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SMITHSONTAN
MISCELLANEOUS COLLECTIONS
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Lee INGTON
“EVERY MAN IS A VALUABLE MEMBER OF SOCIETY WHO, BY HIS OBSERVATIONS, RESEARCHES,
AND EXPERIMENTS, PROCURES KNOWLEDGE FOR MEN’’—SMITHSON
(PUBLICATION 3132)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
1931
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The Lord Galtimore Prees
BALTIMORE, MD., U. S. A.
ADVERTISEMENT
The present series, entitled “ Smithsonian Miscellaneous Collec-
tions,’ is intended to embrace all the octavo publications of the
Institution, except the Annual Report. Its scope is not limited,
and the volumes thus far issued relate to nearly every branch of
science. Among these various subjects zoology, bibliography, geology,
mineralogy, anthropology, and astrophysics have predominated.
The Institution also publishes a quarto series entitled “ Smith-
sonian Contributions to Knowledge.” It consists of memoirs based
on extended original investigations, which have resulted in important
additions to knowledge.
C. Gy ABBOT,
Secretary of the Smithsonian Institution.
(iii)
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CONTENTS
. Aspot, C. G., AND FREEMAN, H. B., Absorption lines of the
infra-red solar spectrum. 17 pp., 5 pls., Aug. 31, 1929. (Publ.
3026. )
Snoperass, R. E., The thoracic mechanism of a grasshopper and
its antecedents. III pp., 54 text figs., Dec. 31, 1929. (Publ.
3027.)
Assot, C. G., The radiation of the planet earth to space. 12 pp.,
2 pls., Noy. 16, 1929. (Publ. 3028.)
MILier, GERRIT S., JR., The Characters of the Genus Geoca-
promys Chapman. 3 pp., I pl., Dec. 9, 1929. (Publ. 3029.)
Miiier, Gerrit, S. Jr., Mammals eaten by Indians, owls, and
Spaniards in the coast region of the Dominican Republic.
TG pps, 2 pls Deci mis 1920.) (Pabk 3030;))
Berry, EpwaArp W., The past climate of the North Polar region.
29 pp., 6 figs., Apr. 9, 1930. (Publ. 3061.)
Ciayton, H. Heim, The atmosphere and the sun. 49 pp., 33
text figs., June 9, 1930. (Publ. 3062.)
Netson, E. W., Four new raccoons from the keys of Southern
Florida. 12 pp.,.5 pls., July 10; 1920. (Publ. 3066.)
HeEron-ALLEN, Epwarp, F. R. S., The further and final re-
searches of Joseph Jackson Lister upon the reproductive proc-
esses of Polystomella crispa (Linné). 11 pp., 7 pls., Nov. 26,
1030; .(Fubl;-3067. )
ScHEDL, Kart E., Morphology of fhe bark-beetles of the genus
Gnathotrichus Bishi 88 pp., 40 figs., Jan. 24, 1931. (Publ.
3068. )
Haury, Emit W., anpD Harcrave, Lynpon L., Recently dated
Pueblo ruins in Arizona. 120 pp., 27 pls., 35 text figs., Aug. 18,
1931. (Publ. 3069.)
BusHNELL, Davin I., Jr., The five Monacan towns in Virginia,
1607. 38 pp., 14 pls., Nov. 18, 1930. (Publ. 3070.)
Miter, Gerrit S., Jr., A note on the skeletons of two Alaskan
porpoises. 2 pp., I pl., Dec. 23, 1930. (Publ. 3107.)
Miter, Gerrit S., Jr., The supposed occurrence of an Asiatic
goat-antelope in the Pleistocene of Colorado. 2 pp., 2 pls.,
Dec. 22, 1930. (Publ. 3108.)
(v)
vi
17a
18.
CONTENTS
Mitter, Gerrit S., Jr., Three small collections of mammals
from Hispaniola. 10 pp., 2 pls., Dec. 24, 1930. (Publ. 3109.)
Reese, A. M., The ductless glands of Alligator mississip piensis.
14° pp. 3) pls. March 9, 193t.. (Publis srr.)
Kennarp, A. S., A. L. S., SALIsBury, A. E., anD WoopWARD,
B. B., F. L. S., The types of Lamarck’s genera of shells as
selected by J. G. Children in 1823. 40 pp., July 11, 1931.
(Publ. 31125)
McInpoo, N. E., Tropisms and sense organs of Coleoptera.
10 pp., 2 pls:, 19 text figs, Apr. 18, 1931. (Publ. 3113")
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SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 82, NUMBER 1
ABSORPTION LINES OF THE INFRA-RED
SOLAR SPECTRUM
(WITH FIVE PLaTEs)
BY
CG. G. ABBOT AND H. B. FREEMAN
(PUBLICATION 3026)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
AUGUST 31. 1929
The Lord Gattimore Press
BALTIMORE, MD., U. S. 4+
ABSORPTION LINES OF THE INFRA-RED SOLAR
SPECTRUM
By C. G. ABBOT ann H. B. FREEMAN
(WitH 5 PLATES)
In the decade 1890 to 1900, the bolometer was used under Langley’s
direction at the Astrophysical Observatory of the Smithsonian Institu-
tion to feel out the positions of lines and bands in the infra-red solar
spectrum. The results were published in Volume I of the Annals of
the Observatory. In the spectral region A to 2, about 550 lines were
recorded as observed in the spectrum of a 60° prism of ordinary
telescope flint.
At Mount Wilson, in the summer of 1928, Dr. H. D. Babcock
urged that further bolographic studies of the infra-red solar spectrum
should be undertaken with apparatus of higher resolving power. Our
Bie. 2.
vacuum bolometer equipment, then on Mount Wilson, presents a sensi-
tive strip of approximately 0.1 mm. width, and the combined outfit
of bolometer and galvanometer was certairily not less than five times
as sensitive as the most sensitive outfit employed at Washington 30
years before.
It appeared practicable to undertake a brief bolographic study of
the upper infra-red solar spectrum from A to © in the time available.
Accordingly we set up a spectroscope (fig. I) comprising a slit 6 cm.
high and (usually) 0.4 mm. wide; a collimating cylindric mirror of
543 cm. focal length; a set of three telescope flint-glass prisms, two
of 60°, the third of 64° angle, and all presenting faces approximately
6 cm. square. From thence a plane silver-on-glass mirror reflected
the spectrum to an image-forming spherical mirror of 230 cm. focal
length. The vacuum bolometer, above mentioned, whose sensitive
strip was 16x 0.1 mm. received the rays at focus. The spectroscope
SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 82, No. 1
2 SMITHSONIAN MISCELLANEOUS COLLECTIONS , VOL. 82
was fed by a two-mirror coelostat with silver-on-glass mirrors. The
solar rays were not concentrated on the slit. Hence they represented
the integrated rays of the entire solar disk.
The infra-red solar energy spectrum was recorded on moving photo-
graphic plates 8 x 24 in. in surface. The clockwork was arranged
so that 4 cm. of plate corresponds to 5’ of spectrum, and the plate
passed the recording light-spot of the galvanometer at the rate of
2 cm, in I minute.
The three prisms were set according to computations so that the
beam of rays of wave-length 1.054 would pass through each one
of the prisms approximately in minimum deviation. This same setting
was continued unchanged in all the observations. The total deviation
of the rays of this wave-length was roughly 180° and the dispersion
from A to 2 was about 5° 25’. Hence we were obliged to use five
61-cm. plates to cover the entire region with overlap sufficient for
identification.
Generally three curves of each of the five regions were impressed
on a single photographic plate. Care was taken to arrange them
vertically in close superposition, so as to facilitate comparison. Plates
I to 5 give reproductions of some of the most satisfactory observations.
Linear scales are drawn on plates I to 5 parallel to the direction
of motion of the recording photographic plate. They have numbers
closely agreeing with those of the extensive table 3 of linear measures
and wave-lengths, given below. In each group of three curves the
air-mass of observation decreases as between the several curves from
the bottom upward and in each curve (except in pl. 5) from left
towards right. In most plates there is a very considerable increase
of air-mass between the upper and lower curves. This will facilitate
the discrimination, by those interested, of solar and telluric lines. De-
tails of times of observation and air-mass and notes on the conditions
are given in table 2.
A very considerable increase of detail appeared in these energy
curves when compared with those taken 30 years ago with a single
glass prism. In the A line, for instance, not only could the doubles
be recognized, but in many of them the individual components were
resolved separately in the energy curve. Some of the bands near wave-
length 0.82 showed as many as five veridical lines in the new curves
where only one broad band could be distinguished in the older work.
The identification of lines was done entirely by Mr. Freeman, and
in the following manner. A series of several bolographs was super-
posed, either on millimeter cross-section paper or on a comparator
NO. I INFRA-RED SOLAR SPECTRUM—ABBOT AND FREEMAN 3
in which a stretched wire was displaceable over a milk-glass back-
ground. Lines were considered provisionally veridical when found
as deflections of similar form and-similar setting in three or more
bolographs. After completing this preliminary study, the positions of
all deflections considered possibly veridical were measured on three
bolographs with the excellent Warner and Swasey comparator de-
scribed on page 64 of Volume I of the Annals. Mean values were
computed of positions on three (or in some cases two) of these
bolographs on which the deflections were found. When found on only
two of the three they were questioned, and rejected unless supported
by further evidence.
In assigning intensities, Mr. Freeman used practically the same
system that was used in Volume I of the Annals. Grades a, b, c, d,
and d? were employed. All lines falling within great bands like
A, por, ¢, w, and © are joined in a bracket and designated as a whole
with “a.” Bands hardly reaching this first-class prominence are
similarly bracketed and marked “ b.” Individual deflections, or com-
posites of several small deflections which altogether make a depres-
sion of 5 mm. or more in bolographs are marked “cc.” Smaller indi-
vidual deflections, whether in the bands or outside of them, are marked
“d.” When considerable doubt remains as to the veridical character
of such a deflection, it is marked “d?.” We do not guarantee that
all the lines included in the table are veridical, but we believe a very
large proportion of them are so. The curves are very free from
accidental deflections as deep as a single half millimeter, and the
repetition on several bolographs of similar configurations of intensity
“d” is regarded as strong presumptive evidence of reality of corre-
sponding solar or terrestrial absorption lines.
To determine the wave-lengths of the lines observed, the advice of
Dr. Babcock was sought. From his studies of all existing laboratory
determinations of infra-red line spectra, amplified by his own extensive
photographic work in the upper infra-red spectrum as far as A=
1.1018 Angstroms, he sent a list of 112 identifications of wave-length
places, given according to our bolographic work in Volume I of the
Annals, as compared to more recent determinations. A curve of cor-
rection to the wave-lengths given in Volume I of the Annals has been
prepared from this material. In summary it is as indicated in table 1.
The data for corrections beyond 1.18 are so scanty and so conflict-
ing that there seemed no justification for applying any corrections in
that region.
4 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
TABLE 1.—Corrections to Wave-Lengths of Annals, Volume I
(Corrections are stated in Angstroms, wave-lengths in microns.)
Wave-lengths...| 0.76 to 0.84 | 0.85 | 0.86 | 0.87 | 0.88 | 0.89 | 0.90 | 0.91
@orrections.-..- O +5 +5 +5 +3 +2 fe) —2
Wave-lengths...| 0.92 | 0.93 | 0.94 | 0.95 | 0.96 | 0.97 | 0.08 | 0.99 | 1.00
Gornections.-5- —2 | —I fe) +2 +2 +1 0 \os=3 1 es
Wave-lengths...| 1.01 | 1.02to 1.06 | 1.07 | 1.08] 1-09] 1.10| 1.11/1.12|1.13
Corrections... ..- —8 —I10 — 9 | —7 | —6|] —5 | —4 | —3 | —1
I.14 to end
0
Wave-lengths...
Corrections.....
In further determination of wave-lengths, Mr. Freeman identified
81 deflections as corresponding each to each in the old and the new
bolometric work. These deflections covered fairly uniformly the range
from 0.76u to 1.804. Having taken out from the tables of Volume I
of the Annals the corresponding wave-lengths, he then applied to these
values the corrections fixed by table 1. He then plotted on a sufficiently
large scale the observed linear places of these 81 deflections as ordi-
nates, and the corrected wave-lengths as abscissae. The curves thus
defined could easily be read off to a single Angstrom. From them
were read all the wave-lengths given in table 3, which contains over
1200 lines.
TABLE 2—Circumstances of Observation
Time Air-mass
Date a
1928 Curve Start Finish Start Finish Notes
Sentara aca I 0: 39 10: 09 Legit 23
2 MOE: 107 10: 47 ee 1.16
10: 52 II: 22 Te is Wo l2
Sentifaaeoe ce I 6: 28 6:58 4.85 3823
2 FE RIO) 8: 29 1.07 1.69
3 8:51 Q: 21 1.54 1.38
Septins. eee. I 6: 34 7:04 4.49 3.05 Sheht earthquake
2 Gj? fil 9: 41 1.43 pitt
3 9:50 10: 20 1.28 1.20
Sepitere ays arerat I 6: 29 6:59 4.70 Bio 1s
2 9:19 9: 49 1.36 Te27)
3 9:58 10: 28 1.24 Tay,
Septreiiacenscee 3 3:34 3:54 1.82 2.18 Ends off plate
2 2:58 a8 BS 1.54 70
I Tens TA 1 17 123
NO lat
INFRA-RED SOLAR SPECTRUM—ARBBOT AND FREEMAN
TABLE 3.—Lines and Bands in the Infra-Red Solar Spectrum
Linear
measures
—34.130
34.035
33-103
32.858
Intensity
VW
aAanmaanananaanaanaAaAAnaAAAAAnaAnaAannananAamA
Wave-
length
7582
7583
7508
7602
7604
7600
7608
7618
7628
7630
7636
7638
7641
7642
7645
7640
7647
7051
7652
7656
7658
7661
7662
7665
7668
7672
7674
7675
7680
7684
7686
7601
7692
7695
7608
7701
7793
7795
7797
7710
Fifi
7712
7714
7718
7722
7725
Linear
measures
— 25.233
22
25.019
24.831
.418
322
-233
153
24.030
23.801
.696
573
461
.176
23.079
22.904
.690
621
.272
ily
22.046
21.906
807
.696
472
343
.240
21.005
20.871
.687
521
318
20.114
19.935
826
.756
.650
547
.226
10.111
18.951
837
755
18.080
17.949
830
Intensity
~w
=!
BaAAaAAAAAAKAAAA A
CEO CW)
wv
QaaannnnaaaanaAAAAAAannAAaAanAananaaamha
“w :
Wave-
length
7727
7728
N
SSS Se ee
DS SS ESS SS SS ee fl i ast
6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
TasLe 3.—Lines and Bands in the Infra-Red Solar Spectrum.—Continued
Linear Wave- Linear Wave-
measures Intensity length measures Intensity length
17720 d 7852 —5.692 d? 8069
16.273 d? 7878 5.570 d 8071
15.621 d 7889 4.806 d 8084
cago id 7804 “Capea idl: 8086
15.136 d 7808 635 d 8088
14.932 d 7901 500 d? 8001
.823 d 7903 .400 d 8003
.644 d 7906 .286 d 8095
470d 7909 Ar53 (od 8007
14.207 d 7914 3.747 d 8106
13.910 d 7919 645 d 8108
625 d 7924 543 d 8109
.500 d? 7926 384 d 8112
.406 d? 7928 .206 d 8114
.196 d 7932 3.105 d? 8118
13.050 d 7934 2.967 d 8120
12.861 d 7937 658 d 8126
704.) od 7939 537 (7d 8129
.676 d 7941 378 d 8132
450 d 7945 2.255 d 8134
1275 d 7948 1.839 d 8142
12.055 d 7952 536 d 8148
11.844 d 7950 .338 lc 8152
.720 d? 7958 238 d 8154
.604 d -c 7900 .142 d b 8156
324 d 7966 1.062 d 8157
208 d 7968 0.812 d 8162
11.004 d 7072 739 d? 8164
10.619 d 7979 0.000 € 8178
9.866 d 7992 .123 d 8181
765 d 7904 | .469 d 8188
688 awd 7906 Or bad 8100
389 d 8001 | 0.775 d 8103
283 d 8003 +1.037 d 819090
0.102 d 8006 | 349 d | 8206
8.098 d 8008 | EL ol ae a! 8214
8.249 d 8022 1.854 d 8216
7.982. d? 8027 2220) ) a? 8224
818 d 8020 .206 d 8225
530 d 8035 392 d 8228
hist d 8038 468 d? b 8220
7.011 d 8044 -700 d 8233
6.927 Gly fete 8045 2.800 d 8235
769 d 8048 3.028 d 8240
647 d 8051 .243 d 8244
6.541 d 8053 336 d 8246
5.055 d 8063 463 d 8249
856 d 8065 5690 d 8251
NO. I INFRA-RED SOLAR SPECTRUM—ABBOT AND FREEMAN
TABLE 3.—Lines and Bands in the Infra-Red Solar Spectrum.—Continued
Linear Wave- Linear Wave-
measures Intensity length measures Intensity length
+3.612 d 8252 +12.086 d 8430
691 d 8253 12.518 d 8440
.820 d | Es 8256 13.006 d 8450
3.927 d 8258 128 d? 8453
4.036 d 8260 .243 d 8455
-706 d 8274 .675 d 8465
794 +~d 8275 13.9048 d 8470
4.892 d 8277 14.416 d 8480
5.108 d 8282 14.509 d 8484
Tor. od 8283 15.042 d | 8404
.421 d 8288 .132 d 8406
.482 d 8280 .231 d ve 8408
506 d? 8201 333 d | 8500
5.829 d 8207 .604 d 8508
6.124 d 8303 15.037 d 8513
.233 al He 8305 16.046 d 8516
304 .- d? 8307 1g2) a ] 8517
.476 d 8310 360 d 8523
S05) . ‘d 8311 16.468 d ¢b 8525
.668 d 8314 17.033 d 8537
6.996 d 8321 218 d 8541
7.163 d 8325 17.947 d 8558
431 d 8330 18.144 d 8562
549 d 8333 260." di? 8565
.666 d | 8335 .456 d 8560
qos. dd 8337 2550", dd 8572
8.003 d 8342 761 d 8576
200), id 8346 18.864 d 8570
.490 d 8352 19.044 d? 8583
743 «2d 8358 138: d 8585
8.888 d 8361 .242 d 8587
9.204 d 8368 19.722 d 8500
317 d 8370 20.251 d 8611
394 d? 8372 398 d 8614
.467 d 8373 20.610 d 8620
575d 8376 21.060 d 8630
o731": d 8379 201 d 8633
10.023 d 8385 .488 d 8640
.260 d 8301 591 d 8642
499 ~d 8306 21.692 d 8645
.697 d 8400 22.256 d? tp 8658
10.814 d 8403 381 d 8662
11.043 d 8407 404 d 8663
.288 d 8413 22.625 d 8667
5900 d 8420 23.056 d? 8677
11.966 d 8428 254 d 8682
8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
TasLe 3.—Lines and Bands in the Infra-Red Solar Spectrum.—Continued
Linear Wave- Linear Wave-
measures Intensity length measures Intensity length
+23.472 d 8687 +32.148 d 8805
562 d 8089 226 d 88907
684 d? 8692 348 d? 8900
23583 4d 8694 5201 «acl? 8904
24.568 d? 8713 733 d ) 8909
25.223 d 8728 32.901 d | 8915
302 «od 8732 33.101 d | 8918
tod 8735 419 d | 8926
506 d? 8737 33.860 d | 8937
1682), \ 4d 8739 34.014 d | 8941
25.823 d 8742 222 © «d? | 8946
26.181 d? 8750 414 d | 8051
288 d 8753 .740 d 8959
.460 d 8757 34.801 d b 8062
705 oad 8763 35.0900 c | 8067
302, - d 8766 448 d | 8076
26.931 d? 8768 35.791 d be 8984
27.052 d 8772 36.049 d 8990
43nd 8774 Bee 8 Gl 9004
261 d 8777 Ges d goo
404 od 8782 36.805 d Qo14
689 d 8787 37.060 +d 9018
27.996 d 8704 680 d 9036
28.105 d 8707 37.819 d 9039
223 «od | 8800 38.004 4d 9047
E332 d 8803 .180 d? 9049
385 d Ve 8804 502 d 9060
ce) | 8811 Beceem 9060
783 d 8813 38.074 d? 9071
.879 d 8816 39.582 d 9087
28.973 d 8818 .783 d bp 9092
20.247 d? 8824 878 d 9095
.402 d 8820 39.892 d 9096
524 d 8832 40.040 d? Q100
.720 d 8837 .192 d P gro4
823 d 8830 1273 d F 9106
29.903 d 8841 40.603 d OII5
30.036 d 8844 AT-GO2.) |) od | 9127
.166 d? 8847 .120 d ‘tc 9120
e3059 \) wal 8852 41.423 d | 9137
.540 d 8856 42.046 d Ik | O154
.667 d 88590 165 d tc Lh bq totex
843 d -c 8864 255 d | | Q160
30.041 d 8866 ‘800. id | 9175
31.128 d 8870 42.985 d 0179
247 d j 8873 43.047 d le oI181
31.881 d 8880 431 d | 9192
32.004 d? 8803 513 d? Q104
NO. I
INFRA-RED SOLAR SPECTRUM—ABBOT AND FREEMAN
TABLE 3.—Lines and Bands in the Infra-Red Solar Spectrum —Continued
Linear
measures
41-43-7 11
43-884
44.101
.278
-347
525
.623
776
44.977
45.101
235
.310
.420
-540
.729
854
45-949
46.221
-337
836
46.907
47.101
3212
533
47-637
48.205
-205
509
48.817
49.124
49.648
50.022
51.122
.204
328
421
51.737
52.107
52.523
53-039
487
53.963
54.060
-174
54.278
55.008
Intensity
GaOoamamanmaaAaA A
—-—- —-———— YY
~w
mw
TAHAaAaAanaanannntAaAhAAhAhAaAaAanhAaAAnAnhAaatamananas
lo)
SS
ao
rb
—
Wave-
length
9200
9204
9210
9215
9217
9222
9225
9229
234
9238
9241
9244
9247
9250
9255
9258
9261
9269
9272
9286
9288
9204
9296
9305
9308
9324
9327
0333
9342
9351
9366
9376
9409
Q412
9415
9418
9427
9439
0451
9467
9480
9495
9498
9502
9506
9529
Linear
measures
+55.674
50.010
369
50.738
57-254
57-503
58.073
-245
.352
534
58.667
59.058
472
568
59.642
60.482
773
60.975
61.069
544
61.931
62.229
.532
634
62.745
63.103
.666
63.956 °
64.252
724
.898
64.085
65.224
342
.464
561
65,603
66.141
285
421
617
66.923
67.250
.486
675
67.772
9
Intensity
d
Q
ae amend
inn
in
BAAOAAAhAAAaAARhnAnhAnmhAAaAAAAAGA
a)
ed
aQ
BAAAAAAAARAARAAAARAAAA
~
Wave-
length
9548
9558
9508
9580
9596
9604
9621
9626
9620
9634
9639
9651
9663
9666
9668
9694
9704
9710
9713
9727
9739
9749
9758
9761
9765
9777
9795
9804
9814
9829
0834
0837
0845
9848
9853
9856
9860
9874
9879
9883
9890
9900
goto
9917
0024
9027
10 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
TaBLe 3.—Lines and Bands in the Infra-Red Solar Spectrum.—Continued
Linear Wave- Linear Wave-
measures Intensity length measures Intensity length
+68.099 d 9937 +74.504 d IOI50
.248 d 9042 .626 d 10154
643 d 9954 -750 d 10159
751 d 9958 74.912 d 10164
1870. | ‘d 9962 75.211 d 10175
68.9907 sd 9066 sat di? 10179
69.0908 d 9909 520 od 10186
235"), 1G 9973 708, id 10192
375...) a 9978 75.830 d 10104
Sy i) od 9083 76.001 d? 10202
615 d 9986 .108 d 10206
772 d QgoI 329 d 10214
863 d 9004 2510) id 10221
69.972 d? 9097 .631 d 10224
70.060 d 10000 76.765 d 10229
.179 d? 10004 77.033 d 10239
401 d? 10010 L227, d 10245
509 «od T0014 1933." od 10249
.634 d 10018 .470 d 10254
-706 d 10024 732 d 10263
70.044 d 10028 77.855 d 10268
71.061 d 10032 78.019 d 10274
452 d 10044 .150 d 10278
.562 d 10048 .289 d 10283
.676 d tb 10053 368 d? 10286
780 d 10056 Ay d 10292
878 d 10060 AGG Cal 10206
71.083 d 10063 757 d? 10300
72.088 d 10067 827 d? 10302
.220 d 10072 78.961 d 10308
natin) d 10075 79.088 d 10312
380 d? 10077 1217 d? 10316
-404 d? TOO8T 346 d 10321
.603 d 10084 534 d 10327
.719 d T0088 Wee d 10334
72.821 d? 10092 842 d 10338
73.023 d 10099 79.908 d? 10344
.139 d 10103 80.219 d? 10352
238 d 10106 .320 d? 10356
317 d? IO1I09 521 d 10363
401 d? 10112 6190 d 10367
514 d IOIIO. .721 d? 10370
691 d 10122 833 d? 10374
73.990 d 10132 80.988 d 10379
74.208 d 10140 81.105 d 10384
323 d 10144 .220 d 10388
NO: I
INFRA-RED SOLAR SPECTRUM—ABBOT AND FREEMAN
Wu
TaBLe 3.—Lines and Bands in the Infra-Red Solar Spectrum.—Continued
Linear
measures
+81.324
.450
.569
81.680
82.016
.120
Intensity
d
d
d?
any,
BA AAAAaAaaohAaAaaahnadahnannhnamaamaanhnanmaaaaaasa
“vu
Wave-
length
10392
10396
10400
10405
10417
10421
10428
10432
10437
10441
10450
10453
10457
10462
10467
10479
10484
10490
10494
10408
10510
10518
10525
10537
10547
10556
10561
10578
10585
10504
TOOOI
10607
IO6II
10621
10623
10636
10643
10650
10662
10673
10680
10684
10688
10604
10607
10708
Linear
measures
+89.892
89.995
90.139
336
441
533
90.753
OI.091
249
344
.467
.566
.667
776
91.953
92.260
472
550
654
782
.866
92.980
93.046
232
-393
.487
Gian
93.962
94.050
.469
554
.679
94.777
95.373
506
625
95.860
96.124
-234
353
96.782
97.003
-119
272
383
.485
Intensity
d?
BaOaOMOaOAMAAaAAAhAaAaAaaAhnoahmAhanaAahnnaaanmanhnanaanaananhamamanhanhanmana
vu
Wave-
length
“10714
10719
10724
10732
10736
10740
10749
10762
10768
10772
10777
10781
10785
10789
10796
10808
10816
10820
10823
10828
10832
10836
10839
10845
10852
10855
10866
10874
10878
10894
10808
10902
10906
10930
10936
10941
10950
10960
109064
10969
10986
10996
11000
T1006
IIOII
IIOIS
[2
SMITHSONIAN
MISCELLANEOUS
COLLECTIONS
VoL. 82
TABLE 3.—Lines and Bands in the Infra-Red Solar Spectrum.—Continued
Linear
measures
+97.673
97.898
08.211
-203
.400
-509
98.737
99.101
-435
.707
99.806
100.387
473
100.900
IO1.102
101.704
102.378
-447
.862
102.960
103.224
761
832
103.934
105.170
105.993
106.242
-595
684
106.790
107.588
107.855
108.607
723
108.808
109.102
380
109.702
110.141
P2R7,
E72
110.912
ITI.116
.587
ITI.669
Intensity
aaamamaa
fv)
~w
AaanannannanaaAntananAAnATFAnAnaanAnananhananamm&Aa
inn
Wave-
length
11023
TIOS2 2
11045
11048
11052
T1057
11066
11081
11004
11106
EEE
T1133
11137
T1155
11165
IIT190
J1217
11220
11238
11242
11254
11278
11281
11285
11340
11376
11388
11404
11407
T1412
11448
11460
11401
11406
11409
TISII
11524
11537
T1555
T1559
11574
T1588
T1596
11616
11620
Linear
measures
+ 112.005
.187
292
112.888
113.051
152
.466
614
113.948
114.119
823
114.916
115.428
115.821
116.084
397
116.914
117.034
.140
117.726
118.034
-154
322
448
.600
720
118.828
119.133
2211
-334
440
119.738
120.059
.232
120.424
T2027;
.200
303
427
821
121.9014
122.022
-134
248
.406
Intensity
BOAAAMAAARARARAAhAAnAAaaAAanAaananaaaana BaqnrRAAhAAnnAnanhnmaAa Aa
Wave-
length
11634
11642
11646
11670
11678
11682
11696
11702
11716
11723
11754
11758
11780
11797
11808
11822
11845
T1850
11855
T1881
11895
11900
11908
IIQI4
11921
11927
11931
11945
11949
11954
T1959
11972
11988
11996
12000
12038
12042
12046
12052
12071
12075
12080
12085
12001
12008
NO. I
INFRA-RED SOLAR SPECTRUM—ABBOT AND FREEMAN
13
TABLE 3.—Lines and Bands in the Infra-Red Solar Spectrum.—Continued
Linear
measures
+122.514
{oni
122.759
123.132
gi
.480
-594
.720
842
123.957
124.044
-177
.438
124.918
125.023
-197
Sirs
396
-495
.627
-707
125.859
126.113
.204
AII
579
.700
-799
126.941
127.047
173,
303
.406
536
644
734
839
127.914
128.025
Heit
.222
-309
415
546
667
783
Intensity
a &
vu
wow “vu
WU}
B@AaaAaAAMARAHRAAAKAAAARAAAAMAAAk Aa
vu
foal
ary
ba
aaaam
Qe Oe
aaadaamdmama
Wave-
length
12104
12108
12115
12133
12140
12150
12156
12162
12168
12173
I2177
12183
121096
12219
12224
12232
12236
12242
12247
12254
12257
12205
12277
12286
12204
12300
12306
12310
12317
12322
12329
12335
12340
12347
12352
12356
12362
12366
12372
12376
12381
12386
12301
12308
12404
12409
|
Linear
measures
+128.914
129.009
.219
319
-437
504
129.903
130.009
Ain)
248
338
545
130.767
131.216
.409
.526
.620
Gin
838
131.927
132.006
-193
agit
307
-473
.600
608
132.799
133.551
134.119
304
134.469
135.049
.201
weil
401
.507
.620
135.961
130.071
186
619
807
136.937
137.031
150
Intensity
a
paar
QaoamnaAanAhnanhtaannaanaahaanhAahaaaaa & &
ao
d
Wave-
length
12410
12422
12432
12437
12443
12450
12468
12473
12478
12485
12489
12501
12512
12536
12540
12552
12550
12562
12568
12572
12577
12586
12593
12506
12601
12608
12613
12618
12658
12688
12608
12707
12730
12748
12753
12759
12764
12770
12788
12794
12799
12822
12833
12839
12845
T2851
[4 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
TABLE 3.—Lines and Bands in the Infra-Red Solar Spectrum.—Continued
Linear Wave- Linear Wave-
measures Intensity length measures Intensity length
+-137.264 d 12858 + 143.603 d | 13198
.4II d 12866 W277, d | 13205
493 1 od 12870 834’ id) 13210
578 d? 12874 143.940 a hg 13217
137.878 d 12891 144.013 d 13221
138.016 d 12898 114 .d 13226
Teen d 12903 .867 d 13267
.231 d 12909 144.959 aq \h 13272
.301 d? 129013 145.434 d 13209
.427 d 12920 508 d 13302
806 d 12940 732 d? 13314
138.930 d 12945 145.820 d 13319
139.048 d 12053 146.484 d 13355
212 d 129062 146.565 d 13360
324 d 12968 147.020 Cc 13385
.489 d 12976 199 c 13306
500 id 12983 147.738 ¢ 13425
7735 d 12990 148.417 Cc | 13464
835 d 129095 .905 d? 13401
139.942 d 13001 148.901 d 13495
140.038 d ] 13006 149.005 d 13501
114 d [ 13010 350 d 13516
210 d rc 13015 .446 d 13522
303: nd 1302¢ 549) iy di rc 13527
424 d 13026 651 d 13532
2025) 13037 149.744 1d 13538
719 d 13042 150.069 d 13555
834 d 13048 .144 d 13560
.926 d 13053 461 d 13578
140.907 d? 13057 508 d le 13580
141.086 d 13061 793 d | 13507
308 d } 13074 150.910 d 13604
.412 d i 13080 151.259 i: | 13623
520 d | 13085 358 doc 13628
617 d 13090 442 d | 13634
141.877 d 13104 815 d | 13654
142.120 d 13118 151.920 d 13660
.236 d 13124 152.024 d? tc 13667
bBRO indus 13120 28a id? 13681
449 42~d 13136 “303° ad 13687
564 d 13142 S45 gids »| 13606
Os th wud 13150 Ts2ee5 ) id i 13716
844 d 13158 153.024 d rc 13724
142.062 d 13163 163 d La772
143.070 d 13170 273 d? 13738
216 aivilre 13178 .467 d 13740
330) aiud 13183 866 d 13772
.446 d 13190 153.956 d 13777
NO. I
INFRA-RED SOLAR SPECTRUM—ABBOT AND FREEMAN
15
TaBLe 3.—Lines and Bands in the Infra-Red Solar Spectrum.—Continued
Linear
measures
+154.208
.206
617
724
154.808
155.010
113
[222
.408
.507
155-936
156.826
157.693
158.425
759
158.859
159.280
587
159.691
160.140
.632
160.969
161.049
-379
534
161.675
162.033
104
162.476
163.155
163.520
164.005
433
.515
755
164.973
165.479
166.341
167.125
167.435
168.031
119
.495
168.033
169.060
765
Intensity
d? |
|
2
aS
mm
SS)
Q
“wu
a TAA nDTAAnHTFAAAAA AAA MA &
Q
loi
TTrTAndqnAn AA rFTrTAAnangnartrtanA
vu
Wave-
length
13781
13796
13815
13821
13833
13838
13843
13850
13860
13866
13890
13942
13094
14030
14057
14060
14086
14103
14100
14135
14164
14184
14188
14206
14216
14225
14245
14250
14271
14312
14333
14366
14385
14390
14404
14416
14447
14495
14542
14560
14594
14590
14621
14646
14655
14606
Linear
measures
+ 169.987
170.701
170.883
171.330
618
171.990
172.090
.501
172.604
173.270
.360
173.918
174.011
win
.436
174.805
175.205
£332
175.987
176.007
.580
842
176.967
177.301
.467
677
177.799
178.131
.200
.408
-597
715
775
883
178.980
179.115
201
Intensity
~~
a
Q
Coy
vu
J)
(or
ow
uy ow,
GS Be as ein Ps) Bk (PsP Pas Oe Ake Ces Bae RS ee BSR Rm BARA AAAAAnRnaAAAaAnAgqgnhnanA
A. Aa A
wm;
Wave-
length
14708
14756
14760
14788
14804
14826
14832
14855
14862
14901
14907
14938
14044
14962
14969
14991
15014
15021
15060
15070
15100
15112
15120
15144
15150
15162
151690
15180
15108
15211
15217
15224
15228
15234
15240
15248
15253
15200
15267
15276
15203
15310
15317
15324
15347
15356
16
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOL. 82
TABLE 3.—Lines and Bands in the Infra-Red Solar Spectrum.—Continued
Linear
measures
+ 181.173
615
-690
824
181.930
182.058
.206
481
182.959
183.182
-207
478
782
183.881
184.420
184.707
185.016
644
185.745
186.107
-353
-493
186.705
187.001
365
187.759
188.458
661
188.908
189.338
190.081
-479
190.754
191.205
313
551
.604
191.889
192.065
.207
563
192.713
193.051
.148
278
.501
Intensity
wu
O45)
QaAaAAaAahhanaAaahanhaaaa ea
fol (ary (eh (oh
aoAmnaanaaqanhmaamama &
=
Wave-
length
15371
15397
15402
15410
15416
15424
15436
15447
15478
15491
15499
15509
15527
15533
155006
15583
15601
15639
15044
15671
15680
150890
15702
15719
15740
15764
15806
15818
15833
15859
15904
15927
15044
15971
15977
15001
16000
16012
16022
16037
16052
16062
16082
16088
16005
16108
Linear
measures
+193.742
193.897
194.049
245
431
.507
682
854
194.981
195.111
308
-557
780
195.958
196.103
.207
196.607
197.052
234
.503
.640
.770
.900
197.994
108.157
212.
-449
825
198.964
190.344
-544
.660
199.923
200.085
-235
305
cr
678
200.950
201.054
282
.695
201.778
202.134
244
202.775
Intensity
wv
Brus, ony
~v
BaBOaAAaAaAanahnhaAahnhamaanahnamanmaha
vu
AaaaaaaaAa
ei] uy nw) =
“vu
jay (ov fel [al fork [al (oly alk fob fal [aly [aly (oly jak
-U
Wave-
length
16122
16132
16141
16154
16165
16170
16180
16190
16108
16206
16223
16232
16246
16256
16265
16272
16301
16322
16333
16350
16358
16362
16373
16379
16390
16309
16407
16430
16438
16462
16473
16481
16407
16507
16516
16524
16534
16543
16560
16566
16580
16605
16610
16632
T6651
16672
NO. I INFRA-RED SOLAR SPECTRUM—ABBOT AND FREEMAN Ly,
TaBLE 3.—Lines and Bands in the Infra-Red Solar Spectrum.—Continued
Linear Wave- Linear Wave-
measures Intensity length measures Intensity length
+203.065 d 166890 +214.210 d | 17380
225 d 166909 214.591 Cc 17404
203.421 d 16711 215.326 c 17449
204.042 d 16748 215.855 d? 17482
3365 d 16768 216.083 d 17495
417 d? 16772 .207 d +c 17504
.550 d? 16780 422 d? 17510
204.851 d 16798 754 d 17537
205.087 d 16812 835 d 17542
.443 d 16834 216.959 d 17550
SY, d 16839 217.123 d 17560
.678 d 16848 217.662 c 17593
789 d 16858 218.336 c 17634
205.982 d 16865 218.958 d 17673
206.140 d 16877 219.168 d 17686
206.630 d 16907 .288 d 17602
207.242 4d 16044 Bao sd 17704
320 d 16950 219.607 Cc 17718
.630 d 16969 220.119 d 17744
207.863 d | 16084 724 c 17782
208.031 d? | 16904 857 d? | 17790
245 d 17006 220.988 d b 17797
492 d 17022 221.105 d | 17805
208.960 d 17051 743 d 17843
200.367 d 17077 221.933 d \ 17855
.490 d 17084 222.043 din® 17862
200.758 de 17102 222.538 d | 17892
210.031 d 17119 223.207 deiac 17937
544 d 17150 223.520 d 17952
.846 d 17168 224.172 Cc 17902
210.965 d Q 17176 .692 d 18024
211.700 d 17223 224.904 Gaaac | 18042
211.822 d a 17220 225.155 d 18052
212.284 Cc | 17258 742 d 18088
880 cL 17200 225.017 d 18100
212.981 de fe 17304 226.318 d 18124
2133537 d 17338 .469 d 18132
.668 d 17347 226.816 d 18154
Gs. dis te 17353 227.152 d | 18174
213.910 d? 17362 227.486 d 18194
oe oe
|
ae
>
-_
- SMITHSONIAN MISCELLANEOUS COLLECTIONS
{ Botocrarns or THE INFRA-Rep PrisMATIC SOLAR SPECTRUM.
The A region, Wave-lengths 7600 to 8600 Angstroms.
VOPR 3 2.eNOreti) peae
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOES S27 NO dG been4:
2 180 |
t.,
7b
my
-
et,
.
i
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOL. 82, NO. 1, PL. 5
Ay
BoLoGRAPHS OF THE INFRA-RED PrisMATIC SOLAR SPECTRUM
The region. Wave-lengths 15200 to 18200 Angstroms
bas
f
a
f
Saeel
:
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P|
3
j
e
I
F
;
1 ,
‘
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ALSTON Sn)
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 82, NUMBER 2
THE THORACIC MECHANISM OF A
GRASSHOPPER, AND ITS
ANTECEDENTS
BY
R. E. SNODGRASS
Bureau of Entomology,
U. S. Department of Agriculture
m SO
7 s\o) b
(PUBLICATION 3027)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
DECEMBER 31, 1929
TBe Lord Baltimore Press
BALTIMORE, MD., U. S. A.
THE THORACIC “MECHANISM OF A> GRASSHOPPER,
AND TYS*ANTECEDENTS
By R. E. SNODGRASS
BuREAU OF ENTOMOLOGY
U. S. Department of Agriculture
CONTENTS
PAGE
IPT CHEMIE STONER yas, a, «= Gok ETN gee eM tae Bee IRAN tLe had es pce ides I
Peet Ali CISCUSSION :.. csee Lie toe enero aes era lets cos, ie elas beg aeons Sd 3
Wie yehoracie tergela ce elvatcre eee eee ear eio cre ee he ese tere Wh techies es 5
hex thoracie ’ pletirasceinae seco ee ee ee eee ie oe 10
Rhemthoracie stertiadaensa cee ee eT eo eee tater eee AA
hieethesthoracic skeleton ot Oissosteuaeeneee isms eine ee eek 33
Whe scervical sclerttes. Steps eerie mes pce eas eee ierk acata cs abs oa 33
hes prothorax > bac aera Cee Cee Her eens, eros ie see es 34
Rhea pterothoraxy je scs se coe IIe Ac enna e es flere orion a 48 37
Diieehesthoractc: muscles) on Mrssosrcoraaeeseneae tee deere eine ae 51
Nimsclessor the neckandsnnothonaxeeemoeieer sini ss oeinesitie asic 53
Murscles: of the pterothiotas sr racrocterat teres crieienetons +t ere psieus creme ake uses 59
eieemesothoracic miutisc esmeennr errata re eerae eis ieee cer 60
ihe smetathoracicmusclessemernememm racemic cei alee: 66
Werine. lees and their niscles soaeesarne see sae ae sheet Aceh e'e als 72
Structire: (of thestlegis.., yp cterertre ths Sneeeteie whe Sis tea eaanese cetera pa aie lov ares 73
Miuscles’of thea least raerere ere aric teen he erecta ce eat chia 78
Veo the wanes and. their mechaitsmenr tame tee. cies 2 hk races, nevaneo vs 84
Structure. ot the ‘winesaa meee ce nose satiate seats toons weds 84
Sih writes mechanisnne tyecuer tre se eee rs ithe an. inetd syagedacadase wars 92
Wile Res spiracles... sad 5 «trclerae- tale meee arse T a ae edn ereuas eal arp slab alee a ares acne 99
Abbreviations .used on the fie tnesa-eeme eerie cereals ete ee ies P= 108
TEC TS eS stele a I eta Seis ig UU bolo om Combe ao Clorecad oem Sod en aaeaoe 109
INTRODUCTION
The principal elements in the motor mechanisms of arthropods are
the muscles and the body wall, though the blood often plays an im-
portant secondary part as a hydraulic medium. All movements, how-
ever, come primarily from muscle contractions. A contracted muscle,
when it relaxes, must be actively extended before it can operate again,
and therefore muscles generally occur in antagonistic sets. But the
muscles of insects are not necessarily opposed by other muscles ; the
counter force may be produced by the elasticity of the part of the
body wall on which a muscle is attached. For this reason it is often
SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL, 82. No. 2.
2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
found in studying the anatomy of insects that a muscle has no an-
tagonist. Moreover, while most insect muscles are muscles of motion,
some are tensors inasmuch as they appear merely to maintain rigidity
between parts that are subject to strain from other muscles.
The ectodermal cell layer of the body wall, or epidermis (commonly
known as the “ hypodermis”’), is covered externally by a cuticula of
which the general constituent is probably chitin, but in which other
substances are deposited to form hardened areas called sclerites. The
nature of the sclerotizing substances in insects is not yet known, but
it appears to be definitely established that the sclerotic areas of the
cuticula are not places where the chitin is thicker or denser (Campbell,
1929). Sclerites are secondary formations in the body wall, and it
would be both interesting and important to know the physiological
processes that produce them, for we should then be better able to
evaluate sclerites as morphological units.
The major plates of the body-wall of an insect are very definite
structures that are consistently reproduced by the deposit of sclerotiz-
ing substances throughout the whole series of insect forms, and some
of them appear to be homologous with corresponding plates in other
arthropod groups. On the other hand, all parts of the insect skeleton
called “sclerites”’ in descriptive entomology are not of equal value.
Many of them are simply areas of larger plates which have become
secondarily demarked by lines of inflection in the cuticula that have
formed internal strengthening ridges. The so-called sclerites in such
cases are in themselves of no significance. The important morpho-
logical features are the endoskeletal structures ; these are the rafters,
the joists, and the upright supports that give strength to the edifice
and enable it to withstand the strain of the muscles pulling on its walls.
It frequently happens, however, that a primary region of sclerotiza-
tion becomes broken up by a discontinuity in the hardening substance
of the cuticula, thus producing true secondary sclerites. The inter-
vening “ membrane ” may take the form of a narrow line of flexibility
(‘‘ suture ’’), or it may cover a large part of the original hard surface
and reduce the primary sclerotization to two or more widely separated
plates. Or again, an original sclerotic area may be contracted to a
relatively small sclerite, or it may be obliterated. It then becomes a
question, if the primary plate has been given a name, whether we are
to apply this name to the area originally occupied by the plate, or
restrict it to the sclerotic remnants or remnant. It is the usual prac-
tice to apply the name only to the sclerite, whatever its extent, and,
if the sclerite is obliterated, to say that the part in question is obsolete
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 3
or lost. This usage has convenience for descriptive purposes, but it is
likely to confuse our morphological conceptions, since an anatomical
part is the same thing regardless of the nature of its surface covering.
I. GENERAL DISCUSSION
If all that has been written about the thorax of insects were true,
or could be made to fit with our present knowledge of insect struc-
Pe’ “ates on ie Pe acs
\ / oe ~ | i
\ / ra oS | Z
Sen Ist
Fic. 1—Diagram of the theoretical structure of a primitive thoracic segment.
The tergum (7) includes the segmental and preceding intersegmental sclero-
tization of the dorsum; the ventral sclerotization consists of a primary seg-
mental sternal plate (Stn) and intersegmental intersternites (Jst) ; the pleural
area is occupied by a basal subsegment of the leg, the subcoxa (Scx), divided
dorsally into a eupleuron (Eupl) and a eutrochantin (Eutn).
acs, antecostal suture; c, d, dorsal and ventral subcoxo-coxal articulations ;
Eupl, eupleuron; Eutn, eutrochantin; Jsg, primary intersegmental line; J/st, in-
tersternite; Mb, secondary intersegmental membrane; Pc, precosta; Scx, sub-
coxa; Sin, primary sternite; 7, tergum.
ture, there would be little need of prefacing a special description of
the thoracic skeleton and musculature of the grasshopper with a gen-
eral discussion. Science, however, is not a collection of facts but a
concept in which to hold the facts. As our collections of facts become
larger, our concepts must be altered and enlarged from time to time.
Moreover, we often think that we have nicely fitted a fact into a
mental container, only to discover presently that it does not fit at all,
or that an important part of the fact has been left out. There is noth-
4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
ing to do then but to discard the container or to remodel it as best we
can to make it serve its intended purpose. The writer, therefore, finds
it necessary to enlarge some general conceptions previously expressed
concerning the nature of the thoracic mechanisms and their apparent
evolutions from simpler origins, in order to accommodate new obser-
vations that must be admitted.
_ The primary intersegmental infoldings of the integument of arthro-
pods are the original lines of attachment of the longitudinal muscles,
and in most cases the principle longitudinal muscles are still attached
on them. When the cuticula becomes sclerotized, the intersegmental
inflections are usually converted into apodemal ridges, and a primary
segmental plate is laid down in the dorsum and generally in the venter
of each segment. In some cases the intersegmental sclerotizations take
the form of narrow intersegmental sclerites alternating with the seg-
mental plates. This condition is found more frequently in the ventral
than in the dorsal region, though it exists dorsally in some insect
larvae.
The typical sclerotization of the dorsum of any segment in an adult
insect consists of a plate (fig. 1, 7) which covers most of the dorsal
area of the segment, and which is continuous anteriorly with the inter-
segmental sclerotization bearing the intersegmental fold or ridge
(fig. 2 A, Ac). The definitive tergum, therefore, occupies a primary
segmental region and the preceding intersegmental region; it bears
anteriorly a submarginal, intersegmental ridge, or antecosta (fig. 2 A,
Ac), marked externally by the antecostal suture (figs. 1, 2 A, acs),
and it terminates anterior to this ridge and its suture in a narrow
lip, or precosta (Pc).
The ventral sclerotization of the segment may take the same form
as the dorsal sclerotization, as in the abdomen of most insects, where
the definitive terga and sterna duplicate each other in structure
(fig. 3). The functional intersegmental rings of the body in such
cases are the posterior, non-sclerotized areas of the primary segments,
and the definitive segmentation is clearly a secondary one. The sternal
sclerotization, however, may preserve a more primitive condition, as
in some of the chilopods (figs. 8, 15) and in the thorax of certain
insects, where the primary sternal and intersternal plates remain inde-
pendent (figs. 1, 2 A, Stn, Ist).
In the membranous areas of the lateral, or pleural, walls of the seg-
ment are implanted the bases of the segmental appendages. In most
arthropods the basis of the appendage (coxopodite) is preserved as
an integral limb segment. In the body segments of the chilopods, the
thoracic segments of insects, and the ambulatory segments of decapod
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 5
crustaceans, however, it appears that the limb basis has become sub-
divided into a coxa and a subcoxa, and that the latter has been incor-
porated into the wall of the body segment (fig. 1, Sc), where it either
forms a “ pleuron”’ supporting the free part of the limb, or also the
base of the wing, or it becomes degenerate and reduced to small
sclerites having little significance or function.
Mp Pe
\ \
‘
Stn
/ i ras
Ph: , DMek B ZPh ZPh
Fic. 2—Diagram showing the relation of the longitudinal muscles to the tergal
and sternal sclerites of the body.
A, three successive segments in which the terga include the primary inter-
segmental regions bearing the intersegmental ridges (Ac, Ac), but in which
the primary sternites (Stn) and intersternites (/sf) are distinct. B, the tergal
region of the thorax in an insect in which the precostae (A, Pc) are enlarged to
form postnotal plates (PN).
Ac, antecosta; ac, antecostal suture; DMcl, dorsal longitudinal muscles; Ist,
intersternites; JT, first abdominal tergum; L, positions of leg bases; Mb, secon-
dary intersegmental membrane; Pc, precosta; 1Ph, 2Ph, 3Ph, the three thoracic
phragmata; PN, postnotal plates; Stn, primary sternite; V Mcl, ventral longi-
tudinal muscles.
THE THORACIC TERGA
The dorsal plates of the insect thorax never retain in all three seg-
ments the simple structure of the definitive abdominal terga, and in
the Pterygota the mesothoracic and metathoracic terga are modified
in various ways correlated with the development of the wings.
The prothoracic tergum (fig. 4, 71) always lacks an antecosta, and
the principal longitudinal muscles (DMcl) that extend forward from
6 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
the anterior phragma of the mesotergum (zPh) run continuously
through the prothorax and the neck (Cv) to be inserted on the post-
occipital ridge of the head (PoR). This ridge, as the writer has else-
where contended (1928), is evidently the intersegmental fold between
the first and second maxillary segments. The neck, therefore, must be
derived from the posterior part of the second maxillary, or labial,
segment and from the anterior part of the prothorax, there being no
satisfactory evidence of the existence of a separate neck segment.
If so, the first postcephalic intersegmental line, or that between the
labial and prothoracic segments (fig. 4, 1/sg), must be contained in the
membranized cervical region, where the protergal costa is lost. By the
suppression of the primary intersegmental line between the head and
Fic. 3.—Diagram of the body segmentation of an insect, and the primitive
relation of the longitudinal muscles to the definitive segmental plates of the body
and to the head; showing the reversed overlapping of the sterna between the
thoracic and abdominal regions.
Cv, cervix; DMcl, dorsal longitudinal muscles; H, head; JS, first abdominal
sternum; JT, first abdominal tergum; Ppt, periproct, or terminal segment;
S1, S2, Ss, thoracic sterna; 71, Tz, Ts, thoracic terga; V Mcl, ventral longitudinal
muscles; XJ, eleventh abdominal segment.
the thorax, giving continuity to the muscle fibers of two segments,
the head acquires a much greater freedom of motion than it could
have if it were attached to the body by an ordinary intersegmental
membranous ring.
The loss of the protergal antecosta deprives the prothorax of the
possibility of being a wing-moving segment, and there is nothing to
suggest that the prothorax ever possessed movable organs of flight.
The reduction of the primitive gnathal region of the body and its con-
densation into the head capsule, accompanying the transfer of the
gnathal appendages to the head, shifted the center of gravity pos-
teriorly in the insect’s body, and the paranotal lobes of the second and
third thoracic segments were developed into movable wings, leaving
the prothorax as a free segment between the head and the pterothorax.
The most conspicuous modifications of the thoracic terga occur in
the mesothorax and the metathorax of winged insects, where clearly
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 7
they are correlated with the part.the terga of these segments play in
the mechanism of flight. In the Apterygota the corresponding terga
are simple plates showing none of the special characters of the wing-
bearing plates of pterygote insects.
The first important tergal modifications connected with the develop-
ment of the paranotal lobes into movable organs of flight pertain to
the ridges upon which the dorsal muscles of the mesothorax and
metathorax have their attachments. These ridges, which are the ante-
costa of the mesotergum, the antecosta of the metatergum, and the
antecosta of the first abdominal tergum, bear each a pair of apodemal
plates, varying in size, that project into the body cavity to give in-
creased surfaces of attachment for the greatly enlarged dorsal muscles
(fig. 2 B, DMcl) which have become depressors of the wings. The
antecostal apodemes, primarily intersegmental, are the thoracic phrag-
mata (1Ph, 2Ph, 3Ph).
The lengthwise pull of the dorsal muscles on the phragmata de-
mands sclerotic continuity in the dorsum, since the function of these
muscles as depressors of the wings depends on their ability to produce
a dorsal curvature in the terga on the relaxation of the antagonistic
tergo-sternal muscles. To insure action by the dorsal muscles the
intersegmental membranes between the mesotergum and metatergum
and between the latter and the first abdominal tergum must be prac-
tically eliminated, and their suppression has been accomplished either
by a fusion of the succeeding terga, or by a forward extension of the
precostal lips of the terga into the territory of the membranes. In the
second case, the precostae become postnotal plates (fig. 2 B, PNo,
PN;), often of large size, lying behind the true tergal plates of the
mesothorax and metathorax (T., T;), where they appear to be parts
of these segments, to which, in fact, they do belong since they lie
anterior to the antecostal sutures (ac, ac) which are the primary
intersegmental lines.
In those insects in which the fore wings are the principal organs of
flight, the second thoracic phragma becomes partially or wholly de-
tached from the metatergum, and both the phragma and the postnotal
plate establish a close association with the mesotergum, while the ex-
tremities of the postnotum commonly unite for security with the pos-
terior dorsal angles of the mesothoracic epimera. In those insects in
which the hind wings have taken on the chief function of flight, the
middle phragma always remains attached to the metatergum, and the
precosta is not enlarged. The third phragma may preserve its connec-
tion with the first abdominal tergum, as it does in the Orthoptera
(fig. 25, PN;), but in most cases it becomes more or less separated
8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
from the abdomen and, together with the precosta, becomes trans-
ferred to the metathorax, where the precosta forms a distinct post-
notal plate united laterally with the epimera. Thus it is usually found
that the segment which assumes the leading role in the flight mechan-
ism is provided with a phragma at both its anterior and its posterior
‘end.
Since the tergal plates of the mesothorax and metathorax are the
intermediary elements in the wing mechanism between the dorsal mus-
Fees /
/ Vi
SA, 5; 19S D2. 208
Fic. 4.—Diagram of the typical relation of the head and the prothorax in
: pterygote insects.
Cu, cervix; Icv, 2cv, first and second lateral cervicai sclerites; DMcl, dorsal
longitudinal muscles; H, head; r/sg, 2Isg, first and second primary interseg-
mental lines; rPh, first thoracic phragma; Poc, postocciput; PoR, postoccipital
ridge; PT, posterior arm of tentorium; Si, S2, thoracic sterna; SA, sternal
epopryees Ss, spinisternites; 71, T2, thoracic terga; V Mcl, ventral longitudinal
muscles.
cles of the segments and the bases of the wings themselves, it is clear
that a proper execution of their function depends upon the ability of
each to respond to the muscle tension at its ends with a dorsal curva-
ture reaching its maximum at the transverse line between the wing
bases. For this reason, as Weber (1924, 1925) has pointed out, the
terga of the wing segments are provided with internal ridges so ar-
ranged that the force of the muscles will not merely deflect the an-
terior and posterior parts of the plates, but will be distributed gradu-
ally toward the middle from each extremity, and thus produce an even
dorsal flexion with its apex between the fulcra of the wings.
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 9
The posterior gradient of an alar tergum usually has the form of a
V-shaped ridge with the apex directed forward and the arms diverging
toward the posterior lateral angles of the tergum (fig. 5 B, VR). The
anterior gradient is less commonly developed than the posterior one,
but, when present, it generally consists of two ridges, the parapsidal
ridges (fig. 5 B, PaR), converging from the anterior margin of the
tergum toward the middle, where they usually terminate without meet-
ing. In some insects the anterior part of the tergum is strengthened
by a transverse prescutal ridge (PR). In addition to these more gen-
eral endoskeletal structures of the tergum, there may be present also
Pe Psc Ph ps acs fe Fse. Ph PR Ac
_Aw mm» XM
E-ANE
Em
—PNP
AxC ae
‘
/
AxC Rd f& B Rd
Fic. 5.—Structure of a wing-bearing tergum, not including a postnotum,
diagrammatic.
A, dorsal. B, ventral. Ac, antecosta; acs, antecostal suture; ANP, anterior
notal wing process; AxC, axillary cord; Aw, prealar process of tergum; Em,
lateral emargination of tergum; Pak, parapsidal ridge; pas, parapsidal suture;
Pc, precosta; Ph, phragma; PNP, posterior notal wing process; PR, prescutal
ridge; ps, prescutal suture; Psc, prescutum; Fd, posterior fold, or reduplication,
of tergum; Scl, scutellum; Sct, scutum; VR, V-shaped ridge; vs, suture of the
V-ridge, or scuto-scutellar suture; WV”, base of wing.
a variety of accessory ridges, or even lines of flexibility in the tergal
cuticula ; but all such features are highly variable in different groups
of insects, and homologies can be traced between them only within
limited groups.
On the outer surface of the tergum the positions of the endoskeletal
ridges are marked by the lines, or “sutures,” of their inflection
(fig. 5 A, ps, pas, vs). The tergal areas defined in this manner by the
more constant of the inner structures can be identified as homologous
in different insects, and some of them have been given distinctive
names used in descriptive works (fig. 5 A, Psc, Sct, Scl). It is quite
impossible, however, to follow the lesser modifications consistently
through the various orders of winged insects, and attempts to do so
have only led to confusion. In any case, it must be recognized that
IO SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
the external “ divisions” of the wing-bearing terga have no signifi-
cance in themselves; they are merely incidental to the formation of
the internal ridges by cuticular inflections, the ridges being the true
functional structures adapting the tergum to its part in the flight
mechanism.
THE THORACIC PLEURA
The lateral walls of arthropod segments, or the areas along the sides
of the body between the terga and the sterna, when dorsal and ventral
plates are present, may properly be designated the pleural regions.
The pleural areas of the segments are primarily membranous, and
Fic. 6—Diagram of the theoretical elementary musculature of the segmental
appendages.
a-b, primitive dorsoventral axis of the appendage; J, tergal promotor muscle;
J, tergal remotor; K, sternal promotor; L, sternal remotor; 7, tergum;
S, sternum.
within them are implanted the bases of the limbs (fig. 6). In some
arthropods, as in many of the Arachnida, each limb basis occupies
almost the entire space between the tergum and the sternum, and may
be articulated to one or the other of these plates, or to both of them.
In most cases, however, a membranous area partially or entirely sur-
rounds the limb base. In this area there are sometimes developed true
pleural sclerites, as in the chilopod family Geophilidae, where there
is a series of lateral plates of the body wall lying between the terga
and the leg bases (fig. 8 A, pl), or in the larvae of some insects where
similar plates occur on the sides of the abdomen. In many arthropods,
however, there are plates in the definitive lateral walls of certain seg-
ments that appear to have been derived from the bases of the appen-
dages. While such sclerotizations are, therefore, not true pleural prod-
ucts, they are generally termed pleurites, and those of each side of each
segment constitute collectively the so-called pleuron of the segment.
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS Il
It is claimed by Becker (1923, 1924) that the pleurites, the coxae,
and the trochanters in the Chilopoda are formed, during the develop-
ment of the individual, from numerous schlerotizations in the lateral
-Eppd
es
WS
ox
Fic. 7.—Maxillipeds and pleuron of a decapod crustacean, Macrobrachium
jamarcensts.
A, first maxilliped, left, posterior surface. B, second maxilliped, right, an-
terior. C, third maxilliped, left, posterior. D, left pleuron, or inner wall of bran-
chial chamber.
Brn, branchia, gill; Cx, coxa; Cxrpd, coxopodite; Endp, endopodite; Eppd,
epipodite; Expd, exopodite; Scx, subcoxa; rTr, first trochanter. :
walls of the body segments, which unite to form the definitive leg
bases and the pleural sclerites of the adult. Though the apparent
facts in the development of the chilopods may be as Becker describes
them, it is difficult to see how they can be interpreted literally as repre-
12 “ SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
senting the phylogenetic origin of the definitive pleural plates and the
leg bases. It would seem more probable that they are ontogenetic phe-
nomena only, and that Becker’s observations really show simply that
the pleurites and the bases of the legs have a common origin.
In the decapod crustaceans the inner walls of the gill chambers,
which are covered externally by lateral folds of the carapace, are
formed of large cuticular plates bearing the gills (fig. 7 D). Each
plate, or pleuron, shows subdivisions (Sca4-Scxg) corresponding with
the body segments of the ambulatory legs, and each subdivision bears
a gill (Brn4-Brng). In the second maxilliped (B) the homologue of
the gill is borne on an epipodite (Eppd) which is distinctly carried by
/ N
A cx. Stn Ist B
Fic. 8—A body segment of Strigamia bothriopus (Chilopoda, Geophilidae).
A, lateral view, with leg removed beyond the coxa. B, ventral view, including
bases of legs. Cx, coxa; /st, intersternite; pl, pleurites between the tergum and
the subcoxa; Scr, subcoxa; Sp, spiracle; Stn, primary sternite; 7, tergum; Tr,
trochanter.
the basal segment of the appendage, or coxopodite (Cxpd). In the
third maxilliped (C), however, the gill arises from a subcoxal part of
the limb basis (Sc). In the ambulatory region (D) the gills on the
pleuron are successively more and more removed from the coxae. It
thus becomes evident that the pleural wall of the branchial chamber in
the decapod crustaceans has been formed from dorsal extensions of
the subcoxal parts of the leg bases, and that the coxae have acquired
special articulations with the subcoxae. In the majority of crustaceans
the leg base is an undivided coxopodite.
In the Chilopoda there is a definitely circumscribed subcoxal area
about the base of each leg, which may be continuously sclerotized, as
in Strigamia (fig. 8 A, Scx), or which may contain one or more
sclerites, as in Lithobius (fig. 9), Scolopendra, or Scutigera (fig. 10).
The coxa is usually articulated to a sclerotized part of the subcoxa
dorsally (fig. 10 A, c), or ventrally (figs. 9, 15, d) ; but since the axis
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 13
Fic. 9—A body segment and leg of Lithobius (Chilopoda), left side.
c, dorsal articular point of coxa; Cx, coxa; d, ventral articulation of coxa with
subcoxa; Fm, femur; Ptar, pretarsus; S, sternum; Scr, subcoxa; Sp, spiracle ;
T, tergum; Tar, tarsus; Tb, tibia; 1Tr, first trochanter ; 2Tr, second trochanter.
Fic. 10.—The leg base of Scutigera forceps (Chilopoda).
A, external view of base of a left leg and part of segment. B, internal view
of base of right leg, showing muscles.
c, dorsal subcoxo-coxal articulation; cpl, supra-coxal plate of subcoxa; Cx,
coxa; Fu, furca; J, tergal promotor muscles; J, tergal remotor muscle; K,
sternal promotor; L, sternal remotor; M, subcoxo-coxal muscle; N, sternal
adductor of coxa; N’, furcal adductor of coxa; Scx, subcoxa; Stn, segmental
sternite; 7, tergum; 17,Y, first trochanter.
I4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
of movement in the coxa is always between its dorsal and ventral
angles, we may presume that the chilopod coxa has had both a dorsal
and a ventral articulation with the subcoxa (fig. 1, c, d), though one
or the other, or both of the articulations (fig. 8) may lose the struc-
ture of definite articulating surfaces.
In the geophilid Strigamia bothriopus (fig. 8) the subcoxal area of
the pleuron has the form of a complete basal limb segment (A, Scr),
though its ventral margin is expanded and united with the sternum
(B), and the coxa turns upon it by an obliquely vertical axis. In
Scolopendra, Lithobius (fig. 9), and Scutigera (fig. 10) the subcoxal
area is mostly membranous, but it contains one or more well-sclerotized
plates.
The tergal muscles of the leg bases in the Chilopoda are inserted not
on the coxae but on the subcoxae. In Scutigera the tergo-subcoxal
muscles are strongly developed, those of each leg comprising a pair
of anterior (promotor) muscles (fig. 10 B, J) inserted upon the dorsal
plate (coxopleure, cpl) of the subcoxal region, and a single large
posterior (remotor) muscle (J) inserted on the posterior dorsal mar-
gin of the subcoxal region.
The structure and musculature of the subcoxal region in the
Chilopoda can leave little doubt that this area is the true base of the
leg, which has become flattened into the lateral body wall, where, in
most forms, its sclerotization has been more or less broken up and
reduced. The sternal muscles of the leg base in the chilopods have
gone over to the ventral rim of the coxa (fig. 10 B). They include
an anterior (ventral promotor) muscle (A) and a posterior (ventral
remotor) muscle (L). In Scutigera (fig. 10) the first of these mus-
cles arises in the anterior lateral angle of the sternum, but in Scolopen-
dra the corresponding muscle arises mesally on the anterior half of
the sternum. The fibers of the posterior muscle in Scutigera (fig. 10 B,
L) are mostly continuous from one coxa to the other, but a small
anterior group on each side arises on the sternum at the base of the
ligamentous endosternal furca (Fu). In Scutigera the coxa has no
ventral articulation with the subcoxa or the sternum, but in those
chilopods in which a ventral subcoxo-coxal articulation is present, the
anterior and posterior ventral muscles (K, L) must act as promotors
and remotors.
The base of the coxa in the Chilopoda is provided also with median
dorsal and ventral muscle. The dorsal median muscle in Scutigera
(fig. 10 B, M) consists of a flat band of short fibers arising on the
dorsal plate of the subcoxa (cpl), and is inserted on the rim of the
coxa just behind the dorsal articulation with the subcoxa. This muscle
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 15
appears to be a remotor of the coxa in Scutigera, though it may have
an abductor function also. The ventral median muscles of Scutigera
comprise two bundles of fibers, one arising medially on the sternum
(fig. 10 B, NV’), the other (.N’) arising on the lateral arm of the en-
dosternal furca. These muscles are coxal adductors since the coxa
has no fixed ventral articulation in Scutigera. The ventral coxal mus-~
cles are covered dorsally by large bands of trochanteral muscles that
take their origin on the sternum.
Fic. 11.—Diagram of the basal musculature of an insect leg.
Bs, basisternum; Cx, coxa; J, tergal promotor muscle; J, tergal remotor;
K, sternal promotor (anterior rotator); k, furcal suture; L, sternal remotor
(posterior rotator) ; M, abductor of the coxa; M’, M”, abductors of the coxa
that become the basalar and subalar muscles in the wing-bearing segments of
adult insects; N, adductor of the coxa; SA, sternal apophysis; Scr, subcoxa;
SI, sternellum; T, tergum; Tn, trochantin.
In the insects the sclerotic areas of the subcoxae of the legs evi-
dently become the pleural plates of the thoracic segments. The tergal
promotor muscle of the leg base (fig. 11, /) retains its connection with
the subcoxa in the more generalized pterygote insects, being inserted
on the trochantinal sclerite of the subcoxa (Tn) except when the
trochantin is lost, the muscle then having its insertion on the anterior
angle of the coxal base. The remotor muscle (/), which may be repre-
sented by several fiber bundles, is always inserted on the coxa or on
coxal apodemes. The anterior and posterior sternal muscles (K, L)
arise on the sterna or the sternal apophysis, or on the spinasteraum.
16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
The median coxal muscles are represented in insects by both dorsal
(MM) and ventral (NV) fibers. In the wing-bearing segments of ptery-
gote insects the first comprise three distinct groups of abductor fibers
(M, M’', M”) arising dorsally on the pleuron, and inserted ventrally
on the coxa both anterior and posterior to the dorsal articulation of
the latter (c). The second and third muscles of this group (/’, M”)
Fic. 12.—Thoracic pleural sclerites of Apterygota.
A, lateral view of left side of mesothorax of Acerentomon doderoit (from
Berlese, 1910). B, thorax, base of head, and base of abdomen of Jsotoma sp.
(from Ewing, 1928). C, left mesothoracic leg turned forward, and lateral re-
gion of mesothorax of Acerentulus barberi (from Ewing, 1928).
c, dorsal articulation of coxa; Cx, coxa; d, ventral articulation of coxa; Fm,
femur; H, head; JT, JIT, first and second abdominal terga; Ptar, pretarsus;
S, sternum; Scr, subcoxa; 7:, Tz, Ts, thoracic terga; Tar, tarsus; Tb, tibia;
Tr, trochanter.
become wing muscles in the adult by the partial or complete detach-
ment of the epipleural areas on which they arise to form the basalar
and subalar plates of the wing base. The ventral median muscle of the
coxa (N) is present in insects that lack a ventral coxal articulation ;
it arises on the sternal apophysis and functions as a coxal adductor.
In the Apterygota the subcoxa becomes rudimentary. In most of
the Protura its sclerotization is reduced to two slender plates arched
concentrically over the base of the coxa (fig. 12 A, C, Sc+), as shown
by Berlese (1910) and by Ewing (1928), though Prell (1913) has
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS Wf
described the pleural sclerotizations of Eosentomon germanicum as
consisting of a number of small sclerites, which, however, fall into
two concentric series. (See Snodgrass, 1927, fig. 8.) In the Collem-
bola, as shown by Ewing (1928), the subcoxal sclerotizations of the
mesothorax and metathorax (fig. 12 B, Sc2, Scv3) consist in each
segment of two slender, supra-coxal arches ; the subcoxal sclerotiza-
tion of the prothorax (Scx,) is a single plate with an incomplete sub-
division. In the Thysanura the subcoxal pleurites likewise take the
form of two arches over the coxal base, or they become reduced to a
single sclerite. The coxal and subcoxal musculature of the Aptery-
gota has been but little studied.
In the thoracic segments of the Pterygota the subcoxae evidently
become the sclerotized parts of the lateral segmental walls known as
the pleura (cf. figs. 1 and 13). The ventral rim of each subcoxa, lying
between the coxa and the sternum (fig. 13 A), may be reduced to a
membranous fold, though in rare cases it contains a large plate (fig. 17,
Ls2, Ls;), and in others a rudimentary sclerite (fig. 16 A, Ls). In the
majority of insects, as has been shown by Weber (1928, 1928 a), the
ventral arc of the subcoxa has apparently fused with the primary
sternite to form a laterosternite of the definitive sternum (figs. 13 B,
TOO). Hes).
The coxa of insects is universally hinged to the subcoxa by a dorsal
articulation (fig. I, c) ; it may also have either an anterior articulation
with the trochantinal piece of the subcoxa (fig. 13 B, e), or a ventral
articulation (A, d) with the ventral rim of the subcoxa or with the
subcoxal laterosternite. The trochantinal articulation of the coxa is
peculiar to certain insects and is, therefore, probably a secondary one.
The ventral articulation, however, so frequently recurs both in the
Chilopoda (fig. 15, d) and in the more generalized insects (fig. 16 A,
B, d) that there can be little doubt that the primitive axis of the
subcoxo-coxal hinge was vertical or approximately so. The writer,
therefore, would retract the opinion, expressed in a former study of
the thorax (1927, pp. 34-36), that the primitive axis of the coxal
movement was a horizontal one between anterior and posterior articu-
lations with the eutrochantinal arch of the subcoxa. The ventral ar-
ticulation of the coxa is highly variable in insects ; it is always absent
in the more generalized Pterygota that have a well-developed trochan-
tin. In the members of the higher orders lacking a trochantin it is
commonly present, but it is to be suspected in such cases that the
articulation is a secondary one developed between the coxa and the
sternum.
2
18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
The usual trochantin of the pterygote pleuron (fig. 13 B, Tm) is
clearly a remnant of a more extensive, primitive, supra-coxal scleroti-
zation (fig. 1, Eutn) carrying the dorsal articulation of the coxa (c),
which Crampton (1914) has named the eutrochantin, and which is
best preserved in the ventral arch of the apterygote pleuron (fig.
12). The eutrochantin is retained as an independent sclerite also
in the prothorax of the Plecoptera, but in all other Pterygota (fig. 13,
Pe acs Ble Pe acs
“a /
SS
/ / : \ , / \ aml
EX Ist Stn $a lst B Ps TES itera Ss¢Ist)
Fic. 13.—Diagrams suggesting the development of the pterygote pleuron from
the subcoxa of the leg basis. (Compare with Fig. 1.)
A, subcoxal sclerotization (Sc) united ventrally with edge of primary ster-
nite (Stn), its dorsal extremity prolonged upward as a wing support (B, WP),
posterior part of entrochantin (fig. 1, Eutn) fused with eupleural arch (fig. 1,
Eupl) of subcoxa.
B, a fundamental structural condition of pleuro-sternal region of a wing-
bearing segment: the area of subcoxa differentiated into an episternum (Eps),
an epimeron (Epm), a precoxal bridge (Acx), a postcoxal bridge (Pcx), a
laterosternite (Ls), and a trochantin (Tn); the definitive sternum includes the
primary segmental sternite (Stn), the following intersegmental intersternite, or
spinasternite (S's), and a subcoxal laterosternite (Ls) on each side; the ventral
coxal articulation (A, d) is lost, and coxa has a secondary anterior articulation
with trochantin (¢).
A, B) its dorsal and posterior parts unite or fuse with the upper arch
of the pleuron (eupleuron), and only its anterior part remains as a
free sclerite (fig. 13 B, Tm) carrying the anterior coxal articula-
tion (e).
The elaborate pterygote pleuron has evidently been developed to
give support to the paranotal lobes, or to the wings evolved from the
latter. It is therefore strengthened by an internal ridge formed from
a linear inflection of its wall, the pleural suture (fig. 13 A, PIS), ex-
tending from the dorsal articulation of the coxa (c) upward to the
wing support (B, WP). The area lying posterior to the pleural suture
is the epimeron (B, Epm), that situated anterior to it and dorsal to
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 19
the trochantin is the episternum (Eps). The sclerotized parts of the
subcoxa lying anterior and posterior to the coxa are the precoxal and
postcoxal bridges (Acx, Pcx), or precoxalia and postcoxalia. The
ventral wall, or infra-coxal arc, of the subcoxa, as already noted, prob-
ably unites in many cases with the edge of the primary sternum (Stn)
to form a laterosternal element (Ls) in the definitive sternum, though
it may be reduced to a separate sclerite or form a membranous fold
between the coxa and the sternum.
All parts of the pleuron are subject to innumerable secondary modi-
fications taking the form of sutures that subdivide the primary areas,
or of membranous lines and spaces that break them up into separate
sclerites. Such modifications are not necessarily homologous between
different orders, but within an order or group of orders they may give
valuable evidence of the evolution and interrelationships of the fami-
lies and genera. An example of this is given by Shepard (1930) in a
study of the secondary pleural sutures of Lepidoptera.
In the wing-bearing segments of the Pterygota two large pleuro-
coxal muscles (fig. 11, M’, MM”) become important muscles of the
wings. These muscles evidently are derived from the abductor system
of the coxa. In nymphal Orthoptera (fig. 27 C) the anterior muscle
(M’) has its origin on the dorsal part of the episternum, the posterior
muscle (.4”) on the dorsal edge of the epimeron. In adult insects,
however, the areas upon which these muscles are attached become par-
tially or entirely separated from the pleuron and intimately associated
with the base of the wing, the first lying before the pleural wing
process, the second behind it. In this way the muscles come to func-
tion as wing muscles, though each retains its ventral attachment on the
coxa.
The epipleurites (“‘ paraptera’’), or sclerites detached from the
pleuron in connection with the coxo-alar muscles, include one or two
episternal sclerites, or basalares, and usually a single epimeral sclerite,
or subalare. The subalare is always completely detached from the
epimeron in adult insects (fig. 14 A, B, Sa). A basalar plate, how-
ever, is not always present as a distinct sclerite ; it frequently occurs as
but an imperfectly separated lobe of the upper end of the episternum
(fig. 14, Ba), and its area is sometimes marked only by the insertion
of the anterior coxo-alar muscle (/’). Even when the basalare is dis-
tinct from the episternum, it is generally hinged to the upper edge of
the latter in such a manner that it is deflected by the contraction of its
muscle. Frequently there is present a second basalar muscle (figs. 27 C,
E) having its origin on the pleuron or on the sternum.
20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
The theory of the origin of the principal pleural sclerites of the
Chilopoda and Hexapoda from subcoxal segments of the legs has
much in its favor. There is little evidence, however, that a subcoxa
is a primary segment in the general arthropod appendage. The limb
bases of the Arachnida, Xiphosura, and most Crustacea are the primi-
tive coxae (coxopodites), for there can be no doubt of the identity
of the coxo-trochanteral articulation in all arthropods. The writer
Fic. 14.—The mesopleuron and base of the middle leg of a scorpion fly,
Panorpa consuetudims.
A, external view. B, internal view, showing muscles. a, accessory sclerite
of basalar lobe; Ba, basalar lobe of episternum; bcs, basicostal suture of coxa;
Bex, basicoxite; Cx, coxa; Epm, epimeron; Eps, episternum; MM’, basalar
muscle; M”, subalar muscle; Mer, meron; O, levator muscle of trochanter ;
PIS, pleural suture; Sa, subalar sclerite; W”7P, pleural wing process.
clearly was mistaken in suggesting in a former paper (1927, p. 33)
that the large basal leg segments of the ticks (Ixodoidea) are sub-
coxal ; and he now believes that the segmentation of the arachnid limb
can be given an interpretation different from that proposed by Ewing
(1928), who would make the basal segment in most cases a subcoxa.
In the decapod crustaceans the inner walls of the gill chambers, as has
already been pointed out, are evidently expansions of the subcoxal
regions of the bases of the ambulatory legs, to which the coxae of the
latter have become articulated ; but there is no evidence of the pres-
ence of subcoxal segments in the limbs of the more generalized Crus-
tacea. In the myriapods and insects, moreover, as the writer has else-
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 21
where shown (1928), there are no true subcoxal segments in the
mouth part appendages. From the evidence at hand, therefore, it ap-
pears most in accord with the known facts to conclude that the subcoxa,
wherever it occurs as a basal leg segment, has been produced by a
secondary subdivision in the primitwe limb basis, or coxopodite.
THE THORACIC STERNA
Sternal plates are by no means so constant a feature in the scleroti-
zation of arthropod segments as are the tergal plates. They may be
present or absent within the same major group, and, where present,
Fic. 15.—Sternites and leg bases of two consecutive segments of Lithobius
(Chilopoda).
Cx, coxa; d, ventral subcoxo-coxal articulation; Jst, intersternite; Scx, sub-
coxa; Stn, segmental sternite; r7r, first trochanter; 277, second trochanter.
they are often highly variable both in form and extent of development
between closely related groups and in the different body regions of
almost any species.
In adult insects the sternal mechanism of the thorax differs in three
important respects from that of the abdomen, and the functional dif-
ferences in the two body regions are reflected as three distinctive struc-
tural features in the sternal parts.
The first distinction to be noted in the sternal structure, as between
the thorax and the abdomen, pertains to the segmental relations of the
intersegmental sternites. In the abdomen of adult insects the inter-
segmental sclerotizations of both the dorsum and the venter are con-
tinuous with the segmental sclerotizations following, and the trans-
verse inflections in the cuticula of the primary intersegmental regions,
22 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
on which the fibers of the longitudinal muscles are attached, become
the antecostae of the definitive terga and sterna (fig. 3). In the
thorax, on the other hand, the ventral intersegmental sclerotizations
either remain as small, free sclerites (fig. 1, 7st), or they unite with
the posterior parts of the segmental plates preceding. The interseg-
mental sternites, or intersternites, of the thoracic region are the
spinasternites (fig. 18 D, Ss), so-called because each usually bears a
small median apodemal process, the spina (fig. 4). A spinaster-
nite occurs typically between the prothorax and the mesothorax, and
between the mesothorax and the metathorax; there is never a free
spinasternite following the metasternum because the corresponding
intersegmental element goes with the first abdominal sternum to form
the antecosta of the latter, except where it is lost as a result of the
degeneration of the first abdominal sternum. The first spinasternite
is more commonly persistent than the second which is usually fused
into the posterior part of the mesosternum, where it may become
entirely obliterated.
The second structural difference between the thoracic and abdomi-
nal sterna accompanies the difference in the relations of the interseg-
mental sclerites to the segmental plates, but is not necessarily corre-
lated with it. It consists of a reversal in the overlapping of the sterna.
The successive abdominal sterna overlap regularly in a posterior direc-
tion, as do the terga of both the abdomen and the thorax (fig. 3). The
sterna of the thorax, on the other hand, overlap anteriorly. The meta-
thoracic sternum, therefore, stands as a dividing plate between the
anteriorly overlapping sterna of the thorax and the posteriorly over-
lapping sterna of the abdomen (fig. 3, S3).
This reversal in the overlapping of the sternal plates as between the
thorax and the abdomen is probably the oldest structural differentia-
tion between the two regions of the body, for it is well shown in some
of the Apterygota, particularly in Japyx, and is exhibited by all
pterygote insects in which the thoracic sterna remain free from each
other. It was probably, therefore, established when the thorax was
first set apart as the locomotor center of the body, and has nothing
to do with the development of the wings. Just what advantage accrues
to the thoracic mechanism from the reversed relations of its sternal
plates is not clear, but presumably it gives a better device for the
movement of the legs or for the movement of the successive seg-
mental plates on each other.
The third distinction between the thorax and the abdomen occurs
in adult pterygote insects, and pertains to the attachments of the ven-
tral muscles. We have assumed that the primitive attachments of the
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 23
fibers of the longitudinal muscles are on the intersegmental folds or
on intersegmental sclerotizations (figs. 2, 3). The dorsal muscles
throughout the length of the body, and the ventral muscles of the
abdomen are thus attached, except where the anterior ends of the fibers
may have migrated to the segmental regions of the definitive terga
and sterna. In the thorax of adult pterygote insects, however, most
of the sternal muscles are stretched between paired apodemal processes
of the segmental sternites (fig. 4, S41, SA2), except that the anterior-
most fibers are inserted anteriorly on the head, while the posteriormost
fibers extend into the abdomen. Only a few slender median muscles
retain a connection with the intersternites (7Ss, 2Ss). The paired
Eps
pee
Fic. 16.—Sternal structure of ephemerid and odonate nymphs.
A, ventral surface of mesothorax, metathorax, and first abdominal segment
of an ephemerid nymph, showing ventral articulations of coxae (d) with sub-
coxal, laterosternal sclerites (Ls).
B, ventral surface of neck and prothorax of an aeschnid nymph, showing direct
articulation of coxae (d) with laterosternal parts (Ls) of the definitive sternum.
apophyses of the thoracic sterna are the so-called furcal arms, which
in the higher orders are united upon a common median base and here
constitute a true furca.
The anterior ends of the ventral muscle fibers, as we have noted, are
attached on the back of the head. In orthopteroid insects the attach-
ment is with the posterior arms of the tentorium (fig. 4, PT), but this
condition is clearly a secondary one since the posterior tentorial arms
are tergal apodemes. In many adult insects, and in most holometabo-
lous larvae, the anterior ventral muscles are inserted on the posterior
part of the head wall. Morison (1927) enumerates three pairs of
prothoracic sternal muscles in the honeybee, all of which are attached
anteriorly on the lateral occipital regions of the head. In the caterpil-
lars the corresponding muscles are inserted on apodemes of the ventral
margin of the foramen magnum. In all such cases the insertion points
of the ventral head muscles must have acquired their present positions
24. SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
by a migration from the true sternal region of the last head segment,
which is the membranous floor of the neck behind the base of the
labium.
The ventral muscles of the thorax retain apparently the primitive
condition in the larvae of most holometabolous insects. In the cater-
pillar, for example, the principal longitudinal ventral muscles consist
of two wide bands of fibers lying to each side of the ventral nerve
cord, extending through the entire length of the body, and attached
regularly on the intersegmental folds as are the dorsal longitudinals.
External to the dorsal and ventral intersegmental muscles of the cater-
pillar there is an intricate complex of small muscles disposed in all
directions against the wall of each segment.
In certain larval forms, as in some Coleoptera, the attachment of the
ventral body muscles shows a condition intermediate between the
usual larval condition and that of the adult. In the larva of Dytiscus,
for example, as shown by Speyer (1922), though most of the ventral
thoracic muscles are intersegmental, being attached either to processes
of the intersegmental folds or to transverse ligaments arising from
the folds, some of the fibers extend between segmental furcal apo-
physes, which are present on each primary sternal region of the
thorax. The ventral muscle bands of the thorax are continued into
the abdomen, some of the fibers of the first abdominal segment being
attached anteriorly on the intersegmental fold behind the metathorax,
others on the furcal arms of the metasternum. In the adult of Dytiscus
(Bauer, 1910) all the ventral muscles of the thorax are interfurcal in
their attachments, and none extends from the thorax into the abdomen.
Ventral muscles from the thorax into the abdomen are absent in the
adult stage of many pterygote insects (fig. 35), though they may be
present in the larval or the nymphal stages. In the nymph of Psylla
mali, according to Weber (1929), two bundles of fibers diverge from
the base of the metafurca to the anterior edge of the second abdominal
sternum, but these muscles, Weber says, are lost in the adult.
In some insects, however, the ventral thoracico-abdominal muscles
are present in the adult stage. They are well developed in the cock-
roach (Blatta orientalis), comprising here three pairs, the first arising
on the second spina, the second on a ligamentous bridge between the
bases of the metasternal apophyses, the third on the apophyses, all of
which are inserted posteriorly on the anterior margin of the second
abdominal sternum. The fibers arising on the metapophyses form the
anterior ends of the ventral longitudinal muscle bands of the abdomen.
In Gryllus, Voss (1905) describes a median pair of muscles arising
on the metafurca which branch posteriorly to the third, fourth, and
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 25
fifth abdominal sterna, and two lateral groups on each side which go
to the parasternal plates of the second abdominal segment. In the
Cicadidae ventral muscles extend from the metathorax to the second
abdominal sternum. In the Tenthredinidae, according to Weber
(1927), a pair of muscles extends from the metafurcal arms to the
second sternum of the abdomen, and in the honeybee Morison (1927)
describes two corresponding pairs of muscles going from the meta-
furca to the anterior margin of the second abdominal segment. Inas-
much as these muscles, which represent the ventral muscles of the first
abdominal segment, have no connection with the first abdominal ster-
num, it is evident, as Morison points out, that their insertions, normally
on the intersegmental anterior edge of the first abdominal sternum,
have been secondarily transferred to the furcal apophyses of the meta-
thoracic sternum. In the Ephemerida, however, Durken (1907)
records the presence of a pair of muscles attached anteriorly on the
bases of the metasternal apophyses and posteriorly on the anterior
margin of the first abdominal sternum. These muscles would appear
to correspond with the furco-spinal muscles, which are present in the
prothorax of the grasshopper (fig. 35, 61).
Even a brief review of the comparative musculature of larval and
adult holometabolous insects thus shows that there takes place during
metamorphosis a rearrangement in the attachments of the ventral
muscles of the thorax, and, in some cases, of those of the first abdomi-
nal segment, as a result of which most of the persisting fibers lose
their intersegmental connections and acquire segmental attachments on
the furcal apophyses of the thoracic sternal plates.
The larval condition of intersegmental muscle attachments is clearly
a more primitive one than that of the adult. The adults of insects with
incomplete metamorphosis resemble those of holometabolous forms in
having the principal ventral muscles attached on the furcal arms.
Therefore, we must suspect that the latter condition is one secondarily
acquired in all pterygote insects, and that it has come about during
the evolution of the thorax as a specialized locomotor region of the
body. Since the transposition of the ventral muscles takes place in
the prothorax as well as in the other two thoracic segments, we cannot
attribute its inception to the development of the wings. As yet, how-
ever, we may draw only tentative conclusions concerning the evolution
of the ventral musculature of the thorax, since our knowledge of the
nymphal muscles in hemimetabolous insects and of both the larval and
adult muscles in the more generalized holometabolous forms is very
incomplete ; but the facts known point strongly to the transformation
suggested above.
26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
An alteration in the attachment of the ventral muscles similar to
that which evidently has taken place in insects may be observed also
in the Chilopoda, here between members of different families. In the
Geophilidae and in Lithobius the ventral longitudinal muscles consist
principally of two flat, widely-separated bands of fibers lying close
against the body wall and inserted on intersegmental sclerotizations.
Fic. 17.—Ventral view of the base of the prothorax, the mesothorax, the
metathorax, and the base of the abdomen of the large South American embiid,
Cylindrachaeta spegazzinii.
d, ventral articulation of coxa with subcoxal laterosternite (Ls); k, furcal
suture; J, secondary suture of mesosternum; Ls, laterosternite; 1S.:, anterior
plate of mesosternum; 252, posterior plate of mesosternum (furcasternite) ;
Ti, ventral fold of protergum.
In Scolopendra and Scutigera, on the other hand, both the longitudinal
ventral muscles and many other muscles of each segment are attached
on two ligamentous supports that arise from the posterior parts of the
segmental sternites. In Scolopendra each ligament has a separate
origin on the sternum ; in Scutigera the two ligaments in each segment
arise from a common base, forming thus a furca-like structure
(fig. 10 B, Fu) suggesting that of the higher pterygote insects.
It is scarcely possible that there is any genetic relation between the
furcal apophyses of insects and the muscle-supporting structures of
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 27
Scolopendra and Scutigera, but it is clear that the muscle attachments
have been altered by a transposition which is of a parallel nature in the
two cases.
