Myelopteris topekensis, n. sp. [microform]

mm

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QKll

McGILL UNIVERSITY

PAPERS FROM THE DEPARTMENT
or

Botany.

iDon

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l.'^'nr.ff.-

No. 5. — Myelopteris Topekensis, n.sp.

>- > BY
D.Pr^'PENHALLOVV.

With Plates II. and III.

[Reprinted from the Botanical Gazette, January, 1897, pp. 15-31]

Montreal, 1897.

MVKI.OITKKIS TOI'KKKNSIS, N. SI'.

A M'W CAKIIOSII'ICKOUS I'r.ANT.

I>. I'. I'K N II A I.I.OW,

(wrni I'l.ATKS II AM) III)

I)uKiN<i the past sixty years a luinibcr of plants, variously
described uiuler the names of Mediillosa' ( 1.S32) . I'aliiiacitcs'
(1845), Myeloxyloii^ ( iS4(j) , Sten/.tliaM 1S64 ),aiul Myelopteris'
(1874), have been obtained from the Carboniferous of France,
Germany and Great Britain, but, so far as I am .uvare, 110
representative of this group has been obtaineil heretofore from
any locality in America.

Recently Professor C. S. I'rosser has sent to me three small
specimens of flattened stems from the upper Carboniferous of
Topeka, Kansas. These fragments are about 6"" long and lie
in a matrix of calcite.^ One specimen represents the full
width of the original structure and is 33""" broad. A second
has the edges broken off, but a natural extension of the curva-
tures of the sides shows the probable breailth to have been about
6'^'". Both of these specimens have been compressed into a
flattened mass having a lenticular transverse section with a
maximum thickness of 5""" and 8'""' respectively. A third
specimen, flattened to an irregularly lenticular mass, represents
thin layers of plant residue adherent to the sides of the matrix,
and obviously but a small part of the original structure. The
dimensions of breadth here given represent very nearly the

'Cotto : Die Deiulmlitlienin liivit'luiiig auf ilircii inneren Uaii. Dresden, 1832.

'Corda : Heitr. zur Flora der \'or\velt. 1 845.

'Hrongniart :'rab. des >,'en. de vdg. foss. Diet. Univ. d' Hist. nal.

* Goeppert : Die foss. Fil. der perm. Form.

5 Renault : Ktude du gen. de^.Myelopteris. Acad, dc Paris aa : — 1875. | no. 10. |

* 1 am much indel)ted to Dr. H. J. Harrington for detenninations of tlie mineral
constituents of these fossils.

1897] IS

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:CTg?iS-S3%i't'-^<'''^»HMtWJiswtfiW«i^,.u'HW!',''

If

Hot.lMi.U u.i/l III.

I I^MAKV

<n.im«tfr tif Itu- Hiriii tiirr iir in original fnrni. Thr ^n MHr.il . ..|..i
!•> iImI <>| Iikiwii I u.iI. Tlir siirl.M r sliows ik ( iisiuii.il iir< ,i«i u|
lliiii ( n,il\' ni;i»l«r iniK li hrnkiii u|» iiiti» Miiall angular Iragiiitiils,
Itiit It IS iliiillv « li.iiailfri/»'il by » HnnH-whal fiiuly sliialti!
a|i|ifaraiUf iliu' Ik ri'imtval hI iIk ...iliral lavi-r, with n»iis(t|ii( nt
ixpnsiiif III iln- iiiKlfilyiiij^ >liafi'l-i u| M'lfrt'iichvriia.

I'lu' tLiiisvi-rsi- scttinii III thi' iiiiir«' |»rrfcrllv |irfs(rv»il
H|H'cmKii sliii\v> an iiiilir /niic i ,5""" tliii k. uliii li is (iiminiiniis
(111 ,ill sidfs. Ci'nlial tn this ami thus Imiiiiii^' tin- axis of tin-
original siriutiiii'. is .1 ili^tiii« tly ilaiktr ami smmwli.it im,ir
l>c>ii»iis mass, r«»iilaiiiiii^. luii- aiiii tlinc. small irr(rj,Milarlv
roiimlfd massis nl |iyrilf. l)'|i<iii Mil)Mi|iiirit mi(ros«ii|ii( al
c'xamiiiatidii, llicsi.' zonal a|i|i('aiaii<t> wnr luiiml tn lir din' tu
wi'll lirfiiH'tl (lirffri'iicrs ol striu tiiri-.

I 111' mil riisn)|iiial lUtails pifsctit iii.iiu li .ituics of iiiti rest
ami, allliiuij;li the m-iifral i-ffi-cts of die av ami ( oinprissioii liavc
iiiiii to complctfly (U-slioy tlu'^tiirral nlalioiis of |iiiits, ami in
many I'ascs, also, to (U-stroy stnutiiral ditails, tlusr last imvc hccn
preserved, in some instaiui's, in a n-markahly ptrfii t manm-r.

