
2. Plate XXIV. Dimidiate, often imbricated forms. The plants
grow on the sides of logs and sticks and on stumps and standing
trunks. They are single or imbricated, and are either stemless or
with short lateral stems. Sometimes a crack in a stump will be
filled with a dense mass of the tough tissue with numerous sessile
pilei on the exposed edge as seen in Plate XXIV, B. These forms
agree so exactly with Lentinus flab elli for mis Fr. that I have re¬
ferred them to that species. The agreement is in shape, size, color,
fimbriate margin of the pileus, nature of the gills, and especially
in the size of the spores. Britzelmayr gives the spores of L. flabel-
liformis as 3-4 x 5-6/a, and those of L. cochleatus as 4-5 x 5-6/a.
They also agree with Britzelmayr ?s illustration (Lentinus 13). On
the other hand, there is no doubt that our plants are forms of
L. cochleatus owing their appearance largely to the place of growth.
TRANS. W1S. ACAD., VOL. XX.
PLATE XXIII.
HARPER- LENTINUS
COCKA.TKE BOSTON
LENTINUS COCHLEATUS FR.
Harper — Species of Lentinus m Great Lakes Region. 381
Pileus/ about 1 inch broad, dimidiate and sessile or with a short
lateral stem, reniform, single or imbricated, tough, pliant, smooth,
even, fimbriate or lacerate and wavy on the margin, fulvous be¬
coming paler on drying. Lamellae broad, subdistant, lacerate,
pallid or whitish. Stem short or none, concolorous or darker than
the pileus, smooth. Spores subglobose or broadly elliptical, 4-6/a
in diameter. On logs, trunks, or buried sticks.
Note 1. Fries reports a still smaller dimidiate form which grows on
stems of herbs.
Note 2. L. piceinus Pk. (N. Y. Mus. Bull. 150: 313), differs from the
plants shown in Plate XXIV only in the fewer and more distant lamellae.
The description given by Peck is as follows:
“Pileus thin, dimidiate, sessile or with a very short stem, broadly con¬
vex or nearly plane, glabrous, pale alutaceous. Lamellae few, distant,
unequal, serrate-dentate, pallid. Stem when present very short. Spores
minute, subglobose, 4-5/a in diameter.
“Pileus 8-12 mm. broad, stem about 2 mm. long. Bark of red spruce
( Picea rubra (Du Roy) Dietr.) A small and rare species. Found but
once.”
Murrill considers it the same as Lentinus cochleatus.
3. Plate XXV. A central-stemmed form with the stem grooved.
Forms like those illustrated in Plate XXV growing singly with
central grooved stems, broadly umbilicate often pervious pilei,
and more or less brown villosity at the base of the stem, are often
found at Neebish on sticks in coniferous woods. They represent
L. ompJialodes Fr.
Pileus 1-2 inches or more broad, thin, tough, deeply umbilicate
and often pervious, repand or lobed, lacerate but not striate on the
margin, pallid to fulvous, drying paler. Lamellae decurrent,
broad, arcuate, pallid or whitish, tom and dentate on the edges.
Stem 1-2 inches long, 1-3 lines thick, tough, flexuous, smooth,
grooved and scrobiculate, concolorous with the pileus. Spores
subglobose 4-6/a. Britzelmayr, 3-4 x 6/a. On buried sticks in coni¬
ferous woods. The European form is found also on the ground.
(Compare under L. umbilicatus below.)
Note 1. L. curtisii Sacc. & Cub. (L. omphalodes B & C.) differs only in
that the gills are said to be entire.
Note 2. L. micheneri B & C is a form of this species with the stem
remarkably spongy-velutine at the base.
4. Plate XXVI, A, B. Central-stemmed forms with even, smooth
stems. The plants are smaller, the pilei more regular, and the
stems more slender, smooth, and even than in the previous form.
382 Wisconsin Academy of Sciences , Arts , and Letters.
They represent L. umbilicatus Pk. (N. Y. State Mns. Rept. 28: 51.
and PI. I, figs. 15-19). The spores are the same as those of
L. cochleatns. Peck’s description in Bulletin 131, page 45, reads:
“Pileus fleshy but thin, tough, glabrous, deeply umbilicate,
hygrophanous, brownish tan color when moist, paler when dry.
Lamellae close, adnate or decurrent, serrate on the edge, whitish.
Stem short, slender, glabrous, nearly even, tough, stuffed or hollow,
central or eccentric, colored like the pileus. Pileus 6-12 lines broad,
stem 8-12 lines long, 1-1.5 lines thick. Ground and decaying wood,
gregarious, July and August. This small species resembles Len-
tinus cochleatus Fr. in texture and color, but it is a much smaller
plant, gregarious in its mode of growth and without furrows on the
stem. It is closely related to Lentinus omphalodes Fr. from which
it has been separated on account of its hollow stem without
elongated furrows or lacunae and its darker color. ’ ’
Note. L. Americanus Pk. (Bull. Torrey Bot. Club 29: 72) differs in the
more marked reddish brown strigose villosity towards the base of the
stem and the rather large spores. It was described from plants collected
in Idaho. It is said to grow most commonly on the ground, but it differs
from the plants shown in Plate XXVI, C in the very short strigose stem.
5. Plate XXVI, C. The plants represent an extreme form of the
small central-stemmed forms of the Lentinus cochleatus group.
The pileus is nearly regular and even on the margin. The stems
are long, relatively thick, even, straight, smooth, hollow, whitish
toward the apex. The plants grew on the ground among moss at
Neebish, Michigan, in September. They might easily have been
taken for a form of Clitocyhe obhata, but the tough substance and
serrate-lacerate gills showed their true relationship. There was
scarcely any villosity at the base of the stems.
The Lentinus ursinus-vulpinus Group
The group consists of large plants with the pilei dimidiate and
reniform, usually imbricated as in Plate XXVII. The gills are
lacerate-toothed. The spores are small, subglobose, 3-4/a in di¬
ameter.
There are two principal forms in our region, L. ursinus and L. vul-
pinus, the former has the pilei even on the margin and dark fuscous-
tomentose behind, and the latter has the pilei ridged and reticulate
on the margin and scrupose behind. They are considered varieties
by some writers, since intermediate forms occur.
Harper — Species of Lentirms in Great Lakes Region. 383
1. Plate XXVII. Lentinus vulpinus (Sow.) Fr. Plants sessile,
imbricate multiplex. Pilei sessile, reniform, conchate, dimidiate,
connate behind, surface corrugated and longitudinally striate,
more or less reticulate or broken up into fibrous tufts on the margin,
alutaceous or whitish, margin extending beyond the gills and in¬
curved, often discolored. Flesh thick, tough, white. Lamellae
broad, crowded, dentate-lacerate on the edges, white. Spores sub-
globose, smooth, 3-4jtt. Stumps and logs, especially of elm.
2. Plate XXVIII. Lentinus ursinus Fr. Like L. vulpinus in
shape and character and habit of growth, but the pilei are smooth
on the thin margin and become coarsely tomentose behind with
reddish brown hairs. The whole plant is darker colored in our
specimens than L. vulpinus. The spores are exactly the same in
both species.
Note 1. L. pelliculosus Schw., found on dead trunks in the Carolinas
and in Ohio, is a small form of L. ursinus. The pileus is said to be fus¬
cous-cervine, thickly strigose like the skin of an animal, with the margin
naked and involute, the lamellae broad and lacerate.
Note 2. L. pectinatus Schw. is a plant of the same character and be¬
longs to the same group if we may judge by the brief description. It is
said to resemble L. reniformis.
Note 3. L. chama Bose, is* a large white form said to have a smooth
pileus. The pilei are said to be 15 cm. broad. It was found on oak
stumps in the Carolinas.
Note 4. L. sulcatus Berk, is a rare plant which grows from crevices
of dry wood and old fence rails and should be found in our territory. It
is described by Peck as follows:
“Pileus fleshy, thin, tough, conic, becoming hemispheric or convex,
cracking on the surface and forming irregular scales, virgate toward the
margin, sulcate on the margin, reddish or tawny, often darker in the
center. Lamellae subdistant, rounded behind or emarginate, slightly
adnexed, obscurely dentate on the edge, white or pallid. Stem central,
short, solid, sometimes narrowed downward, pruinose or slightly fur-
furaceous, white or pallid. Spores oblong, 6-7 y2 x 13-18^.
“Pileus 8-12 lines long, stem 6-10 lines long, 1.5-3 lines thick. Crevices
of dry wood and old fence rails.”
The group of this species is uncertain. The size of the spores places
it in the Lentinus lepideus group, but it is found on deciduous trees.
384 Wisconsin Academy of Sciences, Arts, and Letters.
Description of Plates
Plate XIV. Lentinus lepideus Fr. A. Part of a plant showing the nature
of the edge of the gills in the genus Lentinus. B and D. Surface of
pilei showing the spot-like appressed scales. C. Plant with an an¬
nulus. E. Exannulate plant.
Plate XV. A. Lentinus spretus Pk. A form of L. lepideus. The illustra¬
tion shows the long, slender stem. B. Lentinus obconicus Pk. Also a
form of L. lepideus. The photograph is from plants of the type col¬
lection.
Plate XVI. A large form of Lentinus lepideus. The photograph is re¬
duced one-third.
Plate XVII. A. Plant with nearly smooth, scaly-spotted pileus and
branching dark furfuraceous stem. B. Plants growing from a sclero-
tium. The sclerotium is like a rhizome with the dying end below and
the growing point above.
Plate XVIII. Lentinus tigrinus (Bull.) Fr. A and F show the surface
of the pileus and the gill surface. The plants were preserved in
alcohol, and some of the gills adhere to each other. They were not
parasitized. B and E are from another collection. The gills in E
are partially normal and partially overgrown by hyphae belonging to
the parasite. C shows an enlarged portion of the gills of another
plant which are partially free and partially covered with the parasite.
The photograph was taken from a dried specimen. In the fresh
plant, in the unparasitized part, the gills were straight and normal.
C, D. Still other plants of the same character showing the tendency
of the pileus to split on the margin. The gills are free from parasitfc
hyphae.
Plate XIX. Another form of Lentinus tigrinus. A, B. Gill surface and
surface of the pileus of non-parasitized plants. D-G. The same with
the gills covered with the parasite. C. Two plants grown together
and with the gills parasitized.
Plate XX. A form of Lentinus tigrinus with thick, tufted scales on both
the pileus and stem. The stems long and flexuous.
Plate XXI. A. The form of Lentinus tigrinus shown in Plate XX, but
with the gills covered by the parasite. B-D. Form of the same
species with a thick heavy stem. B is fully parasitized, C only
slightly so- E. Plant very similar to Lentinus adhaesus Britz.
Plate XXII. A, B, C. Various views of Lentinus suavissimus Fr. A has
the form of L. jugis Fr. or L. odorus Vill. D, E, F. Views of Lentinus
haematopus Berk. In D the short red stem is lateral.
Plate XXIII. Lentinus cochleatus Fr. Cluster of large, irregular plants
showing the smooth, even pilei, toothed gills, and grooved stems.
TRANS. WiS. ACAD., VOL. XX.
PLATE XXIV.
LENTINUS FLABELLIFORMIS FR.
HARPER— LENTI NUS
COCKAYNE BOSTON
TRANS. W1S. ACAD., VOL. XX
PLATE XXV
LENTJNUS OMPHALODES FR.
HARPER- LENTINUS
TRANS. W8S. ACAD.. VOL. XX.
PLATE XXVI
A. B. LENTINUS UM88LICATUS PK.
C. FORM WSTH LONG, THICK STEM
HARPER- LENTSNUS
COCKA.TNE BOSTON*
TRANS. WiS. ACAD., VOL. XX.
PLATE XXVII.
LENTSNUS VULP1NUS FR.
HARPER— LEMTiNUS
COCKAYHE BOSTON
TRANS. WIS. ACAD., VOL. XX
PLATE XXVIII
HARPER— LENTINUS
COCKfi-TNE BOSTON
LENTINUS URSINUS FR.
Harper — Species of Lentirms m Great Lakes Region. 385
Plate XXIV. Various views of Lentinus flabelliformis. A-E show the
single pilei with smooth pilei, short lateral stems, and serrate gills.
F shows a cluster of imbricated pilei. G shows a mass of fleshy
fungus growing from a crack in a stump, with the surface covered
with narrow imbricated pilei.
Plate XXV. Various views of Lentinus omphalodes Fr. with smooth
nearly central-stemmed pilei, serrate gills, and the smooth stems more
or less grooved.
Plate XXVI. A, B. Lentinus umbilicatus Pk., showing smooth, orbicular
pilei deeply umbilicate, smooth, even stems with more or less tawny
villous hair at the base. C. Plants with smoother, thicker, and longer
stems growing in moss.
Plate XXVII. Lentinus vulpinus Fr. Cluster of large imbricated pilei
showing the grooved reticulated margins of the pilei and the serrate
gills.
Plate XXVIII. Lentinus ursinus Fr., showing the thin, even margin
and the thick, hairy base of the surface of a pileus.
25— S. A. L.
CYTOLOGICAL STUDIES OF THE LOWER BASIDIOMY-
CETES
I. DACRYMYCES
E. M. Gilbert
The discovery of the fusion nucleus in the hasidium of the
higher Basidiomyeetes, and the further fact that the cells from
which the basidia arise are binucleated, have made it of vital
interest to discover, if possible, the origin of this binucleated con¬
dition.
With this end in view, a morphological and cytological study
has been made of various species of Tremella, Exidia, Dacry-
myces, Calocera, and Auricularia, in an endeavor to work out
the life history from spore to spore, to discover, if possible, at
what stage in the development of the fungus the cells first con¬
tain two nuclei, the manner in which the two nuclei become asso¬
ciated in the cell, and the behavior of the nuclei in subsequent
development.
The work was begun in 1906. In 1911 a progress report (16)
on some of the studies was made. A report, on the nuclear phe¬
nomena as observed in Auricularia, was made at a meeting of
the Wisconsin Academy of Letters, Arts, and Sciences in April,
1911, and the following year a report on the nuclear phenomena
as observed in Tremella and Exidia was given before the so¬
ciety at its annual meeting. The observations made on the vari¬
ous forms have been summarized in a thesis presented in 1914
for the degree of Doctor of Philosophy at the University of Wis¬
consin.
As the writer had not been able to locate definitely the origin
of the binucleate condition, it did not seem that the material was
worthy of publication; but papers which have appeared in re¬
cent years make it seemingly worth while to publish any dis-
387
388 Wisconsin Academy of Sciences , Arts , and Letters.
covery which may finally lead to a solution of this interesting
problem.
Material for this paper has been collected at various places1
during the past ten years and fixed, in the field, in a number of
fixing solutions. The best results have been obtained from the
material killed in Flemming’s weak and Flemming’s medium
solution. Merkel’s solution has also given very satisfactory re¬
sults. Various combinations of stains have been used, but the
most excellent results have been obtained by the use of the triple
stain of Flemming and by the use of Heidenhain’s iron alum-
haematoxylin, the latter giving the best results when studying
the spindle figures. Spores were caught on sterile slides and
some of these were immediately fixed in Flemming’s weak fixa¬
tive and then stippled on slides having a thin layer of egg-albu¬
men fixative. These were allowed to dry and then washed, de¬
hydrated, bleached, and stained in the usual manner.
Some spores were put into tapwater or wood decoction and
fixed from time to time in an effort to get various stages in ger¬
mination, while other spores were plated out in Pe tri-dish and
test-tube cultures in order to secure hyphae in various stages of
development. No culture medium has, however, been found
which will give a continued growth of the mycelium of any of
the species of Dacrymyces studied.
The earlier studies on the cytology of the higher basidiomycetes
have been so thoroughly reviewed by Fries (13, 14), Levine (26,
27), Fitzpatrick (12), and more recently in a most exceptional
thesis by Bensaude (4), that no effort will be made in this brief
paper to refer to any other than more recent studies.2
Studies of Nuclear Phenomena
Spores
The spores of Dacrymyces when thrown from the sterigmata
are one-celled and contain a single nucleus (PL XXIX, fig. 1).
This is the condition typically found in the higher Basidiomy-
1 Most of the collections have been made in the vicinity of Madison ; others
have been made at Superior, Brule, Blue Mounds, and Lake Mills, Wisconsin.
During the spring of 1915 many collections were made in the White Mountains
and in the vicinity of Cambridge, Mass.
2 A fuller discussion of the literature will be found in two papers which are in
preparation, one on the cytology of Auricularia, and the other on the cytology of
Tremella and Exidia species.
Gilbert — Cytological Studies of Lower Basidiomycetes 389
cetes and these spores are quite comparable in this respect with
the spores described by Fitzpatrick (11, 12) and Bensande (4).
The spores may remain in this unicellular condition, but usually,
if moisture be present, will proceed to the formation of eight-
celled spores (figs. 2-8).
It is very possible to find all stages in this process in a single
mount, and the details are readily observable. The primary
nucleus divides, the two nuclei separate, and a cross wall is soon
laid down, giving rise to a two-celled spore. Repeated nuclear
division with consequent formation of cross walls, during a
period varying from four to eight hours, will then result in the
final eight-celled spore (figs. 7, 8).
I have spoken of the one-celled spore as mature, but it is ques¬
tionable which of the described stages is that of a mature spore,
as spores in any of these stages will remain viable for several
days when exposed to the ordinary laboratory conditions.
The nucleus of the spore contains a single nucleole and an
abundance of heavily staining chromatin. In division the typi¬
cal fungal spindle is formed with its very plastic chromosomes.
Because of this plasticity it is not an easy matter to make accur¬
ate counts of the chromosomes, but from the studies made the
chromosomes seem to number three or four as noted on the spin¬
dle. The nucleole is soon crowded to one side and very quickly
breaks down. Centrosomes are present in connection with all
the nuclear divisions within the spore. Astral rays are at times
to be noted (fig. 3), but they are so few in number and so deli¬
cate that they are easily overlooked.
Mycelium
The spores as a general rule do not produce hyphae until the
eight-celled stage has been reached, but many instances of the
production of hyphae by four-celled spores have been noted.
Text figure 1 shows such a four-celled spore with a portion of
the mycelium developed from it. This was taken from a wood
decoction where there was continued growth of hyphae for thir¬
teen days, and the spore with mycelium was taken out at the
end of twenty-one days, killed, and stained. The cells of the
various hyphae were found to have one and only one nucleus
each. There were very few anastomoses, but seemingly no
migration of nuclei during this entire period.
390 Wisconsin Academy of Sciences, Arts, and Letters.
In figure 9 (PL XXIX) is shown the typical method of germi¬
nation. Each cell produces one or more hyphae, each cell of
which will have a single nucleus. Very rarely has a cell been
found, in any of the cultures, which has two nuclei. If, on the
other hand, a study is made of sections of very young fruiting
bodies as found in nature, it is soon observable that a great
number of anastomoses take place, and the greater number of
cells are typically binucleated. It is to be noted, however, that
many of the cells found in the neighborhood of the spore from
which the fruiting body developed, and which may occasionally
be found in sections, are uninucleated, and that at a short distance
from the spore they are found to contain two nuclei. There is
no question but that many of these binucleated cells have arisen
as a result of the fusion of adjacent hyphal cells, but many direct
descendants of uninucleate cells are also found to contain two
nuclei.
Fig. 1. — Germinating spore of Dacrymyces, showing cells having
one nucleus.
It has not been found possible, as a result of these studies on
Dacrymyces, to discover anything quite comparable to the pri¬
mary and secondary mycelia described by Falck (9, 10) and
Bensaude (3, 4), and at no time have clamp connections been
observed in these forms. The hyphae from different parts of an
individual fruiting body vary greatly in diameter, and very
often the cells which develop into basidia are much smaller in
diameter than adjacent cells. Figures 10 and 11 show two bas¬
idia which are derived from such hyphae. The cells immediately
Gilbert— Cyiological Studies of Lower Basidiomyeetes 391
beneath the hymenial layer always contain two nuclei, and it is
impossible at first to decide which cells are to function as basidia
and which are to develop into paraphyses.
The Basidium
The young basidium may at first be smaller than adjacent
cells, but this condition persists for only a very short time. The
cell very rapidly elongates, and as the two nuclei fuse (fig. 12),
it thickens and becomes two to three times the diameter of the
paraphyses surrounding it. As soon as the nuclei have fused,
often before the nucleoles have united (fig. 13), there is a rapid
increase in size of the nucleus. No true resting period ensues,
but a definite synapsis stage (figs. 14, 15) is soon observable.
The chromatin material becomes distinctly massed at one end
of the nucleus, and at this time it is possible to make out a dis¬
tinctly double nature of the chromatin elements. A heavily
staining body is to be found on the nuclear membrane usually at
the point where the chromatin is at its densest. Because of the
association of this body (figs. 15, 16) with the chromatin, and
because of its similarity to the centrosome which soon makes its
appearance, this body has been taken to be the structure from
which the centrosomes arise. It has not been possible to follow
the method of development throughout, but this body is very
evident during later synapsis and persists during the subsequent
stages. The spindle is almost completely formed before the en¬
tire disappearance of the nuclear membrane, and the chromatin
is massing at the center without, however, being in the shape of
distinct chromosomes. The nucleole is pushed out beyond the
spindle fibres and disappears in the cytoplasm with the break¬
ing down of the nuclear membrane (figs 17, 19).
The chromosomes, because of their plastic nature, are not eas¬
ily counted, but four seems to be the number in the species
studied (figs. 18-20).
The daughter nuclei have a very short resting stage, and their
division is quite similar to the preceding division (figs. 21, 22).
The four nuclei now become arranged in a row in the rapidly
growing basidium (fig. 23). The nuclei, however, soon begin
to migrate slowly toward the apex of the basidium where the
sterigmata are being formed (fig. 24). During this migration
they decrease greatly in size and change in shape, becoming dis-
392 Wisconsin Academy of Sciences, Arts, and Letters.
tinctly ovoid with the broader end in advance. The nucleole very
soon takes up its position at the narrower end of the nucleus
(figs. 24, 25). Two of the nuclei are soon observed to lag be¬
hind and plainly indicate by their staining reaction that they
are degenerating (figs. 25, 26). In sections containing basidia
which have already produced spores, it is possible to find traces
of the degenerating nuclei, thus precluding the possibility of a
second set of spores being produced as has been suggested by
some observers.
As the sterigma elongates it gradually decreases in diameter,
and the nucleus at the same time elongates and becomes finally
almost rod-shaped as it passes into the developing spore. There
is at no time any indication of a centrosome or comparable body
aiding in the formation of the young spore as described by Le¬
vine (26, 27), Fries (13, 14), and others for the higher Basidio-
mycetes, but the process is very comparable to that described
by Fitzpatrick (11, 12). As the spore grows to maturity the
nucleus assumes a central position and increases greatly in size.
When fully developed, the spore is seemingly separated from the
sterigma with some force, as is shown by a spore print which
extends to some distance beyond the boundary of the fruiting
body.
Summary
1. These studies have been based upon material from three
distinct species of Dacrymyces.
2. The spores of Dacrymyces when thrown from the sterig-
mata are one-celled and contain a single nucleus each.
3. Under proper conditions the nucleus divides and septa are
formed, producing an eight-celled spore.
4. The spore often produces hyphae in the four-celled stage.
5. The division of the spore nuclei is quite like nuclear division
in other Basidiomycetes.
6. Four chromosomes are typically found, the nucleole is ex¬
truded into the cytoplasm, and centrosomes are present.
7. In cultures, binucleated cells are rarely found, anastomoses
are present but no nuclear migration has been noted. No clamp
connections have been observed.
Gilbert — Cytological Studies of Lower Basidiomycetes 393
8. Anastomoses are numerous in the developing fruiting body
and probably give rise to binueleate cells. Other cells un¬
doubtedly become binueleate due to the division of the single
nucleus without consequent cell division.
9. All cells directly beneath the hymenial layer are binueleate.
10. It has not been possible to count the chromosomes in hy-
phal nuclei.
11. The young basidia are not to be distinguished from the
paraphyses.
12. The nuclei of the young basidium fuse as the basidium be¬
gins to increase in size.
13. There is no distinct resting stage after fusion.
14. The synapsis stage shows the chromatin elements to be
double.
15. A dense, dark-staining body is always present on the nu¬
clear membrane during late synapsis. This is believed to give
rise to the centrosomes.
16. Four chromosomes are found as the typical number pass¬
ing to each pole. The first division is heterotypic.
17. Two sterigmata are formed each bearing a single uninu-
cleated spore.
18. The two remaining nuclei within the basidium soon degen¬
erate.
Bibliography
1. Baum, E. Ueber Zelltheilung in Pilzhyphen. Inaug. Diss. Basel,
1900.
2. Beauverie, M. J. lotude cytologique sur le Merulius lacrymans.
Rev. Gen. Bot. 21: 449-469. 1909.
3. Bensaude, M. Sur la sexualite chez les Champignons Basidio¬
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4. — — . Recherches sur le cycle evolutif et la sexualite ches les
Basidiomycetes. These. Paris, 1918.
5. Brefeld, O. Botanische Untersuchungen iiber Schimmelpilze.
Heft 3, Basidiomyceten I. Leipzig, 1877.
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6. - . Untersuchungen aus dem Gesammtgebiete der Mykologie.
Heft 7, Basidiomyceten II. .Leipzig, 1888.
7. Dangeard, P. A. Memoire sur la reproduction sexuelle des Basid-
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8. - . La reproduction sexuelle des Champignons, etude critique.
Botaniste 7: 89-130. 1900.
9. Falck, T. Die Lenzites-Faule des Coniferenholzes. Hausschwamm-
forschungen 3. Jena, 19 09.
10. - . Die Merulius-Faule des Bauholzes. Hausschwammfor-
schungen 6. Jena, 1912.
11. Fitzpatrick, H. M. The life history and parasitism of Eocronartium
muscicola. Phytopath. 8: 197-218. 1918.
12. — — . The cytology of Eocronartium muscicola. Amer. Jour. Bot.
5: 397-499. 1918.
13. Fries, R. E. Zur Kenntnis der Cytologie von Hygrophorus conicus.
Svensk. Bot. Tidskr. 5: 241-251. 1911.
14. - . Ueber die cytologischen Verhaltnisse bei der Sporenbild-
ung von Nidularia. Zeitschr. Bot. 3: 145-165. 1911.
15. Gilbert, E. M. Studies on the Tremellineae of Wisconsin. Trans.
Wis. Acad. Sci., Arts, and Lett. 16: 1137-1170. 1910.
16. — — . Nuclear phenomena in the basidium and in the germinat¬
ing spores of Dacrymyces and Tremella. Science n. ser. 33:
264. 1911.
17. Guilliermond, A. [Les progres de la cytologie des Champignons.
Prog. Rei Bot. 4: 389-542. 1911.
18. Harper, R. A. Binucleate cells in certain Hymenomycetes. Bot.
Gaz. 33: 1-25. 1902.
19. Hoffman, H. Die Pollinarien und Spermatien von Agaricus. Bot.
Zeit. 14: 137-148; 153-163. 1856.
20. Hone, D. S. (Minn. Bot. Studies. Vol. 4, part 1. 1909.
21. Istvanffi, G. von. Ueber die Rolle der Zellkerne bei der Entwick-
lung der Pilze. Ber. Deutsch. Bot. Ges. 13: 452-467. 1895.
22. Juel, H. O. Die Kerntheilung in den Basidien und die Phylogenie
der Basidiomyceten. Jahrb. Wiss. Bot. 32: 361-368. 1898.
23. Kniep, R. Ueber das Auftreten von Basidien im einkernigen
Mycel von Armillaria mellea. Zeitschr. Bot. 3: 527—553.
1911.
Gilbert— Cytological Studies of Lower Basidiomycetes 395
24. - . Beitrage zur Kenntnis der Hymenomyceten I, II. Zeitschr.
Bot. 5: 593-637. 1913.
25. — - . Beitrage zur Kenntnis der Hymenomyceten III. Zeitschr.
Bot. 7: 369-398. 1915.
26. Levine, M. Studies in the cytology of the Hymenomycetes espe¬
cially the Boleti. Bull. Torrey Bot. Club 40: 137-181. 19.13.
27. - . Review of Kniep’s “Beitrage zur Kenntnis des Hymeno¬
myceten III.” Mycologia 8: 184-186. 1916.
28. Maire, R. Sur la cytologie des Hymenomycetes. Compt. Rend.
Acad. Sci. Paris 131: 121-124. 1900.
29. - — . Recherches cytologiques et taxonomiques sur les Basidio¬
mycetes. Bull. Soc. Mycol. France 18: 1-209. 1902.
30. Nichols, S. P. The nature and origin of the binucleated cells in
some Basidiomycetes. Trans. Wis. Acad. Sci., Arts, and Lett.
15: 30-70. 1905.
31. Rosen, F. Studien liber die Kerne und die Membranbildung bei
Myxomyceten und Pilze. Cohn’s Beit. Biol. Pflanzen 6: 237-
266. 1892.
32. Rosenvinge, L. K. Sur les noyaux des Hymenomycetes. Ann.
Sci. Nat. Bot. VII, 3: 75-93. 1886.
33. Ruhland, W. Zur Kenntnis der intracellularen Karyogamie bei
den Basidiomyceten. Bot. Zeit. 59: 187-206. 1901.
34. Sappin-Trouffy, P. Recherches mycologiques. Botaniste 5: 44-
58. 1896.
35. Tulasne, L. R. Observations sur l’organisation des Tremellinees.
Ann. Sci. Nat. Bot. 19: 193-251. 1853.
3 6. Wager, H. On the nuclei of the Hymenomycetes. Annals Bot.
6: 146-148.
37. - . On nuclear division in the Hymenomycetes. Annals Bot.
7: 489-514. 1893.
38. — . On the presence of centrospheres in fungi. Annals Bot.
8: 321-334. 1894.
39. - — — . The sexuality of the fungi. Annals. Bot. 13: 575-597.
1899.
396 Wisconsin Academy of Sciences , Arts , and Letters .
Explanation of Figures
All the figures were made with the aid of a camera lucida. A Zeiss
2.30 mm. aprochomatic oilimmersion objective and a no. 12 compensating
ocular were used f • figures 13-20. A Leitz 1/16 oil immersion and a
no. 4 ocular were used for the other figures.
Fig. 8. Division of nuclei preceding the development of first germ
tubes.
Fig. 9. A spore with several germ tubes. Some of the tubes contain
Figs. 14, 15. Later spireme stage. The very characteristic massing of
Figs. 18-20. Later stages in primary division, showing typical spindles
TRANS. WIS. ACAD., VOL. XX
PLATE XXIX.
GILBERT — CYTOLOGY OF DACRYM YCES
COCKAYNE BOSTON
Gilbert — Cytological Studies of Lower Basidiomycetes 397
Fig. 25.
Fig. 26.
Fig. 27.
Fig. 28.
Fig. 29.
Basidium with one nucleus beneath the elongating sterigma
preparatory to the passage through the sterigma. The
remaining nuclei show a further decrease in size.
Basidium with two nuclei at the base of the sterigmata, the
other two nuclei distinctly degenerating.
Portion of mycelium at base of fruiting body, showing a
uninucleate cell and a binucleate cell.
Typical binucleate cell in region immediately beneath hy-
menial layer.
Cell in growing portion of fruiting body containing two
nuclei, one of which is dividing preparatory to the mi¬
gration of one of the daughter nuclei which is being
formed.
Department of Botany,
University of Wisconsin
NOTES ON PARASITIC FUNGI IN WISCONSIN— VII.
J. J. Davis
The season of 1918 was not favorable for the parasitic fungus
flora, apparently because of lack of precipitation. Even forms
that occur on marsh or swamp plants seem to require rain for
full development.
Plasmopara humuli Miyabe & Takahashi was found on Humulus
Lupulus in 1918 at Bruce. This adds to the probability that it
is indigenous in Wisconsin.
[Also found in small quantity at Little Suamico on the west
shore of Green Bay in September 1921.]
Of a collection on Smilax hispida referred to Stagonospora
smilacis (E. & M.) Sacc. it was noted that in some of the pycnidia
the sporules were 15-20 x6-7y, somewhat fusoid, hyaline, con¬
tinuous, while in other pycnidia they were elliptical to ovate, sub-
fuligineous, 10-13 x 6-7^ and many of them uniseptate.
In “ Notes” IV (p. 68) reference was made to a Septoria on
leaves of wheat which was referred to S, glumarum Pass. In a
collection on Triticum vulgare (cult.) made at Bailey’s Harbor,
July 28, 1918, the pycnidia are located in the somewhat paler
central portion of more or less elongate brown spots which are
surrounded by a yellow discoloration of the leaf. The somewhat
depressed, ostiolate pycnidia are 90-130/x in diameter, the sporules
15-26 x 3 p, appearing continuous but showing 1-3 divisions of the
endoplasm when stained. Septoria fusispora Died, may be the
same fungus (Kryptogamenflora der Mark Brandenburg 9: 467).
The delicate spore tendrils have a pink tinge.
In the provisional list two species of Septoria were given as
occurring on Viola in Wisconsin. In the Enumeration and De¬
scription of North American Septorias Dr. George Martin gave
399
400 Wisconsin Academy of Sciences , Arts, and Letters.
the spore dimensions of S. violae West, as 40-50x13/2/* (Jour.
My col. 3: 73). The dimensions given by Diedicke in the Krypto-
gamenflora der Mark Brandenburg are 17-20 x 1/*. In Wisconsin
specimens the sporules are 20-36x1/*. The small, circular arid
spots are sometimes surrounded by pale brown discoloration, and
the black perithecia are sometimes borne on such discolored peri¬
pheral areas. Septoria hyalina Ell. & Evht. was described as
having sporules 20-40 x 1 J4/* and black pycnidia on spots having
a dark purple border as contrasted with yellowish brown pycnidia
on spots having a reddish brown border in S. violae West. At
present I am referring all specimens on Yiola to S. violae West.,
considering it to be a variable species.
Septoria intermedia Ell. & Evht., a species founded on a Wis¬
consin specimen, was included in the provisional list. In “Notes”
III (pp. 253-254) it was stated that no similar material had been
collected since, and the opinion was expressed that it is a short-
spored form of S. solidaginicola Pk. In July, 1918, there was
collected at Wild Rose on a large radical leaf of Solidago a speci¬
men in which the sporules are 14-23/* long by 1/* or less in thick¬
ness. The spots are more rounded and somewhat longer than in
the type of S. intermedia Ell. & Evht., and the pycnidial walls
are strongly thickened and blackened about the ostiole, the apex
of the pycnidium being usually papilliform. I have labeled this
Septoria intermedia Ell. & Evht., but think nevertheless that it
is a form of S. solidaginicola Pk.
The conidiophores of Cercospora callae Pk. & Clint, are described
as “short”, but in some specimens they exceed 100/* in length.
It happened that shortly before looking at the description and
figures of Septoriopsis Stevens & Dalbey (Mycologia 11: 4), I had
been examining sections of Ribes leaves bearing Septoria sibirica
Thuem. (Saccardo, Ann. My col. 13: 122). In this fungus, as it
occurs in Wisconsin, the pycnidia are often defective thereby
leading to the proposed name Cylindrosporium ribis Davis (Trans.
Wis. Acad . 16: 759). In the sections at which I had been looking
there had apparently been proliferation of the cells forming the
pycnidial wall, filling the cavity and pushing the sporuligerous
layer to the apex of the pycnidium, resulting in a stromatoid tubercle
bearing scolecospores on its summit. The three figures illustrating
Septoriopsis could be matehed in a single section of the Ribes leaf.
Davis— Notes on Parasitic Fungi in Wisconsin — -VII. 401
This, of course, suggests that the proposed genus may have been
founded upon abnormal material. However, there are fungi that
seem to have normally the structure attributed to Septoriopsis,
i. e., a dark, cellular, tuberculiform pseudostroma emerging from
the leaf and bearing sessile scolecospores. An example of this is
Cercospora leptosperma Pk. Another is Cercospora longispora
Pk. I would not refer these to Tuberculariaceae because the cellu¬
lar pseudostroma is similar to that from which arise the conidio-
phores of many Hyphales, but rather to Mucedinaceae microne-
meae. In this position the genus, if the type is normal, might be
made to fill the gap to which I adverted in “Notes” III (p. 255).
As to Cercospora longispora Pk: As I see it the dark “flocci”
of the author’s description are not conidiophores of the parasite
but instead the conidia arise in tufts directly from small, dark,
tuberculiform pseudostromata.
With this coi ■‘ption of the genus I am writing Septoriopsis
longispora comb.
Cercosn l ngisvova, Pk. 35th "Report p. 141.
SeptoriopSx (Pk.) n. comb.
Cercospora < na tk. 30th Report p. 55.
Cylindrosporu ■; tospermum Pk. Trans. Wis. Acad. 14: 91,
17: 883.
Cercosporella leptosperma (Pk.) Davis Trans. Wis. Acad. 19: 706.
While these fall in Hyphales their phylogenetic relationship is
probably with Septoria and the name is therefore an appropriate
one.
Cylindrosporium tradescantiae Ell. & Kell, forms well developed
pycnidia, and I am therefore referring it to Septoria.
Phleospora oxyacanthae (Kze. & Schm.) Wallr. was erroneously
given as PM. crataegi in the provisional list. There is a note on
this fungus in “Notes” III (p. 254), where the proper name is
used.
In “Notes” II (p. 99), Populus balsamifera was given as a
host of Fusicladium radio sum (Lib.) Lind with the statement that
but a single collection had been made. In July, 1918, another
collection on this host was made in the same locality, Sturgeon
Bay. Like the previous one this collection differs from the form
on Populus tremuloides and P. grandidentata and should, I think,
be kept distinct.
26— S. A. L.
402 Wisconsin Academy of Sciences, Arts , and Letters.
Fusicladium radiosum (Lib.) Lind var. balsamiferae n. var.
Conidia 20-33 x 10-11/*, usually biseptate, the short basal and
apical cells similar and obtuse, central cell elliptic-oblong. On
Populus balsamifera. Sturgeon Bay, Wisconsin. I have not seen
the Yenturia stage.
Aecidium iridis Gerard has been shown by Whetzel and Arthur
to be connected with a rust on Phalaris arundinacea that is ap¬
parently a race of Puccinia sessilis Schneid. The propinquity of
abundant aecia on Iris and uredinia on Phalaris was conspicuous
along the shore of a small lake near Wild Rose in July, 1918.
In “Notes” IY (p. 676) there was a statement that the sorghum
kernel smut upon which was based the record of Sphacelotheca
sorghi (Lk.) Clinton in the preliminary list and thence to the pro¬
visional list, was S. cruenta (Kuehn) Potter as determined by
Potter. This is not to be taken as indicating that S. sorghi (Lk.)
Clint, does not also occur in the state.
Tracy a lemnae (Setch.) Syd. was recorded in the third supple¬
mentary list on the basis of a collection on Spirodela polyrhiza in
southeastern Wisconsin. I did not see it again until September,
1917, when it was collected at Arcadia in western Wisconsin. The
spores are usually crowded and prismatic [This was collected also
at Chetek in September, 1918.]
Telial specimens of Cronartium were collected on Bibes cynos-
bati in 1918 at Schofield (C. E. Allen) and Keshena (R. H. Den-
niston) in central Wisconsin. These locations are far removed
from any known development of aecia.
The station near Sturgeon Bay where Coleosporium sonchi-arven-
sis (Pers.) Lev. was observed in 1912 and 1913 was visited in 1918,
and uredinia were found to be present on leaves of Sonchus asper.
The Aecidium on Euphorbia commutata recorded under the
name Aecidium euphorbiae Gmel. in the provisional list is Aeci¬
dium tithymali Arth. (Bull. Torrey Bot. Club 45: 151, [1918]. It
has been observed in Wisconsin at but the single station, where it
seemed to be nearly extinct when I last saw it.
Aecidium lysimachiae (Schlecht.) Wallr. has been connected
with Puccinia limosae Magn. on Carex.
Davis— Notes on Parasitic Fungi in Wisconsin — VII. 403
Additional Hosts
Synckytrium aureum Schroet.
On Geum strictum. Two Rivers. The material is not mature,
but the parasite is doubtless the one that has been referred to this
species
Peronospora parasitica (Pers.) Fr.
On Lepidium apetalum. Coloma and Hancock. Op Arabis
hirsuta. Fish Creek and Madison. Working mostly with Europ¬
ean material, Gaeumann has divided the mildews on Cruciferae
into upwards of fifty subspecies based primarily on physiological
characters but using morphological characters also. ( Beih . Dot.
Central. 35: 395 et seq.). Field observation in Wisconsin indi¬
cates specialization on the various hosts.
Microsphaera alni (Wallr.) Wint.
To this species I have referred specimens on Baptisia bracteata
collected at Arcadia. The measurements of the asci are 36-51 x
30-36/x, of the spores 12-18 x 9/*.
Frysiphe cichoracearum DC.
On Artemisia serrata. Arcadia.
Phyllachora on Sporobolus cryptandrus has been collected at
Trempealeau by L. S. Cheney. The asci measured were 65-80 x
10-13/a, the spores 10-12 x 6-7/a.
Examination of a collection on Thalictrum dioicum made near
Jacksonport, July 25, 1918, leads me to surmise that it is a micro-
conidial state of Ascochyta clematidina Thuem. var. thalictri
Davis with the pycnidia mostly imperfectly developed and that
Gloeosporium thalictri Davis (Trans. Wis. Acad. 16: 760) is of the
same character. A collection on Thalictrum dasycarpum made at
Bruce August 30, 1918, is of the Ascochyta clematidina type hav¬
ing sporules 12-23 x 3-6/a. The material is probably immature,
as the sporules appear continuous, the bilocular condition show¬
ing only when stained. This supports the suggestion made in
“Notes” Y (p. 698) as to the relationship of the forms on Actaea,
Clematis, and Thalictrum. The group is of the same character
as the forms referred to on succeeding pages of “Notes” Y.
Ascochyta lophanthi var. lycopina Davis.
On Lycopus virginicus. Bruce. Sporules 20-28 x 7-10/a, the
longer ones sometimes 2-3 septate.
404 Wisconsin Academy of Sciences, Arts, and Letters.
Sporules from loculi of Phyllachora on Calamagrostis cana¬
densis have been observed. They were fusoid-cylindrical, guttu-
late, continuous, 16-20 x3/x. Doubtless a spermogonial state.
Septoria bromi Sacc. On Bromus altissimus. Bruce. The
record of the occurrence of S. graminum Desm. on this host in
“ Notes’ ’ V (p. 694) was based on a very poor specimen and should
be canceled.
Septoria verbenae Rob. & Desm.
On Verbena bracteosa. Wild Rose. Sporules 35-75 x
Septoria atropurpWea Pk.
On Aster puniceus. Sullivan and Spooner.
Sacidiwm ulmi-gallae Kell. & Sw.
On TJlmus americana. Wild Rose.
Record has been made of the occurrence of Gloeosporium salicis
West, in Wisconsin on willows of European origin. A correspond¬
ing Melanconiacea on the upper surface of the leaves of native
willows, however, I am referring to Marssonina kriegeriana (Bres.)
Magn.
Collections have been made as follows:
On Salix longifolia. Madison.
S. discolor. Racine.
S. cordata. Shiocton.
S. syrticola. Two Rivers.
S. petiolaris. Arcadia. On this host the acervuli are hypo-
phyllous, sporules 13-17 x 314-6/*,, usually curved, finally
septate toward the narrower end.
Marssonina rubiginosa (Ell. & Evht.) which was recorded in
“Notes” V is apparently merely a form of this. Examination of
a portion of the type collection of Marsonia nigricans Ell. & Evht.
(Proc. Acad. Nat. Sci. PJiila. 1891, p. 84) kindly sent by Pres.
Dearness leads me to believe that the host is Populus balsamifera
and the parasite Marssonina popnli (Lib.) Magn.
Piricularia grisea (Cke.) Sacc.
On Leersia virginica. Blue Mounds and Maiden Rock. Leersia
oryzoides. Millston. Leersia sp. indet. Hixton. Setaria italica
(cult.) Madison. This fungus appears to be divided into several
physiological races. P. oryzae Cav. and P. setariae Nishikado have
Davis — Notes on Parasitic Fungi in Wisconsin — VII. 405
been segregated. (See Nishikado, Ber. Ohara Inst, landivirtsch.
Forsch. 12:171 el seq.) Formosan material on Leersia hexandra
has been segregated by Sawada under the name Dactylaria leersiae
but has been referred to Piricularia by S. Ito. See Mycologia
12:30, 32.
Cercospora rosicola Pass.
On Rosa ~blanda. Wonewoc. A variable species as one might
perhaps expect from the variable character of the hosts. The vari¬
ation is especially marked in the conidiophores. In this collection
they are amphigenous, sometimes scattered or in small tufts, more
or less undulate, usually dilated at base, 40-75 x 3/a. Fungi Co¬
lumbiana 3412 , on Rosa blanda, London, Canada (Dearness) is an
extreme form with slender, undulate conidiophores sometimes at¬
taining a length of 150/a. This form with long, slender, more or
less spreading and undulate conidiophores I am designating var.
undosa n. var. The spots on Rosa blanda are less definite and
orbicular and there is less of the purple discoloration of the sur¬
rounding leaf area, but that is probably determined by the char¬
acter of the tissue of the host.
Cercospora galii Ell. & Hoi.
On Galium asprellum. Westboro, Bruce, and Nekoosa. The
spots are often indeterminate and the conidia tapering. Cercos¬
pora punctoidea Ell. & Hoi. (in lit.) was recorded in A Supple¬
mentary List of Parasitic Fungi of Wisconsin , No. 312 (Trans.
Wis. Acad. 9: 167), but a description was never published pre¬
sumably because Mr. Ellis concluded that it was not distinct.
In “Notes” I (p. 90) mention was made of the occurrence of
a mucedine on leaves of Ribes americanum. What is perhaps the
same fungus was collected at Dexterville on spots on leaves of
Ribes prostratum on the last day of August, 1917. The appear¬
ance of the spots suggests that they were caused by Septoria si-
birica Thuem. but no pycnidia were found. The following notes
were made: Tufts amphigenous, scattered, snow white, variable
in size; conidiophores hyaline, straight or lax or flexuose, contin¬
uous, sometimes branched, 12-45 x1^/a; conidia hyaline, fusiform
to cylindrical, straight, continuous, catenulate, 5—22 x 1-2/a.
Thalictrum revolutum should have been recorded as a host of
Entyloma thalictri Schroet., specimens collected near Kacine be¬
ing upon what I take to be that host species.
406 Wisconsin Academy of Sciences , Arts , and Letters.
Uromyces hyperici-frondosi (Schw.) Arth.
Aecia and uredinia on Hypericum canadense . Wild Rose.
The covered leaf rust that was recorded in the provisional list
under the name Puccinia perminuta Arth. as occurring on Cinna
arundinacea has been collected on Cinna latifolia also at Hixton.
This is now thought to belong to the species bearing aecia on
Impatiens which has been named Puccinia impatientis by Arthur,
P. impatienti-elymi Arth. by Klebahn and P. ely mi-impatient is by
the writer.
Puccinia graminis Pers.
Telia on Agropyron tenerum. Barron (L. S. Cheney).
Puccinia menthae Pers. var. am&ricana Burr.
On Blephilia hirsuta. Wonewoc.
In the summer of 1917, while collecting along the shore of
Lake Michigan at Two Rivers, fasciation of certain branches of
Juniperus horizontalis was observed such as might be produced by
Gymnosporangium but quite different from the galls caused by
G. corniculans Kern which occurs upon this host species. Upon
search through the adjacent woods a Roestelia was found upon
leaves of Amelanchier having a slender peridium like that of the
aecial stage of Gymnosporangium juvenescens Kern, a species
that occurs on Juniperus virgimana in western Wisconsin but
which is not known to occur in the eastern part of the state. In
May, 1918, a trip was made to the locality and telia were secured
with which plants of Amelanchier canadensis were infected in the
greenhouse the infections resulting in the same type of aecia that
had been found the previous summer in the field. In July the lo¬
cality was again visited and Roestelia on Amelanchier was se¬
cured, and the trip extended along the lake shore northward which
showed that the rust exists on the beaches with Roestelia in the
woods as far north as Bailey’s Harbor where the trip ended. Study
of the aecia and telia brought the conclusion that the rust is Gym¬
nosporangium juvenescens Kern and on submitting specimens to
Dr. Kern he concurred in the opinion. As this is not known to ac-
cur on Juniperus virginiana in eastern Wisconsin, the suspicion
arises that there is a race specialized to Juniperus horizontalis .
Coleosporium viburni Arth. Telia on Viburnum pubescens .
Keshena (R. H. Denniston). [Also at Mosinee.]
Davis — Notes on Parasitic Fungi in Wisconsin — VII. 407
Onoclea Struthiopteris should have been given as a host of
Sclerotium deciduum Davis, the pre-selerotial stage having been
collected on that host at Barron in 1907.
Additional Species
While collecting along a branch of Main Creek near Hawkins,
my attention was arrested by the powdered appearance of the
leaves of Laportea. After failing to locate wood borers in a posi¬
tion to be responsible for the appearance, a magnifier was used
which showed the presence of a Synchytrium as the cause. As
it seems somewhat different from the forms on various hosts that
have been referred to Synchytrium aureum Schroet., I have
thought it better to keep it separate.
Synchytrium pulvereum n. sp. (Fig. 1).
Galls epiphyllous, discrete, simple,' prominent (up to 130/*),
contracted at base, at first yellow, then castaneous ; resting spor¬
angia single, spherical, castaneous, wall laminated and about 5/*
thick, 40-90/* in diameter. On leaves of Laportea canadensis .
Fig. 1. Vertical section of gall and resting sporangium of Synchytrium
pulvereum n. sp. Drawn by Charles Drechsler with the aid of camera
lucida. Highly magnified.
408 Wisconsin Academy of Sciences , Arts, and Letters.
Hawkins and Bruce. August and September. There are some¬
times a few galls on the lower leaf surface, especially on the veins
and sometimes on the petiole.
[In examining stained microtome sections of fixed material of
this species Miss Dorothy Bradbury and Dr. E. M. Gilbert, under
whose direction she was working, detected summer sori. The galls
are hypophyllous, prominent, globose, 150-185/* in height and
breadth with walls three cells thick proximally, thinning out dis-
tally; sporangia probably about 30, spherical to elliptical to poly¬
hedral, averaging about 20 /* in diameter.]
Yenturia has been observed on leaves of cranberries and blue¬
berries in Wisconsin, but as the material collected was not ma¬
ture, no determinations and records were made. Shear has re¬
ported Venturia compact a Pk. as occurring on cranberry leaves in
Wisconsin (Bur. Pit. Ind. TJ. S. Dept. Agr. Bull. 110: 45).
Traces of Taphrina ulmi (Fckl.) Johans, were observed in
June, 1918, on leaves of elm shoots growing along the railroad
track in Madison. It has not been seen since.
Septoria polaris Karst.
On radical leaves of Ranunculus rhomhoideus . Plover and Wild
Rose. Of one of the collections referred here the following notes
were made: Spots orbicular to elliptical, brown becoming pallid
with a brown border, 2-8 mm. long, sometimes confluent ; pycnidia
epiphyllous, often numerous, scattered, black, globose to ovoid,
80-130 /* in diameter; sporules filiform, straight or more often
somewhat curved, continuous ( ?), 24-36 x 1/*,. This is the parasite
issued under this name in F. Columbiani 4878 and F. exot. exsicc.
434. It seems closely allied to S. ficariae Desm. S. ficariaecola Sacc.
S. cymbalarina Thuem. and S. ficarioides Pk.
[Since collected also at Caryville]
Septoria hydrocotyles Desm.
On Hydrocotyle americana. Wild Rose and Wautoma.
Septoria coreopsidis n. sp.
Spots circular or marginal and semicircular, cinereous with a
raised margin surrounded by a dark purple zone, 1-2 mm. in
diameter ; pycnidia punctate, black, scattered, innate, globose,
60-90/* ; sporules, hyaline, somewhat curved, sometimes guttu-
Davis — Notes on Parasitic Fungi in Wisconsin — VII. 409
late, 30-50 x l~V/2 y. On leaves of Coreopsis palmata. Hixton,
Wisconsin, September 1, 1917.
Specimens on Chrysanthemum Leucanthemum from Bailey’s
Harbor that I have referred to Septoria chrysanthemi Allesch.
bear spornles mostly 60-70/x long.
Leptothyrium pomi (Mont. & Fr.) Sace.
On fruit of Pyrus Malus. Sparta (L. R. Jones).
Gloeosporium equiseti Ell. & Evht.
On Equisetum sp. indet. Sullivan. The sporules remind one
of the sporidia of Entyloma.
Gloeosporium balsameae n. sp. (Plate XXX).
Acervuli hypophyllous, subepidermal, orbicular to oblong, J/2-l
mm. in length; sporules hyaline, continuous, fusoid, often in¬
equilateral, seldom curved, 16-33 x 4-6/x, borne on hyaline basidia
of about the same length. The affected leaves have the lethal
color throughout, the presence of the fungus being indicated only
by the epidermis being slightly raised over the acervuli. This
epidermal covering apparently becomes cut away at the peri¬
phery and detached in one piece.
On Abies balsamea, Clark lake, Door county, July 22, 1918.
Ramularia destructans Zinssmeister and R. panacicola Zinss-
meister are reported by the author as occurring on roots of Panax
quinquefolium at Wausau (Phytopath. 8: 570).
Ramularia minax n. sp.
Spots small, angular, black, y2-l mm. in diameter ; conidiophores
predominantly hypophyllous, subulate to cylindrical, usually con¬
tinuous, sometimes denticulate near the apex, 6-24 x 2y2-3y2n 5 con-
idia hyaline, straight, fusiform to cylindrical ; acute, 10-23 x 2y2-4ju.
On leaves of Solidago rigida. Buffalo County near Arcadia, Wis¬
consin, September 5, 1917.
Superficial repent hyphae become closely septate and brown,
contorted into brown knots 15-30/*, in diameter, forming superficial
tubercularioid pseudostromata from which the conidiophores arise
as lateral branches of the component cells, the basal portion of
the conidiophore sometimes showing some of the brown color of
the cell from which it sprang. Sometimes, however, the brown,
or sometimes hyaline, septate hypha is extended along the leaf
surface, giving off condiophores as short lateral branches. Often
410 Wisconsin Academy of Sciences, Arts, and Letters.
the hyphae ascend the trichomes, developing lateral conidiophores
and profuse conidia. In the leaf tissue is a fine (l^jt-0 hyaline
mycelium which is apparently nonseptate. Microscopically this
fungus resembles Plasmopara halstedii (Farl.) Berl. & De Toni,
for which it was mistaken in the field. This is another of the
Sporotrichum-like forms but whether it is a saprophyte, a leaf
parasite, or a parasite on Plasmopara, I do not know. [In a
collection made at Chaseburg in August, 1920, the hyphae are all
hyaline and the stromata small, loose, and hyaline. Some of the
longer conidia have a median septum, and one was observed hav¬
ing 3 septa.]
Kriegeria eriophori Bres. (Revue Mycol . 1891, p. 14, tab. cciii,
a-e; Septoglaeum dimorphum Sacc. Syll. Fung. 10:497 (1892) and
Ann. Mycol. 11:550; Platygloea eriophori (Bres.) Hoehn. Sitz.
h. k. Acad. Wiss. Wien 98:1157.)
This was collected a number of times on Scirpus atrovirens at
Kacine but not identified. It occurs also at Madison on the same
host.
Cylindrosporium artemisiae Dearn. & Barth.
On Artemisia serrata. Hixton.
In this collection the sporules are 40-60/x long.
TJstilago panici-miliacei (Pers.) Wint.
On Panicum miliaceum (cult.) Madison (A. L, Stone, 1911; W.
N. Steil, 1912), Baraboo (E. H. Toole, 1918).
Under the name Aecidium smilacinae, (error for maianthae) two
species of Aecidium were confused in the provisional list. One
of them, Aecidium magnatum Arth., has been connected with a
race of TJromyces acuminatus Arth. on Spartina Michauxiana to
which the name TJromyces magnatus Arth. has been given (Myco-
logia 9: 309-12). The aecial hosts of this in Wisconsin as far as
known at present are Smilacina racemosa, S, stellata, and Polygo-
natum. The Aecidium on a leaf of Oakesia sessilifolia collected
at Hancock I am also referring to this species. Of the four races
of TJromyces acuminatus Arth., three occur in Wisconsin. The
least unsatisfactory way of designating these, as it seems to me,
is by the use of trinomials, thus :
TJromyces acuminatus magnatus (Arth.)
TJromyces acuminatus steironematis (Arth.) This form appears
to be rare in Wisconsin.
Davis — Notes on Parasitic Fungi in Wisconsin — VII. 411
Uromyces acuminatus polemonii (Pk.)
The rust on Trifolium hybridum referred to TJromyces trifolii-
repentis (Cast.) Liro is found to be physiologically and to some
extent morphologically distinct by W. H. Davis. He therefore
separates it under the name Uromyces hybrid! W. H. Davis
(Trans. Iowa Acad. Sci. 24: 472).
Puccinia polygalae Paschke (P. pyrolae Cke.).
On Poly gala paucifolia. Casco and Tomahawk.
Coleosporium ribicola (C. & E.) Arth.
On Ribes cynosbati. Shell Lake and Hayward (C. E. Allen).
Price County (P. H. Denniston and W. N. Steil). Uredinia and
telia of this Pocky Mountain rust were collected in the course of
the white pine rust survey in 1918.
Herbarium of the University of Wisconsin,
Madison, Wisconsin, March, 1919
Plate XXX. Vertical section of acervulus of G-loeosporium balsameae
n. sp. Drawn by Charles Drechsler with the aid of camera lucida.
Magnified about 350 diameters.
TRANS, WIS. ACAD., VOL. XX
PLATE XXX
DAVIS— PARASITIC FUNGI
NOTES ON PARASITIC FUNGI IN WISCONSIN— VIII
J. J. Davis
In “Notes” IY (p. 681) it was stated that Synchytrium cellu-
lare was confined to a very limited station from which it had dis¬
appeared and that it had not been found elsewhere. In 1919 it
was rediscovered at Babcock in September, only resting sporangia
being present. It was confined to a very limited area. [This has
since been collected in other localities.]
Coccochora rubi Davis was abundant at a station on Bruce Creek
near Bruce on Rubus canadensis in 1918, but, as was so often the
case that year, the reproductive bodies were not well developed.
In “Notes” VI (p. 712) it was stated that the sporules of Sep-
toria acerina Pk. were borne in acervuli as I had seen the fungus.
Material collected at Mellen in 1919, however shows well de¬
veloped pycnidial walls.
Specimens on Psedera from Wausau and from Nekoosa are re¬
ferable to Septoria ampelopsidis Ellis, the pycnidial walls being
well developed and the sporules long and slender as described for
that species. Septogloeum ampelopsidis (Ell. & Evht.) Sacc.
seems to be the same fungus with imperfect pycnidial walls and
shorter and broader sporules. To the description of Septoria
ampelopsidia Ellis the author added the statement: “This ap¬
proaches Cylindrosporium on account of the imperfectly developed
perithecia”. As the Septoria form may be considered the nor¬
mally developed state, it would be well to use Septoria ampelopsidis
Ellis for the aggregate.
The late E. W. Roark found, as a result of an extensive series
of inoculations, that Septoria rubi West, includes two physiologi¬
cal races: one confined to blackberries (Eubatus), and the other
to raspberries. Because of his untimely death, in the military
413
414 Wisconsin Academy of Sciences , Arts , and Letters.
service, his data have not been published. His death deprived
phytopathology of a promising devotee. The results obtained by
Beach accord with Roark’s findings (Amer. Jour. Bot. 6: 26).
I have not seen an authentic specimen of Ellisiella mutica, Wint.,
but it seems probable that the fungus for which I proposed the
name Collet otrichum silpJiii (Trans. Wis. Acad. 192:686) is con-
specific therewith. If that is the case, of course Winter’s specific
name should be used unless Vermicularia silphn Schw. is the same
thing. Both Peck and Ellis doubted the distinctness of Ellisiella
from Collet otrichum.
In the provisional list Gloeosporium canadense Ell. & Evht. was
included in G. nervisequum (Fckl.) Sacc. Investigation, however,
by H. R. Rosen and by Eleanor J. Murphy indicates that it is
distinct as shown by host relations and by the characters of the
Gnomonia stage.
Gloeosporium leptospermum Pk. collected at Mosinee June 23,
1918, bears sporules but 10-13 x 3/*. The small size of the sporules
might be attributed to immaturity but some of them had already
oozed out in masses. This seems to be connected with Cryptomyces
pteridis (Reb.) Rehm.
While Marssonina as it usually occurs on the aspens bears hypo-
phyllous acervuli and small sporules in material collected at
Mercer on Populus tremuloides the acervuli are epiphyllous and
the sporules are about 18 x 10/*. A specimen from Fish Creek on
Populus balsamifera has epiphyllous acervuli containing sporules
20-26/a long.
Myrioconium comitatum Davis has been collected on leaves of
Populus tremuloides that bear no Sclerotium bifrons Ell. & Evht.
The large, dead, sharply delimited leaf areas are like those on which
the Sclerotium occurs. In this collection the Myrioconium is scat¬
tered more generally over the leaf surface than usual.
In compiling the provisional list the names of four hosts were
placed under Bamularia rufomaculans Pk., namely, Polygonum
aviculare , P. amphibium , P. Muhlenbergii, and P. cilinode. The
plant on P. aviculare is Ovularia rigidula Delacr. ( 0. avicularis
Pk.) with conidiophores varying in length up to 80/* and conidia
10-18 x 4-6/*. The leaves in the specimens on P. cilinode are
Davis — Notes on Parasitic Fungi in Wisconsin — VIII. 415
spotted, but the spots are those caused by Septoria polygonorum
Desm., and the Mucedine, which is apparently superficial, and
perhaps saprophytic, is confined to these spots. I have collected
this several times on Septoria spots on Polygonum cilinode.
Polygonum aviculare and P. cilinode , should therefore be stricken
from the list of hosts of Ramularia rufomaculans Pk. On Poly¬
gonum amphibium and P. Muhlenbergii the conidial fasciculi
appear on the lower surfaces of the leaves before the leaf tissues
die and become brown, that is, before leaf spotting is appar¬
ent. In the leaves of Fagopyrum esculentum the tissue death
proceeds more slowly and the appearance of the leaves would not
lead one to suspect the presence of the fungus until a late stage of
the attack. The same is true of Polygonum scandens which is also
a host in Wisconsin. It was apparently the absence of spotting
that led Peck to give the name Ramularia anomala to the form on
this host. The thin, firm leaves of these hosts seem to succumb
more slowly to the attack of the parasite than do the thicker leaves
of Persicaria. In July 1920 a collection was made at Caryville on
Polygonum Convolvulus that seems referable to Ramularia ano *
mala Pk. The hypophyllous tufts are effused over indefinite areas
or over the entire leaf surface; conidiophores densely fasciculate,
hyaline, straight, 6-15 x 1-2 /z ; conidia hyaline, catenulate, more or
less acute, straight, continuous, 4-20 x lj4-2y2fi- Some of the
shorter conidia are ellipsoid and of the Ovularia type.
In August 1920 there was collected in small quantity at Chase-
burg a Eamularia on Polygonum Persicaria in which the areas
bearing the tufts are not at first discolored but become pale brown
the discoloration showing more plainly after the leaf is dried. The
conidia are but 1-1 n thick. It is quite likely that the forms on
Persicaria, Tiniaria, and Fagopyrum will prove to be distinct in
their host relations. Dr. F. E. Jones failed to infect Fagopyrum
using conidia from Polygonum scandens. The conidiophores and
conidia are sometimes more slender (1%— 2/*) than the descriptions
indicate. Ramidaria occidentalis Ell. & Evht. on Eumex is a
closely related species. I have seen this only on the single species
of host, Rumex britannica, on which it is sometimes abundant.
I offer the following characterization of the growth on Polygo¬
num cilinode:
Ramularia cilinodes n. sp.
Hypophyllous on Septoria spots; conidiophores arising from
more or less intricate superficial mycelial ganglia, erect or as-
416 Wisconsin Academy of Sciences, Arts, and Letters.
surgent, hyaline, straight or somewhat curved, 30-50 x iy2-2y2u ;
conidia apical, hyaline, subacute, straight, continuous, 18-45 x
1^2-2 y2fx. On Polygonum cilinode, Hannibal, July 24, 1920,
(type), Cadott July, 28, 1920, “conidia 15-35 x 2-3/*”; Holcombe,
August 7 and 8, 1920 ; Radisson, July 7, 1906. Also Mountain,
Ellison Bay, Solon Springs, Black River Falls, and Necedah. June
and July. The fungus has been found on Polygonum cilinode only
and confined to the Septoria spots on which it shows as a delicate
white mold.
Ramularia umbrina Davis (Trans. Wis. Acad. 192:714) should,
I think, be considered a synonym of R. diervillae Pk.
The fasciculi of Cercospora rhamni Fkl. are described as oli¬
vaceous, but in Wisconsin on both Rhamnus cathartica and R.
alnifolia they are often a decided rusty brown because of the
color of the conidiophores which are 40-65 /i long.
Cercospora clavata (Ger.) Pk. is a common and variable para¬
site of Asclepias in Wisconsin. What I take to be a form of this
on Asclepias syriaca causing pale orbicular spots is of the char¬
acter of Cercospora asclepiadis Ellis (Am&r. Nat. 16:810, 1890),
C. asclepiadis Henn. (Hedwigia 41: 309, 1902), and C. venturioides
Pk. Conidiophores sometimes exceed 100^.
Ramularia ionophila Davis was given specific rank largely be¬
cause it appeared to be confined to the single species of violet,
Viola canadensis, in Wisconsin. However, a collection on Viola
ocellata made at Paradise, California, by F. R. Jones seems re¬
ferable to this species. Ramularia biflorae Magn. is the European
analogue to which perhaps the American form should be referred.
I have not seen an authentic specimen of Ramularia rubicunda
Bres., but judging from the description Cercospora subsanguinea
Ell. & Evht. can scarcely be distinct.
Arthur and Bisby (Proc. Amer. Philos. Soc. 57:201) found
that Schweinitz’s Caeoma (TJredo) teucrii as represented in the
Schweinitz herbarium is Ramularia racemosa Ell. & Mart., and pro¬
pose the binominal Cercospora teucrii (Schw.) Arthur & Bisby.
As I understand it, the object of rules in nomenclature is to secure
uniformity of usage. As all determined specimens of Cercospora
racemosa were so labeled in all herbaria such uniformity has been
secured and I see no occasion for changing all the labels. This
Davis — Notes on Parasitic Fungi in Wisconsin — VIII. 417
is apart from the matter of the validity of Schweinitz ’s descrip¬
tion which is certainly open to question.
In the provisional list Bubus idaeus aculeatissimus is given as
a host of Gymnoconia peckiana in the Caeoma stage. This is
perhaps an error as I can find no specimen that I am sure is on
a red raspberry and since my attention has been called to the
matter I have seen no instance in the field. The host characters
are usually modified in infected plants and determination often
difficult.
A collection of Aecidium monoicum Pk. on Ardbis lyrata made
by J. M. Holzinger at Trempealeau on the Mississippi River was
referred to in the appendix to the provisional list. It was col¬
lected in 1918 on the same host at Two Rivers on the shore of
Lake Michigan.
Kuehneola uredinis (Lk.) Arth. (K. albida (Kuehn) Magn.),
as I have seen it on Bubus hispidus, forms telia on the over¬
wintered leaves in the spring quickly followed by aecia (Uredo
muelleri Schroet.) on the young leaves of the season. The se¬
quence suggests that it is living mycelium that carries the rust
through the winter. [In 1921 telia were observed on this host in
September.]
Additional Hosts
Upon which the fungi mentioned have not been recorded
as occurring in Wisconsin.
Synchytrium aureum Schroet. On leaves and especially petioles
of Budbeckia laciniata. Bruce. I am applying to this the desig¬
nation that has been used for similar forms on other hosts in Wis¬
consin. The sori lose their yellow color before maturity. They
are often subepidermal on this host as on some others. The de¬
velopment of the sorus in the mesophyll I have thought might be
due to a very firm cuticle, but that explanation does not seem a
very probable one as regards this host. A collection made at
Prentice on Petasites palmatus I have also referred to this species.
Albugo Candida (Pers.) 0. Kuntze. On Arabis glabra. Mosinee.
Sphaerotheca Inumuli (DC.) Burr. var. fuliginea (Schl.) Salm.
On Bidens vulgata. Madison.
27— S. A. L.
418 Wisconsin Academy of Sciences , Arts, and Letters.
Salmon in his studies of the Erysiphaceae encountered speci¬
mens from South Dakota and Wyoming on Yicia for the recep¬
tion of which he proposed the variety ludens of Microsphaera alni
(Wallr.) Wint. Material of similar character has been collected
at Nekoosa on Desmodium paniculatum (Fig. 1) and referred to
this variety. Recurved appendage tips are not abundant.
Fig. 1. Outlines of selected appendage tips of Microsphaera alni ludens
from leaf of Desmodium paniculatum. Camera lucida drawing by Paul
A. Harvey. Magnified about 265 diameters.
Erysiphe graminis DC. Conidia on Dactylis glomerata. Fish
Creek.
Phyllachora was abundant on Andropogon furcatus at Chetek
in September, 1918, but immature. Phyllachora on Melica striata
was collected at Mosinee in September, 1919, also immature, but
the characters of the ascomata are not those of Ph. melicicola Speg.
A collection on Oakesia sessilifolia referred to Phyllosticta
cruenta (Fr.) Kickx bears sporules 15-20x3 %-6/a.
Ascochyta clematidina Thuem. var. thalictri Davis. On Tha-
lictrum dasycarpum Saxon. In this collection the sporules are 10—
17 x 3-4/i, thus approaching the type on Clematis. In the refer¬
ence to this variety in “ Notes” Y the page number should be 557
instead 757 as printed.
Ascochyta pisi Lib. On leaves of Yicia caroliniana. Fish
Creek. In this collection the pycnidia range up to 175/a in diame¬
ter with tenuous, transparent walls, the ostiole hypophyllous, the
sporules 17-30 (mostly about 23) x 3-3*4 /a. In specimens on
Yicia americana the ostiole is epiphyllous.
Davis — Notes on Parasitic Fungi in Wisconsin — Till. 419
Septoria malvicola Ell. & Mart. On Althaea rosea (cult.)
Devil’s Lake (I. Jorstad). From this collection the following
notes were made: Spots blackish brown above, angular, limited
by the veinlets, sometimes confluent, central portion bearing the
pycnidia becoming pale brown or sordid white, paler and less
distinct below, about 2 mm. in diameter; pycnidia epiphyllous,
scattered, globose, 85-100/* in diameter; sporules hyaline, acute
at one end, some of them very faintly triseptate, 24-40 x 1J4-2/*.
This is Septoria fairmani Ell. & Evht., which Beach has found to
be not distinct (Amer. Jour. Bot. 6: 15-16).
Septoria convolvuli Desm. On Convolvulus spithamaeus. Mo-
sinee. Sporules acicular, straight or somewhat curved, 30-40 x
1- 2/*, appearing continuous. (See Beach, loc. cit. 16-19.)
Of a collection on leaves of Aster Tradescanti made at Saxon
and referred to Septoria atropurpurea Pk. the following notes
were made : Spots suborbicular, dark brown with a purple
border above, paler and immarginate below, about 5 mm. in di¬
ameter; pycnidia innate, globose, 125-165/*; sporules curved, 50-
96 x 3/*. What appears to be the same species was collected also
on Aster puniceus.
Phleospora reticulata Ell. & Evht. On Vicia americana. Mosi-
nee. In this collection the dark border lines of the spots are nar¬
rower and less conspicuous or even wanting and the sporules but
2- 3/* thick. A collection on Lathyrus venosus from the same sta¬
tion bears filiform, curved sporules up to 125/* long by about 1 y2\x,
thick. Apparently a poorly developed state.
Ribes Cynosbati , R. gracile, and R, oxyacanthoides should be
added to the list of hosts upon which Gloeosporium ribis (Lib.)
Desm. & Mont, occurs in Wisconsin.
Collet otrichum solitarium Ell. & Barth. On Solidago speciosa
and S. nemoralis Nekoosa.
Cylindrosporium eminens Davis. On H elianthemum majus.
Chetek.
Record was made in “Notes” VI (p. 708) of the occurrence
of Ramularia alismatis Fautr. on Sagittaria heterophylla at Ar¬
cadia. It has since been found on the same species of host at Shioc-
ton and Chetek. As was suggested in “Notes” I (p. 84), this
420 Wisconsin Academy of Sciences , Arts , and Letters.
should not be referred to Ramularia but belongs under the rubric
Micronemeae. An undifferentiated hypha makes its way to a
stoma where it bears upon its extremity a conidium. In surface
view the conidia appear to be standing on end on the leaf sur¬
face. They are easily detached, hence in examining sections they
are seen not in situ but floating free in the mounting fluid. A
similar mode of development is found in the fungus to which
Oudemans gave the name Mar sonia secales and upon which Hein-
sen founded the genus Rhynchosporium. To this genus therefore
I would refer the fungus on Alisma and Sagittaria heterophylla,
considering the bending of the apical portion of the conidium to
one side in the type as of no more than specific import. Diedicke
(Ann. My col. 10:479) states that the specimens of Septoria alis-
matis Oud. that he has examined are of the same character as
Ramularia alismatis Fautr., which is true of those that I have
seen, including North American Fungi , second series, 3371 , col¬
lected in Canada by Dearness. Rhynchosporium Heinsen then,
being emended to include all Mucedinaceae micronemeae hyalo-
didymae, would contain at present two known species :
Rhynchosporium secales (Oud.) n. comb.
Marsonia secales Oud.
Rhynchosporium gramimcola Heinsen
Collected in Wisconsin on the following hosts by A. 0. John¬
son, J. G. Dickson, and C. Drechsler:
JDactylis glomerata
Agropyron repens
Hordeum vulgare
Hordeum distichum
Hordeum hexastichum
Secdle cereale
Bromus inermis
Rhynchosporium alismatis (Oud.) n. comb.
Septoria alismatis Oud.
Ascochyta alismatis Ell. & Evht.
Ramularia alismatis Fautr.
Didy maria aquatica Starb.
On Alisma Plant ago -aquatica and Sagittaria heterophylla .
Cylindrosporium bandy sianum Sacc. on Alisma Plantago (Ann.
My col. 12: 296) judging from the description is probably the same
fungus.
Davis — Notes on Parasitic Fungi in Wisconsin — VIII. 421
The conidiophores of Bamularia sagittariae Bres. were described
by the author as being short branched or forked, septate, 25-26
x 3-5/*,. (Hedwigia, 1896, p. 200) . In my examination of Krieger ’s
Fungi saxonici 1294 issued under this name, I did not find such
conidiophores but instead sessile conidia similar to those that are
found in Wisconsin on Sagittaria heterophylla.
Bamularia impatientis Pk. is also micronematous but the coni¬
dia are not septate and are often clustered reminding one of
Microstroma.
Ovularia pulchella (Ces.) Sacc. var. agropyri Davis. On
Agropyron repens. Fish Creek.
Bamularia rosea (Fckl.) Sacc. On Salix glaucophylla. Ne-
koosa.
Bamularia virgaureae Thuem. On Solidago speciosa. Nekoosa.
Conidia 30-96 x 3-5/*, 1- -5-septate, cylindrical to obclavate.
Of a specimen on Solidago serotina collected at Saxon it was
noted : Conidiophores 20-50 x 4/* ; conidia 40-100 x 4/*, the longer
somewhat tapering. Leaves at first unspotted but showing small
dead areas later.
Piricularia parasitica Ell. & Evht. On Phyllachora on leaves of
Elymus canadensis. Cornell.
Cladosporium nervale Ell. & Dearn. On Bhus typhina. Elli¬
son Bay and Nekoosa. Specimens on Bhus glabra were recorded
under Cladosporium aromaticum Ell. & Evht. in the provisional
list.
Cercospora muhlenbergiae Atk. On Muhlenbergia foliosa. Cor¬
nell.
Cercospora vexans C. Massal. Collected in small quantity on
Fragaria vesca Mellen. This fungus is common on F. virginiama
in the woods in northern Wisconsin.
Cercospora rosaecola Pass. var. undosa Davis. On Bosa humilis.
Nekoosa.
Cercospora violae Sacc. On Viola conspersa. Mosinee. The
spots are reddish brown before becoming pallid.
Cercospora osmorrhizae Ell. & Evht. On Osmorrhiza Claytoni.
Mellen.
422 Wisconsin Academy of Sciences , Arts , afieZ Letters.
Cercospora varia Pk. On Viburnum pubescens. Brnce. In
this collection the conidiophores are 20-30 x 4-6/*, the conidia
60—130 x 3 u- The fasciculi are mostly epiphyllous. As remarked
by Ellis and Everhart (Jour. My col. 1:63), this seems doubt¬
fully distinct from C. tinea Sacc., but I have not seen European
specimens. The petioles of the leaves of the host in this collec¬
tion are 2 cm. long.
Cercospora helianthi Ell. & Evht. On Helianthus strumosus.
Nekoosa. In a collection on H. occidentals from the same lo¬
cality, the conidiophores are mostly epiphyllous on small, im-
marginate purple spots.
Entyloma crastophilum Sacc. On upper leaves and axis and
branches of the panicles of Glyceria pallida. Chetek. This is
the only indigenous grass upon which I have seen this species.
Uromyces fabae (Pers.) DBy. Uredinia and telia on Vida
caroliniana. Fish Creek. Aecia and uredinia were present on
Lathyrus palustris in the same locality but on the opposite side of
the bay. Uredinia on Vicia caroliniana. Nekoosa.
Telial material of Uromyces acuminatus magnatus (Arth.) on
Spartina Michauxiana collected at Hancock in October, 1918, and
wintered out of doors infected Smilacina stellatay S. racemosa ,
Polygonatum commutatum, P. biflorum , and Oakesia sessilifolia in
the greenhouse in May, 1919. Uvularia grandiflora and Conval-
laria majalis proved immune. Two plants of Oakesia were
brought in from the woods and exposed to infection. One was
a vigorous, growing plant having seven leaves. Every leaf of
this plant was thoroughly infected, as shown by the abundant
spermogonia, with the result that they withered and died without
forming aecia except at the base of the lowest leaf. The other
plant was stunted, having but three leaves which seemed old and
firm. This plant was but slightly infected, but bore aecia on a
spot on one of the leaves.
Having some overwintered telial material of Puccinia stipae
Arth. on Stipa spartea which germinated with unusual vigor, a
young plant of Aster multi florus was brought into the greenhouse
and exposed to infection. The result was thought to be nega¬
tive, but before discarding the Aster plant it was examined care¬
fully and two very small aecial spots were found, one bearing one
cup, the other three. These spots were on the leaf edge.
Davis — Notes on Parasitic Fungi in Wisconsin — VIII. 423
Puccinia poarum Niels. Uredinia on Poa triflora. Bruce.
Puccinia agropyri Ell. & Evht. Uredinia on Agropyron caninum.
Mosinee.
Puccinia eriophori Thuem. Telia on Eriophorum viridicarina-
tum. Weyerhaeuser.
Puccinia polygoni-amphibii Pers. A single aecial spot, but a
well developed one, on Geranium Bicknellii. Mosinee.
Epiphyllous telia on Ledum groenlandicum collected at Stur¬
geon Bay have been referred to Melampsoropsis ledicola (Pk.)
Arth. because of their location on the upper surface of the leaf.
Additional Species
Not hitherto recorded as occurring in Wisconsin.
Sphaerotlfieca pannosa (Wallr.) Lev. has been collected a few
times at Madison on cultivated roses, but it does not seem to main¬
tain itself.
A collection of Cercospora comari Pk. on Potentilla palustris
from Nekoosa bears also a Mycosphaerella that I have referred to
M. innumerella (Karst.). The globose black perithecia are
closely and regularly placed on the lower surface of the black
spots and are 70-85 /* in diameter; the asci are 33^t7 x 7/*, and the
fusoid-cylindrical, hyaline, uniseptate spores 13-18 x 2%-3/*. The
collection was made in July and probably at full maturity would
give larger measurements. It appears to be genetically connected
with the Cercospora.
Sphaerulina pallens n. sp.
On large, pale, dead, mostly terminal, leaf areas ; perithecia epi-
phyllous-innate, scattered, black, globose, about 150//, in diameter;
asci elavate-cylindrical, short-stipitate, more or less curved, apara-
physate, 75-85 xlO/*; spores distichous, subfuligineous, fusoid-
cylindrical, 5-9 septate, 30-60 x 3-4/*.
On Carex sp. indet. A large coarse species. Mellen, Wiscon¬
sin, July 31, 1919.
Acantho stigma occidentalis (Ell. & Evht.) Sacc. On leaves of
Cirsium discolor. Black Earth and Madison (I. Jorstad).
424 Wisconsin Academy of Sciences, Arts, and Letters .
Phacidium planum n. sp. (Plate XXXI.)
Apothecia foliicolous, seriate, sometimes confluent, erumpent,
little prominent, overlying epidermis circumscissile ; disk plane,
pale brown or sordid, surrounded by black byphae; asci clavate,
octosporous, 50-80 x 9-12 /* ; spores inordinate, clavate to f usoid-
cylindrical, acute at base, hyaline, continuous, 30-40 x 3/*, para-
physes filiform, about as long as the asci. On languishing leaves
of Pinus Strobus. Mosinee, Wisconsin, July, 1919. The asci and
spores suggest those of Hypoderma.
Phacidium expansion n. sp. (Plate XXXII.)
Apothecia foliicolous, scattered, erumpent, expanding,
mm. in diameter, surrounded by a wall composed of black hyphae
which are adnate proximally but free distally ; asci clavate-
cylindrical, narrow at base, octosporous, 60-80 x 7-9/* ; spores mo-
nostichous, elliptical, hyaline, continuous, 9-12 x 3-5 /* ; paraphyses
numerous, filiform, thicker toward the apex, slightly longer than
the asci. The infected leaves, which are scattered, are killed be-
for the apothecia appear. On leaves of Picea mariuna. Ogema,
Wisconsin, July 29, 1919.
Phacidium balsameae n. sp.
Apothecia flat, circular, subepidermal, exposed by circumscissile
removal of the overlying epidermis, becoming prominent, sur¬
rounded on the sides by a wall of more or less conglutinated paral¬
lel black hyphae, about 350/* in diameter ; asci cylindrical, nar¬
rowed to the base, straight or the peripheral ones somewhat
incurved, 80-100 x 10/* ; spores ovoid, hyaline, obliquely mono-
stichous, 12-14x6-7/*; paraphyses filiform, a little longer than
the asci. On scattered leaves of Abies balsamea, which become
brown and dead. Vilas County, Wisconsin, July 27, 1902.
The three foregoing species are brought together here for the
purpose of calling attention to them rather than to indicate their
exact systematic position.
Lophodermium thuyae n. sp. (Fig. 2, page 425.)
Perithecia epiphyllous, black, hysteriiform, outer wall thickened
toward the sulcus ; asci cylindrical, narrowed to the base, 80-100
x 10/* ; spores 8, filiform, parallel, approximating in length that
of the ascus ; paraphyses filiform, distally flexuous or contorted,
as long as the asci. On Thuya occidentalis. Saxon, Wisconsin,
August 16, 1919.
Hypoderma brachysporum (Rostr.) Tubeuf.
On leaves of Pinus Strobus attached to dead twigs. Mosinee.
I have seen no septation of the spores. The classification of this
group is unsatisfactory.
Davis — Notes on Parasitic Fungi in Wisconsin — Till. 425
Fig. 2. A. Vertical section of an immature perithecium of Lopho-
dermium thuyae n. sp., showing unusual -thickening of the upper wall.
Magnified about 150 diameters. B. Vertical section of a mature peri-
theeium of the same. C. An ascus and paraphysis magnified about 350
diameters. Drawn with the aid of camera lucida by Charles Drechsler.
Phyllosticta platanoidis. Sacc.
On Acer Negundo. Madison.
Phyllosticta bacteriospora Vnill.
On Tilia americana. Mellen.
What is perhaps Asteroma populi Rob. & Desm. has been col¬
lected at Mellen on Populus tremuloides. The material is young
and the “fibrils” are not yet black.
S 'tagonospora arenaria Sacc.
To this species I have referred collections on leaves on Elymus
canadensis from Mellen. The leaves bear blackish brown spots,
bnt the pycnidia are found also in paler portions of the leaves.
The sporules are 20-33 x 3^/i, triseptate. Exceptionally 1-, 4-,
or 5- septate sporules occur.
426 Wisconsin Academy of Sciences , Arts, and Letters.
Stagonospora tetramera n. sp.
Spots pale brown to sordid with a darker brown, indefinite
margin, paler and usually immarginate below, oblong to ellipti¬
cal, about 1 cm. long, sometimes confluent; pycnidia scattered,
epiphyllous-innate, black, ostiolate, globose to depressed-globose,
100-150/a in diameter; sporules hyaline with coarsely granular
cytoplasm, fusoid-cylindrical, obtuse and rounded at both ends,
straight, triseptate, 55-75 x 10-13/a. On leaves of Car ex (riparia?)
Madison, Wisconsin, September 16, 1919.
Stagonospora petasitidis Ell. & Evht.
On Petasites palmatus. Prentice. Sporules about 4/a thick.
Septoria angustifolia Ell. & Evht.
On Kalmia polifolia. Mercer. In the collection referred to this
species provisionally the spots are castaneous above, 2-5 mm. in
diameter or sometimes longer when running along the leaf margin
and the sporules are 30-60 x 1-1%/a. The ‘ ‘ white-glaucous ’ ap¬
pearance of the lower surfaces of the leaves of the host is due
to the numerous small white trichomes.
Septoria xylostei Sacc. & Wint.
Specimens on Lonicera canadensis collected at Saxon have been
referred to this species.
Fig. 3. A. A leaf of Vaccinium canadense bearing Piggotia vaccinii
n. sp. Natural size. B. Vertical section of a pycnidium of the same
magnified about 350 diameters. Drawn by Charles Drechsler with the aid
of camera lucida.
Piggotia vaccinii n. sp. (Fig. 3.)
Spots suborbicular to semi-elliptical, reddish brown becoming
paler and often mottled above, subolivaceous below, 1-2 cm. in
length, sometimes confluent ; pycnidia epiphyllous, scattered,
Davis— Notes on Parasitic Fungi in Wisconsin— VIII. 427
black, flattened, orbicular, astomous, irregularly rugose, 140-200/a ;
sporules hyaline, oblong to short cylindrical, 2-6 x 3^-1/x ; sporo-
phores hyaline, filiform, straight, constipate, about 12/a long. On
leaves of Vaccinium canadense. Tomahawk, Wisconsin, August
25, 1919. The spots usually extend from the leaf margin to the
midrib, but are sometimes terminal and sometimes include nearly
the entire leaf. The clypeus is membranous and often punctulate
as in Sacidium or finely rugulose.
Gloeosporium bicolor n. sp.
Spots circular to irregular, immarginate, olivaceous, sometimes
confluent, *A- 2 cm. in diameter, with a sterile, reddish brown
central portion 2-4 mm. in diameter ; acervuli hypophyllous,
small, subolivaceous, scattered on the olivaceous portion of the
spot; sporules, hyaline, fusoid-cylindrical, rarely curved, contin¬
uous, 7-26 x 3-6/a, basidia short or obsolete. On leaves of Quer-
cus hicolor. Chippewa Falls, Wisconsin, September 14, 1918. The
central portion of the spot resembles that caused by Marssonina
martini (Sacc. & Ell.) Magn.
Gloeosporium ramosum Ell. & Evht.
On Polygala sanguinea. Nekoosa. In this collection the acer¬
vuli are mostly on the stems, speedily killing the host. The spor¬
ules are 8-13/a long, mostly more or less lunate.
Gloeosporium fraxineum Pk.
On Fraxinus pennsylvanica. Mosinee. Gloeosporium irregulare
Pk. is evidently not distinct from G. aridum Ell. & Hoi.
Collet otrichum magnusianum Bres.
On Malva rotundifolia. Madison (A. C. Foster) . Referred to
this species because of the host. In this collection the spots are
not white and arid but brown and subindefinite, the acervuli
bright cinnamon brown, the sporules 10-14 x 4-5/a, the setae less
numerous than in typical C. malvacearum (Br. & Gasp.?) South-
worth. A specimen on the same species of host from Maryland
(C. A. Schwarze, May 22, 1919) bears setae up to 100/a in length.
Spiraea tomentosa was given as a host of Septoria salicifoliae
(Trek) in the provisional list and the character of the fungus on
this host referred to in the appendix (Trans. Wis. Acad. 172 :-
983-984) . It appears to be distinct from Cylindrosp orium salici¬
foliae (Trek) Davis, and should doubtless be referred to Cylindro-
sporium spiraeicolum Elk & Evht. When the sporules become free
the flagelliform basal portion is usually curved. As C. spiraei-
colum was founded on material from Idaho I was led to re-ex-
428 Wisconsin Academy of Sciences, Arts, and Letters.
amine a collection on Spiraea densiflora (given as S. corymb osa f )
made at Wallace, Idaho, and referred to Septoria salicifoliae
Trel. in Trans. Wis. Acad . 152 : 776. In this collection the acer-
vuli are epiphyllous as in C. spiraeicolum Ell. & Evht., but the
spornles are of the C. salicifoliae type but unusually long and
slender (50-100x2-3/*) and mostly but little curved. This may
prove to be distinct.
Eamularia repens Ell. & Evht.
On Aralia nudicaulis. Nekoosa.
Macroscopically this is indistinguishable from Cercosporella lep-
tosperma Pk.
Eamularia magnusiana (Sacc.) Lindau.
On Trientalis americana. Prentice. In this collection the co-
nidiophores spring from black, scattered, stromatoid tubercles.
Cercosporella dearnessii Bubak & Sacc.
On Solidago altissima. Madison. In this collection the conidio-
phores are but 30-50/* long. Very long (up to 180//,) and slender
conidia occur as well as short ones. They are sometimes bent.
This seems to be close to the variable Eamularia virgaureae
Thuem.
Cladosporium astericola n. sp.
Spots small, brown, indefinite, 1-4 mm. in diameter; conidio-
phores mostly hypophyllous, fuligineous, scattered or somewhat
fasciculate, erect or assurgent, straight or curved, denticulate,
1 — 4- septate, constricted or not at the septa, 40-75x3-4/*;; co¬
nidia acro-pleurogenous, fuscous, catenulate, fusoid to subcylindri-
cal, uniseptate, 10-20 x 3-5/*. On upper leaves and upper por¬
tions of stems of Aster umbellatus. Mellen, Wisconsin, August 4,
1919.
Cercospora ranunculi Ell. & Holw.
On Eanunculus septentrionalis. Saxon. In this collection the
conidiophores, which spring from black stromatoid tubercles, are
shorter (30-65/*) and straighter than those of the type as de¬
scribed.
In the provisional list a Cercospora occurring on Spiraea salici-
folia was doubtfully referred to Cercospora rubigo Cke. & Hark.
As I have not seen an authentic specimen of this species and the
description is meager and not quite congruous I give some notes on
the parasite that occurs in Wisconsin.
Davis — Notes on Parasitic Fungi in Wisconsin — VIII. 429
On suborbicular to angular areas which become reddish brown
above, darker and finally granulose below; conidiophores hypo-
phyllous, brown, subdecumbent to erect, bent or more tortuous
and more or less torulose, sometimes branched, occasionally sep¬
tate, 50-100 x 3/a ; conidia hyaline, obclavate to flagelliform, more
or less curved, becoming septate, 65-135 x 3/a. On leaves of
Spiraea salicifolia. Nekoosa, Wisconsin, July 25, 1919. Micros¬
copically the conidial masses are grey. In the provisional list a
specimen on Spiraea salicifolia was doubtfully referred to Cer-
cospora rubigo Cke. & Hark. This specimen was collected at
Spooner July 20, 1911, and in the packet I find the following de¬
scription: Spots angular to suborbicular, limited by the vein-
lets, reddish brown, 5—8 mm. in diameter; conidiophores hypophyl-
lous, fuligineous, assurgent to erect, often arising as branches
from a superficial creeping mycelium, sometimes branched, usually
crooked and denticulate or nodulose, 25-60 x3/a; conidia hyaline,
attenuate from about 10/a above the base where the diameter is
greatest, straight or more often somewhat curved, pluriguttulate,
50-125 x 3-4/a.
Cercospora medicaginis Ell. & Evht.
On Medicago lupulina. Madison (F. R. Jones). The type of
this species was probably immature as the mature conidia become
flagelliform and 100-165/a in length.
Cercospora flagellifera Atk. ?
Of what is perhaps a northern form of this species the follow¬
ing notes were made : Spots few, scattered, definite, angular, dark
reddish brown above, lighter brown below, 2-5 mm. long; conidio¬
phores amphigenous, fuscous, closely fasciculate, usually straight,
simple, continuous, 17-40 x 3/a ; conidia hyaline, obclavate, straight
or somewhat curved, septate, 55-80 x 3-4/a. On leaves of Lespedeza
capitata. Saxon, July 25, 1919. Nekoosa, July 17 and 19, 1919.
Cercospora gaultheriae Ell. & Evht.
On Gaultheria procumbens. Millston.
Cercospora tuberculella n. sp.
On small, angular, somewhat paler areas, limited by the vein-
lets, which finally become black; conidiophores hypophyllous, fas¬
ciculate from scattered, black substomatal pseudo stromata, fuligen-
ous, straight to subundulate, simple, sometimes with a single sep¬
tum, 20-50 x 4—6/a ; conidia dilute fuligenous, cylindrical to obcla-
vate-cylindrical or occasionally fusoid-cylindrical, obtuse, usually
straight, becoming 1--3- septate, 30jf-60 x dr-fi/^. On leaves of
Convolvulus sepium. Madison, Wisconsin, September 16, 1919.
This collection is peculiar because of the very abundant, much
430 Wisconsin Academy of Sciences , Arts, and Letters.
branched, tortnons intrafoliar mycelium which is seldom filamen¬
tous but usually nodulose to moniliform. It is possible that this
will prove to be a form of Cercospora convolvuli Tracy & Earle
(Bull. Torrey Bot. Club 27: 187).
Cercospora tortipes n. sp.
On indefinite leaf areas or entire leaves which finally become
dead; fasciculi small, scattered, mostly hypophyllous ; conidio-
phores fuligenous, usually divaricate, tortuous and denticulate,
sometimes septate, seldom branched, 30-60 x 3 /*, ; conidia hyaline,
obclavate-cylindrical, straight or somewhat curved, 50-130 x 3^.
On leaves of Veronica scutellata. Bruce, Wisconsin, September
4, 1918.
Cercospora tabacina Ell. & Evht.
On Rudbeckia laciniata. Cornell. The translation of “ ab¬
ruptly bent” by “abrupte incurva” in the Sylloge Fungorum is
somewhat misleading. Much longer conidia occur than the de¬
scription indicates, one having a length of 250 p having been meas¬
ured. The leaf tissue of an old spot becomes filled with monili¬
form mycelium, and what appear to be immature epiphyllous peri-
thecia occur. A collection on Prenanthes alba from Mosinee I
have also referred to this species. Collected also at Tomahawk on
this host. The conidiophores are not always bent but are often
undulate instead.
[A collection has since been made on Rudbeckia hirta at Cary-
ville in which the development is less vigorous the conidiophores
being largely scattered and nearly straight, 70-100 x4/u, and the
conidia 25-45 x 3-4y, indistinctly septate or continuous.]
There was collected in small quantity on Trifolium repens at
Madison in the spring of 1916 and of 1917 a rust that I have re¬
ferred to TJromyces flectens Lagh.
Puccinia montanensis Ell. & Evht.
Telia on Elymus canadensis. Hancock. Determined by Ar¬
thur.
Kern has segregated Puccinia canaliculata (Schw.) Lagh. from
P. cyperi Arth. (Mycologia 11:136 et seq.) Cyperus strigosus
and C. Houghtonii are hosts of P. canaliculata in Wisconsin. To
this species Kern refers the Aecidium on Xanthium also.
Puccinia cicutae Lasch.
Telia on Cicuta maculata. Bruce and Cornell.
mm
TRAMS. WIS. ACAD., VOL. MX
PLATE XXXI
DAVIS— -PARASITIC FUNGI
I
Davis — ■ Notes on Parasitic Fungi in Wisconsin — VIII. 431
Puccinia bardanae Cda.
Telia on Arctium minus. Madison. This was first collected in
Wisconsin by Dr. H. S. Jackson in 1918. It rapidly spread
throughout the city. [Aecia and uredinia have since been col¬
lected.]