In the Chilopoda the sternal plates are uniformly developed through-
out the length of the body. In Strigamia, Lithobius, and Scolopendra
there is a series of alternating segmental sternites (figs. 8 B, 15, Stn)
and intersegmental intersternites (Ist). The intersternites are lacking
in Scutigera; they are highly developed in the geophilid Strigamua
bothriopus, where their lateral ends extend upward on the sides of the
body between the subcoxae (fig. 8 A), but they are small and incon-
spicuous in Lithobius (fig. 15). In both the geophilid and lithobiid
as we have seen, the longitudinal ventral muscle bands have their at-
tachments on the intersternites.
The presence of alternating sternites and intersternites in the chilo-
pods might suggest that this condition was the primitive one in in-
sects, and that the intersternites (spinasternites) have remained free
in the thoracic region or have united with the preceding sterna, while
they have fused with the segmental sternites following in the abdomen.
In the odonate larva shown at B of figure 16 there is a long interseg-
mental sclerite (2/st) between the posternum (S,) and the mesoster-
num suggestive of the intersternites of the Chilopoda, and the fold
(Ist) in the ventral side of the neck (Cv), which bears the cervical
sclerites laterally (cv), appears to be likewise an intersternite between
the labial segment and the prothorax. In the Acrididae the spinaster-
nite between the prothorax and mesothorax (fig. 21, Ss) is a well-
formed plate attached to the prosternum (.S) ; that between the meso-
thorax and metathorax is indistinguishably fused into the posterior
border of the mesosternum, though the spina persists (fig. 31, 2S pm).
In many insects the first spinasternite is a free sclerite, and in the Blat-
tidae both the first and the second are distinct plates (fig. 19 A,
Ps 25 Ss).
The definitive thoracic sterna of most insects are undoubtedly com-
posite structures. The first and second intersternites are usually con-
tained in the posterior parts of the prosternum and mesosternum,
respectively, or at least are closely associated with them, though the
first frequently retains its independence. The ventral arcs of the sub-
coxae contribute laterosternal elements in many insects. The evident
union of the ventral rim of the subcoxa with the sternum has been
noted in the Hemiptera (Heymons, 1899, Snodgrass, 1927), but
Weber (1928, 1928a) has given ample reasons for believing that this
fusion of subcoxal elements with the primary sternum has taken place
in the majority of insects. The frequent ventral articulation of the
28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
coxae with the lateral margins of the sternum in generalized insects
(figs. 12 C, 16 B, d) is further evidence that the sternum in such cases
includes the infracoxal parts of the subcoxae, especially since it is
found that, where distinct subcoxal laterosternal sclerites exist (figs.
16 A, 17, Ls), the coxae articulate with these sclerites (d).
It is difficult to find in the insects a good example of a simple pri-
mary sternal plate, comparable with the sterna of the Chilopoda (fig.
15), that does not contain either the following intersternite or sub-
coxal laterosternal elements, or both. In the mesothorax and meta-
thorax of the ephemerid nymph shown in figure 16 A, the sterna
(Sz, S;) may contain the intersternites, but the two small sclerites
in each segment (Ls, Ls) that articulate between the sternum and the
coxa on each side appear to be the only remnants of subcoxal lateroster-
nites. In the large embiid Cylindrachaeta (fig. 17) laterosternal plates
(Ls2, Ls3) likewise are distinct, though the intersternites are clearly
united with the primary sternites. In the prothorax of the aeschnid
larva shown in figure 16 B the intersternite (2/s¢) is independent of
the sternum, but the laterosternites (Ls) are fused into the lateral
sternal margins.
These several forms make it clear that the definitive thoracic ster-
num of insects is typically a compound plate. It consists of a primary
sternite (fig. 18 A, B, Stn), to which may be added the succeeding
intersternite (Ist), which becomes the spinasternum (C, D, Ss), and
a pair of latcrosternites (D, Ls, Ls) derived from the ventral arcs of
the adjoining subcoxae (B, C, Scr).
The possession of paired apophyses, or furcal arms, is character-
istic of the thoracic sterna of all pterygote insects. The apophyses
arise from the sternal plates between the bases of the legs, and their
outer ends are usually closely attached, either by fusion or by short
muscle fibers, to the inner ends of the corresponding pleural apophyses.
Weber (1928, 1928a) advances the view that the sternal apophyses
are primarily invaginations formed on the line of union between the
primary sternites and the subcoxal laterosternites. In some insects,
however, in which there are laterosternal plates not united with the
sterna (figs. 16 A, 17), the origins of the sternal apophyses (s@) are
still well within the sternal margins; and in an aeschnid nymph (fig.
16 B) the apodemal invaginations (sa, sa) are removed from the
apparent margins of the laterosternite sections (Ls) of the definitive
sternum. From evidence of this nature the writer would regard the
sternal apophyses as invaginations in the primary sternal plate itself
(fig. 18 B, Stn), though there is much in favor of Weber’s view. The
mesosternum of wingless females of the black aphis, Aphis fabae,
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 29
Weber (1928a) says, presents a case in which there can be no doubt
that the furca arises at a point between the basisternite, the furcaster-
nite, and the subcoxal laterosternite.
In the higher insects the sternal apophyses approach each other in
each segment and unite upon a common basis produced by a median
Fic. 18.—Diagrams suggesting the theoretical evolution of a thoracic sternum.
A, primitive condition in which the ventral sclerotization consists of alter-
nating segmental sternites (Stn) and intersegmental intersternites (Jst); the
leg basis (LB) is an undivided coxopodite.
B, primary sternite marked by the pits (sa) of a pair of internal apophyses,
intersternite (/st) by the pit (spn) of a median process, or spina; leg basis
(A, LB) subsegmented into subcoxa (Scx) and coxa (Cx), articulated dorsally
and ventrally (d).
C, area of primary sternite (A, B, Stn) divided into basisternum (Bs) and
sternellum (S7) by a furcal suture (k) forming an internal furcal ridge between
bases of sternal apophyses; the following intersternite has become a spinaster-
num (Ss) by union with segmental sternite; subcoxa (Scx) united ventrally
with sternum.
D, typical definitive sternum, composed of primary sternite (A, B, Stn), a pair
of subcoxal laterosternites (Ls, Ls), and the spinasternite (Ss); area of pri-
mary sternite divided into basisternum (Bs) and sternellum (S7) by the furcal
suture (k), and with a narrow presternum (Ps) set off by a secondary presternal
suture (7).
inflection of the sternal wall. In this way is formed the typical, forked
endosternal structure known as the furca, the evolution of which has
been portrayed by Weber (1928). The part of the sternum bearing
the furca lies between the coxae, and is usually much narrowed by
comparison with the region of the sternum anterior to it.
The definitive sternal plate, whether it includes subcoxal lateroster-
nal elements and the following intersternite, or does not, is commonly
subdivided into an anterior and a posterior region. Fundamentally the
30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
dividing line is a transverse inflection, or “ suture,” passing through
the bases of the sternal apophyses (fig. 18 C, k). The inflection is
usually strongly sclerotized, forming an internal ridge evidently de-
signed to brace the sternum and to support the apophyses (figs. 21 B,
31, k) ; it sometimes remains weak, however, and establishes a line
of flexibility in the sternum. In either case the sternum is demarked
by the furcal ridge and its suture (k) into a prefurcal area (fig. 18 C,
Bs) and into a postfurcal area (ST).
The anterior region of the sternum has been variously named
sternum, in a restricted sense, antesternum, mesosternum, basisternwm,
eusternum, and sternannum; the second has been called sternellum,
poststernum, metasternum, and furcasternum. There are objections
to all but one of these terms. “‘Antesternum”’ and “ poststernum ”’
(Amans, 1885) are applicable in some cases, but there is often a pre-
sternal piece before the “‘ antesternum,”’ and very commonly the inter-
segmental spina-bearing plate forms an actual posternal element of the
definitive sternum behind the “ poststernum.” “ Mesosternum ” and
‘““metasternum ”’ (Berlese, 1909) violate the priority of the segmental
prefixes. “ Basisternum ” and “ furcasternum ”’ (Crampton, Ig0g) are
misleading because the part designated by the first is not basal, and that
bearing the second name does not always carry the furcal apophyses.
“ Eusternum ”’ (Snodgrass, 1910) implies that the part so named is
the “true ”’ sternum, which it is not. “ Sternannum”’ (Mac Gillivray,
1923) has no grammatical standing, so far as the writer can find.
“ Sternellum ” (MacLeay, 1830) alone can be given a clean bill. Of
the terms applied to the prefurcal area, however, “ basisternum ”’ ap-
pears to be the least objectionable. In the present paper, therefore,
the writer adopts the following terms for the principal divisions of the
definitive sternum (fig. 18 D): presternum (Ps), basisternum (Bs),
sternellum (S71), and spinasternum (Ss). The first three are secon-
dary subdivisions of the primary segmental sternum; the fourth is the
intersegmental intersternite. To the primary sternal region there may
be added on each side a subcoxal laterosternite (Ls).
The parts of the definitive thoracic sternum as described here fit
exactly with the definitions of the sternal sclerites given by Weber
(1928, pp. 250, 251), with the understanding that the term “ sternel-
lum” is substituted for ‘‘ furcasternum,” and that the poststernite is
the intersegmental spinasternite. This idea of the sternal composition
differs from Crampton’s (1909) conception in that the fundamental
transverse dividing line of the sternum is assumed to be the furcal
suture (k) between the bases of the sternal apophyses, and not a divi-
‘
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 31
sion anterior to the apophyses. A prefurcal division sometimes does
occur (fig. 17, 7), but it is clearly of a secondary nature and is vari-
ously produced.
The furcal suture is subject to much diversity in form, being some-
times produced forward and branched laterally, or curved posteriorly,
thus giving a variety of structure to its apodemal ridge, and often
obscuring the primary line of the sternal division.
The form and size of the sternal plates are frequently altered by a
variation in the extent of the ventral sclerotization in the different
thoracic segments. In the Blattidae it is evident that a partial de-
sclerotization of the sternal cuticula has produced the unusual shapes
and relationships of the sternal sclerites of the thorax (fig. 19). The
prosternum most nearly preserves the typical form (B). It comprises
two median plates (A, Bs,, Sl,) separated by a transverse fold (k)
across the sternal region, from which arise the prosternal apophyses
(SA,). A comparison with the assumed generalized structure of a
thoracic sternum (B) will easily suggest that the transverse fold is the
furcal suture (#), and that the pattern of the prosternal plates (C)
has been produced by suppression of sclerotization in the lateral fields
of the sternal area. In the mesothorax of Blatta (A) the ventral sclero-
tization is reduced to a pair of basisternal plates (Bs.), and a Y-shaped
furca-bearing sclerite (S/.), the two separated by an ample mem-
branous area. In the latter are remnants of the sternal fold (k) from
which arise the sternal apophyses (SA.) at the ends of the sternellar
arms. The diagram D shows more clearly the relation of the mesoster-
nal structure in the roach to the fundamental sternal structure (B),
and again suggests that the peculiar features of the thoracic sterna of
the roach are results merely of a reduction in the extent of the sclero-
tized areas. The metasternum of Blatta (A) is essentially the same as
the mesosternum, but the sternal fold appears to be suppressed and the
apophyses (S'A;) arise from the sternellum (S/,).
In some insects a thoracic sternum may be divided into two parts by
a suture that is quite independent of the furcal suture. A clear case
of this is seen in the thorax of the large embiid Cylindrachaeta (fig.
17), where a suture (/) cuts the mesosternum into an anterior plate
(1S) and a posterior plate (2S,). The second plate is marked by the
usual furcal suture (k) and bears the furcal arms; it is a true furca-
sternite. The metasternum has the usual structure, though the sternel-
lum is reduced to a narrow band behind the furcal suture (2).
Most entomologists have believed that the sternum of a thoracic
segment of an insect is “ composed of ” two principal plates, and the
32 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
pattern of the sternal sclerites in the cockroach (fig. 19) has had much
to do with establishing this idea, for students have not recognized
that the separated plates here are products of sclerotic degeneration,
and that the fundamental structure, as shown best in the prothorax,
is the same as in insects with undivided thoracic sterna. While the
Blattidae undoubtedly retain some relatively generalized structural
sa A
Fic. 19.—The thoracic sterna of a cockroach, Blatta orientalis.
A, general view of the three thoracic sterna and their paired apophyses. B,
diagram of typical structure of a sternum. C, diagram of prosternum of Blatta.
D, diagram of mesosternum of Blatta.
Bs, basisternum; JS, first abdominal sternum; k, furcal suture; SA, sternal
apophysis; sa, external pit of sternal apophysis; S/, sternellum; spn, external
pit of spina; Ss, spinasternum; S, segmental sternum.
characters, they are in many respects highly specialized insects adapted
to a particular kind of habitat, though to one almost universally dis-
tributed. The flattening of the body has been accompanied by a struc-
tural alteration in most of the under parts of the thorax, and there is
every reason to believe that the sterna, covered as they are by the bases
of the legs, have become largely membranized to allow of an inflection
of their posterior parts. We should be on very unsafe ground, there-
fore, if we take the fragmented condition of the sternal sclerotization
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 33
in the thorax of the roach as representative of the primitive structure
of the thoracic sterna in insects.
A search for a generalized thoracic sternum among all groups of
insects, the writer believes, would reveal nowhere the thing sought for,
because it does not exist. A study of the arthropods as a whole, how-
ever, suggests that the original area or areas of sclerotization in the
ventral region of each segment spread into a continuous plate between
the leg bases. The thoracic sterna of insects have been variously modi-
fied by the development of apodemal braces where rigidity is demanded,
and by secondary divisions or by reductions in the areas of sclerotiza-
tion where flexibility is important. This theory recommends itself by
the fact that it permits all kinds of specific structures and sclerotic
patterns to arise, and does not assume that homologies must exist
where none can be established.
If. THE THORACIC SKELETON OF DISSOSTEIRA
The Carolina locust, Dissosteira carolina, is here used as the subject
for a special study of the thorax and its mechanisms because it is an
insect sufficiently large for work on internal structure and is readily
obtained, and because its muscular system is simple and comparatively
easy to dissect. The thorax of the Acrididae is by no means general-
ized, but for this reason it offers a good test for the application of gen-
eral principles to the solution of specific problems. The structural
features of the thorax in the locust, however, are those common to
all insects, and in the musculature there is almost no addition of special
muscles such as are found in most of the higher insect orders and to
some extent in the other orthopteran families.
The thorax of the jumping Orthoptera is so distinctly divided into a
prothorax and a pterothorax that it is scarcely to be regarded as a unit
in the organization of the body. The box-like structure of the com-
bined mesothorax and metathorax, the oblique slant of the pleurites
of these segments, and the firm connection of the first abdominal seg-
ment with the metathorax are characters evidently correlated with the
development of the hind legs as saltatorial organs.
THE CERVICAL SCLERITES
The grasshopper ordinarily keeps its head retracted against the pro-
thorax, in which position the insect appears to have no neck, for the
ample neck membrane (fig. 20 B, Cv), as well as the back part of the
head, is thus concealed within the projecting anterior rim of the
pronotum.
3
34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
The neck skeleton of Dissosteira consists of two pairs of small
cervical sclerites situated ventro-laterally in the membranous walls of
the neck (fig. 20 B, rcv, 2cv). The two sclerites of each pair in the
grasshopper are closely hinged to each other, and form a bridge on
each side between the head and the prothorax. The first sclerite (rcv)
is an irregularly triangular plate articulating with the occipital con-
dyle (fig. 32, g) of the posterior rim of the head, situated just above
the base of the posterior tentorial arm (A, PT). Immediately behind
its articulation this plate bears externally two small lobes that are
conspicuous by their covering of short hairs. The second cervical
sclerite (fig. 20, B, 2cv) is a slender bar articulating posteriorly with
the anterior margin of the prothoracic episternum (Eps) just within
the overlapping edge of the protergum. The two cervical sclerites of
each pair are movably hinged to each other at an angle directed ven-
trally. They are mostly concealed when the head is in the usual re-
tracted position, but they form a small prominence of the neck pro-
jecting just behind the base of the maxilla. The probable function and
mechanism of the cervical sclerites will be described in connection with
the account of the muscles inserted upon them.
The lateral, muscle-bearing cervical sclerites are probably homolo-
gous structures in all insects in which they occur. Dorsal and ventral
neck plates are presnt in some insects, but they are variable in size and
arrangement and are probably secondary sclerotizations of the neck
membrane.
THE PROTHORAX
The prothorax of the grasshopper is a highly individualized segment
of the body, for, though its posterior dorsal and lateral parts widely
overlap the anterior part of the mesothorax, it is separated from the
latter by an ample intersegmental membrane (fig. 20 B, Wb).
The external parts of the prothorax comprise tergal, pleural, and
sternal sclerites. The principal plate is the tergum, a large bonnet-like
piece that covers the back and most of the sides of the segment (fig.
20 A, T). Only a corner of each pleuron shows externally: this is
the small triangular lobe lying anteriorly between the base of the leg
and the lower margin of the tergum (fig. 20 A, Eps). The rest of the
pleuron is deeply invaginated within the lateral wall of the tergum
(B, Eps). The prosternum consists of two sclerites in the ventral wall
of the segment between the bases of the first legs (fig. 21 A, S, Ss),
the anterior one connected by the precoxal bridges (dcx) with the
pleura. The prothoracic legs appear to be inserted between the ster-
num and the lower edges of the tergum, but the lateral connections of
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 35
the legs are really with the inflected pleura covered by the tergal ex-
tensions. The procoxal cavities are “ open” behind, that is, there are
no postcoxal sclerotizations. Lying before the coxa of each leg in the
articular membrane of the leg base is a small trochantinal sclerite
(fic. 20 A, Tn).
The protergun.—The tergum of the prothorax, besides covering
the back and sides of its own segment, projects posteriorly over the
dorsum of the mesothorax in a wide, triangular lobe which fits between
the bases of the folded front wings. The top of the tergal bonnet
Fic. 20.—The prothoracic tergum and pleuron of Dissosteira carolina.
A, outer view of left side. B, inner view of right side, showing episternum
invaginated within the tergum.
a, posterior edge of anterior fold of tergum; b, anterior edge of posterior fold
(B, Rd) of tergum; c, d, e, the external vertical grooves or sutures of the ter-
gum (A) forming the internal tergal ridges (B); f, g, h, 1, horizontal sutures
and their ridges connecting the vertical sutures and ridges.
(fig. 20 A, T) is cut by a deep transverse notch somewhat before the
middle, and the part before the notch is compressed into a median
ridge.
Each lateral area of the protergum is marked by a number of grooves
forming a definite pattern, and by two non-impressed lines. The first
non-impressed line (fig. 20 A, a) lies near the anterior border of the
tergum and runs parallel with it; the second (b) extends downward
in a sinuous course just posterior to the dorsal tergal notch. These
two lines mark the limits of the inner folds of the anterior and pos-
terior inflections of the tergal wall (B, a, b). The grooves of the tergal
surface (A, c-7) lie in the space between the two non-impressed lines.
They have no significance in themselves, but it is important to note
36 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
their positions because they form ridges on the inner surface (B)
which have definite relations to the muscle attachments of the proter-
gum. The first (c) is a short curved line on the upper lateral part of
the tergum ; the second (d) is a longer line extending from the back
almost to the ventral margin of the tergum; the third (e) begins at
the dorsal notch and reaches ventrally just before the second non-
impressed line (b) to the middle of the side. Connecting the three
vertical grooves are four short longitudinal grooves, one (f) lying be-
tween the first and second vertical grooves (c, d), the other three
(g, h, 7) between the second and third vertical grooves (d, e).
A study of the inner surface of the tergum (fig. 20 B) will show
the endoskeletal ridges (c-i) formed by the external grooves. There
AWiaakis
Fic. 21.—The prothoracic sternum and pleura of Dissosteira.
A, ventral view of sternum and lower edges of pleura, showing the spina-
sternum (Ss) united with the segmental sternal plate (S), the latter continuous
with the episterna (Eps) by the antecoxal bridges (Acx). B, dorsal surface of
ee ee bases of sternal apophyses (SA, SA) united by a furcal
ridge (k).
will also be noted the anterior and posterior inflections of the tergal
walls. From the margin of the first inflection (a) the neck membrane
(Cv) is reflected forward, and from the margin (0) of the posterior
fold or reduplication (Rd) the intersegmental membrane (Mb) is
reflected posteriorly to the mesothorax. The first spiracle (Sp2) is
located in this membrane.
The propleura—-We have already noted that each pleuron of the
prothorax appears externally only as a small plate projecting from
beneath the edge of the tergum anterior to the base of the leg (fig. 20 A,
Eps). It is to be seen on the internal surface of either half of the seg-
ment (B), however, that the pleural piece exposed externally is merely
the lower anterior corner of a much larger triangular sclerite (B,
Eps) extended upward within the lateral tergal wall by a deep inflec-
tion of the tergo-pleural membrane. The posterior margin of the
sclerite is turned inward, forming the pleural ridge, which gives off
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 37
the pleural arm (P/A) and ends ventrally in the articular process of
the coxa (CxP). Behind the lower part of the pleural ridge is a small
epimeral piece (Epm) fused with the lower border of the tergum and
concealed just within the edge of the latter. The anterior ventral angle
of the episternum is continuous through the precoxal bridge (Ac)
with the anterior lateral angle of the prosternum (fig. 21 A).
The prosternum.—The sternum of the prothorax in the grasshopper
consists of two distinct plates (fig. 21 A, S, Ss) separated by a trans-
verse suture. The anterior plate (S) is the larger and the more
strongly sclerotized. It is continuous laterally with the precoxal bridges
(Acx) from the episterna (Eps). The definitive sternal plate of the
prothorax, then, evidently includes laterosternal elements derived from
the pleura (subcoxae), but the true pleuro-sternal limits are entirely
obliterated. The anterior rim of the sternum is set off as a narrow
presternal strip (Ps) by a submarginal external suture (A, 7) and
a corresponding internal ridge (B, 7). Posteriorly the first sternal
plate is marked by a deep transverse groove which forms a strong
ridge on its inner surface (fig. 21 B, k) between the bases of the
sternal apophyses (B, SA, SA), which latter are indicated externally
by a pair of pits (A, sa, sa). The first prosternal plate in the grass-
hopper, therefore, is divided in the primitive fashion (fig. 18 C) into
a basisternal and a sternellar region by the suture of a ridge connecting
the bases of the apophyses. The prosternal apophyses are simple arms
(fig. 21 B, SA) diverging dorsally and laterally. Their distal ends are
solidly united with the corresponding pleural apophyses.
The second prosternal plate (fig. 21, Ss) is a spinasternite, and is,
therefore, the intersternite between the prothorax and the mesothorax
which has become closely associated with the primary sternite of the
prothorax. It is mostly overlapped by the anterior margin of the
mesosternum. The spinasternite of Dissosteira is triangular in shape,
and is marked by a deep median impression (fig. 21 A) which forms
the spina internally (B, Spi).
THE PTEROTHORAX
The united mesothorax and metathorax of the grasshopper consti-
tute a unit in the body mechanism. The pleural and sternal walls of
the two segments are solidly united, forming a trough-like structure
perforated only by the openings of the coxal cavities. The leaping
force of the hind legs is thus applied to a rigid middle section of the
body, which also bears the wings. The dorsum of this body section is
covered by the mesothoracic and metathoracic terga, but these plates
38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
are freely attached to the upper pleural margins of the pterothoracic
trough by the ample membranes of the wing bases, and they are mov-
ably joined to each other. As we shall later see, the wing mechanism
demands at least a limited freedom of movement in the wing-bearing
terga.
In the grasshopper the back plates of the pterothorax (figs. 22, 24)
differ somewhat in shape and in details of form and proportion, but
the two have the same essential structure. They are relatively small,
and when the insect is at rest they are hidden beneath the folded wings.
The pleurites are defined externally by distinct grooves (fig. 26)
forming strong ridges internally (fig. 28), which slant posteriorly and
downward in a manner to suggest that they serve thus to brace the
pleural walls against the projectile force of the hind legs. The sterna
of the wing-bearing segments are wide plates fused laterally with the
pleura before the leg bases (fig. 30).
The mesotergum.—tThe tergum of the mesothorax (fig. 22 A) isa
rectangular plate ending posteriorly in a distinct, transverse fold (Rd),
the extremities of which are continued into the posterior thickened
margins, or axillary cords (Axc), of the wing bases. Close to the
anterior margin of the tergum is a deep groove (acs). This is the
antecostal suture, or primary intersegmental inflection which forms
the antecosta of the internal surface of the definitive tergum (B, Ac).
The antecosta bears laterally two wide, flat apodemal plates (1Ph)
projecting into the cavity of the thorax (fig. 25), which are the first
pair of thoracic phragmata.
On the external surface of the mesotergum, two sutures (fig. 22 A,
ps, ps) diverge laterally and posteriorly from the antecostal suture
(acs). They form internally a pair of strong ridges (B, PR) extend-
ing to the bases of the anterior wing processes (ANP). The large, ir-
regular, triangular regions (A, Psc, Psc) forming the anterior lateral
angles of the tergum, set off by the divergent sutures (ps, ps), con-
stitute the prescutal areas of the tergum. In the metathorax the
prescutal sutures do not meet the antecostal suture, and the lateral
prescutal lobes are continuous by a narrow median bridge behind the
antecostal suture (fig. 24). In some other Acrididae, as in Melanoplus,
the continuity of the prescutal area is more pronounced. In other
orthopteran families the prescutum is narrow, but in the Blattidae and
Gryllidae there is a suggestion of its separation from the scutal area.
In any case, however, the prescutum of the Orthoptera must be re-
garded as a secondary differentiation of the anterior part of the tergum.
Its lateral parts become most sharply defined in the mesotergum of
the Acrididae by the strong development of the prescutal ridges (fig.
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—-SNODGRASS 39
22 B, PR) that brace the anterior wing processes. Upon the irregular
surfaces of the prescutal lobes are attached the tergo-sternal muscles
which are the principal elevators of the wings.
A prescutum similar to that of the Orthoptera occurs also in certain
other insects, though very likely it may be an independent differen-
tiation formed as an adaptation to similar demands. In many insects
of the higher orders, however, such as the Hemiptera, Diptera, and
Hymenoptera, a prescutal area of quite a different nature is set off in
the anterior median part of the tergum by the development of two
Fic. 22——The mesothoracic tergum of Dissosteira.
A, dorsal surface. B, ventral surface. Ac, antecosta; acs, antecostal suture;
ANP, anterior notal wing process; Aw, prealar arm of tergum; 14x, first
axillary; 44x, fourth axillary; AxC, axillary cord; Em, lateral emargination
of tergum; Mb, secondary intersegmental membranes; n, lobe of prescutum
articulating with base of subcostal wing vein; 0, lobe of scutum articulating
with posterior part of first axillary; Pc, precosta; 1Ph, first phragma; ps, pre-
scutal suture; Psc, Psc, lateral prescutal areas; Rd, posterior reduplication of
tergum; Scl, scutellum; Sct, principal part of scutum; sct, sct, posterior lateral
subdivisions of scutum; s, s, secondary ridges of tergum; tg, tegular rudiment;
VR, remnant of V-ridge of tergum.
lateral ridges, the parapsidal gradients (fig. 5 B, PaR), which extend
a varying distance posteriorly from the anterior tergal margin, and
usually converge. These ridges and their sutures apparently lie in the
scutal region of the tergum, for there is sometimes present a narrow
transverse prescutal band anterior to their bases. Parapsidal ridges are
absent in the Orthoptera.
The area of the mesotergum of Dissosteira posterior to the ante-
costal and prescutal sutures is differentiated topographically into a
large anterior scutal region (fig. 22 A, Sct), a median, posterior, tri-
angular scutellar region (Sc/), two small, lateral, posterior scutal re-
gions (sct, sct), and a posteriormost, deflected marginal region (Rd).
The structure here presented is quite different in appearance from
that of a typical wing-bearing tergum (fig. 5 A) in which the surface
40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
is divided into scutal and scutellar areas (Sct, Scl) by the suture (vs)
of an internal V-shaped ridge (B, VR), the arms of which are con-
vergent forward from the posterior lateral angles of the tergum.
In the Acrididae the V-shaped endotergal ridge (fig. 22 B, VR)
is almost obliterated, and the tergum is braced by two secondary ridges,
one on each side (s), that converge posteriorly from the posterior
lateral margins of the scutal area and intercept the arms of the rudi-
mentary V-ridge (VR). The altered structure of the acridid tergum
Rd
Fic. 23.—Diagram of the structure of a wing-bearing tergum of Acrididae.
The prescutal suture (ps) is either continuous, or suppressed medially; the
usual V-ridge and its suture (vs) are partially suppressed and subordinated to a
secondary ridge of similar shape but having its arms (s, s) convergent posteriorly.
may be expressed diagrammatically as in figure 23, where the sup-
pressed suture (vs) of the obsolete V-ridge is crossed by the dominant
suture (s) of a secondary ridge of similar shape but having its arms
convergent posteriorly. Thus the scutum consists of a principal an-
terior scutal area (Sct) and of two small postero-lateral scutal areas
(sct, sct) ; and the scutellum is divided into a median scutellar area
(Scl) and two lateral scutellar areas (scl, scl), including the posterior
fold of the tergum (Fd). The evolution of this condition can be traced
in other Orthoptera from the primary structure which occurs in the
Blattidae. A similar modification has taken place in the mesotergum
of Hemiptera and Coleoptera, producing the triangular elevated shield
of the scutellum that lies between the bases of the folded wings.
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 41
It is clear that the external “ divisions’”’ of the wing-bearing ter-
gum are incidental to the development of the internal ridges, which
are adaptations to the part the tergum plays in the mechanism for
moving the wings. The old idea that the tergum is “ composed of ”
sclerites gave undue emphasis to surface features. Though a study
of the latter may have a value for descriptive purposes, the student
must look to the internal characters for a true understanding of the
skeleton of insects.
There is no postscutellar plate in the mesothorax of the grasshopper.
The posterior deflected margin of the scutellum ends in a narrow inter-
segmental membrane (fig. 25, 2b) uniting the mesotergum with
the anterior margin of the precosta of the metatergum. The tergum
of the mesothorax of the grasshopper, therefore, is a typical dorsal
plate of a secondary segment, comprising the primary segmental
sclerotization and the preceding primary intersegmental sclerotization
of the back. In the latter the primary intersegmental fold is marked
by the antecosta (fig. 25, dc) and the antecostal suture (acs).
The lateral margins of the mesotergum are very irregular (fig. 22).
The wings are extended from the tergal edges between the middle of
the prescutal borders and the posterior reduplication of the scutellum.
Anterior to the wing bases the anterior angles of the tergum are ex-
tended as short prealar arms (fig. 22 A, Aw) to which are articulated
the dorsal processes of the first basalar plates (fig. 26, Ba). The lat-
eral margin of the prescutal area forms posteriorly a small process
bearing a socket-like surface (m) in which the base of the subcostal
wing vein turns when the wing is flexed or extended. Posterior to this
process the anterior angle of the scutum is produced to form the large
anterior notal wing processes (ANP), which support the neck of the
first axillary sclerite of the wing base (14). The inner edge of the
first axillary bridges the lateral emargination of the tergum (Em)
and articulates with a marginal lobe (0) behind the latter. There is
no posterior notal wing process in the mesotergum of the grasshopper ;
the fourth axillary (44), which is itself probably a detached piece
of the tergal margin, articulates with the edge of the scutellum.
The metatergum.—tThe tergal plate of the metathorax (fig. 24) is
somewhat longer than that of the mesothorax, since it must support
the wider bases of the hind wings; but in many respects it is more
weakly developed than the mesotergum, there being extensive non-
sclerotized areas in the posterior part of the scutal region.
The precostal rim of the metatergum (fig. 24, Pc) is narrow, except
medially where it forms a conspicuous lip before the deeply inflected
42 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
antecostal suture (acs). The prescutal ridges and their sutures (ps,
ps) are much weaker than those of the mesotergum, but they are not
confluent medially with the antecostal ridge and suture, and the lateral
prescutal triangles (Psc, Psc) appear to be continuous across the back
in a narrow, weakly sclerotized area deflected into the antecostal
suture (acs).
Fic. 24.—The metathoracic tergum and postnotal plate of Dissosteira.
acs, antecostal suture; ANP, anterior notal wing process; 1A», first axillary,
4Ax, fourth axillary; AxC, axillary cord; Em, lateral emargination of tergum;
n, prescutal lobe to which base of subcostal wing vein is attached; 0, tergal lobe
to which posterior end of first axillary articulates; p, tergal arm supporting
anal veins of wing (see fig. 47 B); Pc, precosta; 2Ph, second phragma: PNs,
postnotum; ps, prescutal suture; Psc, prescutum; Fd, posterior fold of tergum
(see fig. 25, Rds); s, s, sutures of secondary tergal ridges; Sci, scutellum; Sct,
principal part of scutum; sct, sct, subdivisions of scutum.
The surface features of the scutal and scutellar regions of the meta-
tergum have even less relation to the generalized structure of a wing-
bearing tergum than do those of the mesotergum, because the tergal
ridges (fig. 23) are here almost completely suppressed, and the exter-
nal characters are the result of secondary inflections which produce a
topographical pattern quite independent of the primary divisions of
the tergum (fig. 5). Most of the scutal region (fig. 24, Sct) and the
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 43
median triangle of the scutellar region (Sc/) are confluent in a large
shield-shaped area that forms the principal part of the tergal plate.
The depressed posterior lateral parts of the scutum (sct, sct) are cut
transversely by the faintly-marked sutures (s, s) of the posteriorly
convergent ridges, which are obsolete in the metatergum, though
strongly developed on the mesotergum (fig. 22 B, s). The posterior
marginal area of the metatergum (fig. 24, Rd), which is a part of the
true scutellar region, is sharply inflected (fig. 25, Ad;) and is continu-
ous with the greatly extended precosta of the first abdominal segment,
which constitutes a postnotal plate of the metathorax (figs. 24, 25,
PNs3).
Ee Set, Sel, Rd Setz Sls
acs
Fic. 25.—Median longitudinal section of mesotergum, metatergum, and meta-
thoracic postnotal plate of Dissosteira, showing the phragmatal lobes of the
right side.
Ac, antecosta; acs, antecostal suture; JT, first abdominal tergum; 1M/b, 2Mb,
secondary intersegmental membranes; z7Ph, 2Ph, 3Ph, first, second, and third
phragmata; PNs, postnotal plate of metathorax, or greatly enlarged precosta of
first abdominal tergum; Fd, posterior reduplication of tergum; Sct, scutum;
Scl, scutellum.
The lateral margins of the metatergum present the same features as
do those of the mesotergum. The posterior angle of each prescutal
area projects as a small marginal process (fig. 24, n) which is con-
nected with the head of the subcostal wing vein by a ligament-like
thickening of the basal wing membrane; it does not articulate with the
vein as in the mesothorax. The anterior notal wing process (ANP)
is a flat lobe of the scutum, to which the first axillary (14x) is closely
hinged. Behind the wing process is a deep emargination (Em) of the
scutellum, posterior to which is a second lobe (0) articulating with
the posterior end of the first axillary. The slender fourth axillary
(4A) articulates with the extreme posterior angle of the lateral scu-
tellar area (sct). Each extremity of the posterior marginal fold of
the tergum (Id) gives off into the anal membrane of the wing a long
arm (p) that supports the anal veins (fig. 47 B).
44 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
The pterothoracic pleura——The pleurites of each side of the meso-
thorax and metathorax are firmly united to form continuous lateral
walls of the pterothoracic region (fig. 26) in which the episterna and
epimera (Eps, Epm) are distinct plates separated by oblique grooves
sloping from above downward and posteriorly. The first principal
groove is the pleural suture of the mesothorax (P/S:;), the second is
the intersegmental line, the third is the pleural suture of the meta-
thorax (P/S;). Each pleural suture terminates above in a large wing
Aer
ZS ee EAS
/ Za
/ \ \ /
s, Tn Cx, 0 Se asAaue a
Fic. 26—The pterothoracic pleura of Dissosteira.
Ba, basalar sclerites; Cx, coxa; Epm, epimeron; Eps, episternum; Fs, base
of hind femur; 7, prepectal suture; P/S, pleural suture; PN3:, lateral arm of
metathoracic postnotum; Ppct, prepectus; r. pleuro-sternal suture; S, sternum;
Sa, subalare; Sp, mesothoracic spiracle; Ss, metathoracic spiracle; Tn, tro-
chantin; Tr, trochanter; 1” P, pleural wing process.
process (WP:, WP:), and below in the pleural articulation of the
coxa. The episternum of each segment (Eps:, Eps) is united ven-
trally before the coxal cavity with the edge of the sternum, the line of
union (7) in the adult insect being obsolete in the mesothorax, but
distinct in the metathorax. In the nymph of Dissosteira and of other
Acrididae (fig. 27 A) the ventral edge of the precoxal part of the
pleuron in both the mesothorax and the metathorax is distinctly sepa-
rated from the sternum ; in the nymph of Gryllus (B) a precoxal plate
(Acx) is separated from the pleuron and intervenes between the
episternum and the sternum. The episternum of the mesothorax of
Dissosteira (fig. 26, Eps.) is marked anteriorly by a submarginal
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 45
suture (7) which is continuous through the anterior part of the meso-
sternum (.S;) and sets off from the sternum and the two episterna a
narrow anterior marginal piece, or prepectus (Ppct), which is analo-
gous to the similar sclerite of the Ichneumonidae and some other
Hymenoptera. To the posterior margin of the epimeron of the meta-
thorax (Epm3) is attached the large lateral extension of the meta-
thoracic postnotum (PN;).
The two pairs of spiracles of the thorax are presumably the meso-
thoracic spiracles and the metathoracic spiracles, each pair being dis-
placed anteriorly. The first spiracle on each side (fig. 26, Sp.) is
PIS
Ve
Fic. 27.—Pterothoracic pleura of orthopteran nymphs.
A, pterothoracic pleura of an acridid nymph, showing laterosternal arms of
pleura (Ls) separated from sterna (S2, Ss) by the pleuro-sternal sutures (r, 1).
B, mesopleuron and coxa of young nymph of Gryllus assimilis, showing a dis-
tinct precoxal sclerite (Acx) between episternum (Eps) and sternum (S).
C, inner view of B, showing the basalar and subalar muscles of the nymph
(M', M") attached dorsally on edges of episternum and epimeron, respectively.
situated laterally in the intersegmental membrane between the pro-
thorax and the mesothorax, where it is covered by the posterior fold
of the protergum. The second spiracle (fig. 26, Sp;) appears in the
adult to lie in the lower posterior angle of the mesepimeron (Epmz)
just above the base of the middle leg, and anterior to the fold between
the mesothorax and the metathorax (fig. 28), but in the nymph
(fig. 27 A) it occurs in the intersegmental fold.
The structural pattern of the internal surface of the pleural wall of
the pterothorax (fig. 28) is a replica of that of the outer surface,
except that the impressed lines of the latter are represented by ridges.
Each pleural ridge (P/R2, PIR;), however, gives off from its lower
end a large pleural arm, or pleural apophysis (P/A2, PIA;), that
projects inward across the coxal cavity, where it is closely associated
with the lateral arm of the corresponding sternal apophysis (fig. 31,
46 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
PIA, SA), and the two are connected by a dense mass of short muscle
fibers (figs. 34, 35, 86, II5).
Of particular interest in the pterothoracic pleuron are the epipleu-
rites, or the small plates situated in the membranes below the bases of
the wings (figs. 26, 28, 29, Ba, Sa). Upon these plates are inserted
the principal so-called direct muscles of the wing mechanism. In the
grasshopper there are three epipleurites in each segment, two (Ba)
Ba pe Sa ee
Fic. 28.—Inner surface of right pterothoracic pleura of Dissosteira, showing the
endoskeletal features.
Lettering as on figure 26, with the following additions: CP, pleural coxal
process; P/A, pleural arm; P/R, pleural ridge.
situated before the wing process and articulated to the episternum, and
one (Sa) in the membrane behind the wing process and above the
epimeron. The episternal epipleurites are distinguished as the
basalares, or basalar sclerites (Ba), the epimeral epipleurite as the
subalare or subalar sclerite (Sa). In most insects there is but a single
basalare. In Dissosteira the basalar sclerites are freely hinged to the
upper margin of the episternum before the wing process (fig. 29, 1Ba,
2Ba) so that they can be turned inward and downward by the muscle
inserted on their inner faces (fig. 49, E, M’'). The function of the
epipleurites in connection with the movement of the wings will be de-
scribed in Section V.
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 47
There can be little doubt that the epipleurites are derived from the
upper parts of the pleura. In a nymphal orthopteron the muscles that
are inserted on the epipleurites in the adult (fig. 49) are attached
directly to the upper edges of the episternum and epimeron (fig. 27 C,
M',M"). In many adult insects the basalare remains as an undetached
lobe of the episternum (fig. 14 A, Ba).
In the membranous corium at the base of each leg there is a small
plate (fig. 26, 7) situated before the coxa and loosely attached by its
lower end to the rim of the coxa. These scierites are evidently rem-
nants of the trochantins (fig. 13 A, B, Tm) since they exactly cor-
2Ba es
Ys \ ——
PIR Epmg
Fic. 29.—Upper edge of the metathoracic pleuron and epipleurites of Dissosteira,
inner view.
1Ba, first basalare; 2Ba, second basalare; Epm, epimeron’; Eps, episternum ;
Sa, subalare; WP, pleural wing process.
respond with the small trochantin of the prothorax (fig. 20 A, Tn),
which is identified as such by the attachment of the promotor leg
muscle upon it (fig. 33 A, 62).
The pterothoracic sterna.—The sternal plates of the mesothorax and
metathorax are united in a broad plastron covering the ventral surface
of the pterothorax, and continuous laterally, in the adult, with the
pleura by a fusion with the precoxal parts of the latter (fig. 30 A).
In the nymph of Dissosteira and of other Acrididae, as already noted,
the pleural plates of the mesothorax and metathorax (fig. 27 A, Plo,
Pls) are distinctly separate from the sterna (S»2, S3), and the pre-
coxal part of each pleuron is extended ventrally and posteriorly as a
slender arm (Ls, Ls) between the sternum and the coxal corium.
These arms are clearly remnants of the infra-coxal arcs of the sub-
48 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
coxae (fig. 18 B). In the adult grasshopper they form the ventral rims
of the coxal cavities, that of the mesothorax becoming a weakly
sclerotized plate, that of the mesothorax a membranous fold. The
definitive sterna of the pterothorax in the Acrididae, therefore, do not
appear to contain subcoxal laterosternal elements as integral parts of
their areas. In the adult of Dissosteira the pleuro-sternal suture
(fig. 30 A, 7) is obsolete in the mesothorax anterior to the coxa, but
remains distinct in the metathorax.
Fic. 30.—Pterothoracic sterna and the base of the abdomen of Dissosteira.
A, general view of pterothoracic sterna and first two abdominal sterna.
B, diagram of probable structure of mesosternum. C, diagram of probable struc-
ture of metasternum. D, diagram of structure of first abdominal sternum.
acs, antecostal suture of first abdominal sternum; Bs, basisternum; C+, coxa;
CaC, coxal cavity; JS, IJS, first and second abdominal sterna; j, prepectal
suture; k, furcal suture; Pc, precosta; Ppct, prepectus; r, r, pleuro-sternal
sutures ; sa, sa, roots of sternal apophyses; SJ, sternellum; ¢, t, infra-coxal lobes
of metasternum.
The mesosternum of Dissosteira is a broad plate (fig. 30 A, Bso, Sl2)
bounded laterally by the obsolete lines of the pleuro-sternal sutures
(vr) and the rims of the coxal cavities. Its anterior edge is slightly
convex ; its posterior border is deeply emarginated to receive a median
rectangular extension of the mesosternum (Bs;) which is dove-tailed
into the mesosternal notch. A prominent transverse suture (k), which
forms internally a ridge through the bases of the sternal apophyses
and extends laterally toward the coxal cavities (fig. 31, k, k) 1s coin-
cident with the posterior edge of the median part of the sternum and
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 49
contains the external impressions of the sternal apophyses (fig. 30 A,
sa, sa). The suture, therefore, is the furcal suture of the mesoster-
num, and the two postero-lateral, quadrate mesosternal lobes (Sls,
SI,) lying laterad of the median projection of the metasternum must
belong to the sternellar region of the mesosternum. A median pit
(spn) opening just behind the furcal suture (k) marks the site of the
internal spina (fig. 31, 2Spu), which normally is intersegmental be-
tween the mesosternum and the metasternum, but which is here fused
with the mesosternal furcal ridge (k). There can be no doubt, there-
fore, that a part of the mesosternum normally intervening between
the furcal ridge and the spinasternum, which is the median area of the
mesosternellum (fig. 30 B, S/), has been obliterated in Dissosteira, and
that the spinasternum itself has been reduced to little more than the
base of the spina.
The mesosternum of Dissosteira is thus to be analyzed into the same
structural elements that are preserved in a less modified form in the
prosternum. The sternellar region of the sternum (fig. 30 B, SJ, S7)
has been cut into a pair of lateral lobes (A, SJz, Slz) by the suppres-
sion of its median area, and the following spinasternum (B, Ss) has
been reduced to the base of the spina (fig. 31, 2Spn), which is united
with the furcal ridge (/).
The endoskeletal features of the mesosternum consist principally of
the strong furcal ridge (fig. 31, k, k) and the two sternal apophyses
(SA, SA). The latter are broad, tapering plates arising from thick
bases and extending laterally beneath the pleural apophyses, to which
they are attached by short muscle fibers (figs. 34, 35, 6). Each has
a triangular basal lobe directed forward. Anteriorly the mesosternum
is marked by the sternal part of the prepectal ridge (fig. 31, 7) which
cuts off a marginal presternal strip continuous laterally with the pre-
episternal areas of the prepectus (figs 26, 30, Ppct).
The metasternum of Dissosteira (fig. 30 A, Bs3) is wider than the
mesosternum and is separated laterally by distinct sutures (r) from
the precoxal parts of the metapleura. Its anterior margin, as just ob-
served, is extended in a large, median, quadrate lobe which is dove-
tailed between the scutellar lobes of the mesosternum. Its posterior
edge is broadly emarginate to receive a corresponding extension of
the first abdominal sternum (Pc). The median scutellar region of the
metasternum, shown diagrammatically at C of figure 30, is suppressed
in the same manner as is that of the mesosternum, and the suture of the
transverse sternal ridge (#) is here also coincident with the transverse
margin of the sternal notch; but the suture does not extend laterad of
the apophyses (sa, sa), and the lateral sternellar lobes are, therefore,
4
50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
not set off by sutures as in the mesosternum. The small triangular
plates (t, t) bordering the coxal cavities appear to be subdivisions of
the sternellar lobes rather than subcoxal laterosternal pieces, since the
ventral arms of the pleurites in the nymph (fig. 27 A, Ls) form only
the membranous folds beneath the coxal cavities in the metathorax.
There is no spina associated with the metasternum. Crampton
(1918) says the spinal pit has disappeared from the metasternum, but
he gives no evidence of its former existence. As we have seen, the
intersternal sclerotization between the metasternum and the first ab-
dominal sternum remains as an integral part of the latter, or disappears
Fic. 31.—Inner surface of ventral pleuro-sternal region of mesothorax.
Bs, basisternum; CrC2, coxal cavity; Epm, epimeron; Eps, episternum;
Isg, intersegmental groove; 7, ridge of prepectal suture; k, furcal ridge; PIA,
pleural arm; P/R, pleural ridge; PIS, pleural suture; Ppct, prepectus; Ss, an-
terior part of metasternum; SA, sternal apophysis; S/, sternellum; 2S pn, second
spina, united with furcal ridge of mesosternum.
when the first abdominal sternum becomes rudimentary. In Dissosteira
the ventral muscles of the first abdominal segment (fig. 35, JS) are
attached anteriorly on a weakly developed ridge (Ac) which crosses
the first abdominal sternum between the angles of the sternellar lobes
of the metasternum. The line of this ridge appears externally as a
faint transverse suture (fig. 30 A, D, acs). The ridge (fig. 35, Ac),
therefore, is the antecosta of the first abdominal sternum, and the
representative of the spinae of the prothoracic and metathoracic sterna.
The median plate dovetailed into the metasternum (figs. 30 A, D, 35,
Pc) is the enlarged precosta of the first abdominal sternum. It cor-
responds exactly with the postnotal plate of the metathorax (fig. 25,
PN;), which is an extension of the precosta of the first abdominal
tergum.
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 51
Il. THE THORACIC MUSCLES OF DISSOSTEIRA
The evolution of insect structure has been largely an evolution of
mechanisms made up of the cuticula and the muscles. Though the
study of the insect skeleton will remain the most important branch of
insect anatomy for purposes of taxonomic description, it is becoming
evident that the morphology of the skeleton is not to be understood
without a knowledge of the relations that exist between the cuticular
modifications and the muscles. Systematists and anatomists have con-
sumed much time and have occupied much printed space with discus-
sions of homologies between sclerites, which, in many cases, are of
little value because the fundamental structure of the parts in question
has not been studied and because mechanical relationships have been
entirely ignored. The time is at hand when we must understand in-
sects as living creatures rather than as museum specimens. Morphol-
ogy must become a basis for the study of function, including both
the physiological processes by which the insect is maintained as a liv-
ing thing, and the mechanisms by which it directs its bodily activities.
A grasshopper furnishes a particularly good subject for the study
of insect musculature. Not only are the individual muscles easily dis-
tinguished in dissections, but the muscles present are principally those
that are common to all generalized insects. Fresh specimens do not
serve well for the purpose of muscle study, but after twenty-four
hours’ immersion in 80 per cent alcohol the fiber bundles become more
compact and are more readily seen as separate muscles. Since most
of the insect’s muscles are arranged laterally, a median sagittal section
of the body will give the best approach to the muscles for an initial
examination ; but eventually it will be necessary to cut specimens into
numerous pieces, for each muscle must be followed from one attach-
ment to the other. Never accept a supposed observation for a fact
until it is seen alike in at least two preparations—not that specimens
differ, but that observations frequently do.
It is customary in describing muscles to follow them from their
origins (fixed ends) to their insertions (movable ends), but the mus-
cles of insects are in general more easily studied by finding the inser-
tion points first and then tracing the bundles of fibers out to their
basal attachments. The origins of muscles are likely to vary more in
different segments and in different species than are the insertions, and
branched muscles are often confusing until their common parts or
apodemes of insertion are determined.
The student will find that the principal thoracic muscles of Dis-
sosteira more nearly correspond with the description of the muscles
of the field cricket, Liogryllus (Acheta) campestris, given by Carpen-
52 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
tier (1923) than with the description of the muscles of Gryllus domes-
ticus given by Voss (1905). The musculature of the cricket is in some
respects more elaborate than that of the locust; but the extra fibers
constitute small and apparently secondary muscles that are not defi-
nitely repeated in insects generally. The account of the musculature of
Gryllus pennsylvanicus given by DuPorte (1920) contains inaccura-
cies, especially with regard to the muscles of the legs ; the leg muscles
of Gryllus are in no essential way different from those of Dissosteira.
No attempt will be made in this paper to homologize the muscles of
Dissosteira with those of other insects, or to correlate them with the
muscles described by other writers, since this would add too much to
the size of the paper. The student, however, should consult the recent
descriptions of the thoracic musculature of insects contained in the
works of Bauer (1910, 1924, adult Dytiscus), Speyer (1922, 1924,
larval Dytiscus), Carpentier (1923, Acheta campestris and Tachy-
cinus asynamorus), Weber (1927, Tenthredinidae; 1924, 1928, Lepi-
doptera ; 1928a, Aphis fabae; 1929, Psylla mali), and Morison (1927,
Apis mellifera). Berlese’s (1909) review of the musculature of in-
sects will need some revision in the light of more extensive compara-
tive studies of insect muscles; but a general myology of insects can
not yet be undertaken since we need more extensive information con-
cerning such groups as Apterygota, Plecoptera, and Neuroptera.
The terminology of insect musculature offers some difficulty for the
reasons that in different species the number of muscles in a functional
group is variable, the attachments may shift from one point to another,
and the functions of muscles undoubtedly homologous are often
changed as a consequence of altered relations in the skeletal parts. In
the following description of the thoracic musculature of the grass-
hopper individual muscles are designated numerically for convenience
of reference only, and the series of numbers (46 to 139) follows the
enumeration of the head muscles of Dissosteira given in a former
paper by the writer (1928).
Dissection of the thoracic muscles is simplified when the general
plan of the segmental musculature is understood. The thoracic mus-
cles of insects fall into a few major groups which, in a general way, are
as follows: (1) dorsal body muscles ; (29 ventral body muscles; (3)
tergo-sternal muscles; (4) special wing muscles; (5) pleuro-sternal
muscles ; (6) coxal wing muscles ; (7) body leg muscles ; (8) muscles
of the leg segments; (9) muscles of the spiracles. In addition there
are the muscles of the neck plates, and often oblique, lateral interseg-
mental muscles.
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 53
MUSCLES OF THE NECK AND PROTHORAX
The prothoracic and neck muscles of the grasshopper are best studied
from the mesal plane of the body. They may be exposed by cutting
into lateral halves a specimen that includes the back of the head, the
prothorax, and the mesothorax. Before removing the alimentary
canal, a branched muscle should be observed going from the side of
the protergum to the crop and the gastric caeca, which is,
| JEST ; ee
Py) SA. Bs..53, 7S B
Fic. 32.—Muscles of the neck of Dissosteira, right side, internal view.
A, muscles extending between head and prothorax, omitting 52, 53, and 54,
shown in B, inserted on first cervical sclerite. B, muscles of cervical sclerites.
Bs,, basisternum of prothorax; c, first ridge of protergum; Cv, neck; Icv,
first cervical sclerite; 2cv, second cervical sclerite; d, second ridge of protergum;
e, third ridge of protergum; Eps, prothoracic episternum; Eps:, mesothoracic
episternum; g, process of head articulating with first cervical sclerite; H, head;
1Ph, first phragma; PoR, postoccipital ridge of head; PT, base of posterior ten-
torial arm; ed, posterior fold of protergum; SA, prosternal apophysis; 1Spn,
first spina; Ss, spinasternum; 71, protergum.
46. Posterior protractor of the crop and gastric caeca (fig. 33 A) .—
A slender, branched muscle arising on lateral surface of protergum
from lower end of first tergal ridge (c) just before base of tergal pro-
motor of coxa (62) ; branching posteriorly to lateral wall of crop and
tips of gastric caeca.
The alimentary canal and fat tissue should now be removed in order
to expose the muscles in the side of the neck and prothorax, some of
which extend from the mesothorax to the head. Functionally there are
three groups of these muscles, namely, those that move the head, those
that move the prothorax, and those that move the fore leg.
54 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
It is impossible to determine, from an anatomical study alone, the
individual action of the muscles attached on the back of the head and
on the cervical sclerites (fig. 32 A, B), since their functions may vary
according to whether opposed sets of them act together or as antagon-
ists. It is evident that the dorsal muscles (fig. 32 A, 47, 48, 49) and
the ventral muscles (55) may tilt the head up or down respectively by
pulling on opposite sides of the fulcrum of the cervical sclerites (g),
or also that they may turn the head laterally if both sets on either side
act as antagonists to those of the other side, while, finally, if they all
act together they would become retractors of the head. The dorsal
muscles of the cervical sclerites (A, B, 50, 52, 52, 53) must be the pro-
tractors of the head, since their combined pull would straighten the
angles between the two sclerites of each pair and thus push the head
forward. The oblique ventral muscles of the cervical sclerites (fig. 35,
54) would appear to be accessory to the lateral movement of the
head.
47. First protergal muscle of the head (fig. 32 A).—A slender
muscle arising dorsally on protergum ; inserted dorso-laterally on post-
occipital ridge of head (PoR).
48. Second protergal muscle of the head (fig. 32 A)=—A larger
muscle arising dorsally on third ridge (¢) of protergum; inserted with
47 on postoccipital ridge of head.
49. Longitudinal dorsal muscle of the neck and prothorax (fig.
32 A).—A broad muscle from first thoracic phragma (1P/) to post-
occipital ridge of head just below 48.
50, 51. Cephalic muscles of the cervical sclerites (fig. 32 A, B).—
Origins on postoccipital ridge below 49; both extend ventrally and
posteriorly, the first (50) inserted on first cervical plate, the second
(51) on second cervical plate.
52, 53. Protergal muscles of the cervical sclerites (fig. 32 B:).—
Origins dorso-laterally on protergum at lower end of first tergal ridge
(c) ; both extend ventrally and anteriorly, crossing internal to 50 and
51, to insertions on first cervical sclerite, the first muscle with a branch
(52a) to articular process (g) of head.
54. Prosternal muscle of the first cervical sclerite (figs. 32 A, B,
33 C, 35).—A horizontal, diagonal muscle arising on prosternal apo-
physis (figs. 32, 35); inserted anteriorly on first cervical sclerite of
opposite side (figs. 32 B, 35), the right and left muscles crossing each
other medially (fig. 35). .
55. First ventral longitudinal muscle (figs. 32 A, 33 A, 35).—A
broad, flat muscle from base of posterior arm of tentorium to apophy-
sis of prosternum (figs. 32 A, 35, SA).
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 55
56. Dorsal lateral neck muscle (fig. 32 A).—A band of slender
fibers from first phragma (zPh) inserted on base of neck mem-
brane (Cv).
57. Ventral lateral neck muscle (fig. 32 A, B)—A short, flat mus-
cle from anterior edge of prothoracic episternum (F/s,), inserted on
base of neck membrane (Cv).
The prothorax is movable on the mesothorax by two oblique, lateral
intersegmental muscles on each side (fig. 32 A, 58, 59), and by three
pairs of ventral intersegmental muscles (figs. 32 A, 35, 60, 87, 88).
58. Tergo-pleural intersegmental muscle (fig. 32 A)—A broad
muscle of several sections, attached anteriorly on protergum behind
upper end of ridge d; extends posteriorly and ventrally to interseg-
mental membrane just before upper end of mesepisternum (EF ps2).
59. Sterno-pleural intersegmental muscle (figs. 32 A, 33 A, 35).—
Attached anteriorly on upper end of prosternal apophysis (figs. 32 A,
33 A, SA) ; extends posteriorly and dorsally to dorsal end of anterior
margin of mesepisternum (Eps.). In some insects this muscle is at-
tached posteriorly on the anterior angle of the mesotergum.
60. Second ventral longitudinal muscle (figs. 32 A, 33 A, 35).—
Extends between prosternal and mesosternal apophyses. Attached
anteriorly by broad base on prosternal apophysis ; tapers posteriorly to
attachment on anterior margin of mesosternal apophysis (fig. 35).
The other two sternal muscles that move the prothorax are the third
and fourth ventral longitudinals (figs. 32 A, 35, 87, 88) attached an-
teriorly on the first spina (7Spn), but they will be described with the
mesothoracic muscles. |
6r. Sterno-spinal muscle (figs. 33 C, 35).—A very small muscle
arising on base of prosternal apophysis (SA) ; the two from opposite
sides converging posteriorly to insertions on anterior end of first spina
(7Spn). Since the spinasternum (Ss) is but little movable on the
prosternum (S,) in the grasshopper, this pair of muscles can act only
as tensors or levators of the spinasternum.
The muscles that move the prothoracic leg of Dissostetra represent
the tergal promotor (fig. 11, J), the tergal remotor (J), and the
sternal remotor (L) of the primitive limb base, and the abductors (M)
and the adductors (N) of the coxa. A representative of the sternal
promotor (K) is absent in the prothorax of Dissosteira. The sternal
remotors function as posterior rotators of the coxa by reason of the
single articulation of the latter with the pleuron only; in Dissosteira
one branch of the sternal remotor arises on the spina.
50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
62. Tergal promotor of the coxa (fig. 33 A).—The largest muscle
of the prothorax. Origin on upper lateral wall of protergum, posterior
to lower end of first ridge (c) ; insertion ventrally on the small tro-
chantin (fig. 33 A, B, D, Tn).
63. Furst tergal remotor of the coxa (fig. 33 A).—Origin on lateral
wall of protergum mesad of upper end of 62 below ridge f; extends
ventrally and posteriorly external to 59 and 60 to insertion on posterior
angle of base of coxa (fig: 33 B, C, D).
64. Second tergal remotor of the coxa (fig. 33 A, B, C, D)—A
short muscle arising on lateral wall of protergum (D) beneath ridge 7;
insertion on posterior angle of coxa (C, D).
65. Third tergal remotor of the coxa (fig. 33 A, B, C, D).—A
slender muscle arising on protergum (D) in angle between ridges e
and h,; insertion ventrally on posterior angle of coxa.
66. First posterior rotator of the coxa (figs. 33 C, D, 35).—Origin
on base of sternal apophysis (figs. 33 C, 35, SA) ; insertion on pos-
terior angle of coxa.
67. Second posterior rotator of the coxa (figs. 33 C, D, 35).—
Origin on side of spina (figs. 33 C, 35, 1S pm) ; insertion on posterior
angle of coxa.
68. Abductor of the coxa (fig. 33, B, D).—A flat, two-branched
muscle arising on inner face of episternum (Eps), the larger branch
(68b) dorsally, the smaller branch (68a) in anterior ventral angle;
both inserted by a common stalk on outer rim of coxa (D) just before
pleural coxal articulation (C+P).
69. Adductor of the coxa (fig. 33 C).—Origin on outer end of
sternal apophysis (SA), or at union of the latter with pleural apo-
physis (P/A) ; insertion on inner rim of base of coxa (Cr).
The following nine muscles (70 to 7S) pertain to the segments of
the telopodite of the prothoracic leg. Three branches of the depressor
of the trochanter (77) have their origins within the body.
zo. Levator of the trochanter (fig. 36 A).—Origin dorsally in an-
terior part of coxa; fibers converge to insertion on tendon arising from
dorsal lip of base of trochanter. This is the lifting muscle of the
telopodite.
71. Depressor of the trochanter (figs. 33 A, B, C, 36 A).—A five-
branched muscle, two groups of fibers arising in the coxa and three in
the prothorax, all converging upon a strong apodeme arising from
ventral lip of base of trochanter (fig. 36 A, 77Ap). The coxal branches
arise anteriorly (77a) and posteriorly in ventral part of coxa; of the
three body branches the first (77D) arises dorsally on anterior margin
NO. 2 .THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 57
/ ae
1Spn 67 66
Fic. 33.—Musculature of the base of the fore leg of Dissosteira.
A, inner view of base of right leg, showing coxal and trochanteral muscles
arising on lateral walls of protergum; tergal ridges lettered as on figure 20 B.
B, same as A but with inner muscles removed, showing coxal and_trochanteral
muscles arising on episternum, and posterior group on tergum.
C, posterior view of prosternum, right pleuron, and right coxa, showing leg
muscles arising on sternum and pleural arm.
D, articular region of base of right coxa, and associated muscles, inner view.
58 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
of episternum (fig. 33 B, Eps), the second (77c) on ventral edge of
pleural arm (fig. 33 C, PIA), the third (77d) on lateral wall of pro-
tergum (fig. 33 A, B) just below ridge h. These groups of fibers con-
stitute the most powerful muscle of the leg and function as the de-
pressor of the telopodite as a whole.
72. Reductor of the femur (fig. 36 A).—A short, broad muscle
in posterior part of trochanter (771) arising on ventral wall of the seg-
ment; fibers extending dorsally and posteriorly to posterior rim of
base of femur, giving the latter a slight posterior flexion.
73. Anterior levator of the tibia (fig. 36 A).—An extremely slen-
der muscle arising anteriorly in base of femur; inserted by long,
thread-like apodeme on a process from anterior side of base of tibia
(as in middle leg, fig. 36 B, r05Ap).
74. Posterior levator of the tibia (fig. 36 A).—Origin dorsally in
proximal part of femur; insertion by a strong tendon on posterior
dorsal angle of base of tibia (as in middle leg, fig. 36 E, 106).
75. Depressor of the tibia (fig. 36 A).—Origin anteriorly (75a)
and posteriorly on ventral wall of femur, with branch (75c) from
base of trochanter (Tr) ; inserted by a strong tendinous apodeme aris-
ing from small ventral plate in membrane of femoro-tibial joint.
76. Levator of the tarsus—Origin on distal third of dorsal wall
of tibia; insertion on dorsal lip of base of tarsus.
77. Depressor of tarsus——Origin on ventral wall of tibia; inser-
tion on ventral lip of base of tarsus.
78. Depressor of the pretarsus: retractor of the claws (fig. 36 A).—
This muscle comprises three branches, the principal one arising pos-
teriorly in base of femur (fig. 36 A, 78), the other two in upper part
of tibia; all inserted on a long tendon extending from femur through
tibia and tarsus to unguitractor plate at base of claws.
The following two muscles are those of the first spiracle, but since
the first spiracle is situated within the region of the prothorax, its
muscles are to be classed as prothoracic. The mechanism of the spira-
cles will be discussed in Section VI.
79. Closing muscle of the first spiracle (fig. 51 B).—Origin on
ventral process of peritreme (/) ; insertion on lever of posterior lip
of spiracle (7).
80. Opening muscle of the first spiracle (fig. 51 B).—Origin on
ventral process of peritreme; insertion on base of posterior lip of
spiracle.
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 59
MUSCLES OF THE PTEROTHORAX
The musculature of the wing-bearing segments differs in many
respects from that of the prothorax, particularly in the great develop-
ment of the dorsal longitudinal muscles (fig. 34, 87, 112), in the pres-
ence of large tergo-sternal muscles (83, 84, 113) and special wing
muscles which are lacking in the prothorax, and in the presence of two
pleuro-coxal muscles that become wing muscles in the adult.
The dorsal longitudinals and the tergo-sternals constitute a group
known as the indirect wing muscles because they effect movements of
Sete Scl, oe es Selz fs
NO ane
By Wye Sea Goher ina ea
Fic. 34.—General view of the musculature in the right half of the pterothorax of
Dissosteira. Median section, seen from the left.
the wings by alternate changes in the curvature of the tergum. There
is but one special wing muscle in Dissosteira connected with each wing :
this is the wing flexor (figs. 37 A, 85; 49, D), a short muscle hav-
ing its origin on the pleuron and its insertion on the third axillary
sclerite of the wing base. In many insects there are several small
muscles from the upper parts of the pleuron to the edge of the tergum
or to the base of the wing, but representatives of these muscles are
absent in the grasshopper. The two pleuro-coxal muscles that become
important wing muscles in the adult are apparently abductors of the
coxa in the nymph (fig. 27 C, M’, M”). The first is the pronator-
60 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
extensor of the wing (fig. 49, M’), having its dorsal insertion on the
basalar plates of the adult ; the second is the depressor-extensor (M”)
with its insertions on the subalar plate. Associated with the first is a
large muscle (£) arising ventrally on the sternum. These three
epipleural muscles (E, M’, M”) together with the wing flexor (D)
constitute the so-called direct wing muscles, though only the flexor is
a true wing muscle.
The ventral muscles of the pterothorax are small ; those of the meso-
thorax (figs. 34, 35, 60, 87, 88) serve to move the prothorax ; those of
the metathorax (176, 777) can have but little motor function, since
the mesothorax and metathorax are immovable on each other, and
they are reduced mostly to tendinous strands. The muscles of the mid-
dle and hind legs are essentially the same as those of the prothoracic
leg, but the muscles of the hind tibia are particularly large and not
of the same relative size as those of the fore and middle legs.
A first dissection of the pterothoracic musculature should be made
from the median plane of the body in a specimen cut into lateral halves
(fig. 34) from which the alimentary canal and other visceral tissues
have been removed.
THE MESOTHORACIC MUSCLES
81. Longitudinal dorsal muscles (fig. 34).—A large mass of fibers
in each side of upper median part of mesothorax, attached anteriorly
on lobes of first phragma (zPh) and posteriorly on middle phragma.
82. Oblique dorsal muscles (not shown in figures).—A small mus-
cle laterad of longitudinal dorsals; arising on lateral part of scutum,
extending posteriorly and ventrally to insertion on outer part of mid-
dle phragma.
8&3. First tergo-sternal muscle (fig. 34.).—Attached dorsally on pos-
terior part of lateral prescutal lobe ; attached ventrally on anterior part
of mesosternum.
84. Second tergo-sternal muscle (fig. 34).—A very large muscle im-
mediately posterior to §3; attached dorsally by inner branch on middle
of lateral scutal area, and by outer branch on marginal lobe of scutum
behind posterior articulation of first axillary; attached ventrally on
mesosternum before inner margin of coxal cavity.
85. Pleuro-alar muscle: flexor of the wing (fig. 37 A).—This mus-
cle lies laterad of the series of dorsoventral muscles in the side of the
segment and may be noted after the latter are removed. It arises by
a broad base on upper part of pleural ridge (PIR), and goes dorsally
and posteriorly between 98 and 99 into wing base where it is inserted
on the third axillary (fig. 49, 34%).
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 61
ZOU
ZED
fe
|
Fic. 35.—General view of the ventral musculature of Dissosteira from the head
to the second abdominal segment.
62 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
86. Pleuro-sternal muscle (figs. 34, 35).—A dense mass of very
short fibers connecting the approximated ends of the pleural apophysis
and the sternal apophysis.
87. Third ventral longitudinal muscle (figs. 34, 35)—Attached
laterally on first spina (1Spm) ; extends posteriorly and laterally over
posterior end of 60 to anterior edge of apophysis of mesosternum
(S'A2).
88. Fourth ventral longitudinal muscle (figs. 34, 35).—A slender
muscle attached anteriorly on first spina (1Spn) and posteriorly on
second spina (2S pn).
The following thirteen muscles (89-101) include the muscles of
the base of the leg and the principal direct muscles of the wing.
89. Tergal promotor of the coxa (fig. 34).—Lies close behind &4
in the innermost series of lateral muscles. Origin on scutum; inser-
tion on stalked disc (fig. 37 A, B, C, 89) arising from articular mem-
brane at anterior angle of coxa close to lower end of trochantin
(B, C, Tn). The representative of this muscle in the prothorax
(fig. 33 A, 62) is inserted on the trochantin, as it is in most insects
in which the trochantin is well developed.
90. First tergal remotor of the coxa (fig. 34).—Origin on scutum ;
goes ventrally posterior to pleural arm to insertion on stalked disc
arising from inner posterior angle of coxa (fig. 37 A, B, 90).
o1. Second tergal remotor of the coxa (fig. 37 A).—A slender
muscle arising on scutum from outer end of ridge s (fig. 22 B) ; goes
obliquely ventrally and posteriorly to slender apodeme arising from
extreme posterior angle of coxa (fig. 37 A, B, 91). This muscle is
the last of the tergal muscles of the mesothorax ; it lies just external
to posterior border of go and is partially visible from median plane
(fig. 34.) between 9o and 173.
The group of mesothoracic muscles attached dorsally on the tergum
includes two segmental branches of the depressor of the trochanter
(103) which will be described later. When the tergal muscles have
been removed there is exposed a second or outer set of lateral muscles
having their origin on the pleuron (fig. 37 A). These muscles include
the abductors of the coxa, and the direct muscles of the wing. The
wing flexor (85) of the latter group has already been described as a
pleuro-alar muscle; the others are pleuro-coxal, with one pleuro-
sternal muscle. Ventrally there will be seen also the sterno-coxal mus-
cles, or rotators of the coxa, a description of which will logically fol-
low that of the tergo-coxal muscles.
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 63
Fic. 36.—Leg musculature of Dissosteira.
A, posterior view of muscles in proximal part of right fore leg, including coxa
(Cx), trochanter (77), and base of femur (Fim).
B, anterior and ventral muscles of tibia arising in femur and trochanter of left
middle leg.
C, cross-section near middle of second left tibia, proximal surface of distal half,
showing positions of tibial muscles.
D, dorsal view of trochanter (77) of left hind leg, showing anterior and pos-
terior coxo-trochanteral articulations (f, g), and levator muscles of trochanter
arising in coxa.
E, dorsal view of femoro-tibial joint of left middle leg, showing anterior and
posterior articulations (J, mm), and bases of levator muscles of tibia.
F, corresponding view of femoro-tibial joint of left hind leg.
64 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
92. Anterior rotator of the coxa (figs. 34, 35, 37).—Origin on
sternellar lobe of mesosternum (figs. 34, 35) ; extends anteriorly and
outward to anterior angle of coxa (figs. 35, 37 A, B, C).
93. Posterior rotator of the coxa (figs. 34, 35, 37).—Origin on
second spina (figs. 34, 35) ; extends outward, above 92, to posterior
inner angle of coxa (figs. 35, 37 A, B).
04,95. First and second abductors of the coxa (fig. 37 A) —Origin
on anterior ventral area of episternum (Eps) ; fibers of each converge
to a pair of long, flat apodemes arising anteriorly on outer margin of
coxa (fig. 37 B, C, 94, 95).
06. Third abductor of the coxa (fig. 37 A)—A wide, flat, fan-
shaped muscle arising on episternal area posterior and dorsal to 95;
fibers converging to insertion on a slender apodeme arising in articu-
lar membrane laterad of base of coxa just anterior to pleural articu-
lation (fig. 37 B, 96).
07. First pronator-extensor of the forewing (fig. 37 A).—A large
muscle inserted dorsally on first basalar plate (7Ba) ; extending ven-
trally to attachment on lateral part of sternum before base of mid-
dle leg.
98. Second pronator-extensor of the forewing (fig. 37 A)—
Insertion dorsally close to 97 on first basalar plate (1Ba) ; attached
ventrally on bases of apodemes of first and second abductors of coxa
(figs 337) B, G08):
99. Depressor-extensor of the forewing (fig. 37 A).—Inserted dor-
sally on subalar plate of wing base (Sa) ; attached ventrally on flat
extension of basicoxal ridge (fig. 37 B, 99) in meral region of coxa
(Mer) posterior to pleural articulation (c).
100. First adductor of the coxa (fig. 37 A).—A broad flat muscle
arising on posterior margin of mesosternal apophysis; insertion on
inner rim of coxa (A, B, 100).
ror. Second adductor of the coxa (fig. 37 A).—A smaller muscle
arising on mesosternal apophysis ; inserted on posterior angle of coxa
(A, B, ror) between attachments of 90 and 91.
The telopodite of the middle leg, or that part of the limb beyond
the coxa, has the same musculature as the telopodite of the first leg;
its muscles are the following :
102. Levator of the trochanter——Origin dorsally in base of coxa;
insertion on dorsal lip of base of trochanter.
103. Depressor of the trochanter.—-A five-branched muscle with all
branches inserted on a tongue-like apodeme arising from ventral lip
of base of trochanter. Two branches arise ventrally in the coxa, one
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 65
anteriorly, the other posteriorly; the others take their origin in the
mesothorax. The first and second body branches arise on scutum, one
medially the other on lateral margin, both pass into coxa anterior to
pleural arm; the third body branch arises on ventral margin of meso-
sternal apophysis. The trochantinal muscles effect the movement of
the telopodite as a whole.
1Ba 2Ba WP, Sa at 95 Mer
easy J ie | 99 91
\ AI ie |
98-1
me \
96—-~ IA WN
Fic. 37.—Coxal musculature of the middle leg of Dissosteira.
A, general view of pleural muscles, right side, inner view, and bases of sternal
coxal muscles.
B, base of coxa, inner view, showing muscle attachments.
C, anterior rim of coxa (Cx), trochantin (Tm), and attachments of associated
muscles, inner view.
104. Reductor of the femur.—A sheet of very delicate fibers in
posterior part of trochanter, arising in base of latter, inserted on pos-
terior rim of base of femur. This muscle is much weaker than the
corresponding muscle of the prothoracic leg (fig. 36 A, 72), the femur
of the middle leg being scarcely movable on the trochanter.
105. Anterior levator of the tibia (fig. 36 B, C).—A delicate, at-
tenuate muscle arising anteriorly in base of femur; inserted by a long
tendon-like apodeme arising from dorsal end of a slender process
from anterior margin of base of tibia (fig. 36 E, 105Ap).
5
66 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
106. Posterior levator of the tibia (fig. 37 C, E).—A long, pinnate
muscle with fibers arising on almost entire length of dorso-posterior
wall of femur ; inserted by strap-like tendon on dorsal angle of base of
tibia (E, 106). .
107. Depressor of the tibia (fig. 36 B, C)—The largest muscle in
the middle femur, comprising three groups of fibers, all inserted on a
long apodeme arising from small plate in ventral membrane of knee
joint. Principal group of fibers (B, z07a) forms a long pinnate mus-
cle arising ventrally in proximal part of femur; second group a small
bundle of fibers (107b) arising in base of trochanter and joining with
those of first group; fibers of third group (107c) arise anteriorly and
dorsally in distal two-thirds of femur and converge ventrally to inser-
tion on base of depressor apodeme.
108. Levator of the tarsus (fig. 42 A).—Origin dorsally in distal
third of tibia ; insertion on dorsal lip of base of tarsus.
tog. Depressor of the tarsus (fig. 42 A).—Origin ventrally in dis-
tal three-fourths of tibia ; insertion on ventral lip of base of tarsus.
110. Depressor of the pretarsus: retractor of the claws (figs. 36 C,
42 A, C).—Fibers arising in femur and tibia ; inserted on long, thread-
like apodeme arising from unguitractor plate at base of claws (fig.
42 C, 110Ap) and extending through tarsus (A) and tibia, and into
femur. Principal group of fibers a long, tapering bundle (fig. 36 C,
110) arising proximally on posterior wall of femur and inserted on
end of tendon ; two smaller groups of fibers in upper end of tibia, one
arising anteriorly in base of tibia, the other dorsally, both inserted on
tendon just above middle of tibia.
Since the second thoracic spiracle lies within the region of the meso-
thorax, its muscle belongs to the same segment.
111. Closing muscle of the second spiracle (fig. 52 B).—Origin on
small lobe (0) of posterior dorsal margin of mesocoxal cavity ; inser-
tion on ventral lobe of spiracular lips (7).
THE METATHORACIC MUSCLES
The musculature of the metathorax almost duplicates that of the
mesothorax, with the principal difference that there are no oblique
dorsal muscles and that there is only one pair of tergo-sternals in the
metathorax.
112. Longitudinal dorsal muscles (fig. 34).—Most of the fibers
extend between middle phragma and third phragma (3Ph), though a
few dorsal ones are attached posteriorly on the postnotal plate (PNs).
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 67
113. Tergo-sternal muscle (fig. 34).—A large muscle, the first of
the inner lateral series in metathorax, attached dorsally on lateral
prescutal lobe, and below by a wide base on lateral part of sternum
before coxal cavity. This muscle corresponds with 83 of the meso-
thorax, a scuto-sternal muscle (84) being absent in the metathorax.
114. Pleuro-alar muscle: flexor of the hind wing (fig. 38, 114).—
This muscle consists of two bundles of fibers in metathorax, one ex-
ternal, the other internal, both arising from upper end of pleural ridge,
and inserted on ventral surface of third axillary sclerite of wing base.
The outer muscle is not visible from mesal plane until the first 1s
removed. :
115. Pleuro-sternal muscle (figs. 34, 35).—A dense mass of very
short fibers connecting pleural apophysis with apophysis of meta-
sternum.
116. Fifth ventral longitudinal muscle (figs. 34, 35).—A strong
fiber, apparently a sclerotized muscle, extending from posterior edge
of mesosternal apophysis (SA) to median anterior angle of meta-
sternal apophysis (SA;).
117. Sixth ventral longitudinal muscle (figs. 34, 35).—A slender
muscle arising on second spina (2Spn), becoming tendinous pos-
teriorly ; extends posteriorly and laterally to inner extremity of pleural
apophysis of metathorax.
The ventral longitudinal muscles of the metathorax have evidently
lost their contractile nature because of the fusion of the mesosternum
and metasternum, and are converted mostly into sclerotic strands to
brace the pull of the mesothoracic ventral muscles (60, 87, 8S) on the
sternal plates of the prothorax.
The following thirteen muscles (118 to 130) are muscles of the
metacoxa and the hind wing.
118. Tergal promotor of the coxa (figs. 34, 38 A).—Lies immedi-
ately behind the tergo-sternal (fig. 34, 7273).’Arises dorsally on lateral
area of scutum (fig. 38 A) ; inserted ventrally on apodemal disc of
anterior angle of coxa (fig. 38 D, F, 178).
119. First tergal remotor of the coxa (figs. 34, 38 A).—A large
muscle arising from posterior scutal margin; goes downward and
posteriorly, behind pleural arm, to apodemal disc on posterior inner
angle of coxa (fig. 38 D, F, 179).
120. Second tergal remotor of the coxa (figs. 34, 38 A).—A slender
muscie lying close behind r79, tapering ventrally to slender apodeme
arising from extreme posterior angle of coxa (fig. 38 B, D, F, 120).
68 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
2Ba WPz Sa
/ /
1B
Fic. 38—Muscles of the hind coxa and trochanter of Dissosteira.
A, tergal muscles of leg base, and the tergo-sternal muscle (773), right side,
inner view.
B, sternal and coxal muscles of basalar and subalar sclerites (1Ba, 2Ba, Sa).
C, abductor muscles of coxa.
D, general view of muscle attachments on base of right coxa, inner view.
E, sternal muscles of leg base, dorsal view.
F, coxal muscle attachments, dorsal view.
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS ° 69
The innermost series of lateral muscles includes two body branches
of the trochanteral depressor (fig. 38 A, 733c) which will be described
later. By removing the muscles attached on the tergum, there is ex-
posed the outer series of pleural lateral muscles pertaining to the leg
and wing, and the sternal muscles of the coxa.
121. Anterior rotator of the coxa (figs. 34, 35, 38 E).—A large
muscle with fibers arising in two groups, one from lateral part of ster-
num before base of sternal apophysis, the other from sternellar lobe
behind the apophysis ; all fibers converge to insertions on anterior angle
of coxa just mesad of stalked apodeme of 178 (fig. 38 D, F, r2r).
122, 123, 124. First, second, and third posterior rotators of the
coxa (figs. 34, 35, 38 E).—Origins on posterior margin of lateral arm
of metasternal apophysis ; insertions posteriorly on base of coxa, the
first (fig. 38 D, E, F, 122) on process of meral region, the second
(123) just within posterior angle of coxa, the third (124) on posterior
part of meral rim of coxa.
The innermost pleural muscles are the large basalar and subalar
wing muscles (fig. 38 B, 127, 128, 129) ; external to them are the
abductors of the coxa.
125. First abductor (accessory promotor) of the coxa (fig. 38 C).—
A small muscle arising from anterior edge of metepisternum just
behind and below second spiracle (2S/) ; insertion anteriorly on ex-
ternal margin of coxa (fig. 38 C, D). Anatomically this muscle evi-
dently belongs to the abductor system of the coxa, but apparently it
functions as an accessory of the tergal promotor (D, 778).
126. Second abductor of the coxa (fig. 38 C).—A large flat muscle
arising on inner face of episternum and on anterior surface of pleural
ridge ; fibers converging to insertion on slender apodeme (fig. 38 D, F,
120) arising in articular membrane at base of coxa just before pleural
articulation.
127. First pronator-extensor of the hind wing (fig. 38 B).—A large
muscle attached dorsally on first basalar plate (7Ba), and ventrally on
lateral part of sternum before coxa and laterad of base of the tergo-
sternal muscle (figs. 34, 38 A, z73).
128. Second pronator-extensor of the hind wing (fig. 38 B).—Lies
close behind 127; attached dorsally on second basalar plate (2Ba),
ventrally on lateral rim of coxa (fig. 38, D, F, 128) anterior to pleural
articulation (c).
129. Depressor-extensor of the hind wing (fig. 38 B).—A powerful
muscle, attached dorsally on inner disc of subalar plate (Sa), and
ventrally on wide basicostal surface of meral region of coxa (fig. 38 D,
F, 129).
7O SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
130. Adductor of the coxa (fig. 38 C, D, E, F).—Origin on pos-
terior surface of lateral arm of sternal apophysis (E) beneath base
of first posterior rotator (122) ; goes posteriorly and downward be-
low 123 and 124 to posterior part of inner margin of coxa (C, D,
Je 82 10))
The following muscles belong to the telopodite of the hind leg. The
total number is the same as in the fore and middle leg, but there are
two distinct levators of the trochanter, and a reductor of the femur
_is lacking.
131. Anterior levator of the trochanter (figs. 36 D, 39).—Origin
on dorsal part of anterior wall of coxa; insertion on anterior lobe of
dorsal rim of trochanter (fig. 36 D).
132. Posterior levator of the trochanter (figs. 36 D, 39). —A two-
branched mus¢le arising dorsally in base of coxa ; both branches (132a,
132b) inserted on levator apodeme and supporting plate in dorsal
articular membrane close to rim of trochanter (fig. 36 D).
133. Depressor of the trochanter (figs. 38 A, E, 39).—This muscle,
as in the other legs, consists of five branches, two of which arise in
the coxa, and three in the metathorax ; all are inserted on ventral rim
of trochanter and together constitute a strong depressor of the telopo-
dite. The coxal branches arise one anteriorly (fig. 39, 133a), the other
posteriorly in ventral part of coxa. Two of the body branches arise
on scutum of metatergum, one from lateral margin, the other (fig.
38 A, 133c) from center of lateral field. These two branches converge
downward and unite before the pleural arm in a broad, tough band of
fibers that curves posteriorly beneath the pleural arm (P/A) to enter
the coxa. The third body branch arises from under surface of lateral
arm of sternal apophysis (fig. 38 A, E, 133d).
134. Anterior levator of the tibia (figs. 36 F, 39)—This muscle
appears to be represented in the hind leg of Dissosteira by only a very
delicaté tendinous strand arising from the anterior angle of the tibial
base (134Ap), and extending proximally for a short distance against
the anterior wall of the distal part of the femur. The writer was un-
able to discover muscle fibers attached to this tendon.
135. Posterior levator of the tibia (figs. 36 F, 39).—This great
muscle occupies most of the cavity of the femur (fig. 39 A, B). The
fibers arise in short, overlapping bundles from anterior and posterior
walls of femur where they are attached on the spaces between the
“fish-bone”’ ridges, with dorsal fibers of posterior set (17350) arising
in dorsal crest of femur. Anterior and posterior fibers converge to
sides of a large, thin, flat apodeme that tapers distally to a thick stalk
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 71
(fig. 39 A, 135AP) arising from dorsal margin of base of tibia. On
the base of thrs apodeme are inserted two short, strap-like branches
(fig. 36 F, 135c, 135d) arising dorsally in distal part of femur.
136. Depressor of the tibia (fig. 39).—A relatively small muscle
with long, slender fibers (1736a) arising in ventral part of femur and
converging to sides of long, tapering apodeme arising in ventral mem-
brane of knee joint. The terminal, strap-like part of this apodeme
slides over a strong, internal process (a) of ventral wall of femur.
—— a
v4 / Ny NS
Il (Gye Aree Abie
Fic. 39.—Musculature of the hind leg of Dissosteira.
A, left leg, anterior (outer) view. B, cross-section through basal half of left
femur, proximal end of distal piece, showing positions of levator and depressor
muscles of tibia and of principal tracheae.
Two small, anterior and posterior bands of fibers, arising on dorsal
wall of femur, are inserted on apodeme near its base, the anterior one
(136b) shown in the figure.
137. Levator of the tarsus (fig. 39)—A very small dorsal muscle
in distal end of tibia, inserted on dorsal rim of base of tarsus.
138. Depressor of the tarsus (fig. 39).—A small muscle, but longer
than the levator, arising ventrally in distal part of tibia, inserted on
ventral lip of base of tarsus.
139. Depressor of the pretarsus: retractor of the claws (fig. 39).—
Comprises three small groups of fibers, one arising posteriorly in ven-
tral part of femur among fibers of tibial depressor, the second (139d)
72 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
in proximal end of tibia, the third (z39c) on ventral wall of basal half
of tibia; all inserted on fine, tendon-like apodeme (139 Ap) arising
from unguitractor plate at base of claws.
IV. THE LEGS AND THEIR MUSCLES
The legs of the grasshopper are all of typical form and segmenta-
tion, but the hind legs, being specially developed as organs of leaping,
are not only of greater size than the others but differ from them in
certain details of structure and in the relative proportions of some
of the muscles. When the grasshopper sits in an ordinary resting posi-
tion it supports itself principally on the first and second pairs of legs,
the tibiae of the hind legs being flexed against the under surfaces of
day, Ne De
tn Gee, ir
Fic. 40.—Middle leg of Dissosteira, anterior surface.
Ar, arolium; Cx, coxa; Fm, femur; Tar, tarsus; Tb, tibia; Tn, trochantin;
Tr, trochanter; Un, claw.
the femora, with the knees usually held low and the tarsi barely touch-
ing the ground. (The grasshopper of illustrations commonly rests on
all three pairs of legs, with the hind knees elevated and the tibiae ex-
tended.) In its natural resting attitude, the insect is always ready for
a leap, the spring being caused by a forcible extension of the hind
tibiae, probably accompanied by a strong depression of the trochantero-
femoral parts of the legs. The chief function of the first and second
legs is the support of the body and the directing of the few movements
of walking or of changing the resting position ; the first legs are ac-
tively used also during feeding for grasping and manipulating the
edge of the leaf. When the grasshopper walks the hind legs are used
with the others in the usual fashion.
In describing the legs it is customary to use terms of orientation
as they would apply if the appendage were extended laterally at right
angles to the body. Preaxial and postaxial surfaces are called anterior
and posterior, and upper and lower surfaces are dorsal and ventral.
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 73
STRUCTURE OF THE LEGS
The general form of a grasshoppper’s leg is shown in the illustra-
tion of the middle leg of Dissosteira (fig. 40). The appendage con-
sists of a cova (Cx), a trochanter (Tr), a femur (Fm), a tibia (Tb),
a three-segmented tarsus (Tar), and a pretarsus comprising a pair of
lateral claws (Un) and a median arolium (Ar). In the articular mem-
brane before the base of each coxa there is a small trochantinal sclerite
(Tn), best developed in the prothorax.
Each leg is set into a membranous area, or coxal corium (fig. 26),
occupying an oval'interruption in the sclerotic wall of the body between
the pleuron and the sternum, known as the coxal cavity, the rim of
which is reinforced by a submarginal inflection. The coxa is hinged
to the body wall by only a single articulation, which is with the pleuron.
The rudimentary trochantin (fig. 40, Tn) does not restrict the move-
ment of the coxa. The anterior and middle coxae are free to move in
any direction, but the hind coxae, which are directed posteriorly, have
a more limited range of motion. The number of muscles inserted upon
the hind coxae, however, suggests that what little movement these
coxae possess is of much importance in the function of the hind legs.
It should be noted that the articulating surfaces of the pleuro-coxal
hinge are formed by inflections of the body and coxal walls, and there-
fore lie on the inner surfaces of the latter (fig. 41 D). In this respect
the basal joint of the leg differs from the basal articulations of the
head appendages with the head wall, for the latter are external sur-
faces of contact lying outside the articular membranes. The peculiar
character of the pleuro-coxal (subcoxo-coxal) articulations attests,
therefore, that these articulations are not homologous with the basal
articulations of the gnathal appendages on the edge of the epicranium.