Tin; ii;NrKAi axis. 1 In- intiif ci-iitral portion of the stiiii
presents a romplete absence of strmtiiial ditail. 'llie whole
central area is occupied i)y a mass of dark colored male-ial so
disposeil as to indicate its prol)al)le derivation from thin walled
tissue, but much altered by tiecay and tlii' sul)se(|uent effects
of extreme compression. Here and there, dark colored masses
appear, possibly the residue of the mucilajfe ori^Miiallv present.
Thiouirhout this rej,non lari^^e rounded o|)eninjfs a|)pear. and
while some of these undoubtedly represent the displacement of
pyrite, many, and probably all, represent the former locations of
vascular bundles, in the dark color and structural character of
this area, we find amjile reason for its evident separation from
the cortical /.one, as ascertained upon microsco])ical examination.
Outwardly, this area is limited by a somewhat well defined but
narrow and irregular darker line, which is obviously composed
of much compressed thin walled cells, but which, nevertheless,
seems to sug)j[est a somewhat definite boundarv line between a

(lAM'MiV

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.Ul/./iU-// fi/.\ /(>/•/ A/. X,\/S

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tii'ral tutor
il an, IS of
Irii^'itit'iils,
ly sliialiil

I MIlM l|lll lit

|iri'sfrv<'i|

( lilltilllMMIS

ixin (»f the
Mliat iimrr
irrt;>j[iilarly
"roscopical

he due t(i

of iiiti rest
Nsion have
rts, ami in
liavi' Iji'i'ii
ii;iiiiii-r.

I till' Slllll

riif wliolf
iiatr "ial so
hill walli'il
•nt rlfccts
t(I masses
y idcsciit.
pear, ami
iccmeiU of
icitioiis of
laracttr of
ition from
;uiiinatioti.

ll'fllK'tl l)llt

composed
I'crthclfss,
between a

central tiicilnlla in wliiih thin vvalli-il (iitiilatiKtilal tis<tiir |>rt'>
<lomiii,it('s, ,tnil .1 soini'what ri^icl, or at least firiiur, out) r /om-.

I 111. M»KirA, No proper tortical struiiiir«' is rrprrsnitt d
II! ihiHf spt'titmiis I he outer liiuitH of the ttiatioitn ar«; fhTiiK il
\>\ more or Ichh liroki ii iIovvii slramlH of HcU'rciuliyina lelK, with
surroiiiiilin)^' parcm liynia tissue, making; it i tear that a certain
,iiiioiiiit of struiture has tieeii ri iiio^ed ; ami thisairords witti
wImI has .ilreadv lieiii noted in spii itiieiis of Myelopteris, that
"tin- tissue layers outside the sclcreni hyma strands are very
rait l\ prcservftl."' In tins ease the thin surface layers of coal
.ilitMfiv ilest rilu'il are in .dl proliahilits to lie re^'anled as
rt presenliiiL,' theitirtiial struiture, wlinh must li.ive lit in < hit fly
or vvholK parent In iii.'itous in eharat-ter, ami of small radi.d
vtilmne.

I'm si 1. I o|< I |( \| I \\ I K. The outer, lontllMloUs /one. i.n"""

thick as alreailv iltsi riheil, li.is iis maertist uph ilitlen nti,ition
from the meilull.i explaim «l l»y the Iar|,(e ;ini<iUiil <if lihioiiH
elements whitli it < ont,iins nuiuLf to tlu- pre-i n« t of these
t lenients, ami tht pet uliar way in uhiih thi:y are tlistrihiitetl,
thev have ser\ed not onlv to prolei:t one .iiiotlu r, Imt they have
,ilso ser\eil to prevent the tl'fti t ol i ttmprission trom f,illiiij,'
with full force upon the interveiiinu fuinlanientai strut lure whit h
in conse(|ue!ice, h,is often retained its strut tur;il ft-aiures in an
exceptitin.illv perfect m.inner ( /ii^'.s. / .iml -»i.

1'ak1'".N( iiVM A. — The i^rouml tissue, tor the irreattr ; .irt, is
much .ilteretl hv tiecav ami ctunpression, so that all structural
features, especi.itlv in the central area, have been pretty com-
pletely elimin.itetl. ( )ccasioiially. however, when prtitected hy
earlier inl"iltralit)n and petrif.ictioii, tir by the resistiiiLT char.icter of
the acct.!n|)anyin,i( ;^tramls of hard sclerenchym.i, this part of the
structure has been preserved in a very be.iutiful manner iji\^s.
I and .?). I''roin these areas it is possible to tlelermiiit; the fact
that this tissue consists of very vari.ible. but chiefly liir-^e ami
thin walled elements of such a character as to reminti me very
forciblv of the fundamental structure in many t)f the lar^^cr ferns.

'Si)lm>l.:iul)acli: r(i>>. H'.t. lOJ.

i

li

HOiMXn.il I, </A/7A

hi«M 411*

l'li> iiiU'ri'i'lliil.ii sjMCCH onliiLirity mu*I \«illi in Hiich tiNNtit* tiru
|ircNi'iil, Intt llit-u' It ID) t-viilciiti' III lilt- I'siHti-nt'f of l.u iiii.i-
St i i him iuma. It l);iH Ih-cii hIiomii .ilitMilv that tln' siilf
curtu.tl /t)iu' \% !.«;""" tlurk. Willuu Ihi-^ region tlu-n- arc
iniiiii-roiiHuval or liiii^t'iiti.illy I'loii^atctl iHinilk'M i>f NtU'rciii'liyiiia,
wliii'l) U»\\\ lon^: ttriiiitlH travi*r>tiii^; tlic stem luii^itiiilinallv lor
^'n-at ilKtaiui's <.//c>' /. •/. inul ./). I'Iuhi' straiuK, wlmli \ii\i'
till* |ii-( (iliarly slriati-d a|i|ii-arai)('f to tlu' Mirlaci- ol tin- >t|ic( iiiuii
wlti-rt'vrr cx|to>*rtl, .ir»' alwav* si'|»ar.itccl Iroiii oiU' aiiulhi i hv
Hcvcral lai^f ,iiitl tiiiii \vallc«l |»iirciitliyma «i'lls {fi}^. i) uliirli
iiri' Hccn to In- vory |tfrlicctly pnv^crvwil in ctrtain ari-as. ||ju
Mi'lcrfniliviiiatoiis clciiu'iits arc always vfr\ tliick wallril in those
strands uhiih lii' mxl the mrtix ( //^,^ >' ). t'»it luruini' nimh
tliiiiiit'i w.illi'd toward tlu- (inter ol llu-siini wlure thes ollcn
appear to III- in a forniative eoiidition. The strands an* Hcparatcd
radially by rather wide areas ol (iindaniental tissue (//</'. j), Itul
in conseipienee of the ^^i-neral and ^reat alteration in relative
positions el'fi-eti d liv lompressiun, it is inipossililc to ditertnine
their ori^Miial distrilmtioii. The radial dislnlitition ol these
strands thron^h a rather wide /one woidd seem to indii .ite that
they ntay ha\e luen developed in more or less well deliiietl con-
eentric layers, a relation whieh is certainly implied liy their
distriliiilitin within eertain areas (//(,^.'). HeNdnd a limit of