Aecidium ingenuum Arth. (Bull. Towey Bot. Club 46: 124).
On Picea canadensis . Fish Creek and Solon Springs. This is
of the type referred to Peridermium in the provisional list. It ap¬
pears to be rare.
Herbarium of the University of Wisconsin,
Madison, Wisconsin, April, 1920
Description of Plates
PLATE XXXI
A. Apothecia of Phacidium planum n. sp. on leaf of Pinus Stratus.
Magnified 35 diameters. B. Vertical section of an apothecium of the
same. C. Asci, paraphyses, and, at the left, four free ascospores. Magni¬
fied about 350 diameters. Drawn with the aid of camera lucida by Charles
Drechsler.
PLATE XXXII
A. Apothecia of Phacidium expansum n. sp. on leaf of Picea mariana.
Magnified 4 diameters. B. Vertical section of an apothecium. C, D.
Asci and paraphyses of same magnified about 350 diameters. Drawn by
Charles Drechsler with the aid of camera lucida.
431a Wisconsin Academy of Sciences , Arts, and Letters.
INDEX
TO HOSTS AND PARASITES REFERRED TO IN
NOTES VII AND VIII.
The names of Fungi are in italic type.
Abies balsamea, 409
Acanthostigma occidentalis (E. & E.)
Sacc., 423
Acer Negundo, 425
Aecidium euphorbiae Gmel., 402
Aecidium ingenuum Arth., 431
Aecidium iridis Ger., 402
Aecidium lysimachiae (Schl.) Wallr.,
402
Aecidium magnatum Arth,. 410
Aecidium maianthae, 410
Aecidium monoicum Pk., 417.
Aecidium tithymali Arth., 402
Agropyron caninum, 423
Agropyron repens, 420, 421.
Agropyron tenerum, 406
Albugo Candida (Pers.) Kuntze, 417
Alisma Plantago-aquatica, 420
Althaea rosea, 419.
Amelanchier canadensis, 406
Andropogon furcatus, 418
Arabis glabra, 417
Arabis hirsuta, 403
Arabis lyrata, 417
Aralia nudicaulis, 428
Arctium minus, 431
Artemisia serrata, 403, 410
Asclepias syriaca, 416
Ascochyta alismatis E. & E., 420
Ascochyta clematidina thalictri Davis,
403, 418
Ascochyta lophanthi lycopina Davis,
403
Ascochyta pisi Lib., 418
Aster multiflorus, 422
Aster puniceus, 404, 419
Aster Tradescanti, 419
Aster umbellatus, 428
Asteroma populi Rab. & Desm., 425
Baptisia bracteata, 403
Bidens vulgata, 417
Blephilia hirsuta, 406
Bromus altissimus, 404
Bromus inermis, 420
Caeoma (TJredo) teucrii Schw., 416
Calamagrostis canadensis, 404
Carex, 423, 426
Cercospora asclepiadis Ellis, 416
Cercospora asclepiadis Henn., 416
Cercospora callae Pk. & Cl., 400
Cercospora clavata (Ger.) Pk., 416
Cercospora comari Pk., 423
Cercospora convolvuli Tr. & Earle, 430
Cercospora flagellifera Atk., 429
Cercospora galii E. & Hoi., 405
Cercospora gaultheriae E. & E., 429
Cercospora helianthi E. & E., 422
Cercospora leptosperma Pk., 401
Cercospora longispora Pk., 401
Cercospora medicaginis E. & E., 429
Cercospora muhlenbergiae Atk., 421
Cercospora osmorrhizae E. & E., 421
Cercospora ranunculi E. & Hoi., 428
Cercospora rhamni Fckl., 416
Cercospora rosicola Pass., 405, 421
Cercospora rubigo Cke. & Hark., 428-9
Cercospora subsanguinea E. & E. 416
Cercospora tabacina E. & E., 430
Cercospora teucrii (Schw.) Arthur &
Bisby, 416
Cercospora tortipes n. sp., 430
Cercospora tuber culella n. sp., 429
Cercospora varia, Pk., 422
Cercospora venturioides Pk., 416
Cercospora vexans C. Massal., 421
Cercospora violae Sacc., 421
Cercospora tinea Sacc., 422
Cercosporella dearnessii Sacc., 428
Cercosporella leptosperma, (Pk.) Da¬
vis, 401
Chrysanthemum Leucanthemum, 409
Cicuta maculata, 430
Cinna latifolia, 406
Cirsium discolor, 423
Cladosporium aromaticum E. & E., 421
Cladosporium astericola n. sp., 428
Cladosporium nervale Ell. & Dearn.,
421
Index to Hosts and Parasites.
431b
Coccochora rubi Davis, 413
Coleosporium ribicola (C. & E.) Arth.,
411
Coleosporium, sonchi-arvensis (Pers. )
Lev., 402
Coleosporium viburni Artli., 406
Collet otrichum magnusianum Bres,.
427
Collet otrichum silphii Davis, 414
Collet otrichum solitarium Ell. &
Barth., 419
Convallaria majalis, 422
Convolvulus sepium, 429
Convolvulus spithamaeus, 419
Coreopsis palmata, 409
Cronarti/um, 402
Cryptomyces pteridis (Reb.) Rehm,
414
Cylindrosporium artemisiae Dearn. &
Barth., 410
Cylindrosporium bandysianum Sacc.,
420
Cylindrosporium eminens Davis, 419
Cylindrosporium leptospermum Pk.,
401
Cylindrosporium ribis Davis, 400
Cylindrosporium salicifoliae ( Trel. )
Davis, 427
Cylindrosporium spiraeicolum E. & E.,
427
Cylindrosporium tradescantiae E. & K,.
401
Cyperus Houghtonii, 430
Cyperus strigosus, 430
Dactylis glomerata, 418, 420
Desmodium paniculatum, 418
Didymaria aquatica, Starb., 420
Diervilla Lonicera, 416
Ellisiella mutica Wint., 414
Elymus canadensis, 421, 425, 430
Entyloma crastophilum Sacc., 422
Entyloma thalictri Schroet., 405
Eriphorum viridicarinatum, 423
Erysiphe cichoracearum DC., 403
Erysiphe graminis DC., 418
Euphorbia commutata, 402
Fagopyrum esculentum, 415
Fragaria vesca, 421
Fragaria virginiana, 421
Fraxinus pennsylvanica, 427
Fu8icladium radiosum (Lib.) Lind,
401, 402
Galium asprellum, 405
Gaultheria procumbens, 429
Geranium Bicknellii, 423
Geum strictum, 403
Gloeosporium aridum E. & Hoi., 427
Gloeosporium balsameae n. sp., 409
Gloeosporium bicolor n. sp., 427
Gloeosporium canadense E. & E., 414
Gloeosporium equiseti E. & E., 409
Gloeosporium fraxineum Pk., 427
Gloeosporium irregulare Pk., 427
Gloeosporium leptospermum Pk., 414
Gloeosporium. ramosum E. & E., 427
Gloeosporium ribis (Lib.) D. & M.,
419
Gloeosporium salicis West., 404
Gloeosporium ' thalictri Davis, 403
Glyceria pallida, 422
Gymnoconia peckiana (Howe) Trot¬
ter, 417
Gymnosporangium juvenescens Kern,
406
Helianthemum majus, 419
Helianthus occidentalis, 422
Helianthus strumosus, 422
Hordeum distichum, 420
Hordeum hexastichum, 420
Hordeum vulgare, 420
Humulus Lupulus, 399
Hydrocotyle americana, 408
Hypericum canadense, 406
Hypoderma brachysporum (Rostr. )
Tub., 424
Impatiens, 406
Juniperus horizontalis, *06
Kalmia polifolia, 426
Kriegeria eriophori Bres., 410
Kuehneola albida (Kuehn) Magn., 417
Kuehneola uredinis (Lk.) Arth, 417
Laportea canadensis, 407
Lathyrus palustris, 422
Lathyrus venosus, 419
Ledum groenlandicum, 423
Leersia oryzoides, 404
Lespedeza capitata, 429
Lepidium apetalum, 403
Leptothyrium pomi (Mont. & Fr.)
Sacc., 409
Lonicera canadensis, 426
Lophodermium thuyae n. sp., 424
Lycopus virginicus, 403
Marsonia secales Oud., 420
Marssonina kriegeriana (Bres.)
Magn., 404
Marssonina nigricans (E. & E.)
Magn., 404
Marssonina populi (Lib.) Magn., 404,
414
Marssonina rubiginosa (E. & E. )
Magn., 404
Medicago lupulina, 429
Melampsoroposis ledicola (Pk.) Arth.,
423
Melica striata, 418
Microsphaera alni (Wallr.) Wint.,
403
431c Wisconsin Academy of Sciences , Arts , and Letters.
Microsphaera alni ludens Salm., 418
Muhlenbergia foliosa, 421
Mycosphaerella innumerella (Karst,),
423
Myrioconium comitatum Davis, 414
Oakesia sessilifolia, 418, 422
Onoclea Struthiopteris, 407
Osmorrhiza Claytoni, 421
Ovularia avicularis Pk., 414
Ovularia pulchella (Ces.) Sacc., 421
Ovularia rigidula De Lacr., 414
Panax quinquefolium, 409
Panicum miliaceum, 410
Peridermium, 431
Peronospora parasitica (Pers.) Fr.,
403
Petasites palmatus, 417, 426
Phacidium balsameae n. sp., 424
Phacidium expansum n. sp., 424
Phacidium planum n. sp., 424
Phalaris arundinacea, 402
Phleospora oxyacantliae (K. & S. )
Wallr., 401
Phleospora reticulata E. & E., 419
Phyllachora, 403, 404, 418
Phyllachora melicicola Speg., 418
Phyllosticta bacteriospora Vuill., 425
Phyllosticta cruenta (Fr.) Kx., 418
Phyllosticta platanoidis Sacc., 425
Picea canadensis, 431
Piggotia vaccinii n. sp., 426
Pinus Strobus, 424
Piricularia grisea (Cke. ) Sacc., 404
Pirictdaria parasitica E. & E.; 421
Plasmopara humuli, M. & T., 399
Platygloea eriophori (Bres. ) Hoehn ,
410
Poa tri flora, 423
Polygala paucifolia, 411
Polygala sanguinea, 427
Polygonatum, 410
Polygonatum biflorum, 422
Polygonatum commutatum, 422
Polygonum amphibium, 414
Polygonum aviculare, 414, 415
Polygonum cilinode, 414, 415, 416
Polygonum convolvulus, 415
Polygonum Muhlenbergii, 414, 415
Polygonum Persicaria, 415
Polygonum scandens,. 415
Populus balsamifera, 401, 402, 404,
414
Populus grandidenta, 401
Populus tremuloides, 401, 414, 425
Potentilla palustris, 423
Prenanthes alba, 430
Psedera, 413
Puccinia agropyri E. & E., 423
Puccinia bardanae Cda., 431
Puccinia canaliculata (Schw.) Lagh.,
430
Puccinia cicutae Lasch, 430
Puccinia ely mi-imp atientis Davis, 406
Puccinia eriophori, Thuem., 423
Puccinia graminis Pers., 406
Puccinia impatientis Arth., 406
Puccinia impatienti-elymi Arth., 406
Puccinia limosae Magn., 402
Puccinia menthae Pers., 406
Puccinia montanensis E. & E., 430
Puccinia perminuta Arth., 406
Puccinia poarum Niels., 423
Puccinia polygalae Paschke, 411
Puccinia polygoni-amphibii Pers., 423
Puccinia pyrolae Cke., 411
Puccinia sessilis Schn., 402
Puccinia stipae Arth., 422
Pyrus Malus, 409
Quercus bicolor, 427
Ramularia alismatis Fautr., 419, 420-
421
Ramularia anomala Pk., 415
Ramularia bi florae Magn,. 416
Ramularia destructans Zinss., 409
Ramularia diervillae Pk., 416
Ramularia impatientis Pk., 421
Ramularia ionophila Davis, 416
Ramularia magnusiana (Sacc.) Lin-
dau, 428
Ramularia minax n. sp., 409
Ramularia occidentals E. & E., 415
Ramularia panacicola Zinss., 409
Ramularia racemosa E. & M., 416
Ramularia repens , E. & E., 428
Ramularia rosea (Fckl.) Sacc., 421
Ramularia rubicunda Bres., 416
Ramularia rufomaculans Pk., 414, 415
Ramularia sagittariae Bres., 421
Ramularia umbrina Davis, 416
Ramularia virgaureae Thuem., 421
428
Ranunculus rhomboideus, 408
Ranunculus septentrionalis, 428
Rhamnus alnifoli^,, 416
Rhamnus cathartica, 416
Rhus glabra, 421
Rus typhina, 421
Rhynchosporium alismatis (Oud. ) n.
comb., 420
Rhynchosporium graminicola Heinsen,
420
Rhynchosporium secales (Oud.) n.
comb., 420
Ribes, 400
Ribes americanum, 405
Ribes cynosbati, 402, 411, 419
Ribes gracile, 419
Ribes oxyacanthoides, 419
Index to Hosts and Parasites.
431d
Ribes prostratum, 405
Rosa, 423
Rosa blanda, 405
Rosa humilis, 421
Rubus, 413
Rubus canadensis, 413
Rubus hispidus, 417
Rubus idaeus aculeatissimus, 417
Rudbeckia hirta, 430
Rudbeckia laciniata, 417, 430
Rumex britannica, 415
' Sacidium ulmi-gallae K. & S., 404
Sagittaria heterophylla , 419, 420
Salix cordata, 404
Salix discolor, 404
Salix glaucophylla, 421
Salix longifolia, 404
Salix petiolaris, 404
Salix syrticola, 404
Scirpus atrovirens, 410
Sclerotium deciduum Davis, 407
Secale* cereale, 420
Septogloeum ampelopsidis (E. & E. )
Sacc., 413
Septogloeum dimorphum Sacc., 410
Septoria acerina Pk., 413
Septoria alismatis Oud., 420
Septoria ampelopsidis Ellis, 413
Septoria angustifolia E. & E., 426
Septoria atropurpurea Pk., 404, 419
Septoria bromi Sacc., 404
Septoria chysanthemi Allesch., 409
Septoria convolvuli Desm., 419
Septoria coreopsidis n. sp., 408
Septoria fairmani E. & E., 419
Septoria fusispora Died., 399
Septoria glumarum Pass., 399
Septoria graminum Desm., 404
Septoria hyalina E. & E., 400
Septoria hydrocotyles Desm., 408
Septoria intermedia E. & E., 400
Septoria malvicola E. & M., 419
Septoria polaris Karst., 408
Septoria rubi West., 413
Septoria salicifoliae (Trel. ), 427
Septoria sibirica Thuem., 400, 405
Septoria solidaginicola Pk., 400
Septoria tradescantiae (E. & E.) n.
comb., 401
Septoria verbenae R. & D., 404
Septoria violae West, 400
Septoria xylostei Sacc. & Wint., 426
Septoriopsis leptosperma (Pk.) n.
comb., 401
Septoriopsis longispora (Pk.) n. comb.,
401
Setaria italica, 404
Smilacina racemosa, 410, 422
Smilacina stellata, 410, 422
Smilax hispida, 399
Solidago .altissima, 428
Solidago nemoralis, 419
Solidago rigida, 409
Solidago serotina, 421
Solidago speciosa, 419, 421
Sonchus asper, 402
Spartina Michauxiana, 410, 422
Sphacelotheca cruenta (Kuehn) Pot¬
ter, 402
Sphacelotheca sorghi (Lk.) Clint, 402
Sphaerotheca humuli fuliginea (Schl. )
Salm., 417
Sphaerotheca pannosa (Wallr. ) Lev.,
423
Sphaerulina pallens n. sp., 423
Spiraea densiflora, 428
Spiraea salicifolia, 428-9
Spiraea tomentosa, 427
Spirodela polyrhiza, 402
Sporobolus cryptandrus, 403
Stagonospora arenaria Sacc,. 425
Stagonospora petasitidis E. & E., 426
Stagonospora smilacis (E. & M.)
Sacc., 399
Stagonospora tetramera n. sp., 426
Stipa spartea, 422
Synchytrium aureum Schroet., 403, 417
Synchytrium cellular e Davis, 413
Synchytrium pulvereum n. sp., 407
Taphrina ulmi (Fckl. ) Johans., 408
Thalictrum dasycarpum, 403, 418
Thalictrum dioicum, 403
Thalictrum revolutum, 405
Thuya occidentalis, 424
Tilia americana, 425
Tracya lemnae (Setch. ) Syd., 402
Trientalis americana, 428
Trifolium hybridum, 411
Trifolium repens, 430
Triticum vulgare, 399
Ulmus americana, 404
Uromyces acuminatus Arth., 410, 422
Uromyces fabae (Pers.) DBy., 422
Uromyces fleet ens, Lagh., 430
Uromyces hybridi W. H. Davis, 411
Uromyces hyperici-frondosi (Schw.)
Arth., 406
Uromyces magnatus Arth., 410
Ustilago panici-miliacei (Pers.) Wint,.
410
Uvularia grandiflora, 422
Vaccinium canadense, 427
Veniuria compact a Pk., 408
Verbena bracteosa, 40 4
431e Wisconsin Academy of Sciences , Arts , and Letters.
V ermicularia silphii Schw., 414
Veronica scutellata, 430
Viburnum pubescens, 406, 422
Vicia americana, 418, 419
Vicia caroliniana, 418, 422
Viola, 399
Viola canadensis, 416
Viola conspersa, 421
Viola ocellata, 416
EXPERIMENTAL WORK ON THE RELATION OF SOIL
TEMPERATURE TO DISEASE IN PLANTS
L. R. Jones
Any branch of science in the making must proceed from the
more obvious to the more fundamental, from observation to ex¬
periment. Plant pathology as the youngest offshoot of botanical
science is in the early stage of its evolution. One of its most ob¬
vious aspects is the relation of weather to the occurrence of plant
disease. Every wheat farmer knows from experience that whether
or not his grain rusts depends upon rain and sun, and the po¬
tato grower fears that a wet autumn may rot his tubers. These
evident relations of weather to rust and mildew were discussed
in the Roman classics, but naturally effect and cause have been
much confused even in the mind of the scientist until within a
century. With the establishment of the germ theory of disease
in animal pathology came, simultaneously, the proof of the causal
relation of parasitic organisms, fungi, and later bacteria, to plant
diseases. Since about 1850 botanists have generally accepted the
evidence that the smuts, rusts, mildews, and similar fungi are
genuine parasites and have held them to be true causes of plant
diseases. But the keenly observant though less scientific farmer
continues to blame the weather when his wheat rusts, his grapes
mildew, or his potatoes blight.
The final analysis requires, of course, consideration of both
factors, the parasite as the active cause, the weather as a possible
controlling factor in its operation. The challenge to the phyto¬
pathologist who aims at working his problem to the finish is to de¬
termine by critical experiment the relation of environment to
parasitism. While the fundamental importance of this has long
been recognized by pathologists, and while we have innumerable
observational data, comparatively slow progress has been made
with exact experimentation. This has been due, not to lack of
28— S. A. L. 433
434 Wisconsin Academy of Sciences , Arts , and Letters .
clearly defined interest, but to the difficulties commonly met in the
attempts to control the essential environmental factors with any
degree of precision. The more obvious parasites like the rusts and
mildews are air-borne, attacking the host foliage. Moisture and
temperature are evidently factors profoundly influencing their at¬
tack, but no experimenter has yet succeeded in exactly measuring
these influences separately for the simple reason that when he at¬
tempts precise regulation of any such factor surrounding the func¬
tioning green plant he necessarily disturbs other variables, and thus
almost hopelessly complicates his problem.
Early in our own studies of plant parasites, we became interested
in those invading the underground structures. This was in part
because the hidden diseases had been relatively neglected and be¬
cause when root attacks occur they cause the most serious types of
maladies. In these studies the importance of environmental factors
soon became obvious. Let us consider, as examples, the yellows
disease of cabbage and the root rot of tobacco, each due to a specific
soil fungus which is locally capable of the complete annihilation of
its host. Both parasites persist and possibly multiply indefinitely
in the soil, rendering it permanently ‘ ‘ sick ’ ’ for that special crop,
Yet it is a matter of common experience in southern Wisconsin
fields, where both occur, that the cabbage yellows is serious only
with a hot dry midsummer, whereas with the tobacco the factors
are exactly reversed. Cool, moist weather favors the root rot, and
with a period of dry heat a tobacco crop sick with this disease
quickly recovers. When we met like conditions with potato and
other 4 ‘root diseases”, the question arose whether the fundamental
problems of the relation of environment to parasitism may not be
handled by more simple and direct experimental methods with
these subterranean parasites than with the aerial ones as hereto¬
fore studied. Certain difficulties of manipulation, and compli¬
cations from the light factor inherent with the leafy shoots, are
lessened or disappear in a study of the organs below ground.
Work upon such problems has been carried on in the Department
of Plant Pathology almost uninterruptedly during some six years
past. The first problem, that of the relation of climate to cab¬
bage yellows, was defined during our early study1 of this disease
in which Dr. J. C. Gilman, then a graduate student assistant,
cooperated. Soil temperature was an evident factor, and Gil-
1 See later list of publications concerning this and other matters referred to
above.
Jones — Relation of Soil Temperature to Disease in Plants. 435
man later determined that about 17° C. (62° F.) constitutes a
critical point below which even in the “sickest” soil the disease
does not develop, whereas above this point its seriousness in¬
creases with rising soil temperature. Although his pioneer trials
were made by relatively crude methods, later repetitions with
more exact apparatus have verified his conclusions. Soon after
this, while Dr. W. H. Tisdale as a graduate student worked upon
the flax-wilt problem in the same greenhouses, he observed that
the plants nearest the heating pipes succumbed first, and, fol¬
lowing this lead experimentally, he determined that with this
disease also there is a critical point, about 16°-17° C., below
which its development is inhibited.
The practical significance of such conditions became evident
when field records were secured which showed that in cool years
like 1915 the soil temperature may remain through midsummer
perhaps 5° C. lower than in hot summers like 1916. The difference
year by year in the advent of higher soil temperatures in the
spring and early summer is even more marked. Such results
naturally stimulated the attempt at improved technique and further
investigation into the relation of soil temperature to still other
cases of soil parasitism. It has been recognized from the outset
that an intelligent interpretation of these problems requires con¬
stant recognition of the fact that each such disease results from the
vital interrelations of two organisms, the parasite and the host,
both of which must be affected by variations in soil temperature.
One must also constantly be mindful of the fact that variations
in soil temperature tend to induce concomitant variations in soil
moisture, aeration, and other environing factors. It is necessary,
therefore, to aim either at avoidance of these other possible variables
or at their adequate evaluation if one is not to be misled. To these
ends considerable time was early given to perfecting experimental
methods.
From the outset these studies have been developed in helpful
and mutually stimulating cooperation in which various staff mem¬
bers and graduate students have taken part. In several cases the
results have reached print, in more, the work is still in progress.
The writer therefore acknowledges the generous approval of several
colleagues and students for the use of such material basic to this
article. The chief problems undertaken thus far are listed below,
together with the pertinent publications, so far as these are in
print, which give the details essential to a fuller understanding.
436 Wisconsin Academy of Sciences , Arts, and Letters.
Publications in Order of Appearance
1. Jones, L. R., and Gilman, J. C. The control of cabbage yel¬
lows through disease resistance. Wis. Agr. Exp. Sta. Res.
Bull. 38. 1915.
(This contains an account of this disease, caused by Fusarium
conglutinans , with a general statement of the relation of tempera¬
ture to its prevalence.)
2. Gilman, J. C. Cabbage yellows and the relation of tempera¬
ture to its occurrence. Ann. Mo. Bot. Gard. 3 : 25-84.
1916.
(This gives in detail experimental evidence for the conclusion
that this disease is limited by low soil temperatures, about 17° C.,
based on field and laboratory investigations of which the first part
was made at Wisconsin, the last at the Missouri Botanical Garden.)
3. Tisdale, W. H. Flaxwilt: a study of the nature and inheri¬
tance of wilt resistance. Jour. Agr. Res. 11:573-605.
1917.
4. - . Relation of temperature to the growth and infecting
power of Fusarium lini. Phytopath. 7 : 356-360. 1917.
(Experimental evidence that flax wilt, caused by Fusarium lini ,
is limited in its development by low soil temperature in about the
same way as cabbage yellows. The first simple soil temperature
regulating apparatus described.)
5. Jones, L. R. Soil temperature as a factor in phytopathology.
Plant World 20:229-237. 1917.
(General outline of problems involved, summarizing progress to
date and describing the first model of the Wisconsin soil tempera¬
ture tank with investigational methods.)
6. Johnson, J., and Hartman, R. E. Influence of soil environ¬
ment on the rootrot of tobacco. Jour. Agr. Res. 17: 41-
101. 1919.
(In cooperative relations with the U. S. Department of Agricul¬
ture, Bureau of Plant Industry, the authors undertook an in¬
vestigation of the various soil factors influencing the occurrence
of tobacco root rot caused by Thielavia basicola. They studied the
Jones — Relation of Soil Temperature to Disease in Plants. 437
relation of amount of infestation, moisture, chemical reaction,
temperature and physical and mechanical composition, reaching
the conclusion that of these factors soil temperature is undoubtedly
the most important.)
Work in Progress, Upon which no Publications or Only Pre¬
liminary Notes have Appeared
7. The relation of soil temperature to the development of the
root tubercles of legumes, caused by Bacillus radicicola. F. ft.
Jones and W. B. Tisdale. In progress 1917 to date. Undertaken
in cooperation with the U. S. Department of Agriculture, Bureau of
Plant Industry. Manuscript submitted for publication in an early
number of the Journal of Agricultural Research.
8. The relation of soil temperature and soil moisture to the de¬
velopment of the yellows disease of cabbage, caused by Fusarium
conglutinans. W. B. Tisdale. In progress 1917 to date. See pre¬
liminary note, Phytopath. 10 : 63. 1920.
9. The relation of soil temperature to the cankering of potato
stems, caused by Rhizoctonia (Corticium vagum). B. L. Richards.
In progress 1917-1919. Completed manuscript filed as doctor’s
thesis in the library, University of Wisconsin. Abstract presented
at the meeting of the American Phytopathological Society, Janu¬
ary 2, 1920. Undertaken with partial financial support of the
U. S. Department of Agriculture, Bureau of Plant Industry. In
connection with this, Dr. Richards and Dr. F. R. Jones determined
the relations of soil temperature to the parasitism of this fungus
upon seedling stems of certain other hosts, including bean, pea,
and cotton. (To appear in the Journal of Agricultural Research.)
10. The relation of soil temperature and other factors to the de¬
velopment of the wilt disease of tomato, caused by Fusarium lyco-
persici. E. E. Clayton. In progress 1917-1920. Completed
manuscript filed as doctor’s thesis in the library, University of
Wisconsin. See also preliminary note, Phytopath. 10 : 63. 1920.
The correlation between air temperatures and soil temperatures
has been studied, also the relation of these to variations in soil
moisture. In cooperation with E. C. Sherwood, attention has also
been given to the influence of soil reaction. See thesis for M. S.
438 Wisconsin Academy of Sciences , Arts, and Letters.
degree deposited by Mr. Sherwood in library, University of Wis¬
consin, June, 1920.
11. The relation of soil temperature to the development of onion
smut, TJ'rocystis cepulae. J. C. Walker with the writer. In prog¬
ress 1918-1920. The influence of soil moisture is also being investi¬
gated. (To be published in the Journal of Agricultural Research.)
12. The relation of soil temperature, soil moisture, and other en¬
vironmental factors to infection and development of certain grain
smuts. Mrs. E. T. Bartholomew and Edith K. Seymour. In
progress under the support of the U. S. Department of Agricul¬
ture, Bureau of Plant Industry. 1917 to date.
13. The relation of soil temperature and other environmental
conditions to infection with certain Fusarium diseases of the cereals.
J. G. Dickson and Helen Johann. In progress 1918 to date, the
major part of the support coming from the U. S. Department of
Agriculture, Bureau of Plant Industry.
14. The relation of soil temperature and other factors to the de¬
velopment of the scab of potatoes caused by Actinomyces scabies.
H. H. McKinney in association with the writer. In progress 1918
to date. See preliminary notes, Phytopath. 9 : 301-302, 1919, and
10:63. 1920.
15. The relation of soil temperature and other factors to the
Fusarium wilt of tobacco. James Johnson. In progress 1918 to
date.
16. The relation of soil temperature to the root and stem rot of
peas caused by species of Pythium and Fusarium (previously
noted). F. R. Jones. In progress 1918 to date.
The above list is intended to show at a glance the varied angles
of approach to what is basically one type of problem and also the
personnel of the cooperating investigators. It will also make it
clear that in the limits of such a paper as this it must suffice
merely to outline the research methods proved most useful and to
attempt a correlated summary of the results obtained in a few
typical cases.
Jones — 'Relation of Soil Temperature to Disease in Plants. 439
Apparatus and Methods op Investigation
In all cases attention has been restricted to cases of parasitism
of subterranean parts. The primary object has been to determine
the influence of various soil temperatures upon the development of
disease. At the outset the attempt was made to do this by growing
the plants in ordinary flower pots in greenhouses held at different
temperatures or in favorable places out of doors, recording the
temperatures and observing the results. Although valuable evi¬
dence was thus secured by Gilman in his pioneer work with cabbage
yellows, the method was not sufficiently accurate for continued use.
Text 'Fig. 1. The first and most simple form of constant temperature
apparatus used in the soil temperature experiments: A is a large earth¬
enware jar with a constant flow of tap water entering through the tube D,
regulated by the faucet E. B is a smaller glass jar resting on a support C
and containing soil in which the experimental plants may be grown at
approximately constant soil temperature. F, control culture jar at room
temperature.
The next step was to grow the experimental plants in a battery
jar which, being water tight, could be immersed in a larger jar of
water of which the temperature could be lowered to the desired
point by allowing cold tap water to trickle through it at a con¬
trollable rate (text fig. 1). This sufficed to give W. H. Tisdale
more accurate data relative to the influence of the lower range of
temperatures upon the flax-wilt disease. When it was proposed
to push the work more aggressively with a larger series of plants
440 Wisconsin Academy of Sciences , Arts , and Letters.
and to include higher temperatures, the more elaborate plan which
has since been followed was quickly evolved. It consists in having
a series of water compartments or tanks in each of which the water
can be held fairly constantly at any desired temperature. The
Text Fig. 2. The second stage in the development of the soil-tempera¬
ture-control tanks. This was installed on a greenhouse bench over the
regular steam heating pipes (G) which served to supplement other heat¬
ing devices such as live steam (F) and electric heating bulbs ( E ). The
lower temperatures were secured by cold running water. Insulation of
the tank was obtained with hair felt (L) between the galvanized iron
lining (M) and board (K), and an asbestos board cover (D). Tempera¬
tures from 5°C. to 40 °C. were obtainable and could be held fairly con¬
stant. In each compartment were sunk four battery jars filled with soil
for the culture of the experimental plants.
experimental plants are grown in glass or metal containers which
can be sunk in these water tanks so as to maintain the soil
temperature desired for the period of trial. To begin with, it
seemed best to try to maintain the graduated temperatures by cir¬
culating the water through the compartments from the cooler to the
warmer until the house temperature was reached. The development
of this idea was undertaken by R. E. Hartman working in associa¬
tion with James Johnson on the tobacco root-rot problem. The
compartment tank evolved, as illustrated in text figure 2, served its
purpose well and has continued in use since 1915. It soon became
Jones — Relation of Soil Temperature to Disease in Plants. 441
evident, however, that more satisfactory results could he secured
by the use of small unit tanks instead of those united in a perma¬
nent series (text fig. 3). These units, which are termed the 11 "Wis¬
consin temperature tanks’’, can then be assembled in any number
Text Fig. ,3. Section of the improved Wisconsin temperature tank,
showing construction and arrangement of apparatus. The soil cans (ca.)
are suspended in the water from the rigid cover (co.). Water for cooling
and filling the tank is run in through water inlet (wa.i.). The alternating
110-volt heating current is carried through heavy wires (he.c.) from
switch (sw.) to relay (re.) and thence to the heating unit (he.u.) en¬
closed in the water-tight copper tube (he.) The direct current operating
the relay is carried from storage batteries through line (ba.c.) from
thermostat (th.) to the magnet on the relay (re.).
or arrangement desired to meet the needs of a particular experi¬
ment. Naturally much attention has been given to the various
possible methods of maintaining a constant water temperature and
also to the size and character of the pots or containers in which
the experimental plants are grown. This has also included the
matter of special methods for securing uniformity in soil moisture
and aeration, as well as devices for covering or insulating the free
442 Wisconsin Academy of Sciences , Arts, and Letters.
surface of the soil about the plants in order to maintain the same
temperature in the surface layer as in the deeper portions. Ex¬
perience has shown that no single method is best for all purposes,
the details being varied to meet particular needs. The main feat¬
ures are illustrated in text figures 2 and 3, and the essential details
are given in the descriptions of these figures. By these methods
the various soil temperatures desired have been maintained with a
sufficient degree of constancy (in general within l°-2°) to meet
our experimental needs while giving the plants otherwise favorable
environmental conditions. In most of the series run the tempera¬
ture range has been as follows, including at the extremes the cold¬
est and the warmest soil temperatures at which the various experi¬
mental host plants will grow satisfactorily : 12°, 15°, 18°, 21°, 24°,
27°, 30°, 33° C. (approximately equivalent to 53°, 58°, 64°, 70°,
75°, 80°, 86°, 91° F.)
Of course, different hosts respond differently as also do the vari¬
ous parasites. Hence temperatures both higher and lower than
any in the series above given have been used in certain cases. It
is to be noted that in all the trials, at least in the initial experi¬
ments, the aerial parts of the plants in each series have developed
at the same air temperatures. The aim has been so to control this
as to secure the best normal host development. This means, for
example, that the experimental tomato and tobacco plants have been
carried in a relatively warm greenhouse with air temperature dur¬
ing the day approximating 27°-30° C. (80°-85° F.) and falling
several degrees lower at night. Potatoes, peas, etc., have been
grown in a relatively cool house where the air temperature has
ranged some 10°-20° F. below that of the warmer house. It is
clearly recognized, of course, that some correlation between soil
and air temperature is essential for the normal plant functions, and
likewise that the degree of soil moisture may have a relation to the
influence of temperature. Attempts have been made in connection
with the work with certain diseases to determine in some degree
these interrelations. While it does not seem wise to introduce the
details in the present paper, it should be noted that the results are
such as to restrain one from too hasty and broad generalizations
concerning the influence of soil temperature as an isolated factor.
As a matter of fact, in nature no one factor varies alone. The
totality of evidence, however, leaves us confident that for certain
types of soil parasitism, and within the usual limits of environ-
Jones — Relation of Soil Temperature to Disease in Plants. 443
mental variation, soil temperature is so dominant a factor as to
deserve the first consideration that we have given it in the present
investigations. Work with the other variable factors should then
follow in natural sequence, and finally some correlations should be
attempted.
Some Typical Results
It must suffice now to select for summary the results from three
such types as will best show the general nature of the problems
met and the evidence secured in these studies concerning the rela¬
tion of soil temperature to disease development. For the present
purposes it seems best to discuss a typical disease on each of three
host plants belonging to one family, the Solanaceae. Two of these,
the tomato and the potato, are so closely related botanically that the
one may be grafted upon the other, yet they differ markedly in
their temperature relations, the potato being favored by a cool
climate and the tomato thriving best at higher temperatures. The
tobacco, which is the third host selected, is also favored by rela¬
tively high temperatures. All of these naturally show great vege¬
tative vigor when free from disease, but all are subject to the at¬
tack of one or another subterranean parasite capable of rapidly
causing serious disease or death. The parasites selected represent
quite different temperature relations and modes of attack. It will
be best, therefore, to consider each of these diseases separately and
then to make certain comparisons.
The Fusarium Wilt Disease of the Tomato
This disease is caused by Fusarium lycopersici , of the mycological
group known as the Imperfect Fungi. It is widespread throughout
the warmer tomato-growing regions of the United States, but
causes little or no trouble in the northern or cooler districts of
commercial tomato culture. Thus, in the eastern Mississippi Val¬
ley states it is highly destructive from Louisiana north through
Illinois and Indiana, but is so rarely observed that it has not be¬
come a recognizable factor in commercial tomato culture even on
the relatively warm trucking soils where tomatoes are so much
grown in southeastern and southwestern Wisconsin. These ob¬
servations, together with the discovery early in this work that the
like Fusarium disease of both cabbage and flax is limited by soil
444 Wisconsin Academy of Sciences , Arts, and Letters .
temperature, led to the clear definition of the question whether or
not soil temperature may be the limiting factor with this tomato-
wilt disease. Dr. E. E. Clayton, as a graduate student, undertook
to determine the facts, and the following summary is taken from
his full report, which is now awaiting publication. To begin with,
he made trials by the agar-plate pure-culture method to determine
the influence of temperature upon the growth of the fungus. The
results, which are shown in Plate XXXIII, figure 3, made it clear
that the parasite is rather sharply inhibited in its development at
both low and high temperatures. Trial culture of the host in non-
inf ested soil held at different temperatures,, the air temperature
being the same for all, showed that the tomato plant has a somewhat
wider range of temperature toleration than has the Fusarium
(PI. XXXIII, fig. 1). It is noteworthy, however, that the optimum
temperature alike for the development of the fungus and of the
host under the conditions of these trials is practically the same,
viz., about 28° C. It is of especial interest now to learn the re¬
sults secured by Clayton when tomato plants were grown in
Fusarium-infested soil held experimentally at different tempera¬
tures ranging from the lower to the higher limits of the heat toler¬
ance of tomato roots. These are illustrated by photograph in Plate
XXXIII, figure 2.
From these data it is evident that the development of the dis¬
ease is definitely and quite sharply conditioned upon favorable
temperature. It is noteworthy also that the soil temperature which
in general seems most favorable for both the tomato and its parasite,
28° C., is approximately that at which the disease appears first
and proceeds most rapidly to the destruction of the host plant. It
is of further interest to note that, although the lower limit or
critical point for the inception of the disease, 19°-20° C., is nearly
the same as for the two related Fusarium diseases of cabbage and
of flax, it is actually sufficiently higher (cabbage and flax be¬
ing about 16°-17° C.) to correspond in a general way with the
difference in temperature relations of the respective hosts, the
tomato being favored by distinctly higher temperatures than are
cabbage and flax. In this connection we may recall that the temper-
ture relations of the three Fusarium parasites are practically
identical. Quite as striking as the sharp delimitation of the dis¬
ease by soil temperature at the lower point is that at the upper
(Plate XXXIII, fig. 2), where it seems that the tomato can live at
soil temperatures well beyond the range of attack of the parasite.
Jones — Relation of Soil Temperature to Disease in Plants. 445
In this respect again the temperature range for development of
the disease is well correlated with that for the growth of the
parasite. In other words, the disease behaves about as one would
anticipate, having knowledge of the temperature ranges of host
and parasite, since, although they have their optimum at ap¬
proximately the same point, the tomato has a somewhat wider
range of temperature tolerance than has its parasite.
The Potato Stem Canker
There is no plant of which the diseases have been more in¬
tensively studied in the last generation than the potato. This is
partly because it is of such widespread economic interest and
partly because potato disease problems are so serious. Yet upon
some of the most common and obvious of these there have been
for years differences of judgment as to the nature and cause.
This has been especially the case with the cankering of the basal
stem and the scurfing of tubers associated with the fungus Rhizoc-
tonia, which in its perfect stage is known as Corticium vagum.
This fungus develops conspicuous resting masses (sclerotia) on
the surface of the tubers in late autumn, aptly described as the
“dirt that won’t wash off” (Pi. XXXI Y, fig. 1).
These sclerotia are, however, purely superficial. Examination
of the surface of the freshly dug tuber under a magnifying glass
shows that they arise from the massed growth of the minute
thread-like strands (hyphae) of the fungus, which ramify in
abundance over the outside of the tuber, but here again always
strictly on the surface and ordinarily causing no apparent harm
to the host. All pathologists are agreed that with such tubers
the Rhizoctonia exists practically as an epiphyte like the moss
or lichens on the sound bark of a tree. If tubers bearing such
sclerotia are planted, the pathological problem begins. The rest¬
ing fungous mass promptly originates an abundant new growth
which creeps over the surface of the tender young potato sprouts
and often finds an especially favorable nidus in the reentrant
angle of the apex of this shoot, which is recurved, or hook-like,
before it breaks through the surface of the soil. The complica¬
tions for the pathologist arise from the fact that, whereas in some
cases the Rhizoctonia apparently behaves as a cankering parasite
(Pl. XXXIV, fig. 2), killing off so many of the shoots before they
emerge as to cause a serious loss of stand in the potato field
446 Wisconsin Academy of Sciences , Arts , and Letters.
(PL XXXI Y, fig. 3), in other cases little or no such harmful
action occurs. The situation later in the season becomes even
more puzzling, since the Rhizoetonia may in late summer develop
as a purely surface growth enveloping the basal part of the
potato stem with no sign of lesion or canker. This fact has led
some pathologists to question whether any of these stem cankers
are due to Rhizoetonia, or whether instead they may not orig¬
inate from other causes, with the Rhizoetonia development as a
purely secondary phenomenon. Our own theory for some time
past has been that the parasitism of the Rhizoetonia must be in
some degree dependent upon environmental factors, and early in
the work with the temperature tanks it was planned to include
studies upon the possible influence of soil temperature. This un¬
dertaking was first projected when R. E. Hartman as a graduate
student worked upon the organism in 1916-1917. His work was
interrupted by war service before greenhouse trials along this
line were inaugurated; but his experience with field plots in 1916
was significant in that, even where he planted seed potatoes
abundantly covered with sclerotia, little or no sprout killing or
stem canker resulted.
Following this, B. L. Richards as a graduate student and re¬
search assistant in this department2 prosecuted these trials dili¬
gently for about two years, presenting the results as his doctor’s
thesis, which was filed in the University Library, October, 1919.
This has not as yet been published,3 but the writer is authorized by
Dr. Richards to cite here certain of his results. In the first place,
the evidence secured was fully convincing that the fungus Rhizoe¬
tonia (Corticium vagum ) is the direct cause of the stem cankering
of the potato, and, conversely, that adequate seed disinfection gen¬
erally serves as a practical preventive for this trouble. The de¬
velopment of the stem cankers is, however, dependent not alone
upon the presence of the fungus but upon favoring environmental
2 Mr. Richards as plant pathologist at the Utah Experiment Station had
worked considerably with the Rhizoetonia disease of the potato before coming
to Wisconsin. In order to enable him to push the work more vigorously
while here, especially in the field during the summer, he was appointed col¬
laborator with Dr. W. A. Orton of the Office of Truck, Cotton, and Forage
Crop Disease Investigations of the U. S. Department of Agriculture.
3 This manuscript is entitled “Soil Temperature as a Factor in Determining
the Pathogenicity of Corticium vagum with Special Reference to the Potato.”
Dr. Richards has returned to his former relations as plant pathologist of the
Utah Agricultural Experiment Station, and the publication of his detailed
results may be expected soon as a bulletin of the Utah Station or in the Journal
of Agricultural Research.
Jones - — Relation of Soil Temperature to Disease in Plants. 447
conditions. Time did not permit Dr. Richards to make a complete
determination of these conditions, including the possible rela¬
tions of soil moisture, aeration, and other variable factors aside
from temperature. He did, however, secure fully convincing
evidence that soil temperature has such a highly important influ¬
ence that under most conditions it would seem to be the domi¬
nant thing in determining the pathogenic results. The details
must be left for Dr. Richards’ full account, but the convincing
nature of this evidence may be judged from two examples.
Potatoes were grown experimentally in a series of like pots
of soil which were held at soil temperatures graduated from 9°C.
to 30°C., the air temperature being for all alike about 18°-22°C.
It was found that if fungus-free seed potatoes were used, the
plants grew at all these temperatures except the very highest,
but the best normal development was at about 18 °C. At lower
soil temperatures their growth was distinctly retarded, and at
higher, although the aerial stem elongation was more rapid, the
leaves were smaller and abnormal and tended to roll. In case
the seed tubers bore sclerotia the fungus developed to some de¬
gree at all temperatures, but the serious stem cankering oc¬
curred at 15°-21°C. with a distinct and rapid lessening of the
diseased condition at higher temperatures.
Evidence of the same import was also secured by Dr. Rich¬
ards in a second way. This was by so varying the date of
planting as to expose the potato plants to different soil tempera¬
tures during the early stages of development. The first plant¬
ing was made April 26 and later ones were made in May and
June, the soil being uniformly inoculated in all trials. Fortu¬
nately for the purposes of this trial the surface soil temperature
remained cool (most of the time below 15 °C. at a depth of 4
inches) until about May 25, when the weather turned suddenly
warmer and the soil temperature at once rose several degrees,
mostly to 22° and above in the upper four inches, and remained
uniformly warmer thereafter. The outcome as to type and de¬
gree of injury to the young sprouts was marked and convinc¬
ing. The tips of the primary shoots of the early planted series
growing in cold soil were badly attacked by the Rhizoctonia and
in many cases they were killed below ground. On the other
hand, with the late planted tubers the sprouts developing at
higher soil temperatures were almost free from this destruction
of growing point. Even more striking, however, was the fact
448 Wisconsin Academy of Sciences, Arts , and Letters .
that with the first planted series, in which the tips of the primary
shoots in the cold soil were cut off or killed by the canker, the
secondary sprouts arising from the bases of these same stems later,
when the soil was warm, developed almost uninjured. (See
Plate XXX Y.)
The influence of soil temperature on this potato canker is,
therefore, as clearly evident as with the tomato wilt but in an
entirely different way. With the tomato disease a soil temper¬
ature below 19 °C. inhibited the attack, and 27 °C. was most fa¬
vorable. With the potato disease at 27 °C. the host is practically
uninjured, whereas the worst attacks occur below 19 °C. If one
seeks the explanation it is natural to recall that the tomato
fungus (Fusarium) developed most aggressively in culture at
about 27° C. and to suspect that the potato fungus (Rhizoctonia) is
favored by the lower temperature. As a matter of fact, however,
although the potato fungus is not as sensitive as the tomato para¬
site to temperature fluctuations, it too grows fastest at the higher
temperatures (24°-26°C.). For adequate explanation of the
escape of the potato shoots at the higher temperature we are,
therefore, forced to turn to one or both of two other theories. The
first was suggested some years ago by the British botanist, Balls4,
for a similar if not identical disease of cotton ; viz., that the fungus
loses its pathogenicity at the higher temperatures as a result of
some form of self -poisoning if the food supply is limited or of
improper character, a condition which may occur in the cells un¬
der these conditions. Richards prefers to consider that it may be
due to inability, at the higher temperatures, to develop certain
enzyms or toxins essential to its parasitism. He has, however,
shown that the more rapid growth of the potato shoots at the
higher temperature offers the important reason and perhaps the
chief one for their escape. Thus in his trials he found that with
rather dormant potato tubers planted 5 inches deep the shoots broke
ground in 18 to 25 days at 24 °C., whereas they were delayed
nearly two weeks longer at 9° to 12°C. In view of the fact that
the chief injury by the fungus results from a rather slow attack
upon the growing tips of the shoots while yet below ground, he
finds evidence for the conclusion that at a temperature of about
21 °C. or above the shoot is able to grow away from the pest in
4 Balls, W. L. The physiology of a simple parasite. Yearbook Khedivial
Agr. Soc. Cairo 1905:184.
Jones — Relation of Soil Temperature to Disease in Plants. 449
these early stages and thus permanently to evade injurious at¬
tack, whereas in colder soils the shoot develops so slowly that
the fungus gets in its deadly work before the tip can escape
through the surface.
The Tobacco Root Rot
Attention may now be directed to the results with the tobacco
root-rot disease. This is caused by the parasitic soil fungus
Thielavia basicola, which is very widespread in its occurrence
and attacks the roots of various plants besides tobacco. Trouble
with this root rot is frequent in the older tobacco sections of
this country, and in southern Wisconsin it is the most serious
tobacco disease. A noteworthy thing about it is that even on
old tobacco soils where the fungus is almost omnipresent the
severity of its attacks varies extremely from season to season.
The farmers might attribute these variations simply to weather
but for the fact that on new soil they do not occur. On virgin
fields a sound crop develops each season regardless of weather,
the failures occurring under certain conditions on the old to¬
bacco soils. Since tobacco is reputed to be a hard crop on the
soil, the growers have naturally attributed their failures to soil
worn out through exhaustive cropping. They have been faced,
however, with the puzzling experience that in certain seasons
the crop starting well would later fail, or, conversely, that when
making a poor promise during June and July, it might suddenly
mend in August and mature a profitable yield in September.
Clearly, the explanation of these anomalies required expert at¬
tention.
Dr. James Johnson, who has been working upon Wisconsin to¬
bacco problems for over a decade, was stimulated by the early
results with cabbage yellows to attempt critical analysis of the
relation of the various environmental factors including soil tem¬
perature to the development of root rot. In this work he had
the cooperation of R. E. Hartman, a graduate student assistant.
The results of these studies, which they have recently published,
show conclusively that while soil moisture and other factors play
a part, soil temperature remains the deciding factor, in most
cases clearly determining the severity of the root rot. Trials
made in the Wisconsin soil temperature tanks have given uni¬
form and convincing evidence as follows : At the lower tem-
29— S. A. L.
450 Wisconsin Academy of Sciences, Arts, and Letters.
peratures the disease occurs and destroys the plants at any and
all temperatures down to the lowest limit at which the tobacco
can grow, viz., about 10° C., showing that at the lower tempera¬
ture the range of parasitic activity for the fungus goes quite as
low as that for the growth of tobacco. The tobacco is favored
by warm soils, and as the soil temperature rises, if in soil free from
Thielavia, the plants show an increasingly vigorous stem and leaf
development up to about 26 °— 30 °C. The optimum for root devel¬
opment occurs a little lower, at 20°-25°C. (PI. XXXVI, fig. 2).
On the other hand, the disease is most serious at about 17 °— 18° C.,
and as the temperature rises the evidence of its injuries is gradu¬
ally lessened until at 26 °C. it ceases to be a noticeable factor in
checking the host plant (PI. XXXVII), while at 30° or above no
further lesions appear on the roots. The tobacco, on the other
hand, with its optimum development at 26 °— 30° and capable of
enduring soil temperatures at least ten degrees higher, makes a
vigorous development at soil temperatures of 28 °C. or above even
in the “ sickest” soil.
These experiments furnish an adequate explanation of the
farmer’s experience in the variations in this disease with soil and
season. In the first place, they show that the direct and sole
cause is the parasitic soil fungus Thielavia. If this fungus is
present in the seed beds, it attacks the roots of the seedlings in the
earliest stages of their growth because of its ability to develop at
the lowest temperatures at which tobacco will start. In old tobacco
fields it is also, as a rule, present in the soils ready to attack the
roots immediately following transplantation, but even with the
newer soils it is very likely to be carried to the fields from the seed
bed on the roots of some of the transplants. It thus gets a foot¬
hold and tends to increase year by year. Soil temperature is low
enough every season for it to make at least some development,
once started in the field, and with a dominance of weather suffi¬
ciently cool to hold the soil temperature below 25° C. during July
and August, it continues to rot off the new roots about as fast as
they originate, so that through midsummer the plants remain sta¬
tionary even if they are not killed outright. When, on the other
hand, the summer continues hot during July and August, the roots
escape disease even in the “sickest” old tobacco fields. The more
usual thing under Wisconsin climatic conditions is to have relatively
cool weather during the first part of the summer with a dry, hot
period during late July or August. When this occurs the tobacco
Jones — Relation of Soil Temperature to Disease in Plants. 451
on old infested fields, no matter how well fertilized and tilled, will
remain stunted by the root rot during the earlier cool weather. With
advent in midsummer of the higher soil temperatures, the root rot
is suddenly checked, and as by magic the plants spring forward
and produce a surprisingly good crop in September. The correct¬
ness of this interpretation and the details of how the temperature
change operates were clearly evidenced by the following experi¬
ments, which Johnson has repeated with a number of plants.
Plants have been grown in the temperature tanks for a period at
the lower temperature, say 18 °C., where the root rot develops so
badly that the plants barely live, then the same pot is transferred
to a higher temperature tank, say at 28°, when the plants quickly
recover. Again Johnson has taken up stunted, sickly plants from
a badly infested field in late summer and placed them in pots so
that the soil temperature might be experimentally controlled. One
half of these have been held at relatively low soil temperature,
about 20 °C., and the other half at a high temperature, about 30 °C.
The plants had practically no roots when set into the temperature
tanks, only the blackened base of the stem remaining. At the
higher temperature new roots promptly developed which remained
white and healthy at the end of the experiment one month later,
whereas at the lower temperature, although new roots continued to
be formed, they survived barely long enough to keep the plant alive
without growth. When at the end of the experiment the root sys¬
tems were washed out for examination, the vigorous healthy roots
which had grown at the higher temperature, in contrast with the
blackened diseased root which had developed at the lower tempera¬
ture, showed strikingly the dominant influence upon tobacco root
rot of a change of only a few degrees in soil temperature (PI.
XXXVI, fig. 1).
Equally conclusive evidence of these temperature relations to
crop production has been secured in a more direct manner. By
means of soil temperature readings taken daily at Madison during
the growing period, it was shown, for instance, that the soil tem¬
peratures in 1915 were relatively very low (average for June, July,
and August 20.3 °C.) as compared with 1916 when the average for
the same period was 27.7° C., or 7.4° C. higher, a difference which
it may be noted almost bridges the gap between the temperature
most favorable for the Thielavia disease and one at which the para¬
site could not function. Correlated field records show that in 1915
452 Wisconsin Academy of Sciences , Arts, and Letters .
Text \Fig. 4. Soil temperatures for July, 1915, and July, 1916, at Mad¬
ison, Wisconsin, at depths of 2, 4, and 8 inches. The days of the month
are indicated by the figures across the top.
the loss from Thielavia root rot in Wisconsin alone ran into millions
of dollars whereas in 1916 relatively unimportant losses occurred.
The evidence seems conclusive as satisfactorily explaining the
practical aspects of the problem. But as a matter of scientific in¬
terest we must persistently ask, “Why?” Why are the tobaco
roots rotted at the lower temperature while they escape at the
higher in soil equally infested? We naturally turn to our experi¬
ences with the other two diseases. In the case of the Fusarium
wilt of the tomato, in which case the disease increased with rise of
temperature to a certain point, we found a possible explanation in
the fact that the optimum soil temperature for the development of
Jones — Relation of Soil Temperature to Disease in Plants. 453
the disease coincides with the optimum for the development of the
fungus. The stimulating effect of the favorable temperature upon
the parasite seems to be the determining factor with tomato wilt.
In the case of the Rhizoctonia canker of the potato, a disease which
is worse at the lower temperature, the condition has been inter¬
preted as resulting from the fact that the lower temperature so re¬
tards the development of the young potato sprouts as to give the
parasite a longer period for its destructive underground action on
the young shoot tips, whereas at the higher temperatures the rapid¬
ity of their growth enables them to escape relatively unharmed.
The retarding effect of the low temperature upon the host seems to
be the most important factor in determining the outcome with
potato stem canker.
Let us now recall the results with the tobacco root rot. Here,
as with the potato canker, the disease is worse at the lower tem¬
perature and is checked at the higher. When we seek the explana¬
tion, however, the facts do not accord with those in either of the
two previous cases. In the first place, the tobacco fungus in pure
culture trials is found to behave almost exactly like the Fusarium,
growing distinctly faster at 25 °C. or above and lessening in growth
rate as the temperature falls. The worst development of the
disease occurs, therefore, at a temperature which seems less favor¬
able for the parasite, whereas the disease disappears at the soil
temperature most favoring growth of the fungus. If now we turn
to the temperature relations of the tobacco root (PI. XXXVI, fig.
2), the case becomes the more puzzling since the most vigorous
root development is found not at the higher temperature where the
root growth is more rapid. Plainly here some explanation must
be sought other than the direct influence of temperature upon the
relative vigor or growth rate of either parasite or host. It is, of
course, easy to make conjectures involving other possible variable
factors which may be influenced by temperature, such as enzym
or toxin development by the parasite or some products or activities
of the host tissue which may contribute to its resistance. We
have already referred to the interesting suggestion by Balls of the
possibility of self-inhibition of the parasitic Rhizoctonia through
auto-intoxication at certain temperatures. However one may in¬
cline toward one or another such possible theory, the truth of the
matter is that we do not in any case know enough as yet about the
intimate relations of parasite and host to justify the formulation
454 Wisconsin Academy of Sciences , Arts , and Letters.
of fully satisfactory explanations. With facts as definite and as
clearly established as are these, however, we may hope that such
explanation may follow in due season.
Discussion and Conclusions
The results here presented have been selected primarily to ex¬
plain what is essentially a new method of attack upon a group of
phytopathological problems. The aim, broadly stated, is to gain a
clearer understanding of the relation of environment to the occur¬
rence of disease in plants. It must be accepted as fundamental
that environment is influential with all types of disease, and in
each case the occurrence of a parasitic disease must evidently be
dependent upon numerous factors. In our attempt to put the
work upon an experimental basis, we have selected, because of
simplicity of definition and practicability of attack, the relation of
soil temperature to certain cases of soil parasitism. The method
which has served well for initial studies has aimed at varying soil
temperature while holding other factors including air temperature
constant. Of course, it is recognized that this cannot give us all
the data necessary to either the full explanation of such variations
as occur in nature or the adequate interpretation of the biological
principles involved. In either case, we must deal with the result¬
ants of a series of variables such as soil moisture and aeration, air
temperature and light, transpiration and nutrition, which rarely
operate singly or simply. Moreover, it is not to be forgotten that
each disease as a biological phenomenon is itself always a resultant
of the vital interrelations of two active organisms, engaged it may
be in a struggle for existence. These relations of parasite and host
are to be conceived of not as static or even as necessarily progres¬
sive in one constant direction, but rather as varying from period
to period. Under these circumstances, one should not only ap¬
proach the problems with due respect for their biological complex¬
ity but perhaps should even present an apology for his temerity in
grappling with them at all. The reason that seems to the writer
adequately to justify an earnest and persistent attempt at least
more clearly to survey the problems and to hack away at their
most protruding angles, is that as a plant pathologist one cannot
intelligently discuss a single disease, whether with plant cultivator
or scientific associate, without consideration of the relation of these
environmental factors to disease development. We cannot dodge
Jones — Relation of Soil Temperature to Disease in Plants. 455
the issues as plant pathologists; we ought not to wish so to do as
biologists. The knottier the problems the clearer the challenge.
We have purposely selected samples where the evidence is relatively
Simple and direct and have aimed to show that it is convincing at
least upon certain points. We have not done this with the inten¬
tion of misleading as to the inadequateness of the progress already
made. Each step has made clearer the complexities of the maze
we are treading. The purpose has been, however, to show, first,
the worth-whileness of tackling the problems, and, secondly, that
worth-while progress is possible by the methods outlined if the at¬
tack is earnest and persistent.
On the other hand, any plant pathologist should wisely hesitate
to undertake work upon such a complex of problems with any
hope of going far working singlehandedly or alone from the path¬
ologist’s viewpoint. As was explained at the outset, much of the
progress made in the work here summarized has been due to the
happy correlation of the efforts of a number of graduate students
as supplementing and greatly extending what any of the depart¬
mental staff members alone could have accomplished. But num¬
bers alone will not get us far if all work at the same angle. Im¬
mediately as the problems take form, it is evident that at bottom
we are dealing with basic questions of the physiology of the host
plant on the one hand and of that of the parasitic fungus on the
other. Here, again, our courage as plant pathologists has been
kept up because of our confidence in our physiologically minded
associates as advisers or co-workers.
We must not, however, assume that by so defining these prob¬
lems as basically physiological we thereby transfer to the already
overburdened plant physiologists the responsibility for the imme¬
diate prosecution of their investigation. If any men specially
trained in plant physiology can be induced to take up work in this
field, it will be immediately to the advantage of pathology. On the
other hand, it cannot be expected that much help will come
through distracting mature physiological investigators from their
own chosen lines of interest. It is, however, evident that this
work thus far has already defined problems which are distinctly
physiological rather than pathological and which have such broadly
attractive scientific interest that we may safely await their further
development by plant physiologists aided in some cases by workers
in other lines of applied botany.
456 Wisconsin Academy of Sciences , Arts , and Letters.
This paper is not the place for details other than for purposes of
illustration. Suffice it, therefore, to cite here a few such problems
which are essentially physiological. (1) The relations already
mentioned of soil temperature to the development and functioning
of the root tubercles of legumes, a matter of keen interest alike
to physiologist, bacteriologist, and plant culturist. (2) The rela¬
tion of soil temperature to the rate and character of root develop¬
ment. (3) The correlation between soil temperature and air
temperature in affecting the extent and type of development of
root and shoot organs. (4) The relation of soil and air tempera¬
tures to the nutritive and reproductive processes of plants, and
consequently the proper correlation of these with the natural se¬
quences in the plant’s development. (5) The occurrence and pos¬
sible significance of bimodality in the growth curves of shoots and
roots, there being evidence that as soil temperature is raised there
occurs in certain cases a check in the rising curve of growth at a
certain point (e. g., it may be about 26°-28° C.), followed by a
subsequent rise. These are not definitely phytopathological prob¬
lems, nor are pathologists, as a rule, well qualified to work them
out fundamentally. "While the phytopathologist is interested in
the outcome in each case, he may well await patiently its develop¬
ment in natural course by ecological physiologists or others inter¬
ested in the relation of environment to the normal plant.
There are, however, certain distinctly pathological problems for
the development of which we cannot so wait for the help of others.
Thus, on the one hand, we have the questions of the evident and
immediate influence of soil temperature and other environmental
factors on the occurrence, severity, and geographical distribution
of certain aggressive introduced parasites. The necessity for im¬
mediate information along this line has been strikingly evidenced
in recent years by the introduction into the northeastern states of
the European parasites causing the powdery scab and black wart
diseases of the potato. It is at once evident that our judgment as
to how serious these are to be and what measures should be taken
for their control must be influenced by what we know about the
relation of our American environment to their aggressiveness.
Other questions of the same sort come out of the recent introduc¬
tion of the Asiatic citrus canker into the southeastern states, the
report last year of something resembling the Australian ‘ ‘ take-all ’ ’
disease of wheat in the Mississippi valley, and the possibility of
serious Indian corn mildews coming from the East Indies. It is
Jones — Relation of Soil Temperature to Disease in Plants. 457
evident in each case that information is needed in order the better
to meet immediate economic situations, but more fundamental than
this is the fact that the plant pathologist cannot even think, much
less act, clearly and intelligently about any of these matters except
as he takes into consideration the influence of environmental fac¬
tors.
Another class of problems in connection with which the plant
pathologists cannot await the help which would come too slowly
and indirectly from outside their ranks deals with the relation of
environment to disease resistance. We are in the habit just now
of saying that the problems of disease resistance or of relative sus¬
ceptibility and immunity are the most vital problems of the day
in phytopathology, whether we think of them in their practical
significance or in their fundamental relations. But, in how far
are these things clearly defined or fixed entities, in how far are
they variables, and what may be the relation of environment to
such possible variations? We do not wish to imply that we are
not, in true disease resistance, dealing with definite and inheritable
factors; the evidence is conclusive that we are. But there can be
no doubt that environment may determine to what degree such
factors dominate or find expression. If, therefore, we are to think
or talk clearly and consistently about disease resistance in a plant
we must have an adequate understanding of the influence of envi¬
ronment upon the development of the disease in question.
For all these reasons we are earnest in the conviction that plant
pathologists must continue to inquire with increasing precision
into the relation of environment to disease development. Not
only must they give to these problems some of their own best ef¬
forts, but they must seek the assistance of special workers in the
related fields, physiologists, ecologists, geneticists, and plant cul-
turists. Working thus in a spirit of correlation and cooperation,
prompt and important progress is assured.
Department of Plant Pathology,
University of Wisconsin
458 Wisconsin Academy of Sciences, Arts , and Let Vers.
Explanation of Plates
plate XXXIXI
Tomato plants and cultures of the tomato- wilt fungus, showing the
effect of different temperatures. (Photographs by E. E. Clayton.)
Pig. 1. Normal tomato plants grown in sterile soil at temperatures of
19°, 22° , 24°, 28°, 31°, 33°, and 35° iC. This shows the effect of these
various temperatures on the development of the tomato plant, the op¬
timum being at about 28°-31°.
Pig. 2, Tomato plants grown at temperatures of 19°, 22°, 24°, 28°,
31°, 33°, and 35° C., in soil infested with the tomato-wilt fungus. Note
that the plants escape the disease at the two extremes, whereas the wilt
appears first and develops most severely at 28°-31°.
Pig. 3. Petri dish cultures of the tomato-wilt fungus showing growth
at temperatures of 8°-38° C. The maximum temperature for the develop¬
ment of the fungus is between 35° and 3'8°, the optimum about 28°, and
the minimum between 8° and 10°. Note the close correlation between
relative vigor of growth of the fungus at these temperatures and the
development of the disease as shown in figure 2.
PLATE XXXIV
The Rhizoctonia disease of potato as shown on the tuber, on the
young stems, and in the field. (Photographs by B. »L. Richards.)
Fig. 1. “Black scurf’ on the potato tuber, this being the dormant stage
of Rhizoctonia. The fungus produces numerous black sclerotia on the
tuber, which are usually strictly superficial and epiphytic. If such a
tuber is planted under certain environmental conditions, the fungus de¬
velops as a serious parasite on the young shoots. (See figure 2.)
Pig. 2. Rhizoctonia injury, or “stem canker,” as it appears on the
young potato sprouts. It kills many of these before they can reach the
surface of the soil and weakens those which survive. (See figure 3.) As
explained in the text, the degree of this injury is largely conditional on
soil temperature. (Compare with Plate XXXY.)
Fig. 3. Experimental potato field, showing the seriousness of Rhizoc¬
tonia damage. The weak plants, row A, grew from untreated, scurfy seed
tubers. Note that many hills are missing. The strong rows on either
side (B, etc.) grew from treated seed tubers, the treatment practically
eliminating Rhizoctonia injury.
Jones — Relation of Soil Temperature to Disease in Plants. 459
PLATE XXXV
Plants which illustrate the relation of soil temperature to the Rhizoc-
tonia injury of potato stems. The primary shoots developing in very
early spring, when the soil was cool (about 15 °C.), were cut off by the
fungus. A little later the soil temperature rose to about 22°C., and the
secondary shoots then developing escaped injury. (Photographs by B. L.
Richards.)
PLATE XXXVI
Normal and diseased tobacco plants showing the effects of different soil
temperatures. (Photographs by James Johnson.)
Fig. 1. These two plants were stunted alike during an entire season
by disease, and were then removed from the field and placed in the
temperature control tanks. The plant placed at a soil temperature of
30OiC. started to make a rapid recovery, whereas the one at 20°C. re¬
mained diseased. This is in accord with the field experience that a low
soil temperature, which is favorable for the Thielavia root-rot disease,
if followed by a high soil temperature, enables the tobacco plant to
recover by sending out new roots which are not affected.
Pig. 2. Root growth of tobacco plants in soil infested with Thielavia
basicola as compared with growth in uninfested soil, at different soil tem¬
peratures. Root to right, in each pair, grown in infested soil. Root to left,
grown in same soil sterilized. Note the reduction in amount of disease as
the soil temperature at which the roots were grown is increased, also the
reduction in size of the normal root system.
PLATE XXX vn
Tobacco plants showing the effects of variation of temperature on the
development of the root-rot disease. (Photographs by James Johnson.)
Pig. 1. a. Tobacco plant grown at 23°iC. in Thielavia-infested soil.
d. Plant grown at 23°C. in same soil sterilized.
Fig. 2. a. Tobacco plant grown at 26°iC. in Thielavia-infested soil.
h. Plant grown at 26°C. in same soil sterilized.
The Thielavia root-rot disease starts on a rapid decline at a soil tem¬
perature of about 26° C., whereas at 23 °C. it is near its optimum for
development.
TRANS. WIS. ACAD., VOL,. XX
PLATE XXXIII
JOXES — SOIL TEMPERATURE
NWW
TRANS. WIS. ACAD., VOL. XX
PLATE XXXIV
JONES-— SOIL TEMPERATURE
TRANS. AVIS. ACAD., VOL,. XX
PLATE XXXV
% ■
JONES— SOIL TEMPERATURE
TRANS. WIS. ACAD., VOL. XX
PLATE XXXVI
JONES — SOIL TEMPERATURE
TRANS. WIS. ACAD., VOL. XX
PLATE XXXVII
JONES — SOIL TEMPERATURE
QUANTITATIVE STUDIES OF THE BOTTOM FAUNA IN
THE DEEPER WATERS OF LAKE MENDOTA1
Chancey Juday
Notes from the Biological Laboratory of the Wisconsin Geological and
Natural History Survey. (XV.
Introduction
A general investigation relating to the biological productivity of
Lake Mendota was begun in the year 1911, and various phases of
this problem have received attention during the intervening period
of time. The first phase of the work involved a study of the plank¬
ton of the lake, both from a quantitative and from a chemical stand¬
point. This part of the investigation was continued from 1911 to
1917, and a bulletin which deals with the results obtained in this
study is now ready for publication.
In the autumn of 1913, Muttkowski (4) began a statistical study
of the bottom fauna in the shallower water of Lake Mendota. Par¬
ticular attention was given to the region between the shore line
and the seven-meter contour line, comprising an area of about 12
square kilometers or slightly more than 30 per cent, of the total
area of the lake. His investigation was continued until Septem¬
ber 1, 1916, thus covering a period of approximately three years,
and the results were embodied in a report which was published in
1918.
In addition to this quantitative study of the bottom fauna of the
shallower water, attention may be called to the fact here that one
of the papers in the present series of notes (no. XVII) deals with
the results obtained in a quantitative investigation of the larger
1 Part of this investigation was done in cooperation with the U. S. Bureau
of Fisheries, and the results are published with the permission of the Commis¬
sioner of Fisheries.
461
462 Wisconsin Academy of Sciences , Arts, and Lett rs.
aquatic plants of this same area, that is, the region between the
shore line and a depth of 7 meters, while another (no. XVIII)
treats of the chemical composition of some of these plants.
For purposes of comparison with the shallow water data, Mutt-
kowski extended his studies in the spring of 1916 to the bottom
fauna of the deeper water also, chiefly the region between 7 meters
and 20 meters. The material from the deeper water was obtained
by means of an Ekman dredge with which samples of the bottom
could be readily secured at various depths, except in places where
the bottom was composed of rock or pure sand. At the conclusion
of Muttkowski’s investigation on September 1, 1916, the study of
the bottom fauna of the deeper water was taken up by the present
writer, and it was continued until the middle of August, 1918. The
statistical data relating to the deep-water area which were obtained
by Muttkowski between the latter part of May, 1916, and the first
of September of that year, have been incorporated with those that
were secured after the completion of his work.
In September, 1916, an extended study of the bottom fauna of
the deepest portion of the lake was begun. Five regular stations
were established within the 20-meter area, and observations were
made at these stations at frequent intervals, usually once a week,
except when the lake was covered with ice ; during the winter sea¬
son the stations were visited only once or twice a month, but in
summer they were sometimes visited as often as twice a week. The
stations were widely separated in order to secure a fair average
for the whole area lying within the 20-meter contour line. The
depth at the various stations varied from 20.5 meters at the shal¬
lowest station to 23.5 meters at the deepest one. The bottom at
these points consists chiefly of marl, the upper part containing a
fairly large admixture of organic material derived from the plank¬
ton as well as from the shallow water and shore vegetation. A
certain amount of clay was found also, particularly at the two sta¬
tions situated in the western part of the lake. Chitinous material
derived from the shells of the plankton Crustacea appeared in the
upper layers of the mud.
A sixth station was established in a depression toward the south¬
ern side of the lake where the water reaches a depth of 18.5 meters.
At this point a somewhat larger proportion of vegetable debris was
found in the mud as well as a slight admixture of sand.
Observations were made, in the latter part of 1916, which ex¬
tended in series from shallow water to deep water, and such ob-
Juday — Quantitative Studies of Fauna in Lake Mendota. 463
servations were continued in 1917 also. The composition of the
bottom in these instances varied from practically pure sand to
material which was characterized as marl. In general, a complete
series was begun at a depth of about 7 meters and was carried out
to a depth of from 18 to 20 meters.
The bottom of Lake Mendota has thus been separated into three
zones for the study of its bottom fauna. The first zone extends
from the shore line to a depth of 7 meters, and it is designated as
the littoral zone; the second lies between the depths of 7 meters
and 20 meters, constituting the intermediate zone, and the third
includes the area within the 20-meter contour line, thus represent¬
ing the deep-water zone. The first or littoral zone has an area of
about 12 square kilometers (4.6 square miles) ; the second or in¬
termediate zone includes an area of about 21 square kilometers
(8.1 square miles) and the third about 6.6 square kilometers (2.5
square miles). Expressed roughly in percentages of the total area
of the lake they are respectively 30 per cent., 53 per cent., and 17
per cent. As already indicated, Muttkowski dealt with the popu¬
lation of the littoral zone in his paper which was published in
1918. The present paper is based upon the studies made upon the
population of the intermediate and of the deep-water zones.
During this investigation 36 samples of mud were obtained from
the outer portion of the littoral zone, that is, at depths between
4 meters and 7 meters ; 303 samples were taken in the second or in¬
termediate zone, but 63 of them were obtained at the regular sta¬
tion in 18.5 meters of water; 276 were taken in the third or deep¬
water zone. With very few exceptions, the samples were secured
with an Ekman dredge which covered an area of 473 square centi¬
meters. Where the bottom consisted of fairly soft material, this
dredge penetrated the mud to a depth of 15 to 20 centimeters, but
where a large percentage of sand was present it did not go so deep.
On fairly pure sand, for example, it reached a depth of only two
to three centimeters, or rarely five centimeters. A few samples
were taken with a smaller dredge, one with an opening 15 centi¬
meters square, covering an area of 225 square centimeters; but
usually two hauls were made for a sample when this dredge was
used, so that the area covered for a catch was almost as large as
that for a sample taken with the large dredge. In the softer mud
the small dredge penetrated to a depth of about 15 centimeters.
The samples of mud were transferred from the dredge to spe¬
cially constructed containers or buckets made of galvanized iron.
464 Wisconsin Academy of Sciences , Arts, and LettVrs.
The opening of these containers is rectangular in outline, being 36
centimeters long by 28 centimeters wide, and they are 30 centi¬
meters deep. The capacity is about double that of the large dredge.
Two handles are fastened to the top of the buckets about five centi¬
meters on either side of the middle. These handles are movable
and are bent so that, when open, they conform to the outline of the
top of the bucket and do not obstruct the opening. Double handles
are used in order to make the containers stable when they are being
carried.
The samples of mud were diluted with about an equal volume
of water, and they were then washed through a gauze net with
meshes fine enough to retain all of the macroscopic organisms. The
material which was retained by the net was taken to the laboratory
where the various forms were enumerated in the living state.
In addition to the numerical work, specimens of the various bot¬
tom inhabitants were picked out and weighed, so that gravimetric
data relating to this population were obtained also. Both the live
and the oven dry weights were ascertained as well as the ash.
Small platinum dishes and a sensitive assayer’s balance were used
for these gravimetric determinations. Several weights of each
form were taken in order to get a general average; these weights
included various sizes of the different forms and also represented
different seasons of the year. Such numerical and gravimetric re¬
sults enable one to make a reasonably good estimate of the produc¬
tiveness of the bottom with respect to the various forms. Petersen
(5) was the first to use this method of ascertaining the productive¬
ness of the bottom, and he has obtained some very interesting re¬
sults for the waters along the coast of Denmark.
A. The Third or Deep-Water Zone
The bottom population of the third or deep-water zone of Lake
Mendota will be considered first. This population consists of the
representatives of three groups of animals, namely: (1) worms
belonging to the genera Limnodrilus and Tubifex; (2) a bivalve
inollusk, Pisidium idakoense Roper; and (3) the larvae of three
Diptera, Corethra punctipennis Say, Chironomus tentans Fabri-
cius, and Protenthes choreas Meigen.
The bottom-dwelling organisms which occupy the area within
the 20-meter contour line have a fairly stable environment in so
far as the temperature and the substratum are concerned. The
Juday — Quantitative Studies of Fauna in Lake Mendota. 465
annual range of temperature at this depth varies from a winter
minimum of about 1°C., or slightly less, to a maximum of approxi¬
mately 14° in late summer or early autumn. In some years the
summer temperature is somewhat less than 10°, which gives a
fairly low mean annual temperature, but the animals living here
do not seem to be affected unfavorably by it. At a depth of three
meters, for example, the variation in temperature is much greater,
ranging from a minimum only a little above freezing in early win¬
ter to a maximum of 25° or more in summer.
This area lies well below the lower limit of wave action so that
the bottom here is not disturbed by the waves. The deposition of
material is relatively slow in this region also because, in general,
only the finer silt and the smaller particles of the larger acquatic
plants are transported so far from shore.
At certain periods of the year there is no dissolved oxygen in
the lower water of Lake Mendota, and these organisms, therefore,
must be able to withstand such a condition in order to survive ; in
other words they must be facultative anaerobes. At depths of 20
meters or more the dissolved oxygen usually disappears before the
middle of July, and it is not found in this stratum again until
early October, so that the summer anaerobic period covers an in¬
terval of about two and a half months. Also during the month of
March in certain years some of the bottom water in the deeper
strata possesses no dissolved oxygen, so that a second anaerobic
period prevails for two or three weeks, perhaps longer, over more
or less of the area within the 20 -meter contour line.
Limnodrilus and Tubifex
These two forms were enumerated in 213 samples taken from the
five regular stations located in the deepest water. In 68 of these
samples the two were counted separately, and the ratio of Limnod¬
rilus to Tubifex in them was 4:1. No attempt was made to keep
the two forms separate in the other catches.
The results of the enumerations are summarized in table 1, which
shows the number of individuals per square meter of bottom at the
different stations. A distinctly larger number was found at the
two stations situated in the western half of the deep-water area;
that is, at those designated as West Bay and North of Second
Point. The maximum, the minimum, and the mean numbers were
larger at the former than at any other station. The difference be-
30— S. A. L.
466 Wisconsin Academy of Sciences , Arts , and Letters.
tween maximum and minimum at the West Bay station is not as
great, however, as it is at two of the other stations, being less than
twelvefold. At the Maple Bluff station the difference is only a
little more than eightfold, while at station II it is slightly more
than twenty-onefold; the other stations fall between these two ex¬
tremes.
Differences in density ranging from tenfold to twentyfold show
that these Oligochaeta are not distributed uniformly over the bot¬
tom in the deep-water area, but these differences are no greater
than might reasonably be expected. Indeed, the population of a
large city, if considered block by block, would probably show as
large differences as these, perhaps even larger, in the various areas.
At station II, where the maximum difference was noted, the samples
were all obtained within a fairly definite area because a buoy was
anchored at this point. The launch was anchored to this buoy
each time when samples were taken, so that the particular area in
the vicinity of this buoy from which a sample was obtained de¬
pended upon the direction, and to a less extent upon the strength,
of the wind.
No buoys were placed at the four other stations, but by means
of landmarks on shore and by soundings it was possible to secure
samples at each of these stations within a reasonably definite area.
It was not desirable, in fact, to obtain samples from too small an
area at each station because the main purpose was to secure a gen¬
eral idea of the density of the bottom population at the various
stations. Such results could then be used to estimate the popula¬
tion of the whole of the deep-water area. Increasing the number
of observations and extending them over a considerable period of
time also made the results more serviceable for this general con¬
sideration.
Table 2 shows the distribution of the Oligochaeta at these five
stations by months. The results do not indicate a seasonal varia¬
tion in numbers, but the monthly averages are higher between
October, 1917, and May, 1918, than at other periods covered by this
investigation. The average number of Limnodrilus and Tubifex
together in the 213 samples amounts to 3,500 individuals per square
meter.
Ekman (3) found two species of Limnodrilus and four species
of Tubifex in the bottom fauna of Lake Yaettern, but the total
number of individuals of both genera was smaller than that found
in Lake Mendota. His results also show that the tubificids were
Juday — Quantitative Studies of Fauna in Lake Mendota. 467
more common than the limnodrilids ; the reverse was true of Lake
Mendota.
There was considerable variation in the size of these two forms,
so that it was necessary to pick out all the individuals in a sample
and weigh them in order to obtain an average weight. The results
of four such determinations are shown in table 3. The average
live weight is 1.65 milligrams per individual, of which 82.4 per
cent, is water, leaving a dry weight of 0.3119 milligram per indi¬
vidual. Applying these results to the numerical average per
square meter at the five stations, we find that the live weight of
these animals is 57.75 kilograms per hectare, or 51.4 pounds per
acre; on the basis of dry weight the amount is 10.9 kilograms per
hectare, or 9.7 pounds per acre. It must be remembered that these
figures show only what may be called the standing crop which was
present during the period of this investigation and that they do
not indicate the amount of this material produced annually.
With respect to the nutritive qualities of this material, it may
be said that the percentage of ash is low in these Oligochaeta, aver¬
aging only 4.25 per cent, of the dry weight, while the percentage
of nitrogen is fairly high, being 7.76 per cent., which is equivalent
to 48.5 per cent, of crude protein in the dry material.
Pisidium idahoense Roper
This mollusk was counted in 216 samples from the five deep¬
water stations. Table 4 gives the range in the number per square
meter at the different stations. The difference between the maxi¬
mum and minimum numbers is greater than in the case of the
Oligochaeta, a fifty-fourfold difference being noted in the samples
from the station North of Second Point. In general, however, the
variation is much smaller than these figures seem to indicate, the
maximum being only about twenty-five to thirty-five times as large
as the minimum. The station off Maple Bluff showed the smallest
mean, namely, -360 individuals per square meter, while station II
gave the highest average, or 690 individuals per square meter. The
mean for the 216 samples is 557 individuals per square meter.
Various sizes of Pisidium were noted in the samples; the range
in size was from shells that were only a little over one millimeter
long up to those that were almost nine millimeters in length. In
134 samples the individuals were grouped into three classes accord-
468 Wisconsin Academy of Sciences, Arts, and Letters.
ing to size, and these groups were enumerated separately. They
were recorded as small, in which the length of the shell varied from
1.5 to 3.5 millimeters, medium with a length of 4 to 6.5 millime¬
ters, and large or those having a length of 7 millimeters or more.
In these 134 samples there were 3,949 individuals, of which 3,343
were recorded as small, 473 as medium, and 133 as large. Stated
in percentages of the total number, they were respectively 84.7
per cent., 12 per cent., and 3.3 per cent. On the basis of these
figures, it appears that only a relatively small percentage of the
individuals reaches what may be called full size, that is, the stage
in which they were classed as large. This seems to indicate a
rather high rate of mortality.
While only 33 individuals per thousand reach what may be
called complete maturity, a much larger proportion become sexu¬
ally mature. In addition to the large ones, those classed as medium
in size were sexually mature, and so were some of the larger ones
belonging to the class designated as small. This provision for
early sexual maturity tends to prevent any decrease in the Pisidium
population which might result from such a high death rate. The
mortality indicated above does not include that which takes place
before the one-millimeter stage is reached, and this would doubt¬
less make an appreciable addition to the above number. There
does not seem to be any special reproductive period, since embryos
in different stages of development are present in the various indi¬
viduals at all seasons of the year. This habit of continuous re¬
production may be a further provision against undue decrease in
numbers.
It has already been stated that there is no free oxygen below a
depth of 20 meters for a period of about two and a half months
during the summer. Pisidium seems to pass this interval in a
quiescent or dormant state ; specimens have been kept under obser¬
vation for a month or more, and no evidences of any activity what¬
ever were detected as long as anaerobic conditions obtained. But
when placed in aerated water they soon resumed their normal ac¬
tivities.
Several gravimetric determinations were made for the purpose
of ascertaining the average weight of the individuals belonging to
the three classes into which this form was separated on the basis
of size. In some instances the animals were removed from their
shells, both were dried and weighed separately, and then ashed.
Juday— Quantitative Studies of Fauna in Lake Mendota. 469
Some of the specimens were put in boiling water for a few seconds
so that the bodies of the animals could be more readily removed
from the shells, while others were removed as completely as possible
without being treated with hot water. Some specimens were
weighed, also, without removing the animals from the shells. The
averages for the different classes are shown in table 5.
With respect to the dry weight it will be noted that the medium
size weighs, on an average, three and a half times as much as the
small, while the large weighs about three and a half times as much
as the medium. The percentage of ash is unusually low in the
animals of large and medium sizes, but much higher in the small
specimens. The percentage of ash in the large and medium classes
is lower than that obtained by Bradley (2) for the larger mollusks
which he collected in the vicinity of Madison. He found that the
ash content of the various tissues and organs ranged from a mini¬
mum of 6 per cent, in the muscle to a maximum of 48 per cent, in
the mantle. The distinctly higher percentage of ash in the small¬
sized Pisidium may be due to a greater activity of the mantle in
the secretion of the shell.
In all three sizes of this mollusk the body portion of the animal
constituted from 17 per cent, to 21 per cent, of the dry weight of
the entire organism, the average being highest in the small size ; in
other words, from 79 to 83 per cent, of the dry weight of the whole
animal consisted of the shell.
By taking the average number of these mollusks per square me¬
ter, namely, 557, and the relative percentages of the different sizes,
the weight per unit area may be estimated. This amounts to 3.7
kilograms of dry matter per hectare, or 3.3 pounds per acre, not
including the weight of the shell. No live weights were determined,
so that the quantity of living material can not be given. These
figures show that Pisidium plays a relatively unimportant role so
far as being a source of food is concerned. No chemical analyses
of this mollusk were made.
Chironomus tentans Fabricius
The larvae of this dipteron were enumerated in 263 samples and
they yielded an average of 593 individuals per square meter of
bottom. The variation in the averages for the five different sta¬
tions are shown in the first part of table 6. The lowest average
was obtained at the station North of Second Point, while the maxi-
470 Wisconsin Academy of Sciences , Arts , and Letters.
mum was found at the Maple Bluff station, the latter being almost
twice as great as the former. The samples from West Bay and
from the Center gave an average of 566 individuals per square
meter, while those at station II numbered 549.
There was a marked difference in the size of these larvae, and
they were separated into three different classes on the basis of size.
In 219 samples those larvae were recorded as small which did not
exceed 12 millimeters in length; from this length up to about 18
millimeters they were designated as medium, and those 20 milli¬
meters or more in length were called large. The largest larvae
noted in this region of the lake were about 27 millimeters in length.
The results of these enumerations for the various stations are
shown in the second part of table 6. Station II yielded the largest
average number of small individuals, and the Maple Bluff station
ranked first in both medium and large sizes. Of the total num¬
ber of Chironomus larvae obtained at the five stations, the small
individuals constituted 56 per cent., the medium size 31 per cent.,
and the large 13 per cent. Thus, the small ones were a little more
than four times as numerous as the large ones, but less than twice
as abundant as the medium size.
The various sizes of this chironomid larva were found during all
seasons of the year. Pupae were most abundant in May and early
June, and again in the latter half of October, the maximum num¬
ber being found during the latter month. This indicates that these
two periods are the chief ones for transformation, but pupation also
takes place between June and October so that eggs are constantly
being deposited during this interval of time. The eggs deposited
by the adults which emerge in late October and early November
give rise to a brood of young which flourishes during the winter
months. The annual distribution of the different sizes is shown in
table 7. The small ones were much more abundant in 1917 than
in 1918, but, up to June, the medium and large sizes were more
numerous in 1918 than in 1917.
Average specimens of the three different sizes were picked out
and their weights were determined; the results are shown in table
8. The large individuals were from 23 millimeters to 25 millime¬
ters long, the medium ones from 14 millimeters to 16 millimeters,
and the small ones from 8 millimeters to 10 millimeters. The live
weight of the large specimens was a little less than five times as
much as that of the medium ones, while the medium ones weighed
Juday — Quantitative Studies of Fauna in Lake Mendota. 471
just a little more than five times as much as the small ones. For
the dry weights, however, the ratio was appreciably less than five
to one in both instances. It will be noted that the percentage of
water was highest in the large individuals and lowest in the small
ones ; the same was true of the ash.
By using the average number of larvae per square meter,
namely, 593, the percentages of the different sizes, and their aver¬
age weights, we may calculate the live and dry weights of the
chironomid population of the deep-water zone. The results of such
a calculation are shown in table 9. While the small individuals
were almost five times as numerous as the large ones, their live
weight was less than one fifth as great, but their dry weight was
slightly more than one fifth as much as that of the large ones.
The live weight for all sizes averages a little more than 75 kilo¬
grams per hectare, or slightly more than 67 pounds per acre, while
the dry weight is a little in excess of 12 kilograms per hectare (10.8
pounds per acre) .
The larvae of Chironomus tentans are regarded as an important
source of food for fishes, and chemical analyses were made for the
purpose of obtaining some idea of their food value. The material
which was used for these analyses consisted of 1,552 individuals
ranging from 20 millimeters to 25 millimeters in length. The re¬
sults obtained in the analyses are shown in table 10.
The percentage of nitrogen given in this table is exclusive of that
in the crude fiber or chitin, namely, 0.23 per cent.; that is, the
total nitrogen amounted to 7.59 per cent, of the dry material.
While the percentage of nitrogen is not as high as in the larvae of
Corethra, nevertheless it is high enough to make these chironomid
larvae an excellent source of protein material. That is, about 46
per cent, of the dry weight of the larvae may be regarded as crude
protein. The fat content is rather low, namely, only 8 per cent.
The ash in these specimens is distinctly higher than that indicated
in table 8 ; this is probably accounted for by the fact that the ash
content depends, in large measure, upon the character of the food
that is present in the alimentary canal.
Corethra pnnctipennis Say
The larvae of Corethra punctipennis may be regarded as bottom
dwellers, in part at least, since the larger ones desert the water
entirely during the daylight hours and remain concealed in the
472 Wisconsin Academy of Sciences, Arts, and Letters.
muddy ooze of the bottom. At night they migrate into the water
and occupy the various strata of the lake, even coming to the sur¬
face. Thus, they show a diurnal periodicity in habitat, occupying
the mud in the daytime and the water at night. The young larvae,
however, are entirely aquatic for a time after they hatch out ; that
is, they occupy the lower strata of the lake, the hypolimnion, in the
daytime instead of the mud, but they come into the upper strata,
also, at night. When they are from one third to one half grown,
they show the same behavior as the full grown larvae, that is, they
occupy the mud in the daytime and migrate into the water at
night.
In table 11 the number of these Corethra larvae per square me¬
ter of bottom is indicated by months for the period covered by these
observations. The first part of this table shows the results obtained
at station II, and the second part gives the averages by months for
the five stations located in the deep water. The data for station
II are given separately because the other stations were not visited
regularly in the winter. Large numbers of the larvae live over
winter; in fact, they are more numerous from November to April
than at any other time of the* year. During this period there is
no loss from pupation, and the losses from other causes are not
large enough to reduce the number very materially, so that the
numbers remain uniformly high during this interval of time.
The results given in table 11 are shown graphically in figure 1.
The solid line in this figure represents station II and the broken
line the averages for the five stations. The vertical spaces in the
diagram indicate the number of individuals per square meter and
range from zero up to 30,000; the months are shown in the hori¬
zontal spaces.
Beginning with the results for May, 1916, there is a marked de¬
cline in the number of Corethra larvae at station II in the month
of June, with a further decrease in July; the decrease during the
latter month was not nearly as great as during the former month.
No observations were made at this station during the month of
August in that year, so that the gap from July to September is
represented by a broken line. In order to complete the curve the
number for August, 1916, has been assumed as about a thousand,
corresponding to the numbers found at this station in August,
1917, and 1918. There was a distinct increase in the number of
Corethra larvae during September, and the rise continued during
Juday—Quantitative Studies of Fauna in Lake Mendota . 473
the months of October and November. In December, 1916, the
number was the same as in the previous month, but there was a
slight decline during the next two months with a distinct rise in
March to the largest average number obtained in the entire series
of observations. This maximum was followed by a uniform de¬
crease to the August minimum of 1917.
Fig. 1. The curves in this figure show the average number of larvae
of Corethra punctipennis in thousands per square meter of bottom for the
different months during the period of these observations. The vertical
spaces range from zero to 30,000. The curve shown as a solid line repre¬
sents the averages for station II; the broken line indicates the monthly
averages for the five stations in deep water combined. The latter curve
is interrupted during the winter because observations were not made
regularly at all of these stations during this season. (See table 11.)
This summer minimum was succeeded by an autumnal rise which
culminated in a December maximum, but the December maximum
of 1917 was appreciably smaller than that of December, 1916. No
samples were obtained at station II in January, 1918, but from
February to May there was a gradual decline in the number of
larvae obtained at this station, followed by a rapid decrease during
the next three months to the August minimum of 1918. The larg¬
est number of larvae obtained in any of the samples was 33,800
individuals per square meter, this number being found at station
474 Wisconsin Academy of Sciences , Arts , and Letters.
II on December 21, 1917 ; the smallest number was also noted at
this station, namely, 295 larvae per square meter, found on August
2, 1917.
The curve shown as a broken line in figure 1 represents the aver¬
ages for the five stations. This curve is interrupted during the
winter each year because station II was the only one visited regu¬
larly during this season of the year. The monthly averages for the
five stations are larger than those of station II during the period
from July to October, but they are smaller than the latter in No¬
vember and in December, and also from April to June. In Decem¬
ber, 1916, however, the two averages were substantially the same.
The maximum differences between the two averages were obtained
in April and in December, 1917 ; on the former date the average
for the five stations was more than 22 per cent, below that of sta¬
tion II, while on the latter date the difference was a little more
than 25 per cent, in favor of station II.
Small swarms of adults appear in May and in early June, but
the great flights do not begin until late June or early July. The
most vigorous period of transformation from larvae to adults lasts
from early July to about the middle of August. During this in¬
terval of time several great waves of adults appear; as a result of
the rapid pupation at this time the larvae decline to the minimum
number of the year in early August. In late August and early
September there is a slackening in the pupation, and correlaated
with this is an increase in the number of larvae. As autumn ad¬
vances pupation becomes less and less frequent; this decline in
transformation, together with the hatching of the vast numbers of
eggs deposited by the later broods, results in a rapid rise in the
number of larvae. This rapid increase begins in late September
and continues until November or December. (See fig. 1.)
Between June, 1916, and April, 1917, more than fourteen thou¬
sand Corethra larvae were sorted out and dried for the purpose of
making chemical analyses of them. The average amount of dry
matter in each individual was 0.251 milligram. In addition to
this, eleven other lots, consisting of 100 to 300 larvae each, were
weighed, and their dry weight ranged from a maximum of 0.311
milligram per individual in June to a minimum of 0.182 milligram
in early September. The results of these weighings are shown in
table 12. The higher average of dry matter in the June material
was probably due to a larger proportion of chitin in the larvae just
Juday - — Quantitative Studies of Fauna in Lake Mendota. 475
before pupation. In August and in early September the larvae
have the appearance of being smaller in size than the winter larvae,
and this is confirmed by the weights. At the height of the pupating
season the summer larvae pupate when they are distinctly smaller
than the winter larvae.
The live weight of the pupa is somewhat greater than that of
the larva of the same date, but its dry weight is nearly twice as
much as that of the larva. This marked difference in dry weight
is probably due to the presence of a larger amount of chitin in the
pupa. Both the water and the ash content of the pupa are smaller
than in the larva.
The adults yielded a much smaller live weight than either the
larvae or the pupae, but they possessed a much smaller percentage
of water. Their dry weight was smaller than that of the pupae
but larger than that of the larvae. The adults were obtained from
a large swarm on June 29, 1918, when pupation was very active,
but there was no means of ascertaining their age. Since, presum¬
ably, the adults do not take food, their weight will decrease with
age, and the adults of this form live from three to five days. The
percentage of ash was about the same in the adults and pupae, but
in both it was distinctly lower than in the larvae.
There is more or less overlapping of the summer broods, which
makes it difficult to estimate the production during this season,
but the winter crop of larvae may be estimated with some degree
of accuracy. During this investigation twenty-two different hauls
were made at the five regular stations during the month of Novem¬
ber, and these samples gave an average of 17,350 Corethra larvae
per square meter. Five samples were also obtained in the month
of December, and they gave an average of 21,900 individuals per
square meter; four of these samples, however, were obtained at
station II, which usually gave a larger yield than the other sta¬
tions. On a conservative basis, then, the early winter population
of Corethra larvae may be estimated as about 18,000 individuals
per square meter in the deep-water zone of Lake Mendota. Be¬
tween October and May the average live weight was 3.1 milligrams
per larva and the dry weight 0.266 milligram. Applying these
weights to the above population gives a live weight of 558 kilo¬
grams of Corethra larvae per hectare (497 pounds per acre) and
a dry weight of 47.9 kilograms per hectare (42.7 pounds per acre).