The coxo-trochanteral joint and the articulations between the seg-
ments of the telopodite, except the trochantero-femoral joint, which is
but little movable, are all of the dicondylic hinge type with anterior
and posterior articulating points on a horizontal axis transverse to the
length of the leg segments. Movement at these joints is approximately
in the same vertical plane. The trochanters are closely attached to the
femora, but the hinge lines lie in a vertical plane, and the presence of a
posterior femoral muscle in the trochanter of the first and the second
leg (fig. 36 A, 72) shows that the primitive motions at the trochantero-
femoral joint were movements of production and reduction. The seg-
ments of the tarsus are movable on each other, but since they have no
musculature, they can be moved only as they are influenced by the
tendon of the ungual retractor which passes through them (fig. 44, ’).
74. SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
The coxae.—The three coxae of the grasshopper differ somewhat in
their positions on the body. The first is the most freely movable; it
projects downward and its base is almost horizontal. The second coxa
is directed outward, downward, and posteriorly; its base lies in an
oblique plane between the pleuron and the sternum. The hind coxa is
directed posteriorly ; its basal aperture is on the inner face and lies in
an approximately vertical, longitudinal plane.
Fic. 41.—Structure of the coxae of Dissosteira.
A, first coxa and base of telopodite, left, anterior surface. B, base of middle
leg, left, anterior surface. C, hind coxa and trochanter, left, anterior (outer)
surface. D, articulation of middle coxa to pleural process, right, inner view.
95Ap, apodeme of second abductor of middle coxa; Bc, basicosta of coxa;
bcs, basicostal suture; c, pleural articulation of coxa; Cvs, hind coxa; cxs, coxal
suture; Epm, epimeron; Eps, episternum; f, anterior coxo-trochanteral articu-
lation; Mer, meron; PIS, pleural suture; 7, trochantin; 77, trochanter.
Each coxa presents a well-marked basal rim, or basicoxite, set off
by a submarginal basicostal suture (fig. 41 A, B, bes) which forms
internally a strong basicosta (D, Bc). Laterally the costa of the mid-
dle coxa (fig. 37 B) and of the hind coxa (fig. 38 D, F) are enlarged
into wide, shelf-like plates for the accommodation of muscle attach-
ments. The basicoxite is very narrow or obsolete on the mesal surface
of the coxa, but on the lateral surface it forms a distinct prearticular
and a postarticular lobe, the latter being known as the meron (fig. 41 B,
Mer). The basicoxal lobes are well developed on the hind coxa (C)
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 75
but they are inconspicuous externally because each is bent outward and
flattened upon the dorsal wall of the coxa. The articular surface (c)
by which the coxa is hinged to the internal coxal process of the pleuron
(D) is strongly inflected mesally in such a manner as to bring the
point of suspension near the central axis of the coxa, thus giving the
coxal muscles a leverage on all sides of it (fig. 43). Each coxa has a
dicondylic hinge with the trochanter (fig. 36 D, f, g), the axis of which
is horizontal and transverse to the length of the leg. The dorsal sur-
tace of the coxa is deeply emarginate between the hinge points, and
the ample articular membrane that occupies the notch allows a free
upward movement of the telopodite. When the latter is deflexed, the
ventral lip of the trochanter passes inside the lower edge of the coxa.
The anterior wall of the prothoracic coxa is marked by a coxal
suture (fig. 41 A, cvs) which extends from the anterior trochanteral
articulation (f) to the basicostal suture at the articulation of the tro-
chantin (Tn). The middle coxa (B) has a similar suture (cvs) ending
at the trochanteral articulation (f), but it begins basally at the pleural
articulation (c) and thus falls in line with the pleural suture (PIS)
of the mesopleuron. The suture is absent in the hind coxa (C). The
coxal suture, when present, forms a ridge on the inner surface of the
coxal wall (fig. 36 A), the purpose of which is evidently to strengthen
the latter.
A coxal structure such as that of the middle leg of the grasshopper
(fig. 41 B), in which the anterior wall is divided by a groove (cs)
continuous with the pleural suture, is likely to be confused with the
quite different structure illustrated by the coxa of Panorpa (fig. 14 A),
in which the postarticular part of the basicostal suture (bcs) bends
distally in the coxal wall and also falls in line with the pleural suture
(PIS). The fundamental differences in the two cases, however, are
quite apparent: in the grasshopper (fig. 41 B) the outer wall of the
coxa itself is divided; in Panorpa (fig. 14 A) the meron (Mer) is
greatly enlarged and is extended into the posterior coxal wall.
The internal ridge of the coxal suture in the prothoracic and meso-
thoracic legs of Dissosteira is continued through the anterior coxo-
trochanteral articulation (fig. 36 A, f), giving a firm but flexible union
between the two articulating segments. The posterior articulation be-
tween the coxa and trochanter of the first and second legs consists of
a condyle on the trochanter opposed by a concave surface on the coxa,
but the two are united by membrane. In the hind leg both coxo-
trochanteral articulations consist of opposing. processes united by
ligament-like thickenings of the articular membrane (fig. 36 D, f, g).
76 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
The trochanters——The trochanteral segments of the prothoracic and
mesothoracic legs have the usual form of the trochanter in insects, each
being a short segment articulating as just noted with the coxa, and
united distally with the femur. The trochantero-femoral union has an
obliquely vertical hinge line and is perhaps slightly movable, since a
femoral reductor muscle is present in each of the first and second legs
(fig. 36 A, 72). The trochanter of the hind leg is a short ring-like
segment (fig. 36 D, Tr) expanded on the posterior (mesal) surface,
but so narrow externally as to be scarcely perceptible here (fig. 39,
Tr) between the coxa and the base of the femur. It is immovably con-
nected with the femur but not fused with it, and there is no trochantero-
femoral muscle in the hind leg (fig. 39 A). The apodemes of the leva-
tor and depressor muscles of the trochanter arise from small sclerites
at the base of the trochanter in the dorsal and ventral articular mem-
branes (fig. 36 D).
The femora.—In the first and the middle leg the femur is a simple
elongate segment (fig. 40, Fm) somewhat flattened in its antero-
posterior diameter (fig. 36 C). At the distal end of the femur the
anterior wall is expanded into a broad lobe that conceals the anterior
femoro-tibial articulation (fig. 36 E, 1); the ventral wall is deeply
emarginate and occupied by an ample articular membrane (fig. 36 B)
which allows a free flexion of the tibia beneath the femur.
The femur of the hind leg (fig. 39, Fm) contains the principal leap-
ing muscles, which are the extensors of the tibiae (135) ; the hind
femur is consequently greatly enlarged and is provided with special
structural features. Its length is more than twice that of the middle
femur, and its greatest vertical diameter is equal to the length of the
prothoracic femur. The flat anterior and posterior surfaces (fig. 39 B)
are ridged longitudinally above and below, and the space between is
marked by the “fish-bone” pattern of a double series of oblique
ridges. The latter separate the lines of attachment of the fiber bundles
of the extensor muscles of the tibia on the inner walls of the femur
(fig. 39, 135a). The distal end of the hind femur (fig. 36 F)
is structurally similar to that of the first and second femora (E),
but its anterior and posterior walls are strengthened by strongly
sclerotized plates.
The tibiae —The tibiae are of similar form and structure in all the
legs, each being a slender shaft used as a lever rather than as a con-
tainer for muscles, and so constructed that it can be folded beneath the
femur. The femoro-tibial articulation is a strong dicondylic hinge
(fig. 36 E, F, 1, m), and the dorsal lip of the tibial base projects well
within the end of the femur to give an efficient leverage to the extensor
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 77
muscles (fig. 44). The ventral, flexor muscles are inserted upon an
apodeme that arises from a small sclerite in the ventral membrane of
the knee joint. The knee mechanism is most strongly developed in the
hind leg (figs. 36 F, 39). The base of the tibia here forms a well dif-
ferentiated articular head bent toward the femur almost at right angles
Fic. 42.Tarsus and pretarsus of Dissosteira.
A, tarsus of middle leg disjointed, showing levator and depressor muscles
(108, 09) inserted on basal subsegment, and tendon-like apodeme (110A)
of retractor of claws (Un) arising on unguitractor plate (Utr) and extending
through tarsus.
B, dorsal view of distal end of tarsus (Tar), arolium (Ar), and claws (Un),
the latter articulated to unguifer (Uf) of tarsus.
C, ventral view of pretarsus and end of tarsus, showing planta (Pin) and
unguitractor plate (Utr) in base of pretarsus.
to the length of the segment (fig. 39), and the dorsal lip of the tibial
base is produced far into the end of the femur by an inflection of the
articular membrane. The first and second tibiae are each provided
with two rows of large, flexible, hollow spines on the distal half of
the under surface, while the hind tibia has two rows of similar spines
on its dorsal surface, but none on the ventral surface except at the end.
The tarsi—tThe tarsi are each composed of three segment-like pieces
(fig. 40, Tar) ; but the tarsal subdivisions, or articles, are clearly not
78 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
segments equivalent to the other parts of the leg, for they are inter-
connected only by infolded membranes in which there are no sclerotic
points of articulation, and none, except the basal one, is ever provided
with muscles (fig. 42 A). The large basal subsegment of the grass-
hopper’s tarsus bears three pairs of cushion-like pads on its under sur-
face ; the middle subsegment has a single pair ; the longer terminal one
has no pads. The presence of three pads on the basal subsegment 1s
suggestive that this piece is a composite of three primary tarsal articles.
The tarsal pads have been termed euplantulae by Crampton (1923).
The pretarsi.—The terminal segment in each leg of the grasshopper
bears a pair of large lateral claws (fig. 42 A, Un), but it is itself
reduced to a simple median lobe, the arolium (B, C, Ar), and has two
sclerites in its ventral wall (C, Pin, Uir). The proximal sclerite is the
unguitractor plate (Utr) ; its base is invaginated into the end of the
tarsus and gives attachment to the tendon-like apodeme (110A) of
the depressor muscle of the pretarsus, known as the retractor of the
claws. A levator of the pretarsus is lacking in all insects. The distal
ventral sclerite, possibly a subdivision of the unguitractor, is distin-
guished as the planta (Pin). The claws arise from the dorso-lateral
parts of the base of the pretarsus and are articulated dorsally to the
unguifer area on the end of the tarsus (fig. 42 B, Uf).
MUSCLES OF THE LEGS
The muscles of an insect’s leg are comprised in three groups: (1)
muscles that move the limb as a whole; (2) muscles that move the
telopodite ; (3) muscles that move the segments of the telopodite upon
each other. The muscles of the first group have their origins entirely
within the body ; they are inserted on the base of the coxa, on the tro-
chantin, or on apodemes arising in thé coxal corium. The muscles of
the telopodite arise in the coxa and within the body ; they are inserted
on the trochanter or on apodemes arising close to the base of the tro-
chanter in the articular membrane of the coxo-trochanteral joint. The
muscles of the individual segments of the telopodite beyond the tro-
chanter arise in the segments proximal to their insertions; they are
inserted either on the bases of the segments they move or on apodemes
arising in the articular membranes.
Muscles of the leg base-——The muscles associated with the coxa that
move the leg as a whole fall into three groups according to their points
of origin ; namely, muscles that arise on the tergum, muscles that arise
on the sternum, and muscles that arise on the pleuron.
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 79
The basal leg muscles arising on the tergum comprise anterior and
posterior groups of fibers, or tergal promotors (fig. 43 A, I) and
tergal remotors (J). The tergal promotors are usually contained in a
single muscle, which is inserted on the trochantin (B, 77) when this
sclerite is present and well developed, otherwise in the articular mem-
brane or on the base of the coxa. The tergal remotors often form a
Pay 2%
Fic. 43.—Diagrams of the cardinal axes of motion in a coxa articulated to the
body by a pleural articulation only, and the coxal musculature in a wing-bearing
segment.
A, mechanism of the coxal movements on the pleural articulation (c), inner
view. The cardinal movements are: (1) promotion and remotion on a trans-
verse axis (c c) by tergal promotor and remotor muscles (J, J); (2) rotation
on a vertical axis (d d) by anterior and posterior sternal rotator muscles
(K, L); and abduction and adduction on a longitudinal axis (b b) by a pleural
abductor muscle (J/), and a sternal adductor muscle (N) arising on sternal
apophysis.
B, diagram of typical musculature of a coxa in a wing-bearing segment freely
movable on the pleural articulation. /, promotor of coxa, tergum to trochantin;
J, remotor, tergum to coxa; K, anterior rotator, sternum to coxa; L, posterior
rotator, sternum or spinasternum to coxa; M, abductor, episternum to coxa;
M’', basalar muscle, basalare to coxa; M”, subalar muscle, subalare to coxa
(M' and M” are pleural abductors of coxa in the nymph, fig. 27 C) ; N, adductor,
sternal apophysis to coxa.
group of muscles. In Dissosteira the tergal promotor is a single muscle
for each leg: that of the prothorax (fig. 33 A, 62) is inserted on the
trochantin (Tm) ; that of the middle leg is inserted by an apodeme
(fig. 37 A, B, C, &9) arising between the trochanter and the coxa;
whereas that of the hind leg (fig. 38 A, 778) is attached directly on the
anterior angle of the coxa (A, D, F, 178). The tergal remotors of the
first leg comprise a group of three muscles (fig. 33 A, B, C, D, 63,
64, 65) inserted on the posterior angle of the coxa; those of the
middle leg include two muscles inserted by apodemes on the posterior
angle of the coxa (fig. 37 A, B, 90, 91) ; and those of the hind leg em-
80 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
brace two muscles (fig. 38 A, 179, 120) similarly inserted (D, F).
The two sets of tergal muscles, promotors and remotors, are clearly
antagonists pulling on opposite extremities of the longitudinal basal
axis of the coxa (fig. 43 A, b-b), the fulcrum of which is at the pleural
articulation (c).
The sternal musculature of the coxa, when complete, includes three
groups of fibers, one inserted on the anterior angle of the coxal base,
one on the posterior angle, and one on the mesal rim. The first two are
the anterior and posterior rotators (fig. 43 B, K, L) serving to turn
the coxa in the plane of its base (A) on the pleural articulation (c).
In the foreleg of Dissosteira an anterior rotator is lacking, but there
are two posterior rotators (fig. 33 C, 66, 67), the first arising on the
base of the sternal apophysis (SA), the other on the spina (1Spn) ;
both are inserted on the posterior angle of the coxa (C, D). The
middle leg has a single anterior rotator (fig. 37 A, B, 92) and a single
posterior rotator (93), the first arising on the sternellar lobe of the
mesosternum (fig. 35, 92), the second (93) on the spina (2Spu). In
the hind leg there is a single, large two-branched anterior rotator of
the coxa arising on the metasternum laterad of the base of the sternal
apophysis (fig. 35, 721), and inserted on the anterior angle of the coxa
(fig. 38 D, 121) ; and there are three posterior rotators (figs. 35, 38 E,
122, 123, 124) all arising from the arm of the sternal apophysis.
The mesal sternal muscle of the coxa (fig. 43 B, NV) is the adductor.
It pulls upward (A, NV) on the inner end of the transverse axis (c-c)
of the coxal base passing through the pleural articulation (c). In each
segment of Dissosteira the coxal adductor arises on the under surface
of the arm of the sternal apophysis (figs. 33 C, 69; 37 A, 100, IOI;
28:6, Di. Hoel 20).
The pleural muscles of the coxa include the functional abductor
fibers (fig. 43 B, 1) which directly oppose the adductor (NV), and, in
the wing-bearing segments, two other muscles (B, ’, M”) that ap-
pear to be derived from the primitive abductor system.
In the foreleg of Dissostcira the abductor of the coxa is a two-
branched muscle (fig. 33 D, 68a, 68b) arising on the inner surface of
the invaginated episternum (Eps). In the middle leg the abductor
group comprises three distinct muscles (fig. 37 A, 94, 95, 96) all
arising on the episternum. The first two are inserted by flat apodemes
anteriorly on the outer rim of the coxa (B, C, 94, 95) and perhaps
function here as accessory promotors. The large third muscle (A,
96), however, is inserted close before and distinctly laterad of the
pleural articulation (B, c) by a slender apodeme (96) arising in the
coxal corium, and it must be the functional abductor of the coxa. In
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 81
the hind leg there are two muscles in the abductor group (fig. 38 C,
125, 126), one being a small anterior muscle (725), apparently acces-
sory to the promotor (A, 178), the other a large posterior muscle
(C, 726) which unquestionably functions as an abductor.
The pleural muscles associated with the functional abductor mus-
cles of the coxa in the wing-bearing segments are attached on the outer
rim of the coxa (fig. 43 B, M’, M@”) and, in adult insects, arise typi-
cally on the epipleural basalar and subalar sclerites (figs. 37 A, 98, 99;
38 B, 128, 129), and function as wing muscles. In some adult insects,
as in Panorpa (fig. 14 B), the first of these muscles (M’) arises on a
dorsal lobe of the episternum (Ba), which is clearly the homologue of
the basalar plate or plates in other insects, such as are present in the
adult of Dissosteira (fig. 26, Ba). The posterior epipleural muscle
(M”) is always attached to the subalar sclerite in adult insects (figs.
14 B, 37 A, 38 B, Sa). Both muscles, however, in the nymph of
Dissosteira (fig. 27 C, M’, M”) and in other nymphal Orthoptera,
arise directly from the upper edge of the pleuron, one on the epis-
ternum, the other on the epimeron, and, if they act together, they
must be abductors of the coxa. The epipleural muscles, therefore,
appear to be groups of coxal abductor fibers that have become specially
developed as secondary wing muscles in the adult. The first is a
pronator-extensor of the wing; the second is the depressor-extensor
of the wing (fig. 49, M7’, M”).
The foregoing analysis of the basal leg musculature of Dissosteira
shows that the coxa is provided with six sets of muscles, including
an anterior and a posterior group of fibers arising on the tergum
(fig. 43 B, I, J), an anterior and a posterior group arising on the
sternum (K, L), a lateral group arising on the pleuron (M, and also
M’' and M” in the wing-bearing segments), and a mesal group arising
on the sternum (NV). The anterior and posterior dorsal and ventral
muscles may be supposed to represent the theoretical primary tergal
and sternal promotors and remotors of a primitive limb basis (fig. 6,
I, J, K, L), which have become transferred to the coxal region (fig.
11) after the subdivision of the basis into subcoxa and coxa. The
lateral and mesal muscles, therefore, are subcoxo-coxal muscles, the
fibers of the first (JZ) retaining their origins on the subcoxal pleuron,
those of the second (N) having been transferred to the sternum,
perhaps by the incorporation of the ventral rim of the subcoxa into
the definitive sternal plate.
Muscles that move the telopodite—The muscles that operate the
telopodite, or that part of the leg beyond the coxo-trochanteral hinge,
6
82 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
comprise the muscles normal to the trochanter, which are a levator
and a depressor arising in the coxa (fig. 44, O, Q), and also special
depressor muscles (P) that have their origin in various parts of the
body segment carrying the leg. The basal lip of the trochanter usually
projects into the coxa well beyond the line of the coxo-trochanteral
hinge, thereby giving a strong leverage to the depressor muscles in-
serted upon it.
The branches of the trochanteral depressor arising within the body
segment vary much in different insects and in different segments of
Fic. 44.—Diagram of the mechanism of the hind leg of a grasshopper.
O, levator muscle of trochanter, or extensor of telopodite, origin in coxa;
P, body branch of depressor of trochanter, or flexor of telopodite, origin on
tergum; QO, coxal branch of depressor of trochanter; S, levator of tibia; 7, de-
pressor of tibia; U, levator of tarsus; , depressor of tarsus; X, X, tibial
branches of retractor of claws; x, tendinous apodeme of retractor of claws aris-
ing on base of unguitractor plate.
the same insect. In the prothorax of Dissosteira there are three body
branches of the trochanteral depressor, one arising on the episternum
(fig. 33 B, 77), the second on the pleural arm (C, 7Zc), and the
third on the tergum (B, 71d). In the mesothorax there are two
body branches of the muscle, both arising on the tergum. In the
metathorax a long outer branch and an inner branch (fig. 38 A, 133c)
arise on the tergum, and a short branch takes its origin on the under
surface of the lateral arm of the metasternal apophysis (fig. 38 A, E,
133d). These muscles ordinarily serve to lift the body on the legs,
but those of the hind legs of the grasshopper are probably accessory
to the extensor muscles of the tibiae in the act of leaping (fig. 44, P).
Muscles of the telopodite segments.—Since the trochantero-femoral
joint usually has but little movement in insects, the muscles of the
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 83
femur arising in the trochanter are small or absent. In Dissosteira
a posterior, or reductor, muscle only is present in the trochanter of
the first leg (fig. 36 A, 72) and in that of the second leg. In the hind
leg there is no movement at the trochantero-femoral joint and femoral
muscles are lacking. The usual flexion between the trochanter and
the femur of insects is anterior and posterior (production and re-
duction), and generally only a reductor muscle is present, called the
“rotator” of the femur by some writers (Morison, 1927; Weber,
1929).
The femur is occupied mostly by the tibial muscles (fig. 44, S, T),
but it contains also the most proximal branch of the flexor of the
claws (X). The tibial musculature comprises levator muscles (S)
and depressor muscles (7). In Dissosteira there are two levator
muscles in each leg, a larger posterior one (fig. 36 E, 106, F, 135),
and a very small anterior one (EF, 105, F, 134). In the fore and middle
legs the depressor of the tibia (fig. 36 B, C, 107) is larger than the
levator, and it has a basal branch arising in the trochanter (B). In
the hind leg the relative proportions of the two muscles are reversed,
the posterior levator, or extensor, of the tibia consisting of the great
masses of fibers arising on the ridged anterior and posterior walls of
the femur (fig. 39, 735a, 1350), and including smaller branches
(135c) arising on the dorsal wall in the distal part of the femur. The
anterior levator of the tibia in each leg consists of a very slender
bundle of fibers arising anteriorly in the base of the femur (fig. 36 A,
73) and inserted by a long, thread-like apodeme on the head of the
tibia (FE, r0o5Ap, F, 134Ap).
The tibiae contain the levator and depressor muscles of the tarsus,
and the tibial branches of the flexor of the claws. The tarsal muscles
are relatively largest in the fore and middle legs of Dissosteira; in the
hind leg they occupy only the distal part of the tibia (fig. 39, 137, 138).
The tarsus contains no muscles, the tarsal segments, as before
noted, being flexible upon one another but not independently movable.
This condition pertains to all insects. The tarsus is traversed by the
“tendon,” or thread-like apodeme, of the flexor of the claws (figs.
39, 139AP, 44, x).
The claws of insects are provided with only a flexor, or rectractor,
muscle, which is the depressor of the pretarsus (fig. 44 X), or the
homologue of the depressor of the dactylopodite in arthropods gen-
erally. The fibers of the claw muscle arise in several groups in the
tibia and femur, and are inserted on a long tendon-like apodeme that
arises from the base of the unguitractor plate (fig. 42 C, Utr) and
84 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
extends through the tarsus and tibia into the femur. In Dissostetra
two small groups of fibers of the claw muscle arise in the upper part
of the tibia (fig. 39, 139, 139c), and one arises posteriorly in the
base of the femur (fig. 36 A, 78). The pull of the muscles on the
tendon retracts the unguitractor plate and flexes the claws ventrally.
The extension of the claws probably results from the elasticity of
their basal connections and the pressure on the supporting surface.
Typically, the muscles of the pretarsus should arise in the tarsus.
It is probable, therefore, that the extension of the fibers into the tibia
and femur in insects (and also in chilopods and diplopods) is a sec-
ondary condition produced by a proximal migration of the primitive
muscle. In Crustacea and Arachnida the pretarsus, or dactylopodite,
is provided with levator and depressor muscles, both of which have
their origin in the tarsus, or propodite. In some Arachnida there are
two pretarsal claws, as in most insects, but the pretarsus has lateral
articulations with the end of the tarsus, and is provided with dorsal
and ventral muscles.
V. THE WINGS AND THEIR MECHANISM
The wing mechanism of the grasshopper is equally developed in
each segment of the pterothorax. The hind wings, though much more
extensive than the forewings, or tegmina, and probably the chief
organs of flight, have no advantage over the latter except in the
stronger development of the flexor apparatus. The forewings, on the
other hand, have a more powerful levator equipment than the hind
wings because of the presence in the mesothorax of the second pair
of tergo-sternal muscles attached dorsally on the scutum (fig. 34, 84).
In structure, the hind wings (fig. 45 B) differ from the forewings
(A) only in the reduction of the costal area and in the great ex-
pansion of the anal area.
STRUCTURE OF THE WINGS
In general structure, articulation, and mechanism the acridid wings
differ little from the wings of other Orthoptera. The tegmina when
at rest are flexed over the body in a manner to form a high roof with
steeply sloping sides (fig. 50 A, W.) covering the back of the abdomen
and inclosing the folded hind wings (W,) in the space above the
latter. The anal areas of the tegmina overlap dorsally in a median
horizontal plane, the left tegmen being usually on top; the pre-anal
areas form the lateral inclines of the tegminal roof. The bend be-
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 85
tween the two wing areas takes place along the anal fold (figs. 45 A,
50 A, AF), and is produced mechanically during the flexion of the
wing. The broad hind wings are folded in a complicated manner, to
eroc kM i Pra
Mar
“ \\
Hi
Ju (RN
\
2G.) SASvIiCu WN yan
i
\
SS \
_ ;——1
SS ———— Oe
—
——
—
7A. B
Fic. 45.—The wings and wing veins of Dissosteira.
A, fore wing, or tegmen. B, hind wing. A, anal veins, anal area of wing ;
1A, first primary anal vein; 74, seventh primary anal; C, costa; 1Cu, first
cubitus ; 2Cu, second cubitus; /, intercalary vein; 71, 7, secondary anal veins of
first anal plait; k, first concave anal vein; Ju, jugal area of wing; /, second con-
cave anal vein; M, media; Pra, preanal area of wing; q, basal support of anal
veins; R, radius; A, first branch of radius; Rs, radial sector; r, first anal plait
of wing; R-+ M, united shafts of radius and media; Sc, subcosta; VD, vena
dividens.
be described later, and when fully flexed are concealed beneath the
overlying tegmina (fig. 50 A).
The area of an insect’s wing presents usually three well-defined
regions, namely, a preanal region, an anal region, and a small, pos-
86 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
terior, basal region, generally membranous, which Martynov (1925)
terms the jugal region. The three wing regions are shown in typical
form in the forewing of the grasshopper (fig. 45 A). The preanal
region (Pra) is that lying anterior to the anal fold; the anal region
is the region of the anal veins (4), the jugal region (Ju) is the
membranous basal fold of the wing. In many insects the jugal region
contains one or two definite veins unconnected basally, or an irregular
network of small veins.
The hind wing of the grasshopper (fig. 45 B), and of other insects
with similar fan-shaped wings, is usually regarded as differing from
the forewing in the great expansion of the anal region. According to
Martynov (1925), however, the true anal region of the hind wing
in Acridium is that part (fig. 45 B, 7) between the anal fold, or vena
dividens (VD), and the first vein springing directly from the basal
support (q) of the anal fan (designated 1A in fig. 45 B). Three
veins (1, 7, k), branching from a common base, lie in this region in
the wing of Dissosteira. The following part of the wing, or that
containing the veins attached directly to the basal support (q) of the
anal fan, Martynov claims is a development of the jugal region of the
more primitive type of wing. A jugal area thus developed into a
functional wing region he calls the “ neala.”
Martynov deduces his interpretations of the morphology of the
acridid wing from a general study of the wings in other orders of
insects. In Dissosteira, however, the vein designated 1A in the hind
wing (fig. 45 B) is so clearly the homologue of 1A in the forewing
(A), considering the basal relations and the connection with the third
axillary sclerite (fig. 47 A, B, 3A), that Martynov’s interpretation
is not convincing. The area (7) of the hind wing (fig. 45 B), lying
between VD and 1A, forms the first fold of the anal region (fig. 50 B)
in which the vein k occupies the position of a “ concave” vein at the
bottom of the fold, while the two preceding veins (7, 7) strengthen
the anterior wall of the fold. The three veins of this region (fig.
45 B, i, j, k) are branches of the first primary anal vein (14). Mar-
tynov’s general study of the wing regions, however, throws much
light on the wing mechanism and morphology.
Venation of the wings—While the venation of the grasshopper’s
wings is comparatively simple, it is difficult to make a satisfactory
interpretation of the homologies of the veins in the posterior parts
of the preanal regions. If the relation of the vein bases to the axillary
sclerites is taken as a guide to the identities of the veins themselves,
the veins of the adult may be named consistently in the two wings, but
their relation to the nymphal wing tracheae is not clear in all cases.
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 87
The forewing (fig. 45 A) has a broad anterior, or costal, area in
which there is no vein represented by a costal trachea in the nymph
(fig. 46 A), though the anterior margin is strengthened by a vein-like
thickening. The first vein (fig. 45 A, C) is evidently the costa branch-
ing from the subcosta (Sc), though the common basal stalk has the
usual relation of the subcosta to the first axillary sclerite (fig. 47 A,
1Ax). The next vein (J) is unquestionably the radius, as shown by
its distal branches and by its basal connection with the second axillary
(fig. 47 A, 2Ax). The media (M) is united proximally with the
radius and with one of the median sclerites of the wing base (m’).
The first long vein following the media is a two-branched cubitus
(1Cu), between which and the basal part of the media is the inter-
calary vein (J). Then comes an unbranched vein, here designated
2Cu, lying close before the anal fold (AF), and finally a group of
three veins (4) connected basally with the third axillary, or flexor
sclerite of the wing base (fig. 47 A, 34x).
If we identify the “anal veins” as those veins lying posterior to
the anal fold and connected basally with the third axillary, there are
then but three anal veins in the forewing of Dissosteira (fig. 45 A,
A, fig. 47 A, 1A, 2A, 3A). An incomplete vein (fig. 47 A, s) lying
just behind the anal fold (AF) is apparently a secondary vein. The
vein immediately before the anal fold (2C) is the “ first anal ”’ of the
Comstock-Needham system, but probably it is the vein regarded as a
part of the cubitus by Tillyard (1919) and others, designated Cuz by
Tillyard and cubital sector by Karny. In Dissosteira the vein in ques-
tion has no basal connections and is here termed the second cubitus
(2Cu). It clearly belongs to the cubital area of the wing. In the nym-
phal wing of an acridid, as illustrated by Comstock and Needham (fig.
46 A), tracheal precursors of the cubitals are not evident, since the
final group of three tracheae springing from a common basal stem
would appear to represent the group of three anals in the adult wing
(fig. 45 A, A). According to Comstock (1918), however, the first vein
of this group is the “cubitus” (fig. 46 A, Cu), and the second the
“first anal” (14). The identity between the nymphal tracheae and
the adult veins in the forewing is certainly not clear, and no solution
of the problem can be offered here.
The vein tentatively called “second cubitus”’ in this paper (figs.
45 A, 47 A, 2Cu) is, by nomenclatural priority, the true anal vein,
and the name “anal,” though a poor designation, should be retained
for it, while a new term should be devised for the veins lying posterior
to the anal plica associated with the third axillary sclerite. The fan-
,
88 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
like wing region between the anal and jugal plicae might appropriately
be called the vannus (Latin, fan), and its veins termed the vannal
veins. This region plays a passive part in flight. The pre-vannal part
of the wing is the true remigium (Latin, oar) of the flight mech-
anism, being the region of the wing directly productive of motion. We
might then say that the area of the wing distal to the basal axillary
region is divided into a remigial, a vannal, and a jugal region. The
separating folds, when present, would then become the vannal and
the jugal plicae. The jugal region expanded is the neala of Martynov.
Fic. 46.—Wings of an acridid nymph. (From Comstock, after Comstock
and Needham.)
A, fore wing. B, hind wing. The tracheal identifications as given by Com-
stock and Needham: 14, first anal; Cu, cubitus; MM, media; FR, radius; Sc,
subcosta.
In the hind wing (fig. 45 B) the costa (C) forms the anterior
margin of the wing and is united basally with the subcosta (Sc).
The base of the subcosta (fig. 47 B, Sc) does not reach the first
axillary sclerite (14), evidently by reason of the reduction of the
anterior process of the latter, neither does it articulate with the pre-
scutal lobe of the tergum (fig. 22 A, 7), but it is connected with the
latter by a ligament-like thickening of the wing membrane (fig. 47 B,
d). The radius (R) is well developed, branched distally, and con-
nected basally with the second axillary (fig. 47 B, 247). The ap-
parent media is united proximally with the radius; its free part con-
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 89
sists of a single branch (J) given off from the radial sector (Rs).
Between the basal radio-medial shaft (A + MM) and the first anal
fold, there are two veins in the cubital area (1Cu, 2Cu) ; the first
(fig. 47 B, rCw) is united proximally with the radio-media, the second
(2Cw) has no basal connections. In the hind wing, therefore, as in
the forewing, there are two distinct cubitals, here named the first
cubitus (zCu) and the second cubitus (2Cu). Each of these veins
is represented by a trachea in the hind wing of the nymph (fig. 46 B) ;
the first is the “cubitus” (Cw) of Comstock, and the second the
“first anal”? (14). Since both of these veins lie anterior to the anal
fold in the adult wing (figs. 45 B, 47 B), however, the writer would
agree with Tillyard (1919) that the second is a cubital rather than
an anal vein. The orthopteran wing suggests that the second cubitus
has the status of an independent vein rather than that of a basal
branch of the first cubitus.
The anal fold of the hind wing is double (fig. 50 B), consisting of
two plicae, or lines of flection in the wing membrane, between which
lies the vena dividens (figs. 45 B, 47 B, 50 B, VD). According to Till-
yard the vena dividens is the “ first anal,” but since it has no basal con-
nection with the other anals (fig. 47 B), the writer would regard it
as a secondary, interpolated vein. The incomplete vein of the fore-
wing lying just behind the anal fold (fig. 47 A, s) may represent the
vena dividens of the hind wing, but it appears rather to correspond
with the vein 7 of the hind wing (figs. 45 B, 47 B).
The anal veins of the hind wing form a distinct group lying pos-
terior to the anal fold. All the primary anals spring from a basal sup-
port (fig. 47B, q) which is attached anteriorly to the distal arm of
the third axillary (34%), and which, in the grasshopper, is braced
posteriorly by an arm from the tergum (figs. 24, 47 B, p). There
are ten primary anal veins. A fork from the first (714) divides into
three branches (7, j, k) lying in the first lap of the wing that folds
beneath the preanal region when the wing is flexed (fig. 50 B). Al-
ternating with the primary, or “ convex,” anal veins are nine secon-
dary “concave” veins lying in the troughs of the folds between the
primary anals, while the vein (#) branching from the first anal is the
concave vein of the fold between the vena dividens and the first anal
(fig. 50 B, Rk).
Articulation of the wings—In the membrane of each wing base
are four axillary sclerites. The first and the fourth (fig. 47 A, B, 1Ax,
4Ax) are hinge plates articulating with the edge of the tergum; the
second (24x) is the pivotal sclerite of the wing base ; the third (347)
TH
7A
Fic. 47.—The wing bases of Dissosteira.
A, base of tegmen. B, base of hind wing. 14, 2A, 3A, 7A, first, second, third,
and seventh primary anal veins; AF, anal fold; 1Ax, 2Ax, 3Ax, 4Ax, first,
second, third, and fourth axillary sclerites ; C, costa; Cu, first cubitus; 2Cu,
second cubitus; d, attachment of base of subcosta to prescutal lobe of tergum;
I, intercalary vein; i, secondary vein of first anal plait; k, first concave anal
vein; J, second concave anal vein; M/, media; m, m’, median plates of wing base;
p, posterior arm of tergum supporting the anal veins; g, basal support of anal
veins; R, radius; R + M, united basal shafts of radius and media; s, secondary
vein of fore wing behind anal fold; Sc, subcosta; tg, tegular rudiment; ’D,
vena dividens.
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS gli
is the flexor sclerite. In addition to the axillaries there are two plates
in the median area of the forewing (A, m, m’), and a single median
plate in the hind wing (B, m).
The first axillary intermediates between the edge of the tergum
and the second axillary, with each of which it is movably connected,
and usually, by an anterior process, it articulates with the base of the
subcostal vein. The first axillary is confined to the dorsal membrane
of the wing base. In the forewing of Dissosteira the first axillary
(fig. 47 A, rAx) is a flat plate with a narrow anterior process curved
outward to meet the base of the subcosta. The sclerite bridges the
lateral emargination of the tergum (fig. 22 A, Em) ; its anterior end
is supported on the anterior notal wing process (ANP), and its pos-
terior part is hinged to the lobe of the scutum (0) behind the emargi-
nation ; its oblique outer margin articulates with the second axillary
(fig. 47 A). The first axillary of the hind wing of Dissosteira (B,
1Ax) is exceptional in the reduction of its anterior process which
does not meet the base of the subcosta (Sc).
The second axillary presents an exposed surface in both the dorsal
and the ventral membranes of the wing base. Its dorsal part forms
a triangular plate (fig. 47 A, B, 2A) lying lateral of the first axillary,
and closely hinged to the oblique outer margin of the latter; its pos-
terior outer margin articulates with the proximal median sclerite (#7) ;
to its anterior end is attached the base of the radius (/¢). The ventral
part of the second axillary forms a strong, concave plate (fig. 48,
2Ax) resting by its lower edge on the pleural wing process (WP).
The second axillary differs somewhat in shape in the two wings of
Dissosteira, as shown in the figures, but its structure and associations
are the same in both.
The third axillary is developed principally in the dorsal wing mem-
brane (fig. 47 A, B, 34%), but it includes also a small sclerotization
in the ventral membrane (fig. 48, 341). The dorsal part of the third
axillary (fig. 47) has the form of a strong bar extending outward, in
the fully-expanded wing, from the small fourth axillary (44x) to
the anal veins, which latter it supports by an arm bent forward from
its distal end. The mesal part of the sclerite bears a strong, elevated
process on its anterior margin upon which is inserted the flexor muscle
of the wing. Distal to the muscle process, the proximal median plate
(m) is firmly attached to the third axillary and is functionally a part
of it. In the forewing (fig. 47 A) the distal median plate (m’) is
hinged to the outer margin of the proximal plate.
g2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
The fourth axillary (fig. 47 A, B, 44x) is a plate of the dorsal
wing membrane only. It is small in each wing and serves merely as a
connective between the edge of the tergum and the third axillary.
It is probably a detached lobe of the tergum, since it is usually absent
in insects that have a posterior notal wing process.
Beneath the base of each wing are the epipleurites, or small scler-
ites derived from the pleuron, which are intimately associated with
the wing mechanism in the adult insect. In Dissosteira there are in
each segment two episternal epipleurites, or basalares, (fig. 48, IBds,
2Baz, 1Baz, 2Ba;), and a single epimeral epipleurite, or subalare
(Sa). The basalares are hinged to the upper edge of the episternum
(Eps), and are connected with the subcostal region of the wing base
by a ligamentous thickening (a) of the ventral wing membrane. The
subalare (Sa) lies free in the subalar membrane behind the wing
process, but it is connected with the ventral plate of the second ax-
illary (2Ax) by a thickening (b) of the intervening membrane.
THE WING MECHANISM
Flying insects are unquestionably descended from wingless an-
cestors. When paranotal lobes were first evolved on the thoracic seg-
ments, the insect was already organized for terrestial locomotion—
there was no provision for future organs of flight. When movable
wings were evolved from the paranotal lobes, they had available for
their purposes only a motor mechanism developed for other pur-
poses. It needed but an area of flexibility at the base of each paranotal
extension to convert the lobe into a movable flap. The dorsal ends of
the pleura, previously supporting the bases of the paranotal lobes,
easily became fulcra on which the wing flaps could rock up and down.
A contraction of the longitudinal muscles of the dorsum could now
give a down-stroke to the wing flaps by producing an upward curva-
ture in the tergal plates of the wing-bearing segments, and probably
at first the elasticity of the terga sufficed to produce the up-stroke.
Thus, apparently, by the simple device of becoming flexible at their
bases, the paranotal lobes became wings that could be weakly flapped
up and down by the simple motor equipment already at hand.
Modern insects, however, have added much to the primitive wing
mechanism. In each of the wing-bearing segments there are powerful
tergal-depressor muscles, which, since they do not occur in the pro-
thorax or in the segments of the abdomen, are probably specially
developed wing muscles, though they may be supposed to have been
evolved from small, lateral tergo-sternal muscles such as are usually
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 93
present in the abdomen. Being attached ventrally on the sternum,
these muscles indirectly impart a strong up-stroke to the wings by
flattening the dorsal curvature of the tergum. The down-stroke of
the wings produced by the contraction of the longitudinal dorsal
muscles has been strengthened in two principal ways: first, by the
obliteration of the secondary intersegmental membranes between the
terga, thus eliminating lost motion ; and second, by the great enlarge-
ment of the dorsal muscles themselves in the wing-bearing segments.
hea
a
WP, _ 2 Nap bane
a
~ _1Baz
--2Baz
Bal fos
2Bay Eps, \ Epm,\ Epsz \Epm;
paaey
Fic. 48.—Ventral surface of the base of the left tegmen, and upper part the
pleuron of Dissosteira.
a, thickening of membrane uniting basalar sclerites with humeral angle of
wing ; 2A x, ventral plate of second axillary; 34x, ventral plate of third axillary ;
b, connection between subalar sclerite and ventral plate of third axillary; 1Ba,
first basalare; 2Ba, second basalare; Sc, base of subcostal vein; Sa, subalare;
WP, pleural wing process.
The suppression of the intertergal membranes has been accomplished
by a fusion between the successive tergal plates, or by a forward
extension of the precostal lip of the tergum until it meets the pos-
terior edge of the preceding tergum. Thus are produced the post-
notal plates between the mesothoracic and metathoracic terga, and
between the metathoracic and first abdominal terga. The enlargement
of the dorsal muscles has been accompanied by the development of
supporting plates (phragmata) from the ridges of the muscle attach-
ments on the primary intersegmental folds. Furthermore, each tergal
plate has been strengthened and better adapted to its function in the
flight mechanism by the development of internal ridges, the principal
94 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
ones of which are so arranged as to bring the peak of the curvature
in the tergum on a transverse line between the wing bases.
The Odonata are commonly said to have a wing mechanism quite
different from that of other insects. On the basis of von Lendenfeld’s
(1881) description of the odonate wing muscles, the dragonflies have
been supposed to be equipped with a special set of muscles inserted
directly on the wing bases. A study of the thoracic musculature in
either the Anisoptera or the Zygoptera, however, will show that there
are ony two small muscles that can be regarded as special wing
muscles ; one of these is accessory to the pronator of the wing, the
other to the depressor. The large pronator and depressor muscles,
though they arise ventrally on the lower edge of the pleuron and are
inserted directly on the two basal plates of the wing, are evidently the
homologues of the basalar and subalar muscles of other insects. Two
smaller muscles lying mesad of the pronator are clearly leg muscles
since they have their origins on the coxa and their insertions on the
extreme lateral edge of the tergum. Von Lendenfeld ascribed these
muscles to the wings ; he describes them as arising on the pleuron and
as inserted on the wings. Each wing has a homologue of the flexor
muscle in other insects, though it does not function as such because
of the lack of a flexor mechanism in the base of the odonate wing.
The tergo-sternal muscles are highly developed, their ventral attach-
ments are on the sternum and their dorsal attachments on the antero-
lateral lobes of the tergum. The dorsal longitudinal muscles are re-
duced to a pair of small, divergent fiber bundles attached anteriorly
on the median apodemal spine of the tergum, and posteriorly on the
anterior margin of the following tergum. The wing mechanism of
the dragonflies is thus merely an extreme modification of that common
to all insects.
A wing, in order to be an efficient organ of progressive flight, must
be capable not only of an up-and-down movement, but also of anterior
and posterior movements accompanied by a partial rotation on its
long axis. The anterior margin of the wing must be brought forward
and deflected during the down-stroke, and lifted with a posterior
movement during the up-stroke. The rotary movement of the insect’s
wing is caused partly by the structure of the wing itself and its re-
sponse to air pressure, and partly by the nature of the wing articula-
tion on the body, but it is greatly augmented by muscles that pull
downward on the base of the wing, one before the pleural fulcrum,
the other behind it. These muscles are inserted on the basalar and
subalar sclerites beneath the wing base (fig. 48, Ba, Sa). Two of them
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 95
are evidently muscles of the leg that have been taken over into the
service of the wing, for they are attached ventrally on the coxa (fig.
49, M’, M”) ; the other (£) arises on the sternum, or in some insects
on the pleuron, and is perhaps a specially developed wing muscle. The
two muscles of the basalar sclerites (E, M’) are called pronators be-
cause they deflect the costal wing margin. The muscle of the subalar
sclerite (M”) not only deflects the posterior part of the wing, but it
acts as a powerful depressor of the entire wing by reason of its con-
nection (b) with the ventral plate (c) of the second axillary (24+).
These muscles probably also enable the insect to alter its course during
flight, and, by changing the plane of the wing movements, to hover in
the air, or to fly sidewise or backward.
Finally, most insects have found it advantageous to fold the wings
posteriorly over the body. The folding of the wings has involved
the development of a mechanism for their flexion and extension. The
ability of the wing to be flexed depends upon the mechanism of its
axillary region, but the flexing is caused by one or more flexor muscles
arising on the pleuron and inserted on the third axillary sclerite (fig.
49, D). The extension of the wing is produced by the basalar and
subalar muscles (F, M’, M”). Considering the other functions of
these muscles, the first, therefore, is a pronator-extensor of the wing,
the second a depressor-extensor.
The special features in the mesothorax and the metathorax of the
grasshopper that contribute to the mechanism of the wings have been
described in Section II of this paper. It was there shown that the
fusion of the pleurites and sterna of the mesothorax and metathorax
converts these segments into a strong trough-like structure covered
dorsally by the two wing-bearing terga. The union of the pleural and
sternal elements in the pterothorax is probably a direct adaptation to
the leaping function of the hind legs, but the resulting structure also
gives a strong framework for the support of the wings and the wing
muscles. The tergal plates are separated from the edges of the pleuro-
sternal trough by the ample membranes of the wing bases, and they are
thus free to respond to the downward pull of the tergo-sternal muscles.
The close union of the terga (fig. 25) and the great size of the dorsal
muscles (fig. 34) give efficiency to the latter as elevators of the wings.
When the wings are spread they are pivoted on the pleural wing proc-
esses by the second axillary sclerites of their bases, and, being
closely hinged to the terga by the first and fourth axillaries, they are
sharply thrown upward when the tergal plates are depressed, and
are turned downward when the terga are elevated.
96 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
The mechanism for extending and flexing the wing is highly com-
plex. The muscles that produce the movements of extension and
flexion depend for their effect on the details of shape and inter-rela-
tionships in the axillary sclerites, on the articulation of the sclerites
with the tergum and pleuron, on their connections with the bases of
the wing veins, and on the structure of the wings themselves.
MeN
! | ED ( ae
Ax
ay) Ua
V7 | ———
Fic. 49.—The pleural elements of the wing mechanism in the mesothorax
of Dissosteira.
a, thickening of cuticular membrane uniting basalar sclerites with humeral
angle of wing (see fig. 48) ; 2Ax, second axillary; 34.4, third axillary (first and
fourth axillaries removed) ; b, thickening of cuticular membrane uniting subalar
sclerite (Sa) with ventral plate (c) of second axillary; Ba, first basalare; c,
ventral plate of second axillary resting on pleural wing process (see fig. 48) ;
D, flexor muscle of wing, inserted on third axillary; E, pleuro-sternal muscle,
or first pronator-extensor of the wing, inserted on basalar sclerite; 1’, episternal
pleuro-coxal muscle, or second pronator-extensor of the wing, inserted on
basalar sclerite; 1”, epimeral pleuro-coxal muscle, or depressor-extensor of the
wing, inserted on subalar sclerite; P/R, pleural ridge; Sa, subalare; tg, tegular
rudiment; WW2, base of tegmen, showing dorsal surface.
During extension and flexion the wings do not simply turn forward
and backward on the pleural wing processes, since each wing is at-
tached to the tergum by its entire basal width. The-horizontal move-
ments of the wings are made possible mainly by the flexible lines in
the wing bases and by the articulations of the axillary sclerites on one
another. The working of the parts involved may be easily observed in
a freshly killed specimen if the extended wing is slowly flexed.
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 97
In the fully exended wing of Dissosteira the axillary sclerites lie
approximately flat and in the same plane as the general wing surface
(fig. 47 A, B). When the wing is turned posteriorly, however, the
axillaries take different positions. In the living grasshopper it is
probable that the first movement of flexion is produced by the elas-
ticity of the wing base when the extensor muscles are relaxed, for the
wing of a dead specimen automatically assumes a partly flexed position.
The fully flexed and folded condition, however, undoubtedly depends
on the pull of the flexor muscle (fig. 49, D) on the third axillary.
On the relaxation of the wing, the initial flexing causes the outer
end of the third axillary to turn upward, and the pull of the flexor
muscle brings this sclerite to a vertical position. The movement of
the third axillary turns the attached median plate (m) likewise to a
vertical position on its hinge with the second axillary (24%). In the
forewing (A), the revolution of the first median plate (mm) draws the
second median plate (m’) inward. The second median plate, however,
is firmly attached to the united bases of the median, radial, and sub-
costal veins, and the head of the radius (2) is flexibly attached to the
anterior end of the second axillary. As a consequence, the movement
of the first median plate turns the entire anterior part of the wing
posteriorly on the hinge between the radius and the second axillary.
But, since the basal connection of this part of the wing forms an
oblique line from the head of the first axillary to the articulation
between the two median plates, the entire preanal area of the wing
is deflected as it turns posteriorly. At the same time, the anal area is
lifted but maintains its horizontal plane as the third axillary assumes
a vertical position. When the wing finally comes to a longitudinal
position over the back, therefore, the anal area is uppermost and the
preanal area slants downward on the side (fig. 50 A). During the
final revolution of the wing the first axillary turns upward on its
hinge with the tergum, the second axillary rotates slightly on the
pleural wing process, and the third axillary revolves posteriorly in its
vertical position on the fourth axillary.
In the hind wing the mechanism of flexion is in general the same
as that of the forewing, but, in addition to the posterior turning of
the wing, the great anal area is folded fan-like into many plaits. The
third axillary of the hind wing (fig. 47 B, 3A) is relatively much
larger than that of the forewing (A), its muscle process stands out
prominently from the shaft, and the flexor muscle inserted on it con-
sists of two bundles of fibers. A distal median plate is lacking in
the hind wing, but the single plate (A, m) attached to the third ax-
7
98 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
illary affects the anterior group of veins in the same manner as does
the corresponding plate of the forewing.
When the distal part of the third axillary is lifted by the pull
of the flexor muscle, the median plate turns the preanal area of the
wing posteriorly and toward the body, and at the same time deflects
it to an almost vertical position, with the costal margin downward.
The wing surface makes a double fold along the vena dividens (figs.
45 B, 50 B, VD), and the area between the vena dividens and the
AR VD
Fic. 50.—Positions of the flexed wings of Dissosteira.
A, vertical cross-section through fourth abdominal segment, with wings folded
over body, seen from behind. B, section of right hind wing more enlarged.
IA, 2A, 3A, 7A, first, second, third, and seventh primary anal veins; AF,
anal fold; C, costa; 1Cu, first cubitus; 2Cu, second cubitus; /, intercalary vein;
1, 7, secondary veins of first anal plait; k, first concave anal vein; J, second con-
cave anal vein; M/, media; R, radius; R + M, combined basal shafts of radius
and media; S, sternum; 7, tergum; /D, vena dividens.
first principal anal vein (1A) is folded outward beneath the preanal
area, with the secondary vein k in the ventral angle of the fold (fig.
50 B). While these maneuvers are taking place in the anterior and
middle parts of the wing, the anal fan is bent downward as it comes
against the side of the abdomen, and its ventral surface is turned
outward beneath the deflected preanal area. The membrane of the
fan is plaited between each two of the first seven principal anal veins
(fig. 50 B, rd-7A), with the secondary veins occupying the ventral
lines of the folds. The posterior part of the fan spreads out against
the upper part of the side of the abdomen (A).
e
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 99
A careful study of the forms of the folded wings of the grass-
hopper, as seen in transverse section (fig. 50), will suggest that many
details of structure, both in the tegmina and in the hind wings, are
adaptations to the passive state of flexion rather than to the active
phases of flight.
The extension of the wings is effected probably by the action of
both the basalar and the subalar muscles (fig. 49 E, M’ and M”).
The basalar sclerites (fig. 48, rBa, 2Ba) are connected by a tough
membranous fold (a) with the base of the wing anterior to the wing
process. A depression of these sclerites on their episternal articula-
tions, caused by the contraction of their muscles (fig. 49, E, M’), must
therefore release the flexed wing from its position over the body and
turn it outward. The principal extensor of the wing, however, ap-
pears to be the muscle of the basalar sclerite (fig. 49, M”). In the
flexed wing, the second axillary sclerite is elevated between the first
axillary on the one hand, which now stands in a vertical plane on its
tergal hinge, and the median plate (mm) on the other, which rises ver-
tically from its hinge on the second axillery. The ventral plate of the
second axillary is connected with the subalar sclerite by a thickening
of the intervening membrane (figs. 48, 49, b). The downward pull of
the basalar muscle (fig. 49, M’’) is therefore exerted on the second
axillary. It is easy to demonstrate that a downward pressure on the
second axillary flattens the entire wing base by restoring the first
axillary, the median plate, and the third axillary to, the horizontal
plane, and thereby extends the wing.
When the wings are extended, the mechanism of flight becomes
operative. This includes the direct and indirect muscles, which ac-
complish the movements of levation, depression, and rotation, and
which have already been described.
VI. THE SPIRACELES
The generalized ancestors of modern insects possibly had a pair
of tracheal invaginations on each of the 17 body segments between
the primitive head, or procephalon, and the periproct. Evidence of
the existence of such invaginations has been found, however, on only
14 segments, namely, the second maxillary segment, the three tho-
racic segments, and the first ten abdominal segments.
Tracheal invaginations of the second maxillary segment have been
reported by Nelson (1915) to be present in the embryo of the honey-
bee. They arise, Nelson says, on the lateral surfaces of the anterior
half of the segment above the bases of the rudiments of the second
100 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
maxillae, shortly behind the boundary between the first and second
maxillary segments. The second maxillary spiracles have thus the
same relative position on their segment as have all the body spiracles
in the embryo, or the abdominal spiracles in adult insects. The em-
bryonic invaginations of the labial segment, according to Nelson,
give rise to a part of the tracheal system of the head, but are later
closed and leave no external trace of their existence in the adult insect.
Prothoracic spiracles are known to exist as functional organs of
the adult only in some of the Sminthuridae (Collembola). They are
situated laterally in the neck membrane close to the posterior margin
of the head, but Davies (1927) claims that the region bearing the
spiracles belongs to the prothorax. These cervical or prothoracic
spiracles are the only spiracles present in the Sminthuridae, and no
other collembolan is known to possess either spiracles or tracheae
in any part of the body. Temporary prothoracic spiracles, followed
by the usual series of spiracular invaginations, have been described
in the embryo of Blattella by Cholodkowsky (1891), and in the em-
bryo of Leptinotarsa by Wheeler (1889).
The usual first pair of thoracic spiracles of adult, nymphal,and larval
insects is always situated either in the posterior part of the prothorax
or in the intersegmental membrane between the prothorax and the
mesothorax. In the embryos of most insects, however, these spiracles
are said to lie anteriorly in the mesothorax; they would appear,
therefore, to be the true mesothoracic spiracles which have become
prothoracic in position by a secondary forward migration. The usual
second pair of adult thoracic spiracles are the embryonic meta-
thoracic spiracles, and they sometimes occur on the anterior part
of the metathorax in the adult, though more commonly they lie in
the membrane between the mesothorax and the metathorax, or in
the posterior part of the mesothorax. The segmental relations of the
thoracic spiracles is somewhat complicated by the fact that the muscles
of their closing apparatus have their origins in the segments on which
the spiracles are situated in the adult. Since, however, the musculature
of the thoracic spiracles is not alike in different groups of insects and
is often different in the two spiracles of the same insect, it is probably
of secondary development in all cases.
Contrary to the embryological evidence of the segmental relations
of the spiracles, there are many points in the anatomy of the tracheal
system, and in the innervation of the spiracular muscles, that suggest,
as now claimed by several writers, that the spiracles are primarily
intersegmental invaginations, and that their definitive positions are
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS IOI
the result of migrations either forward or rearward into the segmental
regions of the body.
The abdominal spiracles are situated, with few exceptions, on the
anterior lateral parts of the abdominal segments, where they lie
in the tergal plates, between the terga and the sterna, or in the edges
of the sterna. There are usually eight pairs of abdominal spiracles
in adult and larval insects, though the number may be variously
reduced. There is evidence, however, of more than eight spiracles
having been present on the abdomen of primitive insects. Cholod-
kowsky (1891) reports the existence of a pair of tracheal invaginations
on the first nine abdominal segments of Blattella,and Heymons (1897)
says there are apparent rudiments of spiracles on the tenth abdominal
segment of Lepisma. In certain insects the spiracles of the first
abdominal pair are situated very close to the base of the metathorax,
and long discussions recur as to whether these spiracles belong to
the thorax or to the abdomen. In all cases, however, it will be found
that the spiracles in question lie posterior to the third phragma, which
marks the intersegmental line between the metathorax and the first
abdominal segment, or behind the lateral extensions of the postnotal
plate in the metathorax. The spiracles are therefore abdominal, as
is shown also by the destination of their tracheae.
The external aperture of a spiracle may be a simple opening leading
directly from the exterior into the trachea. In most cases, however,
there is a pre-tracheal chamber, or atrium (fig. 53 A, B, Atr), formed
by an inflection of the body wall, from the inner end of which arises
the trachea (Tra). The atrium of the spiracle, therefore, appears
to be a secondary invagination of the body wall, which has carried
the mouth of the original tracheal invagination to a more protected
position beneath the surface. In some cases the edges of the atrial
orifice are elevated to form a pair of protruding lips guarding the
entrance (fig. 53 A, c, d), in others the opening is fringed with
opposing brushes of hairs, usually thickly branched, or it is itself
reduced to a very small diameter.
Spiracles are usually provided with a closing apparatus. In the
Apterygota the spiracles are said to lack an occlusor mechanism
(Du Buisson, 1926; Davies, 1927), and the thoracic spiracles of
Plecoptera are simple apertures giving open passages into the tracheae.
In general, however, the spiracles have either a device for closing
the outer lips of the atrial chamber, or an apparatus for blocking the
passage from the atrium into the tracheae. The occlusor mechanism
of the abdominal spiracles is of the second type; that of the thoracic
102 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
spiracles may be of similar structure (caterpillars and other larvae),
but usually the closing apparatus of the thoracic spiracles in adult
insects effects a movement of one or both of the outer lips of the
atrial chamber.
The Acrididae possess the two usual pairs of thoracic spiracles and
eight pairs of abdominal spiracles. The first thoracic spiracle on
each side is situated laterally in the ample intersegmental membrane
between the prothorax and the mesothorax (fig. 20 B, 26, Sp.) where
it is covered externally by the lateral part of the large posterior fold
of the protergum (fig. 20 B, Rd). The second spiracle lies in the
posterior ventral angle of the mesothoracic epimeron just above the
base of the middle leg and immediately before the intersegmental
groove between the mesopleuron and the metapleuron (fig. 26, Sps3).
The abdominal spiracles are carried by the first eight abdominal terga,
each being placed in the lower anterior angle of the corresponding
tergal plate. The first of the series, therefore, lies in the tympanal
cavity of the first segment, where it is situated on the small triangular
area before the tympanal membrane and just in front of the support
of the chordotonal organ. All the spiracles are well developed, and
each is provided with an efficient closing apparatus, the mechanism of
which presents the usual two types of structure, the first pertaining
to the thoracic spiracles, the second to the abdominal spiracles. The
details of structure, however, are quite different between the two
thoracic spiracles.
The first thoracic spiracle-—The first spiracle of the thorax of Dis-
sosteira carolina is contained in a small, irregular plate, or peritreme
(fig. 51 A, Ptr), lying laterally in the intersegmental membrane
between the prothorax and mesothorax (fig. 20 B, Sp2), covered
externally by the overlapping fold of the protergum. The lower end
of the peritreme is produced posteriorly and upward in a small, free
process (fig. 51 A, a), bearing on its base a flat-topped, pale-colored
tubercle (b) projecting outward. The tubercle is a little higher than
the lips of the spiracle and evidently serves as a stop to prevent the
covering flap of the protergum from resting too closely against the
spiracle. The spiracular opening is an obliquely vertical slit with a
slight italic curve and strongly protruding anterior and posterior
lips (c, d). The length of the slit is about 0.60 mm. in the male
grasshopper, and about 0.75 mm. in the female. The anterior lip (c)
is a rigid elevation of the wall of the peritreme; its inner face is
soft and deeply grooved parallel with the outer edge. The posterior
lip (d) is a weaker and freely-movable flap, but it has a sharp,
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 103
strongly-sclerotized marginal band (¢) which, when the spiracle is
closed, fits into the groove of the anterior lip.
The cleft of the first spiracle opens into a shallow atrium from
which are given off two tracheae, a larger dorsal one (fig. 51 B, f)
and a smaller ventral one (g). From without, therefore, the first
spiracle appears to have a double opening (A, f, g,). In some of the
Orthoptera that have tympanal organs on the front legs, the trachea
of the ventral (or posterior) opening of the spiracle appears to have
become specialized as an “ acoustic” trachea since it goes only to the
Fic. 51.—First thoracic spiracle of Dissosteira.
A, outer view of left spiracle. B, inner view of right spiracle. a, ventral lobe
of peritreme; b, process of peritreme protecting spiracle from covering flap of
pronotum; c, anterior lip of spiracle; d, posterior lip of spiracle; e, hard edge
of posterior lip; f, dorsal trachea; g, ventral trachea; h, internal lever of pos-
terior lip forming a septum between the tracheae; 7, head of lever on which
closing muscle (79) is inserted; /, ventral internal process of peritreme on
which spiracular muscles arise; m, external pit forming internal process 1; Ptr,
peritreme ; 79, opening muscle; 80, closing muscle.
front leg, where it branches into the two tracheae of the tympanal
organ. This fact led Graber to the conclusion that the double structure
of the first spiracle in Orthoptera originated from the separation of
an “‘acoustic”” trachea from the general respiratory tracheae of the
prothorax. Carpentier (1924, 1925), however, has shown that the
double first spiracle is a character of Orthoptera in general, whether
tympanal organs are present in the front legs or not, and that in most
forms the tympanal trachea is not isolated from the rest of the
respiratory system. The specialization, he says, is carried to its highest
degree in the tettigoniid Phasgoneura viridissima, where the spirac-
ular orifice of the leg trachea is enormously enlarged. Here, ap-
104 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
parently, is a case of an advantage derived by a specific organ from
a general structure first developed for some other reason.
In the septum between the two spiracular openings in Dissosteira
(fig. 51 A, h) is a strong internal bar (B, h) projecting anteriorly
and ventrally from the posterior lip of the spiracle, and terminating
in a free process (7) that extends anterior to the spiracular opening.
Upon this process is inserted a short muscle (79) which has its origin
ventrally on an inner process (/) of the lower angle of the peritreme,
the site of which is marked externally by a pit (A, m). A second
muscle (B, So) arises from the base of the same process (/) and
extends. dorsally and posteriorly to its insertion on the base of the
posterior lip of the spiracle behind the ventral trachea. The first
muscle (79) is the occlusor of the spiracle; the second (80) is evi-
dently its antagonist. A downward pull on the head of the septal arm
(i), where the anterior muscle is inserted, closes the spiracle by
rotating the movable posterior lip forward on its dorsoventral axis
and bringing thus its sharp free edge into the groove of the anterior
lip. Conversely, a downward pressure on the base of the posterior
lip, at the point where the posterior muscle (80) is inserted, rotates
the lip in the reverse direction and opens the spiracle. The differential
action of the two muscles results from the opposition of their two
points of insertion on either side of the long axis of the posterior
lip, and is accentuated by the difference in their points of origin
on the ventral process (/) of the peritreme. Vinal (1919), Lee
(1925), and other writers have regarded both muscles of the first
spiracle in Acrididae as occlusors.
The second thoracic spiracle—The second thoracic, or metatho-
racic, spiracle of Dissosteira is located in the lower, posterior angle
of the mesothoracic epimeron of the adult (fig. 26, Sp3), where it
is surrounded by a narrow membranous area (fig. 52 A, mb).
Externally this spiracle presents two thick, elongate oval, valve-like
lips, (fig. 52 A, c, d) separated by a sinuous vertical cleft having a
length of about 0.50 mm. in the male insect. Both lips of the second
spiracle are movable, though they are united ventrally in a broad lobe
(n). The spiracular lips stand out prominently from the body wall
(fig. 53, A, c, d), and between them is a shallow atrium (4ér) from
which arises a single large trachea (Tra) that soon divides into a
dorsal and a ventral branch. The closing mechanism of the second
spiracle includes but a single short occlusor muscle (fig. 52 B, ELD) e
The muscle arises ventrally from a small process (0) on the posterior
dorsal margin of the mesocoxal cavity, and is inserted on the ventral
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 105
lobe (7) of the spiracle. There is no special device for opening this
spiracle ; the lips diverge by their own elasticity, as is shown by the
fact that the spiracle is always open in a dead insect. The occlusor
muscle brings the edges of both lips together.
The abdominal spiracles—tThe eight spiracles of the abdomen in
Dissosteira are quite different from either of the thoracic spiracles.
They are not provided with projecting external lips (fig. 53 B),
the body wall being directly inflected in each spiracle to form an open
atrial chamber (Atr). The atrium leads by a narrowed aperture at
Fic. 52.—Second thoracic spiracle of Dissosteira.
A, outer view of left spiracle. B, inner view of right spiracle. c, anterior lip
of spiracle; d, posterior lip of spiracle; Epme, epimeron of mesothorax; Eps:,
episternum of metathorax; /sg, intersegmental fold; mb, membrane surrounding
spiracle; n, ventral lobe of spiracle uniting the lips and giving insertion to
spiracular muscle (z1rr); 0, internal lobe on rim of coxal cavity on which
spiracular muscle arises; 7, internal intersegmental fold; Tyra, trachea; IZ1,
closing muscle of spiracle.
its inner end into the spiracular trachea (Tra), and the occlusor
mechanism regulates this opening.
The longer axis of the first abdominal spiracle is obliquely hori-
zontal (fig. 54 A) with the anterior end a little higher than the
posterior. The other spiracles (C, D) are placed more nearly vertical,
so that the dorsal end of each corresponds with the anterior end of
the first spiracle. In each spiracle one wall of the atrium is rigid
(fig. 53 B, t), and the other (s) is movable. The rigid wall is dorsal
in the first spiracle (fig. 54 A, t) and posterior in the other spiracles
(C, D). It is strengthened by a thickening in the external body wall
(figs. 53 B, 54 A, #) from which it is inflected. The movable wall
of the atrium (s), which is ventral in the first spiracle (fig. 54 A)
106 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
and anterior in the others (C, D), is flexible because the body wall
immediately external to it 1s weak, and because the two end walls
of the atrial chamber are membranous. The posterior or ventral end
of the movable wall is produced into a long, free manubrium (fig.
54 B, D, q) that projects into the body cavity and gives attachment
to two muscles, one dorsally, the other ventrally. These muscles,
acting antagonistically, either close or open the passage from the
atrium into the trachea (fig. 53 B) by means of their attachments
on the movable wall of the atrium.
The short dorsal muscle of the first abdominal spiracle (fig. 54 B,
CMcl) arises on the rim of the tympanum (/) above the spiracle;
?
Fic. 53.—Sections of spiracles of Dissosteira.
A, longitudinal section through second thoracic spiracle, showing anterior and
posterior lips (c, d) as projecting folds of body wall (BW) inclosing an atrium
(Atr), or entrance to trachea (Tra).
B, vertical section through first abdominal spiracle, showing direct inflection
of body wall to form atrial chamber (A?r), of which anterior wall (s) is moy-
able, and posterior wall (¢) immovable.
Atr, atrium; BW, body wall; c, anterior lip of spiracle; d, posterior lip of
spiracle; mb, membrane surrounding lips of spiracle; g, manubrium or muscle
process of ventral wall of atrium; 7, intersegmental fold; s, ventral wall of
atrium; ¢t, dorsal wall of atrium; 7a, trachea; wu, plate in tergal wall support-
ing dorsal wall of atrium.
the long, slender ventral muscle (OMcl) arises ventrally on an in-
flection of the integument mesad of the hind coxa and posterior to the
triangular coxal plate of the metasternum (fig. 30 A, ¢). It is easy
to demonstrate that the dorsal muscle (fig. 54 B, CMcl) is the closer
of the spiracle and the ventral one (Omcl) the opener. A dorsal pull
upon the manubrium (q) of the movable ventral wall of the atrium
(s) brings the inner edge of the latter against the inner edge of the
fixed dorsal wall (¢) and thus closes the passage from the atrium
into the spiracular trachea. By a counter movement the passage
is opened.
The mechanism of the other abdominal spiracles is the same as that
of the first. The short, fan-shaped occlusor muscle (fig. 54 C, D,
NO. 2 THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS 107
CMcl) arises on the wall of the tergum immediately behind the
spiracle, and is inserted on the manubrium of the movable wall of
the atrium (gq), which projects ventrally and posteriorly. The long
opening muscle (OMcl) arises on the lateral edge of the correspond-
ing sternum and extends posteriorly and dorsally to its insertion on
the manubrium.
Fic. 54.—Abdominal spiracles of Dissosteira.
A, first abdominal spiracle, left, outer view. B, inner view of the same spiracle,
showing muscles. C, second abdominal spiracle, right, inner view, with tergo-
sternal muscles of second abdominal segment. D, eighth abdominal spiracle,
right, inner view.
CMcl, closing muscle of spiracle; Sp, first abdominal spiracle; J/.S, second
abdominal sternum; J/Sp, second abdominal spiracle; OMcL, opening muscle
of spiracle; p, anterior margin of tympanal cavity; g, manubrium of ventral or
anterior wall of atrial chamber; s, movable ventral or anterior wall of atrial
chamber; ¢, fixed dorsal or posterior wall of atrial chamber; u, thickening of
tergal wall supporting dorsal or anterior wall of atrial chamber; wv, anterior
apodemal arm of abdominal sternum; IIS, eighth abdominal sternum; V JJ] Sp,
eighth abdominal spiracle; w, posterior angle of tympanal cavity.
The grasshoppers are abdominal breathers. A discussion of the
mechanism of respiration would, therefore, lead too far beyond
the anatomical limits of the present paper. Recent studies on the
breathing of Orthoptera give such varied and conflicting results that
we must conclude either that the subject still needs a critical investi-
gation or that the insects have no fixed methods of respiration.
The weight of evidence is rather in favor of inconsistancy on the
part of the insects.
108
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOL. 82
ABBREVIATIONS USED ON THE FIGURES
A, Anal veins. 1A, 2A, etc., first anal,
second anal, etc.
Ac, antecosta.
acs, antecostal suture.
Acx, precoxal bridge.
AF, anal fold.
ANP, anterior notal wing process.
Ap, apodeme.
Ar, arolium.
Atr, atrium.
Aw, prealar arm of tergum.
Ax, axillary sclerite. 14x, 2Ax, 34x,
4Ax, first, second, third, and
fourth axillaries.
AxC, axillary cord.
Ba, basalare.
Bc, basicosta, basal ridge of coxa.
bcs, basicostal suture of coxa.
Bcx, basicoxite.
Brn, branchia, gill.
Bs, basisternum.
BW, body wall.
CMcl, closing muscle of spiracle.
cpl, supra-coxal plate of subcoxa.
Cu, cubitus. 1Cu, 2Cu, first and second
cubitus.
Cv, cervix, neck.
Icv, 2cv, first and second lateral cer-
vical sclerites.
Cx, coxa.
CxC, coxal cavity.
CxP, pleural coxal process.
Cxpd, coxopodite.
cxs, coxal suture.
D, flexor muscle of wing.
DMcl, dorsal longitudinal muscles.
E, basalar-sternal muscle.
Em, lateral emargination of tergum.
Endp, endopodite.
Epm, epimeron.
Eppd, epipodite.
Eps, episternum.
Eupl, eupleuron.
Eutn, entrochantin.
Expd, exopodite.
Fim, femur.
Fu, furca.
H, head.
/, intercalary vein.
tergal promotor muscle of coxa.
/-XI, abdominal segments.
[S-XIS, abdominal sterna.
Isg, intersegmental fold.
ISp, IIS p, first and second abdominal
spiracles.
Ist, intersternite.
IT-XIT, abdominal terga.
J, tergal remotor muscle of coxa.
Ju, jugal area of wing.
K, sternal promotor, anterior rotator
of coxa.
[legs
sternal remotor, posterior rotator
of coxa.
LB, leg basis.
Ls, laterosternite.
M, media.
abductor muscle of coxa.
m, m’, distal median plates.
M', basalar-coxal muscle.
M”, subalar-coxal muscle.
Ib, secondary intersegmental mem-
brane.
mb, membrane
Mer, meron.
N, N’, adductor muscle of coxa.
O, levator muscle of trochanter.
OMcl, opening muscle of spiracle.
P, body branch of depressor muscle of
trochanter.
Pak, parapsidal ridge.
pas, parapsidal suture.
Pc, precosta.
NO. 2
Pcx, precoxal bridge.
Ph, phragma. rPh, 2Ph, 3Ph, first,
second, and third phragmata.
PI, pleuron.
pl, pleural sclerites between tergum
and subcoxa.
PIA, pleural apophysis.
Pln, planta.
PIR, pleural ridge.
PIS, pleural suture.
PN, postnotum.
PNP, posterior notal wing process.
Poc, postocciput.
PoR, postoccipital ridge.
Ppct, prepectus.
Ppt, periproct.
PR, prescutal ridge.
Pra, preanal area of wing.
Ps, presternum.
ps, prescutal suture.
Psc, prescutum.
PT, posterior arm of tentorium.
Ptar, pretarsus.
Ptr, peritreme.
Pw, postalar arm of postnotum.
QO, coxal branch of depressor muscle
of trochanter.
R, radius.
Rd, posterior fold, reduplication, of
tergum.
S, levator muscle of tibia
sternum.
SA, sternal apophysis.
Sa, subalare.
sa, external pit of sternal apophysis.
Sc, subcosta.
Scl, scutellum.
scl, subdivision of scutellum.
THORACIC MECHANISM OF A GRASSHOPPER—SNODGRASS
109
Sct, scutum.
sct, subdivision of scutum.
Scx, subcoxa.
ST, sternellum.
Sp, spiracle. Sp, Sps, first and second
thoracic spiracles.
Spn, spina.
spn, external pit of spina.
Ss, spinasternum.
Stn, primary segmental sternite.
T, depressor muscle of tibia.
tergum.
Tar, tarsus.
Tb, tibia.
tg, tegular rudiment.
TmMcl, tympanal muscle.
Tn, trochantin.
Tr, trochanter. r7r, 277, first and
second trochanters.
Tra, trachea.
U, levator muscle of tarsus.
Uf, unguifer of tarsus.
Un, unguis, claw.
Utr, unguitractor plate.
l’, depressor muscle of tarsus.
VD, vena dividens.
VIIISp, eighth abdominal spiracle.
VMcl, ventral longitudinal muscle.
VR, ridge between scutum and scu-
tellum.
vs, scuto-scutellar suture.
W, wing.
WP, pleural wing process.
X, depressor muscle of pretarsus, re-
tractor of claws.
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SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 82, NUMBER 3
Hodgkins Fund
THE RADIATION OF THE PLANET
BARTH TO SPACE
(WitTH Two PLATEs)
BY
C. G. ABBOT
(PUBLICATION 3028)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
NOVEMBER 16, 1929
The Lord Baltimore Preas
BALTIMORE, MD., U. 3. A.
bovgkins Fund
THE RADIATION OF THE PLANET EARTH TO SPACE
By C. G. ABBOT
(Wira Two Prates)
In an illuminating series of papers, G. C. Simpson recently has
approached the subject of terrestrial and atmospheric radiation to
outer space. The first of these papers is entitled “Some Studies in
Terrestrial Radiation.” * Here Simpson makes the unsatisfactory
assumption that the atmospheric water vapor behaves like a “ grey
body ” in absorbing radiation. That is, he assumes that general coefh-
cients of absorption and of transmission may be employed, without
regard to the wave length of the radiation considered. Arriving in
this way at unexpected and questionable results, Simpson then modi-
fied his procedure in a second paper entitled “ Further Studies in Ter-
restrial Radiation.” * Here he makes the following important assump-
tions: (a) The stratosphere contains 0.3 mm. of precipitable water.
(b) The absorptive properties of atmospheric water vapor may be
regarded as so similar to those of steam that Hettner’s * observations
of the absorption of a layer of steam may be taken as representing the
coefficients of absorption of atmospheric water vapor between wave
lengths 4» and 34.’ (c) “The stratosphere absorbs all radiation
between wave lengths 54 and 7p, and from wave length 14, to the
end of the spectrum.”
As the Smithsonian Institution has hitherto published considerable
evidence relating to these three subjects, it has occurred to me to see
whether the use of our independently derived data would check well the
‘
* Mem. Roy. Meteorol. Soc., Vol. 2, No. 16, 1928.
* Ibid., Vol. 3, No. 21, 1928.
*Hettner, G., Ann. Physik. Leipzig, 4th Folge., Band 55, p. 476, 1918.
“Simpson nevertheless calls attention to the incomplete similarity between
the absorption of concentrated and unconcentrated vapors, and therefore cor-
rects Hettner’s curve between 8 and t1u from other data derived from atmos-
pheric experiments.
SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 82, No. 3.
2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
results of Simpson in this important field. Fowle’ carried on for
several years, 1908 to 1917, experiments on the absorption of radiation
of long wave lengths by the atmosphere contained in tubes of large
diameter and up to 800 ft. in length. These tubes were laden with
water vapor ranging from 0.2 up to 2.5 mm. of precipitable water,
and of carbon dioxide content ranging from 7 grams up to 160 grams
per meter cross-section at normal temperature and pressure.
In his early experiments, Fowle had established means for deter-
mining the quantity of precipitable water in atmospheric air by means
of measurements on the bands por, ¢, and w of the upper infra-red
solar spectrum. These experiments are fortunately very definite as
to the determination of water vapor equivalent to 0.3 mm. of pre-
cipitable water.
In the summer of the years 1909 and 1910, Abbot observed the
infra-red solar spectrum from Mount Whitney, California, altitude
4,420 m. Bolographs of the spectrum were obtained, having very
satisfactory quality as far as the delineation of the bands por, ¢, and y
is concerned.” From these, Fowle determined the quantity of total
precipitable water in a vertical path of atmosphere above Mount
Whitney. On August 14, 1910, he observed 0.6 mm. Considering the
moderate altitude and the summer season, this small observed water-
vapor content hardly prepares one to accept Simpson’s assumption
that the stratosphere, which begins at 12,000 m., and is at a tempera-
ture about 50° C. lower than that which prevailed at the summit of
Mount Whitney on that occasion, can contain half of the precipitable
water above that station. We have other evidence leading to the same
view.
At Mount Montezuma, Chile, altitude 2,710 m., we have observed
spectroscopically the total precipitable water in a vertical column
above the station almost daily for about 10 years at all seasons. The
following table gives average values for the 12 months, and also
extreme values for each of these months, together with associated
surface temperatures.
In illustration of the great alterations in the appearance of the solar
energy spectrum depending on the quantity of atmospheric humidity,
we give reproductions of two days’ observations at Montezuma,
plates 1 and 2. Note the bands por, ¢, y and Q.
*Fowle, F. E., Ann. Astrophys. Observ., Vol. 4, pp. 274-286. Astrophys.
Journ., Vol. 38, p. 303, 1913; Vol. 42, p. 304, 1915. Smithsonian Misc. Coll.,
Vol. 68, No. 8, 1917.
* See Annals, Vol. 4, fig. 50.
ise)
ABBOT
EARTH TO SPACE
RADIATION OF
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4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
From the tabular data, it is clear that values of total precipitable
water are frequently observed at Montezuma closely approaching the
value assumed ,by Simpson for the stratosphere. These values are
found in winter, with a surface air temperature of +9° C., on the
edge of the tropics at 2,710 m. altitude.
In view of these observations at Montezuma, and considering the
rapid decrease of humidity with temperature (the vapor pressure at
—50° and o° C. being respectively 0.03 and 4.58 mm.) and also the
fact that three-fourths of the superincumbent atmosphere lies’ between
Montezuma station and the bottom of the stratosphere, one is forced
to conclude that the value of the precipitable water contained by the
stratosphere is vanishingly small, rather than 0.3 mm. as assumed by
Simpson. This materially affects his argument, especially that part
which relates to cloudy skies.
As an independent approach, instead of Simpson’s two other basic
assumptions, which we have designated as (b) and (c), we have em-
ployed Fowle’s two summaries of the results obtained in his long-tube
experiments. To make these results of Fowle’s applicable to the
problem of atmospheric radiation and absorption, as set by Simpson,
we have prepared a large scale plot, reproduced in reduced size in
figure 1. From this plot we take table 2. In choosing the quantities
of precipitable water to be used, we have doubled the values given by
Simpson for successive layers in the table he designates as “ Fig. I,”
page 72, of his paper “‘ Some Studies in Terrestrial Radiation.” This
doubling we do because of the following consideration. .
We are proposing to ascertain the radiation which certain layers of
the free atmosphere, containing natural loads of water vapor and car-
bon dioxide, will send upwards in all directions within a solid angle
filling a complete hemisphere. We assume, as does Simpson, that for
monochromatic rays the emission of such a layer bears the same pro-
portion to the emission of the perfect radiator that the absorption of
the layer in question bears to unity. While some rays are emitted ver-
tically, most rays are emitted obliquely, so that the average emission
and absorption of a layer exceeds that which corresponds to the pre-
cipitable water vapor and carbon dioxide found in a vertical path. It
is readily proved by performing the integration over a complete
hemisphere that the average upward path is double the vertical one.
Hence we have doubled Simpson’s figures for the precipitable water
contained in the layers he has chosen. These data appear in table 2.
*See Annals, Vol. 4, Table 102, p. 286; also Smithsonian Physical Tables,
7th Rey. Ed., 4th reprint, p. 308.
RADIATION OF EARTH TO SPACE—ABBOT
Say Seas or 2
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ion of Radiation by Air Laden
m
6 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
As a second step, we consider the spectral distribution and intensity
of emission of the perfect radiator at different temperatures.’ By
interpolation on large scale plots we have prepared table 3. This gives
the approximate ° intensity of emission of the perfect radiator at tem-
peratures corresponding to the mean temperatures of Simpson’s lay-
ers, and to those of his selected latitudes of the earth’s surface.
Multiplying the values in table 2 by corresponding ones in table 3,
we obtain the emission of radiation outwards from each Simpson at-
mospheric layer towards a complete hemisphere. The values are given
in table 4.
Again interpolating in the plots (fig. 3) we next obtained the trans-
mission coefficients for each superincumbent atmospheric mass lying
above the respective Simpson layers. Allowance is made for the ozone
absorption between Qu and I1p. These values are given in table 5.
Multiplying these values by corresponding ones in tables 3 and 4,
we obtained the contributions of the Simpson atmospheric layers and
also of the earth’s surface * at the latitudes go°, 70°, 60°, 50°, 40°,
and o° to the intensity of emission of the earth as a planet towards
outer space.’ These results are given in table 6.
All of these results apply to cloudless skies. We now assume, with
Simpson, that the earth is 50 per cent cloudy; that the clouds totally
absorb all radiation arising from beneath them ; that they radiate quite
as efficiently as the perfect radiator; and that their upper surfaces
maintain the same average temperature as the earth at 70° latitude.
Weare not able to compute their radiation in Simpson’s manner, since
we have shown reasons to believe that the stratosphere is almost desti-
tute of water vapor, instead of containing 0.3 mm. of precipitable
water as he supposes. We simply assume that the combined emission
of clouds and atmosphere during one-half the time at all latitudes is
the same as that of the earth’s surface and the superincumbent atmos-
phere at latitude 70°. That is: For the atmosphere 0.151 cal. per
cm. per min. ; for the cloud surface 0.100 cal., giving a total for com-
pletely overcast sky of 0.251 cal. During the other half of the time,
*See Smithsonian Physical Tables, p. 248.
*We do not guarantee these values to within 2 per cent.
*We assume, with Simpson, that the earth’s surface may be regarded as a
perfect radiator.
* Notwithstanding our previous evidence that the water-vapor content of the
stratosphere is vanishingly small, we have thought best to estimate 30 per
cent of black-body efficiency as applicable to the stratospheric radiation in the
wave-length region 13” to 504, where water vapor is so very active. We have
allowed 16 per cent of black body efficiency to the ozone band, ou to 11m.
OF EARTH TO SPACE—ABBOT
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I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
the values at different latitudes are as given in table 6 for clear skies.
For half-cloudy skies we take the mean of the two conditions.
We are now prepared to assemble our results and compare them
with those of Simpson (table 7).
It is clear that our employment to a considerable extent of inde-
pendent data and methods has made no very great difference in the
totals from those of Simpson. The range of our totals for half-
cloudy sky is indeed considerably greater than his as between the
equator and the poles. Our method has enabled us to segregate the
contributions of the atmosphere and of the earth’s surface, which
in Simpson’s second paper are not computed separately. We find the
earth’s surface almost equally contributing at all latitudes, but the
TABLE 7.—Radiation of Earth and Atmosphere to Space
Calories per cm.” per min.
Smithsonian results Simpson results
Latitude Clear sky Half-cloudy sky Atmosphere plus surface
Atmos-} Sur- Atmos- | Surface Over- | Half-
phere face Total phere jor cloud Total | Clear cast | cloudy
ORS aiseeee 0.220 | 0.105 | 0.325 | 0.186 | 0.102 | 0.288 | 0.316 | 0.213 | 0.264
AO": 0.192 | 0.107 | 0.299 | 0.171 | 0.103 | 0.274 | 0.307 | 0.243 | 0.275
BO ne. eee 0.182 | 0.105 | 0.287 | 0.166 | 0.102 | 0.268 | 0.291 | 0.249 | 0.270
GOo teres 0.162 | 0.104 | 0.266 | 0.156 | 0.102 | 0.258 | 0.274 | 0.252 | 0.265
[Ae seeks heecyt 0.151 | 0.100 | 0.251 | 0.151 | 0.100 | 0.251 | 0.253 | 0.253 | 0.253
QOS ae 0.129 | 0.096 | 0.225 | 0.140 | 0.098 | 0.238 0.245
atmosphere, which contributes much more than half the total (even
more than two-thirds the total on cloudless days at the equator) emits
very much lesser proportions as we approach the poles. The two
sources are very different as regards wave lengths of principal contri-
bution ; the atmosphere emitting mostly in the region exceeding I6p
in wave length, the surface emitting principally in the region Op to I3u.
If we sum up the results in the seventh and tenth columns, which
represent our own and Simpson’s totals for half-cloudy sky, and as-
sign weights to them proportional to the areas of earth which they
respectively represent, we find that the earth as a planet radiates aver-
ages of 0.277 or 0.265 cal. per square centimeter per minute according
as our results or Simpson’s are taken. If we compute the same quan-
tity from the solar constant, 1.94 cal., and Aldrich’s albedo, 43 per
Ray ; :
cent, the result aa <0.57=0.276 cal. The discrepancies are very
small and far within the probable error of the determinations.
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SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 82, NUMBER 4
THE CHARACTERS OF THE GENUS
GEOCAPROMYS CHAPMAN
(WitTH ONE PLATE)
BY
GERRIT S. MILLER, JR
Curator, Division of Mammals, U.S. National Museum
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(PUBLICATION 3029)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
DECEMBER 9, 1929
The Lord Baltimore Press
BALTIMORE, MD., U. 8. A.
THE CHARACTERS OF THE GENUS GEOCAPROMYS
CHAPMAN
By GERRIT S. MILLER, JR.
CURATOR, DIVISION OF MAMMALS, U. S. NATIONAL MUSEUM
(WitH ONE PLATE)
In his “ Revision of the Genus Capromys”’ (Bull. Amer. Mus. Nat.
Hist., Vol. 14, pp. 313-323, Nov. 12, 1901) Mr. Frank M. Chapman
established a sub-genus Geocapromys (p. 314) to include Capromys
brown J. B. Fischer, C. thoracatus True and C. ingrahami J. A. Allen,
animals that were supposed to have skulls and teeth essentially like
those of the species of true Capromys, but to have unusually short
tails and poorly developed thumbs. Sixteen years later Dr. Glover M.
Allen raised Geocapromys to generic rank and added to its characters
the presence of a small supplemental reentrant angle near the front
of the lingual side of the first mandibular molariform tooth (Bull.
Mus. Comp. Zool. Vol. 61, p. 9, Jan., 1917). In 1919 Mr. H. E.
Anthony noticed that the course of the upper incisor of Geocapromys
is clearly shown on the face of the maxillary as a prominent swelling
on the wall of the antorbital foramen, while in Capromys no such
swelling is present (Bull. Amer. Mus. Nat. Hist., Vol. 41, p. 631,
Dec. 30, 1919). In his 1917 paper Dr. Allen, misled by Chapman’s
imperfect specimens of Geocapromys columbianus, made his own
better material of the Cuban animal the basis of the new name G.
cubanus (p. 9), and proposed (p. 5) the generic name Synodontomys
for the original C. columbianus. These errors he later recognized and
corrected (Bull. Mus. Comp. :Zool., Vol. 62, p. 145, May, 1918).
When preparing the copy for my “ List of North American Recent
Mammals 1923” I concluded that the dental features pointed out
by Allen and Anthony did not warrant the generic separation of the
group from Capromys. Not knowing of any other characters I
relegated Geocapromys to subgeneric rank again. More recently, while
examining broken skulls from caves in Cuba, I found that there are
important and constantly present features of both skull structure and
tooth arrangement that fully justify the generic separation of the
‘two groups. The diagnostic characters may be tabulated as follows:
* SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 82, No. 4
2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
Preorbital bar of maxillary sloping obviously forward; root capsule
of upper incisor terminating in contact with outer half of an-
terior border of alveolus of pm*; bases of alveoli of right and
left pm* separate, not encroaching on floor of narial passage ;
pm: with only two reentrant angles on lingual side.............. Capromys
Preorbital bar of maxillary vertical or sloping slightly backward;
root capsule of upper incisor terminating above and ectad to
anterior half of outer border of alveolus of pm‘; bases of alveoli
of right and left pm* in contact, encroaching on floor of narial
passage; pm, with a small third reentrant angle on lingual
Side psoas Rhtes Soke era edi eked east epee Ra ea RY 3 ate eels oar Geocapromys
REMARKS ON GEOCAPROMYS
Skull—The ascending branch of the maxillary dividing the orbit
from the antorbital foramen is vertical (G. ingraham) or backward-
sloping (G. brownii and G. thoracatus) in relation to alveolar line
instead of conspicuously forward-sloping as in Capromys (pl. I, figs. I
and 2). By this character alone any one of the three living species
can be distinguished from any of the four living Capromys. (I have
not seen a specimen of the extinct G. columbianus in which the as-
cending branch is preserved). The backward slope in Geocapromys
is never so strong as the forward slope in Capromys, but the difference
is obvious when the general direction of the ascending branch is com-
pared with the line of the alveolar margin.