1.3 Iroiii the siirlace the development ol the strands a|ipears

to lie wholly arrested.

\'\^i riAU inMH IS. riu- vaseular bundles are not Ireipieiitly
represented, siuie in uiust cases they have been removed by
decay, or other causes, and their former positions are then
markeil by the presence of rather broail, irre^nilarly rounded
openinj^s of variable dimensions, which appear throuj^hout the
transverse section ( y/X""-*. /. -'. -md ,,'). and particularly internal to
the sclerenchyma /one. Occasionally the bundles are preserved
in a very perfect manner, and exhibit all their essential structural
features with tfreat clearness {y/^,^ /). The outermost uf the
two bundles seen in //{,'". /, when much enlaru;eil (//i,'-. y). is found
to consist of several broad scalariform ducts enclosed 011 two

I MM tut

llHHtif tire

,1111.1'

till' Hllll-

tlUTf AtK.'

renchyina,

iiially lor
liH li ^ivc
>»|H'i iiiifii
Itillirr l)V
/I uliitli

CJIH, I III

li ill tlioNC
illir lllticll
lu'\ ullcn
Ht'paiati'il

(C- ■'I . 'it'l
II relative
(K-tiTinitu*

m| tlu'Hi'
liratc that
fllU'il ruii-

l>y I heir
I limit III
Is appears

■re(|iicntly
uoveil l)y
are then
V riiundcd
,^hniit the
nternal to
|)reservtHl
stnictiiral
St ul the
), is I'oiiiul

li 1)11 two

• ■•»7 1

.U r A A r»/7/ A7.S / 1 »/'A A A \.S/M

»'»

Hlili'H l>y tttws )ii lliick walleii I'lhruiiH elritHMIts, The phlucrii,
rtither Hiiiall tii vuliinie, in hen.* iiituh hrnki ii ilowii. hut it it
•titiiiiteii riiiiiiilly outwanl, while in the other hiittille { Jijf. th
where it in rather mure perleetly prcHervcil, it •« situated radially
iiuv.iri!. The prntuxvli Ml in luri' sicn as n iiTnu\> of siiialU r
elt iiienis iiiiuli altered l»y n»iiipre""<i«in ( /f),'. /). or i»i otlnr
iiisiiiiices mure perfectly preservi il (///;. /), sometimes un the
uuter fuce uf the veNsels, {irid sumetiiiicH un the inner face, hut
Hlways hetween them and the phloem. While the hundles vary
con><ideral)lv in si/e, tin v all < unlnrm to the en" iteral type and
it IS ol interest to note ili.it in .ill their structur.il features, they
H^:ree very cli»scly with the iKindlcii of n Hpeclen of Myclox\loi)
tltserilied l»y Solnis |,.uil).ich," and xhtt hy Seward.^

l*'rom the present material I have heeii wholly un.ihie to
oliiiiin ^atisf.utorv det.iiN of th«' structure (»l the bundle m
loii^'itiuliii.il section, he;, Olid the latt that the vessels are dis
tiiu tlv scalarifornt, and in this respect they conform to the type
j^eiierally ohservi d in (mis.

The peculiar situation td these bundles is not altoj^ether easy
to .lecount for. Tlu \ icrtainlv appe.ir to lie between, ;ind are
therefore iiiitifrliil Willi, the strands of si K reiichynia, from which
eircumstiinee I was at first led to siippos*- them to he collateral,
as in the case of I'hoeiii\ .iiid titlier p.iliiis, but .t very careful
examination fails to disclose any satisfactory eviilence of such
relationship, while in some cases at least the vascul.ir bundle is
separ.ited from the nearest sclerenchyma stranil by a broad zciie
ni fundamental tissue. Indeed, the evidence, so far as obtain-
able from the present material, seems icj indicite that these
bundles and the sclerenchyma are altot,'ether independent of or\c
another; but in the present unsatisfactory comlition of the
niateri.^I now available, :io fin.il conclusion can be drawn. Kroiii
the eviilence at hand, however, it would seem that the vascular
bundles have their extreme outwanl distribution in the central
portion of the .sclerenchyma zone. I-'mm this position they

•^■o^!^. It'll. ii>i. //f. / / /i,

•.\nn. lint. 7: />.'. / .111.1 //, /ii't. /. V. /y.

/.'(»// \/l.l/ ( :.!//■ Ill:

( I A M \ K V

ilUllMSi- ill IIIIHlhlT illUMld tin- 1 I'lltlT .111(1 1)C( lilllr llll>->t iiuiiu'r-

oiis uilliin tlu' ci'iitiii' rii^JDii.