This estimate does not take into consideration the summer gener¬
ations, of which Muttkowski (4) says there may be two. Since
476 Wisconsin Academy of Sciences, Arts, and Letters.
the summer larvae average somewhat smaller than those of the
winter generation, it may be conservatively estimated that the sum¬
mer broods are equal in weight to the winter generation. That is,
the total annual production of Corethra larvae in this region of
Lake Mendota may be estimated as about 1,200 kilograms of live
material per hectare (a little over 1,000 pounds per acre), or some¬
what more than 100 kilograms of dry matter per hectare (about 90
pounds per acre). These figures, however, apply only to that por¬
tion of Lake Mendota which lies within the 20-meter contour line,
or to 16.8 per cent, of the total area. In the shallower water the
larvae are found in much smaller numbers, as indicated on a sub¬
sequent page.
The results of the chemical analyses of the larvae are shown in
table 10. It will be noted that the percentage of nitrogen is espe¬
cially high, amounting to 10.7 per cent, of the dry material, and
this means a correspondingly high percentage of crude protein,
namely, a little more than 67 per cent, of the dry sample. The
figure for nitrogen does not include that in the crude fiber or
chitin, which amounted to 0.46 per cent, of the dry material ; that
is, the total nitrogen amounted to 11.2 per cent. The Corethra
iarvae yielded a distinctly higher percentage of nitrogen than
either the Oligochaeta or the larvae of Chironomus tentans, thus
showing that they are a better source of nitrogenous material.
The Corethra larvae also yielded a fairly large percentage of
fat or ether extract. The crude protein and the fat together con¬
stituted more than 76 per cent, of the dry weight of these organ¬
isms. From the standpoint of quality, therefore, this large propor¬
tion of these two excellent food materials gives these larvae a very
1 igh rank as a source of food for other forms.
Protenthes choreus Meigen
The larvae of Protenthes choreus were present in the mud of the
deeper portion of the lake in comparatively small numbers. They
were somewhat irregular in their distribution also, so that there
were relatively large variations in numbers from time to time. The
distribution by months is shown in table 13. There were marked
differences in numbers during the winter, due mainly, perhaps, to
the small number of observations made during this season. Seven¬
teen samples taken in April yielded an average of 400 Protenthes
larvae per square meter, and 32 samples in May gave an average
Juday — Quantitative Studies of Fauna in Lake Mendota. 477
of 390 larvae per square meter. These averages indicate that about
400 larvae per square meter survive the winter. The number re¬
mains above 300 individuals per square meter until June; follow¬
ing this is a marked decrease in July, with a further decline in
numbers in August. The minimum number was found in Septem¬
ber. Following this, there was an increase, with the number ris¬
ing above 300 per square meter in December. The majority of the
samples obtained in September contained no Protenthes larvae, and
similar results were obtained for many of the samples procured in
late August and early October.
Table 14 gives data concerning the weight of these larvae, to¬
gether with the percentages of water and of ash. The results for
the large specimens (9 millimeters to 10 millimeters long) repre¬
sent the averages of four sets of weighings. The rest of the table
shows the results obtained when all of the specimens found in a
sample were weighed. These individuals ranged from 5 millime¬
ters to 10 millimeters in length, the minimum length increasing
somewhat between autumn and the following summer. The growth
of the smaller individuals is well illustrated m the four weighings
that were made between December 21, 1917, and June 10, 1918;
the average weight of the mixed sizes almost doubled during that
period of time. The maximum weight of the mixed sizes (June
10, 1918) was less than half as much as the average weight of the
large size. The percentages of water and of ash were also smaller
in the mixed sizes than in the large specimens.
The mean of the weights obtained for the mixed sizes in Febru¬
ary and May gives a fair average for the larvae which survive the
winter; that is, live weight 1.4 milligrams per larva and dry weight
0.238 milligram. On the basis of these weights, the April crop of
larvae, namely, 400 individuals per square meter, represents a live
weight of 5.6 kilograms per hectare (5 pounds per acre) and a dry
weight of approximately one kilogram per hectare (0.8 pound per
acre). Whether there is more than one generation per year has
not been determined, so that no estimate of the annual production
can be given for this form. The winter larvae, however, repre¬
sent a comparatively small amount of material, so that an addi¬
tional generation or two would not increase the food supply in
this region very materially. The above figures, also, do not in¬
clude the various losses sustained during the winter, but these
losses doubtless represent only a small percentage of the annual
crop of larvae.
478 Wisconsin Academy of Sciences, Arts, and Letters.
While the larvae of Protenthes choreas are eaten extensively by
fishes, they do not play nearly as important a role in the diet of
these animals as the other two dipterous larvae found in the deeper
water, because they average much smaller in size than the chiron-
omid larvae and are also less abundant, and because they are pres¬
ent in very much smaller numbers than the larvae of Corethra.
The gravimetric data for the various organisms inhabiting the
bottom in the deeper water of Lake Mendota are summarized in
table 15. For the live weights the totals do not include Pisidium
idahoense because such data were not obtained for this form. The
weights given for the larvae of Corethra show only the April crop,
but, even on this basis, these larvae yield about four times as much
dry material as the next form in importance, namely, Chironomus
tentans. Just the April crop of Corethra larvae, in fact, contains
more dry material than all the other forms combined. If the sum¬
mer broods of this form be added, it shows a very marked excess
over the others. The larvae of Protenthes yielded the smallest
amount of material.
The annual production of the various forms has not been esti¬
mated except for the Corethra larvae. It would probably not be
far from the actual production to assume that each of the other
forms produces annually a quantity of material equivalent to the
average amount found in this investigation, while Corethra yields
twice as much as was noted for this form in April. On this basis,
the total annual yield of these bottom forms within the area under
consideration would amount to somewhat more than 1,255 kilo¬
grams per hectare live weight, including Pisidium (1,120 pounds
per acre), or a dry weight of about 125 kilograms per hectare (112
pounds per acre). The Corethra larvae are responsible for about
77 per cent, of this quantity of dry material, and the larvae of
Chironomus tentans for about 10 per cent., while Limnodrilus and
Tubifex come third with somewhat less than 9 per cent. For the
deep-water zone, which comprises 664 hectares, this would mean
an annual crop of slightly more than 833 metric tons of these or¬
ganisms in the living state, or 83 metric tons of dry material.
B. Station I
In addition to the five stations located in the deep water, a
sixth station was visited regularly. The latter is situated in the
southern portion of the deeper water, between Picnic Point and
Juday — Quantitative Studies of Fauna in Lake Mendota. 479
the lakeside laboratory of the University of Wisconsin, and it has
been designated as station I in the various observations that have
been made here. The water has a depth of 18.5 meters at this
point. The data obtained at this station are discussed separately
for three reasons: (1) Because the station is isolated somewhat
from the main body of the deeper water and lies well toward its
southern edge; (2) because the water is somewhat shallower here
than at the other stations ; and ( 3 ) because the results show some
characteristic differences. The mud at this station contains a
somewhat larger proportion of coarse vegetable debris than at the
other stations, which is due, doubtless, to the fact that it is situ¬
ated near shallow areas supporting fairly dense growths of the
larger aquatic plants.
Limnodrilus and Tubifex. Sixty-four samples were taken at
station I. The Oligochaeta were counted in 52 of them, and an
average of 2,320 individuals per square meter was obtained. This
number is distinctly smaller than the mean of any of the stations
within the 20-meter area (table 1). The average for the Maple
Bluff station comes nearest this figure, while the mean for West
Bay is twice as great.
Pisidium idahoense. This form was enumerated in 52 samples
from station I and a mean of 570 individuals per square meter was
found (table 4). This average was exceeded by only two of the
stations in the deeper water; both the maximum and the minimum
numbers were larger than at any of the other stations. The dif¬
ferent sizes were counted separately in 33 samples, and in them
were found 14 large individuals, 124 of medium size, and 742 small.
Stated in percentages of the total number, these figures represent
1.7 per cent., 14 per cent., and 84.3 per cent., respectively. As
compared with the five stations in deep water, the percentage of
small individuals at station I is substantially the same, but that for
the medium size is larger and that for the large size is smaller.
Chironomus tentans. Table 6 shows that station I ranked sec¬
ond in the average number of the large and medium sizes of this
chironomid larva, but it was first in the small size and second in
the general average. The different sizes in 54 samples consisted
of 11 per cent, of large individuals, 32 per cent, medium, and 57
per cent, small. These percentages are substantially the same as
those for the deep water stations, which are, respectively, 13 per
cent, large, 31 per cent, medium, and 56 per cent, small.
480 Wisconsin Academy of Sciences, Arts, and Letters.
Corethra punctipennis. The average number of Corethra larvae
is shown for the different months of the year in table 11. The win¬
ter population of this form at station I was much smaller than at
the other stations. In 20 samples obtained from October to April,
inclusive, at station I, the average number of these larvae is 9,605
individuals per square meter. During these same months 30 sam¬
ples taken at station II yielded an average of 21,170 larvae per
square meter, while, in the same period, 74 samples from the five
regular stations in the deep water gave an average of 17,425 Cor-
thra larvae. In general, then, it may be said that the population
of Corethra larvae at station I was about half as large from Octo¬
ber to April as it was at the stations in the deeper water. The
August population at station I was also lower than the general
average of the other stations combined, but it was larger than the
August population of station II in 1917 and smaller than the
August population of 1918.
Protenthes choreus. These larvae were much more abundant at
station I than at the deep water stations (table 13). These larvae
were not counted in the January sample from station I, but in the
other months the averages were distinctly larger at station I than
at the other stations, except in April.
The live weight and dry weight of the various forms found at
station I are shown in the second part of table 15. The amounts
have been calculated on the same basis as those for the five stations
in the deeper water, that is, on the general average of Oligochaeta,
Pisidium, and Chironomus, and on the April crops of Corethra
and Protenthes. The results for the two regions are thus directly
comparable. The dry weight of Pisidium and Protenthes was sub¬
stantially the same at station I as at the five stations in the deeper
water, but that of the Corethra larvae was very much smaller and
that of Limnodrilus and Tubifex was smaller. Chironomus, on the
other hand, gave a larger amount of dry material at station I than
the average for the other stations. The total quantity of dry mat¬
ter in the bottom population at station I amounted to 53.8 kilo¬
grams per hectare, while the other stations yielded an average of
76.6 kilograms per hectare. The former is only about 70 per cent,
as large as the latter.
Juday — Quantitative Studies of Fauna in Lake Mendota . 481
C. Intermediate Zone
In the intermediate zone the bottom shows much variation in its
composition, varying from gravel and sand, or even rocks, at one
extreme to the fine mud of the deeper water at the other. In cer¬
tain parts of the shallower regions of this zone there are fairly
large areas of sandy bottom, and this compact material does not
furnish a very good habitat for many forms, such as the Corethra
larvae, for example. While such areas are not entirely devoid of
inhabitants, they are by no means as densely populated as the re¬
gions which have a softer bottom. In the more protected parts of
the lake, mud bottom is found at a depth of 5 to 6 meters, while
along the more exposed shores sand may extend to a depth of 12
meters or more. On the steeper slopes practically no loose mate¬
rial accumulates, so that the bottom is compact even where it con¬
sists of marl. These steep slopes also possess a sparse population.
In the study of this zone an attempt was made to get samples from
the various types of bottom in fair proportions so that the results
would constitute a good general average for the entire intermediate
zone.
Table 16 shows the general averages for the different forms that
were found between the depths of 8 and 20 meters. The results
shown for the 5--7-meter area are taken from Muttkowski’s report
(4) ; for further comparisons, also, the general averages for station
I, station II, and the general average for all samples from the five
deep-water stations are included in this table. All of the results
given in table 16 for Corethra and Protenthes represent general
averages for the entire series of samples and not the spring crops
of these larvae. This method of computation gives a lower average
than that noted for the vernal generation because the number of
larvae is so much smaller during the summer than during the other
three seasons of the year. The averages have been computed on
this basis for this table in order to obtain results which are directly
comparable with each other.
No samples were obtained from the intermediate region in the
month of April, so that the vernal crop of larvae of Corethra and
Protenthes has not been determined directly. The vernal weights
have been estimated for these larvae, however, by means of the
general averages for all samples from this region ; this general av¬
erage has been multiplied by the ratio of the April crop to the
31— S. A. L.
482 Wisconsin Academy of Sciences , Arts , and Letters.
general average for all samples from the five deep-water stations.
This quotient has been taken as the vernal crop of these two forms
in the intermediate zone.
The 5- -7 -meter region is very sparsely populated as compared
with the areas a meter or two deeper. The averages of the 21 sam¬
ples obtained at a depth of 8 meters are from eight times to seventy-
five times as large as those shown by Muttkowski for the 5—7-meter
area. The differences are still more marked when the latter is com¬
pared with the data for 9 meters and 10 meters, or for greater
depths.
Limnodrilus and Tubifex. These two forms show a fairly regu¬
lar increase in numbers with the increase in depth; the average
between 16 meters and 20 meters is about eight times as large as
that at 8 meters. At 17 meters, however, the difference is almost
fourteenfold, but the average for this depth is probably larger than
it should be because three of the five samples here contained unus¬
ually large numbers of these Oligochaeta. The samples from sta¬
tion I (18.5 meters) gave substantially the same mean as those
taken at a depth of 18 meters. The averages for station II and for
the five deep-water stations combined are appreciably larger than
the average for 20 meters, while the mean of all samples from the
intermediate zone is less than half as large as that of the deep
water zone.
Pisidium iddkoense. The most marked increase in Pisidium in
the intermediate zone was between 8 meters and 9 meters, the latter
depth yielding more than twice as many as the former depth. The
averages between 10 and 20 meters show considerable irregularity,
but there is a general tendency for the numbers to increase some¬
what with depth. The largest average for this zone was found at
19 meters. The general average for the whole zone is only a little
more than 63 per cent, as large as that for the deep-water zone.
Chironomus tentans. The larvae of Chironomus show a rather
irregular increase with depth in the intermediate zone, but their
maximum point is reached at 18 meters. Beyond this depth the
number decreases, and the average for the whole zone is a little
more than twice as large as that of the deep-water zone, thus show¬
ing a preference for the former area.
Corethra punctipennis. This larva shows a decided preference
for the deeper water ; the general average for the intermediate zone
is a little less than one third as large as that of the deep-water zone.
Juday — Quantitative Studies of Fauna in Lake Mendota. 483
Protentlfies choreus. The larvae of Protenthes are like those of
Chironomns in that they show a distinct preference for the inter¬
mediate depths; the average number for this zone is about 30 per
cent, larger than that for the deep-water zone.
In addition to the above mentioned forms, which constitute the
regular macroscopic bottom fauna of the deep water, several others
are found in the intermediate region, more especially in the 8- -IO¬
meter area. Leeches were noted in four samples from this zone,
but the number was too small to be of any great importance in the
general population of this area. Also some specimens of Hyalella
were found in two samples obtained at a depth of 15 meters.
Aquatic snails were noted in 22 samples taken from the inter¬
mediate depths, that is, in about 10 per cent, of the samples from
this zone. With very few exceptions they were not found beyond
a depth of 10 meters, but four specimens of Lymnaea were noted
in one sample from a depth of 15 meters. One sample from a depth
of 8 meters yielded 1,100 snails per square meter, but the average
for the other 21 samples in which snails were found was 50 in¬
dividuals per square meter. Beyond a depth of 10 meters, snails
constitute a very unimportant element of the population, from the
numerical standpoint.
Some larvae of Chironomus lobiferous were noted in four catches
from the 8- -10-meter area, but they were not found at greater
depths.
Larvae of Palpomyia longipennis were present in 13 samples
from the intermediate zone, but the average number for these sam¬
ples was only 30 individuals per square meter. A specimen was
found in one sample taken at a depth of 18.5 meters, but these
larvae were obtained most frequently between 8 meters and 12 me¬
ters.
Larvae of Sialis infumata were found in 28 samples from the
intermediate zone. They were obtained at all depths in this region,
but they were not noted in any of the catches made at a greater
depth than 20 meters. The average number found in the 28 sam¬
ples amounted to 72 individuals per square meter, or about 10 per
square meter on the basis of the total number of samples taken from
the intermediate zone. Muttkowski (4) found them in the littoral
zone only in the 5-— 7-meter area, and his average for this area is
approximately five individuals per square meter, or half the num¬
ber for the entire intermediate zone.
484 Wisconsin Academy of Sciences, Arts, and Letters.
Baker (1) states that the larvae of Sialis infumata were rare in
Oneida Lake and that they were found only where the water was
between 2.5 feet and 8.5 feet deep. In Lake Mendota they were
found regularly at a distinctly greater depth, that is, from 5 meters
to 20 meters.
Table 17 shows the live weight and the dry weight of those forms
inhabiting the intermediate zone for which gravimetric data have
been obtained. Such determinations have not been made for the
leeches, snails, Hyalella, Palpomyia, and Chironomus lobiferous,
but these forms were found in such small numbers that the addi¬
tion of their weight to the data given in this table would not in¬
crease the general average for the whole zone very materially.
These weights are based upon the numerical results obtained for
the intermediate depths and upon the mean weights of the various
forms. The data given in this table should be compared with those
given for the deep-water area in table 15.
As might be expected from the numerical results, the Oligochaeta,
Pisidium, and Corethra yield a smaller quantity of dry material
in the intermediate than in the deep-water zone. On the other
hand, Chironomus and Protenthes show a larger amount in the
former than in the latter. Chironomus tentans yielded nearly
twice as much dry material in the intermediate area as it did in
the deep-water region, while Protenthes was about one-third
greater in the former zone. The dry weight of the Oligochaeta
was a little more than twice as large in the deep-water zone as in
the intermediate zone ; Pisidium gave somewhat less than twice as
much in the latter region and Corethra substantially three times
as much.
With respect to the larvae of Sialis infumata, it may be said that
the weight of the different sizes of individuals found in the samples
was not determined, but gravimetric data were obtained for the
full-grown larvae when they migrated to the shore for the purpose
of pupating. About 4,500 individuals of this size were secured for
a chemical analysis, and the dry weight per individual for this
number was 8.4 milligrams. Thus an average of 10 larvae per
square meter in the intermediate zone, on the basis of full grown
larvae, would mean 0.8 kilogram per hectare, or about three quar¬
ters of a pound per acre. The mixed sizes found in the samples
would yield a smaller quantity of dry matter, however, probably
not more than half a kilogram per hectare, but the maximum
amount has been indicated in the table.
Juday — Quantitative Studies of Fauna in Lake Mendota. 485
The results of the chemical analyses made on the larvae of Sialis
are shown in table 10. Substantially 69 per cent, of the dry weight
of these larvae consists of crude protein and ether extract (fat),
while relatively small percentages of crude fiber (chitin) and ash
were found. The total nitrogen amounted to 8.52 per cent, of the
dry weight, of which 0.45 per cent, was derived from the crude
fiber, or chitin.
On the basis of the spring crop of Corethra larvae, the total
quantity of dry material in the various forms shown for the inter¬
mediate zone in table 17 amounts to about 63 per cent, as much
as is indicated for the deep-water zone in table 15. Allowing for
two crops of Corethra larvae per annum, the total amount of dry
material in the intermediate zone becomes 64.2 kilograms per hec¬
tare (57.3 pounds per acre) as compared with 125 kilograms per
hectare (112 pounds per acre) in the deep-water area. The latter
figures may be regarded as representing the annual crops of bot¬
tom fauna in these two zones on the assumption that the Corethra
larvae produce twice the April crop each year and that the other
forms produce yearly a quantity of material equivalent to the
average crop found in this investigation.
Literature
1. Baker, Frank O.
1916. The relation of mollusks to fish in Oneida Lake. N. Y. State
Coll. For. Syracuse Univ. Tech. Pub. 4: 15^366.
1918. The productivity of invertebrate fish food on the bottom of
Oneida Lake. N. Y. State Coll. For. Syracuse Univ. Tech.
Pub. 9: 11-233,
2. Bradley, H. C.
1910-11. Manganese of the tissues of lower animals. Jour. Biol.
Chem. 8: 237-249.
3. Ekman, S.
1914. Die Bodenfauna des Vaettern. Internat. Rev. 6: 146-425.
4. Muttkowski, R. A.
1918. The fauna of Lake Mendota. Trans. Wis. Acad. Sci., Arts,
and Lett., 19: 374-482.
5. Petersen, C. G. J.
1911. Valuation of the sea. I. Rept. Danish Biol. Sta. 20: 1-76.
1913. Valuation of the sea. II. Rept. Danish Biol. Sta. 21: 1-68.
1918. The sea bottom and its production of fish food. Rept. Dan¬
ish Biol. iSta. 25: 1-62.
486 Wisconsin Academy of Sciences , Arts , and Letters.
Table 1. Results obtained for Limnodrilus and Tubifex at the five
deep-water stations. The figures for maximum, minimum, and mean indi¬
cate the member of individuals of these two forms per square meter of\
bottom.
Table 2. The number of Limnodrilus and Tubifex per square meter
during the different months of the year. The' numbers represent the
monthly averages for the five stations in deep water combined.
Table 3. The average weight of Limnodrilus and Tubifex stated in
milligrams per individual, together with the percentages of watdr and of
ash in them. The results for the ash are given in percentages of the dry
weight.
Juday — Quantitative Studies of Fauna in Lake Mendota. 487
Table 4. Numerical results obtained for Pisidium at the five deep¬
water stations and at station I. The figures given for maximum, mini¬
mum, and mean indicate the number of individuals per square meter of
bottom.
Table 5. This table shows (1) the average dry weight of the different
sizes of Pisidium in milligrams, (2) the percentage of ash in the body
part, (3) the percentage of the total weight constituted by the body of the
animal, (4) the percentage of the total weight consisting of shell, and
(5) the percentage of organic matter in the shell.
Table 6. The average number of larvae of Chironomus tentans per
square meter of bottom at the regular stations, and the average number |
of the different sizes in the samples in which they were counted sepa¬
rately.
488 Wisconsin Academy of Sciences, Arts, and Letters.
Table 7. The number of the three sizes of Chironomus larvae noted
at the five deep-water stations during the different months of the year.
With the exception of January and February 1917, the figures for each
month represent averages per square meter , of three to twenty-three
samples.
Table 8. The average weight in milligrams of a single individual of
the three different sizes of Chironomus larvae, together with the percent¬
ages of water and of ash.
Table 9. The live weight and dry weight of Chironomus larvae per
unit area of bottom.
Juday — Quantitative Studies of Fauna in Lake Mendota. 489
Table 10. Results of the chemical analyses of the different insect
larvae. The results are! stated in percentages of the dry weight.
Table 11. The first part of this table shows the average number of
Corethra larvae per square meter of bottom at station II during the
different months of the year. In all months except January and Febru¬
ary the numbers represent averages of two to nine samples . The second
part gives the averages by months for the five deep-water stations. The
third part indicates the results for station I.
Monthly Averages for Station II
Monthly Averages for Five Deep-water Stations
Monthly Averages for Station I
490 Wisconsin Academy of Sciences, Arts, and Letters.
Table 12. Average weight in milligrams of a single individual of Co-
rethra punctipennis, together with the percentages of water and ash.
Table 13. The average number of larvae of Protenthes choreus per
square meter of bottom during the different months of the year. The first
line shows the averages for the five stations in deiep water and the second
line the averages for station I.
Table 14. Live weight and dry weight of the larvae of Protenthes
choreus, together with the percentages of water and of ash. The weights
are stated in milligrams per individual.
Juday— Quantitative Studies of Fauna in Lake Mendota. 491
Table 15. Summary of the live weight and of the dry weight of the
various forms found at the five stations in deep water and at station I.
The weights for the deep water area are based on the averages for- the
five stations. The live weight of Pisidium was not determined, so that it
is not included in the totals under live weight.
Deep-water Stations
Station I
492 Wisconsin Academy of Sciences , Arts , and Letters,
Table 16. The average number of individuals per square meter at the
various depths in the intermediate zone and at the regular stations. The
data for the 5- -7- meter region are taken from Muttkowski’s table 5
a, p. m>.
Juday — Quantitative Studies of Fauna in Lake Mendota. 493
Table 17. Live 'weight and dry weight of the various forms found in
the intermediate zone of Lake Mendota. These results are hosed on thes
averages numbers indicated for the 8 - - 20 - meter region in table 16, with
the ' exception of Corethra. The figures given for Corethra larvae are
based upon the April crop. The live weights of Pisidium and Sialis were
not determined , and they are not included, therefore, in the total under
live weight. Compare with table 15.
A SURVEY OF THE LARGER AQUATIC PLANTS OF
LAKE MENDOTA1
R. H. Denniston
Notes from the Biological Laboratory of the Wisconsin Geological and
Natural History Survey. XVI.
The work included in this report was done during the month
of August, 1912, for the Wisconsin Geological and Natural His¬
tory Survey. The purpose of the investigation was to ascertain
the location and the distribution of the various species of the
larger plants growing in the lake at this season of the year.
Lake Mendota, located at Madison, Wisconsin, covers an area
of 15.2 square miles and has a shore line of about 22 miles. It
is irregularly oval in shape, the east end being considerably larger.
The length is about six miles and the width about four miles.
There are five main points of land which project into the lake,
namely, Picnic Point (station 1), Second Point (station 5), Far-
well’s Point (station 22), Governor’s Island (station 26), and
Maple Bluff (station 29). There are a number of small bays
with marshy shores, and two of considerable size, University Bay
(station 40) and Catfish Bay (station 20). The former is sup¬
plied by a small creek from the University grounds, the second
by the Catfish River and Six Mile Creek. At a number of sta¬
tions, as at Maple Bluff, Eagle Heights, Fox’s Bluff (station 19),
and Baskerville ’s Landing (station 11), the shore is precipitous
and rocky and the bottom slopes off abruptly.
The work was carried on in a small launch and the plants were
gathered with a close-toothed iron rake, with an extensible handle
6 meters long. At most stations, vascular plants were not found
1 This survey was made in cooperation with the U. S. Bureau of Fisheries,
and the results are published with the permission of the Commissioner of
Fisheries.
495
496 Wisconsin Academy of Sciences , Arts , and Letters.
growing beyond a depth of 5-5.5 meters. Where the shore is
protected by points of land from the action of the waves, plants
were found close up to the beach; at exposed points they did not
grow at a depth of less than 1.5 meters.
For convenience the lake was divided into 44 sections or sta¬
tions (see map), and the plants were gathered and recorded at
the different depths for each station.
Pig. 1. Map of Lake Mendota with shaded area along shore showing
distribution of plants. The 6-meter depth is shown by a
broken line. The numbers indicate the stations referred
to in table 2.
The distribution of plants in the lake is governed, as would be
expected, by the conditions at the different stations. Plants along
the exposed shores were few, both in number of species and of in¬
dividuals, while in protected bays the number of individuals was
high.
The character of the bottom also determines the kind and num¬
ber of species to some extent. Chara grows best in the sand, al¬
though Pieters (1) states that in Lake St. Claire, Michigan, it
Denniston — The Larger Aquatic Plants of Lake Mendota. 497
grows more abundantly on clay bottoms. Castalia and Nymphaea
appear to prefer a mur bottom. West (2) found Castalia speciosa
growing equally well in mud or sand in the Scottish lakes.
The Potamogetons, Elodea, Myriophyllum, Najas, and Vallis-
neria grow apparently as well in mud as in sand. Upon the rocks
no flowering plants were found. Only an alga, Cladophora, was
present.
In University Bay a strip of Scirpus lacustris extends along a
bar about 200 meters from shore; behind this bar are found
Ranunculus aquatilis, Nymphaea advena, Castalia odor at a, Lem-
na minor , Lemma trisulca, Ceratophylhcm demersum, Myriophyl¬
lum verticillatum var. pectinatum, Vallisneria spiralis, Wolffla
Columbiana , Elodea canadensis, Najas flexilis, and five species of
Potamogeton.
Another station where plants were found in abundance and
variety was at the outlet of Pheasant Branch Creek (station 10),
at the west end of the lake. Here a few plants of Potamogeton al-
pinus were found and also Utricularia vulgaris, in addition to the
same species found in University Bay, with the exception of
Wolffia which was not found at this station.
A sand bar extends for fully 1600 meters across Catfish Bay.
This is exposed in one small area only. In all other places ex¬
cept at the channel, the water covers the bar to a depth of from
5 to 60 centimeters. There is a passage or channel through the
bar near the east end, where the bottom drops suddenly to a depth
of a little over 5 meters. In this deep hole were found Cerato-
phyllum demersum, Najas flexilis, and Vallisneria spiralis grow¬
ing on a mud bottom.
In the bay between Maple Bluff and the State Hospital (sta¬
tion 28), there was a considerable accumulation of dead vegetable
matter which prevented the growth of vascular plants near the
shore. The same was true to a less extent opposite Morris Park
(station 19).
The prevailing summer wind from the southwest keeps this
matter in the bays at the north side of the lake fairly constantly.
On days when there is little wind, the odor from this decaying
vegetable matter is very disagreeable. In certain Scottish High¬
land lakes (3) great quantities of dead vegetable matter accumu¬
late and prevent the growth of water plants both near shore and
in deeper water. In these lakes, however, there is no unpleasant
odor, as the plant remains are preserved from decay by the pres-
32— S. A. L.
498 Wisconsin Academy of Sciences , Arts , and Letters.
ence of humic acid in the water. This is brought to the lakes by
streams filtering through immense peat beds.
Opposite Governor’s Island the larger plants occur in a nar¬
row zone at a depth of about 1.6 meters. Toward the shore, the
plants are prevented from growing by wave action on the exposed
point, and away from the shore by the stony character of the
bottom.
At stations 34 and 35 a rock bottom limits the lake ward de¬
velopment of vegetation. Outside the bar the lake is shallow for
a distance of more than a kilometer, with a depth of from 1 to 4.5
meters. Here were found Ceratophyllum demersum and Potamo-
geton RicJiardsonii in abundance, their stems reaching nearly to
the surface. Somewhat less abundant were Najas flexilis, Vallis-
neria spiralis , and Potamogeton pectinatus. Inside the bar the
plants were not studied, since they belong rather to the river than
to the lake.
At only one point around the entire lake shore was there a break
in the border of flowering plants, and this was opposite Maple
Bluff (station 29), where the bottom is covered with large rocks.
These rocks had quantities of Cladophora, an alga, clinging to
them.
The total number of flowering plant species found was 19.
Chara was also found on almost all sandy bottoms at all depths
up to 4 meters.
The principal factor determining the depth limit of vegetation
in Lake Mendota is probably that of light. The water of the
lake in the month of August is not at all clear, mainly because of
the immense quantities of plankton in suspension.
Ceratophyllum was found at a depth of six meters. According
to Warming (4), Spermatophyta descend at most to thirty meters.
It may be that these are detached, since, according to Pond (5),
six meters is given by Schimper as the maximum depth for this
group.
Chara and Nitella have been found at a depth of thirty meters.
West (6) distinguishes three zones according to the intensity of
the light received: First, the photic zone, where there is sufficient
light for the development of higher flora: in some lakes this in¬
cludes depths of twelve feet or less, in others depths of as much
as forty feet (Lake Lismore in Scotland). Second, the dysphotic
zone, deeper than zone one. A few higher plant stragglers are
found here. It is normally occupied by members of the lower
De.nniston — The Larger Aquatic Plants of Lake Mendota. 499
cryptogamic flora. Third, the aphotic zone, where no light-de¬
manding organisms are to be found.
Magnin (7) names five zones according to the main plants found
in each. Beginning at the shore, they are: (1) Phragmitetum,
(2) Scirpetum, (3) Nupharetum, (4) Potamogetonetum, (5)
Characetum. The first two zones extend to a depth of 3 meters,
the third to 4 meters, the fourth to 6 meters, the fifth to 12 me¬
ters.
In Lake Mendota there appears very little distinct zonation of
plants. Except in the bays, they are mostly of the submerged
type. Near the mouth of Pheasant Branch Creek, Scirpus oc¬
curred near the shore, farther out were Castalia and Nymphaea,
followed by several species of Potamogeton and Najas. These cor¬
respond closely with the species found in the four last-named
zones of Magnin.
Flowers or fruits were found on all the species of Potamogeton
with the single exception of P. alpinus. Vegetative methods of
reproduction are also common, such as runners, winter buds, and
fragments of stems. All the flowering plants in this list are
perennials with the exception of Najas flexilis.
Two of the larger plants found in the lake are usually not at¬
tached to the substratum, namely, TJtricularia vulgaris in shallow
water, and CeratopJiyllum demersum in deeper water. The lat¬
ter plant often appears to be rooted when the stems become bur¬
ied in the bottom.
Calcium carbonate was found incrusting the stems and leaves of
four of the six species of Potamogeton. Pond (5) states that this
coating probably does not injure the plants and may offer some
protection from aquatic animals.
The five most abundant species, in the order of their abundance
are: Vallisneria spiralis L., Najas flexilis (Willd.) Kostk. &
Schmidt, Potamogeton Richardsonii (Benn.) Rydb., Potamogeton
zosterifolius Schumacher, Potamogeton pectinatus L.
In addition to these there were found in less abundance : Pot¬
amogeton luceus L., Potamogeton alpinus Balbis, Potamogeton
amplifolius Tuckerm., CeratopJiyllum demersum L., Myriophyllum
verticillatum L. var. pectinatum Wallr., Scirpus validus Vahl,
Elodea canadensis Michx., Ranunculus aquatilis L. var. capilla-
ceus DC., Nymphaea advena Ait., Castalia odorata (Ait.) Wood-
ville & Wood, Lemna minor L., Lemna trisulca L., Wolffla Colum¬
biana Karst., TJtricularia vulgaris L. var. americana Gray, CJiara
species, Cladophora species.
500 Wisconsin Academy of Sciences, Arts, and Letters.
TABLE! 1. DEPTHS IN METERS AT WHICH DIFFERENT SPECIES ARE FOUND
0 1 5 3 4 5 0
1. Potamogeton pectinatus . .
2. Potamogeton zosterifolius ..... -
3. Potamogeton Richardsonii . — -
4. Potamogeton lucens . .
5. Potamogeton alpinus . .
6. Potamogeton amplifolius . .
7. Najas flexilis . . .
8. Vallisneria spiralis . — •- -
9. Ceratophyllum demersum - -
10. Myriophyllum verticillatum
var. pectinatum ............. - - - -
11. Scirpus validus . . —
12. Elodea canadensis . . . .
13. Ranunculus aquatilis . .
14. Nymphaea advena . . .
15. Cast alia odor at a . — — —
16. Chara sp . . — —
17. Lemna minor . . .
18. Lemna trisulca . .
19. Cladophora sp . . .
20. Woljfia columbiana . .
21. Utricularia vulgaris
var. americana . .
References
lc Pieters, A. J. The plants of Lake St. Clair. Bull. Mich. Fish Comm.
2. 1894.
2. West, G. S. Flora of Scottish lakes. In “Bathymetrical survey of
the fresh water lochs of Scotland” 1: 166.
3. West, G. S. (Above cited) 1: 162.
4. Warming:, E. Oecology of plants (Eng. transl.), p. 150. Oxford,
1909.
5. Pond, R. H. The larger aquatic vegetation. In Ward and Whip¬
ple “Fresh-water biology”, p. 19 6.
6. West, G. S. (Above cited) 1: 164.
7. Magnin, A. Contributions a la connaissance de la flore des lacs
du Jura. Bull. Soc. Bot. France, 41: CXXV. 1894.
abundant.)
TABLE 2. PLANTS AT DIFFERENT STATIONS
(Abundance indicated as follows: /, small numbers; x, common; *, abundant.)
A QUANTITATIVE STUDY OF THE LARGER AQUATIC
PLANTS OF LAKE MENDOTA1
H. W. Rickett
Notes from the Biological Laboratory of the Wisconsin Geological and
Natural History Survey. XVII.
Introduction
The object of this study is to determine as nearly as possible
the quantities of the various species of attached plants growing
on the bottom of Lake Mendota. Incidentally the data furnish
a basis for comparisons of the growths on different types of lake
bottom and at different depths.
The work was done for the Wisconsin Geological and Natural
History Survey, under the direction of Professor Chancey Juday.
For the identification of the plants collected I am indebted to the
work by Professor R. H. Denniston embodied in the preceding
paper (Note XVI). No work having been done, so far as I am
aware, that is exactly comparable to that which forms the subject
of the present report, the method followed was of necessity origi¬
nal and experimental, and consequently subject to modifications
as the work progressed. Reference may be made to similar studies
on aquatic animal life (4), to quantitative studies of marine fauna
and flora (3,6), and to less exact ecological work on lake floras
(1,5). Many of the data relating to the physical characteristics
of Lake Mendota were obtained from Muttkowski’s work on the
animal life of the lake (4).
1 This investigation was made in cooperation with the U. S. Bureau of Fisher¬
ies, and the results are published with the permission of the Commissioner of
Fisheries.
501
502 Wisconsin Academy of Sciences, Arts, and Letters.
Obtaining Samples
Apparatus. The different parts of the lake were visited by
means of a small rowboat with an attached motor. In this way
it was possible to reach all parts of the lake in a short time, and
to bring back a considerable quantity of collected plants.
In order to obtain quantitative data, it was necessary to gather
plants from measured areas of the lake bottom. After several
more or less unsuccessful attempts to measure under water a plot
of ground which had been stripped of its vegetation, the follow¬
ing method was resorted to : a square frame of galvanized iron
was constructed, measuring m. on a side, and about 7 cm. in
height ; this was lowered to the bottom, and all plants enclosed by
it were gathered. This made it possible to obtain samples all over
the lake from areas of the same size, or differing by simple mul¬
tiples, and having a simple relation to the unit of area used, name¬
ly, 1 sq. m. This method is subject to some error, inasmuch as the
frame may include the heads of some plants whose roots are out¬
side the measured area, and also inasmuch as it is difficult to pluck
and save every plant without spending an unreasonable amount of
time on one sample.
Another source of error lies in the fact that it is difficult to pull
up entire plants under water, and therefore the roots were in
most eases left in the soil. This is more frequently true in muddy
than sandy soil.
As the results must be in the nature of an estimate rather than
an exact calculation, such errors may be considered as compensated
by the large number of samples — 221 — which were obtained.
In water not more than 2 m. deep, the plants were reached by
swimming, using as a guide the cord to which was tied the meas¬
uring frame, and as anchorage a cement block with an inset
handle, let down near the frame. An area of %. sq. m. could be
cleaned off in this way in three or four trips to the bottom, de¬
pending upon the density of the vegetation.
In deeper water, a diving hood was used. This was supplied
with air by a hand pump which could be conveniently carried in
the boat, and descent and ascent in the water were made easily
by an anchor line. Depths as great as 7 m. were visited in this
way.
The collected plants were placed in galvanized iron boxes, with
enough water to cover them, and taken to the laboratory where
Rickett—The Larger Aquatic Plants of Lake Mendota. 503
they could be separated according to the species, weighed, and
spread out to dry.
Method of Collecting. The attached plants of the lake, of which
all but two species are seed plants, are found in a strip along the
shore varying from 100 to 1000 m. in width. The vegetation
covers the bottom, in varying degrees of density, at depths up to
about 6.5 m. in some places. The average limit of the plant zone
is at a depth of about 5 m. In almost all places, growth is con¬
tinuous from this outer limit up to the shore. Near the water’s
edge is a strip of Cladophora,. varying in width from 1 m. to 40
m., extending around the lake in almost all places where there are
rocks to which it may attach itself.
As one follows the shore line around the lake, it is easy to di¬
vide it into stretches which differ in character as regards the na¬
ture of the banks and of the bottom. It is natural to expect that
the vegetation should show corresponding differences in amount
and character, and such is found to be the case. Collections were
therefore made on this basis, the distinctive regions referred to
being designated as stations, of which there were 41. A station
in general represents a region of fairly uniform physical and
floral characteristics. It was originally intended to make collec¬
tions from all these stations, but, owing chiefly to unfavorable
weather, the time was too limited to accomplish this end, and
several stations were omitted. The number of stations (35) from
which samples were obtained, however, includes every type of
growth found in the lake. The shallow water was worked over
first, enough samples being obtained in each station to represent
every type of growth found within that station. Later, descents
were made in deeper water in as many stations as possible. It
soon became evident that, although there is no definite zonation
of plants in regard to the distribution of certain species, the char¬
acter of the vegetation differs as a whole at different depths ; i. e.,
certain species are found to predominate at certain depths which
are not abundant at other depths. This fact has formed the basis
for subsequent handling of the data.
Keferenee may be made to the accompanying map for the iden¬
tification of the various stations, the width of the plant zone at
various points, and the general character of each station. The
designation of the limits of the plant zone is reproduced without
change from Denniston’s map (preceding paper, fig. 1). Wherever
this was checked it was found to be accurate.
504 Wisconsin Academy of Sciences, Arts, and Letters.
It is obvious that the amount of any one species varies greatly
at different times during its growing period. An effort was made
to make collections within a short time from widely scattered
stations, and to choose places where the plants were at or near
maturity, so that results obtained from all stations would be com¬
parable and would indicate the average amount for the summer as
a whole. Plants in shallow water usually reach maturity before
Fig-. 1. Map of Lake Mendota showing stations and limits of the plant
Rickett — The Larger Aquatic Plants of Lake Mendota. 505
those in deeper water, and, partly for this reason, collections from
shallow regions were completed before any were made from
deeper points.
In the cases of the several shallow bays, usually separated from
the main body of the lake by bars (see map), the problem is differ¬
ent. The character of the flora is entirely different, the growth
is very dense, and collection is difficult. The best that could be
done in the limited time available was to estimate by eye the pro¬
portions of various plants or types of growth, and to take repre¬
sentative samples from each.
It was necessary to handle Cladophora separately. The period
of greatest growth of this plant extends roughly over the first
weeks of June, before the other plants of the lake have reached
their full size. This time was therefore spent in a detailed sur¬
vey of the entire shore with reference to the distribution of Clado¬
phora, and in the obtaining of samples. Some Cladophora occurs
on rocky bottoms in deep water, but the amount is too small to
be taken into account.
Data Obtained. With the exception of the two special cases
last described, the following observations were made in the field
in the case of each sample: the number of the station; the num¬
ber of the sample; the date; the depth of water; the distance
from shore ; the character of the bottom ; and the area from which
the sample was obtained. A specimen set of data for one station,
involving several samples, is shown in table 2.
Weighing the Samples
Separation of Species. The samples were brought back to the
laboratory and separated into the various species of which they
were composed. The ease with which this could be done varies
greatly in different cases. Some plants, such as Chara and Cera-
tophyllum, grow in some places very intimately mingled with each
other and with other species, and absolute separation is next to
impossible.
The large number of the sub-samples separated in this way — 730
— is the best argument for the approximate accuracy of the final
results.
Wet Weight. For obtaining the wet weight, an ordinary
household scale, graduated so as to read in fractions of a kilo-
506 Wisconsin Academy of Sciences, Arts, and Letters.
gram, was used. This proved, on calibration with known weights,
to be sufficiently accurate. Weights were read to the nearest 5
grams. The difficulty in obtaining the wet weight lies in the im¬
possibility of separating the surface water adhering to submerged
plants from the contained water. The method followed was to
allow the plants to drain for a short time, and then to shake off
as much of the surface water as possible. Another error in the
weighings is due to the fact that the plants on removal from the
water become matted together so that they sometimes retain a con¬
siderable quantity of sand or mud. For these reasons it is prob¬
able that the wet weights listed in the tables represent values
slightly above the true weights.
The case of the Potamogetons is peculiar. These plants are
heavily incrusted with a deposit of calcium carbonate (2, p. 171).
It is especially noticeable on the broad-leaved forms. An effort
was made to conserve as much as possible of this material but it
was impossible to pluck the plants without disturbing a certain
quantity of it. On shaking the plants, it is dislodged in the form
of fine flakes of considerable size, which, of course, become im¬
mediately distributed through the water. On drying, it takes the
form of a fine powder, which is difficult to conserve.
Dry Weight. After being weighed wet, the samples were spread
out on sheets of paper until air-dry. They were left in this condi¬
tion until the end of the collecting period, so that all had reached
about the same degree of dryness. A number of samples of various
species were then selected and dried in ovens at various tempera¬
tures until a constant weight was reached. It was determined
that after 48 hours at 60° C. further loss of weight was so small —
a small fraction of 1 per cent. — that for the purposes of these de¬
terminations the samples could be considered as dry. Accordingly,
all the samples collected were dried at this temperature for 48
hours, and weighed. Weights were read to the nearest 0.1
gram. It was found necessary to weigh each individual sample
because of the great variation in the ability of different samples
even of the same species to retain water in an air-dry condition,
making it impossible to obtain the air-dry weights and then to
deduct a percentage calculated by oven-drying a small number of
samples. This variation among the samples depends probably to
a large extent upon the different types of bottom from which the
samples come.
Rickett — The Larger Aquatic Plants of Lake Mendota . 507
A few samples were lost before their dry weights were ob¬
tained; the latter were calculated in these cases on the basis of
the average percentage of water obtained from all the other
samples of the species in question.
The method of recording the weights, wet and dry, is illustrated
in table 2.
Calculations
General Averages. The first step in the utilization of the data
obtained was to summate the weights, wet and dry, of each spe¬
cies, and to compute from these figures the average percentage of
water for each species. For this purpose only those samples were
used for which both wet and dry weights had been obtained. The
dry weights of those samples that were lost in drying were cal¬
culated from the average percentage of water for the species in
question. The totals, thus corrected, for all species, were then
summated, and to the total were added the weights of a few sta¬
tions in which it was impossible to separate the various species.
From this final total was determined the average percentage of
water for all species, and, by a comparison with the total area
from which collections were made, an average weight per square
meter. The total area of the plant zone being known (see be¬
low), it was possible to calculate on this basis the total quantity
of plants in the lake. These results are shown in table 3.
Zones. Each station was divided into zones according to
depth, as follows : zone 1, 0 to 1 m. ; zone 2, 1 to 3 m. ; zone 3,
3 to 7 m. This division was based on an examination of the
samples gathered from various depths. It was evident, for in¬
stance, that in water 1 meter or less in depth the prevailing
species were Potamogeton pectinatus, P. Richardsonii, and Vallis -
neria spiralis ; while in water deeper than 1 m., P. amplifolius
predominated, with large masses of Ceratophyllum and Myrio-
phyllum in places. In water deeper than 3 m., vegetation was
for the most part scanty, and was characterized by the absence
of many species found in shallower water. Of course the line
of division between the zones is arbitrary, but the samples were
gathered well within the zone limits.
Weight per Square Meter of Stations. From the actual weights,
wet and dry, of each sample, was calculated the weight per square
meter of each species for that particular spot. The samples within
508 Wisconsin Academy of Sciences, Arts, and Letters.
a station were then grouped together according to the zones in
which they were gathered, and for each zone, within the station,
the average weight, wet and dry, of each species per square me¬
ter was calculated.
Not all zones were thus treated in all stations. In some places,
e. g., stations 1 and 13, the slope of the shore is so steep that there
is practically no shallow zone, the vegetation at the water’s edge
being composed entirely of Cladophora. Not so many samples
were obtained from the deepest zone as from the others, owing
partly to lack of time, and partly to the fact that the growth here
is comparatively uniform, so that fewer samples were necessary
in order to obtain an average.
Weight per Square Meter of Zones. Considering, then, one
zone as a whole, the average weight per square meter was com¬
puted from the values calculated for that zone in each station.
For this purpose, the portions of the zone within the various sta¬
tions were first grouped according to the type of bottom found,
and the average weight per square meter calculated for each type.
Two types of bottom may be distinguished, consisting of sand or
gravel, or of mud. A third type is found only in one place (sta¬
tion 34). Here there is a floor of clean rock, entirely barren.
Only in rare cases is there a clean sand floor, and there are all
gradations from this to one of pure mud, with varying amounts
of gravel or rocks ; the characterization of each station as ‘ ‘ sandy ’ 1
or “ muddy” means, therefore, that it is more sandy than muddy
or vice versa. Within one zone in one station the bottom is prac¬
tically constant, and, of course, there is a gradual transition into
the next type. These differences, with the above noted limita¬
tions understood, are represented on the map.
All three zones were handled in this way. It should be borne
in mind that the grouping of stations according to their soil char¬
acter has reference only to the zone then under consideration;
the zones themselves may be different in this respect within a
single station. The results of these calculations are shown in
tables 4, 5, and 6. The average weight per square meter of all
species taken together is obtained by adding together the figures
for the individual species in each case.
In calculating the average weight per square meter for a whole
zone, no account was taken of station 34, since the area of this
station is so small when compared with the total of that of the
other 40 stations.
Rickett — The Larger Aquatic Plants of Lake Mendota. 509
Total Weights . The total areas of the different zones were ob¬
tained from Mnttkowski ’s work (4, p. 470). For the third zone,
it was assumed that the outer limit of the vegetation corresponds
on the average to the 5-meter contour line; that is, that the area
of plants without this line equals the plantless area within it.
The total weights of the different species in each zone and in the
plant zone as a whole were determined from the weight per square
meter and the total area. These figures are listed in table 7. The
final figure arrived at for the total weight of plants in the lake is
close to that already arrived at in table 3, the difference being of
course due to the fact that samples have different amounts of
weights in the various calculations according to zones which they
represent — the latter being unequal in area.
Comparative Tables. Table 8, derived from the preceding table,
shows the amount of each species, in each zone, and in all zones
together, in terms of the total weights. The relative amount of
each species found in each zone is given in table 9, also calculated
from table 7.
The Shaelow Bays
The data obtained so far have reference only to the large bulk
of the lake flora which is found in the uninterrupted zone that
follows the shore. It remains still to consider the case of Clado-
phora and that of the vegetation of the shallow bays.
Bays of different sizes are found in stations 4, 27, 29, and 30.
In all but that in station 29, the bays are separated from the main
body of the lake by bars. In the first two cases the bars are com¬
posed mainly of sand and gravel, in the last of rocks. In all cases
the ordinary plant zone extends continuously outside the bars.
Within the bars, the growth is of an entirely different character.
In University Bay (station 4), almost all of the species of plants
found in the lake are present in a dense, tangled growth. Col¬
lections were made in the central portion of the bay from the
shore to the bar, at intervals of about 50 meters. In this way,
representative samples of each type of vegetation present were
obtained in a proportion approximating their distribution. From
these samples was calculated the average weight per square meter
of each species for the bay. The area of the bay was measured
on the map by means of a planimeter. The total weight of each
species could then be determined from these figures.
510 Wisconsin Academy of Sciences , Arts, and Letters.
Some species, occurring only in scattered patches, could not be
dealt with in this way. Their occurrence is reserved for later
discussion.
The remaining bays were not so easily handled. The bulk of the
growth is very dense, matted together, and mixed with large
quantities of filamentous algae. Under such conditions, separa¬
tion of species is next to impossible, and the best that could be
done was to estimate the percentage of the total weight of each
species in each sample. Owing to the extremely varied character
of the flora in these bays, it was not possible to designate stations
having each a homogeneous nature, but samples were taken of all
the different types of growth, and the relative amounts of these
types were judged by eye. Using these estimates, the weight per
square meter of the several species was calculated for this type
of growth as a whole, and the total weights were obtained by mul¬
tiplying by the area of the three bays, measured as in the previous
case. Collections were made only in the small bay in station 30.
Lack of time prevented samples being obtained from the other
two bays, and it was necessary for the purposes of calculation to
assume that all three were of the same nature, as was suggested
by their general appearance.
Data for all of the shallow bays are shown in table 10.
Cladophora
Three types of growth of Cladophora were distinguished, the
nature of which will be discussed later. From notes taken in the
field, the distribution of these types was plotted on the map, and
measurement was made of the area of each. The total weight
found in each area was then calculated, as in the cases of other
species, from samples of each type. These data are given in
tables 11, 12, and 13.
Discussion
Species of Plants. A list of the plants of the lake is given in
table 1. This is not to be understood as a complete list of the
attached flora of the lake, since only those types are listed which
were represented in the samples, and these did not include some
of the rarer species or those found only in the inlets or in the
marshes.
Not all the species of Potamogeton were separated from each
Rickett — The Larger Aquatic Plants of Lake Mendota. 511
other. This would have been an impossible task in the time avail¬
able. It was fairly easy, however, to separate P. zosterifolius, P.
amplifolius, and P. Richardsonii. The differences between these
three species are very apparent in the shape and size of the
leaves. The two narrow-leaved species, P. pectinatus and P. in¬
terior, were grouped together. Of these, the former is by far the
more abundant; the second occurs in scattered patches in deep
water. Another species which is found occasionally is P. lucens,
which is a broad-leaved form, but readily distinguishable from P.
amplifolius and P. Richardsonii . Mixed with this there may have
been a little P. alpinus, which has been reported by Denniston in
Lake Mendota.
The plant listed as Heteranthera dubia could not be identified
with certainty. It does not appear in any previous lists of plants
found in the lake. It is found in comparatively small quantities,
and was at first confused with Potamogeton zosterifolius. The
discovery of the fruits in the axils of the leaves proved this to be
an error, but by this time the plant was well past its prime, flowers
could not be found, and identification was impossible. It answers
to the general description of Heteranthera more nearly than to
that of any other plant.
The first ten and the last two species in the list compose the
bulk of the vegetation of the lake. The other species are found
only in scattered patches, chiefly in the shallow bays and mouths
of streams. One plant, Elodea camadensis, is listed although it
was found only in minute quantities in the samples. This species
is abundant in the streams that flow into the lake, especially in
that which empties into University Bay.
In some places there was an abundant growth of floating and
attached filamentous algae (notably several species of Spirogyra).
This was found mostly on muddy shores and in the bays. It was
impossible to separate this from the larger plants, and in some
cases it made the separation of the latter impossible, so that no
account can be given of the amount of this algal growth in the
lake. In other cases, some plants, especially Potamogeton pectina¬
tus and Najas flexilis, were covered with globular masses of a
blue-green alga, Gloeotrichia, which also was impossible of separa¬
tion. These algae, of course, disturb to some extent the values
for the weights of the plants with which they were found.
The Yield of the Plant Zone. Table 14 represents the total
weights of all species, brought together from the ordinary plant
512 Wisconsin Academy of Sciences , Arts , and Letters.
zone (table 7), from the shallow bays (table 10), and from the
Cladophora belt (tables 11, 12, 13). The total yield, in round
numbers, is 18,500,000 kilograms wet, 2,100,000 kilograms dry.
The total area covered by this vegetation is 10,400,000 square me¬
ters, or 1,040 hectares. The yield per unit area is therefore 17,788
kilograms per hectare wet, 2,019 kilograms per hectare dry, — or
14,867 pounds per acre wet, and 1,801 pounds per acre dry.
Reference to table 9 makes it evident that, of the total yield,
nearly one half is found in depths of water of from 1 m. to 3 m.
More than one quarter is found in water shallower than this, and
almost exactly one quarter in deeper water. In terms of weight
per unit area, this means in zone 1 about 16,000 kilograms per
hectare wet, 1,800 kilograms per hectare dry; in zone 2, about
24,000 kilograms per hectare wet, 2,700 kilograms per hectare dry ;
in zone 3, about 13,000 kilograms per hectare wet, 1,500 kilograms
per hectare dry. Converted to pounds per acre, the figures are
as follows: zone 1, 14,000 wet, 1,600 dry; zone 2, 21,000 wet, 2,400
dry; zone 3, 11,000 wet, 1,300 dry.
Comparison of the Quantities of Different Species v Table 8
shows that, of the above stated total weight, almost half the en¬
tire wet weight of plants is made up of Yallisneria. The dry
weight of this species amounts to about one third of the total
weight (compare its relatively high water content, table 3). Of
the remaining species, Potamogeton amplifolius is the most abund¬
ant, composing about one quarter of the total. P. Richardsonii
and P. pectinatus each compose about 10 per cent, of the total
weight, and all the remaining species are present in much smaller
quantities, none forming more than 4 per cent, of the total.
Considering separately the different depth zones, a different situ¬
ation is found in each one, except for the common predominance
of Yallisneria. In the shallow water, the vegetation, exclusive of
Yallisneria, is composed of about equal quantities of P. pectina¬
tus and P. Richardsonii , these two species making up about 30
per cent, of the total. Najas, Ranunculus, and Chara each com¬
pose about 6 per cent. Ceratophyllum and Myriophyllum are
present only in small quantities. In the middle zone, Potamogeton
amplifolius replaces to a large extent the other species of Pota¬
mogeton, and it is here that Ceratophyllum and Myriophyllum are
most abundant. Two plants, Potamogeton lucens and Ranuncu¬
lus , are not found beyond the shallow zone; and one new species,
Rickett — The Larger Aquatic Plants of Lake Mendota. 513
Potamogeton zosterifolius, appears only in the two deeper zones.
In the deepest zone, the number of species present is still smaller.
Chara, Lemna, and Heteranthera are no longer found. Vallis-
neria here forms a lower percentage of the total, and Potamogeton
amplif alius composes about half of the entire flora. An interest¬
ing incident in this zone is the occurrence of several patches
(stations 40, and 41) of the latter plant growing very luxuriantly
and reaching the surface in 4 meters of water. All other
species, except P. zosterifolius , are present in smaller relative
quantities than in the shallower water.
The proportion of these plants varies slightly according as the
wet or the dry weight is under discussion, owing to the fact that
the different species vary considerably in their water content
(table 3). When, however, all species are taken together, as in
table 9, in a comparison between the different zones, this varia¬
tion is corrected, and wet and dry weights form practically the
same percentage of the total.
Variation in Amount and Kind of Vegetation . The data pre¬
sented in the two preceding sections have reference to averages,
which, of course, conceal a great deal of regional variation. The
latter is especially marked in the shallow waters of the lake.
For instance, sample 74, station 8, was collected from an
area of 3 square meters, but yielded a total wet weight of only
100 grams, composed of Najas and Chara. Such almost barren
regions are found here and there in places where there is an ex¬
tensive sandy shallow region. On the other hand, there are places
correspondingly much more productive than the average. Sample
116, station 37, shows a wet weight of 1,300 grams from an area
of 0.3 square meter. The shallow bays also rank high in yield
(table 10), producing many plants, especially emerged plants,
not found elsewhere in the lake. Local variation exists not only
in the amount, but also in the character of the vegetation. Refer¬
ence to table 2 will serve as an illustration. Among the first six
samples of the station, all coming from about the same depth,
there is a marked variation in the species of which they are com¬
posed, though most of them contain the predominating species,
Potamogeton pectinatus and P. Richardsonii.
In water from 1 meter to 3 meters deep, there is much more
uniformity both in quantity and in character within a station,
but between the different stations there are considerable differ-
33 — S. A. L.
514 Wisconsin Academy of Sciences , Arts , and Letters.
ences in the species which make np the bulk of the vegetation.
This is evident from a glance at table 5.
In the deepest waters in which plants are found, a different situ¬
ation exists. Here the flora is made up of relatively few species
uniformly distributed, but it is characterized by the occurrence
of luxuriant growths of Potamogeton amplifolius in more or less
scattered patches. In this zone, as in the shallowest zone, there
are several places where the vegetation is extremely scanty, com¬
posed of Najas or Potamogeton zosterifolius in very small quanti¬
ties.
The Depth of each Species. The abundance of several species
has a definite relation to the depth of water (table 9). Most of
Potamogeton zosterifolius is found in water deeper than 3 meters,
while P. lucens and Ranunculus are entirely limited to shallow
water. About 75 per cent, of the Ceratophyllum and Myriophyl-
lum of the lake is found in water from 1 meter to 3 meters deep.
Almost exactly half of the Vallisneria is also found within the
same limits, and since this plant forms a high percentage of the
total yield of each zone (table 8), it is evident that this zone ex¬
ceeds the other two in total yield.
Several species are limited either to the first, or to the first two
zones, so that a smaller number of species is present in the third
zone. Most of the species, however, show no definite depth limi¬
tations in their occurrence, but grow more abundantly at certain
depths than at others.
Comparison between Different Types of Bottom. In tables 4
and 5 are shown the relative amounts of each species found on
each type of lake bottom. Since the classification of the bottom
into the two types, “ sandy’ ’ and “muddy”, is very approximate,
these figures have not an exact significance, but they may serve
to indicate the kind of soil on which each species flourishes. In
order to classify the plants according to the type of bottom on
which they are most abundant, it is necessary to keep in mind
their relative abundance in the different depth zones (table 9).
Thus, in table 5 (zone 2), Potamogeton pectinatus is represented
as occurring in about equal quantities on both types of soil,
whereas in table 4 (zone 1), about 80 per cent, of this plant is
found in sandy regions. Reference to table 9 shows that 75 per
cent, of this species is found in zone 1. Hence, P, pectinatus may
be classified as flourishing on a sandy substratum. In this way
Bickett — The Larger Aquatic Plants of Lake Mendota. 515
the plants of the lake may be classified as follows: plants flour¬
ishing on sandy soil, Potamogeton pectinatus, P. Bichar dsonii,
Najas, Ranunculus, Chara; plants flourishing on muddy soil,
Potamogeton zosterif olius , P. amplifolius, P. lucens, Ceratophyl-
lum, Myriophyllum. Yallisneria and the plant listed as Heter-
anthera thrive equally well in sand or mud. The remaining plants
are found in the bays, where the bottom consists chiefly of mud.
That the substratum is of importance to aquatic plants in ways
other than merely as an anchorage, is shown by the work of
Pond (7).
The Shallow Bays. There are two large and two small bays,
having distinctive floras. The two small bays, one on either side
of Governor’s Island, are of the same general nature as Catfish
Bay, and a study of one of them (station 30) was made as a type
of all three. The remaining bay (station 4), is of a different
type. It is shallower, the bottom is composed of mud which is
not so rich and deep as that of the other bays, and its flora is en¬
tirely different.
The type of flora represented by Catfish Bay consists mainly of
Potamogeton pectinatus , Yallisneria, and Ceratophyllum, growing
in a dense, tangled mass, and mixed with a considerable growth
of both green and blue-green algae. There are in addition sev¬
eral patches of Castalia and Scirpus and a fringe of Typha grow¬
ing along the shore, as the bay proper merges into a swamp.
In University Bay are found almost all of the species present
in the lake. The amount of the commoner submerged forms is
indicated in the data (table 10), but several species occur only in
patches, and no account can be given of their abundance, save
that these plants form a very small percentage of the vegetation
of the bay as a whole. In this class are Zizania, Nymphaea,
Lemma minor. Scirpus occurs in a fairly definite strip across the
bay on the bar, the area of which could be estimated, so that the
total bulk of this plant could be calculated. University Bay also
is surrounded by a strip of marsh, where occur several typical
marsh plants.
The vegetation in the bays compares more closely with that
of small, shallow lakes of which ecological studies have been made
(4), while the great mass of the vegetation of the lake is of an
entirely different kind.
Cladophora. The growth of this plant is very varied. In gen¬
eral, it is possible to distinguish between three types, as follows:
516 Wisconsin Academy of Sciences , Arts , and Letters.
a very dense and luxuriant growth on tumbled masses of sub¬
merged or half-submerged rocks ; a similar dense growth on rocks
scattered much more thinly over the bottom; and a scanty growth
on small pebbles or single, isolated large rocks.
In the case of the first of these types (Type A, table 11),
samples were collected from individual rocks, a whole rock being
cleaned off and the area covered by Cladophora measured. From
a number of such samples, the weight per square meter of rock
surface was arrived at. By cleaning off all the rocks from a small
section of ground, and by then measuring the area of this ground
and the total surface of all the rocks that came from it, the rela¬
tion of ground surface to rock surface was determined. And by
comparing the average ratio of the surface of a rock to the amount
covered by Cladophora, it was possible to estimate roughly the
relation between a measured area of ground and the Cladophora-
bearing surface upon it. On one square meter of ground, there
were nearly 2 square meters of Cladophora-bearing surface. Twice
the weight per square meter of rock surface is therefore an ap¬
proximation of the weight per square meter of ground surface.
The second type (Type B, table 12) includes samples of very
different values per square meter. Since these differences would
not be readily distinguished in the field, so that the relative area
occupied by each sort could be determined, it was necessary to
group them all together, and to obtain an average. Samples were in
five cases collected from measured areas of ground. Three came from
rocks of average character, representing a large area in which the
total number of rocks was counted. The first two samples shown
in the table came from station 1, which, however, is usually char¬
acterized by growth of Type A. The explanation is that, as the
main crop of Cladophora dies down toward the end of June, the
water of the lake rises slightly, and on the rocks now for the first
time submerged a new crop arises. This, however, is not of so
luxuriant a character as the earlier growth, and for this reason
is not classed under Type A.
Type C, representing those places where there is a very scanty
and scattered growth of Cladophora, is illustrated by one sample,
shown in table 13. From this sample the total weight of Clado¬
phora of this type — relatively a very small quantity — was cal¬
culated.
Cladophora is a significant element in the lake vegetation dur¬
ing only a small part of the growing period. Early in June, the
Rickett — The Larger Aquatic Plants of Lake Mendota. 517
bulk of the plant growth of the lake has not yet reached its full
growth, while at this time Cladophora is at its prime. At this
time, therefore, it forms a considerable part of the whole lake
flora, while afterward it dies rather rapidly, and during the rest
of the summer it is present only in small quantities.
Comparison with Previous Work . Of the similar studies that
have been made on lake floras, that on the vegetation of Oneida
Lake, New York (1), is the most closely comparable. This lake
is more varied in character than Lake Mendota, and a larger num¬
ber of aquatic plants are found there. There is more of the
marshy type of shore vegetation, with numbers of emerged species ;
and the submerged plants are comparatively less abundant, none
being found beyond the 6-foot contour. Mention is made, by the
author, of several lakes in which the plant zone extends to greater
depths. The problem of quantity in this study is approached
from a different angle, the number instead of the weight of plants
per unit area being determined, and the data used directly to cal¬
culate the animal population found associated with the plants. An
exact comparison with the present study is therefore impossible
from the data presented.
Literature Cited
1. Baker, F. C. The productivity of invertebrate fish food on the
bottom of Oneida Lake, with special reference to mollusks.
N. Y. State College For. Tech. Publ. 9. 1918.
2. Birge, E. A., and Juday, C. The inland lakes of Wisconsin. Dis¬
solved gases of the water. Wis. Geol. and Nat. Hist. Survey
Bull. 22. 1911.
3. Jensen, P. B. jStudies concerning the organic matter of the sea
bottom. Rept. Danish Biol. Sta. 22: 1-3 9. 1914.
4. Muttkowski, R. A. The fauna of Lake Mendota — a qualitative and
quantitative survey with special reference to the insects. Trans.
Wis. Acad. Sci., Arts, and Lett. 19*: 374-482. 1918.
5. Pammel, L. H. The vegetation of Iowa lakes. Rept. State High¬
way Comm, on Iowa Lakes and Lake Beds, pp. 162-189. 1917.
6. Pettersen, C. G. J. A preliminary result of the investigation on
the valuation of the sea. Rept. Danish Biol. Sta. 22: 29-32.
1915.
7. Pond, R. H. The biological relation of aquatic plants to the sub¬
stratum. U. S. Fish Commission Rept. 1903: 483-526.
Wisconsin Academy of Sciences, Arts, and Letters.
Table 1.
List of Plants
1. Potamogeton zosterifolius Schumacher.
6. Heteranthera dubia (Jacq.) MacM. (?)
7. Najas flexilis (Willd.) Rostk. & Schmidt.
8. Vallisneria spiralis L.
9. Ceratophyllum demersum L.
10. Myriophyllum verticillatum D. var. pectinatum Wallbr.
11. Scirpus lacustris L.
12. Elodea canadensis Michx. (Philotria canadensis Britt.)
13. Ranunculus aquatilis L. var. capillaceus DC.
14. Nymphaea advena Ait.
15. Castalia odorata (Ait.) Woodville & Wood.
16. Lenina minor D.
17. L. trisulca L.
18. Typha latifolia D.
19. Zizania aquatica L.
20. Ohara crispa L.
21. Cladophora glomerata L.
Table 2.
Specimen Set of Data from a Station (Station IS)
Bickett — The Larger Aquatic Plants of Lake Mendota. 519
Weights
Distances and depths are recorded in meters, areas in square meters, weights in grams,
x indicates a trace.
In recording dates, Roman numerals refer to the months, Arabic numerals to the days
of the month.
Numbers in parentheses are calculated on the basis of the water content for each species
given in table S.
Species are designated by numbers according to their place in table 1.
520 Wisconsin Academy of Sciences, Arts, and Letters.
Table 3
Total Weigjit Collected and Percentage of Water of Each Species
Ail weights are recorded in grams, areas in square meters.
I
Table 4
Average Weight per Square Meter of Each Species , Zone 1
Wet Dry
Wet Dry
Wet Dry
Wet Dry
Wet Dry
Sandy and Gravelly Stations
352 I 53. {
419 44.1
Muddy Stations
Sandy..
Muddy.
69 6.6 !
73.9 19
All Stations
Total .
Average. .
Sandy..
Muddy.
Percentage on Each Type Bottom
Species are designated by numbers according to the places they occupy In table 1. All weights are recorded in grams.
0.4
44.0
340 i 96.0
Table
Average Weight per Square Meter of Each Species , Zone 2
Wet Dry
Wet Dry
Wet Dry
Wet Dry
Sandy Stations
Sandy..
Muddy. .
Species are designated by numbers according to the places they occupy in table 1. All weights are recorded In grams.
Species are designated by numbei
All weights are recorded in gram
*This species (Elodea) probably
Table 6
Average Weight per Square Meter of Each Species, Zone 3
Total Weights of Plants (From tables 4, 5, and 6)
Rickett—The Larger Aquatic Plants of Lake Mendota. 521
Weights are recorded in kilograms, areas in square meters.
522
Wisconsin Academy of Sciences , Arts , and Letters,
Table 8
Percentage of Each Species in the Total Weight (From table 7)
Rickett — The Larger Aquatic Plants of Lake Mendota. 523
Table 9
Percentage of Weight of Bach Species Found in Each Zone (From
Talle 7)
Table 10
Weights of Plants in Shallow Bays
524 Wisconsin Academy of Sciences, Arts, and Letters.
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Bickett — The Larger Aquatic Plants of Lake Mendota. 525
table ll
Cladophora , Type A
Numbers in parentheses are calculated on the basis of the percentage of water (table 3).
Areas are recorded in square meters, weights ini grams, except total weight which is in
kilograms.
526 Wisconsin Academy of Sciences , Arts, and Letters,
Table 12
Cladophora, Type B
Samples from individual rocks
Areas are recorded in square meters, weights in grams, except total weights which are
in kilograms.
Rickett — The Larger Aquatic Plants of Lake Mendota. 527
Table 13
Cladophora, Type G
Areas are recorded in square meters, weights in grams, except total weights which are
in kilograms.
Table 14
Estimated Total Weights of all Species (From tables 7, 10, 11, 12, 13)
Weights are recorded in kilograms.
NOTES ON THE CHEMICAL COMPOSITION OF SOME OF
THE LARGER AQUATIC PLANTS OF LAKE MENDOTA.
I. CLADOPHORA AND MYRIOPHYLLUM
Henry A. Schuette and Alice E. Hoffman
Notes from the Biological Laboratory of the Wisconsin Geological and
Natural History Survey. XVIII.
It is the purpose of this paper to present the results of a
chemical analysis of several of the larger vegetative forms grow¬
ing upon the bottom of Lake Mendota. This information sup¬
plements, in a measure, similar data obtained by one of us1 in¬
vestigating the biochemical nature of the lesser and unattached
forms which live at or near the surface of the lake.
The green material used for this study was hand-picked from
the rocks along the shore of Lake Mendota. It was spread out
to dry and frequently aerated in order to prevent fermentation.
When “ cured”, it was cut up into shreds and spread out again
until thoroughly dry after which it was comminuted in a drug
mill until it passed through a sixty-mesh sieve. The resulting
powder was then subjected to analysis.
Cladophora was selected because of its relative abundance and
comparatively easy harvest. It grows quite abundantly in Lake
Mendota, where it may be found along the shores, having been
washed in by the waves. The plant is attached to rocks in shallow
water by rhizoids. When undisturbed, its filamentous growth fre¬
quently reaches a foot in length.
It is an alga belonging to the Siphonocladiales, an order which
includes plants of a filamentous and usually branched growth.
This plant is made up of large multinuclear cells. Its filaments
have a typical apical growth. Branching takes place from the
1 Schuette, H. A. A biochemical study of the plankton of Lake Mendota.
Trans. Wis. Acad. Sci., Arts, and Lett. 19: 594-613. 1918.
34— S. A. L. 529
530 Wisconsin Academy of Sciences , Arts , and Letters.
upper ends of the cells by the formation of a protrusion. This
is then cut off and forms the first cell of the branch.
Reproduction takes place either sexually or asexually. Asexual
reproduction is by means of biciliate zoospores which arise in num¬
bers from the upper cells of the filaments and escape from these
sporangia by a lateral opening in the cell wall. Sexual repro¬
duction is isogamous. Sexual swarm cells or planogametes are
formed by the division of cells in a manner similar to that of the
formation of zoospores, but in much greater numbers. By con¬
jugation of planogametes in pairs, zygotes are produced which,
after drawing in their cilia, become spherical and then surrounded
by a thick cell wall. After a period of rest, each zygote gives rise
to several swarm spores. These swarm spores develop into new
filaments.
Myriophyllum is more abundant than Cladophora. A quan¬
titative survey of the flora of this lake by Rickett2 shows that
Lake Mendota contains, when calculated by weight of the fresh
plant, about eleven times as much Myriophyllum as Cladophora.
This Myriophyllum is a perennial aquatic plant which supports
a vertical portion of the stem considerably above the water sur¬
face, and on the emersed portion ordinary aerial leaves are
borne. The submerged leaves are finely cut. The leaves are
often crowded and sometimes whorled. The flowers are borne
usually above water in summer.
The methods of analysis used for both the organic and the in¬
organic plant constituents were, in the main, those of the Associa¬
tion of Official Agricultural Chemists3. Inasmuch as these are
recognized and standard methods, no description will be given of
them. All results are calculated on an air-dry basis.
2 See the preceding- paper, Note XVII.
3U. S. Dept. Agr. Bur. of Chem. Bull 107 (revised): 21-24; 38-39. 1912.
Schuette and Hoffman — Aquatic Plants of Lake Mendota. 531
SUMMARY OF ANALYSES
Analysis of the ash of these two plants gave the following re¬
sults, which are stated in percentage of the air-dry weight of the
original sample.
COMPOSITION OF THE ASH
A SECOND REPORT ON LIMNOLOGICAL APPARATUS
Edward A. Birge
Notes from the Biological (Laboratory of the Wisconsin Geological
and Natural History Survey. XIX.
This paper on limnological apparatus continues one published
by Mr. Juday in these Transactions (18:566-592, 1916). It
describes certain pieces of apparatus constructed and employed
by this Survey, chiefly those employed in studying the lake tem¬
peratures.
I. Mud Thermometer (Text figures 1, 2, 3)
In March, 1908, several observations were made to determine
the temperature of the mud at the bottom of Lake Mendota. The
results showed temperatures which rose from 2.6° C. at the sur¬
face of the mud to 8.8° or 9.0° about 3 m. below the surface. For
this purpose there was used an iron pipe about 3 m. (10 ft.)
long and 10 cm. (4 in.) in internal diameter. The pipe was open
at the bottom, covered by a cap at the top; the cap bore a ring
to which was attached the rope for suspending the apparatus.
This instrument was used while the lake was covered by ice. It
was lowered to the bottom by a rope which was attached to a
windlass ; and it was driven into the mud by a weight which rested
on the top of the pipe when it was lowered. The weight was
raised by means of two small lines and then dropped, sliding down
the rope and striking the top of the pipe; and the pipe with the
contained mud was then drawn to the surface and laid on the ice.
At regular intervals along its side there were small holes through
which thermometers could be pushed into the mud so as to ascer¬
tain its temperature. The mud is a fine-grained, pasty material,
through which water passes very slowly, practically not at all
during the few minutes necessary to raise and take it from the
533
534 Wisconsin Academy of Sciences , Arts , and Letters.
water ; the temperature of the mud could therefore be ascertained
with sufficient accuracy.
By this method, however, it is not possible to determine
the depth below the surface of the mud from which is derived any
Fig. 1. The mud thermometer on the ice. This is the first one made,
which was 3.5 m. long, with a hammer in one piece. The insulated
wires and rope are seen attached to the top. The hammer with its two
lines can also be seen. The thermometer is driven into the mud as far as
the point where the hammer rests.
given portion of the column of mud in the pipe, since the column
is always shorter than the depth to which the pipe is driven. The
upper strata of the mud are compressed, and as the pipe is driven
down the mud is partly displaced and partly forced up into it.
The deeper the pipe is driven the greater is the proportion of
Birge — A Second Report of Limnological Apparatus. 535
displacement, and the greater the difference between the height
of the column of the mud in the pipe and the depth to which the
pipe is driven. When, therefore, in 1915 it appeared advisable
to ascertain the annual heat exchanges of the mud, a special ap¬
paratus had to be devised.
This apparatus is an electrical resistance thermometer, which
is mounted on the end of an iron pipe 5 cm. (2 in.) in diameter
and about 4.7 m. (15.5 ft.) long. The case of the thermometer is
of brass, about 9 cm. long and 5 cm. in diameter. It is conical
at the end and provided with a steel point. The resistance coil
is contained in the cavity of the conical tip and is sealed into
place with water-tight cement. The whole thermometer is con¬
nected to the iron pipe by a piece of wood about 25 cm. long, so
as to interpose a non-conducting material between the thermome¬
ter and the pipe. The entire instrument is about 5.0 m. long, and
it can be driven into the mud to that depth.
The resistance coil is connected with the surface by an insu¬
lated cable which is carried up through the pipe. The resist¬
ance is read by the box of a Whipple thermophone, whose scale
is graduated into degrees centigrade. The temperature of the
mud, therefore, is read directly in degrees.
This pipe with the thermometer at its tip must be driven into
the mud. From the top of the 5.0 cm. pipe extends a smaller
pipe about 3.5 cm. in diameter and about 120 cm. long. The
insulated cable from the resistance thermometer passes through
this pipe, and to its upper end is attached the stirrup by which
the whole apparatus — over 6 m. long — is suspended from this
pipe. On this smaller pipe slides the hammer, a cylindrical iron
weight of about 10 kg. The hammer is made of two pieces (text
fig. 3) so that it can be put into place around the pipe after the
thermometer is suspended in the water from its rope, and it can
be removed again before the instrument is taken out of the wa¬
ter. When the two pieces of the hammer are in place on the pipe
they are held together by two long pins (text fig. 3, A) which
are passed through holes. These pins are longer than the diame¬
ter of the hammer; they are provided with a joint about 3.0 cm.
from the end so that the tip of the pin turns down after it has
been put in place. This arrangement prevents the pin from being
jarred out when the hammer is worked. Two ropes are attached
to lugs (text fig. 3, B) on opposite sides of the hammer, by which
it can be raised and dropped so as to slide down the smaller pipe
and strike the top of the larger one.
536 Wisconsin Academy of Sciences , Arts, and Letters.
Fig. 2. The mud thermometer suspended from the tripod through
a hole in the ice. The hammer appears just above the ice. The insulated
cable leads to the thermophone box which is on the sled.
In the first form of the thermometer (text fig. 1) the hammer
was a simple cylindrical iron weight sliding on the smaller pipe.
In this form the hammer is permanently attached to the ther¬
mometer and constitutes a source of some difficulty in handling it,
especially when the thermometer is used in a launch. A heavy
weight near one end, and that the upper end, when the instru¬
ment is put into place, increases the difficulty of handling it.
When the thermometer is used through the ice, it is suspended
from a tripod composed of three pieces of wood, 8 ft. long and 2
in. by 4 in. These are connected at the top by a chain passed
through a hole in each piece (text fig. 1 and 2). The launch from
which the thermometer is used during the open season is also
provided with a frame from which the thermometer can be sus¬
pended. The thermometer is suspended by a %-in. braided cot¬
ton rope. A twisted rope can not be used, since, when the ther¬
mometer is suspended from such a rope, it turns and winds up
together the main rope, the wire cable leading to the galvanome¬
ter, and the two ropes which are used for working the hammer.
With the braided cotton rope we have no trouble from this cause,
though the instrument has been used in water as deep as 25 m.
Birge—A Second Report of Limnological Apparatus. 537
In taking the temperature of the mud, the depth of the water
is first exactly determined by means of a wire sounding line, and
the depth is marked on the insulated cable connected with the
thermometer. Since this cable is not under any strain, it is easy
to know the exact depth to which the thermometer is lowered,
and it is easy to he certain that the thermometer has been low¬
ered so that its point just reaches the mud before beginning the
readings of temperatures. Readings of the mud are usually taken
at half-meter intervals, though other distances can readily be
used. The change of temperature is very considerable, often as
much as 2.0° or even nearly 3.0° in a half meter and since the
Fig. 3. Hammer for mud thermometer. A, Pin for holding the two
parts of the hammer together. B, Lug for attaching ropes by which
the hammer is worked. The hammer itself is about 9 cm. long and about
6.5 cm. in diameter.
538 Wisconsin Academy of Sciences, Arts , and Letters.
mud in contact with the thermometer is chilled or warmed by the
brass point, it is necessary to drive slowly and to allow the in¬
strument to acquire the temperature of the mud as it goes down,
so that the difference between mud and thermometer will be mini¬
mal when the depth is reached at which the reading is to be taken.
Our regular method has been to drive the thermometer down to
a point about 10 cm. short of the depth desired. We then wait
until the thermometer has acquired the temperature of the mud
as nearly as possible, and then slowly drive it down to the full
depth. In this way the final rise or fall of the temperature is
small.
This thermometer was used during the winters from 1916-17,
to 1919-20 through the ice. Observations were made during the
summers of 1918, 1919, and 1920 from a launch, so that records
were secured of the gain and loss of heat by the mud during the
open season. For such observations days were chosen when there
was little wind. The launch was secured by four anchors put
out with long ropes in different directions, and there was no diffi¬
culty in keeping the boat in one spot or in working the instru¬
ment from the launch.
The thermometers first made were used without accident — one
of them for three years — until March, 1919. In that month both
were lost in trying to extract them from the mud at the same sta¬
tion — 12 m. depth of water. The mud here seems exceptionally
sticky, and the thermometer was pulled out with difficulty. The
hammer was used to help the process, by pulling it up so as to
strike from below on the fastening of the stirrup to the top of the
small pipe on which the hammer slides. In both cases the top
of the pipe broke off. In constructing a new instrument, care has
been taken to use extra heavy pipe so as to avoid similar acci¬
dents in the future. Steel pipe, if obtainable, would probably be
even better.
Table 1 is given in order to show the temperature of the mud as
taken by the mud thermometer in winter, spring, and summer,
in relatively shallow water.
Birge — A Second Report of Limnological Apparatus. 539
TABLE 1
Temperature of mud. Lake Mendota, 1910. Depth of water, 8 meters. “Surface” means
temperature of water at surface of mud.
II. Apparatus for Determining Transmission and Absorption
of Radiation by Lake Water
Our attention was first called to this line of investigation by
an experiment recorded by Forel (Le Leman 2: 294). He placed
a thermometer with blackened bulb in a bottle, lowered it to the
depth of 1 m. below the surface of the lake, and exposed it to the
sun. He noted the rise of the temperature which came before
the reading of the instrument remained stationary. It occurred
to us that by using a black-bulb thermometer in vacuo and ex¬
posing it to the sun for known intervals of time and at different
depths, some notion could be gained of the relative amount of
heat absorbed by various strata of water.
A. Black-Bulb Thermometer in Vacuo (Plate XXXIX,
figures 1, 2)
Experiments were first made in 1901. A black-bulb minimum
alcohol thermometer in vacuo was employed. This instrument
could be raised from small depths to the surface and read with
no appreciable change of the readings during the process. It was
attached to a frame so that it was nearly horizontal when low¬
ered into the water, and the bulb was protected by a sliding cover
which could be operated by cords leading to the surface.
540 Wisconsin Academy of Sciences, Arts, and Letters.
The thermometer was lowered to the proper depth — say 1 m. —
and allowed to remain until it had acquired the temperature of the
water. In order to ascertain this fact, the instrument was drawn
to the surface and examined from time to time until the ‘index
ceased to fall When it was finally returned to its place, the slide
was drawn from the bulb; the thermometer was exposed to the
sun; after five minutes it was drawn to the surface, and the rise
of the alcohol above the index was noted. The instrument was
then lowered to another depth — say 2 m. — and the process was
repeated.
This instrument was successfully used by both Mr. Juday and
myself on a considerable number of lakes ; but it was very slow in
its operation, requiring 15-20 minutes for a single reading. Many
series of observations were lost because of clouds coming on dur¬
ing the long time required even for two readings. In all cases
the lapse of time was so great as to give the sun ample opportunity
for change.
The second instrument was a great improvement on the first.
Its centra] part is a sensitive black-bulb mercurial thermometer
with a scale 25 cm. long extending from -2° to 32°, divided to
0.2° C. and mounted in vacuo. It is fastened into a frame 65 cm.
long and 10.5 cm. wide, made of 12-mm. brass tubing. On this
frame is a sliding cover for the bulb, operated by cords which
lead to the surface. The thermometer is read from the surface
by a telescope inserted into a carrier which is attached to the
top of a brass tube 5 cm. in diameter. This tube is composed of
pieces of tubing, each about 85 cm. long; these can be screwed
together with water-tight joints and are added as the depth of
observation increases. The bottom joint of the tube is screwed
into a base piece which has a glass center. This base is movable
on the frame so that it can be fastened securely above that part
of the thermometer scale which corresponds to the temperature of
the water.
Two persons are employed with this instrument, one to read
and one to record. The temperature of the water at the point
of observation is first ascertained by an ordinary deep-sea ther¬
mometer. Ice water is poured over the vacuum bulb, and the
black-bulb thermometer is thus cooled down a degree or more be¬
low the temperature of the water. The instrument is then lowered
into place, and after the outer bulb has acquired the temperature
of the water the slide is drawn and the thermometer is exposed
Birge — A Second Report of Limnological Apparatus. 541
to the sun. The readings are taken on each side of the tempera¬
ture of the water in order to eliminate the effect of loss or gain
of heat due to difference bewteen the thermometer and the sur¬
rounding water. At the proper point the observer notifies the
recorder, who starts a stop-watch. Each 1/5 degree of rise of
temperature is then called, and the recorder notes the correspond¬
ing number of seconds. When the mercury has risen as much
above the temperature of the water as it was below it at starting,
the recorder is notified and stops the watch. The important record
is the total number of seconds between starting and stopping with
the total rise of temperature in that time. The smaller intervals
noted show whether the rise during the period is approximately
uniform. The total exposure varies ordinarily from one-half min¬
ute near the surface to two minutes, or even three or more minutes
at depths of 2 m. to 5 m.
A good deal of work was done with this instrument, and it was
regarded as decidedly successful. It was soon replaced by the
electrical instrument to be described later; and therefore it was
not employed so extensively as had been expected when it was
constructed. I describe it so fully because I believe that it may
easily find a useful place in limnology. It is less complex than
an electrical instrument. It can be used without trouble from any
kind of a boat or anywhere on a lake if the boat is anchored. It
is easy to be sure that the bulb is properly exposed to the sun,
and the instrument is reasonably rapid and accurate. It has been
used without difficulty to depths of 5 m. in inland lakes, though
at depths lower than 2 m. or 3 m. the rise is so slow as to make
it inadvisable to go to such depths except in very transparent
lakes. The ordinary small waves of an inland lake do not cause
trouble, but the instrument can not be used in a rough sea. We
attempted once to use it on Lake Superior when the water was
smooth and there was only a slight swell, not very noticeable when
the boat was moving. When, however, the boat was anchored, it
lay parallel to the swell and rolled so that the eye could not be
kept at the telescope.
In such an instrument as this it is not practicable to employ
the method of alternate warming and cooling and noting the times
of both processes. The glass of the vacuum bulb and the water
about it behave differently toward the light rays from the sun and
the longer heat rays emitted by the black bulb of the thermometer.
542 Wisconsin Academy of Sciences , Arts, and Letters.
In the process of cooling the thermometer lost per minute about
1/18 of the difference between its temperature and that of the
surrounding water. Its cooling, therefore, at temperatures near
that of the water was very slow. If observations are taken on both
sides of the temperature of the water as indicated above, no cor¬
rection is needed for loss or gain from this source.
The record of a single series of observations is given (table 2)
in order to show the kind of results which may be expected from
the black-bulb thermometer.
TABLE 2
Observations with black-bulb thermometer in vacuo Lake Mtendota, September 5, 1912
These observations show that 27 per cent, of the radiation pres¬
ent at a depth of 50 cm. was transmitted to the depth of 150 cm.,
and that 28 per cent, of that at 100 cm. was found at 200 cm.
The transmission measured by the pyrlimnometer between 11:00
and 11 : 15 a. m. was 26 per cent, between 50 cm. and 150 cm.,
and 30 per cent, in the 100-200 cm. interval.
B. The Electrical Pyrlimnometer (Plate XXXIX, figures 3, 4;
Plate XI, figure 5; text figures 4, 5)
The term pyrlimnometer has been recently devised as the name
for an instrument which this Survey has used since 1912. It is
essentially an arrangement of electrical thermal couples for meas¬
uring the intensity of solar radiation below the surface of a lake,
and the name given above is intended to indicate these facts.
The Receiver. The instrument was designed for this Survey
by Professor C. E. Mendenhall, of the department of physics of
the University of Wisconsin, and was constructed by Mr. J. P.
Foerst, mechanician of the same department. The radiation of
Birge—A Second Report of Limnological Apparatus. 543
the sun is received by 20 iron-constantan thermal couples, whose
junctions are covered by blackened silver disks about 1.5 mm. in
diameter. They are arranged in two series of ten each, making
a double row about 20 mm. long and 3 mm. wide. They are cov¬
ered by a hemispherical glass dome about 65 mm. in diameter,
which is cemented into the top of a brass box about 75 mm. square
and 20 mm. thick. This box is made of two pieces of brass closely
fitted together so as to exclude moisture. In a receptacle on one
side of the box is placed some phosphorus pentoxide, so that all
moisture may be removed from the air within the box. From
one side of the box leads a tube about 18 cm. long, into which are
sealed the connections of the wires from the thermal couples and
the wires of the cable leading to the galvanometer.
This box containing the thermal couples is mounted on a frame
46.5 cm. long and 15.5 cm. wide, made of solid brass bars 12.5 mm.
square. The frame is therefore heavy, weighing about 2 kg. It
is made long so as to carry the lines by which it is suspended in
the water to such a distance that their shadow will not be likely
to fall on the thermal couples. It is made heavy so that its
equilibrium will not be disturbed by opening and closing the cover
of the thermal couples in order to admit or exclude the sun. This
cover is hemispherical and made of thin brass; it turns on an
axis attached to the frame and is operated by a pinion on the axis
and a rack which slides on one of the longer sides of the frame.
The rack is pulled back and forth by two cords, one attached to
each end. These pass through small pulleys on the frame and so
lead up through the water to the boat. Two small spirit levels
are attached to the frame at right angles to each other so that
the instrument may be held in a horizontal position. When in
use this receiver is suspended from a small braided rope. From
each corner of the frame a strong cord about 75 cm. long extends
to the rope. Near the upper end these cords pass through a brass
tube about 5 cm. long. This is fastened to the cords, which are
cemented into it, and on each side is attached an eyelet through
which passes one of the cords for operating the shutter. This ar¬
rangement prevents these small cords from becoming entangled
with the various parts of the receiver and frame or twisted with
the suspension cords.
544 Wisconsin Academy of Sciences, Arts, and Letters.
Fig. 4. Galvanometer of the pyrlimnometer.
Fig. 5. Small receiver for electrical pyrlimnometer, about full size
This receiver shows the form used in the first year, with flat glass cover
for the thermal couples instead of the dome as shown in Plate XL,
figure 5. This receiver was later used for measuring the absorption of
radiation by snow and ice and by filtered water in a tank.
The Galvanometer. The electrical currents set up by the sun
in the thermal couples are measured by d’Arsonval galvanometer.
This is of the marine type in which the suspended coil is attached
to a wire under tension, so that it is less sensitive to the motion of
the boat. As used at present, the galvanometer is kept on shore;
but the type of construction makes it readily portable without get¬
ting out of adjustment.
The appearance of this instrument is seen from text figure 4.
Its base is 24 cm. long, 17.2 cm. wide at the front, and 8.5 cm.
wide at the back; its height is 17 cm. The scale is in the front
of the box and is 16 cm. long, divided into millimeters. The figure
shows the windows by which the scale is illuminated.
The galvanometer is provided with a shunt coil, and when this
is included in the circuit the swing of the galvanometer is about
one half as great as when the shunt is cut out. It is therefore
possible to use the galvanometer for determining the full intensity
of the sun. A rate of 1 cal. cm.2 min. would cause a deflection
Birge — A Second Report of Limnological Apparatus. 545
of about 90 divisions of the scale if the shunt coil is in the cir¬
cuit.
The mirror and coil are contained in the tube which projects
from the top of the galvanometer at the rear. It can be drawn
out and the coil locked into place for transportation.
Method of Using the Instrument. In all of the earlier work
with the machine the galvanometer was taken on the boat and
operated there. Two persons are required to use it, one to operate
the receiver and the other to read the galvanometer. This is little
affected by the ordinary motion of the boat. It is not seriously
disturbed by motion in the line of its main axis, but responds
quite readily to a motion at right angles to this. It can be used
even in a moderate breeze by bringing the boat to the windward
side of the lake and anchoring it head to the wind, since the short
pitching motion under these conditions does not disturb it. But
such side motion as is caused for instance by the swell from a
launch, even coming from a distance of a mile, sets the mirror
swinging in a way that quite stops reading until it becomes quiet
again. In the summer, therefore, it was often almost impossible
to get readings on Lake Mendota, since there are at all times
launches in motion somewhere on the lake and their swells are
propagated over the entire surface with enough energy to disturb
the galvanometer. It was found better, therefore, to do most of
the work with the aid of an insulated cable, 100 m. long, and thus
keep the galvanometer on shore. The amplitude of the swing of
the galvanometer is practically the same whether it is connected
with the receiver directly or through the longer cable. The ease
of operating is much greater and the readings are more accurate
when the galvanometer is on a stable foundation rather than in
the boat. There is no reason why a longer cable than 100 m.
should not be used, but in most of our small lakes 100 m. is quite
long enough to get outside the weeds and to give sufficient depth
of water. The instrument has been operated to depths of 10 m.,
though commonly not below 5 m. In most of our lakes the de¬
flection at 5 m. depth is very small — less than one per cent, of the
deflection in air. Ordinarily it is quite sufficient to extend read¬
ings to the depth of 3 m. In more transparent lakes greater depths
must be reached.
The person operating the receiver lowers it to the proper
depth, fastens the rope, and takes the opening cord in one hand
35— S. A. L.
546 Wisconsin Academy of Sciences, Arts, and Letters.
and the closing cord in the other. The receiver is so suspended
that the long side of the frame is turned toward the sun and the
side with the opening and closing cords is next to the boat.
At the signal of the recorder, the operator pulls the cord which
turns back the shutter so that the thermal couples are exposed to
the sun. The full deflection is reached in a few seconds. The
recorder notes it and then signals to close the cover. When the
galvanometer has returned to zero, the reading is repeated. Two
or more swings of the galvanometer are ordinarily recorded from
each depth. Readings are usually made at half -meter or meter
intervals. There is no difficulty, but little profit, in using smaller
intervals. However, the hemispherical form of the glass cover
of the thermal couples is not suitable for use in determining ab¬
sorption by thin strata of water that lie close to the surface.
With this instrument readings can be made very rapidly and
very accurately. It is not hard to estimate to tenths of a divi¬
sion of the scale. With the coils in use one division of the scale
equals about 0.011 calories per square centimeter per minute.
This is when the shunt is included in the circuit. When the shunt
is cut out, the value of one division is 0.0059 cal. cm,2 min.
Ordinary errors in estimating tenths of a division make little
difference in the results.
A portion of a set of readings is added (table 3) in order to
show the general results obtained from the pyrlimnometer.
The pyrlimnometer was first used in 1912. In its original form,
the glass cover of the thermal couples was flat. Two receivers
were made, one to read in the water and the other to read in the
air. The plan was to read the instrument in the air alternately
with that in the water. This method, however, proved unneces¬
sarily complex. If the sun’s radiation is changing rapidly, as
by haze or drifting clouds, the changes are too rapid to be fol¬
lowed by the instrument. If the sun is fairly steady, as on an
ordinary sunny day, the second instrument adds little to accuracy
and much to trouble.