Teeth—Root of upper incisor encapsuled in the lower half of the
maxillary wall of the antorbital foramen (see pl. 1, fig 1), the dis-
tance between the outer surfaces of the very obvious incisor capsules
of opposite sides greater than that between the outer sides of the
basal capsules of the opposite first molars. In Capromys the root of
the incisor terminates opposite the antero-inner edge of the lower
lip of the antorbital foramen (pl. I, fig. 2), and the transverse diam-
eter of the rostrum through the scarcely evident capsules is less
than that through the bases of the first molars. The base of pm‘,
which is hidden by the incisor capsule in Geocapromys, often forms
an obvious external swelling in Capromys (as in pl. 1, fig. 2).
These characters indicate that the members of the two genera have
been developing along consistently different lines. In Capromys the
incisor root has pushed back to a position where more advance is
prevented by contact with the base of pm‘; in Geocapromys its posi-
tion is such that it could be extended much farther back in a capsule
lying along the outer surface of the molar shafts as in Spalacopus.:
The Capromys condition is nearly paralleled in Octodontomys. In
correlation with the position of the incisor roots the molar roots are
NO. 4 THE GENUS GEOCAPROMYS CHAPMAN—MILLER 3
farther apart in Capromys than in Geocapromys. This character is
not visible in complete skulls, but is evident in the broken-away palates
so often found in caves. The upper surface of such a fragment of
the maxillary (lower floor of nares) in the region between the anterior
zygomatic roots is traversed by a deep median sulcus in Capromys
occupying the space between the rather widely separated bases of the
opposite premolars ; in Geocapromys there is no median sulcus between
the premolars, but the maxillary rises as a broad flat plate to the
level of the connate bases of these teeth. Immediately behind this
level the groove begins, passing backward to the posterior nares
between the progressively more separated roots of the molars.
The genus Geocapromys contains four species—the living G. brownii
(Fischer) of Jamaica, G. thoracatus (True) of Little Swan Island,
Gulf of Honduras, G. ingrahami (Allen) of Plana Keys, Bahamas,
and the extinct though geologically Recent G. columbianus (Chap-
man) of Cuba (with its synonym G. cubanus G. M. Allen).
EXPLANATION OF PLATE
All figures natural size
Fic. 1. Geocapromys brownti (Fischer). Adult female. No. 143851, U. S. Nat.
Mus. Jamaica. i= base of incisor capsule.
Fic. 1a. Geocapromys browniit (Fischer). Adult male. No. 141908, U. S. Nat.
Mus. Jamaica. Palate cut away from skull and viewed from above.
pm = base of premolar, i= base of incisor capsule.
Fic. 2. Capromys pilorides Desmarest. Small individual, No. 103884, U. S. Nat.
Mus. Cuba. pm —=capsule at base of premolar, i= base of incisor capsule.
Fic. 2a. Capromys pilorides Desmarest. Large individual. No. 253232, U. S.
Nat. Mus. Palate cut away from skull and viewed from above. pm = base
of premolar, i= base of incisor capsule.
Fic. 2b. Capromys sp. No. 254679, U. S. Nat. Mus. Cuba (cave deposit). Palate
cut away from skull and viewed from above. pm = base of premolar, 1 = base
of incisor capsule.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLE S2 NO 4 on Pleeed
1. Geocapromys.
2. Capromys.
(All figures natural size)
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 82, NUMBER 5
MAMMALS EATEN BY INDIANS, OWLS, AND
SPANIARDS IN THE COAST REGION OF
THE DOMINICAN REPUBLIC
(WiTH Two PtatTEs)
BY
GERRIT S. MILLER, JR.
Curator, Division of Mammals, U.S. National Museum
qnaenneseeee
Zant NCRes
<2 OCEAN
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(PUBLICATION 3030)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
DEGEMBER 11, 1929
The Bord Baftimore Press
BALTIMORE, MD., U. 8. A.
MAMMALS EATEN BY INDIANS, OWLS, AND
SPANIARDS IN THE COAST REGION OF
THE DOMINICAN REPUBLIC
By GERRIT S. MILLER, JR.
CURATOR, DIVISION OF MAMMALS, U. S. NATIONAL MUSEUM
(WitTH Two PtatTes)
In February and March, 1928, I visited the Samana Bay region,
northeastern Dominican Republic with the special object of obtaining
remains of mammals in the Indian deposits that had been previously
examined by Gabb in 1869-1871 and Abbott in 1916-1923. I was
accompanied by Mr. H. W. Krieger, who had charge of the strictly
ethnological side of the work. Together or separately we obtained
material from six localities: four on the south shore of Samana Bay ;
one, Anadel, near Samana town, on the north shore of the bay; and
one, a large Indian site at the mouth of the Rio San Juan, on the
Atlantic coast, across the peninsula from Samana.’ Mr. Krieger re-
turned alone the following winter and revisited the places that we had
previously worked. He also made excavations in two village sites
not far from Monte Cristi at the northeastern extremity of the
Republic.
At all of these localities we obtained many bones of mammals from
the heaps of Indian refuse. Only once, however, in a lateral recess
about half way up the sloping floor of the cave that occupies most
of the islet of San Gabriel, off the south shore of the bay, did we
find an owl-made deposit of extinct mammals. Here, as at St. Michel,
Haiti, the small living barn owl had plentifully bestrewn the surface
with dejecta containing bones of bats, small birds, and the introduced
European rats and mice. Immediately beneath its surface the cave
floor material was intermingled with the bones of the larger native
rodents that had been devoured by the great extinct owl. This deposit
was not more than two feet deep, and, unlike the kitchenmidden
lying in the lower level of the cave, it was considerably hardened
*A general account of this work was published in Expl. and Field-Work
Smithsonian Inst., 1928, Smithsonian Publ. No. 3011, pp. 43-54, March 22, 1929.
SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 82, No. 5
2 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
by infiltrated lime drip.* Other owl deposits of the same kind may have
once existed in the neighboring caves, but if so, they all appear to
have been removed years ago by guano diggers. The specimens
that we obtained in these caves and village sites form the subject
matter of the present paper.
While no hitherto unknown species are represented in our col-
lections the material proves to be of much interest. It throws ad-
ditional light on the characters and distribution of the two species of
Plagiodontia that I recognized in 1927 as occurring in the Dominican
Republic (Proc. U. S. Nat. Mus., Vol. 72; Art. 16, pp. 1-8, Sept. 30;
1927) ; it furnishes the means to identify all four of the native mam-
mals, the hutia, the quemi, the mohuy and the cori, that Oviedo said
were habitually eaten by both natives and Spaniards during the early
years of the sixteenth century ; and finally it shows beyond the possi-
bility of reasonable doubt that this recently extinct fauna included a
ground sloth.
The identity of only two of the mammals that Oviedo ascribed to
the island of Hispaniola remains to be determined—the “ dumb dog ”’
and the indigenous rat. The few dog bones collected appear to differ
in no way from the corresponding parts of European dogs, and there
is nothing to prove that they represent the native breed. Hence the
status of the famous “ perro mudo,” the dog that was unable to bark,
is still as much of a mystery as ever. Equally obscure is the question
as to whether or not there were rats on the island at the time of its
discovery. Oviedo relates that on inquiring into this matter he found
those who told him that “mures 6 ratones” did in fact then exist;
a circumstance that appeared to him quite believable because these
animals were so well known to be generated, like flies, mosquitoes,
wasps, and grubs, anywhere, out of any kind of putrifying matter,
a not unnatural belief at the time when he wrote, more than 125
years before Francisco Redi had published his “ Esperienze Intorno
Alla Generazione Degl’ Insetti.” Nevertheless our search has failed to
reveal a trace of rats or mice other than the European species that
could have easily been brought by the Spaniards on their ships. After
enumerating the specimens that we obtained I shall return to the sub-
ject of Oviedo’s mammals in greater detail.
DESCRIPTION OF COLLECTING STATIONS
1. Railroad cave—A large cave situated about 15 minutes walk
inland along the abandoned railroad on the south shore of San Lorenzo
Bay. There is an extensive kitchenmidden at the entrance. I was
not able to find any trace of a bone deposit made by the extinct owl,
NO. 5 MAMMALS FROM DOMINICAN REPUBLIC——-MILLER 3
though pellets of the living bird were abundant in one of the chambers.
This may be the “ Cueva del Templo” of Rodriguez (Geografia
fisica, politica e historica de la Isla de Santo Domingo o Haiti, p. 367,
Santo Domingo City, 1915).
2. Boca del Infierno—Two large caves, one in each of the pro-
jecting points at the locality marked Pta. de Boca del Infierno on the
Hydrographic Office chart of Samana Bay.
The larger cave is in the smaller, inner point. It has been extensively
worked for guano, but some of the original floor material remains.
Near the outer entrance there is a small kitchenmidden. A few leg
bones of extinct rodents were found in this cave, but no skulls or jaws.
At the inner entrance to the other cave we found the remnant of
a kitchenmidden left intact by the guano diggers. From this deposit
we unearthed bones of both the “quemi” and the ground sloth,
mammals whose remains have not been found elsewhere among the
Indian refuse.
These caves appear to be, respectively, the “ Boca del Infierno”’
and the “Cueva del Infierno”’ of Rodriguez. It is probable that in
one or the other of them Gabb collected the bones of Plagiodontia
that I recorded in 1916.
3. San Gabriel—An islet about two miles west of Boca del Infierno.
Most of its interior is occupied by a large cave, the floor of which
slopes rather steeply upward from an opening on the south side facing
the shore to another on the north side overlooking Samana Bay.
There is a large kitchenmidden near the lower entrance, and a deposit
made by the extinct owl on the left side of the passage leading up
to the north aperture.
4. Rio Naranjo Abajo—A kitchenmidden was found on a nearly
level rock ledge, perhaps one-fourth acre in extent, on a key lying
about half a mile east of the stream mouth.
These four localities are all on the south shore of Samana Bay in
the region known as the Playa Honda coast. Rodriguez describes
the caves under the general title: “ Cuevas de los Haitis.”
5. Anadel.—A large village site at a stream mouth on the north
shore of Samana Bay about 14 miles east of Santa Barbara de Samana.
6. Rio San Juan—Another large village site on the Samana Penin-
sula. It lies at the point where the Rio San Juan flows into the
Atlantic Ocean, almost directly north of Santa Barbara de Samana.
7and 8. Kilometer 2 site and Kilometer 4 site-——Two very extensive
village sites in the foothills of the mountains southeast of Monte
Cristi. Both of these localities differ from those in the Samana Bay
region in being situated in the semiarid portion of the island.
4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
LIST OF MAMMALS COLLECTED
NESOPHONTES PARAMICRUS Miller
Railroad cave-—Humerus, I.
San Gabriel (owl deposit )—Mandibles, 3; humeri, 4; femora, 6;
innominate, I.
It is impossible to determine whether or not the presence of the
humerus in the Railroad cave kitchenmidden indicates that Neso-
phontes was eaten by the Indians. In this cave such a bone might
as well have been dropped by an owl as by a man.
NESOPHONTES HYPOMICRUS Miller
San Gabriel (owl deposit ).—Mandibles, 4; humerus, 1; femur, 1;
innominate, I.
The remains of both species of Nesophontes agree perfectly in size
and other characters with topotypes from St. Michel, Haiti.
SOLENODON PARADOXUS Brandt
Railroad cave—Perfect right humerus, 1.
Naranjo Abajo—Mandible, 1; distal half of left humerus, 1.
Rio San Juan.—Mandible, I.
Kilometer 2 site—Distal half of humerus, I.
These specimens do not differ in any way from the corresponding
parts of the living animal. Their presence in three widely separated
kitchenmiddens is sufficient indication that Solenodon was customarily
eaten by the Indians.
EPTESICUS HISPANIOLAE Miller
MACROTUS WATERHOUSII WATERHOUSII Gray
ARTIBEUS JAMAICENSIS JAMAICENSIS Leach
A few remains of these common Dominican bats were found in
the owl deposit in San Gabriel cave.
CANIS FAMILIARIS Linnaeus
Kilometer 2 site——Right mandible, immature, 1; both mandibles
of a very young individual, 1 pair; separate milk pm‘, 1; adult m’,
2 (not from same individual); adult canine, 1; adult incisor, 1;
auditory bulla, 1; vertebrae, 8; ribs, 2; fragments of pelvis, 4 (repre-
senting at least 2 individuals) ; tibia, proximal end, 1; tibia, distal end
(probably of another individual), 1; calcanea, 2 (opposites) ; astraga-
NO. 5 MAMMALS FROM DOMINICAN REPUBLIC——-MILLER 5
lus, I (apparently belongs with the calcanea) ; metapodials, 9; im-
perfect scapula, 1; ulna, 1; radius (perfect) 1; radius (proximal end
only), I.
The remains pertain to at least two adult dogs and two puppies.
Taking the radius (total length 127.6 mm.) scapula, and teeth as
guides, the animal must have been about the size of a Scotch terrier
whose skeleton is now in the National Museum, No. 21997 (total
length of radius 120.2 mm.).
I am unable to find characters in any of these specimens that sug-
gest specific or racial peculiarities as compared with domestic dogs
of European origin or with pre-Columbian dogs from either North
or South America. Furthermore, as the middens near Monte Cristi
yielded bones of both pig and cow, there is no reason to suppose that
the dog had any other than European origin. The apparent absence
of dog bones from all the other deposits of Indian refuse is a clear
indication that the natives did not habitually use these animals as food.
CERCOPITHECUS ? sp. ?
Plate 2, fig. 4
Naranjo Abajo.—Distal end of tibia, 1.
The well preserved distal end (42 mm.) of a monkey’s tibia was
found among the miscellaneous long bones dug from the kitchen-
midden on the Naranjo Abajo key. The exact level at which it lay
was not determined. In state of preservation the bone is essentially
like the rodent leg bones from the same deposit.
I cannot identify this fragment with the corresponding part of
any American primate, chiefly because the shaft of the bone, im-
mediately above the articular enlargement is too robust. By this
character the fragment (pl. 2, fig. 4) can at once be distinguished
from specimens of Cebus (pl. 2, fig. 6) Ateles, and Alouatta, the only
common genera containing species large enough to approach it in size.
When compared with Cercopithecus (pl. 2, fig. 5), however, the dis-
crepancy is less obvious, though I have not been able to find an African
tibia that I should regard as certainly pertaining to the same species.
As members of this genus were early introduced into the Lesser
Antilles I am inclined to believe that the monkey of the Naranjo
Abajo key had been brought over alive before the Indians abandoned
the coast of Samana Bay.
BROTOMYS VORATUS Miller
Plate 1, fig. 3
Railroad cave——Skull, lacking braincase and all teeth except pm*,
1; fragments from interorbital region, 2; mandibles, 7.
6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
Boca del Infierno—Humerus, I.
San Gabriel (owl deposit ).—Lower incisor, 1; humeri, 3.
San Gabriel (culture deposit ).—Mandible, 1.
Naranjo Abajo.—Right side of palate with pm* and m! in place, I;
mandibles, 3.
Anadel.—Mandibles, 14.
Rio San Juan.—Right side of palate with all four teeth in place, 1;
fragment of premaxilla with incisor in place, 2; mandibles, 43.
Kilometer 2 site——Right premaxilla and anterior portion of palate
with pm‘ in place, I; palate with all the alveoli and pm* left and pm*
and m1? right in place, 1. Left side of palate with alveoli of all four
teeth, I ; mandibles, 61.
Kilometer 4 site-—Mandibles, 16.
The frequency with which the bones of this animal occur in the
Indian deposits indicates that Brotomys must have been abundant
and generally distributed in pre-Columbian days. It was probably
much like the living South American spiny-rats in size and general
form, but with heavier, less elongated head. I have little doubt that
this animal was the mohuy described by Oviedo as the most eagerly
sought for of the native edible quadrupeds (see p. 13).
This material agrees in all essential features with the original
specimens from San Pedro de Macoris and with those that have been
collected in the Haitian cave deposits. Except for individual pecu-
liarities that appear to be due to age the jaws are very constant in all
their characters. I can detect no differences between those collected
in the humid Samana Bay region and those from the semiarid country
near Monte Cristi.
In one jaw from Kilometer 2 site, the premolar is in a stage of wear
to show that the small enamel “lake ” usually present in the anterior
lobe of the crown is the remnant of a reentrant fold penetrating from
the outer side of the tooth.’ In two others from Rio San Juan, the lake
has been joined to the tip of the anterior inner reentrant fold, while
in one specimen from the same locality, the crown, though not exces-
sively worn, shows no trace of the anterior “lake,” its pattern thus
resembling that of the molars.
ISOLOBODON PORTORICENSIS Allen
Plate 1, fig. 6
Railroad cave —Imperfect skulls, 4; left half of rostrum, 1; right
premaxillary with incisor, 1. Left half of palate, without teeth, 1;
right half of palate, m? in place, I; mandibles, 20.
* An even better specimen in the same stage was collected by Arthur J. Poole
in the small cave near St. Michel, Haiti.
4
NO. 5 MAMMALS FROM DOMINICAN REPUBLIC—MILLER Hi
Boca del Infierno—Imperfect palate with left m* in place, 1;
mandibles, 7.
San Gabriel (culture deposit).—Palate, 1; fragment of left pre-
maxilla with incisor, I ; mandibles, 3.
Naranjo Abajo.—FPalate with right m? in place, 1; upper molar of
a larger individual, 1; mandibles, 3.
Anadel.—Palate with all teeth, 1; fragments of rostrum with in-
cisor, 5; occipitals, 1; mandibles, 41. Numerous odd teeth.
Rio San Juan.—Broken skull, 1; complete palate with all teeth, 1;
palate lacking left m*, 1; fragments of palate, 9; fragments of rostrum
with incisor, 10; mandibles, 184.
Kilometer 2 site—Imperfect skull, 1; fragments of palate, 3;
mandibles, 10.
ISOLOBODON LEVIR (Miller)
Plate 1, fig. 5
San Gabriel (owl deposit )—Imperfect skulls, 2; palate with right
molars in place, 1; separate maxillary teeth, 2; mandibles, 13.
Kilometer 2 site——Palates and fragments, 21; mandibles, 281.
These specimens agree with the original series from caves near St.
Michel, Haiti, and differ obviously from the remains of [solobodon
portoricensis recovered from the kitchenmiddens in the Samana region.
Among 15 jaws selected for large size, the length of mandible from
articular process ranges from 44.6 to 48 mm., height of ascending
ramus through articular process from 20.6 to 23 mm., and alveolar
length of toothrow from 16 to 17.6 mm. In 11 jaws of J. portoricensis
from the San Juan River, also selected for large size, the extremes
of the same measurements are respectively 50 to 52.6 mm., 24 to
26.6 mm. and 19 to 20.8 mm.
After examining the entire series of Santo Domingan Isolobodon
remains I am still as unable to distinguish the large form from the
Porto Rican J. portoricensis as I was in 1918 on the basis of the very
few specimens then collected. It seems improbable that such a
distribution could exist without human intervention. No other species
of rodent has been found to be common to the two islands and
no species could be expected to remain constant in two areas that
have been separated as long as these two land masses. Finally,
Porto Rico and the eastern part of the Dominican Republic, together
with the Virgin Islands, where the same large Jsolobodon also occurred,
are in a region known to have been freely traded over by pre-
Columbian man in his sea-going canoes. It must be admitted, however,
that the hypothesis of human transportation meets with a difficulty
& SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
not easy to dispose of, namely, the fact that no Plagiodontia or
Brotomys seems to have been carried in the opposite direction to
Porto Rico or the Virgin Islands.
At only two localities have the large and small forms of /solobodon
thus far been found together, in the kitchenmidden at the Kilometer 2
site near Monte Cristi and in San Gabriel cave. In the kitchenmidden
the remains of the two were mingled together—ten jaws of the large
animal among a total of 290. In the cave they occurred separately—
the large animal in the culture deposit at the main (south) opening,
the small one in the owl deposit near the middle of the long, ascending
passage that leads up to the aperture facing north. All the bones in
the owl deposit have the appearance of much greater age than those
in the midden. The material in which they were found is heavily
and uniformly impregnated with lime, while that in the midden, like
that in the human deposits in all the neighboring caves, shows no
such infiltration except at spots where actual drip from the ceiling
is now taking place. The presence of Aphetreus montanus among
the owl refuse may also be an indication of greater age, as this rodent
has not yet been found in any midden, though it is the second most
common species in the owl-made cave deposits near St. Michel, Haiti.
On the assumption that Jsolobodon portoricensis was introduced
by man in the Samana Bay region, these facts would be explained by
supposing that the San Gabriel owl deposits were formed before the
importation of this larger species and the subsequent extermination
of the smaller indigenous form. The process of replacement of the
smaller animal by the larger would have afterward become so com-
plete throughout the Samana region that no remains of the native
species have been found in the deposits left by the Indians. Extending
its range westward, Jsolobodon portoricensis would have just begun
to establish itself near Monte Cristi when both it and the Indians
became extinct.
APHATREUS MONTANUS Miller
San Gabriel (owl deposit )—Mandibles, 3.
These specimens show no peculiarities as compared with jaws from
the type locality, near St. Michel, Haiti.
No bones of Aphetreus have yet been found in any culture deposit.
PLAGIODONTIA ADIUM F. Cuvier
Plate 1, fig. 2
Anadel—tImperfect skulls, 2; fragments of palate, 2; mandibles,
20; odd teeth, 9.
NO. 5 MAMMALS FROM DOMINICAN REPUBLIC—MILLER 9
Rio San Juan—Rostrum with incisors and first two cheekteeth, 1;
fragments of premaxilla with incisor, I; mandibles, 30; odd teeth, 25.
Kilometer 2 site—Fragments of palate, 2; complete mandible, 1;
fragments of mandibles, 2; odd teeth, 3.
Kilometer 4 site-—Fragment of palate, 1 ; mandibles, 6; odd teeth, 5.
PLAGIODONTIA HYLAUM Miller
Plate 1, fig. 1
Railroad cave——Imperfect skulls, 2; right side of rostrum with
incisor, 1; palate lacking m* of both sides, 1; mandibles, 4.
San Gabriel (owl deposit )—Mandible, 1 young; left lower incisor,
adit, 2.
The specimens now at hand enable me to confirm the original
diagnosis of Plagiodontia hyleum and also to add two important
characters.
That the living animal is decidedly smaller than Plagiodontia edium
is abundantly shown by comparison of the skulls and jaws from
Guarabo and the south shore of Samana Bay with the remains of the
larger animal collected on the Samana peninsula and near Monte
Cristi. The 12 jaws of P. hyleum whose measurements are given in
the original description range from 51 to 55.2 mm. in length. An
additional specimen from the Railroad cave is slightly imperfect but
its length must have been about 51 mm. One mandible of P. edium
from San Pedro de Macoris was recorded as slightly more than 62 mm.
long. Unfortunately most of the jaws from the Samana Peninsula are
injured at one end or the other, so that their length cannot be de-
termined, but two from Anadel give measurements of approximately
61 and 62 mm. A measurement that is more useful, because mandibles
are seldom so badly broken that it cannot be taken, is the depth from
the alveolar margin to the protuberance made by the root of pmg.
In 10 jaws of Plagiodontia edium this depth averages 16.3 mm. with
extremes of 15.4 and 17.4 mm. In an equal number of jaws of the
smaller animal the average depth is 13.2 mm., the extremes 12.2 and
14.0 mm. Similarly obvious and constant is the difference between
the alveolar length of mandibular toothrow in the two species. Ten
specimens of each give the following averages and extremes: P.
hyleum, 19.8 mm. (18.6 to 20.6 mm.) ; P. edium, 24.2 mm. (23.2 to
25.4 mm.).
The most important character brought to light by the new material
is, however, the difference in relative length of the first and second
maxillary cheekteeth. In Plagiodontia hyleum the crown length in-
10 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
creases gradually and rather uniformly from the fourth tooth to the
first; in P. edium there is the same gradual increase from fourth to
second, and then an obviously and abruptly greater increase from
second to first (see pl. 1, figs. I and 2). The relative lengths of the
first and second teeth, measured along the median line of the grinding
surface is as follows in seven specimens of each species: P. hyleum,
pm‘, average 6.1; m+, average 5.1; ratio of premolar to molar 119.8;
P. edium, pm‘, average 7.6; m', average 5.6; ratio of premolar to
molar 135.7.
CAPROMYS PILORIDES Desmarest
Plate 1, fig. 4
San Gabriel (culture deposit).—Complete nasals and turbinates,
1; right mandibles (all toothless), 5; upper incisors, 2.
These specimens were found near together in the San Gabriel
kitchenmidden at a depth of about three feet. They do not differ
from the corresponding parts of Cuban skulls of Capromys pilorides
in any way that I can discover. Consequently I have no doubt that
the animals to which they pertained were brought to the cave as food,
either by the Indians or by early European sailors.
QUEMISIA GRAVIS Miller
Plate 2, fig. 3
Boca del Infierno—Distal half of right femur, 1; proximal ex-
tremity of left ulna, I.
Both fragments (pl. 2, fig. 3) were found at a depth of about four
feet in the kitchenmidden near the south entrance to the outermost
of the two caves.
As compared with the corresponding part in the Porto Rican
Elasmodontomys the distal extremity of the femur has a reduced
antero-posterior diameter (ratio to lateral diameter about 78 instead
of 92.5 and 93.6 in two Elasmodontomys) ; the shaft is more flattened
on its anterior aspect and less flattened on its posterior aspect; and
the antero-posterior diameter at middle of shaft is less in proportion
to the transverse diameter.
As compared with the femurs of /solobodon and Plagiodontia from
the Samana region this fragment is at once distinguishable by its
strikingly greater size. It appears to correspond perfectly with the
opposite end of the femur of Quemisia that I found in one of the
caves near the Atalaye Plantation, St. Michel, Haiti; and its presence
No. 5 MAMMALS FROM DOMINICAN REPUBLIC—MILLER II
in a kitchenmidden confirms my belief that this large rodent is
Oviedo’s “ quemi.”
CAVIA sp.
Anadel_—Mandibles, 2 (opposites but not from one individual).
I cannot distinguish these jaws from specimens of Cavia porcellus.
They present every appearance of having been buried as long as the
remains of Brotomys voratus and Plagiodontia edium with which they
were associated.
ACRATOCNUS COMES Miller ?
Plate 2, fig. 2
Boca del Infierno—Penultimate phalangeal bone, probably of sec-
ond or fourth pedal digit, 1.
This bone was found in the kitchenmidden at the south entrance
to the outermost of the two caves. It was unearthed at a depth of
not more than four feet, near the femur of Quemisia, with which it
agrees in its perfect and seemingly unmodified condition, of preserva-
tion. Both bones, in fact, seem to be, so far as it is possible to de-
termine from superficial inspection, in essentially the same state as
bones of the living species of Plagiodontia with which they were
associated. There appears to be no longer the slightest reason to
doubt that a ground sloth was a member of the recently man-exter-
minated fauna of Hispaniola.’
This bone (pl. 2, fig. 2) is similar in general form to the second
right pedal phalanx of the Patagonian Hapalops elongatus as figured
by Scott (Rep. Princeton Univ. Exped. Patagonia, Vol. 5, Palaeont, 2,
pl. 41, fig. 2), but it is about 2 mm. longer and its proximal extremity
appears to be deeper. It also resembles in a general way an isolated
phalangeal bone of Acratocnus from Porto Rico figured by Anthony
(item: miuer. Mus. Nat. Hist...n'-s. Vol:’2,.Pt 2. fie. 53 f, p. 425,
1918). From an imperfect specimen that may represent the cor-
responding bone in Acratocnus comes it differs rather noticeably in the
less diameter of the distal articular region (compare pl. 2, figs, 1 and
2) and the more abrupt deepening toward the proximal end.
TRICHECHUS MANATUS Linnaeus
Rio San Juan—Fragments of palate, 2 (large and small) ; im-
perfect ribs, 2.
*T have already discussed the evidence to this effect furnished by the conditions
existing in the caves near St. Michel, Haiti (Smithsonian Misc. Coll., Vol. 81,
No. 9, pp. 25-26, March 30, 1929).
12 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
Kilometer 2 site-——Palate, 1; odd teeth, 8; vertebrae, 2.
Kilometer 4 site-—Fragments of occipital region, 2; fragments of
mandible, 2; broken ribs, 13; humerus, 1; distal end of humerus, 1;
fragment of scapula, 1; fragment of femur, I.
These specimens do not differ appreciably from Florida material,
except that the alveoli in the palate found at the Kilometer 2 village
site appear to be exceptionally large.
THE MAMMALS DESCRIBED BY OVIEDO
Gonzalo Fernandez de Oviedo y Valdés (1478-1557), the first
European chronicler of things West Indian, was alcalde of Santo
Domingo City from January, 1536, to August, 1546. In his Historia
GENERAL y NATURAL DE Las Inp1As, Book 12, Chapters 1 to 6 (pp.
389-392 of the edition issued by the Royal Academy of History,
Madrid, 1851) he described the following mammals as known or
believed by him to inhabit the island of Hispaniola: the hutia, the
quemi, the mohuy, the cori, the dumb dog (“perro mudo”) and
the mice (‘‘mures 6 ratones’’).
Hitherto there has been much doubt as to the exact identification
of these animals, for the reason that Plagiodontia edium and Soleno-
don paradoxus were, up to a few years ago, the only indigenous
mammals known, other than bats and sea-cows. It now seems possible,
however, to allocate all of Oviedo’s names, with the exception of
the “dumb dog.” I shall take them up in order.
THE HUTIA
Oviedo writes that there occur in this island of Hispaniola, and
in others lying in the seas near it, animals called hutia, four-footed,
and resembling a rabbit, but smaller sized, smaller eared and rat-tailed.
The natives, he says, kill them with small dogs that they have in
domestication, dumb and not knowing how to bark; and the Christians
do this much better with the dogs they brought from Spain. “ These
animals are grizzled gray (pardo gris) in color according to the
evidence of many who have seen and eaten them and who praise
them as food; and there are now many persons in this city of Santo
Domingo and in this island who say so. But at present these animals
are no longer found except very rarely.”
This account would apply so well to the species of Plagiodontia,
and presumably also to the Isolobodons, that there seems to be no
reason to doubt that these were the animals that Oviedo had in mind.
By the present day Dominicans the name seems to have been trans-
NO: 5 MAMMALS FROM DOMINICAN REPUBLIC—MILLER 13
ferred to Solenodon; at least, such persons whom I met as knew of
an animal called hutia expatiated on the great length and pointedness
of the creature’s snout. The very few who were acquainted with
Plagiodontia hyleum happened to be English speaking descendants
of negroes from the United States. They always spoke of the animal
as the “muskrat,” and they told me that many of these creatures
had been killed by the workmen who cleared the narrow San Lorenzo
Peninsula for cocoanut planting 20 or more years ago.
THE QUEMI
The quemi resembled the hutia in color and general appearance,
but was much larger, its size equaling that of a medium-sized hound.
Oviedo did not see it himself, and he believed it to be extinct. How-
ever, he assures his readers that: “‘ There are many persons in this
island and in this city who have seen and eaten these animals and
who declare that they were good food; but in truth, according to
what has been said and known about the hardships and deprivations
that the first colonists endured in this island it can be presumed that
everything that could be eaten must have then appeared to them
very good and delicious, even when it was not.”
The qualifications of an animal resembling the hutia, good to eat,
and as big as an ordinary hound seemed to me to be fulfilled by the
large rodent whose remains I found in the caves near St. Michel,
Haiti, in 1925. Consequently I proposed for it the generic name
Quemusia. The presence of the same creature in the Boca del Infierno
kitchenmidden appears to confirm my guess.
THE MOHUY
“The mohuy is an animal somewhat smaller than the hutia: its
color is paler and likewise gray. This was the food most valued and
esteemed by the caciques and chiefs of this island; and the character
of the animal was much like the hutia except that the hair was denser
and coarser (or more stiff), and very pointed and standing erect or
straight above. I have not seen this animal, but there are many who
declared it to be as aforesaid ; and in this island there are many persons
who have seen it and eaten it, and who praise this meat as better
than all the others we have spoken about.”
There can be little if any doubt that the animal Oviedo thus de-
scribed was Brotomys voratus. This rodent was smaller than either
Plagiodontia hyleum or Isolobodon levir, and its remains have been
found in every kitchenmidden that has been examined in the Domini-
IA SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
can Republic (the type specimen came from San Pedro de Macoris),
a fact that shows how universally it was liked as food. Finally the
account of stiff, pointed, erect-standing hairs of the back seems
especially applicable to a relative of the South American spiny-rats.
THE CORI
Oviedo had first-hand knowledge of the cori. Consequently his
description of it is more detailed and accurate than in the case of
the three preceding animals. He writes: “ The cori is a small quad-
ruped, the size of a half grown young rabbit. These coris appear to be
a species of the rabbit kind although they have a muzzle like a rat but
not so pointed. They have very small ears which they hold so close
that it appears as if they lacked them or did not have any. They
have no tail whatever ; they are very slender as to feet and hands from
the joints or hams downward; they have three fingers and another
smaller, and very slender. They are wholly white, and others every-
where black, and the most of them spotted with both colors. Also
some are wholly reddish and some spotted with reddish and- white.”
Continuing his account he says that the coris are kept in the house
and fed on grass, with some cassava to fatten them. He has eaten
them and found them to taste like young rabbit. When he wrote
they were plentiful in Santo Domingo City. They were also to be
found on other islands and on the mainland.
It is not difficult to recognize the guineapig in this account of the
cori; but if any doubts might have existed, in the absence of more
tangible evidence, they are disposed of by Mr. Krieger’s discovery
of the two Cavia jaws in the kitchenmidden at Anadel. It remains
an open question whether the guineapig was introduced by the Span-
iards or by native trade with South America. I incline to the first
alternative, chiefly because remains of the animal have been found
in only one midden. Bones of cow, horse, and pig, as well as artifacts
of European origin occasionally occur in the Indian deposits, showing
that the native village sites continued to be used for some time after
the Spanish conquest began, and that material brought in by the
newcomers found its way to the aboriginal refuse heaps. Such might
easily have been the history of the guineapig jaws at Anadel.
THE DUMB DOG
In his account of the hutia we found Oviedo alluding to a native
dog that could not bark, but which was, nevertheless, very useful
as a game getter. On pages 390-391 of the 1851 edition of his book
NO. 5 MAMMALS FROM DOMINICAN REPUBLIC—MILLER I5
he gives an extended account of the dogs formerly and at the time
of his residence (1536-1546) occurring on the island of Hispaniola.
Parts of this account I translate as follows: “ Domestic cur dogs
were found in this island of Hispaniola and in all the other islands
of these seas (inhabited by Christians). They were bred by the Indians
in their houses. At present there are none; but when they had them
the Indians used them to capture all the other animals [that is, the
hutia, the quemi, the mohuy and the cori] that have been spoken of
in the preceding sections. These dogs were of all the colors that dogs
have in Spain ; some of a single color and others spotted with white and
blackish or reddish or ruddy or any color that the coat is accustomed
to have in Castile. Some woolly, others silky, others short-haired ;
but the most of them between silky and short-haired, and the hair
of all of them more harsh than our dogs have, and the ears lively
and alert like those of wolves. All of these dogs, here in this island
and the other islands, were mute, and even though they might be
beaten and killed they did not know how to bark: some of them yelped
or whined when they were hurt.”
Continuing, he tells us that he has seen dogs of the same kind on
the mainland in the province of Santa Marta as well as in Nicaragua,
and that in the latter country the natives regularly used them as food.
He makes no mention of the eating of dogs by the natives of
Hispaniola, and the complete absence of bones of this animal from
the collections made by us in the Samana region and by Theodoor
de Booy at San Pedro de Macoris makes it seem probably that this
habit did not exist, or at least that it was not very general. If the
dumb dog was anything else than a special breed of Canis familiaris
we have as yet no evidence of the fact.
The five animals thus described are, Oviedo insists (p. 391), the
only furred terrestrial quadrupeds, other than rats or mice, native to
Hispaniola. It therefore seems evident that he knew nothing about
Solenodon or the ground sloth.
With regard to the mice, which he believed to be indigenous, there
is no reason to suppose that they were not brought over by the
Spaniards themselves. No native mammal the size of a mouse, except
Nesophontes, has been found in any owl deposit or kitchenmidden on
the island, and it seems improbable to the highest degree that this
small insectivore could have been the animal known to Oviedo and
supposed by him to have been spontaneously generated from some
kind of corruption in this remote part of the world.
16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
EXPLANATION OF PLATES
PLATE I
All figures natural size
Fic. 1. Plagiodontia hyleum Miller. Adult male. No. 239801, U. S. Nat. Mus.
The largest specimen of the living animal collected by Dr. W. L. Abbott.
Guarabo, Dominican Republic, Nov. 24, 1923.
Fic. ta. Plagiodontia hyleum Miller. Adult female. No. 239888, U. S. Nat.
Mus. Guarabo, Nov. 23, 1923.
Fic. 2. Plagiodontia edium Desmarest. Adult. No. 254376, U. S. Nat. Mus.
Anadel, Dominican Republic.
Fics. 3 and 3a. Brotomys voratus Miller. Nos. 254683 and 254684, U. S. Nat.
Mus. Railroad cave, San Lorenzo Bay, Dominican Republic.
Fic. 4. Capromys pilorides Desmarest. Adult. No. 254449, U. S. Nat. Mus.
San Gabriel cave, Samana Bay, Dominican Republic. (Observe spacing of
ridges in alveoli as compared with that of the ridges in alveoli of Jsolobodon,
fig. 6.)
Fic. 5. Isolobodon levir (Miller). Adult. No. 254686, U. S. Nat. Mus. Near
Monte Cristi, Dominican Republic.
Fic. 6. Isolobodon portoricensis Allen. Adult. Railroad cave, San Lorenzo Bay,
Dominican Republic.
PLATE 2
All figures natural size
Fic. 1. Acratocnus comes Miller. Phalangeal bone from cave near St. Michel,
Haiti. No. 253210, U. S. Nat. Mus.
Fic. 2. Acratocnus comes Miller ? Phalangeal bone from kitchenmidden in Boca
del Infierno cave, Samana Bay, Dominican Republic. No. 254680, U. S. Nat.
Mus.
Fic. 3. Quemisia gravis Miller. Parts of femur and ulna from kitchenmidden in
Boca del Infierno cave, Samana Bay, Dominican Republic. No. 254681, U. S.
Nat. Mus.
Fic. 4. Monkey. Lower end of tibia from kitchenmidden on Naranjo Abajo Key,
Samana Bay, Dominican Republic. No. 254682, U. S. Nat. Mus.
Fic. 5. Cercopithecus pygerythrus. Lower end of tibia. Changamwe, British
East Africa. No. 163327, U. S. Nat. Mus.
Fic. 6. Cebus capucinus. Lower end of tibia. North Ecuador. No. 113418, U. S.
Nat. Mus.
=? eae. eee ee ee ee ee
Ee ———
———e——— ss. lhc ee ee
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOEMS2 NOs Si rled
1. Plagiodontia hyleeum.
2. Plagiodontia zedium.
3. Brotomys voratus.
. Capromys pilorides.
. Isolobodon levir.
. Isolobodon portoricensis.
one
>
Co
(All figures natural size.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS Who 25 INOs Be FIL, 2
uu
; es
5 Fa Ta
1. Acratocnus comes. 4. Monkey (not identified ).
2. Acratocnus comes ? 5. Cercopithecus pygerythrus.
3. Quemisia gravis. 6. Cebus capucinus.
(.\ll figures natural size.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 82, NUMBER 6
THE PAST CLIMATE OF THE NORTH
POLAR REGION
BY é
EDWARD W. BERRY
The Johns Hopkins University
(PUBLICATION 3061)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
APRIL 9, 1930
The Lord Baltimore Press
BALTIMORE, MD., U. 8. A.
A
OL am. | a JA ee
ak PASI CEivVAtTE OF THE NORTH POLAR REGION *
By EDWARD W. BERRY
THE JOHNS HOPKINS UNIVERSITY
The plants, coal beds, hairy mammoth and woolly rhinoceros; the
corals, ammonites and the host of other marine organisms, chiefly
invertebrate but including ichthyosaurs and other saurians, that have
been discovered beneath the snow and ice of boreal lands have always
made a most powerful appeal to the imagination of explorers and
geologists. We forget entirely the modern whales, reindeer, musk ox,
polar bear, and abundant Arctic marine life, and remember only the
seemingly great contrast between the present and this subjective past.
Nowhere on the earth is there such an apparent contrast between the
present and geologic climates as in the polar regions and’ the mental
pictures which have been aroused and the theories by means of which
it has been sought to explain the fancied conditions of the past are
all, at least in large part, highly imaginary.
Occasionally a student like Nathorst (1911) has refused to be
carried away by his imagination and has called to mind the mar-
velously rich life of the present day Arctic seas, but for the most
part those who have speculated on former climates have entirely
ignored the results of Arctic oceanography. Recently, Kirk * has mar-
shalled some of the evidence of the abundance of the present marine
life in the Arctic, and he concludes from this survey that marine
organisms are not dependable as indicators of geologic climates. |
think this conclusion is impregnable, and therefore if we are ever to
get any information regarding past climates, the evidence will be fur-
nished by fossil plants, and not too precisely either. Here again
prudence is the watchword ; imagination must be entirely suppressed,
and the distribution of recent plants must be understood and used.
A correct solution of the problem is not only of prime interest to
geologists and paleontologists but it offers assurance to geophysicists
confronted with the now fashionable belief in wandering poles, and
"Given in summary before the Paleontological Society at the December, 1928,.
meeting.
* Kirk, Edwin, Fossil marine faunas as indicators of climatic conditions. Ann.
Rep. Smithsonian Inst. for 1927, pp. 299-307, 1928.
SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 82, No. 6
2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
likewise comfort to meteorologists confronted with the traditional
view of a lack of climatic zones during most of the eons of earth
history. I propose to pass in review what we know of the past dis-
tribution of plants in the Arctic, after which I will endeavor to evaluate
what they mean in terms of climate.
Aside from some very scrappy plant fragments from the Silurian
of Norway, the oldest traces of land plants in the north occur in rocks
of Devonian age. Devonian plants have been discovered within the
Arctic Circle at the three localities shown on the accompanying map,
(fig. 1). These range froma few scraps, such as those found in
Ellesmere Land and Spitzbergen, to the extensive flora found on
Bear Island which embraces 31 named forms. These three floras
are of upper Devonian age but not necessarily synchronous, since an
earlier and a later horizon is represented on Bear Island and probably
on Spitzbergen.
I have shown on the map (fig. 1) the occurrence of some other
Devonian floras outside the Arctic Circle and some in lower latitudes
in order to give an idea of the known geographical range of Devonian
plants in the present North Temperate Zone. The oldest of these is
the Lower Devonian flora of Roragen, Norway, embracing eight
very interesting forms. Of particular interest are the Middle Devonian
plants found in silicified peats at Rhynie in northern Scotland and
the flora described recently from Germany, since these give us our
first considerable insight into the structure of these ancient plants.
In looking over the list of identifications from Bear Island, all
except Pseudobornia are seen to belong to widely distributed types,
several are identical with species from the south of Ireland, and similar
forms occur rather generally in lower latitudes. There are several
seams of coal at both the older and younger horizons, to which
Bothrodendron contributed a large amount of material. Beneath the
coal seams are underclays with roots in place and the plant remains
show no sorting—that is, delicate material is mixed with stems and
branches of all sizes—both facts indicating conclusively that the bulk
of the material was not transported but grew in the immediate vicinity.
The same statement is true of the Devonian of Ellesmere Land.
The plants of the Devonian are so remote from living forms that
I do not feel that any conclusions regarding the climate are warranted
beyond the statement that they show that there were no climatic
barriers to prevent most of the types found in Latitude 45° to 50°
extending northward to Latitude 75°. There are, however, certain
types which have not yet been found in the north, such as Eosper-
—
No. 6 PAST CLIMATE OF NORTH POLAR REGION—BERRY 3
SCALE FOR AREA
100.000 SQUARE MILES LAMBERTS AZIMUTHAL, EQUAL AREA PROVECTION,
For clase use in Geography, History, Civics, Economics, etc. Prepared by J. Paul Goode Published by The University of Chicago Press, Chicago, Ill
Copyright 1920, by the University of Chicago
Fic. 1—Location of Devonian and Lower Carboniferous northern floras.
Melville Island. Devonian
Ellesmere Land. Devonian
New Brunswick, Maine, etc. Devonian
Northeast Greenland. Lower Carboniferous
Spitzbergen. Devonian and Lower Carboniferous
Bear Island. Devonian and Lower Carboniferous
West Norway. Devonian
Rhynie, etc. Scotland. Devonian
Nova Zembla. Lower Carboniferous
Northern Urals. Lower Carboniferous
. Siberia. Devonian.
SO RIANA Y NO
Leal
|
4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
matopteris from New York and Cladoxylon and Aneurophyton from
Germany, that may possibly indicate more genial climates in those
places than obtained farther north, and Pseudobornia seems to be a
northern type, but until Devonian floras become much better known
no adequate conclusions can be reached.
There are one or two points that deserve emphasis in this connection.
These northern Devonian floras all consist of plants belonging to the
Pteridophyte, Arthrophyte, Psilophyte, Lepidophyte and Pteridosperm
phyla, and such existing representatives of these phyla as have sur-
vived to the present, though few and not directly filiated, such as
Equisetum and Lycopodium, are singularly unaffected by temperature.
For example, there are now two species of Equisetum and one
of Lycopodium found within 10 degrees of the pole in northwestern
Greenland (Ostenfeld, 1925). To be sure these modern Greenland
forms do not reach the size of their Devonian relatives, but this is
true of all existing members of these genera irrespective of latitude.
Moreover, all of these northern Devonian plants appear to have
been bog types. This conclusion is indicated by their forming coal in
place and by the structures disclosed in the silicified peats of Rhynie.
Therefore, we conclude that the chief climatic factor was moisture
rather than temperature. The fact that many of the Devonian plants
were palustrine also gives force to an observation which I have
elaborated in another place * that these Devonian plants while ancient
and simple were not primitive and ancestral, but were the reduced
descendants of more highly organized ancestors. Since speculation
was to have no part in this discussion I refrain from elaborating my
own belief regarding the more precise character of Devonian climate.
LOWER CARBONIFEROUS
(DINANTIAN oR CULM)
Fossil plants have been found in the Lower Carboniferous, or
Mississippian as Americans prefer to call it, at five or six localities
within or near the Arctic Circle. These floras range in extent from
a few doubtful specimens at some localities to the 59 nominal species
described by Nathorst from Spitzbergen. The latter extend to 79°
North Latitude, and a considerable flora of similar species to the
number of ten at least is found between 80° and 81° North Latitude
in northeast Greenland.
1 Berry, Edward W., Devonian Floras. Amer. Journ. Sci., Vol. 14, pp. 109-120,
1927.
No. 6 PAST CLIMATE OF NORTH POLAR REGION—BERRY 5
The Spitzbergen flora comprises 12 fernlike plants, 5 pteridosperms,
I arthrophyte, 25 lepidophytes, 1 cordaites (wood), and 15 of un-
certain botanical affinities. Stigmarias and various roots occur in
place beneath the coal seams, showing that the vegetation was pre-
served essentially in place; and Lepidodendron stems have been col-
lected up to 16 inches in diameter. There are no peculiar Arctic types
in this most extensive known Culm flora nor are there any genera
that are not common to floras of the same age from lower latitudes.
The single wood, Dado. ylon spetsbergense Gothan, fails to disclose
any seasonal growth changes, which might be expected to result from
the Arctic night. No other traces of the Cordaitales other than this
wood have been discovered here, which leads Nathorst to suggest
that the wood may have been carried by currents from some more
southern clime, where also the woods fail to show growth rings.
This may be true, but on the other hand there is great specific varia-
tion in the degree to which growth rings develop in existing conifers,
as Antevs has pointed out, and they tend to be absent under fairly
uniform conditions of humidity. That this is an individual trait of
this particular species and is probably without climatic significance is
shown by the presence or absence of rings in Devonian and Mississip-
pian Dadoxylon woods from lower latitudes. For example, Dadoxylon
beinertianum Endlicher from Silesia, Dadoxylon Tchichatche fianium
Endlicher from Russia, and Dadoxylon vogesiacum Unger from the
Vosges, all of the same age as the Spitzbergen species, show distinct
seasonal rings, but other contemporaneous European species fail to
show them.
I cannot see any very conclusive indications of climate in these
Lower Carboniferqus floras, other than the fact that they extended
in places to within 10 degrees of the pole. Palustrine types pre-
dominate as in the case of the Devonian, and more than half the
known forms are Lepidophytes which we have reason to believe show
little response to temperature. Sphenophyllums are entirely wanting
in Spitzbergen, but are found farther north in Greenland and occur
on Bear Island, so that their absence in Spitzbergen is merely an
accident of preservation or discovery. In general Arthrophytes are
much rarer in the far north than in middle latitudes at this time and
the same seems to be true of a number of genera of large fronded
fern-like plants, which is taken to indicate differences due to latitude.
ERIASSIC
Triassic plants except in the latest or Rhaetic stage are scarcely,
if at all known in the north polar region. There is a species of
6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
Schizoneura recorded from the New Siberian Islands which may be
Rhaetic and there are scattered Rhaetic plants in Greenland and
Spitzbergen; and somewhat farther south in northwestern Norway
(Ando) and southern Sweden.
The most extensive northern Rhaetic flora is that from Scoresby
Sound, East Greenland, between 70° and 71° North Latitude. This
comprises 51 named forms and several additional ones which are not
named. Cycads and ferns predominate, and so far as we can judge
at this lapse of time all belong to cosmopolitan Rhaetic types. Harris,
who has given an excellent account of these plants concludes that
they indicate a temperate climate, largely on the ground of the pre-
dominance of certain forms indicative of relatively pure stands and
the absence of mixtures such as occur in recent tropical assemblages.
He concludes also, from a study of the cuticles of many of the species,
that moisture was plentiful. The wood of Dadoxylon in the Rhaetic
of Spitzbergen has very feebly marked seasonal rings.
JURASSIC
Supposed Jurassic floras completely surround the pole and are
extensively developed throughout Siberia, in Alaska, Greenland, Spitz-
bergen, Franz Josef Land, New Siberian Islands, and elsewhere.
Formerly, many of these, as those in Siberia, were considered Middle
Jurassic, but Nathorst is the authority for the statement that all of
the more northern ones are post Oxfordian, and several, such as that
of Spitzbergen, are on the border between the Jurassic and the Lower
Cretaceous.
The Spitzbergen flora is the most extensive and, according to
Nathorst, includes 2 horizons, one Portlandian and the other possibly
as young as Neocomian. A combined list of these comprises 57
species, including 11 fern-like plants, 1 lepidophyte (Lycopodites),
1 arthrophyte (Equisetites), 4 cycadophytes, 4 Ginkgoales, 23 conifers
and 13 of uncertain affinities. Nine different types of coniferous
woods have been described and all show pronounced seasonal growth
rings. Most of the generic types have a very great geographical range,
but several, such as Phoenicopsis, Torellia and Drepanolepis, appear
to be distinctly northern, and the predominance of conifers suggests
a cool temperate climate. They are found in sandstones associated
with coal seams and freshwater mollusks (Lioplax, Unio) and evi-
dently grew in the vicinity of their burial place.
No. 6 PAST CLIMATE OF NORTH POLAR REGION—BERRY 7
SCALE FOR AREA
100.000 SQUARE MILES
LAMBERT'S AZIMUTHAL, EQUAL AREA PROJECTION
For class use in Geography, History, Givica, Economics, etc. ‘ Prepared by J. Paul Goode. Published by The University of Chicago Press, Chicago, Ill.
Copynght 1920. by the University of Chicago
Fic. 2.—Location of Triassic and Jurassic northern floras.
1. Manchuria. Triassic and Jurassic 14. Solitude Island. Jurassic
2. Ussuri. Jurassic 15. Franz Josef Land. Jurassic
3. Amur. Jurassic 16. King Charles Land. Jurassic
4. Trans Baikal. Jurassic 17. Spitzbergen. Triassic and Jurassic
5. Irkutsk, etc. Jurassic 18. Northeast Greenland. Jurassic
6-10. Alaska. Jurassic (?)} 19. East Greenland. Jurassic
11. Bathursy Island. Jurassic 20. Scoresby Sound. Triassic
12. New Siberian Islands. Triassic (?) 21. Ando, Norway. Triassic
and Jurassic 22. Scania. Triassic and Jurassic
13. Mouth of Lena. Jurassic 23. Scotland. Triassic and Jurassic.
8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
LOWER CRETACEOUS
Lower Cretaceous floras are found along the east coast of Asia,
in Alaska, Greenland, and King Charles Land. From the last a
number of coniferous woods have been described by Gothan. These
show pronounced growth rings, said to be more prominent than in
woods of the same age from central Europe. Nathorst records an
incomplete trunk 32 inches in diameter and showing 210 seasonal
rings. The most extensive Arctic flora of Lower Cretaceous age is
that described by Heer from the Kome beds of western Greenland,
but this, although generally considered to be of Barremian age, is
subject to doubt as to age and content because collectors appear to
have mixed several Cretaceous horizons. As it stands in the literature
it comprises over 100 species, including 46 ferns (no less than 15
are referred to Gleichenia, and although these surely represent that
genus they are artificially multiplied), 1 marsilea, 1 lycopod, 3 equise-
tums, 13 cycads, 20 conifers, 2 ginkgos, 5 monocotyledons, 3 or
4 dicotyledons, and 6 of uncertain identity. The abundance of ferns
indicates a humid climate as does the presence of coal. This flora
differs very little from those of corresponding age in lower latitudes
(e. g., the Kootenai of western Canada and Montana).
UPPER CRETACEOUS
Strictly Arctic Upper Cretaceous floras are limited to Alaska and
Greenland but others of this age are found in northern Europe and
eastern Asia. The most extensive is that from the two horizons in
West Greenland known as the Atane and Patoot beds. These have
in large part been described by Heer and there is a great and un-
warranted multiplication of species. That from the Atane beds has
184 recorded species. It includes 31 ferns, I equisetum, I selaginella,
I marsilea, 12 cycads, 2 ginkgos, 25 conifers, 4 monocotyledons, 94
dicotyledons and 14 of uncertain affinities.
The seemingly most incompatible plant is the authentically deter-
mined Artocarpus and this raises a question which cannot be decided
without prejudice. If a genus which is tropical at the present time
is found fossil associated with a preponderatingly temperate flora,
which is to be given the most weight? The one or the many, bearing
in mind the latitude where they occur? My own feeling is that the
majority are less likely to have altered their environmental require-
ments than the minority, but this falls short of actual proof.
The Patoot flora includes 19 ferns, I equisetum, 19 conifers, 2
monocotyledons, 80 dicotyledons, and 2 uncertain.
NO. 6 PAST CLIMATE OF NORTH POLAR REGION——BERRY 9
SCALE FOR AREA
100.000 SQUARE MILES
LAMBERT'S AZIMUTHAL EQUAL AREA PROJECTION
For clase use in Geography, History, Civics, Economics, etc. Prepared by J. Paul Goode. Published by The University of Chicago Press, Chicago, Ill.
Copyright 1920, by the University of Chicago
Fic. 3.—Location of Cretaceous northern floras.
1. Japan tr. Vancouver Island
2. Sakhalin Island 12. Kootenai -
3-4. Siberia 13. Mattagami, Ontario
5. Klin 14. West Greenland (Kome, Atane,
6-8. Alaska Patoot)
9. Spitzbergen 15. Scania
10. Queen Charlotte Islands 16. Wealden.
10 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
TERTIARY
Tertiary plants from the Arctic have been encountered at very
many localities, usually associated with coal. This, and plants with
their roots in place as in the case of Equisetum in Spitzbergen; the
association with fresh water mollusks, as in Greenland; or aquatic
beetles, as in Spitzbergen and Iceland; as also the presence of fresh
water diatoms in the matrix and the mixtures of branches and delicate
foliage, prove conclusively that these Arctic floras and the associated
coals cannot represent drift material from lower latitudes as some
have supposed.’
The similarity in facies and their mode of occurrence, as well
as the similar petrographic character of the intimately associated
basalts suggest that all of these Tertiary Arctic floras are essentially
similar in age, although it is clear that in Spitzbergen, Alaska and
probably elsewhere, more than a single horizon is represented. Heer,
the pioneer in this field, called them Miocene, just as Lesquereux
called the Fort Union and Wilcox floras Miocene, but the Arctic Ter-
tiary floras are certainly older than Miocene and younger than Ft.
Union. This is indicated by the determination of the so-called Kenai
flora of Alaska as of upper Eocene age, and if any one of them 1s
proved to be upper Eocene none of the others can be older than
middle Eocene or younger than Oligocene. Collateral evidence of their
age is furnished by the age of the greatest extension of subtropical
floras into the Temperate Zone, which is in upper Eocene ( Jackson)
to middle Oligocene (Vicksburg) time.
Plants or coal of Tertiary age are found at the numerous widely
distributed localities shown on the accompanying sketch map (fig. 4).
These completely encircle the pole and reach to within 85° of it
(Grinnell Land). These will be treated at greater length than the
older floras because in some cases they are more extensive and also
because they consist very largely of species belonging to existing
genera, and hence can be discussed more intelligently than the older
floras.
It may be well at the start to dispose of an oft quoted assertion,
as for instance “most of Heer’s determinations were based upon
leaves, which give no data for generic identification’ (Gregory, op.
cit., p. 413). I would readily admit that much of Heer’s material was
fragmentary, that he was over sanguine in some of his determinations,
* Gregory makes much of this idea, which as we have seen is easily disproved.
Gregory, J. W., Congres Géol. Intern. Compte rendu Xéme Session Mexico,
1906, p. 413, 1907.
NO. O PAST CLIMATE OF NORTH POLAR REGION—BERRY [I
SCALE FOR AREA
100,000 SQUARE MILES
LAMBERT AZIMUTHAL FQUAL AREA PROJECTION
For class use in Geography, History, Civics, Economics, etc. Prepared by J. Paul Goode. Published by The University of Chicago Presa, Chicago, Ill.
Copyright 1920, by the University of Chicago
Fic.4.—Location of Tertiary northern floras.
1. Commander Islands 21. Prince Patrick Island
2. Japan 22. Melville Island
3. Sakhalin Island 23. Bathurst Island
4-8. Eastern Siberia 24. North Devon
g-11. Northern Siberia 25-26. Ellesmere Land
12. Central Siberia 27-28. West Greenland
13. Vancouver Island 29. New Siberian Islands
14. British Columbia 30. Nova Zembla
15-18. Kenai 31. Spitzbergen
19. Mouth of the Mackenzie 32. Iceland
20. Banks Land 33. North Ireland and Mull.
TZ SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
and described a great many more species than he should have done.
Some genera do not have a characteristic leaf form, but to make such
a statement of genera such as Liquidambar, Betula, Corylus, Ulmus,
Platanus, Sassafras, Liriodendron, Acer, Potamogeton, Cornus, and
Nymphza, to mention but a few of those recorded from the Arctic
Tertiary, is the height of misunderstanding. Moreover, as I pointed
out in 1922 (op. cit., p. 4): “ Plant fossils have this merit aside from
any question of botanical identification, and this feature seems to have
been lost sight of by numerous critics of paleobotanical practise:
that the size and form of leaves, their texture, the arrangement and
character of their stomata, and the seasonal changes in wood, afford
criteria that are quite as valuable climatically even though the species
or genus to which they belong remains undetermined.” Furthermore,
a great many of the generic determinations are corroborated by fruits
and seeds, as for example, the genera Vitis, Acer, Nyssa, Hicoria,
Juglans, Liriodendron, Fraxinus, etc.
As recorded in the literature the number of species varies from
the single Pinus recorded from Bathurst Island, 5 species from Elles-
mere Land, 6 species from Banks Land to 55 species from Iceland,
168 species from Spitzbergen, and 283 species from Greenland, the
last being greatly overelaborated. I have shown * that Heer’s 30 species
of fossil plants from Grinnell Land (Lat. 81° 42’) represent not more
than half that number; and that Viburnum, Alnus, Ulmus and Tilia
represent Populus and Corylus. As thus revised the Grinnell Flora
contains nothing extraordinary unless it be the supposed Nymphzea
rootstock and this may really belong to one of the plants represented
by fragments of grasses or sedges.
I will consider only the four most extensive of these floras in any
detail. These are Iceland, Spitzbergen, Greenland and Alaska.
The Icelandic flora is preserved in tuffs, along with fresh-water
diatoms, Unios, Potamogeton; and the wood and branches appear
to have been broken off and buried by showers of ashes. The woods
show sharply marked seasonal rings; and conifers, willows, alders,
birch, and hazel are prominent. The only plants certainly determined
that might not justly be considered cool temperate are the following:
Platanus, Liriodendron, Acer, Juglans, Ginkgo, Fraxinus, Hicoria.
Representatives of all of these except Ginkgo, which is not a native,
and Liriodendron, which reaches its northern limit in southern New
England, are hardy in northern New England (Platanus) or eastern
Canada (Acer, Juglans, Hicoria, Fraxinus) at the present time.
1 Berry, Edward W., Proc. Amer. Phil. Soc., Vol. 61, pp. 8-9, 1922.
NO. 6 PAST CLIMATE OF NORTH POLAR REGION——BERRY I
ios)
The Spitzbergen flora comes from two horizons and the two total
108 species and are not essentially different in facies. They are as-
sociated with coal seams and are clearly continental palustrine associa-
tions. There are 4 ferns, a Ginkgo, 27 conifers, 27 monocotyledons
and 80 dicotyledons. Three woods described by Gothan show marked
seasonal rings. The warmer elements are Taxodium, Platanus, Jug-
lans, Nymphza, Magnolia and Nyssa. Here also oaks, hazels, willows,
poplars and conifers predominate. There is not a single tropical or
subtropical type and not one justly considered warm temperate.
The Greenland Tertiary flora comprises 283 nominal species and
includes 8 fungi, 1 moss, 1 lycopod, 1 equisetum, 19 ferns (all tem-
perate types), 1 Ginkgo, 28 conifers, 21 monocotyledons and 202
dicotyledons. The petrified coniferous woods show well marked
seasonal rings and the only genus that is seemingly out of place in
the far north is Taxodium, whose abundance in all Arctic floras and
in proved temperate floras of other regions and other horizons shows
that it was not out of place here. The monocotyledons include mostly
miscellaneous leaf fragments, not generically determinable, as well
as two supposed palms (Flabellaria). It has frequently been pointed
out by others as well as by myself that the nature of the last cannot
be considered as proving the presence of palms. The dicotyledons
are very much overelaborated. Probably 100 species is nearer the
correct figure than the 202 which Heer differentiated.
In Greenland as in all known Tertiary Arctic floras the leaves of
willows, poplars, birches, and hazels predominate, but there are many
other genera whose identification cannot be disputed, such as Liquid-
ambar, Alnus, Fagus, Quercus, Ulmus, Platanus, Sassafras, Fraxinus,
Cornus, Liriodendron, Acer, etc. Vitis is represented by both leaves
and seeds, and other genera also show fruits. The genera that appear
to me to be highly questionable are the following: Castanea, Juglans,
Pterocarya, Benzoin, Laurus, Myrsine, Apeiobopsis, Pterospermites,
Zizyphus, Colutea, Dalbergia, Diospyros, Sapindus, and several others.
I base this conclusion on the fossils and not on the probabilities of
their presence. Some, such as Zizyphus and Ficus clearly do not
represent those genera, in fact Heer’s discussion shows his lack of
conviction of the latter and he queried his determination.
Heer devoted considerable space to a discussion of the climatic
significance of this as well as other Arctic floras and concluded that
the Greenland plants indicated a mean annual temperature of 53.6° F.,
or a considerably lower figure than he estimated by the same methods
for the supposed contemporaneous flora of Switzerland, thus clearly
recognizing a climatic zonation.
14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 8&2
The so-called Kenai flora of Alaska was originally described by
Heer and additions to it have been published by Lesquereux and
Knowlton. Hollick has been engaged in a revision of this and related
floras from Alaska for a number of years, but his results are not
yet published. That from the type locality as listed by Hollick in
1915 comprised but 40 named species and contains not a single
tropical or subtropical type. Associated with the plants are thick
coal seams and fresh water mollusca (Unio, Anadon, Amicola, Mel-
ania), as well as beetle elytra.
He states in a recent letter that localities in the southeastern coastal
region of Alaska (Alexander Archipelago) have yielded a Tertiary
flora that is distinctly indicative of warmer climatic conditions than
those from farther north, including cycads, palms, and such dicoty-
ledonous genera as Anona, Dillenia, etc., but he has not yet determined
whether they are the same or different in age. In either case they sup-
port the conclusion that climatic zoning is indicated.
As listed by Knowlton* in 1919 the Kenai flora (so called)
comprised about 120 species. The most abundant forms are willows,
oaks, poplars, walnuts, beeches, birches, hazels, and alders—dis-
tinctly temperate, and cool rather than warm temperate types. Per-
haps the most abundant plants individually, certainly the widest rang-
ing geographically in northern latitudes (Holarctica), are the leaves
of hazel bushes (Corylus). Of the 54 genera of Knowlton’s list, the
following nine are not present in the existing flora of North America:
Ginkgo, Glyptostrobus, Taxites, Hedera, Paliurus, Elaeodendron,
Pterospermites, Trapa, and Zizyphus,
It may seem that I am juggling the evidence in omitting these nine
genera from further consideration, but let me point out that the three
of these about which there seems to be no doubt regarding their iden-
tity, namely, Ginkgo, Trapa, and Glyptostrobus, are all temperate
types in the existing flora. The remaining six genera are under more
or less suspicion of quite a different order from any differences of
opinion among paleobotanists regarding the identification of the hazels,
birches, alders, etc., with which they are associated. Opinion might
differ as to whether a particular species of the latter was a Betula or
Alnus, an Ulmus or a Carpinus, or a Planera; or whether one or
several species of Corylus should be recognized as distinct species ; but
opinion is unanimous that the choice is thus narrowed, whereas in the
case of such things as Tavites—all any one knows is that it represents
* Hollick, A., U. S. Geol. Surv. Bull. 587, pp. 88-89, 19015.
* Knowlton, F. H., U. S. Geol. Surv. Bull. 696, pp. 786-780, 1910.
No. 6 PAST CLIMATE OF NORTH POLAR REGION—BERRY I5
some Conifer. Why waste time trying to explain the climatic signifi-
cance of Paliurus, a mostly extinct genus, when the particular fossil
is probably not a Paliurus; or why concern oneself with an Arctic
species of Zizyphus when the form in question is probably a Ceano-
thus? I ask, can any one prove that the form-genus Pterospermites
is genetically related to the existing genus Pterospermum? or that
Elaeodendron is a sound botanical identification? I think not!
On the other hand, the great mass of not only the Kenai but of all
the Arctic Tertiary floras are the readily recognizable, normal units of
a natural assemblage, which individually leave but slight room for dif-
ferences of opinion regarding their identity. If fruits chance to be
found in association with the leaves, they are such things as birch or
alder cones, never the fruits of the “suspects” above mentioned.
Of the remaining genera listed in the Kenai flora, all but the follow-
ing six are represented in the existing flora of Canada: A¢sculus,
Diospyros, Ficus, Liquidambar, Sequoia, and Taxodium. It may be
said of these that the #sculus may not be an Zsculus, but a Hicoria;
that the two species that have been referred to Ficus do not belong
in that genus; and that Sequoia is on the verge of extinction at the
present time and its modern range bears little relation to its former
range. The case of Sequoia is of especial interest in its bearing on my
thesis. Formerly a Holarctic type, it survives today in a most re-
stricted area particularly favored by humidity.
The remaining genera of the Kenai flora appear to be determined
with reasonable certainty. Not only are 39 of these represented in
the existing flora of Canada, but the following are still represented in
the existing flora of Alaska, or adjacent areas in northwestern
Canada, or as far north as Labrador and Hudson Bay in eastern
Canada: Abies, Acer, Alnus, Alnites, Andromeda, Betula, Carex,
Corylus, Equisetum, Fraxinus, Myrica, Osmunda, Phragmites
(grass), Picea, Pinus, Populus, Prunus, Pteris, Quercus, Sagittaria,
Salix, Spiraea, Thuites, and Vaccinium.
Seventeen of the Kenai species are conifers, and the only types that
would seemingly be out of place in a cool temperate climate with
well-distributed moisture are Liquidambar, Paliurus, Taxodium, and
Zizyphus. I have already given reasons for discrediting the deter-
minations of some of these, and all of them have frequently been
found fossil in temperate assemblages.
The significant feature about these Eocene Arctic floras is that
they show a comparable northward swing of not alone their northern
limits, but also of their southern limits, which in turn is comparable
to the northward advance of the Jackson flora that I have considered
16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
to be of the same age. The Jackson flora reaches Latitude 37° North.
The most similar existing flora to that of the Jackson does not extend
above Latitude 26° North, and then only under especially favorable
conditions of situation with respect to warm ocean currents. This is
a difference of 11 degrees. The flora of the Jackson was, moreover,
a coastal flora, and I have not the slightest doubt but that had the
Mississippi embayment extended five degrees farther North, its shores
would have been clothed with the same Jackson flora, for at that time
similar floras are found in the Paris Basin in Latitude 49° North,
in southern England in Latitude 51° North, and along the expanded
Mediterranean sea of the Old World. ;
The southern limit of the contemporaneous “ Arctic flora” is about
Latitude 45° North in North America (British Columbia), and about
57. North in Europe (Isle of Mull). It seems to me that the essential
concordance of these facts is significant, and whatever may be thought
of them, it would certainly seem to be difficult for any one to claim that
these various Eocene floras mentioned do not show a climatic change
in passing northward from the equator toward the pole. Moreover, at
present—a time of, in many ways, an abnormal climate in a geologic
sense; with rather sharp zoning, although not nearly so sharp as the
textbooks would have us believe; a time of almost, if not quite, un-
precedented land expansion in the Northern Hemisphere, which I
believe expresses a casual relationship—the reliable members of these
IXocene Arctic floras range much farther southward than they did
in late Eocene time.
EXISTING ARCTIC FLORAS
Greenland is the most illuminating of Arctic Lands because it is
much the largest, and therefore more likely to preserve endemic
species, and to receive immigrants from other Holarctic lands. Al-
though mostly covered by ice which rises to an altitude of more than
8,000 feet in the interior, it has island peaks (nunataks) with recent
plants. Moreover the northeastern part appears never to have been
glaciated.
About 400 species of recent vascular plants have been recorded
from Greenland and at the south trees may reach heights of 10 or
12 feet. North of the Arctic Circle the number of plants is fewer,
but Ostenfeld (1925) records 125 species north of Latitude 76° and
108 between Latitudes 78° and 80°, including 2 equisetums, a lycopod,
3 ferns, 32 monocotys and 70 dicotys, including Salix and Vaccinium.
In an earlier paper (Ostenfeld, 1923) this author describes the flora
of the north coast and records 70 species of plants from Latitude 82°.
NO. 6 PAST CLIMATE OF NORTH POLAR REGION—BERRY 17
This brief statement will be sufficient to indicate that there are other
and more important factors than cold. The almost entire absence
of vascular plants (a single species as I recall it) from Antarctica
shows the part geography plays in the problem. The absence of trees
in Lapland (Kihlman) shows the part taken by cold desiccating winds.
The northern limits of many tree species in coastal Alaska and Nor-
way will indicate the ameliorating climatic effects of warm ocean cur-
rents and humidity.
EXISTING ARCTIC CLIMATES
This is a complex subject which cannot be discussed in this con-
nection beyond pointing out certain observed facts which support the
thesis of the present discussion. These are the slower heating and
cooling of water bodies as compared with land areas, with their respec-
tive influence on air temperatures and pressures, their influence on
the amount of water vapor in the air and the resulting effect of
humidity on equability.
The climatic influence of the northward drift of oceanic waters
may be illustrated by the course of the present day isotherms over the
north Atlantic, a somewhat hackneyed illustration but nevertheless the
most striking. I am showing a few of the isotherms for January and
July in figures 5 and 6. Those for January which show the full effect
of the rapid radiation and quick cooling of the land, contrast most
markedly with the slow radiation and cooling of the ocean. At this
time the zero isotherm reaches Latitude 35° in Asia and about Lati-
tude 74° north of Norway, a difference of 39°. Much the coldest place
is in northern Siberia which is 10° to 20° colder than at the pole itself.
The — 30° isotherm reaches almost to the pole north of the Atlantic
and swings to approximately Latitude 55° in Siberia—a difference of
about 35° of latitude. The midsummer isotherms naturally smooth
out these curves somewhat but even at this season the isotherm of 5°
swings from about 62° in southern Greenland to 80° just west of
Spitzbergen and the oceanic effect is clear as far eastward as Nova
Zembla.
A few figures quoted from Sir John Murray’s calculations will
serve to emphasize the relations referred to. The energy radiated
by the lowering of the temperature of a cubic meter of water 1° is
sufficient to raise the temperature of more than 3,000 cubic meters of
air 1°, and a second calculation shows that the heat released by lower-
ing by 1° a stratum of water 200 meters deep and of 700,000 square
kilometers area would suffice to raise the temperature of a stratum
18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
of air 4,000 meters deep over the whole of Europe on an average
Or 10
I have not attempted to evaluate the effects of the present ice cap
on Greenland or of the present altitude of the land surface, as all
Fic. 5 Midwinter isotherms at the present time.
I wish to do in this brief discussion is to emphasize in a graphic way
the major thermal effect of land and water.
The Arctic is an oceanic basin and shows a remarkable climatic
contrast with the elevated glacier-covered Antarctic continent. A few
of the probable climatic effects which would follow if the Arctic re-
4
No. 6 PAST CLIMATE OF NORTH POLAR REGION—BERRY 19
ceived ocean waters from the Pacific or across Eurasia from ancient
Tethys, or became ice free during the summer are discussed very
briefly in a subsequent section of this paper.
Fic. 6—Midsummer isotherms at the present time.
SOME PALEOBOTANICAL MISCONCEPTIONS
Although I have on previous occasions emphasized the lack of
climatic value of most of the plant types which paleobotanists have
relied upon as indicating tropical climates, this subject should not be
passed over without some comment in the present connection.
The principal evidence upon which tropical climates have been pred-
icated falls into three somewhat dissimilar categories. First it rests
upon tradition which never had any basis. For example the concep-
20 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
tion that the flora of the Carboniferous grew in a supertropical
climate with a humid atmosphere charged with carbon dioxide, of
which Koppen and Wegener make such specious use, had as its
original basis the 18th century idea that the strange plants associated
with the coal had been swept to Europe from the tropics by Noah’s
flood and the further fact that the habit and venation of certain fern-
like Carboniferous plants, now referred with strong probability to
the Pteridosperms, resembled certain existing tropical ferns.
European students accustomed to the modern accumulations of
peat in high latitudes concluded quite as illogically that peat could
not accumulate in the present equatorial region because of the rapid
oxidation there, so added carbon dioxide to make growth extraor-
dinarily rapid and great moisture to prevent rapid oxidation. The
carbon dioxide stimulation would also conveniently account for the
enormous size of some of the calamites and lepidodendrons as com-
pared with their diminutive survivors the equisetums and clubmosses.
Then Koorders and Potonié described a peat bog from Sumatra and
others have been subsequently described from other tropical lands,
and there has been much readjustment of views, which might have
been accomplished much earlier if the experts on geological climates
had ever visited the tropics or even consulted the report on the peat
deposits of Florida published by the Geological Survey of that state.
There is not space at my disposal to follow the vagaries of opinion,
but it may be stated in the most positive way that temperature or the
position of any region with respect to the equator, that is between
hot or cold climate, is not a factor in the formation of either peat or
coal. Second the tropical idea relies on representatives of long lived,
vigorous groups with very many species, which either in the past or
ii the present have become adapted to a variety of habitats, as is
usually the case in large vigorous groups of all kinds of organisms.
As outstanding examples I may cite just a few types such as the
palms, and figs, or such genera as Cinnamomum and Zizyphus. The
great bulk of the existing palms are tropical and they are one of the
first types of plants visualized when we think of tropical climates,
whether we picture the Arab and his date palms or the South Sea
Islander and his cocoanut palms. Nevertheless certain palms extend
to approximately 39° South in Chile, 44° South in New Zealand,
34° North in California, 35° North in North Carolina and 36° North
in Japan, and commonly are hardy several degrees north of their nat-
ural limits, as in the Sacramento valley in California, or in southern
* Not considering the subtropical arid belts of high pressures.
No. 6 PAST CLIMATE OF NORTH POLAR REGION—BERRY 21
France. The greater limits of cultivated forms is usually not a result
of cultivation so much as it is of selecting the species that will grow
in a particular environment. In nature the proper species is subject
to the historical factor of there having been ancestors in the region
or in a region offering access to the particular region. For example our
native Californian palm (Neowashingtonia) is a plant of sandy alka-
line. soils whose range seems to be conditioned by the geologically late
submergence of the Colorado Desert area, and to bear no relationship
to latitude. In the present tropics certain palms range upward to
nearly 10,000 feet, as in the wet parts of the northern Andes
(Ceroxylon, Geonoma, etc.).
The genus Ficus, to which the cultivated fig belongs, is one with
upwards of 600 existing species of a great variety of habitats, and
with probably as many fossil species, extending back to the dawn of
the Upper Cretaceous. Various members range well into the temper-
ate zone, both geographical and altitudinal. The cultivated fig gener-
ally ripens its fruits in Baltimore. I have seen it in the temperate
altitudinal zone in Bolivia, and Weberbauer* records an altitudinal
range for it through 8,255 feet in Peru.
Cinnamomum is the genus to certain members of which the names
cinnamon and camphor trees are applied. The genus is large and
ranges from the Upper Cretaceous to the present. Although the ma-
jority of existing species are confined to the tropics some extend for
considerable distances into the Temperate Zone, in fact the com-
mercial supply of camphor comes in large part from Formosa and
Japan, and the tree is hardy in the southern parts of the latter country.
Introduced into Florida it has been widely seeded by birds and is
perfectly hardy throughout that state.
Zizyphus is a large genus also going back to the Upper Cretaceous,
whose present center of population is southern Asia and the Sunda
Islands. The new world species are practically confined to the tropics,
but in the old world there are distinctly temperate species in southern
Europe and eastern Asia. It has run wild in Louisiana, and charac-
teristic fruits occur in the Pleistocene of the Atlantic coastal plain as
far north as Long Branch, New Jersey. Obviously as a fossil.
Zizyphus entirely lacks a tropical significance.
A third source of error is the common assumption that because a
particular type of plant has its home in the equatorial zone it is neces-
sarily a tropical plant. The type most frequently alluded to in fossil
Arctic floras as indicative of a once tropical climate is the tree ferns,
the term embracing a variety of species in several genera,
* Weberbauer, A., Archiv. Asoc. Peruiana Progreso Ciencia, tomo 2, p. 60, 1922.
i)
i)
SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
As a matter of fact tree ferns reach their maximum development
in temperate rain forests, as in New Zealand (Lat. 40° S.), or in
similar situations in tropical uplands, as was pointed out by Alexander
von Humboldt over 100 years ago. They reach their greatest pro-
fusion in South America in the temperate part of the montafia zone
of the eastern Andes. They grow luxuriantly on the mountains of
central Africa at altitudes where they are buried in snow for part of
each year, and as fossils their climatic significance is wet temperate
and not tropical.
There are a great many other genera or species in the same category.
I have seen Anonas and Ingas (cultivated) at 10,000 feet in the Andes
perfectly hardy, and a large number of generic types that are com-
monly thought of as lowland tropical above the tropical altitudinal
zone—such things as Dodonaea viscosa, Sapindus saponaria and
Swietenia mahagoni. In fact it was my own observations in the Andes
that first turned me from the paleobotanic tropical tradition.
Another misinterpreted type is the Gleichenia type of ferns (now
segregated in several genera) very common in the Cretaceous floras
of Greenland, but largely absent from the northern hemisphere in
recent floras. Although commonly confined to low latitudes at the
present time, it is by no means confined to the tropical altitudinal
zone ; in fact, where I have seen it (Yungus of Bolivia) it is promi-
nent above the tropical zone, as it is also in Hawaii, Peru, Ecuador,
Asia, etc. Representatives reach 54° South in Chile and 40° South
in New Zealand.
All this is related in any account of fern distribution (e.g., Die
naturlichen Pflanzenfamilien, 1902), and still Gleichenias, along with
palms, cycads, and tree ferns always appear in the paleobotanists
tropical repertoire.
I suppose that constant reiteration of facts like the foregoing will
have to be continued over many years before the news reaches those
who write on paleoclimatology, and at least another generation will
elapse before writers of geological text books cease to talk about the
tropical climate of Tertiary Greenland.
Juniperus communis Linné is found as far north as the North Cape,
which is at least 20° farther north than any other member of the
family Cupressinaceae is found in the Eastern Hemisphere (Nat-
horst, 1911). Sassafras, of the mostly tropical family Lauraceae, ex-
tends northward to southern Maine, or about 13° beyond the bulk
of the family. Diospyros, of the mostly tropical family Ebenaceae,
extends northward to southern Connecticut, or about 12° beyond the
bulk of the family.
No. 6 PAST CLIMATE OF NORTH POLAR REGION—BERRY 23
Nor must it be lost sight of that at those times in the past when
certain groups were varied and abundant, as were the seed ferns in
the Paleozoic or the cycads in the Mesozoic, they were quite likely
to have shown the features of dominant organisms, both plant and
animal, and to have occupied more environmental niches than the
depleted survivors of the cycad phylum do at the present time.
In Newfoundland and western Labrador the larch (Larix ameri-
cana), the balsam poplar (Populus balsamifera), the paper birch
(Betula papyrifera), and the balsam (Abies balsamea) fail to reach
the Straits of Belle Isle (52°) whereas they all extend far above
Latitude 60° in Alaska and the first crosses the Arctic Circle. Podo-
carpus just fails to reach the Tropic of Cancer in Cuba. A Chilean
species reaches 42° South Latitude in Chile. The northern limit of
forests crosses the Arctic Circle in Alaska and reaches 70° North
Latitude in Norway, the latter 20° north of the tree line on the
Atlantic coast of North America.
EXPLANATION, OF PAST ARCTIC CLIMATES
It is perhaps fatuous to point out that climate, either present or
past, depends upon a variety of factors, both cosmic and terrestrial.
Of the former the only one that is of practical importance is solar—
that is, radiant energy from the sun, since it is inconceivable that
other heavenly bodies or the introduction of kinetic energy by meteor-
ites exert any appreciable effect.
The amount of solar energy reaching the earth depends upon the
sun’s activity, which is variable; on the distance of the earth from the
sun, which is also variable ; and more practically in so far as terrestrial
climates are concerned, on the condition of the earth’s atmosphere,
especially with respect to the amount of ozone, water vapor, carbon
dioxide, and dust present, all of which again are variable. The lati-
tude, determining the angle of incidence of the sun’s rays, is an obvi-
ous factor, as is also the geographic pattern and the topography, in-
cluding altitude under the latter. The geography determines whether
the sun’s energy falls on the land or the water, it determines the
temperature gradient between the equator and the poles and the con-
sequent force of the planetary winds and ocean currents, and in less
obvious ways is of the greatest significance, as the following illus-
tration will make clear.
The North and South Equatorial currents in the Atlantic are so
situated that the South Equatorial, the stronger and the larger of the
two, is divided by Cape San Roque into a larger, northern or Guiana
current; and a smaller, southern or Brazil current. Some authors,
24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
e. g., Guppy, are inclined to consider the South Equatorial as bipartite
throughout, calling the Guiana current the Main Equatorial current.
The point is immaterial in the present connection since all I desire
to show is that the shape of eastern South America and the latitude of
Cape San Roque are purely fortuitous in so far as their relation to
climate is concerned, and yet if the latter had happened to lie a few
degrees north of its present position much of the water that ultimately
contributes to the Gulf Stream would have turned southward to aug-
ment the Brazilian current, and the climate, especially of Europe and
the Arctic, would be profoundly modified. It has been estimated that
if Cape San Roque were 2° north of its present position there would
be a shift of 4o% of the Equatorial current which would be deflected
southward instead of northward. The same results would be attained
if the southern trades were not stronger and more constant than the
northern trades, because of the relative amounts of land and water
in. the northern and southern hemispheres.
Scant attention will be devoted to the various theories that have
been advanced to explain geological climates. These range from that
of Croll, in its original or modified form, based upon the eccentricity
of the earth’s orbit and the obliquity of the ecliptic, which was doubt-
less a factor at all times, but hardly a controlling one ; through those
theories that rely on changes in the atmosphere, such as alterations
in the amount of carbon dioxide (Tyndall, Arrhenius, Chamberlin)’
amounts of volcanic dust (Humphreys), to the extreme form of the
hypothesis advanced by Manson, and elaborately defended by Knowl-
ton, that a combination of cloudiness progressively diminishing dur-
ing earth history, and a terrestrial control due to a cooling earth,
instead of a solar control as at present, are the primary factors which
explain past climates. Finally there are those highly speculative
hypotheses such as Chamberlin’s reversal of the oceanic circulation,
and a group which predicate a wandering of the poles in various
ways, now fashionable in the revived form put forward by Wegener.
I have quite possibly omitted other proposals that might be men-
tioned, and I have now to mention the theory, if it can be called a
theory, which is the main thesis of the present paper, namely: that
it seems to me possible to interpret geological climates in the light of
demonstrated changes in topography and geography, including under
the latter differences in the distribution of land and water and the
transfer of energy by currents.
‘It is of interest to note that Neumayr in 1883 pointed out that excesses of CO,
would be impossible since the absorption by the oceans would maintain an almost
perfect balance.
NO. 6 PAST CLIMATE OF NORTH POLAR REGION—BERRY 25
This idea, as applied to the Pleistocene glaciation, was first advanced,
I believe, by Lyell, and in its more general application has been re-
cently put upon a scientific basis by Brooks, with whom I am in perfect
agreement to the extent of the evaluation of these as major factors,
but also in my firm conviction that arm chair philosophy with its
fondness for highly speculative and catastrophic hypotheses, has no
place in a uniformitarian world or in 20th century science, but be-
longs in the medieval age of human thought.
Climate, in a uniformitarian geology, occupies a somewhat anoma-
lous position, which the scientific world has been slow to recognize,
namely, that the history of the human race has been run under cli-
matic conditions which, from the point of view of earth history, are
exceptional. Man was evolved subsequent to the relative elevation
and the great extension of the continents which ushered in the Pleis-
tocene glaciation, and therefore what is normal in human experience,
is abnormal for the bulk of geological climate.
While, therefore, we recognize that the climatic factors and the
meteorological elements are the same now as always, their combina-
tion to form actual climates has depended upon a great many factors,
among the chief of which was the size, shape, position, and relative
elevation of the land masses. It may be remarked parenthetically
that numerous theories of the causes of, or descriptions of geological
climates have been advanced by students ignorant of meteorology,
and also usually ignorant of the relationship of organisms to their
environments, and the last is strikingly true of Koppen & Wegener’s
recent Die Klimate der geologischen Vorzeit (1924).
In attempting, a few years ago, to explain the extension of floras
nearly to the poles during the late Eocene, I relied chiefly on the sub-
mergence of continental areas in the middle Eocene and the resulting
free oceanic connections at that time between equatorial and Arctic
waters, pointing out that these Arctic floras were coastal floras and
therefore under the régime of an oceanic climate.’ Essentially the
same explanation was put forward independently in connection with
Jurassic climates a few months later by Kerner von Marilaun.’ An
additional and important factor has since been brought forward by
Brooks,’ who points out that the temperate gradient is a simple func-
tion, whereas the influence of the ice increases as the square of the
* Berry, Edward W., A possible explanation of upper Eocene climates. Proc.
Amer. Phil. Soc., Vol. 61, pp. 1-14, 1922.
2 Kerner von Marilaun, F., Sitz. k. Akad. Wiss. Wien, 1922.
* Brooks, C. E. P., The problem of mild polar climates. Quart. Journ. Roy.
Meteor. Soc., Vol. 51, pp. 83-94, 1925.
26 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
radius. Hence a coincidence of minor factors sufficient to effect an
overturn in the one or the other direction, that is, toward ice forma-
tion or melting, would suffice to induce a wide extension of polar
ice, or to prevent the polar regions from maintaining a permanent
ice cap. If this is true then it seems quite probable that there was little
polar ice during those times already enumerated when temperate
floras invaded the polar regions. This would mean profound changes
in the distribution of barometric pressures and consequent wind cir-
culation, and in fact, in all of the elements which constitute climate.
It would mean that in western Greenland, for example, where the
most extensive late Eocene Arctic flora has been found, the present
day glacial anti-cyclonic winds would be replaced by westerly or south-
westerly winds blowing from the relatively warmed waters of Baffins
Bay, and this would satisfactorily explain the details of the floral
facies. This does not mean that there would be tropical climates
in the Arctic or that the region would not be ice bound in the winter
season. The protective effect of snow, and cold sufficient to cause
a cessation of plant activity during the Arctic night are a physiological
necessity. Otherwise most vascular plants could not maintain them-
selves. They tend to die either if active in darkness or if exposed
to desiccation by air and wind when the ground water is frozen.
Regarding the general history of discussions of geologic climates
I believe that most paleontologists who have written on this topic,
especially those dealing with the pre Cenozoic periods, have had little
basis in fact for their speculations. They seem to me to be utterly
oblivious to the great amount of modern work on the distribution
of marine organisms; and their ideas of the climatic significance of a
trilobite, eurypterid, or ammonite is purely a tradition inherited
from the distant past when all strange organisms were associated with
torrid climates.
In stating my belief in a greater uniformity of climate during the
past than obtains at the present I do not wish to be understood as
advocating such unsound beliefs as the entire absence of zonation,
such as many paleobotanists have defended (Jeffrey, Knowlton), or
a similar uniformity throughout all time. Both are equally disproved
both by geological observations and meteorological principles. Jeffrey,
for example (Anatomy of Woody Plants, Chapter XXX, 1917), holds
that the more ancient the epoch the warmer the climate, and that there
has been a gradual and progressive refrigeration during geologic
time ; that the organization of secondary wood in extinct plants furn-
ishes the most reliable evidence of climatic conditions; that toward
the end of the Paleozoic, growth rings appeared in woods in high
latitudes ; that in the Triassic, growth rings were developed ten degrees
No. 6 PAST CLIMATE OF NORTH POLAR REGION——BERRY 27
nearer the equator than had been the case during the Paleozoic ; that in
the Jurassic, the tracheids first developed tangential pitting which
was at the end of the annual ring, and accompanied by storage ele-
ments (wood parenchyma).