Si cKi.TiiKv oK.iAN.-.. A ii()t,il)U- fi-at'irr ^li llu" prcsfiil fossil
is tlir orciirronci- of mmuTous largi. iiiiicila^f passa^fs. As a
nik' tln'SL- stniiluifs ait' imuli altiTiM hv ili-iay ami c-nmpiis-
sioii, hilt ill two instances tlu'V wire foiiiul in a vi^y piTfi'Ct
state ol preservation ( //;,'-. J\. So far as it is at pii'scii possible to
deteriniiK', these organs occur tlirounliuut the siih-cortical region
where thev are in nion- or less intimate association with the
sclerencliyni.\ stran('s. i-Jsewhere it is not possible 1o determine
the (listrihiition satisfactorily, l)nt, fiom our kii iwledi^c ol their
occurrence in recent |<laiils, it is a fair inference tliat they must
also be distributeil throui'li Die entire l)ody of the fniidaiiuntal
structure.

Measurements of such of these passages as were in a
sufficient state of prcservati.m for such a pu'pose showed them
to ha\e the lollowinn' dimensions: i.;;xi00jw,; JC);xi3;/x;
2 1 1; ■ 143^. From these results it is possible to deduce an
averaj^e dimension of 127 I(j2/x. I-'-om this ayain it appears
that these passa<.jes mavbe described as of elliptical form, in which
the minor and major axes have a ratio ol i : 1.5. The very
great size of these structures, unusual except in a few irroui)s of
plants, seems to su'^^'nest a comparison with both Lyciclaceie and
Marattiace;e. In structure they are siniijle. LoiiL,n'tudinally
thev form liMig tubular passaf^es which traverse the stem for
great distances. l:i transverse section they consist of large ellip-
tical openings bounded by a very regular w.iU composed of par-
en:hyma cells often iliffering but little from those of the sur-
rounding tissue. Thev are mc>re commonly somewhat elongated
tangential Iv to the central canal, and by analogy with similar
structures in recent plar.ts we may infer that they contained
active proto])lasm. Thev thus forn: the secretory cells, or an
epithelium which is not .-.pecially differentiated ^fig. _^). A
comparison of the two canals {fig. 2) will serve to show, how-
ever, that the secretory cells often show little or ikj dexiation
from llie general character of tlie fundamental structure.

( I \ M \ H V
st IIUIIUT-

HMit fossil

t"-. As ;i

cumprcs-

\- [HTli'Ct

iussil)|i' to
cal rr'^ioii
I witli tlu'
(Iftfiiniru'
(• ol tlicir
tlicv must
idaiiu'iilal

icrc in a
wed tlitin
D5X i3;/x;
IcdiKc an
it a|)|)(.-ais
1, in whicli

The \rrv
groups ol
l.iCLM' and
;itudinallv

stem for
art^c el lip •
ed of par-
)f the sur-
eloiigatcd
itli similar
contain. -d
A\s, or an

:^- /)■ A

how, how-
dexiation
structure.

iK>7|

MVi.i.ori hRis hiI'I-.k i:\sis

31

Another important feat'ire of tliese laiials is t«. Ix- found in the
fact that tliev are always devoid of contents. llii-. ap|)ears to
justily tlie view tliat wliatever they may liave contained origi-
nallv was of a soiul)ie nature and thus |)assed out ol the ixuly
of tin- plant during the process of petrilactioii.

In all tlieir principal structural aspects these canals hear a
stronj,' reseml)lance to those of Ani^iof^fcris ivcctii (they are of
the same tyi)e), and it mav al^o l)e |)ointed out that they are
similar to tliose fourd in Riuliiophris / 1 '////</w,sw/// vvhi( h .Seward
has recently separated honi M velowlon," as also to tiiose of
Myeloxylon itself."

'rhroutfhoul the transverse section of the Topeka specinuii
there are numerous resinous or coaly masses of very variable
si/.e, l)Ut cvidentlv originalls contained in special channels or
cells, which have heconu' much disorifani/ed, aiul the details ol
which cannot now he made out. In huinituilinal section these
masses are (;f indefinite lenjfth. but ratiier freiiuently septate.
From these features it is |)ossil>le to refer them to the residue ol
resin masses which the plant oriifinally contained, and they are,
therefr)re, directlv c(jmparaljle with the similar resin bodiis
fouiul in recent plants, particularly those of Ani^iopkns tVCiUi.

It is tiui^- fairlv certain tiiat in the Topeka plant there were
orii(inally at least two. and possibly niore, kinds of secretory
origans, tiie one holding mucilage, the other resinous matter,
and in these resjiects our plant is once more comparable with
certain recent forms.

The general view of tiie internal structure thus obtained
permits us to gain some conception of the real composition of
this stem or stipe, from which we may infer that a restoration
would show a cortical layer of parenchyma at least several layers
of cells thick, containing numerous mucilage passages. P'ollow-
ing this is a sub-cortical zone distinguished by the presence of
nuiiicrous rather widclv separatetl strands of sclcrenchyma, the
elements of which are very thick walled toward the cortex, but

■'.\nn. Hot. 8: />/. .V///./(V.f. 'i' // t.'"-
" .\nn. Uui. 7; //. I, figs. 1,1).