The regular receiver was built in 1913 and was then
furnished with the hemispherical glass cover shown in the figure.
This change made it unnecessary to compute the varying amounts
of radiation reflected from the glass at different altitudes of the
sun.
The smaller receiver (text fig. 5) was kept in its original form
and used in determining the absorption of the sun’s radiation by
Birge — A Second Report of Limnological Apparatus. 547
TABLE 3
Observations with pyrlimnometer. Hector Point, Seneca Lake, New York, August 1, 1918
snow and by water which has been filtered or otherwise treated,
and whose transmission of the sun’s radiation can be studied in
a tank.
III. The Kemmerer Water Bottle, Modified Form (Plate XL,
figures 6, 7)
In 1910 this Survey had a water bottle constructed after the
well known design of Petterson. This was loaned at a later date
to the United States Bureau of Fisheries for investigations by Dr.
G. I. Kemmerer who used it in the deep lakes of the western
United States. It was lost in one of those lakes by the breaking
of the line, and in order to go on with his work Dr. Kemmerer
devised and had made a simple form of bottle, employing a brass
tube with rubber stoppers for its ends. The parts were brought
548 Wisconsin Academy of Sciences , Arts, and Letters.
together by springs and there was a positive release not unlike
that of the Petterson bottle. This bottle has been described by
Dr. Kemmerer in his report to the Bureau of Fisheries.
The Survey needed in 1918 another bottle and one was devised
and made by Mr. J. P. Foerst, mechanician of the department of
physics, University of Wisconsin. It resembles the Kemmerer
bottle in using a brass tube and rubber stoppers, but differs in
employing gravity instead of springs to bring the parts together
and in using a friction release instead of a catch.
The tube, or bottle proper, is 43 cm. long and 6.7 cm. (2.5 in.)
in diameter and holds about 1400 cc. Through its center extends
a small tube 56 cm. long, on which the movable parts slide. The
suspension line is passed through this tube and is knotted below
it. When the parts of the bottle are separated, the upper stop¬
per is held in place by two small pins which are pressed by springs
into a shallow groove near the top of the small inner tube. The
larger tube when raised is held in a similar way by pins attached
to one of the two wheel-like guides inside the large tube. These
guides keep the large tube centered on the small one and so con¬
trol its descent to the lower stopper.
The ends of the tube are ordinary rubber stoppers. Each is
held between two brass discs screwed together, so that the stop¬
per can be easily removed when it becomes hard. A short, heavy
brass cylinder is attached to the top of the upper stopper in
order to give it more weight and so to make sure that it will re¬
lease the large tube and cause the bottle to close. Through each
stopper passes a small tube on one side ; this ends in a rubber
tube closed by a spring clip. The lower tube serves to allow the
water to flow from the bottle ; the upper one serves to admit air
as the water flows out.
On the top of the whole apparatus when in use there is placed
a rubber stopper to receive the blow of the messenger and to pre¬
vent this from battering the brass of the tube or the stopper.
The operation of the bottle and the method of release are plain
from the figures.
Such a bottle operates very satisfactorily. We have found
it simpler than the original Kemmerer bottle and equally satisfac¬
tory. It can not take the place of more elaborate bottles for
certain types of investigation, but it is entirely adequate for most
purposes of lake studies. It is convenient, light, and inexpensive,
Birge — A Second Report of Limnological Apparatus . 549
either in its original form as designed by Kemmerer or in that
which we have used.
IV. Ekman Dredge with Modified Release (Plate XL,
figures 8, 9)
This dredge was built according to the figures and description
of Ekman1. It is used to obtain mud from the lake bottom. We
have employed it in two sizes; in one of these the box was about
15.3 cm. (6 in.) square, and in the other about 23 cm. (9 in.)
Both worked equally well, but of course the smaller form is the
more portable. The body and jaws of the dredge are made of
stout sheet brass ; the covers are made of very thin spring sheet
brass so attached that they open readily as the dredge descends
and permit the water to flow through it, but close when the dredge
is drawn up.
As is well known, the purpose of the Ekman dredge is to obtain
a sample of mud from a known area of the bottom, so that the
animals living in it may be determined. Thus information may
be secured regarding the kind and number of the bottom popula¬
tion. The dredge is lowered until it rests on the mud; the jaws
are then released, which bite into the mud and ordinarily secure
enough to fill the entire dredge about two thirds full. It can be
operated only on a soft bottom, since the springs are not strong
enough to force the jaws into sand bottom, nor will the jaws close
properly if sticks or stones or large animals like clams are pres¬
ent. But it is most successful and useful in exploring that great
area of an ordinary lake bottom which is covered with lake mud
and vegetable debris, and the Survey has made very extensive use
of it in such studies.
Ekman used a form of release held in place by friction and
operated bj^ a vigorous pull on the line after the dredge had
reached the bottom. As he states, great care had to be exercised
to prevent the machine from “going off” prematurely. We there¬
fore adapted to the instrument a simple form of release like that
described by Mr. Juday in his former paper. Its construction
is clear from the figures, as also is the way in which the jaws are
released by a brass messenger sent down the line.
1 Ekman, S. Neue Apparate zur qualitative!! und quantitative!! Erfor-
schung der Bodenfauna der Seen. Internat. Rev. Ges Hydrobiol. u. Hydrogr.
a: 558. 1011; 7: 165.1915.
550 Wisconsin Academy of Sciences , Arts , and Letters.
V. Water Sampler for Bacteria (Text fig. 6)
This instrument is modified from one described in 1892 by Dr.
H. L. Russell, as used by him for work in the Mediterranean Sea.
It has received more than one modification in the course of the
Fig. 6. Water sampler for bacteria. A, Messenger. B, Line. C'lf C2,
Movable jaw. D, Spring clamp for holding sample tube. E, Break¬
ing pin.
work of this Survey. Its latest form is that devised by Mr. Frank
C. Wilson and fully described by him in the Journal of Bacteri¬
ology, vol. 5, pp. 103-108 (1920). This article also gives reference
to the papers which describe other forms of water samplers for
bacteriological purposes.
Birge — A Second Report of Limnological Apparatus.
551
The construction of the sampler is plain from the figure. The
base of the apparatus is made of heavy sheet brass, about 30 cm.
by 7 cm. The sample tubes are ordinary hard-glass test tubes,
2.5 cm. by 20 cm., fitted with a rubber stopper which is pierced
with one hole. The tube is held in place by a rubber-covered
spring clamp. The sample tube is sterilized and then exhausted
with an air pump, and the small inlet tube is sealed off with the
blow pipe. The point of the inlet tube rests between two jaws
and is broken by the blow of the brass messenger which is sent
down the line and strikes the movable jaw.
In all of our work we have used sample tubes closed by rubber
stoppers. We have found these entirely satisfactory for depths
up to those of Lake Mendota — 25 meters — and we have used them
without difficulty up to a depth of 60 m. in Green Lake. In
deeper water a glass sample tube can be used, which has been
drawn out and sealed; but unless the depth makes such an ar¬
rangement necessary, the stoppered tube will be found much more
convenient.
TRAN S. WIS. ACAD., VOL. XX
PLATE XXXIX
DIRGE — LIMNOLOGICAL APPARATUS
TRANS. WIS. ACAD., VOL. XX
PLATE XL
BIRGE— -LIMNOLOGICAL APPARATUS
THE ROTIFER FAUNA OF WISCONSIN
H. K. Harking and F. J. Myers
Notes from the Biological Laboratory of the Wisconsin Geological and
Natural History Survey. XX.
Introduction
Some years ago Mr. Chancey Juday, biologist of the Wisconsin
Geological and Natural History Survey, suggested to the writers
the advisability of undertaking a study of the rotifer fauna of
Wisconsin. As few states of the Union are more favored by na¬
ture with opportunities for collecting material, we gladly promised
to do all that we could to make a preliminary survey, sufficiently
detailed to serve local students as a basis for continuation of the
work. Mr. Juday enlisted the cooperation of Dr. E. A. Birgc,
president of the Board of Commissioners of the Survey and of the
University of Wisconsin. We take this opportunity to express
our gratitude to Mr. Juday and Dr. Birge, not only for essential
material aid, but for their deep personal interest and ever ready
cooperation, without which our part of the Avork could not have
been done.
While field work is still in progress, so much material has al¬
ready been accumulated that a preliminary report is desirable.
We hope eventually to be able to publish figures and complete de¬
scriptions of all the rotifers found in Wisconsin. This is made
all the more desirable by the fact that the North American rotifer
fauna has never been made the subject of an intensive study, and
the determination of material is consequently an exceedingly
laborious undertaking, necessitating access to, and study of, publi¬
cations from all over the world. Moreover, a new standard for
553
554 Wisconsin Academy of Sciences, Arts, and Letters.
rotifer work was established by de Beauchamp when he published
his Recherches sur les Rotiferes in 1909, which suddenly made the
earlier literature on the group all but worthless. We are endea¬
voring as far as possible to follow the path which he has pointed
out in the re-description of the central group of the Notommatid
rotifers which follows the list of described species so far found in
Wisconsin. When preparations were begun for this paper we
planned to include all the genera of Notommatids, but the under¬
taking proved even more laborious and time-consuming than we
had anticipated, and the number of plates required is so large
that it was decided to divide the paper into two parts, of which
this is the first instalment.
List of the Rotifers of Wisconsin
Order PLOIMA
Notommata aurita (Muller).
Notommata cerberus (Gossee).
Notommata collaris Ehrenberg.
Notommata contorta (Stokes).
Notommata copeus Ehrenberg.
Notommata cyrtopus Gosse.
Notommata galena new species.
Notommata lenis new species.
Notommata pachyura (Gosse).
Notommata peridia new species.
Notommata pseudocerberus de Beauchamp.
Notommata saccigera Ehrenberg.
Notommata silpfia (Gosse).
Notommata tripus Ehrenberg.
Notommata trypeta new species.
Proales decipiens (Ehrenberg).
Proales sordida Gosse.
Proales parasita (Ehrenberg).
Pleurotroeha petromyzon (Ehrenberg).
Proalinopsis caudatus (Collins).
Drilophaga judayi new species.
Taphrocampa annulosa Gosse.
Taphrocampa selenura Gosse.
Lindia Candida new species.
Lindia fulva new species.
Lindia pallida new species.
Lindia truncata (Jennings).
Tetrasiphon hydrocora Ehrenberg.
liarring and Myers — Tine Rotifers of Wisconsin.
555
Birgea enantia new genus, new species.
Eosphora anthadis new species.
Eosphora ehrenbergi Weber.
Eosphora melandocus (Gosse).
Enteroplea lacustris Ehrenberg.
Eothinia elongata (Ehrenberg) new genus for Eosphoi'a elongata Ehren¬
berg.
Sphyrias lo fuana (Rousselet).
Cephalodella catellina (Muller) = Diglena catellina (Muller).
Cephdlodella forficula (Ehrenberg) = Furcularia forficula Ehrenberg.
Diaschiza auriculata (Muller) — Diaschiza lacinulata (Muller).
Diaschiza derby i Dixon-Nuttall and (Freeman.
Diaschiza eva (Gosse).
Diaschiza exigua (Gosse).
Diaschiza forficata (Ehrenberg) = Diaschiza caeca (Gossee).
Diaschiza gibba (Ehrenberg).
Diaschiza globata Gosse.
Diaschiza gracilis (Ehrenberg).
Diaschiza hoodii Gosse.
Diaschiza megalocephala (Glasscott).
Diaschiza paeta Gosse.
Diaschiza parasitica (Jennings) = Pleurotrocha parasitica Jennings.
Diaschiza tenuior Gosse.
Diaschiza tenuiseta (Burn).
Dory stoma caudata (Bilfinger) new genus for Proales caudata Bilfinger.
Rousseletia corniculata Harring.
Tylotrocha monopus (Jennings) new genus for Notommata monopus Jen¬
nings.
Monommata orbis (Muller )=Furcularia longiseta (Muller).
Dicranophorus auritus (Ehrenberg) — Eosphora aurita (Ehrenberg).
Dicranophorus caudatus ( Ehrenberg )— Diglena biraphis (Gosse).
Dicranophorus corystis new name — Arthroglena rostrata von Hofsten,
not Dicranophorus rostratus (Dixon-iNuttall and Freeman.
Dicranophorus forcipatus (Muller) = Diglena forcipata (Muller).
Dicranophorus liitkeni (iBergendal) = Arthroglena liitkeni Bergendal.
Dicranophorus uncinatus (Milne) == Diglena uncinata Milne.
Encentrum aper Harring.
Encentrum circinator (Gosse) =Diglena circinator Gosse.
Encentrum felis (Muller) = Proales felis (Muller).
Encentrum ricciae Harring.
Erignatha clastopis (Gosse) new genus for Diglena clast opis Gosse.
Epiphanes clavulata (Ehrenberg) = Notops clavulatus (Ehrenberg).
Cyrtonia tuba (Ehrenberg) .
Microcodon clavus Ehrenberg.
Mikrocodides chlaena (Gosse).
Mikrocodides doliaris (Rousselet).
Mikrocodides robustus (Glasscott).
556 Wisconsin Academy of Sciences , Arts, and Letters.
Brachionus angularis Gosse.
Brachionus angularis caudatus Barrois and Daday.
Brachionus calyciflorus Pallas — Brachionus pala Ehrenberg.
Brachionus capsuliflorus Pa\\a,s=Brachionus bakeri Muller.
Brachionus havanaensis Rousselet.
Brachionus mollis Hempel.
Brachionus patulus Muller = Brachionus militaris Ehrenberg.
Brachionus poly acanthus Ehrenberg.
Brachionus serious Rousselet.
Platyias quadricornis (Ehrenberg) — Noteus quadricornis Ehrenberg.
Keratella cochlearis (Gosse) = Anuraea cochlearis Gosse
Keratella paludosa (Lucks) = Anuraea paludosa Lucks.
Keratella quadrata (Muller) = Anuraea aculeata Ehrenberg
Anuraeopsis fissa (Gosse) — Anuraea hypelasma Gosse.
Notholca bostoniensis Rousselet.
Notholca longispina (Kellicott).
Notholca striata (Muller).
Mytilina macrocera (Jennings) = Salpina macrocera Jennings.
Mytilina mucronata (Muller) = Salpina mucronata (Muller).
Mytilina trigona (Gosse) = Diplax trigona Gosse.
Euchlanis deflexa Gosse.
Euchlanis dilatata Ehrenberg.
Euchlanis lyra Hudson.
Euchlanis propatula (Gosse).
Euchlanis pyriformis Gosse.
Euchlanis triquetra Ehrenberg.
Lecane aeganea Harring.
Lecane arcula Harring.
Lecane brachydactyla (Stenroos) = Cathy pna brachydactyla Stenroos.
Lecane Clara (Bryce) == Distyla Clara Bryce.
Lecane compta Harring.
Lecane crepida Harring.
Lecane flexilis (Gosse) = Distyla flexilis Gosse.
Lecane hornemanni (Ehrenberg) = Distyla hornemanni (Ehrenberg).
Lecane inermis (Bryce) = Distyla inermis Bryce.
Lecane leontina (Turner) == Cathy pna leontina Turner.
Lecane ligona (Dunlop) = Cathy pna ligona Dunlop.
Lecane ludwigii (Eckstein) = Distyla ludwigii Eckstein.
Lecane luna (Muller) = Cathypna lima (Muller).
Lecane mira (Murray) = Cathypna uiira Murray.
Lecane nana (Murray) = Cathypna nana Murray.
Lecane ohioensis (Herrick) — Distyla ohioensis Herrick.
Lecane papuana (Murray) = Cathypna papuana Murray.
Lecane ploenensis (Voigt) = Cathypna ploenensis Voigt.
Lecane pusilla Harring.
Lecane stokesii (Pell) — Distyla stokesii (Pell).
Lecane tenuiseta Harring.
Lecane ungulata (Gosse) — Cathypna. ungulata Gosse.
Harring and Myers — The Rotifers of Wisconsin .
557
Monostyla acus Harring.
Monostyla bulla Gosse.
Monostyla closterocerca Schmarda.
Monostyla cornuta (Miiller). ..
Monostyla crenata Harring.
Monostyla furcata Murray.
Monostyla hamata Stokes.
Monostyla lunaris Ehrenberg.
Monostyla monostyla (Daday) = Diarthra monostyla Daday.
Monostyla pygmaea Daday — Monostyla turbo Murray.
Lepadella acuminata (Ehrenberg) = Metopidia acuminata Ehrenberg.
Lepadella amphitropis Harring.
Lepadella apsida Harring.
Lepadella benjamini Harring.
Lepadella cristata (Rousselet) .
Lepadella cyrtopus Harring.
Lepadella ehrenbergii (Perty).
Lepadella ovalis (Muller) = Metopidia solidus Gosse.
Lepadella patella (Miiller) — Metopidia lepadella Ehrenberg.
Lepadella quinquecostata (Lucks).
Lepadella rhomboides (Gosse).
Lepadella rhomboidula (Bryce).
Lepadella triptera Ehrenberg.
Colurella bicuspidata (Ehrenberg).
Colurella defiexa (Ehrenberg).
Colurella obtusa (Gosse).
Colurella sulcata (Stenroos).
Colurella tesselata (Glasscott).
Squatinella bifurca i (Hudson) = Stephanops bifurcus Hudson.
Squatinella leydigii (Zacharias) — Stephanops leydigii Zacharias.
Squatinella longispinata (Tatem) = Stephanops longispinatus Tatem.
Squatinella mutica (Ehrenberg) — Stephanops muticus Ehrenberg
Squatinella stylata (Milne) = Stephanops stylatus Milne.
Squatinella tenella (Bryce) = Stephanops tenellus Bryce.
Squatinella tridentata (Fresenius) = Stephanops intermedins Burn.
Trichotria intermedia (Bergendal) = Dinocharis intermedia Bergendal.
Trichotria pocillum (Muller) = Dinocharis pocillum (Muller).
Trichotria similis (Stenroos )= Dinocharis similis Stenroos.
Trichotria tetractis (Ehrenberg) — Dinocharis tetractis Ehrenberg.
Wolga spinifera (Western) = Distyla spinifera Western.
Lophocharis oxysternon (Gosse) = Metopidia Oxysternon Gosse.
Lophocharis salpina (Ehrenberg) = Metopidia salpina Ehrenberg.
Macrochaetus collinsii (Gosse) — Dinocharis collinsii Gosse.
Macrochaetus subquadratus Perty =Polychae tus subquadratus (Perty).
Scaridium eudactylotum Gosse.
Scaridium longicaudum (Muller).
Trichocerca bicristata (Gosse) = Rattulus bicristatus (Gosse).
Trichocerca bicuspes (Pell) — Rattulus bicuspes (Pell)
558 Wisconsin Academy of Sciences , Arts , and Letters.
Trichocerca cristata Harring = Rattulus carinatus (Ehrenberg).
Trichocerca cylindrica (Imhof ) = Rattulus cylindricus (Imhof).
Trichocerca elongata (Gosse) = Rattulus elongatus (Gosse).
Trichocerca flava (Voronkov) = Rattulus flavus Voronkov.
Trichocerca iernis (Gosse) — Rattulus gracilis (Tessin).
Trichocerca lata (Jennings) — Rattulus latus (Jennings)
Trichocerca macera (Gosse) = Rattulus macerus (Gosse).
Trichocerca mucosa (Stokes) — Rattulus mucosus (Stokes).
Trichocera multicrinis (Kellicott) = Rattulus multicrinus (Kellicott).
Trichocerca pusilla (Jennings) = Rattulus pusillus Jennings.
Trichocerca rattus (Muller) — Rattulus rattus (Muller).
Trichocerca scipio (Gosse) — Rattulus scipio (Gosse).
Diurella brachyura (Gosse).
Diurella collaris (Rousselet).
Diurella insignis Herrick.
Diurella dixon-nuttalli Jennings.
Diurella porcellus (Gosse).
Diurella stylata Eyferth.
Diurella sulcata (Jennings).
Diurella tenuior (Gosse).
Diurella tigris (Muller).
Diurella weberi Jennings.
Elosa worrallii Lord.
Chromogaster ovalis (Bergendal) = Anapus ovalis Bergendal.
Oastropus minor (Rousselet).
Gastropus orbicularis (Kellicott).
Gastropus stylifer Imhof.
Ascomorpha ecaudis Perty.
Ascomorpha saltans Bartsch.
Ascomorpha volvocicola (Plate) —Hertwigia volvocicola Plate.
Synchaeta longipes Gosse.
Synchaeta oblonga Ehrenberg.
Synchaeta pCctinata Ehrenberg.
Synchaeta stylata Wierzejski.
Synchaeta tremula (Muller).
Polyarthra trigla Ehrenber g — Polyarthra platyptera Ehrenberg.
Polyarthra euryptera Wierzejski
Ploesoma lenticulare Herrick.
Ploesoma truncatum (Levander).
Filinia longiseta (Ehrenberg) == Triarthra longiseta Ehrenberg.
Filinia passa (Muller) = Triarthra mystacina Ehrenberg.
Pedalia mira (Hudson) — Pedalion mirum Hudson.
Asplanchna brightwellii Gosse.
Asplanchna herrickii de Guerne.
Asplanchna priodonta Gosse.
Asplanchnopus hyalinus Harring.
Asplanchnopus multiceps (Schrank).
Harringia rousseleti de Beauchamp.
Harring and Myers — The Rotifers of Wisconsin.
559
Testudinella incisa (Ternetz) = Pterodina incisa Ternetz.
Testudinella parva (Ternetz) — Pterodina parva Ternetz.
Testudinella patina (Hermann) = Pterodina patina (Hermann).
Pompholyx sulcata Hudson.
Trochosphaera solstitialis Thorpe.
Order FLOSCULARIACEA
Floscularia conifera (Hudson) = Melicerta conifera Hudson.
Floscularia janus (Hudson) = Melicerta janus Hudson.
Floscularia ringens (Linne) — Melicerta ringens (Linne).
Limnias ceratophylli Schrank.
Limnias melicerta Weisse = Limnias annulatus Bailey.
Liinnias myriophylli (Tatem).
Ptygura brachiata (Hudson) = Oecistes brachiatus Hudson.
Ptygura crystallina (Ehrenberg) = Oecistes crystallinus Ehrenberg.
Ptygura longipes (Wills) = Oecistes umbella Hudson.
Ptygura mucicola (Kellicott) = Oecistes mucicola Kellicott.
Ptygura pilula (Cubitt) = Oecistes pilula (Cubitt).
Ptygura velata (Gosse) = Oecistes velatus Gosse.
Pseudoecistes rotifer Stenroos.
Sinantherina socialis (Linne) = Megalotrocha alboflavicans Ehrenberg.
Sinantherina semibullata (Thorpe) = Megalotrocha semibullata Thorpe.
Lacinularia flosculosa (Muller) =Lacinularia socialis Ehrenberg.
Octotrocha speciosa Thorpe.
Conochilus hippocrepis (Schrank) = Conochilus volvox Ehrenberg.
Conochilus unicornis Rousselet.
Conochiloides dossuarius Hudson.
Conochiloides natans (Seligo.)
Order COI LOT HKCA CE A
Collotheca algicola (Hudson) = Floscularia algicola Hudson.
Collotheca ambigua (Hudson) = Floscularia ambigua Hudson.
Collotheca campanulata (Dobie) = Floscularia campanulata Dobie.
Collotheca cornuta (Dobie) = Floscularia campanulata Dobie.
Collotheca coronetta (Cubitt) = Floscularia coronetta Cubitt.
Collotheca edentata (Collins) = Floscularia edentata Collins.
Collotheca mutabilis (Hudson) = Floscularia mutabilis Hudson.
Collotheca ornata (Ehrenberg) = Floscularia ornata Ehrenberg.
Collotheca pelagica (Rousselet) = Floscularia pelagica Rousselet.
Collotheca trilobata (Collins) = Floscularia trilobata Collins.
Stephanocerus fimbriatus ( Goldfuss )— Stephanoceros eichhornii Ehren¬
berg.
Cupelopagis vorax (Leidy) = Apsilus vorax (Leidy).
Acyclus inquietus Leidy.
560 Wisconsin Academy of Sciences, Arts, and Letters .
Order BDELLOIDEA
Adineta gracilis Janson.
Adineta vaga Davis.
Philodina citrina Ehrenberg.
Philodina megalotrocha Ehrenberg.
Philodina plena (Bryce).
Philodina roseola Ehrenberg.
Philodina vorax (Janson).
Rotaria citrina (Ehrenberg) = Rotifer citrinus Ehrenberg.
Rotaria macroceros (Gosse) == Rotifer macroceros Gosse.
Rotaria macrura (Ehrenberg) = Rotifer macrurus Ehrenberg.
Rotaria neptunia (Ehrenberg) =Actinurus neptunius Ehrenberg.
Rotaria rotatoria (Pallas) = Rotifer vulgaris Schrank.
Rotaria sordida (Western) =Callidina sordida Western.
Rotaria tardigrada (Ehrenberg) = Rotifer tardus Ehrenberg.
Machrotrachela habita (Bryce) = Callidina habita Bryce.
Macrotrachela multispinosa Thompson.
Macrotrachela nana (Bryce) = Callidina nana Bryce.
Macrotrachela plicata (Bryce) — Callidina plicata Bryce.
Macrotrachela quadricornifera Milne.
Macrotrachela zickendrahti (Richters) = Callidina zickendrahti Richters.
Dissotrocha aculeata (Ehrenberg).
Dissotrocha macrostyla (Ehrenberg).
Mniobia russeola (Zelinka).
Habrotrocha augusticollis (Murray).
Harbrotrocha constricta (Dujardin).
Habrotrocha lata (Bryce).
Habrotrocha munda Bryce.
Scepanotrocha rubra Bryce.
A Review of the Central Group of Notommatids
One of the most difficult tasks connected with this survey proved
to be the specific determination of the Notommatids, and as a first
step towards a revision we undertook a re-examination of the
central group of genera, as far as material was obtainable. Our
original intention was to include all the Notommatids and to pub¬
lish the results in this paper. We were aware that this was quite
a task, and when the war interrupted scientific work and inter¬
course it became necessary to restrict it to what has usually
been considered the central group. Even so we have been unable
to include the Proales-Pleurotrocha assemblage, not having access
to essential material, and only a few species have been described
without any attempt at determining their systematic position.
Harring and Myers — The Rotifers of Wisconsin. 561
What was supposed to be a fairly homogeneous group turns out
to be quite otherwise upon closer examination, as is frequently
the case. There remains consequently but a somewhat haphazard
aggregation of Notommatids. But even so, we feel that it will
still be useful to publish the descriptions and figures here
presented, as the existing information on this group is so scanty
and largely antiquated. It is our hope to take up the remainder
of the group in a subsequent paper, which will also deal more
fully, with the taxonomy. We have refrained from any but the
most obviously necessary generic rearrangements; the greater the
number of species studied, the more impressive becomes the fact
that the species of such a very old and, in its larger features at
least, relatively homogeneous group, have in their development
followed a great many and diverse crossways and byways, and the
unraveling of these is a very difficult task, which certainly ought
not to be attempted without the fullest possible knowledge of all
the available species.
Special efforts have been made to give characteristic figures of
the trophi, as it developed that no two species are identical in this
respect, and since determination of species is possible from the
trophi alone, which is very fortunate in view of the difficulties
of preservation. A word of caution concerning the use of the
figures is necessary; the sclerified parts of the Notommatid mastax
are generally extremely complicated, and an exhaustive study,
such as that made by Zelinka of Synchaeta atlantica, would have
consumed more time than we could command. We have there¬
fore figured as faithfully as possible the salient features. Many
minor details in the internal ribbing and bracing provided in the
rami to resist or to transmit the stresses of the muscles have thus
been omitted, but we hope that enough has been given to serve
the purpose of identification, which has been our principal aim,
and that it may also be of assistance in disentangling the inter¬
relations of genera.
36— S. A. L.
562 Wisconsin Academy of Sciences , Arts, and Letters.
Notommata Ehrenberg
notom mata copeus Ehrenberg
Plate XLI, figures 1-4
Notommata copeus Ehrenberg, Abh. Akad. Wiss. Berlin (for 1833)
1834: 213; Infusionsth. (1838), p. 434, pi. 51, fig. 1. — de Beauchamp,
Arch. Zool. Exp. IV, 10 (1909): 97, 195, 334, figs. XIX, LVI, PI. 8,
figs. 99-105.— Harking, U. S. Natl. Mus. Bull. 81 (1913): 78.
Notommata centrura Ehrenberg, Abh. Akad. Wiss. Berlin (for 1833)
1834: 211, 333, PI. 9, fig. 1; Infusionsth. (1838), p. 435, PI. 51, fig. 2.
— Eeydig, Zeitschr. Wiss. Zool. 6 (1854): 33, PI. 3, figs. 21-25, 27. —
Weber and Montet, Cat. Invert. Suisse, fasc. 11 (1918): 107.
Copeus notommata Ehrenberg, Infusionsth. (1838), p. 441.
Copeus ehrenJ)ergii Gosse, Hudson and Gosse, Rotifera (1886) 2: 28;
ibid., Suppl. (1889), p. 19, PI. 32, fig. 17. — Calman, Ann. Scott. Nat.
Hist. 1892: 242, PI. 8, fig. 2. — Skorikov, Trav. Soc. Nat. Kharkow
30 (1896): 288. — Rousselet, Jour. Royal Micr. Soc. 1906: 403.
Copeus labiatus Gosse, Hudson and Gosse, Rotifera (1886) 2: 26, PI. 16,
fig. 1. — Weber, Rev. Suisse Zool. 52 (1897): 92, PI. 4, fig. 6; ibid.
54 (1898): 453, PI. 18, figs. 8-10. — Stenroos, Acta Soc. Fauna et
Flora Fennica 171 1898) : 127, PI. 3, fig. 10. — Wesenberg-Lund,
Vid. Medd. Nat. For. Kjobenhavn 1899, PI. 2, fig. 31. — Montet,
Rev. Suisse Zool. 23 (1915): 321.
Copeus americanus Peel, Microscope 10 (1890) : 144, fig. 1.
Copeus copeus Collin, Deutsch-Ost-Afrika 415 (1897): 5, fig. 3. — Jen¬
nings, Amer. Nat. 35 (1901): 741, PI. 4, fig. 69.
Copeus centrurus Daday, Zoologica 44 (1905): 95; ibid. 59 (1910): 68.
— von Hofstein, Arkiv. Zool., Stockholm 61 (1909): 14. — Voigt,
Siisswasserfauna Deutschlands, pt. 14 (T912): 94, fig. 169.
The body is elongate, nearly parallel-sided and truncate poster¬
iorly; its greatest width is a little less than one third of the total
length. The integument is leathery and the outline quite constant ;
it is a fairly transparent species.
The head segment is short, broad, and truncate anteriorly;
its width is one half the greatest width of the body. The neck
segment is fairly long and slightly enlarged posteriorly. The
anterior transverse folds are well marked. The abdomen increases
gradually in width for about two thirds of its length; from this
point it diminishes rapidly and becomes truncate and rounded
posteriorly. The tail is a long, round, conical projection, ab¬
ruptly reduced at mid-length and ending in a blunt point. The
foot has two fairly long and broad joints; at the base of the
toes the terminal joint has a small, knob-like projection with a
COCKfcTNE B0STOTST
Harring and Myers — The Rotifers of Wisconsin.
563
tuft of minute setae. The toes are moderately long, about one
sixteenth of the total length, slender, conical, and slightly de-
curved.
The dorsal antenna is spindle-shaped and very long ; it is
sharply constricted at the base and has a tuft of setae at the
tip. The lateral antennae are short tubules with a few extremely
long setae.
The corona has a median, trough-like depression throughout its
length; the post-oral portion projects from the body as a long,
conical, bluntly pointed chin. The auricles are tubular and ex¬
tremely long; only the ends are ciliate. The width across the
fully extended auricles is more than twice the normal width of
the head segment.
The mastax is of the normal virgate type, with strongly asym¬
metric trophi, much more robust on the left side than on the
right. The fulcrum is very long and stout, tapering from the
base to the posterior end, which is slightly incurved. The posterior
half of the fulcrum is Y-shaped in cross-section, as two thin lamel¬
lae, gradually increasing in width, project diagonally outwards
and provide increased surface for the muscles of the piston. The
rami are irregularly triangular in ventral view and are strongly
curved longitudinally, so that their posterior edge is nearly at a
right angle to the fulcrum. Near the base of the rami there is a
rather prominent, triangular median projection; immediately be¬
hind this the right ramus has a broad, lamellar tooth projecting
diagonally towards the left; the margin is coarsely denticulate.
The left ramus is deeply excavate opposite this tooth. The alulae
are very large and well developed. The left uncus has a very
large, ventral tooth, clubbed at the tip, and followed by three
much smaller, slightly divergent teeth; at the dorsal margin of
the lamella uniting all the teeth there is a rudimentary fifth
tooth. The right uncus has four teeth, unequally developed, but
much less so than the mating teeth on the opposite side. The
manubria are very long and broad, the posterior ends curving
inwards and towards the dorsal side. Immediately below the
posterior margin of each ramus a curved rod is imbedded in the
walls of the mastax ; the ventral end rests on the inner surface of
the ramus, and the dorsal ends meet below the tips of the rami.
The oesophagus is very long and slender. The gastric glands,
stomach-intestine, and ovary are normal. The bladder is formed
564 Wisconsin Academy of Sciences, Arts, and Letters.
by an expansion of the cloaca. The foot glands are very long,
slender, and slightly club-shaped.
The retrocerebral sac is extremely long, clear, and vacuolate;
the subcerebral glands are less than half the length and usually
include aggregations of baeteroids, rendering them partly opaque.
The very large eye-spot is at the posterior end of the small, sac¬
cate ganglion.
Total length 750-1000/*,; toes 50-60/*,; trophi 90-100/*.
Notommata copeus is common in weedy ponds everywhere. In
certain places it is found inhabiting a jelly-case, secreted by the
integument; occasionally a blue-green alga is attached to the in¬
tegument and grows through the jelly-case. This species is usually
regarded as being strictly a plant-feeder; this may be a matter
of necessity rather than choice. Some time ago some material
was forwarded by Dr. Whitney from Lincoln, Nebraska, consist¬
ing mainly of colonies of the free-swimming Lacinularia ismai-
loviensis (Poggenpol), but containing also some individuals of
Notommata copeus. These were feeding vigorously on detached
individuals from the Lacinularia colonies, to which they held so
tenaciously that the captor could be transferred to a slide without
letting go its hold on the captive. It was then discovered that
the unci are, contrary to the general impression, very mobile and
capable of being opened out sufficiently to make them parallel
with each other, and also that they are very effective in capturing
and tearing the prey.
The specific name copeus has been used in preference to cen¬
trum, partly because it appeared to receive general acceptance
after Gosse’s names labiatus and ehrenbergii had been discarded,
and also because it was originally attached to the more correct
description. Ehrenberg’s citation in Infusionsthierchen under N.
centrura of Abh. Akad. Wiss. 1832 ; 438 is incorrect ; both centrum
and copeus were published simultaneously and for the first time
in the Abhandlungen for 1833 (published in 1834). Centrum
has page priority, but copeus has a better description; as is well
known, Ehrenberg described the animal with retracted auricles
as N. centrura and with extended auricles as N . copeus. The
recommendations of the International Code of Zoological Nomen¬
clature is that, “all other things being equal, page precedence
should obtain”, but “show preference to the best described, best
TRANS. W8S. ACAD., VOL. XX,
PLATE XL1 5 .
HARRiNG and MYERS.— ROT8FERS
COCKAYNE BOSTON
TRAMS W IS. ACAD., VOL. XX
PLATE XLI 1 !
HARRING and MYERS.— ROTIFERS
COCKAYNE BOSTON
Harring and Myers — The Rotifers of Wisconsin .
565
figured (etc.) species”. It seems therefore that both rules and
established usage are being obeyed in accepting the name Notom-
mata copeus.
NOTOM MATA PACHYURA (GrOSSe)
Plate XLII, figures 1, 2 ; Plate XLIII, figures 1-5
? Notommata ansata Ehrenberg, Abh. Akad. Wiss. Berlin (for 1831)
1832: 131; Infusionsth. (1838), p. 430, PI. 52, fig. 5.
Copeus pacTiyurus Gosse, Hudson and Gosse, Rotifera (1886) 2: 31,
PL 16, fig. 4. — Dixon-Nuttall, Jour. Quekett Micr. Club II, 5
(1894): 3i3i3, PL 15, figs. 1, 2. — Skorikov, Trav. Soc. Nat. Kharkow,
30 (1896): ,288.— Weber, Rev. Suisse Zool., 5 (1898): 458, PL 18,
figs. 11, 12. — Stenroos, Acta Soc. Fauna et Flora Fennica 171
(1898): 129. — Lie-Pettersen, Bergens Mus. Aarbog (for 1909)
101015: 42. — Voigt, Siisswasserfauna Deutschlands, pt. 14 (1912):
94, fig. 170. — Mola, Ann. Biol. Lac. 6 (1913): 242. — Montet, Rev.
Suisse Zool. 23 (1915): 321.
Copeus quinquelobatus Stokes, Jour. Royal Micr. Soc. 1896: 277, PL 6,
figs. 10, 11.
Copeus triangulatus Kirkman, Jour. Royal Micr. Soc. 1906: 264, PL 12,
figs. 1, 2.
Notommata brachiata Daday, Math. Term. Ert. 26 (1908) : 31, text fig.;
Zoologica 59 (1910): 67, fig. 2, PL 3, figs. 18, 19.
Notommata (Copeus) quinquelobatus de Beauchamp, Arch. Zool. Exp.
IV, 10 (1909): 86.
Notommata pachyura Harring, U. S. Natl. Mus. Bull. 81 (1913): 79;
Proc. U. S. Natl. Mus. 46 (1913): ,392. — Weber and Montet, Cat.
Invert. Suisse, fasc. 11 (1918): 109.
The body of this species is spindle-shaped, rather short, stout,
and gibbous posteriorly ; its greatest width is more than one third
of the total length. The integument is leathery and the outline
quite constant. It is a moderately transparent animal.
The head segment is very small; its width is very nearly equal
to the length, about one third of the greatest width of the body.
The neck is relatively long, one sixth of the total length, and in¬
creases rapidly in width towards the abdomen. The anterior
transverse folds are well marked. The abdomen is fusiform,
gibbous dorsally and rounded posteriorly. The tail is very large
and projects dorsally far above the first foot joint; it has only a
single lobe. The foot has two short, ^ery broad joints; at the
base of the toes the terminal joint has a small, dorsally projecting,
knob-like spur; apparently it does not have any setae. The toes
566 Wisconsin Academy of Sciences, Arts, and Letters.
are long and conical ; there is a slight constriction near the middle
and the proximal portion seems to be soft, while the distal por¬
tion has the appearance of a normal, slightly decurved toe. The
entire length is about one twelfth of the total length of the body.
The dorsal antenna is a small, setigerous papilla in the normal
position; the lateral antennae are small tubules with a tuft of
minute setae.
The corona extends down on the ventral side of the body about
one fourth of the total length ; the post-oral portion projects from
the surface of the body as a prominent chin. The auricles are
very large and stout ; when extended they nearly double the width
of the head. The ciliation is continuous with the corona.
The mastax is of the virgate type, with strongly asymmetric
trophi, much more robust on the left side than on the right. The
fulcrum is very long, stout, and slightly curved throughout its
length ; the anterior half is nearly parallel-sided, while the
posterior tapers gradually to the somewhat abruptly incurved
point. Beginning near mid-length there is on the ventral edge a
Y-shaped groove of gradually increasing width; its sides are
formed by two thin, corrugated lamellae, which provide increased
surface for the attachment of the muscles of the piston; the cross
section of the posterior portion of the fulcrum thus becomes
Y-shaped. The anterior end of the fulcrum has a blunt, angular
projection at the base of the rami; immediately behind this the
right ramus has a broad, lamellar tooth with coarsely denticulate
margin curving diagonally towards the left. Opposite this tooth
the left ramus is deeply excavate. The alulae are very large, but
also very unequally developed, the left being much the larger,
very broad and strongly curved, so that it continues backwards
beyond the base of the rami for a distance almost equal to one
third the length of the fulcrum. The left uncus has a huge,
blunt, club-shaped ventral tooth; closely appressed to this is a
somewhat shorter, very slender second tooth ; the third and fourth
teeth are rudimentary and do not project beyond the crenulate
margin of the basal plate uniting all the teeth of the uncus, which
converge somewhat towards the inner ends; the fifth tooth is on
the dorsal margin and projects beyond the inner edge as a spear-
shaped point; the crenulate margin extends slightly beyond it
dorsally. The ventral tooth of the right uncus is as broad as the
mating tooth, but somewhat blunter and only half the thickness;
it is followed by three straight, slender, slightly convergent teeth,
Harring and Myers — The Rotifers of Wisconsin.
567
all projecting beyond the crenulate margin, which is much nar¬
rower than on the opposite side. The mannbria are very long and
broad; a broad, thin lamella extends nearly to the posterior end
on the dorsal margin. Below the posterior edge of the rami
there is a curved rod imbedded in the walls of the mastax; the
ventral end rests on the inner surface of the ramus, and the
dorsal ends meet below the tips of the rami.
The oesophagus is long and slender. Gastric glands, stomach-
intestine, and ovary are normal. The bladder is formed by an
expansion of the cloaca. The foot glands are very long, slender,
and slightly club-shaped.
The retrocerebral sac is extremely long, pyriform, clear, and
vacuolate ; the subcerebral glands are small, barely one fourth the
length of the sac, and usually contain bacteroids, especially toward
the posterior end. The ganglion is small and saccate, with the
large eye-spot at its posterior end.
Total length 600-800/x; toes 75/x; trophi 75 jjl.
A variant of this species, described by Kirkman as C opens
triangulatus and later by Daday as Notommata brachiata, develops
three triangular humps, two lateral and one dorsal. As noted
by Harring, a complete series of intermediates may be found
between the humpless form and the fully humped form which is
illustrated on Plate 3, figures 1 and 2. There is little doubt
of their specific identity. The humpless form appears early in
the spring, while the intermediate and fully humped do not oc¬
cur until June at Washington. The humpless form continues
to occur throughout the summer, but in reduced numbers; the
humped variant is far more abundant and continues until late in
the fall. The humped form has not been reported from Europe;
we have not had an opportunity to study typical European ma¬
terial, and it is possible that the American and African forms
may be a distinct species.
Notommata pachyura is common in weedy ponds in the United
States, wherever collections have been made.
568
Wisconsin Academy of Sciences , Arts , and Letters.
notom mata collaris Ehrenberg
Plate XLIV, figures 1-5
Notommata collaris Ehrenberg, Abh. Akad. Wiss. Berlin (for 1831)
1832: 131, PL 4, fig. 11; ibid, (for 1833) 1834: 333, PL 9, fig. 2;
Infusionsth. (1838), p. 428, PL 52, fig. 1. — Leydig, Zeitschr. Wiss.
Zool. 6 (1854): 38. — ? Hudson and Gosse, Rotifera (1886): 2: 27,
PI. 17, fig. 6. — Wierzejski, Rozpr. Akad. Umiej., Wydz. Mat.-Przyr.
Krakow II, 6. (1893): 228. — Biufinger, Jahresh. Ver. Naturk. Wiirt-
temberg 50 (1894): 43. — Levander, Acta Soc. Fauna et Flora Fen-
nica, 123 (1895): 30. — de Beauchamp, Arch. Zool. Exp. IV, 10
(1909): 157. — Lie-Pettersen, Bergens Mus. Aarbog (for 1909)
1010ir>: 41. — Weber and Montet, Cat. Invert. Suisse, fasc. 11 (1918) :
108.
Copeus collaris Voigt, Siisswasserfauna Deutschlands, pt. 14 (1912) :
93, figs. 167, 168.
The body of this species is fusiform, short, stout, and gibbous
posteriorly; its greatest width is more than one third of the to¬
tal length. The integument is leathery and the outline fairly
constant. It is a moderately transparent animal.
The head segment is very small; its width is only one third
of the greatest width of the body, and the length is still less. The
neck is fairly long, about one seventh of the total length, and
considerably enlarged posteriorly to provide room for the very
large mastax. The anterior transverse folds are well marked.
The abdomen increases to twice its anterior width at a point
somewhat beyond mid-length; it is rounded posteriorly. The tail
is short and very broad; it projects dorsally considerably above
the foot and has only a single lobe. The foot has two very short,
broad joints; on its dorsal side the terminal joint has a small
knob-like papilla bearing a tuft of setae. The toes are very short,
straight, conical, slightly blunted at the tips and with an indis¬
tinct enlargement at the base ; their length is about one thirtieth
of the total length.
The dorsal and lateral antennae are small, setigerous conical
elevation in the norma] positions.
The corona extends down on the ventral side somewhat less
than one fourth of the length of the body; the post-oral portion
projects from the surface of the body as a fairly prominent chin.
The auricles are moderately long and stout; the ciliation is con¬
tinuous with the corona.
TRANS. WIS. ACAD,
VOL. XX
PLATE XL.IV
HARRING and MYERS.— ROTIFERS
COCKRYNE BO STOTT
Harring and Myers — The Rotifers of Wisconsin. 569
The mastax is virgate, with strongly asymmetric trophi, and,
in proportion to the size of the animal, larger than in any other
species belonging to this genus. The fulcrum is very long, stout,
and slightly tapering, the curvature gradually increasing towards
the posterior end. Beginning at a point near mid-length there
is on the ventral edge a Y-shaped groove of gradually increasing
width; its sides are formed by two thin, corrugated lamellae,
which provide increased surface for the attachment of the muscles
of the piston and give it a Y-shaped cross section. The anterior
end of the fulcrum has a blunt, angular projection at the base
of the rami ; immediately behind this the right ramus has a broad,
lamellar tooth with coarsely denticulate margin curving obliquely
towards the left. Opposite this tooth the left ramus is deeply ex¬
cavate. The alulae are very large and somewhat asymmetric;
their external angles are a considerable distance below the base of
the rami. The unci are roughly square, and each has three teeth,
united by a plate-like web; only the ventral tooth in each uncus
is developed. The inner margins of the unci are thickened and
marked with irregular, longitudinal striae. The left tooth is
extremely large, broad, blunt, and clubbed at the point; the right
tooth is a little smaller but of the same general form ; on the ven¬
tral edge there is near the point a spirally twisted projection,
corrugated like the inner margin of the unci, and opposing the
left tooth. The second and third teeth in each uncus are rudimen¬
tary and do not project beyond the inner margin. The manubria
are very long and broad; a wide, double-curved, thin lamella ex¬
tends nearly to the posterior end on the dorsal margin. Be¬
low the posterior edge of the rami there is a curved rod imbedded
in the walls of the mastax; the ventral end rests on the inner
surface of the ramus, and the dorsal ends meet below the tips
of the rami.
The oesophagus is very long and slender. Gastric glands and
stomach-intestine are normal. The ovary is elongate and broadly
ribbon-shaped. The bladder is formed by an expansion of the
cloaca. The foot glands are long, slender, and slightly club-
shaped.
The retrocerebral sac is very long, pyriform, clear, and vacuo¬
late. The subcerebral glands are small and somewhat variable in
size ; the dorsal and lateral views show the two extremes ; a few
bacteroids are scattered through the posterior portion of the
570 Wisconsin Academy of Sciences, Arts, and Letters.
glands. The ganglion is small and saccate, with the transversely
elongate eye-spot at the posterior end.
Total length 600-750/a; toes 25-32/a; trophi 100 y.
Notommata collaris does not appear to be common; we have
found it only among submerged mosses in Lake Kathan, about 5
miles from Eagle River, Yilas County, and in a small pond near
Minocqua, Oneida County, Wisconsin; at the time the last-named
collection was made, there was very little water in the pond, but
there were great masses of algae in conjugation.
This species is closely related to Notommata pachyura; it is
readily distinguishable by the much smaller tail and the very short
toes.
Notommata falcinella Harring and Myers, new species
Plate XLV, figures 8-12
The body is moderately slender and spindle-shaped, its greatest
width being somewhat less than one third of the total length. The
integument is moderately rigid and the outline remains fairly
constant. It is not very transparent.
The width of the head segment is nearly equal to its length,
approximately one half the greatest width of the body; the neck
segment is of the same length, but slightly wider, nearly two
thirds of the body width. The anterior transverse folds are well
marked. The abdomen increases very slightly in width to a point
about mid-length, and from there tapers gradually to the tail,
which is three-lobed, with a large median lobe, rounded poste¬
riorly, and two small lateral lobes. At the base of the indenta¬
tion separating the lobes there is a minute additional lobe. The
foot has two joints, the posterior considerably smaller than the
anterior; it terminates in a short, recurved spur, projecting over
the toes, which are rather slender, conical, and obtusely pointed;
their length is about one sixteenth of the total length.
The dorsal and lateral antennae are small setigerous papillae
in the normal positions.
The corona extends down on the ventral side about one third
the length of the body; the post-oral portion projects from the
body as a fairly prominent chin. The auricles are rather short
and quite stout, with an abundance of robust cilia, continuous
with the coronal eiliation.
TRANS. WIS. ACAD., VOL. XX. PLATE XLV.
HARRING and MYERS.— ROTIFERS
COCKAYHE BOSTON
Harring and Myers — The Rotifers of Wisconsin. 571
The mastax is of the normal virgate type, with decidedly asym¬
metric trophi, much more robust on the left side than on the
right. The fulcrum is long and stout, nearly parallel-sided for
two thirds of its length, and from there tapering to the slightly
incurved point, this terminal portion having a Y-shaped cross
section, giving additional surface for attachment of the muscles of
the piston. The rami are nearly semicircular in a ventral view,
and are bent nearly at a right angle at the extreme anterior point.
The right ramus has a moderately deep, ventral transverse groove,
in front of which there is a broad, lamellar tooth, projecting
diagonally towards the left; the margin of this tooth is coarsely
denticulate. The left ramus is excavate opposite this tooth ;
posteriorly the alula is continued as a stout, incurved spur. The
left uncus has one very large, club-shaped ventral tooth; there
are three additional rudimentary teeth radiating fanwise from
the basal plate of the uncus. The right uncus has two well de¬
veloped teeth, the ventral one much larger than the second, but
itself only half the bulk of the left tooth ; these are followed by two
small, pointed teeth and a blunt, blade-like dorsal tooth, the basal
plate uniting all nearly to the tips. The manubria are very
robust; the median section is very broad and nearly parallel¬
sided, slightly incurved posteriorly and expanded into a roughly
triangular plate at the anterior end. The oesophagus is long and
slender. The gastric glands, stomach-intestine, and ovary are
normal ; the bladder is formed by an expansion cf the cloaca, as
in Notommata pseudo cerberus and in a few other species. Two
long, slightly club-shaped foot glands are present.
The retrocerebral organ consists of an extremely long, clear,
vacuolate sac, one third the length of the body, and two very
short subcerebral glands, which usually include bacteroids, but
not in great numbers. The eye-spot is at the posterior end of
the small, saccate ganglion.
Total length 500-550/a; toes 30-33/a; trophi 75/a.
Notommata falcinella was collected some years ago in great
numbers among submerged sphagnum at the head of Furnace
Creek, near Baltimore, Maryland, in company with Notommata
saccigera. It has never been found since, although the same spot
has been searched every year at the same season.
This species is closely related to the other large species of the
genus, but is readily distinguishable by the small spur at the base
of the toes.
572 Wisconsin Academy of Sciences , Arts , and Letters.
NOTOM MATA CERBERUS (GoSSe)
Plate XLVI, figures 5-8
Covens cerberus Gosse, Hudson and Gosse, Rotifera (1886) 2: 34, PL
16, fig. ,3. — ? Skorikov, Trav. Soc. Nat. Kharkow 30 (1896): 289. —
L.ib-’Pettersen, Tromso Mus. Aarsh. 33 [1911 (for 1910)]: 57.
Notom mat a cerberus be Beauchamp, Zool. Anz. 33 (1908): 401, figs.
1-3. — Voigt, ;Siisswasserfauna Deutschlands, pt. 14 (1912): 97,
fig. 174. — Weber and Montet, Cat. Invert. Suisse, fasc. 11 (1918):
lOS.
The body is fairly slender, its greatest width being from one
fourth to one fifth of the total length. The integument is rather
flexible, but the outline is quite constant.
The head and neck are rather narrow and slender; there is a
slight transverse fold between the head and the neck, and also one
between neck and abdomen. The abdomen is elongate and very
slightly tapering to a point about two thirds its length, from
where it decreases gradually in width to the base of the foot. The
tail is rounded and has a small but distinct median notch and a
shallow, rounded lateral sinus which separates the median lobe
from the rudimentary lateral lobes. The foot has two short joints,
the posterior one carrying a very small tuft of setae. The toes
are of moderate length, about one fifteenth of the total length;
they are slender, tapering, slightly compressed laterally, and end
in acute points ; the ventral edge is almost straight, and the dorsal
slightly decurved.
The corona extends down on the ventral side about one fourth
the length of the body ; the post-oral portion forming a small chin.
The auricles are short and stout, with robust tufts of cilia, not
continuous with the corona.
The dorsal and lateral antennae are small setigerous pits in the
normal positions. On the second foot joint there is, on the dorsal
side, immediately above the base of the toes, a small circular pit
with a few setae in its center.
The virgate mastax is of the normal, pumping type, and the
trophi are robust and somewhat asymmetric. The fulcrum is of
moderate length, straight, and tapering towards the posterior
end, which is slightly crutched and formed of two lamellae
joined in a Y-shaped section, to which the muscles of the piston
are attached. The right ramus has a fairly deep, ventral, trans¬
verse groove, in front of which there is a broad, subsquare
Harring and Myers — The Rotifers of Wisconsin.
573
lamellar tooth, projecting diagonally towards the left; the margin
of this tooth, as well as the inner edge of the dorsal portion of
both rami, is striated, but not denticulate. Opposite this tooth
the left ramus is excavate ventrally, and at its highest anterior
point there are six small, blunt teeth. The right uncus has a
strong, club-shaped ventral tooth, followed by a second, much
smaller tooth, and a third tooth which is again smaller than the
second ; following these there is a rudimentary, almost linear,
diagonal tooth, starting from the base of the third tooth and
crossing to the tip of a linear tooth, just inside the dorsal margin
of the plate uniting all the teeth. The left uncus has a strong
ventral tooth, similar to the opposing right tooth, and is followed
by a much smaller second tooth and a linear rudiment of a third
tooth ; from the base of this third tooth a diagonal, linear tooth
crosses the uncus to the tip of the linear tooth near the dorsal
margin of the uncus. To the ventral edge of the first, large tooth
there is loosely attached a very slender, linear tooth, which
separates readily from the uncus when the trophi are treated with
hypochlorite ; there is no counterpart on the right side. The
manubria are very robust, with a broad, central, nearly straight
stem and a lamellar, subsquare anterior portion, the width of
which is nearly one half the length of the entire manubrium. A
pair of curved rods, attached at their ventral ends to the inner
surface of the rami, pass under the manubria and terminate be¬
low the dorsal tips of the rami; they are imbedded in the walls
of the mastax and aid in their support during the pumping ac¬
tion. The oesphagus is long and slender. Stomach and intestine
are not distinctly separated. The gastric glands are of normal
size and form. The ovary is slightly elongate transversely to the
axis of the body and somewhat irregular in outline. A bladder
of normal size is present. There are four foot glands, one pair
fairly large and the other quite small.
The retrocerebral sac is pyriform and reaches nearly to the end
of the mastax ; the subcerebral glands are only half the length of
the sac. Bacteroids are very numerous in the glands, where they
are generally collected in irregular clumps; in the vacuolate sac
there are only a few granules scattered among its contents. The
eye-spot is a large lenticular body at the posterior end of the
ganglion.
Total length 450— 500/*,; toes 30-33y; trophi 55/x.
574 Wisconsin Academy of Sciences , Arts, and Letters.
This species, described by de Beauchamp from a pond in the
forest of Fontainebleau (Seine et Marne), and also found in the
swamps of the Dombes (Aisne), France, appears to be rare. We
have collected it only at Mamie Lake and Eagle River, Yilas
County, Three Lakes, Oneida County, and Mather, Juneau
County, Wisconsin.
Notommata cerberus is closely related to Notommata galena,
from which it differs in being much larger, and from all other
known Notommatids it differs in having a median notch in the
posterior margin of the tail.
Notommata Galena Harring and Myers, new species
Plate XLYI, figures 1-4
The body is slender, its greatest width being about one fourth
the length. The integument is fairly stiff and the general outline
relatively constant. The entire body has a light orange-brown
tint.
The rather small head and the neck are separated by a slight
fold, and a second, more strongly marked fold divides the neck
from the abdomen. The neck is slightly larger posteriorly; the
abdomen increases gradually to its greatest width at a point a
little beyond mid-length ; it is then rapidly reduced to the base of
the foot, the first joint of which is considerably wider than the
second and nearly twice as long. The tail has a large, rounded
central lobe and two minute lateral lobes. The toes are about one
fifteenth of the total length, their inner edges straight and outer
edges slightly curved; viewed laterally they are rather large at
the base, tapering gradually to the point, and very slightly de-
curved.
The dorsal and lateral antennae are small setigerous pits and
in the normal positions. On the second foot joint there is a
minute circular pit, which appears to have a few short setae in
its center.
The corona extends down on the ventral side about one fourth
the length of the body; the post-oral portion projects slightly
from the body, forming a “chin”. The auricles are of medium
size and provided with strong tufts of cilia, continuous with the
corona.
The mastax is of the normal, pumping type and has the usual
trilobate appearance when seen from the ventral side. The trophi
HARR1NG and MYERS.— ROTIFERS
COCKATNTE BOSTON
Harring and Myers — The Rotifers of Wisconsin. 575
are very stout and somewhat asymmetric. The fulcrum is of
moderate length, straight, and slightly tapering towards the tip,
which curves inward and is formed of two lamellae joined in a
Y-shaped section serving for the attachment of the muscles of the
piston. The right ramus has four transverse grooves directly be¬
low the teeth of the unci, all very shallow with the exception of
the first, ventral groove, which is quite deep and separates off a
broad, subsquare, lamellar tooth, curving to the left; the margin
of this tooth, as well as the entire inner edges of both rami, is
striated, but not denticulate. To make room for the broad,
curved tooth of the right ramus, the left ramus is excavate ven-
trally; it has three additional transverse grooves, in which the
teeth of the mallei rest; at its highest, anterior point there are
five or six small teeth, the remainder of the inner edge being
striated like that of the right ramus. The right uncus has a
strong, club-shaped ventral tooth, followed by two slender teeth
and a curved, diagonal rib crossing the uncus from the lower ven¬
tral edge to the tip of a fourth, dorsal tooth ; the left malleus has
one tooth less. At the point of the first tooth of the right uncus
there are four minute, slender supplementary teeth ; the left uncus
has four similar but much larger teeth. These supernumerary
teeth are not very firmly attached to the unci and are detached
quite readily when the tissues are dissolved in potassium hypo¬
chlorite; they are found in quite a number of species of the genus
and may possibly be considered a local specialization of the sub¬
uncus or a remnant of it, as the subuncus proper is not present
in the virgate mastax. The manubria are nearly straight, with
a broad, subsquare lamella at their upper ends. A pair of curved
rods, attached to the inner surfaces of the rami and terminating
below the tips, assist in supporting the walls of the mastax dur¬
ing the pumping movement. The oesphagus is long and slender.
Stomach and intestine are only indistinctly separated. The
gastric glands are small and rounded. The ovary is slightly
elongate transversely to the axis of the body and of somewhat ir¬
regular outline. A small bladder is present. The foot glands
are club-shaped and rather small.
The retrocerebral sac is pyriform and reaches beyond the
mastax; the subcerebral glands are only half the length of the
sac. The sac presents the usual vacuolate appearance; a few
opaque granules are scattered through its substance. The glands,
on the contrary, are crowded with bacteroids, especially at their
576 Wisconsin Academy of Sciences , Arts, and Letters.
posterior ends. The eye-spot is a relatively large lenticular body
at the posterior end of the ganglion.
Total length 250-300//,; toes 16-20 //,; trophi 40//,.
As far as our observations go, this species is more widely and
abundantly distributed in the United States than any other Notom-
matid.
The nearest known relative of this species is N. cerberus , from
which it differs in being considerably smaller, about two thirds the
size, and not quite so slender. The trophi are nearly identical
with the exception of the pre-uncial teeth, which N. cerberus does
not have. In N. galena the tail is rounded and entire posteriorly,
while in N. cerberus it has a small median notch, not found in any
other known Notommatid.
Notommata Peridia Harring and Myers, new species
Plate XLYIII, figures 1-5
The body is elongate, fusiform, and very slender; its greatest
width is one fifth of the total length. The integument is very
flexible, but the outline is fairly constant. It is a very trans¬
parent species.
The head segment is short and narrow, its width being about
one half the greatest width of the body. The neck segment is
very little wider, but nearly twice as long. The anterior trans¬
verse folds are well marked. The abdomen increases very slightly
in width for two thirds of its length and then tapers gradually to
a slight constriction at the base of the tail, which forms a blunt,
rounded projection; on the dorsal side there are two wart-like,
fairly prominent papillae. The foot has two small joints. The
toes are long, about one twelfth of the length of the body, taper¬
ing and very slender.
The dorsal antenna is a small setigerous papilla in the normal
position ; the lateral antennae are very long, tubular, slightly
swollen at the base and knobbed at the ends; they are provided
with a few very long setae.
The corona extends down on the ventral surface for about one
fourth of the total length. The post-oral portion projects as a
fairly prominent chin. No auricles are present ; at the anterior
lateral angles are two V-shaped, grooved, ciliated appendages,
which may be rudimentary auricles.
The mastax is virgate with strongly asymmetric trophi; the
TRANS. WIS. ACAD., VOL. XX.
PLATE XLVII.
HARRING and MYERS.— ROTIFERS
COCKA.TNE B03T0K
Harring and Myers — The Rotifers of Wisconsin.
577
two sides are nearly equally developed. The fulcrum is very
long, stout, and slightly tapering for two thirds of its length;
the posterior third is obliquely truncate and slightly incurved
at the tip. The oblique ventral edge has a Y-shaped groove, the
sides of which are formed by two thin, corrugated laminae, which
provide increased surface for the attachment of the muscles.
The anterior end of the fulcrum has a rounded projection at the
base of the rami; immediately behind, the right ramus has a very
broad, lamellar tooth curving diagonally towards the left; the
free edge has a finely striated border. The left ramus is deeply
excavate opposite this tooth and has at its extreme anterior point
two large, blunt teeth, interlocking with two similar teeth on the
right ramus; the decurved dorsal section of the rami beyond the
anterior teeth is not denticulate. The left alula is fairly large and
strongly curved ; the right is a small, conical knob. The left uncus
has a very large ventral tooth, clubbed at the point and with a
rudimentary pre-uncial tooth, lamellar at the tip, attached to its
ventral edge; the second tooth is very much smaller and shorter;
the third and fourth teeth are closely appressed and, like the
fifth tooth, linear. The basal plate unites all the teeth nearly
to their tips and projects considerably beyond the ventral tooth.
The right ramus has a linear pre-uncial tooth and the first and
second teeth of nearly equal size, but much smaller than the left
ventral tooth; the third and fourth teeth are but little more than
half as long. The basal plate projects very little beyond the pre¬
uncial tooth. The manubria are very broad and lamellar an¬
teriorly, terminating in a rather narrow, straight median branch.
Below the posterior edge of the rami there is a long, incurved rod
imbedded in the walls of the mastax ; the ventral end rests on the
inner surface of the ramus, and the dorsal ends meet some distance
below and behind the tips of the rami.
The oesophagus is long and slender. Gastric glands, stomach-
intestine, and ovary are normal. The bladder is very small. The
foot glands are rather small, very slender, and slightly club-
shaped.
The retrocerebral sac is extremely large, nearly circular, flat¬
tened and disk-shaped ; the duet is a very long, slender tube. The
subcerebral glands are large and pyriform, reaching to the level
of the eye-spot; they usually contain bacteroids. The ganglion
is elongate and quite slender.
Total length 300-375/*; toes 25-30/*; trophi 65/*.
37— S. A. L.
578 Wisconsin Academy of Sciences , Arts , and Letters.
N otommata peridia appears to be rare. It has been collected
only at Oceanville, near Atlantic City, New Jersey, and in Star¬
vation Lake, Loon Lake, and swampy ponds at Eagle River, Yilas
County, Wisconsin; Starvation Lake, about 5 miles southwest of
Eagle River, is the only station where it it has been found in
abundance. The animal is invariably found inhabiting an exceed¬
ingly tough jelly-case, even more resistant than the case of Te-
trasiphon hydrocora, and this affords much-needed protection
against enemies, as it appears to be unable to swim at all, on ac¬
count of the feeble development of the corona and the absence of
auricles.
This species has a superficial resemblance to Tetrasiphon hydro-
corn, but apart from its much smaller size is readily distinguish¬
able by the two small papillae on the tail, as well as by the retro-
cerebral organ and the trophi.
NOTOMMATA AURITA (Muller)
Plate XLVIII, figures 6-9
Vorticella aurita Muller, Anim. Inf us. (1786), p. 288, PI. 41, figs. 1-3.
FurciMaria aurita Lamarck, Hist. Nat. Anim. sans Vert. 2 (1816): 38.
Ft otommata aurita Ehrenberg, Abh. Akad. Wiss. Berlin 1830: 46; ibid.
(for 1831) 1832: 131, PI. 4, fig. 12; Infusionsth. (1838), p. 430, PI.
52, fig. 3. — Gosse, Trans. Micr. ,Soc. London 3 (1852): 93, PI. 12,
figs. A, B, PI. 15; Phil. Trans. Royal Soc. London 146 (1856): 430,
PI. 16, figs. 16-21. — Leydig, Zeitschr. Wiss. Zool. 6 (1854): 38, PI.
3, fig. 30. — Eckstein, Zeitschr. Wiss. Zool. 39 (1883): 360, PI. 25,
figs. 23-27. — Plate, Jenaische Zeitschr. Naturw. 19 (1886): 21. —
Hudson and Gosse, Rotifera (1886) 2: 21, Ft. 17, fig. 6. — Wierzej-
ski, Rozpr. Akad. Umiej., Wydz. Mat.-Przyr. Krakow II, 6 (1893):
|227.- — Skorikov, Trav. Soc. Nat. Kharkow 30 (1896): 285.- — Weber,
Rev. Suisse Zool. 5 (1898): 439, PI. 17, figs. 16-18. — Wesenberg-
Lund, Yid. Medd. Nat. Poren. Kj0benhavn 1899, PI. 1, fig. 14. — von
Hofsten, Arkiv Zool., Stockholm 61 (1909): 28, fig. 4. — de Beau¬
champ, Arch. Zool. Exp. IV, 10 (1909): 158, fig. XII C. — Lie-Pet-
tersen, Bergens Mus. Aarbog (for 1909) 191 015: 39. — Hirsch-
felder, Zeitschr. Wiss. Zool. 96 (1910): 326, PI. 13, figs. 41-46. —
Voigt, Siisswasserfauna Deutschlands, pt. 14 (1912): 98, fig. 177. —
Montet, Rev. Suisse Zool. 23 (1*915): 320. — Weber and Montet,
Cat. Invert. Suisse, fasc. 11 (1918) : 111, fig. 31.
Cycloglena lupus Ehrenberg, Abh. Akad. Wiss. Berlin 1830: 48; In¬
fusionsth. (1838), p. 454, PI. 56, fig. 10.
N otommata lupus Eyferth, Mikr. iSusswasserbew. (1877), p. 49.
TRANS. WIS. ACAD., VOL. XX.
PLATE XLVf i I
HARRING and MYERS— ROTIFERS
eOGKSTPTE BOSTON
Earring and Myers — The Rotifers of Wisconsin. 579
The body is comparatively short and thick-set, the greatest
width being about one third of the total length. The integument
is moderately flexible and the outline of the animal quite constant.
It is one of the most transparent species of the genus and very
favorable for the study of the internal organs.
The head and neck are short and broad, and there is a well
marked transverse fold between the head and neck, as well as one
between neck and abdomen. The abdomen increases very gradu¬
ally in width to a point about two thirds its length and is rounded
posteriorly. The tail is distinctly marked off from the body;
it has only a single, median lobe, rounded posteriorly. The foot
has two joints; the basal joint is broad and short, its length
barely exceeding that of the tail; the second foot joint is also
short, but only half the width of the first. The toes are short,
about one twentieth of the length, conical, slightly blunted at the
tips, and very slightly decurved. The dorsal and lateral antennae
are small setigerous pits and in the normal positions.
The corona extends down on the ventral side about one third
the length of the body ; the post-oral portion forms a fairly
prominent chin. The auricles are short and stout, with close-set
robust cilia continuous with the corona.
The mastax is virgate and appears trilobate in a ventral view
on account of the muscles attached to the posterior ends of the
fulcrum and manubria. The trophi are noticeably asymmetric,
the left side being more strongly developed than the right. The
fulcrum is long and quite stout, the posterior end being incurved
and of a Y-shaped section, providing a larger surface for the at¬
tachment of the muscles of the piston. The rami appear sub¬
square in a ventral view and are bent nearly at a right angle at
the extreme anterior point. The right ramus has a large basal,
median projection ending in a blunt tooth; behind this projection
it is slightly excavate and then inclines gradually towards the
teeth of the unci. This ventral portion is rather coarsely denticu¬
late; beyond the anterior angle the dorsal portions of both rami
are finely, but distinctly, denticulate. The left ramus has a basal
projection similar to that of the right, but smaller, and it is more
deeply excavate in front of the uncus; the ventral denticulation
is somewhat finer than that of the right ramus. The unci have
only a single well developed tooth; to this is attached a basal
plate with a rudimentary tooth close to the main tooth ; a diag¬
onal rib crosses the uncus from the posterior ventral angle
580 Wisconsin Academy of Sciences, Arts, and Letters.
to the median dorsal angle. To the point of the right uncial
tooth are attached two very small pre-uncial teeth, and to the
left three still smaller teeth; it is possible that these supplemen¬
tary teeth perform the functions normally belonging to the
atrophied dorsal teeth of the unci. The anterior half of the
manubria is a broad, angular plate, continued posteriorly as a
nearly straight rod, the tip of which curves outward and toward
the ventral side. A pair of curved rods, gradually widening
toward their dorsal ends, are attached to the inner ventral surfaces
of the rami and meet dorsally some distance below the tips of the
rami, thus supporting the external walls of the mast ax during
the pumping action. The oesophagus is fairly long; stomach and
intestine are not separated by any constriction. The gastric
glands are rather smaller than usual. The ovary is of the usual,
slightly irregular form. A small bladder is present. The foot
glands are of medium size and pyriform.
The retrocerebral sac is large and almost spherical; it opens
on the corona through a long, tubular duct and is so crowded
with baeteroids as to appear black by transmitted light. The sub¬
cerebral glands are rudimentary and fused with the ganglion.
The eye-spot is large, but is usually visible only from the side on
account of the opacity of the sac.
Total length 325-350 /x; toes 16— 20/x ; trophi 36/x.
Notomma aurita is widely distributed; it seems to be most
abundant in the spring, immediately after the ice leaves the ponds ;
later in the year it is not very common.
IMotommata Augusta Harring and Myers, new species
Plate LII, figures 9-12
The body is elongate, nearly parallel-sided, and quite slender,
its greatest width being only one fifth of the total length. The
integument is very flexible, but the outline is nevertheless fairly
constant. It is a moderately transparent species.
The width of the head and neck segments is slightly greater
than their length and very little less than the greatest width of
the body. The head is strongly convex or bluntly pointed an¬
teriorly, and on ithe ventral side the mouth region forms a de¬
cided elevation above the general surface of the corona. The
Harring and Myers— The Rotifers of Wisconsin. 581
anterior transverse folds are not very conspicuous. The abdomen
increases very slightly in width for about one half of its length
and then tapers very gradually to the tail, which has a single,
broad, rounded lobe. The two joints of the foot continue the
general outline of the body without any abrupt reduction. The
toes are rather short, about one twentieth of the total length,
slender, conical, and acutely pointed.
The dorsal and lateral antennae are minute setigerous papillae
in the normal positions.
The corona extends down on the ventral side nearly one third
of the length of the body ; the post-oral portion projects from the
body as a very small chin. The auricles are small, but strongly
ciliate and continuous with the corona.
The mastax is of the virgate type and slightly asymmetric. The
fulcrum is very long, slender, and slightly tapering; the posterior
end is expanded and incurved. The rami are approximately tri¬
angular in ventral view and bent at a nearly right angle at the
extreme anterior point ; near their base there is a blunt projection,
bifid at the tip and continuing the ventral line of the fulcrum.
Behind this projection the rami are slightly excavate, and the
inner margin of the right ramus is concave and minutely denti¬
cular, while the left has a convex margin with six or seven small
teeth, continued diagonally downwards on the convex lamella as
fine striae. The alulae are unusually large and project outwards
and downwards from the rami proper. The unci have an ap¬
proximately subsquare basal plate with a large ventral tooth and
a very slender second tooth immediately behind it; a diagonal
rib crosses the basal plate from the external end of the second
tooth to the anterior dorsal angle. The left uncus has three
small pre-uncial teeth at the tip of the principal tooth, and the
right has four similar, but slightly smaller and more pointed
teeth. A short and nearly straight lateral supporting rod is em¬
bedded in the walls of the mastax. The manubrium is broadly
expanded anteriorly as a subsquare plate, continuing as a long,
slender rod, slightly incurved at the tip. The oesophagus is very
long and slender, and the gastric glands are fairly large. The
stomach and intestine are separated by a well marked constriction.
The ovary is normal and the bladder large. The foot glands are
pyriform and rather small.
The retrocerebral organ is unusually well developed. The sac
is more than one third as long as the body; the posterior portion,
582 Wisconsin Academy of Sciences , Arts, and Letters.
or sac proper, is somewhat triangular in outline, and squarely
truncate behind; it then contracts abruptly to a fairly large
cylindrical section, which at the eye-spot again contracts to a
very slender tube, opening on the corona. The truncate posterior
end is faintly lobate and opaque, with bacteroids outlining the
lobes. The subcerebral glands are long and slender, reaching
some distance beyond the eye-spot, which is at the posterior end
of the slightly elongate, saccate ganglion.
Total length 300-325/*,; toes 15/x ; trophi 36/*.
Notommata august a was found rather sparingly in a polluted
brook at Brick Haven, south of the Potomac River, near Wash¬
ington, District of Columbia ; it has not been found elsewhere.
This species is readily recognized by the slender body, the un¬
usual outline of the head, and the structure of the retrocerebral
organ.
NOTOMMATA CYRTOPUS (Gosse)
Plate XLIX, figures 5-8
Notommata cyrtopus Gosse, Hudson and Gosse, Rotifera (1886), 2: 22,
PI. 17, fig. 7. — Wierzejski, Rozpr. Akad. Umiej. Wydz. Mat.-Przyr.
Krakow II, 6 (1893): 228. — Bilfinger, Jahresh. Ver. Naturk. Wiirt-
temberg 50 (1894): 44. — von Hofsten, Arkiv Zool., Stockholm 61
(1909): 30, fig. 5. — Lie-Pettersen, Bergens Mus. Aarbog (for 1909)
191015: 39. — Voigt, Susswasserfauna Deutschlands, pt. 14 (1912):
100, figs. 182, 183. — Mola, Ann. Biol. Lac. 6 (1913): 241. — Weber
and Montet, Cat. Invert. Suisse, fasc. 11 (1918): 113.
? RattUMs cimolius Gosse, Hudson and Gosse, Rotifera (1886), 2: 66,
PI. 20, fig. 14.
? Notommata distincta Bergendal, Acta Univ. Lundensis 28 (1892),
sect. 2, no. 4, p. 61, Pis. 3, 4, fig. 23L — Voigt, Forschungsber. Biol.
Stat. Plon 11 (1904): 39, PI. 3, fig. 18; Susswasserfauna Deutsch¬
lands, pt. 14 (1912): 100, figs. 184, 185.
The body is fairly stout and spindle-shaped, its greatest width
being a little less than one third of the total length. The integu¬
ment is soft and flexible, but the outline remains fairly constant.
The stomach is usually so crowded with food that it is difficult to
study the internal organs, even though the animal is really fairly
transparent.
There is no separation between the head and neck segments;
the wudth of the two fused segments is about three fourths of the
HARRING and MYERS. — ROTIFERS
TRANS. WIS. ACAD.. VOL XX.
COCKAYNE BOSTON
Barring and Myers — The Rotifers of Wisconsin. 583
greatest width of the body, and their length is but little more.
The single anterior transverse fold is well marked. The abdomen
increases gradually in width to a point about two thirds the
length and is rounded posteriorly. A tail is not present. The
foot has two joints, both short and fairly stout. The toes are
fairly long, about one seventh of the total length, slender, taper¬
ing, outcurved and decurved.
The dorsal and lateral antennae are small setigerous papillae
in the normal positions.
The corona extends down on the ventral side about one third
the length of the body; the post-oral portion forms a slight chin.
The auricles are short and stout, and the ciliation is continuous
with the corona.
The mastax is virgate, with slightly asymmetric trophi. The
fulcrum is long, moderately broad, slightly tapering, and fairly
stout; the posterior end is slightly expanded and curves inward.
The rami are approximately triangular in ventral view and are
bent nearly at a right angle at the extreme anterior point; on
their inner, ventral edges there is a blunt projection, continuing
the line of the fulcrum and bifid at the anterior tip; this cor¬
responds to the basal apophysis of the mallcate mastax. Behind
this projection the rami are slightly excavate, and the inner edge
of the right ramus is concave, while the left ramus is convex;
both are coarsely denticulate, and the left ramus is also striate.
The well developed alulae end in curved, horn-like prolongations;
their tips are only partly sclerified, so that the actual length is
somewhat uncertain. The unci have a single well developed ven¬
tral tooth and a rudimentary second tooth immediately behind
it ; the basal plate is subsquare and crossed by a diagonal rib from
the base of the ventral tooth to the anterior dorsal angle. The
right ramus has four short, curved pre-uncial teeth loosely at¬
tached to its tips, and the left has five similar, but more slender,
teeth. Some distance below the posterior edge of the dorsal sec¬
tion of the rami, a slender, nearly straight rod is imbedded in
the walls of the mastax as additional support during the pump¬
ing action. The anterior half of the manubrium is a rather nar¬
row plate and the posterior a slender rod, curving towards the
ventral side at the tip. The small, slender, bent rods in front of
the rami support the edges of the mouth. The oesaphagus is
short and rather slender. The gastric glands, stomach-intestine,
584 Wisconsin Academy of Sciences , Arts, and Letters.
ovary, and bladder are normal. The foot glands are large, pyri¬
form, and compressed laterally.
The retrocerebral sac is pyriform and fairly large; it opens
on the corona through a long, tubular duct and is filled with bac-
teroids, which render it opaque to transmitted light. The sub¬
cerebral glands are vacuolate and partly fused with the fairly
large, saccate ganglion, which has a small eye-spot at its posterior
end.
Total length 150-175/a; toes 22-26/a; trophi 32/a.
Notommata cyrtopus is common everywhere in weedy ponds.
It is readily recognized by the opaque retrocerebral sac and the
long, curved toes. Considerable variation is found in the stout¬
ness of the body ; the specimen figured may be taken as represent¬
ing the maximum, and it is usually much more slender.
Notommata Telmata Harring and Myers, new species
Plate XLIX, figures 1-4
The body is slender and spindle-shaped, its greatest width be¬
ing about one fourth of the total length. The integument is
moderately rigid and the outline remains fairly constant. It is
not very transparent.
The width of the head segment is slightly greater than its
length, or very nearly two thirds of the greatest width of the
body; the neck segment is of the same length, but slightly wider.
The anterior transverse folds are well marked. The abdomen
increases very slightly in width for about two thirds of its length
and is rounded posteriorly. The tail is inconspicuous and has a
single, median, rounded lobe. The foot is two-jointed; the ter¬
minal joint is considerably smaller than the first joint. The toes
are of unusual form ; the basal section is straight and extremely
short, and the posterior portion is long and slender, curved out¬
wards and downwards, with a distinct heel at its junction with
the basal section. Their length is about one twelfth of the total
length.
The dorsal and lateral antennae are small setigerous papillae
in the normal positions.
The corona extends down the ventral side about one fourth of
the length of the body; the post-oral portion projects from the
Harring and Myers — The Rotifers of Wisconsin. 585
body as a slight chin. The auricles are rather small, and the
ciliation is continuous with the corona.
The mast ax is of the virgate type, with slightly asymmetric
trophi. The rami are nearly triangular in ventral view and are
bent nearly at a right angle at the extreme anterior point. The
right ramus has on its median ventral edge a blunt, knob-like
projection, which is a continuation of the ventral edge of the ful¬
crum; behind this projection it is slightly excavate and then in¬
clines gradually toward the teeth of the unci. This section of
the ramus is finely denticulate. The left ramus has a ventral
projection similar to, but smaller than, the one on the right ramus;
it is also more deeply excavate, and the ventral, inclined surface
is coarsely denticulate and striate on its inner edge. Beyond the
anterior angle the dorsal portions of both rami are minutely den¬
ticulate on their opposing edges. The fulcrum is long, broad,
and stout; the posterior end is enlarged and curves inward. The
unci have a single, well developed ventral tooth and a rudimen¬
tary dorsal tooth at the edge of a broad basal plate ; from the root
of the ventral tooth to the tip of the torsal tooth there is an
inconspicuous diagonal rib. The left ramus has four very slender
pre-uncial teeth attached ventrally to the tip, while the right
ramus has three much larger teeth in this position. The anterior
half of the manubria is a very broad, thin lamella, and the poste¬
rior half a slender, curved rod. Besting with their anterior ends
on the inner surface of the rami and meeting dor sally at a point
somewhat below their tips, a pair of slender, slightly curved rods
pass diagonally under the anterior ends of the manubria ; they are
imbedded in the walls of the mastax and assist in its support dur¬
ing the pumping action.
The oesophagus is rather short and slender. The gastric glands,
stomach-intestine, ovary, and bladder are normal. The foot glands
are fairly long and pyriform.
The retrocerebral sac is pyriform and moderately large; it
opens on the corona through a long, tubular duct and is filled
with bacteroids, which make it appear black by transmitted light.
The subcerebral glands are long and pyriform, reaching down well
below the eye-spot, which is at the posterior end of the large,
saccate ganglion; the glands do not contain bacteroids.
Total length 250 /*; toes 20/* ; trophi 42/*.
586 Wisconsin Academy of Sciences, Arts , and Letters.
Notommata telmata was collected at Gravelly Run, near Atlan¬
tic City, New Jersey; it is not known from any other locality.
This species is closely related to Notommata cyrtopus , but is
easily distinguished by the peculiar toes, as well as by the more
slender body and the characteristics of the retrocerebral organ.
Notommata Lenis Harring and Myers, new species
Plate L, figures 9-13
The body is fairly slender and spindle-shaped; its greatest
width is about one fourth of the total length. The integument is
moderately rigid, and the outline remains fairly constant. It is
not very transparent.
The head segment is comparatively short and broad, its width
being about three fifths of the greatest width of the body. The
neck segment is very little wider, but nearly twice as long. The
anterior transverse folds are well marked. The abdomen in¬
creases very slightly in width to about mid-length, and from there
tapers gradually to the base of the foot, ending in a three-lobed
tail; the median lobe is unusually long and narrow, rounded
posteriorly and separated from the small lateral lobes by a slight
indentation. The foot has two joints, the anterior fairly long
and robust, the posterior divided by a median notch into two
papillae bearing the toes, which are short, conical, and abruptly
reduced to short, acute points; their length is only one twenty-
fifth of the total length. The anterior foot joint has a small seti-
gerous papilla on its posterior dorsal margin.
The dorsal antenna is remarkable for being double; the two
tufts of sensory setae are about 10y apart and united by a slender
muscle as in Asplanchna. The lateral antennae are in the normal
position.
The corona extends down on the ventral side of the body about
one fourth the length of the body; the post-oral portion projects
from the body as a slight chin. The auricles are small, but have
numerous long cilia, continuous with the corona.
The mastax is virgate with somewhat asymmetric trophi. The
fulcrum is very long and slender, broad at the base and tapering
gradually to the expanded posterior end. The rami are roughly
semicircular in ventral view, and are bent at a nearly right
angle at the extreme anterior point. The right ramus has a
Harring and Myers — The Rotifers of Wisconsin . 587
moderately deep, ventral, transverse groove, in front of which there
is a short, broad, lamellar tooth, projecting obliquely towards
the left. The ventral portion of the right ramus is denticulate
and striate from the transverse groove to the anterior bend. The
left ramus is excavate ventrally, opposite the denticulate margin
of the right ramus. The unci are nearly symmetrical; each has
two prominent teeth; to the larger, ventral tooth of each ramus
there are attached two fairly large pre-uncial teeth. The basal
plate of the unci is rounded dorsally and has three slender, ob¬
lique ribs crossing it, differently spaced on the two unci. The an¬
terior ends of the manubria are expanded into very broad, thin
lamellae; the posterior section is slender and curves slightly for¬
ward at the extreme end. Under the upper ends of the manubria,
and starting from the dorsal tips of the rami, are two slender,
strongly curved rods, imbedded in the walls of the mastax and
aiding in its support during the pumping action. The oesophagus
is of moderate length and very slender. The gastric glands,
stomach-intestine, ovary, and bladder are normal. The foot
glands are small and slightly club-shaped.
The retrocerebral organ consists of a very large, clear, vacuolate
sac, nearly circular in outline and strongly compressed dorso-
ventrally; no subcerebral glands are present. The eye-spot is
at the posterior end of the large, pyriform ganglion.
Total length 300/a; toes 12/x; trophi 35/x.
Notommata lenis is common In weedy ponds all over the United
States, at least wherever we have been able to make suitable col¬
lections.
This species is readily recognizable by the divided terminal
foot joint and the double dorsal antenna.
Notommata Placida Harring and Myers, new species
Plate XLII, figures 3-6
The body is fusiform and slender, its greatest width being
about one fourth of the total length. The integument is very
flexible, but the outline is fairly constant. It is a very trans¬
parent animal.
The length of the head segment is nearly equal to its width,
or about two thirds of the greatest width of the body; the neck
588 Wisconsin Academy of Sciences , Arts , and Letters.
segment is of the same length, but slightly wider posteriorly.
The anterior transverse folds are well marked. The abdomen in¬
creases very slightly in width for one half of its length and then
tapers gradually to the foot. The tail projects very slightly on
the dorsal side. The foot has but a single, very stout, joint. The
toes are short, about one fifteenth of the total length, stout, coni¬
cal, and acutely pointed.
The dorsal and lateral antennae are minute setigerous papillae
in the normal positions.
The corona extends down on the ventral side nearly one third
of the length of the body ; the post-oral portion projects from the
ventral side of the body as a slight chin. The auricles are small,
and the ciliation is continuous with the corona.
The mastax is virgate, and the trophi are of relatively simple
form without conspicuous asymmetry. The fulcrum is relatively
short, slender, and slightly tapering; the posterior end is some¬
what incurved and slightly enlarged for the attachment of the
muscles. The rami are nearly semicircular in ventral view and
very slightly curved toward the dorsal side at the anterior ex¬
tremity. The inner edges are smooth, without any trace of den-
ticulation. The right uncus has five, and the left uncus four,
straight teeth, slightly clubbed at the point and gradually de¬
creasing in size toward the dorsal margin. The anterior end of
the manubrium is moderately expanded, the posterior slender and
nearly straight, with an external, membranous keel. Two rela¬
tively large, roughly triangular epipharyngeal pieces aid in the
support of the mouth during the pumping action.
The oesophagus is rather short and slender. The gastric
glands, stomach-intestine, and ovary are normal. The bladder
appears to be formed by an expansion of the cloaca. The foot
glands are very large, elongate-ovate, and terminate in a very
small mucus reservoir at the base of the toes. The retrocerebral
sac is moderately large and ovate, while the subcerebral glands
are unusually large and pyriform; no bacteroids are present. The
eye-spot is at the posterior end of the saccate ganglion.
Total length 130/a; toes 9/a; trophi 30/a.
Notommata placida has been found only in small numbers in
weedy ponds at Kenilworth, near Washington, District of Colum¬
bia. While without any striking external characteristics, it
should be readily recognized by the general form of the body,
Earring and Myers — The Rotifers of Wisconsin. 589
the well-developed retrocerebral organ, and the short, conical
toes; it has a superficial resemblance to Proales decipiens, but
the corona and auricles are distinctive. An examination of the
trophi is in any case sufficient for determination.
notommata tripus Ehrenberg
Plate XLIX, figures 9-13
? Vorticella felis Muller, Verm. Terr. Fluv. I1 (1773): 108; Anim.
Infus. (1786), p. 301, PI. 43, figs. 1-5.
? Eeclissa felis Schrank, Fauna Boica 32 (1803) : 109.
? Furcularia felis Lamarck, Hist. Nat. Anim. sans Vert. 2 (1816): ,39.
? Distemma felis Ehrenberg, Isis (Oken) 26 (1833): col. 247.
Notommata tripus Ehrenberg, Infusionsth. (1838), p. 434, PI. 50, fig. 4.
— Leydig, Zeitschr. Wiss. Zool. 6 (1854): 37, PI. 3, fig. i28. — Plate,
Jenaische Zeitschr. Naturw. 19 (1886): 24. — Hudson and Gosse,
Rotifera (1886) 2: 22, PI. 17, fig. 4. — Wierzejski, Rozpr. Akad.
Umiej. Wydz. Mat.-Przyr. Krakow II, 6 (1893): 228. — Skorikov,
Trav. Soc. Nat. Kharkow 30 (1896): 286. — Weber, Rev. Suisse
Zool. 5 (1898): 443, PI. 17, figs. 19, 20. — von Hofsten, Arkiv Zool.,
Stockholm 61 (1909): 34. — Lie-Pettersen, Bergens Mus. Aarbog
(for 1909) 191015: 39. — Voigt, Susswasserfauna Deutschlands, pt.
14 (1912): 97, fig. 176.— Mola, Ann. Biol. Lac. 6 (1913): 241.—
Weber and Montet, Cat. Invert. Suisse, fasc. 11 (1918) : 113.
? Plagiognatha felis Dujardin, Hist. Nat. Zooph. (1841), p. 652.
? Notommata onisciformis Perty, Mittheil. Naturforsch. Ges. Bern
1850: 19; Zur Kenntn. kleinst. Lebensf. (1852), 39, PI. 1, fig. 3.
Notommata pilarius Gosse, Hudson and Gosse, Rotifera (1886) 2: 23,
PI. 17, fig. 5. — Levander, Acta iSoc. Fauna et Flora Fennica 123
(1895): 30. — Lie-Pettersen, Bergens Mus. Aarbog (for 1909)
191015: 40.
Notommata miraMlis Stokes, Ann. Mag. Nat. Hist. VI, 18 (1896): 26,
PI. 8, figs. 20, 21.
The body is very short and broad, its greatest width being
about two fifths of the total length. The integument is very
rigid, almost semi-loricate, and the outline constant. It is fairly
transparent.
The head and neck form a single short and very broad seg¬
ment, its width being about three fourths of the greatest width
of the body; dorsally it has a very large hump, which makes the
head appear almost squarely truncate anteriorly. The single an¬
terior transverse fold is well marked. The abdomen increases
gradually in width to a point about mid-length and is rounded
590 Wisconsin Academy of Sciences , Arts , and Letters.
posteriorly, with a projection carrying the tail and toes; no true
foot is present. The abdomen is flattend ventrally and has two
longitudinal keels or lateral expansions, beginning at the anterior
transverse fold and most prominent at about two thirds the length,
where the cross section of the body is nearly semicircular; from
this point it decreases in size and disappears at the base of the
posterior projection. The rounded tail has dorsally a very large
spur with a knob-like enlargement at the base and a conical point.
The toes are fairly long, about one tenth of the total length, very
robust, acutely pointed, and slightly curved; they are carried
wide apart and appear to be incapable of any lateral motion.
The dorsal and lateral antennae are small setigerous papillae
in the normal positions.
The corona extends down on the ventral side about one third
the length of the body; the post-oral portion forms a fairly
prominent chin. The auricles are very short and stout, and the
ciliation is continuous with the corona.
The mastax is virgate, and the trophi are highly asymmetric.
The fulcrum is very long, slender, and tapering; the posterior
end is slightly expanded and incurved. The rami are roughly
triangular in ventral view and are bent at an approximately
right angle at the extreme anterior point. Continuing the line
of the fulcrum, the rami have near their base a blunt projection,
bifid at the tip, which represents the basal apophysis of the
malleate mastax; behind this projection the rami are slightly ex¬
cavate. The right ramus is minutely denticulate and finely
striated; the left ramus has four blunt teeth on its inner margin.
The alulae are very unequally developed; the right is of normal
size, ending in an acute point, while the left is of huge propor¬
tions, very broad and strongly curved, and continuing backward
until nearly opposite the mid-point of the fulcrum. The unci
have a triangular basal plate ; the left uncus has a large, strongly
developed ventral tooth, which is clubbed at the point and has a
fairly large pre-uncial tooth attached to its ventral edge; closely
appressed to this is a slightly shorter and much more slender
tooth, from the clubbed tip of which a diagonal rib crosses the
basal plate to the external dorsal angle. The ventral tooth on the
right ramus is slightly shorter than its mate on the opposite side ;
it is followed by two very slender teeth, and from the tip of the
third tooth a diagonal rib crosses the basal plate to the external
dorsal angle. The anterior half of the manubrium is a very
Harring and Myers — The Rotifers of Wisconsin. 591
broad, roughly triangular lamella; the posterior half is a slender
rod, slightly expanded at the tip and curving towards the ventral
side. The oesphagus is unusually short. The gastric glands,
stomach-intestine, ovary, and bladder are normal. The foot glands
are fairly large and pyriform.
The retrocerebral sac is moderately large and nearly spherical ;
it usually contains a sufficiently large number of bacteroids to
render it virtually opaque. The ganglion is large and saccate
with the eye-spot at its posterior end.
Total length 175-200/*,; toes 18-20/*,; spur 16-22/*,; trophi 30y.
Notommata tripus is common everywhere among the minute
algae covering submerged water-plants.
Ehrenberg cites as a doubtful synonym Vorticella felis Muller
and its later transformations at the hands of Schrank and
Lamarck ; it has been included here also, for the sake of complete¬
ness, although Muller’s figure shows perhaps more resemblance to
a Mytilina (= Salpina) than to Notommata tripus.
Notommata Venusta Harring and Myers, new species
Plate L, figures 1-4.
The body is rather short and moderately stout without any
very prominent external characteristics. The integument is soft
and flexible, but the outline is nevertheless quite constant.
The head and neck are not distinctly separated ; an in¬
conspicuous fold marks the junction of the neck and abdomen.
The latter is slightly enlarged posteriorly and tapers rather ab¬
ruptly to the foot, which is composed of two short, slender joints,
the anterior one a little larger than the posterior. The tail is
small and indistinct, with a single rounded, median lobe. The
toes are about one fifteenth the length of the body, almost straight
and unusually slender, tapering to very acute points. . The
dorsal and lateral antennae are small setigerous pits in the normal
positions.
The corona continues down the ventral side of the body about
one fourth the length and terminates in a slightly projecting chin.
The auricles are small and continuous with the corona.
The mastax is virgate with unusually slender and strongly
asymmetric trophi. The moderately long fulcrum is large at the
592 Wisconsin Academy of Sciences, Arts, and Letters .
base, but tapers rapidly to the slender posterior portion, the end
of which is slightly incurved. The right ramus is produced pos¬
teriorly as an acute-angled point; on its inner edge there is a
blunt, tooth-like projection, fitting into a corresponding depression
on the inner edge of the left ramus, the external angle of which ends
in a rather blunt lobe. This tooth correponds to the subsquare,
shear-like blade on the rami of N. cerberus and other species. Both
rami are strongly curved in the median plane, and the inner edges
of the dorsal points are striated. The unci have a single promi¬
nent ventral tooth, followed dorsally by a slender diagonal rib
and a second small tooth ending in a blunt knob. The manubria
are very slender ; from a point just below the anterior end a very
long spur curves inwards, which, with a long and slender rod at¬
tached to the rami at its ventral end, serves to support the walls of
the mastax when functioning as a pump. The oesophagus is com¬
paratively short; there is no very distinct separation between
stomach and intestine, and the gastric glands are small and ovate.