None of the statements in the foregoing paragraph are facts of
observation. There is no geological or paleontological evidence indi-
cating a progressive climatic cooling during geologic time, and the
Permo-Carboniferous glaciation was admittedly more extensive than
that of the Pleistocene. The presence or absence of growth rings
exhibits what might be called constitutional variations quite indepen-
dent of climate, not that they really are independent, but two associated
species under an identical climate will behave differently with respect
to this feature of their anatomy. Growth rings appear in some Pale-
ozoic woods many degrees nearer the equator than Jeffrey admits,’
and in marine formations deposited off low coasts so that they cannot
be considered to have been upland types. Several Lower Carbonifer-
ous examples have already been cited. The Paleozoic genus Mesoxy-
lon shows tangential pitting, which, according to Jeffrey, first appeared
in the Jurassic; and the citation of a wood from the Triassic of
Arizona as an argument for the advance equatorward of cooler cli-
mates during the early Mesozoic is particularly disingenuous, as it
is perfectly clear that the growth rings in this case have nothing to do
with temperature, but are due to periodic lack of moisture in that
region, as exemplified by the contemporaneous gypsum deposits.
Similarly in the recent elaborate work on geologic climates by Kop-
pen & Wegener, already alluded to, these authors offer explanations
to account for climates during the successive geologic periods, which
climates have not been proved to have ever existed.
As I have pointed out on previous occasions, paleobotanists in
general have entirely lacked objective experience outside the temper-
ate zone, and have invariably overestimated temperatures. They have
been prone to use the present distribution of the fancied or real rela-
tives of their fossil forms as if temperature were the sole factor in the
environment, and have stopped with the geographic occurrence, with
the apparently simple trust that all lands in the equatorial zone were at
sea level and wet tropical. A sojourn in the Arctic climate beneath
the equator on the backbone of South America would do much to
correct this misapprehension, as would also some experience in the
temperate rain forests of different regions.
I had intended to indicate current conceptions of contemporaneous
paleogeography on the maps showing the plant localities but have
* Several have been named in the preceding paragraph devoted to Mississippian
Arctic plants and others could be added.
28 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
not done so although I did publish such a map for the Eocene in 1922.
This intention was abandoned for the reason that it was not possible
to compile maps that did not cover too much time nor in which the
extrapolation was not so great as to destroy any real value.
Arldt has compiled maps which represent a synthesis of opinions
and showing the areas of agreement and disagreement among special-
ists and to these the reader is referred. The debatable North Atlantic
continent and the Gondwana continent would, if they ever existed,
have had a profound effect on climate. Whether or not they were
ever realities [ am not prepared to say. I can, however, make the
following statements with a considerable degree of certainty, namely :
That there was a wide extent of land in the Northern Hemisphere from
late Mississippian through the Permian. That the Arctic was Jand-
locked in early and middle Triassic and that there was a wide trans-
gression of the sea in the Neo Triassic. That the maximum Jurassic
transgression was about Oxfordian; that of the Lower Cretaceous was
in the Neocomian ; that of the Upper Cretaceous was in the Emscher-
ian; that the late middle Eocene was a time of wide sea transgression
and low lying lands; and that during the Miocene, the age to which
Heer assigned the Arctic Tertiary floras, the amount of land in the
Northern Hemisphere was nearly as great as it is at the present time.
It will be seen that there is a correspondence between times of sea
extension and Arctic floras and times of land extension and no traces
of Arctic floras. This correspondence is not exact, and so little of
paleogeography is objective, that I would not want to appraise it for
more than it is worth, but in so far as it 1s known it does offer
corroboration of my thesis.
I had expected to attempt an estimate of the meteorological condi-
tions at the various times at which fossil floras are found in the Arctic,
but after abandoning any hope of getting reliable paleogeographic
data I have also abandoned the former. Brooks has published some
interesting meteorological estimates using as a basis those parts of
Arldt’s maps where authorities agree, but it should be pointed out that
majorities are quite as likely to be wrong in science as in politics, and
if generalizations are valid (which of course they are not) then
minorities are usually right.
There are, however, a few considerations that may be put forward
as having a high degree of validity, namely the importance of ice as
a third factor, added to the long recognized rotational (planetary) and
geographic (distribution of land and water and altitude) factors
in influencing the distribution of pressures and consequently of pre-
vailing winds. And also the effect of the volume of fresh water car-
No. 6 PAST CLIMATE OF NORTH POLAR REGION—BERRY 29
ried into the Arctic basin by rivers in the formation of ice and the
effect of current-borne ice in maintaining subnormal density and con-
sequently the identity of the present day cold currents as they move
southward. Once they become of normal density they disappear below
the surface and lose their climatic influence.
Another factor of considerable importance climatically, especially
in connection with the theory of Brooks, is the amount of reflection
from the earth’s surface. I do not have the exact figures, but estimates
given to me orally by W. J. Humphreys, are about 7 per cent from
land or water and about 70 per cent, or ten times as much, from the
surface of snow and ice. If there has been the wide fluctuations in
polar ice as Brooks predicts, then reflection is a factor which can not
be safely neglected.
At the present time in high latitudes the prevailing wind circula-
tion is easterly with a southward moving component at the surface.
If the ice cap were gone we would have westerly winds in high lati-
tudes with a poleward component at the surface.
If Bering Strait was open and less shallow, a great volume of warm
Pacific water would pour into the Arctic and greatly ameliorate the
climate, as would also be the case if a Cretaceous seaway bisected
North America, or a Devonian or Eocene seaway bisected Eurasia,
such as are shown on current paleogeographic maps. If the best avail-
able sources are utilized in plotting Eocene seaways nearly all the
Tertiary coal occurrences and floras in the Arctic range themselves
along the easterly coasts of such seaways.
CONCLUSIONS
The major factor in the polar extent of temperate floras is
not primarily the direct effect of temperature so much as it is the fact
that above 32° F. water is a liquid and below 32° F. it is a solid. Asa
Gray said “ Plants are the thermometers of the ages.” I have no
doubt that terrestrial vegetation when properly interpreted is the safest
guide to geological climates, but as thermometers they are pretty
poor and we have no means of calibrating them,
There is no unequivocal botanical evidence of tropical or subtropical
climates at any time in the Arctic. There is no evidence from paleo-
botany of a lack of climatic zonation at any geological period from
which fossil plants are known, although at such times the evidence
points to a relative mildness and a lack of sharp zonation, as com-
pared with the present.
The distribution of the known fossil Arctic floras with respect
to the present pole proves conclusively, as Seward pointed out in
1892 (p. 53), that there could have been no wandering pole.
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 82, NUMBER 7
THE ATMOSPHERE AND THE SUN
BY
H. HELM GLAYTON
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(PUBLICATION 3062)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
JUNE 9, 1930
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The Lord Baltimore Press
BALTIMORE, MD., U. S. A.
tHE ATMOSPHERE AND “AE SUN
By H. HELM CLAYTON
CONTENTS
PAGE
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SUMmimmatiyewscnts Siests Sela e eee ee eee oie Se oom pie carseat 48
INTRODUCTION
This paper is the fifth of a series giving the results of investiga-
tions of the relation of solar activity to atmospheric changes. The
earlier ones were published as Smithsonian Miscellaneous Collec-
tions, Vol. 68, No. 3; Vol. 71, No. 3; Vol. 77, No. 6; and Vol. 78,
No. 4. The author has been stimulated to continue these researches
because he believes in their great importance. The interest of
Dr. C. G. Abbot and the sympathy and aid of Mr. John A. Roebling
have encouraged him in the task and enabled him to undertake much
work that otherwise would not have been possible. Miss M. I. Rob-
inson has aided in the calculations needed for the discussion.
1 SOLAR CHANGES
It has long been known that spots appear on the surface of the
sun and that the number and size of these spots varies from day
to day, from month to month, and from year to year. More recently
it has been discovered by Dr. C. G. Abbot and his associates that the
radiation coming from the sun varies ; so that, in general, it is known
that the sun is hotter when there are many spots on its surface than
when there are few or none.
There is also evidence that the heat of the sun varies from day
to day and from week to week in short cycies of change. The most
convincing evidence of this fact is the comparison of measurements
SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 82, No. 7
Z SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
of solar radiation made at observatories thousands of miles distant
from each other, one in the northern hemisphere and the other in
the southern hemisphere; so that the chance of both being affected
by the same weather changes becomes very small. The solar radia-
tion reaching the earth is measured in calories per square centimeter
per minute, and averages about 1.940 calories. Table 1 shows a com-
parison of observations of solar radiation made simultaneously in
northern Chile and in the United States (first in California and then
in Arizona) during the years 1918 to 1924. The table shows the
frequency of different values observed in the United States for each
increase of .010 calorie in Chile.
TABLE I1.—Comparison of Solar Radiation Values in Chile and the United States
(Number of Cases)
Values in Values observed in Chile
United —— A —
States I.910-9 1.920-9 1.930-9 1.940-9 1.950-9 1.960-9
TAOOOHO sis yrecs aerate ousve I 6 6 6 fe) 0
T#QOO-O ste se ersichevsie sie II II I a) O 0)
TQLO=Ol wei tastne ee rs 20 25 II 5 4 (0)
TOZO=Oie thoy enters a 18 38 21 5 4 2
THOS O=OCraihe ee Steere 7 23 29 II II (0)
THOYOHOe Sse ae 4 6 12 15 16 4
TO5O=Onie,c selehret ee (a) 4 10 10 13 6
THOOO=O lenis xe ete eh () I 3 4 7 5
TO70-Oma nance see (a) I (6) I 3 2
If there were no relation between the measurements at the two
stations, the observed values would be scattered through the dif-
ferent classes at random. The tabulation shows that a random dis-
tribution does not exist ; but for each group of observations in Chile,
there is a maximum near the same values in the observations in the
United States. There is, therefore, a progressive displacement of
the maximum frequency as the solar values increase from 1.g10-9
to 1.960-9, or nearly three per cent of the mean value. The probable
error of the measurements is +.006 calorie; so that the solar varia-
tion during the interval covered by the observations was more than
eight times the probable error of each group of observed values.
Since variability in solar radiation has been questioned by some
investigators, it is well to state that the evidence of this variability
rests on three fundamental and independent facts:
(1) The changes in radiation are alike when measured at two
widely separated stations, allowing for variations from a middle
value due to errors of observation.
NO. 7 THE ATMOSPHERE AND THE SUN—CLAYTON 3
(2) The changes both of short period and of long period in solar
radiation are related to visible changes in the number and area of
spots, faculae and flocculi seen on the sun.
(3) The changes in solar radiation are correlated with other phe-
nomena such as certain changes in terrestrial magnetism, in radio-
receptivity, and meteorological changes which are known by other
evidence to be related to solar conditions.
The critics of solar variability have pointed out that the measured
variations have decreased as the accuracy of the observations in-
creased and that in the earlier observations the effects of water vapor
DAYS BEFORE DAYS AFTER
G 4 2 0 i i f i ie i i a. re 20
1.952 | | | Ceriter |
of sun
1.951
(
1
\
1.950 wogeedeee.,
F SHOVE eseStesocc: Se
ss Neakk= nese ssaeeoer el
aii am
CAAA ES ——s unit
Re aces as Glam ae) Be
CET. (ate
a — {ff SAH a
Fic. 16.—Pressure in 3.77-year period.
ress here indicated would carry the wave from pole to Equator in
one period.
In figure 16 is found a period in which the unit of time is years
instead of days. The period of oscillation in pressure is about four
years. It was taken to be 3.75 years, or one-third of a sun-spot
period of 11.3 years, and averages were made for each three suc-
cessive oscillations of 3.75 years. The continuous curves show the
NO. 7 THE ATMOSPHERE AND THE SUN—CLAYTON 25
averages. The letters a, b, c, etc., show successive maxima. The data
were derived from ‘“ World Weather Records” and cover the con-
tinent of Asia where the data are more complete for different lati-
tudes than in North America.
It is seen from the plot that the maxima and minima of the period
occur first in high latitudes and successively later at stations nearer
the Equator, at least down to about 30° latitude, taking about three
years to move from Obdorsk, 66° N., 66° E., to Ley, 34° N., 77° E.
In the equatorial belt between 20° N. and 20° S. the maxima and
minima occur simultaneously at all stations as shown by the results for
Madras and Batavia. However, from figure 16 it is seen that the
pressures at Alma Ata, 43° N., and at Batavia, near the Equator, are
opposite in phase, which is further evidence that this wave traversed
go° of latitude in one period of about 3.75 years.
A recent study of 2- and 34-year waves in temperature by Ernest
Rietschel * shows a rather complex movement indicating a combina-
tion of standing and moving waves.
That the law of wave progress quoted above holds true in the
Southern Hemisphere as well as in the northern is shown by the
rate of progress of a temperature wave of about 18 days shown
plotted on page 223 of “ World Weather.” “ This wave progressed
from Santa Cruz, 50° S., to Cuyaba, 16° S., in seven days, a rate
which would carry it from pole to Equator along a meridian in one
period of 18 days.
The rate of progress of a 7.5-year wave is indicated in figure 22
where the maxima and minima of the waves occur successively later
at Stykkisholm, Rome, and Calcutta, the minima and maxima at
Calcutta being about 7 years later than at Stykkisholm,
These facts render it evident that the rate of latitude displace-
ment is a general law for periodic oscillations of all lengths. This
law may be stated as follows:
Law of latitude displacement of periodic waves—Periodic oscil-
lations in atmospheric conditions progress in latitude from point to
point along a meridian at a rate that would carry the wave from
pole to Equator in one period, whatever the period of oscillation.
It is probable that the law of displacement in longitude is equally
simple. Figure 14 shows that the 7-day wave progressed in longi-
tude about 180°, or half around the world, in seven days.
*Die 3-34 jahrige und die 2 jahrige Temperaturschwankung, von Ernst
Rietschel. Geographical Institute of the University of Leipzig, Vol. 1V, No. 1,
1920.
VoL. 82
SMITHSONIAN MISCELLANEOUS COLLECTIONS
‘0061 ‘€6gr ‘Sggr sysody ‘porsad se9k-$°Z Ul URS WOT dInssaid Jo soinjredaq—ZI ‘dIq
LEG)
/ f- U
Lh
NO. 7 THE ATMOSPHERE AND THE SUN—CLAYTON 27
A proportional rate of progress appears to occur in the periodic
wave of about 7.5 years. Figure 17 shows the centers of oscillation in
a 7.5-year wave on a world map. This map is derived from har-
monic values computed from groups of three periods between 1883
and 1913 at 117 stations scattered over the world. It shows the
centers of oscillations at the epochs, 1885, 1893, 1900, etc. Con-
tinuous lines show equal values above normal and broken lines show
equal values below normal. It is not possible with available data
to follow the progressive movement of all the centers, but the center
over Greenland shows a distinct progress from west to east. This
progress will be evident from figure 18 which shows the centers of
oscillation in the area between 50° W. and 120° E. north of the Equa-
tor when the epochs are taken successively two years later. The
results in figure 18 are derived from the data of 48 stations taken
from ‘‘ World Weather Records.”
In 1885 there was a marked excess of pressure over Greenland
(see fig. 17) ; in 1887 this center of excess pressure is displaced to
Norway; in 1889 this center is over the northern part of central
Siberia; two years later, in 1891, it is over the northern part of
western Siberia. The progress of the centers is shown by small cir-
cles in the upper chart of figure 18. The circles show that the center
was displaced eastward about 180° in a period of 7.5 years or at a
rate which would carry it around the world in two oscillations of
this period.
In his study of the 2}-year period Mr. Clough’ found that the
epochs at Portland, Oregon, preceded those at Toronto by about
0.75 year. The difference in longitude is 43°. At that rate the epoch
would move about 150° of longitude in one period, or approximately
around the world in two periods.
The charts given by Dr. Weickmann in his study of the 24-day
period referred to previously do not show the drift in longitude so
clearly as the drift in latitude. However, in his charts there are
found centers of maximum departure which show a drift in longi-
tude. A center in the Aleutian Islands on December 10, 1923, moved
eastward across Canada to Labrador in 11 days, which is at the rate
of about one period for 180° of longitude; but a center near Green-
land moved eastward to northern Siberia and then retreated.
The longitude drift of the waves is, hence, not so clearly defined
as the latitude drift ; but there is undoubtedly a trend which may be
stated as follows:
* Monthly Weather Review, Vol. 52, No. 1, p. 30, Jan., 1924.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
p
ne:
q re
y £3 t
a
eH)
5-yea ae ete een
|
rg. 18.—Departures of pre
1880, 1 ae
NOS 7, THE ATMOSPHERE AND THE SUN—CLAYTON 29
Law of longitude displacement of periodic waves.—Periodic waves
tend to drift eastward at a rate of 180° of longitude in one period,
whatever the length of the period. The centers of greatest departure
are found in high latitudes, 60° to 80° from the Equator.
There are several factors which make this drift toward the east
difficult to follow. First there are the factors depending on solar
changes described in the latitude effect and which are nearly instan-
taneous with solar changes. There are also seasonal factors and
probably others which influence the results.
Examining the successive charts in figure 18 it is found that the
magnitude of the departures in the 7.5-year period decreased rap-
idly as the central areas passed into Siberia and increased again
over Kamchatka. This enhanced intensity in the departures coincided
with a maximum of solar activity as will be seen later.
Another disturbing factor is the formation of centers of distur-
bance moving at right angles to the normal waves. When waves of
high pressure and low temperature are advancing from the north-
west, low pressure areas form in front of them and advance from
southwest to northeast. These disturbances advancing toward the
northeast are particularly frequent over the warm ocean waters to
the east of Asia and of North America. These cross currents greatly
complicate the normal movement of atmospheric waves and make
analysis of the data difficult.
V. RELATION OF THE WEATHER WAVES TO SOLAR CHANGES
If the values of solar radiation observed by the Smithsonian
Astrophysical Observatory simultaneously with the pressure waves
are treated in the manner just described they show in each case
wavelike changes of the same length as the pressure waves.
Figure 19 shows the successive means of four periods of seven
days in solar radiation during November and December, 1927, com-
pared with the atmospheric pressure observed at the same time at
Eagle, Alaska, and treated in the same manner as in table 6. The
dotted curves in each case show the harmonic values of the 7-day
wave computed from the data. Compare this diagram with the plots
in figure 12.
Figure 20 shows the means of successive values of a period of 13.6
days in solar radiation and in pressure derived from the means of
two periods. This diagram may be compared with the plots in fig-
ure 13. The dotted curves in figure 20 show harmonic curves com-
puted from the data.
30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
NOV. 5, 1927
Fic. 19.—7-day period in solar radiation and pressure.
MAR. 2l, 1927 MAY 10
Pa oe a
Mead asa
a ee
aa
SSURE EAGLE ,65 IN., 141 Ww.
eevee
Fikes AALS Rages
IG. 20.—13.6-day period in solar radiation and pressure.
NO. 7 THE ATMOSPHERE AND THE SUN—CLAYTON 31
Figure 21 shows the observed values of solar radiation during
December, 1923, and January and February, 1924. These values are
compared with the observed values of pressure at Spitzbergen and
at Hamburg. A 24-day period of oscillation is evident in each case
and this oscillation is shown by the dotted curves computed from
the data in each case by harmonic analysis. Pressure data from all
DECEMBER 1923 JANUARY 1924 FEBRUARY
| 9
10 20
FAN W Ged |
1 Da a a
ASSURE IN SPITSBERGE!
Fic. 21.—24-day period in solar radiation and pressure.
over the northern hemisphere were treated in this way for a period
of 24 days by Dr. Weickmann and showed a systematic wave
movement from the polar basin southward.
For the study of long periods, no values of solar radiation are avail-
able; but the 7.5-year period shows a distinct relation to sun-spot
changes. Figure 22 shows a plot of consecutive means of three
periods of 7.5 years. This period is one-third of Hale’s sun-spot
period of 22.5 years, and the mean of the three periods eliminates
the I1.3-year sun-spot period which is one-half of Hale's period.
32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
Pressure curves are plotted for five widely separated stations. These
plots show distinctly an oscillation in the atmosphere of the length of
7.5 years and a progress southward from high latitudes.
1880
ee Saas
BMAive oa Sa
a a
PRESSURE
STYKKIS|HOLM , 65 NI, 23°W.
Wi. a Sey 6 NEAT NG Le Ga 8 ya
eT NE Ad AE Sn A
pf __¥ |__| Nf
ne i A
enna eM a7 Berea o
Faas
Fic. 22.—7.54-year period in sun spots and pressure means of 3 periods.
VI. SOLAR CYCLES. AND WEATHER (GYCLES
From the preceding investigation it is evident that atmospheric
and solar conditions show wavelike changes of a periodic nature.
The question has long been a challenge to investigators, as to whether
there are fixed and regular cycles in weather and in solar changes.
If such regular cycles could be found, it would greatly assist in
unraveling the complexities of the weather and in forecasting future
occurrences. There is a dominating period of about II years in sun-
spot numbers, and many efforts have been made to find this same
NOle 7 THE ATMOSPHERE AND THE SUN—CLAYTON 33
dominating period in weather changes. Such a relation has not been
found and the reason appears to be that weather changes follow
changes in solar radiation more closely than they do sun-spot num-
bers, and solar radiation is more variable and shows a more complex
periodicity than do sun spots.
When the 11-year period 1917 to 1928 is analysed harmonically
for sun spots and solar radiation, the results in table 7 are obtained.
TasLe 7—Harmonic Terms for 11%4-Year Period in Sun Spots and Solar
Radiation
Sun-spot numbers ; Solar radiation
(om A aren ar ae \ oe 8 SE
Amplitude Amplitude
Epoch 1917.5 in numbers Epoch 1917.5 in calories
Ax = 104° Gis= 35:3 Aa = 64° di = .009
Ag 274 aZ—70 AG 2385 a2 = .004
Ag 305" a3 = 10.6 AG 505 a; = .008
AEE Ee a; = 6.0 As = 330° cJ—005
These results show that in a general way the oscillation in the num-
ber of sun spots and in the intensity of solar radiation are in the
same phase—that is, when one increases the other increases ; but the
amplitudes of the changes are very different. The amplitude of the
primary oscillation, a,, in the sun spots (the 114-year period) is
decidedly predominant while in the solar radiations the amplitudes
of the harmonics of 5, 4, and 4 of 11.3 years, a2, a3, and ay are almost
as large as the primary a. The pressure data for tropical stations for
the 11 years 1917 to 1928 are not available at present, so that a com-
putation of pressure changes was made by going back two periods of
I14 years to January, 1890, and computing the harmonic terms from
the mean pressure of nine equatorial stations extending from Quix-
eramobim in Brazil eastward across Africa and the Indian Ocean to
Malden Island in the Pacific. The data covered two 11-year periods,
1890-1913, and the epochs were taken at 1895.0= 1917.5.
Taste 8.—Harmonic Terms for 11%4-Year Period in Pressure, 1890-1913
Mean Pressure of 9 Equatorial Stations
Epoch 1895.0 = 1917.5
INS 57 ds = 0.36 mb.
200 a2 = 0.32 mb.
As = 265° ds = 0.25 mb.
AGT 50 as = 0.35 mb.
This comparison indicates that in the 1t14-year period in pres-
sure in the Tropics, the phase is in general terms opposite to that
of sun spots and solar radiation, and hence when these increase the
34 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
pressure decreases. This fact is also made very apparent by com-
paring the individual periods in sun spots with pressure from 1870
to 1920. It is also evident that the amplitude a, of the 11-year period
is not dominant as in the sun-spot period ; but as in the case of solar
radiation the subharmonic terms a», a3, and a4, are almost as large as
the primary a.
In order to compare the harmonic terms of the 114-year period in
pressure in equatorial regions with those in other latitudes, the mean
pressure was obtained for each 10° of latitude in the northern hemis-
phere for each year from 1890 to 1913. A period of 23 years was
taken because from Hale’s observations of magnetism in sun spots
the complete period of the sun spots is about 22.6, so that 11.3
years becomes the second harmonic of this period. From the data
thus obtained harmonic terms were computed for each zone of lati-
tude and are given in table 9. The phases of the periods varied for
TABLE 9.—Amiplitudes of the Harmonics of a 22.6-Year Period in Pressure
dz as a4 a6 as do4 a48 a7z2
Zones of No. of 11.3 7.54 5-65 3-77 2.83 Tes 5.7 3.8
latitude sta. yr. yr. yr. mo. mo. mo.
yr. yr. :
70°-80° N. 2 0.60mb. 0.65 mb. 0.94mb. 0.92 mb. 1.12mb. 2.40 mb. 2.50 mb. 3:75 mb.
60°—70° 14 0.59 0.61 0.77 0.88 0.86 2.09 2.78 Pray
5SOn-60+) 2 10:34 0.38 0.46 0.61 0.76 1.32 1.60 1.28
40°-50° 25 0.32 0.42 0.43 0.45 0.51 1.22 1.10 123
30°—40 30 0.24 0.33 0.40 0.31 0.31 0.67 1.02 0.88
20°—30° i3) 036 0.21 0.29 0.24 0.23 0.62 0.63 0.53
10°—20° 15 0.42 0.25 0.35 0.21 0.33 0.42 0.45 0.40
0°-10° 6 0.36 0.27 0.32 0.25 0.35 0.39 0.25 0.38
Norre.—In computing the harmonics in Table 9 the means of three or of four periods
were used in each case except the case of a2 where two periods were used. The observed
values from which az were computed covered the entire interval of 22.6 years, while the
values from which a72 were computed covered only one twenty-fourth of this interval.
°
each latitude as it was evident they must do from the preceding inves-
tigation of wave movement. The striking facts brought out are:
(1) The amplitudes of the periods increase greatly in high latitudes
where they are much greater than in low latitudes and, (2) the
amplitudes of the smaller subharmonics in high latitudes are much
greater than that of the period of 11.3 years.
This last finding is of the utmost importance to meteorology,
because it shows that the shorter periods are of much more impor-
tance in the meteorology of high latitudes than the longer periods of
II years or more. These meteorological and solar periods are all
believed to be harmonics of longer solar periods.
Clough found solar periods of 300, 11.3, 7, and 2.5 years, and an
analysis of the sun-spot data by Schuster disclosed a number of other
NO. 7 THE ATMOSPHERE AND THE SUN—CLAYTON 35
periods besides the 11-year period. Turner found evidences of a
period of 260-280 years from a study of tree rings, Nile floods, Chi-
nese earthquakes, and sun spots. (Mon. Not. Roy. Astron. Soc., 1919
and 1920.) According to a recent analysis of the Wolfer sun-spot
data made by Dinsmore Alter, published in the Monthly Weather
Review of October, 1928, there are solar periods of more than 200
years in length, and the 11-year sun-spot period is a subharmonic of
much longer periods. This view agrees with that put forward by
Ellsworth Huntington and S. S. Visher in “ Climatic Changes,”
1922, p. 45. My own investigations are in accord with this view, ex-
cept that recently the longer periods seems somewhat greater than that
given by Alter.
Beginning with a period of go years, instead of 84 as given by
Alter, I find periods of approximately the following length: Length
of solar periods in years: 90, 56, 45, 35, 30, 28, 22.5, 13,85; E20;
14, 10, 9, 8.2, 74, etc. All of these shorter periods are subharmonics
of go years, except 56, 35, and 28, which are harmonics of a longer
period.
They agree very well with meteorological cycles found by Prof.
A. E. Douglass * from rings indicating the annual growth of trees in
the southwestern part of the United States where rainfall is the most
essential factor in growth. The periods found by Professor Douglass
pee 2h, 40,26, 22:5-24.0, 20:5, 17-2, 14:2, P1.2-11:7,.10.2,,0.0;.7:6, 0.8
years.
A study of periodicities in the Nile floods by C. E. P. Brooks *
leads him to pick out the following periods in years: 76.8, 64.6-67.4,
30.85, 33-49, 24.43, 21.81-22.43, 18.32, 16.68, 14.87, 12.50, 10.86-
11.36, 8.33, 7.33, 6.83, 5.52, 3.66, 2.86. It is pointed out that 11 out
of 16 of these periods are multiples or submultiples of a period of
22.12 years. This period is somewhat shorter than Hale’s period of
22.6 years; but the difference may be due to the fact that the period
actually was shorter during the intervals covered by Brook’s data
which go back to the year 641. His data indicate a systematic varia-
tion in the phase of this period, so that at the end of about 200
years the phase is inverted as regards epochs 200 years earlier.
The researches of D. Brunt* also indicate that there are a great
many meteorological cycles, or else there are none. His periods in
years derived from the Greenwich temperatures are: 23, 17.5, 15, 8.17,
‘ Climatic cycles and tree growth, Vol. 2, p. 123. Carnegie Inst. of Washington,
1928.
* Mem. Roy. Meteorol. Soc., Vol. II, No. 12, 1928.
* Quart. Journ. Roy. Meteorol. Soc., Vol. 53, No. 221, Jan., 1927.
36 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
7.24; and in ‘months, 64; 60, 42;°37, 20,°25; 214) °10:3-10:5,, as5-
14.7, 13, 124. The researches of Dinsmore Alter* published in the
Monthly Weather Review also bear testimony to the multiplicity of
meteorological cycles.
My own researches have dealt largely with shorter periods of days
and months rather than years, principally because there was a much
larger mass of data available for discussion. In my earlier studies
of pressure and temperature data in the United States.” I found the
following periods in days: 3, 3.6, 4.6, 5.45, 6.14, 7.24, 9.1, 11, 18, 22,
29, 44, 58, etc. My recent studies indicate that there are many more
cycles and that all are probably harmonics of the sun-spot cycle.
A. Defant* in a world-wide study in 1912 found the following
periods in days: 4.4, 7.9-8.7, 12.0-13.0, 16.8, 24.5, 31.2-31.5. Arec-
towski, Turner, Simpson, Wallén, Myrback, Wasserfall, Schosta-
kowitsch, and Kidson have all found short meteorological cycles of
various lengths Even the short period cycles of a few days are
probably submultiples of much longer solar cycles, the most promi-
nent of which is the 11-year sun-spot cycle, or its double value, the
22.5-year cycle.
In most cycles the subharmonics of small length are not important,
but it has been shown in table 9 that in high latitudes the subhar-
monics of the 11-year period in meteorological cycles are of greater
amplitude than the primary period of 11 years and that the ampli-
tude increases with decreasing length of the harmonic. The sequence
has not been followed through for the entire Northern Hemisphere
beyond the period of about four months, but the amplitudes of
meteorological cycles at stations in the northern United States and
Canada apparently increase down to a length of about three days.
These shorter periods determine the origin and movement of the
ordinary cyclones and anticyclones seen on the weather map.
Most investigators of meteorological cycles assume at the begin-
ning of their work that any cycle which may exist is constant in
amplitude and phase and may by repetition be separated from other
changes by which it is masked. This belief is the basic assumption
underlying the analysis by the Fourier series or the Schuster peri-
odogram. Prolonged investigation usually convinces the research
worker that this assumption cannot be maintained. I early became
*Monthly Weather Review, Vol. 54, p. 44, and Vol. 55, pp. 60 and 263.
* Amer. Meteorol. Journ., Feb., 1895, p. 376; also Amer. Journ. Sci., March,
1894.
* Sitzungberichte d. Wiener Akad., Bd. 121, Heft 3.
NO. 7 THE ATMOSPHERE AND THE SUN——-CLAYTON 37,
convinced that meteorological cycles change both in amplitude and
phase. (Science, 1898, p. 243.)
Figure 23 shows an analysis of the Wolfer sun-spot numbers
between 1890 and 1913 into a period of 22.6 years and its harmonics.
It is seen that the chief period is one of 11.3 years, but some of the
other periods show a fairly large amplitude of oscillation.
1890 1895 {900 1905 1910
y cA a
=) ORE RENE Ce Ce eweee
EEE EE PPE PEELE EET
Fic. 23.—Harmonic analysis of 22.6-year sun-spot period, 1890-1913.
The meteorological data at more than a hundred stations in various
parts of the world were analyzed in the same manner. Figures
24, 25, 26, and 27 show lines of equal departure of pressure for the
various periods at the time of maxima of the solar periods of the
same length. A chart showing the departures at the time of the
solar maxima of the 7.5 year period is given in figure 17.
38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
Certain common features stand out clearly in all these charts.
First, in the equatorial belt, except possibly over parts of the Pacific
Ocean, the pressure is lower than normal at the time of maximum
ZZ WRX NA .
Jeo = = SS SN
fi a a + cee RS
| eee ae KY
i ees a
7 see aff, tere t See ENN)
ces eee hr
Semi Si Ea
LLL LX
= fy Ea
z
Fic. 24.—I1.3-year period in pressure = 4 of 22.6 years. Departures at time
of maximum of solar eae of same length.
solar activity in each period. Second, in middle latitudes of the
Southern Hemisphere there is a tendency to a belt of pressure above
normal which cannot be well outlined on account of insufficient
Fic. 25.—5.65-period in pressure = + of 22.6 years. Departures at time
of maximum of solar period of same length.
observations. Third, in the Northern Hemisphere in high latitudes
there is a tendency for the departures to form centers of positive and
negative departures, usually two centers of positive departure, and
NOS 7 THE ATMOSPHERE AND THE SUN—CLAYTON 39
two centers of negative departure. Fourth, these centers are not in
the same geographical position for the different periods and do not
remain fixed for successive epochs of the same period. The reasons
Fic. 26.—3.77-year period in pressure = + of 22.6 years. Departures at time
of maxima of solar period of same length.
for these shifting centers are not clear. They are associated with
changes in the phase and amplitude of the cycles.
Changes in amplitude are both apparent and real. Apparent
changes occur where two periods of nearly the same length first
pales \
a SP
ieee
DHE |
ILE
WZ a
Ez
Sema
Fic. 27.—2.82-year period in pressure = § of 22.6 years. Departures at time
of maxima of solar period of same length.
strengthen each other when they are in the same phase and then
weaken each other when they are opposed in phase. This change will
be familiar to most readers from diagrams to illustrate beats in sound
40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
waves. The beats are even more complicated when there are three or
more periods of nearly the same length. In such a case there may be
an apparent change of phase in one of the periods.
Real changes in amplitude are brought about by the influence of
longer periods on shorter periods. An example of this is the influ-
ence of the annual period on shorter weather cycles. All weather
changes are most intense in winter, because then the contrasts in
temperature between Equator and pole, between ocean and continent,
and between adjacent bodies of land and water are at a maximum
intensity and the general atmospheric circulation is increased.
Also all periodic changes in the atmosphere are more intense when
solar activity increases. The reason for this increased intensity will
be clear, first from the fact shown in the early part of this paper
that increased contrasts of temperature and pressure in the atmos-
phere result from increased solar activity, and second from the fact
that the amplitude of the solar cycles increases with increased solar
activity.
An example of the increased amplitude of solar periods with
increased solar activity is shown in figure 22 where the amplitudes of
the 7.5-year sun-spot period is distinctly greater during the inter-
val 1865 to 1875, when the general level of solar activity was higher,
than during the interval 1885 to 1895, when it was lower. The
increase of amplitude during the first of these intervals and decrease
during the second was also evident in the sun-spot cycle and in its
harmonics of 5.65, 3.75, 2.82 years, etc.
An example of increased amplitude of meteorological cycles with
increased solar activity is shown in figure 28 where a period of 74
months in pressure at Chicago shows a marked increase in ampli-
tude at the time of maximum of sun spots in 1917 and a diminished
amplitude during the intervals of minima of sun spots in 1913 and
1923-1924. The data for this curve are the means of 10 overlapping
periods of 74 months obtained in the manner indicated in table 6.
The dotted curves are sine values computed for each individual
period.
That meteorological cycles change in phase as well as in intensity
is also evident. These changes of phase appear to arise from several
different causes. First, the solar periods themselves change phase. In
most cases this change occurs suddenly and appears to be about 180°
AND THE SUN—CLAYTON AI
THE ATMOSPHERE
NO.
‘(uimutxeu jods-uns 1vou apnyydue jo asevaruT SsulMoys) OsPoIy) Ul ainyesodwia} ur ported syyuow-¢£— gz “DIY
| |
iS ae ee as | aE
FAW ALAIN ADA TAT ETA TA Din atta
LSS eae Avera Eee 1 ee
Mii ea see ae
Yn oe Ee ee ee ee ee
a a a Ma E:T TT RT: 1) SR GES Ta 1
42 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
or a complete reversal in phase. Figure 29 shows what appears to be
a reversal in phase in the sun-spot cycle. The average length of this
cycle is about 11 years, so that two cycles occur in 22.5 years. If the
cycles are plotted in 22-year periods as in figure 29 it is seen that in
the period 1770 to 1792 the cycle is nearly inverted in phase to the
cycles occurring 22 years earlier and 22 years later. It is, however,
quite possible that this result is due either to interference of periods
of different lengths, or to lack of accuracy in the early observations.
No such apparent inversion has occurred since 1800.
10 11 12 13 14 15 16 17 18 19 20 21 22
iS S2NeRRha
aaa coal et a
Fic. 29.—11-year sun-spot cycle, showing apparent inversion of phase.
In figure 30 is given what appears to be a reversal of phase in the
74-year period. This cycle was in one phase from 1848 to 1870,
as shown by the broken curve in figure 30, but appears to have been
in an opposite phase from 1825 to 1847, as will be seen by the con-
tinuous curve in figure 30. This type of change is found in every solar
and meteorological period. Brooks and Clough seem to think that
shiftings of phase are gradual; but my own researches lead me to
the opinion of Professor H. H. Turner that the changes are sudden
and of the nature of discontinuities.
The change of phase in meteorological cycles is not brought about
entirely by changes in phase of solar cycles; but is in part, at least,
due to shifting of centers of action in the atmosphere. In the case
NO. 7 THE ATMOSPHERE AND THE SUN—CLAYTON 43
of pressure, when it rises in one part of the world, there is an equiva-
lent fall in other parts. These centers of rise and fall are not fixed
in position, but shift their position to some extent as illustrated in
the case of a 25-month period in a preceding paper of this series.’
The variations in intensity and phase of solar and meteorological
cycles makes the investigations of the separate cycles difficult. The
use of the Fourier series and of the Schuster periodogram are not
well adapted to such work. In order to meet these difficulties I
devised the correlation periodogram* which is to a considerable
extent independent of variations in intensity of the periods; but
does not overcome the difficulty of shifting of phase. The best
EAR 2.5.4 5 6 7 6 9 10.00 12 13.4 15.16. 17.16 19 20 2) 22
Ul: NA PERU UT NUT
WIZE SE58
il
a
CPR EERE
Fic. 30.—7}-year sun-spot cycle. Means of 3 cycles, showing apparent inversion.
N@ N
method appears to be to use trial periods of successively greater and
greater length and harmonic analysis for each individual oscilla-
tion *; then to combine the results for each period into groups of
3, 5, 10 or more. By this method the curve in figure 28 was
obtained. This method of research, using groups of 10, has made it
possible to analyze and to follow the changes of a great number of
meteorological periods and to recombine them by synthesis for a
trial in practical forecasting. Such analyses of more than 100 cycles
have convinced me that these cycles follow solar cycles of the same
length and that they are, mostly at least, harmonics of long solar
cycles.
"Smithsonian Misc. Coll., Vol. 78, No. 4, p. 48, 1926.
* Smithsonian Misc. Coll., Vol. 71, No. 3, p. 15, 1920.
* Clayton, H. H., World Weather, p. 376, New York, Macmillan & Co., 1923.
44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 62
VII. THE USE OF WEATHER CYCLES IN PORECASTING
Having developed methods of separating and studying various
conditions which make up the weather, it seemed important that a
test be made of the possibility of using them in practical forecasting.
Forecasting future weather conditions in the present state of knowl-
edge may be undertaken in at least three different ways: (1) By
tracing out the results which follow the increase or decrease in the
general circulation of the air with changes in solar activity. (2)
By analyzing and following weather waves of different classes. (3)
By computing the amplitudes and phases of different cycles found
in solar and weather changes and projecting these forward into the
future.
In regard to the use of the first method, since increased solar
activity is attended by a fall of pressure in equatorial regions and
by increased contrasts of pressure in higher latitudes, there is brought
about an increased atmospheric circulation and certain general con-
ditions follow :
(1) The cloudy and clear belts of the world are intensified and
thus alter the incoming and outgoing radiation.
(2) The increased air circulation means an increased flow of ocean
waters which brings an increased northward flow of warm water
along the east coast of the United States and Japan and an accumula-
tion of warmer water in the North Atlantic and North Pacific. The
accumulation of warmer waters in these regions especially in autumn
brings increased cloudiness and increased rainfall. The increased
cloudiness reacts by diminishing radiation losses from the earth and
thus further modifying weather conditions. On the other hand
the increased oceanic circulation brings increased cold water to the
shores of North Africa and southern California, and produces a
chain of atmospheric conditions which affect the northern shores of
South America and the West Indies and extend well out into the
Pacific. A parallel set of changes is produced in the Southern Hemis-
phere in an opposite way on the east and west sides of the con-
tinents. When solar activity diminishes the reverse conditions
prevail.
(3) Increased solar activity brings also an increased flow of air
over the continents and with it an increased rainfall in certain regions
and a decreased rainfall in other regions. The distribution of pres-
sure and attendant conditions is to a large degree influenced by the
seasons.
NO. 7 THE ATMOSPHERE AND THE SUN—-CLAYTON 45
Hence, to follow the sequences of weather resulting from increased
solar activity it is necessary to consider the month or seasons sepa-
rately and to work out expected conditions for different intensities of
solar activity.
In regard to the use of the second method, forecasting weather as
ordinarily practiced at the present time depends on anticipating for
a day or two at a time the drift of weather conditions. Such fore-
casts can be improved and extended in time by analyzing weather into
waves of different lengths and forecasting the progress of the
stronger waves. Even long range forecasts can be made on this basis,
as I have demonstrated by actual tests.
The third method of forecasting is by means of the periodic
vibrations in the sun and atmosphere. Any pulsation in solar condi-
tion will be attended by similar pulsations in the earth’s atmosphere.
The shorter pulsations will be felt relatively more in high latitudes
of the earth and the longer pulsations relatively more at low lati-
tudes, but all will be repeated to some extent in every part of the
atmosphere. An analysis of the periodic terms in the weather at any
point on the earth would make it possible to project the periodic
terms ahead to any length of time desired, were there no variations
in the amplitude and phase of the periods. But there are variations
and for this reason it is necessary to redetermine the periodic terms
at short intervals and to limit the time in advance which they are
made to cover. When these variations in the periodic terms become
calculable, this method of forecasting will probably replace all others.
Already considerable progress has been made along this line.
In practical forecasting at present it is desirable to consider all of
the three methods mentioned and to use them as checks on each
other. Forecasting in words has but little meaning to the average
expert, because the meanings of words can be interpreted in various
senses and there are no accepted rules for verifying such cases.
Quantitative forecasts can, however, be verified by accepted stand-
ards ; so that from the beginning of my experiments in forecasting,
both verbal and quantitative forecasts were made. These quantitative
forecasts were made first for about a week in advance, then for
longer intervals up to a month. Figure 31 gives one of the more
recent of these forecasts of pressure made on November 24, 1929, for
27 days in advance beginning on November 26 and ending on
December 21. The forecast was made up from a combination of
cycles varying in length from 3 days to 13 days. The correlation of
the forecasted with the observed pressure is 0.64 + 0.06.
46 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
By computing pressure in this way for a network of stations,
weather maps bearing unmistakable resemblance to observed weather
maps may be computed in advance. In March, 1929, values of pres-
sure were computed for one week in advance for 23 selected sta-
tions forming a net over the United States and from these computed
values lines of equal pressure departures were drawn, The maps
NOVEMBER, 1929 DECEMBER
2627026029150 Nae SAS 6 BO NOMIIZ Sta IS enim Snl9 scored
CHICIA
: 4H
fis ZAIN
ae ai
calsT
Fic. 31.—8 a. m. pressure.
thus forecasted are compared with the observed pressure distribu-
tion in figure 32. The close resemblance of the two sets of maps is
apparent. This degree of accuracy can be obtained, however, only
when the meteorological cycles are comparatively steady. It is never-
theless the goal toward which research is leading and to which it
will undoubtedly attain.
In April, 1929, a diagram was sent to a number of persons,
including the Secretary of the Smithsonian Institution, giving a fore-
cast of departures from normal temperature by weeks from April 2
to September 3 for New York City and for two other stations. Fig-
ure 33 gives a copy of this plot for New York City. The broken
curve shows the forecast and the continuous curve shows the ob-
served departures from normal. The correlation coefficient for the
23 weeks is 0.37+0.12. This correlation taken alone is inconclusive
as to the possibility of such forecasts, except in the light of other
data indicating its possibility. It is believed that forecasts by months
and years are feasible on the same basis and by the same methods,
but no prolonged test is yet available.
If the conclusions presented in this paper are verified and accepted
by other research workers, as I feel they must be in time, it will
NO: 7 THE ATMOSPHERE AND THE SUN—CLAYTON 47
PRESSURE MARCH [3 OBSERVED
-—. | aS
+ ——rr—
Fic. 32.—Pressure forecasted from a combination of meteorological cycles.
APRIL MAY JUNE JULY AUGUST
2 9. 16.25.50 7 14.21.26 4 W186 25. 2.9 16.25.30 6 15 20 27 3
PPP
UN ALALN ALLL
5 = a, \Y/ Ghz S a
CUNVETETELL EY TT
) /
: /
+---~- FORECASTED
OBSERVED
Fic. 33 Weekly temperatures 1920.
48 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
mean a revolution in present methods of weather forecasting. The
forecasting of pressure and temperature will be made in much the
same way that ocean tides are now predicted, except that the periods
used will be solar periods rather than lunar periods and will need
to be treated in a special way owing to changes in phase and
amplitude.
Such a successful forecast as that shown in figure 31 seems con-
clusive evidence that day to day weather is not a haphazard occur-
rence as many persons believe, but is subject to calculation. It is
evident that changes of pressure are calculable to some extent now,
and the calculations will, no doubt, in the future be made with
increasing accuracy for weeks and perhaps months in advance. Proc-
esses will be simplified and machinery like the tidal machines will be
introduced in order to handle the immense amount of data which will
be needed for world-wide forecasts, or even for forecasts over a large
area like the United States.
SUMMARY
This paper contains evidence pointing to the following conclusions :
Solar activity varies in complicated pulses. These pulses or varia-
tions in intensity are attended by variations of pressure in the earth’s
atmosphere. When solar activity, as indicated by spots and radia-
tion values, increases, the latitude contrasts of pressure in the earth’s
atmosphere are increased and atmospheric circulation speeded up.
The pressure falls in the equatorial belt, rises in middle latitudes
and falls in the polar regions. When solar activity decreases the
reverse conditions occur. The zonal regularity of these changes is
interfered with by the distribution of land and water and by seasonal
changes.
Immediately following the decrease of pressure in the polar region
with increased solar activity, a wave of decreased pressure moves
toward the Equator. With decreased solar activity the pressure in
polar latitudes increases and a wave of increased pressure travels
towards the Equator. These waves move with a speed proportional to
the length of the solar pulse or period causing them. If the period of
oscillation is seven days the wave moves from pole to Equator, when
measured along a meridian, in seven days. If the length of the
oscillation is 27 months, or 24 years, the time of the wave movement
from pole to Equator is 27 months and if the length of the period is
74 years the time of movement from pole to Equator is 74 years, or
one period of oscillation in each case.
NO. 7 THE ATMOSPHERE AND THE SUN—CLAYTON 49
There are also east to west movements of the waves, and there
are probably returning waves toward the poles of less intensity ; so
that the observed phenomena are extremely complex. The analyzed
wave movements are subject however to apparently simple laws, and
can, therefore, probably be computed and combined to produce ob-
served conditions.
The observed data of sun-spot numbers and solar radiation values
when subjected to harmonic analysis for the 11-year period 1917 to
1928 show that the dominating period of about II years in sun
spots is no more marked in solar radiation values than the subhar-
monics of 4, 4, 4, etc., of the 11-year period which have ampli-
tudes nearly as large as the 11-year period itself.
When the pressure observations in the Tropics are subjected to
harmonic analysis they show periods resembling in amplitude those
of solar radiation values and not those of sun spots. The analyses of
the data in higher latitudes show that the amplitudes of the subhar-
monics increase with latitude, so that in high latitudes in the neigh-
borhood of the pole the subharmonics become vastly more important
than the primary period.
A study of the possibility of analyzing the data at each particular
part of the earth with the view to discovering fixed periodic cycles
indicates that if such cycles exist, the amplitudes are subject to
wide variations and even to inversion of phase from time to time.
However, when the complex cycles are analyzed individually and
averages taken for a small number of successive cycles, it is possible
to project them into the future and combine and plot them in a curve
which at times has a striking resemblance to observed data. As knowl-
edge of methods and laws of change progress, this kind of fore-
casting will undoubtedly be done with increasing accuracy.
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 82, NUMBER 8
FOUR NEW RACCOONS FROM THE KEYS
OF SOUTHERN FLORIDA
(WiTH Five PLATEs)
BY
E. W. NELSON
(PUBLICATION 3066)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
JULY 10, 1930
ey ys f a an i ny
“I eae Ay aad A Sl kil ;
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oT ie !
i : 7 wy
if
I
The Lord Battimore Press
BALTIMORE, MD., U. & Ae :
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4 4 ee ee no
FOUR (NEW RACCOONS FROM. THE KEYS OF
SOUTHERN FLORIDA
By E. W. NELSON
(WitH Five Pirates)
Between the last of February and late in March, 1930, the author
visited the keys lying about the southern end of Florida and collected
on them a series of ‘61 specimens of raccoons. The keys, or islands,
visited proved to be segregated into four rather well-defined groups
and the specimens collected show very definitely that each group is oc-
cupied by a subspecies of Procyon lotor peculiar to it, and all differing
from Procyon lotor elucus of the neighboring mainland.
The main islands of each group are named below but, in addition,
each group includes many smaller islets practically all of which are
covered with mangroves.
Ist. Ten Thousand Islands Group forms a broad compact belt of
mangrove keys lying for about I00 miles along the southwestern
coast of the peninsula, from a little south of Naples down to Shark
River. The width of this belt varies from one to several miles, its
exact width and some other details not being as accurate as desirable
in published maps I have seen.
2d. Key Largo Group, as here considered, lies along the south-
eastern border of the peninsula and includes Virginia and Biscayne
Keys just north of the entrance to Biscayne Bay, and Elliott Key,
Key Largo, Plantation Key with Upper and Lower Matecumbe Keys
to the south of the entrance.
3d. Key Vaca Group lies southwesterly from the preceding group
and begins with Long Key on the north and extends south to include
Duck, Grassy, and Fat Deer Keys, Key Vaca, and Knights Key.
4th. Big Pine Key Group still farther to the southwest includes
No Name, Big and Little Pine Keys, Torch Keys, Ramrod, Cudjoe,
Summerland, Saddlebunch, and Boca Chica Keys, and Key West.
The isolation of raccoons of the Procyon lotor type in islands on
the Atlantic coast side of the continent from Cozumel Island, off the
peninsula of Yucatan, to the Bahamas and the coast of southern
Florida, has tended toward the production of depauperate forms as
all of them are smaller than the animals of the adjacent mainland.
On the Pacific side of the continent the raccoons of the Tres Marias
SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 82, No.8
2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
Islands are about the same size as the animals of the neighboring
Mexican mainland. These islands are high and wooded and fresh
water occurs on them.
Conditions are such, on practically all of the keys off the coast of
southern Florida, that the raccoons living on them have no access to
fresh water beyond the fortuitous opportunities that may occur
during a heavy rain. To this circumstance, many of the local trappers
believe, is due the smaller size of the animals of the islands, for they
say the larger, better furred animal of the mainland occurs where
permanent fresh water exists back from the shore. Along the south-
western shore of the peninsula these large fresh water “coons”’ are
said usually to keep about the fresh water but to come down to the
mangrove swamps along shore at certain seasons. The trappers
agreed, however, that they never cross to the adjacent keys, even in
the Ten Thousand Islands, some of which have only narrow, shallow
channels cutting them off from shore.
In addition to their tendency to reduction in size the Key raccoons
have a distinct tendency toward duller and paler colors than the animals
of the mainland. The paleness is especially marked in the forms from
the Key Vaca and Big Pine Key Groups.
The abundance of crabs, shellfish and other food een by the
sea for the raccoons, would appear to remove the possible factor of
food shortage from the list of possible causes for the smaller size
among these animals, At the same time the absence of natural enemies
evidently gives favorable conditions for their increase and the ac-
counts given me of the amazing number of them trapped on the
keys, when the high prices of furs gave the incentive, make it ap-
parent that they must have been excessively numerous for an animal
of their size. Under natural conditions in the past, therefore, com-
petition for food may have been very strenuous for long periods.
On the Ten Thousand Islands some of the trappers told me that
when the high prices for fur began single trappers sometimes took the
skins of more than 800 raccoons on the keys in a season but that,
owing to this severe trapping, it is difficult now for a man to get even
one-third of that number. Similar accounts of the former abundance
of raccoons on other keys off the coast were given me by trappers
elsewhere. Some of the trappers put out more than 100 traps and
settings of from 40 to 60 are common. The total catch of raccoons
for southern Florida and the keys must be very large.
The measurements and weights in the flesh of the series collected
by me gave direct evidence that the female raccoons of that region,
in the flesh, weigh more than 20 per cent less than the males, and
No. 8 NEW RACCOONS FROM FLORIDA—NELSON 3
the more smoothly rounded, smaller skulls of the females give the
same evidence. Measurements, weights, and skulls of the series
collected by Dr. E. A. Mearns in Polk County, the type region of
P. 1. elucus, show similar size differences between the sexes of that
form.
The study of my series of specimens and comparisons with a large
number of specimens of Procyon lotor elucus from the Florida main-
land and of representatives of P. maynardi and P. minor from the
Bahamas makes it appear that a natural laboratory of evolution is
obviously at work on the Florida Keys with the generally distributed
and abundant raccoons as, at least, one of the principal subjects.
It is a pleasure to acknowledge my indebtedness to Dr. Thomas
Barbour and to Mr. H. E. Anthony for the use of material under
their charge in the Museum of Comparative Zoology and the American
Museum of Natural History, respectively.
THE RACCOON OF THE TEN THOUSAND ISLANDS
More than two years ago a friend told me that a raccoon smaller
than that of the mainland was reported to exist on the chain of keys
bordering the southwest coast of Florida, known as the ‘“ Ten
Thousand Islands.” This suggestive information was kept in mind
and the last of February, 1930, I arrived at Fort Myers, on my way
to investigate the rumor. Being delayed there for a day I located a
fur buyer and in reply to my inquiries he promptly confirmed the
truth of the statement which had caused my quest. He said that.a
very small, rather pale colored, raccoon is well known to the trappers
and fur buyers of this region, where it occupies the islands, or “ keys ”
along the coast. He added that owing to its small size and the inferior
quality of its fur “key coon”’ skins bring only about one-half to two-
thirds the price paid for the larger, better furred skins from the
mainland. The trader then produced about 30 “ key coon” skins from
Marco Island, which he had recently purchased, and I was delighted
to note their distinctive peculiarities.
The next morning I proceeded by rail to Marco Island, one of the
larger of the Ten Thousand Islands, near the northern end of the
group. It is several miles in both length and breadth and is bordered
by a belt of mangrove swamp where the raccoons live. The interior
is sandy, several feet above high tide mark, and covered with a thin
forest of scrubby, slender pines with scattered undergrowth.
Soon after my arrival I found resident trappers who agreed that
“key coons’’ were common on the island, but that those found on
the keys near Chokoloskee Bay farther south were the smallest of all.
4. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
Desiring to secure the most marked expression of this scientifically
unknown form, I left Marco Island at once and proceeded by auto-
stage to the town of Everglades, located about the middle of the
east side of Chokoloskee Bay.
The next morning I went to the fishing village of Chokoloskee, near
the southern end of the bay, where a trapper with a small motor boat
was employed and at the expiration of about ten days we had a
series of 16 specimens, all taken on the small mangrove keys lying
between Chokoloskee Bay and the open waters of the Gulf of
Mexico. The delay in getting these animals was due to the fact
that I had arrived at the end of the trapping season and the number
left was comparatively small.
Previously I had always thought of raccoons as animals dependent
on available fresh water and it surprised me to find them living in
great numbers among the mangrove islands, both in the Ten Thousand
Islands and on other Florida keys, without any possible source of ©
such water. Their food consisted of an abundant supply of fish,
crustaceans, and shellfish left exposed on the mud at each low tide.
The haunts of the raccoons among the mangrove roots of the Ten
Thousand Islands were shared by great numbers of roof rats (Rattus
rattus alexandrinus). These were so numerous that they interfered
with our success in trapping the raccoons, dozens of them being
caught. Another interference was the frequency with which fish and
crabs ate the bait above the traps while they were submerged during
high tide. As a result of these marine visitants the traps sometimes
capture curious prey. Several kinds of fish and crabs, in wallowing
about when tugging at the bait, now and then spring the trap and
are caught. My trapper said that on one occasion he caught a small
shark about 18 inches long. His method of trapping, which he in-
formed me was the regular practice among these keys, was to make
a little U-shaped enclosure by sticking pieces of dead mangrove roots
into the mud in a small opening among the mangrove roots at the
head of small bay-like indentations of the shore line, where the
animals patrolling the bare mud at low tide would find it on their way
from point to point. A piece of fish for bait was impaled on a small
stick, the other end being stuck in the mud at the inner part of the
enclosure. The steel trap was then set on the bare mud at the entrance
of the enclosure guarding the bait, without the slightest effort to
conceal it—a stick thrust through the ring at the end of the chain and
deep in the mud serving to hold any animal caught (see pl. 1, fig. 2).
At high tide this trap would be more than two feet under water.
Trapping among these keys is practically all done by using small
boats with outboard motors, or small gasoline engines.
No. 8 NEW RACCOONS FROM FLORIDA—-NELSON 5
The common name for this group well indicates the great number
of little keys of which it is formed. They are closely grouped and of
- most irregular outlines, being separated from one another, and from
the mainland, by a network of tortuous, shallow tide channels varying
in width from a few yards to several hundred yards. Enclosed within
this mass lie Chokoloskee and other narrow land locked bays. Cho-
koloskee Bay is the largest, having a length of about eight miles and
a width of from a half mile to a mile and a half. Nearly all the
keys of the group, like those where we trapped the raccoons, are
overgrown with tangled masses of the aerial roots of the red man-
grove above which rises the low wall of their dense, green tops.
These keys, in general, are covered with from one to about three
feet of salt water at the daily high tide and are devoid of fresh water.
The Ten Thousand Islands form, in fact, a great mangrove swamp
based mainly on the flat tops of old oyster beds. The red mangrove
thickets rise about 25 feet and are interspersed with larger black
mangroves and a few other salt loving small trees or bushes. In the
Shark River section, to the south, the mangroves are the largest I
have ever seen, forming closely set forests of trees 40 to more than 60
feet high. The mainland adjacent to these keys is bordered by a
fringe of mangrove swamps limited by the end of salt water. The
accompanying photograph gives an idea of the typical water front of
the mangrove islets near Chokoloskee (see pl. 1, fig. 1).
I have included the mangrove keys of the Shark River area within
the range of the small Chokoloskee raccoon wholly on statements of
several trappers that the raccoons there are the same as those about
Chokoloskee Bay. No specimens have been seen from there but
several from the mainland a few miles away at Cape Sable and Flam-
ingo are unmistakably P. 1. elucus.
RACCOONS AND THE FLORIDA KEYS
After observing the effect of environment on the raccoons of the
“Ten Thousand Islands ” it appeared probable that similar influences
may have produced modifications of the animals living on the great
series of islands or “keys” extending from Biscayne Bay to Key
West, commonly known as the Florida Keys. To determine the facts
concerning this I crossed the Everglades, over the motor highway,
from the town of Everglades to Miami. There, through the friendly
assistance of Mr. E. J. Brown, some trappers were promptly located
who were well acquainted with all the keys. The information they
gave confirmed my belief that the raccoons there would differ from
those of the mainland. Two trappers were employed as field assistants
6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
and by the use of motor boats and an automobile we made a rapid
reconnaissance of the entire chain of keys to Key West. Raccoons
occur abundantly on all the keys and we obtained a sufficient series to
indicate their characteristics. As at the Ten Thousand Islands this
work was done soon after the close of the regular trapping season,
during which thousands of raccoons had been caught, so that much
more time was required to capture the animals needed than would
have been necessary earlier in the year.
These keys are connected by the Over-Seas Railway from Miami
uniting them by fills where the channels are narrow and shallow
and by cement viaducts and bridges where they are more broadly
and deeply separated. In recent years the Over-Seas motor highway
has been constructed parallel to the railway from Miami, striking
the middle of Key Largo and extending thence down the keys to
the southern point of Lower Matecumbe, whence a ferry carries
passengers and motor cars across about 40 miles of water to the
northern end of No Name Key. There the motor road begins again
and continues southward crossing various keys to Key West. In
addition to the railway fills which unite some of the keys in this chain,
other fills made for the highway broaden the land bridge made by
the railway embankments uniting a number of keys, thus destroying
their individuality.
About midway in the 40-mile gap between the northern and southern
groups of keys lies a smaller group, the main one of which is Key
Vaca. A practically disused motor highway, extending north and
south about 12 miles, joins the main keys of this group except Long
Key which lies isolated by sea channels nearly midway between Key
Vaca and Lower Matecumbe. As shown below, the raccoons living
on each of these natural groups of keys, the northern, the middle, and
the southern, has its local subspecies.
The Florida Keys differ very much in formation and vegetation
from the Ten Thousand Islands. Most of these keys, except the
more recent small ones, are of limestone with nearly level surfaces,
elevated well above tide water, with borders of varying width of
mangrove swamp which are covered at high tide. Back of the tidal
area on the Key Largo and Key Vaca Groups is a strong growth of
deciduous tropical shrubs and trees often forming a dense and al-
most impenetrable jungle, from 15 to 30 feet high, where it has not
been destroyed by man. The northern half of Key Largo has the
heaviest forest growth I saw on the keys, rising to 50 or 75 feet high,
or perhaps more, in places. The trees appear to be the same species
as those elsewhere but grow on a better soil.
Roe a Ae ae
No. 8 NEW RACCOONS FROM FLORIDA—NELSON 7
The western, or Gulf, shores of all the large keys are bordered by
a practically continuous fringe of mangrove swamp but on the
Atlantic side occur sandy beaches often bordered by irregular series
of coconut palms, while scattered fan palms, often with trunks ten
to more than 25 feet long, are generally distributed in the interior.
A large part of Big Pine Key is covered with a thin growth of
slender dwarfed pines and palmettos with scattered areas of deciduous
trees and bushes. The neighboring Little Pine, No Name, Torch, and
Ramrod Keys have smaller areas of pine forest. The interior of the
other keys to Key West have the deciduous species common to the
entire chain of keys, with the usual marginal mangrove swamps and
occasional sand beaches.
The trappers informed me that the raccoons of the Florida Keys
go back inland from the mangrove swamps, where they generally
live, to feed on any ripening fruit that occurs. At the time of my
visit to Key Largo, a shrub about three feet high was laden with
clusters of dark fruit appearing, in color and shape, like huckle-
berries. Raccoons were feeding extensively on it.
PROCYON LOTOR MARINUS subsp. nov.
Chokoloskee Raccoon
Type—From near Chokoloskee, Florida. No. 254989, J adult,
U. S. National Museum, collected by E. W. Nelson, February 28,
1930.
General characters ——A very small subspecies, the largest old male
in the series taken weighing 8 pounds. Duller grayish than P. /. elucus,
of adjacent mainland, with skull much smaller, more depressed on
frontal area and molariform teeth proportionately much heavier.
Color.—Not very different from typical elucus but averaging duller,
more grayish on upperparts owing to smaller amount of black tips to
overlying hairs; rusty buffy nape patch averaging less strongly
marked, often obsolescent ; light rings on tail paler, less buffy yellow-
ish; black mask more restricted with remainder of top and sides of
head paler. No indication of the generally rusty or dark buffy suf-
fusion covering entire upperparts frequently present, and sometimes
strongly marked, in specimens of elucus from the type region.
Skull—Much smaller and more delicately proportioned than in
elucus; frontal area much more depressed, braincase more rounded ;
last upper premolar and carnassial relatively, and sometimes actually,
larger ; palatal shelf about the same.
8 SMITHSONIAN MISCELLANEOUS COLLECTIONS. VOL. 82
Measurements——Type: Total length, 665 mm.; tail vertebrae,
222; hind foot, 105. Skull (type): Greatest length, 105.9; condylo-
basal length, 101.8; zygomatic breadth, 64.8; interorbital breadth,
22.3; least width of palatal shelf (between last molar and interptery-
goid fossa), 13.9; upper canine-molariform tooth row (alveolar
length), 40; weight of type in flesh, 7 pounds. See page 12 for tables
of measurements and weights.
Remarks.—This is one of the smaller subspecies of raccoons that
have developed on the keys about the southern end of the peninsula
of Florida, not differing much in size from the form on the Key Vaca
Group. It appears to be limited to the great mass of mangrove covered
or bordered islands, or keys, known as the “ Ten Thousand Islands ”
where they exist in great numbers. Most of these islets are completely
covered by the sea to a depth of from about one to three feet at each
high tide, and are totally devoid of fresh water. As most of these keys
have no large trees to afford hollows and no dry land the raccoons
must make their homes on top of the mangrove roots where they are
forced to retreat by the incoming tide.
Specimens examined—t17: from type locality, 16; from Ten
Thousand Islands (exact locality unknown), 1.
PROCYON LOTOR INESPERATUS subsp. nov.
Matecumbe Raccoon
Type.—From Upper Matecumbe Key, Florida. No. 255037, J
adult, U. S. National Museum, collected by E. W. Nelson, March 19,
1930.
General characters—In weight and color resembling typical P. 1.
elucus of adjacent mainland but both body and skull measurements
smaller, especially length of hind foot; frontal area of skull much
more depressed.
Color—Much as in typical elucus, strongly washed with black on
upperparts and well marked nuchal patch of dark rusty or buffy, dark
rings on tail well marked and light rings often strongly buffy.
Skull—Differs from that of elucus in smaller size and marked
depression of frontal area. From P. /. marinus skull may be distin-
guished by its much larger size, more massive proportions and smaller
molariform teeth.
Measurements—T ype: Total length, 730 mm.; tail vertebrae, 250;
hind foot, 115. Skull (type); Greatest length, 110.4; condylobasal
length, 108.1; zygomatic breadth, 68.2; interorbital breadth, 23.1 ;
least width palatal shelf, 15.2; upper canine-molariform tooth row, 41.
Weight of type in the flesh, 8.5 pounds. See page 12 for tables of
measurements.
No. 8 NEW RACCOONS FROM FLORIDA—NELSON 9
Remarks.—The present subspecies occupies the group of keys be-
ginning with Virginia and Biscayne Keys on the north side of the
entrance to Biscayne Bay and ranges south to the southern point of
Lower Matecumbe Key. Key Largo, the median island, is by far the
largest of this group and broad mangrove swamps bordering its
western side extend out, about the middle of its length, until only a
comparatively narrow channel separates them from the similar swamps
which extend eastward from the mainland, at the south end of
Biscayne Bay. The railroad and motor highway fills, and viaducts
extend across these swamps, from Miami to the middle of Key Largo
on their way down the keys to Key West.
The comparatively short distance separating the raccoons living on
these keys from those of the adjacent mainland, with the size and
color of the island animals, made me doubt any strong differentiation
when I was collecting them. Fortunately I was able to secure two
good males and a female on the adjacent shore of the mainland to
determine the question. The skulls of these specimens are typical
P. 1. elucus, with characteristic high arched frontal areas from which
all the skulls of the series from the various keys of this group may at
once be distinguished by their appreciable smaller size and more
flattened frontals.
The largest male taken on Key Largo weighed 12 pounds in the
flesh, the same as old males collected and weighed by Dr. E. A. Mearns
in Polk County, the home of typical elucus, but the skull of the present
form is smaller and flatter. It may be added also that specimens taken
on Upper Matecumbe and especially those from Lower Matecumbe
Key, the farthest point in the group from the mainland, show gradation
toward a smaller animal than those of Virginia Key and Key Largo.
Specimens examined.—15, all from the Key Largo Group, as
follows: Virginia Key, 2; Key Largo, 3; Plantation Key, 2; Upper
Matecumbe Key, 1; Lower Matecumbe Key, 7.
PROCYON LOTOR AUSPICATUS subsp. nov.
Key Vaca Raccoon
Type.——From Marathon, Key Vaca, Florida. No. 255080, ¢ adult,
U.S. National Museum, collected by E. W. Nelson, March 28, 1930.
General characters—Very small, about the same size as marinus
from which it may be distinguished by its grayer upperparts, more
brownish yellow pale rings on tail, more depressed frontal area on
skull and shorter palatal shelf. Its small size distinguishes it at once
from the other forms described here from the Florida Keys.
Io SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
Color.—General coloration rather paler grayish than in marinus
with tendency to a smaller black mask and more whitish head, rusty
nape patch brighter, light rings on tail broader, more brownish buffy,
black rings relatively narrower.
Skull——Similar in its delicate proportions to that of marinus but
more flattened over frontal area, braincase more rounded, and palatal
shelf shorter.
Measurements —Type: Total length, 644 mm. ; tail vertebrae, 214;
hind foot, 99. Skull (type): Greatest length, 99.8; condylobasal
length, 95.5 ; zygomatic breadth, 65.5 ; interorbital breadth, 19.4; least
width palatal shelf, 13.9; upper canine-molariform tooth row, 37.4;
weight of type in the flesh, 5.5 pounds. See page 12 for tables of mea-
surements and weights.
Remarks.—The small raccoon from the Key Vaca group in con-
nection with other raccoons of Florida requires comparison only with
P. 1. marinus. It is abundant in the mangrove swamps of Key Vaca
and the immediately adjacent keys. The range occupied by the Key
Vaca raccoon is the smallest and most compact of that of any of the
forms I found among the keys.
Specimens examined.—13, all from type locality.
PROCYON LOTOR INCAUTUS subsp. nov.
Torch Key Raccoon
Type—From Torch Key, Big Pine Key Group, Florida. No.
255000, ¢' adult, U. S. National Museum, collected by E. W. Nelson,
March 24, 1930.
General characters—Slightly smaller than inesperatus, upperparts
palest, most dingy gray of any of the forms described here; skull
comparatively narrow interorbitally with elevated frontal area more
like elucus than in the other key forms described here.
Color.—The palest gray of the key raccoons, black mask more
restricted, sometimes obsolescent, and elsewhere top and sides of head
whiter ; pale rings on tail broader, dark ones narrower and usually
dusky brown; rusty rufous nape patch usually present and sometimes
strongly marked. As the season advances the colors commonly bleach
until many are wholly dirty yellowish or dingy whitish.
Skull.—Differs from other key forms described here by greater
interorbital compression and distinctly more highly arched frontal
area, resembling that of elucus but less strongly arched ; zygomatic
breadth widest of the key forms; molariform teeth smaller propor-
tionately.
No. 8 NEW RACCOONS FROM FLORIDA—NELSON II
Measurements —T ype: Total length, 694 mm. ; tail vertebrae, 263 ;
hind foot, 218. Skull (type): Greatest length, 110; condylobasal
length, 105.3; zygomatic breadth, 67.3; interorbital breadth, 19.8;
least width palatal shelf, 15.1; upper canine-molariform tooth row,
38.8 ; weight of type, 8.5 pounds. See page 12 for tables of measure-
ments and weights.
Remarks.—The home of this form of raccoon is on the group of
Florida keys farthest from the mainland. As in the case of the other
forms described here they live mainly, and sometimes entirely, in
mangrove swamps without access to fresh water except during rains.
The brilliant light of their environment has affected their general
color more than in the others, as shown by their pale, faded tints.
Specimens examined.—z20, all from the keys of the Big Pine Group,
as follows: No Name Key, 5; Big Pine Key, 6; Torch Key, 2 (type
locality) ; Ramrod Key, 1; Boca Chica Key, 2; Stock Island 2; Key
West, 1.
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SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82, NO. 8, PL.
Z.
Fic. 1.—Procyon lotor clucus, 3, typical skull, Polk County, Florida.
Fic. 2.—Procyon lotor inesperatus, 3, type skull, Upper Matecumbe Key, Florida.
Skulls natural size.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLE. 82; NO:-8),.PE. 3
2.
Fic. 1—Procyon lotor auspicatus, 3, type skull, Marathon, Key Vaca, Florida.
Fic. 2—Procyon lotor incautus, 3, type skull, Torch Key, Florida.
Skulls natural size. The subspecies reproduced in plates 2 and 3 follow in the
geographic sequence of their occurrence from P. /. elucus of the mainland down
the three groups of the Florida Keys.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL; 182, INO: <8, (PEas4:
Fic. 1.—Procyon lotor elucus, 3, typical skull, Polk County, Florida.
Fig. 2—Procyon lotor inesperatus, 3, type skull, Upper Matecumbe Key, Florida.
Skulls natural size.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOENS 2; a NO mone Leno
Fic. 1—Procyon lotor auspicatus, 3, type skull, Marathon, Key Vaca, Florida.
Fic. 2.—Procyon lotor incaulus, 3, type skull, Torch Key, Florida.
Skulls natural size. Subspecies reproduced in plates 4 and 5 follow in their
geographic sequence as in plates 2 and 3.
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 82, NUMBER 9
THE FURTHER AND FINAL RESEARCHES OF
JOSEPH JACKSON LISTER UPON THE REPRO-
DUCMIVE: PROCESSES OF POLYSTOMELLA
CRISPA (LINNE)
(An Unpublished Paper Completed and Edited from
His Note-Books)
(WITH SEVEN PLATES)
BY
EDWARD HERON-ALLEN, F.R.S.
(PUBLICATION 3067)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
NOVEMBER 26, 1930
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The Lord Baltimore Press
BALTIMORE, MD., U. 8. A.
fot PURTHER AND FINAL RESEARCHES OF JOSEPH
JACKSON LISTER UPON THE REPRODUCTIVE
PROCESSES OF POLYSTOMELLA CRISPA
(LINNE)
(An Unpublished Paper Completed and Edited from His Note-Books)
By EDWARD HERON-ALLEN, F.R.S.
(WitH SEVEN PLaTEs)
INTRODUCTORY NOTE
For over 30 years Lister’s work upon the reproductive processes
of the megalospheric form of Polystomella crispa (Linn.) by means
of flagellispores has been familiar to protozoologists, and especially
to students, both in text-books and in the lectures of university pro-
fessors, and it recurs incessantly in examination papers. For over
30 years it has been well known that his equally, if not more, impor-
tant work on the reproductive processes of the microspheric form by
what may not improperly be called viviparity had been completed,
but with the exception of the short postscript to his paper published
in 1895 it has not been available to protozoologists. The reason for
this is difficult to fathom for anyone who did not know him inti-
mately. We know that his nervous sensibility was such as to reach
at times a pathological condition, and he could never be persuaded to
publish anything that he took it into his head to keep to himself.
I suppose that, so far as the Foraminifera were concerned, I was his
most intimate friend and fellow-worker, but, though we frequently
visited each other and continually corresponded, he never showed
me any of these preparations, photographs, or lantern slides.
I esteemed it therefore as a great privilege that I was allowed, after
his death, to sort and arrange the whole of his microscopical materials
and preparations, and the papers and note-books relating to them,
and having found the whole of his material, his slides and his own
unpublished paper upon it, I was permitted by the late Mrs. Lister,
herself a noteworthy zoologist, to give to the world of science the
information that it had been awaiting for 30 years.
When the last and the youngest of the men that he taught have
died; when the problems attaching to Astroclera have been solved ;
SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 82, No. 9
2 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
when his teaching in the Zoological Schools of Cambridge have be-
come a mere tradition; when his patient, laborious work upon the
geology, ethnology, and ornithology of the South Seas has become -
but a brick in the vast edifice of human knowledge; indeed so long
as men shall devote labor and research to the science of protozoology,
the name of Joseph Jackson Lister, like that of his illustrious uncle,
Lord Lister, and of Harvey, Jenner, Leeuwenhoek, Redi, and
Schwann, will live in the hearts and on the lips of men, and the dis-
covery which will spring to the mind when his name is mentioned
will be that of the reproductive processes of the reticularian Rhizopods,
especially those of Polystomella, and the meaning and significance of
dimorphism.
Those of us who were privileged to know him, and especially
those of us who in our researches have followed in his footsteps,
know how delicate and polite was his use of the results of his
amazing industry, patience, and almost universal interests—industry
and interests pursued and recorded indefatigably in the face of physi-
cal difficulties beneath which a lesser man would have early succumbed.
How much more profound must be the appreciation of one who,
like myself, has been privileged to examine his laboratory note-books
and the mass of valuable papers that he left behind him at his early
death—left in a condition of such exquisite perfection that the ex-
aminer is never once checked by the reflection: “‘ If only he were here
to explain exactly what he meant by that.”
I should hesitate, for fear of being accused of extravagance of
speech, to describe the note-books of Lister, were it not that they
lie before me as I write and that Mrs. Lister has presented them to
form part of the Heron-Allen and Earland library and collections
at the Natural History Museum (London), where they will be always
available, for the instruction and emulation of future generations of
protozoologists. He made notes, not merely for his own use in the
kind of persona] cryptogram which we are all apt to adopt in recording
our own observations, but for the guidance of any research-workers
that should come after him. Every sketch or drawing that he made
was as highly finished as though it had been prepared for publication,
and the systematic dating of his notes enables us to follow his work
step by step, and, for practically the whole of his working life, day by
day.
It was in the course of the examination that I have been privi-
leged to make, that I came across his own manuscript record of his
work at Plymouth, extending from June 1 to August 9, 1894, a
NO. RESEARCHES OF JOSEPH JACKSON LISTER—HERON-ALLEN
J
record of such vital interest and importance that no apology is needed
for publishing it in extenso. It is the record of his observations upon
the reproductive processes of the microspheric form of Polystomella
crispa (Linn.), and I have been able to compare this manuscript with
his original laboratory note-books, and with the records of his later
work in 1895, and in 1904-5.
Lister’s researches upon this organism appear to have commenced
in March-May, 1893, and it was during this period that he observed
the reproductive processes of the megalospheric form by means of
flagellispores, which observations he recorded, with a masterly study
of the dimorphism and nuclear conditions of the species, in his world-
famous paper read in June, 1894, and published in the Philosophical
Transactions of the Royal Society of Great Britain in 1895." By one
of those coincidences that are curiously far from uncommon in
science, Fritz Schaudinn was pursuing an identical line of inquiry at
the same time, and the same results were arrived at and published
almost simultaneously by both of these great men though working
quite independently and unknown to each other.”
But when Lister’s Philosophical Transactions paper was virtually
on the press, or at least awaiting publication, he was again at Plym-
outh, checking his former observations, and making further dis-
coveries which may well be described as epoch-making. Let him
tell us the story in his own words, ‘but first let us consider for a
moment the organisms in question: they are:
(a). Polystomella crispa (Linn.): one of the commonest known
species, and one of practically world-wide distribution.
(b). Polystomella macella (Fichtel & Moll) : a depressed and closely
allied species, found associated with P. crispa but somewhat rarer
north and south of the sub-tropical areas. I have recorded it myself
from Selsey, Torbay, Clare Island, Galway, St. Mawes, and St.
Andrews and the North Sea, so it may well have been among Lister’s
material: he had no time to be an accurate systematist.
Lister does not seem to have recorded observations upon
(c). Polystomella striato-punctata (Fichtel & Moll) until 1904. He
was again at Plymouth in June-July, 1904, as he says in his note-book,
“ hoping to maké some progress in the life-history of the Foramini-
fera.” He records his observations upon the reproduction of the
* Lister, J. J., Contributions to the life-history of the Foraminifera. Phil.
Trans., Vol. 186 B., p. 401, 1895.
* Schaudinn, F., Die Fortpflanzung der Foraminiferen und eine neue Art der
Kernvermehrung. Biol. Centralbl., Bd. 14, No. 4, February, 1894.
4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
megalospheric form of P. striato-punctata by flagellispores on June 29
and July 1. There is no mention of the reproduction of the micro-
spheric form by the processes we are now describing, though it is of
world-wide distribution, and nearly always found with P. crispa, and
it was the first Foraminifer ever figured—by Hooke in 1665 * and by
Leeuwenhoek in 1702.”
He was there again in the same months in 1905, but added nothing
of moment to his previous records, but records the fact that in
several of his dishes left behind in 1904, Foraminifera and other
organisms were still alive.
He printed a highly condensed abstract of these later observations
as a postscript, dated August 3, 1894, to his Philosophical Transactions
paper, and this postscript, slightly elaborated, appears im this still
contracted form in his later works, viz: in Lankester’s Treatise on
Zoology in 1903, his address as president of Section D at the York
Meeting of the British Association in 1906, and in his Evening Dis-
course at the Royal Institution in 1907.” But, for some reason which
we shall never know, but which was doubtless connected with his
state of health, he never published his paper. No apology is needed
therefore for giving to the scientific world his own description of
his researches, and their results. It is as follows: °
LISTERS NOLES
July 7, 1894. Plymouth—For the last three weeks or so, I have
kept Polystomella in tall glass jars,’ the water in which has been
renewed frequently by a jet playing through a muslin cover over the
mouth of the jar“. The Polystomellas crawl up the sides from the
* Hooke, R., Micrographia, p. 80, pl. v, fig. x, London, 1665.
*Van Leeuwenhoek, A., Sevende Verfolg der Brieven, etc., p. 105, pl. (opp.
PTOL) patio NGO RGD .elitmm7O2:
* Lister, J. J.. The Protozoa, Sect. 1. The Foraminifera, in E. Ray Lankester
“A treatise on zoology,” Pt. 1, 2nd Fasc., pp. 47-149, 1903.
* Lister, J. J., The life history of the Foraminifera. Brit. Assoc., York, 1906,
Section D, Presidential Address.
*Lister, J J., The Foraminifera. Proc. Roy. Inst. Great Britain, Evening
Meeting, February 15, 1907, p. 489, London, 1900.
“T have carefully compared this manuscript statement with Lister’s laboratory
note-books, and the further details to be derived from these are of such value
that I have transcribed such parts of his notes as bear upon his statement, in
the foot-notes.
*The Polystomella were dredged in 5 fms. on Zostera grass, inside Drake
Island in Plymouth Sound, and the experiments commenced on June 17.
* When this spraying was not in progress the jars stood on the sill of a window
in the sun. J. J. L.
ae
NO. 9 RESEARCHES OF JOSEPH JACKSON LISTER—-HERON-ALLEN 5
bottom and may be clearly seen when in bright light, with a black
background. Usually one or two of the terminal chambers contain
little protoplasm and appear empty. In the ordinary condition their
pseudopodia are inconspicuous, but they may generally be detected
with a pocket lens, extending in sheaves from the shell,” and with
a microscope they have the usual appearance of delicate straight threads
with granules.
In some 50 cases I have seen examples in the reproductive phase.
The first sign of this phase is seen in the character of the pseudopodia.”
Instead of being disposed as above described, the pseudopodia are
limited to a circular or oval area immediately about the shell, the
center of the area being the mouth of the shell, which is nearly always
flat against the glass. This area is covered completely by a close web
of radiating and interlacing pseudopodia, and its limits are fairly
sharply defined. The pseudopodia are so thick as to intercept the
light, giving rise to the appearance of a semi-transparent milky
halo about the shell. This may be called the “ premonitory halo.” It
is readily seen by the naked eye.
The animals are generally found in this condition in the early
morning.
In nearly all the cases that I have observed the halo has been
established by 6: 30 a.m.”
After the animal has remained in this condition for some hours,
the protoplasm begins to leave the shell, the strands of the pseudopodia,
especially the radiating ones, become thicker, and the protoplasm
withdraws itself, first from the peripheral ends of the outer whorl of
chambers, being massed in the terminal chambers. Gradually the pro-
toplasm leaves the shell, passing through the direct communications
between the chambers, and also, as is seen in specimens preserved at
this stage, through the canals which exist in the walls separating the
chambers from one another. The protoplasm is generally brown, owing
to the presence of brown granules.” As the protoplasm emerges into
the area of the halo, this becomes streaked with brown, at first in
*When they are observed immediately after the jet of water has been playing
into the jar, a long sheaf of the pseudopodia is seen pointing towards the current
set up in the water. After the water has become still, they are shorter and point
in various directions. J. J. L.
* This was first observed on June 21, 10: 30 a. m.
* While these observations were in progress, Lister frequently remained all
night in the Laboratory of the Marine Biological Association.
palate Til.
6 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
thin radiating streaks, which become thicker and fuse with one another
until the area is, to a greater or less extent, filled with the dark brown
mass.”
Specimens which are removed from the wall of the jar while
surrounded by the premonitory halo, and placed on cover-slips with
a small drop of water, will generally send out fresh pseudopodia and
attach themselves to the cover-slip. They may then be placed over
holes in filter paper through which a current of water flows, and the
hole be completely filled with water. In this way the animal is im-
mersed in circulating water. In some cases the pseudopodia thrown
out are of the short reticulated character of those forming the halo,
and these indicate that the process is going on uninterruptedly ; but
it frequently happens that though the animal attaches itself to the glass,
the pseudopodia are of the long, little anastomosing character of the
ordinary condition. In this case the protoplasm remains in the shell—
the animal having, owing to the disturbance, returned to the ordinary
state. If however the reproductive phase is continued, the process
may be observed under the microscope.
When the protoplasm has emerged from the shell, the whole mass
undergoes amoeboid changes of shape, and under the microscope may
be seen to be in a turmoil of movement,’ the protoplasm coursing
along in broad interlacing streams. The streams may be seen to
pursue a definite course, the protoplasm in any one part of the mass
moving in the same direction for many minutes without interruption.
When newly emerged, the brown granules are uniformly scattered
through the protoplasm. Gradually a mottled arrangement is pro-
duced owing to the appearance of clear spaces (regions) free from
brown granules. As the process continues these clear spaces (regions)
become larger and more defined, and they are then seen to be more
stationary than the remainder of the protoplasm, the streams of
granules flowing past them. They are not however entirely stationary.
In one instance the area free from brown granules first became
evident one and a half hours after the protoplasm had emerged from
the shell.
Gradually the streaming movements of the protoplasm become less,
as the clear regions attain greater prominence, and in about half an
hour after their appearance the whole mass becomes broken up into
lS Spent:
2 The mass which is at first diffuse, gradually draws together into a more
compact mass, the pseudopodia being almost entirely withdrawn leaving branch-
ing lines of fine granules marking the positions they had occupied. J. J. L.
*230) p.) tt:
NO. Q RESEARCHES OF JOSEPH JACKSON LISTER—HERON-ALLEN a
rounded spheres, some 50 p» in diameter, whose center consists of
clear granular protoplasm (in which I thought I detected a radiate
arrangement) the peripheral part being dark with the coarse brown
granules.’
In some cases part of the protoplasm remains in the terminal
chambers of the shell and divides into spheres in this position. This,
however, is not usual.
The spheres form at first a compact mass. Shortly, each becomes
surrounded by a close felt of very delicate anastomosing pseudopodia,
which when seen with a simple lens has the appearance of a mass of
white granules * and I supposed at first that a deposit of lime granules
had taken place about the group of spheres. They gradually draw
apart from one another, so that each can be seen as a distinctly iso-
lated mass. :
I have killed and stained a number of specimens at different stages
of this process. Those in the stage before the protoplasm has left the
shell, as well as those at the beginning of the later stage, have the
central chambers still filled with protoplasm. In all these specimens
it is obvious that it is a muicrospheric individual that is undergoing
the process. Again, the specimens mentioned above which, having
shown a “ premonitory halo,” returned to the vegetative condition, on
being removed to a cover-slip, are found on being decalcified and
stained, to belong also to the microspheric form. Altogether I sup-
pose I have some I5 specimens whose central chambers are still filled
with protoplasm and hence furnished evidence as to the form to which
they belong.
In a batch of specimens of Polystomella (some 200-odd) megalo-
spheric and microspheric specimens were nine to one. This is good
evidence that the reproductive phase I am dealing with is that of the
microspheric form.
The specimens killed in the first stage present, when stained, the
appearance I have often seen, the many nuclei being of irregular
shapes, and surrounded by numbers of “ stained strands.” In some
cases no definite nuclei can be detected, and only few of the strands.
When nuclei are present they do not extend into the terminal chambers
of the shells.
In specimens whose protoplasm has begun to emerge, faintly stained
round nuclei 10 » in diameter are found in the clear protoplasm
of the terminal chambers, while the innermost protoplasm still has
the diffused flush and contains strands of stained matter.
*2:40 p.m.
Sas 300). mm.
8 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
In some cases the protoplasm immediately surrounding these nuclei
has a yellowish red color, as though, perhaps, it contained material
which was passing in towards the nucleus. The nuclei 10 p» in di-
ameter are found in the later stages now described. They acquire
stronger staining powers as the sphere stage is approached.
Continued July 21—The duration of the stage in which the pro-
toplasm remains in the condition of closely aggregated spheres, and
which I call stage iii, varies from two to eight hours. The aver-
age duration in a number of cases is about four hours. The most
marked change which can be seen in the living condition during this
period is in the disposition of the brown granules in the spheres.
At first the center of the spheres is clear, the granules lying at their
periphery, while in the later part of the stage the granules occupy
the sphere leaving only a clear layer at the periphery.
After stage iii the development may pursue one of two courses:
I. In by far the greater number of cases that I have watched the
spheres lose their sharply defined outline and gradually fuse with
one another into one or a few lumps.” Sometimes long pseudopodia
are thrown out during the process of fusion but this does not always
occur. The lumps so found generally divide, forming smaller lumps
of varying size which move apart by means of pseudopodia, and also
exhibit amoeboid changes of form. It frequently occurs that one of
these lumps attaches itself to the empty shell, spreading over part
of the exterior and interior.’
The protoplasm remains in this condition for weeks. The smaller
lumps appear gradually to break up into granules, losing their definite
outline and apparently dying. The large lumps may retain their sharply
defined outline for weeks, the movement of the protoplasm becoming
gradually more sluggish, and the volume of the mass apparently de-
creasing. What their ultimate fate may be I have not seen—but it
appears probable that it is the same as that of the smaller divisions
(lumps) into which the protoplasm divided—gradually disintegrating.”
*7: 30-9: 30 p.m.
211-330 p. mi.
* These conditions lasted until July 1, many other specimens being kept under
observation. On this date Lister tabulates the stages of the process as follows:
Stage 1. Premonitory halo formed but protoplasm still in shell. Stage 2. Proto-
plasm emerged from shell but still in one mass. Stage 3. Protoplasm divided
up into isolated spheres. Stage 4. Spheres fused to form a common mass
sending out pseudopodia. Stage 5. The mass divided up into amoeboid lumps
with pseudopodia. Lister’s preparations of shells in all of these stages are in our
collection at South Kensington.
i
NO. 9 RESEARCHES OF JOSEPH JACKSON LISTER—-HERON-ALLEN 9
2. In some ten cases (out of say 150 which I have now seen) the
spheres have separated after some hours, sending out long and abun-
dant pseudopodia, and moving rapidly away from one another.
Within 24 hours a second chamber is added to the sphere,’ and a
calcareous shell is developed.’ In this condition they are readily recog-
nized as young megalospheric individuals.
July 22.—After writing the above yesterday it occurred to me that
the first of the two processes described above was an abnormal one—
the result of the impurity of the water. I therefore changed the water
1 This was first observed at 7:10 p. m. on July 2. Lister’s note (note-book, vol.
ii, p. 91) reads “ The empty shell is now surrounded by a host of small bodies.