-r*jSJ^^^^|J(^||flffff^^BnW^?J^''"'

HO/AXJCl/. UA/./.J //:

I.IANIARY

Cdiuimially tliiiitu"' walled toward the mciliilla. Tlicsc strands
iisuall)' liavf an accompunvin^ iiuicilajfc passaj^c on the oiiter
lace, inul are inconstant {':) process of formation tow.ird tlie
center. Within this zone, vascular bundles, distinf^uislied by
tiieir broad scalariform vessels, appear, anil increase in number
toward the central re^non. The central axis consists of a rather
broad tract of parenchyma tissue, throuj^h which tlie vascular
bu.ndles are distributed in larj, numbers.

l*rom this point of view, and with ilue allowance for the
effects of compression, it is possible to trace a strikinj^ similarity
in several respects to a species of Myeloxvlon describeil by
Solms-Laubach," and i lore |)articularly in certain respects to
specimens of Myelopte is described bv Williamson.'^ The evi-
dence is both clear and direct that this plant must be considered
as belontrinn' to that peculiar trroup for which the name
Myelo|)teris, pro])osed by Renault, has been most jrcnerally
emploved.

In 1S32, Cotta ilescribed certain fossils from the Carbonifer-
ous of Kuro|)e under the name of Medullosa, which has more
recently become mer^red in that of Myelopteris. Williamson,
however, informs us that Cotta's figures of M. clegans are wholly
misleading, the structure being represented in a much exagger-
ated form, while his two .>^pecies, J/, stiilata and .]/. porosa,
remain too obscure to be depended upon without further evi-
dence than has come down to us.'^ The genus Medullosa,
nevertheless, constitutes the basis of that group of i)lants
which, ])assing under several names, ha;; finallv come to be known
under that of Mvelopteris.

In 1845, Corda assigned to his genus Falmacites two plants
from the Coal Measures of Bohemia, under the names of P.
ntrboiiigciiiis ami P. Icptoxylon.^^ An examination of Corda's
figures shows that there is no very great resemblance, although

"Fos.s. Hdt. l6i, fiji. 14A.

■'Fossi, plants of tlic Coal Measures. I'hil. Trans. 166: fin!. /,.?, 4.
'■•Koss. plants of the Coal Measures. I'hil. Trans. 166'.
'SFIora tier Vcjrweit 4c, 41. //. /9, .?(). 1845,

I.IAMARY

c Strands
the outer
•ward the
lislicd bv
II iuinil)cr
if a rather

■ vascular

L' for the
simihirity
:ribecl l)y
rspects to
The evi-
onsitlcred
he name
j^cne rally

irbonifer-
lias more
illiamson,
re wholly
exagger-
'/. poros(7,
rther cvi-
IcduUosa,
)f plants
be known

\\o ])lants
lies of P.

■ Corda's
although

lSi>7

M) / lor/ / h/S /<)/7A/:XS/S

M

there is a suggestion of similarity to our plant in the general
eiiaracter of the fundamental tissue, and tiie presence of numer-
ous mucilage passages. These latter, however, are small and
a|)|iarentlv altogether separated from the vascular bundle.s.

Subsecpient observers have not been unmimlful of certain
structural aspects in these plants, which ha\e seemed to suggest
their possii)le relationship to the palms, and more particularly to
that tvpe of structure represented in the genus IJrac;ena, but much
doubt has always been entertaineil as to the possibility of mono-
cotyledons occurring so far back as the Carboniferous. These
doubts w-ere first ijromiuently expressed by lirongniart as the
result of comparing with the plants figuied by Cotta and Corda,
new material obtained from Autun, France."' He says "il y ait
des differences fort essentielles et (|ue rendent tres difficile
d'etablir des rapports entre ces fossiles et les vegetaux vivants."
He therefore preferred to regard Cotta's Midullo.ui i/t\!^'-a//s as
the representative of a new genus, for which he proposed the
name Mveloxylon. which thus seemed to indicate the leading
structural features indicated by the former name, the signifi-
cance of which was thereby jjcrpetuated.

Fifteen years later, Goeppert, in reviewing Cotta's species,
regartled Midiil/osa cligtiiis as possessing characters which were
variously represented in the gymnosperms, in i)alms, and in the
ferns. As a generalized type, he applied to it the name of
.Stenzelia.

In 1873, Williamson first drew attention to the belief that
the relations of these fossils had not been correctly interjireted,
and expressed the view that tiiey were really ferns allied to the
Marattiace;e.''

In 1874. Renault reviewed the fossils obtained from the
Carboniferous beds at Autun, as a result of which he supports
the conclusions reached by Williamson, and while he regards the
name proposed by Corda to be wholly untenable, and those of
Cotta and Goeppert to be insufificiently indicative, he views that

"'Tab. des gen. ile Veg. l-'oss. Oo. 1849. (Quoted from Williamxm.)
"Hril. A.-is. .\<X\. Sci. 1873.

I

34

/U)/.l\/l'.t/. (,.l/./:/r/:

I ia.\i;akv

of Hroii^Mjiart \vit!i lavor, l»iil ictfards a tlinVrcnt form as more-
expressive of tlie rclati()nslii|) wliicli lie (litirniliud.'** He
tluit lore says: " I'oiir coiiserver le iioiii, |ireiiiier eii tiale, doiiiie
|iar M. Hron^niiart a ces [jortioiis de plaiUes, et eii iiieine temps
|toiir rappeler leiir nature, je les desi^iierai sous !e iiom tie
M yeloptcris."

riie yet more recent studies of tliese plants h\ Williamson
led him to admit the force of the ari^uments employed Uy
Renault and the appropriateness of his nanie.'^ Referenct- to
Willianisou's figures discloses several points of resemblance
between his specimens and my own. This is to be noted (irsl
in a i,n-eat similarity with res|)ect to the Lfeneral distribution of
tissues, particuhiily as exhibited in his Jii^s. j and ./, as likewise
in the very general removal of the vascular inmdles. The vas-
cular bundle i,nven by him ( Williamson, y/^'-. 7*) is closely similar
to that derived from the Topeka s|)ecinien [ Jii^. 7), but differs
materially from his other representation (Williamson, y/i,''- 7)
taki'U from the uj)|)er end of a rachis, which is closelv simil.ir to
bundles observed by me in Dioon cdulc, whereby it offers some
basis of C(jmparison with the Cvcadacea;.