The ovary is somewhat elongate and irregular in outline. A small
bladder is present. The foot glands are small and of the usual
club-shape.
The retrocerebral sac is pyriform and small, its length being
but little more than that of the ganglion. No subcerebral glands
are present in this species. The eye-spot is a rather small, lenticu¬
lar body at the posterior end of the ganglion.
Total length 200-225/*; toes 15/*; trophi 24/*.
A few specimens of this species were collected in a swampy
region near English Creek, west of Atlantic City, New Jersey.
It seems to be rare, as it has not so far been found elsewhere.
Notommata venusta does not appear to be very closely related
to any other described Notommatid, at least not to the point of
possible confusion. The straight, very slender toes and the small
auricles distinguished it from other small species of the genus,
and the dorsal spur on the anterior portion of the manubrium is
distinctive.
Earring and Myers — The Rotifers of Wisconsin.
593
Notommata Pygmaea Harring and Myers, new species
Plate LYI, figures 5-8
The body of this species is short, stout, and spindle-shaped;
its greatest width is about one third of the total length. The in¬
tegument is soft and very flexible, but the outline remains fairly
constant. It is a moderately transparent species.
The head and neck segments are short and very broad; their
length is about two thirds of the width, which is nearly three
fourths of the greatest width of the body. The anterior trans¬
verse folds are not strongly marked. The abdomen increases
slightly in width for one half its length and then tapers gradually
to the tail, which has a single, inconspicuous, rounded median
lobe. The foot has two short, broad joints. The toes are short,
straight, and conical; at mid-length they are abruptly reduced to
slender, acute points. Their length is about one twelfth of the
total length.
The dorsal and lateral antennae are small, setigerous papillae
in the normal positions.
The corona extends down on the ventral side for about one
fourth of the length of the body; the post-oral portion projects
as a slight chin. The auricles are short and stout, and the cilia-
tion is continuous with the corona.
The mastax is virgate, and all its parts are of very simple form.
The rami are approximately triangular in ventral view and have
well developed alulae ; at the extreme anterior point they are bent
at a nearly right angle, and the dorsal portion is obscurely den¬
ticulate. The edges of the rami are produced as thin lamellae,
giving the effect of a semi-domed structure, which effectively sup¬
ports the mastax during the pumping action. The fulcrum is a
long, slender, straight, and slightly tapering rod, without any
posterior expansion or crutch. The unci have a short, broad basal
plate and a single, well developed ventral tooth. The manubria
are somewhat curved rods, slightly expanded at both ends. The
gastric glands, oesphagus, stomach-intestine, ovary, and bladder
are normal. The foot glands are large and elongate-pyriform.
The retrocerebral organ consists of a nearly spherical sac open¬
ing on the corona through a long tubular duct, and two subcere¬
bral glands partly fused with the very large, pyriform ganglion.
No bacteroids are present in either sac or glands. The eye-spot
38— S. A. L.
594 Wisconsin Academy of Sciences , Arts , and Letters.
is at the posterior end of the ganglion; it is very large, flattened,
and disk-shaped.
Total length 100/a; toes 8/a; trophi 22/a.
Notommata pygmaea occurs rather sparingly in swampy ponds
near Atlantic City, New Jersey.
This very small species is readily recognized by its short, stout
body and by the form of the toes.
notommata saccigera Ehrenberg
Plate XLVXI, figures 1-5
Notommata saccigera Ehrenberg, Abh. Akad. Wiss. Berlin (for 1831)
1832: 133; Infusionsth. (1838), p. 434, Pi. 50, fig. 8. — ? Hudson and
Gosse, Rotifera (1886) 2: 24, PI. 17, fig. 2. — Wierzejski, Rozpr.
Akad. Umiej. Wydz. Mat.-Przyr. Krakow II, 6 (1893): 228. — Ln>-
Pettersen, Bergens Mus. Aarbog (for 1909) 191 015: 42. — Mola, Ann.
Biol. Lac. 6 (1913): 242.
The body of this species is fusiform, short, stout, and strongly
gibbous posteriorly; its greatest width is more than one third of
the total length. The integument is leathery and the outline
constant. The color is slightly brownish, and the animal is not
very transparent.
The head segment is short and very broad; its width is fully
two thirds of the greatest width of the body. A transverse fold
separates the head from the neck, but there is no separation be¬
tween neck and abdomen; the latter increases very gradually in
width for about half its length and is truncate and strongly gib¬
bous posteriorly, falling off abruptly to the very small tail, which
has only a single, rounded, median lobe. The foot has two very
short and narrow joints. The toes are minute, conical, slightly
decurved, and blunted at the tips ; their length is less than one
twenty-fifth of the total length.
The dorsal and lateral antennae are small setigerous papillae
in the normal positions.
The corona extends down on the ventral side more than one
third of the entire length; the post-oral portion projects from
the surface of the body as a fairly prominent chin. The auricles
are short and very stout, almost triangular; the ciliation is con¬
tinuous with the corona.
j Earring and Myers — The Rotifers of Wisconsin. 595
The mastax must probably be considered as belonging to the
virgate type, but the pumping action has been completely lost.
The trophi are strongly asymmetric and very robust. The fulcrum
is a short and very broad plate without any posterior enlargement.
The rami are lyrate in ventral view and are bent nearly at a right
angle at the extreme anterior point; the dorsal branch is rather
short. The right ramus has on its inner edge a very large lamellar
and semicircular projection, the margin of which is marked with
radial striations and very faintly denticulate; at the apex of the
ramus there are two broad and very blunt teeth. The left ramus
is strongly concave on the inner ventral edge, which is relatively
broad and marked with numerous very narrow transverse striae,
giving it the appearance of being denticulate; the large apical
teeth of the right ramus are lacking. The alulae are fairly
prominent. The ventral tooth of the left uncus is extremely large,
rather short and strongly clubbed at the point ; this is followed by
three minute, comblike, rudimentary teeth and a fairly large,
rather blunt dorsal tooth at the junction of the dorsal rib and
a curved diagonal rib crossing the basal plate from the base of
the large ventral tooth. The right uncus has a large ventral
tooth, blunt and clubbed at the tip; from its base radiate two
similar teeth, nearly as large; on the dorsal margin is a small
tooth which, like the opposing tooth on the left uncus, is formed
by the junction of the dorsal rib and a diagonal rib starting
from the base of the ventral tooth. The manubria are unequally
developed, the right being only three fourths of the length of
the left and its posterior end nearly straight, while the left
manubrium curves strongly inward and toward the dorsal side.
The anterior ends of both manubria are broadly expanded, the
thin lamina on the dorsal side continuing for three fourths of
the entire length. Below the posterior edge of the dorsal branch
of each ramus there is a short, robust, curved rod imbedded in
the walls of the mastax; its ventral end rests on the inner sur¬
face of the ramus and the dorsal ends meet below the tips of the
rami; in the normal virgate, pumping mastax this rod without
doubt aids in the support of the walls of the mastax during the
pumping action, but it is not evident what its function, if any,
may be in this case.
The oesophagus is long and slender; gastric glands, stomach-
intestine, and ovary are normal. The bladder is formed by a
596 Wisconsin Academy of Sciences , Arts , and Letters.
slight expansion of the cloaca. The foot glands are very small
and pyriform.
The retrocerebral organ reaches in this species its extreme
development. At the end of a very slender, tubular duct, one
third as long as the body, there is a very large, pyriform, vacu¬
olate sac, containing bacteroids in large numbers, especially at
the margin, and strongly compressed dorso-ventrally ; the combined
length of sac and duct is more than one half of the entire length
of the animal. The subcerebral glands are short, reaching but
little beyond the posterior end of the ganglion, where the large
eye-spot is situated.
Total length 325-350/* ; toes 12-14/* ; trophi 75/*.
Notommata saccigera is common in submerged or very wet
sphagnum, but it does not occur in ponds unless strayed from
its normal habitat.
This species is readily recognized by its stout, truncate body
and very short foot and toes, as well as by the remarkably de¬
veloped retrocerebral sac. It seems improbable that the animal
here described is identical with Gosse’s, but it agrees fairly well
with Ehrenberg ’s rather meager description.
NOTOMMATA SILPHA (GoSSe)
Plate XLY, figures 1-7
? Notommata forcipata Gosse, Hudson and Gosse, iRotifera (1886) 2:
23, PI. 18, fig. 1; not Notommata forcipata Ehrenberg, 1838. — Mon-
tet, Rev. Suisse Zool. 23 (1915) : 321. — Weber and Montet, Cat.
Invert. Suisse, fasc. 11 (1918) : 115.
Biglena silpha Gosse, Jour. Royal Micr. Soc. 1887: 2, PI. 1, fig. 2;
Hudson and Gosse, Rotifera, Suppl. (1889) , p. 30, PI. 31, fig. 22.
Notommata silpha Harring, U. S. Natl. Mus. Bull. 81 (1913) : 79.
The body is elongate, fusiform, and very slender ; its greatest
width is about one sixth of the total length. The integument is
very flexible, but the outline is fairly constant. It is a very
transparent animal.
No anterior folds are present to indicate the boundaries of the
head and neck ; the body is nearly parallel-sided for about two
thirds of its length ; posteriorly it is much reduced and wrinkled.
The tail is very small and has only a single, rounded lobe. The
Harring and Myers — The Rotifers of Wisconsin. 597
foot is very short and appears to have only a single joint. The
toes are very short, conical, and slightly decurved; the extreme
tips are somewhat abruptly curved inward and upward. At the
base of the toes there is a very small, blunt knob-like projection
on the posterior margin of the terminal foot joint; the length
of the toes is about one twentieth of the total length.
The dorsal and lateral antennae are minute setigerous papillae
in the normal positions.
The corona has two strongly ciliated areas corresponding to
the auricles of other species, but not evertile. Close to the dorsal
margin there are two small papillae, near the openings of the
ducts of the retrocerebral sac. The buccal field is evenly ciliated
and projects strongly in the region around the mouth, giving the
corona in lateral view a decidedly angular appearance ; the mouth
is situated immediately above the most prominent part of the
corona.
The mast ax is apparently a highly modified form of the virgate
type ; the pumping action has been completely lost, and the robust
and strongly asymmetric trophi have become adapted to prehen¬
sion. They can be thrust out through the mouth quite as ef¬
fectively as in the forcipate type of mastax. The fulcrum is
very short and lamellar and tapers from the base to less than
half the with at the posterior end. The rami are deeply bifur¬
cate and very dissimilar in form. The right ramus has near the
base a large, rounded, ventral projection, and behind this a
roughly parallel-sided ventral branch, obtusely pointed at the tip ;
the inner margin is thickened and marked with numerous very
narrow transverse striae, giving it the appearance of being den¬
ticulate ; the striae are continued as a broad border on the ventral
side. The dorsal branch is approximately triangular and has a
fairly prominent tooth on its inner edge, just above the bottom
of the deep transverse groove separating it from the ventral
branch. The left ramus has a knob-like or tooth-like, slightly
recurved ventral projection near the base; it is, like the right
ramus, divided by a deep, transverse groove into a ventral and
a dorsal branch. The ventral branch is fairly broad and lamellar ;
on the anterior, obliquely truncate margin are two prominent
triangular teeth; the dorsal branch has two relatively long, sharp
teeth on the inner margin and a long, tapering, curved projection
from its anterior end towards the dorsal side of the mastax; this
projection is totally lacking on the right ramus. The right alula
598 Wisconsin Academy of Sciences , Arts , and Letters.
is a narrow, membranous plate, while the left is very large and
roughly triangular; near the external angle there is a prominent,
hooklike posterior projection. Each uncus has three functional
teeth, slightly clubbed at the tips and united by the basal plate
for a considerable part of their length. The ventral tooth of the
left uncus does not reach the manubrium, but is joined to the
very large second tooth in its entire length. The manubria are
broadly triangular anteriorly; the median rib is nearly straight.
In front of the ventral branch of the left ramus there is a slightly
curved rod, indented at the anterior, lamellar end, and imbedded
in the ventral wall of the mastax ; its function is no doubt to sup¬
port the mouth, but it is without any counterpart on the right
side.
The oesophagus is long and slender. The gastric glands, sto¬
mach-intestine, ovary, and bladder are normal. The foot glands
are moderately long, slender, and slightly club-shaped.
The ganglion is large and saccate ; the eye-spot is at the
posterior end. The retrocerebral sac is nearly hemispherical,
vacuolate, and fairly large ; only infrequently does it contain bac-
teroids. There is no trace of the subcerebral glands.
Total length 160-200/x; toes 8— 10/x ; trophi 24/>t.
Notommata silpha is widely distributed, but usually in small
numbers.
This species is readily recognized by the unusual form of the
corona, the much reduced, wrinkled abdomen, and the shape of
the toes. Whether Gosse ’s Notommata forcipdta belongs here or
not is uncertain; if it really has auricles, as stated by Gosse, it
is obviously different, but the rest of his description fits N. silpha
fairly well, and it is possible that he may have mistaken the frontal
papillae for auricles.
notommata pseudocerberus de Beauchamp.
Plate XL VII, figures 6-9
'Notommata cerberus de Beauchamp, Arch. Zool. Exp. IV, 6 (1907): 4,
figs. 1, 2; Zool. Anz. 31 (1907): 905, figs. 1-3; not Notommata cer¬
berus Gosse, 1886.
Notommata pseudocerberus de Beauchamp, Zool. Anz. 33 (1908): 400;
Arch. Zool. Exp. IV, 10 (1909): 101, 116, 198, 337, figs. VII B, XX,
PI. 1, figs. 1, 2, PI. 4, fig. 33, PI. 8, figs. 106, 107.
Hawing and Myers — The Rotifers of Wisconsin. 599
The body is fusiform and moderately slender, its greatest width
being somewhat less than one third of the total length. The in¬
tegument is leathery and the outline fairly constant. It is a
moderately transparent animal.
The width of the head segment is equal to its length and a
little more than one half the greatest width of the body. A trans¬
verse fold separates the head from the neck, but there is no con¬
striction between neck and abdomen; the latter increases very
regularly in width for nearly two thirds of its length and then
tapers gradually to the tail, which has a small, rounded median
lobe and two very small lateral lobes. The foot has two joints,
the terminal but half the length and width of the anterior. The
toes are moderately long, conical, slender, and very slightly de-
curved; their length is about one twelfth of the total length.
The dorsal and lateral antennae are small setigerous papillae
in the normal positions.
The corona extends down on the ventral side for about one
fourth of the entire length; the post-oral portion projects from
the surface of the body as a slight chin. The rostrum or cuticu-
lar fold above the mouth is well marked. The auricles are moder¬
ately long and stout, and the ciliation is continuous with the
corona.
The mastax is virgate, but feebly developed, and the trophi are
of very simple form, adapted to support the walls of the mastax
only ; it is at the end of a short pharyngeal tube. The fulcrum is
long, slender, and slightly tapering, without any posterior en¬
largement. The rami are symmetrical, very broad, lamellar
throughout, and bent at an obtuse angle at the extreme anterior
point; there are no marginal teeth or other irregularities on the
inner edges. The unci have a membranous basal plate without
any definite inner border, and a single, very slender tooth; on
each side there are four or five very minute pre-uncial teeth.
The manubria are very short and expanded anteriorly into thin
plates. Nearly level with the lower edges of the rami, two slender,
curved rods are imbedded in the walls of the mastax; their ven¬
tral ends rest on the inner surface of the rami, and the dorsal
ends meet at a considerable distance below the tips of the rami.
Two vestigial, non-functional salivary glands occupy the posterior
portion of the mastax.
The oesophagus is very long and slender. Gastric glands,
stomach-intestine, and ovary are normal. The bladder is formed
600 Wisconsin Academy of Sciences, Arts, and Letters.
by an expansion of the cloaca and is transversely elongate. The
foot glands are moderately large, slender, and slightly club-shaped ;
at the base of the toes are two very small accessory foot glands.
The retrocerebral organ is well developed; the sac is large
and pyriform and usually contains numerous bacteroids, collected
principally immediately behind the eye-spot ; the subcerebral
glands are fully as long, and nearly as large, as the sac. The
ganglion is fairly large and saccate ; at its posterior end is the
large eye-spot.
Total length 500-600/*; toes 30-35/*; trophi 45/*.
Notommata pseudocerberus is widely distributed in the United
States, but in very small numbers. It was first described by de
Beauchamp from ponds at Chaville (Seine-et-Oise) and later col¬
lected at other localities in France; at first identified with Notom¬
mata cerberus (Gosse), it was later recognized as a different species
and given a new name. Dr. de Beauchamp has kindly sent us
material from the type locality for comparison with our American
specimens, which seem to be on the average somewhat smaller in
size.
Notommata contorta (Stokes)
Plate LVIII, figures 5-8
Diglena contorta Stokes, Ann. Mag. Nat. Hist. VI, 19 (1897): 630, PI.
14, fig. 5.
Notommata contorta Harring, U. S. Natl. Mus. Bull. 81 (1913): 78.
The body is elongate, sprindle-shaped, and very slender; its
greatest width is only about one eighth of the total length. The
integument is very flexible and the outline constantly changing.
No anterior transverse folds are present to indicate the bounda¬
ries of the head and neck; the frontal margin of the head pro¬
jects as a blunt knob, corresponding to the rostrum of the forcipate
Notommatids, Notommata pseudo cerberus and a few other species.
The width of the head is nearly equal to the greatest width of the
body, while the neck is only about two thirds as wide. The ab¬
domen increases very gradually in width for about two thirds of
its length and then tapers rapidly to the base of the minute,
three-lobed tail. The very small foot appears to have only a single
joint. The toes are very short, about one thirtieth of the total
Earring and Myers — The Rotifers of Wisconsin. 601
length, and abruptly reduced to acute points, one eighth the
length of the toe.
The dorsal and lateral antennae are minute setigerous papillae
in the normal positions.
The corona has two strongly ciliated areas corresponding to the
auricles of other species, but not evertile. The buccal field is
evenly ciliated and continued down on the ventral side of the
body for nearly one third of its length as a very narrow, parallel¬
sided band, resembling the chin of Notommata copeus. The cir-
cumapical eiliation has completely disappeared.
The mastax is of a highly specialized virgate type; it is re¬
markable for being at the end of a very long pharyngeal tube, and
its functions are consequently limited to suction alone, a fact
which is made still more evident by the extremely simple form
of the trophi, adapted exclusively to the support of the walls of
the mastax. The fulcrum is a long, slender rod, incurved at
its posterior end and disconnected from the incus, or at least
very loosely connected to it; under the action of solvents it sep¬
arates almost at once from the rami. These are strongly curved
and have prominent alulae, the general outline being lyrate with
the tips nearly meeting dorsally. The incus is approximately at a
right angle to the fulcrum. The manubria are long, very slender
rods, with a slight ventral curvature at the posterior end, and
an anterior crescent-shaped expansion. The unci are two straight,
very slender, divergent rods, resting with one end on the rami
and the other on the median, rod-like portion of the manubrium.
The posterior portion of the mastax is occupied by the rudimen¬
tary salivary glands, which are apparently no longer functional,
thus recalling the mastax of Notomma pseudocerberus, described
by de Beauchamp. The gastric glands, stomach-intestine, and
ovary are normal; a bladder does not appear to be present. The
foot glands are long, slender, and slightly club-shaped.
The retrocerebral organ consists of a small, rounded sac and
two large subcerebral glands, more than twice the length of the
sac; no bacteroids are present. The ganglion is large and sub¬
square ; there is no eye-spot.
Total length 220-25 0/a; toes 7-8/a; trophi 15/a.
Notommata contorta is not rare in swampy ponds, although it
does not occur in large numbers.
602 Wisconsin Academy of Sciences , Arts , and Letters .
This species is easily recognizable by the elongate form of the
body and by its never ceasing, worm-like contortions, which evi¬
dently suggested the specific name to Stokes.
Notommata Trypeta Harring and Myers, new species
Plate L, figures 5-8
The body is short and stout, its greatest width being nearly
one third of the total length. The integument is soft and flex¬
ible, but the outline is fairly constant.
The head and neck together form a single segment; its length
is nearly equal to the width, about three fourths of the greatest
width of the body. The anterior transverse fold is well marked.
The abdomen increases very slightly in width for three fourths
of its length; posteriorly it is rounded and ends in a small tail
with a single rounded median lobe. The foot is relatively long,
fairly stout, and obscurely two-jointed. The toes are short, about
one sixteenth of the total length; at the base they are very large
and then abruptly reduced, recurved and incurved, resembling the
chela of a decapod crustacean.
The corona has two lateral tufts of strong cilia, resembling
auricles, but neither retractile nor evertile; it continues down on
the ventral side with a few scattered, weak cilia.
The dorsal and lateral antennae are minute setigerous papillae
in the normal positions.
The mastax is of a modified virgate type. The rami are
elongate, slightly curved, triangular plates without any denticu-
lation. The fulcrum is a straight, slightly tapering rod, enlarged
at the posterior end. The unci are extremely small, triangular
plates, resting on the tips of the rami. The manubria are very
slender, double-curved rods; at the posterior end there is an ir¬
regular enlargement with a ventrally projecting, oval lamella. The
epipharynx consists of two large, very thin, triangular lamellae;
the points project very slightly through the mouth. The oesoph¬
agus, gastric glands, and ovary are normal; a sharp constriction
separates the stomach from the intestine. No bladder is present.
The foot glands are long, slender, and slightly club-shaped.
The ganglion is large and ovate; the only indication of a re-
trocerebral organ is a cup-shaped cell, filled with pigment and
surrounding the eye-spot at the posterior end of the ganglion.
Total length 150/x; toes 9/x; trophi 1 6^ long, 22[x wide.
Harring and Myers — The Rotifers of Wisconsin.
603
Notommata trypeta is parasitic in the Cyanophycean Gompho ■-
sphaeria ; the material on which this description is based was
collected in pools on the southwestern shore of Lac Yieux Desert,
Vilas County. Nearly every colony had its parasite, but no ani¬
mals were found free-swimming; when removed forcibly from the
algal host, they swim rapidly about in search of a new colony.
It was not discovered how the animal manages to penetrate the
tough gelatinous matrix; the feeble trophi seem to be very poorly
adapted to this purpose.
Proales Gosse
proales decipiens (Ehrenberg)
Plate LI, figures 5-8
Notommata decipiens Ehrenberg, Abh. Akad. Wiss. Berlin (for 1831)
1832: 132; Infusionsth. (1838), p. 431, PI . 52, fig. 6. — Tessin, Arch.
Naturg. Mecklenburg 43 (1890) : 143, PI. 1, fig. 5.
Notommata vermzcul\aris Dujardin, Hist. Nat. Zooph. (1841), p. 648,
PI. 21, fig. 7. — Eckstein, Zeitscbr. Wiss. Zool. 39 (1883): 362, PI.
25, fig. 29. — Plate, Jenaische Zeitschr. Naturw. 19 (1886) : 23.
Proales decipiens Hudson and Gosse, Rotifera (1886) 2: i36, PI. 18, fig.
6.-— Skorikov, Trav. Soc. Nat. Kharkow 30 (1896): i290. — Weber,
Rev. iSuisse Zool. 5 (1898): 466, Pi. 18, figs. 16, 17. — Voigt, For-
schungsber. Biol. Stat. Plon. 11 (1904) : 41; Susswasserfauna
Deutscblands, pt. 14 (1912): 90, fig. 158. — de Beauchamp, Arch.
Zool. Exp. IV, 10 (1909): 160. — Weber and Montet, Cat. Invert.
Suisse, fasc. 11 (1918): 100.
? Taphrocampa clavigera Stokes, Ann. Mag. Nat. Hist. VI, 18 (1896):
18, PI. 7, fig. 2.
Pleurotrocha decipiens von Hofsten, Arkiv Zool., Stockholm 61 (1909):
12.
The body of this species is elongate, slender, and fusiform; its
greatest width is one fourth of the total length. The integument
is soft and flexible, but the outline is quite constant. The entire
body is very transparent.
The length of the head segment is a little greater than its
width, which is approximately one half of the greatest width of
the body; it is rounded anteriorly, and this portion is separated
from the head proper by a slight transverse fold ; this section cor¬
responds to the rostrum of the forcipate Notommatids. The neck
segment is rather short and slightly wider than the head. The
604 Wisconsin Academy of Sciences , Arts , and Letters.
anterior transverse folds are well marked. The abdomen increases
very gradually in width for half its length and then tapers to the
foot; it has a transverse fold on the posterior portion, behind the
lateral antennae. The foot is short and broad, continuing the
general outline of the body without marked reduction ; it has only
one joint. The toes are comparatively short, stout, and conical,
ending in acute points; their length is about one sixteenth of the
total length.
The dorsal and lateral antennae are minute setigerous papillae
in the normal positions.
The corona is oblique and has laterally two strongly ciliated
areas, corresponding to the auricles of other Notommatids. The
ciliation of the buccal field does not extend beyond the mouth ;
as in the forcipate Notommatids, the circumapical band has dis¬
appeared.
The mastax is virgate, but preserves much of the general ap¬
pearance of the malleate type. The incus is nearly straight; the
fulcrum is triangular and terminates in a knob-like enlargement.
The rami have a large basal apophysis and, near mid-length, two
prominent teeth ; the dorsal ends are pointed and also act as teeth.
The right uncus has five, and the left four well developed teeth,
gradually decreasing in size toward the dorsal side and united by
a plate-like web. The manubria are broad, roughly triangular
plates, reinforced by prominent ribs. In front of the rami are
two epipharyngeal rods, bent at a right angle and with a knob¬
like enlargement at the apex.
The oseophagus is long and slender. The gastric glands,
ovary, and bladder are normal. The stomach is separated from
the intestine by a marked constriction. The foot glands are very
large and club-shaped.
The ganglion is very large and saccate. The eye-spot is asym¬
metrically placed; in all the specimens we have examined it has
been on the right side, but Yoigt reports that specimens collected
by him in the Kleine Madebrocken See near Plon had the eye-
spot on the left side. The sac is pyriform and appears to be
somewhat variable in size ; the figure represents average condi¬
tions. Some years ago material was collected in the Potomac
River at Washington which agreed perfectly with Stokes’s descrip¬
tion of Taphrocampa clavigera; the sac was filled with bacteroids,
and was jet black and opaque. When the animals were pre¬
served, the black color of course disappeared, and they were then
Harring and Myers — The Rotifers of Wisconsin. 605
seen to be indistinguishable from normal specimens of Proales
decipiens. It is therefore probable that under certain conditions
the sac may be filled with bacteroids, but normally it is perfectly
transparent. No subcerebral glands are present.
Total length 175-250/a; toes 12-16/x; trophi 15/x long, 23/x wide.
Proales decipiens is common everywhere in weedy ponds and is
always abundant in wet sphagnum.
This species is closely related to Proales sordida , but is
distinguishable by its smaller size, somewhat stouter form, re¬
latively larger toes, and the oblique corona.
PROALES SORDIDA GoSSe
Plate LI, figures 9-12
Proales sordida Gosse, Hudson and Gosse, Rotifera (1886) 2S: 37, PI.
18, fig. 7. — Wierzejski, Rozpr. Akad. Umiej. Wydz. Mat -Frzyr.
Krakow II, 6 (1893): 229. — Skorikov, Tray. Soc. Nat. Kharkow 30
(1896): 291. — Voigt, Siisswasserfauna Deutschlands, pt. 14 (1912):
90, fig. 159.— Mola, Ann. Biol. Lac. 6 (1913): 243.— Weber and
Montet, Cat. Invert. Suisse, fasc. 11 (1918): 100.
Pleurotrocha sordida Harring, U. S. Natl. Mus. Bull. 81 (1913): 85.
The body is elongate, fusiform, and slender; its greatest width
is somewhat less than one fourth of the total length. The integu¬
ment is soft and flexible, but the outline is fairly constant. The
entire body is very transparent.
There is no separation between the head and neck segments;
the width of the two fused segments is about three fourths of
the greatest width of the body, and their length is equal to the
body width. There is a single, well marked, anterior transverse
fold. The abdomen increases slightly in width for one half its
length and then tapers gradually to the base of the foot, where
there is a marked transverse fold. There is no distinct tail. The
foot is very robust and fairly long, continuing the general out¬
line of the body without any marked reduction in size; no di¬
vision into joints is distinguishable. The toes are short, straight,
and conical; the tips are abruptly reduced to acute points. The
length of the toes is about one twentieth of the total length. At
their base is a small, rounded, knob-like spur, projecting dorsally.
The dorsal antenna is in the normal position ; the lateral an¬
tennae are somewhat farther back than usual.
606 Wisconsin Academy of Sciences , Arts, and Letters.
The corona is frontal and has a strongly ciliated lateral area,
serving for locomotion; this corresponds to the auricles of the
genus Notommata.
The mastax is of a primitive virgate type and has considerable
resemblance to the malleate form. The incus is nearly straight
and the fulcrum triangular. The rami have a large basal
apophysis of complicated form; their inner edges are without
teeth. The right uncus has five, and the left four, well developed
teeth, decreasing in size from the ventral to the dorsal side
and united by a plate-like web. The anterior portion of the
manubrium is a broad plate of irregular outline; posteriorly it
is prolonged as a slightly curved, short rod. In front of the
mastax there is an epipharynx, the two parts of which are asym¬
metric and consist of an irregularly curved plate with a short,
rod-like lateral projection.
The oesophagus is long and slender. The gastric glands are
normal in form and position. The stomach and intestine are
separated by a marked constriction. The ovary is very large and
somewhat elongate. The bladder is small. There are two pairs
of foot glands, one pair rudimentary and the other quite small;
they discharge through a mucus reservoir at the base of the toes.
To the posterior end of the large ganglion is fused a hemis¬
pherical retrocerebral sac, which presents the usual vacuolate ap¬
pearance and is ductless. The eye-spot consists of a clear globule
with a disc of red pigment behind it, and it is placed some distance
to the right of the median line, at the junction surface of gang¬
lion and sac.
Total length 275-300/*; toes 15 /*; trophi 48/*.
Proales sordida is not very common; we have collected it at
several stations in Wisconsin, but always in small numbers.
This species is recognizable by the terminal corona, slender,
fusiform body, asymmetric eye-spot, and the small spur over the
foot. It is uncertain how many of the literature references
really do concern this species, as Gosse’s description is so incom¬
plete. We submitted specimens to Mr. Rousselet, who verified
our identification and informed us that this is Gosse’s species, ac¬
cording to tradition. The animal has a habit of curving the body
toward the dorsal side, and nearly always dies in this position.
Marring and Myers — The Rotifers of Wisconsin. 607
proales parasita (Ehrenberg)
Plate LI, figures 1-4
Notommata parasita Ehrenberg, Infusionsth. (18i38),p. 426, PI. 50, fig. 1.
— Cohn, Zeitschr. Wiss. Zool. 9 (1858): 291, PI. 13, figs. 8, '9.
Proales parasita Rousselet, Proc. Royal Irish Acad. 31, pt. 51, (1911) : 8.
— Voigt, iSusswasserfauna Deutschlands, pt. 14 (1912): 89, fig. 155.
The body is short, stout, and fusiform; its greatest width
is nearly one third of the total length. The integument is very
flexible, but the outline is fairly constant. The stomach is usually
so filled with chlorophyl that it is difficult to make out the in¬
ternal organs.
The width of the head segment is nearly equal to its length,
and but little less than the greatest width of the body; it is
truncate anteriorly. The neck segment is only half the length of
the head and a trifle narrower. The anterior transverse folds
are well marked. The abdomen increases very slightly in width
for one half its length and then tapers gradually to the foot. The
tail is not very prominent ; it has a single, rounded, median lobe.
The foot is vevry broad and obscurely two- jointed and continues
the general outline of the body without any marked constriction.
The toes are large at the base and taper abruptly to acute points.
The dorsal and lateral antennae are small setigerous papillae
in the normal position.
The corona is frontal and has two lateral, strongly ciliated
areas, corresponding to the auricles of the genus Notommata.
The mastax is of a modified virgate type. The fulcrum is a
long, straight, slender, tapering rod; the rami are somewhat asym¬
metric, the right side being more developed than the left. The
basal apophysis is a nearly semicircular plate, thickened at the
edge and directed towards the ventral side. The rami are ir¬
regularly pentagonal in ventral view. The unci have three rudi¬
mentary teeth, united by a thin lamella. The manubria are ex¬
panded into broad plates at the anterior end and continue as
slender, slightly curved rods. Imbedded in the walls of the mas-
tax, immediately behind the mouth, are two slender, curved rods,
which represent a rudimentary epipharynx.
The oesophagus is long and slender. The gastric glands,
stomach-intestine, ovary, and bladder are normal. The foot glands
608 Wisconsin Academy of Sciences , Arts, and Letters.
are large and ovate, terminating in a small mucus reservoir at
the base of the toes.
The ganglion is large and saccate. The eye-spot is an aggre¬
gation of red pigment granules in a small, hemispherical vesicle,
which probably represents a rudimentary retrocerebral sac. There
is no trace of the subcerebral glands.
Total length 140-160/*; toes 10/*; trophi 15/*.
Proales parasita is parasitic in Yolvox colonies. The name
has very frequently been used for another Volvox parasite, Asco-
morpha volvocicola (Plate), which is much more common. Our
material was collected in the “Widespread” of the Yahara River,
between Lake Monona and Lake Kegonsa, south of Madison, Wis¬
consin. The two parasites are occasionally found inhabiting the
same Yolvox colony in apparent peace and amity. The confusion
of names was started by Gosse in 1852, when he found the animal
later described by Plate as Hertwigia volvocicola and thought he
had Ehrenberg’s species; the error was continued in the Rotifera
and was recently corrected by Rousselet in the Proceedings of the
Royal Irish Academy, volume 31.
Proalinopsis Weber
PROALIN OPSIS CAUDATUS (Collins)
Plate LII, figures 1-5
Notommata caudata Collins, Science Gossip 1872: 11, text fig- — Mon-
tet, iRev. Suisse Zool. 23 (1915): 320, PI. 13, fig. 32.
Covens caudatus Hudson and Gosse, Rotifera (1886) 2: 33, PI. 16, fig. 5.
— Bergendal, Acta Univ. Lundens. 28 (1892), sect. 2, no. 4, p. 81,
PI. 4, fig. 25. — Bilfinger, Jahresh. Ver. Naturk. Wiirttemberg 50
(1894): 45. — Weber, Rev. Suisse Zool. 5 (1898): 461, PI. 18, figs.
13-15. — Stenroos, Acta Soc. Fauna et Flora Fennica 171 (1898):
130. — de Beauchamp, Zool. Anz. 31 (1907): 910. — Lie-Fettersen,
Bergens Mus. Aarbog (for 1909) 191015: 42. — Voigt, Siisswasser-
fauna Deutschlands, pt. 14 (1912): 95, fig. 171 — Lucks, Rotatorien-
fauna Westpreussens 19*12: 50.
Proalinopsis caudatus Weber, Weber and Montet, Cat. Invert. Suisse,
fasc. 11 (1918): 98.
The body is fusiform, moderately slender, and gibbous poste¬
riorly; its greatest width is less than one fourth of the total
length. The integument is very flexible, but the outline remains
■I
TRANS. WIS. ACAD., VOL. XX.
PLATE L.
HARRSNG and MYERS. — ROTIFERS
COCKAYNE BOSTON
j Earring and Myers— The Rotifers of Wisconsin. 609
quite constant. It is the most transparent of all the Notommatid
rotifers.
The head segment is very small; its width is only one half of
the greatest width of the body. The neck is nearly the same
width as the head, long and slender, without clearly defined
posterior limit. A single anterior transverse fold separates the
head and neck. The abdomen is oval in outline and gibbous
dorsally, ending posteriorly in a short, collar-like tail. The foot
is spindle-shaped and has two joints, the proximal much larger
than the posterior and with a knob-like dorsal projection bearing
a few long, deflexed setae. The toes are moderately long, about
one twelfth of the total length, lanceolate, acutely pointed, and
slightly decurved.
The dorsal antenna is a small setigerous papilla in the normal
position; the lateral antennae are minute tubules with a few very
long setae.
The apical plate of the corona is frontal, with a marginal wreath
of cilia and a lateral, auricle-like tuft of very long cilia; the buc¬
cal field is oblique and evenly ciliated.
The mastax is virgate, but retains a strong resemblance to the
malleate prototype. The fulcrum is long, slender, and slightly
tapering with a posterior enlargement for the attachment of the
muscles. The rami are triangular and form a nearly right angle
with the fulcrum; on the upper surface of the right ramus there
are nine short transverse ridges continuing over the inner edge
as blunt, knob-like teeth, interlocking with eight similar teeth
on the left ramus. The right uncus has eight, and the left seven,
straight teeth, resting in the grooves between the transverse
ridges of the rami; the ventral tooth in each uncus is more
robust than any of the others and clubbed at the point; the re¬
maining teeth gradually decrease in size towards the dorsal,
marginal tooth, which is linear. The manubria are triangular
anteriorly, and the median branch curves posteriorly towards the
ventral side.
The oesophagus is very long and slender. The gastric glands
are elongate and pyriform. Stomach and intestine are separated
by a slight constriction. The ovary is normal and the bladder
very large. The foot glands are very small and slender, reaching
only to the middle of the anterior fooot joint.
39 — S. A. L.
610 Wisconsin Academy of Sciences, Arts , and Letters .
The ganglion is large and saccate; the only remnant of the
retrocerebral organ is a minute sac behind the eye-spot ; it is
rendered semi-opaque by included bacteroids.
Total length 200-225/a ; toes 17-20/a ; trophi 13/a.
Proalinopsis caudatus is fairly common in weedy ponds, but
readily escapes observation on account of its excessive transpa¬
rency. It is easily distinguishes from other Notommatids by the
small corona, gibbous abdomen, and the unusual form of the foot.
The setigerous papilla on the anterior foot joint probably cor¬
responds to one frequently found on the terminal foot joint at
the base of the toes; it is, however, more prominent than in
other species. The toes are always carried closely appressed and
have never been seen to separate ; whether they are really immov¬
able is not known.
BIRGEA Barring and Myers, new genus
Birgea enantia Harring and Myers, new species
Plate LVIII, figures 1-4 '
The body of this species is short, broad, and truncate ante¬
riorly; the width is one third of the length of the body proper.
The integument is flexible, but the outline is fairly constant.
No transverse folds are present to indicate the limits of the
head and neck. The head is very broad, as wide as the widest
part of the abdomen. The neck is represented by a slight con¬
striction behind the dorsal antenna, about three fourths the width
of the head. The abdomen is ovate in outline and somewhat ab¬
ruptly reduced posteriorly to a short, bulky tail. The foot is ex¬
tremely slender, and its length is one fourth of the length of the
body; it has three joints, of which the posterior equals in length
the two anterior joints. The toes are moderately long and
lanceolate, ending in long, acute points; their length is one tenth
of the total length.
The dorsal antenna is a very large, knob-like projection at
the base of the head with a small tuft of setae at the center;
the lateral antennae are very slender and relatively long tubules
with a few very long setae.
The corona is frontal and has a circumapical band of short
cilia, interrupted dorsally and passing in a curve to lateral, au-
HARRING AND MYERS. - ROTIFERS
COCKAYNE BOSTON
Earring and Myers— The Rotifers of Wisconsin. 611
ricle-like areas of strongly developed cilia; the buccal field is
closely set with short cilia.
The mastax is of a remarkable type and without any parallel
among known species of rotifers. The normal apparatus for
seizing or triturating the food, the trophi, has in this instance
become very nearly, if not completely, atrophied, and its func¬
tions taken over by a pair of ‘ 1 pseudunci ’ These have appar¬
ently been developed from the secondary epipharyngeal structures,
which are frequently present in the Notommatoid mastax, but
always in a very subordinate role: to spread the mouth open, as
in the forcipate type, or to support the anterior region of the
head during the pumping action, as in Madia. The large, hooked
pseudunci are expanded at their posterior ends into broad lami¬
nae, which are imbedded in the walls of the buccal canal. When
at rest the points meet at a right angle, with the apex dorsal, and
protrude slightly through the mouth. The trophi are relatively
large, but the various parts are very feebly developed; they lie
obliquely in the buccal canal, with the fulcrum directed toward
the mouth. The fulcrum is long, rod-shaped, and divided for
more than half its length. The rami are elongate and rather
narrow, gradually attenuated toward the tips, and imbedded in
the soft parts of the mastax; they do not appear to possess either
abductor or adductor muscles. The left uncus has five, and the
right four, long, linear teeth, united by a very thin, membranous
basal plate. The manubria are heart-shaped. The action of the
mastax has not been studied in the living animal, but from its
structure it is not unreasonable to assume that the trophi are
capable of making a partial revolution around a transverse axis,
as in AsplancJma, which at the same time thrusts the pseudunci
out through the mouth, brings their points into the same plane,
and spreads them apart. This could all be accomplished by re¬
latively simple muscular arrangements. It would be necessary to
effect an inward movement of the external edges of the basal
plates of the pseudunci, which are connected to the bases of the
rami, in order to bring the points into the plane of the rami;
continuing the turning movement in this plane would open the
pseudunci. A pair of adductor muscles attached to the rami
near mid-length would be sufficient to bring this about.
The oesophagus is short. The walls of the stomach are crowded
with symbiotic zoochlorellae, and, as is usually the case among
rotifers sharing this relationship, no gastric glands are present,
612 Wisconsin Academy of Sciences, Arts, and Letters. *
but the stomach is produced into a number of gastric caeca;
there are four of these on the anterior wall, and four ventral and
two small posterior ones in the tail. The ovary and bladder are
normal. The foot glands are nearly cylindric and somewhat
longer than the foot.
The ganglion is small and has a large eye-spot on the ventral
side, which appears disk-shaped in dorsal view. No retrocerebral
organ is present.
Total length 240-275/a; length of foot 35— 40/x ; toes 24^28/a;
trophi 50/a.
Birgea enantia was found in considerable numbers in a swampy
pond near Eagle Biver, Vilas County, Wisconsin. A single speci¬
men was collected at Bargaintown, near Atlantic City, New
Jersey, some years ago.
The taxonomic position of this species is very uncertain. While
the trophi proper have a certain resemblance to those of the
Asplanchnids, this is probably fortuitous; at any rate there is
nothing else in the anatomy pointing in this direction. It may
with some plausibility be connected with the Proales group ; while
this is as yet without definite limits, the three species described
here show some points of resemblance to Birgea; the incus is
nearly straight, rather elongate, and of relatively simple form,
the teeth of the unci resemble the primitive malleate type, and
a fairly well developed, paired epipharynx is present in all three
spcies, notably in Proales sordida. The structure of the corona
does not disprove this assumption; if anything it is in favor of
it. The form of the body is very different, but the importance
of this in the taxonomy of rotifers is still an open question; it
may be of unequal value in the different groups.
We take great pleasure in naming this genus for Dr. E. A.
Birge, president of the University of Wisconsin and of the Wis¬
consin Geological and Natural History Survey.
Drilophaga Vejdovsky
Drilophaga Judayi Harring and Myers, new species
Plate LII, figures 6-8
The body is elongate and spindle-shaped, with some indistinct
transverse folds. The cuticle is soft and flexible and the entire
body very transparent. The anterior portion of the head is
TRANS. WIS. ACAD., VOL. XX.
PLATE Li!.
HARRSNG and MYERS.— ROTIFERS
cockatnt:
BOSTON-
Harring and Myers — The Rotifers of Wisconsin. 613
cylindric and elongate; the mouth is at the center of a slight,
strongly wrinkled elevation, about one fifth of the body-length
from the anterior end. The abdomen is not strongly marked off
from the head ; its greatest width is about mid-length of the body.
A small tail projects slightly over the base of the foot, which is
very short and apparently unjointed. The toes are minute, coni¬
cal, and very slightly decurved; their length is about one twen¬
tieth of the length of the body.
The dorsal antenna is a small setigerous pit in the normal po¬
sition; the lateral antennae have not been observed.
The corona is a simple, circumapical ring of cilia, all very
nearly of the same length.
The trophi are virgate and less specialized than in the two
other species of the genus.The fulcrum is a slender, slightly taper¬
ing rod without any lateral extensions for the attachment of
muscles; the rami are strongly curved and their inner edges
slightly thickened, but without any teeth; at their bases they are
separated by a fairly large, lyrate opening. The large, triangu¬
lar, and very thin alulae distinguish this species from D. delagei
and D. bucephalus. The manubria are slender, double-curved
rods, crutched at the posterior ends; the unci are two oval plates
with an elongate opening at their inner ends; it is possible that
this opening is not really such, but closed with a very thin web.
The salivary glands of the mastax are large, but do not reach
the extreme development of D. delagei ; there is no obvious long¬
itudinal division.
With the exception of the large germary, the digestive and
reproductive organs call for no special remarks. The foot glands
are extremely small, with a mucus reservoir nearly equaling them
in size.
The ganglion is very large and saccate, filling nearly the entire
head anterior to the mastax. No trace of a retrocerebral organ or
eye-spot has been found.
Total length 240-275/*; toes 8/*; trophi 20/*.
This species occurred sparingly among decaying sphagnum
from a roadside ditch near Mamie Lake, Yilas County. We take
great pleasure in naming it for Mr. Chancey Juday, of the Wis¬
consin Geological and Natural History Survey.
While the two previously described species are parasitic, D.
judayi is, as far as now known, free-living; the opportunity for
614 Wisconsin Academy of Sciences, Arts, and Letters.
observation was rather limited, but everything seemed to justify
this conclusion. It was found among sphagnum so far decayed
that no other animals were present which might serve as hosts ;
in fact, the collection was barren of animal life with the exception
of a single specimen of Monostyla lunaris, two specimens of Le-
padella rliomboides, and a few Protozoa. The animal is extremely
sluggish, changing its outline very little at any time. The corona
is normally extended, and the cilia are in constant motion, but they
are so feeble and move so slowly that they appear to be of very little
assistance in locomotion. As a matter of fact, we did not see
the animal effect any considerable displacement at all; the ob¬
served motion was limited to rare and extremely slow bending
movements.
In spite of being non-parasitie, D. judayi agrees in so many
respects with D deldgei and D. bucephalus that it must be in¬
cluded in this genus, and it also shows that Drilophaga is simply
an otherwise fairly normal Notommatid genus which has lost
all but the eircumapical ciliation. It is very unfortunate that
no observations are at hand concerning the biology of this species,
as it would be very interesting to know how an animal apparently
so severely handicapped by nature manages to obtain its food
and to avoid its enemies.
Lindia Dujardin
Lindia Candida Harring and Myers, new species
Plate LIV, figures 10-14
The body of this species is fairly slender, its greatest width
being a little less than one fourth of the total length. The in¬
tegument is very flexible, and the animal is highly contractile.
The entire body is very transparent.
The head and neck segments are of nearly equal length and
width, about two thirds the greatest width of the body. The an¬
terior transverse folds are not strongly marked. The abdomen
increases very little in width for about two thirds of its length;
from this point it tapers rather rapidly to the tail, which has a
single, rather small, rounded median lobe. The foot has two
short joints, the posterior much smaller than the basal joint. The
toes are short, slender, conical, and slightly incurved ; near the
Harring and Myers — The Rotifers of Wisconsin. 615
tip they are abruptly reduced to needle-like points. Their length
is about one sixteenth of the total length.
The dorsal and lateral antennae are small setigerous papillae
in the normal positions.
The corona extends down the ventral side for about one fourth
the length of the body; the post-oral portion projects as a slight
chin. The auricles are relatively small and carry a tuft of close-
set, moderately long cilia, not continuous with the corona.
The mastax differs from the typical form of the genus in the
absence of an epipharynx, as well as in other minor details. The
rami are slightly asymmetric and oval in outline, produced later¬
ally as thin lamellae ; at the anterior angles, where the dorsal pro¬
longation branches off, are two large, blunt, projecting teeth on
each ramus. The fulcrum is rather short, slightly curved, and
tapering to a small posterior expansion. The unci have each
two functional teeth; the left uncus has a large, robust ventral
tooth, slightly clubbed at the tip, which has a decided bend for¬
ward; the second tooth is more slender and a little shorter, with
a similar forward inclination; from its tip two diagonal, slightly
divergent ribs go to the base of the uncus; the dorsal tooth is
very slender, almost linear. A plate-like web unites all the teeth
of the uncus nearly to their tips. Attached to the ventral tooth
at the point is a short, but fairly large, pre-uncial tooth. The right
uncus is slightly different from the left; the two principal teeth
are much more slender, and of the two diagonal ribs starting at
the tip of the second tooth, the dorsal one is absent on the right
uncus. The ventral branch of the manubrium is large and of
very irregular form; its general direction is parallel to the axis
of the body; the median branch is a slender, slightly curved
rod, forming an angle of about 45 degrees with the ventral branch
and united to it by a thin lamella extending well down on both
branches. The dorsal branch is a slender rod, turning back and
joining the median branch with its threadlike point; it acts as
a reinforcement to the dorsal, lamellar extension of the median
branch of the manubrium. As stated above, no epipharynx is
present. There appears to be a rudimentary piston.
The oesophagus is moderately long and slender. The gastric
glands, stomach-intestine, and ovary are normal; a large bladder
is present. The foot glands are small and club-shaped.
The ganglion is small and saccate. The retroeerebral organ
is limited to a very small, ductless sac at the posterior end of
616 Wisconsin Academy of Sciences , Arts , and Letters.
the ganglion; it encloses the eye-spot and is crowded with bac-
teroids, especially at its anterior end, so that it is nearly opaque
by transmitted light.
Total length 250/*; toes 15/x ; trophi 32 /* long, 32/x wide.
Lindia Candida is known only from shallow, weedy areas of
Lac Yieux Desert, Yilas County, Wisconsin, and from a swampy
brook at Oceanville, near Atlantic City, New Jersey.
This species is distinguished from other members of the genus
by its general outline, hyaline body, and the unusual form of
the toes.
Lindia Producta Harring and Myers, new species
Plate LIII, figures 9-14
The body of this species is elongate, spindle-shaped, and
slender, its greatest width being only one fifth of the total
length. The integument is moderately flexible and the outline
quite constant. It is fairly transparent.
The head and neck segments are of nearly equal length and
width, about three fifths of the greatest width of the body. The
anterior transverse folds are not strongly marked. The anterior
portion of the abdomen is nearly cylindrical ; posteriorly it tapers
very gradually to the tail, which is three-lobed, with a small,
rounded median lobe and two minute lateral lobes. The foot
has two joints of very unequal size, the proximal being but little
smaller than the posterior end of the abdomen, while the last
joint is very small and tapers toward the toes. On the posterior
dorsal edge of the first foot joint there is a minute tuft of sen¬
sory setae. The toes are minute, nearly cylindrical, and end in
blunt points; their length is about one thirtieth of the total
length.
The dorsal and lateral antennae are small setigerous papillae
in the normal positions.
The corona extends down the ventral side about one fourth
the length of the body; the post-oral portion forms a slight chin.
The auricles are of the normal Notommata-type, with long, close-
set cilia, continuous with the corona.
The mastax is a somewhat simplified form of the specialized
virgate type of this genus. The rami are of the usual lyrate
Earring and Myers— The Rotifers of Wisconsin. 617
form with well-developed alulae and without the dorsal right-
angled extension usually present. The external edges of the
rami have a thin, lamellar extension of somewhat different form
on the two sides. The width of the rami is variable; the ex¬
tremes are shown on Plate LIII, figures 11 and 12. This is
the only instance of noticeable variation that we have found in
the trophi of the Notommatid rotifers studied. These extremes
are somewhat rare; the majority closely approximate the mean
between the two variations figured. The fulcrum is a fairly
large, subsquare plate, as long as the rami. The unci have two
functional teeth and one rudimentary tooth. The principal tooth
is long, slender, and acutely pointed, with a decided curve near
the tip, to which is loosely attached the rudimentary tooth, about
one third the length of the main tooth. The second functional
tooth is very slender, with a very much enlarged, barb-like tip,
closely appressed to the point of the principal tooth. The manu¬
brium is of the normal form, with a large, crescent-shaped an¬
terior branch forming an approximately right angle with the
median branch, which is rather slender and very slightly de-
curved; the dorsal branch is lamellar and curves inwards. No
epipharynx is present.
The oesophagus is moderately long and slender. The gastric
glands are of moderate size and ovate outline. Stomach and in¬
testine are indistinctly separated. The bladder is large and pyri¬
form. The ovary is of the normal, elongate-ovate form and rather
large. The foot glands are moderately long and slightly club-
shaped.
The ganglion is moderately large and somewhat elongate. The
retrocerebral organ is reduced to a small, ductless sac enclosing
the eyespot at the posterior end of the ganglion and filled with
red pigment granules, most abundant at the anterior surface.
Total length 500-550/a ; toes 12-14/a; trophi 25/a long, 36/a wide.
Lindia producta is fairly common in swampy ponds and bogs
in Polk County, Florida. It has not been found elsewhere.
This species is readily recognized by its large size, general
form, and the very small toes.
618 Wisconsin Academy of Sciences , Arts , and Letters.
lindia torulosa Dujardin
Plate LIII, figures 5-8
Lindia torulom Dujabdin, Hist, Nat. Zooph. (1841), p. 653, PI. 22, fig. 2.
— ? Cohn, Zeitschr. Wiss. Zool. 9 (1858): 288, PI. 13, figs. 1-3 —
Plate, Jenaische Zeitschr. Naturw. 19 (1886): 25.
Notommata roseola Perty, Mitth. Nat. Ges. Bern 1850: 18; Zur Kenntn.
kleinst. Lebensf. (1852) p. 39, PI. 1, fig. 2.
Notommata tardigrada Leydig, Zeitschr. Wiss. Zool. 6 (1854): 39, PI. 3,
fig. 31.
Notommata torulosa Eyferth, Einf. Lebensf. (1878), p. 81. — Hudson
and Gosse, Rotifera, Suppl. (1889), p. 22, PI. ,32, fig. 20. — Bilfinger,
Jahresh. Ver. Naturk. Wurttemberg 50 (1894): 45.— Levander,
Acta Soc. Fauna et Flora Fennica 12P (1895): 32.— Voigt, Siiss-
wasserfauna Deutschlands, pt. 14 (1912): 101, fig. 187. — de Beau¬
champ, Deuxe Exped. Antarct. Frang., Sci. Nat., Doc. Sclent., Roti-
feres (1913), p. 108, fig. 1. — Penaed, Rev. iSuisse Zool. 22 (1914) : 9.
—Weber and Montet, Cat. Invert. Suisse, fasc. 11 (1918): 115.
? Notommata rubra Glas scott, Proc. Royal Dublin Soc. n. ser. 8 (1893) :
48, PI. 3, fig. 7.
? Notommata vorax Stokes, Ann. Mag. Nat. Hist. VI, 19 (1897): 628,
PI. 14, figs. 1-3.
The body of this species is elongate, spindle-shaped, and slen¬
der, its greatest width being only one fifth of the total length.
The integument is fairly flexible and the outline variable. A
rather faint, reddish-orange color extends to all parts of the body.
The head and neck segments are of nearly equal length and
width, a little less than the greatest width of the body. The an¬
terior transverse folds are fairly well marked. The abdomen is
nearly parallel-sided anteriorly; from a point near mid-length
it tapers gradually to the inconspicuous tail, which has a single,
median lobe, rounded posteriorly. The foot is obscurely two-
jointed. The posterior portion of the body has a number of
partly telescopic annulations. The toes are small and nearly
cylindrie ; posteriorly they contract abruptly to minute tubules;
their length is about one fortieth of the total length.
The dorsal antenna is in the normal position; in the material
at our disposal it was not possible to locate the lateral antennae.
Not having had an opportunity to study the living animal, we
have no information on the corona.
The mastax is of a highly specialized virgate type. The rami
are of a lyrate form closely resembling the normal forcipate type.
COCKAYNE BOSTON
Harring and Myers — The Rotifers of Wisconsin. 619
Their tips are bent rather abruptly toward the dorsal side, and
the alulae are well developed. The fulcrum is a roughly triangu¬
lar plate, rounded posteriorly. The unci have three teeth, the
ventral longest and the dorsal about half as long, while the middle
tooth is intermediate in length. The first or ventral tooth is well
developed; the basal two thirds of its length is fairly stout,
straight, and cylindric, while the distal third is enlarged as a
barb-like or sickle-shaped point. The second tooth is of the same
form, but much more slender. The dorsal tooth has a very
slender basal portion, about one half the entire length, and the
terminal portion is of nearly the same size as the tip of the ven¬
tral tooth, but more curved. All three teeth are united by a thin
lamella, extending to the curved tips. The manubria have a
strongly developed, crescent-shaped ventral branch and a nearly
straight median, or principal, branch, forming an angle of about
45 degrees with the ventral branch. The dorsal branch is a thin
lamella, slightly curved toward the median plane. The epipharynx
is composed of two rod-like, obtusely bent pieces, whose inner
ends meet in a straight line immediately behind the mouth, and
two very thin, rhomboid lamellae with a slight double curvature.
From the external angles of these lamellae radiate 12 to 15
evenly spaced striae or very slender ribs. The inner sections of
the epipharynx are very resistant to solvents, but the external
lamellae are much less so; evidently they are only partly scleri-
fied.
The oesophagus is short and slender. Stomach and intestine are
not distinctly separated. The gastric glands are moderately large
and ovate. A large bladder is present. The ovary is of the nor¬
mal elongate-ovate form. The foot glands are small and discharge
into a minute mucus reservoir.
The ganglion is large and slightly alongate. The retroeerebral
organ consists of a rather small, elongate, apparently ductless sac
at the posterior end of the ganglion ; it encloses the eye-spot and is
nearly opaque on account of the numerous bacteroids contained,
which are most abundant at the anterior end.
Total length 350-400/4,; toes 10/4,; trophi 32 /x long, 40/t wide.
Lindia torulosa has not so far been found in the United States ;
we are indebted to Dr. P. de Beauchamp for the material from
which this description has been made; it was collected in the
Canal de Bourgogne at Dijon, France. As the description has
been made entirely from preserved material, it may need modifi-
620 Wisconsin Academy of Sciences , Arts , and Letters.
cation in some details ; the corona could not be studied at all and
will have to be described from living specimens; on this subject
Dr. de Beauchamp writes:
“J’y joins des specimens de Notommata torulosa ou du
moins de 1’espece que j’ai appellee ainsi jusqu’a ce jour.
Peut-etre en effet n’a-t-elle pas droit a ce nom: j’ai ete
etonne en examinant les individus vivants de trouver l’annu-
lation moins nette et les oreillettes moins arrondies que je n’en
avais garde le souvenir”.
"Whether this species is really the one studied by Dujardin and
subsequent authors it will probably forever be impossible to de¬
cide; it does not, however, do violence to the original descrip¬
tion to assume that it is, especially as it is found in France. In
our opinion it ought to retain the name and to be considered the
type of the genus Lindia , of which very little was known until
de Beauchamp described and explained the action of this pe¬
culiar modification of the virgate mastax, which was then com¬
pletely unknown. We are gratified to be able to show that this
modification is not a unique instance, but occurs in six other
species of Notommatids.
Lindia Pallida Harring and Myers, new species
Plate LIII, figures 1-4
? Lindia torulosa Cohn, Zeitschr. Wiss. Zool. 9 (1858) : 288, PI. 13, figs.
1h3.
The body is elongate, almost vermiform, and very slender, the
greatest width being only one sixth of the total length. The
integument is very flexible and fluted longitudinally and also has
several transverse folds, so that it presents an annulate appear¬
ance, almost like a Bdelloid rotifer. It is a very transparent
species.
The head segment is short and narrow, about two thirds of the
greatest width of the body; the neck is nearly twice as long, but
very little wider; in addition to the two transverse folds separat¬
ing head and neck and neck and abdomen there is an additional
faint transverse fold on the neck segment. The abdomen is very
nearly parallel-sided and cylindric; it contracts rather abruptly
to a conspicuous tail, with a single median lobe, rounded poste-
Earring and Myers — The Rotifers of Wisconsin. 621
riorly. The foot has two short, broad joints. The toes are short,
conical, and slightly decurved; their length is one eighteenth of
the total length.
The dorsal and lateral antennae are small setigerous papillae
in the normal positions.
The corona extends down the ventral side about one sixth the
length of the body; the post-oral portion forms an indistinct chin.
The auricles are unusual; at the end of a slender peduncle there
is a tuft of long, closely set cilia, all of the same length, so that
when the auricles are everted they present a striking illusion of
a revolving sphere.
The mastax is of the highly specialized virgate type common
to the entire genus. The rami are of a lyrate form which bears
a considerable resemblance to the forcipate type; the anterior
points are, however, bent somewhat abruptly at an obtuse angle
toward the dorsal side, and at this point are reinforced by a dia¬
gonal, lamellar brace. The alulae are well developed. The ful¬
crum is a flat plate, slightly curved in the median plane. The
unci have two functional teeth; the ventral tooth is long and
slightly clubbed at the point, while the dorsal is short and has
a separate tip, which seems to be very loosely joined to it. At
the tip of the main tooth there is a striated lamella, which evi¬
dently corresponds to the pre-uncial teeth present in the mastax
of a number of Notommatid rotifers. The anterior branch of the
manubrium is very large and broadly crescent-shaped, its posterior
tip curving toward the end of the principal, or median, branch,
which is nearly straight, with a slight ventral curvature at the
extreme end. The dorsal branch of the manubrium is long and
curves toward the median plane; expanded posteriorly by a tri¬
angular lamella, it contributes the dorsal section of the approxi¬
mately hemispherical or dome-shaped arrangement of the trophi.
The epipharynx consists of two ribbon-like pieces, meeting be¬
low the mouth and gradually twisted through an angle of nearly
180 degrees, at the same time curving dorsally and posteriorly.
The oesophagus is short; stomach and intestine are not separated
by any constriction. The gastric glands are very small and nearly
spherical. The bladder is formed by an expansion of the cloaca,
as in Notommata pseudo cerberus. The ovary is of the normal
elongate-ovate form. The foot glands are rather small and slightly
club-shaped.
622 Wisconsin Academy of Sciences, Arts, and Letters.
The ganglion is unusually long and slender. The retrocerebral
organ is limited to a small and apparently ductless sac at the
posterior end of the ganglion; it is usually crowded with bac-
teroids and opaque by transmitted light. The eye-spot is fairly
large and at the posterior end of the ganglion.
Total length 250-300//,; toes 14-1 6/a ; trophi 20//, long, 25/a wide.
Lindia pallida is present wherever wet or partly submerged
sphagnum is to be found; in such locations it may be collected
at any time and any place. We have no information as to its
presence outside of the United States, but there can be little doubt
that it will prove just as common elsewhere. It does not, as far
as known, occur in ponds.
Cohn’s figure and description have been with some doubt re¬
ferred to this species, as they seem to agree better with it than
with the French species, to which de Beauchamp has definitely
limited the name torulosa.
Lindia Annecta Harring and Myers, new species
Plate LIY, figures 6-9
The body is elongate, sprindle-shaped, and very slender, the
greatest width being only one sixth of the total length of the
body. The integument is soft and very flexible; it has a number
of transverse folds, giving it an annulate appearance. It is mo¬
derately transparent.
The head and neck segments are of nearly equal length and
width, but slightly less than the greatest width of the body. The
anterior transverse folds are not strongly marked. The abdomen
is nearly cylindrical, tapering slightly and very gradually to the
tail, which is three-lobed, with a small, rounded median lobe and
two minute lateral lobes. The foot has two rather short, broad
joints, continuing the general outline of the body without any
marked constriction. The toes are short, conical, and slightly
pinched to acute points; their length is about one twentieth of
the total length.
The dorsal and lateral antennae are small setigerous papillae
in the normal positions.
The corona extends down the ventral side about one fifth the
length of the body; the post-oral portion forms an indistinct chin.
TRANS. WIS. ACAD., VOL. XX
PLATE L1V
HARRING and MYERS —ROTIFERS
Harring and Myers- — The Rotifers of Wisconsin. 623
The auricles are approximately of the normal Notommata- type,
with a small tuft of close-set, long cilia.
The mastax differs only in details from the specialized virgate
type common to the genus. The rami are of a lyrate form with
well-developed alulae and without the dorsal right-angled ex¬
tension, thus resembling very strongly the normal forcipate type.
The fulcrum is a short, nearly square plate. The unci have three
unequally developed teeth; the central tooth is long, slender, and
slightly curved; the lateral are only half as long. All three teeth
are joined by an elongate, oval plate with a longitudinal rib; the
ventral tooth does not appear to be connected to the manubrium
except through the plate and the two dorsal teeth of the uncus.
The manubrium is of approximately normal form; the anterior
branch is very large and broadly crescent-shaped; its general di¬
rection is nearly parallel to the longitudinal axis of the body and
approximately at a right angle to the median branch, which is
very slender and slightly expanded at the tip. The dorsal branch
is short, and the walls of the mastax are not supported dorsally
by the trophi. The epipharynx consists of two thin, lamellar
pieces, meeting below the mouth; the anterior portion is rather
narrow, while the posterior is somewhat abruptly expanded into
a broad plate. Together the two sections of the epipharynx form
approximately a semicircle. Rudiments of the ventral glands
of the mastax are present, but they do not appear to be func¬
tional.
The oesophagus is long and slender; it is frequently looped.
Stomach and intestine are indistinctly separated. The gastric
glands are moderately large and rounded. The bladder is of nor¬
mal form. The ovary is large and elongate-ovate in outline. The
foot glands are large and ovate ; they discharge into a small mucus
reservoir.
The ganglion is large and saccate. The retrocerebral organ
consists of a moderately large, ductless sac at the posterior end
of the ganglion; it encloses the eye-spot, and red pigment granu¬
les are scattered throughout its contents, most abundantly at the
anterior surface.
Total length — 300-350/a ; toes 15-18/a ; trophi 18/a long, 30/a wide.
Lindia annecta occurs in sphagnum bogs and ditches near At¬
lantic City, New Jersey, usually in very large numbers. It has
not so far been found elsewhere.
624 Wisconsin Academy of Sciences, Arts, and Letters.
This species may be distinguished from other members of the
genus by the general form, as well as by the robust, acutely
pointed toes.
Lindia Tecusa Harring and Myers, new species
Plate LY, figures 1-4
? Notommata gravitata Lie-Pettersen, Bergensi Mus. Aarbog 190510:
29, PI. 2, figs. 3-5.
The body of this species is elongate, spindle-shaped, and slender,
the greatest width being only one fifth of the total length. The
integument is leathery and the form fairly constant. The color
of the entire animal is a light orange-brown.
The head segment is short and strongly rounded in front, al¬
most semicircular; the neck segment is as long as the greatest
width of the body and considerably narrower. The anterior trans¬
verse folds are well marked. The abdomen is nearly cylindric
for two thirds of its length, the posterior third tapering gradu-
ually to the toes; there is no true foot. Over the cloaca there is
a rounded hump with the outline of a tail on its posterior end.
The toes are very short, broadly conical, and somewhat compressed
laterally; they are not sclerified, as in the case of the great ma¬
jority of rotifers, but are very soft and may be completely in¬
verted and withdrawn within the body under the action of the
longitudinal muscles.
The dorsal antenna is in the normal position ; the lateral an¬
tennae are somewhat farther back than usual.
The corona extends down on the ventral side about one fourth
of the length of the body ; the post-oral portion projects as a chin.
The rostrum or cuticular fold immediately above the mouth is
well marked. No auricles are present; two slight antero-lateral
depressions are somewhat more strongly ciliate that the rest of
the corona. The very long, pendant, bottle-shaped cells of the
corona are conspicuous.
The mastax differs only in minor details from the typical form
of the genus. The rami are broadly lyrate with very large alu¬
lae and a short, nearly right-angled dorsal extension, on which
the unci rest. The fulcrum is of nearly the same length as the
rami, rather slender, and tapers to a blunt point. The unci con¬
sist of two parts, firmly united to each other but not integral; a
Marring and Myers — The Rotifers of Wisconsin. 625
thin plate rests on the manubrium, and at its extremity a stout,
slightly curved piece is joined to it at a right angle. The inner
surface of this curved section of the uncus is very closely set with
minute, very slender spines, which no doubt play a part in tri¬
turating the food. The manubrium is less abnormal than usual
in the genus ; the ventral branch is very slender, but little shorter
than the median branch and nearly parallel to it; the dorsal
branch is broad and plate-like and curves strongly toward the
median line.
The epipharynx consists of a single piece of somewhat com¬
plex form; it is only imperfectly sclerified and dissolves rather
rapidly in a solution of potassium hypochlorite of very moderate
strength. The ventral section is a broad lamella, slightly rein¬
forced at the anterior edge and curved nearly in a semicircle,
at the ends of which there is a deep sinus, reducing its width
nearly to one half; at this point there is a rather sharp outward
and downward bend of the external portion.
The oesophagus is fairly long. The gastric glands and stomach-
intestine are huge and nearly fill the entire body-cavity. The
ovary is elongate and very large; the animal is remarkable for
being viviparous, the only known instance among Notommatid
rotifers; just before birth the embryo crowds all the intestines
out of their normal positions, as it is fully half the length of the
parent. The bladder is very small and a simple expansion of
the cloaca. The foot glands are very small and terminate in a
mucus reservoir almost as large as the gland.
The ganglion is an enormous, broadly pyriform sac reaching
fully half way down on the mastax. At its posterior end there
is a very large eye-spot enclosed in a ductless, hemispherical re-
trocerebral sac, which is tinted a deep red and filled with minute
pigment granules.
Total length 1000-1500/z; trophi 45 /i long, 48/x wide.
Lindia tecusa occurs in abundance in a stagnant, shallow pool
of brackish water at Margate, New Jersey. The pool is only a
short distance from the ocean and receives salt water by seepage
through the ground; during the summer it is filled with dense
mats of Vaucheria thuretii, among which the rotifer lives.
It is possible that this may be the animal described by Lie-
Petterson as Notommata gravitata ; with the small amount of ma¬
terial available to the author it is not surprising that his account
40— S. A. L.
626 Wisconsin Academy of Sciences , Arts , and Letters.
should be incomplete. The large size, brackish-water habitat, and
the sketch of the trophi suggest close relationship to L. tecusa,
but the figure of the foot and toes indicates a somewhat different
species, and until more material is obtained at the type locality
it would seem advisable to treat them as two distinct species.
In this connection attention may be called to Dipodina arctis-
con Ehrenberg, also said to be viviparous, described in a paper
read before the Gesellschaft der Naturforschenden Freunde in
Berlin on November 15, 1842. A report of the meeting was
published in the Spenersche Zeitung, Berlin, for November 24,
1842, and copied in Froriep ’s Neue Notizen zur Nat nr- und- Heil-
kunde, series 2, volume 24 (1842), page 184. The proceedings of
the society for 1839 to 1859 were not published until 1912 (Sitz-
ungsber. Ges. Naturf. Freunde, Berlin, 1839-1859. Berlin, 1912).
On page 47 occurs the following:
“Herr Ehrenberg. . . . Endlicli zeigte derselbe Abbildnngen und
getrocknet erhaltene Exemplare eines unbekannten Raderthierchens der
Ostsee bei Wismar vor, welches sich durch der beiden Zangenhalften
seines Zangenfusses in zwei ganz getrennte Scheiden sehr auszeichnet,
im uebrigen aber dem Genus Notommata ganz ahnlich ist. Er sah es
lebendig gebahrend und hat ihm den Namen Dipodina arctiscon gegeben,
wegen seiner Aehnlichkeit mit dem kleinen Wasserbaren, Arctiscon.”
As far as known, no further description was ever published
of this animal, nor have we been able to locate the drawing ex¬
hibited by Ehrenberg at the meeting; see page 633.
lindia truncata (Jennings)
Plate LIY, figures 1-5
Notommata truncata Jennings, Bull. Mich. Fish Comm. 3 (1894): 16,
figs. 10, 11. — Stenroos, Acta Soc. Fauna et Flora Fennica 171
(1898) : 125, PL 3, figs. 11, 12. — Lie-Pettersen, Bergens Mus. Aarbog
(for 1909) 191015: 41.
The body of this species is elongate, spindle-shaped, and slen¬
der, the greatest width being about one fourth of the total length.
The integument is moderately rigid; a number of transverse folds
give it a annulate appearance. The color of the entire animal
is a deep orange-brown.
The head and neck segments are short and broad, their width
being but little less than the greatest width of the body. The
TRANS. WIS. ACAD., VOL. XX
PLATE LV
HARRING and MYERS.— ROTIFERS
Earring and Myers — The Rotifers of Wisconsin. 627
anterior transverse folds are not strongly marked. The abdomen
is nearly cylindric, tapering slightly and gradually to the in¬
conspicuous, rounded tail, which is not lobate. The foot has two
rather short, broad joints, continuing the general outline of the
body without any marked constriction. The anterior foot joint
has a minute, blunt median lobe, projecting slightly over the last
foot joint. The toes are very short and conical, slightly com¬
pressed laterally, and slightly pinched just below the base; the
tips are slightly blunted. Their length is about one twentieth of
the total length.
The dorsal antenna is somewhat further back than usual; the
lateral antennae have not been observed.
The corona extends down the ventral side about one third the
length of the body; the post-oral portion projects as a slight chin.
The auricles are very robust and carry a tuft of close-set, long
cilia, continuous with the corona.
The mastax differs somewhat from the typical form of this
genus. Seen from the ventral side, it has a strikingly triangular
aspect, due to the presence of two large, pyriform salivary glands,
lying in the ventral angles of the mastax and crowding the pos¬
terior ends of the manubria outward. The rami are lyrate, with
very large alulae and a fairly long, right-angled, dorsal extension,
on which the unci rest. The fulcrum is as long as the rami,
slender and tapering. The unci have three teeth, united by a
plate-like web. The ventral tooth is only half the length of the
uncus, but very large and obtusely conical. The second and third
teeth rest with their base on the manubrium and the tip on the
dorsal extension of the rami ; the second tooth is well developed ;
the third is very slender, almost linear, and forms the dorsal
edge of the uncial plate. The ventral branch of the manubrium
is of the normal crescent shape but somewhat smaller than usual,
and forms an approximately right angle with the nearly straight
median branch. The dorsal branch is short, and forms the an¬
terior margin of a broad lamella projecting dorsally from the
median branch of the manubrium.
The epipharynx consists of two symmetrical, roughly hammer¬
shaped pieces. The crutched ends are immediately behind the
mouth; the main stem extends diagonally backward and toward
the dorsal side, tapering gradually to a thread-like tip ; near the
middle, on the inner side, is a tooth-like projection. The oesopha¬
gus is short and slender. The gastric glands, stomach-intestine,
628 Wisconsin Academy of Sciences , Arts , and Letters.
and ovary are normal ; a large bladder is present. The foot glands
are very small.
The ganglion is fairly large and somewhat elongate. The re-
trocerebral organ is limited to a ductless hemispherical sac en¬
closing the eye-spot at the posterior end of the ganglion ; it is ap¬
parently filled with a liquid, very dark red pigment, almost
opaque by transmitted light, so that the eye-spot is usually in¬
visible in the living animal; in preserved material the color of
the sac clears up somewhat.
Total length 300/*; toes 15/*; trophi 30/* long, 50/* wide.
Lindia trwncata is widely distributed in swampy ponds and
bogs, at times in very large numbers.
This species is readily recognizable by the color of the body
and the peculiar retrocerebral sac. Through the kindness of Dr.
Jennings we have had the opportunity to study an abundance of
preserved material of this species, which he collected around Ann
Arbor, Michigan.
Lindia Fulva Harring and Myers, new species
Plate LVT, figures 1-4
The body is elongate, spindle-shaped, and moderately slender,
the greatest width being about one fourth of the total length.
The integument is very flexible and the outline somewhat vari¬
able. The color of the animal is a light brownish red.
The head segment is strongly rounded anteriorly; its width is
about two thirds of the greatest width of the body and its length
a little less. The neck segment is of nearly equal length and
width. The anterior transverse folds are well marked. The ab¬
domen is very slightly larger at mid-length than at the anterior
and posterior ends and terminates in a very broad, not very promi¬
nent, rounded tail. The foot has two joints, continuing the gen¬
eral outline of the body and marked by slight constrictions ; the
terminal joint is less than half the length of the anterior and car¬
ries the two short, bluntly conical toes, about one twenty fourth
the length of the body; they are reduced to small tubules at the
tips.
The dorsal and lateral antennae are in the normal positions.
COCKA.TKE BOSTON
Earring and Myers — The Rotifers of Wisconsin. 629
The corona extends down on the ventral side about one fourth
of the length of the body; the post-oral portion projects as a
prominent chin. The auricles are very large and powerful; in a
frontal view they appear as nearly circular discs of a diameter
equal to their length and much compressed in the longitudinal
direction of the body. The coronal cells are very conspicuous
through the rather sparse ciliation of the buccal field.
The mastax is of the type of L. truncata (Jennings) and has
the same triangular outline due to the presence of two large,
ventral salivary glands that occupy the space in the ventral angles
of the mastax, between the fulcrum and the manubria. The rami
are of the normal, lyrate form with large alulae and a right-angled
dorsal extension of moderate length, serving for the support of
the unci. The fulcrum is of nearly the same length as the rami
and tapers gradually to the posterior, rounded end. The unci
are large, thin, roughly triangular lamellae with a single, super¬
posed tooth, strongly clubbed, and terminating in an acute point;
a diagonal reinforcing rib crosses the uncus to the external dor¬
sal angle, where there is a rudimentary second tooth. The ven¬
tral branch of the manubrium is nearly crescent-shaped and about
half the length of the median branch, which is very slightly de-
curved. The dorsal branch is fairly long and curves strongly in¬
wards in order to give additional support to the dorsal wall of
the mastax.
The epipharynx consists of no less than four separate pieces;
the principal pair are approximately straight, thin lamellae with
a reinforcing rib at the anterior edge and expanded at the oral
end into a roughly subsquare plate. The second pair are elongate,
nearly parallel-sided, and rounded at the ends; they are twisted
throughout their length, so that the ends lie nearly in the same
plane.
The oesophagus is long and slender. The gastric glands are
remarkable; inserted approximately in the normal positions on
the anterior portion of the stomach they extend for nearly two
thirds of the length of the abdomen as slightly tapering, sinuate
tubes; their contents is a blood-red, opaque, granular plasma
mass, in which the nuclei are not visible, probably on account of
the lack of transparency. At the death of the animal the glands
contract to about one third of their normal length. The stomach
and intestine are normal. Bladder and foot glands are very
i
630 Wisconsin Academy of Sciences, Arts, and Letters.
large; the glands terminate in a mucus reservoir extending into
the cavity of the toe.
The ganglion is very large and ovate; at its posterior end is
the huge eye-spot, enclosed in a ductelss retrocerebral sac, ap¬
parently filled with a liquid, dark red, granular pigment.
Total length 1000-1200/*; toes 40-48 /*; trophi 56/* long, 90/*
wide.
Lindia fulva was found in great abundance in sheltered spots
along the shore of Loon Lake, about one mile south of Eagle
River, Vilas County, Wisconsin. It also occurs in a pond at
Oceanville, near Atlantic City, New Jersey.
This species is in bulk the largest of all known rotifers. The
enormous gastric glands are sufficient to distinguish it from re¬
lated species.
Tetrasiphon Ehrenberg
tetrasiphon hydrocora Ehrenberg
Plate LVII, figures 1-4
Tetrasiphon hydrocora EUrenberg, Ber. Verh. Akad. Wiss. Berlin 1840:
219; Sitzungsber. Ges. Naturf. Freunde Berlin (for 1839-1859)
1912: 12.
Notommata spicatia Hudson, Jour. Royal Micr. Soc. 1885: 612, PL 12,
fig. 5.
Copeus spicatus Hudson and Gosse, Rotifera (1886) 2: 29, PI. 16, fig. 2,
PI. 30, fig. 7. — Stenroos, Acta Soc. Fauna et Flora Fennica 171
(1898): 128. — Lucks, Rotatorienfauna Westpreussens (1912), p. 50,
fig. 7.
'Notommata hydrocora Harking, U. iS. Natl. Mus. Bull. 81 (1913): 78.
The body is elongate and fusiform; its greatest width is less
than one fourth of the total length. The integument is leathery,
and the outline is constant. It is a very transparent species.
No anterior transverse folds are present to indicate the limits
of head and neck; the anterior margin of the head is rounded,
and its width is about one half of the greatest width of the body;
the neck is very slightly narrower. The abdomen is fusiform and
slightly constricted at the base of the tail, which forms a rounded,
blunt, dorsal projection. The foot has two joints, the basal one
fairly large and robust, the terminal only half as wide and long.
The toes are long, about one twelfth of the total length, conical
PLATE LVIL
TRANS. W1S. ACAD., VOL. XX.
HARRING and MYERS.— ROTIFERS
COCKAYNE BOSTON
Harring and Myers — The Rotifers of Wisconsin. 631
and tapering for one half their length, ending in very slender,
acute points.
The dorsal antenna is double ; two fairly large, conical tubules,
ending in a rounded knob with a tuft of long sensory setae, pro¬
ject from the dorsal surface of the head; their distance apart is
about one half the width of the head and they are united by a
conspicuous muscular band, as in Asplamchna. The lateral an¬
tennae are very far back on the body, near the base of the tail;
they are very long, tubular, knobbed at the ends, and carry a
few excessively long setae.
The corona is oblique, and its length is a little more than one
fifth of the total length. It has a median, trough-like depres¬
sion throughout its entire length, and the post-oral portion pro¬
jects as a fairly prominent chin. No auricles are present; at the
anterior lateral angles are two V-shaped, concave, ciliated ap¬
pendages of the corona, which may be rudiments of auricles. The
ciliation of the corona is very faint and may serve only to con¬
vey food to the mouth ; at least it appears to be entirely too feeble
to be of any use in locomotion.
The structure of the mastax is peculiar and very aberrant.
While it is of the pumping type, it has noting else in common
with the normal form of the virgate mastax. Its correct inter¬
pretation is still a matter of doubt, and the affinities of the genus
are therefore unknown ; there is a certain resemblance to the
mastax of the genus Lindia, but it does not extend beyond gene¬
ralities. In spite of long continued efforts, the animal has never
been observed while feeding, and it appears to be necessary to
await some fortunte accident to elucidate fully the action of the
mastax.
The fulcrum is a very small, thin lamella. The rami at first
sight resemble the forcipate type, but instead of being straight
they are curved nearly in a semicircle and enlarged by a broad,
thin lamella attached to the external edges. At the mouth there
are two large, blunt, conical teeth which appear in lateral view
as superposed knobs. The alulae are large. The uncus has only
a single long and slender tooth, resting with its tip on the ramus
immediately below the toothlike projection; the basal plate of the
uncus is represented by a crescent-shaped, extremely thin lamella.
The manubrium is a curved rod with two anterior and two poste¬
rior lamellar, small apophyses. In addition to the incus and
632 Wisconsin Academy of Sciences, Arts, and Letters.
mallei, there is a rather complex supplemental oral armature;
two thin, trapezoidal plates with upturned external and posterior
edges are superposed on the rami and meet at the inner posterior
angles; at the anterior angles are hinged two curved rods, slightly
expanded at the opposing anterior ends. The function of this
complicated apparatus is no doubt to enlarge the oral opening;
the food of the animal consists mainly of desmids and it is able
somehow to swallow entire the very large Micrasteiria torreyi, which
is often found unbroken in the stomach. An enormous piston
very nearly fills the entire interior of the mastax; while it is
bulky, it seems to be very weak, and the points of attachment of
the feeble muscles have not been definitely ascertained. It is
certain, however, that the powerful muscle attached to the fulcrum
in the normal virgate mastax plays but a very subordinate role in
this instance, which fact is clearly indicated by the small size of
the fulcrum.
The oesophagus is very long and muscular and is capable of
being greatly distended. The gastric glands are elongate, con¬
stricted near the middle, and strongly curved, so that in certain
positions four glands appear to be present. The stomach and
intestine are separated by a relatively long, tubular constriction;
the walls of both are syncytial, and the only provision for storing
food reserves is a single ring of spherical cells, attached by a
slender tube to the walls of the stomach at the opening of the
tubular constriction. The ovary is very long, slender, and rib¬
bon-like, and contains from 20 to 30 nuclei. The spinous eggs
seen by Hudson are without doubt the ordinary summer eggs
and not the resting eggs ; the hooked spines serve to attach them
to the water plants. The bladder is large. Four pyriform foot
glands of nearly equal size are present.
The retrocerebral sac is small, projecting but little beyond the
ganglion ; the subcerebral glands are nearly twice as long, slender,
and pyriform; they do not contain bacteroids. The eye-spot is
large.
Total length 750-1000y; toes 60-80 /z; trophi 80/l.
T etrasiphon hydrocora is rather rare and seems to be limited
to non-calcareous regions with very soft, acid waters. It is nearly
always covered with a very firm jelly-case, much more resistant
than the protective covering of any other rotifer except Notom-
mat a peridia, and almost rivaling the egg-cases of the pond-snails
in toughness.
Harring and Myers — -The Rotifers of Wisconsin. 633
We have used Ehrenberg’s name for this species, as it does not
appear that there can be any reasonable doubt about its identity.
Ehrenberg’s description is as follows (Ber. Yerh. Akad. Wiss.
Berlin 1840: 219) :
“274. Tetrasiphon hydrocora Nov. Gen. — Tetr. hydroc. berl. naturf.
Gesellsch. 18 Juni 1839; T. maximus, hyalinus, organis duobus prominu-
lis tubulosis occipitalibns, duobus aliis prope finem dorsi positis, glandu-
lis pancreaticis quatuor globosis, maxillis bidentatis, organo rotatoria
Pleurotrochae obliquo. Pedis digitis gracilibus acutis longis, ocello
occipitali. Long. y8 linea superat. Berolini.'’
The same publication states further (ante, p. 197) :
“Herr Ehrenberg legte hierauf 274 Blatter von ihm selbst ausgefiihrter
Zeicbnungen von eben so vielen Arten in dem 1838 erschienenen grossen
Infusorienwerke noch nicht abgebildeter Infusorien vor, und sprach iiber
die auffallend rasche Entwickelung dieser Kenntnisse. . . .”
We have tried in vain to locate these drawings; as they con¬
tain figures of several rotifers of doubtful identity, such as Cal-
lidina rediviva, Philodina hirsuta, Otoglena papillosa, and Tetra¬
siphon hydrocora , it would be of considerable interest to obtain
all available information concerning them. Dp. Anton Collin
very kindly made a careful search in the Zoological Museum, Ber¬
lin, where a large number of Ehrenberg’s original drawings are
still preserved, but none of the species mentioned above were to
be found. The secretary of the Academy of Sciences in Berlin,
Dr. Waldeyer, replied to our inquiry that Ehrenberg had exhib¬
ited the drawings in question at the meeting of November 9, 1840,
but had not presented them to the Academy, and that their pres¬
ent location, if they were still in existence, was unknown.
The note in the Sitzungsherichte der Gesellschaft Naturfor-
schender Freunde (for 1839-1859), p. 12, cited above, is as follows:
“Herr Ehrenberg .... Ferner legte derselbe mehrere Blatter
detaillierter Zeichnungen von neuen Infusorienformen vor, darunter 4
neue Genera aus der Umgegend Berlins: . . . 4) Tetrosyphon
(sic) hydrocora, ein Raderthierchen mit einem Nackenauge aus der
Familie der Krystallfischchen (Hydatinaea). Letzteres scheint durch
zwei vordere und zwei hintere hervorstehende Rohren dafiir immer mehr
zu entscheiden, dass diese Rohren, deren gewohnlich immer nur eine im
Nacken der Raderthiere vorhanden ist, nicht dem Fortpflanzungssystem,
sondern dem Respirations-Organismus angehoren. Es wurde lebend
vorgezeigt. Eine neue Art von Kletterthierchen ( Xanthidium Echinus)
mit mehrfach verasteten Stacheln schliesst sich zunachst an X. racemo-
sum der Feuersteine an und wurde lebend zum Theil im Leibe des Tetra¬
siphon, als genossene Speise liegend, vorgezeigt . u
634 Wisconsin Academy of Sciences , Arts , and Letters.
Hudson lists Tetrasiphon liydrocora in the Supplement (p. 60)
among the rejected genera ; while admitting that 1 1 this may have
been Copeus spicatus”,, he offers as excuses that “the trophi of
C. spicatus are not bidentate ; moreover no mention is made
of the gelatinous covering in which spicatus is so often enveloped ’ \
As to the trophi, they are unquestionably bidentate, and in some
localities the animal occurs without the jelly-case.
Eosphora Ehrenberg
eosphora najas Ehrenberg
Plate LX, figures 7-11
Eosphora najas Ehrenberg, Abh. Akad. Wiss. Berlin 1830: 47, 84, PL 7,
fig. 3; ibid, (for 1831) 1832: 140, PI. 4, fig. 13; Infusionsth. (1838),
p. 451, PI. 56, fig. 7. — Leydig, Zeitschr. Wiss. Zool. 6 (1854): 40,
PL 3, fig. 29. — Hudson and Gosse, Rotifera, ,Suppl. (1889), p. 27,
Pl. 33, fig. 9. — Bergendal, Acta Univ. Lundens. 28 (1892), sect. 2,
no. 4, p. 91, BL 5, fig. 29. — Skorikov, Trav. Soc. Nat. Kharkow 30
(1896): 297. — Lie-Pettersen, Bergens Mus. Aarbog (for 1909)
191015: 47. — Voigt, Susswasserfauna Deutschlands, pt. 14 (1912):
115, fig. 220.
Eosphora digitata Ehrenberg, Infusionsth. (1838), p. 452, PI. 56, fig. 8.
— Tessin, Arch. Freunde Naturg. Mecklenburg 43 (1890): 145. —
Hudson and Gosse, Rotifera, Suppl. (1889), p. 27, PI. 33, fig. 10.
— de Beauchamp, Arch. Zool. Exp. IV, 3 (1905): CCXXV (notes et
revue), figs. 1-3; ibid., IV, 6 (1907): 22, fig . 12 G; ibid., IV, 10
(1909): 219, fig. XXXI, PI. 13, fig. 26. — Hirschfelder, Zeitschr.
Wiss. Zool. 86 (1910): 217, Pis. 9-11, PL 12, figs. 10-25. — Lie-Pet¬
tersen, Bergens Mus. Aarbog (for 1909) 1910: 47. — Voigt, Siiss-
wasserfauna Deutschlands, pt. 14 (1912): 114, figs. 217-219. — Pen-
ard, Rev. Suisse Zool. 221 (1914): 7, PL 1, figs. 3, 4. — Weber and
Montet, Cat. Invert. Suisse, fasc. 11 (1918): 121, figs. 3i3, 34.
Furcularia najas Dujardin, Hist. Nat. Zooph. (1841), p. 650.
Furcularia digitata Dujardin, Hist. Nat. Zooph. (1841), p. 650.
Notommata eosphora Bartsch, Jahresh. Nat. Wurttemberg 26 (1870):
339.
The body is large and robust; its greatest width is about two
fifths of the total length. The integument is comparatively firm
and the outline of the animal fairly constant. The entire body
is usually of a light orange color.
The head and neck are short and broad, and there is a well
marked transverse fold between the head and neck, as well as
Hcirring and Myers — -The Rotifers of Wisconsin. 635
between neck and abdomen. The abdomen is broad and oval in
outline. The tail is not very large ; it has a median lobe, rounded
posteriorly, and two small lateral lobes. The foot is very nearly
cylindric and obscurely three-jointed; its length is about one
eighth of the total length. The toes are long, straight, and slen¬
der, ending in conical points ; their length is about one fourteenth
of the length of the body.
The dorsal antenna is a small setigerous papilla in the normal
position ; the lateral antennae are small, short tubules with a few
setae at the tip; they are somewhat farther back on the abdomen
than is usually the case.
The corona is frontal and consists of a marginal wreath of
cilia, interrupted dorsally and passing in a curve to the lateral
angles, where it joins an inner circle starting also from the dor¬
sal gap ; from the angles the marginal wreath continues as a single
band, closed ventrally and passing immediately below the mouth.
The buccal field is not ciliated.
The° mastax is of a modified virgate form, in which the pump¬
ing action has been partly lost and the trophi have become adapted
to the seizure of prey, as the animal is carnivorous. The rami
have the form of a nearly equilateral triangle in ventral view;
there are two strong, recurved lateral apophyses for the attach¬
ment of the powerful muscles which open the rami. Near mid-
length they are bent at a right angle, and at this point the left
ramus has a single, large tooth, which interlocks with two similar
teeth on the right ramus. There is an elongate-ovate ventral op¬
ening between the rami, and a narrow space dorsally, between the
frontal teeth and the tips of the rami; the inner edges of this
dorsal extension are minutely denticulate. From the lateral
apophyses a thin lamella passes in a very complicated curve to
the dorsal points of the rami, thus limiting the cavity of the mas-
tax. It should be noted that the only openings in the rami are
the median dorsal and ventral; the irregularly oval areas on the
ventral face are merely thin sections of the same material as the
ribbed structure. The fulcrum is a very broad, subrhomboidal
plate, slightly rounded posteriorly. The unci consist of a rather
small, subsquare basal plate with a single, very robust, club-shaped
ventral tooth; from a point near the middle of the tooth a diago¬
nal rib crosses the basal plate to a point opposite the dorsal mar¬
ginal rib of the manubrium. At the median dorsal angle of the
unci there is a very loosely attached small plate of irregular out-
636 Wisconsin Academy of Sciences , Arts , and Letters.
line, striated on the inner margin. The fuction of this accessory
structure is not clear ; the external end of the plate appears to be
attached to the uncus and the inner end to the dorsal tips of the
rami ; it may possibly act simply as a hinge during the movement
of prehension. The manubrium has a nearly straight central
section, very broad anteriorly, with a thin lamella at each side, and
tapering gradually to the posterior end, which ends in a small
rounded plate or knob. In front of the rami, immediately be¬
hind the mouth, there are two slender, right-angled pieces, serv¬
ing for the support of the lips. The ventral salivary glands are
very conspicuous, the right one nearly twice as long as the left.
The oesophagus is long and begins high up on the mastax. The
gastric glands are small and round. There is a distinct constric¬
tion between the stomach and the intestine ; the latter is very small.
The foot glands are very long, slender, and slightly club-shaped.
The ovary is large and of irregular, elongate outline ; the nuclei are
unusually large.
The ganglion is rather small and saccate. The retrocerebral
organ consists of a small, clear sac and two subcerebral glands
of the same size as the sac. The eye-spot is at the posterior end
of the ganglion, and two pigment spots or so-called accessory
eye-spots are situated on two small projections on the corona, not
far from the openings of the ducts leading from the sac.
Total length 500/x ; toes 36/^ ; trophi 75^.
Eosphora najas appears to be rare in the United States; a few
specimens have been collected at Los Angeles, California, and so
far it has not been found elsewhere.
It seems highly probable that the two names given by Ehren-
berg represent the same animal, the only difference noted being
that E. najas is supposed to have shorter toes than E. digitata ,
and this impression may very well have been obtained from ob¬
servations of animals with partly retracted toes. As najas is
the older specific name, and also the type of the genus, we have
used this in preference to digitata ; the two names appear to have
been used rather indiscriminately by later observers without any
very good reasons being given for their preference.
Harring and Myers — The Rotifers of Wisconsin. 637
EOSPHORA EHRENBERGI Weber
Plate LXI, figures 6-9
Notommata najas Ehrenberg, Abh. Akad. Wiss. Berlin (for 1831)
1832: 132; Infusionsth. (1838), p. 429, PI. 52, fig. 2. — Leydig,
Zeitschr. Wiss. Zool. 6 (1854): 38. — Eckstein, Zeitschr. Wiss. Zool.
39 (1883): 363, PI. 25, fig. 28.— Hudson and Gosse, Rotifera (1886)
2: 25, PI. 18, fig. 2. — Wierzejski, Rozpr. Akad. Umiej. Wydz. Mat.-
Przyr. Krakow II, 6 (1893): 228. — Skorikov, Trav. Soc. Nat. Khar-
kow 30 (1896): 287. — Weber, Rev. Suisse Zool. 5 (1898): 448, PI.
18, fig. 4. — Wesch£, Jour. Quekett Micr. Club II, 8 (1902): 327,
PI. 17, fig. 2. — Lie-Pettersen, Bergens Mus. Aarbog (for 1909)
191<P5: 40. — Daday, Zoologica 59 (1910): 67, PI 3, fig. 17. —
Voigt, .Siisswasserfauna Deutschlands, pt. 14 (1912): 96, figs. 172,
173.— de Beauchamp, Bull. Soc. Zool. France 38 (1914) : 326.
Furcularia najas Dujardin, Hist. Nat. Zooph. (1841), p. 650.