These all possess a globular mass containing yellow protoplasm. In many cases a
second mass shaped like the second chamber of a megalospheric form, and more
transparent than the globular mass is present. In other cases two transparent
4-oval masses are present. Some of them have extended pseudopodia.” On July 3,
at 9 a. m. these second chambers had highly accentuated themselves. (See draw-
ing note-book iii, p. 27.) All the stages were observed on this day in another
specimen between 9 a. m. and 9: 40 p. m. (drawing in note-book, vol. ii, p. 92).
This had acquired its second chamber at 8: 30 a. m. on July 4.
* This was first observed at 12:20 a. m. on July 5. At 9:10 a. m. the former
specimen had its young “with a rough shell-hyaline material outside this.”
(Note-book, vol. 11, p. 90.)
*On July 7 (the day Lister began the account we are studying) at 10:15 a.m.
he writes (note-book, vol. ii, p. 101) ‘‘ The specimen which yesterday morning
at 10:30 a. m. was in stage 3, and whose spheres then separated, I have now
transferred to picric with several of the spheres attached. They are ‘young
megalospheric forms with a second chamber formed.” These observations were
continued and repeated day after day with the same patient observations and
accurate records. On July 11 Lister was able to construct a table, founded upon
23 specimens which had been observed, at the times passed in the stages, the
average being: Stage I, 3 hours. Stage 2, 2 hours. Stage 3, 7 hours. (On the
12th he observed the same phenomena in Rotalia beccarii (Linn.) of which he
made a beautiful drawing (note-book, vol. ii, p. 106).) The average for 12
specimens watched on July 13 was: Stage I, 4 hours. Stage 2, 2 hours. Stage 3,
8 hours. (The young Rotalia (66 specimens) gave an average of stage 1, 3 hours,
20 minutes. Stage 2, 1 hour, 30 minutes. Stage 3, 4 hours, 20 minutes. The
observations went on as before, thereby confirming the accuracy of his previous
records up to the 21st; on the 22nd he made the discovery as to the water which
has been recorded in his own words, which now follow. When making dredgings
at Plymouth, while this paper was awaiting publication (in April, 1929), we
had the good fortune to find, in a dredging from inside Drake Island (the
locality from which Lister obtained his material) a very large number of young
Polystomellae, in exactly the condition here discribed—i. ¢., a megalospheric
chamber, followed by one or two succeeding chambers. See our paper, “ The
Foraminifera of the Plymouth District,” E. Heron-Allen and A. Earland, Journ.
Roy. Micro. Soc., 1930, vol. 50, p. 194.
se) SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
in two of the jars, putting instead of the aquarium water, water from
beyond the Breakwater which I had in a bottle.
Today six individuals have passed through the reproductive phase
and all have pursued the second course—giving rise to young megalo-
spheric individuals. Those (4) in which the protoplasm separated in
the afternoon have already (9:35 p.m.) added a second chamber
to the megalosphere, while those which have separated since 7 o'clock
consist only of a spherical body, with abundant ray-like pseudopodia.
These six individuals pursued the course described above up to
the end of stage iii—the most noticeable feature about them being the
large area covered by the protoplasm in the second and third stages—
filling the area of the halo.
This clears up the process in a most satisfactory manner.”
To the abstract of the above given in his later works he merely
adds: ‘The whole of the protoplasm of the parent is used up in
the formation of the brood of young, the shell being left empty. The
process from the first appearance of the halo to the dispersal of the
young is complete in about 12 hours. ... . Each of the spheres was,
in fact, a megalosphere. The microspheric parent has given rise to,
indeed it has become, a brood of megalospheric young.”
Lister was an expert micro-photographer, and in April, 1895,
he obtained fresh material from Plymouth, and repeated his obser-
vations in his own laboratory, at Cambridge. He heads his notes:
“ Polystomella material started in dishes, April 30. Several micro-
spheric individuals have reproduced in the manner seen at Plymouth
last summer. In a few cases the fusion has occurred, but many have
formed normal young.” His observations, like those of 1894, were
made by the “hanging drop” method in the use of which he was
an expert. His notes at this time relate to the flagellispore method
of reproduction by the megalospheric specimens, but from May 22
to 24 he made a wonderful series of micro-photographs, 24 in number
from a single microspheric individual of which 15 were made into
lantern slides, which slides Mrs. Lister has presented to our collection.
A selection of these together with certain of the photographs (which
are now being published for the first time) are now available, for all
time, for the use of students of Protozoology. In all, 20 photographs
are here reproduced. They may be tabulated as follows:
*On July 25 he writes ‘ There is reason to believe that the aquarium water
is affected by the asphalt lining of the tanks.” He tested this with further
Polystomellae on the 26th in “outside water” between 8:30 a. m. and 7 p. m.
with brilliantly successful results. His notes on Polystomella cease, for this
time, here.
NO. 9 RESEARCHES OF JOSEPH JACKSON LISTER—-HERON-ALLEN neat
Lantern slide,
Heron-Allen
and Earland
Figure Series May 22, 1895 Collection
I aL 1:00 p.m 12/32 | The “ premoni-
2 Bee Brea, Ue © Tae Aimy poets te tory halo.”
3 1B 3) Amos a? 12/33 Stage ii.
4 E. 4 AvAQma. 12/34 ey
5 E.5 Gz 12/35 a
6 E.6 O700l 12/36 #
7 B.7 G2 sau. 12/37 i
8 E. 8 TAOS ee MURA iy a mere ence Stage iii.
9 E.9 SATS ee a) VP Nahar seers -
10 E. 10 8:55 7 12/38 ”
EE Bont On60--%* 12/39 uy
12 E. 12 Ors O mee 12/40 e
Tes ets MOR 12/41 2
14 E. 14 DES) oe 12/42 Final stage.
15 E.15 nits Age 12/43 ag
16 E. 16 EY SO 12/44 a
U7, a7, Tha55. 4 12/45 -s
May 23, 1895
18 E. 19 OAL ay | ours e y
May 24, 1895°¢
19 E. 19 iS oy 12/49 a
20 F.r&2
" Deep view.
» Nearer view.
© There is something wrong here. J. J. L. dated this slide May 23, 1895,
II: 50 a. m. but it obviously belongs here as a second chamber is in process of
formation. I think it should have been dated May 24. After E. 18 (which is
not reproduced) the slides are not numbered, but his register of photographs
gives the numbers 109 et seq.
The last of these, figure 20, is one of the two slides in the collection,
showing different magnifications of the same object, which are regis-
tered as Ser. F. 1 & 2 but no dates or times are given. They are from
another specimen and represent the post-final stage (probably 25-26
May) when a second chamber has been added. They are registered
in the Heron-Allen and Earland Lantern-slide Collection as 12/46-47.
In conclusion I may be allowed to pay a tribute of warm gratitude
to Mrs. Lister, whose early death occurred shortly afterwards, for
the public spirit which led her to present the Lister Collections in
general, and these priceless note-books, preparations (there are nearly
450 of P. crispa alone in our collection at South Kensington), and
lantern slides, to the nation for the use of future generations of
students. I desire, also, to express my gratitude to Dr. J. A. Cushman
for correcting the proofs of this paper.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82, NO. 9. PL. 1
For explanation, see page IT.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82, NO
For explanation, see page II.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLE S25 NO oy rie.
For explanation, see page IT.
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LZ “1d ‘6 “ON ‘28 “10A SNOILO31100 SNOANVITISOSIN NVINOSHLINS
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 82, NUMBER 10
MORPHOLOGY OF THE BARK-BEETLES OF THE
GENUS GNATHOTRICHUS EICHH.
BY
KARL E. SCHEDL
(PUBLICATION 3068)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
JANUARY 24, 1931
The Lord Baltimore Press
BALTIMORE, MD., U. S. A.
MORPHOLOGY OF THE BARK-BEETLES OF THE
GENUS GNATHOTRICHUS EICHH.
By KARL E., SCHEDL
CONTENTS
‘Liciynaahvie itor ead c Geen Ganictnc 8 Gisie ote re a Ce BOR eS oie eirs SO nes numa eee
EMstractsvoithe literature tp) tondates as sacs nese ies nee. ok
Numer Ie seCIItl Senate core eucyescterswstevel vias tae yamsatnithe Ceara a ae aseiatnn wsla ores
B: Gnathotrichus matertarius Fitch............0.+.2..+-+eceeess>
PVE COLE VATI CUE apet a cpeter ie eke icy ete il « ase icvokean te onsitucnete hens lst Netane ie snc reree @RGS SoS
Debrstantaa (OTe TIDINGS imetete rare saver al cay stave eis serern ereial onset eevee Rhee Rae Gites
General appearance, vestiture, color and size.....................
“TTESUNEETA SS 5s eee SEROMA CCR A Oi cic: teas De Oa SORES ets
sihnesapperidaresuor the headiss svar ceeraiecie st ates Seine eesti waver
MIB eeaATItEN TAGs fecere\s, 5 Sccie ciseno ey ORES RIO ee eS rae al sd ee telist ie ohs
BINTE STOUT PALES 1; tera ected sreyey otabeeear ares euros Son eats oy arcane aires ons
“OLAV ARG) eb Sieben oP RiP MC RRR SIO RN i tics ecat ye Aum ia I
PRhetprothorax wane aie errr t ee AC re ak nan eee
silseemesothoraxps Hopkins; Aw Di(a4):
The author examined one specimen from Chaleo and another from
Michocan, Mexico, and suggested that Blandford’s Gnathotrichus
consentaneus is identical with sulcatus Lec.
1905... CURRIE, Jka Po (32)
Copy of Hopkins, A. D. Fir is replaced by lowland fir.
LO07. FALL, EC. and Cockerrrn, DAG G@s6)r
Gnathotrichus sulcatus (Hopkins MS.). —— Beulah (Skinner).
1909. SwaIne, T. M.(40).
References to literature only.
1910. Haceporn, M.(42).
References only.
1914. SWAINE, J. M.(47).
See retusus Lec.
1918. SwaIne, J. M.(49).
No new data concerning the description and the biology. Host
trees: Grand fir, western hemlock, Douglas fir, western white
pine. Distribution: Generally distributed throughout southern
British Columbia, extending southwards. In sap-wood and heart-
wood of dying and recently killed trees, and more rarely in those
apparently sound.
1922. Hoppine, R.(5r).
According to the author Gn. sulcatus breeds in the following host
trees: Pinus monticola Dougl., Abies concolor Parry, Abies mag-
nifica Murr., Abies grandis Lindl., Pseudotsuga taxifolia Britt.,
and Tsuga heterophylla Raf.
SYNONYMA
A. Gunathotrichus materiarius Fitch
Tomicus materiarius Fitch (Fitch, Schwartz, 1890, Packard, 1890).
Crypturgus materiarius Fitch (Zimmerman, Packard, 1869).
Gnathotrichus corthyloides Eichh.
Pityophthorus materiarius Fitch (Leconte, Packard, 1881).
Cryphalus materiarius Fitch (Provancher).
B. Gnathotrichus retusus Lec.
Cryphalus retusus Leconte, 1868.
Pityophthorus retusus Lec. (Leconte, 1876).
NO. I0 MORPHOLOGY OF GNATHOTRICHUS—SCHEDL 15
C. Gnathotrichus sulcatus Lec.
Cryphalus sulcatus Leconte, 1868.
Pityophthorus sulcatus Leconte, 1876.
Common NAMES
A. Gn. materiarius Fitch.
Pine timber-beetle (Fitch).
Timber-beetle (Hopkins, 1893).
Eastern pine wood stainer (Hopkins, 1904, Felt, 1906).
B. Gn. retusus Lec.
Timber-beetle (Hopkins, 1893, Swaine, 1914).
Pine hole borer (Swaine, 1914).
C. Gn. sulcatus Lec.
Western hemlock wood stainer (Hopkins, 1904).
Tue CHITINOUS SKELETON OF THE ADULT
GENERAL APPEARANCE, VESTITURE, COLOR, AND SIZE
The general form of all three species is slender in both sexes,
cylindrical, with the head concealed from above by the pronotum. The
pronotum is longer than wide with the sides parallel on more than
the caudal half and broadly rounded in front. The elytra are slightly
narrower or as wide as the pronotum, subparallel as far as the origin
of the declivity, sometimes slightly tapering posteriorly ; moderately
broadly rounded behind.
Length and relative proportions. (Dorsal aspect) .—
Gnathotrichus materiarius Fitch.
Length of the body (elytra and pronotum; head concealed
from above), 3.06 mm.
The body is 3.11 times as long as the width of the pronotum.
Width of pronotum, 0.98 mm.
The elytra are 1.46 times as long as the pronotum.
Examined specimens, 12.
Gnathotrichus retusus Lec.
Length, 3.72 mm.
The body is 3.23 times as long as the width of the pronotum.
Width of the pronotum, I.15 mm.
The elytra are 1.54 times as long as the pronotum.
Examined specimens, 10.
VOL. 82
MISCELLANEOUS COLLECTIONS
SMITHSONIAN
16
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NO. 10 MORPHOLOGY OF GNATHOTRICHUS—SCHEDL V7,
Gnathotrichus sulcatus Lec.
Length, 3.59 mm.
The body is 3.38 times as long as the width of the pronotum.
Width of the pronotum, 1.07 mm.
The elytra are 1.60 times as long as the pronotum.
Examined specimens, 10.
_——
UNW—ulsw——yWd
Fic. 2—Gnathotrichus materiarius Fitch, adult male, ventral aspect.
An, antenna; Cc, coxal cavity; Co, coxa; Ep, epistoma; Epm, epimeron; Epst, episternum;
Ey, compound eye; Gu, gular area; Hst, hypostoma; La, labium; Msst, mesosternum; Msth,
mesothorax; Md, mandible; Mtst, metasternum; Mtth, metathorax; Mz, maxilla; Pr,
pregula; Prst, prosternum; Prth, prothorax; st, sternite; St/, sternellar area; a, median line.
Color —The color ranges from a reddish-brown to nearly black in
mature specimens; it is of no importance in this genus.
V estiture —In general it can be said that all three species are with-
out extraordinary hair characters; few large bristles occur on the
declivity. However, by careful examination under the microscope it
was found that all external parts are more or less covered with fine
hairs.
VOL. 82
SMITHSONIAN MISCELLANEOUS COLLECTIONS
18
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NO. I0 MORPHOLOGY OF GNATHOTRICHUS—SCHEDL 19
Sculpture —The front and the elytral declivity are the only bearers
of specific modifications. The sculpture of the pronotum and _ the
elytra, which is very useful in distinguishing the species in many other
genera of the Scolytidae, does not vary to any extent in this genus.
Secondary sexual characters—-Secondary sexual characters were
found in the development of the hairs on the antennae, the number of
fully developed tergites and in the number of spiracles mainly.
THE HEAD
The head capsula or cranium (figs. 4-7) is dorsally divided by the
epicranial suture. The sutura fronto verticale Berlese or coronal su-
ture (figs. 5, 6, Cos) is distinct in all three species examined. In
Gnathotrichus retusus Lec., and less pronounced in Gn. materiarius
Fitch, this suture and a short piece of the sutura metopica da Miall
and Denny or frontal suture become elevated near the junction, form-
ing a Y-like ridge. The slightly raised line (figs. 4, 5, a) which origi-
nates at the upper margin of the antennal groove (figs. 4, 6, b, 7, c)
and which is directed toward this junction may be considered a rem-
nant of the frontal suture. These anterior remnants of the frontal
suture are never connected with the posterior portion.
Front.—The area between the frontal sutures is largely occupied
by the frons (figs. 4, 5, 7, Fr). The shape of the front is plano-
convex ; the median line is slightly raised, and there is a shallow de-
pression on each side of the latter. The front is sculptured by scratches
which radiate from the center of the anterior margin, and which ex-
tend over the entire sclerite, becoming less distinct and less dense
towards the outer margin. These scratches look as though they had
been made with a needle point, and therefore have been called accicu-
lation by several authors. Scattered over the acciculate area are small
but deep, sparse punctures. Bristles varying from short to moderately
long occur over the entire front; most of these originate in punctures.
They are more numerous in the antero-lateral corners.
Specific modifications :—
A—Acciculation strongly developed, close; front with punctures near
the outer margin and antero-lateral region only, with bristles
very sparse in the acciculate, area (fig. 5, Ac).
Gn. sulcatus Lec.
AA—Acciculation weakly developed or obsolescent, more or less re-
stricted to the median anterior area; front elsewhere punctu-
late, bristles more numerous.
Gn, retusus Lec. and materiarius Fitch.
20 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
Labrum, clypeus and epistoma—The labrum and clypeus are not
evident as separate sclerites. The epistoma (figs. 4, 5, 7, ep), probably
also containing elements of the labrum and clypeus, is not separated
from the front by a distinct suture or line. It is present as a ridge and
forms the anterior margin of the front. This region is widened later-
ally, and is connected with an elevation which surrounds the antennal
Fic. 4.—Gnathotrichus sulcatus Lec.: Head, lateral aspect.
An, antenna; Ep, epistoma; Ey, compound eye; Fr, frons; Ge, gena; La, labium; Md,
mandible; Mz, maxilla: Oc, occipital area; Ve, vertex; a, frontal suture; b, antennal
groove.
Fic. 5.—Gnathotrichus sulcatus Lec.: Head, dorsal aspect.
Ac, acciculation; An, antenna; Ep, epistoma; Ey, compound eye; Fr, frons; Md, mandible;
Mx, maxilla; Cos, coronal suture; Ve, vertex; a, frontal suture.
scrobe. In the middle of the anterior margin, an evenly rounded
emargination is visible which may be called serratus epistomalus
(fig. 7, SE) and which corresponds to Hopkin’s median impression.
From the serratus epistomalus the acciculation of the front radiates.
The epistoma is without sculpture and bears a row of bristles on its
posterior margin. The mandibles articulate with the underside of the
epistoma.
iN(GE 106} MORPHOLOGY OF GNATHOTRICHUS—SCHEDL 21
Gula.—The gula is bounded by the two very closely placed gular-
sutures (figs. 6, 7, Gus) and widens anteriorly to form the pregula.
Fic. 6.—Gnathotrichus sulcatus Lec.: Head, ventral aspect.
An, antenna; Cos, coronal suture; Ey, compound eye; Ge, gena; Gus, gular sutures; Hy,
hypostoma; La, labium; Md, mandible; Mx, maxilla; Oc, occipital area; i
bi ( i ca, occipital
apodeme; Oct, occipital foramen; Pr, pregula; a, dorsal articulation of the mandible; },
ventral articulation of the mandible; c, antennal groove.
R
Fic. 7.—A, Gnathotrichus sulcatus Lec.: Head, oral aspect. B, Gnathotrichus
retusus Lec.: Pregula.
An, antenna; Ep, epistoma; Ge, gena; Gus, gular sutures; Fr, frons; Hy, hypostoma;
La, labium; Mx, maxilla; Pr, pregula; SE, serratus epistomalus; a, dorsal articulation of
mandibles; b, ventral articulation of mandibles; c, antennal groove.
These two distinct sutures appear to be exceptional in this family.
Hopkins apparently did not find any double sutures in his investi-
gations.
22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
In Gnathotrichus, the pregula (figs. 6, 7, Pr) is always without
sculpture or hairs. The very distinct specific modifications are de-
scribed in the following key:
A—Pregula flat, triangular, not produced anteriorly, its anterior mar-
gin continuous with those of the géna.
Gn. sulcatus Lec.
AA—Pregula convex, produced anteriorly, its anterior not continuous
with those of the gena.
B—Pregula very convex, extending far beyond the genal
margin.
Gn. retusus Lec.
BB—Pregula feebly convex, extending slightly beyond the
genal margin.
Gn. materiarius Fitch.
E picranium.—The remaining lobes of the cranium situated between
the epicranial suture, the gula, and the foramen bear the compound
eyes (figs. 4-6, Ey), the antennae and the articulations of the mouth-
parts. For merely descriptive purposes these lateral areas of the
cranium have been divided into several regions. The occipital area
(figs. 4, 6, Oc) surrounding the occipital foramen is not limited an-
teriorly in Gnathotrichus. However, there occur obscure lines which
may be regarded as homologous with the occipital suture. The epi-
cranium or parietals (Crampton), the gena, and the vertex do not
show any sculpture which exhibits specific differences. The hypo-
stoma (Hopkins) (figs. 6, 7, Hy), an area corresponding to the epi-
stoma, which like the former belongs morphologically to the epicra-
nium, is well developed. It is in the form of a semicircular band and
bears at its extremities the ventral articulations (figs. 6, 7 b) of the
mandibles. From the oral aspect it is visible; from the ventral, it is
hidden by the pregula and the gena.
Other regions, such as the pregena (Hopkins), etc., are not at all
distinct or limited and are therefore of no interest in this discussion.
THE APPENDAGES OF THE HEAD
THE ANTENNAE
The antennae present good generic characters ; the species modifica-
tions are less important. The sexual differences are distinct also. All
the longer setae of the antennae are feathered.
1 The hypostoma, as the term is used here, or the ventral angles of the postgena,
is the bearer of the maxillare and not the labium. A submentum, as this term
was used by Hopkins in Dendroctonus, is not defined by sutures.
NO. 10
MORPHOLOGY OF GNATHOTRICHUS—SCH EDL 23
The scape (fig. 8, A, S) is slender and clavate toward the apex.
Hairs and punctures are rather sparse. The scape is about as long
as the funicle and the club together. The funicle (fig. 8, A, F) is
Fic. 8, A.—Gnathotrichus retusus
Lec., adult female: Antenna, interno-
lateral aspect.
C, antennal club; F, funicle; S, scape; p,
pedicle; 2, 3, 4, 5, joint of the funicle; 1’, 2’,
3’, joints of the club.
Fic. 8, C—Gnathotrichus sulcatus
Lec., adult male: Antenna, externo-
lateral aspect.
pill
=e
vy
h
fi
Fic. 8, B—Gnathotrichus retusus
Lec., adult female: Antenna, externo-
lateral aspect.
Fic. 8, D.—Gnathotrichus sulcatus
Lec., adult male: Antenna, interno-
lateral aspect.
five-jointed and distinctly shorter than the club. The pedicle (fig.
8, A. p) the first and longest of all joints, is as long as joint two
and three together. The setae and punctures are more concentrated
on the apical half of the pedicle, only two small bristles occurring
near the basal articulation (fig. 8, B). The other joints decrease in
24 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
length but increase in width towards the club. All the joints bear
numerous punctures and hairs except the second which has neither.
The club (fig. 8, A, C) is from 1.21 to 1.35 times as long as wide,
egg-shaped in outline, widest near the apex and strongly compressed.
Two nearly continuous septate sutures divide the club into three
joints. The first and third joints are nearly equal in length; the
second is distinctly shorter. The septae are arcuate and distinctly
visible from the externo-lateral aspect (fig. 8, B, C). On the interno-
lateral side (fig. 8, A, D) they are indicated by single sutures. Each
of the joints is covered by numerous hairs and punctures. Externo-
laterally these are closely placed and arranged in arcuate rows ; interno-
laterally, they are sparse, and the punctures are more numerous.
The females (fig. 8, A) bear, moreover, on the interno-anterior
margin of the club a few very long hairs. The anterior setae of the
third, fourth, and fifth joints of the funicle are longer in this sex.
These are the only external characters by which the sexes may be
distinguished when the elytra are kept in the closed position.
Specific modifications :—
A—Septae in the form of continuous bands of equal width through-
out, slightly less pronounced medially; externo-lateral side
of the club with minute, transverse wrinkles and small
punctures producing a slightly roughened surface (fig. 8,
ENG IE Ne
Gn. retusus Lec.
AA—Septae wider laterally, indistinct and narrow medially ; externo-
lateral side of the club smooth or with very minute, sparse
wrinkles; club stouter.
B—Externo-lateral side of the club smooth, interno-lateral
side with numerous hairs and punctures (fig. 8, C).
Gn. sulcatus Lec.
BB—Externo-lateral side of the club with minute wrinkles,
interno-lateral side with very few hairs and few
punctures ; club very stout.
Gn. materiarius Fitch.
THE MOUTHPARTS
' The mouthparts present generic as well as specific characters. They
include the mandibles, the first maxilla and the second maxillae or
labium.
NO. IO MORPHOLOGY OF GNATHOTRICHUS—SCHEDL 25
Mandibles—The mandibles (fig. 9) are very much alike in all
three species, so much so that it is not possible to distinguish them
by mandibular characters. The shape is stout and triangular as in
most Scolytidae. The apical tooth (fig. 9, a) and the subapical tooth
(fig. 9, b) are well developed and separated by a deep emargination.
In contrast to other genera, two median teeth (fig. 9, c) occur. The
latter are rather feebly developed and the separating emargination is
shallow. The molar tooth (fig. 9, d) is evenly rounded. There are
one lateral and two dorsal setae (fig. 9, f, g). Other details are
illustrated in figure 9. No specific differences have been found in
these.
Fic. 9.—Gnathotrichus materiarius Fitch: Mandible, dorsal aspect.
a, apical tooth; b, subapical tooth; c, median teeth; d, molar tooth; e, dorsal area; f, lateral
bristle; g, dorsal bristles; h, transverse ridge; i, anterior fossa; k, condyle; /, posterior
fossa; m, posterior impression; n, condyle of ventral articulation; 0, extensor tendon; »,
retractor tendon; q, extensor disk; r, retractor disk.
The Maxilla—The maxillae (figs. 2, 6) are exposed on each side
of the labium. Each is held in such a way that the cardo is parallel
with the long axis of the head; the stipes, lacinia, and galea are at
right angles to the cardo; the second and third palpial joints are
slightly curved outwards. The galea and lacinia are represented by
two lobes which are fused except at the apex where they are sepa-
rated by a moderately deep notch. The stipes is separated from these
fused lobes by a very fine, feeble suture on the outer side; on the
inner side, no suture is evident. Another paper will deal with the
generic differences in the maxilla of Guathotrichus Eichh. and re-
lated genera. Nevertheless, it should be noted that, as far as known
at present, the labium and the maxillae in the genus Pityophtorus
Eichh. always bear at least a few feathered bristles ; in Gnathotrichus
26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
Eichh. these bristles are simple. This is important, as it has been
seen that the character “ maxilla spinose ” in Pityophthorus Eichh.
and “ maxilla pilose”? in Gnathotrichus Eichh., as used by several
Fic. 10.—Gnathotrichus sulcatus Lec.: Maxilla, outer aspect.
C, cardo; G, galea; P, palpus; 1, 2, 3. first, second and third joint of the palpus; a,
articulation of the maxilla; b, stipes; c, palpi re area; 1, anterior emargination separating
galea and lacinia; k, dorsal setae of the galea; /, median setae of the galea; m, cardol setae.
} | -
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SiG me
Ni
Fic. 11.—Gnathotrichus sulcatus Lec.: Maxilla, inner aspect.
C, cardo; L, lacinia; P, palpus; 1, 2, 3, first, second and third joint of the palpus; a,
articulation of the maxilla; b, stipes; c, palpiferal area; d, subgaleal area; e, posterior setae
of the lacinia; f, dorsal setae of the lacinia; g, median ‘setae of the lacinia; h, papilla.
authors to separate Gnathotrichus Eichh. from the rest of the Pityoph-
thorinae does not hold, as will be shown in the discussion of the
lacinia and galea. The maxillae exhibit no specific differences of
importance in Gnathotrichus. The maxillae are illustrated in figures
1OanGd iL.
NO: LO MORPHOLOGY OF GNATHOTRICHUS—SCHEDL 27
Cardo.—The cardo (C) is similar in shape in all three species and
is articulated with the ventral angles of the postgena in such a way
that the maxilla can be moved laterally as well as in a dorsal-ventral
plane. The longitudinal axis of the cardo and that of the stipes (b)
enclose an angle of about ninety degrees or even a little less. The
articulation of the stipes and the cardo is similar to that of other
genera of the family. There is also an articulation between the sub-
galea and the cardo. In figure 12, which illustrates this fact, the
cardo and the rest of the maxilla are stretched to show the connection.
The inner side of the cardo is smooth; the outer side shows from five
to seven hairs which may be called the cardol setae (m).
Stipes.—On the outer side the stipes is defined as an elongate, sub-
parallel sclerite; on the inner side it is fused with the lacinia. The
Fic. 12—Gnathotrichus materiarius Fitch.: Maxilla, stretched, the subgalea
showing.
setae are not very numerous, about six to eight being present. These
are more concentrated anteriorly near the base and near the apex. The
latter, situated on the apical area of the stipes, may correspond to
the palpiferal setae of some authors. The palpifer (c) being a topo-
graphical area of the stipes only, is not limited by lines or sutures.
Lacinia and galea.—The lobus internus or the lacinia and the lobus
externus or the galea are largely fused to form a single large lobe.
Apically they are distinctly separated by a moderately deep emargina-
tion (i) or notch. The inner free lobe is distinctly larger than the
outer one. Posteriorly they are fused. It will need much further
investigation to decide the exact areas of these two lobes. In the
meantime it is proposed to call the inner lobe and the inner surface
of the combined lobes the lacinia and the outer free lobe and the
outer surface of the fused part the galea.
28 SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOL. 82
The lacinia bears on the anterior portion of the free dorsal margin
short, stout, blunt setae (f) which are slightly incurved at their tips.
Fic. 13, A.—Gnathotrichus retusus
Lec.: Labium, ventral aspect.
L, ligula; M, mentum; Pp, palpifer; a,
base of ligula; b, neck of mentum; c, ventral
setae of the palpifer; d, anterior part of
ligula; e, first joint of palpus; f, second joint
of palpus; g, third joint of palpus.
Fic. 13, B.—Gnathotrichus retusus
Lec.: Labium, dorsal aspect.
P, palpus; L, ligula; a, articulation of the
labium; b, base of ligula; c, dorso-lateral
setae of the palpifer; d, dorso-anterior setae
of the ligula; e, f, g, first, second, and third
joint of the palpus.
Fic. 13, C—Gnathotrichus sulcatus
Lec.: Labium, ventral aspect.
Fic. 13, D.—Gnathotrichus sulcatus
Lec.: Labium, dorsal aspect.
These setae gradually become longer, more slender, and pointed pos-
teriorly (e). Near the subgaleal area (fig. 11, d) they are about three
times as long as on the dorsal margin. Besides these setae on the outer
NO. I0 MORPHOLOGY OF GNATHOTRICHUS—SCHEDL 29
margin of the lacinia numerous bristles (g) are scattered over the
entire lobe ; the surface is slightly reticulate. Basally, the united lacinia
and galea end in a lobe, the subgaleal area (d), which is distinctly
defined on the inner side by an impressed line. The galea bears a
short row of slender setae (k) on its free dorsal margin; the surface
is armed with rather sparse, long hairs (e).
Adopting the terms of other authors, it may be said the lacinia is
pilose on its dorsal margin and spinose on its posterior and that the
galea is spinose.
Palpi—tThe palpi (P) are three-jointed. The first joint is the
longest, the second and the third are subequal in length. Anteriorly
joints one and two bear setae, joint three punctures only. There occur
slight differences in the size and the shape of the joints in the species,
but they are not distinct enough to be of importance.
Labium—tThe labium (fig. 13), consisting of the mentum, the
ligula, and the palpi mainly, presents the most important generic and
specific characters of the mouthparts.
The shape of the mentum (fig. 13, M) is similar in all three species.
It is flask shaped, the palpifera always being wider than the neck
(fig. 13, b) and the base of the mentum and the latter always nar-
rower than the ligula (fig. 13, L). The basal portion of the mentum
is feebly widened in Gn. retusus Lec. (fig. 13, A) and strongly widened
in Gn. materiarius and sulcatus (fig. 13, C). In Gn. sulcatus and
materiarius, the mentum and the ligula are separated by deep sutures
throughout, even at the extreme base of the ligula; in retusus they are
similarly distinct except at the extreme base before which they com-
pletely disappear. The sculpture of the mentum on the ventral side
(fig. 13, C), except for a few distinct punctures, is smooth in Gn. sulca-
tus. Gn. materiarius is very similar but it shows slight evidences of
transverse wrinkles also. In Gn. retusus the punctures are obsolete ; the
wrinkles are strongly developed and become on the neck of the mentum
and the basal half of the palpifer more or less toothlike (fig. 13, A). All
the setae are simple bristles (c), and there are eight to ten of these
bristles on each half of the ventro-lateral side of the apical third of
the palpifer. In Gn. retusus and materiarius there are ten of these
bristles ; in sulcatus eight is the usual number.
Ligula—lIn this genus, the ligula (L) is much larger and more
prominent than is usual; it is wider than the mentum in its basal
half and is distinctly limited by sutures at the extreme base of the
labium except in retusus. The ligula (fig. 13, L) extends in all three
species a little farther anteriorly than the first joint of the palpus. Its
30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82 »
shape is elongate ; it is subparallel with two more or less distinct con-
tractions when viewed from the dorsal aspect. The basal portion is
semicircular in cross-section, while the anterior, unfused part is
strongly compressed. In Gn. retusus Lec., the ligula is more parallel
and the contractions less distinct than in the other species.
The anterior margin is very feebly, shallowly emarginate or evenly
rounded ; however, this does not seem to be constant in individuals
of the same species. On the ventral aspect, there is on each side near
the anterior margin a row of setae (d). No specific differences have
been found either in the shape or in the number of these setae which
vary in number from three to seven in each row. The sculpture of the
ligula on the ventral aspect in all three species is very much alike,
being nearly smooth or feebly transversely wrinkled. Dorsally Gn.
retusus shows similar transverse, toothlike wrinkles on the mentum;
the basal portion only is smooth. Gn. materiarius and sulcatus have
the corresponding area smooth with slight indications of transverse
wrinkles on the sides.
Palpi—trhe palpi are directed ventrad. Each is composed of three
segments, decreasing in size toward the apex. Segments two and
three bear dorsally a plush-like arrangement of hairs, while the first
joint has a few scattered hairs only. Ventrally, the setae are more
sparse and are intermixed with punctures on segments one and two;
segment three has no setae but a few punctures.
Summary of the specific characters :—
A—Base of the mentum scarcely widened basally, fused with the
ligula before the basal margin of the latter; mentum and
ligula when seen from above with strongly developed, tooth-
like, transverse wrinkles ; ligula subparallel. Gn. retusus Lec.
AA—Base of the mentum distinctly widened basally, extending to
the basal margin of the labium; mentum and ligula when
seen from below smooth or with slight indications of trans-
verse wrinkles ; contractions of the ligula well developed.
B—Ventral side of the mentum smooth, with few punctures.
Gn. sulcatus Lec.
BB—Ventral side of the mentum with transverse wrinkles.
Gn. materiarius Fitch.
THE THORAX
The three segments of the thorax, pro-, meso-, and metathorax
_ (figs. I, 2 and 3) are clearly defined in this genus as is usual in this
family. Each of these segments has as a basis three primitive ele-
NO. IO MORPHOLOGY OF GNATHOTRICHUS—SCHEDL 31
ments, the dorsal plate, or tergum, the ventral plate, or sternum, and
the lateral area, or pleuron. The thoracic segments and their elements
vary considerably in shape and size. The protergum, or pronotum, is
nearly twice as long as the tergum of the metathorax. The mesoter-
gum, still shorter than the latter, is represented by a short triangular
area only. The prosternum is about half as long as the metasternum
and about one-third the length of the pronotum, giving the pleural
area of the prothorax the shape of a trapezium. The mesosternum is
present as a plate nearly equal in size to the mesotergum; the meso-
pleura are also developed in proportion, giving the mesothorax the
shape of a short tube. The metathorax, more complicated in struc-
ture, represents the segment in which all three primitive elements are
well developed and defined.
THE PROTHORAX
In the prothorax the tergal, pleural and sternal areas (fig. 14) are
fused, forming a continuous chitinous tube. However, corresponding
elements to other thoracic segments are visible due to the different
kinds of sculpture. Taxonomically the prothorax bears generic char-
acters but none of specific importance. The two openings of the tube-
like prothorax, the anterior and posterior foramina, are bordered by a
fringe of closely placed hairs which arise from the inner margin.
Pronotum.—The pronotum (figs. 1, 2, 3 and 14) is one solid plate
by which the head is concealed. The dimensions are as follows:
Gn. materiarius Fitch., length, 1.23 mm., width, 0.98 mm.
Gn. retusus Lec., length, 1.44 mm., width, 1.15 mm.
Gn. sulcatus Lec, length, 1.35 mm., width, 1.07 mm.
The measurements are the average of 10 specimens from each
species.
Seen from above, the sides of the pronotum are subparallel on the
posterior margin, while the anterior margin is broadly rounded. The
anterior margin bears 10 to 16 low serrations which are only slightly
longer toward the median line and sometimes fused at their base.
On the anterior area, that is, the area in front of the summit which
consists of a short slightly curved ridge and is placed a short distance
before the middle of the pronotum, occur many comparatively small
and low serrations arranged in concentric rows and decreasing in size
towards the summit. Behind the summit the surface of the pronotum
is covered with punctures. There are two kinds of punctures; the
one comparatively coarse in contrast to the other but fine compared
to those in other nearly related genera; the other, minute and only
3
.
32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
visible under higher magnification. The coarser punctures are sparse,
deep and distinct, the minute ones closely placed giving this part of
the pronotum a more opaque appearance. The posterior margin is
slightly arcuate. Immediately anterior to the posterior margin there
is an impressed line giving the intermediate space the appearance of
a low ridge. This raised margin served as one of the main characters
in placing this genus near Pityophthorus and allied genera. The pro-
notum is covered with inconspicuous hairs which are longer and
C
Fic. 14.—Gnathotrichus sulcatus Lec.: Prothorax, ventral aspect.
a, anterior foramen; b, posterior foramen; c, anterior margin of pronotum; d, posterior
margin of pronotum; CC, coxal cavities; Epm, epimeral area; Eps, episternal area; ICP,
intercoxal process; Peps, preepisternal area; Pst, poststernal area; St, sternal area; T,
tergum or pronotum.
coarser anteriorly. Posterior to the summit occurs a shallow trans-
verse impression. The lateral limitations of the pronotum are not
clearly defined but near the postero-lateral angle is a longitudinal ridge
which may be considered as a remainder of the pleuro-notal suture.
Pleural area.—The propleural area (fig. 14) is represented as one
continuous plate in the shape of a trapezium of which the base is
formed by the remainders of the pleuro-notal suture. Ventrally the
pleural area is completely fused with the prosternum (St). Externally
no sufficiently distinct lines are present to justify the distinction of
subdivisional plates as episternum, epimeron, etc. Nevertheless there
can be distinguished three parts of different structure which one may
call the preepisternal (Peps), the episternal (Eps) and the epimeral
NO. I0 MORPHOLOGY OF GNATHOTRICHUS—SCHEDL 8)
area (Epm). The preepisternal area is flattened, slightly depressed
with a surface smooth except for a few minute wrinkles parallel to
the longer axis. The episternal area is entirely covered by continua-
tions of the serrations which occur on the anterior half of the pro-
notum. A narrow strip along the posterior margin of the propleuron,
quite distinctly limited to the surrounding plates by its transverse
wrinkles, may be called the epimeral area.
Fic. 15.—Gnathotrichus retusus Lec.: Prosternum and fore legs.
C, coxa; F, femur; ICP, intercostal process; Pst, poststernal area; St, sternal area; Ta,
tarsus; 7%, tibia.
Sternal area.—The lateral ill-defined sternal area is largely occupied
by the coxal cavities (CxC). The intercoxal process (ICP) be-
longing to the intercoxal or sternellar plate is clearly defined and
very short so that the coxae touch each other. The sternum proper
and the presternal area are again not limited by sutures but by dif-
ferences in sculpture.
Prothorax.—The distinguishing characters of the prothorax are:
I. The pronotum of Gn. materiarius Fitch is a little more slender
than that of Gn. retusus Lec. and sulcatus Lec.
2. The punctures of the posterior half of the pronotum of Gn. retusus
Lec. are somewhat coarser than in the case of the others.
34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
THE MESOTHORAX
The mesothorax is the shortest of the three thoracic segments. The
form is that of a ring with the anterior diameter smaller than the
posterior one. The meso- and metathorax are very closely connected,
giving the appearance of one single unit. Seen from above (dorsal)
the scutellum only is visible, while in the lateral and ventral aspect
the mesopleura and the mesosterna can be distinguished. The two
Fic. 16.—Gnathotrichus retusus Lec.: Mesothorax, view from anterior foramen.
The parts are disconnected by stretching under the coverglass.
A, external aspect; B, internal aspect; Co, coxa; El, elytra; Ep, epimeron; Pr, preepi-
sternum; Psc, prescutum; Eps, episternum; Pst, presternum; Pstr, poststernellum; Sct,
scutellum; St, sternum; St/, sternellum; Stu, scutum; Tr, trochanter; a, articulation of
elytra; b, clavicola; c, pleural hook of scutum; d, pleural clavicola.
pairs of appendages are implanted between the pleura and tergum
and the pleura and sternum respectively.
Mesonotum.—F rom the mesonotum or tergum only the scutellum
is exposed dorsally. The rest of the notum is covered by the elytra
laterally and is hidden by the pronotum anteriorly.
The scutellum.—tThe scutellum (fig. 16, Sct) is a triangular plate,
heavily chitinized, covered with short bristles, and externo-anteriorly
fused with the prescutum. There occur slight differences in shape in
the different species but they are of no importance,
NO. 10 MORPHOLOGY OF GNATHOTRICHUS—SCHEDL 35
Scutum.—The scutum (fig. 16, Stu) is represented as two slightly
chitinized lobes which are anteriorly fused with the prescutum. The
posterior ends are free and lie under the scutellum. A long slender
pleural hook (c) originates near the extreme lateral angle.
Prescutum—tThe largest area of the mesonotum is occupied by
the triangular prescutum (fig. 16, Psc). Externally it forms with
the scutellum a smooth continuous plate. The inner side consists of
two large and shallow excavations which are surrounded by heavily
chitinized ridges. The median ridge is anteriorly divided by two
closely placed sutures. Obscure remainders of these sutures are also
visible externally and on the scutellum. The extreme anterior corners
of the prescutum are produced into two prominent hooks which may
correspond to Hopkins’ “ Lateral arm of prephragma and prescutum ”
or Korschelt’s “ Zapenfortsatz des Mesonotums,” or Berlese’s “ clavic-
ola.” The clavicola (b) is hinged in parts of the pleural claviculas.
At a short distance behind the clavicola is a well developed prealar
process which embraces the third axillary of the elytra. On the
anterior two-thirds of the prescutum, numerous small punctures are
visible externally and near the lateral margin occurs a single row of
minute hairs.
The prephragma and the postscutellum are not represented by sepa-
rate plates. Remainders of the former are probably the wide anterior
ridge of the prescutum.
Mesopleura—The mesopleura are distinctly defined from the ter-
gum and sternum. The largest area of each is occupied by the epi-
sternum (Eps) which is a strongly chitinized plate, elongate, with the
lateral dorsal area smooth and with a fine pubescence on the lateral
ventral area. A narrow strip in front of the episternum, which is
defined externally by a suture, and a continuation of it towards the
ventral posterior angle of the episternum, may represent the preepi-
sternum. Dorsally the preepisternum is produced into the clavicolar-
disk. Under the ventral half of the episternum projects a narrow
plate, the epimeron (Ep). Preepisternum and epimeron are struc-
tureless but the latter bears on its posterior margin a row of fine hairs.
Mesosternum: Presternum.—A narrow ridgelike plate which is
fused with the preepisternum (fig. 16, Pr) of the pleura represents
the presternum (Pst). The rest of the sternum is externally defined
from this by a suture. Internally the suture is obsolate.
Sternum.—The sternum (St) is a rectangular plate with its poste-
rior side produced into an angle, externally fused with the sternellar
area or intercoxal process (Stl) but internally defined by a suture.
The outer surface is smooth and bears a row of bristles and also a
few punctures.
30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
Sternellar area—vThe sternellar area is strongly produced poste-
riorly and internally defined by a suture from the postero-sternellar
piece.
Poststernellar area.—This piece (Pstr) is a narrow continuation
from the sternellar area. Externally both are covered with long
bristles.
THE METATHORAX
Metatergum.—The general appearance and structure is illustrated
in figures I, 2 and 3, which also show the two main parts, namely
the notum and the postnotum, or pseudonotum, as the postnotum is
often called. The latter is well developed and connected with the
notum by a transparent membrane (a). The dorsal aspect of the
metatergum is shown in figure 17; the inner in figure 19.
Metanotum.—The metanotum (figs. 17, 19) is typical in Gnatho-
trichus in so far as the prescutum (Psc) is only loosely connected
with the scutum (Sct). The connection consists medially of a trans-
parent membrane (b); laterally the extreme ends of the posterior
prealar process (d) are fused with the lateral margin of the scutular
lobes. In this connection it should be mentioned that Hopkins’ pre-
scutal lobe seems more likely to be a part of the scutum than of the
prescutum. Other modifications are the overlapping of the scutum
beyond the posterior prealar process and the development of the
scutellum.
The metanotum is clearly defined into three transverse divisions
which are the prescutum, the scutum and the scutellum.
Prescutum.—tThe prescutum (Psc) forms a well developed trans-
verse band extending from pleuron to pleuron. Medially it is bent
downwards, forming a ventral reflected lobe which may correspond
to the prephragma in other insects. A precosta is not present as a
plate defined by lines or sculpture. On the level of the interior origin
of the anterior apodeme a suture extends interiorly separating the
posterior prealar process from the prescutum proper. This process
extends laterally, is covered by the scutular lobe and is fused with
it at the extreme end. From the antero-lateral corners of the prescu-
tum proper originate two prominent hooks, the anterior prealar proc-
esses (f). Laterally to the anterior prealar process are two disklike
formations on each side (g, h) connected with the posterior prealar
process. According to Hopkins, the posterior disk (prescutal disk)
serves for the small muscles connecting it with the pleural clavicula.
A rather intensive investigation would be necessary to determine the
purpose of all these formations and homologize them with similar
equivalent parts in other groups.
37
OF GNATHOTRICHUS—SCHEDL
MORPHOLOGY
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38 SMITHSONIAN MISCELLANEOUS COLLECTIONS volt. 82
Scutum.—The largest plate of the metanotum is the scutum ex-
tending backward from the anterior membrane (b). The limitation
between the scutum and the scutellum, the scuto-scutellar suture, is
visible as a ventrally elevated ridge which becomes indistinct laterally.
The anterior apodeme divides the scutum in two subdivisions, the
scutum proper and the scutular lobes. The latter extend beyond the
posterior prealar process of the prescutum as was stated above and
cover the process dorsally. Externally the scutum is one continuous
half spherical plate medially divided by the scutular groove. In-
ternally projections of the scutellum extend far into the scutum. From
the inner externo-lateral corners of the scutum proper arise two pairs
of parapsidal ridges (k) which converge in a slightly curved line
medially.
Scutellum.—tThe scutellum, which is posteriorly limited by a mem-
brane (a), encloses the scutum in a half circle. The scuto-scutellar
suture is produced anteriorly forming the lateral limitations of the
scutellar groove. It extends to the anterior margin of the scutum.
The space between the produced suture is strongly depressed ex-
ternally forming a wide scutellar groove. The posterior end of the
mesoscutellum rests in this groove. From the posterior lateral corners
of the scutellum arise two armlike formations which converge an-
teriorly meeting at about the middle of the scutum. These forma-
tions correspond to the endodorsum (Amans), V-shaped ridge (Snod-
grass), or median apodeme (Hopkins). Where they join they are
connected with the scutellar groove. It is likely that the intensive
study of the endodorsum of the Scolytidae will bring out many new
features for the systematic arrangement of this group.
Postnotum.—The postnotum or pseudonotum is subdivided into
three transverse divisions, the precosta (Pc), with the prominent
prealar bridges (1) and the postphragma (Pph). The precosta is a
simple band separated from the postphragma by the postnotal apodeme.
Laterally it is produced into prominent prealar bridges. The inner
margin is developed as a heavily chitinized ridge which is posteriorly
divided, forming a ring and externally produced into a slender hook,
the prealar hook (m).
The postphragma is a slightly ventrally bended band. The post-
notum is visible externally, giving the appearance of a reduced tergum
at the first abdominal segment.
Metapleura—The pleuron of the metathorax (fig. 17) consists
chiefly of two plates, the episternum (fig. 17, Eps) and the epimeron
(Epm). The pleural suture (Ps) is externally represented as a dis-
tinct line, internally as a ridgelike structure. From the pleural suture
NO. 10 MORPHOLOGY OF GNATHOTRICHUS—SCHEDL 39
branch three other sutures or ridges. Anteriorly there is a short
suture dividing the parapterum from the wing process; posterio-
dorsally another ridge separates the postepimeron (Pepm) from the
rest of the epimeron. Ventrally another ridge indicates that part
of the episternum which is covered by the sternum and the metacoxa.
Episternum.—The episternum is an externally continuous plate
having the shape of a narrow triangle with the base facing forward.
Ventro-anteriorly it is produced into a hooklike angle, the sternal
hook (c,) or anterior sternal hook of Hopkins. The sternal hook
fits into an emargination of the sternum, the clavicula (fig. 18, d).
Dorsally the extreme angle of the episternum with the pleural suture
(probably also containing elements of the epimeron) is produced
into the parapterum (e), or coracoid process, and the wing process,
or clavicular process (d). From the underside of the parapterum
originates a well-developed pronator, or muscle disk (fig. 17, t:). The
episternum is heavily chitinized and covered by numerous large punc-
tures which are interlaced by a network of fine ridges. See also fig-
ure 17, h,. When the elytra are kept in the closed position the meta-
pleura are nearly completely hidden by them.
Epimeron.—Between the pleural suture and the lateral margin of
the metanotum a more flexible, partly membranous sclerite is in-
serted. Posteriorly it is subdivided by a branch of the pleural suture
separating the postepimeron from the epimeron proper. While the
epimeron proper is more membranous and flexible, the postepimeron
is more heavily chitinized. The postepimeron bears dorso-posteriorly
a recurved hook, the postepimeral hook (j,), which articulates with
the clavicula of the sternum of the first abdominal segments. The
hypopleurite (Hpp) extends forward up to a shallow emargination on
the dorsal margin of the postepimeron. The anterior margin of the
hypopleurite is double reflexed, fitting in a reflexed ridge of the inner
layer of the elytra. The area between the anterior margin of the
hypopleurite and the dorsal nob of the pleural suture is deeply im-
pressed. Epipleurite 1 of the abdomen is situated in front of the
hypopleurite closely attached to the scutellum of the postnotum.
Metasternum.—tThe metasternum is represented as a rectangular
continuous plate divided by remainders of a median line. It is illus-
trated in figure 18. The anterior margin is medially produced into
an angle which extends far between the mesocoxae. This angle and
the thickened anterior margin may correspond to the presternum of
other segments. The anterior lateral angles (a) are modified to re-
ceive the sternal hooks of the metaepisternae. Therefore these emar-
ginations should be called the anterior claviculae of the metasternum.
Posteriorly the sternum is infolded producing a narrow plate, the
40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
sternellar area. Normally the sternellar area is hidden by the coxae
except for the two prolongations between the coxae. These are bent
dorsally, giving the support for and articulation of the furca. The
Fic. 18—Gnathotrichus materiarius Fitch: Metasternum, inner aspect.
C, coxa; St, sternum; Ji, tibia; Tr, trochanter; a, sternellar area; b, median line; c, pre-
sternellar area; d, clavicula; e, sternal hook of metapleuron; t, head of postepimeron.
Fic. 19.—Gnathotrichus materiarius Fitch: Metatergum, inner aspect.
APh, prephragma; Pc, precosta of postnotum; PN, postnotum; Pph, postphragma; Psc,
prescutum; Sct, scutum; Sc/, scutellum; a, posterior membrane; b, anterior membrane; c,
scutellar lobe; d, posterior prealar process; e, anterior apodeme; f, anterior prealar process;
g, anterior prescutellar disk; h, posterior prescutellar disk; 7, scuto-scutellar suture; 7, median
apodeme or endodorsum; k, parapsides; /, prealar bridge of postnotum, or postscutellum; m,
prealar hook of postnotum; 7, postnotal apodeme; o, scapular hook.
sculpture of the metasternum is minutely rectangulate. The punctures
which occur are sparse, the hairs of medium length and more numer-
ous laterally.
The metathorax shows little specific and no sexual modifications.
NO. LO MORPHOLOGY OF GNATHOTRICHUS—SCHEDL AI
THE ABDOMEN
The chitinous skeleton of the abdomen shows little specific modifica-
tion but it bears characters which are of importance in separating
the higher groups. The differentiation is mainly in the number of
dorsal plates or tergites and the development of the spiculum ventrale
in the females. The structure and the relative proportions of the dif-
ferent sclerites are illustrated in figures 1, 2, 3, 20, 21, 22, 23, and 24.
Dorsal plates or tergites——All the tergites are normally covered by
the elytra. In the females seven and in the males eight tergites are
well developed. The first six tergites are more or less membranous
Fic. 20.—Gunathotrichus materiarius Fitch, male: Abdominal tergites, ventral
aspect.
I, 2, 3, 4, enlarged sections showing details of structure on the dorsal side; Epp, epipleurites;
Epp 7 and 8 fused with tergites; Sp, spiracles; T, tergites.
and flexible in both sexes, yellowish in color, and covered with spinous
to toothlike armature on the external surface. The spinelike armature
is more medial and posterior on the plates. Near the anterior margin
the armature becomes more toothlike, forming broad plates armed
with minute spines posteriorly (figs. 20, Ti, 2,3,4). These plates near
the median line on the second, third and fourth tergites are stouter
and not so densely placed. In the same tergites there is a reduction
of the chitinization, forming a membranous emargination. Still
another type of armature is found on the pleurites as illustrated in
figures 20, 4. The purpose of these armatures is doubtful. The in-
tersegmental membranes are colorless without any markings. Heavily
chitinized bands are situated near the posterior margin of plates two
42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, 82
to six. These bands or ridges resemble the parapsides of the meta-
thorax both in structure and in position. Punctures are numerous in
all the plates and the lateral limitations are always irregular. The
seventh and eighth tergites in the males and the seventh in the females
are heavily chitinized and lack the above mentioned armature but
have numerous hairs and punctures.
Lateral plates or pleurites——vThe pleural suture seems to be the
line which divides the heavily chitinized hypopleurites from the mem-
branous epipleurites. The pleural suture is not visible in the seventh
and eighth pleurites because the epipleurite is heavily chitinized here
and completely fused with the tergite. The second hypopleurite which
Fic. 21.—Gnathotrichus materiarius Fitch: Abdominal sternites, ventral aspect.
St, sternites; a, coxal cavity; b, clavicula; c, intercoxal process.
extends beyond the postepimerum of the metathorax is ventrally de-
fined by a suture while the others are fused ventrally with the sternites.
The former corresponds to the second tergite. The hypopleurite be-
longing to the first tergite is not represented as a plate but fused with
the epipleurite. The hypopleuritic areas are covered by the elytra when
they are kept in the closed position, and form a vertical plane, while the
sternites make an angle of about 120 degrees with them. As was men-
tioned before, the epipleurites are membranous except those which cor-
respond to the last tergites. The hypopleurites corresponding to the
last seventh and eighth tergites are present only as narrow membranes.
Ventral plates or sternites—tIn both sexes only five ventral plates
or sternites are distinctly defined by sutures as is illustrated in figure
21 and they represent the sternites three to seven. The sternites are
NO: TO MORPHOLOGY OF GNATHOTRICHUS—SCHEDL 43
widest at the base, decreasing in width towards the apex. The first
visible sternite is the longest and is separated from the second by a
suture which is only visible under high magnification. It is very
likely that the first visible sternite contains elements of more than
one abdominal segment but they are not indicated by sutures or lines.
Medially the first sternite is produced into a long process.called the
intercoxal process, which extends far between the metacoxae. From
Fic. 22.—Gnathotrichus retusus Lec.: Seventh tergite of female.
A, dorsal aspect; B, ventral aspect; Spz, seventh spiracle.
the base of this process a ridge arises which extends laterally, sepa-
rating the part of the sternite covered by the metacoxae from the
remainder. At the antero-lateral corners this ridge is developed into
a clavicula. In this clavicula (b) articulates the ventral part of the
postepimeral hook. The surface of the uncovered part of the first
sternite bears numerous hairs which are arranged in concentric rows
encircling the metacoxae. Punctures are sparse. All the other ster-
nites are separated from each other by deep septae and they have the
shape of short rectangular plates of nearly equal length. They are
all heavily chitinized and armed with long hairs arranged in a trans-
44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
verse row. The last sternite is infolded on the apex. There occur no
striking differences in either different species or in the two sexes.
Spiculum ventrale —In the females of all three species the spiculum
ventrale is well developed. By careful dissection it can be seen
fastened to the Fortsatzlappen Verhoeff. Without any doubt we
are dealing here with a true spiculum laterale clausum (Fuchs). The
median line commissura, which gives by certain modifications the
spiculum ventrale opportum, is especially well defined in Gn. materi-
Fic. 23, A.—Gnathotrichus retusus Lec.: Sixth and seventh sternites of a female
with the spiculum ventrale.
Ca, caput; Coj, conjunctus lateralis; Com, commissura; Br, brachium; Ra, radix; st, sternites,
Fic. 23, B.—Gnathotrichus retusus Lec.: Seventh and eighth sternites of a male.
arius Fitch and retusus Lec. For further studies on this subject it is
proposed to name the different parts of the spiculum. The part
where the spiculum joins the seitlichen Ansatzlappen (Verhoeff),
seitlichen Lappen (luchs), may be called radix spiculorum ; the more
or less chitinized, sometimes membranous bands extending anteriorly,
brachium ; and the apically thickened part, caput. The seitlichen Lap-
pen (Fuchs), Fortsatzlappen (Verhoeff) should be called the con-
junctus lateralis. The spiculum ventralis of Gn. sulcatus Lec. may be
easily distinguished from the others by its slimmer brachiae. Gn.
retusus Lec. and Gn. materiarius Fitch show no striking differences
in this regard.
NO. 10 MORPHOLOGY OF GNATHOTRICHUS—SCHEDL 45
THE SPIRACLES
Corresponding to the number of the epipleurites, there are eight
spiracles in the male and seven in the female. These are implanted
in the membranous epipleurites and the apical epipleuro-tergite re-
spectively. The eighth pair of spiracles in the male are always rudi-
mentary. Gn. materiarius Fitch shows comparatively the best de-
velopment in this regard. The seventh epipleurotergite of the female
bears a pair of spiracles which are well developed and only very
slightly smaller than the others. There also occurs one pair of spiracles
on the prothorax. They lie under the produced caudad-lateral angles
of the pronotum.
Fic. 24.—Spiculum ventrale in female: A, Gnathotrichus sulcatus Lec.,
B, Gnathotrichus materiarius Fitch. .
Br, brachium; Ca, caput; Com, commisura.
THE LEGS
The three pairs of legs are illustrated in figures 25 and 26. They
do not vary strikingly from species to species neither in form nor in
sculpture. Therefore the drawings were made from Gnathotrichus
materiarius only. The legs present all the typical segments common
in Coleoptera, which are the coxa, the trochantin, the femur, the tibia,
the tarsus and the pretarsus.
Coxva.—The coxa (fig. 25) differs considerably in shape in the three
pairs of legs. The coxa of the prothorax (A) is very stout, ball-like,
the mesocoxa (B) is slightly longer and the metacoxa (C) is nearly
twice as long as the forecoxa. The proximal end of the fore- and
mesocoxa show distinct indications of a basicostal suture (a). In the
fore-coxa the basicostal suture is formed into an external ridge
ventrally, which becomes lower and indistinct laterally. In the meso-
coxa a simple suture (a) indicates the limitation of the basicosta. The
basicostal area of the forecoxa (Bc) is about twice as long as the
same structure in the mesocoxa. The basicoxite (Bex) is present as
46 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
a marginal flange and is visible in both the fore- and the
mesocoxa. The metacoxa does not show the separation of a basicosta
by external lines or internal ridges. The extreme proximal end bears
a rather indistinct, internal marginal ridge only.
The three articular surfaces (Snodgrass) are largely modified in
the different legs. The pleural articular surface of the fore-coxa is
present as the medial, proximal margin of the basicosta only. This
part is slightly more heavily chitinized. In the mesocoxa the pleural
articular surface (b) is more strongly developed. It consists of a
Fic. 25.—Guathotrichus materiarius Fitch: A, forecoxa, lateral aspect; B, meso-
coxa, C, metacoxa, both ventral aspect.
Bc, basicosta; Bcx, basicoxite; F, femur; Fo, fossa; Tr, trochanter; a, basicostal suture;
b, pleural articular surface; c, outer ring of fossa; d, inner ring of fossa; e, anterior coxal
suture; f, internal ridge; g, basicostal ridge of trochanter; h, condyle of trochanter; 1,
trochantero-femoral suture.
projection of the basicosta. In the metacoxa a hooklike structure (b),
which is basally fused with the anterior coxal suture, may be con-
sidered as the pleural articular surface.
The articulation of the trochantin is monocondylic with the fossa
in the coxa. The fossa (Fo) is a circular, conelike, impression on
which two main parts are clearly defined; the outer ring (c) with a
ridgelike elevation on the external surface of the coxa and the inner
ring (d) or bottom which bears a heavily chitinized knob on the inner
surface. It is most probable that the outer ring corresponds to the
anterior, the inner ring to the posterior or distal articular surface of
Snodgrass. Each coxa has opposite to the fossa a circular opening in
NO. 10 MORPHOLOGY OF GNATHOTRICHUS—SCHEDL 47
the heavily chitinized wall of the coxa which is covered by a thin
membrane only. The extreme tip of the trochantinal condyle touches
this. Ridges to strengthen the coxal walls do not occur in the fore-
and mesocoxa. The metacoxa bears such a ridge on the ventral side
Fic. 26.—Gnathotrichus materiarius Fitch: A, fore leg, B, mesothoracic leg,
C, metathoracic leg.
F, femur; Ptar, pretarsus; Tar, tarsus; Ti, tibia; Tr, trochanter; a, trochantero-femoral
joint; b, basicostal ridge; c, condyle of trochanter; d, dorsal femoral fossa; e, ventral
femoral fossa; f/, anterior groove; g, tibial condyle; h, outer margin of tibia; 7, marginal
tooth; j, apical tooth; k, /, m, n, tarsal segments; 0, apical segment; gq, arolium; p, claw.
which forms a well developed internal ridge (e); this ridge may
correspond to the anterior coxal suture (Snodgrass). Another more
feebly developed ridge (f) occurs on the dorsal coxal wall.
The hair armature is best explained by the figures. No differences
have been found from species to species.
4
48 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
Trochantin.—The trochantin (fig. 26, Tr) is a small structure and
is closely attached to the femur. On the ventral or outer side, when
the femur is kept close to the body, a deep groove, the trochantero-
femoral groove (a) separates the femur from the trochantin. On
the dorsal, or inner side, the separation is indicated by a suture only.
Structurally, three parts may be distinguished on the trochantin.
These are the basicostal ridge (b), the articulatory condyle (c) and
the small apical piece (r). The basicostal ridge is a broad, stuffed,
ringlike structure which gives the articulation with the coxa an ex-
ternal rest. For the same purpose a hooklike process on the postero-
median angle of the basicosta is used. The prominent, cone-shaped
articulatory condyle originates basally. The shape of the condyle is
alike in all three pairs of legs; the slight differences showing in the
plate are due to the different angles from which the drawings were
made. The apical piece projects over the basicosta when viewed from
above, dorsally, and is fused with the basicosta when seen from below.
It bears a long slender spine in all three pairs of legs. The basicosta
is externally armed with a few small hairs. From the kind of con-
nection between the femur and the trochantin it can be concluded that
but little movement is possible between them.
Femur.—The femur (fig. 26, F) is the strongest segment of the
leg and is about equal in length with the tibia (Ti). Basally it is con-
nected with the trochantin; apically, the tibia articulates. The femur
is long, oval to rectangular in outline and strongly compressed. The
articulation of the tibia is bicondyle (d, e), the femur containing two
half circular fossa. There occur no striking differences in the three
examined species or in the three pairs of legs in one species. The
inner or dorsal surface of the femur is smooth without hairs or
bristles ; the exposed surface bears numerous rather fine, long hairs
which are directed transversely. A deep groove (f) with two lateral
winglike extensions occurs latero-apically. This groove allows the
tibia to be flexed closely against the femur.
Tibia—The tibia (fig. 26, Ti) is about as long as the femur, tri-
angular in outline and strongly compressed anteriorly. The proximal
end is widened, half circular in outline, and bears the two articulatory
condyles. The proximal quarter of the tibia is slightly bent laterally.
The dextral margin is without teeth or armations; the sinistral margin
bears four to six low serrations and three marginal teeth (1) which
are imbedded in sockets. The apical tooth (j) is straight, rather stout
and not imbedded in a socket; the subapical tooth is present as a low
elevation only. The articulation of the tarsus is membranous,
NO. I0 MORPHOLOGY OF GNATHOTRICHUS—SCHEDL 49
Tarsus.—The tarsus (fig. 26, Tar) is composed of five joints or seg-
ments. They are not articulated by hinges with each other but they
are movable by means of inflected connecting membranes. The first
three segments (k, 1, m) are subequal in length and shape. The basal
segment or basitarsus does not show any special armations such as
occur in other genera. The fourth segment (n), the smallest of the
tarsus, resembles somewhat in shape the trochantin. This segment
which is often highly modified in the Scolytidae, is short and knob-
like in Gnathotrichus. The apical segment (o) is longer than seg-
ments one to three, more slender and slightly curved. Except the
fourth, all tarsal segments bear at least a few hairs scattered over the
entire length. The third segment also bears a plushlike arrangement
of hairs apically.
Pretarsus: the terminal segment.—The terminal foot structure (fig.
26, Ptar), which has been called praetarsus, Krallenglied, unguis,
ungula and pretarsus by different authors, bears two simple claws (p).
The areolium (q) is membranous and heartshaped.
THE WINGS
As in all the Scolytidae, Gnathotrichus Eichh. has well developed
elytra or mesothoracic wings and hind or metathoracic wings. The
development of the metathoracic wings would indicate, as has been
found to be the case, that the species of this genus are good fliers. No
attempt will be made in what follows to speculate on the functions
of the different sclerites of the articulation of the wings since the
author has had no opportunity to make observations on them. Merely
a description of the different parts will be given which it is hoped may
be of value for future taxonomic and physiological studies.
MESOTHORACIC WINGS OR ELYTRA
The mesothoracic wings consist as usual of two layers of integu-
ment, both of which are chitinized, the outer layer however being
much thicker than the inner layer. The space between these two
layers contains the tracheal and circulatory systems. The punctures
which mark these wings externally indicate, according to Hopkins,
the points of union between the two layers. The form and general
structure are shown in figures 1, 3, 16 and 27. The elytra when closed
and viewed from above are as wide as or slightly narrower than the
pronotum, with the sides subparallel, slightly tapering toward the apex
and broadly rounded behind, the extreme external margin subacute.
50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
Basal area.—The basal area (fig. 29), as here interpreted, is the
basal part of the elytra itself and the sclerites which form the con-
nection with mesonotum and the mesopleura respectively. The articu-
latory elements of the elytra itself are made up chiefly of the pro-
jected costa (Co) and subcostal veins (Sco) and the costal (a) and
the subcostal heads (b). These heads articulate with the mesopleural
clavicola (fig. 16, d). There are also four distinct axillaries (fig.
29, aX; to ax,) or pteraliae. The first axillary (ax,) partly encloses
Fic. 27.—Gnathotrichus retusus Lec.: Left elytron, ventral aspect.
An, anal; Cu, cubitus; Me, media; Ra, radius; Sc, subcosta; W1/, lateral wing lock; a,
dorsal or sutural margin; b, costal margin; c, lateral convexity; d, stridulating scraper;
é, apex.
the tegula and fuses at its apex with the second axillary. The second
axillary (ax) articulates at its base with the prealar process (a) of
the prescutum. The apex of the second axillary forms a heavily
chitinized clamp in which the elytra fits as a tongue. The third
axillary (ax;) corresponds to Hopkins’ flexor plate. It commences on
the internal surface of the second axillary and ends on the external
surface of the elytra. It is believed that a sclerite (ax,) projecting
from the posterior margin of the tegula is the fourth axillary. The
tegula (Te) is well developed, having the form of a hairy pad.
—
NO. IO MORPHOLOGY OF GNATHOTRICHUS—SCHEDL SI
Tracheation—All six primary veins (fig. 27) are present and
these run roughly parallel and equidistant from each other the whole
length of the wing. The costal vein is fused with the corrugated and
thickened anterior margin (b) forming an anterior ligature of the
wing. The subcostal (Sc), medial (Me), cubital (Cu) and anal veins
(An) are simple ; the radius is split into two branches (Ra,, Ra.). The
finer divisions of the tracheae permeate the spaces between the veins
producing a fine network. Between the adjacent veins there are situ-
ated two fairly well defined rows of punctures.
Sculpture and vestiture —The occurring punctures are as previously
mentioned arranged in approximate rows, being more confused
A i
Fic. 28.—Gnathotrichus materiarius Fitch: Mandibles of the larva.
a, apical tooth; b, subapical tooth; c, median tooth; d, molar tooth; e, lateral bristle, or
seta (seta mandibulae lateralis); f;~», dorsal setae, or bristles (seta mandibulae lateralis) ;
g, retractor tendon; gi, extensor tendon; /:, retractor disk; 7, extensor disk.
laterally and on the declivity. The interspaces are minutely recticu-
late and punctulate and in all three species about alike. From the
punctures originate minute hairs which are slightly longer on the
declivity. There also occur longer bristles but they are scarce and
scattered over the entire surface, being more numerous on the declivity.
Lateral wing lock—On the inner lateral side of the elytra at the
level of the anterior margin of the hypopleura a short transverse ridge
(fig. 27, W1) is situated. That ridge is recurved, fitting into a similar
formation which is formed by the anterior margin of the hypopleura
(fig. 1, f, fig. 17, i,). These two parts interlock and are apparently
intended to keep the elytra closed.
Declivity—The extreme lateral margin of the elytra is subacute.
The declivity itself is sloping with a more or less distinct sulcus on
52 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
each side of the median suture. The lateral convexities (fig. 27, c)
bear at least faint traces of granules from which bristles arise. There
also occur a few bristles near the apex of the declivity.
Stridulating accessories—In both sexes the left elytron bears on the
declivity a well developed lobe which lies under the right elytron
Deere
Prescutum
\
ie sae
Fig, 29.—Gnathotrichus retusus Lec.: A, articulation of the elytra, ventral as-
pect; B, stridulating apparatus on the apex of the elytra, ventral aspect.
Cal, callus; Co, costal ridge; Sco, subcostal vein; Te, tegula; a, costal head; b, subcostal head,
ax1, @%2, aX, axs, the axillaries; c, lateral arm of prephragma Hopk., or pleural hook of
scutum; d, prealar process; e, stridulating rasp; f, stridulating scraper.
when the elytra are kept in the closed position. This lobe, the stridu-
latory scraper (fig. 27 d, fig. 29 f) is finely transversely sulcate on
the dorsal side. The corresponding area on the right elytron (fig. 29, c)
is also rasplike but only on a narrow strip. If these two parts are
grated against each other, it is most probable that they will produce
a chirping sound. Up to date this sound has not been heard by the
NO. IO MORPHOLOGY OF GNATHOTRICHUS—SCHEDL 53
author and therefore this statement is merely an attempt to explain
these two parts of the elytra.
Specific modifications of the elytra——While Gnathotrichus shows
the strongest development of the declival sulci, with lateral convexities
and granules on the latter, these characters decrease in development
in Gn. sulcatus Lec. and are faint in Gn. materiarius Fitch. There also
octur many variations from specimen to specimen so a determination
cannot be based on these characters only.
METATHORACIC WINGS OR HIND WINGS
The metathoracic wings (figs. 3, 17) are homologous with the meso-
thoracic wings but differ considerably in structure and development
due to their use as flying apparatus. In the genus Gnathotrichus
Eichh. no specific modifications of importance have been observed.
As in all genera of this family the wings are folded together and
covered by the elytra when the beetles are at rest. The hind wings
are twice as long as the elytra and three times as long as wide. Near
the base on the inner side occurs a deep emargination separating a
distinct lobe from the wing—the posterior wing lobe (fig. 3, WL,
17, LL). The shape of the wings is very much alike in all three species,
long oval with the anterior and posterior margins nearly parallel.
The wings consist of two layers of integument enclosing the tracheal
system. In the hind wings both of these layers are membranous ex-
cept on the veins and the basal sclerites. Externally they are covered
with microscopic hairs, except on the basal heavy parts of the veins
and the axillaries (fig. 17).
V eins—The venation of the wings shows considerable reduction.
The only visible veins are the costa (fig. 3, Co), subcosta (Sco), radius
(Ri+R.2), media (Mi, Mz) and cubitus (Cu,). The radial and the
medial veins are split into two branches.
Costa——As Hopkins has shown for the genus Dendroctonus, the
costal vein (fig. 3, Co, 17, C) is also in Gnathotrichus Eichh. confined
to a short basal piece. Apically it does not join another vein but is
reduced so that the costal margin is occupied by a membrane only up
to the point where the radial vein is bent forward and structurally
replaces the costa. The base of the costa is produced into the so-called
costal head (fig. 30, CoH). It consists of two parts, the costal condyle
(fig. 30, a), and the costal pocket (b). The former articulates with
the clavicle process of the metaepisternum (fig. 17, d,). From the
base of the costal condyle a pocket stretches to the subcosta. At about
the center of the pocket a projection of the subcostal head, the sub-
54 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
costal hinge (fig. 30, c), is attached by tendons giving the motion of
a hinge joint. This attachment enables the subcostal head to move in
the pocket.
Subcosta—The subcosta (figs. 3, Sco, 17, Sc, 30, Sco) consists of
a chitinous band extending apically as far as the costa and strengthened
by two ridges of dense chitin. The basal part, or subcostal head (fig.
30, ScoH), as it was called by Hopkins, is anteriorly formed into a
S00 Go
Fic. 30.—Gnathotrichus retusus Lec.: Basal area of the wing, wing articulation.
Co, costal vein; CoH, costal head; Cu, cubital vein; Me, medial vein; Sco, subcostal
vein; ScoH, subcostal head; az, axillary plates; a, costal condyle; 6, costal pocket; c,
subcostal hinge; d, subcostal labrum; e, condyle of first axillary; f, subcostal fossa; 9,
membranous fold connecting Me and ars; h, membranous fold connecting Me and Cu; 1,
membranous fold connecting Cu and azz; j, epicondyle of ax; k, scapula arm; /, axillary
horn; m, axilla; n, anterior process of azz; 0, articulatory margin; 9, scapular hook of
prescutum; q, apical margin; r¢, lateral groove; s, axillary tendon; t, uw, lateral emarginations;
v, posterior part, or handle; w, anterior part, or blade.
projection which is composed of a circular band of chitin (fig. 30, c),
articulating with the costal pocket. The posterior part forms a fossa,
the subcostal fossa (f), which articulates with the condyle (e) of
the first axillary. The labrum (d) of the fossa is deeper on the outer
side than on the inner. The outer side of the labrum is divided at
its free edge by a V shaped groove.
Radius.—The radius (figs. 3, R, Ri, Re, 17 R) is not directly fused
with any of the axillary plates but is basally closely connected with
the subcosta. From the point of fusion with the subcosta the radius
NO. 10 MORPHOLOGY OF GNATHOTRICHUS—SCHEDL 55
extends to the folding hinge on the wing gradually increasing in
width. The anterior border of the radius becomes the anterior border
of the wing just after the reduction of the costa and subcosta. The
folding hinge occurs as a V-shaped plate, the point of the V being
anterior. From the folding hinge the radius is divided into two
branches (R, and R,). Radius 1 extends as a broad chitinous band
along the anterior border of the wing to its apex gradually diminishing
in width. Radius 2 is slightly narrower than R,, running two-thirds
with, and parallel to it.
Media—The media (figs. 3, M, 17, M and 30, Me) is connected
to axillary four (fig. 30, ax,) by a membranous fold (g) of L shape.
Another fold (h) runs posteriorly to the cubitus (Cu). From the
base to the level of the folding hinge it is continued as a single vein.
At the latter point it is divided into two branches (fig. 3, Mi and M2),
both of which extend to the anal margin. The connection between the
single basal part of the media and M, is membranous, while M, is
a direct continuation of the former.
Cubitus—The cubitus (figs. 3, Cu, 17, Cu and 30, Cu) is con-
nected with axillary three (fig. 30, ax,;) by a membranous fold (1)
which runs from the base of the cubitus anteriorly. From its base
the cubitus proceeds toward the anal margin which it does not reach.
No other branches of the cubitus nor an anal vein are present.
Wing articulation—The articulation of the wings (figs. 3, 17 and
30) is brought about by means of the costal and subcostal heads, the
axillaries of the wings, the clavicle and coracoid process of the meta-
episternum, the scapular hook of the posterior prealar process of the
prescutum and a series of muscles and tendons connecting and moving
these parts. The connection of the costa and subcosta to each other
and the costa with the metapleuron was discussed before. In the
following lines the axillaries and their connection with the meta-
tergum and metapleuron will be explained.
The axillaries are chitinous plates, differing in number in the dif-
ferent orders and also it seems in the genera of the Scolytoidea, which
function as articulatory accessories. In the genus Gnathotrichus
Eichh. four of such plates are distinctly developed. The heads of the
costal and subcostal veins are here not counted as axillaries because
they are fused with the veins in such a way that separations seem
unnecessary.
First axillary —The first axillary (fig. 30, ax,) or scapular plate,
as it was called by Hopkins, is very similar in shape to that of Den-
droctonus valens Lec. as it was illustrated by Hopkins. Anteriorly it
is produced to a condyle (e) with a distinct epicondyle (j), articulat-
56 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
ing with the fossae (d) of the subcosta. The slender part posterior
to the condyle, which was called the scapular arm by Hopkins (k),
bears a hornlike process (1), the axillary horn. The horn forms with
the scapular arm an axilla (m) in which the anterior process (n) of
the second axillary rests. The margin towards the notum or articu-
latory margin (Hopkins) (0) is connected with the posterior prealar
process of the prescutum by the scapular hook (p) and tendons. The
scapular hook is a heavily chitinized hooklike plate on the lateral
margin of the prealar process. The margin towards the apex (q) of
the wing fits into the lateral groove (r) of the second axillary. The
base of the scapular plate is deeply emarginated.
Second axillary—vThe second axillary (fig. 30, ax.) or subscapular
plate has the shape of an equilateral triangle with its base anteriorly.
The side towards the first axillary bears a deep groove, the lateral
groove (r), which encloses the apical margin of the first axillary (q).
With axillary four it is connected by a membrane only while a well
developed tendon (s) connects axillaries two and three on its pos-
terior end. By means of this tendon a strong connection is brought
about from the posterior prealar process to the first, second and third
axillaries.
Third axillary—rThe third axillary (fig. 30, ax3) or flexor plate
(Hopkins) has the shape of a sickle with two emarginations (t, u) on
its inner side. The anterior one (t) ends in the tendon connecting
axillaries two and three. The handle of the sicklelike plate (v) is
without special characters. The blade (w) shows on its posterior
margin near the apex a membranous fold (1) connecting this plate with
the cubital vein.
Fourth axillary—The median plate (Hopkins) corresponds to the
fourth axillary (fig. 30, ax,). It is triangular in shape, connected
to the median vein by a membranous fold (g) and to the other axil-
laries by membranes only.
When the radial plate (Hopkins) is not considered as a distinct
plate but as a connecting tendon between the second axillary and
the radial vein only, the shape and structure of the other axillaries
are very much alike in the widely separated genera Dendroctonus Er.
and Gnathotrichus Eichh. Further studies will show if this means
a parallel modification or if we have to deal with a character common
to the superfamily of the Scolytoidea.
Lateral impression—An area distinctly impressed on the meta-
scutum (fig. 17, q), according to Hopkins, accommodates the flexor
plate at rest when the wings are closed.
NO. I0 MORPHOLOGY OF GNATHOTRICHUS—SCHEDL 57
Lateral emargination.—The lateral emargination (Hopkins) (fig.
17, p) is an emargination on the scutum on the lateral margin of the
scutellar lobe in which is implanted the inner posterior lobe of the
scapular plate and the scapular hook. The latter connects the pos-
terior prealar process with the scapular plate and the scutellar lobe,
respectively.
Tue Mare REPRODUCTIVE ORGANS
The male reproductive organs consist, as shown by Nuesslin for
this family, of elements of endodermal and of ectodermal origin.
These two groups of elements are separated in the larvae and be-
come connected during the pupal stage. Of endodermal origin are
the testi (fig. 31, Te), the vasa deferentia (fig. 31, Vd) and the
Fic. 31.—Gnathotrichus materiarius Fitch: Male reproductive organs.
Dej, ductus ejaculatorius; Gl. acc, accessory glands; Pen, penis; Te, testi; Vd-d, vasa
deferentia, duplex; Vd-s, vasa deferentia, simplex.
mesadenic, or mucous glands (fig. 31, Gl. acc). On the other hand,
the seminal vesicle (in part, fig. 33, Rec.), the ductus ejaculatorius
(fig. 31, Dej) and the penis (fig. 3, Pen) are of ectodermal origin. The
point of connection of these elements is plainly visible in the adults.
The distal part of the vasa deferentia and the mucous glands join a
short, usually narrow tube, ‘“ Zunge’’ (Nuesslin) ; the latter connects
with the ductus ejaculatorius. Externally the Zunge, an endodermal
structure, is encircled by a wider tubelike or sphericle envelope, which
consists of two parts, the distal end or “ Mantel’? (Nuesslin) of en-
dodermal, and the basal part or “ Becher” (Nuesslin) of ectodermal
origin. These last mentioned structures are seen commonly as a ball-
like widening of the ductus ejaculatorius and are usually called the
seminal vesicle.
The following description is based on -Gnathotrichus materiarius
Fitch only.
58 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
The testes consist of two oval structures which are closely connected
medially. The vasa deferentia is Y shaped. The mucous glands are
wide and stout and about as long as the testi. The seminal vesicle is
represented by a subsphericle structure to the upper side of which the
glands are joined. The ductus ejaculatorius is comparatively short,
stout, and as long as the testi. In Nuesslin’s key to the larger groups
of Scolytidae, based on the male reproductive organs except the penis,
Gnathotrichus falls near to the Ernoporinae.
Penis.—In order to consider the penis of Gnathotrichus, it is first
necessary to discuss in a general way the Scolytid penis.
Lindeman, who was the first investigator of the S colytid penis, dis-
tinguished two main groups of elements, the primary and the acces-
sory. He states that the primary elements, which comprise the body
(Koerper), the fork (Gabel) and the stalk (Stengel), are constant
throughout the family Scolytidae, while the accessory elements, which
form together the so-called Aufsatz, vary considerably in the dif-
ferent genera as well as from species to species in one genus. Ver-
hoeff, the second to deal with the subject, did not agree with this
classification, while Nuesslin supported Lindeman’s opinion. The
last of the more important investigators of the Scolytid penis, Dr.
Fuchs, constructed a new system for such a classification, without
consideration of the phylogeny, and distinguished covers (Huellen),
inclosed parts and parts external to the covers. The author cannot
agree with Lindeman and Nuesslin that the primary elements always
included in the Scolytid penis consist of the fork (parameren Ver-
hoeff) and the stalk (spiculum ventrale (Fuchs)). Already Fuchs
has shown how far the reduction of the parameren and the spiculum
ventrale in the European Hylesinidae has gone and the author is con-
vinced that more intensive investigations of this subject will bring
up many new facts. Without doubt more than three-quarters of the
genera and ninety per cent of the species of the Scolytidae have not
been studied at all or not thoroughly enough. For example, it was
found in the present study that the spiculum ventrale is absent in
Gnathotrichus, and it will not be long until genera are found in which
the complete loss of the parameren occurs. If it is desirable to dis-
tinguish between primary and accessory elements, the author would
prefer that the primary elements be considered the body only, the
accessory elements all the parts outside of it. For further studies of
this subject, the author will adopt Dr. Fuchs’ classification and nomen-
clature until a time when sufficient new material is available to fur-
nish new conclusions.
NO. 10 MORPHOLOGY OF GNATHOTRICHUS—SCHEDL 59
The penis presents, in the genus Gnathotrichus, excellent generic as
well as specific characters. The following discussion is based on
slides which were made from dried material treated with 10% sodium-
hydroxide solution. Dr. Fuchs distinguished two layers of covers, the
outer and the inner. These two layers were also found to be present
in Gnathotrichus. The outer layer consists of a membranous tube
Fic. 32—A, Gnathotrichus sulcatus Lec.; B, Gnathotrichus retusus Lec.; C, D,
Gnathotrichus materiarius Fitch.: A, B, C, lateral aspect; D, dorsal aspect.
Ca, caput; Fep, femora penis; Lad, laminae dorsales; Lav, laminae ventrales; Me, metula;
Pal, pallidium; Pa, parameren; Prp, preputial sac; Ro, rostrum; a, ventral nob of the
parameren; b, tegmen furcae; c, internal ridge on Lad.
in which the inner cover slides forward and backward. A ringlike
part of this tube is heavily chitinized, and when dried specimens are
used only this structure is obtained. It was called by Lindeman the
gabel, the parameren by Verhoeff and the tegmen by Hopkins. The
inner cover consists of a second tube which is heavily chitinized
throughout and which shows remarkable variations from species to
species. Figure 32 illustrates the peni of the genus under exami-
nation.
60 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
Parameren.—In Gnathotrichus the parameren (Pa) is a heavily
chitinized ringlike structure. It is entire, without a dorsal transverse
suture or knoblike projection cephalad, as in Pityogenes Bedel, and
was called umbellicus by Fuchs. Ventrally, a faint suture is visible in
Gn. sulcatus Lec. and retusus Lec. only. A prominent, heavily chiti-
nized hook extends cephalad from the ventral suture. This structure
was called the metula (Me) by Fuchs and the apodemal process by
Hopkins. In Gn. retusus Lec. and sulcatus Lec. a second, caudad,
but much smaller knob (a) is present. The tegmen furcae (Fuchs)
(b), which are a lateral continuation of the metula, are weakly de-
veloped. Lateral widenings of the parameren, Seitenfluegel (Fuchs),
are absent.
Specific modifications :—
A—Parameren without small knob ventro-caudad; metula long but
slender. Gn. materiarius Fitch.
AA—Parameren with small knob ventro-caudad; metula variously
modified.
B—Metula short, weakly developed. Gn. retusus Lec.
BB—Metula long, strongly developed. Gn. sulcatus Lec.
Inner covers—The inner cover (Fuchs), the body (Hopkins), the
penis tube (Nuesslin) is a tubelike structure, bilateral-symmetric. On
the inner covers three main parts may be distinguished which are the
lamina dorsales (Lad), the laminae ventrales (Lav) and the peduculi
penis (Fep). The dorso-caudad portion of the inner covers was called
the Endplatten by Lindeman, the laminae dorsales by Fuchs and the
dorsal plates by Hopkins. The laminae ventrales (Fuchs) or ventral
plates (Hopkins) are the corresponding ventral portion of the laminae
dorsales. In the genera Pityogenes Bedel, Ips de Geer, Pityokteines
Fuchs, Neothomicus Fuchs, and others, the laminae dorsales and ven-
trales are largely separated by deep emarginations caudad giving the
laminae the shape of four free projections which are connected basally
only. The basal, fused, entire part of the laminae ventrales is the
pallidium (Fuchs (Pal)). The sometimes narrow band connecting
the two dorsal plates is the jugum or Steg (Fuchs). In Gnathotrichus
the laminae dorsales and ventrales are fused laterally. Dorsally, the
two laminae dorsales are separated by an obscure suture only. The
laminae ventrales are fused ventrally, open on the extreme caudad
portion. The laminae dorsales as well as the laminae ventrales bear
numerous sensory pores on the caudad half. The latter is strengthened
by a stronger chitinized band, the caudad limitation of which (c) is
strongly emarginate. This chitinous strengthening was called the
NO. IO MORPHOLOGY OF GNATHOTRICHUS—SCHEDL 61
radius by Fuchs and this is well developed in all three species of
Gnathotrichus. A corresponding strengthening on the laminae dor-
sales, which as the manubrium (Fuchs) is well developed in the genus
Pityogenes is obscure in Gnathotrichus. The caudad portion of the
laminae ventrales, the caput (Fuchs), is variously modified, sometimes
bearing a beaklike projection dorsally which was called the rostrum by
Fuchs. The caput and rostrum vary considerably in shape and de-
velopment in the genus Gnathotrichus. The area of the pallidium,
from which the peduculi penis originate, the radix (Fuchs), is not
characterized by a heavier chitinization. The peduculi penis were also
called Fuesschen by Lindeman, femora penis by Verhoeff and body
apodemes by Hopkins. In the normal position the peduculi penis are
parallel; when mounted on slides they usually cross each other.
Cephalad the peduculi are slightly widened, their connection with the
pallidium is solid, not hingelike.
Enclosed parts-——The enclosed parts are a short part of the ductus
ejaculatorius, the preputial sac and chitinous strengthenings of the
latter. The ductus ejaculatorius is easily recognized by its enclosing
muscle structure. The author was able to trace this structure as far
caudad as the radius (c) extends. The preputial sac (Prp) consists
of a colorless membrane without any chitinous strengthenings. It
seems to be connected with the ductus ejaculatorius at the anterior
emargination of the radius. Chitinous structures such as the Rinne
(Fuchs) do not occur in the genus Gnathotrichus.
Parts outside of the covers——It was stated in the introduction to
the discussion of the penis that no indications of the spiculum ventrale
are present in the genus Gnathotrichus. While in Xyloterus Er. and in
some genera of the European Hylesinidac, the reduction of the Rinne
(Fuchs) mostly is followed by a stronger development of the spicu-
lum ventrale, Gnathotrichus presents a complete reduction of both.
Generic characters of the penis—Spiculum ventrale absent; para-
meren an entire ring, metula well developed; laminae dorsales and
ventrales fused laterally, the laminae dorsales dorsally separated by
an obscure suture, the laminae ventrales fused except on the extreme
caudad portion, the laminae forming a tube, radius distinct, manu-
brium obscure, jugum and pallidium not clearly defined ; sensory pores
on the caudad half of the laminae dorsally as well as on the laminae
ventrales ; the latter always extending farther caudad than the former ;
the peduculi penis slender, slightly widened cephalad, about as long
as the laminae dorsales, connection with the pallidium solid not hinge-
like; Rinne absent, preputial sac about as long as half of the laminae
ventrales.
62 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
In the classification of the Scolytidae, based on the chitinous skele-
ton of the penis by Nuesslin, Gnathotrichus would have been in con-
trast to all the other genera. This is easily explained by the fact that
only this one character was used in placing the genera. The author
has no reason to believe that Gnathotrichus is not a highly specialized
genus of the Pityophthorinae.