In longitudinal section the resemblance is rather close, but
in this aspect the To|)eka specimen offers little evidence of a
satisfactory nature beyond the general relations of jiarts, anil
the structural markings of the vessels winch are seen to be
scalariform, as in the ferns.

Finally, the relation of the mucilage passages to the vas-
cular bundles (Williamson, y?^. 14) and of the very large, elli])-
tical mucilage passages to the sclerenchvma strands (William-
son, Aa''- A))> ^^ '^'^" i''*-' ^■'-"'■y thin walled elements of the funda-
mental structure, all |)resent features almost identical with those
observed in the Topeka specimens {Jigs. /, ^, .?).

Williamson's specimens appear to differ from mv own chiefly
with respect to the particular distribution of the sclerenchvma
strands in the cortical region, a difference which, however, is

■' Reclierclies .sur les veKctaiix silicitics d'.Viitun. ( l-'rom Willjumxtn.)
" Fos.sil plants of the Coal .Measures. I'liil. Trans. i66'.

I i\\i;akv

II as more
ncl/" ifc
late, (Idiiiu'"
einc temps
L' iKim <lc

A'illiamsnn
ployed by
■tereiice to
;scnil)Iaiice
noted first
iil)iiti(iii of
as likewise
The vas-
^ely similar
but dil'lers

t^'i.A'-- 7)
• similar to

Ifers some

■ close, but

deuce of a

parts, and

ieeti to be

o the vas-
arife, ellip-
(Williani-
the funda-
with those

)vvn chiefly
ereiichvma
lowever, is

n.)

IM.,7

.M\ I i.oi'i r.ias i{<i'i Kissis

35

more speeilk: than y;eneric, l)iit my material has been so altered
bv compression tliat I slimdd liesitate t<t place much reliance
upon tliese aspects oi >triuture, preferring- ratiur to establish
the aflinity i)y nicans of the more perfectly prestrved slriutmal
elements.

The distribution of the vascular l)imdles in concentric zones,
as described by Williamson, may also be a feature of the Topeka
specimen, but for reasons alreaily stated this cannot lie asserted
with anv def^ree of confidence.

More recently Solms-I.aubach* has reviewed the entire
relations of this tfrouj) of i)lants, and while lie rejects Renault's
name because he rei^ards the evidence as not altogether satis-
factory, he prefers to retain HronLjni.irt's name (jf Myeloxylon
••rather than .Sten/elia, because it is better known." He \(\\c^
two il^nires, one of a i,a-neral transverse section, the other of a
separate vascular bundle, and it is r)f considerable interest to
note that this latter is almost the e.\act counterpart of a vascular
bundle obtained from the Topeka fossil (y/!,^ /i. His ^a-neral
vii-w of tlie structure is not so satisfactory, but it nevertheless
exhibits a close similarity to my own material in all its principal
features.

Solnis-Laul)ach dissents from the conclusions of both Renault
and Williamson, holdini( that there are stront^ reasons, on ana-
tomical trrounds. for considerin^r the alliance to be with the
CN-cadace;e, and cites Mcdnllosa Lciulun-ti as pnjbably affording
important evidence in support of this view.

The most recent contribution to our knowledL^e of these
|)lants is that offered i)y Mr. A. C. Seward, who has not only
reviewed tlie material originally described by Williamson, but
has made a detaileil study of specimens contained in the Hinncy
collection of the Woodwardian Museum. Cambrid;^^'. as well as
of new naterial derived from the Millstone grit of Lancaster."
The o.'r, ^.')ses show that his material is generically the same as
tiiat represented by the Topeka specimens as described. In a

^'Koss. lii)t. 161. I^'OI.
=" Ann. liot. 7:1.

It

a6

MO TAX/C.t I. GA/.E TTE

[jANi-A«V

scCDiid |);i|)cr, tlic same authority tuakcs a stiulv of certain spec-
imens containetl in the Williamson collection and originally
incliulcd by Williamson in the ^rcniis Myelopteris, hut which he
fiiuls to be in reality ([uite distinct. lie therefore He|)arateH
them under the name of Riuhhptcris WiUinnisoni." This species
is quite tlistinct from our Topeka specimen with res|)ect to the
character of '.he vascular bundles, which are concentric, and thus
show a tlistinct approach to the type represented iit Aui:;iof>tins
evccta. On the other hand, the mucilage passages, which are
also of the type found in Angiopteris, are essentially the same
as those of the Topeka specimen, differing only in distri-
bution.'3

l'>om the review thus presented, it is tpiitc clear that our
specimen must be regarded as a s|)ecies of myelopteris, accord-
ing to the name adopted by Renault anil Williamson, and
retained by me as expressing its probable relations, but that it
differs s|)ecifically from any of the specimens heretofore
described. It may be concluded further that the present mate-
rial represents the stipe of a frond, rather than the stem
proper.