? Notommata pot\amis Gosse, Jour. Royal Micr. Soc. 1887 : 365, PI. 8,
fig. 10. — Hudson and Gosse, Rotifera, Suppl. (1889), p. 21, PI. 31,
fig. 9.
? Notommata najas thermalis Issel, Atti Soc. Ligustica 17 (1906): 29,
PI. 1, figs. 7, 8.
Eosphora ehrenbergi Weber, Weber and Montet, Cat. Invert. Suisse,
fasc. 11 (1918): 123.
The body of this species is broad; nearly parallel-sided and
very robust; its greatest width, at mid-length, is fully one third
of the total length. The integument is comparatively rigid and
the outline very constant. The entire body is tinted a light
orange-brown.
The head is short and broad; its width is but little less than
the greatest width of the abdomen. The neck is as wide as the
head, but somewhat shorter; there is a rather indistinct trans¬
verse fold between head and neck, as well as between neck and
abdomen. The abdomen is very broad and nearly parallel-sided
for the greater part of its length; it is rounded posteriorly and
terminates in a short, very broad tail with two small lateral lobes.
The foot is cylindrical, two- jointed, and fairly long. The toes
are rather short, about one eighteenth of the total length, coni¬
cal, and acutely pointed; the ventral edge is straight and the
dorsal slightly curved.
The antennae are small setigerous pimples in the normal po¬
sitions.
The corona is frontal and consists of a marginal wreath of
cilia, interrupted dorsally and passing in a curve to a lateral,
638 Wisconsin Academy of Sciences, Arts, and Letters.
auricle-like area of strongly developed cilia; the ventral arcs
meet immediately below the mouth. The buccal field is faintly
ciliate.
The mastax is of a specialized virgate type, which has lost
the pumping action and become adapted to the seizure of prey,
as this species is carnivorous. The trophi differ but slightly
from thos of Eosphora tJCerina; they are, however, still more
robust. The rami are approximately triangular, and the two
sides are nearly alike. They are bent at a nearly right angle at
a point near mid-length, where they are provided with two strong
teeth on each ramus. There is a large, irregularly lozenge¬
shaped ventral opening between the rami, and they are provided
with intricately shaped bars, with ribbed edges, to resist defor¬
mation under the pull of the very robust muscles. The fulcrum
is a very broad plate ; the posterior ventral edge is cut off dia¬
gonally, approximately at a right angle to the pull of the power¬
ful abductor muscles, which are attached here. The unci have
a large, subsquare basal plate with a single, very large and
strong ventral tooth, the free length of which is nearly equal
to the length of the basal plate ; from the junction of the tooth
and the basal plate a diagonal rib crosses to a point near the
dorsal edge, opposite the dorsal rib of the manubrium. At the
median dorsal angle of the basal plate there is a triangular,
lamellar, striated expansion of the marginal rib, which probably
acts as a shearing tooth. The manubrium has a strong, nearly
straight central section, curving slightly toward the ventral side
at its extreme posterior end; anteriorly it is expanded into a
broad triangular plate, and it is directed diagonally toward the
dorsal side. In front of the rami, immediately behind the mouth,
there are two tripod-like selerified pieces, serving for the sup¬
port of the mouth. The mastax has two large ventral salivary
glands, the left gland nearly as long as the fulcrum, the right
somewhat longer.
The stomach is very large and without distinct separation from
the intestine. The gastric glands are large and ovate. A con¬
tractile bladder is present. The foot glands are fully as long as
the foot and slightly compressed laterally, no mucus reservoir being
present. The ovary is irregularly ovate and fills the ventral por¬
tion of the body from the mastax to the bladder ; the nuclei are very
large and of irregular polygonal outline.
Earring and Myers — The Rotifers of Wisconsin. 639
The ganglion is rather small and saccate. The retrocerebral
organ consists of a small, clear sac and two subeerebral glands,
as large as the sac; their contents is granular, but they do not
contain bacteroids. The eye-spot is large and very dark red:
it is at the posterior end of the ganglion.
Total length 350-450// ; toes 24-30// ; trophi 65//.
Eosphora ehrenbergi is not common; when it is found, it gen¬
erally occurs in large numbers.
Eosphora Therina ITarring and Myers, new species
Plate LIX, figures 1-5
The body of this species is broad, nearly parallel-sided, and
very robust ; its greatest width, at mid-length, is a little less than
one third of the total length. The integument is very flexible,
but the outline remains fairly constant. The entire body is very
transparent.
The head and neck are not separated by any external fold;
their combined length is somewhat more than one fourth of the
total length, and the anterior width is a little less than the greatest
width of the body; a well marked transverse fold is present be¬
tween the neck and the abdomen. The abdomen is slightly nar¬
rower at the anterior end than at mid-length ; from there it tapers
gradually to the very small tail, which projects very little be¬
yond the base of the foot. There is no marked reduction in width
at the junction of the abdomen and the foot ; the latter is ob¬
scurely three- jointed and conical, its posterior width being only
one half of the width at the base. The toes are rather short,
about one eighteenth of the total length, conical, and slightly com¬
pressed laterally ; posteriorly they end in a minute, but well
marked claw-like tip.
The dorsal antenna is unusually far back on the anterior por¬
tion of the body ; the lateral antennae are approximately in the
normal position. They are all small, setigerous pimples.
The corona is frontal and consists of a marginal wreath of
cilia, interrupted dorsally and passing in a curve to a lateral,
auricle-like area of strongly developed cilia ; the ventral arcs meet
immediately below the mouth. The buccal field is faintly ciliate.
The mastax is of a modified virgate form, in which the pump¬
ing action has been almost completely lost and an adaptation
640 Wisconsin Academy of Sciences , Arts , and Letters.
to the seizure of prey has taken place in accordance with the
carnivorous mode of life of this species. The rami are roughly
triangular and nearly symmetrical. Near mid-length they are
bent at a nearly right angle and armed with two well developed
teeth on each ramus. There is a large, lozenge-shaped ventral
opening between the rami, and they are abundantly supplied
with ribs and webs to transmit the pull of the very strong mus¬
cles. The fulcrum is a very broad plate, serving for the attach¬
ment of the powerful abductor muscles; the posterior ventral
edge is cut off diagonally. The unci have a large, subsquare
basal plate with a single very large and strong ventral tooth, and
from the junction of the tooth and basal plate a diagonal rib
crosses to a point near the dorsal edge, opposite the dorsal rib of
the manubrium. At the median dorsal angle of the basal plate
there is a triangular, lamellar, striated expansion of the marginal
rib; this probably assists, with its mate, in subdividing the food.
The manubrium has a strong, nearly straight central section,
expanded anteriorly into a broad triangular plate, and it is di¬
rected diagonally toward the dorsal side. In front of the rami,
immediately behind the mouth, there are two tripod-like sclerified
pieces, serving for the support of the lips. The salivary glands
are rudimentary.
The oesophagus is long, wrinkled longitudinally, and very dis¬
tensible, permitting a large organism to pass into the stomach. The
gastric glands are large and nearly spherical. There is no con¬
striction between the stomach and the intestine. The bladder is
normal. The foot glands are long and slightly club-shaped, dis¬
charging into a moderately large mucus reservoir at the base of
the toes. The ovary is very large and irregularly ovate ; the nuclei
are deeply buried in its tissues, so that they are not easily seen.
The ganglion is rather small and saccate. The retrocerebral
organ consists of a small, clear sac and two subcerebral glands,
as large as the sac; they do not contain bacteroids. The eye-spot
is a large, lenticular or saucer-shaped body at the posterior end
of the ganglion; it is on the inner side, and thus appears as a
circular disc when seen from the dorsal side.
Total length 450-475/*,; toes 25-28/*,; trophi 72/*,.
Eosphora tJierina was found in a small puddle containing only
a couple of gallons of water, which had gathered in a slight de¬
pression in the mud pumped up from the Potomac River at the
TRAMS. WJS. ACAD., VOL. XX.
PLATE LVII!
MARRING and MYERS.— ROTIFERS
COCKAYNE BOSTON
Earring and Myers — The Rotifers of Wisconsin. 641
time of the construction of Potomac Park, in Washington, District
of Columbia. The surface was covered with Euglena in an un¬
broken layer; the only other rotifer present was the spinous form
of Epiphanes (= Notops) brachionus, but both occurred in enor¬
mous numbers.
Eosphora therina swims with remarkable speed and in an al¬
most straight line; it will dart from the surface to the bottom of
a column of water five inches high in a fraction of a second. While
it has a considerable external resemblance to E. ehrenbergi and
almost identical trophi, the two species are not easily confused;
E. ehrenbergi is of an almost opaque orange-brown color, while
E. therina is as transparent as an Asplanchna, and the small tail
and tapering foot with clawed toes are sufficient to distinguish it
from the former, which has a nearly cylindric foot, a prominent
tail, and clawless toes.
Eosphora Anthadis Harring and Myers, new species
Plate LVIII, figures 9-13
The body of this species is broad and very robust; its greatest
width, at mid-length, is one third of the total length. The in¬
tegument is soft and flexible, but the outline is quite constant.
The entire body is very transparent.
The head is short and very broad; the neck is a slight constric¬
tion, marked with an indistinct fold, between the head and the
abdomen. The very large and stout abdomen tapers slightly from
a point about mid-length to the base of the very broad foot. There
is no distinct tail. The foot is conical and without any joints,
but with some faint wrinkles. In a dorsal view the external
border of the very short toes forms almost a semicircle; they are
strongly compressed dorso-ventrally, so that when seen from the
side they appear as nearly normal, conical toes; the length is
about one twentieth of the total length.
The corona is frontal and consists of a marginal wreath of
cilia, interrupted dorsally and passing in a curve to the lateral
angles, where it joins an inner circle starting also from the dorsal
gap ; from the angles the corona continues as a single band, closed
ventrally and passing immediately below the mouth. The buccal
field is very faintly ciliate.
41— S. A. L.
642 Wisconsin Academy of Sciences , Arts , and Letters.
Tlie dorsal antenna is just in front of the transverse neck fold;
the lateral antennae are placed well forward and nearer the dorsal
side than is usually the case.
The mastax of this species is a modification of the virgate type,
intermediate between Enteroplea (= Triphylus) and Eosphora
and still retaining the pumping action. The rami are symmetri¬
cal and elongate-triangular with a right-angled bend near mid¬
length. There is an elongate, rather narrow ventral opening be¬
tween the rami, and a much larger, ovate opening dorsally be¬
yond the angle. Between these two openings there are four or
five very small teeth in each ramus. The fulcrum consists of two
plates with their dorsal edges joined to form a V. The unci have
only a single, strong, slightly curved tooth; at its base there is a
very small, subsquare plate with two faint striae as the last rudi¬
ments of the normal 4 to 6 teeth. The manubrium is a nearly
straight rod, with a small triangular expansion at its anterior
end. The piston is large and well-developed. Salivary glands
have not been observed; the gastric glands are unusually large.
The stomach is joined to the intestine without constriction. A
large bladder is present. The foot glands are large, elongate, and
almost cylindrical, terminating in a large mucus reservoir at the
base of the toes. The ovary is large and slightly elongate; the
nuclei are irregularly polygonal in outline.
The ganglion is comparatively small and rounded. The retro-
cerebral organ consists of a small sac and two subcerebral glands,
nearly as large as the sac; they do not appear to include bacte-
roids at any time, either in young or in full-grown animals. No
eye-spot is present.
Total length 350-400/*; toes 20-22/*; trophi 33/*.
Eosphora anthadis appears to be widely distributed, although
it is usually found in small numbers.
Eosphora Gelida Harring and Myers, new species
Plate LX, figures 1-6
The body is elongate, nearly parallel-sided, and quite slender;
its greatest width is only one fourth of the total length. The
integument is unusually flexible, and the animal is highly con¬
tractile. The body is very transparent.
The head and neck are not separated by any external fold;
Harring and Myers — The Rotifers of Wisconsin. 643
their combined length is one fourth of the total length, and the
anterior width is almost equal to the greatest width of the body.
The abdomen is slightly narrower anteriorly than at mid-length;
from this point it tapers gradually to the rudimentary tail. The
foot is rather long and tapers gradually to the toes ; there is no con¬
striction at its junction with the abdomen, and it has no apparent
joints. The toes are short, about one twentieth of the total
length, slender and tapering, with slightly blunted points.
The dorsal antenna is unusually far back on the anterior por¬
tion of the body; the lateral antennae are in the normal position;
all are small, setigerous pits.
The corona is frontal and consists of a marginal wreath of
cilia, interrupted dorsally and passing in a curve to the lateral
angles, where it joins an inner circle starting below the dorsal
gap; from the angles the corona continues as a single band,
closed ventrally and passing immediately below the mouth. The
inner circle has a median dorsal tuft of long cilia. The buccal
field is very faintly ciliate.
The mastax is of a modified virgate form, in which the pump¬
ing action is of secondary importance and an adaptation to the
seizure of prey has taken place. The rami are broad and roughly
triangular; there is only a trace of asymmetry. Near mid-length
they are bent at a right angle and have a single, well developed
tooth in each ramus. There is a small, lozenge-shaped ventral
opening between the rami; the inner edges of the upper half are
finely striate. The supporting framework of the mastax consists
of rather slender, round rods of intricate form. The fulcrum
is fairly long and slightly tapering toward the posterior end,
where it passes abruptly into a quadrant-shaped expansion, some¬
what flattened at the tip and with a roughened surface for the
attachment of the muscles of the piston. The unci are reduced to
a single, rather slender tooth with three minute pre-uncial teeth
on the left and two on the right side; at the base of the uncus
there is a diagonal supporting piece of complicated form, with
one end connected to the manubrium and the other to the tooth.
The manubrium is fairly slender, its posterior end curving slightly
toward the ventral side ; anteriorly there is a large, rounded ven¬
tral lobe and a triangular dorsal lobe curving inwards. A slen¬
der, slightly curved rod, resting with its ventral end on the in¬
ner surface of the rami, is imbedded in the walls of the mastax
and aids in their support during the pumping operation. The
644 Wisconsin Academy of Sciences, Arts, and Letters.
salivary glands are rudimentary. The long and slender oesophagus
starts high up on the mastax. The gastric glands are very small
and rounded. Stomach and intestine are not distinctly separated.
The ovary is fairly large and ovate in outline. A bladder of
normal size and form is present. The foot glands are long and
slightly club-shaped.
The ganglion is large and saccate. The retrocerebral organ is
limited to a small, apparently ductless sac at the posterior end
of the ganglion. No true eye-spot is present; but the sac con¬
tains a large number of red pigmented granules, the greater num¬
ber being heaped up in a dense cluster against the posterior end
of the ganglion, where the eye-spot would normally be found,
and the rest scattered through the contents of the sac.
Total length 350-400/x; toes 18-20/*; trophi 50 /*.
Eosphora gelida has been found in ponds around Washington,
District of Columbia; it seems to occur only in the very early
spring, at the time the ice is breaking up ; it is then often found
in considerable numbers.
The elongate, slender body without projecting tail, the short
toes and the absence of subcerebral glands sufficiently distinguish
this sepcies from other members of the genus. In addition, the
integument is marked with fine, closely spaced, longitudinal rows
of minute circular dots. No other rotifer has similar markings,
as far as known; the dots are not all of the same size, but vary
in regular succession from large to small in approximately equal,
short spaces ; the lines of dots are straight and parallel, and there
is only a small space between successive dots. Figure 6, Plate
LX shows the arrangement.
EOSPHORA MELANDOCUS (GoSSe)
Plate LIX, figures 6-10.
Furcularia melandocus Gosse, Jour. Royal Micr. Soc. 1887: 2, PL 1,
fig. 4. — Hudson and Gosse, Rotifera, Suppl. (1889), p. 27, PL 31,
fig. 18. — Bilfinger, Jahresh. Nat. Wiirttemberg 50 (1894) : 47. —
Voigt, Siisswasserfauna Deutschlands, pt. 14 (1912): 102, fig. 189.
Notommata melandocus Harking, U. S. Natl. Mus. Bull. 81 (1913): 79.
The body of this species is rather slender; its greatest width,
at mid-length, is about one fourth of the total length. The in¬
tegument is soft and flexible, and the outline undergoes consid-
TRANS- W IS- ACAD., VOL. XX.
PLATE LS X.
HARRING and MYERS. — ROTIFERS
COCKAYNE BOSTON
Earring and Myers — The Rotifers of Wisconsin. 645
erable variation with the contractions of the animal. The entire
body is very transparent and indistinctly fluted longitudinal.
The head and neck segments are of about equal length and
width, approximately three fourths of the greatest width of the
body; the transverse folds separating head and neck as well as
neck and abdomen are not very distinct. The abdomen increases
gradually in width for one half its length, and then tapers gently
to the base of the foot, which is as broad as the posterior end
of the abdomen. The tail is very indistinct and visible only in a
lateral view. The foot is very short and broad, without any
joint. The toes are peculiar; the base is a very broad, flattened,
bulbous enlargement, passing abruptly into the slender, slightly
decurved and incurved posterior section. It is possible that the
basal bulb really is a second foot joint, but there is no external
indication of this.
The dorsal and lateral antennae are small setigerous papillae
in the normal positions.
The corona is very slightly oblique and consists of a simple
marginal wreath of cilia, very short dorsally and somewhat longer
laterally, closed on the ventral side immediately behind the mouth.
The buccal field is covered with short, close-set cilia; the apical
plate is unciliated.
The mastax is of a type intermediate between the normal Notom-
mata and the Enteroplea type; while still retaining the pump¬
ing action of the virgate mastax, it has become adapted to
attack upon living prey. The rami appear triangular in ven¬
tral view; at a point near mid-length they are bent at a right
angle toward the dorsal side and provided with two fairly large
teeth in each ramus. There is a small, lozenge-shaped opening
in front of the angle, and an elongate, narrow opening dorsally.
The rami are intricately ribbed and webbed to resist deformation
through muscular action. The fulcrum is straight and tapers
gradually to the posterior end, where it is slightly expanded for
attachment of the muscles of the piston. The unci have only
a single tooth; a very thin and quite narrow lamella attached
to the dorsal side represents the normal additional teeth. To the
tip of the left tooth are attached two minute pre-uncial teeth, and
to the right a single similar tooth. The central portion of the
manubrium is nearly straight and tapers gradually to the poste¬
rior end; the dorsal and ventral extensions are thin lamellae of
moderate size. A slender rod, pointed at both ends and bent
646 Wisconsin Academy of Sciences, Arts , and Letters.
nearly into a semicircle, is imbedded in the walls of the mastax
on each side, immediately under the upper ends of the manubria,
the dorsal ends meeting in the median plane, approximately at
the level of the anterior end of the fulcrum; these rods serve for
the support of the mastax during the pumping operation. Two
ventral salivary glands are present; the left gland is small, while
the right is extremely large and curves under the fulcrum.
The oesophagus is slender and of normal length. The gastric
glands are large and ovate. Stomach and intestine are not dis¬
tinctly separated. The ovary is rather small and irregularly ovate
in outline. A rather small bladder is present. The foot glands are
ovate and unusually large; they terminate in a large mucus re¬
servoir, which is partly in the basal portion of the toe.
The ganglion is moderately large and saccate. The retrocere-
bral organ is limited to a long, fairly large sac, so crowded with
bacteroids as to be completely opaque. No eye-spot is present.
Total length 225-275 /u.; toes 28/a, reduced portion 22/u,; trophi
38/a.
Eosphora melandocus is widely distributed, but usually in
small numbers. Bilfinger states that it lives chiefly on Bdel-
loids, and it is unusual to find a , specimen that has not at least
one pair of the characteristic trophi in its stomach.
This species is readily recognizable by the peculiar toes, as
well as by the opaque sac and the general outline of the body.
Eothinia Harring and Myers, new genus
eothinia elongata (Ehrenberg)
Plate LXI, figures 1-5
Eosphora elongata Ehrenberg, Abh. Akad. Wiss. Berlin (for 1831)
1832: 140; Infusionsth. (1838), p. 452, PI. 56, fig. 9. — Eckstein,
Zeitschr. Wiss. Zool. 39 (1883): 366, PI. 25, fig. 30. — Plate, Jen-
aische Zeitschr. Naturw. 19 (1886): 28.— -Hudson and Gosse, Roti-
fera, Suppl. (1889), p. 27, PI. 33, fig. 8. — Wierzejski, Rozpr. Akad.
Umiej. Wydz. Mat.-Przyr. Krakow, II, 6 (1893): 231, PI. 5, fig. 38.
—Hood, Proc. Royal Irish Acad. Ill, 3 (1895): 683, PI. 21, fig. 3.—
Voigt, Siisswasserfauna Deutschlands, pt. 14 (1912) : 115, fig. 221.
Weber and Montet, Cat. Invert. iSuisse, fasc. 11 (1918) : 122.
Notommata elongata Bartsch, Jahresh. Naturk. Wiirttemberg 26
(1870): 339.
Eosphora striata Glasscott, Proc. Royal Dublin Soc. n. ser. 8 (1893):
57, PI. 4, fig. 5.
HARRiNG and MYERS.— ROT! FERS
COCKAYNE BOSTON
Harring and Myers — The Rotifers of Wisconsin . 647
The body is elongate, fusiform, and slender; its greatest width,
at mid-length, is one fourth of the total length. The integument
is very flexible, but the outline is quite constant. The entire
body is very transparent.
The head and neck segments are of nearly equal length and
width, approximately three fourths of the greatest width of the
body; the transverse folds separating head and neck as well as
neck and abdomen are not very distinct. The abdomen increases
gradually and very slightly in width for one half its length,
and then tapers gently to the tail, which is very broad and in¬
distinctly three-lobed. The foot is fairly long, two- jointed, and
slightly tapering; its length is about one fourth of the total
length. The toes are straight and slender, cylindrical for two
thirds of their length, and conical at the tips; their length is one
tenth of the total length.
The dorsal and lateral antennae are small setigerous papillae
in the normal positions.
The corona is frontal and consists of a marginal wreath of
cilia, much reduced dorsally and passing in a curve to a lateral,
auricle-like area of strongly developed cilia; the ventral arcs
meet immediately below the mouth. The buccal field is faintly
ciliate.
The mastax is a specialized form of the virgate type and very
closely resembles the structure of the mastax in Sphyrias lofuana
(Rousselet).1 The rami are roughly triangular and symmetri¬
cal; at the base they are expanded into broad, somewhat de-
curved lamellar alulae, and near mid-length they are bent rather
abruptly at a nearly right angle toward the dorsal side. The in¬
ner edges are strongly reinforced and enclose an elongate lyrate
opening, which is widest near the middle of the ventral portion;
from this point to the tips the rami are armed with numerous, very
close-set, pointed teeth, increasing in size toward the dorsal ends.
The fulcrum is long and straight; it is formed of two plates
joined at the dorsal edges, so that the cross-section is Y-shaped;
the Y is narrow at the base and spreads gradually, so that the
posterior two thirds of the fulcrum is right-angled in section. The
posterior end is dissected into fine fibrillae, imbedded in the mus¬
cles of the piston. The unci have only a single, strong tooth; at
the base there is a minute lamella with two or three striae, repre-
1 Harring-, Proc. U. S. Natl. Mus. 46 (1913) : 400, PI. 37, figs. 4-8.
648 Wisconsin Academy of Sciences, Arts, and Letters.
senting the rudiments of the normal additional teeth. The manu¬
brium is a straight rod, very slightly larger at the base. Two
straight rods, expanded at their dorsal ends into triangular plates,
pass transversely across the mastax to the dorsal ends of the
rami; they are imbedded in the walls of the mastax and support
them during the pumping action. Two well developed ventral
salivary glands are present; the right gland is considerably larger
than the left.
The oesophagus is unusually long and slender. The gastric
glands are small and rounded. The stomach and intestine are not
distinctly separated. The ovary is large and ovate in outline. A
normal bladder is present. The foot glands are long and slightly
club-shaped.
The ganglion is small and saccate. The retrocerebral organ
consists of a small, clear sac and two subcerebral glands, as large
as the sac. Neither glands nor sac contain bacteroids. The eye-
spot is large and at the posterior end of the ganglion ; two pig¬
ment spots or accessory eye-spots are situated on the corona, im¬
mediately under the integument, near the openings of the ducts
of the retrocerebral organ.
Total length 350-400/a ; toes 35-40 /a ; trophi 50/a.
Eothinia elongata is not very common; we have collected it in
small numbers at Lac Vieux Desert, Vilas County, Three Lakes,
Oneida County, Wisconsin, and at Four-mile Run, in Virginia,
near Washington.
It is necessary to create a new genus for this species, as the
mastax is entirely different from the normal Eosphora- type; the
great dissimilarity in external form between Eothinia elongata
and Sphyrias lofuana prevents its being referred to the latter
genus.
Notes on Collecting and Mounting Rotifers
The methods for collecting the limnetic rotifers living away
from plants in the larger bodies of water are very simple; either
a throwing or a dip net will do. It is necessary to have it made
of material sufficiently thin to allow the water to pass through,
and at the same time fine enough to retain the rotifers; these con¬
ditions are readily fulfilled by several varieties of moderate priced
cotton goods, india linen, nainsook, lawn, etc. A convenient size
for the net is a diameter of about six inches with a length of ten
Harring and Myers — The Rotifers of Wisconsin. 649
inches. The finished net should be sewed to a metal ring with
a socket (text fig. 1) for attachment to a pole or to a jointed fish¬
ing rod of the stout marine type ; this can be taken apart and
packed in a small space. A wide-mouthed bottle or an aluminum
Text Pig. 1. — Collecting net.
centrifuge shield of 50 cubic centimeters’ capacity should be at¬
tached to the lower end of the net; the centrifuge shield is more
satisfactory, as it will not break on coming in contact with stones,
tree-stumps, etc. The concentrated portion of the collection should
be poured through a metal strainer which will allow the rotifers
to pass through but will keep out the larger organisms, plants,
and trash; 40 to 60 meshes per inch is a very suitable screen.
By far the largest number of species of rotifers are to be found
among the leaves of submerged plants; in such places the net
method of collecting is rather inefficient, and far better results
are obtained by simply bringing home the plants. Several bottles
should be about half filled with plants, without crowding, holding
the bottles close to the surface of the water while filling in order
650 Wisconsin Academy of Sciences , Arts, and Letters .
to avoid loss of material ; the bottle should then be filled with clear
water from the same source. Many species will be obtained in
this way which are not to be had with the net. If the plants are
not injured by being packed too tightly, and if the containers are
kept in a subdued light, rotifers may be taken from them for days,
as photosynthesis will be sufficiently active to keep up the oxygen
supply and the absence of strong sunlight will retard the proces¬
ses of decay. Twenty-four hours after weed collections have been
made, about one third of the water should be removed with a
bulb pipette, keeping the opening as near the bottom as possible ;
an equal amount of fresh water should then be added. By doing
this daily the collections may be kept for a week or more.
When making weed collections it is a great convenience to have
a metal hook, which can be screwed into the socket of the pole,
replacing the net. A weighted, three-pronged grapple attached
to a strong cord is desirable for collecting plants out of reach of
the hook. Any kind of wide-mouthed bottle will answer for the
field collections; a screw-top jar of 500 cubic centimeters ’ capacity
is very convenient. If several hours must elapse before the col¬
lections can be examined, thermos bottles are very good, as they
keep the water at a constant temperature, and sudden changes are
fatal to many rotifers.
After reaching home the collections should be allowed to stand
for some time in order to let everything settle. As the oxygen
held in solution gradually decreases the rotifers work their way
towards the surface, and will be found swimming or attached by
the toes to the glass in the meniscus on the side toward the light;
they can then be removed with a pipette. Nearly all the free-
swimming species can be secured from the collections in this man¬
ner. The Rhizota must of course be searched for on the plants
to which they are fixed.
The rotifers are next transferred to watch glasses about 75
millimeters in diameter. A convenient ring for holding these is
made of a narrow circle of brass tubing with three holding clips
soldered to the outside (text fig. 2). These rings keep the glasses
from tilting and hold them securely so that they can be safely
moved about on the stage of the dissecting microscope.
After considerable experimenting, the following method of
narcotization has been found to give the best results. To the
water in the watch glasses containing the rotifers add at once at
least an equal amount of a 5 per cent, solution of cocaine hydro-
Earring and Myers — The Rotifers of Wisconsin . 651
chlorate in 10 to 12 per cent, alcohol ; the rotifers at once contract
strongly, but expand again on recovering from the shock, swim
feebly about, and soon sink to the bottom. The motion of the
cilia should now be watched, and when it has nearly stopped, a
BRASS WATCH-GLASS holder
Text Figure 2
drop of 1 per cent, osmic acid should be dropped on the animals
and the water agitated for a moment, in order to distribute the
acid through it. This kills and fixes at once; the animals should
now be passed through several changes of water in order to re¬
move the osmic acid; if blackened, the material may be bleached
in a weak solution of potassium hydroxide, stopping the action
when the desired clearing has been attained. For nearly all the
free-swimming forms this method gives very good results.
The Rhizota are very sensitive to the cocaine solution ; it should
be added drop by drop to the water containing the animals ; they
will then be narcotized very slowly.
Some Notommatids may be killed with auricles fully extended
by “cooking”, which fixes the protoplasm with very little dis¬
coloration but swells the animals somewhat. The rotifers should
be in a small quantity of water in a watch glass, just enough to
allow them to swim about freely, and while they are in motion a
considerable quantity of boiling water should be added suddenly.
No satisfactory way has yet been found for narcotizing and
killing the Bdelloids in an extended condition; but they have a
tendency to die extended, and this tendency may be taken ad¬
vantage of by allowing them to die slowly in a large amount of
water to which has been added a small quantity of narcotic, not
enough to cause them to contract violently; fixing should be done
as soon after death as possible. By this method fair results may
be obtained with at least some species.
All the loricate rotifers should be killed in 10 per cent, formalin
without previous naerotizing, as the specific characters are more
readily seen in contracted specimens.
Preserved material should be kept in 2 per cent, formalin so¬
lution; formalin is here taken to mean the usual commercial so-
652 Wisconsin Academy of Sciences , Arts , and Letters.
lution, containing nominally 40 per cent, formaldehyde. It is
well to add a small amount of aqueous eosin and to decant after
a few hours ; the eosin stains the rotifers pink and has little ef¬
fect on wood fibres and minute trash, so that the animals are
more readily picked out. Rotifers can be removed from vials
without fear of loss if they are not poured out but taken out
in the following manner: first, be sure that everything has settled
to the bottom if the water has been previously disturbed, then
draw off about two thirds of the water, being careful not to take
up any rotifers; next agitate the remainder with the pipette, but
do not blow air through it, draw all into the pipette and transfer
to a watch glass. The vial is next filled one quarter full of water
and gently agitated; the entire contents is then drawn into the
pipette and added to the collection in the watch glass. From this
the animals may be picked out under a dissecting microscope or
binocular with a low power.
A very convenient form of pipette is made with a brass top,
drilled to fit a readily obtainable size of glass tubing, which is
cemented in and drawn out at the free end to a fine point; a
commercial test-tube cap on the brass top completes the pipette
(text fig. 3). Its principal advantage is that it is easily repaired;
the rubber caps are obtainable in most cities, and a broken point
can be repaired by almost anyone.
•-»))/)>
Text Fig. 3. — Pipette.
Temporary mounting of living or preserved material should be
done on slides with concave, ground centers; the cavity ought
not to be over 10 millimeters in diameter or small specimens will
be hard to find, and if it is over millimeter deep the higher
power objectives will not work through the center of the cell.
The cover glass should never be lowered into place directly over
the cell; place a drop of liquid at the side of the cell and lower
the cover on this drop; then push it over the cell. Covers 18 or
20 millimeters square are most convenient. When the slide is
examined on an inclined instrument, the rotifers gradually sink
to the lower edge of the cell, touching both cover and slide ; they
can then be turned into any desired position by pushing the cover
Earring and Myers — The Rotifers of Wisconsin. 653
about. If the slide is to be kept for a day or more, the edges of
the cover should be sealed with vaseline to prevent evaporation.
For permanent mounts either glycerin or glycerin jelly may
be used; a cement cell should be spun on the slide if glycerin
is used as a mounting median. A very convenient and strong slip
invented by Dr. N. A. Cobb, of the Bureau of Plant Industry, is
made of tin with a round hole punched out of the center; the
edges are folded over to form grooves into which a 25 millimeter
square cover slides freely (text fig. 4). Mounting is done on the
cover glass, which is held in place by two square pieces of card¬
board fitting tightly in the grooves ; if necessary they can be secured
by pinching the corners of the slip.
F/sin fin s//p end & finished s/ide cover-yfess in poi/pon
Text Figure 4
Whether glycerin or glycerin jelly is to be used as the mount¬
ing medium, the rotifers should first be transferred to a 10 per
cent, solution of glycerin in distilled water and set aside in cov¬
ered watch glasses to keep out the dust. In a few days the water
will have evaporated, leaving the animals in glycerin, which has
gradually penetrated the body cavity and displaced the water; the
change will have been so gradual that ordinarily no shrinkage
will take place. If difficult species, such as Notommatids or
Asplanchnids, do shrink and become too much wrinkled during
the process, reconcentration on a drop culture slide becomes neces¬
sary. Place several drops of glycerin in the hollow of the slide
and transfer the shrunken rotifer to it; add just enough water,
drop by drop, to expand the rotifer, and slide a square cover
654 Wisconsin Academy of Sciences , Arts , and Letters.
glass over the cell. Wipe the four corners of the cover with a
cloth moistened in alcohol and secure it with a small drop of al¬
coholic cement at each corner. In a few days an air bubble will
appear in the cell and will increase in size until all the water is
evaporated, which may take several weeks. Concentration will
have been so gradual that the animals will generally be free from
wrinkles. The cement may now be scraped from the corners of
the cover, permitting it to be moved aside, and the rotifers picked
out and mounted.
A pipette should never be used for handling rotifers in gly¬
cerin, as they will invariably stick to the inside and be lost. They
are very easily picked up with a rather flat bristle mounted in
a holder; bristles from an old and much worn tooth brush are
especially good for this purpose, as the ends seem to wear to just
the right shape, a wedge-shaped, slightly blunt point. Removing
a rotifer from glycerin should be done under a dissecting micro¬
scope ; the specimen must be brought to the surface and then lifted
out by getting the bristle under it. If once lifted out, the rotifer
may be left on the bristle with perfect safety until wanted ; slides
and covers may be cleaned, if necessary, without fear of loss of
the specimen. Glycerin used for mounting should be removed
from the bottle by inserting the bristle for its entire length and
quickly withdrawing; the glycerin will then run down the bristle
and form a drop, the size depending on how rapidly it was with¬
drawn.
Rotifers may be kept in glycerin for observation and study by
temporarily mounting them in “stock” slides: cavity slides with
square covers held at the corners with a drop of alcoholic cement.
A collection that is in active use is perhaps more useful if mounted
in this way than in any other. The rotifers may be manipu¬
lated in any way by simple scraping the cement from the corners
and pushing the cover about; if necessary, they can be trans¬
ferred to water and put back again.
All strongly loricated species may be mounted in glycerin
jelly. No cell is required, and they are mounted directly on
plain slides or on the 25-millimeter-square cover glasses of the
Cobb thin slides. Place a very small drop of glycerin in the
center of one of these cover glasses and transfer the rotifers with
the bristle. Warm some jelly until it is of about the same con¬
sistency as the glycerin and place a drop directly on top of the
glycerin drop containing the rotifers. Orient the rotifers and
Harring and Myers — The Rotifers of Wisconsin. 655
cover with an inverted watch glass to keep out dust until the
jelly sets; then trim the jelly with a scalpel as close to the roti¬
fers as possible, leaving them oriented in a small block of the
hardened jelly. Next place a drop of the warmed jelly in the
center of a round cover glass, invert it and place it in position
on the mount. As the glycerine jelly containing the rotifers is
set, this operation will not disturb their position or crush them.
Apply gentle heat for a moment until the jelly spreads to the cir¬
cumference of the cover glass. Lay the mount aside until the
jelly is set; then remove the excess with a camers-hair brush
moistened in water. Do not attempt to wipe the mount with a
cloth before it has been ringed, but use the moistened brush and
let the moisture surrounding the cover glass evaporate; then re¬
wipe with a brush moistened with alcohol until all traces of jelly
are removed. After the first ring of cement hardens, the mount
may be gently wiped with a cloth before applying the succeeding
rings.
If the Cobb slides are to be used for glycerin mounts, the roti¬
fers are mounted on a 25-millimeter-square cover glass on which
a cement ring of the necessary thickness is spun and allowed to
dry thoroughly. The cover glass is now fastened to a larger and
thicker piece of glass by pressing it down hard on a small drop
of water and wiping thoroughly; this will hold it in place and
also prevent breakage during the mounting operations; the cover
is too small and fragile to be easily handled and moved about on
the stage of the dissecting stand. A small drop of glycerin is
placed in the center of the cement cell, and the rotifers are trans¬
ferred to it with the aid of the mounted bristle. Another small
drop is next deposited directly on the cement ring of the cell, and
a round cover glass a little smaller than the outer edge of the
cement ring is lowered on this drop and pushed over the roti¬
fers. If there is not enough glycerin to fill the cell, more should
be added as minute drops on the end of the bristle as the cover
is pushed along; care should be taken not to use too much, but
just sufficient to fill the cell. After the cover glass is in place,
the excess glycerin, if any, should be removed with a small piece
of filter paper laid flat over the mount and pressed down gently
around the edges. Unless a considerable excess of glycerin has
been used, this gentle pressure will not disturb the position of
the cell contents to any great extent. The edge of the cover glass
is now carefully wiped with a moistened cloth, holding down one
656 Wisconsin Academy of Sciences , Arts, and Letters.
side while the other is being wiped. The cover glass and slide
are next carefully cleaned with a camel’s-hair brush moistened
in alcohol to remove all traces of glycerin. The mount may now
be finally sealed in the usual manner; about six coats of cement
should be used, and each one should be allowed to dry thoroughly
before applying the next. The first coat should always be an
alcoholic cement, as this will absorb any small quantity of glycerin
present and the covers will stick much better.
If the transfer to glycerin can not be accomplished and the
animals persist in shrinking, they should be mounted in 2 per
cent, formalin in hollow ground slides. It is extremely doubt¬
ful whether really permanent mounts in a watery medium can
be made ; no cement is known that is absolutely impermeable to
water. A slide made according to the following method, originated
by Rousselet, will last 5 or 10 years, seldom longer. A ring of
benzol lammar, about 1 part dammar in 15 parts benzol, should
be spun on the slide, about 1 millimeter outsile the edge of the
cavity; this ring must be very thin, or the rotifers will get be¬
tween the cover and the flat top of the slide, and as evaporation
proceeds they will become crushed. The first ring of cemeuf
should be made with Windsor and Newton’s “Picture Copal
Varnish”, and this should be followed by 5 or 6 coats of gold size.
If a small air bubble is included in the cell at the time of mount¬
ing, the slide will probably last much longer, as this allows the
liquid to expand and contract without straining the cement ring.
All slides mounted with watery media or glycerin should be
stored lying flat, or the specimens will gradually work down to
the edge of the cell and either be caught in the cement, from
which they are removed only with great difficulty, or get under
the cement ring on top of the cover and thus become invisible.
PLATE LXI.
TRANS. W!S. ACAD., VOL. XX.
HARPING and MYERS.— ROTIFERS
COCKAYNE BOSTON
j Earring and Myers — The Rotifers of Wisconsin. 657
Explanation of Plates
All the figures are highly magnified. For actual measurements see text.
The epipharynx has been tinted dark.
PLATE XLI
Fig. 1. Notommata copeus, dorsal view; page 562.
Fig. 2. Notommata copeus, lateral view.
Fig. 3. Notommata copeus, trophi, ventral view.
Fig. 4. Notommata copeus, trophi, lateral view.
PLATE XLI i
Fiq. 1. Notommata pachyura, dorsal view; page 565.
Fig. 2. Notommata pachyura, lateral view.
Fig. 3. Notommata placida, dorsal view; page 587.
Fig. 4. Notommata placida, trophi, lateral view.
Fig. 5. Notommata placida, trophi, ventral view.
Fig. 6. Notommata placida, trophi, lateral view.
plate xliii
Fig. 1. Notommata pachyura, form triangulata, dorsal view; page 565.
Fig. 2. Notommata pachyura, form triangulata, lateral view.
Fig. 3. Notommata pachyura, form triangulata, trophi, ventral view.
Fig. 4. Notommata pachyura, form triangulata, trophi, lateral view.
Fig. 5. Notommata pachyura, form triangulata . unci, frontal view.
plate XLIV
Fig. 1. Notommata collaris, dorsal view; page 568.
Fig. 2. Notommata silpha, lateral view.
Fig. 3. Notommata collaris, trophi, ventral view.
Fig. 4. Notommata collaris, trophi, lateral view.
Fig. 5. Notommata collaris, unci, frontal view.
Fig. 1.
Fig. 2.
Fig. 3.
Fig. 4.
Fig. 5.
Fig. 6.
Fig. 7.
Fig. 8.
Fig. 9.
Fig. 10.
Fig. 11.
Fig. 12.
PLATE XLV
Notommata silpha, dorsal view; page 596.
Notommata silpha, lateral view.
Notommata silpha, trophi, ventral view.
Notommata silpha, trophi, lateral view.
Notommata silpha, incus, lateral view.
Notommata silpha, incus, dorsal view.
Notommata silpha , unci, frontal view.
Notommata falcinella, dorsal view; page 570.
Notommata falcinella, lateral view.
Notommata falcinella, trophi, ventral view.
Notommata falcinella, trophi, lateral view.
Notommata falcinella, unci, frontal view.
42— S. A. L.
658
Wisconsin Academy of Sciences , Arts, and Letters.
Fig. 1.
Fig. 2.
Fig. 3.
Fig. 4.
Fig. 5.
Fig. 6.
Fig. 7.
Fig. 8.
Fig. 1.
Fig. 2.
Fig. 3.
Fig. 4.
Fig. 5.
Fig. 6.
Fig. 7.
Fig. 8.
Fig. 9.
Fiq. 1.
Fig. 2.
Fig. 3.
Fig. 4.
Fig. 5.
Fig. 6.
Fig. 7.
Fig. 8.
Fig. 9.
Fig. 1.
Fig. 2.
Fig. 3.
Fig. 4.
Fig. *5.
Fig. 6.
Fig. 7.
Fig. 8.
Fig. 9.
Fig. 10.
Fig. 11.
Fig. 12.
Fig. 13.
plate XLVI
Notommata galena, dorsal view; page 574.
Notommata galena, lateral view.
Notommata galena, trophi, ventral view.
Notommata galena, trophi, lateral view.
Notommata cerberus, dorsal view; page 572.
Notommata ceberus, lateral view.
Notommata cerberus, trophi, ventral view.
Notommata cerberus, trophi, lateral view.
plate xlvii
Notommata saccigera, dorsal view; page 594.
Notommata saccigera, lateral view.
Notommata saccigera, trophi, ventral view.
Notommata saccigera, trophi, lateral view.
Notommata saccigera, unci, frontal view.
Notommata pseudo cerberus, dorsal view; page 598.
Notommata pseudo cerberus, lateral view.
Notommata pseudocerberus, trophi, ventral view.
Notommata pseudocerberus, trophi, lateral view.
PLATE XL VIII
Notommata peridia, dorsal view; page 576.
Notommata peridia, ventral view.
Notommata peridia, trophi, ventral view.
Notommata peridia, trophi, lateral view.
Notommata peridia, unci, frontal view.
Notommata aurita, dorsal view; page 578.
Notommata aurita, lateral view.
Notommata aurita, trophi, ventral view.
Notommata aurita, trophi, lateral view.
PLATE XLIX
Notommata telmata, dorsal view; page 584.
Notommata telmata, lateral view.
Notommata telmata, trophi, ventral view.
Notommata telmata, trophi, lateral view.
Notommata cyrtopus, dorsal view; page 582.
Notommata cyrtopus, lateral view.
Notommata cyrtopus, trophi, ventral view.
Notommata cyrtopus, trophi, lateral view.
Notommata tripus, dorsal view; page 589.
Notommata tripus , lateral view.
Notommata tripus, trophi, ventral view.
Notommata tripus, trophi, lateral view.
Notommata tripus, unci, frontal view.
Barring and Myers — The Rotifers of Wisconsin. 659
Fig. 1.
Fig. 2.
Fig. 3.
Fig. 4.
Fig. 5.
Fig. 6.
Fig. 7.
Fig. 8.
Fig. 9.
Fig. 10.
Fig. 11.
Fig. 12.
Fig. 13.
Fiq. 1.
Fig. 2.
Fig, 3.
Fig. 4.
Fig. 5.
Fig. 6.
Fig. 7.
Fig. 8.
Fig. 9.
Fig. 10.
Fig. 11.
Fig. 12.
Fig. 1.
Fig. 2.
Fig. 3.
Fig. 4.
Fig. 5.
Fig. 6.
Fig. 7.
Fig. 8.
Fig. 9.
Fig. 10.
Fig. 11.
Fig. 12.
PLATE L
Notommata venusta, dorsal view; page 591.
Notommata venusta, lateral view.
Notommata venusta , trophi, ventral view.
Notommata venusta, trophi, lateral view.
Notommata trypeta, dorsal view; page 602.
Notommata trypeta, lateral view.
Notommata trypeta, trophi, ventral view.
Notommata trypeta, trophi, lateral view.
Notommata lenis, dorsal view; page 586.
Notommata lenis, Zateral view.
Notommata lenis, trophi, ventral view.
Notommata lenis, trophi, lateral view.
Notommata lenis, unci, frontal view.
PLATE LI
Proales parasita, dorsal view; page 607.
Proales parasita, lateral view.
Proales parasita, trophi, ventral view.
Proales parasita, trophi, lateral view.
Proales decipiens, dorsal view; page 603.
Proales decipiens, lateral view.
Proales decipiens, trophi, ventral view.
Proales decipiens , trophi, lateral view.
Proales sordida, dorsal view; page 605.
Proales sordida, lateral view.
Proales sordida, trophi, ventral view.
Proales sordida, trophi, lateral view.
PLATE LII
Proalinopsis caudatus, dorsal view; page 608.
Proalinopsis caudatus, lateral view.
Proalinopsis caudatus, trophi, ventral view.
Proalinopsis caudatus, trophi, lateral view.
Proalinopsis caudatus, trophi, frontal view.
Drilophaga judayi, lateral view; page 612.
Drilophaga judayi, trophi, ventral view.
Drilophaga judayi . trophi, lateral view.
Notommata angusta, dorsal view; page 580.
Notommata angusta, lateral view.
Notommata angusta, trophi, ventral view.
Notommata angusta, trophi, lateral view.
660 Wisconsin Academy of Sciences , Arts, and Letters.
PLATE LI II
Fig. 1. Lindia pallida, dorsal view; page 620.
Fig. 2. Lindia pallida, lateral view.
Fig. 3. Lindia pallida, trophi, ventral view.
Fig. 4. Lindia pallida, trophi, lateral view.
Fig. 5. Lindia torulosa, dorsal view; page 618.
Fig. 6. Lindia torulosa, trophi, ventral view.
Fig. 7. Lindia torulosa, trophi, lateral view.
Fig. 8. Lindia torulosa, unci, frontal view.
Fig. 9. Lindia producta, dorsal view; page 616.
Fig. 10. Lindia producta, lateral view.
Fig. 11. Lindia producta, trophi, ventral view.
Fig. 12. Lindia producta, incus, ventral view.
Fig. 13. Lindia producta, trophi, lateral view.
Fig. 14. Lindia producta, unci, frontal view.
PLATE LIV
Fig. 1. Lindia truncata, dorsal view; page 626.
Fig. 2. Lindia truncata, lateral view.
Fig. 3. Lindia truncata, trophi, ventral view.
Fig. 4. Lindia truncata, trophi, lateral view.
Fig. 5. Lindia truncata, unci, frontal view.
Fig. 6. Lindia annecta, dorsal view; page 622.
Fig. 7. Lindia annecta, lateral view.
Fig. 8. Lindia annecta, trophi, ventral view.
Fig. 9. Lindia annecta, trophi, lateral view.
Fig. 10. Lindia Candida, dorsal view; page 614.
Fig. 11. Lindia Candida, lateral view.
Fig. 12. Lindia Candida, trophi, ventral view.
Fig. 13. Lindia Candida, trophi, lateral view.
Fig. 14. Lindia Candida, unci, frontal view.
PLATE LV
Fig. 1. Lindia tecusa, dorsal view; page 624.
Fig. 2. Lindia tecusa, lateral view.
Fig. ,3. Lindia tecusa, trophi, ventral view.
Fig. 4. Lindia tecusa, trophi, lateral view.
PLATE LVI
Fig. 1. Lindia fulva, dorsal view; page 628.
Fig. 2. Lindia fulva, lateral view.
Fig. 3. Lindia fulva, trophi, ventral view.
Fig. 4. Lindia fulva, trophi, lateral view.
Fig. 5. Notommata pygmaea, dorsal view; page 593.
Fig. 6. Notommata pygmaea, lateral view.
Fig. 7. Notommata pygmaea, trophi, ventral view.
Fig. 8. Notommata pygmaea, trophi, lateral view.
Earring and Myers — The Rotifers of Wisconsin.
Fig. 1.
Fig. 2.
Fig. 3.
Fig. 4.
Fig. 1.
Fig. 2.
Fig. 3.
Fig. 4.
Fig. 5.
Fig. 6.
Fig. 7.
Fig. 8.
Fig. 9.
Fig. 10.
Fig. 11.
Fig. 12.
Fig. 13.
Fig. 1.
Fig. 2.
Fig. 3.
Fig. 4.
Fig. 5.
Fig. -6.
Fig. 7.
Fig. 8.
Fig. 9.
Fig. 10.
Fig. 1.
Fig. 2.
Fig. 3.
Fig. 4.
Fig. 5.
Fig. 6.
Fig. 7.
Fig. 8.
Fig. 9.
Fig. 10.
Fig. 11.
PLATE LVII
Tetrasiphon hydrocora, dorsal view; page 630.
Tetrasiphon hydrocora , lateral view.
Tetrasiphon hydrocora , trophi, ventral view.
Tetrasiphon hydrocora , trophi, lateral view.
PLATE LVIII
Birgea enantia, dorsal view; page 610.
Birgea enantia , lateral view.
Birgea enantia, trophi, oblique ventral view.
Birgea enantia, trophi, lateral view.
Notommata contorta, dorsal view; page 600.
Notommata contorta, lateral view.
Notommata contorta, trophi, ventral view.
Notommata contorta, trophi, lateral view.
Eosphora anthadis, dorsal view; page 641.
Eosphora anthadis, lateral view.
Eosphora anthadis, trophi, ventral view.
Eosphora anthadis, trophi, lateral view.
Eosphora anthadis, trophi, frontal view.
PLATE LIX
Eosphora therina, dorsal view; page 639.
Eosphora theirina, lateral view.
Eosphora therina, trophi, ventral view.
Eosphora therina, trophi, lateral view.
Eosphora therina, rami, oblique frontal view.
Eosphora melandocus, dorsal view; page 644.
Eosphora melandocus, lateral view.
Eosphora melandocus, trophi, ventral view.
Eosphora melandocus, trophi, lateral view.
Eosphora melandocus, trophi, oblique ventral view.
PLATE LX
Eosphora gelida, dorsal view; page 642.
Eosphora gelida, lateral view.
Eosphora gelida, trophi, ventral view.
Eosphora gelida, trophi, lateral view.
Eosphora gelida, trophi, oblique ventral view.
Eosphora gelida, pattern of integument.
Eosphora najas, dorsal view; page 634.
Eosphora najas, Zateral view.
Eosphora najas, trophi, ventral view.
Eosphora najas, lateral view.
Eosphora najas, trophi, oblique ventral view.
662
Wisconsin Academy of Sciences, Arts, and Letters.
Fig. 1
Fig. 2
Fig. 3
Fig. 4
Fig. 5
Fig. 6
Fig. 7
Fig. 8
Fig. 9
PLATE LXI
Eothinia elongata , dorsal view; page 646.
Eothinia elongata, lateral view.
Eothinia elonggta, trophi, ventral view.
Eothinia elongata, trophi, lateral view.
Eothinia elongata, rami, oblique ventral view.
Eosphora ehrenbergi, dorsal view; page 637.
Eosphora ehrenbergi, lateral view.
Eosphora ehrenbergi, trophi, ventral view.
Eosphora ehrenbergi, trophi, lateral view.
THURE KUMLIEN
Publius V. Lawson
To Mrs. Angie Kumlien Main of Fort Atkinson, Wisconsin,
granddaughter of the naturalist, Thure Kumlien, who has in¬
herited her distinguished ancestor’s love of birds and all nature,
is due very much of the success of this study; since, but for her
willing and ceaseless energy and industry in seeking out and re¬
covering the muniment material of his life, his paintings, draw¬
ings, and writings, this life story would still remain untold. Other
appreciation and credit has been given throughout this paper ; and
to these good people who have so kindly aided in the several in¬
vestigations our thanks are due.
When William D. Hoard resided at Lake Mills about 1870, and
long before he became governor of the state, he one day read in
the Chicago Tribune
“a brief note to the effect that Prof. Agassiz of Boston had said that
Thure Kumlien of Bussey ville, Wisconsin, was the greatest authority in
the world on bird nests”.
He started out at once to make the acquaintance of a man who
was so well known in scientific circles and unknown to him, who
lived in the same county. He found the lover of nature.
“occupying an eighty-acre farm with a log house, and engaged in plow¬
ing with a pair of oxen. He preferred oxen to horses, for if a rare bird
came within his vision, he could leave them in the furrow and hunt the
bird. I found he was a graduate of the University of Upsala in Sweden
and a correspondent of most of the scientific societies of Europe and
America. . . . His life work as a naturalist brought fame to his adopted
state. His son Ludwig Kumlien was also a famous naturalist (*) .”
Several plants have been named for him. He was an expert
taxidermist. There was no one in America so well informed on
663
664 Wisconsin Academy of Sciences , Arts, and Letters.
our native birds. Every bug, fly, moth, butterfly, mole, rat, or
game animal within miles around his favorite Lake Koshkonong,
with its habits and life story, was perfectly known to him. The
life story and habits of every freshwater fish of the lake and of
Rock River he thoroughly understood. He was master of every
phase of the zoology and botany of this state. As an artist he
had few equals. He drew plants and animals in water colors.
Few men were adept in more languages. One writer has said he
knew almost every language in Europe. He was master of Latin
and was well versed in Greek. He could speak and write classic
English before he came to this country. His predominant quali¬
ties were strict honesty, purity of conduct, and exactness in scien¬
tific research. He was the highest authority on oology in this
country. No one has furnished so many learned institutions and
well known museums with so many or such fine mounting and
exact correct designation as our modest, lovable old blonde natur¬
alist of Jefferson County.
Governor Hoard in the notice of the death of Thure Kumlien
writes of him in his paper (2) that he was an “eminent natur¬
alist” and that he was
“known as one of the most prominent scientists of the world. His con¬
tributions to science were many and of great importance”.
Mr. Wheeler, in writing of the sudden death of Thure Kumlien
while conservator at the public museum at Milwaukee, speaks of
him
“as one who was present at the institution’s small beginnings and who
has diligently assisted in raising it to its present flourishing condition” (2) .
The late Prof. Edward Lee Greene wrote (4) that
“A purer, nobler type of the naturalist of the reserved and quiet non-
advertising class,, there probably was not in his day, in America, than
Thure Ludwig Theodor Kumlien”.
He was contemporary, friend, and correspondent of Dr. P. R.
Hoy of Racine and of those other pioneer scientists who in 1870
came together to found the Wisconsin Acadamy of Sciences, Arts,
and Letters. In 1875 he read a paper before this society on the
rapid disappearance of Wisconsin wild flowers (5). Professor
Greene says:
“It was evident not only from the friendly correspondence -which was
always kept up between them, but also from many a pleasing anecdote
which we were wont to hear of life and study and travel in intimate com-
Lawson — Thure Kumlien.
665
panionship with his revered master, that Mr. Kumlien has been, while at
Uusala, a very special favorite among the botanical pupils of Professor
Elias Fries (6). How thoroughly worthy the youth must have been, of the
particular attention of the great Swedish botanist of the nineteenth cen¬
tury, was still manifest in Mr. Kumlien when I first made his acquaint¬
ance, some sixteen or eighteen years after his arrival in this country.
He was then a sort of a second and American edition of jFries, in his
almost equal familiarity with each of the following departments of bot¬
anical study; phanerogams, ferns and their allies, mosses, lichens, and
fungi. He had, in 1860, and I know not how long before, so well mastered
the extensive and varied flora of southern Wisconsin that there was no
indigenous tree or shrub, flower, grass, or sedge, or moss, or hepatic,
lichen, or mushroom, the scientific name of which was not at his tongue’s
end for you at any moment. I am confident that, notwithstanding our
considerable list of worthy names in American botany, no state in our
union has ever had so complete a master of its whole flora as Wisconsin
had in this extraordinary man, whom our eastern botanists seldom heard
anything of; whom, with his low stature, muscular frame, rather stoop¬
ing shoulders, light hair and keen blue eyes, a stranger might have mis¬
taken as he passed along the country roads for an ordinary farmer from
the Scandinavian settlement; who, in the most polished society would
have been recognized as an intelligent, refined and almost courtly gentle¬
man; in whom any scholar would have found a finished collegian of
the old Swedish school; whose pen could indite Ciceronian Katin and
whose tongue could address a foreigner in, I believe, any one of the
languages of Europe spoken between Spain and (Sweden. But that which
makes his thorough familiarity with so many branches of botany seem
more remarkable, more unmistakably indicative of uncommon natural
gifts, is the fact that, even from boyhood, his specialty appears to have
been ornithology. It was to the birds, yet not so as to exclude other
branches of zoological study, with which he was also vary familiar, that
he gave most of his time. On his vacation tours in college days, he had
penetrated to some of the remoter parts of the Scandinavian peninsula,
and had visited the islands of the Baltic; and, although he gave us charm¬
ing word pictures of the flora of those more secluded places, it was
plain that what had pleased him most had been the new gains thus made
in the knowledge of his particular favorites, the birds. Even the fame,
which he could not seek, but which was thrust upon him at last, in no
small measure, was that of an ornithologist. It was with reference to its
probable facilities for ornithological work, that, under the guidance of a
map only, and from afar, he made choice of the locality in Wisconsin
where he would build his cottage and consecrate his home”(4).
Although his full baptismal name was Thure Ludwig Theodor
Kumlien, he always signed himself with the first name, Thure.
He was born at Hertorp in Harlunda parish, Skara stift (district),
Skaraborg lan (county), Vest ergo tland, Sweden, on November 9,
666 Wisconsin Academy of Sciences, Arts, and Letters.
1819 (6). His father and grandfather bore the name Ludwig
Kumlien (8). His father owned and operated several large
estates. Thure Kumlien, in a letter to the Hofintendant (Royal
Minister), says:
“My father, Ludwig Kumlien, was born in Uppland, Sweden, March 4,
1790; he died, July 7, 1839. He married Johanna Petronelle Rhodin,
daughter of a minister in Broddetorp, born June 25, 1800, died April 28,
1838. My father passed the royal examination in Upsala. Served after
that for some time in Sweden’s Royal Court. After that he was ap¬
pointed regimental secretary to Skaraborg’s regiment, with the title,
honor, and worthiness of assessor” (7) .
“Thure, the oldest of fourteen children, was early entrusted to a pri¬
vate tutor, and soon entered the gymnasium at Skara and subsequently
graduated from the University of Upsala in 1843” (3).
“His undaunted desire for zoology and botany manifested itself so early
that already as a school boy he had a whole room packed full from floor
to ceiling with birds that he had shot and mounted so skillfully and true
to life that an old accustomed conservator could not have done it better,
wherefore, also at the time of his leaving the Skara institution of learn¬
ing, all this was received with gratefulness as a gift from him to the
zoological collection of the school” (7).
Upsala had no professor of ornithology, and young Kumlien in
his study of wild life was obliged to encompass it almost alone (9).
The aid and encouragement of his father enabled him to acquire a
considerable amount of knowledge of art and natural history aside
from his studies ( 10 ) . His parents had designed that young Kum¬
lien should follow medicine, and proposed after his graduation at
Upsala that he should attend the best medical schools.
“While a student at Upsala, he was the bosom friend of the elder Fries.
Among his classmates were men who have since become some of Sweden’s
most eminent statesmen, poets, musicians, scientists, and literati. With
many of these he corresponded for many years” (3).
In a letter to President Twombly of the University of Wiscon¬
sin, Thure Kumlien writes of himself:
“Having gone through the regular course of studies at the schools and
gymnasium, I studied four years at Upsala University. In 1842 I made a
collecting tour of some of the Islands of the Baltic [iSea] and found many
rare specimens both of plants and birds. Among the latter was a gull
that had not been found in Sweden since Linnaeus found it.”
There lived near Upsala the most beautiful girl of all this na¬
tion of handsome women. She was Margretta Christina Wall-
Lawson — Thure Kumlien.
667
berg, born in the Parish of Tibia in Uppland, Sweden, April 21,
1820. Her father was an officer in the Swedish army and in
charge of the training of horses for the cavalry in Sweden (11).
His family consisted of eleven children. Subsequently he lived
to exceed the age of eighty years. Christina had gone through the
only school open to girls. This was similar to our domestic
science schools.
Young Kumlien, while a student at Upsala, had met this
splendid girl, who was of about his own age ; and a mutual at¬
tachment led to a betrothal. Her people were agreeable to the
union, but his people objected; and so, as they had other plans
for him, no marriage was possible under the laws of church and
state in Sweden. Both were communicants in the Lutheran faith
and church. Thure Kumlien ’s mother died when he was twenty
years old, and his father died one year later.
In this dilemma it was determined that they would sail for
America. Her family sent Sophia Wallberg, her sister, with her
as companion.
“With the girl he loved and her sister as her companion, he took pas¬
sage in an old sailing vessel for the United States. The old vessel, which
he afterwards learned had been condemned, was ten weeks at sea. While
they were becalmed at mid ocean for several weeks, their drinking water
gave out, and the passengers suffered of thirst. When the storm came
they were nearly shipwrecked” (10).
On their arrival in America, August 20, 1843 (4), they pro¬
ceeded at once to Milwaukee, where they were married September
5, 1843, pursuant to a license, before William A. Prentiss, Justice
of the Peace (12). They arrived at Milwaukee by boat and
walked seventy miles to Lake Koshkonong, where Kumlien selected
and bought a government forty acres, afterward adding another
forty acres of virgin forest in the town of Sumner, Jefferson
County, on the shore of Lake Koshkonong. When his log house
was finished,
“it was the nicest log house around here, for we had an extra bedroom
besides our one big room. Under the stairs we had a pantry, which was
more than most of the pioneers had.”
The newly married pair were very happy in their new home.
They lived in the log house until 1874, when a frame house was
erected near the site. Thure Kumlien brought with him many
beautiful water-color paintings of the flowers and birds of Swe-
668 Wisconsin Academy of Sciences , Arts , and Letters.
den, which he had painted while at Upsala. These were used to
decorate the walls of their new home. Aunt Sophia lived and
died there. After the death of his wife Christina, September 22,
1874, from cancer, Sophia took charge of the family, and after a
long, noble, and useful life she died at ninety years of age and
sleeps in the family cemetery. She was beloved by all the coun¬
tryside. The old log house was removed in 1908.
The location of the house on the shore of the beautiful lake
proved a paradise for the young naturalist.
“The region abounded in game of all kinds common to this section.
Even Buffalo horns were found by the early settlers. The lake was a
favorite resort for water fowl, and these, with the fish and the animals,
were new to him. The grand old forests, as yet untouched by the settlers,
were swarming with birds. A great treat was in store for him in the
study of the flowers which at that time had only been disturbed by the
grazing of the wild deer. The entomology of this country was an end¬
less source of pleasure to him”(10).
Gustaf Unonius, a Swedish Episcopal clergyman, who came to
this country in 1841, and who visited Koshkonong in 1845, says
in his “Minnen”, in reference to Thure Kumlien:
“It was quite remarkable to see how he divided his time between farm¬
ing and scientific researches. Necessity bound [his] hands to the plough
and to the hoe, while interest and desire held [his] thoughts on flowers,
birds, and insects. A rich herbarium and an ornithological cabinet, of no
mean importance, but on account of its small capacity, not arranged in
order, possibly also bears evidence to a greater field of work as a natural¬
ist than as an agriculturist. It could very well happen that the oxen
would be permitted to stand in the furrow a little longer than was nec¬
essary for their rest, in case the ploughman’s eye accidentally caught a
glimpse of some rare insect or of some flower of the field that had not as
yet been analyzed. He united with a real scientific education also an ex¬
cellent ability for mounting birds and other animals, and worked in this
manner in order to bring himself a small income” (13).
On August 28, 1843, eight days after Thure Kumlien landed on
our shores, and as soon as he arrived in Milwaukee, he sought
out the clerk of the circuit court for the county of Milwaukee
and swore out his first papers in the process of naturalization.
Five years later, on October 23, 1849, the circuit court of Jeffer¬
son county issued to him his second papers, declaring him a full
citizen of the United States (14).
In his earliest days in his new home, Kumlien was obliged to
meet and overcome difficulties that for him were very formidable.
In a letter to Dr. Brewer he writes of this :
Lawson — Thure Kumlien.
669
“I am poor, Sir, I have to work hard to support my family and I see
money but seldom. I was not brought up to work with make [which
makes it] come harder for me; still I can live well here being content
with little. I have bought another 40 acres of land and when I get that
paid for (nearly 200$) and some more improved I calculate to let out
on shares and I hope I would be able to live, with the addition of some
work, on half what the field will yield and then I will have time for birds
& flowers of which two things I have been passionately fond ever since
a child and ‘quo semel est imbuta recens servabit dorem test diu.’ ”
In another letter he writes Dr. Brewer an apology for not
finding as many eggs to send him as was expected. He adds :
“Having been obliged to work hard for a living, and not being used to
hard labor, as I have passed my days at school and college, I have had
but little time for hunting and explorations”.
To come to America he had borrowed money of Baron Carl
Grustaf Lowenhjelm, a nobleman, who was splendid in patiently
awaiting the return of the loan, which he finally received entire.
In one of Kumlien ’s letters to the Baron he mentions something
of the serious difficulties for him in this very new country. The
land he had settled on was covered with a dense forest of hard¬
wood, and one can see the hardships of a pioneer who did not un¬
derstand how to swing an axe. He writes:
“When I came here I did not understand anything about farming, or
how to handle an axe, or a plow, and in such a country as this”.
In a letter to Dr. Brewer he writes:
“I am glad to get fifty cents apiece for yellow-headed blackbird skins,
and I wish I could sell many for that price. It is easier for me to kill and
skin a bird than it is to go out and work hard for fifty cents a day for
a farmer”.
To these pioneer struggles of the naturalist the writer of a
paper in The Auk thus refers:
“He was a zealous collector and acute observer, a man of high intel¬
lectual culture, and most amiable and unassuming in character. His youth¬
ful love for scientific pursuits persisted through life, but in consequence
of his untoward surroundings and isolation from large museums and li¬
braries, his investigations were necessarily limited to the products of the
woods and prairies of his immediate vicinity. His early pioneer life was
thus unfavorable to the spirit of research and he has consequently left no
published works or papers of any great importance. His influence, how¬
ever, upon the rising generation of naturalists with whom he came in con¬
tact was most efficient and encouraging. Ornithology and botany were
670 Wisconsin Academy of Sciences, Arts, and Letters.
his favorite fields of study, and he is said to have early made himself
familiar with all of the species of birds and plants found about his wil¬
derness home”(5).
Lake Koshkonong,
“when reached, must fully have answered every expectation of the
young naturalist. The lake, some eight or nine miles long and four in
breadth, as I remember it, is but an expansion of iRock River, its sinuous
shore line touching the bases of a hundred low hills covered with oaks or
overrun with hazel, with many a fair interval of open grassy slope, or
widespread lowland meadows. The larger estuaries, sheltered by neigh¬
boring groves, their still and shallow waters bordered with green fields of
reed and wild rice, were twice in each year the resort of great flocks of
wild geese, pelicans, and swans, and indeed of all the tribes of water fowl
and wading birds, not excepting many that are usually maritime only.
And the wooded hills and open meadow lands were equally the home of
the whole concourse of spring and summer song-birds, of grouse, and
pheasant, and other larger wild fowl. While the region remained almost
unsettled, and while wild birds so abounded, an ornithologist might have
been pardoned had he forgotten more or less of his botany. Rut this one
did not. So ardent a lover as he was of all things beautiful in nature,
could not but have been enraptured with the floral splendors of wild wood¬
lands and unbroken prairie as they must have appeared to his eye in
that early day. Even as late as 1858, when I first saw that land, after
multiplied settlements had sprung up everywhere, and the prairies had
been converted into fields of waving grain, and the open woods turned
over to the destructive teeth and hoofs of the domestic flocks and herds,
there still remained in many a protected spot charming traces of the
primeval floral wealth, in pink and azure banks of phlox and Pole-
monium, violets, Dentarias and Dielytras, lupines, wild peas, and vetches;
extensive yellow beds of Caltha and Ranuculus; meadow patches of scarlet
and yellow Castilleias; fence corners filled with grassy-leaved Hypoxis,
Tradescantia, Camassia, and Zygadenus; hazel borders all undergrown
with Erythroniums, Trilliums, Orchis, and nodding wood anemone; thick¬
ets of wild rose and shad bush, wild plums and cherries; groves of white-
barked aspen and fragrant, rosy-blooming crabapple.
“The building site which Mr. Kumlien chose at first, and whereon he
dwelt to the end of his life, was, for the work and the pleasure of a poet-
naturalist — and such was he — admirably selected; lying back from Lake
Koshkonong, to the northward, upon a pleasant elevation, forth from
which one looked down across a mile or more of moist meadow, to the
shores of the lake. A considerable extent of oak woods enclosed the
place northward and westward; to the eastward lay a stretch of open,
undulating arable land, suitable for farming purposes. The pristine quiet
and seclusion of the place was always retained; for when other settlers
had taken possession of all the country round about, and regular public
roads had been laid out, the naturalist’s home was left about equally
distant from every public highway; so that one reached the place by
Lawson — Thure Kumlien.
671
either of two by-roads, closed by gates, and leading circuitously about
among the woods.
“Lake Koshkonong proved not to be the only naturalists’ paradise in
that immediate region. Some two miles to the westward lay, deep down
among the wooded hills, a chain of three lesser lakes. Among the bluffs
above these lakes there flowered in earliest spring, almost as soon as the
ice had melted, such rarities as Anemone patens, Ranunculus rhomboidus,
Braba Caroliniana and Arabis lyrata, plants not then to have been found
in Wisconsin except on the bleak summits of such hills; and they are
probably all now extinct even there. In the reedy margins of the smaller
lakes there grew in summer time such interesting aquatics as Pontederia
cordata and Brasenia, and also every kind of water lily indigenous to
the northern states; Castalia tuberosa and Nelumbo lutea, both in rich
abundance” (4).
“Koshkonong Lake proved to be a paradise for this enthusiastic young
naturalist, abounding as it did in rare species of water birds, such as
swans, pelicans, wild geese, snipe, gulls, terns, a great many of which it
would be impossible to procure at this time, and ducks were no rarity.
Canvas backs were plentiful, also the wild pigeon, which now lives only
in the memory of the older inhabitants. In the early forties they did not
have the prairie chicken in this section; the pintail grouse was common,
but in a few years it disappeared, the prairie chicken taking its place;
a great number of our plants and birds have now disappeared from this
section” (10).
In 1867 lie was appointed professor of botany and zoology in
Albion Academy, a collegiate institution located at Albion, a few
miles from his home, just over the line in Dane County. He also
taught languages in the academy. He filled this position with
great credit to himself and to the school, and with much benefit
to the students under his charge, until 1870, when mismanage¬
ment and want of funds practically interfered with the school’s
success, and he severed his connection with the institution.
Among the distinguished students of this academy may be
named the late Edward Lee Greene, professor of botany at the
University of California and at Notre Dame University, and Lud¬
wig Kumlien, son of Thure Kumlien, professor of physics and
natural history in Milton College at the time of his death. Both
of these men of science admit the great influence in their studies
of their careful instructor, Thure Kumlien.
About the time that Kumlien began his engagement as professor
at Albion Academy, a new hall was erected. This was at once
named Kumlien Hall. The building is still in use.
Frank Bardeen, a member of Kumlien ’s class at Albion, shot
a strange bird on the campus, and on taking it into his study
672 Wisconsin Academy of Sciences , Arts , and Letters.
Kmnlien exclaimed: “I have not seen that bird since I was a
boy in my native land. It is the Passer domesticus, or English
sparrow.” This was the first appearance of this busy bird in
Jefferson county, about 1870 (10).
Professor Greene, who carried on an intimate correspondence
with Kumlien up to the time of his death, says of his work:
“During the first twenty years of his residence in America, Mr. Kum¬
lien was engaged in forming collections in all branches of natural his¬
tory for such celebrated institutions as the Stockholm, Leyden, and Brit¬
ish museums in Europe, and the Smithsonian in this country; and also
for many private individuals on both sides of the Atlantic.”
In a letter to Dr. John H. Twombly, President of the Univer¬
sity of Wisconsin, written between 1871 and 1874, Kumlien says :
“During the time I have lived in this country (since 1843) I have sent
many large collections of birds, insects, etc., to the museums of Leyden,
Holland, Prof. H. Schlegel; Berlin, Prof. Peters; Stockholm, Prof. Sunde-
vell; Upsala, Pr. Lilljeborg; and many smaller collections to Engand and
east in the United States; and to the Smithsonian, Philadelphia (Cass in)
and to Boston Society of Natural History, of which I have been a member
since 1854.
“Of plants I have sent large collections to Prof. E. Fries, Upsala, among
which plants several are by him considered different from those described
in our American works. I have consequently had a great deal of experi¬
ence in collecting in almost every branch of natural history.”
He was employed by the state of Wisconsin in forming and
arranging collections in natural history and botany for the Uni¬
versity of Wisconsin, all of which were destroyed in the burning
of the first Science Hall. He was also employed to make collec¬
tions of fauna and flora for several of the Wisconsin normal
schools.
William M. Wheeler, an intimate friend, connected with the
museum at Milwaukee, says of his work:
“From his arrival in Wisconsin to 1863 he devoted his time to forming
natural history collections for the Stockholm, Leyden and British Muse¬
ums, and for many museums and private parties in the United States.
This work necessitated considerable correspondence. Mr. Kumlien was
in constant communication with Dr. T. M. Brewer of Boston, from 1851
till Brewer's death in 1880, and was one of the largest contributors to
the History of North Ameircan Birds, published by Brewer, Baird, and
Ridgeway. Other correspondents were Professors E. Fries, Sundeval,
Nielson, and Von Ehlen of iSweden ; Steensrup, Sars, and Loven of Nor¬
way; Prof. Peters of Berlin, Count Turati of Milan, Prof. H. Schlegel of
Lawson — Thure Kumlien.
673
Leyden, and Professors J. E. Gray, Alfred Newton, and H. E. Dresser of
England. In this country he corresponded with nearly every prominent
ornithologist and botanist up to within the last six or eight years of his
life. In 1867 he was appointed teacher in Albion Academy, a position
which he held till 1870. Later he formed the collections for the State
Normal Schools and for the State University. From 1881 to 1883, Mr.
Kumlien was employed by the Wisconsin Natural History Society. For
the past five years he has been Conservator to the Milwaukee Public
Museum" (3).
In Birds of Wisconsin , by Ludwig Kumlien and N. Hollister,
mention is made in the sources of information:
“Added to this, and perhaps or even greater value has been the use of
the extended, accurate and perfectly authentic notes of the late Thure
Kumlien, covering a period of constant residence in the state of nearly
forty-five years, from 1843 to 1888’ (16).
In Studies in Plant Distribution by Ernst Bruncken (Bull.
Wis. Nat. Hist. Soc. 2: 151, 1902), there is a list of plants from
Jefferson County collected by Thure Kumlien for the Milwaukee
Museum, a work “mostly done forty and more years ago” — 1862
and before. The Two Rivers list names one plant .found there
by Thure Kumlien.
Spencer Fullerton Baird, a noted American ornithologist and
author, wrote Kumlien, August 30, 1880 :
“Would you consider twenty dollars sufficient compensation for your
swan? I shall be very glad to have it as an important addition to the
Museum of the Smithsonian Institute".
i,
The offer was accepted, and the trumpeter swan is still pre¬
served in the National Museum at Washington.
Kumlien furnished to the herbarium of the Public Museum of
the city of Milwaukee 163 identified specimens of the plants and
flowers of Wisconsin included under numbers from 26 to 1726,
“thereby contributing greatly to the value of the museum’s her¬
barium” (17).
“A new museum is being built in Stockholm (1913) which is to have
many of Thure Kumlien’s specimens’^18).
He wrote on one occasion in 1861 that,
“having more than twenty years’ experience in collecting and prepar¬
ing specimens of natural history for museums”,
43— S. A. L.
674 Wisconsin Academy of Sciences, Arts, and Letters.
he offered
“his services in collecting whatever in the several departments of zoolo¬
gy was desired, such as birds, small mammals, reptiles, Crustacea, and in¬
sects, especially insects from Wisconsin, Illinois, Iowa, and Minnesota.”
J. E. Gray of the British Museum obtained specimens of mam¬
mals, birds, freshwater fish, reptiles, shells, and insects. Dr.
Brewer writes, March 25, 1855, that Dr. Baird wanted bird skins
for himself and the Philadelphia Academy of Science. W. Peters
of the Zoological Museum of Berlin, Germany, ordered, on No¬
vember 4, 1852, and February 20, 1864, from Kumlien, snakes,
cotton rat, harvest mouse, soft-tailed mouse, gopher, prairie dog,
kangaroo rat, tortoise, 500 insects, reptiles, and squirrel.
F. M. von der Walp, The Hague, on June 5, 1867, wrote Thure
Kumlien in part:
“In the government’s museum of natural history at Leyden were
received, one or two years ago, North American diptera, which were sent
there by you. Having applied myself since several years on the study of
that order of insects, the conservator of the entomological part of the
museum, Mr. Snellen van Vollenhoven, has brought these diptera under
my eyes, and I have helped him in determining them. As I was occupied
with this work, I found among them several very interesting species.
Since the last time also the North American diptera have attracted much
the attention of European entomologists, and this has excited my desire
to become acquainted with these insects. It is with this purpose that I
take the liberty to write you the present letter.’”
Kumlien was asked by Prof. E. Fries, prefect for the Royal
Botanical Garden at the University of Upsala, Sweden, to collect
plants, seeds, and specimens of natural history. He furnished a
cabinet of natural history “ birds and animals, mounted by him,
to the Albion Academy (19).” In his earlier days, before enough
land was cleared and the wild-cat money became reliable, he sought
to exchange eggs, skins, and other natural history specimens for
books on ornithology.
Mr. William Brewster, in 1880, assistant in charge of the col¬
lection of birds and animals for the Boston Society of Natural
History, and in 18855 curator of ornithology at the Museum of
Comparative Zoology, Cambridge, obtained a collection of bird
skins labeled with dates, locality, and sex, as was the usual prac¬
tice. Kumlien collected for and sent to Mr. Edward August
Samuels, Boston, in 1864, many wild bird skins, eggs, and nests.
Lawson — Thure Kumlien.
675
There was a warm friendship and an intimate correspondence
between Kumlien and Dr. Thomas Mayo Brewer, naturalist and
author of bird books, who was also connected with the Smithsonian
Institute. Dr. Brewer died in Boston January 24, 1880. He
edited an edition of Wilson’s Birds of North America in 1839.
It was this edition of which Kumlien secured a copy about 1849.
Brewer, with Baird and Ridgeway, published a History of North
American Birds in 1874. After March 30, 1851, a correspondence
was begun between these two naturalists which ended only with
the death of Dr. Brewer thirty years later. The subjects included
the habits of birds, their descriptions, and eggs and nests. Brewer
took care of the sale of bird skins for Kumlien, and of eggs and
nests and all other subjects of natural history, and found books
and reports for him. He procured pins for insects and glass
bird-eyes for mounting. He mailed seeds of a new squash and
bought butter from Kumlien. He advised about the wealth of
prospective customers for natural history objects.
In 1859, in a letter to Dr. Brewer, Mr. Kumlien wrote :
“I am very fond of natural history and have been since a school boy,
but my means do not allow me to follow my inclinations in this respect,
and to increase my collections as I should like, so that I sometimes, plow
in hand, make observations.”
In 1851, he wrote Brewer :
“Not being enough acquainted with the English language to write the
same any ways proper, it is with some hesitation I undertake to write to
you. I am a native of Sweden and emigrated to America in 1843, but
have ever since been shut up in the woods, and my time most wholly
devoted to hard work.”
It is evident from the language and the beautiful handwriting
of this, among the earliest of his letters that I have seen, that
Thure Kumlien had no reason for excuses on either score. By the
possession of the Birds of America for the previous two years,
he had at that early date made himself familiar with the proper
scientific names of all the birds in his region, and had already
made a catalogue of the birds seen about him. He had mentioned
this catalogue to Mr. Holt, and noV offered to mail a copy to
Brewer. The warm friendship of these two naturalists was
fortunate for both of them. As for Kumlien, it kept him con¬
stantly in touch with the work and thought of other naturalists.
To Brewer it gave the most expert and educated field observa¬
tion, research, and information.
676 Wisconsin Academy of Sciences , Arts , and Letters.
Brewer and Kumlien were made acquainted with each other
through Mr. Charles Holt (20) of the Janesville Gazette, early in
1851. Brewer wrote, March 31, 1851, that Mr. Holt had informed
him that Kumlien could help him to illustrate the eggs of North
American birds. To his offer to pay for the eggs, Kumlien said
he would have collected them without pay if he could afford the
time. He knew no “rule by which to measure the value ’ \ As
to the time required, “one may not find desirable eggs in days,
and again find many in a few hours.” He left the matter of the
value to Brewer. This was the system he ever followed with all
his customers, for all his collecting. Brewer also wanted birds’
nests. He had no place then to accommodate skins, but Kumlien
could send them to him and he would sell them for him. This
was the custom for many years afterward.
Kumlien had written Brewer for books on quadrupeds and
botany, and about glass eyes. In reply, after writing a long
letter detailing the bird skins, eggs, and nests he would like to
obtain if it could be, ‘ ‘ without it interfering with your other
duties”, he concludes:
‘‘With regard to the books you ask about, there is no book on our
quadrupeds that conies at a low price. Audubon is to be $30.00, and is
not yet finished. There was one published some time ago, but it is out
of print. I believe it was Goodman’s and that cost considerable. Gray’s
Manual of Botany, a friend tells me, is the best book on botany. It costs
somewhere between one and two dollars. I think I can get one for less
than the bookstore prices at a second hand store, which I will as soon as
I can, and send you by mail. A friend of mine wishes me to ask you
Whether you would like to exchange dried plants of New England for those
of Wisconsin. He will send them to you if you do wish for them, for
specimens of your neighborhood. Glass eyes cannot be readily procured
here. Some time since I had to purchase a few from a friend in New
York, but I do not remember the price. I am very glad to learn that you
like my paper, and be assured I am happy to continue it. Let me hear
from you soon. Write me whether you think you can procure any of the
skins for Mr. Lawrence.”
Brewer wrote, August 2, 1854, of the election of Kumlien as
corresponding member of the Boston Natural History Society,
saying he had mentioned before the meetings on several occasions
his good work, and
“facts of interest I had derived from you, and our secretary Dr. Ab¬
bott asked me one day why I did not propose your name as a correspond¬
ing member. I assented, and he nominated you at the next council meet¬
ing.”
Lawson — Thure Kumlien .
677
The following intimate and interesting letter, from its contents
and date, is supposed to have been written to Dr. Brewer at
Boston. In his correspondence Kumlien wrote each letter over
twice, thus preserving a copy. If these copies could all be found
now, one might easily write the intimate story of his life.
“September 29th, 1856. Dear Sir! In hopes that during the summer I
would obtain some more eggs and skins I have postponed writing to you.
Never, it seems, did I get so small a lot of eggs and skins. True, I have
been obliged to work at farming this season and that has hindered my
egging and hunting operations to some extent, but when I have been out
I have had very little success indeed. Some of my boys, that is, my egg-
boys, have furnished any quantity that you don’t want and the prairie
boys have been down with the smallpox during egg season. They had
promised Prairie Lark eggs and eggs of the Lanius. The Lake that 2
or 3 years back has been very high in summer has this summer been so
low that, where I used to pick eggs of Gallinules, Black Terns, Wrens
and Yellowheaded blackbirds it was perfectly dry and not a nest of those
birds to be seen and but very small number of the birds themselves. I
will mention what I have obtained of eggs and skins.
Eggs. Sterna fissipes a few.
Crex Carolinus 3 or 4.
Rallus virg. 8.
Rallus elegans 6.
Muscicapa 3 eggs in the nest (I keep one);
it may be nothing else but the wood pewee which it greatly resembles
having under bill yellowish, but I think perhaps it is not. Wilson says
the wood pew. eggs are white; those are white too, but on the place where
the egg is thickest or rather a little nearer the big end is a ring of dark
spots some large and some very small, a marking I think Wilson would
have mentioned did those of the wood pew. have such marks. I have
prepared skins of both the male & female. I think this bird not alto¬
gether rare here, I observed 2 more nests but could not get at them; they
too were in Burr oaks but very high and out on the weakest limbs. I
have found a nest of Muscicapa minima but the boy I had to climb the
tree saw only 2 eggs so I concluded to wait a few days and get more
eggs; some other boys learned where the nest was and in hopes of a
good pay took the eggs and afterwards quarreling about their prey broke
the eggs. This bird seemed not to be very rare this summer and I may
have better luck another year. I have two eggs of Coccyz — erythroph-
thalm — lot of Qwails — some Blue winged teals. This is about all except 5
nests of small birds that I do not know for certain where they belong
. If you think those eggs and skins are of sufficient value to
send I will send them else I will keep them till another season in hopes
of getting a greater variety and number. A few days ago I got Talco
fuscus 2 & tennessee warbler 1, all skins. The warbler I procured last
fall and could not then make out, is nothing else but S. icterocephala in
immature plumage . Your obt. servant Thure Kumlien” C20).
678 Wisconsin Academy of Sciences , Arts, and Letters.
He writes to Brewer, August 11, 1861, in detail of all the
birds, eggs, and nests he had seen that season and what he had
secured, then adds that he had received an advertisement of a
splendid work with plates by Baird and Cassin on North American
birds: “to purchase such a work is for me out of the question”.
He requests Brewer to ask the editors if he could get the work
“by exchanging birds and animals. I am willing to collect a good deal
to obtain a good book.”
“I am about collecting insects for the Royal Museum in Stockholm,
Prof. Boheman, and know of no way to get insect pins here. May I ask
you to assist me in this matter. Are such articles kept for sale in Bos¬
ton? If so can I trouble you with sending me by express a lot of 5 to
6000, if they are not too high prices I would like 10,000 (altogether of
2 or 3 sizes, most of the small numbers). I know not the price here,
but in Copenhagen they cost about one dollar for 1000. As soon as possi¬
ble I can get hold of Eastern money I will repay you, for our money here
is very bad indeed, most of it uncurrent everywhere. If I can obtain
no pins in U. S. I will have to order some from Europe, but then I would
have to put off collecting insects till next year. I know I am taxing your
kindness to the utmost, but forgive! You have always been so kind to
me and I have.no friend or acquaintance at all in the East! Yours very
respectfully, Thure Kumlien.”
The Museum of Comparative Zoology, founded in 1859 at Cam¬
bridge, with Louis Agassiz as its curator, was the repository of
Agassiz’ immense accumulation of scientific collections and was
commonly called the Agassiz museum. In a letter dated October
23, 1873, Dr. Brewer writes:
“Mr. Allen of Prof. Agassiz’ Museum is to spend the evening with me.
I can then tell from him what he will want for the museum of skins,
nests, and eggs; and I shall went those separately.”
On the order which was received, some fifty birds’ nests, eggs,
and skins were shipped to the museum of comparative zoology,
Cambridge. It was this and other information that came to Louis
Agassiz before and after that led him to make the remark pub¬
lished in the public press that Thure Kumlien of Busseyville was
the greatest authority in the world on birds’ nests.
We have no space to print the mass of information and ques¬
tions put to Kumlien in this correspondence, but to illustrate
here is one in a letter from Dr. Brewer dated November 22,
1873:
“I shall be glad of all the notes you or your son can let me have in
regard to water birds. What you have? When they visit you? How
Lawson — Thure Kumlien,
679
long they stay with you, and if you can tell me in regard to them while
With you?”
A small white alpine anemone that disdains to exist below five
thousand feet altitude, but lives, thrives, and blossoms on the rug¬
ged rock escarpment of the Sierra Nevada Mountains, just at the
perpetual snow line, above the clouds, was named, for our natural
historian, Kumlienia. Few men but the most daring mountain
climbers will ever enjoy the sight of this beauty of the clouds.
But all who, know the story of this hardy survival of the fittest,
this charming white emblem of purity blossoming up there against
perpetual snow caps, will never look on those picturesque ever¬
lasting granite hills rising one above another into the birthplace
of the winds, without a kindly thought for Kumlienia.
Edward Lee Greene, in his life of Thure Kumlien, writes of it:
“A rare and still but little known Ranunculaceous plant of the middle
Sierra Nevada in California, a flower with calyx of a Caltha, corolla of
Helleborus, and an utricular carper not like that of any genus of its fam¬
ily, a clear generic type, was dedicated to him two or three years since
(1885) as Kumlienia, but these small tokens, like our worded tributes,
are all inadequate to speak the praises, or worthily perpetuate the memory
of a man so pure, so simple, so noble, and so well beloved” (4).
“Among the purple autumnal Asters, as they grew around him there,
at least in the earlier time, there was one species, which received from
Prof. Elias Pries the name Aster Kumlieni” (4) .
Aster Kumlieni is represented in the herbarium of the Public
Museum at Milwaukee by specimens marked 456 and 457, col¬
lected in 1858-60 in a ditch three quarters of a mile of Bussey-
ville, at a place known as Beneworth’s collected by Thure Kum¬
lien and originally labeled by him i( Aster amethyst inns Nutt,”
and “ Aster oblongifolius Nutt” (17). He mailed specimens to
Fries, at Upsala, who pronounced it a new species, and gave it
the name Aster Kumlieni in 1860.
The Kumlien vireo (Vireo Philadelphicus ) is referred to by
Kumlien in a letter to Prof. H. Schlegel, of the Royal Museum of
the Netherlands, February 13, 1862, as “Viris PJiiladelphica. I
was the first who found that bird.” Why this rare and little
known beauty of the orchard and forest was not properly given
the name “Kumlien Vireo” for its discoverer, the author cannot
now ferret out. This late justice is accorded our modest natur¬
alist.
680 Wisconsin Academy of Sciences , Arts , and Letters.
From Dr. Brewer’s letter of November 20, 1854, it appears
that Kumlien had discovered the new vireo in 1854, or earlier.
Ornithologists in Boston became interested in the find. It was
supposed by Kumlien to be the Vireo Belli of Audubon. Others
thought the same.
Dr. Brewer had written Kumlien that Mr. John Cassin had de¬
scribed this vireo in the Proceedings of the Philadelphia Academy,
and Brewer wrote that he had prepared a paper for the Boston
Society of Natural History, “in which I mean that you shall have
all due credit.” After this paper was published, Mr. Cassin wrote
Dr. Brewer to compliment him on the paper.
The case is thus stated in Birds of Wisconsin;
“In the early forties Thure Kumlien procured specimens of a vireo
which he named Belli of which he could find no description. This led to
some confusion with Lewrence, Baird, and others who had not seen the
specimens. The bird referred to was later described by Cassin as Vireo
Philadelphicus” (16> .
Kumlien was not a traveler. In 1865 he had planned to make
a zoological and botanical exploration through Kansas, and a
visit to his friend Greene, then in the northern army in Tennes¬
see; but I cannot find any evidence of these journeys having been
taken. He frequently made visits to Madison. He went by boat
to Two Rivers. He lived for the last seven years of his life in
Milwaukee. He may have made journeys to other parts of Amer¬
ica and Wisconsin, and doubtless he did. The herbarium of the
Milwaukee Public museum has collections made by him in Wau¬
kesha County. In a number of letters he proposed to make a
six months’ tour of the region of the upper Mississippi River and
its principal tributaries and of the Lake Superior region in 1861.
Some memoranda given the date on which he shot a certain bird
as June 17, 1861, and the place as Sauk City. He may have ac¬
complished the contemplated excursion, as Sauk City is on the
Wisconsin River. One of his letters mentions having returned
after a six months’ absence from home.
“Krom miles about, people came to Tbure Kumlien to consult bim on
all manner of domestic questions because of bis general reading, informa¬
tion, and education. They consulted about their taxes, and got bim to
survey their land lines.”
He was well known and respected and looked up to by all his
neighbors for many miles about his home. He was a God-fear¬
ing, honest man, and never had a lawsuit.
Lawson — Thure Kumlien.
681
From a letter which Kumlien wrote his son Frithiof, we get
an account of the crowds of people that came to him within the
space of a few days.
“Last Monday Currin was here with 75 or 100 followers of both sexes.
Now for two or three days I have had a visitor by name of iStiles from
Lake Mills, a man much interested in natural history” ( 23 ). “Two other
visitors today. Next Saturday I expect 40 boys and I don’t know how
many girl students from Whitewater Normal under the guidance of Prof.
Wooster” (20) .
This is the story of one week, and we do not wonder that he
writes in the same letter “the tobacco plant bed is quite good
except where the cows have perambulated.” These mobs of people
who came and went all summer, every season enjoyed the expert,
kind naturalist, and he gave them real pleasure and inspiration.
In the host of letters we have seen the writers all love to recall
the good naturalist with warmest love and admiration.
Kumlien was the recipient of honorary degrees from several
institutions of learning, and was corresponding member of a
number of learned societies in Europe and America (4).
He joined the American Ornithologists’ Union in its first year
(1883) (15). He was elected corresponding member of the Bos¬
ton Society of Natural History June 7, 1854. He was a member
of the Wisconsin Natural History Society, and of the Wisconsin
Academy of Science, Arts, and Letters, founded in 1870.
Kumlien amused himself all his life with the flute. At an
early age back in Sweden he took up that delightful instrument.
There are three notebooks still in possession of his family, that he
made for the flute in 1839, when he was twenty years old.
“One male oriole was known to return to the Kumlien homestead five
years in succession with a mate, and build its nest on the same limb each
year. The bird was accustomed to answer a certain note on the flute,
and seemed happy to try to imitate different flute notes” (16).
Ludwig Kumlien took wild mourning doves from the nest when
young. They were raised at the old home, in 1872, and became
quite tame. At the time a letter was written by Thure Kumlien
to Dr. Brewer, the doves had been captive two years:
“The male is so tame, he will light on my head, allow me to take him
in my hand and stroke him. He will coo whenever I ask him, unless
strangers are present. He will coo evenings after dark. He is sure to
coo if I play a melancholy tune on the flute.”
682 Wisconsin Academy of Sciences , Arts, and Letters.
Kumlien was a close observer of all fauna and flora; he knew
all the birds, animals, and insects, and through years of observa¬
tion found out all their habits. The plants and flowers were all
intimately familiar to him. All who wrote on any of these sub¬
jects of natural history had free and welcome access to all this
information, and many failed to give due credit for its source.
For forty years he furnished much of the detail of papers and
books on natural history subjects published in America. It is
to be regretted that he went to his rest with all or most of that
wonderful story of detail of natural history accumulated through
the years unpublished. Mr. W. M. Wheeler has written of this
looting of the brains of the expert:
“Mr Kumlien’s arrival in our country at a time when many of our na¬
tive species were still very imperfectly understood opened a wide field
for work. Too modest to think his own often very valuable observations
sufficiently important to publish, he devoted many years of his life to
helping other naturalists by sending them large and carefully prepared
collections. Mr. Kumlien was a singularly accurate observer, though his
powers of observation were stimulated by an intense and childlike love of
natural objects rather than by any interest in their importance from a
speculative standpoint. Hence his conversation teemed with interesting
facts, but seldom rose to wide generalizations. He was satisfied to ob¬
serve, to collect and prepare plants and birds because they were full of
marveolus beauty and offered endless material for comparative study” (3).
Prof. E. A. Birge had spoken to Kumlien several times about
writing out his information on natural history, and on Septem¬
ber 17, 1879, wrote him as follows:
“I want to urge again on you what I spoke of to you sometime ago,
vis., that you give the Wisconsin Academy a paper on some subject next
winter. You know a great deal about the habits, etc., of our birds and
mammals which no one else knows and much of which no one else can
ever know now. It ought to be put on record and I hope that you will
find time to set down some of it before next Christmas.”