Specific modifications :-—
A—Parameren without ventro-caudad knob, metula long but slender ;
laminae dorsales with their dorsal margin straight, the dorso-
caudad angle obtuse and broadly rounded, the posterior mar-
gin is directed ventro-caudad from this angle; the laminae
ventrales with their ventral margin straight, only slightly
projecting caudad farther than the laminae dorsales, caput
weakly developed, nearly in line with the dorso-caudad mar-
gin of the laminae dorsales, rostrum obsolate ; peduculi penis
about as long as the laminae ventrales, very slender, only
very slightly widened at the cephalad end, basal part narrow.
Gn. materiarius Fitch.
AA—Parameren with ventro-caudad knob opposite the metula, the
latter variously modified; laminae dorsales with the dorsal
margin broadly arcuate, the dorso-caudad angle variously
modified either slightly acute but rather broadly rounded or
almost evenly arcuate with the dorsal margin of the laminae
dorsales; the laminae ventrales with their ventral margin
distinctly incurved, much farther projecting caudad than the
laminae dorsales, caput strongly developed, rostrum distinct ;
peduculi penes vary in length, stouter, more strongly widened
at the cephalad end, basal part distinctly widened.
B—Metula short, weakly developed ; the dorso-caudad angle
slightly acute but rather broadly rounded, the pos-
terior margin is directed ventro-cephalad from this
angle, caput not evenly rounded caudad, more tube-
like, ventro-caudad margin oblique, the dorsal angle
of the rostrum extending to about half of the
width of the combined laminae dorsales and ven-
trales, between the cephalad margin of the rostrum
and the caudad margin of the laminae dorsales,
a wide, rectangular space membranous; peduculi
penis as long as the laminae dorsales on the dorsal
margin. Gn. retusus Lec.
NGOS YO) MORPHOLOGY OF GNATHOTRICHUS—SCHEDL 63
BB—Metula long, strongly developed; the lamina dorsales
with their posterior and caudad margin almost
evenly rounded throughout ; caput evenly rounded
caudad, half sphericle, the membranous space be-
tween the cephalad margin of the rostrum and the
caudad margin of the laminae dorsales narrow,
slitlike, the dorso-cephalad angle of the rostrum
and the dorso-caudad angle of the laminae dorsales
in one level; peduculi penis distinctly longer than
the laminae dorsales. Gn. sulcatus Lec.
THE FEMALE REPRODUCTIVE ORGANS
The female reproductive organs (fig. 33) were examined from
Gn. materiarius Fitch only. As in all Rhychophora, two pairs of
Fic. 33.—Gnathotrichus materiarius Fitch.: Female reproductive organs.
A, the complete organs: B, receptaculum seminis; Ceg, cement glands; Ov, ovaries;
OvD, paired oviducts; Ree, receptaculum seminis; Tc8, eight tergites, reduced; Spg,
spermathecal gland; Spv, spiculum ventrale; Vag. vagina.
ovaries (Ov) are present which are connected distally. The paired
oviducts (OvD) are short and stout. Fhe vagina (Vag) is distinctly
shorter than the ovaries. Near the basal end of the latter two cement
glands (Ceg) originate. They are rather weakly developed, short
and ball-like in outline. A bursa copulatrix is not present. The re-
ceptaculum seminis (Rec) originates from the vagina at about the
5
64 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
same level as the cement glands; it has the shape of a pipe and bears
on its distal end the spermathecal gland (Spg). The receptaculum
was examined in all three species but no differences have been found.
In the classification of the Scolytidae, based on the female repro-
ductive organs by Nuesslin, Gnathotrichus would fall in a group to-
gether with the genus Thamnurgus Eichhoff.
THe ALIMENTARY CANAL
The alimentary canal was fully investigated in Gn. materiarius
Fitch, the proventriculus in Gu. retusus Lec. and sulcatus Lec., also.
The whole alimentary canal of Gnathotrichus is illustrated in figure 34,
the proventriculus in figure 35.
The alimentary canal is a tube extending from one end of the
body to the other. As in most of the mandibulary insects, three main
divisions are clearly defined. These are termed the fore, mid, and
hind intestine (fig. 34, A, B, C). The ectodermal origin of the fore-
and hind intestine are well illustrated by the occurring chitinizations.
Fore-intestine —On the fore-intestine the following consecutive
divisions are well defined: The Pharynx ( ?), Oesophagus (Oes), Crop
(Cr), and the proventriculus (Pve). The well developed proven-
tricula are the characteristicum of the adults.
The pharynx is not distinctly defined from the mouth cavity.
The oesphagus is about as long as the crop and the proventriculus
united. It consists of a simple tube, widened distally where it gradu-
ally passes over to the crop.
The strongly widened tube situated apically of the proventriculus
may be designated as the crop.
Proventriculus—The proventriculus is a highly specialized organ
in which the food is prepared before it enters the more delicate ventric-
ulus. The characteristic features of the proventriculus are a remarka-
ble development of the chitinous intima into folds and teeth and a
considerable increase of the size and development of the muscles of
this region. On account of the importance of the proventriculus as
a taxonomical characteristicum and of the general morphological in-
terest of this structure, a more detailed discussion seems to be
necessary.
Lindeman, who was the first investigator of the Scolytid proven-
triculus, distinguished two main parts or longitudinal divisions. The
anterior part he called the Sack, the posterior, or caudad part, the
Kaumagen. Nuesslin and Fuchs followed Lindeman’s nomenclature.
The Sack (Lindeman) corresponds to the crop (Hopkins) and forms
the intermediate part between the oesophagus and the Kaumagen
(Lindeman). Hopkins used the term proventriculus for the Kau-
NO. IO MORPHOLOGY OF GNATHOTRICHUS—SCHEDL 65
Fic. 34.—Gnathotrichus materiarius Fitch: Adult, alimentary canal and its
appendages.
A, fore intestine; B, mid-intestine; C, hind-intestine; Cr, crop; Mav, malpighian vessels;
Oes, oesophagus; Pve, proventriculus; Re, rectum; Ve, ventriculus; a, large intestine;
b, posterior tube of the mid-intestine; c, small intestine.
Fic. 35.—Gnathotrichus materiarius Fitch: Proventriculus.
a, median line; 6, sutural teeth; c, anterior closing teeth; d, dentation of masticatory
teeth, all together, masticatory brush; e, femora of the masticatory teeth, Abdachungszaehne
(Nuesslin); #, posterior closing teeth; A, masticatory plate; B, anterior plate.
66 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
magen (Lindeman) only. In the following discussion the interpreta-
tions of Hopkins of the terms crop and proventriculus are used.
The proventriculus consists of eight plates arranged to form a
tube, as illustrated in figure 35. On each plate two longitudinal divi-
sions are clearly defined. Lindeman called the cephalad part the Plat-
tenteil (A), the caudad part the Ladenteil (B) ; the corresponding
terms of Hopkins are anterior plate and posterior or masticatory
plate. The latter terms are adopted in this paper. In Gnathotrichus,
both of these plates are divided by a median suture (a) which is dis-
tinctly visible on the masticatory plate and indicated by a row of
bristles (b) on the anterior plate. These bristles, which are of taxo-
nomic importance, have been called Zaehne am medianen Kauplatten-
rand by Nuesslin and sutural teeth by Hopkins. In Gnathotrichus
they are present as slender, sharply pointed, and simple bristles. The
anterior plate is not quite one-sixth as long as the whole proventriculus.
The masticatory plate (A) bears a symmetrical arrangement of teeth
which presents the true chewing apparatus of the proventriculus. On
each tooth two elements are clearly defined, the instep and the den-
tation. The totality of the insteps corresponds to the Abdachung (c) ;
that of all dentations to the Buerste (a) of the German authors. Hop-
kins called the first mentioned the femora of the masticatory teeth,
the latter the masticatory brush. The masticatory teeth are all similar
in shape and very numerous in the genus Gnathotrichus. Cephalad,
they are bordered by a few (8-12) longer teeth (c) which differ
greatly in shape and which are directed toward the center of the
proventriculus. These teeth are apparently intended to regulate the
entering of the food. They have been called Sperrborsten by Nuess-
lin and closing teeth by Hopkins. To distinguish them from a similar
arrangement of teeth which occur in Gnathotrichus and other genera
on the caudad end of the masticatory plate ({) it is proposed to call
the former cephalad closing teeth, the latter caudad closing teeth. In
Gnathotrichus, the following armatures are not present :
(a) Hackenzaehne (Nuesslin), or apical teeth of the anterior plate;
the designation of a row of short, often curved teeth on the
apical margin of the anterior plate.
(b) Ersatzperrborsten (Nuesslin), marginal bristles or marginal
fringe (Hopkins) ; a longitudinal row of bristles along the
lateral margin of the anterior plate.
(c) Kreuzlinie (Nuesslin), a row of short, stout teeth arising from
the lateral margin of the anterior plate and converging pos-
teriorly.
NO. 10 MORPHOLOGY OF GNATHOTRICHUS—SCHEDL 67
(d) Abdachungszaehne (Nuesslin), masticatory teeth which have
two dentations, one, the totality of all composing the mastica-
tory brush, and a second smaller tooth on the instep, the
totality of which forms a second brush consisting of a single
row of teeth only.
In the classification of the Scolytidae based on the proventriculus,
Gnathotrichus should be placed with Xyleterus Er., Xyleborus Eichh.,
Anisandrus Ferr. and other ambrosia beetles together in one group.
There is little doubt that the similarity of the digestive systems in
ambrosia beetles of the Superfamily Scolytoidea is of no importance in
the classification. These are merely parallel] modifications of groups
deriving from very different ancestors.
The proventriculus is very similar in all three species of Guathotri-
chus and it is not possible to distinguish them by characters of this
part.
Mid-intestine —The mid-intestine is about one-third of the whole
length of the alimentary canal. According to Nuesslin the proportions
are the same as in Amsandrus dispar Fabr. and Xyloterus lineatus
Oliv.
In Gnathotrichus two subdivisions of the mid-intestines are well
defined. The anterior part, which presents the widest part of the whole
alimentary canal, has the form of a ball and is covered with short,
half spherical gastric coeca. This part is here designated as the ven-
triculus proper. The posterior, much narrower tubelike part, is here
called the posterior tube of the mid-intestine. It bears much smaller
gastric coeca which distinctly decrease in size toward the origin of
the Malpighian vessels and which are always more filiform in shape.
The origin of the Malpighian vessels marks the posterior limit of the
mid-intestine. These are rather narrow, long tubelike vessels, strongly
entangled around the posterior part of the mid-intestine and do not
vary noticeably in size.
Hind-intestine —While the fore- and mid-intestine are situated as
a straight tube in the body, the hind intestine forms a distinct loop.
The latter does not bear any gastric coeca but is characterized by the
muscles which enclose it. The subdivisions, the small and large in-
testine and the rectum, are not so clearly separated. The ovaries of
the female reproductive organs are always situated above the hind
intestine ; the testi of the male lie under and ventral to it.
THE LARVAE
On account of the difficulty in getting material of the western
species, the following discussion is based on Gnathotrichus materiarius
Fitch only.
VOL. 82
SMITHSONIAN MISCELLANEOUS COLLECTIONS
68
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70 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
The structure and general appearance of the larva are shown in
figure 36. The larva is legless, subcylindrical, white in color except
the heavily chitinized and therefore reddish-brown headcapsula. The
full grown larva is about 3.7 mm. long. Three thoracic and nine
abdominal segments are well developed. The anal lobes may be re-
garded as a tenth abdominal segment. The three thoracic segments
are nearly equal in size and only little larger than the first abdominal
segment. The abdominal segments decrease slightly in width and
length toward the apex. All segments and the head are armed with
constant setae. The statement of Hopkins (38) that “ with the ex-
ception of scattering hairs on the head and on the scutellar lobes of
the thoracic and abdominal segments the body is without distinguish-
ing vestiture ” in Dendroctonus has been found to be not true. There
occur at least in Dendroctonus valens distinct setae. That this is not
exceptional in the Scolytidae has been proved by Russo (57) and the
author. The latter has studied many species and has found that every
examined species shows distinct setae. The following discussion is
based on the full grown larvae only. The fact that the present study
was made in a private home did not allow of rearing work and there-
fore the question of molds, etc., cannot be discussed at present.
THE CHITINOUS: SKEEETON
THE HEAD
The head of the larva is more simple in structural details than
that of the adult. It is distinctly narrower than the first thoracic seg-
ment when seen from above, but nearly equal in length and in width
to the latter when viewed from the side. The general structure is
shown in figure 36, the anatomical details in figure 37. The more
striking differences in the larval head are found in the presence of
a clearly defined front, clypeus, labrum, and a well developed sub-
mentum.
Epicranial suture -——The sutura metopica as well as the sutura
fronto-verticale are well developed and double lined (a, b). They are
not raised or padded as in the adults.
Front.—The front (fig. 37, Fr) is triangular in outline, plano-con-
vex and clearly defined by the epicranial suture. The lateral sides are
bordered with six strong bristles each (d). These setae are here
called setae fronto-lateralis. The base of the front is smooth without
any armation.
In the discussion of the larval setae it seems to be very useful to
compare the results obtained with those of other authors. From the
NO. 10 MORPHOLOGY OF GNATHOTRICHUS—SCHEDL FL
literature available, only two species have been found to be studied
at the present. These are Dendroctonus valens Lec., investigated by
Hopkins, and Chaetoptelius vestitus Fuchs studied by Russo. The
ea Pst
Fic. 37.—Gnathotrichus materiarius Fitch: Head of fully-grown larva; A, fron-
tal aspect, B, ventral aspect, C, lateral aspect, D, occipital aspect.
An, antenna; C/, clypeus, seta clypei; Eg, entogular plate; Ep, epicranium; Eph, epi-
pharynx; Est, epistoma; Fr, front; Ge, gena; Gu, gula; Hst, hypostoma; Lar, labrum,
setae labralis; Li, ligula; Md, mandible; Me, mentum; Mz, maxilla; Occ, occiput; Ocd,
occipital apodeme; Pr, pregena; Pst, pleurostoma; SM, submentum; Ve, vertex; a, sutura
metopica; b, sutura fronto verticale; c, seta geno mediana; d, seta fronto lateralis; e, seta
epistomalis; f, seta verto mediana; g, seta verto lateralis; h, seta geno lateralis; 7, seta
epicrano-lateralis; 7, gular suture; k, maxillare foramen; stl, sternal lobe.
comparison of the setal arrangement with the latter species will be
given at the end of each chapter.
Clypeus—As mentioned before, the clypeus (fig. 37, Cl) is well
developed in the larvae. It consists of a nearly rectangular plate
which is heavily chitinized, distinctly emarginate anteriorly and
broadly rounded antero-laterally. Basally it is separated from the
72 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL, 82
epistoma by a faint suture. A pair of bristles and two punctures
occur near the basal margin. The setae are called setae clypei.
Labrum.—The labrum (fig. 37, Lar) is basally separated from the
clypeus by a distinct suture. It consists of a heavily chitinized half
circular plate. It is as long as, and about one-third narrower, than
the clypeus and bears several papilla apically. Half way of its length
occur two long bristles, the setae labralis.
E pistoma.—Between the front and the clypeus, a narrow thickened
transverse band occurs which may be designated as the epistoma
(Est). Laterally this structure is slightly bent forward and this
part bears the dorsal articulation of the mandibles. On the latero-
basal angles, near the suture, a long bristle is situated. The proposed
name is seta epistomalis.
Pleurostoma.—A faint suture hems the lateral portion of the oral
foramen, parallel to the lateral exposed part of the mandibles, which
cuts off a narrow area of the cranium. This area, the pleurostoma, is
slightly raised externally; internally it is ridgelike, and connects the
epistoma with the hypostoma.
Hypostoma.—The hypostoma (fig. 37, Hst), which bears the ven-
tral articulation of the mandibles, is rather ill-defined externally. In-
ternally it is ridgelike and the articulatory condyles and fossa are
submerged. Towards the occipital foramen it is extended in another
ridge from which the connecting membranes of the maxilla and the
submentum arise.
Occipital foramen.—The occipital foramen (fig. 6, Oct) is situated
on the caudad face of the head capsula. It is heart-shaped and is bor-
dered by a ridgelike rim, the occipital apodeme (fig. 37, OcA). The
latter is interrupted ventrally. The entogular plate extends into the
foramen under the occipital apodeme giving the open space of the
foramen the shape of a triangle, the sides of which are broadly
rounded.
Gula.—The gula (Gu) is present as two small lobes, each situated
along the ventro-lateral angles of the occipital apodeme. The lateral
limitation is indicated by an obscure suture.
Entogular plate—A subchitinous plate (Eg) connects the genal
areas and extends ventrally up to the hypostoma. It was called the
entogular plate by Hopkins. The open space of the oral foramen is
also reduced by the lammella-like extension of this plate. The entogu-
lar plate is not visible externally but hidden from the submentum.
Pregena.—A narrow area along the ventral extensions of the hypo-
stoma, and laterally limited by obscure lines, may be considered as
the pregena (Pr).
NO. IO MORPHOLOGY OF GNATHOTRICHUS—SCHEDL 73
Other topographical regions of the cranium are not defined by
sutures but they are somewhat limited by the occurring setae.
V ertex —The area on both sides of the sutura fronto-verticale, the
vertex (Ve), bears two groups of setae. Eleven bristles ({) are situ-
ated along each side of the sutura fronto-verticale. Four of them
are arranged in a single row along the suture; the rest occur in the
angle formed by the sutura fronto-verticale and the sutura metopica.
They are called setae verto-mediana. A single bristle widely separated
from the setae verto-mediana, the seta verto-lateralis (g) presents the
second group. Sometimes a minute hair is also visible near the latter
mentioned bristle.
Gena.—The area between the sutura metopica and the occipital
apodeme ventro-laterally to the vertex is here designated as the
gena (Ge). It bears two groups of setae, one on the level of, and close
to, the antennae (c) and one laterally (h) to them but widely sepa-
rated from them. The former consists of five, the latter of six long
bristles. The corresponding names are the setae geno-mediana and
the setae geno-lateralis.
Epicranium.—tThe area dorsally to the genae and laterally to the
vertex is called the epicranium. A single bristle (i) sometimes asso-
ciated with a minute hair, is situated near the occipital apodeme. It
is called the seta epicrano-lateralis.
THE APPENDAGES OF THE HEAD
THE ANTENNAE
The antennae (fig. 37, An) are present as small membranous lobes
next to the ventral end of the sutura metopica. Each bears one papilla
and numerous minute hairs.
THE MOUTHPARTS
Mandibles —The mandibles (fig. 28) differ somewhat from those
of the adults, in contrast to which only one median tooth is present
and the shape of the mandible is more slender and triangular. The
setae are present in the same number as in the adults but their position
in relation to each other is different. Proposed names: setae mandi-
bulae dorsalis and setae mandibulae lateralis.
Mavzillae-—The maxilla of the larva (fig. 38, A) is much simpler
in structure than that of the adult. However, all parts present in the
adult maxilla are also distinguishable in the larva.
The cardo (Ca) is present as a distinct sclerite, triangular in out-
line and connected with the extended hypostomal ridge, the mentum
and the stipes respectively. This is, as in the adult, not the only con-
74 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
nection of the maxillae with the mentum. The subgaleal area is also
connected with the mentum along its interno-lateral margin. The
stipites (St) are distally not subdivided into a palpifer, and are fused
internolaterally with the subgalea.
A single bristle (b) occurs near the externo-lateral margin. It was
called the setole laterale dello stipite by Russo. In the present paper
the name is modified into seta stipitis maxillaris. The palpiferal area
Fic. 38.—Gnathotrichus materiarius Fitch: A, labium and maxilla, ventral as-
pect: B, clypeus and labrum, ventral aspect.
Ca, cardo; Fr, front; Eph, epipharynx; Ga, galea; La, lacinia; Lab, labium; Lar, labrum;
Lpp, labial palpi; Li, ligula; M, mentum; Ma, maxilla; Oes, oesophagus; Pp, maxillar palpi;
Ppf, maxillar palpifer; SM, submentum; St, stipes; C, clypeus; a, seta palpifera maxillaris;
b, seta stipitae maxillaris; c, seta ligulae distalis; d, seta submento-lateralis; e, papilla labro-
apicalis; #, papilla epipharno-distale; g, papilla clypei; h, seta menti; 7, seta ligulae mediana;
7, seta labio-palpiferis, and palpiferal area; k, seta ligulae basalis, and basal area; /, seta
lacinio-maxillaris; m, epistomal apodeme; n, apodeme between lateral angle of epistoma and
dorsal angle of pleurostoma.
bears two bristles (a) and one puncture; the outer corresponds to
the setola laterale dello palpifera, and the inner to setola mediana
dello palpifera (Russo), here called seta palpiferae (maxillaris). The
palpus is two-segmented, telescopic, the first segment armed with a
short fine hair and two punctures, the second with punctures only. The
bristle on the first segment is the setola palpiale (Russo) but to dis-
tinguish it from possible setae on other joints, it should be called
seta palpo-maxillaris 1°. The lacinia is present as a well developed
lobe distally armed with two long bristles (setae lacinio-maxillaris )
NO. 10 MORPHOLOGY OF GNATHOTRICHUS—SCHEDL 75
and a few papillae. The galea is largely fused with the lacinia. Dis-
tally a rather shallow fold indicates the separation of these two
sclerites.
The setae of the maxillae have not been studied thoroughly enough
to decide if they are of taxonomic importance or not. In Den-
doctonus valons, no setae or punctures occur on the palpus and none
of them on the lacinia, according to the drawings published by Hop-
kins. On the other hand it was found that the setal arrangement of
TABLE I.—Setae of the head, nomenclature.
Nomenclature of Dr. Russo Used on New nomenclature Used on
Setole mediane-distale .....
Setole submediane ....... BronGh acess saee Seta fronto-lateralis ........ Frons.
SGUGIS TENGTEAIN Ceo toge conan : ;
Ratner basalinescadcas cee o cents Frons-epistoma .Seta epistomalis ....... .--. Epistoma.
Setole basali clypeali....... Glypeus! =. tsicee Setar clypeit casvsrcieisiecieiare ...-Clypeus.
Setole mediane distali.....
Setole premediane ........ ILeiayebir Go oadac Seta wlabraliisaeeveysettelatetete t=! Labrum.
Setole sublaterali .........
Setole basali-laterali ...... A
Seta verto-mediana ...-..-) vortex
Seta verto-lateralis ....... )
Setole laterale del vertice..
Setole mediane ........... Genae ye ciieer -Seta geno-lateralis ....... .. Genae.
Setole basale?.. ja: <2.» S908
eee He a del vertice..
Setole della gena..........
Setole mediane ... oat WGENAG | sertarsrers Seta geno-mediana ......... Genae.
Setole esterna
Setole basale Y Seta epicrano-lateralis ...... Epicranium.
Setole dorsali ...1.+-12+++++Mandible ..1.Seta mandibuiae lateralis. . { Mandible.
Rereialaterale dello stipite. . . 5tiPes aoc Oso Seta stipitae maxillaris...... Stipes.
Setole mediane del palpifero. | Palpifer .......Seta palpiferae maxillaris....Palpifer.
Setole laterale del palpifero. ' First joint of | Seta palpo-maxillaris 1°..... First joint of
eLole palpiale) occ. cleccie «0 palpus. te ; . palpus.
Seta lacinio-maxillaris ..... Laciniae.
aa eee Ug ae Ges ae [Sabena ...Seta submento-lateralis ..... Submentum.
1 ; :
Setole subbasali ......... ; Seta menti .....-...-.--.0. Mentum.
Retalemenbhasalie.scsocce. _Mentum Seaeicrie etal lablo palpitenisircijeretelelere Beer ee
: Seta ligulae distalis.
Setole distale ............ Wei puila, smcen Seta ligulae mediana.
Setole mediane ..:.....0ce if Seta ligulae basalis.
Gn. materiarius Fitch is exactly the same as that of Chactoptelius
vestitus Fuchs, illustrated and described by Russo.
Labium.—tThe labium (Lab) of the larva is very different in struc-
ture from that of the adult. Indications are that the labium of the
larva as well as that of the adult will become more and more im-
portant as the bearer of taxonomic characters in the Scolytidae.
The submentum (SM) is present as a large sclerite and is trapezoid
in outline. The shape is more like that of Chaetoptelius vestitus Fuchs.
It is slightly chitinized, laterally connected with the maxillae and
bears three pairs of setae (d), the same number and in a similar ar-
rangement as in Chaetoptelius vestitus and Dendroctonus valens.
Russo called these setae setole subbasale, setole mediane and setole
distale in order proceeding distad. The anterior margin of the sub-
6
76 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
mentum does not extend as far as the origin of the palpi as in the
two other species mentioned above.
The mentum (M) is a submembranous triangular plate the base
of which is anterior. The posterior angle is narrow and strongly
produced. Anteriorly it is fused with the palpiferal area of the labium.
A single bristle and a puncture (h) are situated on the antero-lateral
angles. The name proposed is seta ment.
Tas_e I].—Comparison of head setal arrangement.
Chaetoptelius Dendroctonus Gnathotrichus
Seta vestitus valens materiarius
Fuchs. Lec. Fitch.
HM TOntoslatetallismme sera 4 4 6
Epistomaliswer rn... acs aerate I I I
Gly peices cc gecesmnew oc Oca: 2 I I
JEallp palliS Rese erctcrmtarsrs snscore Teel ones 5 2° I
Werto-medianay qaceie secs sor (@) iF 12
Nertolatenalisne esta sce eer (a) ? 2
Geno lateralismeeee eee meio 8 pit 6
Geno=medianaue eee ene einen 4 @ 5
E\picrano-lateraliss apes (0) ? I
Mandibulae-dorsalis .......... 2 2 2
Mandibulae-lateralis .......... I I I
Stipitissimeaxallaris/eerserei cern I iP I
Palpiterae maxallarisy sos... 2 R 2
Palpo-meaxalllanicneeenneeeeere I (a) I
acinio-meaxallanismeer reece I (0) 3
Submento-lateralis..-5..--- 3 3 3
Mientti Sas tinesind fs icisraie saa atest 0) 0 I
Lalor “sococnconocduec I I 2
Wigtlae=distalismecemeee ret I I I
ioulae-medianaw eee rier (a) 0) I
ieulaesbasalicuem mermtaeie I I I
The ligula (Li) consists of a circular plate largely surrounded by
the mentum. From subdivisional parts the base of the ligula is visible
from which it is well defined by faint sutures and the palpi (Pp).
The base bears a single pair of setae. They are present as two short
bristles and are called the setae ligulae basalis. The palpiferal area
is situated on both sides of the base. The part anterior to the base of
the ligula and between the palpi may be designated as the distad end
of the ligula. The palpi are two-jointed. The palpiferal area bears
two setae (j) which are called setae labio-palpiferis. The distal area
of the ligula is armed with two pairs of setae. Proposed names: setae
ligulae mediana and distalis.
NO. IO MORPHOLOGY OF GNATHOTRICHUS—SCHEDL WG)
THE THORAX
The thoracic segments are somewhat larger in size than those of the
abdomen. They do not differ from each other in shape but are dis-
tinguished by the structure and the development of the setae. The
thoracic segments are legless as in the whole superfamily but the foot
calli are distinct.
Dorsally, the prothorax is distinguished from the other two thoracic
segments by the lack of the suture which divides the prescutum
(fig. 36) from the fused scutal-scutellar area. However, the prescu-
tum is indicated by the corresponding setae. On the meso- and meta-
thorax, the prescutum is present as a narrow transverse sclerite (f),
near the anterior margin of which the prescutal setae (f,4) are
situated. The scutum (e) and the scutellum are in all three segments
fused. The pleural area is fairly well defined by a longitudinal fold
ventrally. Latero-dorsally it is fused with the scutal-scutellar area.
Another longitudinal fold divides the pleural area into two distinct
parts. The part next to the scutal-scutellar area represents evidently
the epipleurite (d), the ventral part the hypopleurite (c). The former
has a smooth surface bearing setae only; the latter has the surface
covered with minute spines beside the setae. On the sternum two
subdivisions are plainly visible, the sternal (a) and the sternellar
area (b). The sternellar area or sternellum consists of two large
lateral lobes which are connected by a very narrow band medially.
The lateral lobes which bear the foot calli are covered with minute
spines similar to those in the hypopleurites. There is not enough evi-
dence to speak about a poststernellar area in Gnathotrichus as it
should be present in Dendroctonus according to Hopkins. The setal
arrangement will be discussed with that of the abdomen.
THE ABDOMEN
The abdomen (fig. 36) consists of ten segments, the tenth of which
is strongly reduced and present as the anal lobes. The segments de-
crease in size slightly towards the apex. Segments one to seven in-
clusively are alike in structure and setal arrangement. They differ
in structure from the meso- and metathorax in having longitudinal
folds which separate the epipleural area from the sternum and the
scutal-scutellar area. Also the sternal-sternellar suture is restricted
to a membranous fold. The two lateral lobes of the sternellum are
apparently not connected medially. The eighth segment does not show
signs of the prescutal-scutal suture. Still more reduced is the ninth
segment ; it has no sutures or folds but the different sclerites can be
78 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
determined very plainly from the position of the setae. The anal or
tenth segment is separated from the ninth by an obscure suture. It
consists chiefly of the four anal lobes. There is no difference in the
structure of these lobes and also armations do not occur as in other
genera.
THE SPIRACLES
Nine pairs of spiracles are present, eight of which are situated on
the epipleurites of the first eight abdominal segments. The ninth
spiracle is on the same sclerite of the prothorax very close to the
mesothorax.
THE THORACIC AND ABDOMINAL SETAE
It was found that the setal arrangement is very constant from
specimen to specimen. The number of setae varies in the different seg-
ments. The smallest number of setae was found to be present in
the anal segment (2 x 2) and the eighth abdominal segment (2 x IT) ;
the greatest number was borne by the meso- and metathorax (2 x 22).
The number of setae in the different segments is best explained by
Tasie III].—Setae of thorax and abdomen, nomenclature.
Used on the Used on the
Nomenclature of Dr. Russo segments New nomenclature segments Fig. 36
Setole tergali mediana.......... Ti, UES JU re : F Tae t
Sefoleuprotercalistemeiteerorsieterts 1-8 {Seta praescutl ...++- 1-9 t
Setole posttergali .............. I, IT, III, 1-8 Li. Wa
Setole tergali-laterali .......... I Setay scutilul cyvsieiete iiet) ?
Setole tergali ee ienicoee ont 9
Setolemepiplerri cremrecteratcieielerslerer I, 1-8 I Wut
Setole pleuri-sternali .........-. 1 Seta epipleuricum ... } EM ot) , t d
Setole fereal pleural: dsisistinieret 1 a 1-9
Setole upopleuralt, (eee res) nn sle'es ar
Setole pleurali-sternali ......... z set hypopleuricum .. { I, 1, WII, ¢ c
Setoleepipleunit ie ocr isereleleiaieletee 15 ANE 38)
Setole sternali-anteriori-externe. . I
Setole sternali-posteriori-externe.. I
Setole sternali-mediane ........ I Oncor
Setoleipopleurales -r-teretejers cle cievele Mis Seta sternellaris 2 hee Ai b
Setole sternali-laterali .......... its LAU 1-9
Setole sternali mediane......... his 1H
Setole stermaliy i
ra a > wel ¥ Lie i pate a ee (uel ehaiv:
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 82, NUMBER 11
RECENTLY DATED PUEBLO RUINS
IN ARIZONA
(WITH 27 PLaTEs)
BY
EMIL W. HAURY and LYNDON L. HARGRAVE
(PUBLICATION 3069)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
AUGUST 18, 1931
a. eed
———
j The Lord Wattimore (Press
RALTIMORE, MD., U. S. A.
RECENTLY DATED PUEBLO RUINS IN ARIZONA
By EMIL W. HAURY anp LYNDON L. HARGRAVE
(WITH 27 PLATEs)
FOREWORD
In 1921 the National Geographic Society began excavation and
study of Pueblo Bonito, a prehistoric ruin in Chaco Canyon, north-
western New Mexico. It was the privilege of the undersigned to
direct these archeological investigations from their inception to the
conclusion of actual field-work in the autumn of 1927.
Pueblo Bonito is unquestionably the finest extant example of that
phase of southwestern history known to archeologists as Pueblo III—
the period during which small, isolated villages were drawn together
into large, complex communities. After a time these communities
began to disintegrate; to separate into lesser groups that spread far
and wide in search of more fertile fields and’ freedom from the attacks
of nomadic enemies. Then came, in 1540, Coronado and his fellow
adventurers to initiate the Spanish conquest of the Southwest; to
bring about still further disintegration of the peaceful Pueblo tribes
and their unique social organization. This Spanish-dominated phase
of Pueblo history, from 1540 to the present, is commonly designated
Pueblo V. .
In undertaking exploration of Pueblo Bonito, one of our prime
desiderata was the determination of a method whereby this famous
ruin could be correlated accurately with the recorded history of the
Southwest. To be sure, the relative age of the ruin had previously
been ascertained through stratigraphic methods and the study of
pottery types. It was known to be older than certain ruins; younger
than others. Our hope, notwithstanding, was to discover an absolute
date—the very year, if possible, during which its prehistoric walls
were building.
This desire was finally realized by a novel but altogether convinc-
ing method. As I have explained in a short paper published in
“Explorations and Field-Work of the Smithsonian Institution in
1929” (Dating our prehistoric Pueblo ruins, pp. 167-176, Washing-
ton, 1930), Dr. A. E. Douglass, Director of Steward Observatory,
SMITHSONIAN MISCELLANEOUS COLLECTIONS, VoL. 82, No. 11
2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
University of Arizona, accepted an invitation from the National
Geographic Society to extend his researches in dendro-chronology
to include such beam material as might be provided by the Society’s
expeditions. While the age of Pueblo Bonito was the Society’s sole
objective Dr. Douglass’ chief interest lay in the evidence of sun-
spot influences and climatic variations as revealed by the annual
growth rings of the timbers.
Three separate collecting parties, under the general supervision
of Dr. Douglass and myself, were sent out by the Society in the
summers of 1923, 1928 and 1929. Mr. Haury has briefly reviewed
the purpose and results of these successive expeditions in his intro-
duction. It should be emphasized, however, that we had a definite
plan constantly in mind; that we worked as directly as possible
toward our objective. When Dr. Douglass had brought his ring-
record into two separate sequences we sought to join them into a
single series. It was the Third Beam Expedition, that of 1929,
which finally crowned this unique adventure with success.
As Mr. Haury states, a certain transient phase of Pueblo pottery
provided the clue to ruins which immediately antedated Oraibi, the
inhabited Hopi village from which Dr. Douglass had secured his
oldest historic timbers. From collections in the United States Na-
tional Museum and elsewhere a list was prepared of 20 prehistoric
villages from which that particular type of pottery had previously
been gathered. Our 1929 reconnaissance was undertaken for the
purpose of eliminating from that list those ruins in which there
seemed little likelihood of finding charred fragments of pine ceiling
beams, for only thoroughly charred timbers could have resisted seven
centuries of decay in an exposed site. Of the ruins visited on that
preliminary survey four only were selected for partial examination :
Kin Tiel and Kokopnyama, north of the Little Colorado River, and
Pinedale and Showlow, in the forested area to the south. Mr. Har-
grave, from his more intimate knowledge of early Hopi cultures, was
placed in charge of excavations at the two former sites; Mr. Haury,
at the latter two.
In the pages which follow, Messrs. Haury and Hargrave describe
their individual efforts in the four ruins above named. Since these
are more or less well known to all students of Pueblo archeology, it
is felt that the authors’ observations will form a welcome addition
to the rapidly growing literature on the Southwest, especially in view
of the fact that each of the ruined villages has now been correlated
definitely with our own calendar.
NO. TL PUEBLO RUINS IN ARIZONA—-HAURY AND HARGRAVE 3
In behalf of the National Geographic Society, Dr. Douglass and
myself, I wish to take this opportunity publicly to acknowledge our
joint appreciation of the zeal and industry exhibited by Messrs.
Haury and Hargrave in pursuing the tasks individually assigned
them; to further acknowledge our obligations to those citizens of
Arizona who, by granting us permission to excavate on their lands
and in other ways, contributed so directly to the success of this
important undertaking. Especially are we grateful to the Directors
and to Dr. Harold S. Colton, President of the Museum of Northern
Arizona, at Flagstaff, for so generously placing at our disposal the
valued services of Mr. Hargrave. Dr. Colton aided us still further
by personally leading our 1929 reconnaissance party; by providing
Dr. Douglass with temporary laboratory space in his museum; by
expediting in every way possible the purpose of our concluding
expedition.
Net M. Jupp.
U. S. National Museum,
November 17, 1930.
SHOWLOW AND PINEDALE RUINS
By Emit W. Haury
INTRODUCTION
The year 1923 marks the inception of a new method for ascertaining
the actual ages and determining the chronological sequence of South-
western ruins—a method devised by Dr. A. E. Douglass and based
upon the annual ring-growth of certain coniferous trees. A brief sum-
mary of the results of this seven-year investigation has recently been
published by him in the National Geographic Magazine.’
Doctor Douglass has conclusively shown that the width of annual
rings of pine in the Pueblo area is conditioned by the amount of
precipitation; thus, in wet or favorable years, ring-growth will be
normal, while in drought years the growth will be sub-normal, the
width of the rings decreasing with the severity of the drought. He has
shown also that practically all trees over a large area record the
periodical fluctuations in moisture in identically the same way. Com-
menting further on this point, Doctor Douglass says in his recent
article:
The same succession of drought and plenty appears throughout the forest.
.... Certain sequences of years become easily recognized from tree to tree,
county to county, even from State to State.
Furthermore, it has been shown to be highly improbable that a
given ring-sequence with its characteristic narrow rings will ever be
exactly duplicated. In the present continuous calendar which extends
over a period of 1,200 years, duplications in even short ring-records
have not been discovered.
With the above facts in mind, it should be possible by a method of
cross-dating or over-lapping the inner rings of one beam with the
outer rings of another, first in living trees and then in old timbers cut
by man before the living trees started their record, to build up a
chronology which would extend far into the past.
In June, 1923, the First Beam Expedition of the National Geo-
graphic Society entered the field for the express purpose of recovering
* The secret of the Southwest solved by talkative tree rings. Nat. Geogr. Mag.,
Vol. 56, No. 6, pp. 737-770, December, 1929. Doctor Douglass will elaborate
his methods and results in a paper to accompany the report on the Pueblo Bonito
explorations, by Neil M. Judd, now in preparation.
4
NO. II PUEBLO RUINS IN ARIZONA—HAURY AND HARGRAVE 5
beams which would contribute to the historical sequence established
shortly before from living trees in the forest about Flagstaff, Arizona.
This chronology had been extended back to about A. D. 1260. Pre-
historic beams which were brought in by the first field party and
additional timbers sent in from Pueblo Bonito, Aztec, Mesa Verde,
and other ruins, provided enough material for an extended exami-
nation. By 1927 two prehistoric chronologies independent of the
historical ring-record had been evolved, the one from Pueblo Bonito
and Aztec beams, and the second from timbers obtained in Citadel
ruins, Mesa Verde, and others. In 1928 the Pueblo Bonito and
Citadel chronologies were united by specimens from Betatakin and
Keet Seel, two northern Arizona cliff dwellings. Thus a prehistoric
but independent sequence of more than 580 years was established.
During early spring of 1928, the Second Beam Expedition started
its search for timbers to link the two existing chronologies. A careful
survey of the Hopi villages indicated that available beams there were
not sufficiently old. Then, in order to discover such ruins as were
neither too old nor too recent, recourse was taken to pottery to serve
as a guide in the further selection of sites for examination. Prelimi- .
nary studies by Doctor Douglass and Mr. Lyndon L. Hargrave of pot-
tery types, in which a sequence of development was evident, and of
associated dated timbers soon revealed the fact that for a certain
form of decorated orange-red pottery no dates were available,
whereas older and more recent pottery types could be assigned either
relative or absolute dates. Supposedly, then, the undated orange-red
pottery fell in the gap which separated the two chronologies. Hence,
ruins where such pottery was plentiful might possibly supply the
needed timbers.
_A reconnaissance of sites in the Little Colorado River drainage in
the spring of 1929 resulted in the selection of the following four ruins
for investigation: Kin Tiel to the northeast of Holbrook, Arizona;
Kokopnyama in the Jadito Valley ; Showlow and Pinedale ruins in the
pine forest of the Mogollon Plateau 50 miles south of Holbrook. On
these sites the Third Beam Expedition centered its activities, Mr.
Lyndon L. Hargrave working first at Showlow ruin with the writer
and later in Kin Tiel and Kokopnyama, while the writer continued ex-
cavations in Showlow and Pinedale ruins. The findings of the two
units of the expedition, collateral to the search for beams, are given
in this joint paper.
The Third Expedition was successful in recovering timbers which
tied together the historic and prehistoric chronologies, thus carrying
6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
the tree-ring calendar back to A. D. 700. Beam HH-39, found at
Showlow, proved to be the key beam that converted the relative dating
series of more than 580 years into the absolute chronology by con-
firming what was found to be a short over-lap instead of a gap. The
outer rings of the specimen in question could be read to about A. D.
1380, while its central ring dated A. D. 1237. The innermost rings
coincided with the last rings of the prehistoric chronology and the outer
rings were readily identifiable with the 13th and 14th century records
of the historic sequence. Subsequent excavations yielded additional
timbers which covered practically the same period, thus substantiating
the record of the key log and relieving all feeling of uncertainty that
might accompany the placing of so much reliance on a single beam.
The relation of tree-ring studies to archeology is obvious with the
realization that beams showing true outsides found in ancient dwellings
will not only give the cutting date of the timber, but will also strike
near the time of the construction of the dwelling. Eventually we may
be able to interpret the entire development of Pueblo architecture in
terms of actual dates and even trace minor changes of architecture
within a single pueblo. Needless to say, the artifacts associated with
dated ruins become more instructive in view of the added data con-
cerning their antiquity.
The future of tree-ring research as applied to archeology is ex-
tremely promising. The present status of the study, which has enabled
the dating of approximately 40 pre-Spanish ruins, is the bare beginning
of what will eventually be accomplished. There remains yet the dating
of many ruins which have escaped thorough investigation ; the back-
ward extension of the tree-ring calendar beyond the present terminal
date of A. D. 700, so that structures older than Pueblo Bonito may be
assigned to their respective places in the scale of development ; and the
dating of the many impressive ruins in the Rio Grande drainage, New
Mexico, which, as yet, have gone undated. Other phases of the prob-
lem, now unthought of, are certain to follow.
In the following report, an attempt is made to correlate the archeo-
logical observations derived incidental to the search for charcoal with
the data obtained from the charcoal specimens themselves. For the
use of the latter information I am deeply obligated to Doctor Douglass.
In view of the rather limited excavations, it is desirable to point out the
trend of cultural development in the region under consideration rather
than to venture many positive conclusions. Furthermore, it is too early
yet to speak with finality concerning the complete relations of the
datahle charcoal to the ruins, for that depends upon continued research.
NO. II PUEBLO RUINS IN ARIZONA—HAURY AND HARGRAVE 7
The two ruins, the first located at Showlow and the second at Pine-
dale, Arizona, and the artifacts recovered from each, are considered
in the order named. Since both ruins are situated on privately owned
land, we are indebted to the liberality of the owners in permitting us
to work. At Showlow we wish to thank Mr. Edson Whipple who owns
the major part of the Showlow ruin, Mr. W. Whipple and Mrs. L.
Stratton, owners of the extreme south and north ends of the pueblo,
respectively, for their cooperation. To Mr. Owen Cheney we wish also
to express our appreciation for permission to conduct work in the
Pinedale ruin. A single day was spent in investigating a large ruin on
the ranch of Mr. Geo. W. Bailey, located about 15 miles northwest of
Pinedale. Although no charcoal was obtained, the ruin is a most
promising one for future archeological work. The very commendable
attitude of the owner to prevent its despoliation by pot-hunters is
largely responsible for its good condition.
The Showlow and Pinedale ruins are situated in the area drained by
Silver Creek, one of the important southern tributaries of the Little
Colorado River. The sources of this stream are in the northern slopes
of the White Mountains almost due south of Holbrook. It flows
northward past Taylor and Snowflake, Arizona, and then becomes
confluent with the Little Colorado about two miles south of Woodruff.
Its principal tributary from the west is Showlow Creek, about three
miles south of Taylor. Cottonwood and Morterson Washes are also
contributory from the west. Eastern affluents are all of a minor
character. The area drained by the Silver Creek system embraces
approximately 800 square miles. The land ranges from about 8,000
feet elevation on the south to 5,000 feet at the Little Colorado on the
-north. Vegetation varies from heavy pine timber in the high altitudes
to treeless expanses in the lower regions. Both ruins investigated lie in
the southern timbered part of the drainage area. (See map, fig. 1.)
SHOWLOW RUIN
Showlow ruin is located about 55 miles south of Holbrook in
Showlow, Navajo County, Arizona (fig. 1). It is situated on a low
elevation marginal to a narrow valley formed by Showlow Creek
which lies less than a quarter of a mile to the east. The surrounding
country is thickly wooded with western yellow pine, several species of
juniper, and oak. The proximity of the ruin to pine was one of the
determining factors in its selection for this investigation, for the pre-
cision and sensitiveness with which pine registers the passage of years
by annual rings makes it the ideal timber with which to work in build-
VOL. 82
SMITHSONIAN MISCELLANEOUS COLLECTIONS
SSS Ss
: AVON
Scale 230;000
Contour Interval 200 feet
Fic. 1—Showing location of Showlow and Pinedale ruins. From United States Geologi-
cal Survey map, Holbrook sheet, Arizona.
No. II PUEBLO RUINS IN ARIZONA—HAURY AND HARGRAVE 9
ing up a tree-ring chronology. The accessibility of pine caused much
of it to be incorporated into the roof structures of the pueblo. Hence
rich returns of wood either in the normal state or charred could be
expected.
The ruin is first mentioned in literature by Mr. A. F. Bandelier ’
who examined it briefly in April, 1883, during his extensive survey of
ruins in the Southwest. His description of the site is as follows :
.... The ruin.... is that of a communal pueblo consisting of two houses
with one circular estufa. The walls are 0.20 m. (about 8 inches) thick, built of
sandstone, and only the foundations remain. Situated on a rise above a fertile
bottom, this pueblo occupied a good position both for agriculture and defence.
In July, rgor, Dr. Walter Hough,’ Director of the Museum-Gates
Expedition, spent several days in excavation at Showlow ruin. He
refers to it, however, as the Huning ruin, as at that time and when
Bandelier was there as well, it was on the ranch of Mr. Henry Huning.
Hough, in a brief description of his work at Showlow, says:
The Huning ruin is a good example of the rectangular pueblo, showing con-
siderable skill in laying out a village. The masonry exposed during the excava-
tions is good; the material is of blocks of Carboniferous sandstone. .... It
seems probable that the pueblo was inhabited only for a short time.
The change in name from Huning to Showlow ruin, as it is here
called, seems legitimate in view of the fact that Bandelier referred to
it as the ruin at Showlow, rather than naming it after the man on whose
ranch it was located. Furthermore, its identity will be longer retained
when connected with Showlow instead of a ranch name which has
almost been forgotten.
Prior to 1901, a residence had been built on the extreme south end
of the ruin. In 1903, the land on which it is located was acquired by
Mr. Edson Whipple, who owns the major part of the pueblo at the
present time. In the same year Mr. Whipple erected his house on the
northern extremity of the ruin, and ‘subsequently put up a barn,
workshop, blacksmith shop and other sheds. A few years ago, the third
house was erected on a razed space near the center of the pueblo.
In still other sections, crumbled walls have been leveled for garden
spaces and for a roadway. Many wagon-loads of rock have been
removed for building purposes since the Showlow settlement was
started. Hough notes that “ during this process (of removing stone)
a room at the south end of the pueblo was found to contain a large
amount of charred corn, beans, etc.”” This room was later roofed
‘Bandelier, A. F., 1892, pp. 392-393.
* Hough, Walter, 1903, p. 301.
Io SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
and made into a vegetable cellar. Another room in the south end was
excavated and converted into a cistern from which the seepage has
completely saturated the rooms to the north for 50 or more feet.
(See fig. 2.) In addition to all this, the owner of Showlow ruin has
been more or less actively engaged the last 25 years in recovering
artifacts through diggings of his own. A remarkable collection of
pottery and other objects was thus accumulated which has recently
been acquired by Gila Pueblo,’ Globe, Arizona. Needless to say, the
ruin has suffered tremendously under the march of our 2oth century
civilization, but nevertheless, from the few intact rooms opened by us
were obtained the charred timbers that definitely joined the two sec-
tions of the tree-ring calendar and extended it back to A. D. 700.
The ruin is roughly rectangular in shape, the longest axis running
north and south. Our excavations did not verify Bandelier’s descrip-
tion of a two-unit pueblo, but instead, as Hough indicated, the struc-
ture is continuous. Several tiers of rooms occupy the western side,
from which three short salients extend eastward to form an E. The
intervening spaces thus created were used as plazas, the southernmost
one having a large depression. Apparently Bandelier had this in mind
when he said there was “ one circular estufa.’”’ As to the probability of
a circular ceremonial chamber in Showlow ruin comments are made
later.
Hough mentions the fact that the débris covering the ruin was so
thin that a plan was not difficult to make out. A further advantage
to him was the fact that only a single modern house existed on the site
and very little excavation and leveling had been done. While the
general plan of a ruin may be observed from surface indications of
walls and contour levels, some doubt must always remain as to precise
room arrangement and dimensions until actual excavation has been
accomplished. To identify rooms uncovered by us from previously
drawn plans,” which were based on superficial examination only, was an
impossibility. Hence, it seemed advisable to replot the ruin in rough
outline and place thereon all tests and rooms uncovered by us and to
show their relation to the present superimposed dwellings. (See fig. 2.)
In pursuing the search for charcoal it seemed most advantageous to
put down test-pits which could be abandoned if old diggings or rooms
unproductive of charcoal were encountered. Frequently the shovel
‘Mr. Gladwin, Director, has kindly permitted us to examine the collection and
to utilize any data obtained therefrom in the preparation of this report.
> Bandelier, A. F., 1892, pl. 1, fig. 38; Hough, Walter, 1903, pl. 21.
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would bring up from depths of 3 to 6 feet an old rusty horse-
shoe, baling wire, a tobacco can or a fragment of a modern glazed
dish, all evidences of a former disturbance. During the course of our
operations, 16 major tests were made. Some of these extended over a
part of a room only, whereas others included as many as three or four
adjacent dwellings.’
We explored, in whole or in part, 29 rooms; of these 22 had been
destroyed by fire and consequently contained greater or lesser quanti-
ties of the desired charred timbers. All unburned rooms were located
in the northeast section of the pueblo which had apparently been
abandoned and covered with débris while the dwellings to the west
were yet occupied. At a time subsequent to the abandonment of the
old northeast section it seems that the entire remaining inhabited part
of the pueblo was destroyed by fire, for apparently the same fate befell
room after room from the extreme north to the extreme south ends.
This wholesale burning is very probably an indication of the work of
marauders, the domestic utensils and stores of corn found in nearly
every room forming additional evidence of the fact that the occupants
of the pueblo were forced to a hasty evacuation.
Approximately 1,200 specimens of charcoal were collected at Show-
low. A large proportion of these are small fragments of roof timbers,
sections of branches, or pieces of split pine from roof members. Large
sections of timbers, less frequently found, are the more desirable, since
from them the actual cutting dates of the trees may usually be derived.
Cutting dates, furthermore, under normal circumstances signify con-
struction dates. This is especially true when several timbers in the
same room terminate with the same year. The value of cutting dates
will be recognized at once, for they not only provide the actual con-
struction time of given rooms or parts of pueblos, but they also furnish
fairly reliable dates concerning the associated artifacts.
To intimate, as we already have, that the true outside of a tree or the
first ring beneath the bark indicates the cutting date, may be assuming
too much without further explanation. Is it probable that the Pueblo
builders used the dead wood in forests in preference to felling living
trees?
The present Pueblo Indians, if unable to salvage beams from
deserted habitations, go to the forest for fresh timber when this is
not furnished by the government. Several years ago in northeastern
*On the plot of the Showlow ruin, fig. 2, the tests are designated by the
letter T and the rooms by the letter R.
NO: ET PUEBLO RUINS IN ARIZONA—HAURY AND HARGRAVE 13
Arizona, I observed an Indian bringing in a wagon-load of poles fora
new dwelling and all were freshly cut. In ancient times when trees
had to be felled with stone axes, the choice of green over dead wood
can easily be understood ; the seasoning that a dead tree gets considera-
bly increases its resistance against blows of a stone axe.
Doctor Douglass has found that large areas of dead timber, killed by
some natural cause, are rare in the Southwest. It is improbable that
the ancient Pueblo Indian was accustomed to search for such regions
which might be far from his pueblo, or to hunt isolated dead trees
in the forest when the more easily cut living trees were available on all
sides. That an occasional dead tree was utilized cannot be questioned,
but facts derived in the study of timbers from many ruins indicate the
common use of trees cut while growing. .
Several pieces of charcoal with bark intact were found in Showlow
and Pinedale ruins during our excavations. This is a good indication
of green wood, as bark on dead trees in the forest soon falls off. On
the other hand, it was customary at Pueblo Bonito * and undoubtedly
in many other pueblos to remove the bark from roofing beams before
they were used. Such a custom would readily account for the lack
of more bark-covered wood from Showlow and Pinedale.
Where a number of beams in the same room yield the same year of
cutting, it is far more plausible to suppose that the trees were cut
simultaneously while growing than that they were collected when dead.
In the latter event their ring-records would not terminate with the same
year, unless gathered in a large area where trees were killed simul-
taneously. Such regions Doctor Douglass has found to occur but
rarely.
On these grounds, we are safe in saying that the final rings in an
overwhelming majority of beams from ruins indicate the actual year
of cutting.
It would be useless to go into the detail of circumstances under
which the numerous fragments of charcoal were found, but of the
larger specimens brief record seems desirable.
Five rooms opened in test 11, in the extreme north end of the
pueblo (pl. 1, fig. 1), yielded a number of precious beam fragments.
Fortunately, this section was undisturbed except for the removal some
years previously of the shallow surface soil. Among the wood speci-
mens from room 4 of this test is one bearing the field catalogue number
HH-39. It is a beam section 7 inches in diameter and approximately
10 inches long, charred to a point at one end and internally decayed at
* Judd, N. M., September, 1925, p. 237.
2
14 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
the other, owing to incomplete burning. It rested near the northwest
corner of the room, about a foot below the surface and approximately
the same distance above the floor. By slowly working around it in a
vertical cut and wrapping the end with string as it was exposed, the
specimen was removed and turned over to Doctor Douglass who was
present to witness its removal. The initial study which the specimen
immediately received brought out its importance and historic value.
Its central ring dated A. D. 1237; its outer gave a cutting date at about
1380. The inner rings cross-dated with the last rings of Doctor
Douglass’ relative chronology; the outer agreed with his modern
ring series, extending from 1929 to A. D. 1260." The archeological
importance of this particular specimen, therefore, lay in the fact that
it definitely and convincingly joined the Douglass modern and pre-
historic ring chronologies and thus made possible the absolute dating
of this and other pre-Spanish ruins.
From room 4 there were recovered 30 other specimens, mostly
small fragments of charred pine, of which 15 have been dated. A
majority of these do not give actual cutting dates, but their broken,
outer rings end somewhere in the early 14th century. The most
recent identifiable year of the pieces giving terminal dates from this
room is 1378 which apparently marks the beginning of a short build-
ing period in the north end of the pueblo. One specimen, giving a
true cutting date of 1279, or approximately Ioo years earlier than
the other pieces from the room, is probably a fragment of timber sal-
vaged from an abandoned dwelling and re-used. Doctor Douglass has
found that such a custom still exists in the Hopi town Oraibi where
beams cut as early as the 14th and 15th centuries are in use in present-
day dwellings and kivas.
From room 2 of test 11 (pl. 1, fig. 2), a total of 243 pieces of
charcoal were obtained. Of these, 110 have been matched into the
established calendar ; 44 pieces registering cutting dates and 66 near
cuttings.’ Among the pieces are two beam sections shown im situ in
plate 2, figures 1 and 2, both giving 1378 as the cutting date. The
inverted bowl over the charred timber in plate 2, figure I, was on the
roof of the dwelling when the fire took place. Thirty-one fragments,
some possibly parts of the logs pictured, also gave 1378 as the true
outside. Eight other pieces dated 1382, and the years 1356, 1369, 1375,
and 1381 are represented by one specimen each.
1 Douglass, A. E., December, 1929, Nat. Geogr. Mag., pp. 766-767.
*On specimens giving near cutting dates the immediate outside has either
been worn or broken away.
NO. II PUEBLO RUINS IN ARIZONA—-HAURY AND HARGRAVE 15
Rooms 1 and 3 of test 11, yielded 75 fragments of charred pine;
those which have been dated give either 1378 or 1380 as the year of
cutting.
The number of wood specimens recovered from rooms 1, 2, 3, and 4,
test 11, totals 348; of these 106 have been matched into the tree-ring
calendar but do not show the true outsides of the logs, while 62 other
beam sections record the actual year of cutting. This latter group
represents possibly 20 to 25 individual trees, the majority of which
were felled either in 1378, 1380, or 1382. It seems permissible, there-
fore, to assume that the block of rooms in which these dated specimens
were found, was erected during the five-year period beginning in 1378.
Apparently construction ceased in this section with the year 1382. A
few earlier dates, namely, 1175, 1179, 1279, 1282, and 1356 are
undoubtedly re-used timbers.
Charcoal fragments from the various rooms of tests 1, 2, 3, and 12
(fig. 2) collectively, show a building period that extended, roughly,
from 1360 to 1375. Numerous other pieces fall generally into two
previous periods, ending respectively about 1204 and 1272. Unfortu-
nately few of these show terminal dates so that the above years are of
relative value only. As is set forth later, this part of the pueblo showed
an unmistakable double occupation. While some disturbance of the
débris between the two floors was apparent, much of the charcoal
coming therefrom dated in the neighborhood of the earlier period
suggested. Positive association, however, cannot be claimed, hence
1204 remains only a tentative date for the lower level of occupancy.
Tests 15 and 16, in the southeast quarter of the ruin, disclosed no
evidence of superposed dwellings. With but few exceptions, all beams
dated from this section were felled in the late 14th century. Ina
number of pieces from test 15 the final ring is 1383, indicating that the
tree was cut sometime during the winter of 1383-4. This is the
most recent cutting date found by us at Showlow and probably marks
the end of construction in the pueblo.
A comparative study of all dated charcoal specimens verifies an
anticipated condition in Showlow ruin, viz., that the last structural
additions were made at both extremities of the pueblo, the most recent
at the south end. This longitudinal expansion was apparently con-
trolled by the contour of the elevation on which the ruin rests. It seems
that maximum lateral expansion was reached first ; then a few rooms
were added at the north end filling out all available space at the point
of the elevation, and the last rooms were appended at the south end,
the only direction in which the pueblo could be easily enlarged. If the
16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
same factor entered into the construction of other pueblos, as it un-
questionably did, it might be possible to locate the most recently built
rooms of a given pueblo by an examination of its periphery with
respect to topographical surroundings.
ARCHITECTURAL FEATURES
Inasmuch as building stones have been removed from Showlow
ruin for many years and excessive alteration has taken place, it was
difficult to tell how much, if any, of the pueblo was originally more
than one story in height. The former existence of a two-story structure
was noted in but one of several tests; from conversation with Mr.
Whipple it was gathered that he rarely encountered two-story remains
during the 20-odd years in which he was actively engaged in local
excavations. With a few exceptions, therefore, the rooms of Showlow
ruin appear to have been only one story in height.
Showlow masonry is not all of the same type, due to the fact that it
is not all contemporaneous. It has already been intimated that rooms
opened in tests I, 2, 3, and 12 (pl. 4, fig. 1) evidenced two levels of
occupation. A tentative dating of A. D. 1204 has been assigned to the
lower horizon ; the upper level of occupancy dating about 1375. Walls
related to both levels were uncovered. Lower-level masonry, composed
of comparatively large, well-selected stones, and usually chinked, is
superior in workmanship to upper-level walls which are poorly
constructed of ill-chosen building stones.
A study of potsherds from Showlow ruin indicates the possibility
of an even older horizon in the northeast part of the pueblo. Walls
exposed here were formed by a basal row of large vertical slabs above
which small blocks were placed in rude, horizontal courses (pl. 3,
fig. 1). Although inferior in composition, this masonry is similar to the
lower level type exposed in tests I, 2, 3, and 12. This relationship, and
the greater antiquity suggested by potsherds for the northeast quarter,
leaves little doubt as to the early sequence of wall types.
In test 12 was uncovered an exceptionally good section of lower-
level wall, 13 feet in length by 5 feet in height. Here large blocks were
substituted for the vertical slabs of the lowest course displayed in
the northeast section of the ruin. It was then continued upward by
alternating layers of large blocks and small spalls (pl. 3, fig. 2).
In some cases the spalls were employed primarily in chinking; in
others, it is obvious they served to provide a base for the succeeding
layer of large stones. The exposed faces of these larger units were
dressed by pecking with hammer stones. This type of masonry,
NOs EE PUEBLO RUINS IN ARIZONA—HAURY AND HARGRAVE 17
although much inferior in workmanship, is reminiscent of the second
type of Pueblo Bonito stonework, described by Judd.’ That this
section of Showlow ruin was abandoned and subsequently reoccupied
is shown by the fact that walls of different masonry abut the wall just
described on a higher level.
The most noteworthy instance of superposition was found in test
2,room 1. This dwelling measured 94 by 104 feet ; near its center was
the usual slab-lined firebox (pl. 4, fig. 2). In the northern part of
the room, a foot below its floor, a second level, with related fireplace,
was encountered (pl. 5, fig. 1). Further excavation revealed the fact
that the upper firebox was built upon and almost directly above a
partly razed, earlier wall (pl. 5, fig. 2; also consult text fig. 3 for
ground plan and cross-section of this room). The north and south
walls of the upper room did not go below their related floor, but the
east and west walls extended to and rested on the lower level.
Since similar evidence of remodeling and re-use of old walls was
noted also in neighboring dwellings it may be supposed that this section
of the pueblo experienced general desertion between the two occupa-
tions. Whether or not the entire village was abandoned could not be
learned from our rather limited work. The difference in pottery from
the two levels suggests the lapse of a considerable period of time
between occupations. For information concerning the actual length
of this period, we resort to charcoal. Accepting for the moment the
tentative dating of A. D. 1204 for the lower horizon, the difference
between this date and 1375, which has been assigned to the upper
level, is 171 years. This is only an approximation because of the
readily foreseen difficulty of ascribing definite dates to culture hori-
zons. The abandonment, however, provides a convenient break in the
continuity of development, thus emphasizing the time element. That
the estimated 170-year period is correct, or nearly so, is indicated by
the discovery of pottery types at Pinedale which were developed
during this time from types present in the !ower Showlow level.
Timber associations with Pinedale types place them as post-129g0.
They in turn developed into the dominant forms of the Showlow
upper level. In this way, pottery sequence and tree-ring records were
found to support each other in their peculiar ways of recording
elapsed time.
The masonry of all the upper-level dwellings, which were occupied
to the final abandonment of the pueblo, is of nondescript form.
Unworked rocks of all sizes were laid down with no attempt whatever
* Judd, N. M., March, 1922, p. 326.
18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
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Fic. 3.—Plan and section of room 1, test 2.
INOS: EE PUEBLO RUINS IN ARIZONA—-HAURY AND HARGRAVE 19
at coursing. The walls frequently show flat curves or slight offsets,
but, whatever their appearance, they were invariably plastered with
clay.
The building stone in Showlow ruin is exclusively of buff-colored
sandstone of Carboniferous age, broken from out-cropping ledges
nearby. In all types of masonry, the walls averaged a foot in thick-
ness * and were laid up with an abundance of clay mortar.
The living rooms are approximately square; 11 by 12 feet are
measurements frequently recorded. The largest room opened by us was
rectangular, measuring 11 by 17 feet. Floors are uniformly of hard
packed clay and unbroken except for a slab-lined firebox, about a
foot square and 8 or 10 inches deep, located near the center of each
room. The rooms apparently were entered through hatchways as no
side entrances were found. In several rooms we encountered frag-
ments of large sandstone slabs perforated with a hole large enough to
admit the passage of a man. Their position in the fallen roof material
suggests that they were used as door frames. In one instance a
rounded slab, large enough to close the hatchway, was found in asso-
ciation with the fragments of such a door frame.
Details of roof construction could not be learned with any degree of
accuracy since in most cases the charred fragments of ceiling beams
were too limited and in an extremely disorderly condition. If we may
judge, however, from the impressions in burned roof clay, two methods
of covering the principal supporting beams were pursued. In one case,
reeds laid at right angles over the main beams formed the filling
material, on top of which a final layer of clay was put down. In the
second method, pine planks were substituted for the reeds. These
planks, roughly split from logs, measure from 4 to 8 inches in width
and 1 to 2 inches in thickness.
Kivas were not encountered by us, nor does Mr. Whipple, from
our description to him, recall that he uncovered any in the course
of his excavations. The depression in the south plaza was apparently
judged to be a circular estufa by Bandelier.”. An exploratory test in
this depression failed to show the flagstone floor which is almost
invariably associated with kivas. If, however, this low place actually
marks the site of a kiva, in all probability it is rectangular instead of
round. All kivas uncovered thus far in late pre-Spanish ruins in this
* Bandelier’s statement that the walls were 0.20 m. (about 8 inches) thick
was probably based on few measurements, as he did not excavate in Showlow
ruin.
* Bandelier, A. F., 1892, p. 392.
20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, 82
region’ and also in the Jeddito Valley* have been found to be of
rectangular form. The apparent lack of kivas in Showlow ruin need
not necessarily imply their entire absence as they may yet be found
in several parts of the pueblo which have not been thoroughly
examined. .
MATERIAL CULTURE
Although the ensuing consideration of cultural material may ap-
pear to be a repetition of notes presented some years ago by other
investigators, I feel that repetition is justified by the pre-Spanish
dates recently fixed for Showlow ruin. By dating the building itself,
the artifacts found within it may also be dated.
Foods.—All vegetal products recovered were found in a charred
state, obviously preventing accurate and complete description. Practi-
cally every room yielded some corn, either shelled or on the cob.
Shelled corn was ordinarily stored in large ollas, while ear corn was
piled up row upon row, as is the custom of the modern Pueblo Indians.
Such piles, when burned, generated a heat so intense as to fuse together
large masses of the corn and even vitrify the adobe wall plaster.
Although beans likewise were stored in ollas we encountered them
much less frequently than corn. A single small variety, Phasealus sp.
is represented.
Berry-yielding bushes are found abundantly in this timbered region,
especially along the water courses. It was not surprising, therefore,
to find among the stores of food stuffs, small quantities of berries.
These have been identified as the fruit of the manzanita, Arctostophy-
los pungens. This berry is about the size of a pea and has a large
crenulated seed. The meaty parts are edible either raw or cooked.
Black walnuts (Juglans major) were also uncovered. The nuts are
small but palatable, and probably were extensively used by the Indians.
Walnut thickets still fringe the banks of Showlow Creek and the
annual crop of nuts is relished by the children of the neighborhood.
Mammals.—Showlow ruin is located in what was formerly a well
stocked game area. Deer and turkeys still range the nearby forests and
+A rectangular room with flagstone floor and platform in Four-mile ruin
described by Fewkes (1904, pp. 137-138) is apparently a kiva; the Hawikuh
kiva (Hodge, F. W., 1922, pp. 9-10), and the kiva uncovered in Pinedale ruin
described in this report were rectangular.
* See the chapters on Kokopnyama and Kin Tiel, herein, by Hargrave.
°FRor the identification of vegetal products we are indebted to Prof. J. J.
Thornber, botanist, Agricultural Experiment Station, University of Arizona,
Tucson.
NO. II PUEBLO RUINS IN ARIZONA—HAURY AND HARGRAVE 20
it is only natural that their ancestors should have contributed most
generously to the quantity of mammal bones recovered during our
excavations. Lower jaw bones of dog, Canis familiaris,’ and Arizona
badger, Tasxidia laxus, were found in room 1 of test 2 and test 13,
respectively. Hough reports that he collected “bones of dog, two
species of rabbits, turkey, and deer ” in Showlow ruin.
OBJECTS OF STONE
Metates—tThe milling stones used for grinding maize and other
food stuffs are relatively numerous at Showlow. Most of these were
manufactured of a porous basaltic lava, but a few were shaped from
indurated sandstone. Two types of metates are represented in the
collection :
(a) Trough-shaped specimens with both ends open and sometimes
worn to a depth of from 4 to 6 inches. These are seldom more than
to inches wide; their sides are usually carefully shaped by pecking.
Without exception these metates were found free in the rooms, 7. e.,
independent of bins. From the position of some in the débris, it
was judged they were originally on the house tops.
(b) Flat or slightly concave metates, with grinding surface measur-
ing from 14 to 18 inches long and 10 to 12 inches wide. Mills of this
kind are usually found in mealing bins either singly or in series. In
room I, test 3, a set of three contiguous mealing bins was encountered,
and Mr. Whipple informed us that he had found during the course
of his work as many as five adjacent bins. In a few cases the lower,
embedded end of the positioned metate was narrower than the other
by several inches.
Manos.—The movable hand stones with which grinding is actually
done, are readily classifiable into two broad types, according to the
shape of the metate on which they were employed :
(a) Manos accompanying the first type of metate described are
rectangular in form with rounded corners. They measure from 7 to
10 inches long by 3 to 5 inches wide and are made of igneous rock,
rarely of granite or sandstone. Only one surface was used for grinding
and this is slightly convex lengthwise of the implement.
(b) Manos associated with the second type of metate are charac-
teristic of those usually found in Pueblo IV ruins of northern Arizona.
In cross-section they are triangular or nearly so, owing to the double-
faceted grinding surface. This distinctive feature is shown by the
*Tdentified in the Department of Biology, U. S. National Museum.
22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
fragmentary specimen in plate 6, figure 1, a, which is 8 inches long and
3 inches wide. The mano shown in plate 6, figure 1, b, is of the same
class but the grinding face is not so pronouncedly angular. Its dimen-
sions are 124 by 5 inches. The obverse sides of both manos here
pictured are rough and unfinished. Their length corresponds to the
width of the metates on which they were used. Instead of being made
of volcanic rock as are the manos of the first type, these are invariably
composed of hard sandstone.
As to the relative time of the above two forms of grinders, we can
say that in the northeast quarter and in the lower level of occupation
the trough variety only were found, while in rooms dating after about
A. D. 1375 both types were coexistent, a greater proportion of the
second type being present.
Paint metates—Still in the category of metates are several speci-
mens of rough volcanic blocks which are unmodified except for a
shallow depression in one side. In these hollows the traces of paint
materials which were ground in them are still visible. Oval handstones,
some also showing color, were no doubt used in these. The basinlike
depression suggests a rotary motion instead of one away from and
towards the body. ;
Hammerstones.—These were used for pecking, chipping, and in
countless other ways about the home, and are plentiful in Showlow
ruin. They vary from small discoidal stones weighing a few ounces to
large ones weighing a pound or more, and are usually formed of lava,
diorite, or quartzite.
Stone axes.—-The stone axes recovered by us were all short-bitted
(pl. 6, fig. I, c), none being over 6 inches in length. All, however,
were carefully finished and possessed the three-quarter groove or the
straight back which is typical of Middle Gila axes. Long-bladed axes
of the true Gila type were found by Mr. Whipple.
Polisher —While this fragmentary specimen (pl. 6, fig. 1, d@) seems
to have been used primarily in polishing arrowshafts, it also suggests
other uses. It comes from the northeast quarter of the ruin and is
made of a fine-textured basaltic lava, carefully worked even to the
smoothing of the sides and ends. Its dimensions are 25 inches in
height from the bottom to the crest of the medial shoulder and 2%
inches in width. The groove to the right of the shoulder was
apparently used in finishing arrow shafts but the opposite side does
not show this, although it is well worn. A short lateral groove cuts
the shoulder 1 inch from the unbroken end. The bottom of the
implement also indicates wear.
NO; DE PUEBLO RUINS IN ARIZONA—HAURY AND HARGRAVE 23
Polishing pebbles—Figures e and f, plate 6, figure 1, show two
typical polishing pebbles used in the surfacing of pottery. Both have
the very smooth facets which result from long use.
Perforated sandstone plates—F¥or the two objects represented in
plate 6, figure 1, g and h, we can suggest no definite use. Both are
made of sandstone not exceeding $ inch in thickness ; both are carefully
smoothed. Figure g is 64 by 53 inches and has rounded corners as well
as trimmed edges. Along one edge there are three perforations,
drilled from both sides of the plate. Specimen h is rounded, 6% inches
in longest dimension. It has but a single perforation placed 4 inch
in from the short straight edge.
These objects are not greatly dissimilar to perforated boiling stones
which resemble large pendants frequently found in ancient sites in
southern California.’ Boiling stones of this type are heated in the
fire and suspended in vessels containing liquids for cooking purposes.
Both of the specimens represented here show the effects of contact
with fire, but whether the result from their use or from the burning
of the rooms in which they were found, cannot be determined. The
custom of using boiling stones, however, is a trait quite foreign to
Pueblo culture ; hence, their use as such must be questioned.
Loom block.—A single loom block was encountered in one of the
living rooms. It is made of a soft friable sandstone and measures
about 10 inches in height by 7 inches in width. The base and one side
are entirely flat, while the rest of the block is more or less of rounded
form. In the flat side near the top is a small depression less than an
inch in depth which engaged one end of a stick or rod which formed
a part of a loom. Similar objects found in old Hopi’ and present-
day Hopi kivas are usually shaped into the forms of rectangular
blocks with the depressions in one end.
Potter’s kneading slab—A sandstone slab, not unlike those used
at Zufii* at the present time by potters for kneading clay after the
ingredients have been ground and mixed, is shown in plate 6, figure 2.
It was found with a quantity of raw clay, yellow ochre, and a collection
of sherds which had been gathered for pulverization to form tempering
material. The slab is rectangular in outline, 33 by 17 inches. Its
edges are chipped and pecked to a rough finish and the working surface
bears two shallow worn depressions which still show traces of clay.
1Handbook of the American Indian, Bull. 30, Bur. Amer. Ethnol., Pt. I,
pp. 126-127.
* Hargrave, Lyndon L., p. 108 herein.
*Guthe, C. E., 1925, footnote p. 20.
24. SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
Chipped implements—Relatively few chipped stone objects were
recovered. Projectile points are both plain and tanged (see fig. 13
for representative specimens from Pinedale), and a few have serrated
edges. Knives and scrapers of chert and obsidian flakes show com-
paratively little secondary chipping. Figure 7, plate 6, figure I, is a
chert knife on which the cutting edge has been slightly retouched.
The large implement shown in plate 6, figure 1, 7, is lanceolate in
shape, 74 inches long and 3} inches wide. It is crudely chipped from a
spall struck from a lava block. Near the center of the object on each
edge, are shallow notches which probably aided in hafting it to a
handle.
Obsidian cache.—F rom the northwest corner of room 1, test 15, was
taken a cache of 17 obsidian nodules which were probably intended to
supply the material for projectile points. The nodules vary from
} to 14 inches in diameter. They are somewhat angular and covered
with a thin veneer of opalitic substance.
OBJECTS OF BONE AND HORN
Awls.—Specimens a, b, c, and d, figure 4, are representative of the
bone awls from Showlow ruin. Awls a and d were split from large
bones after longitudinal groovings had been made and then trimmed
down, while b and c were formed of natural bones with little modi-
fication. Incising was noted on several awls, probably representing
crude attempts at decoration.
Dagger (?).—A large broken implement (fig. 4, e) is tentatively
identified as a dagger. The fragment shown is 6 inches long with an
inch or more broken from each end. A comparison of its size with
normal awls makes it quite evident that it was intended to be used in
some other way, the most logical use being that of a dagger. In
his excavations at Turkey Hill Pueblo, near Flagstaff, Dr. Byron
Cummings recovered excellent examples of these. In one instance
several were found at the waist of a burial as though they had been
suspended from or tucked beneath a girdle. Hodge’ also pictures
quite similar implements from Hawikuh, although he lists them as
awls.
Incised bone.—Figure 4, f is a section of a mammal leg bone 7 inches
long and 1 inch in diameter from which a portion has been severed
by a circumambient incision. Below the cut end is another scoring
made in preparation for the removal of a short section.
* Hodge, F. W., 1920, pls. 10, 11, 12.
\
{
|
|
i}
NO] Lt PUEBLO RUINS IN ARIZONA—-HAURY AND HARGRAVE 25
Horn implement.—This specimen (fig. 4, g) consists of a prong of
a deer antler 6 inches in length. It is unaltered except for slight wear
at the blunt point and several hackings at the base apparently made
with a flint tool in the process of its removal from the major antler.
Panga
ae
Sa ~
~————- >
Fic. 4.—Bone tools from Showlow ruin. (% actual size.)
POTTERY
The Silver Creek drainage (see map, fig. 1) is a region which in past
ages was on the periphery of several important southwestern sub-
culture areas. To the north lies the center of Hopi development with
its very distinctive yellow decorated pottery ; to the east, the old Zuni
pueblos with their variable types of glaze-painted vessels; to the
southeast, the Upper Gila; and to the south, the Middle Gila culture
areas, each with characteristic pottery. Ceramic types from all of these
areas are represented in Showlow ruin. Thus, a treatment of the
pottery of a region, especially with reference to its sequential occur-
rence, where a considerable mixing of local and foreign types has
taken place, is a problem of some complexity and one that can be
solved only by prolonged and careful research.
Although the limited nature of our excavations does not warrant
the drawing of fine distinctions between the several recognized culture
26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
horizons at the present time, the data procured from stratigraphic
evidence at Showlow should prove of interest to archeologists. It is
also possible here to include some of the invaluable results which have
been obtained from the study of tree rings displayed in charcoal. The
reliable stratigraphical method of studying prehistory bids fair to be
surpassed by this newer means of establishing succession, which
embraces the association of artifacts with datable timber. Results
obtained from stratigraphy will not only be checked but will be supple-
mented with the precision that only accurate dates can give. However,
precaution must be exercised in its use, for the associated circumstances
are of utmost importance. For instance, the finding of newly developed
ceramic types in very old rooms, or in rooms where structural timbers
were re-used, would be certain to give misleading results unless
corroborated by other finds.
At Showlow ruin, the association of pottery with datable charcoal
was relied upon to a certain extent in an attempt to establish a
sequence. This is true especially of the pottery from the more recently
occupied section of the pueblo. In arriving at ceramic correlations
between Showlow and Pinedale ruins, charcoal proved to be of in-
valuable assistance.
Hough’ observed that potsherds on the surface at Showlow ruin
were exceedingly scarce. This reminded him of the practice of modern
Zui potters, who gather from the rubbish heaps of old ruins sherds for
pulverization to be used as tempering material in clays for new vessels.
That this practice obtained in more remote times is well known to all
students of southwestern prehistory. We found it also at Showlow.
In the corner of room 1, test 12, was encountered a polychrome-on-
red bowl filled to the brim with sherds which had apparently been
picked up on the trash pile. There is little question but that these
had been set aside for future grinding. Almost all types of pottery
known at the pueblo up to the time of collection were represented.
This has a slight bearing on the technology to which we refer later.
Not far from the sherd collection and leaning against the wall was the
kneading slab pictured in plate 6, figure 2. The several slight depres-
sions in the working surface were caused by much mixing of ground
sherds with clay and the kneading of the paste to obtain the proper
consistency. Behind the slab was stored a quantity of raw clay of
gray color just as it had been mined, and yellow ochre for use as
coloring matter. Worthy of mention too, are several fragments of
unfired vessels which were recovered. These clearly show the gray
* Hough, Walter, 1903, p. 30T.
NO. II PUEBLO RUINS IN ARIZONA—HAURY AND HARGRAVE 27
paste with the admixture of pulverized sherd tempering (see pl. 10,
fig. 2). Exterior and interior surfaces of the sherds bore the yellow
slip applied in wash form, which, upon firing turns to a rich red.
Other tools such as scrapers, formed of broken fragments of pottery,
and pebble polishers also show the similarity of ancient and modern
methods of making pottery. While styles of decoration and vessel
forms are continually undergoing changes, there seems to have been
no great departure in method since the close of the 14th century, the
approximate age of the objects found.
Two sharply defined levels of occupation were found to exist in the
Showlow ruin. All of the rooms opened in tests 1, 2, 3, and 12 had
two floor levels (see fig. 3 for plan of room I, test 2). Pottery types
from all lower level rooms were in entire agreement and a corres-
pondence of sherds gathered in upper level rooms was also noted.
Comparisons of the sherds from the two levels, however, introduced
the fact that there was a considerable difference of time between the
two occupations. This difference we have provisionally placed at about
170 years, based upon data derived from datable charcoal.
Lower level types—Potsherds from the lower level include the
following types: Black-on-white, black-on-red, an orange-red ware
decorated in both black and white paint, corrugated, and a small
amount of intrusive material.
The black-on-white sherds roughly fall into two groups: (a) those
showing definite Chaco Canyon affinity, and (b) those obviously
related to the black-on-white of the now known Upper Gila culture
area. Chaco-like sherds (pl. 7, fig. 1, nos. I, 2, 3, 4, 5) are fragments
of bowls with direct rims having a chalky white slip and rough
exteriors. The decoration is in dull black paint and the designs, while
they bear certain similarities to Chaco black-on-white, are neverthe-
less somewhat different. In the hatched elements, for instance, the
framing lines are of the same width as the filling lines. The rims,
however, are usually painted black and tapered, both typical Chaco
features.
Sherds bearing similarities to vessels better known from the south
and southeast are preponderantly of ollas and smaller, full-bodied
vessels. The paint is dull black, but in rare cases it has a lustrous
silvery appearance. The designs (pl. 7, fig. 1, nos. 6, 7, 8, 9, 10, II,
12) consist of alternating solid and hatched elements, opposed stepped
figures and interlocking elements. The canteen and fragmentary
bowl, figured in plate 7, figure 2, were found in a firebox in the lower
level over which the later occupants had erected a wall. The canteen is
28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
6 inches in horizontal diameter and the orifice formed by a small
vertical neck is but 3 inch in diameter. The lugs are squared and
the sides of the vessel are depressed immediately below the lugs
to allow more space for the passage of a cord. The black paint is
dull, applied in an all-over design except for a small circular area
at the bottom. The rim is edged with black dots. Canteens of a
similar type have also been found on the Gila River in the region of
San Carlos and in the Tonto Basin."
The bowl is not round but elongated ; apparently it was compressed
before or during the firing process. Its greatest diameter is 10 inches,
The heavy walls terminate in a flattened rim and a slightly over-turned
lip. The background is gray rather than a dead white as in the
canteen, and the design is in a flat black paint.
The Chaco-like black-on-white ware is most certainly the older of
the two black-on-white types from the lower level. The terminal date
for Pueblo Bonito given by Doctor Douglass * is A. D. 1127, but it
reached its heyday in 1067. Hence, possibly by 1067, certainly before
1127, the Chaco influence was extended southwestward as far as the
Silver Creek drainage. Its presence should therefore be expected in
sites which antedate the lower Showlow level. That this condition
actually exists was clearly demonstrated by Roberts during the
summer of 1929 in his work on a pit house and early pueblo site on
the old Long H ranch, 20 miles north of St. Johns. He reports * that
the pottery from both the pit houses and the surface pueblo is distinctly
related to the Chaco Canyon cultures. In the Showlow lower level, the
pottery of Chaco affinity was decidedly on the wane and pottery
suggesting Upper Gila influence was springing into prominence. By
about 1290, as we found at Pinedale, the former had entirely lapsed
and the latter, whose exact relation needs yet to be established, was
strongly reflected in the dominant black-on-white ware. In all
probability the culture represented by it, first in the lower Showlow
level and later more strongly at Pinedale, is subordinate in this region
and an extension from the parent stock to the southeast.
Fragments of black-on-red pottery are extremely rare in lower
level débris. Rim sherds of a number of individual bowls (fig. 5),
however, illustrate the existence of a fairly uniform type. Several
of the sherds were found built into the walls of the dwellings of the
first occupation. This may signify that a still older horizon to which
* These specimens are to be found in the Arizona State Museum, Tucson.
* Douglass, A. E., December, 1920, p. 767.
* By personal letter of January 27, 1930.
NO. II PUEBLO RUINS IN ARIZONA—HAURY AND HARGRAVE 29
these sherds belong is to be found elsewhere in the ruin. Features
which correspond in all sherds are: A rather coarse-textured paste is
used which burned red with a dark core ; tempering consists of crushed
rock of light color and possibly a small amount of pulverized pot-
sherds; the slip is of a deep red color applied both inside and outside
of bowls; vessel forms consist of bowls only, and these are unusually
deep * with slightly incurved rims and rounded or squarish lips; the
Fic. 5.—Black-on-red sherds from Showlow ruin which probably antedate the
lower horizon. (4 actual size. )
decoration is in a thin black paint’ which has a slaty appearance and in
some places is obliterated. Brush-work is very crude; lines were over-
carried and variable in width. A narrow black line below the rim on
the inner surface is characteristic of all the fragments in the collection.
The designs consist of rudimentary hatching, dentate elements,
terraces, and triangular figures. These are separated from the rim-
line by a narrow unpainted zone. There is no exterior decoration.
If the inference is correct that this type is aberrant to the lower
horizon, belonging to an older culture stratum, its age is unquestionably
pre-1200. While the provenance of this ware is not clear, I am of
‘The Showlow collection in the possession of Gila Pueblo, Globe, has several
complete examples.
3
30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 282
the opinion that it is parental to the later orange-red decorated
ware which bears the white exterior and black interior decorations.
This latter form is nearly as abundant as the black-on-white in the
refuse of the first occupation. It has commonly been referred to as
Little Colorado polychrome and was undoubtedly the most ubiquitous
pottery known to the Southwest. In this instance it is an orange-red
phase instead of the deeper red which it usually takes. Its presence
was one of the guides used by Doctor Douglass in selecting sites
which would provide the gap material for his tree-ring chronology.
The paste is of a gray color, which shades to a light gray towards the
surfaces of the vessels, Tempering material of coarsely pulverized
sherds is abundant. The slip varies from a dark to a light orange-red
color and is unusually thick. By actual measurement it was found to
be one-twentieth of an inch thick in one case. Vessel shapes are
exclusively bowls with pronounced incurving rims. Interior orna-
mentation is in dull black pigment (pl. 8, no. 1) consisting mostly of
repeated interlocking or opposed solid and hatched figures. These
are confined in a broad horizontal zone by two narrow framing lines,
one below the rim and the second towards the center. The center of
the bowl is thus left undecorated. Exterior decoration is made up of
broad, rudely drawn, white stepped lines or terraced figures (pl. 8,
nos. 2, 3, 4). These are also limited to a horizontal zone below the
rim but never as wide as the one on the interior. A very small number
of pottery fragments of this type show the use of white as a comple-
mentary color to the black on bowl interiors, and, conversely, the use
of black on exteriors. These are believed to be local prototypes of
an upper level pottery and will therefore be considered later.
Corrugated ware of the first horizon appears to consist almost
solely of large ollas. The paste is gray to nearly black, quartz
tempered and crumbles easily. The coils are not very fine and the
indentations are shallow (pl. 9, fig. 1). Attempts at decoration
by leaving a series of coils unindented as in 5 and 7, or other tech-
niques, are seldom met with. Fragments of finely corrugated bowls
bearing white exterior designs (pl. 9, fig. I, nos. 4, 6, 10) and
blackened interiors were rare in the lower level but occurred fre-
quently in the northeast section of the ruin which we believe to be
older still than the first horizon. This form is possibly best known
from the Upper Gila. Its abundance at Showlow and at Pottery Hill *
suggests local manufacture and thus may indicate a direct link between
the southeastern sub-culture.
‘Hough, Walter, 1903, p. 300.
NO. II PUEBLO RUINS IN ARIZONA——-HAURY AND HARGRAVE 31
Trade pieces are represented by a few scattered fragments of Pinto
polychrome bowls,’ 7. e., black-on-white interiors and undecorated red
or reddish-brown exteriors.
Upper level pottery—Turning now to the upper level pottery we
find that radical changes took place during the interval of time
represented by the abandonment.’ Lower level forms had ceased to
exist; new forms had been invented and the area was being pene-
trated by trade vessels from several adjacent regions. This second
horizon, to which we can give the general dating of A. D. 1375,
existed through to the devastation of the village by fire.
In the upper level, black-on-white is practically absent. The few
scattered sherds probably represent survivals from the preceding
period when that form was in vogue. Black-on-red is entirely lacking
as is also the form of Little Colorado polychrome here described.
Four-mile polychrome.—In speaking of the artifacts at Showlow,
Bandelier remarks that he found “ nothing unusual except the pottery,
which resembles that at Tule.* There are specimens with glossy
decorative lines, but the glaze is more carefully applied, the designs
more perfectly executed... .”
In 1897, Fewkes conducted excavations in Four-mile ruin,’ situ-
ated 4 miles from Snowflake, and about 25 miles west of Taylor,
Arizona. The predominating pottery type recovered by him was a
“redware with black decorations having a margin of white.”
The pottery thus briefly characterized by both Bandelier and Fewkes
was found in abundance during our work at Showlow. It formed
approximately 75 per cent of all the pottery during the time of its domi-
nance. This type is not new to those familiar with the archeological
literature of the Little Colorado drainage. Fewkes gives excellent
color plates ° of it; other later investigators in that field also picture it.
As far as we have been able to ascertain, however, little has been
done toward a careful description of the ware or an analysis of its
derivation and relationship to other types. Likewise it lacks a suitable
name to accord it the distinction which it merits.
* The Medallion, 1930, pp. 4-5, pl. II.
* By the abandonment we refer only to the rather restricted section where
double occupation was found to exist. It is likely that the pueblo was occupied
continuously but with small local movements within the structure.
* Located about 14 miles east of St. Johns.
*Fewkes, J. W., 1904, pp. 136-164.
*Idem, pls. XXI, XXII, XXIII, XXIV, XXV, XXVI, XL, XLVII.
32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
Upon the suggestion of Dr. A. V. Kidder, at the 1929 Archeologi-
cal Conference at Pecos, New Mexico, we here refer to this form
as “ Four-mile polychrome.” Fewkes’ Four-mile ruin seems to be the
type site for Four-mile polychrome. There it is found in great abun-
dance and in what appears to be its most highly developed stages.
_ Furthermore, Four-mile ruin is the approximate focus of the known
distribution of this pottery. Its distribution is roughly conterminous
with the area drained by Silver Creek and its affluents. Notable sites
not included in this drainage but where Four-mile polychrome is
found in some abundance are Forestdale, Chavez Pass, Homolobi, and
Chevlon. All of these, however, are peripheral to the drainage area in
question. Trade pieces have been found as far south as Bylas,
Arizona, on the Gila River; in the Tonto Basin; at Casa Grande,
and as far north as the old Hopi ruins in the Jeddito Valley.
Technology.—A study of the technology of Four-mile polychrome
reveals the traits on which the creation of this type is based. The
constancy with which some of the characteristics occur is worthy of
note.