Heretofore the representatives of this genus have been
derived wholly from the Carboniferous of Kuropc. The material
now at hand from the Upper Carboniferous of Kansas thus
afforils important evidence as to the wider geographical range
of these plants, while the well ])reserved condition of portions
t)f its structure permits a further discussion of its possible affin-
ities. I have, therefore, carefully passed in review such species
of living plants as are available in the botanic (iardens of
McGill University, as affording a possible solution of this ques-
tion. In jjrosecuting these studie;N I have had in view the sug-
gestions of earlier investigators, as well as those which natu-
rally arose in my own mind upon making a preliminary
examination of these fossils. I have, therefore, carefully exam-
ined Cordylinc tcrminalis. Phoenix dactylifcra , Kcntin Fosteruvia,

=-■ Anil. Hot. 8 : 207.
-■'.■\nn. Bot. 8://. /.^A^ *.

f lAMAKV

•rtain spoc-

orijfinally

t which he

; SL-paratcs

his spt'cit's

)ect to the

c.and thus

/hii^io/ifin's

wliiclj are

the same

in tlistri-

r that our
is, accord-
uson, and
)iit tliat it
heretofore
scnt niate-
tlie stem

have been
ic material
msas thus
ical ranjj^e
if portions
sible aflfin-
ch species
ardens of
this ques-
w the sug-
hich natu-
reliminary
Lilly exani-
Fosteriano,

I8.J7I

MM. I A tpTKKts ror/:Ki:\s/s

>1

LiitiNiiii borbonha, Cyciis irroliitii, IHoon cdulc, /.lonio intn^ri/oliti,
Cibotiiiin tr^iih, and .■hij;^io/'/iii.\ ivcitii.

A close comparison of the Uracicna type, as represented by
Cordvline, shows that any suj,fgeslion of resemblance which
might at first appear, has no real basis in structural characters,
while in manv essential respects there is a \ery wide difference.
Noteworthy points of reseniblance being absent, it is wholly
uiujecessarv to enter into a more iletailed consideration of the
structural aspects of this type. \'ery nearly the same observa-
tions arc applicable to the palms. In this grou|> of |)lants, how-
ever, there is a somewhat closer point of contact to be found in
the mucilage passages. Here these structures appear as tubular
channels of great length, and in this respect, as well as in their
distribution and great number, there is a general resemblance to
the Topeka fossil. Their detailed structure is, on the other hand,
(|uite different, anil it jxiints to a want of aflinity which is sup-
ported bv the structure and distribution of the vascular bundles,
as also the character of the funilamental structure, and no very
searching comparison is required to establish the fact that the
affinities of our fossil must be sought elsewhere.

Hv several authorities the Cycadace;e ha\e been suggested
as affording a satisfactorv basis of comparison, a view which, in
more recent times, appears to have been particularly urged by
Solms-Laubach,^* although he elsewhere agrees with other
observers that certain exceptions which have been taken to the
cycadaceous character of the Medullos;e are well fouiuled."

Mr. Seward, vet more recentlv. has given expression to the
same view, basing his opinion upon a very critical examination
of a large amount of material.-''^ While admitting the many
points of resemblance to ferns, he holds that in the position of
the protoxvlem and in the structure of the fnucilage passages,
as also in the distribution of the sub-cortical sclerenchyma,
there arc strong reasons for considering the affinity to be with

^*l-'<)ss. Hot. Ibl.
'ilMii. 100.
"'^Aiin. Hot. 7: iS.

!

«l

*.

fi

in

I

i

.1

I!

.'S

/mr.iMc.if. i;.\/.i- 1 II:

(lAMAHV

tin- cyi.uls ratlur than with thi- firns. Without liopiii^- to
settle this i|iicsti<tn at ihf prcsiiit tiim-, it may l)r prufitalilr to
coiisiilcr .some of the arguments ailvancctl by Mr. Seward in
the light of tvideiice obtained froii) an examination of material
ilerivcd from existing species, as also from the Topeka specimen
itself.

Vasili.ak iiiM)i.i;s. — An examination of both tyeadaceoiis
jilants and ////i,w/'/<m cvccto afforiis but littU- evidence contrary
I') the view urgeil bv Mr. .Seward. The evidence obtained shows,
as he contemls, that the position of the |)rotoxylem in these
plants is certainlv an argimieiit in favor of the cycadaceous
character of .Mveloxylon. On the other hand, the collateral
character of the vascular bundles in the latter cannot be taken
as final evidence of afViiiity either with the ferns or with the
cvcads, as Mr. Seward himself points out. Althougli the longi-
tudinal sections of the Topeka specimens havi' given far from
satisfactorv results, the evidence to be derived from them indi-
cates a much closer resemblance to Angiopteris than to any t)f
the cvcads 1 li.ive been able to study.

.Si;( KKiuKV ()i<(;.\Ns. — lu the Cycadacea', ;is represented by
Cyciis nvoliitii, /.oniui iiili'i^rifolid, and Dioon ciiiile, the secretory
organs ap|)ear to be all of j>ne kind as represented by mucilage
canals. These structures are distributicl throughout the funda-
mental tissue and are represented by broad canals v/hich are
chiefly limited by tangentially elongatetl parenchyma cells.
These latter, therefore, differ somewhat conspicuously from the
cells of the surrounding tissue, as already shown by Mr. .Sew-
ard.'' .So far as apj)ears from the species above irulicated, how-
e^'cr, these canals are alway.s lined with a layer of very thin-
walled epithelium cells, which become ru|)tured with age and,
shrinking back upon the main wall of the canal, give it a thick-
ened and very ragged ajjpearance.

In Angiopteris cvecta there are three distinct kinds of secre-
tory organs: {a) tannin sacs, \,b) resin canals, and U) mucilage
canals.

=?Ann. l{<it. 8: 214.