He wrote again in 1882. It is much to be regretted now that
these kind invitations were not accepted.
The magnificent museum at Milwaukee had been founded by
the Wisconsin Natural History Society in the days of Increase
Allen Lapham. In the year 1881, Kumlien was engaged as taxi¬
dermist and conservator at this museum, a position he held under
that society until May, 1883, when the museum was taken over
by the city of Milwaukee. Kumlien was engaged at once by the
new management and remained in that position until his acci¬
dental death by poison (3).
Lawson — Thure Kumlien.
683
While in the museum, for the last seven years of his life, he
employed himself,
“as taxidermist and conservator, to take care of the collections in the
museum. His first work among the museum collections was the exam¬
ination and classification of the mounted specimens of birds and mam¬
malia, which was followed by an examination of the skin collections
stored away in the museum” (24).
During his incumbency the museum was located in the origi¬
nal Exposition Building, which was later burned.
While at the museum, Kumlien employed himself with stuffing
and mounting some few birds, and also in sorting over geological
specimens and arranging them in the cases. In the afternoon he
gave much time and attention to visitors that crowded the place,
and assisted when the crowds were greatest during annual exhi¬
bitions in the main exhibition rooms. He had finished the ex¬
amination and repair of birds and animals then mounted and in
place in the museum, and had got up the boxes of skins that had
been shipped to the museum and stored away until such time as
his duties permitted him to restore and mount them.
The exact facts as to his death are a bit hazy at this late date ;
but Mr. Carl Thai, who has been for twenty-seven years librarian
at the museum, and was engaged in the museum at the time and
knew Kumlien, who lived with Mr. Thai, recalls
“that it was the impression at the time of Professor Kumlien’s death,
that it was due to the effec of corrosive sublimate poisoning through in¬
halation. He was working over a large collection of prepared bird skins,
which had been poisoned with corrosive sublimate, and it was assumed
that he inhaled a considerable amount of this deadly poison and that it
was this that caused his death” C25).
Kumlien died at Passavant Hospital (now Milwaukee Hospital)
at 3:30 in the afternoon, of Sunday, August 5, 1888, the same
day he was taken to the hospital. The hospital record reads :
“K. Kumlien, 69 years old. Protestant, German nationality, admitted
Aug. 5, 1888, diagnosis chronic diarrhoea. Died same day. Physician
Dr. Nicholas Senn.”
The “K”, should be “ Thure” : “ German” should be “Swe¬
dish”. It is gratifying to know that he had as attending phy¬
sician the renowned Dr. Nicholas Senn. As the disease was re¬
ported as chronic, it is safe to assume that long working with and
inhaling small quantities of deadly poisons finally undermined his
684 Wisconsin Academy of Sciences, Arts, and Letters.
constitution and that he succumbed a martyr to his profession.
Dr. J. J. Kaumheimer, secretary to the medical staff of the hospi¬
tal where he died, writes the author that there is only one per¬
son now living who was connected with the institution at that
time. In the same letter Dr. Kaumheimer explains the action of
these poisons:
“You say the man was a naturalist. This, with the diagnosis given,
leads me to the following explanation which, however, is purely personal
and speculative. In the preparation and preservation of shipped speci¬
mens, large quantities of arsenical and mercurial preparations are used
as preservatives. It iS" possible that by constant association with these
poisons he inhaled enough to cause poisoning, chronic looseness of the
bowels being a not infrequent symptom of such poisoning, if not acute.”
He was buried Tuesday, August 7, at the cemetery near his
home (2).
“(From 1881 to the time of his death he held the place of conservator
to the Milwaukee Public Museum; a position which he was about to re¬
sign simply in order that he might retire and pass a peaceful old age at
his quiet and secluded home near Lake Koshkonong. The dear com¬
panion of all his early and long years of frontier life had passed to her
rest in 1874. Four of his children, three of them sons and all adult,
were spared to him until early in the present year, when Frithiof, the
youngest, died, and the father’s bereavement was most distressing. But
there was no indication that his own end was approaching. A young
man, indeed, for one who had so nearly filled out his three score years
and ten; neither mind nor body yet showing the infirmities of age. He
was making ready for a return to the birthplace of his children, and now
of his grandchildren, to be with them thenceforward. .His own death
came speedily, from accidental poisoning, and that after long years of
experience with the deadly chemicals of the botanical and zoological
laboratory” (4).
“Mr. Kumlien was no narrow man. He was passionately fond of paint¬
ing, music, and poetry. I have heard him repeat with a glow of delight
verses from Buneberg, and from Frithiof’s Saga, rendering the wonder-
full rhythm of the latter with exquisite grace and precision. He was a
man of most refined tastes, without any of the extravagant desires which
such tastes often engender. He was satisfied to live most simply a life
which philosophers might envy.
“None who met Mr. Kumlien ever forgot his kind address, the lack
of all affectation and the modesty and ease of his conversation. He was
very fond of the young and always ready to put at their disposal his long
experience as a practical ornithologist and botanist. Such qualities are
not to be underestimated in a naturalist, for they are the means of charm¬
ing the young and making good naturalists of youths who would be re¬
pelled by a cold exterior. Many of our rising botanists and zoologists
owe much to Mr. Kumlien’s warm and sympathetic enthusiasm, which
was as contagious as hearty mirth” (3).
Lawson — Thure Kumlien.
685
Governor Alva Adams of Colorado, who knew him in youth,
wrote :
“Kindness is the characteristic I best recall. I was but a boy when I
knew him. As I remember him, he was such a man as I would today
like as a companion to go up Into the mountains to tread its ways of
river and wilderness streams, to study the trees, flowers and birds. He
knew and loved nature. He was her interpreter. Simple as a child
among men, he was wise in the works of God. As a professor of its
‘“Out-ofdoors” he ranks with the masters” ( 27 ) .
NOTES
a Document in possession of Mrs. Angie Kumlien Main, Fort Atkinson, Wis¬
consin.
2 Jefferson County Union, August 17, 1888.
3 Necrologue by W. M. Wheeler, as a supplement to the annual report of the
trustees of the Milwaukee Public Museum, 1888. “Life of Thure Kumlien”.
See notes 5 and 8. Sundevall, Boheman, and Loven were connected with the
Royal Academy of Science, Stockholm, Sweden.
4 “Sketch of Life of Prof. Thure Kumlien, A. M.” by Prof. Edward Lee
Greene, Pittonia, vol. 1, part 5, October 1888, reprinted in Albion Campus, June,
1891. Greene was professor of botany at the University of Notre Dame when he
died, and left his library and collections to that institution.
8 “On the rapid disappearance of Wisconsin wild flowers”, etc. Paper read
by Thure Kumlien before the Wisconsin Academy of Sciences, Arts, and Letters,
published in the Transactions of the Academy, vol. 3, p. 56, 1876. A typo¬
graphical error in this paper makes his name read “Kumlein”.
6 Th. M. Fries, whom I suppose to be a son of Prof. Elias Fries, writing to
O. A. Linder of Chicago, from Upsala, Sweden, December 6, 1908, speaks
“concerning Thure Kumlien whom I well remember since the time that he
(in the beginning of the 1840’s) visited in my parents’ home.” He also writes
that there are biographical notations on Thure Kumlien “preserved in the na¬
tional archives at Vestgotha.” Also “a letter is found there from — ”. As the
letter did not complete the sentence, the information is incomplete. He adds:
“Besides this I have searched through my father’s (Elias Fries’) scientific
correspondence file which has been donated to the library of the University
of Upsala. In this are found eight letters, all quite lengthy, but were written
during the beginning of the 1850’s and are almost exclusively concerned with
collected plants. As far as I know, the correspondence of Prof. C. J. Sundevall
has been destroyed ; otherwise there would undoubtedly have been found con¬
siderable of interest in it.”
7 From unpublished paper by Oliver A. Linder, Chicago, and memoranda fur¬
nished him from Sweden. See note 4.
8 Letter to the author by T. V. Kumlien, Fort Atkinson.
9 Letter by Thure Kumlien written to his son in reference to Gerhard Von
Yhlen.
10 A private paper by T. V. Kumlien in possession of Mrs. Angie Kumlien
Main.
n “Commemorative biographical record of counties of Rock, Green, Iowa,
Lafayette, Wisconsin”, pp. 89-90, Chicago, 1901.
12 Certificate of marriage.
686 Wisconsin Academy of Sciences, Arts, and Letters .
13 Reminiscences (Minnen) of seventeen years’ wanderings in northwestern
America”, Gustof Unonius, vol. 11, p. 221. 1861.
14 Original certificate.
15 Dr. T. S. Palmer, 1939 Biltmore St. N. W., Washington, D. S., 1920. Bio¬
graphical notices of Thure Kumlien appeared in the “Auk”, vol. 6, pp. 204-205,
1889,
16 Bulletin of Wisconsin Natural History Society, new series, vol. 3, nos. 1, 2,
3, pp. 1—143, January— July, 1903. The author cannot now locate these notes.
His natural history collection and library was willed to his son Theodore, who
has transferred it to his daughter Mrs. Angie Kumlien Main. Letter from
Theodore Kumlien.
11 Letter to the author by Huron H. Smith, Curator of Botany, Public Mu¬
seum, Milwaukee, 1920.
18 Letter, Eva Theleen, September 22, 1913, to Mrs. Angie Kumlien Main.
19 Letter to the author by Sylvanius Ames, 1920.
20 See life of Charles Holt in “Fathers of Wisconsin” ( Tenney and Atwood,
Madison, 1880) dated June 2, 1881.
21 “Wild asters of Wisconsin”, Charles E. Monroe, Bulletin of the Wisconsin
Natural History Society, vol. 3, September, 1913.
22 “Bird guide. Part 2 : Land birds east of the Rockies,” Chester A. Reed, 1906.
23 E. H. Stiles of Richland Center, 1888.
24 Recollections of Carl Thai, obtained for the author by Dr. S. A. Barrett.
26 Letter to the author by Dr. S. A. Barrett, Anthropologist, Public Museum,
Milwaukee, March 7, 1920.
26 The cemetery where Thure Kumlien is buried is over the line in Dane
county, three and one-half miles from his old home and one and one-half miles
from Albion. It is a Protestant cemetery used by two M. E. and P. M.
churches.
27 Children of Thure and Christine Kumlien, all born in the town of Sumner,
Jefferson county, Wisconsin : Augusta Christina Sophia, born March IT, 1845,
died October 24, 1845 ; Ludwig Aaron, born March 15, died December 4, 1902 ;
Theodore Victor, born January 7, 1855, resides at Fort Atkinson, Wisconsin
(1920) ; Frithiof Axel, born December 19, 1859, took the degree of B .S., Uni¬
versity of Wisconsin, 1882, died January 6, 1888 ; Svea Maria, born August 8,
1857, died August 2, 1891, married Robert S. Martin ; they resided at Cedar
Mills and other places in Minnesota and both died in Colorado.
EXPLANATION OF PLATES
Plate lxii
Plate lxiii
Frame dwelling of Thure Kumlien completed in 1874. The original log
cabin of 1843 is partly seen in the rear. Log cabin destroyed 1903. The
frame dwelling has been removed to the highway.
Plate lxiv
(1) Birth place of Thure Kumlien. Hertrop, Harlunda parish, Sweden,
from water color painting made by himself about 1860.
(2) Thure Kumlien when a student in Sweden from a pencil drawing
made by his brother.
(3) Thure Kumlien about 1880 from an old photograph.
TRANS. WIS. ACAD., VOL. XX
PLATE LXII
LAW S O \ — K U M LIE N
TRAAS. AVIS. ACAD., VOL. XX
PLATE LXIII
LAWS OX — KU3ILIEX
TRANS. AVIS. ACAD., VOL. NX
PLATE LXIV
(1) Birth place of Thure Kumlien. Hertrop, Hiirlunda parish, Sweden, from water color
painting- made by himself about 1860.
(2) Thure Kumlien when a student in Sweden
from a pencil drawing made by his brother.
(3) Thure Kumlien about 1880 from
an old photograph.
LAW S O N— -KUMLIEN
PROCEEDINGS OF THE ACADEMY,
1919 AND 1920
FORTY-NINTH ANNUAL MEETING, 1919
The forty-ninth annual meeting of the Wisconsin Academy
of Sciences, Arts, and Letters, in joint session with the Wiscon¬
sin Archeological Society, was held at Milwaukee, in the Trus¬
tees’ Room of the Milwaukee Public Museum, on Thursday and
Friday, March 27 and 28, 1919.
In the absence of the Secretary and Treasurer overseas, Mr.
R. H. Whitbeck assumed the duties of Secretary, and Mrs. Ar¬
thur Beatty had charge of the finances. There was no business,
with the exception of that of a routine nature, the most im¬
portant of which was the reports of the Secretary and Treas¬
urer, copies of which are appended to these minutes. President
Birge made announcement of the forthcoming Fiftieth Anni¬
versary of the Academy in 1920, and stated that steps were being
taken to observe the occasion in a fitting manner.
Under the direction of the President the following programme
of papers w^as presented:
First Session, Thursday, March 27, 10:00 A. M.
Introductory Remarks. President E. A. Birge.
General Business.
Presentation of Papers:
1. Psychological Tests of Aptitude for Flying. V. A. C. Henmon.
2. Milton’s Character as Reflected in His History of Britain. Harry
Glicksman.
3. The Problem of Controlling Atmospheric Temperature and Hu¬
midity for Plant Growth. W. E. Tottingham.
4. The Construction of a Sensitive Eight Coil Galvanometer. L. R.
Ingersoll.
5. The Construction of Power Vacuum Tubes and their use in Radio
Telephony. E. M. Terry.
687
688 Wisconsin Academy of Sciences , Arts, and Letters.
Second Session, Thursday, March 27, 2:00 P. M.
6. The Stratigraphy of the Sparta and Tomah Quadrangles. W. H.
Twenhofel and F. T. Thwaites.
7. Contrasts in the Economic Development of the Michigan and
Wisconsin Shores of Lake Michigan. R. H, Whitbeck.
8. The High Terrace Gravels of the Driftless Area of Wisconsin.
F. T. Thwaites.
9. The Wisconsin Soil Survey. A. R. Whitson.
10. Changes in Drainage Lines in Southeastern Wisconsin due to
Glaciation. W. O. Blanchard.
11. Transmission of Eye-effects produced by sensitized sera. M. F.
Guyer.
12. The Development of the Frenulum on the Wax Moth. W, S.
Marshall.
13. The Fishes of Lake Valencia, Venezuela. A. S. Pearse.
14. The Artificial Fertilization of Queen Bees. G. H. Bishop.
15. Chemical Warfare Work at the University with Particular Ref¬
erence to Phosgene. W. J. Meek, J. A. E. Eyster.
Third Session, Thursday, March 27, 8:00 P. M.
16. 8:00 — Public Lecture. Lecture Hall of Public Museum. En¬
trance on Eighth Street.
Subject: Gas Warfare. Major J. H. Mathews.
Fourth Session, Friday, March 28, 9:30 A. M.
17. The Proportions of the Human Body During Post-natal Growth.
C. R. Bardeen.
18. Fungous Parasites of Fern Prothallia. E. M. Gilbert.
19. Further Studies on the Mechanics of Sap-Flow. J. B. Overton.
20. The Fixation of Magnesium by Aspergillus niger. J. P. Ben¬
nett.
21. Notes on Parasitic Fungi in Wisconsin, VII. (By title.) J. J.
Davis.
22. The Distribution of Plankton Desmids in Wisconsin Lakes.
(By title.) G. M. Smith.
23. The Development of the Ovule in Oentiana procera. R. H.
Denniston.
24. Sex Differences and their Inheritance in Sphaerocarpos. (With
lantern slides.) C. E. Allen.
25. Regeneration from Antheridia in polypodium iroides. W. N.
Steil.
26. Temperature and Respiratory Exchange of Gases in Relation to
the Storage and Shipping of Potatoes. E. T. Bartholomew, J. P. Ben¬
nett.
Proceedings of the Academy.
689
27. The Summer Temperature of the Mud at the Bottom of Lake
Mendota. E. A. Birge.
28. The Behavior of CorethrU Punotipennis Say. C. Juday.
29. A Bacterial Blight of the Soy Bean. .Florence Coerper.
30. Progress with Disease Resistance in Plants. L. R. Jones and
J. C. Walker.
31. Dry Heat Sterilization of Seeds. D. Atanasoff and A G. John¬
son.
Fifth Session, Friday, March 28, 2 :00 P. M.
32. En Early English Map of the Great Lakes. Louise P. Kellogg.
33. An Archaeological Survey of Adams County. (By title.) H. E.
Cole.
34. Wisconsin Stone Gouges. Charles E. Brown.
35. Wisconsin Pottawatomi. Publius V. Lawson.
36. The Wisconsin War History Commission. Albert O. Barton.
37. Sheboygan and Manitowoc County Pottawatomi Chiefs. Al¬
phonse Gerend.
38. Manufacture of Indian Stone Axes. H. L. Skavlem.
39. Milwaukee War Gardens. L. D. Peaslee.
40. The Indian Mounds of Buffalo Lake. (With lantern slides.)
S. A. Barrett.
41. Upper-Middle Devonian Rocks in Wisconsin. Ira Edwards.
42. Glacial Lakes in the Menominee and Rock River Valleys. Ira
Edwards.
43. The Colby Meteorite of July 4, 1917. H. L. Ward.
44. The Establishment of the Cardinal in Various Wisconsin Lo¬
calities. H. L. Ward.
At the close of the programme the acting Secretary presented
the following applications for membership. On motion he was
instructed to cast the ballot in their favor :
Hugo William Albert, Madison
Dimitar Atanasoff, Madison
Augustus Lawrence Barker, Ripon
Edward Bennett, Madison
William Oscar Blanchard, Madison
Mrs. Ellen P. Butterfield, South Milwaukee
William Henry Coleman, Madison
John Rogers Commons, Madison
Charles Ives Corp, Madison
Henry Coe Culbertson, Madison
Charles A. Culver, Beloit
Charles Drechsler, Madison
August Frederick Fehlandt, Ripon
Charles Lewis Fluke, Jr., Madison
Ira Franklin, Milwaukee
44— S. A. L.
690 Wisconsin Academy of Sciences, Arts, and Letters.
Eloise Gerry, Madison
Harry Glicksman, Madison
Clark Graham, Ripon
M. C. Gugler, Madison
James Garfield Halpin, Madison
Josephine Ruth Hargrave, Ripon
Anna Heise, Ripon
Fred Reuel Jones, Madison
Delos O. Kinsman, Appleton
Otto Louis Kowalke, Madison
Olaf Lars ell, Madison
Publius V. Lawson, Menasha
Herdis L. Lewis, Madison
Genivera Edmunds Loft, Madison
Harold H. McKinney, Madison
Joseph Howard Mathews, Madison
Daniel Webster Mead, Madison
Eleanor J. Murphy, Madison
William James Mutch, Ripon
iF. Louise Nardin, Madison
Max Charles Otto, Madison
Bert Lorin Richards, Madison
Edith Catharine Seymour, Madison
Charles Mervin Slagg, Madison
Elizabeth Anita Smith, Madison
Walter Henry Snell, Madison
William N. Steil, Madison
Ruth A. Tillotson, Madison
William B. Tisdale, Madison
William Henry Twenhofel, Madison
Ray Hughes Whitbeck, Madison
Wilson Robb Woodmansee, Ripon
The acting secretary then presented his report for the past
year as follows. On motion it was adopted :
Report of the Secretary for the Year 1918.
Present Membership March 28, 1919
Honorary Members ........................ 6
Life Members ............................. 11
Corresponding Members .................... 41
Active Members . . 210
Total
268
Proceedings of the Academy. 691
Changes since Last Annual Report
New Memberships taken . . . . . 8
Active members in 1917 . 208
216
Resignations . . . 2
Dropped for non-payment of dues . 4 6
Present Active Membership . 210 210
New Application to be acted on at this Meeting . 48
I very much regret to report the death of Charles Richard Van
Hise, who served the Academy as President from 1894 to 1896, and
was elected to a Life Membership December 29, 1897. He died No¬
vember 19, 1918.
R. H. WHITBECK,
Acting Secretary.
The annual report of the Treasurer was then presented and
adopted :
Report of the Treasurer for the Year 1918
Receipts
Received from Dues and Initiations . . $187.17
Received from Sale of Transactions . - 14.39
Received from Interest on Liberty Bonds . 14.44
216.00
Balance on hand April 9, 1918 . 549.33
Disbursements 765.33
One-half of Secretary-Treasurer’s Allowance . $100.00
Safety-Deposit Box Rent . 3.00
Purchase of Bonds of Third Liberty Loan . 400.00
503.00
Balance on hand March 20, 1919 . 262.33
765.33
Present Permanent Investment consists of 25 City of Madison Bonds,
and 4 bonds of the Third Liberty Loan of the United States.
ARTHUR BEATTY,
Treasurer .
The meeting then adjourned.
692 Wisconsin Academy of Sciences, Arts, and Letters.
FIFTIETH ANNUAL MEETING, 1920
Fiftieth Anniversary Meeting
In accordance with the announcement made at the forty-ninth
meeting and in fulfilment of plans made earlier, the President
appointed a General Committee of twenty-five to consider plans
for the proper celebration of the completion of fifty years of
service by the Academy. The committee was as follows:
Rufus iM. Bagg, Appleton
W. Harley Barber, Ripon
Samuel A. Barrett, Milwaukee
Arthur Beatty, Madison, Secretary
Edward A. Birge, Madison, Chairman
Melvin H. Brannon, Beloit
Charles E. Brown, Madison
Charles N. Brown, Madison
Alfred C. Clas, Milwaukee
William C. Daland, Milton
John J. Davis, Madison
Lewis R. Jones, Madison
Edward Kremers, Madison
Publius V. Lawson, Menasha
Virgil E. McCaskill, Superior
William A. P. Morris, Madison
Samuel Plantz, Appleton
John M. Olin, Madison
John B. Sanborn, Madison
Charles S. Slichter, Madison
Erastus G. Smith Beloit
Halsten J. B. Thorkelson, Madison
Frederick E. Turneaure, Madison
Charles H. Vilas, Madison
Henry L. Ward, Milwaukee
This committee, under the direction of its chairman, President
E. A. Birge prepared a programme which represented the work
and relationships of the Academy, so far as they could be pre¬
sented in a single day.. The 23rd of April was chosen as the
day, as April was the month in which the first Bulletin of the
Academy was issued in 1870, and the 23rd was the date on
which it was possible to bring those persons together whom the
committee most desired to have present.
The following programme was projected and carried out.
Proceedings of the Academy.
693
First Session, 11 :00 A. M.
Auditorium, Biology Building, University of Wisconsin.
This session was a combined meeting of the Academy and a
University Convocation, presided over by Edward A. Birge, in
his combined capacity as the President of the Academy and
President of the University of Wisconsin. The object of the ses¬
sion was to hear an address by Professor Thomas Chrowder
Chamberlin, of the University of Chicago, formerly President
of the University of Wisconsin, President of the Wisconsin Acad¬
emy of Sciences, Arts, and Letters, 1885-1887, and one of its
original incorporators ; and to confer upon him the Degree Doc¬
tor of Science.
Professor Chamberlin was introduced to the session and de¬
livered the following address:
THE FOUNDING OF THE WISCONSIN ACADEMY OF SCIENCES,
ARTS, AND LETTERS
The event we are met to commemorate was a quiet one in itself.
It took place without noise or pagentry. But none the less, in the
intellectual history of the people of this commonwealth, it was a sig¬
nal event. The founding of the Wisconsin Academy of Sciences, Arts
and Letters, tho quite without the paraphernalia of a great event,
was yet a most distinctive step in the passage from the first stage in
the intellectual evolution of our people into this, the second stage.
The first stage, it is needless to say, was that of pioneer develop¬
ment. It began with the coming of our forefathers into this goodly
land between the Great Lake and the Great River. The territory was
then in its virgin state, tenanted by the wild life that had taken pos¬
session of it on the retreat of the Great Ice Invasion. This first stage
was a period of pioneer struggle and this struggle almost necessarily
delayed certain forms of scientific and cultural development. This
pioneer stage continued not only until the virgin prairies, the wild
meadows, the park-like groves, and the trackless forests of Wiscon¬
sin had been replaced by cultivated fields, comfortable dwellings and
prosperous towns, but until all these had been bound together by a
net-work of roadways and railways that united the whole into an in¬
tercommunicating cooperative community ready to enter upon a com¬
mon organized career in pursuit of its higher interests.
The second stage could really begin only when the conditions were
thus ripe for unified efforts to develop the higher intellectual, ethical
and aesthetic interests of the community. I think you will agree with
me that no step toward this higher evolution could be more fun da-
694 Wisconsin Academy of Sciences, Arts, and Letters.
mental than the beginning of a concerted endeavor to search out
rigorously, to test and to make known the basal truths that condi¬
tioned the lives of the Wisconsin people: — our habitat, the native life
of the land, our material inheritances, our climatic and other physical
surroundings, our social and moral conditions, our political institu¬
tions, as well as the arts and the literatures that made it possible
to use these most effectually. I do not think that the partiality of
the occasion leads us beyond the realities, when we regard the found¬
ing of the Academy as at least the most representative step in this
new development. It was of course by no means the only step, nor
was it the pioneer step in the transition from primitive conditions to
the more mature civilization to which the state has since attained;
for, in addition to the effective work of the schools and the churches,
which had taken on broader aspects and become more efficient as the
passing of primitive conditions permitted, the State Historical So¬
ciety, the State Agricultural Society, the State Teachers’ Association,
and other organizations had already taken up their special tasks and
had become effective agencies of progress; but, none the less, the
founding of the Academy was the most representative event in the
turn to the new order of things, for, better than any other single
event, it typified the coming of a higher order of endeavor, in that
its distinctive feature was cooperative research for the common good,
and this, I think you will agree, is the most basal and truest index
of real progress.
The movement furthermore was a comprehensive one, and altru¬
istic; it was unrelated to special interests. It was entered upon
spontaneously in full realization of the sacrificial labors that
would be necessary to make the enterprise a real success. And so,
in its high purpose and in its sacrificial spirit, this coming together,
fifty years ago, of good men from all parts of the state to found an
Academy whose chief purpose was to facilitate a concerted search
for truth for the common good, stands forth as an altogether signal
event in the intellectual development of our people.
The Pioneer Preparatory Stage
But before we pass on to review with gratitude and appreciation
the work of the founders of the Academy, let us pay a passing word
of respect to the pioneers who paved the way for the later era. Let
us also not altogether pass in silence the native conditions which be¬
came our inheritance and which contributed more than perhaps we
realize to what Wisconsin now is and is likely to be.
To one who saw the primitive wildness of this region as it was
vanishing and who played his little part in the early struggle to re¬
place the unbroken sod with cultivated land, it is a pleasure to recall
this early epoch and all that it meant to the founders of the state.
The primitive wildness had a charm which no one who saw it can
easily forget, and the struggle with this wildness, strenuous as it was,
had in it such an imperative call for personal resourcefulness and
Proceedings of the Academy.
695
such a toughening of physical and mental fiber as one would not
wish to have escaped. It brought its hard lessons of self-dependence,
of adaptation, of courage and of tenacity. It would be a pleasure
to dwell at length upon the primitive aspects of Wisconsin clothed in
the charm of its untouched nativity, but I must confine myself to that
one phase which stimulated some of the special intellectual activities
which led up to the event we celebrate.
Virgin Wisconsin was a paradise for the naturalist. Its situation
gave it rare advantages. Its latitude placed it in the mid-zone of
the teeming life that migrated annually between the high north and
the genial south, while its longitude placed it in a peculiarly rich
tract of that zone. The great lake on its eastern border served as a
broad blunt wedge which parted the migrating host into two great
divisions: on the one hand, the forest lovers who sought the wooded
regions of the northeast in summer and the like regions of the south¬
east in winter; on the other hand, the prairie lovers that preferred
the great open plains. Between these there was a middle zone and
a middle host formed in part of the overlap of the two other hosts,
but in part also of those species which distinctly preferred the border
tract of “openings,” the parks of interspersed prairie, meadow and
woodland, lying between the great forests and the great plains. The
southern and western part of Wisconsin was one of the most charm¬
ing sections of this great border tract of natural parks. Thru this
parkway there swept northward each spring and southward each fall
a mixed multitude of winged life that now, in its depleted state, seems
really incredible. The great woods of the north and northeast, with
Lake Superior in their rear, tended to shunt this host to the northwest
and caused congestion of their front. If I were to try to tell you
in specific terms of the richness and variety of life in springtime, as
I remember it, I fear you would feel impelled to call into service
the famous mot of Von Buch: “I am glad you saw that; for if I had
seen it, I would not have believed it.”
Out of the irresistible attractions of the native life of the air, the
woodlands, the grove-encircled prairies, the meadows, the marshes,
the limpid streams, and the charming lakes of Wisconsin, there grew
the first notable stage of spontaneous scientific activity, the stage of
the enthusiastic naturalist. It was quite in the natural order of
things that where personal conditions favored, as among surveyors
like Lapham and amony doctors of wide country practice like Hoy,
there should arise enthusiastic students of the rich fauna and the
flora of the region as also of the land that lay beneath and of the
sky that hung overhead. This stage of naturalistic enthusiasm
reached its climax somewhat before the general conditions in the
state were ripe for the founding of the Academy; and so the pioneer
naturalists of Wisconsin, particularly Lapham and Hoy, may be re¬
garded as the forefathers of the Academy quite as truly as its found¬
ers. Though the naturalistic stage had already somwhat declined
when the time for the inauguration of the Academy had come, it was
a very essential preliminary to the founding of the Academy.
696 Wisconsin Academy of Sciences, Arts, and Letters.
The Immediate Pre-Period of Preparation
The thirties, the forties, and the early fifties of the last century
were eminently pioneer days. With the sixties came the Civil War,
and with the mid-sixties, its close. It left the natural aftermath of
war, diverse currents and counter currents of thought and feeling
setting in devious directions — on the one hand, a desire for peace and
rest, for cessation of serious thought, for physical, mental and even
moral relaxation; on the other hand, when these first desires were in
some measure satisfied, a resumption of the tension that had become
habitual in the war, a new impulse to tenacious pursuit, a new will
to victory. The larger vision that came with the wider interests and
experiences of the war, visions of that which was national rather
than personal, entered into the new mental attitude. The man whose
pre-war thoughts had centered on his farm, his town, or his county,
had been forced to dwell on his state and his county at large and
he could not permanently shrink back to his former limitations of
interest. The man who had marched shoulder to shoulder with his
fellows could not well relapse into personal isolation. And so the
half decade following the war became the generative period of those
broader views and those generous instincts of coordination that led
to the organization of a common effort for the intellectual develop¬
ment of the state. This was the immediate pre-period of the found¬
ing of the Academy.
The Formal Founding of the Academy
During this half-decade, voluntary organizations were formed here
and there for the promotion of science and for personal culture, and
some futile efforts of a more general order were made, all of which
were more or less tributary to the coming general movement. En¬
couraged by these symptoms of readiness. Dr. J. W. Hoyt, Secretary
of the State Agricultural Society, worked out a comprehensive scheme
for a State Academy. He sent printed copies of this to such citizens
of the state as were thot to be interested in such a movement,
whether or not they were likely to be able to engage in research or
to make contributions to any phase of science, arts, or letters. He
also proposed that a convention be called to organize such an Acad¬
emy. The proposals met with a cordial response and a special call
for the proposed convention was issued bearing the signatures of 105
representative men of various callings and intellectual interests. In
explanation of my presence here today and my effort to serve you
as requested by your President, I may be permitted to say that my
name formed the tail end of the list, and that is perhaps why “the
rider of the pale horse” has thus far overlooked me in his frequent
and fateful visitations. If he shall continue to feel that the vanish¬
ing end of the long list is too immaterial to require any notice on his
part, his good judgment will meet with my most hearty concurrence.
Proceedings of the Academy.
697
The Convention met on February 16, 1870, and proceeded with
great unanimity to organize the Wisconsin Academy of Sciences, Arts,
and Letters. A constitution was adopted, officers elected, provision
made for incorporation, and for the other requirements of a new or¬
ganization. The Constitution provided for three Departments, em¬
bracing respectively the Sciences, the Arts, and Letters. Only the
first of these was organized at the initial meeting, but a fuller or¬
ganization was effected during the ensuing year.
The general purpose of the Academy was declared to be the encour¬
agement of investigation and the dissemination of correct views of
the various phases of Science, Literature and the Arts. The special
purposes of the Department of the Sciences were declared to be gen¬
eral scientific research, a progressive and thorough scientific survey
of the state under the direction of the officers of the Academy, the
formation of a scientific museum, and the diffusion of knowledge by
the publication of original contributions to science; that of the De¬
partment of the Arts to be the advancement of the useful arts through
the application of science and the encouragement of original inven¬
tion; the encouragement of the fine arts and the improvement of the
public taste by original contributions to art and by the formation of
an art museum; that of the Department of Letters to be the en¬
couragement of philological and historical research, the improvement
of the English Language, the collection and preservation of historic
records, and the formation of a general library.
Thus took place, fifty years ago, the formal founding of the Acad¬
emy.
The Substantial Establishment of the Academy
As already implied, this formal inauguration of the Academy rep¬
resented rather the ideals and aspirations of those who planned it,
and the good will and fond hopes of those who gave it countenance,
than a substantial banding together of real workers in science or
scholarship. It is unnecessary to say that the future of the Acad¬
emy as a vital working institution depended almost wholly upon the
persistent and sacrificial endeavors of men personally devoted to re¬
search and to culture. Scarcely a dozen of those who signed the call
for the convention were productive workers in any of the fields em¬
braced within the purposes of the Academy. The more comprehensive
clientele sought for the Academy at the outset was altogether laudable
and the sympathy and encouragement of this larger body was very
helpful, but I assume that you who now form the working members
of the Academy and are to hand it on to the next generation, care
most to learn who were the real leaders in giving working vitality
to the Academy in those earliest days, all the more so because certain
vital phases of this essential feature of the enterprise linger only in
vanishing impressions and fading memories and will soon be lost if
not now recorded.
698 Wisconsin Academy of Sciences , Arts, and Letters.
The important part played by Dr. Hoyt in planning so broadly and
in urging so successfully the initial steps, has already been indicated.
This service was recognized by choosing him first President of the
Academy. He was thus enabled to round out the formal organiza¬
tion of the Academy on the comprehensive plan adopted. He had the
merit of assiduity in calling into activity the latent as well as active
talent available in the state at the time. Though not a special worker
in any line of research, his intellectual sympathies were wide, his
aspirations were high; his dream for the Academy was ambitious.
The working nucleus of the Academy at the start was the group
of enthusiastic naturalists who had grown up under the stimulus of
the pioneer conditions. Among these I beg to include those who
studied the strata beneath and the sky above, as well as those de¬
voted to the plants and animals that tenanted the surface. Fore¬
most among these, by common consent, was Dr. I. A. Lapham of
Milwaukee, then already a veteran scientist. By profession a civil
engineer, he had become at an early day a faithful collector, ob¬
server and recorder of natural phenomena in nearly all leading lines
from bed-rock to sky. He was at once a botanist, a zoologist, an
archeologist, a geologist, and a meteorologist. He was a distinguished
example of the best order of the old school of all-around students of
natural science. Probably we owe to Dr. Lapham, more than to
any other single individual, the establishment of our Weather Serv¬
ice. He served as the first General Secretary of the Academy.
Scarcely less active and influential in giving vitality to the Acad¬
emy at the start was Dr. H. P. Hoy, of Racine, an intimate friend and
co-worker of Lapham’s in early naturalistic work. He had already
become a veteran student of birds, insects, and fishes, and was also
an enthusiastic collector of plants and of fossils from the ancient
crinoid fields of Racine. He was also an eager student of the relics
of aboriginal life. Lapham was quiet and modestly demonstrative,
but Dr. Hoy so bubbled over with enthusiasm that he easily set the
pace in demonstrative interest. He was chosen as the second Presi¬
dent of the Academy. Dr. J. G. Knapp, of Madison, was a frequent
contributor in several naturalistic lines, as was also Dr. Engelmann,
of Milwaukee, but the former soon moved from the state and the lat¬
ter was removed by death.
In the physical sciences, Dr. John E. Davies, of the State Univer¬
sity, was at first perhaps the leading contributor, with Dr. J. H.
Eaton, of Beloit, and Drs. R. Z. Mason and J. C. Foye, of Appleton,
as almost equally active co-workers. Dr. Eaton was perhaps the only
original member of the Academy who had any notable academic train¬
ing in technical research. A graduate of Amherst, he had won a
Fh. D. at Gottingen by his researches on the compounds of manganese.
In the field of political economy and social science, at the outset,
advancement was sought more by rational discussion than by rig¬
orous determinations of basal data; and so there was more general
participation in the discussions than in the more specific sciences.
The most active leaders were President A. L. Chapin, of Beloit,
Proceedings of the Academy.
699
(chosen third President of the Academy), President G. M. Steele, of
Appleton, Superintendent Samuel Fallows, the Reverend Charles Cav-
erno, Professor A. O. Wright, and later President John Bascom, Rev¬
erend Dr. Holland and others.
Though not active at the very outset. Dr. Wm. F. Allen, of the
State University, soon began a memorable series of papers replete
with specific historical research. These set a high standard of true
original investigation in humanistic lines. From his scholarly papers
some of us caught our first realizing sense of what constitutes orig¬
inal research in history.
Dr. Feuling, of the State University, was at the start a rather
lonesome leader in philological research but the fewness of workers
in this line was offset by the quality of the papers offered.
An attempt was made to give speculative philosophy a distinct place
in the work of the Academy under the leadership of Dr. S. H. Car¬
penter, of the University, but the effort scarcely survived his early
death.
Diversity and picturesqueness were given to the heavier parts of
the program by the sprightly literary contributions of the inimitable
Dr. Butler.
Earlier and Later Trends of the Academy
As already noted, the formal organization of the Academy was dis¬
tinctly broad, and there was a general desire and a definite effort to
preserve an appreciative and balanced attitude toward all phases of
research and of culture. None the less almost inevitably distinct
trends disclosed themselves almost from the start, and new trends
appeared in close succession, partly due to the new men that came
to the state, and partly to the development of young talent within it.
Of the papers presented during the first two years, 35 per cent re¬
lated to geological subjects, 23 per cent to biological, 17 per cent to
physical and mathematical science, 1.5 per cent to political and so¬
ciological subjects, and the remaining 10 per cent to historical and
philological subjects or to topics not readily classified. A distinct
geological trend at the outset is thus disclosed and the preponder¬
ance grew for a time. This special activity was due partly to char¬
ter members, particularly Lapham, Eaton and Chamberlin, but also,
in a quite notable degree, to the advent of Professor R. D. Irving,
who came to the state in the year following the founding of the
Academy. He came with excellent training and the advantage of
some field work, and at once took an active part in leading geological
inquiry along sound scientific lines. Irving was chosen fourth Presi¬
dent of the Academy. Two years later a systematic Geological Sur¬
vey was instituted by the state, largely through the influence of mem¬
bers of the Academy, and this not only gave unusual opportunities
for productiveness in this line, but helped to develop young talent that
made itself felt in the later activities of the Academy.
700 Wisconsin Academy of Sciences , Arts , and Letters.
Soon after the founding of the Academy, the great movement to¬
ward a higher order of things in agricultural science and practice
began and at first was most definitely represented by the chemical
work of Professor W. W. Daniells. The developments in Agirculture
were more closely connected with the State Agricultural Society and
particularly with the State University than with the Academy, but the
Academy claims some little merit for this most signal development.
About the same time also Major Nicodemus and Captain Nader took
the lead in developing interest in engineering themes by notable and
stimulating discussions.
There has been occasion to lay emphasis on the type of study of
plants and animals most familiarly known as Natural History, which
prevailed at the founding of the Academy and in the preceding pioneer
stage. The career of the Academy was scarcely more than under
way before this began to give place to modern biological inquiries and
this led on to those important ecological and other studies that char¬
acterized the later official surveys and that mean so much to the in¬
tellectual and material welfare of the people of the state. This was
perhaps the most notable change of trend in the intimate work of
the Academy. It was led by a young man who came to the state in
the fifth year of the Academy, and who has given the Academy one
of its most prolonged and valued series of papers. Then a young man,
we now delight to honor and revere him as President at once of the
Academy and of our State University, President Birge. The syste¬
matic and ecological phases of this modern departure were a little
later admirably illustrated by the important contributions of Profes¬
sor and Mrs. Peckham.
By the end of the first decade of the Academy’s life, it had under¬
gone further changes and had taken on much more distinct diver¬
sity. It thus began the better to represent the varied intellectual
development which the state was rapidly coming to enjoy, and which
it has more fully realized in these later years.
By the end of the second decade the divergencies toward the later
phases of the Academy became still more marked. The distinctions
of departments, that were rather formally defined at the outset, be¬
gan to fade away, though the departments themselves grew more di¬
vergent. A more cosmopolitan spirit arose which made less of sub¬
jects and more of method and real intellectual advance. The forma¬
tive period was being merged into what now seems to a founder
“the Golden Era” of the Academy. Doubtless intrinsically, it was no
better than later stages — perhaps not so good — but these are the days
of relativity and to one who felt the struggle and the weakness of
the start, it seemed golden.
With it there came rapid changes in the personnel. The veteran
naturalists passed away and other losses were many and grave. But
the chief changes came from two other sources. The educational in¬
stitutions of the state were rapidly developing in research lines and
there came to the state many able men, well equipped and productive.
It would be easy to begin the list — for there was Trelease and the
Proceedings of the Academy.
701
lamented Barnes — and to go on at length, but where could I end it?
Besides it is not my function to deploy the Golden Age of the Acad¬
emy, but merely its founding. The other source of change came even
closer to the hearts of the founders, the coming of choice youth of
the state into productive membership in the Academy, the children
of the Academy. They were equally and perhaps more the children
of the educational institutions of the state, but we claim them as
children of the Academy none the less. Very notable among these
was President Van Hise, who rapidly rose to leadership in the state,
in the nation, and beyond. It would be a delight to name many
others, but how could the parental affection of a founder permit him
to stop short of naming all the children of the Academy? The di¬
lemma is in itself evidence that the formative stage of the Academy
had already passed away. The founding of the Academy had really
taken place.
At the close of the address Professor Chamberlin was pre¬
sented to President Birge by Professor Charles K. Leith for the
degree of Doctor of Science. President Birge conferred the
degree in the following words :
Dr. Chamberlin:
I assume today the privilege of adding to the official formula which
it is at once my pleasure and my duty to pronounce; and my words
are of a personal character rather than a supplement to the review
of your scientific achievements so well set forth by Dr. Leith.
I can not forget that our friendship began at a meeting of the
Wisconsin Academy of Sciences, Arts, and Letters, something more
than forty-four years ago; nor will either of us ever forget the friend
through whom we were brought together — the lamented Irving. You
were then a young professor in Beloit College, just entering on your
first great scientific task — the direction of the Wisconsin Geological
Survey; I was a still unfledged instructor, beginning my teaching at
the University.
Through all the years that have passed since 1876, my life and
my work have lain close to yours, as a teacher and student in a
cognate department of a neighboring college, and, not least, during
five happy years, as a member of the faculty over which you presided.
I have therefore been so placed that I have not only known the long
record of your labor and success in the fields of education and
science; I have seen and I have in some degree shared the influences
which for so many years came to this state through your presence
here as a living and working personality.
And when the central place of your work was removed from our
university to its next great neighbor, when your scientific outlook en¬
larged so as to include not only the earth but the heavens, I could
still appreciate, perhaps more justly than younger men could do, the
702 Wisconsin Academy of Sciences , Arts , and Letters.
increasing fruit of your labors for our state and our university and
the continued presence with us of your personality.
In you, Sir, the Wisconsin Academy recognizes eminence and dis¬
tinction in science; we take pride in the achievements of a fellow cit¬
izen; we are grateful to him for increased knowledge, for a wider vis¬
ion, for a deeper insight into nature. But today and on this occasion
there is something more: We look back on a half-century of your
life among us; we see not only new fields won for science during
those years; we see also in you constantly present with us the spirit
and temper of science; and as you spoke to us just now we heard
not merely, or indeed chiefly, the record of scientific work told us
by one of the workers, we hear rather the voice of one through whom
science had for a half-century been exerting a vital influence among
us; we felt, as we have felt so often before, the presence of the man
and the influence of his life.
Such thoughts and such feelings were present when the Wisconsin
Academy proposed that some special recognition be made of the com¬
pletion of your half-century in the Academy which you helped to
found. With this sense of personal gratitude the University acted —
the University so greatly advanced and enriched by you.
It is therefore with a peculiar pleasure that I now comply with the
request of the Wisconsin Academy of Sciences, Arts, and Letters, with
the recommendation of the faculty of the University of Wisconsin,
and the vote of its regents. By the authority committed to me, I
confer upon you the degree of Doctor of Science, and in testimony
thereof, I present you with this diploma.
Second Session, 2:30 P. M.
University Armory.
This session was also a University Convocation, which was
addressed by President Nicholas Murray Butler, of Columbia
University, New York City.
Third Session, 4:00 P. M.
Room 301, Biology Building.
President Birge took the chair and introduced Professor John
M. Coulter, of the University of Chicago, who spoke on the re¬
lations of the local Academy to the American Association for
the Advancement of Science.
Professor Charles E. Allen, of the University of Wisconsin,
reported on the plan for the affiliation of the local Academies
Proceedings of the Academy.
703
and the American Association for the Advancement of Science,
as formulated at the St. Louis meeting of the American Asso¬
ciation.
Considerable discussion followed these addresses, and Profes¬
sor Allen made clear the following features of the proposed affili¬
ation :
1. The American Association for the Advancement of Science is
ready to enter into an agreement with the Wisconsin Academy of
Sciences, Arts, and Letters, whereby membership in both organiza¬
tions may be held by a payment of five dollars a year, subject to
certain conditions.
2. Those who are now members of both organizations will pay $5.00
to the Wisconsin Academy, $4.00 of which will be transmitted to the
American Association.
3. Members of both organizations, and members of the American
Association who joined the Academy in 1920, and who paid $5.00 to
the American Association, will receive membership in the Wisconsin
Academy by the return of $1.00 to the Academy by the American
Association.
4. Present members of the American Association who join the
Wisconsin Academy after 1920, and those who after 1920 join first
the American Association, paying $5.00 to the American Association
before April 1st, must pay an additional $1.00 if they wish to be¬
come members of the Wisconsin Academy for that year. Thereafter
they pay $5.00 to the Wisconsin Academy, $4.00 of which is sent to
the American Association.
5. Persons of class 4 who have not paid $5.00 before April 1st of
their first year of membership may pay $5.00 to the Wisconsin Acad¬
emy. The reason for this rule is that such persons are regarded as
delinquent, and encouragement is given to the Academy to collect the
dues.
It was moved by Professor Allen that affiliation with the Amer¬
ican Association for the Advancement of Science be effected on
the terms outlined. This motion was put, and was carried unani¬
mously.
The Secretary presented the name of Mr. George B. Merrick
of Madison as Honorary Member of the Academy. President
Birge seconded the motion of the Secretary in these terms:
We who have lived in Madison a long time know Mr. Merrick very
well. We have known him as a fellow citizen engaged in the work
of the University, and as one who in his younger days was pilot on
the Mississippi, and who has collected an immense amount of in¬
formation in those early days when the steamboat was the means of
travel up and down the Mississippi Valley, and has published consider¬
able on that subject. One ought not to omit to say that his health
704 Wisconsin Academy of Sciences, Arts, and Letters.
has failed; it is not likely that this honor which we are proposing to
confer upon him can be long enjoyed by him, and if I may as chair¬
man second the motion of the secretary, I will do so.
The motion was carried unanimously.
The Secretary then presented the report of the committee on
membership, proposing the following new members ;
Robert Thomas Aitken, Milwaukee
William E. Alderman, Madison
Edward William Blakeman, Madison
Harold William Browning, Madison
Harry E. Cole, Baraboo
Ezra Jacob Kraus, Madison
Herman W. Kunz, Milwaukee
E. W. Lindstrom, Madison
Mrs. Angie Kumlien Main, Fort Atkinson
Harold William Rickett, Madison
Gilbert E. Seaman, Milwaukee
E. J. B. Schubring, Madison
Selma L. Schubring, Madison
Warner Taylor, Madison
Mrs. E. H. Van Ostrand, Madison
Edward Strong Worcester, Madison
On motion the Secretary was instructed to cast the ballot in
their favor.
The Secretary then presented his report for the year 1919;
Report of the Secretary for the Year 1919
Honorary Members . 6
Life Members . 10
Corresponding Members . 39
Active Members . 244
Total . 299
Changes since last report:
Active Members reported for 1918 . . . 210
New Members enrolled in 1919 . . . 46
256
Deaths . . . . . 4
Resignations . . . 5
Dropped for nonpayment of dues . 3
_ 12
Present Active Membership . . 244
New Memberships acted on at this meeting . . 16
Proceedings of the Academy.
705
I regret to have to report the loss of four Active Members by death:
Mr. D. P. Blackstone, Mr. C. E. McLenegan, Mr. John Nader, and Mr,
Wm. Finger. Arthur Beatty,
Secretary.
It was moved that the Secretary’s report be adopted. Car¬
ried.
The Secretary then read acknowledgments of invitations and
letters of congratulations from numerous individuals and from
the following institutions.
The Academy of Natural Sciences of Philadelphia, represented by
Professor J. M. Coulter, of the University of Chicago.
The American Academy of Sciences, Boston, represented by Profes¬
sor Edwin B. Frost, of the University of Chicago.
The American Museum of Natural History, New York, represented
by Professor A. S. Pearse, of the University of Wisconsin.
The American Philosophical Society, Philadelphia.
The Boston Society of Natural History.
The Buffalo Society of Natural Sciences.
The California Academy of Sciences.
The Connecticut Academy of Arts and Sciences, New Haven.
The Elisha Mitchell Scientific Society, Chapel Hill, N. C.
The Illinois Academy of Science, Chicago, represented by Profes¬
sor William Trealease, of the University of Illinois.
The Iowa Academy of Sciences, Des Moines.
The Kansas Academy of Science, Lawrence.
The National Academy of Sciences, Washington, represented by
Professor Thomas C. Chamberlain, of the University of Chicago.
The National Research Council, Washington.
The New York Academy of Sciences, New York, represented by
Professor Armin K. Lobeck, of the University of Wisconsin.
The Ohio Academy of Science, Delaware.
The Royal Canadian Institute, Toronto.
The Washington Academy of Sciences, Washington.
The Secretary presented a letter of congratulations and a set
of resolutions on the part of the Wisconsin Archeological Soci¬
ety, which read as follows:
Milwaukee, Wisconsin,
Prof. Arthur Beatty, April 19th, 1920.
Secretary, Wisconsin Academy of Sciences, Arts & Letters,
Madison, Wis.
My dear Professor Beatty:
As secretary of the Wisconsin Archeological Society I take great
pleasure in forwarding to you herewith the resolutions adopted by
this Society in honor of the fiftieth anniversary of the founding of
the Wisconsin Academy of Sciences, Arts, and Letters.
45—8. A. L.
706 Wisconsin Academy of Sciences, Arts, and Letters.
We trust that these resolutions will, in some measure at least, ex¬
press to your Academy the high regard in which we hold it, and our
hopes for its future.
Very sincerely yours,
Charles E. Brown,
Sec'y, Wisconsin Archeological Society.
Resolutions Adopted on the 50th Anniversary of the Wisconsin
Academy of Sciences, Arts and Letters
The Wisconsin Archeological Society in meeting assembled at the
Milwaukee Public Museum, Milwaukee, on Monday evening, April 19,
1920, adopts the following resolutions:
Whereas the officers and members of The Wisconsin Archeological
Society are aware that the Wisconsin Academy of Sciences, Arts and
Letters, with which it has been closely identified in joint annual meet¬
ings for some years past will on Friday, April 23rd celebrate at Madi¬
son with an appropriate commemorative programme, the Fiftieth
Anniversary of its foundation;
Therefore be it Resolved: That The Wisconsin Archeological So¬
ciety extends to the Wisconsin Academy of Sciences, Arts and Letters,
its officers and members, most hearty congratulations on this, its
Golden Anniversary, and with these the most sincere hope that its
great usefulness in its chosen fields of research and publication may
continue for many years to come to the honor and glory of our state.
And be it further Resolved: That as an added evidence of our
appreciation of this Anniversary there be, and they are hereby elected
to honorary membership in The Wisconsin Archeological Society, Ed¬
ward A. Birge, president and Arthur Beatty, secretary of the Wis¬
consin Academy of Sciences, Arts and Letters.
Lee R. Whitney,
President.
Charles E. Brown,
Secretary.
S. A. Barrett,
Charles G. Schoewe.
William H. Vogel,
E. J. W. Notz,
Directors.
Milwaukee, Wisconsin,
April 19, 1920.
The Treasurer presented his report for the year as follows:
April 23, 1920.
Proceedings of the Academy .
707
Report of the Treasurer of the Wisconsin Academy of Sciences,
Arts, and Letters for the Year 1919.
Receipts
Received from Dues and Initiations . $290.00
Received from Sale of Transactions . 22.54
Received from Interest on Bonds, April 1, 1919 . 140.00
Received from Interest on Bonds, April 1, 1920 . 123.00
Received from Interest on Liberty Bonds . 25.13
Received from 3 Bonds matured, April 1, 1919 . 300.00
Received from 1 Bond matured, April 1, 1920 . 100.00
Balance on hand, March 20, 1919 . 262.33
1, 263.00
Disbursements
Secretary-Treasurer’s Allowance . 200.00
Safety Deposit Box Rent . 3.00
Stenographic Work, etc . 7.05
Victory Bonds purchased . 300.00
Certificates of Deposit . 600.00
1, 110.25
Balance on hand, April 21, 1920 . 152.95
$1, 263.00
Arthur Beatty,
Treasurer.
On motion this report was adopted.
The auditing committee was called upon to report, and cer¬
tified as follows:
April 22, 1920.
We have examined the accounts of the Treasurer, and find them
correct; we have also examined the securities belonging to the Acad¬
emy, and find them to be as follows:
Madison Street Improvement Bonds . $2100.00 (par)
Third Liberty Loan . 400.00 (par)
Victory Loan . 300.00 (par)
Certificates of Deposit . 600.00
Total . $3400.00
(Signed)
J. J. Davis
George Wagner
On motion duly seconded the Auditors’ report was adopted.
The meeting then adjourned until 7 :00 o ’clock.
708 Wisconsin Academy of Sciences, Arts, and Letters.
Fourth Session, 7:00 P. M.
The University Club.
This, the closing session of the meeting, took the form of a
commemorative dinner, at which 125 guests were present, Presi¬
dent Birge presiding as toastmaster.
Each guest was presented with a reprint of the first Bulletin
of the Academy, which was issued in April, 1870.
The general topic of the session was the Relation of the Acad¬
emies to Research; and the speakers were
Bishop Samuel Fallows, of the Reformed Episcopal Church, Chi¬
cago;
President Melvin A. Brannon, Beloit College, Beloit;
Professor William Trelease, University of Illinois, Urbana;
Superintendent Joseph Schafer, Wisconsin State Historical Society,
Madison.
In his opening remarks the President announced that the
Academy would have two memorials to mark this turning point
in its history. First, a memorial volume is provided for by the lib¬
erality of the state. At the last meeting of the State Legisla¬
ture a special appropriation of $2000.00 was made, to enable
the Academy to publish a volume of transactions which would
be more worthy of the occasion than those which we are ordi¬
narily able to issue.
The second memorial is a medallion to be struck in bronze,
which commemorates fifty years of science in the state. A full
account of this will be found on pages 711 to 716 of this volume.
The President then introduced Bishop Fallows, who warned
against thing-mindedness, which is a besetting vice of the age,
and advocated person-mindedness as a solution of the difficul¬
ties into which the society of today has been brought by the too
great emphasis which has been placed upon mere things.
President Brannon, of Beloit College was next introduced, and
spoke on the great importance of research, using as illustrations
the work of the Wisconsin Academy in the last fifty years.
Professor Trealease, of the University of Illinois, was next
introduced by the President, and he spoke of his former connec¬
tion with the University of Wisconsin, furnishing some delight¬
ful reminiscences of the town and university of the year 1883,
He then proceeded to speak of the function of the Academy in
Proceedings of the Academy.
709
encouraging research by the student who is unknown and in
developing him ; in encouraging a love of knowledge for its own
sake ; of developing a knowledge of nature, and so keeping peo¬
ple in touch with the things about them. In this way comes a
mastery of nature. Of the scientist it may he said that getting
down and putting other people in touch with nature is the surest
way of making himself a master of nature.
Superintendent Schafer, of the Historical Society, dwelt on
the function of encouraging the amateur in research and of con¬
serving knowledge by publication. In these two things the
Academy had well justified its existence during the last fifty
years.
President Birge concluded the programme, declaring the meet¬
ing adjourned:
“I think we shall all agree that our speakers have stuck pretty close
to the text assigned them, and that they have presented us with vari¬
ous illuminating instances of the relations of the Academy to Re¬
search.
The hour is late, and I am not going to add anything. Yet there
is one thing which I should like to say, one note in some of the
speeches which we have listened to, to which I should like to recur.
We have been warned here against thing-mindedness, and I agree
in the necessity and in the value of the warning. Yet I feel that the
attitude which we of the Academy hold toward the world in which
we are to work during the next half century on which we enter, that
that attitude is of no small importance, and I feel that we may easily
over-rate the thing-mindedness, the commercialism of the world at
the present day; not perhaps in a sense that the world is not so
commercial as we think it is perhaps; and yet when we feel that
the world of today is peculiarly indifferent to our motives don’t we
make a mistake? Of whom do we seek when we desire to get that
expression? We go back to Wordsworth; “The world is too much
with us”. Well, that was written nearly one hundred and twenty
years ago, and it was written I think at a breathing spell in one of
the great heroic periods of English history, at the pause in the great
Napoleonic warfare, just before the struggle of a dozen years which
terminated at Waterloo. The world was too much with them.
Wordsworth was right. And yet after all as we look back upon that
period we say “Those were heroic days”, and we contrast them with
the commercialism of our own day. Yet how will people look back
upon our own day? How will people look back upon the history of
the last five years? Will they look back upon those years as years
of commercialism, or as years of heroism? We have seen in the far
east — not the technical far east, but the far east of Europe — not
individuals, but nations, not highly educated, but people of every
grade, by the million, ready to die the most cruel deaths in behalf
710 Wisconsin Academy of Sciences, Arts, and Letters.
of their religion. And on the other side of that continent we have
seen nations willing to suffer in the same way for the sake of na¬
tionality. Is that a world in which commercialism is rampant, in
which thing-mindedness dominates the world? Is that a world in
which things are in the saddle, or are we living today in a world in
which ideas may be in the saddle? And if the ideas which we rep¬
resent are not in control is the fault in our system, or is it in our¬
selves?
With that spirit and with that temper I would go forward into the
new era, the new life of the Academy of Sciences, Arts and Letters.
And so we close our celebration, with greetings, with thanks to the
representatives of other learned bodies, to the members of our Acad¬
emy, who have come in to us from more distant parts of the state,
and to all members and friends and guests of the Wisconsin Acad¬
emy.”
Fiftieth Anniversary Exhibit.
In connection with the fiftieth anniversary celebration of the
Wisconsin Academy of Sciences, Arts and Letters an extensive
special exhibit was made by the State Historical Museum. This
exhibit occupied the greater part of its south exhibition hall,
filling ten table cases. It consisted of a large number of photo¬
graphs of early officers and prominent members, copies of its
early and recent publications, programs, and announcements of
past meetings, the minutes and scrapbooks of the early secre¬
taries, copies of early correspondence and other materials con¬
nected with the history of the Academy. Some of these were
kindly loaned by the University and State Historical libraries.
A photograph of the medal struck in commemoration of the an¬
niversary occupied a prominent place in the exhibit.
In several cases were shown the fine collection of medals
awarded to the University of Wisconsin during the past twenty-
five years of its history, and of the more important publications
of the Wisconsin Archeological Society and Wisconsin Natural
History Society two state organizations which for many years
have cooperated with the Academy in its meetings held in Madi¬
son and Milwaukee.
The exhibit continued in place for several weeks and was
greatly appreciated by many members who visited the Museum
for the purpose of seeing it.
The exhibit was inaugurated and arranged by Mr. Charles E.
Brown, Curator of the Academy and of the State Historical So¬
ciety. Arthur Beatty,
Secretary .
TRANS. WIS. ACAD. VOL. XX
PLATE LXV
ACADEMY MEDALLION 1870-1920
THE MEDALLION OF THE ACADEMY, 1870-1920
E. A. Birge.
The Wisconsin Academy has commemorated its Semi-centen¬
nial by a medallion bearing the portraits of six representatives of
its distinguished members. The medallion was made possible by
friends who provided the funds necessary for designing it and
sinking the dies.
The artist is Leonard Crunelle, the sculptor, of Chicago. He
has succeeded in the difficult task of furnishing a permanent
memorial of the semi-centennial in a form which unites great
historical value with high artistic excellence. The medallion is
72 mm. high and 47 mm. wide. The obverse bears the figure of
Minerva tending the lamp of learning and a motto from Lucre¬
tius, Naturae species ratioque; the reverse carries the inscription,
The Wisconsin Academy of Sciences, Arts and Letters, 1870~
1920 , and the portraits of the six members, namely:
William Francis Allen, Historian
Thomas Chrowder Chamberlin, Geologist
Philo Romayne Hoy, Physician, Naturalist
Roland Duer Irving, Geologist
Increase Allen Lapham, Naturalist, Geologist
George Williams Peckham, Zoologist
These men were selected as representative members of the
Academy, out of a much longer list of persons who are well
worthy of mention, as may be learned from the memorial address
of Dr. Chamberlin (pp. 000-000). They are chosen in part for
their intellectual eminence and for their services to the Acad¬
emy, and in part for the periods in which their lives and activi¬
ties fall. Three of them, Chamberlin, Hoy, and Lapham, were
charter members of the Academy, and the others became mem¬
bers very early in its history. Each of them served as president
of the Academy, except Lapham, who was its secretary from the
711
712 Wisconsin Academy of Sciences, Arts, and Letters.
beginning until his death in 1875. They fittingly represent three
groups of the members of the Academy and three eras in its
history : those of the pioneer days, those who helped to establish
modern learning in the State, and those who have contributed
largely to the advance of science in its more recent phases nearly
or quite to the close of the first half-century of the Academy’s
life.
Dr. Hoy and Dr. Lapham represent the pioneer period of
science in the State. Dr. Lapham, the surveyor, was also botan¬
ist, meteorologist, and geologist; Dr. Hoy, the busy country
physician, was also a naturalist, especially on the side of zoology.
Professor Allen and Professor Irving, in turn, were among the
first men to bring modern scholarship to the state, Professor
Allen in the departments of Latin and history and Professor
Irving in geology. Both were members of the faculty of the
University of Wisconsin, and the work of each was cut short
by an early death so that their influence was limited to the first
twenty years of the history of the Academy. Finally Dr. Peck-
ham as zoologist and Dr. Chamberlin as geologist represent the
men whose large contributions to science have continued nearly
or quite to the present time.
These six members of the Academy may be grouped in still
another way : Three — Allen, Chamberlin, and Irving — were
members of the university and their great contributions to
knowledge are in the field of their professional work; the other
three — Hoy, Lapham, and Peckham — reached large results by
giving to science the free hours of lives whose principal duties
were in other lines.
Such are some of the considerations which underlie the selec¬
tion of the men chosen to represent the Academy during fifty
years of its life. A brief sketch is given of each of them and
of his relations to the Academy.
WILLIAM FRANCIS ALLEN, 1830-1889.
Professor Allen was born in Massachusetts; be graduated from
Harvard in 1851 and studied in Europe, 1854-1856. He served with
the Freedmen’s Aid Commission and the Sanitary Commission during
the Civil War. He came to the University of Wisconsin in 1867 as
professor of ancient languages and history, at the time of the reor¬
ganization of the University under President Chadbourne. His title
was changed to professor of Latin and history in 1870; he devoted
Birge — The Medallion of the Academy.
713
more and more time to the latter subject until 1886, when he was
wholly relieved from teaching Latin and became professor of history.
His death in 1889 cut short his labors in that department.
Professor Allen was at once a great teacher and a great scholar.
He represented the humanities in the faculty of the university and
in the state with a fullness and power beyond any other member. He
was a recognized authority in the fields of Latin and history.
He joined the Academy in its first year; all of his numerous con¬
tributions to its Transactions are concurred in with history and are
found in the first seven volumes of the Transactions. At the time of
his death he was president of the Academy, having been elected in
1887 for the three-year term.
THOMAS CHROWDER CHAMBERLIN, 1845-
Dr. Chamberlin was born in Illinois and graduated from Beloit
College in 1866. He was professor of geology at Beloit, 1873-1882,
and assistant state geologist of the Wisconsin Geological Survey,
1873-1876. He was director of that Survey, 187 6-1882, and was re¬
sponsible for its very excellent report. Turning more especially to
glacial geology, he has contributed greatly to the knowledge and in¬
terpretation of the phenomena of the glacial period. From 1882 to
1887, he was in charge of the Glacial Division of the United States
Geological Survey, and from 1887 to 1892 he was president of the
University of Wisconsin. In 1892 he went to the newly-established
University of Chicago as head of the department of geology and re¬
mained in that position until he retired as emeritus professor in 1919.
For the past twenty years his chief scientific work has been on
fundamental geological problems connected with the origin and de¬
velopment of the earth and the solar system, and he is still actively
engaged in these studies. The contributions to science made during
this half century, so largely given to fruitful research, brought him
many academic and scientific honors.
Dr. Chamberlin was a charter member of the Wisconsin Academy
in 1870; he was the principal speaker at its semi-centennial in 1920
(see p. 000). He was president of the Academy, 1884—1887, and was
a constant contributor to its meetings during his stay in Wisconsin.
Several geological papers from him are found in its earlier volumes,
the latest in Vol. VIII, but his principal geological work has been pub¬
lished elsewhere.
PHILO ROMAYNE HOY, 1816-1892.
Dr. Hoy was born of Revolutionary ancestry in Ohio and his early
life was spent among pioneer surroundings. He graduated from the
Ohio Medical College in Cincinnati in 1841. In 1846 he came to Ra¬
cine and there he remained for the rest of his life, engaged in active
practice as a physician.
714 Wisconsin Academy of Sciences , Arts, and Letters.
In the first year of his residence at Racine, Dr. Roy made the
acquaintance of Dr. Lapham, and the two men remained devoted
friends and fellow students of nature until Dr. Lapham’s death nearly
thirty years later. Dr. Hoy was interested in every side of natural
history but gave most attention to animal life. His collection of
birds, insects, fishes, etc., was very large and yielded important in¬
formation regarding the fauna of the newly-settled region. He was
also the first to pay special attention to the biology of Lake Michigan
and its fish fauna.
Like Dr. Lapham, Dr. Hoy gave himself to advancing natural
science in Wisconsin at a time when very few members of a frontier
population were occupied with anything outside of the labors and the
profits of a pioneer life. These two men are in a peculiar sense rep¬
resentatives of those few who initiated the study of nature in the
state and carried it on with a devotion and enthusiasm that made
possible the organization of science in the Academy and a little later
in the Geological Survey.
Dr. Hoy was a charter member of the Academy and was its presi¬
dent, 1875-78. He contributed numerous short papers, mainly on
zoological subjects, to the earlier volumes of the Transactions.
ROLAND DUEIR IRVING, 1847-1888.
Professor Irving was a native of New York City, a graduate of
Columbia College and of its School of Mines. He was appointed
professor of geology in the University of Wisconsin in 1870, when he
was twenty-three years old, and he held this position until his death.
In 1873 he became assistant geologist of the State Geological Survey.
In this position he contributed much to the knowledge of the diffi¬
cult geology of the Lake Superior region and he also introduced the
then new methods of microscopic petrology. After the close of his
field work on the State Survey in 1880 he continued work in the Lake
Superior district as a member of the United States Geological Survey,
and became the first authority in. the country on the intricate prob¬
lems of pre-Cambrian geology.
Professor Irving was as definitely the leader of the university on
the side of science as was Professor Allen on that of the humanities.
His capacity for investigation made him the first member of the fac¬
ulty of the University of Wisconsin to attain a position among Amer¬
ica’s leading men of science.
He entered the Academy in 1870 and was its president, 1881-84.
Several papers by him are found in the early volumes of the Transac¬
tions; but, as is the case with Dr. Chamberlain, most of his geological
work was published in the reports of the Wisconsin Geological Survey
and the United States Geological Survey.
Birge — The Medallion of the Academy.
715
INCREASE ALLEN LAPHAM, 1811-1875.
Dr. Lapham was born at Palmyra, New York. His father was a
civil engineer and he followed the same profession. From boyhood
he had a thirst for science and he had a singular capacity for the
study of nature. At the age of sixteen he attracted the attention of
Professor Benjamin Silliman, of Yale, and thus began a correspond¬
ence with scientists far and near which continued with increasing vol¬
ume to the end of his life.
He came to the then frontier town of Milwaukee in 1836 at the
invitation of Byron Kilbourn. He there exercised his profession but
gave much of his time to scientific studies. He spent much of his
energy in collecting and cataloguing the plants and the shells of his
new home. He surveyed its Indian mounds; he investigated its geol¬
ogy and collected its fossils; and in short he eagerly devoted him¬
self to the development of the knowledge of nature, and so became
in his generation the first scientific scholar of Wisconsin. He was
state geologist, 1873—5, but his death came before he could take a
large part in the work of the Survey. After his death the state pur¬
chased his scientific collections for the University. The geological
specimens were destoryed by fire in 1884, but the collection of plants
forms the nucleus of the University herbarium.
Dr. Lapham was a charter member of the Academy and its secre¬
tary from its foundation to his death. He contributed several short
papers on natural history and geology to the first two volumes of
the Transactions.
GEORGE WILLIAMS PECKHAM, 1845-1914.
Dr. Peckham was born in Albany, New York, but his life from
boyhood on was spent in Milwaukee. He was in the United States
Army during the Civil War, being mustered out in 1865 with the
rank of First Lieutenant of Artillery. He graduated from the Uni¬
versity of Michigan with the degree of doctor of medicine in 1873.
He was teacher of biology in the Milwaukee High School, then prin¬
cipal of the High School, Superintendent of City Schools, 1890-96,
and after 1896 was head of the Milwaukee Public Library. Along
with the duties of these positions he was always actively engaged in
scientific work, chiefly on the habits and classification of insects.
These zoological studies were carried on jointly with his wife, Eliza¬
beth G. Peckham, and their results, when published, always bear both
names. The instincts and habits of wasps and of spiders and the
classification of spiders were their two departments of special re¬
search, and in both fields their work bears an international reputa¬
tion.
Dr. Peckham joined the Academy in 1877 and was its president,
1890-93. He was one of the largest as well as one of the most valued
contributors to its Transactions. His first paper is published in Yol-
716 Wisconsin Academy of Sciences, Arts, and Letters.
ume VI, on the temperature of Pine Lake. He contributed eight
papers on the classification and habits of spiders, chiefly from the
family Attidae. These appeared between 1885 and 1909, and aggre¬
gate about 800 pages with nearly 50 plates.
The persons who subscribed to the fiftieth anniversary fund
of about $1,200 and thus provided the medallion are the follow¬
ing:
Dr. EL A. Birge, Madison,
Hon. T. E. Brittingham, Madison,
Hon. F. P. Hixon, La Crosse,
Hon. A. J. Horlick, Racine,
Dr. Charles K. Leith, Madison,
Hon. Frank A. Logan, Chicago,
Mrs. Charles W. Morris, Milwaukee,
Dr. M. S. Slaughter, Madison,
Dr. C. S. Slichter, Madison.
The Academy recognizes a peculiar debt of gratitude to these
donors who have enabled it to put forth so beautiful and signifi¬
cant a memorial. The reverse of the medallion displays in word
and symbol the spirit of the Academy; the obverse commem¬
orates the first half century of its life and also preserves in en¬
during form the features of six citizens of the state and mem¬
bers of the Academy who during that time contributed so much
to the advancement and development of knowledge. The Acad¬
emy has placed these donors on its list of life members in recog¬
nition of the service thus rendered.
The Academy has also received gifts in the form of subscrip¬
tions for copies of the medallion, so that it might be more wisely
distributed. About 50 members and friends of the Academy
subscribed for one copy each and several subscribed for more
than one, the additional copies being sent to institutions.
Among the latter are to be named Miss Katharine Allen (5
copies), Dr. E. B. Van Vleek (5 copies), and Dr. J. J. Davis
(2 copies), all of Madison; and Mr. A. C. Clas (2 copies), of
Milwaukee. In this connection also especial mention must be
made of Hon. A. J. Horlick, of Bacine, who added largely to his
original gift so that this memorial of Wisconsin history might
become more generally known. These gifts have made it pos¬
sible for the Academy to place many copies of the medallion in
Wisconsin institutions and also to send it out to a greater num¬
ber of learned societies.
LIST OF OFFICERS AND MEMBERS
CORRECTED TO JURY 1, 1931.
Officers.
President , Melvin A. Brannon, Beloit.
Vice-President , Sciences , Samuel A. Barrett, Milwaukee.
Vice-President, Arts, Grant Showerman, Madison.
Vice-President, Letters, Karl Young, Madison.
Secretary , Chancey Juday, Madison.
Treasurer , Chancey Juday, Madison.
Curator, C. E. Brown, Madison.
Librarian, Walter M. Smith, Madison.
Committee on Publication.
The President, ex officio,
The Secretary , ex officio,
W. E. Tottingham, Madison.
Council.
The President, Vice-Presidents, Secretary, Treasurer, Librarian
and Past Presidents retaining their residence in Wisconsin.
Committee of Library.
The Librarian, ex officio,
George Wagner, Madison.
W. H. Barber, Ripon.
A. A. Trever, Appleton.
Committee on Membership.
The Secretary, ex officio,
P. W. Boutwell, Beloit.
V. E. McCaskill, Superior.
H. H. Smith, Milwaukee.
G. S. Bryan, Madison.
717
718
Wisconsin Academy of Sciences, Arts, and Letters.
Past Presidents.
Honorable John W. Hoyt, M.D., LL.D.,1 Washington, D. C.,
1870-75.
Dr. P. R. Hoy, M.D.,1 1876-78.
President A. L. Chapin, D.D.,1 1879-81.
Professor Ronald D. Irving, Ph.D.,1 1882-84.
Professor Thomas C. Chamberlin, Ph.D., Sc.D., LL.D., Chicago,
Ill., 1885-87.
Professor William F. Allen,2 1888-89.
Professor Edward A. Birge, Ph.D., Sc.D., LL.D., Madison,
1889-90.
Librarian George W. Peckham, LL.D., Milwaukee, 1891-93.1
President Charles R. Van Hise, Ph.D., LL.D., Madison, 1894-96.
Professor C. Dwight Marsh, A.M., Ph.D., Washington, D. C.,
1897-99.
Professor Charles S. Slichter, M.S., Madison, 1900-1902.
Dr. John J. Davis, M.D., Racine, 1903-1905.
Professor Louis Kahlenberg, Ph.D., Madison, 1906-1909.
President Samuel Plantz, Ph.D., D.D., LL.D., Lawrence College,
Appleton, 1910-1912.
Professor Dana C. Munro, A.B., A.M., Princeton, New Jersey,
1913-1915.
Director Henry L. Ward, Milwaukee, 1915-1918.
President Edward A. Birge, A.B., A.M., Ph.D., Sc.D., LL.D.,
Madison, 1918-1921.
HONORARY MEMBERS.
Chamberlin, Thomas Chrowder, Hyde Park Hotel, Chicago, Ill.
A.B. (Beloit) ; Ph.D. (Wisconsin. Michigan) ; LL.D. (Michigan, Beloit,
Columbia, Wisconsin, Toronto) ; Sc.D. (Illinois, Wisconsin). Profes¬
sor Emeritus, Geological Department, University of Chicago,
Consulting Geologist U. S. Geological Survey ; Consulting
Geologist, Wisconsin Natural History Survey ; Geologi¬
cal Research Associate, Carnegie Institution of
Washington. D. C. ; Editor, Journal of Geology.
Fairchild, Mrs, Lucius, 302 Monona Avenue, Madison, Wisconsin.
1 Deceased.
’Deceased December 9. 1889. Professor Birge elected to fill unexpired term.
List of Members.
719
Garland, Hamlin,
New York, N. Y.
Vice-President, International Institute of Arts and Letters. Chairman
of Cliff-Dwellers, of Chicago.
Jordan, David Starr,
Chancellor Emeritus of Stanford University, Stanford Univer¬
sity, Cal.
Chancellor Emeritus of Stanford University, Stanford University, Cal. ;
M.S.. Cornell University. 1872 ; M.D., Indiana Medical College, 1875 ; Ph.D.,
Butler College, 1878 ; LL.D., Cornell University, 1886, Johns Hopkins Uni¬
versity, 1902, Illinois College, 1903, Western Reserve, 1915, University
of California, 1915, Indiana University, 1909 ; Instructor in Botany,
Cornell University, 1871-72 ; Professor of Natural History, Lom¬
bard University, 1872-73; Principal of Appleton (Wis. ) Collegi¬
ate Institute, 1873-74 ; Lecturer in Marine Botany at Penikese,
1873-74; Teacher of Natural History, Indianapolis High
School, 1874-75 ; Professor of Biology, Butler College, 1875-
79 ; Instructor in Botany, Harvard Summer School, Cumber¬
land Gap, 1875-76 ; Assistant to U. S. Fish Commission,
1877-81 ; Professor of Zoology, Indiana University, 1879-
85 ; President of Indiana University, 1885-91 ; President
Stanford University, 1891-1913, Chancellor, 1913 ;
President of the California Academy of Sciences, 1891-
98, 1901-03, 1908-1910; U. S. Commissioner in
charge of Fur Seal Investigations, 1896-98; of Sal¬
mon Investigations, 1903 ; International Commis¬
sioner of Fisheries, 1908-1910; President of the
American Association for the Advancement of
Science, 1902.
Merrick, George B., 850 West Washington Avenue, Madison
Author of Genealogy of the Merrick— My rick Family of Massachusetts,
1686-1902. Madison, 1902; Old Times on the Upper Mississippi, The
Recollections of a Steamboat Pilot from 185 4-186 3, Cleveland,
1909 ; Genesis of Steamboating on Western Rivers, with a
Registry of Officers on the Upper Mississippi (with
William R. Tibbals), State Historical Society,
Madison, 1912.
Urbana, Ill.
Trelease, William,
B.S. (Cornell) ; S.D. (Harvard) ; LL.D. (Wisconsin, Missouri, Washing¬
ton University) ; Professor of Botany University of Wisconsin,
1883-5 ; Professor of Botany Washington University, 1885-1913 ;
Director Missouri Botanical Garden, 1889-1912 ; Professor
of Botany University of Illinois, 1913.
Wheeler, W. M.,
Forest Hills, Boston, Mass.
PH.D. Professor of Economic Entomology, Harvard University.
720
Wisconsin Academy of Sciences , Arts , and Letters.
LIFE MEMBERS.
*Birge, Edward Asahel, 772 Langdon St., Madison
A.B., A.M. (Williams); Ph.D. (Harvard); Sc.D. (Western University of
Pennsylvania) ; LL.D. (Williams, Wisconsin, Missouri). Professor of
Zoology, Dean of College of Letters and Science, and President of the
University of Wisconsin ; Secretary of Commissioners of Fisheries,
Wisconsin ; Director, Superintendent, and President, Wisconsin
Geological and Natural History Survey ; Member Wisconsin
State Board of Forestry, Wisconsin Conservation Commis¬
sion, Wisconsin Free Library Commission ; President,
United Chapters Phi Beta Kappa ; President Wiscon¬
sin Academy of Sciences, Arts, and Letters ; Presi¬
dent National Association of State Universities.
Brittingham, Thomas Evans, Dunmuven, Old Sauk Road, Madison
Lumberman.
# Davis, John Jefferson, 419 Sterling Place, Madison
B.S. (Illinois) ; M.D. (Hahnemann). Physician. Curator of
Herbarium, University of Wisconsin.
*Flint, Albert S to well, 1311 Morris St., Madison
A.B. (Hard) ; A.M. (Cincinnati). Astronomer Emeritus, Washburn
Observatory, University of Wisconsin.
Hobbs, William Herbert, 1705 Hill St., Ann Arbor, Mich.
B.S. (Worcester Polytechnic Institute); A.M., Ph.D. (Johns Hopkins).
Professor of Geology and Director Geological Laboratory, Univer¬
sity of Michigan ; Secretary, Wisconsin Academy of Sciences,
Arts, and Letters, 1891-1893 ; President, Michigan Acad¬
emy of Sciences, 1916-1917.
Horlick, A. J., Racine
President Horlick's Malted Milk Company, Racine. Regent, University
of Wisconsin.
Hxxon, Frank B., La Crosse
Lumberman.
Leith, Charles Kenneth, Moraine, Old Sauk Road, Madison
B.S., Ph.D. (Wisconsin). Professor of Geology, University of Wisconsin.
Logan, Hon. Frank A., Chicago
Banker.
• Member of the American Association for the Advancement of Science resident
In Wisconsin.
List of Members.
721
Marsh, Charles Dwight,
1882 Monroe St., N. W., Washington, D. C.
A.B., A.M. (Amherst) ; Ph.D. (Chicago). Physiologist in Charge of
Poisonous Plant Investigations, United States Department
of Agriculture.
Morris, Mrs. Charles W., Milwaukee
Member of Board of Visitors, University of Wisconsin.
Plantz, Samuel, 545 Union St., Appleton
A.M. (Lawrence) ; Ph.D. (Boston) ; D.D. (Albion) ; LL.D. (Baker,
Wisconsin). President, Lawrence College.
Sharp, Frank Chapman, 27 Mendota Court, Madison
A.B. (Amherst) ; Ph.D. (Berlin). Professor of Philosophy,
University of Wisconsin.
^Skinner, Ernest Brown, 210 Lathrop St., Madison
A.B. (Ohio) ; Ph.D. (Chicago). Professor of Mathematics,
University of Wisconsin.
Slaughter, Moses Stephen, 633 Frances St., Madison
A.B., A.M. (De Pauw) ; Ph.D. (Johns Hopkins). Professor of Latin,
University of Wisconsin.
*Slichter, Charles Sumner, 636 Frances St., Madison
B.S., M.S. (Northwestern). Professor of Applied Mathematics and Dean
of the Graduate School, University of Wisconsin.
Van Cleef, Frank Louis, 39 Fort Greene Place, Brooklyn, N. Y.
A.B. (Oberlin, Harvard) ; Ph.D. (Bonn). Chief of Sixth Division and
Translator in Office of Commissioner of Records, Kings County.
ACTIVE MEMBERS
Aitken, Robert Thomas, Mount Hamilton, Calif.
A. B. (California). Research Associate, Bishop Museum, Honolulu, T. H.
Albertz, H. W., Madison
M.S. (Wisconsin). Instructor in Agronomy, University of Wisconsin.
Alderman, William E., 718 Church St., Beloit
Ph.B., A.M., Ph.D. (Wisconsin). Professor of English Literature,
Beloit College.
* Allen, Charles Elmer, 2014 Chamberlin Ave., Madison
B. S., Ph.D. (Wisconsin). Professor of Botany, University of Wisconsin.
46— S. A. L.
722
Wisconsin Academy of Sciences, Arts, and Letters.
* Anderson, John Arlington, 211 Prospect Ave., Madison
B.Sc. (Agr. ) University of Wisconsin. Assistant in Agricultural
Bacteriology, University of Wisconsin.
Arzberger, Emil Godfrey, Washington, D. C.
Ph.B. (Wisconsin). Bureau of Plant Industry.
#Bagg, Rufus M. Jr., 7 Brokaw Place, Appleton
B.A. (Amherst); Ph.D. (Johns Hopkins). Professor of Geology and
Mineralogy; Curator of Museum, Lawrence College.
*Baird, Edgar A., 316 S. Henry St., Madison
B.A., M.A. (University of Wisconsin). Instructor in Botany, University
of Wisconsin.
Barber, W. Harley, 416 Woodside Ave., Ripon, Wis.
A.B., M.A. (University of Wisconsin). Dean and Professor of Physics,
Ripon College, Ripon, Wis.
#Bardeen, Charles Russell, 23 Mendota Court, Madison
A.B. (Harvard) ; M.D. (Johns Hopkins). Professor of Anatomy, and
Dean of the Medical School, University of Wisconsin.
Barker, Augustus Lawrence, 200 Elm St., Ripon
B.S., M.S. (Alabama). Professor of Chemistry, Ripon College.
#Barrett, S. A., Public Museum, Milwaukee
B.S., M.S., Ph.D. (University of California). Anthropologist;
Director, Public Museum, Milwaukee.
* Barrett, Storrs Barrows, Williams Bay
A.B. (University of Rochester). Assistant Professor of Astrophysics,
University of Chicago ; Secretary, Yerkes Observatory.
Barth, George P., 2120 Grand Ave., Apt. 8, Milwaukee
Physician. Director School of Hygiene.
Bascom, Lelia, 419 Sterling Court, Madison
B.A.. M.A. (Wisconsin). Assistant Professor of English,
University of Wisconsin.
Beatty, Arthur, 1824 Yilas St., Madison
A.B. (Toronto) ; Ph.D. (Columbia; Officier d’Acad§mie ; Assistant
ProfessoPof English, University of Wisconsin.
^Bennett, Edward, 1919 Jefferson St., Madison
E.E. (Western University of Pennsylvania). Professor of Electrical
Engineering, University of Wisconsin.
List of Members.
723
#Bertschy, Adaline F., 532 32nd St., Milwaukee
Instructor, La Farge High School.
Blakeman, Bey. E. W., 1121 University Ave., Madison
A.B., M.A. (Wisconsin); D.D. (Lawrence). Pastor, University Methodist
Church, Madison.
Blanchard, W. 0., 2610 Stevens Ave., Madison
Instructor in Geology, University of Wisconsin.
Bleyer, Willard Grosvenor, 423 N. Carroll St., Madison
B.L., M.L., Ph.D. (Wisconsin). Director, Course in Journalism,
University of Wisconsin.
*Boutwell, Paul Winslow, Beloit College, Beloit
B.S. (Beloit)-; M.A. and Ph.D. (Wisconsin). Assistant Professor of
Chemistry, Beloit College.
•Brannon, Melvin Amos, 846 College Ave., Beloit
B.A., M.A. (Wabash) ; Ph.D. (Chicago). President, Beloit College.
Braun, Adolph B., 832 38th St., Milwaukee
Graduate of National German-American Teachers’ Seminary, Milwaukee.
Teacher of Modern Languages, North Division High
School, Milwaukee.
Brown, Charles E., Waban Hill, Nakoma, Madison
Secretary and Curator, Wisconsin Archaeological Society ;
Chief, State Historical Museum.
Brown, Charles Newton, 41 Boby Boad, Madison
LL.B. (Wisconsin). Lawyer.
Brown, Eugene Anson, Langdon Apts., Madison
M.D. (Hahnemann). Physician and Surgeon; Secretary of Board of
Federal Pension Examiners, Madison District.
Brown, Mabel Mary, Platteville
Assistant in Botany.
•Browne, Frederick Lincoln,
223 Clifford Court, Madison
B.Chem. (Cornell); M.S., Ph.D. (Wisconsin). DuPont Fellow,
University of Wisconsin.
Browning, Harold William, Kingston, Bhode Island
B.S. (Rhode Island State College) ; M.S. (University of Wisconsin, 1916) ;
Professor of Botany, Rhode Island State College.
724 Wisconsin Academy of Sciences, Arts, and Letters.
#Bryan, G. S'., 803 State St., Madison
Assistant Professor of Botany, University of Wisconsin.
#Bryan, Ollie Clifton, University of Wisconsin, Madison
B.A. (Alabama Polytechnic Institute). Fellow, University of
Wisconsin.
Buehler, Henry Andrew, Rolla, Mo.
B.S. (Wisconsin). Geologist; State Geologist of Missouri.
^Bunting, Charles Henry, 2020 Chadbourne Ave., Madison
B.S. (Wisconsin) ; M.D. (Johns Hopkins). Professor of Pathology,
University of Wisconsin.
Burke, Rush Pearson, 615 Bell Building, Montgomery, Ala.
M.Sc., M.D. Physician and Surgeon.
Bussewitz, M. A., 435 Kenwood Blvd., Milwaukee
Professor, Milwaukee State Normal School.
Butterfield, Mrs. Ellen F.,
713 Milwaukee Ave., South Milwaukee
Chairman, Museum Committee of South Milwaukee Public Museum.
Cahn, A. R., 4720 Greenwood Ave., Chicago, Ill.
B.S. (Cornell) ; M.S. (Wisconsin). Associate Professor of Biology,
A. and M. College, College Station, Texas.
Cairns, William B., 2010 Madison St., Madison
A.B., Ph.D. (Wisconsin). Associate Professor of American Literature,
University of Wisconsin.
Campbell, 0. J., Jr., Ann Arbor, Mich.
Ph.D. (Harvard). Professor of English, University of Michigan.
Carter, Sylvester J., 850 Newhall St., Milwaukee, Wis.
B.A., B.L.S. Reference Librarian, Milwaukee Public Library.
Chandler, Elwyn Francis, University, N. D.
A.B., A.M. (Ripon). Professor of Civil Engineering, University of
North Dakota.
Chase, Wayland J., 141 Summit Ave., Madison
A.B., A.M. (Brown). Associate Professor of History, University of
Wisconsin.
List of Members.
725
* Clark, Paul F., 2136 Van Hise Ave., Madison
Professor of Bacteriology, University of Wisconsin.
Clas, Alfred Charles, 445 Milwaukee St., Milwaukee
Architect (Ferry & Clas), 419 Broadway, Milwaukee. Member,
, Board of Park Commissioners.
Clawson, Arthur Brooks,
1884 Monroe St., N. W., Washington, D. C.
A.B. (Michigan). Department of Agriculture, Washington.
Cole, Harry E., 908 Ash St., Baraboo
Editor of Baraboo News.
*Cole, Leon J., 1915 Keyes Ave., Madison
A.B. (Michigan) ; Ph.D. (Harvard). Professor of Genetics,
University of Wisconsin.
^Collie, George Lucius, 48 Sherwood Drive, Beloit
B.S. (Beloit) ; M.A., Ph.D. (Harvard) ; LL.D. (Beloit). Professor,
Beloit College.
Commons, John R., 816 University Ave., Madison
Professor of Economics, University of Wisconsin.
Conklin, G. H., Suite 201-202 Board of Trade Building, Superior
Practicing Physician.
Corp., C. I., 2114 West Lawn Ave., Madison
Professor of Hydraulic and Sanitary Engineering, University of
Wisconsin.
^Crawford, John Forsyth, 726 Milwaukee Road, Beloit
A.B. (Princeton) ; A.M. (Princeton) ; B.D. (McCormick) ; Ph.D.
(Chicago). Professor of Philosophy, Beloit College.
Culbertson, Henry Coe, Ripon
Culver, C. A., Beloit
Professor of Physics, Beloit College.
* Culver, Garry Eugene, 301 Center Ave., Stevens Point
A.M. (Denison). Chair of Geology and Chemistry, State Normal School.
Dancey, Lloyd Slote, 125 Charles St., Waukesha
B.A., M.A. (Illinois). Professor of Physics and Applied Mathematics;
Dean and Acting President, Carroll College.
726 Wisconsin Academy of Sciences , Arts , and Letters.
^Daniels, Farrington, 1162 Emerald St., Madison
Assistant Professor of Chemistry, University of Wisconsin.
#Davis, Carl Henry, 141 Wisconsin St., Milwaukee
A.B. (Oregon) ; B.S. (Chicago) ; M.D. (Rush Medical College).
Dean, Alletta F., 87 West St., Mansfield, Mass.
Ph.B., Ph.M. (Wisconsin).
^Delwiche, Edmond Joseph, R. F. D. 3, Green Bay
B.S. A., M.S. (Wisconsin). Professor of Agronomy, University of
Wisconsin.
Dennis, Alfred Lewis Pinneo, 518 Wisconsin Ave., Madison
A.B. (Princeton) : Ph.D. (Columbia).
#Denniston, Rollin Henry,
1414 West Washington Ave., Madison
Ph.G., B.S., Ph.D. (Wisconsin). Assistant Professor of Botany,
University of Wisconsin.
Dernehl, Paul Herman, 717-718 Majestic Building, Milwaukee
B.S. (Wisconsin) ; M.D. (Johns Hopkins). Physician.
Dodge, B. 0., New York, N. Y.
Ph.B. (Wisconsin); Ph.D. (Columbia). Instructor in Botany;
Secretary-Treasurer Torrey Botanical Club, Department
of Botany, Columbia University.
Dodge, Robert Elkin Neil, 15 W. Gorham St., Madison
A.B., A.M. (Harvard). Associate Professor of English,
University of Wisconsin.
^Doolittle, Sears P.,
Department of Plant Pathology, University of Wisconsin, Madison
B.S., M.S. (Michigan Agricultural College) ; Ph.D. (Wisconsin).
Assistant Pathologist, University of Wisconsin.
Dowling, Linnaeus Wayland, 2 Roby Road, Madison
Ph.D. (Clark). Associate Professor of Mathematics,
University of Wisconsin.
Downes, Robert Hugh, 2434 Jefferson Ave., Norwood, Ohio
B.L. (Wisconsin). General Manager, Norwood Sash and Door
Manufacturing Company.
List of Member's.
727
*Drechsler, Charles, 978 Washington Ave., Brooklyn, N. Y.
B.S., M.S. (Wisconsin) ; Ph.D. (Harvard). Assistant Pathologist,
Bureau of Plant Industry, Department of Agriculture.
^Dresden, Arnold, 2114 Vilas St., Madison
S.M., Ph.D. (Chicago). Associate Professor of Mathematics,
University of Wisconsin.
^Dudley, William Henry, 148 Breese Terrace, Madison
Chief, Bureau of Visual Instruction, Extension Division,
University of Wisconsin.
Du Mez, Andrew Grover,
25th and East Streets, N. W., Washington, D. C.
Associate Pharmacologist, United States Public Health Service.
Edwards, Ira, Austin, Texas
Adjunct Professor, Universil y of Texas.
Ely, Richard Theodore, 205 Prospect Ave., Madison
A.B., A.M. (Columbia) ; Ph.D. (Heidelberg) ; LL.D. (Hobart).
Professor of Political Economy, University of Wisconsin.
*Evans, Clarence Turner, 3415 Sycamore St., Milwaukee
M.E. in E.E. (Ohio State University). Development Engineer,
Cutler-Hammer Mfg. Co., Milwaukee.
^Farley, John Herbert, 482 South St., Appleton
A.M. (Lawrence). Professor of Philosophy and Psychology, Lawrence
College.
Fehlandt, August F., Ripon
A.B. (Wisconsin) ; B.D. (Yale). Professor of Economics and Sociology,
Ripon College.
*Finch, Vernon C., College Hills, Madison
Associate Professor of Geology, University of Wisconsin.
Finkler, Adolph, 612 Commerce St., Milwaukee
Secretary, Adolph Trostel and Sons Company ; President, Board of
Trustees, National German-American Teachers’ Seminary ; Presi¬
dent, Board of Trustees, German-English Academy.
^Fischer, Richard, 119 East Johnson St., Madison
Ph.C., B.S. (Michigan) ; Ph.D. (Marburg). Professor of Chemistry,
University of Wisconsin.
728
Wisconsin Academy of Sciences , Arts , and Letters.
Fish, Carl RussELLr 244 Lake Lawn PL, Madison
A.B. (Brown) ; A.M., Ph.D. (Harvard). Professor of American
History, University of Wisconsin
*Fisk, Emma Luella, Madison
B.A. (Wellesley College). Assistant in Botany, University of Wisconsin.
Fling, Harry R., 601 Jackson St., Oshkosh
A.B. (Bowdoin). Professor of Biology, State Normal School.
#Fluke, Charles Lewis, Jr., 1219 Lee Court, Madison
M.S. (Wisconsin). Instructor in Economic Entomology,
University of Wisconsin.
*Fracker, Stanley Black 1632 Adams St., Madison
A.B. (Buena Vista College) ; M.S. (Iowa State College) ; Ph.D.
(University of Illinois). State Entomologist.
Franklin, Ira, M. D., 204-205 Majestic Bldg., Milwaukee
Oculist and Aurist.