1 i

I I A M \ K V

liopiim to
(ifitalili' III
Seward in
il iD.'itfrinl

I sprciilun

■ciulfiiruiis
;c contrary
ncd allows,

II in these
,'ca(laccoiis

coilatLial
)t be taken
ir \vitl\ the

tliu longi-
•II far from
tliein inili-
II to any of

cscnttil l)v
; secrctorv
y nuicilaife
the funda-

which are
yma cells.
V from tlic
y IMr. Sew-
atcd, how-
very thiii-
1 age and,

it a thick-

s of sccrc-
) mucilage

|H.)7|

.1/ ) A l.Ont-./i/S Ton. A7. .\.s7.s

29

Toniiin stus. Ill U-iiisverM- section the tamiin sarn .ire ultrii
harely ilistiiigiii^hahle frtnii the resin canals, In reason of their
structural similarity. They occur al>un<lantly in the cortex and
^l^roll^dlout the fundamental tissue, and especially in ilose |»r«).\-
imity to or within the limits of the vascular hundles. To me
these appear to he the structures rcfcrieil to by Mr. Seward in
his description of Kmhiopkns W'illiiiinsoni, when he says, "there
are smaller canals in the peripheral part of the phloem of each
bundle."'** In longitudinal section tliesc sacs are seen to be of
about the same diamet<'r as in the transverse section. e.Ncept in
the cortex, where thev assume the loMii of cylindrical cells about
three or four limes longer than broad. The contents are much
lighter colored than those of tlu- resin canals, and often present
a well defined granular appearance. They readily yield the
characteristic reactions for tannin,

Rfsin <v///^//.v. — Throughout the sub-corticd zone, scattered
among the sclereiichyma cells ami also central t«) each of the
isolated strands, are rather broad canals of indefinite length.
Throughout the fundamental tissue, particularly in the neighbor-
hood of the vascular bundles, there are also numerous c ;iii.ils
which differ but slightly in their structural aspects from the
surrounding cells. In all cases, however, they are at once
recognizable by the r.ither ilark reil resinous mass which each
contains. In longitudinal section the canals ;ire of indefinite
length. The contents are often septate. These structures
appear tt) mc to be comparable with the black, resinwus m.isses
of variable size to be met with not only in the Topeka specimen,
but in most of the Kuro|)ean specimens of Myeloxylon.

Miiciltigv Kimtls. — Throughout the ground tissue of Angio-
pteris there may be seen broad openings bounded by more or
less taiigeiitially elongated cells. These are the canals from
which issue the very large volume of mucilage freely liberated
when the stipe is sectioned. These canals are always limited by
cells which differ but little from those of the surrounding tissue,
except that they are more or less elongated taiigentially. Here

•-•'Ann. Hot. 8: 214.

fl

i

!

I

fli

r I

I*

M

m>t'i\n u I, i/h lit

\\SHV\*H

llun* in no «<|»»cl;illv iliff«niiliattil t|iithiliiim, ;int! in tlii«« rtN|ntt
w.' unit will) a Icitiiii' wliitli sirv«s tn sIi.m|iIv M|iai.ili- tlus«'
.stiiiitmi's Iroin Uiuhi- hI tin- l.yiatlai «',r. < >ii llir^f uniiimh I
mIiouIiI ti'il m» hositalioii iii ^l^•t•i(lill^; as tn wlnllitr a ^;iviii plant
wi'rc cvi"iulai"iM)iiH m filimiil iti its alVinitits. l-'inm llii', |miiiiI ul
vifW, tlu-n, It \\i>ultl sitin that tlir lt>|i«'ka s|iii iimii in inun-
lu-arlv alli«'<l to Iitiih. and tin* same wniild Imlil (riu- ul Myrlu.s-
vliin. il \vr an- tn haso an i>|iinii>n ii|Min tlir i'.\«illrnl li,niin-> u\
Mr. Si'waitl.

Sin-i iiKin \i SI iiiKi'MiiVMA, — Tlu' ilistrilKilioii <>t tin- sclri-
i'lu-hyma lan liaidly l>o taki-ii as an arj,Minu-nt uiu- way m tin-
..till r. siiui- lii luilli liTiis and cyfails tln-ir is siuli widf variation.
I .should cons'clcr this a sprcilii latluT than a i-fniiu' i liar ictrr.
In all of tiu' njycloxylons so lar studiid. tlu' m lirfiu hyiii.i is
dislributi'd in st-paratc strands. In the i)iads studinl l>\ inc
this tissue forms a continuous i)and in all lasi-s wht-rr slrnn^ly
di'\clo|n"d. In .\n>jio|itiTis it forms a rontiniions /oiu' of ron-
sidcrahli.' lliickiu'ss, with si'|>aralc stramls lyin^^ aloii^f tin.- iiiiur

face.

A rcsunu' i>f the results above- dctailo<i shows that in tin-
To|>i.'l<a s|ii'cinicn tluMV arc characters which directly connect it
with Kitiliioptcrii Willi, nnsoni, and aUo with other l-airopean spe-
cies of Mvelo.\ylon, and tlu- ividence would seem to indiiate
that few of these can be separateil ifeiierically. yUlmittinu the
force of sonii- of the objections raised by Mr. Sewaid respecting'
the tilicoid character of Myeloxylon, there are, nevertheless,
Strom,' arguments in favor of this \iew. which seem to me to jire-
poiulerate and thus to justify the retention of the name Myelop-
teris as a name expressive of this possible relationship; while
the fact that these plants do not conform closely It) any modern
tvpe would seem to raise a ipiestion as to the possible correctness
of the view oriifinally expressed by (ioeppert that these plants
in realitv represent a ^Generalized type occupyini;- a position
between the cyads and the ferns.

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