* Frost, William Dodge, 1010 Grant St., Madison
B.S., M.S. (Minnesota) ; Ph.D. (Wisconsin). Professor of Bacteriology,
University of Wisconsin.
Gay, Lucy Maria, 216 N. Pinckney St., Madison
B.L. (Wisconsin). Assistant Professor of Romance Languages,
University 'of Wisconsin.
# Gerry, Eloise 419 Sterling Place, Madison
A.B. and A.M. (Radcliffe College, Harvard University). Microscopist
and Lecturer in Forest Products, University of Wisconsin.
* Gilbert, Edward Martinius, 25 Spooner St., Madison
A.B. (Wisconsin). Associate Professor of Botany, University of
Wisconsin.
* Gilman, Albert F., Waukesha
S.B., A.M. (Amherst) ; Ph.D. (Denver). Professor of Chemistry,
Carroll College.
Glicksman, Harry, 1601 Adams St., Madison
B.A., LL.B., Ph.D. (Yale). Instructor in English; Assistant Dean,
College of Letters and Science, University of Wisconsin.
List of Members.
729
Gloyer, Walter 0., Geneva, N. Y.
B.A., M.A. (Wisconsin). Associate Botanist, New York Agricultural
Experiment Station.
*Graul, Edward John, 8 Lathrop St., Madison
B.S.A. and M.S. (Wisconsin). Assistant Professor of Soils,
University of Wisconsin.
Greene, Howard,
1416 First Wisconsin National Bank Bldg., Milwaukee
Gregory, John Goadby, 717 Jefferson St., Milwaukee
Member, Wisconsin History Commission.
Griggs, Horace William, 2421 Sycamore St., Milwaukee
Recording Inspector, C., M. & St. P. Ry. Co.
^Groves, James Frederick, Ripon
Ph.B. (Chicago). Professor of Biology, Ripon College.
Gutsch, Milton R.,
Professor of History, University of Texas.
Austin, Texas
#Guyer, Michael F., 138 Prospect Ave., Madison
Professor of Zoology, University of Wisconsin.
Haase, Ewald, 182 Wisconsin St., Milwaukee
Secretary-Treasurer, Milwaukee Gas Light Company.
Haessler, Luise, 100 Morningside Drive, New York, N. Y.
A.B. (Chicago). Associate Professor of German, Hunter College.
Halpin, J. G., 2500 University Ave., Madison
Professor of Poultry Husbandry, University of Wisconsin.
* Hargrave, Josephine R., 415 Thorn St., Ripon
A.B. (Ripon) ; S.B. (Simmons). Librarian of Ripon College Library.
*Harmer, Paul M., Milwaukee
B.S. (Carleton) ; M.S. and Ph.D. (University of Minnesota).
730 Wisconsin Academy of Sciences, Arts, and Letters.
Harper, Robert Aylmer, New York City
Professor of Botany, Columbia University.
Harper, Mrs. Robert Aylmer, New York City
^Hawkins, Pliny H., 1910 Regent St., Madison
*Heddle, John R., 822 W. Johnson St., Madison
Assistant in Botany, University of Wisconsin.
*Heineman, Paul G., Woodworth
B:S. and Ph.D. (University of Chicago). Director of Laboratories,
United States Standard Products Company.
#Henmon, Vivian Allen Charles, 18 Mendota Ct., Madison
A.B. and A.M. (Bethany College) ; Ph.D. (Columbia). Director, School
of Education ; Professor, School of Education, University
of Wisconsin.
HIodgson, A. J., Waukesha
M.D. (Rush) ; Sc.D. (Carroll College).
Hohlfeld, Alexander Rudolph, 124 Breese Terrace, Madison
Ph.D. (Leipzig). Professor of German, University of Wisconsin;
President, Modern Language Association of America ; Member
of Board of Administration, National German-American
Teachers’ Seminary, Milwaukee.
^Holden, Eugene Davenport, Madison
Instructor in Agronomy, University of Wisconsin.
Holmes, Samuel Jackson, Berkeley, California
B.S., M.S. (California) ; Ph.D. (Chicago). Professor of Zoology,
University of California.
^Hotchkiss, W. 0., College Hills, Madison
Director, Wisconsin Geological and Natural History Survey ; Geologist,
State Highway Commission.
Hubbard, Frank Gaylord, 141 W. Gilman St., Madison
A.B. (Williams) ; Ph.D. (Johns Hopkins). Professor of English,
University of Wisconsin.
Hubert, Ernest Everett, Old Soils Building, Madison
M.S. and B.S. (University of Montana) ; Scientific Assistant in Plant
Pathology, Bureau of Plant Industry.
List of Members,
731
^Humphrey, Clarence J., 1714 Adams St., Madison
A.B., B.Sc. in Forestry (Nebraska). Pathologist, Forest Products
Laboratory.
Ibsen, H. L., Manhattan, Kansas
Assistant Professor of Genetics, State College, Manhattan.
*Ingersoll, Leonard R., 1933 West Lawn Ave., Madison
B.S. (Colorado College) ; Ph.D. (Wisconsin). Associate Professor of
Physics, University of Wisconsin.
Jackson, Hartley H. T., Washington, D. C.
U. S. Biological Survey.
*Jana, Ashutosh, Haria, Bengal, India
M.Sc., M.E.E., LL.B. (Intercontinental) ; LL.B. (Potomac). President,
The Bengal Institute of Arts and Sciences ; Managing Director, Tam-
luk Commercial Industries, Ltd. ; Managing Proprietor, Jana
Research Laboratory ; Proprietor, The Birulia Scientific
Library.
*Jansky, Cyril M., 2117 Jefferson St., Madison
A.B. (Valparaiso) ; B.S. (Michigan). Professor of Electrical
Engineering, University of Wisconsin.
^Jastrow, Joseph, 237 Langdon St., Madison
A.B., A.M. (Pennsylvania) ; Ph.D. (Johns Hopkins). Professor of
Psychology, University of Wisconsin.
* Johnson, Aaron Guy, 1713 Chadbourne Ave., Madison
Plant Pathologist, University of Wisconsin.
* Johnson, James, 2005 Madison St., Madison
Associate Professor of Horticulture, University of Wisconsin.
* Jones, Fred Reuel,
Bureau of Plant Industry, Washington, D. C.
B.S. (Maine) ; M.S. and Ph.D. (Wisconsin). Pathologist, Bureau of
Plant Industry.
* Jones, Lewis R., 146 Prospect Ave., Madison
Ph.B., Ph.D. (University of Michigan) ; Sc.D. (Honorary, University of
Vermont). Professor of Plant Pathology, University of Wisconsin.
* Jones, Sarah Yan Hoosen, Madison
Assistant in Experimental Breeding, University of Wisconsin.
732
Wisconsin Academy of Sciences, Arts, and Letters.
5*Juday, Chancey, 35 Lathrop St., Madison
A.B., A.M. (Indiana). Biologist, Wisconsin Geological and Natural
History Survey.
*Kahlenberg, Louis, 234 Lathrop St., Madison
B.S. and M.S. (Wisconsin) ; Ph.D. (Leipzig). Professor of Chemistry,
University of Wisconsin.
#Keitt, George Wannamaker, 803 State St., Madison
B.S. (Clemson) ; Ph.D. (Wisconsin). Professor of Plant Pathology,
University of Wisconsin.
*Kemmerer, George I., 916 Van Bnren St., Madison
B.S., M.A. (Wisconsin). Assistant Professor of Chemistry, University
of Wisconsin.
*Kiekhoefer, William Henry, 1919 Arlington Place, Madison
B.A., Ph.D. (Wisconsin). Professor Of Economics, University of
Wisconsin.
Kinsman, Delos 0.,
Professor of Economics, Lawrence College.
Appleton
^Koehler, Arthur, 1819 Adams St., Madison
B.S. (Michigan). Lecturer in Forest Products, United States Forest
Service.
*Kowalke, Otto Louis, 2012 Jefferson St., Madison
B.S. and Ch.E. (Wisconsin). Professor of Chemical Engineering,
University of Wisconsin.
*Kraus, Ezra Jacob, 803 State St., Madison
B.S. (Michigan Agricultural College) ; Ph.D. (Chicago). Professor of
Applied Botany, University of Wisconsin.
^Kremers, Edward, Highlands, Madison
Ph.G., B.S. (Wisconsin) ; Ph.D. (Gottingen) ; D.Sc. (Michigan).
Director of Course in Pharmacy and Professor of Pharma¬
ceutical Chemistry, University of Wisconsin.
Kutchin, Mrs. Harriet Lehman, Green Lake, Wis.
A.B. (Ripon) ; A.M. (Northwestern).
Langenhan, H. August, 1821 West Lawn Ave., Madison
Ph.G., Ph.C., B.S., Ph.D. (Wisconsin). Assistant Professor of Pharmacy,
University of Wisconsin.
List of Members.
733
Larsell, Olaf, ' Evanston, Ill.
Associate Professor of Zoology, Northwestern University.
^Lawson, Mrs. Publius V., 327 Nayrnut St., Menasha
*Lee, Oliver Justin, Yerkes Observatory, Williams Bay
A.B. (Minnesota) ; M.Sc. and Ph.D. (Chicago). Instructor in Practical
Astronomy, University of Chicago.
^Lenher, Victor, 158 Summit Ave., Madison
Ph.D. (Pennsylvania). Professor of Chemistry, University of Wisconsin.
^Leonard, William Ellery, 2015 Adams St., Madison
A.B. (Boston University) ; M.A. (Harvard) ; Ph.D. (Columbia).
Associate Professor of English, University of Wisconsin.
Lewis, F. F., 115 Jackman St., Janesville
President, Lewis Knitting Company, Janesville.
*Lindstrom, Ernest W., 803 State St., Madison
A.B. (Wisconsin) ; Ph.D. (Cornell). Assistant Profesor of Genetics,
University of Wisconsin.
*Lobeck, Armen Kohl, 431 Hawthorne Court, Madison
Assistant Professor of Geology. University of Wisconsin.
Loft, Genivera E., 20 Lathrop St., Madison
B.A., M.A. Instructor in Geology, University of Wisconsin.
#Lowe, John N., 610 N. Front St., Marquette, Michigan
Head of Department of Natural Science, Marquette Normal School.
McAllister, Fred, 3205 West Ave., Austin, Texas
A.B., A.M., Ph.D., Associate Professor of Botany, University of Texas
McCaskill, Virgil E., Superior
President, State Normal School.
McKenna, Maurice, 152 S. Main St., Fond du Lac
Lawyer ; President Bar Association of Fond du Lac County.
#McKinney, H. H., 24 Lathrop St., Madison
B.S. and M.S. Assistant Pathologist, Department of Agriculture.
734 Wisconsin Academy of Sciences, Arts, and Letters.
McLeod, Andrew Friedley, Northfield, Minn.
Ph.D. (Wisconsin). Professor of Organic Chemistry, Carleton College.
•McMinn, Amelia, State Normal School, Milwaukee
B.S. (Wisconsin) ; M.A. (Columbia). Instructor in Hygiene, State
Normal School, Milwaukee.
Main, Mrs. Angie Kumlien, Fort Atkinson
•March, Herman William, 115 Summit Ave., Madison
A. B. (Michigan) ; Ph.D. (Munich). Associate Professor of Mathematics,
University of Wisconsin.
•Marlatt, Abby Lillian, 612 Howard Place, Madison
B. S., M.S. (Kansas State College). Director of Course and Professor of
Home Economics, University or Wisconsin.
Marquette, William George, New York, N. Y.
Ph.G. (Northwestern) ; B.S., Ph.D. (Wisconsin).
•Marschall, A. J., 14 Proudfit St., Madison
Proprietor, Marschall Dairy Laboratory.
Marshall, Ruth, Rockford, Ill.
B.Sc., M.S. (Wisconsin) ; Ph.D. (Nebraska). Head of Department of
Biology, Rockford College.
•Marshall, William Stanley, 139 E. Gilman St., Madison
B.S. (Swarthmore) ; Ph.D. (Leipzig). Associate Professor of
Entomology, University of Wisconsin.
•Martin, Ella May, 1717 Monroe St., Madison
Assistant in Botany, University of Wisconsin.
•Mason, Max, 1902 Arlington Place, Madison
B.S. (Wisconsin). Professor of Mathematical Physics, University
of Wisconsin.
•Mathews, Joseph Howard. 128 Lathrop St., Madison
B.S., A.M. (Wisconsin) ; A.M., Ph.D. (Harvard). Professor of
Chemistry and Director of the Course in Chemistry.
•Maurer, Edward Rose, 167 Prospect Ave., Madison
B.C.E. (Wisconsin). Professor of Mechanics. University of Wisconsin.
List of Members.
735
Mayor, J. W., Schenectady, N. Y.
Professor of Zoology, Union College.
*Meachem, John Goldesbrough, Jr., 745 College Ave., Racine
M.D. (Rush). Physician; President, St. Luke’s Hospital.
*Mead, Daniel W., 120 W. Gorham St., Madison
C.E. (Cornell) ; Professor of Hydraulic and Sanitary Engineering,
University of Wisconsin.
*Mead, Warren J., 922 Yan Bnren St., Madison
Associate Professor of Geology, University of Wisconsin.
Meyer, Balthasar Henry, Washington, D. C.
B.L., Ph.D., LL.D. (Wisconsin). Member, Interstate Commerce
Commission.
* Miller, William Snow, 2001 Jefferson St., Madison
M.D. (Yale) ; Sc.D. (Cincinnati). Professor of Anatomy,
University of Wisconsin.
Moe, Carl, International Falls, Minn.
Chief Chemist, Minnesota and Ontario Paper Company.
Monroe, C. E., 512 Van Bnren St., Milwaukee
A.B. (Oberlin College) ; LL.B. (Michigan University). Lawyer.
Moore, Samuel, 1503 Cambridge Rd., Ann Arbor, Mich.
A.B. (Princeton) ; Ph.D. Harvard). Associate Professor of English,
University of Michigan.
Morris, H. H., 1426 Yan Buren St., Wilmington, Delaware
Executive Secretary, E. I. du Pont de Nemours & Co.
*Mullenix, Rollin Clarke, 461 Washington St., Appleton
A.B. (Wheaton College) ; Ph.D. (Harvard) ; Professor of Zoology,
Lawrence College.
Murphy, Ray Y., Bloomington, Illinois
B.S. (Illinois Wesleyan University) ; M.S. (Wisconsin).
Illinois Wesleyan University.
Mutch, William J., 221 Watertown St., Ripon
A.B., A.M. (Wisconsin) ; B.D., Ph.D. (Yale). Professor of Philosophy
and Education, Ripon College.
736
Wisconsin Academy of Sciences , Arts, and Letters.
Muttkowski, Richard Antony, Moscow, Idaho
A.B., Ph.D. (Wisconsin). Assistant Professor of Zoology and
Entomology, University of Idaho.
#Nagler, Mrs. Ellen Torelle, 2548 E. Johnson St., Madison
#Nardin, F. Louise, 612 Howard Place, Madison
B.S., A.M., Ph.D. Dean of Women, University of Wisconsin.
Naylor, Wilson Samuel, Appleton
Professor, Lawrence College.
#Neilson, Walter Hopper, 114 Garfield Ave., Milwaukee
M.D. (Rush). Dean of the Medical Faculty and Professor of the
Principles and Practice of Medicine and Clinical Medicine,
Milwaukee Medical College.
Oberholser, Harry Church,
2805 18th St., N. W., Washington, D. C.
A.B., M.S., Ph.D. Assistant Biologist, U. S. Department of Agriculture.
Olin, John Myers, 130 Prospect Ave., Madison
A.B., A.M. (Williams) ; LL.B. (Wisconsin). Lawyer ; Professor of Law,
University of Wisconsin.
O’Shea, M. Vincent, 140 Langdon St., Madison
B.L. (Cornell). Professor of the Science and Art of Education,
University of Wisconsin.
Otto, Max, 1155 Edgewood Ave., Madison
Professor of Philosophy.
^Overton, James Bertram, 512 Wisconsin Ave., Madison
Ph.B. (Michigan) ; Ph.D. (Chicago). Professor of Plant Physiology,
University of Wisconsin.
Owen, Edward Thomas, 614 State St., Madison
A.B., Ph.D. (Yale). Emeritus Professor of French and Linguistics,
University of Wisconsin.
Owen, Ralph W., 1501 State St., Eau Claire
Litt.B. (Princeton) ; M.A. (Wisconsin).
Pammel, L. H., Ames, Iowa
Professor of Pathology, General and Systematic Botany, Iowa State
College.
List of Members .
737
Parker, Fletcher Andrew, 14 W. Gilman St., Madison
Professor Emeritus of Music, University of Wisconsin.
♦Parkinson, John Barber, 516 Wisconsin Ave., Madison
A.B., A.M., LL.B. (Wisconsin). Vice-President and Professor Emeritus
of Constitutional and International Law, University of Wisconsin.
Paxson, Frederic L., 2122 Van Hise Ave., Madison
Ph.D. (Pennsylvania). Professor of American History, University of
Wisconsin.
♦Pearse, A. S., 2240 Rowley Ave., Madison
B. A., A.M. (Nebraska) ; Ph.D. (Harvard). Professor of Zoology,
University of Wisconsin.
Peltier, George L., State Farm, Lincoln, Neb.
Professor of Plant Pathology, University of Nebraska.
*Peterson, William Harold, College Hills, Madison
B.S. (Wesleyan) ; M.A. (Columbia) ; Ph.D. (Wisconsin). Associate
Professor of Agricultural Chemistry, University of Wisconsin.
♦Phillips, James David, 1925 West Lawn Ave., Madison
B.S. (Illinois). Professor of Drawing, University of Wisconsin.
Pierson, Merle, 115 Park Terrace, Ann Arbor, Mich.
Pitman, Annie, 414 N. Henry St., Madison
B.A., Ph.D. (Wisconsin). Assistant Professor of Latin, University
of Wisconsin.
Pope, Thomas Edmund Burt, 724 Newhall St., Milwaukee
Ph.B., A.M. Curator of Invertebrate Zoology, Public Museum.
♦Public Museum,
Milwaukee
Reed, George Mathew, Brooklyn, N. Y.
A.B. (Geneva) ; A.M., Ph.D. (Wisconsin). Curator, Brooklyn Botanic
Garden.
♦Reid, Mary Elizabeth, 148 Breese Terrace, Madison
Fellow in Botany, University of Wisconsin.
Rice, Ole S., Madison
B.S. (Wisconsin). State Supervisor of School Libraries, State
Department of Public Instruction.
47— S. A. L.
738 Wisconsin Academy of Sciences , Arts , and Letters.
* Richards, C. Audrey, Madison
Assistant in Forest Pathology, Forest Products Laboratory, Madison.
Rickett, Harold W., 625 Mendota Court, Madison
#Roark, Raymond Jefferson, 433 N. Murray St., Madison
B.S. in Civil Engineering, M.S. in Civil Engineering (Illinois). Assistant
Professor of Mechanics, University of Wisconsin.
Rockwook, Stanley Walker, 232 East Ave., Waukesha
B.S. (Dartmouth) ; A.M. (Chicago). Professor of Romance Languages,
Carroll College.
^Roebuck, John Ransom, 2210 Hollister Ave., Madison
A.B., Ph.D. (Toronto). Associate Professor of Physics, University of
Wisconsin.
Roedder, E. C. L. C., 315 S. Orchard St., Madison
A.B., A.M., Ph.D. (University of Michigan). Associate Professor of
German Philology, University of Wisconsin.
Rogers, James Elcana, 300 East Ave., S., Waukesha
B.A., M.A. (Maryville) ; B.D. (Union) ; Ph.D. (Liege). Professor of
Education, Carroll College.
*Rohde, Hugo W., Shorewood
Chief Chemist, Schlitz Brewing Company.
Rosenberry, Mrs. Lois K., 1717 Kendall Ave., Madison
#Sammis, J. L., 234 Breese Terrace, Madison
Associate Professor of Dairying, University of Wisconsin.
Sanborn, John Bell, Wisconsin Building, Madison
B.L., M.L., Ph.D. (Wisconsin). Lawyer; Treasurer, Wisconsin State
Bar Association; Lecturer, University of Wisconsin Law School;
Member, Wisconsin Council, American Bar Association.
Schinner, Augustine, Right Reverend,
238 East 13th Avenue, Spokane, Wash.
DD. Bishop of Spokane.
Schlundt, Herman, Columbia, Mo.
Professor of Chemistry, University of Missouri.
List of Members.
739
Schorger, A. W., 2021 Kendall Ave., Madison
Ph.B., M.A., Ph.D. Burgess Laboratories, Madison.
Schubring, E. J. B., 415 N. Park St., Madison
B.L., LL.B. (Wisconsin). Attorney.
*Schubring, Mrs. E. J. B., 415 N. Park St., Madison
B.A., M.A., Ph.D. (Wisconsin).
#Schuette, Henry A., 401 Wisconsin Ave., Madison
Assistant Professor of Chemistry, University of Wisconsin.
* Schultz, Alfred R., 800 Vine St., Hudson
B.S. (Wisconsin); Ph.D. (Chicago). Manager, Burkhardt Milling and
Electric Power Company.
Seaman, Gilbert E., Goldsmith. Bldg., Milwaukee
M.D., F.A.C.S. Physician; Regent of the University of Wisconsin.
# Seymour, Edith Katharine, College of Agriculture, Madison
A.B. (Miami); M.S. (Wisconsin). Scientific Assistant, Bureau of
Plant Industry, U. S. Department of Agriculture.
#Showalter, Amos Martin, 314 N. Warren St., Madison
B.A. (Goshen College); M.A. (Wisconsin). Assistant in Botany,
University of Wisconsin.
Showerman, Grant, 410 N. Butler St., Madison
A.B., A.M., Ph.D. (University of Wisconsin). Professor of Latin,
University of Wisconsin.
Sieker, William Christian, 1542 Prospect Place, Milwaukee
B.S. (Wisconsin). Principal, Milwaukee Vocational School.
*Smith, Elizabeth Anita, 417 Sterling Place, Madison
B.A. (Cincinnati); M.A., Ph.D. (Wisconsin). Assistant Professor of
Zoology, University of Wisconsin.
* Smith, Gilbert Brown Lorenzo,
207 W. Washington Ave., Madison
A.B. (Colgate); M.A. (Wisconsin). Assistant in Chemistry,
University of Wisconsin.
* Smith, Gilbert Morgan, 113 Spooner St., Madison
Associate Professor of Botany, University of Wisconsin.
740 Wisconsin Academy of Sciences, Arts, and Letters.
* Smith, Huron Herbert, Milwaukee
Curator of Botany, Public Museum.
Smith, Walter McMynn, 127 Langdon St., Madison
A.B. (Wisconsin). Librarian, University of Wisconsin.
Smythe, Sidney T., Delafield
A.B., A.M. (St. Stephen’s); B.D. (Nashotah) ; D.D., Ph.D (Hobart).
S.T.D. President, St. John’s Military Academy; Member, Commit¬
tee on Canons, Protestant Episcopal Church.
Snell, Walter H., 19 Nisbet St., Providence, Rhode Island
A.B. (Brown); A.M. (Brown); Ph.D. (Wisconsin). Assistant Pro¬
fessor of Botany, Brown University.
*Snow, Benjamin Warner, Langdon Apartments, Madison
Ph.D. (Berlin). Professor of Physics, University of Wisconsin.
Squier, George Hull,
Dairyman.
Trempealeau
Starr, Harry Linn, 348 Wisconsin Ave., Waukesha
B.A., M.A. (Wabash). Professor of English and Secretary of the
Faculty, Carroll College.
*Starr, William J., 135 Marston Ave., Eau Claire
LL.B. (Columbia). Member, Board of Commissioners of Fisheries,
Wisconsin; President, Eau Claire Public Library.
#Steenbock, Harry, 2117 West Lawn Ave., Madison
B.S., M.S., Ph.D. (Wisconsin). Professor of Agricultural Chemistry,
University of Wisconsin.
*Steidtman, E., 2002 Monroe St., Madison
A.B., A.M., Ph.D. (University of Wisconsin). Assistant Professor of
Geology, University of Wisconsin.
*Steil, W. H,, 115 S. Warren St., Madison
Ph.B., Ph.D. Instructor in Botany, University of Wisconsin.
*Stickney, Gardner, 864 Summit Ave., N., Milwaukee
Stickney, M. E., Granville, 0.
A.B. (Bates); A.B., A.M. (Harvard). Professor of Botany, Denison
University.
List of Members.
741
* Stockhausen, Regina Emma, 511 N. Carroll St., Madison
B.A. (Smith); M.A. (Mount Holyoke). Assistant in Botany,
University of Wisconsin.
Stout, Arlow Burdette, New York City
A.B. (Wisconsin). New York Botanical Gardens, Bronx Park.
Talbert, George A., Ripon
B.S., M.S. (Ohio Wesleyan). Professor of Biology, Ripon College.
Taylor, Warner, 219 Clifford Court, Madison
Assistant Professor of English, University of Wisconsin.
Teller, Edgar Eugene, 66 Highland Ave., Buffalo, N. Y.
Real Estate.
#Thorkelson, Halsten Joseph Berford,
1526 W. Washington Ave., Madison
B.S. M.E. (Wisconsin). Business Manager, University of Wisconsin.
Th waites, F. T., Turvillwood, Madison
Curator of Geological Museum, University of Wisconsin.
* Tisdale, William B., 145 Iota Court, Madison
M.S., Ph.D. Instructor in Plant Pathology, University of Wisconsin.
Titus, W. A., 54 Oak Ave., Fond du Lac
Manufacturer; Member of Board of Visitors, University of
Wisconsin; State Senator.
Tomlinson, Charles Weldon, 5070 E. 25th Ave., Denver, Col.
B.A., M.A. (Wisconsin); Ph.D. (Chicago). Geologist, Gypsy Oil
Company, Denver.
Toole, Eben H., Washington, D. C.
Assistant Pathologist', Bureau of Plant Industry, U. S. Department
of Agriculture.
*Tottingham, W. E., 2206 West Lawn Ave., Madison
Associate Professor of Agricultural Chemistry, University of
Wisconsin.
Trever, A. A., 466 Alton St., Appleton
Ph.D. (Chicago). Professor of Ancient and Medieval History,
Lawrence College.
742
Wisconsin Academy of Sciences , Arts, and Letters.
*Truog, Emil, 803 State St., Madison
B.S.A., M.S. (Wisconsin). Professor of Soils, University of Wisconsin.
Turneaure, Frederick Eugene, 166 Prospect Ave., Madison
C.E. (Cornell). Professor of Engineering and Dean of the College
of Engineering, University of Wisconsin.
Turner, Clarence L., Beloit
Professor of Zoology, Beloit College.
#Twenhofel, William Henry, Lake Forest, Madison
A.B., M.A. Ph.D. (Yale). Professor of Geology, University of
Wisconsin.
*Uehling, Edward A., 3226 McKinley Boulevard, Milwaukee
M.E. (Stevens Institute of Technology). Former President Uehling
Instrument Company, 71 Broadway Ave., New York.
Van Ostrand, Mrs. E. H., 214 W. Gilman St., Madison
National Vice-President of Service Star Legion; Honorable State
Regent, D.A.R.
#Van Yleck, Edward Burr, 519 N. Pinckney St., Madison
A.B., A.M. (Wesleyan); Ph.D. (Gottingen); LL..D. (Clark). Frofessor
of Mathematics, University of Wisconsin; Editor, Transactions
of the American Mathematical Society.
Vaughan, R. E., 1708 Regent St., Madison
Associate Professor of Plant Pathology, University of Wisconsin.
Vogel, Mrs. Guido Charles, 409 Terrace Ave., Milwaukee
Vorhies, Charles Taylor, Salt Lake City, Utah
B.S. (Iowa Wesleyan). Professor of Entomology, University of
Arizona.
Voss, Ernest Karl Johnann Heinrich,
175 Virginia Terrace, West Lawn Heights
Ph.D. (Leipzig). Professor of German Philology, University of
Wisconsin; Vice-President, Germanic Museum Association.
Wadmond, Samuel C., Delavan
Manager of Statistical Department, Bradley Knitting Company.
List of Members.
743
#Wagner, George, 1901 Jefferson St., Madison
Ph.C. (Michigan); A.B. (Kansas); A.M. (Michigan). Assistant
Professor of Zoology, University of Wisconsin; Icthyologist,
State Geological and Natural History Survey.
Wakeman, Nellie A., 1814 Ray St., Madison
Instructor in Pharmacy, University of Wisconsin.
# Walker, J. Charles, Madison
Assistant Professor of Plant Pathology, University of Wisconsin.
* Warfield, Louis Marshall, 221 Grand Ave., Milwaukee
A.B., M.D. (Johns Hopkins).
Weidman, Samuel, 410 North Henry St., Madison
B.S., Ph.D. (Wisconsin). Geologist, Wisconsin Geological and Natural
History Survey.
# Werner, Fred W., 991 — 16th St., Milwaukee
Instructor in Biology, North Division High School.
West, George A., 97 Wisconsin St., Milwaukee
Lawyer; President, Board of Trustees, Milwaukee Public Museum.
*Whealdon, Albert D., 817 N. 16th St., Superior
A.B., A.M. (Wisconsin). Professor of Chemistry, State Normal
School, Superior.
Whitbeck, R. H., 105 Spooner St., Madison
Professor of Geography, University of Wisconsin.
Whitson, Andrew Robinson, R. 7, Madison
B.S. (Chicago). Professor of Soils and Drainage, University of Wis¬
consin; Field Agent, United States Department of Agriculture.
Whyte, William F., 1108 Garfield St., Madison
M.D. Physician. President, State Board of Health of Wisconsin.
# Wilson, H. F., 425 Sterling PL, Madison
Professor of Economic Entomology, University of Wisconsin.
#Winchell, Alexander N., 200 Prospect Ave., Madison
B.S. and M.S. (University of Minnesota); D.Sc. (University of Paris).
Professor of Mineralogy and Petrology, University of Wisconsin,
Geologist, Oregon Bureau of Mines and Geology.
744 Wisconsin Academy of Sciences , Arts, and Letters.
*Windesheim, Gustave, 453 Durkee, Kenosha
M.D. Physician.
Wolfenson, Louis B., 1113 W. Dayton St., Madison
Assistant Professor of Hebrew and Hellenistic Greek, University
of Wisconsin.
Woll, Fritz Wilhelm, Berkeley, California
B.S., Ph.B. (Christiana); M.S., Ph.D. (Wisconsin). Professor in the
California State Agricultural College.
Woodmansee, Wilson Robb, 649 S. Grove St., Ripon
A.B. (Indiana); A.M. (Wisconsin). Professor of Mathematics,
Ripon College.
#Worcester, Edward Strong, 148 Langdon St., Madison
A.B. (Princeton); B.D. (Hartford Theological Seminary). Pastor of
the First Congregational Church, Madison.
*Wright, William Harmon, 1905 West Lawn Ave., Madison
B.S. (Purdue); M.S. (Wisconsin); M.S. (Cornell). Assistant Professor
of Agricultural Bacteriology, University of Wisconsin.
*Yeaton, Chester HJenry, 373 Broadway, Milwaukee
A.B. (Bowdoin); A.M. (Harvard); Ph.D. (Chicago). Professor of
Mathematics, School of Engineering, Milwaukee.
Young, Karl, 619 N. Frances St., Madison
A.B. (Michigan); A.M. and Ph.D. (Harvard). Professor of English,
University of Wisconsin.
Zdanowicz, Casimir Douglass, Madison
Associate Professor of Romance Languages, University of Wisconsin.
ZlMMERMANN, OLIVER BRUNNER,
International Harvester Corporation, Chicago, Ill.
B.S., M.E. (Wisconsin). International Harvester Corporation.
I
List of Members.
745
CORRESPONDING MEMBERS
Abbott, Charles Conrad, Trenton, N. J.
M.D. (Pennsylvania).
Armsby, Henry Prentiss, State College, Pa.
B.S. (Worcester Polytechnic); Ph.B., Ph.D. (Yale); LL.D. (Wiscon¬
sin). Director of Institute of Animal Nutrition; Expert in Ani¬
mal Nutrition, United States Department of Agriculture.
Bennett, Charles Edwin, 1 Grove Place, Ithaca, N. Y.
A.B., Litt.D. (Brown). Professor of Latin Language and Literature,
Cornell University.
Bridge, Norman, Auditorium Building, Los Angeles, Cal.
A.M. (Lake Forest); M.D. (Northwestern, Rush). Emeritus
Professor of Medicine, Rush Medical College. Physician.
Coulter, John Merle, University of Chicago, Chicago, Ill.
A.B., A.M., Ph.D. (Hanover); Ph.D. (Indiana). Professor of Botany
and Head of Department, University of Chicago.
Crooker, Joseph Henry,
820 South St., Roslindale, Boston, Mass.
D.D. (St. Lawrence, Nashville). Minister, Unitarian Church.
Davis, Floyd,
317 Iowa Loan and Trust Building, Des Moines, Iowa
Ph.B., C.E., E.M. (Missouri); Ph.D. (Miami). Analytical and
Consulting Chemist.
Eaton, Edward Dwight, Beloit
A.B., A.M. (Beloit) ; B.D. (Yale) ; LL.D. (Wisconsin) ;
D.D. (Northwestern, Yale).
Eckels, William Alexander, Easton, Pa.
A.B., A.M. (Dickinson); Ph.D. (Johns Hopkins). Associate
Professor of Greek, Lafayette College.
Fallows, Samuel, 2344 Monroe St., Chicago, Ill.
A.B., A.M., LL.D. (Wisconsin); D.D. (Lawrence, Marietta).
Presiding Bishop, Reformed Episcopal Church; President,
Board of Managers, Illinois State Reformatory.
746 Wisconsin Academy of Sciences , Arts, and Letters.
Hendrickson, George Lincoln,
68 Trumbull St., New Haven, Conn.
A.B. (Johns Hopkins); L.H.D. (Western Reserve). Professor of
Latin, Yale University.
Hoskins, Leander Miller, 365 Lincoln Ave., Palo Alto, Cal.
M.S., C.E. (Wisconsin). Professor of Applied Mathematics,
Leland Stanford Jt. University.
Iddings, Joseph Paxon, 5730 Woodlawn Ave., Chicago, Ill.
Ph.B. (Yale). Professor of Petrology, University of Chicago;
Geologist, United States Geological Survey.
Kinley, David, Urbana, Ill.
A.B. (Yale) ; Ph.D. (Wisconsin). President, University of Illinois.
Leverett, Frank, 312 N. Thayer St., Ann Arbor, Mich.
B.Sc. (Iowa Agricultural). Geologist, United States Geological
Survey; Lecturer in Geology, University of Michigan.
Libby, Orin Grant, Grand Forks, N. D.
B.L., M.L. (Wisconsin). Professor of History, University of North
Dakota, State Historical Society of North Dakota.
Lurton, Freeman Ellsworth, Fergus Falls, Minn.
B.S., M.S. (Carleton) ; A.M. (Upper Iowa); Ph.D. (Gale). Superin¬
tendent of Public Schools; Member, Board of Directors,
Fergus Falls Public Library.
Luther, George Eimer,
262 South College Ave., Grand Rapids, Mich.
Cashier, People’s Savings Bank; Treasurer, Historical Society of
Grand Rapids.
Marx, Charles David, Palo Alto, Cal.
B.C.E. (Cornell); C.E. (Karlsruhe). Professor of Civil Engineering,
Leland Stanford Jr. University.
McClumpha, Charles Flint, 56 Church St., Amsterdam, N. Y.
A. B., A.M. (Princeton); Ph.D. (Leipzig). Treasurer, McClumpha
Company; Member, Fort Johnson Club; Treasurer, Amsterdam
Free Library; Historian, Montgomery County Historical
Society; Member, New York State Historical Society.
Moorehouse, George Wilton, 2069 East 96th St., Cleveland, 0.
B. L. M.L. (Wisconsin); M.D. (Harvard). Physician to the Dis¬
pensary of Lakeside Hospital and Western Reserve University.
List of Members.
747
Munro, Dana Carlton, Princeton, N J.
A.B., A.M. (Brown). Professor of History, Princeton University.
Olive, Edgar W., Brooklyn, N. Y.
Curator, Brooklyn Botanic Garden.
Potter, William Bleecker, 1225 Spruce St., St. Louis, Mo.
A.B., A.M., M.E., Sc.D. (Columbia). Mining- Engineer and
Metallurgist.
Power, Frederick Belding, 535 Warren St., Hudson, N. Y.
Ph.G. (Philadelphia College of Pharmacy); Ph.D. (Strassburg).
Director of Wellcome Chemical Research Laboratories,
London, England.
Salisbury, Rollin D., 5730 Woodlawn Ave., Chicago, Ill.
A.M., LL.D. (Beloit). Professor of Geographic Geology, Head of the
Department of Geography and Dean of the Graduate School
of Science, University of Chicago; Geologist, United States
Geological Survey and State Geological Survey
of New Jersey.
Stone, Ormond, University Station, Charlottesville, Ya.
A.M. (Chicago). Director of the Leander McCormick Observatory and
Professor of Practical Astronomy, University of Virginia.
Tolman, Albert Harris, 5750 Woodlawn Ave., Chicago, Ill.
A.B. (Williams); Ph.D. (Strassburg). Professor of English
Literature, University of Chicago.
Tolman, Herbert Cushing, Nashville, Tenn.
A. B., Ph.D. (Yale); D.D. (Nashville). Professor of Greek, and Dean.
Vanderbilt University; Canon, All Saints’ Cathedral.
Townley, Sidney Dean, Ukiah, Cal.
B. S., M.S. (Wisconsin); Sc.D. (Michigan). Astronomer in Charge of
International Latitude Observatory; Lecturer in Astronomy,
University of California; Editor of Publications, Astro¬
nomical Society of the Pacific.
Turner, Frederick Jackson, Cambridge, Mass.
A.B., A.M. (Wisconsin) ; Ph.D. (Johns Hopkins); LL.D. (Illinois);
Litt.D. (Harvard). Professor of American History, Harvard
University; President, American Historical Association;
Member, Massachusetts Historical Association; Ameri¬
can Antiquarian Society; Colonial Society of Mas¬
sachusetts; Wisconsin Historical Society; Mis¬
sissippi Valley Historical Society, etc.
748 Wisconsin Academy of Sciences , Arts , and Letters.
Van de Warker, Ely, 404 Fayette Park, Syracuse, N. Y.
M.D. (Albany Medical and Union). Surgeon, Central New York
Hospital for Wom.en; Consulting Physician, St. Ann's Mater¬
nity Hospital; Senior Surgeon, Women’s and Children’s
Hospital; Commissioner of Education, Syracuse.
Verrill, Addison Emery, 86 Whalley Aye., New Haven, Conn.
B.S. (Harvard); A.M. (Yale). Professor of Zoology, Yale Univer¬
sity; Curator of Zoology, Yale University Museum; President,
Connecticut Academy of Arts and Sciences.
Young, Albert Adams, 531 South Claremont Ave., Chicago, Ill.
A.B., A.M. (Dartmouth); B.D. (Andover). Clergyman.
MEMBERS DECEASED.
Information of whose decease has been received since the issue of
Volume XIX.
Blackstone, Dodge Pierce, Berlin
Deceased , November 4, 1919.
D aland, William Clifton, Milton
President of Milton College.
Deceased , June 21, 1921.
Finger, Wiliam, Milwaukee
Broker.
Deceased , March 28, 1917
Harper, Edward T., Geneseo, Ill.
Deceased , January 24, 1921
Kunz, Herman W., Milwaukee
Teacher of Biology.
Deceased , 1918
Lawson, Publius V., Menasha
Manufacturer.
Deceased , December 1, 1920
McLenegan, Charles E.
Librarian.
Milwaukee
Deceased, March 17, 1920
List of Members.
749
Morris, William Augustus Pringle, Madison
Lawyer.
Deceased, September 15, 1920
Nader, John, Rosebank, N. Y.
Architect and Civil Engineer.
Deceased , June 29, 1919
Van Hise, Charles Richard Madison
President, University of Wisconsin.
Deceased , November 19, 1918
Vilas, Charles H., Madison
Retired Physician.
Deceased, December 23, 1920
CHARTER OF THE ACADEMY
An Act to incorporate the Wisconsin Academy of Sciences, Arts and
Letters.
The people 0/ the state of Wisconsin, represented in. senate and assem¬
bly, do enact as follows:
Section 1. Lucius Fairchild, Nelson Dewey, John W. Hoyt, Increase
A. Lapham, Alexander Mitchell, Wm. Pitt Lynde, Joseph Hobbins, E. B.
Wolcott, Solon Marks, R. Z. Mason, G. M. Steele, T. C. Chamberlin, James
H. Eaton, A. L. Chapin, Samuel Fallows, Charles Preusser, Wm. E.
Smith, J. C. Foye, Wm. Dudley, P. Engelmann, A. ,S. McDill, John Mur-
rish, Geo. P. Delaplaine, J. G. Knapp, S. V. Shipman, Edward D. Holton,
P. R. Hoy, Thaddeus C. Pound, Charles E. Bross, Lyman C. Draper, John
A. Byrne, 0. B. Smith, J. M. Bingham, Henry Bsetz, LI. Breese, Thos. S.
Allen, S. S. Barlow, Chas. R. Gill, C. L. Harris, J. C. Squires, George Reed,
J. G. Thorp, William Wilson, Samuel D. Hastings, and D. A. Baldwin,
at present being members and officers of an association known as “The
Wisconsin Academy of Sciences, Arts, and Letters,” located at the city
of Madison, together with their future associates and successors forever,
are hereby created a body corporate by the name and style of the “Wis¬
consin Academy of Sciences, Arts, and Letters,” and by that name shall
have perpetual succession ; shall be capable in law of contracting and being
contracted with, of suing and being sued, of pleading and being impleaded
in all courts of competent jurisdiction; and may do and perform such
acts as are usually performed by like corporate bodies.
Section 2. The general objects of the Academy shall be to encourage
investigation and disseminate correct views in the various departments
of science, literature, and the Arts. Among the specific objects of the
Academy shall be embraced the following;
1. Researches and investigations in the various departments of the
material, metaphysical, ethical, ethnological, and social sciences.
2. A progressive and thorough scientific survey of the state with a
view of determining its mineral, agricultural, and other resources.
3. The advancement of the usual arts, through the applications of
science, and by the encouragement of original invention.
4. The encouragement of the fine arts, by means of honors and prizes
awarded to artists for original works of superior merit.
5. The formation of scientific, economic, and art museums.
751
752 Wisconsin Academy of Sciences , Arts , and Letters.
6. The encouragement of philological and historical research, the
collection and preservation of historic records, and the formation of a
general library.
7. The diffusion of knowledge by the publication of original contri¬
bution to science, literature, and the arts.
Section 3. iSaid Academy may have a common seal and alter the
same at pleasure; may ordain and enforce such constitution, regula¬
tions, and by-laws as may be necessary, and alter the same at pleasure;
may receive and hold real and personal property, and may use and
dispose of the same at pleasure; provided, that it shall not divert any
donation or bequest from the uses and objects proposed by the donor,
and that none of the property acquired by it shall, in any manner, be
alienated other than in the way of exchange of duplicate specimens,
books, and other effects, with similar institutions and in the manner
specified in the next section of this act, without the consent of the
legislature.
Section 4. It shall be the duty of the said Academy, so far as the
same may be done without detriment to its own collections, to furnish,
at the discretion of its officers, duplicate typical specimens of objects
in natural history to the University of Wisconsin, and to the other
schools and colleges of the state.
Section 5. It shall be the duty of said Academy to keep a careful
record of all its financial and other transactions, and, at the close of each
fiscal year, the President thereof shall report the same to the Governor of
the State, to be by him laid before the Legislature.
Section 6. The constitution and by-laws of said Academy now in force
shall govern the corporation hereby created, until regularly altered or
repealed; and the present officers of said Academy shall be officers of
the corporation hereby created, until their respective terms of office shall
regularly expire, or until their places shall be otherwise vacated.
Section 7. Any existing society or institution having like objects
embraced by said Academy, may be constituted a department thereof,
or be otherwise connected therewith, on terms mutually satisfactory to
the governing bodies of the said Academy and such other society or
institution.
Section 8. For the proper preservation of such specimens, books and
other collections as said Academy may make, the Governor shall prepare
such apartment or apartments in the Capitol as may be so occupied with¬
out inconvenience to the State.
Section 9. This act shall take effect and be in force from and after
its passage.
Approved March 16, 1870.
Charter of the Academy.
753
STATUTES OF 1898.
Transactions of the Academy.
Section 341. There shall be printed by the state printer biennially
in pamphlet form two thousand copies of the transactions of the Wis¬
consin Academy of Sciences, Arts, and Letters, uniform in style with
the volumes heretofore printed for said society.
Note. — Under a ruling- of the printing commissioners of the state of Wis¬
consin, made in response to a presentation by a committee of the Academy
appointed December 29, 1897, each volume of the Transactions might be issued
in two consecutive parts ; so that a publication might thus be issued each year
covering the papers accepted after the previous annual meeting. The Academy
allows each author one hundred separate reprints of his paper from the Trans¬
actions without expense, except a small charge for printed covers when desired.
Additional copies are charged for at the actual cost of printing and binding.
Of the Distribution of Public Documents.
Section 365. The transactions of the Wisconsin Academy of Sciences
Arts, and Letters shall be distributed as follows: One copy to each
member of the legislature, one copy to the librarian of each state insti¬
tution; one hundred copies to the State Agricultural Society; one hun¬
dred copies to the State Historical Society; one hundred copies to the
State University, and the remainder to said Academy.
Section 366. In the distribution of books or other packages, if such
packages are too large or would cost too much to be sent by mail, they
shall be sent by express or freight, and the accounts for such express
or freight charges, properly certified to, shall be paid out of the state
treasury.
STATUTES OF 1901
OHAPTEiR 447.
Binding of Exchanges.
Section 1. Section 341 of the revised statutes of 1898 is hereby
amended by adding thereto the following: The secretary of state may
authorize the state printer to bind in suitable binding all periodicals
and other exchanges wihch the Society shall hereafter receive, at a
cost not exceeding one hundred and fifty dollars per annum. The
secretary of state shall audit the accounts for such binding.
STATUTES OF 1921.
Section 35.32. That part of section 35.32 of the statutes relating to
printing for the Wisconsin Academy of Sciences, Arts, and Letters is
amended to read: “of each number as issued, of the transactions of
the Wisconsin Academy of Sciences, Arts, and Letters, not more than
two thousand copies * * * together with suitable binding at a cost
not exceeding one hundred and fifty dollars per annum of all periodi¬
cals and other exchanges which said academy shall hereafter receive.”
48— S. A. L.
754 Wisconsin Academy of Sciences, Arts , and Letters.
CONSTITUTION
OF THE WISCONSIN ACADEMY OF SCIENCES, ARTS, AND
LETTERS.
[As amended at various regular meetings.]
Article I. — Name and Location.
This association shall he known as the Wisconsin Academy of
Sciences, Arts, and 'Letters, and shall he located at the city of Madison.
Article II. — Object.
The object of the Academy shall be the promotion of sciences, arts,
and letters in the state of Wisconsin. Among the special objects shall
be the publication of the results of investigation and the formation of
a library.
Article III. — Membership.
The Academy shall include four classes of members, viz.: life mem¬
bers, honorary members, corresponding members, and active members,
to be elected by balolt.
1. Life members shall be elected on account of special services ren¬
dered the Academy. Life membership in the Academy may also be
obtained by the payment of one hundred dollars and election by the
Academy. Life members shall be allowed to vote and to hold office.
2. Honorary members shall be elected by the Academy and shall be
men who have rendered conspicuous services to science, arts, or letters.
3. 'Corresponding members shall be elected from those who have
been active members of the Academy, but have removed from the state.
By special vote of the Academy men of attainments in science or letters
may be elected corresponding members. They shall have no vote in
the meetings of the Academy.
4. Active members shall be elected by the Academy or the council
and shall enter upon membership on the payment of an initiation fee
of two dollars which shall include the first annual assessment of one
dollar. The annual assessment shall be omitted for the president,
secretary, treasurer, and librarian during their term of office.
Article IV. — Officers.
The officers of the Academy shall be a president, a vice-president for
each of the three departments, sciences, arts and letters, a secretary, a
librarian, a treasurer, and a custodian. These officers shall be chosen
by ballot, on rcommendation of the committee on nomination of officers,
by the Academy at an annual meeting and shall hold office for three
Constitution.
755
years. Their duties shall be those usually performed by officers thus
named in scientific societies. It shall be one of the duties of the presi¬
dent to prepare an address which shall be delivered before the Academy
at the annual meeting at which his term of office expires.
Article V. — Council.
The council of the Academy shall be entrusted with the management
of its affairs during the intervals between regular meetings, and shall
consist of the president, the three vice-presidents, the secretary, the
treasurer, the librarian, and the past presidents who retain their resi¬
dence in Wisconsin. Three members of the council shall constitute a
quorum for the transaction of business, provided the secretary and one
of the presiding officers be included in the number.
Article VI. — Committees.
The standing committees of the Academy shall be a committee on
publication, a library committee, and a committee on the nomination
of members. These committees shall be elected at the annual meeting
of the Academy in the same manner as the other officers of the Academy,
and shall hold office for the same term.
1. The committee on publication shall consist of the president and
secretary and a third member elected by the Academy. They shall
determine the matter which shall be printed in the publications of the
Academy. They may at their discretion refer papers of a doubtful
character to specialists for their opinion as to scientific value and
relevancy.
2. The library committee shall consist of five members, of which the
librarian shall be ex officio chairman, and of which a majority shall not
be from the same city.
3. The committee on nomination of members shall consist of five
members, one of whom shall be the secretary of the Academy.
Article VII. — Meetings.
The annual meeting of the Academy shall be held at such time and
place as the council may designate; but all regular meetings for the
election of the board of officers shall be held at Madison. Summer
field meetings shall be held at such times and places as the Academy
or the council may decide. Special meetings may be called by the
council.
Article VIII. — Publications.
The regular publication of the Academy shall be knoAvn as its Trans¬
actions, and shall include suitable papers, a record of its proceedings,
and any other matter pertaining to the Academy. This shall be printed by
the state as provided in the statutes of Wisconsin. All members of the
Academy shall receive gratis the current issues of its Transactions,
the Transactions.
756 Wisconsin Academy of Sciences, Arts, and Letters.
Article IX. — Amendments .
Amendments to this constitution may be made at any annual meeting
by a vote of three-fourths of all the members present; provided, that
the amendment has been proposed by five members, and that notice has
been sent to all the members at least one month before the meeting.
RESOLUTIONS
REGULATIVE OF THE PROCEEDINGS OF THE ACADEMY.
THE TRANSACTIONS OF THE ACADEMY.
[. By the Academy, December 28, 1882.]
2. The secretary of the Academy shall be charged with the special
duty of overseeing and editing the publication of future volumes of
the Trancactions.
3. The Transactions of the Academy hereafter published shall con¬
tain: (a) a list of officers and members of the Academy; (b) the
charter, by-laws and constitution of the Academy as amended to date;
(c) the proceedings of the meetings; and (d) such papers as are duly
certified in writing to the secretary as accepted for publication in ac¬
cordance with the following regulations, and no other.
6. In deciding as to the papers to be selected for publication, the
committee shall have special regard to their value as genuine, original
contributions to the knowledge of the subject discussed.
9. The sub-committee on publication shall be charged with insisting
upon the correction of errors in grammar, phraseology, etc., on the
part of authors, and shall call the attention of authors to any other
points in their papers which in their judgment appear to need revision.
[ By the Academy, June 2, 1892.]
The secretary was given authority to allow as much as ten dollars
for the illustration of a paper when the contribution was of sufficient
value to warrant it. A larger amount than this might be allowed by the
committee on publication.
[ By the Academy, December 29, 1896.]
The secretary was directed to add to the date of publication as printed
on the outside of author’s separates the words, “Issued in advance of
general publication.”
Resolutions.
757
Fees of Life Members.
[By the Academy , July 19, 1870.]
Resolved, That the fees from members for life be set apart as a per¬
manent endowment fund to be invested in Wisconsin state bonds, or
other equally safe securities, and that the proceeds of said fund, only,
be used for the general purposes of the Academy.
Annual Dues.
[By the Academy, December 29, 1892.]
Resolved, That the secretary and treasurer be instructed to strike
from the list of active members of the Academy the names of all who
are in arrears in the payment of annual dues, except in those cases
where, in their judgment, it is desirable to retain such members for a
longer time.
Arrears of Annual Dues.
[By the Council, December 29, 1897.]
Resolved, That the treasurer be requested to send out the notices of
annual dues as soon as possible after each annual meeting and to ex¬
tend the notice to the second or third time within a period of four
months where required.
Secretary’s Allowance.
[By the Academy, December 27, 1902.]
Resolved, That the Academy hereby appropriates the sum of seventy-
five dollars per annum as an allowance for secretary’s expenses, for
which a single voucher shall be required.
Secretary’s Allowance.
[By the Council, April 5, 1912.]
Resolved, That the Academy appropriates the sum of two hundred
dollars per annum for the secretary-treasurer’s allowance.
Election of Mrs. Lucius Fairchild as Honorary Member.
[By the Council, April 12, 1918.]
Resolved, That because of the honorable and leading part that was
played by the late General Fairchild in the founding of this Academy,
his widow, Mrs. Lucius Fairchild, of Madison, be voted an honorary
member of the Wisconsin Academy of Sciences, Arts, and Letters.
GENERAL INDEX OF AUTHORS OF PAPERS
IN THE BULLETINS AND TRANSACTIONS OF THE
ACADEMY
Bulletins 1—5.
Transactions, Vols. I— XX.
Allen, Charles E. Some Hepaticae of the Apostl© Islands. . . .XIV., 484
Allen, Ruth F. Studies in spermatogenesis and apogamy of
ferns . XVII., 1
— — See also Jolivette, Hallie D. M.
Allen, William F. The rural population of England as classified
in Domesday Book......... . . . ....... I., 167
- The Rural classes of England in the 13th century. ..... .II., 220
- Ranks and classes among the Anglo-Saxons _ ... _ ..II., 234
- United States’ sovereignty: whence derived and when
vested . . III., 125
- - Peasant communities in France . . . . .... IV., 1
- The Origin of the Freeholders . . IV., 19
- The English cottagers of the Middle Ages . . . .V., 1
- The primitive democracy of the Germans. . . .VI., 28
- Village community and serfdom in England _ ... - .VII., 130
- Town, township, and tithing. . . .VII., 141
Andrews, Edmund. Discoveries illustrating the literature and re¬
ligion of the Mound Builders . ...IV., 126
Andrews, Joy E. Some experiments with the larva of the wax
moth, Galleria mellonella L . XX., 255
Armitage, Right Rev. W. E. The German iSunday . . . I., 62
Atwood, William Henry. The visceral anatomy of the garter
snake. (With five figures) . XIX., 531
Austin, Louis W. and C. W. Eastman. On the relation between heat
conductivity and density in some of the common
woods . . XIII., 539
Baker, Frank Collins. The molluscan fauna of Tomahawk lake,
Wisconsin . . .XVII., 200
Balg, G. H. The science of the English language in the light of the
Gothic . . . . VIII., 187
Barnes, Charles R. Artificial keys to the genera and species of
mosses recognized in Lesquereux and James’s Manual of
the Mosses of North America . . . VIII., 11
759
760 Wisconsin Academy of Sciences , Arts , and Letters .
Barnes, Charles R. Artificial keys to the genera and species of
mosses recognized in Lesquereux and James’s manual of
the Mosses of North America — Additions and Correc¬
tions . VIII.,
Barnett, James D. The state administration of taxation in Wis¬
consin . XV.,
Bartlett, James L. The cold-waves of south-central Wisconsin. XVI.,
Bascom, John. Freedom of will empirically considered . VI.,
Beatty, Arthur. The St. George, or Mummers’ plays: A study in
the protology of the drama . XV.,
Bennett, Chas. E. Some new theories of the Greek KA-Perfect.VIIl.,
Bille, John H. A history of the Danes in America . XI.,
Birge, Edward A. Notes on Cladocera . IV.,
- On the motor ganglion cells of the frog’s spinal cord _ VI.,
- List of Crustacea cladocera from Madison, Wisconsin _ VIII.,
- Notes on Cladocera, III . IX.,
- O. A. Olson, and H. P. Harder. Plankton studies on Lake
Mendota . X.,
- Plankton studies on Lake Mendota. II. The Crustacea of
the Plankton from July, 1894, to December, 1896 . XI.,
- and C. Juday. A summer resting stage in the development of
Cyclops bicuspidatus Claus . XVI.,
- An unregarded factor in lake temperature . XVI.,
- On the evidence for temperature seiches . XVI.,
- Notes on Cladocera. IV . XVI.,
- The heat budgets of American and European lakes. With
three Figures, and five Tables . XVIII.,
- The work of the wind in warming a lake. (With Plates
I— X) . XVIII.,
- A second report on limonological apparatus. (With Plates
XXXIX-XL, and five Figures) . . XX.,
- The Academy Medallion . XX.,
- See also Juday, Chancey.
Blackstone, ID. P. The variation in the attraction due to the figure
of the attracting bodies . VI.,
Blaisdell, J. J. Some suggestions concerning methods of psycho¬
logical study . IX.,
- The methods of science, as being in the domain of logic. .XI.,
Blake, William P. The progress of geological surveys in the State
of Wisconsin. A review and bibliography . IX.,
- Arizona diatomite . XIV.,
Brown, Chas. E. Wisconsin’s quartzite implements . .XV.,
Brues, Charles T. and Beirne B. The grasses of Milwaukee county,
Wisconsin . XVII.,
Bruncken, Ernest. The ues of parties in municipal government. XI.,
- On some differences between private and public business. XII.,
Buck, Solon J. The settlement of Oklahoma . XV.,
Buckley, E. R. Ice ramparts . XIII.,
163
163
289
2
273
141
1
77
51
379
275
421
274
1
989
1005
1017
166
341
533
711
197
33
49
225
107
656
57
225
325
325
141
General Index of Authors of Papers.
761
Buckley, E. R. Sleet storm in the Ozark region of Missouri . . .XVI., 307
Buell, Ira M. The corals of the Delafleld . . . V., 185
— — Geology of the Waterloo quartzite area . IX., 255
- Boulder trains from the outcrops of the Waterloo quartzite
area . X., 485
Bundy, W. F. A List of the Crustacea of Wisconsin . . . .V., 177
Burd, Henry A. Eight unedited letters of Joseph Ritson . .XIX., 1
Butler, James D. The naming of America . II., 203
- Copper-tools found in the state of Wisconsin . . III., 99
- First French foot-prints beyond the Lakes . V., 85
- The “'Aitofe hEydooera” of iShakespeare . . V., 161
- Household words: their etymology . XIII., 366
- Phases of witticism . X., 41
- Dante, His quotations and his originality: The greatest imi¬
tator and the greatest original. . XI., 149
— - Codfish. Its place in American history . XI., 261
- Personal names: Their signficance and historical origin.XIII., 475
- The vocabulary of Shakespeare . XIV., 40
Cahn, Alvin R. Notes on the vertebrate fauna of Houghton and
Iron counties, Michigan. (With Plates I-V) . XIX., 483
Carlton, G. P. See Miller, W. S.
Carpenter, Prof. S. H. The metaphysical basis of science . II., 23
- The philosophy of evolution . II., 39
Caverno, Rev. Charles. Abstract of a paper on the relations be¬
tween social and moral science . Bulletin IV., 59
- iSocial science and woman suffrage . I., 72
- The people and the railroads . III., 143
- The abolition of the jury system . IV., 7
- Life insurance, savings banks, and the industrial situation. V., 21
Chamberlin, Thomas C. Suggestions as to the basis for the grada¬
tion of vertebrata. (Abstract.) . . . Bulletin V., 76
- On the 17-year cicada, its geographical distribution and time
of appearance in this State. (No Abstract.) . .Bulletin V., 81
- Facts relating to the local geology of the Whitewater region.
(Abstract.) . Bulletin V., 81
- Suggestions as to a basis for the gradation of the verte¬
brata . I., 138
- Some evidences bearing upon the method of the upheaval of
the quartzites of Sauk and Columbia counties . II., 129
- On fluctuations in level of the quartzites of Sauk and Co¬
lumbia counties . II.» 133
- On the extent and significance of the Wisconsin kettle mo¬
raine . IV., 201
- On a proposed system of lithological nomenclature . ...V., 234
- Observations of the recent glacial drift in the Alps . V., 258
- Some additional evidences bearing on the interval between
the glacial epochs . VIII., 82
762 Wisconsin Academy of Sciences, Arts, and Letters .
Chamberlin, Thomas C. The founding of the Wisconsin Academy
of Sciences, Arts, and Letters . . XX., 693
Chandler, Charles H. Notes and a query concerning the
Ericaceae . VIII., 161
- An improved harmonograph . X., 61
- Transcendental space . XI., 239
- An historical note on early American railways . XII., 317
- The inter-generation period . XII., 499
— — A problem of longevity . XIII., 384
- A study in longevity . XIV., 56
- See also, Doudna, P. E.
Chapin, A. L. The relation of labor and capital . . . I., 45
- The nature and functions of credit . V., 57
Chase, Ruth. The length of life of the wax moth, Galleria rnellon-
ella L., in its different stadia. (With one Figure) . .XX., 263
Cheney, L. S. and R. H. True. On the flora of Madison and vicin¬
ity, a preliminary paper on the flora of Dane County, Wis¬
consin . IX., 45
- A Contribution to the flora of the Lake Superior Region. .IX., 233
- Spaghna of the upper Wisconsin valley . X., 66
- Parasitic fungi of the Wisconsin valley . X., 70
- Hepaticae of the Wisconsin Valley . X., 70
Christman, A. H. Observations on the wintering of grain rusts. .XV., 98
- The nature and development of the primary uredospore. .XV., 517
Collie, George L. Geology of Conanicut Island, R. I . X., 199
Comstock, G. C. The present condition of the latitude prob¬
lem . VIII., 229
Comstock, Elting H. The real singularities of harmonic curves
of three frequencies . XI., 452
Congdon, Russell T. Saskatchewan birds . . . XIV., 569
Copeland, Edwin iB. and Louis Kahlenberg. The influence of the
presence of pure metals upon plants . XII., 454
Crooker, Erin E. Aluminum alcoholates . XI., 255
Culver, G. E. Notes on a little known region of northwestern
Montana . VIII., 187
Culver, G. E. and Wm. H. Hobbs. On a new occurrence of olivine
diabase in Minnehaha County, South Dakota . VIII., 206
Culver, G. E. Some New Jersey eskers . X., 19
- The erosive action of ice . X., 339
Daniells, W. W. Note on the absorption of arsenic by the human
liver . II.* 128
- On the analysis of the water-flowing artesian well at Mari¬
nette, Wisconsin . XI., 112
Davies, John E. Abstract of a paper on the importance and practi¬
cability of finding a unit of force in physics that shall be
of universal application . Bulletin II., 34
- On the kinetic measures of force. (No Abstract.). Bulletin IV., 67
General Index of Authors of Papers.
763
Davies, John N. On potentials, and their application in physical
science . . . I,
— — Recent progress in theoretical physics. . . III.,
— — - Recent progress in theoretical physics. Part II. Magnetic
rotatory polarization of light. ...................... .IV.,
— — On some analogies between the equations of elasticity and
electro-magnetism . . IX.,
Davis, J. J. A supplementary list of parasitic fungi of Wiscon¬
sin . . . . . . 1 ......... . IX.,
- - Second supplementary list of parasitic fungi of Wisconsin. XI.,
- - Third supplementary list of parasitic fungi of Wisconsin.XIV.,
— — Mycological narrative of a brief journey through the Pacific
Northwest . . . . . . . . . . . .XV.,
— — The Academy: Its past and future - ...... - ....... .XV.,
- Fourth supplementary list of parasitic fungi of Wisconsin. . . .
. . XVI.,
- - Notes on parasitic fungi in Wisconsin. . . .XVIII.,
— — Notes on parasitic fungi in Wisconsin. .............. .XVIII.,
- Notes on parasitic fungi in Wisconsin — III. . . . . . XVIII. ,
- North American ascochytae . . .XIX.,
- Notes on parasitic fungi in Wisconsin, IV. (With two Fig¬
ures) . . .XIX.,
— — Notes on parasitic fungi in Wisconsin, V . .XIX.,
- Notes on parasitic fungi in Wisconsin, VI . .XIX.,
- Notes on parasitic fungi in Wisconsin, VII. (With Plates
XXX-XXXII and one Figure) . . . .XX.,
- Notes on parasitic fungi in Wisconsin, VIII. (With three
Figures) . XX.,
Day, F. H. On the fauna of the Niagara and upper iSilurian Rocks,
as exhibited in Milwaukee county, Wisconsin, and in
counties contiguous thereto. . . IV.,
De Hart, J. N. The antiquities and platycnemism of the mound
builders of Wisconsin. . . .IV.,
Denniston, Rollin ,H. The Russulas of Madison and vicinity. . .XV.,
- The structure of the starch grain. . . . . ..... .XIV.,
- The growth and organization of the starch . .XV.,
- A Survey of the larger aquatic plants of Lake Mendota.
(With one Figure) . . XX.,
Desmond, Humphrey J. The sectional feature in American poli¬
tics . . ......VIII.,
Doudna, P. E. and E. F. Chandler. On the errors with which loga¬
rithmic computations are affected.. . XIII.,
Downes, Robert H. Economic and social development of Kenosha
county . . .XIII.,
Du Mez, Andrew G. History of the United States pharmacopoeia, I.
The Galenical Oleoresins .... . .XIX.,
Eastman, C. W. See Austin, Louis W.
Eaton, James H. On the formation of certain new compounds of
manganese. (Abstract.) . . .Bulletin II.,
155
205
241
3
153
165
83
775
887
739
78
93
251
653
671
690
705
399
413
113
188
71
527
664
495
1
427
545
907
35
764 Wisconsin Academy of Sciences, Arts , and Letters .
Eaton, James H. On the geology of the region about Devil’s Lake,
iSauk County, Wisconsin; being a report of observations
made at the request of the Academy. (Abstract) .
. Bulletin IV.,
- Report on the geology of the region about Devil’s Lake. _ I.,
Relation of the sandstones, conglomerates and limestones of
Baraboo valley to each other and to the axoic quartz¬
es . .
Elmendorf, Prof. J. J. Nature and the supernatural . V.,
- Aristotle’s physics . VIII.,
- Nature and freedom . jy
Engelmann, Mrs. P. On the importance of more attention to the
preservation and culture of forest trees in Wisconsin.
(Abstract.) . Bulletin II.,
- - Some Observations upon the fauna of Mammoth Cave. (Ab¬
stract.) . Bulletin III.,
Everest, Kate A. Early Lutheran immigration to Wisconsin. .VIII.,
Feinberg, S. M., and Mavor, J. W. Lymphocystis vitrei, a new
sporozoan from the pike-perch, Stizostedion vitreum
Mitchell. (With Plate IX.) . XIX.,
Feuling, John iB. On the place which the study of the Indian
languages should hold in ethnology. (No Abstract.)
. . . Bulletin VI.,
— — Of the place of the Indian languages in the study of Eth¬
nology . I.,
- The Etymology of church . . . II.,
- Studies in comparative grammar . III.,
Fillmore, John C. Some forms spontaneously assumed by folk¬
songs . XI.,
Finch, A. J. On metallic veins and the deposition of minerals.
(Abstract.) . Bulletin IV.,
Fish, Carl Russell. Table illustrating the progress of rotation in
office to 1835 . XV.,
Fluke, Charles L. Syrphidae of Wisconsin. (With Plates
V-VI) . XX.,
Flint, A. S. Note on the progress of meridian transit observations
for stellar parallax at the Washburn Observatory . X.,
Folkmar, Daniel. The duration of school attendance in Chicago
and Milwaukee . XII.,
55
124
123
€6
169
62
30
39
288
559
68
178
82
117
119
50
709
215
64
257
Gifford, Elizabeth M. and Peckham, Geo. W. Temperature of Pine,
Beaver, and Okanchee Lakes. . . V., 273
Gilbert, Edward M. Studies on the tremellineae of Wisconsin. XVI., 1136
- Cytological Studies of the lower basidiomycetes. I. Dac-
rymyces. (With Plate XXIX) . XX., 387
Graenicher, S. The relation of the Andrenine bees to the ento-
mophilous flora of Milwaukee county . XV., 89
General Index of Authors of Papers.
765
Gregory, Charles N. Political corruption and English and Amer¬
ican laws for its prevention . X., 262
Gregory, John G. Negro suffrage in Wisconsin . . . XI., 94
Grossenbacher, J. G. The periodicity and distribution of radial
growth in trees and their relation to the development of
“annual” rings . . . XVIII., 1
Grundtvig, F. L. On the birds of Shiocton in Bovina, Outagamie
county, Wisconsin, 1881-3 . X., 73
Hadden, Clarence B. History of early banking in Wisconsin _ X., 159
Hardenberg, C. B. Comparative studies in the trophi of the
Scarabaeidae . XV., 548
Harder, H. P. iSee Birge, E. A.
Harper, Edward T. The pholiota of the region of the Great Lakes
. XVII., 470
- - Additional Species of pholiota, stropharia, and hypholoma in
the region of the Great Lakes. (With Plates XI-XXVI).
. XVIII., 392
- Species of lentinus in the region of the Great Lakes. (With
Plates XIV-XXVIII) . XX., 365
Harper, Robert A. Nuclear phenomena in certain stages in the
development of the smuts . XII., 475
- See also Holden, R. J.
Harring, H. K., and Myers, F. J. The rotifers of Wisconsin. (With
Plates XLI-LXI, and four Figures.) . XX., 553
Hastings, Hon. S. D. The common jail system of the country - 1., 90
Higley, W. K. Reptilia and batrachia of Wisconsin . VII., 155
Hillyer, Homer W. On the action of aluminum chloride on satu¬
rated hydrocarbons . X., 367
Hobbs, Wm. ,H. Notes on a trip to the Lipari Islands in 1889. . .IX., 21
- On Some metamorphosed eruptives in the crystalline rocks of
Maryland . VIII., 156
- The correlation of fracture systems and the evidences of plan¬
etary dislocations within the earth's crust . XV., 15
— — ■ See also, Culver, G. E.
Hoffman, Alice E., and Schuette, H. A. Notes on the chemical com¬
position of some of the larger aquatic plants of Lake
Mendota. I. Cladophora and Myriophyllum . XX., 529
Holden, R. J. and R. A. Harper. Nuclear divisions and nuclear
fusion in Coleosporium sonchi-arvensis, Lev . XIV., 63
Holmes, S. J. Description of a new subterranean amphipod from
Wisconsin . XVI., 77
- Description of a new species of eubranchipus from Wisconsin,
with some observations on its reaction to light . XVI., 1252
Holland, Rev. F. M. Vexed questions in ethics . II., 35
- Records of marriages . II., 72
- Industrial education . ..III., 136
- The boa-constrictor of politics . .III., 151
766 Wisconsin Academy of Sciences , Arts, and Letters.
Holland, Rev. F. M. Were the stoics utilitarians? . III., 179
Hoskins, L. M. Maximum stresses in bridge members . X., 24
Hoy, Dr. P. R. Abstract of a paper on the fauna of the Lake Mich¬
igan off Racine . Bulletin II., 34
- The mammalia of Wisconsin. (Abstract.) . .Bulletin IV., 62
- - Deep water fauna of Lake Michigan . I., 98
- Insects injurious to agriculture — Aphides . I., 110
- Natural history as a branch of elementary Education . II., 105
- iSome of the peculiarities of the fauna of Racine . II., 120
- Fish culture . III., 37
- - Extent of the Wisconsin fisheries . III., 65
- The catocalae of Racine County . III., 96
- How did the aborigines of this country fabricate copper im¬
plements? . IV., 132
- Why are there no upper incisors in the ruminantia? . IV., 147
- Menobranchus lateralis . V., 248
- The larger wild animals that have become extinct in Wis¬
consin . V., 248
- Who built the mounds? . VI., 84
— — - Who made the ancient copper implements? . VI., 101
Hoyt, Dr. John W. Requisites to a reform of the civil service of the
United States . II., 89
- The Formal commendation of government officials . III., 133
— — The revolutionary movement among women . III., 161
- Some personal recollections of Abraham Lincoln . XVI., 1305
Hubbel, Herbert P. An examination of Prof. S. H. Carpenter’s
theory of evolution . III., 196
Huntington, Ellsworth. Experiments with available road-making
materials of southern Wisconsin . XI., 249
Irving, Roland. On the age of quartzites, schists, and conglomerates
of Sauk County . I., 129
- On some points in the geology of northern Wisconsin . II., 107
- On a hand specimen showing the exact junction of the pri¬
mordial sandstones and Huronian schists . II., 139
- On the occurrence of gold and silver in minute quantities in
quartz from Clark county. . . . II., 140
- Kaolin in Wisconsin . III., 3
Jegi, John I. Auditory memory-span for numbers in school chil¬
dren . XIV., 509
Jewell, J. S. Mind in the lower animals . . . IV., 164
Johnson, Arden R. Electrolytic production of iodoform . XVI., 253
- The chemistry of boron and some new organic-boron com¬
pounds . XVII., 528
Johnson, Roswell H. Axial bifurcation in snakes . XIII., 523
- The individuality and variation of the pyloric caeca of the
Centrarchidae . XV., 713
General Index of Authors of Papers.
767
Jolivette, Hallie D. M. Spore formation in Geoglossum Glabrum
Pers . . . XVI.,
— — and Ruth F. Allen. A study of the light reactions of philo-
bolus . XVII.,
Jones, Edward D. The relation of economic crises to erroneous
and defective legislation, with special reference to bank¬
ing legislation . . X.,
— — Chartism — a chapter in English industrial history . XII.,
Jones, L. R. Experimental work on the relation of soil temper¬
ature to disease in plants. (With Plates XXXIII-
XXXVII, and four Figures.) . XX.,
Juday, Chauncey. The diurnal movement of plankton Crus¬
tacea . XIV.,
- * Studies on some lakes in the Rocky and Sierra Nevada
mountains . XV.,
- Some aquatic invertebrates that live under anaerobic con¬
ditions . XVI.,
- and George Wagner. Dissolved oxygen as a factor in the
distribution of fishes . XVI.,
- iSome European biological stations . XVI.,
- Limnological apparatus. (With Plates XXXIV-XXXVIII
. XVIII.)
- Limnological studies on some lakes in Central America.
(With four Figures and three Tables.) . .XVIII.,
- Qantitave studies of the bottom fauna in the deeper waters
of Lake Mendota. (With one Figure) . XX.,
- See also Birge, E. A.
Kahlenberg, Louis. Action of metallic magnesium upon aqueous
solutions . XIV.,
- On the nature of the process of osmosis and osmotic pressure
with observations concerning dealysis . XV.,
- Some factors in the progress of scientific research. The Presi¬
dent’s Address, 1909 . XVI.,
- See also Copeland, Edwin B.
Keene, Mary Lucille. Studies of zygospore formation in phycomy-
cetes Nitens Kunze. (With Plates XVI-XVIII).. .XIX.,
King, Charles I. Boiler explosions . IV.,
King, F. ,H. On the suspension of solids in fluids and the nature
of colloids and solutions . XVI.,
Kinley, David. The direction of social reform . IX.,
Knapp, J. G. Abstract of a paper on the coniferae of the Rocky
Mountains, and their adaptation to the soil and climate
of Wisconsin . . . Bulletin II.,
- Abstract of a paper on the isothermal lines of the North¬
west . Bulletin III.,
- On the climatic relations of the flora of Wisconsin. (No
Abstract.) . Bulletin IV.,
1171
533
370
509
000
524
781
10
17
1257
506
214
000
299
209
1289
1195
151
275
137
30
40
62
768 Wisconsin Academy of Sciences , Arts , and Letters.
Knapp, J. G. On the ancient lakes of Wisconsin. (No Abstract.) . .
. Bulletin V., 76
- On the rocks and mines of the upper Wisconsin River. (No
Abstract.) . Bulletin V., 81
- Coniferae of the Rocky Mountains and their adaptation to
the soil and climate of Wisconsin . I., 117
- Ancient lakes of Wisconsin . I., 151
Kremers, Edward. The limonene group of terpenes. . . . VIII., 300
- On the classification of carbon-compounds . X., 310
Krey, A. C. John of Salisbury’s attitude towards the classics. XVI., 948
Kuhl, Ernest P. Chaucer’s burgesses . XVIII., 652
Kumlien, Thure. The rapid disappearance of Wisconsin Wild-
flowers . III., 56
Lap ham, Dr. I. A. On the age of the quartzite of Baraboo. (Ab¬
stract.) . . . . . Bulletin II.,
- On the classification of plants. (Abstract.) _ Bulletin III.,
- On the classification of plants . I.,
- Oconomowoc and other small lakes of Wisconsin . III.,
- Embryonic development the same in plants as in ani¬
mals . III.,
Langenhan, H. A. The arsenical solutions. No. I. Liquor
Potassii* Arsenitis. (Fowler’s Solution) . . .jXX.,
Lawson, Publius V. Thure Kumlien. (With Plates LXII-LXIV)
. XX.,
Legler, Henry E. Origin and meaning of Wisconsin place-names
with special reference to Indian nomenclature - XIV.,
- A Wisconsin group of German poets . XIV.,
Lehmann, Harriet. Variations in forms and size of Cyclops
hrevispinosus Herrick and Cyclops Americanus
Marsh . XIV.,
Lenher, Victor. Fluoride of gold . XIV.,
- and A. W. Homberger. The gravimetric determination of
tellurium . XVI.,
Leverett, Frank. Raised beaches of Lake Michigan . VII.,
- On the correlation of moraines with raised beaches of Lake
Erie . VIII.,
Libby, O. G. Economic and social development of Kenosha and
La Fayette counties— Introduction . XIII.,
- A study of the greenback movement, 1876-84 . XII.,
- Stanton, F. Bolle, Palmer, Bernard M„, and Smith, Allard J.
An economic and social study of the lead region of Wis¬
consin, Illinois, and Iowa . XIII.,
- Some pseudo-histories of the American Revolution. .. .XIII.,
Lincoln, Azariah T. The electrical conductivity of non-aqueous
solutions . ? . XII.,
Loomis, H. B. The effect of changes of temperature on the distri¬
bution of magnetism . . . VIII.,
35
47
102
31
110
141
663
16
471
279
312
1278
177
233
643
530
188
419
395
273
General Index of Authors of Papers.
Lueders, Herman F. Vegetation of the town Prairie du Sac. . . .XX.,
- Floral structure of some gramineae . XI.,
Lurton, Freeman A. A study of retarded children in a group of
Northwestern schools . XVII.,
Lutman, Benjamin F. Some contributions to the life history and
cytology of the smuts . XVI.,
McAllister, Frederick. The cytology of the convallariacae. . .XVII.,
McLeod, Andrew F. The Walden inversion. . XVII.,
McMIurphy, J. G. Rotation as a factor of motion . IV.,
Marquette, W. Concerning the organization of the spore mother-
cells of Marsilia quadrifolia . XVI.,
Marsh, C. Dwight. On the deep water Crustacea of Green
Lake . VIII.,
- Notes on the depth and temperature of Green Lake _ VIII.,
- Cyclopidae and calanidae of central Wisconsin . IXX.,
- On two new species of Diaptomus . X.,
- On the limnetic crustaces of Green Lake . XI.,
- On some points in the structure of the larvae of EpiscJiura
lacustris, Forbes . XII.,
- The plankton of fresh water lakes . XIII.,
- On a new species of canthocamptus from Idaho . XIV.,
- A revision of the North American species of diaptomus. XV.,
- A Revision of the North American species of cyclops. .XVI.,
- (Structural abnormalities in copopeda . XVII.,
- Diaptomus coloradensis, sp. Nov . XVII.,
Marshall, Ruth. Ten species of Arrunuri belonging to the sub-genus
megalurus Thon . XIV.,
- A new Arrenurus . XIV.,
- The vegetation of Twin Island . XVI.
- The American water mites of the genus Neumania. (With
Plates II-IV) . XX.,
Marshall, W. S. Entocy there camftaria. (nov. gen. et nov. spec.
a parasitic ostracod . XIV.,
- and Henry Severin. Some points in the anatomy of Ranan-
tra fusca . XIV.,
- The reproductive organs of the female maia moth, Hemileuca
maia (Drury.) . XV.,
- The development of the frenulum of the wax moth, Galleria
mellonella Linn. (With Plate 10) . XX.,
Mason, R Z. Abstract of a paper on the nebular hypothesis in astron¬
omy . Bulletin 111.,
- On the clay deposits and the fossils found therein, in the
region about Appleton. (Abstract.) . . Bulletin V.,
- The duty of the state in its treatment of the deaf and
dumb, the blind, the idiotic, the crippled and deformed,
and the insane . IV.,
769
510
109
275
1191
599
503
235
81
211
214
189
15
179
544
163
112
381
1067
195
197
145
520
773
205
117
486
1
199
44
73
25
49— S. A. L.
770 Wisconsin Academy of Sciences , Arts, and Letters.
Mavor, James W. and Strasser, William. On a new myxosporidian,
Henneguya Wisconsmensis, n. sp., from the urinary
bladder of the yellow perch, Perea flavescens. With
three Figures) . XVIII., 676
* - and Strasser, W. Studies on myxosporidia from the urinary
bladders of Wisconsin fishes. (With Plates VI-
VIII) . . . XIX., 553
- and Feinberg, S. M. Lymphocystis vitrei, a new sporozoan
from the pike-perch, Stizostedion vitreum Mitchell.
(With Plate) . XIX., 559
Merrell, Edward H. The relation of motives to freedom . XII., 389
Merrell, Harriet B. On Bunops Scutifrons, Birge . IX., 318
Meyer, Balthasar H. The adjustment of railroad rates in Prus¬
sia . XI., 78
- Early general railway legislation in Wisconsin, 1853-
1874 . XII., 337
Miller, W. S. The anatomy of the heart of cambarus . X., 327
- and E. P. Carlton. The relation of the cortex of the cat’s
kidney to the volume of the kidney and an estimation
of the number of glomeruli . X., 525
- Variations in the distribution of the cat Felis domesti-
cus . XIV., 621
Morris, H. H. The preparation of selenic acid . XIX., 869
Munro, Caroline Walker. A preliminary study of the digestive se¬
cretions of pickerel and perch . XX., 269
Munro, Dana Carleton. Some tendencies in history. Presidential
Address, 1915 . XVIII., 695
Murrish, John. Abstract of a paper on the origin of the Potsdam
sandstone. (Abstract.) . Bulletin II., 32
- Abstract of a paper on the results of recent investigations
in the lead regions of Wisconsin . Bulletin IV., 62
- Further results of the examination of the lead region of
Wisconsin. (Abstract.) . .Bulletin V., 80
Muttkowski, R. A. The fauna of Lake Mendota — A qualitative and
qualitative survey with special reference to the in*
sects. (With one Figure) . XIX., 374
Myers, F. J., and Harring, H. K. The rotifers of Wisconsin. (With
Plates XLI— LXI, and four Figures.) . . XX., 553
Nader, John. On the strength of materials as applied to engi¬
neering . II., 153
- The improvement of the mouth of the Mississippi River. III., 84
- Leveling with the barometer . III., 68
- The Tides . V., 207
- A chapter on foundations . V., 282
Nebel, Catherine Elizabeth. The amount of food eaten by the spi¬
der, Aranea sericata . XIX., 524
Nicodemus, W. J. L. On the Wisconsin River improvement . II., 142
- Railway gauges . II., 101
General Index of Authors of Papers.
771
Nicodemus, W. J. L. History of the science of hydraulics . II.,
- The ancient civilization of America . III.,
Nichols, Susie P. The nature and origin of the binucleated cells in
some Basidiomycetes . . . XV.,
Norton, Richard G. An investigation into the breaking of watch
mainsprings in greater numbers in the warm months of
the year than in the cold months . XV.,
Oberholser, Harry C. A review of the plover genus Ochthodromus
Reichenbach and its nearest allies . XIX.,
Oldenhage, Prof. H. Remarks on the descent of animals . IV.,
Olive, Edgar W. Notes on the occurrence of Oscillatoria prolifica
(Greville) Gomont in the ice of Pine Lake, Waukesha
county, Wisconsin . XV.,
- Cytological studies on ceratiomyxa . XV.,
Olson, E. A. See Rirge, E. A.
O’iShea, M. V. The parts of speech in the child’s linguistic develop¬
ment . XV.,
Overton, James Bertram. The organization of the nuclei in the root
tips of podophyllum peltatum. (With Plate VII) _ (XX.,
Owen, Edward T. The meaning and function of thought connec¬
tives . XII.,
- A revision of the pronouns. . . XIII.,
- Interrogative thought and the means of its expression. . .XIV.,
- Hybrid parts of speech; a development of this proposition:
In a single sentence, and though unrepeated, a word may
operate as different parts of speech . XVI.,
- Relations expressed by the passive voice . XVII.,
Paetow, Louis J. The neglect of the ancient classes at the early
medieval universities . XVI.,
Palmer, Bernard M. See Libby, O. G.
Parkinson, J. B. Wealth, capital and credit . V,.
Patten, Harrison E. Action upon metals of solutions of hydrochloric
acid in various solvents . XIV.,
Paxson, Frederic ,L. The railways of the old Northwest before the
Civil War . XVII.,
Payne, Alford. Art as education . IV.,
Peckham, Elizabeth G., and George W., Genera of the family atti-
dae, with a partial synonomy . VI.,
- Attidae of North America . VII.,
- land Wm. H. Wheeler. Spiders of the sub-family lysso-
manae . VII.,
Peckham, George W. and Elizabeth G. The sense of sight in spiders,
with some observations on the color sense . X.,
- Spiders of the Phidippus group of the family Attidae - XIII.,
- New species of the family Attidae from South Africa, with
notes on the distribution of the genera found in the Ethiopian
region . XIV.,
193
58
30
654
511
138
124
753
178
275
1
1
353
107
77
311
46
316
243
31
255
1
222
231
282
173
772 Wisconsin Academy of Sciences, Arts, and Letters.
Peckham, George W. and Elizabeth G. The Attidae of Borneo. .XV., 603
- [Revision of the Attidae of North America . . . XVI., 355
Peckham, George W. See also Gifford, Elizabeth M.
Peet, Stephen D. Primitive architecture in America . V., 290
- Ancient villages among emblematic mounds. . VI., 154
- The so-called elephant mound in Grant County and effigies in
the region surrounding it . VII., 205
- The clan centers and clan habitat of the effigy builders. .VIII., 299
- The animal effigies of Wisconsin and the totem system. .XVI., 320
Pengra, Charlotte E. On the conformal representation of plane
curves, particularly for the cases p — 4, 5, and 6 - XIV., 655
Pierson, Merle. The relation of the Corpus Christi procession to the
Corpus Christi play in England . XVIII., 110
Pinetenpol, W. D. Selective absorption in the visible spectrum of Wis¬
consin Lake waters. (With thirteen Figures) . XIX., 562
Plumb, Ralph G. Early harbor history of Wisconsin . XVII., 187
Puls, Arthur J. The need of a medical faculty in connection with
the State University . XI., 236
Reed, George M. Infection experiments with Erysiphe graminis,
D. C . XV., 135
- Infection experiments with the mildew on cucurbits. Erysiphe
cichoracearum D. C . XV., 527
Regan, Katherine P. Economic and social development of La Fayette
county . XIII., 582
Rickett, !H. W. A quantitative study of the larger aquatic plants of
Lake Mendota. (With Plate XXXVIII) . jXX., 501
Roedder, Edwin C. Richard Wagner’s “Die Meistersinger von
iNurnberg” and its literary precursors . XX., 83
Safford, Truman H. On the present state of our knowledge of Stellar
motion . VI., 145
- On the employment of the method of least squares in the reduc¬
tion of transit observations . .VII., 193
- Combinations of Pythagorean triangles as giving exercises in
computation . ...XII., 505
Salisbury, R. D. Notes on the dispersion of drift copper . VI., 42
Sanborn, John Bell. Railroad land grants . XII., 306
Sands, M C. Nuclear structure and sport formation in Microsphaera
alni . XV., 733
iSawyer, W. C. Letters an embarrassment to literature . IV., 50
- The philosophy of F. H. Jacobi . V., 146
Schlundt, Herman. The radioactivity of some spring waters at
Madison, Wisconsin . XVI. 1245
Schorger, A. W. Contribution to the chemistry of American conifers.
(With one Table) . XIX., 728
Schuette, H. A. A biochemical study of the plankton of Lake Men¬
dota . XIX., 594
Schuette, ,H. A., and Hoffman, Alice E. Notes on the chemical com¬
position of some of the larger aquatic plants of Lake Men¬
dota. I. Cladophora and Myriophyllum . . . XX., 529
General Index of Authors of Papers.
Scott, Jonathan F. An investigation in regard to the condition of
labor and manufactures in Massachusetts. 1860-
1870 . XVII.,
Secrist, Horace. The anti-auction movement and the New York
workingmen’s party of 1829 . XVII.,
Severin, Henry, and Severin, Harry. Anatomical and histological
studies of the digestive canal of Cimbex americana
Leach . XVI.,
- Habits of the American saw-fly, Cimbe americana Leach, with
observations on its egg-parasite, Trichogramma pretiosa
Riley . XVI.,
Severin, Henry. See also Marshall, W. S.
Sharp, Frank C. The personal equation in ethics . X.,
Shrosbree, George. The scientific development of taxidermy and its
effect upon musums . XVI.,
Skinner, Ernest Brown. The determination of the value of the right
of way of Wisconsin railroads as made in the appraisal
of 1903 . XV.,
Sherman, W. H. The production of sulphide of mercury by a new
process, and its use in photography . I.,
Simmons, H. M. Mr. Spencer’s social anatomy . IV.,
Simons, Algie M. Railroad pools . XI.,
Slichter, Charles iS. Harmonic curves of three frequencies. .... .XI.,
- Recent Criticism of American scholarship . XIV.,
Smith, Allard J. See Libby, O. G.
Smith, Erastus G. On the determination of chlorine in natural
waters, its accuracy and significance . XIII.,
Smith, Gilbert Morgan. A Monograph of the algal genus Scenedes -
mus, based upon pure culture studies. (With Plates
XXV-XXXIII) . XVIII.,
- A preliminary list of algae found in Wisconsin lakes. .XVIII.,
- - A second list of algae found in Wisconsin lakes. (With Plates
X-XV) . XIX.,
- The phytoplankton of the Muskoka Region, Ontario, Canada.
(With Plates VIII-XII, and one Figure) . . XX.,
Smith, John Y. Abstract of a paper on the laws which govern the
configuration of comets . Bulletin IV.,
- Effect of the duty on imports on the value of gold . II.,
Smith, Leonard S. An experimental study of field methods which
will insure to stadia measurements greatly increased ac¬
curacy . X.,
Smith, O. R. On the duty of the state to its idiotic children. (No
Abstract.) . Bulletin IV.,
Squire, George Hull. Peculiar local deposits on bluffs adjacent to
the Mississippi . XVI.,
773
167
149
38
61
299
343
794
165
56
66
449
1
359
422
531
614
323
64
77
539
67
258
774 Wisconsin Academy of Sciences , Arts, and Letters .
Stanton, F. Belle. See Libby, O. G.
Steele, Dr. G. M. Population and sustenance . II., 59
Stewart, Alban. Some observations concerning the botanical condi¬
tions on the Galapagos Islands . . . XVIII., 272
Stewart, K. Bernice, and Watt, Homer A. Legends of Paul Bunyan,
Lumberjack . XVIII., 639
Stout, Arlow B. A biographical and statistical analysis of the vege¬
tation of a typical hay meadow . XVII., 405
Strasser, William. iSee Mavor, J. W.
Strong, Edgar F. The legal status of trusts . .^1., 127
Stuart, J. It. The harmonic method in Greek art . IV., 44
Sweet, E. T. Notes on the geology of northern Wisconsin . III., 40
- On kerosene oil . III., 77
Sweezy, G. D. On some points in the geology of the region about
Beloit . V., 194
Teller, Edgar E. An operculated gastropod from the Niagara forma¬
tion of Wisconsin . XVI., 1285
Timberlake, Hamilton G. Development and structure of the swarm-
spores of Hydrodictyon . XIII., 486
Todd, James E. A description of Some fossil tracks from the
Potsdam sandstone . V., 276
Tolman, A. H. English surnames . X., 1
Trelease, William. Preliminary list of Wisconsin parasitic fungi.. VI., 106
- The morels and puff-balls of Madison district . VII., 105
- The working of the Madison Lakes . VII., 121
- Remarks at Fifteenth Anniversary Banquet . XX., 708
Tolman, H. C. The cuneiform inscriptions on the monuments of the
Achaemenides . VIII., 241
Thompson, George. The Gothenburg method of regulating the liquor
traffic, 1892-8 . XIII., 387
True, R. ,H. See Cheney, L. iS.
Urdahl, Thomas K. The fee system in the United States . XII., 49
Van Cleff, F. L. The pseudo-Gregorian Drama in its relation to the
text of Euripides . VIII., 363
Van Hise, C. R. Origin of the iron ores of the Lake Superior re¬
gion . VIII., 219
- The origin of the dells of the Wisconsin . X., 556
- Earth movements . XI., 465
- Discussion of ice ramparts . XIII., 158
Vickery, R. A. See Wilson, H. F.
Vorhies, Chas. T. Habits and anatomy of the larvae of the caddis fly,
Platyphysax designatus (Walker.) . XV., 108
- Studies on the trichoptera of Wisconsin . XVI., 647
Voss, Ernst. Jacob Wympfflinger’s “Tutschland.” . XV., 823
- An ordinance of the city of Nuremberg, adopted in the year
1562 . XV., 874
General Index of Authors of Papers. ' 775
Voss, Ernst. The regulations of the University of Wittenberg,
1540 . . . . . XVII., 397
- - - A true bit of instruction showing why we are under obliga¬
tions to pay taxes and tithes for the preservation of Chris¬
tian peace and the avoidance of trouble . XVIII., 683
Wadmond, Samuel C. Flora of Racine and Kenosha counties, Wis¬
consin: A List of fern and seed plants growing without
cultivation . XVI., 798
Wagner, George. Notes on the fish fauna of Lake Pepin . XVI., 23
- See also Juday, Clancey.
Wakeman, Nellie A. Pigments of flowering plants . XIX., 767
Walker, Milo iS. Some uses of the potential alternating current in
a chemical laboratory . XI., 114
Wann, Louis. The Influence of the French farce on the Towneley
Cycle of Mystery Plays . XIX., 356
Ward, Henry L. A study in the variations of proportions in bats,
with brief notes on some of the species mentioned. .XIV., 630
- Modern exhibitional tendencies of museums of natural history
and ethnography designed for public use . XVI., 325
Watt, Homer A. See Stewart, K. Bernice.
Wheeler, W. H. On the appendages of the first abdominal segment
of smbryo insects . VIII., 87
- See also Peckham, E. G. and W. H.
Willard, S. W. Migration and distribution of North American birds
in Brown and Outagamie counties . VI., 177
Williams, F. E. The passing of a historic highway . (XX., 131
Wilson, H. F., and R. A. Vickery. A species list of the Aphididae
of the world and their recorded food plants . XIX., 22
Wingate, U O. B. The scientific importance of more complete vital
statistics of the state of Wisconsin . XI., 102
Wolff, Henry C. The unsteady motion of viscous liquids in capillary
tubes . XII., 550
Woodman, E. E. The pipestone of Devil’s Lake . V., 251
Wright, Albert C. Abstract of a paper on the classification of the
sciences . Bulletin II., 27
- The metamorphic rocks in the town of Portland, Dodge Co.
(Abstract.) . Bulletin III., 38
- The Metamorphic rocks at Devil’s Lake. (Abstract.) .
. Bulletin III., 39
- The mineral well at Waterloo, Wisconsin. (Abstract) .
. Bulletin III., 46
- On the mineral well at Waterloo, Wis . I., 154
- The Philosophy of history . V., 12
- Distribution of profits, a new arrangement of that subject. .V., 38
- The increase of insanity . VI., 20
- The defective classes . VIII., 176
0
776 Wisconsin Academy of Sciences , Arts, and Letters.
Young, Karl. The Harrowing of Hell in liturgical drama . XVI., 889
- Officiurn Pastorium: A study of the dramatic development
within the liturgy of Christmas . XVII., 299
- William Gager’s defence of the academic stage . XVIII., 593
- Ordo Prophetarum . . . XX., 1
Zimmerman, Oliver ;B. A treatment of instant angular and linear
velocities in complex mechanism . XIV., 513
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