The paste is light gray in color, usually merging into a dark gray core
towards the inner part of the vessel walls, and sometimes appearing
reddish outwards. The paste fires to a hardness sufficient to give breaks
with fairly sharp and smooth edges. Tempering material consists of
ground-up potsherds and small rounded grains of sand, these in-
gredients occurring in about equal proportions. There is also a
relatively small amount of dark particles, apparently ground basalt.
In plate 10, figure 1, is shown a quantity of tempering, enlarged four
times, which was removed from a sherd of an unfired vessel. Small
angular bodies of white slipped vessels can be detected. Plate Io,
figure 2, shows an unfired sherd (four times normal size) containing
a sizable fragment (circled) of a former red-slipped vessel. As
may be supposed, the addition of tempering derived from pulverized
sherds of various sorts would be apt to introduce a considerable
variety of extraneous inorganic substances.’ The quartz grains as well
as particles which appear to be crushed basalt were probably used as
tempering material in previous vessels.
The slip is an even red color unless over-fired, when it turns to
dark brown or almost black. Before firing, the slip is yellow, being
made of the yellow limonite. This is to be seen in the upper half
The sherd collection, which has already been noted as having been made for
the purpose of grinding, is ample evidence of this.
>
NO: DT PUEBLO RUINS IN ARIZONA—HAURY AND HARGRAVE 33
of the unfired sherd in plate 10, figure 2. The slip is thin, fairly well
pebble-polished and rather soft, so much so in fact, that on the bottoms
- of vessels and other surfaces exposed to wear, it has been entirely
worn away. In a good many cases, the surface is seamed with minute
cracks.
Shapes ——Vessel forms of Four-mile polychrome are almost ex-
clusively ollas and bowls, the latter predominating. Several olla shapes
are noted:
(a) The largest ollas (fig. 6, a) are shaped somewhat like the
polychrome ollas from the Middle Gila. The body is compressed
vertically and the neck rises at a steep angle to form the mouth
Se
ACCC
c
d OS NEG! aie eel hs wa
Fic. 6— Vessel forms of Four-mile polychrome.
which is smallest at the rim. The lip is direct. As far as we know,
only fragments of these exist.
(b) The most prevalent form of olla is much like the first excepting
that the body is usually more rounded and the neck rises more abruptly
(fig. 6, b and pl. 11). The neck is from I to 2 inches high and the
lip always direct. The horizontal diameter seldom exceeds 12 inches.
(c) The third type (fig. 6, c) consists of small round-bodied ollas
whose diameters vary from 5 to 8 inches. The neck is short or absent
and the lip terminates in a decided flare. The mouth is larger in
proportion to its size than was noted in the foregoing types.
Bowls (fig. 6, d) vary from 7 to 14 inches in diameter and are
always less than one-half the diameter in depth. The rims are gener-
ally gently incurved and carried to the lip with or without thinning of
34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
the vessel wall. The lips show a variety of treatment (fig. 6, e to k)
in rounded, squarish, and bevelled forms. Bowls with rounded lips
appear to be the rule.
Pigments—tThe paints employed in carrying out the designs on
Four-mile polychrome are black and white. The black paint is basically
a lead glaze. Where thin it usually has a metallic luster and where
thickly applied it is apt to be dull, sometimes vitreous and showing
small granular bodies. It bit deeply into the slip, giving the black
patterns permanency. The apparently gritty texture of the paint made
it more difficult to manipulate than a free-flowing pigment, hence the
brushwork does not show the clear-cut edges that it otherwise might
have. Brush dips are clearly visible.
Quantitative analysis" of this glaze paint show that the main
constituents are lead, copper, and usually some manganese. As one
would expect in primitive pottery, where the pigments were not
mixed by exact formula, the ingredients vary in proportion in the
paint on different vessels. This fact possibly accounts for the slight
difference in appearance of the paint. It is also likely that the length
of time the pottery was subjected to fire and the intensity of the fire
affected the final product. Individual paint determinations showed
that lead and copper were almost always present in considerable
amounts. A composite test of the paint on 12 sherds of Four-mile
polychrome indicated a ratio of 1:2+ of lead to copper and the
presence of a negligible amount of manganese. If the lead was in the
form of an oxide when used, it would, upon heating, produce a silicate
or glaze of light brown to yellow color. This silicate would promote
the fusing of the copper and manganese compounds which impart the
black color. The common occurrence of blue and green copper carbon-
ates in ruins suggests that the copper element was added to the paint
mixture in the form of a powdered carbonate. Heating would convert
the carbonates to an oxide of copper of black color, which, being less
fusible than the lead component, would tend to remain as the gritty
particles already mentioned. The small amount of manganese present
could have been combined with either the lead or copper ores when
mined. Sodium salts which give a glaze similar to that of lead may
also be present in the paint.
The white is a soft chalky paint that can be readily scratched off
with a knife. As a result the white parts of the designs are often
The chemical tests of glaze paints on pottery from Showlow and Pinedale
pueblos were kindly made by Mr. F. G. Hawley, Chief Chemist, International
Smelter, Miama, Arizona.
NO. II PUEBLO RUINS IN ARIZONA—-HAURY AND HARGRAVE 35
partly obliterated. While the white pigment may be considered to be
of secondary importance in its use as an outliner for the heavier
black lines, it was the first to be applied to the vessel surfaces in
blocking out the designs and later supplemented with black.
Designs.—In the ornamentation of Four-mile polychrome, a stylistic
divergence is noted from the usual trend of Southwestern pottery.
The conventional repetition of elements in orderly zones and the
involved interlocking elements noted on some of the lower level pottery
are devices of the past. Instead, the field of design is broken up into
irregular units and treated with a freedom and boldness previously
unknown. In the latest forms, pure geometrical figures give way to
conventional adaptations of life forms.’
Olla decorations —The several types of ollas are decorated much
in the same manner. The neck and a small part of the upper-body are
covered with a white slip which is carried well down on the inside of
the neck. The white is seldom pebble-polished, hence rough and
cracked. The remainder of the body is covered with a red slip and
fairly well polished. There are two zones of decoration: (a) the white
upper part and neck, and (b) from the lower edge of-the white to a
point not far below the maximum diameter of the vessel. The designs
of the two fields are entirely different. On the white field the elements
in black are simple and very often used independently. Crosses (fig.
7,a and b), “ turkey tracks ” (c), dots, stepped and paired lines (d, e,
f, and g) are favorites. Elements represented in figure 7, h, 1, and j,
are appended to a continuous band placed just below the rim. The sec-
ond field of design with patterns in black and white on red is a broad
horizontal band bordered above and below with heavy black lines (pl.
11, a, b, c, and d). It is divided into panels by broad black lines or
otherwise divided off into recurrent units. White is used as a
complementary color to the black. The design elements are essentially
the same as those occurring in bowls, which are described later.
Bowls——Decoration was applied to both interiors and exteriors of
bowls, the principal design being on the inside. A very constant
feature of inner decoration is a black band, 4+ to $ inch in width,
placed immediately below the rim. It is invariably bordered by a
narrow white line on the lower side only. In the majority of cases
this band completely encircles the bowl. Where interruptions were
made, the white lines are continued around the ends of the black band
and extended upwards to the rim. The persistent occurrence of
this rim-band, even in bowls where the rest of the interior was left
*For color plates see Fewkes, J. W., 1904, pls. XXV and XXVI.
36 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
unpainted, and a corresponding pair of lines on the rim exterior, may
be considered as helpful marks of identification.
Separated from the black and white border by an undecorated zone
from 4 to 2 inches in width is the design area proper. It is nor-
mally of circular form. Only in extreme cases are the decorative
elements appended directly to the black band at the rim. For an
adequate understanding of these interior designs, we must resort to
illustrations. In plate 12, a, the field of decoration is quadrate. Op-
posing quarters are paired and treated similarly. The whole is
encircled by a heavy black border, outlined in white. In b, the field
> Sa
vee ~~~ [|
eo
Fic. 7.—Design elements on Four-mile polychrome olla necks.
is divided into two opposing sectors which are connected by a
medial line joining extensions of the arcs on opposite sides of the
sectors. The designs in each of the latter are identical. The bilateral
symmetry exhibited in these two bowls seems to be general in the
early phases of Four-mile polychrome. In later forms the central
field is treated with spirals, triangles, rectangles, sectors, or life
forms, all parts of the design being connected, thus forming a con-
tinuous though not balanced arrangement. The design represented in
figure 10, c, is typical of this feature.” The bowl figured in plate 12, c,
is ornamented with a horizontal band 2 inches in width placed
"For additional examples see Fewkes, 1904, pls. XL, XLVIIa.
NO. II PUEBLO RUINS IN ARIZONA—HAURY AND HARGRAVE 6i7/
3 inches below the rim. Four white lines dotted at regular intervals
with paired rows of black spots are diametrically drawn across the
central bare area. Specimen d is an excellent example of the unique-
ness of design found in many Four-mile polychrome bowls. From the
central black spot, a number of white lines terminating in black dots
radiate rimwards. It reminds one of a child’s drawing of the sun.
It will be noted from the preceding vessels shown that rectilinear and
curvilinear styles are used with equal skill, and that transitions from
one to the other are made within the same vessel.
Bowl exteriors are treated in a very uniform manner. The orna-
mentation is confined to a horizontal zone beginning immediately
below the rim and extending downwards for from 14 to 3 inches. This
zone is enclosed by two parallel, heavy black lines. The upper one 1s
outlined in white on the lower side only, while the lower one is framed
on both sides. The designs introduced into the zone are in white except
in the more elaborate instances where black also appears. Figure 8,
no. I, shows a simple continuous line pattern, and in nos. 2 and 3
concentric parallelograms and triangles are represented. Modifications
of the fret or ‘“‘ dentiform ”’ figures * are characteristic as in nos. 4 and
5. The zone is often panelled by one or more diagonal or vertical lines,
or merely by the enclosing lines as in no. 5. Elements are usually
repeated in each panel but sometimes with slight variations. In no. 6
we see a more elaborate treatment. Black diagonal, terraced, and
plain vertical elements connect the top and bottom borders. To these
further embellishment is added in white.
Elements of design.—The units of decoration in the main are not
dissimilar to those generally employed in Southwestern pottery.
The distinction of Four-mile polychrome designs, however, is based on
the very singular treatment of the elements, a freedom from the con-
ventional equating of the field of design. Obviously this paved the
way for greater elasticity in the expression of the relatively few ele-
ments into innumerable variations.
Of the elements, the following appear to be of primary importance :
(a) triangles, (b) terraced or stepped figures, (c) spirals, and
(d) frets.
In figure 9, nos. 1 and 2, triangles are shown as they occur in their
simplest forms, while in no, 3 a triangle is embellished in white, and in
no. 4 may be seen a common modification of the triangle combined
with another element. Frequently in the center of this geometric
figure a small rectangular area is left unpainted, which is then either
* Spier, L., 1919, pp. 367-8.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
RED
Fic. 8.—Typical bowl exterior decorations of Four-mile polychrome.
NO. II PUEBLO RUINS IN ARIZONA—-HAURY AND HARGRAVE 39
Fic. 9—Design elements of Four-mile polychrome pottery.
40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
dotted in the center or outlined in white. Terraced elements (nos.
5, 6, and 7) are characteristic. They are used singly, in pairs, or in
complex patterns and, less often, in opposed positions as in nos. 8 and
9. Spirals fall into two classes: (a) large curvilinear forms consti-
tuting a unit of the entire design (fig. 10, ¢) which are reminiscent of
similar devices occurring commonly in Upper Gila black-on-white and
in some Little Colorado polychrome but without the solid interlocking
element; (b) small angled spirals used as filling elements (fig. 9,
nos. 10, 11, and 12). The latter is an uncommon form of the inter-
locked spiral. Variant forms of the fret (fig. 9, nos. 13 and 14) also
(fig. 8, nos. 4, 5, and 6) predominate on bowl exteriors. Combinations
of the fret with the above listed elements are frequently used in bowl-
interior and olla patterns.
Structure of designs—The initial step in applying the decoration
was to block out the area to be covered with narrow white framing
lines (fig. 10, a). It is evident that the artist had a fair conception of
the combination of elements that were to be used before the first
pigment was applied. Where panels were to be filled with white only,
as in the case represented, that was done before the next color was
taken up. .
The second step consisted of applying the heavy black lines inside
the white borders (fig. 10, b). The width of these varies somewhat.
If drawn within areas later to be filled with other elements, they are
seldom more than twice as heavy as the white, but where the white
framers connected parts or surrounded the design, the black lines are
heavier.
In the final stage, all bordered areas are filled with coarse hatching
(fig. 10, ¢) which ordinarily parallels the longest side of the block.
A variant of hatching is to be noted in the spiral where sets of parallel
lines are offset by appended dots. Heavy stepped lines are also used
with hatching as fillers.
The dating of Four-mile polychrome is made possible by a number
of cases of its association with timbers that yielded cutting dates. The
last big construction period in the Showlow ruin extended over
approximately 25 years prior to 1383. The latter date is the most
recent cutting date found in over 1200 specimens gathered from
various parts of the pueblo. It appears, then, that no major building
was going on after 1383. In 15 rooms belonging to the period in
question, Four-mile polychrome was invariably present. This means
that the pottery found in a room constructed in, say, 1375, would likely
NO. II PUEBLO RUINS IN ARIZONA—-HAURY AND HARGRAVE 4!
Fic. 10.—Steps in the application of a design in a Four-mile polychrome bowl.
42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
be post-1375, provided the associated pottery was not exotic but repre-
sentative of local types at that time. The finding of Four-mile poly-
chrome in these 15 rooms places its time unquestionably as shortly
after 1383, although it was present in rooms dating 1375 and may even
have been in existence 25 or 30 years earlier. That it had not been
developed by A. D. 1300 is shown by the results of excavations at
Pinedale, where an antecedent form was in existence at that time
and Four-mile polychrome was absent. Nor do we know exactly
how long it survived after 1383. Showlow ruin was probably aban-
doned at the close of the 14th or early in the 15th century just before
the Hopi Jeddito black-on-yellow penetrated the region in any ap-
preciable amount. In addition to its characteristic polychrome,’ Four-
mile ruin contains an abundance of the Hopi yellow (Jeddito black-on-
yellow) which would place the abandonment of that site after Show-
low. But how long after, we are not ready to say. The absence of
Sikyatki polychrome at Four-mile ruin implies its abandonment prior
tora 5ocAe Ds
Plain redware.—In the later period at Showlow plain red bowls
and ollas were made and used. Sherds were not found in the lower
deposits.
A nearly complete, plain red olla seems representative as far as our
sherd collections show. Its paste is gray and tempered with coarse
sand ; its outer surface bears a red slip. Firing clouds are common on
these plain red ollas. The shape is similar to the Four-mile poly-
chrome ollas except that the upper body is flatter and the neck more
cylindrical. Plain red bowls are technologically the same as Four-
mile polychrome bowls except that the decorations are omitted.
Several nearly complete specimens and numerous sherds indicate
that they were seldom over 8 inches in diameter.
Corrugated. —In the upper Showlow level, corrugated pottery occurs
only as ollas. The paste is gray, quartz tempered, and crumbly. Cor-
rugations and indentations are much the same as in vessels of the
older horizon (pl. 9, fig. 2, nos. 1 and 2), but a new treatment, not
found in the lower rooms, has come into general use. In this the coils
were so manipulated as to give the surface shallow horizontal flutings
or ribs (pl. 9, fig. 2, nos. 3, 4, 5, and 6).
The finely corrugated vessels bearing exterior decoration are absent
in this level.
“Type specimens of Four-mile polychrome vessels are to be seen in the U. S.
National Museum; Gila Pueblo, Globe, Arizona; Arizona State Museum,
Tucson; and the Museum of Northern Arizona, Flagstaff.
NO.. LT PUEBLO RUINS IN ARIZONA—HAURY AND HARGRAVE 43
MISCELLANEOUS POTTERY OBJECTS
Parching plates ( ?).—Utensils formed of fragments of broken clay
vessels were occasionally encountered in the diggings. From room 2,
test 16, two small plate-like vessels were collected. The larger of the
two (pl. 13, fig. 1, a@) is 74 inches in diameter and 14 inches deep.
It is made from the bottom part of what appears to have been a large
smooth red olla. The edge, although irregular, has been ground down
to remove all rough places. The smaller plate (pl. 13, fig. 1, b) is
53 inches in diameter and made from the base of a corrugated olla.
Its edge is not finished as in the former specimen. Fragments of a
larger but similar vessel came from room 2, test 3. It is approximately
12 inches in diameter and 4 inches deep, and at one time formed the
bottom of a thin, plain red olla.
As to their use, it is not unlikely that they served as parching plates
for corn. The dark irregular centers and light edges of both plates
pictured show that they were used over hot coals since their manu-
facture from the original vessels. That they were also put to other
uses, such as containers for pigments, etc., is suggested by an incrusta-
tion of fine clay-like material on the inner surface and on the under
side along the edge of specimen a. This substance is apparently the
same as the material used for red slips on vessels.
Pottery scrapers.—Several objects made of potsherds and used in
scraping the surfaces of unfinished clay vessels were recovered in
Showlow ruin. The nature of the vessel on which a scraper was to be
used apparently governed its size and shape. One specimen is
ovate, measuring 4% by 3 inches; another is of semi-circular form
and considerably smaller. The wearing on all, however, indicates that
the convex surfaces advanced in the scraping stroke.
Intrusive pottery types——In rooms from which the latest cutting
dates were recovered, there appeared occasional fragments of vessels
obviously foreign to the Showlow district. Some of these undoubtedly
came from the Zuni region, not a great distance to the east. And
Middle Gila pottery" is represented by a very few bow! sherds
showing the characteristic red on the outside and the black-on-white
internal decoration.
The following Zuni types, as established by Hodge in his work at
Hawikuh * are represented in our collections from the upper Showlow
level: Type C, by a sherd of a full-bodied vessel with black glaze on
* Gladwin, H. S., September, 1928, p. 20.
* Hodge, F. W., 1923, p. 20.
44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
a white slip and by a bowl fragment with greenish glaze on a cream
slip; Type D, by a cream-colored bowl fragment ornamented with
green glaze and red, a non-glaze color. Hodge’s Period E is probably
represented by the fragmentary bowl pictured in plate 13, figure 2, a,
in which the designs are rudely executed in a dark buff, non-glaze
color on a light buff background. A similar bowl, with black glaze
decoration, is to be found in the Showlow collection now at Gila
Pueblo, Globe, Arizona. All of these Hawikuh types came in well
towards the end of the occupation of Showlow ruin.
Jeddito black-on-yellow, the early forms of which appear to have
come into use soon after 1300 in the great ruins of -Jeddito Valley, was
almost unknown at Showlow as late as 1383. A single sherd of this
type was found in room 2, test 2, with timber that dated approximately
1375, but that it was better known than this one sherd would indicafe is
evidenced by what we have interpreted as local imitations of Jeddito
ware. The bowl figured in plate 13, figure 2, b, has a yellowish surface
decorated in black by an uninterrupted line below the rim on the
inside and by simple angled figures attached to the rim line on the
outside. Olla fragments are also noted. In all specimens the base clay
is dark and coarse, surfaces pebbly, and the designs are in black glaze
paint. While the designs do not exactly duplicate those found on the
Jeddito yellow ware, they show points of similarity.
PINEDALE RUIN
Pinedale ruin is situated in Navajo County, Arizona, about half
a mile southeast of Pinedale, and 16 miles west of Showlow ruin.
Less than a mile to the east of the pueblo is Morterson Wash, a
tributary of Silver Creek.
The ruin consists of two units. The first is a large rectangular area
surrounded by single tiers of one-story rooms. The second is a
structure of compact form, the rooms of which are grouped about a
central rectangular plaza (see fig. 11 and pl. 14, fig. 1). The east,
west, and south sides bear proof of having been two stories in height,
terraced away from the court, while the north end of the plaza appears
to have been open or partly closed by single-storied rooms. Extending
eastward for approximately 100 feet from the northeast corner of the
main pueblo is an additional wing of rooms several tiers in breadth.
It was in the large compact unit that the search for charcoal was
continued after work had stopped at Showlow.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VO 2 NO eat elt pete nant
1. The extreme north end of the Showlow ruin, looking west. Beam HH-39
was found at the point where the man with the straw hat is at work.
2. Showlow. Room 2 of test 11, built between A. D. 1378 and 1382.
4
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLE. 82> NO= d1)) (PE
1. Showlow. Charred roof timber (below bowl) in test 11, room 2, which
dated A. D. 1378. The bowl was on the roof when the fire occurred.
2. Showlow. A second timber in test I1, room 2, which also gave A. D. 1378
as the terminal date.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLE. 82; NO= 11. (PEs 3
1. Showlow. Basal part of an old wall showing the use of large blocks sur-
mounted by small stones in poorly constructed horizontal courses.
2. Showlow. Exceptionally good section of lower-level wall uncovered in
test 12, composed of alternating layers of large blocks and small spalls.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82, NO. 11, PL. 4
1. Showlow. The diggings at the right are tests 1, 2, 3, and 12 where the
excavated rooms were found superimposed upon older structures.
2. Showlow. Room 1 of test 2 showing the firebox near the center of the room.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOPR S22 NOG titi amleS
1. Showlow. Room 1 of test 2 showing a second firebox in a lower floor.
2. Showlow. Complete excavation revealed the latest firebox imposed upon a
wall foundation of the older structure.
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SNOILO3S11090 SNOANVTISOSIN NVINOSHLINS
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOER825 (NOM til, RE 7
1. Lower level black-on-white sherds, Showlow ruin.
>
2. Black-on-white canteen and fragmentary bowl recovered from a lower level
firebox, Showlow ruin. (Diameter of canteen 6 inches. )
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOL. 82, NO. 11, PL.
Interior (1) and exterior (2, 3, 4) patterns of decorated orange-red ware, commonly known as Little
Colorado polychrome, Showlow ruin.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOPR S 2 aNOe ities
1. Lower-level corrugated ware, Showlow ruin.
2. Upper-level corrugated ware, Showlow ruin.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOE. 82, NO= ity, PES 10
1. Showlow. Tempering material consisting of pulverized sherds, quartz
grains, and a small proportion of crushed basalt found in the paste of Four-
mile polychrome pottery. (Enlarged four times.)
2. Showlow. An unfired sherd of Four-mile polychrome pottery showing an
inclusion (circled) of a fragment of a fired vessel. (Enlarged four times. )
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOL. 82, NO. 11, PL
11
Four-mile polychrome ollas dating about
/a\.
D. 1375 from Showlow ruin.
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOL. 82, NO. 11,
PES is
1. Parching plates (?) from Showlow ruin. (Diameter of a, 74 inches. )
5 me
“Werner eset?
a
2. Fragmentary bowl of Hawikuh type E (a) and imitation bowl of old Hopi
black-on-yellow (>), upper-level, Showlow ruin.
No. II
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PUEBLO RUINS IN ARIZONA—HAURY AND HARGRAVE
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Fic. 11.—Plot of Pinedale ruin showing Beam Expedition excavations.
46 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
Before the present operations were started the surface was found
to be pitted here and there by the work of previous excavators and
pot-hunters, and by shallow trenches where rocks had been removed
for building purposes. Residents of Pinedale informed us that
as late as 1887 sections of walls were still standing to a height of
from 3 to 4 feet above ground. These as well as parts of some walls
not exposed were subsequently removed by the early settlers for
building stones. Rubbish heaps are located on the east and west sides
of the main unit, and lesser deposits of débris are to be found along
the north side of the east wing. Fewkes* worked the east midden
extensively for burials in Igor.
Our search for charcoal in the Pinedale ruin showed the existence
of a different condition from that noted at Showlow. Out of a total
of 21 rooms opened, only four produced charcoal or otherwise indi-
cated that they had been destroyed by fire. From all appearances, the
pueblo was evacuated voluntarily some years before the Showlow
ruin as we will endeavor: presently to show. These burned rooms
are no doubt the results of accidents; in only one instance did we
observe what appears to be intentional firing.
For the initial test, a place was selected in the southeast section
near the plaza. In an older excavation near this spot, charcoal was
in evidence. This test, designated in figure 11 as Pr (P is equivalent
to the symbol T of the Showlow workings), was eventually extended
into three adjoining rooms, two of which were burned. The east
half of room I was excavated and the remainder was left undisturbed
because of the presence of a growing pine tree. Small charred beams
showing short ring records were recovered from this room. Room 2
proved to be of interest because of its use as a depository for rubbish
both before and after its burning. Several good beam sections, EH-56
(see pl. 15, fig. 1, for photograph of this beam in situ) and EH-62
were recovered whose outer rings dated 1286 and 1273 respectively.
Room 3 was unproductive of charcoal, hence only partially excavated.
Tests 3, 4, and 5 all showed an unburned condition. Test 6 devel-
oped into a kiva which to our good fortune was burned. All charcoal
was either directly on the slab floor or on the platform along the east
wall, indicating that its destruction probably took place while it was
in use. Here two good beam sections were encountered, as well as
numerous small fragments. Specimen EH-68 is a section of a pine
plank about 18 inches long, 10 inches wide, and 2+ inches thick. It
rested on the platform about midway between the east and west wall
(pl. 15, fig. 2) and may originally have been a part of the frame of
*Fewkes, J. W., 1904, pp. 164-167.
NO. II PUEBLO RUINS IN ARIZONA—HAURY AND HARGRAVE 47
the hatchway. The specimen does not give a true cutting date, but
the last ring, obviously near the outside, reads 1131. The second
charred beam, EH-69, was found in the south end of the kiva, partly
overlying the platform. It consisted of a short section of a timber
about 7 inches in diameter, but unfortunately was charred only exter-
nally and was badly decayed on the interior. Its outermost ring gave
the year 1207.
Test pits 7 to 19, inclusive, all failed to produce charcoal.
ARCHITECTURAL FEATURES
The walls in Pinedale pueblo were, on the average, far better than
those described from Showlow. In spite of the fact that much of the
structure had been two stories high, the walls are seldom more than
a foot thick. An exceptionally good section of wall exposed in P-16
may be seen in plate 14, figure 2. This wall, uncovered to a depth of
7 feet, was uniform throughout and chinked with small thin spalls.
The larger stones average 4 inches in thickness and are carefully
dressed both on the exposed surface and on the ends. Unusually large
slabs were employed, the second one below the whisk broom (pl. 14,
fig. 2) measuring 4 feet in length. Others of 3-foot lengths were not
uncommon.
Room 2, in the first test, was the largest excavated. Its dimensions
are 15 feet north and south by 13 feet east and west. The floor level
was found over 10 feet below the surface, which is ample evidence
of the existence of a two-story structure on that spot. That weaknesses
of construction occasionally developed is noted in the east wall. Its
lowest 5 feet are made up of large blocks of sandstone placed in
position without any effort at coursing. Above these, smaller blocks
are put down in courses and chinked with small spalls wherever the
irregularities demanded. The weight of the superimposed story ap-
parently caused this wall to buckle inwardly (pl. 16, fig. 1). Occupants
of the room, in an attempt to strengthen the wall, blocked the doorway
which leads into the room beyond. This had little effect and eventu-
ally the threatening wall caused abandonment of the room. Subse-
quently it was used as a dumping place for trash which consisted of
broken pottery, ash, discarded implements, etc.
The doorway, unlike the usual pre-Spanish Pueblo door, is at the
floor level. Its dimensions are 22 by 28 inches. Near the center of
the room was a fire-pit, oval in shape, dug into the clay floor.
KIVA
Unquestionably the most interesting architectural feature uncovered
at Pinedale is a rectangular kiva, one of the few known of that form
48 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
south of the Santa Fe railroad. The kiva site was marked by a
surface depression at the southwest corner of the plaza. A test pit
revealed a flagstone floor at a depth of 5 feet; above this, burnt soil
and charcoal encouraged our hope that we might find here suitable
timbers from which the room could be dated. This hope was subse-
quently realized; from several fragmentary timbers and numerous
small pieces of charcoal, Doctor Douglass has been able to determine
the approximate construction date of this ceremonial chamber.
The kiva was not perfectly rectangular nor was it accurately
oriented as to the cardinal points. The long axis of the chamber had
an approximate bearing 20 degrees east of north. The inner dimen-
sions are as follows: * north wall, 13 feet 3 inches; south wall, 13 feet
3 inches ; east wall, 17 feet 3 inches; west wall, 15 feet 5 inches.
Masonry.—The kiva masonry was inferior to that in neighboring
dwellings. Its building stones were generally unshaped although a
great many carefully dressed blocks, apparently fallen from the ad-
jacent two-storied rooms, were removed from the débris which filled
the kiva. Adobe mortar was plentifully used, the walls were never
more than a foot thick and horizontal coursing of building stones was
practically absent. |
The west wall, 6 feet from the northwest corner of the kiva (see
fig. 12), curves slightly to the east and then back to the west again.
At this same point is a distinct vertical separation in the masonry
which we interpret as the place of juncture of two walls of a former
room. This suggests, as several other points did also, that the kiva
was remodelled from previous living rooms.
The central part of the east wall had collapsed. We rebuilt this
portion in 1929 in a manner readily distinguishable from the original
masonry.
In the west half of the south wall is a shallow offset (pl. 16, fig. 2;
text fig. 12, a) 1 foot 10 inches wide and 4 inches deep. Its significance
is not known to us.
Plaster —Originally the inner walls of the kiva and the face of the
platform were covered with adobe plaster. Small patches still adhering
to the walls showed upwards of 11 coats or separate applications with a
total thickness of # inch. There was considerable variation in the
color of the several plaster layers; some were excessively smoke-
blackened while others were less so, probably denoting long or short
elapses of time between renewals of plaster.
*In treating of the descriptive material of the kiva, the sides will be referred
to as being either north, south, east or west, although these were not exactly
oriented to those points.
PUEBLO RUINS IN ARIZONA—HAURY AND HARGRAVE 49
LM
NO.
SCALE
Fic. 12.—Plan and section of Pinedale kiva.
50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
Platform.—This feature, which is typical of the rectangular kivas
of the Little Colorado drainage, extends the full length of the east
wall (fig. 12, b) ; the normal old Hopi kiva, however, is benched at
the south or southeast end across the shortest dimension of the room.
Its maximum width is 3 feet; its height at the south end is 2 feet
2 inches, but at the north only 1 foot Io inches. The front of the
platform, made of the same nondescript masonry as that found else-
where in the chamber, supported a fill of loose earth at the north end
and, at the south end, of clean coarse sand. This filled space was then
capped with thin sandstone slabs to form the bench floor.
Referring to plate 17, figure 1, it will be noted that the capping
slabs at the south end of the platform had collapsed under the weight
of the debris forced upon them. This may denote the existence of a
hollow space there at the time of destruction and also suggests that the
sand may have been contained there for a definite purpose. What
that may have been we do not positively know, but judging from
modern practices, the sand could well have been used in forming the
ground work of sand paintings constructed during ceremonial observ-
ances. The platform would make a convenient storage place for that
material and could be easily reached by removing the slab covering.
Hargrave’s’* finding of two large sand-filled ollas buried under the
floor in a kiva at Kokopnyama probably signified a similar custom.
Bench.—The kiva is without a true bench or banquette. The
merest suggestion of one exists along the south wall (pl. 16, fig. 2;
text fig. 12, c) from the east side of the offset to the platform. It
measured 7 inches in width and 2 feet 4 inches in height.
Floor.—The floor is completely paved with sandstone slabs from
I to 2 inches in thickness. These vary in size from slabs 3 and
4 feet in length to small pieces which were fitted into corners and
crevices. The larger flagstones were carefully placed, leaving only
small cracks which were later filled with clay.
Fire-pit—Two feet 9 inches from the platform and midway be-
tween the north and south walls is the fire-pit (fig. 12, d), a circular
opening in the floor, 16 inches in diameter and 11 inches in depth.
The bottom of the pit was formed by the convex surface of a large
water-worn boulder. Wood ashes completely filled the fireplace.
Deflector —No true deflector or fire screen, such as those usually
associated with early Pueblo kivas, was found in the Pinedale
chamber. Less than 2 feet east of the fire-pit, however, a stone
Io inches wide by 4 inches thick protruded 6 inches above the flag-
stones (fig. 12, e). Inasmuch as this stone was not directly in line
See page 112, hereinafter.
NO. DT PUEBLO RUINS IN ARIZONA—-HAURY AND HARGRAVE 5!
with the fire-place and the ventilator opening, and also because of its
low height, its function as a deflector must be regarded as doubtful.
Ventilator —The ventilator shaft connects with the room through
an opening in the front of the platform. This opening, a foot square
at the floor level, is located 7 feet from the northeast corner (fig. 12, g;
pl. 17, fig. 2). Leaning against the platform immediately north of
the vent was a squared slab which fitted the opening. The duct passes
under the platform at an angle; it is stone-lined and its floor rises
slightly to pass under the east wall of the chamber. While two wooden
beams (fig. 12, h) supported the wall at this point, there was no indi-
cation that the platform slabs above the passage had been similarly
supported. As already mentioned, the east wall at about this point
had fallen away, making it impossible to determine whether or not
the shaft continued vertically to the surface.
Sub-floor depository—Situated between the problematical de-
flector and the platform is a rectangular hole (fig. 12, f) measuring
Io by 6 inches by 13 inches in depth. The sides are formed of rock
slabs which protrude slightly above the floor. This pit was filled with
black earth.
Floor perforations—Six holes pecked through the floor flags
(fig. 12, 7) may next be mentioned. When the first one was dis-
covered, west of the fire-pit and in alignment with the usual kiva
features, it was considered to be the sipapu. Later, however, five other
similar openings were exposed, leaving some doubt as to the presence
of a true sipapu. The first opening west of the fire-pit is 1 foot
IO inches away and the second a foot from the west wall. Both are
1} inches in diameter. The two pairs to the north and the south are
about equidistant from the center of the kiva but not equally spaced.
Although not in alignment, as in Hopi kivas, it is not improbable that
these holes once served as loom anchors. A seventh opening, plugged
with a chipped stone (fig. 12, 7), was found in the southern part of
the room. The depth of these holes could not be determined. ~
The presence of two rounded stones in the floor (fig. 12, k), one on
either side of the deflector, must also be recorded. They are somewhat
thicker than the average floor flags and of a different material. Their
significance, if any, is not known.
Roof.—vThe positions of a few sizable timbers lying across the
kiva platform, indicate that the principal roof supports spanned the
chamber across the short dimension, as would be expected. Further
details could not be learned although numerous sections of branches
a few inches in diameter were presumably fragments of cross pieces.
Relation to living rooms—Exploratory tests into contiguous rooms,
two to the east and one to the west, revealed the fact that the kiva
52 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
floor was only 4 inches lower than those of adjoining rooms. Thus
the structure was not subterranean as is usually the case. The guiding
depression in reality was due to the greater amount of accumulated
débris from two-story structures to the north and south of the kiva.
A single-storied room separated the kiva from the plaza. The bulge
and separation in the west wall which has already been mentioned,
the fact that the floors of the adjacent living rooms and that of the
kiva were almost on the same level, and the proximity of the latter to
the domiciles, lead to the inference that the kiva was incorporated
into that part of the village by remodeling existing living rooms.
Corroborative evidence of this is derived from the dates of charcoal.
A series of pieces which cover a period of time between 1132 and 1231
are probably parts of beams that were originally in the rooms and
subsequently re-used in the kiva. The latest dates ranging between
1293 and 1330 probably cover the actual construction time.
Thus, by the aid of datable charcoal, we know that this rectangular
kiva was in use soon after 1300. It is too early yet to say when the
transition from the round to the rectangular form took place. We
gain a comparative idea from the two circular pre-Hawikuh kivas
excavated by Hodge. Associated with these he found black-on-white,
black-on-red often with white exterior patterns, and finely corrugated
pottery." These types correspond with those found in the lower
stratum at Showlow, for which we gave a tentative minimum date of
1204. Hence, in ruins which were occupied between about A. D. 1200
and 1300, we might expect to find the transitional forms.
In its general features the Pinedale kiva has points in common with
those uncovered by Hargrave * at Kokopnyama. The most outstanding
difference is in the position of the platform. In the old Hopi kivas,
built before the coming of the Spaniard, this feature usually occurs
on the southerly side and across the shortest dimension of the
chamber, whereas in the Pinedale kiva the platform is along the
longest side to the east. The Pinedale kiva does not show the jogs at
the platform end which Hargrave believes to be a characteristic
feature of Hopi kivas. The rectangular kiva in Hawikuh which was
abandoned and filled after the arrival of the Spaniards and later
uncovered by Hodge * also has the platform and the air duct on the
south side.
Artifacts —Comparatively few artifacts were recovered from the
kiva. Two loom blocks (pl. 18, fig. 1) found on the platform at the
* Hodge, F. W., 1923, p. 28.
* See pages 103-116, hereinafter.
* Hodge, F. W., 1922, pp. 9-10.
NO. II PUEBLO RUINS IN ARIZONA—HAURY AND HARGRAVE 53
south end are considered in the description of stone objects. The
occurrence of loom blocks in kivas is quite in keeping with modern
Hopi practices.
No complete pieces of pottery were recovered. Fragments of several
black-on-white ollas and one black-on-red olla are in the sherd col-
lections. The black paint in all cases is a thin glaze. As the sherds
from the kiva are wholly typical of those recovered from the other
diggings in the ruin, we may omit the consideration of pottery at
this point.
OBJECTS OF STONE
The stone implements recovered at Pinedale largely duplicate those
found at Showlow. Certain objects, however, are worthy of special
notice.
Metates and manos.—Concerning these, the description given of
Showlow milling stones will also apply, although the correlation of
the two types was less distinct. This is due in part to the limited
number of rooms excavated completely and possibly to the fact that
Pinedale ruin was abandoned a half century or more earlier than
the Showlow pueblo, before the second type of grinders came into
general use.
Mortar.—A mortar made of a shaped block of rhyolite was found
in the refuse-filled room of test 1. The grinding cavity was approxi-
mately 6 inches deep and the same distance in diameter at the top.
Stone axes —In the same room were found six or eight stone axes
on which the cutting edge had been completely battered away or
broken off. There were also present in the refuse several sandstone
building blocks which bore grooves, transverse to the long sides.
On one block the grooves were cut directly opposite each other on
the two sides, obviously to facilitate the breaking of the stone at those
points. The character of the grooves suggests that they were cut
with stone axes, and if so, the presence of so many battered imple-
ments can be accounted for. All axes recovered, both broken and
complete, are of the short-bitted, three-quarter groove type.
Stone hammer.—Only one stone hammer is contained in the col-
lection. It is made of diorite and is 33 inches long. Unlike the stone
axes, the hafting groove entirely surrounds the implement.
Loom blocks.—On the south end of the kiva platform were found
the pair of loom weights pictured in plate 18, figure 1. Both are
made of coarse-grained sandstone, the bases are flat, and the holes
which engaged the loom stick are near the top of the blocks and less
than an inch deep. The one block shows a long groove, evidently
where weaving tools were sharpened.
54 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
Chipped implements—A representative collection is shown in
figure 13, a tom. Specimens a and bare chert knives which have been
struck from a core much in the same manner as were the famous
Aztec knives of obsidian. The triangular tool c is also of chert,
24 inches long, and was probably used as a knife or scraper. Objects
d, f, and g are drills, the first one being of especial interest because of
the bevelling on opposite sides of the point (see cross section e¢). The
entering edge was thus made sharper than it would have been other-
wise. Arrow points are of two types, plain (fig. 13, h, 7,7) and tanged
(k, 1, m). The plain points usually have slight concave bases and are
either short or long and slender, the latter being like the points com-
monly found in the Middle Gila. The bases of the tanged points are
flat or nearly so.
Stone pipe.—The pipe represented in figure 13, n and 0, is made of
a very even-grained slatelike rock. It is 13% inches long and 1 inch
in maximum diameter. All outer surfaces are highly polished. The
bowl is formed by a conical drilling ? inch in diameter at the top
which converges into a 3-inch boring at about the middle. The boring
then extends from the juncture with the bowl proper to the base of
the object. Just above the base on the outside a large chip was knocked
off sometime during its use, for the broken surface shows much wear.
A small hole perforates the side wall at this point which looks as
though it had been made intentionally because of the small cup-shaped
depression about the hole. As to its significance we cannot be sure.
The small depression may have received the end of a stem fastened
into place by means of pitch or some other substance. True elbow
pipes, however, are the exception from southwestern ruins so that the
trait in question may have served another purpose or the object may
even have been discarded after the break occurred and put to some
secondary use.
Pendant.—Figure 13, /, illustrates a plummet-shaped pendant re-
covered in room 2, test I. It is 2% inches long, made of slate, and
perforated at one end for suspension. A single notch on each side of
the perforation represents the only efforts at incised decoration.
Ring.—The small fragmentary stone ring shown in figure 13, q,
was probably intended to be worn by a child. The material appears
to be the same as that of the pipe figured here. The protuberance
suggests a copy of shell rings made of the Glycymeris, from which
the beak is usually not removed.
Miscellaneous objects——Crystals of quartz (fig. 13, r) are not
uncommon in the ruins of central and southern Arizona. One was
owe ow
NO. II PUEBLO RUINS IN ARIZONA—HAURY AND HARGRAVE 55
ES
—
=e
SS
EE
—
Fic. 13.—Stone objects from Pinedale ruin. (7/10 natural size.)
56 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
recovered at Pinedale. It was probably used as a fetish or formed a
part of a medicine man’s outfit. Crinoid stems and asbestos which
were also found belong in the same class of articles.
Pigments——Hematite is by far the most prevalent of pigment
materials. It was found not only in its normal state, but also in
micaceous and specular forms. Limonite, used in pottery making,
kaolin, and copper carbonate were also found.
Painted stone slab.—Just above the floor of room 2, test I, in
rubbish which had been cast into the abandoned room were found the
shattered parts of a painted stone slab. The restored object, pictured
in figure 14, is made of a pinkish-colored sandstone. It measures
16 by 19 inches and is # inch in thickness. The central portion of the
stone has been neatly trimmed out to forma circular opening, 10 inches
in diameter. Prior to its painting, one side of the slab was smoothed
down to provide a better surface for the designs. A thick white kaolin
wash was first applied, not only to the dressed surface but to the
reverse side and the edges as well. The black and yellow pigments,
both apparently of mineral origin, are evanescent. Nevertheless, in
the best preserved parts, the paints are still vivid and full of life.
Nowhere is the loss of figures so great as to cause doubt as to the
continuity of design.
From the top of the slab extend a series of 11 black triangles,
14 inches high with apexes pointed downwards. Below these, travers-
ing the full width of the slab, is a heavy black line made by two
parallel strokes of the designer’s brush. Still lower and just above
the aperture is a yellow band 1 inch wide framed at both edges by
narrow black lines. Underneath the upper framing line a dark red
pigment shows up clearly, apparently a part of a previous design. To
the right and left of the opening are free zig-zag patterns in yellow
outlined in black except for the ends which are left open. The inner
edge of the hole is finished with black pigment, now almost wholly
obliterated. A single black line near the base completes the decoration.
As to its use, we believe it safe to say that the slab formed a part
of an altar decoration or was otherwise used in religious rites. This
inference is not wholly without foundation for painted slabs are still
used by the Hopi Indians in the construction of the Antelope altar in
the Snake Ceremonial." Matilda Coxe Stevenson, in her treatise of
Zuni mythology and ritual, records a ceremony * in which a wooden
1 Fewkes, J. W., 1804, p. 43.
* Coming of Ko’loowisi (Plumed Serpent) and Involuntary initiation into the
Ko’tikill. Twenty-third Ann. Rep. Bur. Amer. Ethnol., pp. 94-102, 1901-02.
—
NO. II PUEBLO RUINS IN ARIZONA—HAURY AND HARGRAVE 57
Fic. 14.—Painted altar slab from Pinedale ruin. (4 actual size.)
58 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
tablet bearing a large opening plays an important part (see fig. 15,
introduced here for comparison). The tablet is ornamented with cloud
terraces cut into the wood and outlined in color. In the center is a
circular hole 9 or Io inches in diameter through which the head of
the fetish, Ko’loowisi (plumed serpent), is passed at specified times
during the ceremony. The serpent effigy is about 8 inches in diameter.
The tablet is either carried or used in a vertical stationary position.
It is not improbable that the stone slab found by us, is precursory
to the wooden tablet in use today and that both are a manifestation of
f
p r
Fic. 15.—Wooden tablet used in a modern Zuni ceremony. Introduced for
comparison with figure 14. After pl. XIV, 23rd Ann. Rep., Bur. Amer.
Ethnol., 1901-02.
a ceremony that has survived for 600 or more years. While the
wooden tablet is the larger, the opening in both is almost identical in
size. Applying the present Hopi interpretation of symbols, the black
triangles on the ancient specimen are rain-clouds and the zig-zag
patterns are symbolical of lightning. The modern tablet, however,
does not have the latter symbols and the rain-cloud symbols are in
forms of terraces, which is the common method of representation
today. The change from stone to wood in the material of the slab
would naturally be accompanied by certain modifying features, such
as the increase in size and the cutout terraces of the present-day piece.
NO; ole PUEBLO RUINS IN ARIZONA—HAURY AND HARGRAVE 59
It will be noted that the base of the Pinedale slab is irregular and
less carefully finished than the other three edges. It has been sug-
gested that this unfinished side was imbedded in the ground to the
depth of the black line during its use. This would hold the object in a
vertical position so that a fetish such as the plumed serpent could be
drawn back and forth through the opening with facility as in the
recorded ceremony.
Relatively few painted slabs of this nature have been recovered,
and those nearly all from the ruins in the Silver Creek drainage or its
environs. Fewkes found an excellent specimen of rectangular form in
a grave at the Chevlon ruin, and Mr. Whipple at Showlow uncovered
three or four well-made slabs in living rooms in his ruin. These are
squared at the base and rounded at the upper end. Fewkes also notes
the presence of one in a grave at Sikyatki,’ which is, as far as we
know, their northernmost occurrence. None other, however, has been
recorded with the large opening.
A section of a charred timber found near the Pinedale tablet gave
an unmistakable cutting date of A. D. 1286, which, with other speci-
mens of comparable age from the same room, yields the approximate
building date of the room. Consequently, the slab was discarded at a
somewhat later time.
OBJECTS OF BONE
Awls.—Representative specimens are shown in figure 16, a, b, c.
The first one is fashioned from the proximal end of an ulna, probably
of the antelope, and c is a similar bone from some other animal.
Bodkins (?).—In his work at Chavez Pass ruin, Fewkes * recovered
13 bone implements which he termed bodkins. One complete and one
fragmentary specimen were found by us (fig. 16, d and e) at Pinedale.
These and one figured from Chavez Pass by Fewkes display such
similar characteristics that they must represent a type. They seem
always to be made of about 6-inch sections of the metatarsal bones of
the deer or antelope, including the joint. The maximum diameter of ¢
is 3 inch. The hollow of the bone is exposed for 2 inches below the
blunt point and from there to the distal end the implement is highly
polished. The articular faces of the joint were removed in order to
maintain a more uniform diameter. Hodge* pictures one from
*Fewkes, J. W., 1904, pp. 104-5, pl. XLVI.
* Ibid., p. 162.
* Tbid., p. 94.
“Hodge, F. W., 1920, pl. XX, a.
60 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
Hawikuh on which this modification was not made, although it is
identical in every other respect.
The natural furrow on the reverse side of the complete specimen
figured here, terminates in a foramen which passes through to the
base of the bifurcation of the condyle. It is apparent that the cleft
served a definite function. If used as a bodkin, the thread could
either be inserted through the foraminal passage from the base and
then knotted, or it could be brought back down and tied to the trailing
end of the thread. In the latter case, the knot formed would slip
Fic. 16.—Pinedale bone implements. (% actual size.)
conveniently into the cleft, thus eliminating the possibilities of its
catching in other threads during operation.
Problematical tool—The bone object in figure 16, f, is made of a
tubular bone 7 inches long and 4 inch in diameter. The pointed end
is too soft and spongy to have been used as an awl or a punch.
Incised bones.—Specimens g, h, and i (fig. 16) are further examples
of bones showing the cutting process. The first two were apparently
segmented in the manufacture of beads but the last disjointments
were not made. Example 7 is the discarded articular end of a leg bone.
Perforated antler—A curious fragmentary piece for which we
can suggest no utilitarian use is shown in figure 16, j. It is made of a
forked section of a deer antler 54 inches long split longitudinally. The
No. II PUEBLO RUINS IN ARIZONA—-HAURY AND HARGRAVE 61
more porous structural material on the inner side was removed,
leaving the object a mere shell $ inch thick. The edges are trimmed
down and smoothed well to the inner side of the antler. The shell
bears evidence of 11 perforations which vary from +} to 2 inch in
diameter and which are placed at more or less regular intervals length-
wise along the object.
OBJECTS OF SHELL
Ornaments of sea shells were apparently not plentiful in pueblos of
the Little Colorado drainage occupied in immediate pre-Spanish times,
for the collection from Pinedale is sparse indeed. The genera repre-
sented are: Glycymeris, Olivella, Conus, and Turritella.
POTTERY
The few complete vessels and the great quantity of sherds recovered
from the Pinedale ruin provide much needed data concerning certain
aspects of the Silver Creek drainage pottery complex. It will be
recalled that at Showlow, two levels of occupation existed and that
the pottery types from the levels differed. The great mass of the
Pinedale material is not analogous to the pottery of either of the
Showlow levels, but nevertheless small proportions show unmistakable
affinity with both. Hence, it is inferred that the period of greatest
productivity at Pinedale was intermediate in relation to the two stages
found in the former ruin. This relation was strongly suggested by a
cursory test in the refuse heap where the supposedly later types were
found to overlay the decorated orange-red phase of pottery recog-
nized in the lower level at Showlow. Corroborative evidence was sub-
sequently obtained from datable charcoal with which the dominant
Pinedale types were associated. This association was provided by the
refuse-filled room of test I and the kiva, both of which contained an
abundance of sherds and ample charcoal. No doubt exists, therefore,
as to the exact position of the Pinedale pottery in the sequence of
development supplied by the two ruins.
As has already been mentioned, the most recent date from room 2,
test I, is 1286, and from the kiva about 1330, although the largest
timbers of the latter gave earlier dates. Generally speaking, the
construction period of these two chambers was nearly 100 years earlier
than the last building period at Showlow. This would also place a
corresponding difference of time on the pottery. To the lower level
at Showlow we have tentatively given 1204 as the most recent date,
6
62 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
pending further examination of charcoal specimens. This, then, would
signify an interval of fully 75 years between the earliest Showlow
level and the major Pinedale occupation, and a difference of more
than 75 years between the latter and the final Showlow occupation.
Or, applied to pottery, which would be post-datum in all cases, the
early Showlow wares would be 1204 (?), Pinedale about 1290, and
late Showlow, 1375. Thus pottery types are not only assigned to their
respective positions in the chronology but we also obtain a relative
idea as to the amount of time elapsed during which wares acquired
new traits or even nearly lost their original identity.
The Pinedale pottery which is treated in the following paragraphs
was dominant in the late 13th and early 14th centuries. It can be
broadly classified into the following types: black-on-white, black-on-
red, black-and-white-on-red, plain, corrugated, and intrusive.
Black-on-white——This type is apparently the result of a blending
of the two black-on-white types observed at Showlow which evidenced
both northern and southeastern contacts. In the Pinedale black-on-
white, features of both older types were retained in modified form
and new ones added producing quite a distinct pottery. In properly
fired vessels, the paste is nearly white, otherwise it is darker in color.
The tempering is almost pure pulverized sherds. In some cases there
is a slight admixture of quartz grains and a few dark particles which
may be basalt. The slip is thin, light gray in color, and the surfaces as
a rule are well pebble-polished. The black paint is thin, either dull
or a near glaze; when the latter, it is frequently translucent. Whether
two pigments were used is doubtful, as in a number of pieces the dull
paint may merge into the shiny, the difference being due apparently
to the amount of paint present and the length of firing. Overfiring
seems to have eliminated the incipient glaze qualities, as in extreme
cases of overfiring the paint is a dull brown.
Ollas—Judging by the abundance of sherds, black-on-white ollas
were very common. The bodies are full and of greatest diameter
horizontally ; the orifices are small, seldom exceeding 5 inches in diam-
eter. The neck is almost invariably vertical, in some cases rising at
right angles from the body to a height of 1 to 2 inches. The lips are
direct or slightly flared.
Decoration is applied in two zones, on the neck and on the major
portion of the body. There seems to have been no fixed manner of
decorating the necks. Sometimes the lip is painted black to which
other elements are attached as in plate 19, figure I, nos. I, 2, and 3.
Again the patterns may be banded horizontally, framed by narrow or
NO: LL PUEBLO RUINS IN ARIZONA—HAURY AND HARGRAVE 63
broad lines as nos. 4, 5, 6, 7, and 8, or they may be used as independent
units as represented by no. 9. A similarity of vessel form and neck
ornamentation will be noted between these ollas and the Four-mile
polychrome ollas already described. Although black-on-white pottery
was practically non-existent at the time the latter were made, the
traits of likeness are probably survivals.
The body decoration consists of a single broad field beginning a few
inches from the base of the neck and extending well down towards
the bottom of the vessel. The field is emphasized by two heavy black
lines which border it above and below. Set apart from these are
narrow lines which form the actual framers of the patterns. In
plate 19, figure 2, is given a representative collection of olla sherds.
The designs are almost universally of contrasted heavy solid and
hatched elements. The solid figures are either continuous triangles,
terraced, or spiral elements (nos. 1 to 15). The latter, which always
interlock with a hatched counterpart, it will be recalled, are typical of
the Tularosa or Upper Gila black-on-white. Sometimes small fields
are blocked off and treated as in nos. 16, 17, 18, and 19. The hatch-
work is well drawn, much better than that of the black-on-white found
in the lower stratum at Showlow. The framing and filling lines are of
equal width and the latter are always straight. Sherds 20 and 21 show
an unusually fine brush technique. The filling lines average 25 to the
inch. Not infrequently the ends of the lines are so brought together
as to give a herring-bone effect (nos. 22, 23, and 24). Perhaps most
characteristic of the hatching is the change in direction of the fillers
in almost every small unit (nos. 8, 9, and 25). Cross-hatching was
found on one sherd only (no. 12).
Bowls——Two types of black-on-white bowls are present. The one
form is obviously related to the Showlow lower stratum black-on-
white bowls and therefore older than the second type, which on stylistic
evidence and nature of paint is judged to be contemporary with the
ollas just described. They are comparatively rare, for their place was
taken by the decorated redware bowls.
The bowls are small, very rarely more than 8 inches in diameter.
The bottoms are rounded and the rims gently incurved. Both interiors
and exteriors are polished and slipped. Ornamentation consists of
all-over patterns on interiors, two of which are shown in figure 17,
a and b, and independent elements on bowl exteriors (fig. 17, c toh).
An unusual feature of the inner design in some vessels is its lack of a
framing line at the rim. On several rim sherds, the lip is painted
black in the manner of the Chaco bowls. As in the ollas the design
64 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
units are largely contrasted solid and hatched figures. The exterior
patterns are either zoomorphic or geometric. The animal (fig. 17, c)
and the combined birds, d, appeared on opposite sides of the bowl
from which the pattern a was obtained. The tail feathers of the left
bird of the combination represented in e joined the lip-line on one
bowl and hung downward in the rakish angle shown. Figures f, g,
and h are characteristic of the geometric units. A continuous decora-
tion was noted in a single case only.
Fic. 17.—Black-on-white interior and exterior bowl decorations, Pinedale.
Black-on-red—A true black-on-red was apparently little used
although a polychrome on red was common. The sherds of the few
black-on-red vessels at hand are probably the by-products of the
early stages of the manufacture of polychrome ware. The paint is
usually dull and the designs call to mind those found on the orange-red
Little Colorado polychrome.
Black-and-white-on-red.—The polychrome on redware presents a
considerable variety of features. Despite the variety, however, the
NO. LE PUEBLO RUINS IN ARIZONA—HAURY AND HARGRAVE 65
great amount of it has an unmistakable unity. On the basis of the
chronological checks that have been provided by datable charcoal, this
dominant form was in vogue before and after 1290. Those specimens
that cannot be classed in the above group either approach the decorated
redware at the lower end of the scale represented at Showlow, or
they fall in the upper end of the sequence, also represented at Show-
low. Since the predominating type of decorated redware at Pinedale
is clearly a lineal descendent from the orange-red Little Colorado
polychrome and adumbrates the development of Four-mile poly-
chrome, we shall refer to it as a distinct sub-type, “ Pinedale poly-
chrome.”
The base clay does not differ greatly in texture from that used in
the black-on-white, although it usually burns darker. The tempering
is almost pure ground sherds. In plate 20 a small amount of sherd
tempering is shown (enlarged six times) just as it was washed from
unfired clay. The largest fragment distinctly shows the scorings found
in olla interiors. On other particles, bits of black-on-white designs
may be seen, and the dark particles reveal either black paint or the
red slip of former vessels. The fragments of the latter are rarely
found in the paste of the black-on-white pottery. Quartz grains and
other extraneous material occur in such negligible amounts that their
presence is probably accidental.
The slip is fairly thin and varies in color from red to an orange-
yellow, the former color being the more common.
The black paint on this pottery shows a gradation from a dull to a
distinct glaze finish. The lack of a glaze technique in the lower Show-
low level, which we have indicated as being older than the Pinedale
horizon represented by this pottery, and the gradual shading into
glaze decoration are strongly suggestive of the fact that paint com-
pounding was passing through an experimental stage. This, however,
does not imply local invention of glaze, as stimuli from a focus not
yet determined could readily have affected ceramic decoration in
Pinedale pueblo.
While the glaze generally resembles that of Four-mile polychrome,
in the better examples it is more lustrous, often displaying greater
relief, and usually freer of gritty particles, On firing, the glaze often
ran, thus blurring the sharpness of the lines. These differences, how-
ever, because of the variability of the paint, can scarcely be used as
determinants in distinguishing the types.
Quantitative determinations of the glaze materials by Mr. F. G.
Hawley again show lead, copper, and some manganese to be present.
66 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
In a composite quantitative test, practically the same ratio of lead to
copper obtained as was noted in the glaze of Four-mile polychrome.
The greater copper content in both cases makes it quite evident that
that element was added to the compound to produce the black color.
The latter is usually opaque, the degree of opacity depending upon the
thickness of the glaze. On some sherds, the glaze is brownish and
translucent ; and it may even have a purple cast which is possibly due
to the manganese content. The occurrence of green glaze is noted
rarely, in fact the few sherds which show it are considered to be
indicative of Hawikuh relations where green glaze was prevalent.’
In this connection we wish also to mention the occurrence of a dull
brown paint on a few sherds. It appears to be much like that used on
Jeddito yellow ware, although in the specimens under consideration
it is present on normal Pinedale types.
The white paint is chalky, and as in Four-mile polychrome, it is
much less tenacious than the black.
Ollas—Ollas are represented by a few scant sherds which do not
permit a full description. Their dearth can be explained by the
abundance of black-on-white ollas. White is used sparingly, in fact
in some cases it appears never to have been used, thus making a black-
on-red product, although belonging technically in ‘this group. Olla
shapes agree with the standard black-on-white form.
Bowls —These must have been very abundant for our collection has
in it more than 150 rim sherds of individual vessels. The sizes vary
from 7 to 10 inches in diameter, seldom larger. The bowls are con-
spicuously shallower than the preceding Little Colorado polychrome
prototype. The rims are gently incurved and the lips are almost
invariably rounded.
The variability of the decoration of bowls probably is the best
expression of the transition through which this form was passing.
To recount all the modifications would take us far afield for little
gain, especially with rather limited data. It may be said, however,
that the trend in design is distinctly approaching the results gained in
the later Four-mile polychrome.
Interior patterns of four bowls are given in figure 18 with their
corresponding exterior designs. In all but d, only black was used in
the interior while white was brought into play on the outsides. Such
is the case with approximately go per cent of the vessel fragments in
the collection, while in the remaining 10 per cent white was employed
in the capacity of outliners for the heavy black elements. In pattern a
* Hodge, F. W., 1923, p. 20.
NO. II PUEBLO RUINS IN ARIZONA—HAURY AND HARGRAVE 67
a circular area in the bottom of the bowl is unpainted; in b the field
is divided into quadrants, both designs have broad rim bands. In c
the design area is tripartite and lacking the border at the rim. Two
opposing quarters in d have a white background on which the black
ad
Fic. 18.—Interior and exterior bowl decorations of Pinedale polychrome.
elements are superimposed. The use and non-use of rim bands is
about equally divided.
Additional designs appearing on bowl exteriors are shown in
figure 19. Figures of birds (a, b, and c) and dentate elements (d, e,
and f) are characteristic. The latter are probably conventional adap-
tations of bird wings, for this is well illustrated by g where one such
68 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
figure occurs in combination with what appears to be a bird’s beak as
it is often drawn. The last three designs j, k, and / (fig. 19) are
continuous and placed from 4 to ? of an inch below the rim. Other
continuous patterns as in figure 17, b, begin to take on the appearance
of the exterior decoration of Four-mile polychrome (see fig. 8).
Plain ware——The utter lack of plain cooking vessels is one of the
surprising features of the Pinedale pottery. Decoration was lavished
on practically every vessel made except the corrugated. The only other
unpainted vessels in the collection are the shallow platelike objects
pictured in plate 18, figure 2, which we believe were used as bases in
k
covons: MMM evack (waite ES] reo
Fic. 19.—Exterior designs of Pinedale polychrome bowls.
the process of manufacturing pottery. To the depressed surface of
the smaller plate, a small quantity of tempered clay is still adhering.
Both vessels are unslipped but highly polished on the inner surface.
The exteriors are rough, but show no corrugations. The larger
specimen I is 164 inches in diameter and 4 inches deep while 2 is
I1$ inches in diameter and 14 inches deep. About the periphery of
the latter are two rows of perforations 4 inch apart. The holes were
made from the inside outward before the surface was polished, for
the latter process nearly closed some of the perforations. Similar
objects have been recovered in Marsh Pass* and in the old Hopi
ruins. As to the significance of the marginal perforations, there
* Kidder, A. V., and Guernsey, S. J., 1919, p. 143; also by Doctor Cummings.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82, NO. 11, PL. 14
2. A typical section of Pinedale masonry. The second slab below the whisk
broom 1s 4 feet long.
‘RAIN O[Bpoulgq JO wuo0yeId uo ‘SY -Id Url punos ‘ogzi “q “Vy palep yoiyM nye wi OS-FAA
uolisod ur (WOOIg YSIYM MOTEq) gO-F{A UsutIadS °z wesq jno suljulod ssejsnoq “y -y Aq ‘aTepauld ‘1
Gl “ld “LI “ON ‘28 “10A SNOILOA1100 SNOANVIISOSIN NVINOSHLIWS
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOES (825 NO} 11, PE 16
1. Pinedale. Pi-R2 after excavation. Note bulge in far wall and the door-
way on level with the floor.
e
’
is)
Pinedale. The offset in the south wall of the kiva and the suggestion of a
banquette extending from the offset left to the platform.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLTS 25NOs til, eens
1. The Pinedale kiva looking south. Note the collapsed platform at the
south end.
2. Pinedale. East view of kiva looking into ventilating shaft and firepit.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLES 2s NO. IM) Pea is
1. Loom blocks found in the Pinedale kiva.
2. Platelike vessels from Pinedale. (Diameter of figure 1, 164 inches.)
‘UINA aTepeul gq
‘UOZIIOY OOTI JNoqe WOT, Sp1OYS LI]O 9}IYM-UO-YoR]q ‘Zz ‘Unt aepoulg ‘uozlsoy OOZI JNoqe Wor} ‘sprays-WII eI[O 9}1YM-UO-YyOeLG “1
61 “Id LL “ON “28 “10A SNOILO3A1100 SNOANVIISOSIN NVINOSHLIWS
SMITHSONIAN MISCELLANEOUS COLLECTIONS
Pinedale. Pulverized potsherds used
showing former designs. (Enlarged six times.)
as tempering material.
VOLES 2 INO} lithe Ple
Note particles
f
is
NO. II FUEBLO RUINS IN ARIZONA—HAURY AND HARGRAVE 69
seems to be no satisfactory explanation, although in the vessel illus-
trated here they appear to have served no other purpose than orna-
mentation.
Corrugated.—Corrugated ware does not differ appreciably from
that found in the lower horizon at Showlow. Finely corrugated ware
is almost entirely lacking at Pinedale.
MISCELLANEOUS POTTERY OBJECTS
Knobbed vessel.—Figure 20 illustrates a fragment of a small globu-
lar black-on-white vessel found in test 5, Pinedale ruin. It has, in
addition to the paint decoration, two horizontal rows of protuberances
Fic. 20.—Portion of a knobbed and painted vessel. (Actual size.)
spaced at about half-inch intervals. They were made by sticking
small pointed pellets of clay to the vessel wall and emphasized further
with touches of black paint. Broad black lines border the two rows
of knobs.
Vessel fragment showing molded hand.—The object pictured in
figure 21 illustrates another attempt at adding plastic features to a
vessel exterior. Here a crude four-fingered hand is portrayed, evi-
dently a part of a more elaborate production. The top of the hand
and fingers originally were painted black but the paint is now almost
obliterated. The thickness of the sherd, excluding the hand, is % inch,
and, judging by its curvature represents a vessel that was fully a foot
in diameter.
Pottery scrapers—In addition to several pottery scrapers similar
to those found at Showlow, one was recovered in Pinedale ruin that
7O SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
is of unusual interest. It is made of the handle and part of the body
of a small black-on-red pitcher (fig. 22). The handle is decorated in
imitation of an animal. Two small projections at the rim, each tipped
with black, are probably intended to portray the eyes. Fewkes”* re-
covered two black-on-white pitchers in Kin Tiel, the handles of both
showing similar treatment.
Intrusive pottery types—Fragments of vessels which are charac-
teristic of the Middle Gila were found in considerable numbers in
Pinedale ruin. These occur contemporaneously with the types of
pottery described in the preceding pages. The abundance of this non-
local ware need not imply, however, that all of the vessels were
Fic. 21.—Vessel fragment showing molded hand. (Actual size.)
actually acquired in trade relations, but rather that some of it was
made locally in imitation of trade pieces or possibly even made by
women coming from the southern district. Examination of the
sherds leads to a conviction that such was the case for there are some
specimens identical in every respect with vessels of similar style found
on the Gila, while others show local sherd-tempered paste and local
treatment of the designs.
The fragments fall into two groups: (a) those of bowls with
black-on-white interior decoration and plain red or brown exteriors,
and (b) fragments of bowls with black interior designs on red. The
former type is well known and was found to be a comparatively late
arrival at Casa Grande by Mr. H. S. Gladwin.”
*Fewkes, J. W., 1904, pp. 130-131.
* Gladwin, H. S., September, 1928, p. 20.
No. II PUEBLO RUINS IN ARIZONA—-HAURY AND HARGRAVE Fifi
The second group is perhaps less known and as far as we can find,
has never been described. Vessels of the same character have been
recovered by Dr. Bryon Cummings with cremations in ruins on the
Gila near San Carlos. Technologically it is identical with the two-color
decorated ware except that a light red or salmon-pink slip was sub-
stituted for the white. Sometimes the slip was dispensed with, the
natural color of the base clay serving as the background.
Hopi relations are shown by two fragmentary vessels of Jeddito
black-on-yellow, one found near the surface and the other with a late
burial made in the fill in a room. Two bowls of Pinedale polychrome
Fic. 22.—Pottery scraper made of a black-on-red pitcher fragment. (Actual size.)
were found with the latter. Sherds of local imitation of Hopi yellow
pottery, such as noted from Showlow, were not encountered here.
Contacts with old Zufi ruins are not clearly indicated. Hodge’s
early Hawikuh Period C may be represented by a few fragments of a
black-glaze-on-white olla.
THE BAILEY RUIN
During the course of excavation at Pinedale, we were informed of
a ruin of considerable extent, situated in the Phoenix Park district,
approximately 15 miles west and slightly north of Pinedale. The ruin
stands on land owned by Mr. Geo. W. Bailey and locally is known as
“ Bailey’s ruin,” although some years ago it was known as the “ Stott
72 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
Ranch ruin.”’* In the late ’80s the ranch was a notorious center from
which horse thieves operated.
The ruin is located a few hundred yards southwest of the ranch
buildings on the south edge of a long narrow park. The higher part
of the ruin is covered with large pine trees.
A sherd collection made on the first visit to the site contained, among
other types, a small proportion of the black-on-orange-red phase of
pottery which fell in the gap-period of the Douglass chronology. On
these grounds a day was spent at the close of the field-work in putting
down three tests to ascertain whether or not a prolonged search for
charcoal would be justifiable. The three pits, sunk at widely separated
points in the northern extension of the ruin were unproductive of char-
coal, consequently further search was postponed.
The pottery at this site indicates contemporaneity with Pinedale.
Late 14th century types are but meagerly represented, hence its
history may be quite analogous to that of Pinedale.
The Bailey ruin is one of the few, if not the only one, of consider-
able size in this region which has not been pilfered. Less than a half-
dozen rooms have been touched, and the trash mound which is
extensive and superficially appears to have depth, is undisturbed.
From it much valuable stratigraphic data could undoubtedly be ob-
tained which would greatly clarify and contribute to the present
meager knowledge of human activity in the Silver Creek drainage.
CONCLUSION
We may now turn our attention away from the purely archeological
aspects of the problem and consider it on an historical basis. Dates
obtained from charcoal have already been given in the several sections
of this paper, but there still remains the formulation of the process
of development as shown by the present data. Spier and Hodge * have
contributed materially to the establishment of the sequent stages of
progress in this southern part of the Little Colorado drainage. The
present reconstruction follows the general outlines set by the above
investigators, but it applies only to the area drained by Silver Creek
and its immediate environs in which some local variations have been
noted.
*So named by Fewkes, who visited it in the summer of 1897 but did not ex-
cavate due to its isolation from supplies. Fewkes, J. W., 1904, p. 167.
* Spier, L., 1919; Hodge, F. W., 1923; note also Kidder’s résumé, 1924, p. 04.
NO. II PUEBLO RUINS IN ARIZONA—-HAURY AND HARGRAVE 73
The Showlow ruin and possibly the one at Pinedale originated as
small, loosely arranged buildings of probably not more than a dozen
rooms. The remains of small sites of this description are plentiful
in the region, especially along Showlow Creek. Their existence is
indicative of a time when the settlements were generally small; the
buildings were apparently not more than one story high and not
closely knit. Tests in the northeast section of the Showlow ruin
brought to light the foundations of one of these early settlements.
The associated pottery types are black-on-white of San Juan (Chaco)
affiliation, a very small percentage of redware with dull black
interior and white exterior designs, and corrugated ware of relatively
fine technique. These types are in marked agreement with those
found in the small communities not far afield.
Judging by the pottery found in them, the small sites seem to have
been evacuated almost simultaneously not long before the decorated
redware became plentiful. Such concordant action is almost cer-
tainly the result of a disorder, the cause of which might well be
attributed to the influx of nomadic and predatory people. Just when
this disorder, or whatever it was that motivated the change, took
place, we cannot yet say positively as none of the small sites has been
explored for wood specimens. However, on the strength of the evi-
dence secured in the lower level at Showlow, we venture the assertion
that it was prior to A. D. 1200, for by about that time a greater
amount of decorated redware and black-on-white of southern deri-
vation is present. The paucity of these types in the small ruins would
place their abandonment at a somewhat earlier time.
The concentration of the population in a few chosen pueblos intro-
duced problems before unknown. As a consequence, the move-
ment was attended by an accelerating force and a stimulus in the
development of certain cultural traits. Extensive structural additions
to the villages were demanded in order to accomodate all. Rooms
were compactly arranged side by side and one above the other. Thus,
it seems, the true pueblo came into existence in this region.” In
ceramics, the few types which showed exterior influences were modi-
fied and fused and native types underwent relatively rapid changes
to produce highly localized forms.
We find that the Showlow and Pinedale pueblos attained their
largest size during the 14th century. The Pinedale pueblo experienced
a major building period in the last decades of the 13th century but
*The pueblo idea, however, was not original, for structures of that type had
been in existence for several hundred years previous in the San Juan.
74 SMITHSONIAN MISCELLANEOUS : COLLECTIONS VOL. 82
apparently grew very little after that time. A century later, the Show-
low pueblo was still growing, then suddenly stopped and from all
indications was involuntarily abandoned. The highest refinement in
masonry is shown at Pinedale in the rooms dating towards the close
of the 13th century. Several shallow late walls at this site and all of
the late 14th century structures at Showlow indicate a marked retro-
gression in construction. This is in accord with the general trend of
Pueblo architecture manifested in the ruins built after the Great
Period (Pueblo III). The length of the occupation at Showlow,
which Hough believed to be short, is approximately 200 years as
indicated by datable charcoal, but, since charcoal is lacking from the
old northeast quarter which appears to be still older than the first
definite Showlow horizon, we can assume a considerably longer period
of occupancy. The greatest exodus at the Pinedale pueblo seems to
have taken place early in the 14th century, at least before the develop-
ment of Four-mile polychrome in its fullest form. The presence of a
small percentage of the latter indicates, however, that a few indi-
viduals remained or returned subsequent to the abandonment.
Concerning the development of ceramics during the lapse of time
represented by the present diggings, we find it necessary to correlate
the data from both ruins in order to place the material in its sequential
order. Three periods are registered which pass almost imperceptibly
into each other. For the oldest horizon represented in tests 1, 2, 3,
and 12 at Showlow, we have ascribed the tentative date of 1204; the
second period manifested at Pinedale is 1290; the last phase at Show-
low again, is 1375. First period forms of decorated ware are black-
on-white which evidences foreign influence, and black-on-orange-red
with white exterior designs, possibly of local derivation. This phase
is probably broadly coincident with Hodge’s pre-Hawikuh Period A.’
It is worthy of note that the pottery of Chaco traits existed in this
region possibly a hundred years after the Chaco Canyon culture
ceased.” This is a good example of the survival in marginal areas of
a trait which has died out in the center of origin.
3y about 1290 the black-on-white, which was still abundant, had
been more or less standardized into a form typical of the region but
with the retention of definite traits exhibited in black-on-white found
along the Salt River. It represents probably the last survival of black-
on-white pottery south of the Santa Fe railroad. Decorated redware
* Hodge, F. W., 1923, p. 29.
? Judd, N. M., (in preparation).
NOS DE PUEBLO RUINS IN ARIZONA—HAURY AND HARGRAVE 75
shows a signal increase in quantity and several rather abrupt depar-
tures from features of the lowest horizon form. The bowls have
become shallower, black plays an important part in exterior designs,
and white sometimes in interior patterns. This form, referred to as
Pinedale polychrome, has been shown to be antecedent and leading
up to the typical Four-mile polychrome as here described. Its time
of appearance is certainly not later than the last decades of the 13th
century. This period also marks the invention or the introduction
of black glaze paint which was used on both white and red wares,
without an appreciable change in decorative style. During this second
stage practically all of the pottery made, except the corrugated, was
decorated. External relations were largely to the south with the people
of the Middle Gila, and to a lesser degree to the east with old Zuni
and north with the old Hopi cultures.
By 1375, black-on-white pottery was practically non-existent. Its
rather sudden disappearance may.be explained by an intense local
specialization of the decorated redware or Four-mile polychrome. The
basic differences between the latter and its ancestral form are: a
deeper red slip covering the paste; a poorer, gritty black paint but
still basically a glaze ; exterior continuous patterns in black and white ;
and generally unbalanced geometric units and life patterns on the
interior fields of design, also executed in black and edged with white.
Guided by Fewkes’ finds at Four-mile ruin, we may say that the use
of life patterns in bowls became more prevalent in the most recent
forms of Four-mile polychrome. Relatively few are noted in bowls
from the Showlow ruin and more from Four-mile which was aban-
doned after Showlow. The last phases of Four-mile polychrome at its
type site are contemporary with Jeddito black-on-yellow and the three-
color Tonto polychrome, and then, rather suddenly apparently, it
passed out of existence. Interrelations at the close of the last period
were increasing with the Zuni and Hopi areas although contacts with
the Gila are still represented.
At this point it is well to insert a brief discussion of the age
of lead glaze in the Pueblo region in the light of the expedition’s
discoveries. Some investigators are of the opinion that the glaze
technique is acultural and not indigenous; that it was obtained from
the Spaniards or even from Mexico later than the Conquest. Other
explorers, however, have given glaze decoration a pre-Spanish status
on the basis of stratigraphy. To this latter contention our evidence is
directly corroborative. Not only does stratigraphy at Showlow and
* The Medallion, 1930, pp. 8-9, pl. VI.
76 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
Pinedale ruins, both totally abandoned before the advent of the
Spaniards, uphold this, but the newest science for age-determination
of southwestern ruins, namely, dendro-chronology, confirms the strati-
graphic claim. And more, it hints strongly at the Jength of time that
elapsed between the development of a glaze technique and the first
Spanish contacts.
As previously suggested, the apparent gradual merging of the dull
black paint into one with glaze properties on Pinedale polychrome
does not bespeak a sudden appearance of glaze but rather of a
slow development in or not far from the center of invention. The
beginnings of Zufi and Pinedale glaze development appear to parallel
each other, both seemingly responding to the same stimulus. That this
stimulus was pre-Spanish and consequently native, is indicated by the
presence of lead glaze on Pinedale polychrome coming from the late
13th century horizon at Pinedale, and the continuance of its use on
Four-mile polychrome from the upper Showlow level dating 1375.
Thus, glaze paint was known and used fully 200 years before the ar-
rival of the Conquistadores. Indeed, it had already become decadent
by the time of their arrival and soon after the Pueblo potters reverted
completely to the dull, flat colors.”
These facts do not agree with Hough’s statement, therefore, when
he says* ““ . . . . that the Pueblo potter could take the step to glaze
which appears to have arisen in the line of smelting metals is doubt-
ful.” Or, “ . .. . acritical examination of the question as to the use
in prehistoric times of lead glaze decoration by the Pueblo Indians
brings forward a number of points which seem to render the assump-
tion doubtful if not improbable.” He states further that glaze deco-
rated wares are never associated with black-on-white. Numerous tests
in Pinedale ruin always showed glazed wares and black-on-white to be
synchronous, the latter also frequently, but not always, decorated with
glaze. This contemporaneity of glaze and black-on-white wares either
signifies that glaze was invented earlier than has been supposed, or,
that the Pinedale black-on-white was a late survival.
Additional checks on the age of lead glaze may be had by the
presence of intrusive glazed sherds in ruins outside of the glaze area.
Thus, Four-mile polychrome sherds and Zuni glazes found in Kokop-
nyama which has yielded no trace of Spanish influence and no datable
wood more recent than A. D. 1416, and the occurrence of Four-mile
* Kidder, A. V., 1924, p. QI.
* Hough, Walter, 1928, pp. 248-249.
NO. II PUEBLO RUINS IN ARIZONA—-HAURY AND HARGRAVE 77
polychrome at Casa Grande ruin,’ show that glazed pottery entered
the trade channels and was carried far afield before 1520.
The arguments for a post-Spanish origin for glaze in the Pueblo
region can hardly be considered tenable in view of the unerring
evidence of stratigraphy and dendro-chronology combined.
It is of especial interest to note that in the Pinedale horizon two
types of glaze occurred contemporaneously for a short time. The
indications of Mr. Hawley’s paint tests are that the redware invariably
carried a lead glaze with a relatively high copper content, while the
black glaze on white ware contained neither lead nor copper but was
apparently of a salt compound. The latter was earlier than the lead
glaze, for a small percent of black-on-white pottery from the lower
Showlow stratum bore a black shiny paint, foreshadowing its later
development. It seems to have passed out of existence, however, with
black-on-white ware, being survived by the lead glaze. It may there-
fore have contributed in some measure to the development of the
superior lead glaze paint.
The sequential development of the decorated redware of the first
Showlow horizon to Four-mile polychrome of the final stage makes
an interesting disclosure concerning the spatial element involved in
ceramic progression. If the date ascribed to the oldest culture period
is correct, namely, A. D. 1204, then approximately 200 years elapsed
to effect the changes. The transitional form, Pinedale polychrome,
comes about midway between the two terminal dates. Thus approxi-
mately each century, basic changes were made in pottery, so that
by the end of a 200-year period, the resultant form had but little
resemblance to its earliest forerunner. These figures are not given as
generalities, as the rate of development undoubtedly depended upon
the nearness to a manufacturing nucleus and upon external influences,
but they seem to hold true for the region under consideration.
The accurate date-checks of Four-mile polychrome which have
been secured make it invaluable as an indicator of time in those
ruins where it is found. Thus, Four-mile ruin, a site near Shumway,
Homolobi, Chavez Pass, Chevlon, and a few others were occupied as
late possibly as the beginning of the 15th century. The occurrence of
Four-mile polychrome in Gila Pueblo, Globe, and at Casa Grande,
west of Florence, shows trade relations and, hence, life in those sites
as late as about 1400. Mr. H. S. Gladwin’s recovery of four or five
sherds of this ware in the Late or Classic horizon at Casa Grande is
convincing as to the recency of occupation there.
*The Medallion, 1929, pl. IV.
Z
78 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
The following composite tabulation of the several recognized hori-
zons and the related local ceramic types in Showlow and Pinedale
ruins is necessarily incomplete and must be enlarged upon at a
subsequent time.
First Horizon: A. D. 1204 (?) Showlow
Black-on-white very abundant; shows both Chaco Canyon and southern
affinity.
Black-on-orange-red with white exterior patterns abundant; designs usually
balanced solid and hatched elements in dull paint. (PI. 8.)
Corrugated ware crude to fine, some with exterior white decoration. (PI. 9,
figs De)
Second Horizon: 1290 + Pinedale
Black-on-white abundant; vessel forms mainly ollas; decorations are dis-
tinctly local but retain certain foreign features; thin black glaze paint
commonly used. (PI. 10, figs. 1 and 2, text fig. 17.)
Pinedale polychrome very abundant, appearing almost exclusively as bowls;
is directly derived from the first horizon orange-red phase; white seldom
used on interiors and almost invariably in association with black on
exteriors in independent units or continuous patterns; interior designs
geometric and balanced but of great variety; black paint is preponder-
antly a lead glaze, seldom merging into other colors. (Figs. 18 and 19.)
Corrugated ware crude and not very abundant.
Third Horizon: 1375 = Showlow
Black-on-white ware rare or absent, apparently no longer in vogue.
Four-mile polychrome predominating decorated ware; slip a darker red and
generally softer than antecedent stage; glaze paint is decadent, may lack
luster and be vitreous and gritty; exterior patterns are in black and
white and almost without exception continuous; interior designs are
also in black and white in specialized geometric and zoomorphic ele-
ments. (Pls. 11 and 12.)
Corrugated ware more abundant, some shows horizontal flutings or ribs not
evident before. (PI. 9, fig. 2.)
BIBLIOGRAPHY
BAnpbeE.IER, A. F.
1892. Final report of investigations among the Indians of the Southwestern
United States, Part II. Papers Archaeol. Inst. America, American
Series, No. IV, Cambridge, 1802.
Douctass, A. E.
1929. The secret of the Southwest solved by talkative tree rings. Nat.
Geogr. Mag., December, 1929. Vol. LVI, No. 6, pp. 737-770, Wash-
ington, 1929.
FEwKEs, J. W.
1894. The Snake ceremonials at Walpi. Journ. Amer. Ethnol. and Ar-
chaeol., Vol. IV, Cambridge, 1894.
1904. Two summers’ work in pueblo ruins. Twenty-second Ann, Rep., Bur.
Amer. Ethnol., Pt. I, pp. 3-195, Washington, 1904.
NO. TE PUEBLO RUINS IN ARIZONA—HAURY AND HARGRAVE 79
GLapwIn, H. S.
1928. Excavations at Casa Grande, Arizona. Southwest Museum Papers,
No. 2, Los Angeles, 1928.
GuTHE, Cart E.
1925. Pueblo pottery making. Papers of the Southwestern Expedition,
No. 2. Published for the Department of Archaeology, Phillips
Academy, Andover, Mass., by the Yale University Press, New
_ Haven, 1925.
Honce, F. W.
1920. Hawikuh bonework. Mus. Amer. Ind., Heye Foundation; Indian
Notes and Monog., Vol. III, No. 3, New York, 1920.
1922. Recent excavations at Hawikuh. El Palacio, Vol. XII, No. 1,
pp. I-11, Santa Fe, 1922.
1923. Circular kivas near Hawikuh, New Mexico. Contr. Mus. Amer. Ind.,
Heye Foundation, Vol. VII, No. 1, New York, 1923.
Houcu, WALTER.
1903. Archeological field-work in northeastern Arizona, the Museum-
Gates Expedition of t901. Ann. Rep. U. S. Nat. Mus. for roo1,
pp. 279-358, Washington, 1903.
1928. The lead glaze decorated pottery of the Pueblo region. Amer. An-
throp., N. S., Vol. 30, No. 2, April, 1928, pp. 243-249; Menasha,
1928.
Jupp, N. M.
1922. The Pueblo Bonito Expedition of the National Geographic Society.
Nat. Geogr. Mag., March, 1922, Vol. XLI, No. 3, pp. 322-331,
Washington, 1922.
1925. Every-day life in Pueblo Bonito. Nat. Geogr. Mag., September,
' 1925, Vol. XLVIII, No. 3, pp. 227-262, Washington, 1925.
Kipper, A. V., and GUERNSEY, S. J.
1919. Archeological explorations in northeastern Arizona. Bull. 65, Bur.
Amer. Ethnol., Washington, 1910.
Kipper, A. V.
1924. An introduction to the study of southwestern archeology. Papers
of the Southwestern Expedition, No. 1. Published for the Depart-
ment of Archaeology, Phillips Academy, Andover, Mass., by the
Yale University Press, New Haven, 1924.
MEDALLION, THE
1929. The red-on-buff culture of the Gila Basin, Pasadena, 1920.
1930. Some southwestern pottery types. Globe, 1930.
Spier, L.
1919. Ruins in the White Mountains, Arizona. Anthrop. Papers Amer.
Mus, Nat. Hist., Vol. XVIII, Pt. 5, New York, 1919.
STEVENSON, MatiLpA Coxe.
1904. The Zufi Indians, their mythology, esoteric fraternities, and cere-
monies. Twenty-third Ann. Rep., Bur. Amer. Ethnol. for 1901-
1902, Washington, 1904.
EXCAVATIONS AT KIN TIEL AND KOKOPNYAMA
By Lynpon L. HARGRAVE
Following a superficial examination of several ruins in the Little
Colorado River drainage early in the summer of 1929, the Third Beam
Expedition of the National Geographic Society centered its initial
search for datable fragments of charred ceiling timbers in the two
pre-Spanish Indian villages at Showlow and Pinedale, Navajo County,
Arizona. After working a few weeks with Mr. Haury at the former
site the present writer proceeded to Kin Tiel ruin; later, to Kokop-
nyama.’ Since both these villages lie north of the Little Colorado and
show definite Hopi affinities, the observations presented herein may
prove helpful to those archeologists interested in that period of Hopi
history which shortly preceded the coming of the Spaniards in 1540.
The writer begs to remind his readers, however, that the 1929 explora-
tions were undertaken solely for the purpose of collecting ancient
beams that might contribute to the completion of the Douglass tree-
ring chronology and thus disclose the actual age of Pueblo Bonito, in
Chaco Canyon, New Mexico. The data herein recorded are, therefore,
to be regarded merely as by-products of the expedition.
KIN TIEL RUIN
Wide Ruin, or Kin Tiel as the Navajo call it, is a well-known ruin
situated on an eastern tributary of LeRoux Wash 18 miles north of
Chambers, Arizona. On early maps the ruin is indicated as Pueblo
Grande ’* and was first given prominence through the work of Minde-
leff *and Fewkes.* At the time of their respective observations, the old
village must have presented an inspiring sight, for many portions of
its broken walls were standing two stories high. Today these have
been reduced to a low mound which, from its shape, is sometimes
referred to as ‘“ The Butterfly Ruin.” The appropriateness of this
term is at once apparent from the accompanying ground plan (fig. 23).
*The author wishes to thank Mr. E. C. Greene, Jr., for his services as field
assistant, and for drawing the plans.
*8th Ann. Rep., Bur. Amer. Ethnol., 1886-7, p. 91.
*8th Ann. Rep., Bur. Amer. Ethnol., 1886-7. A study of Pueblo architecture,
Tusayan and Cibola, by Victor Mindeleff.
*22nd Ann. Rep., Bur. Amer. Ethnol., part I, 1900-01, p. 124.
80
NOs Dt PUEBLO RUINS IN ARIZONA—HAURY AND HARGRAVE 81
Unlike other known ruins in the district, the outer wall of Kin
Tiel was unbroken save for narrow passageways.’ Terraced dwellings
looked down upon open courts as in the case of Pueblo Bonito; these
courts were separated by a stream channel which appears to have been
crossed by extensions of the outer wall of the village, if we may judge
from Mindeleff’s carefully prepared plan.
Toward the west end of the pueblo and crossed by the wash, or
stream channel, a masonry-walled spring furnished the village folk
with an abundance of clean pure water. Today this spring is used by
Mrs. D. W. Balcolm, the present owner, who has installed a pump
to supply domestic water for her trading post, as well as water for the
flocks of Navajo sheep that range the district. When Mindeleff made
his reconnaissance the location of this spring was unknown,’ and he
comments upon the apparent lack of an adequate water supply. The
spring was discovered about 40 years ago by Mr. Hawthorn, who
settled and built a trading post nearby, and who unfortunately de-
stroyed most of the outer wall of the ruin in his search for suitable
building material. This destruction has been completed within the
past two years, for the foundation stones at Kin Tiel were quite
naturally preferred in the construction of modern dwellings. In con-
sequence, no primitive masonry today stands above ground. The
Butterfly Ruin has been levelled utterly; its once terraced chambers
have been reduced to a low, wide-spread mound of sandstone blocks
and adobe mortar.
KIVA KT-I
Since the sole object of the National Geographic Society’s 1929
expedition was to secure datable beam material, and since previous
experience had taught us that the largest beams were likely to be
found in kivas, the well-known subterranean ceremonial chambers
of the Pueblo tribes, our initial efforts in Kin Tiel were directed
toward discovering burned rooms of this type. A large circular de-
pression was therefore chosen for excavation. This, because of its
location and diameter, was thought to indicate a kiva of Pueblo Bonito
type. Certain local resemblances to Bonitian architecture and pottery
had greatly influenced this belief, in spite of the traditional connec-
*8th Ann. Rep., Bur. Amer. Ethnol., 1886-7, p. 92.
*We wish to acknowledge Mrs. Balcolm’s willing cooperation in the purpose
of the expedition, her generous permission to excavate and her warm hospitality
which was extended to the members of our party.
*8th Ann. Rep., Bur. Amer. Ethnol., 1886-7, p. 92.
VoL. 82
SMITHSONIAN MISCELLANEOUS COLLECTIONS
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NOs LE PUEBLO RUINS IN ARIZONA—-HAURY AND HARGRAVE 83
tion with Zufi.” Excavation, however, proved the absence of charred
timbers and we turned to another depression of similar appearance.
Contrary to expectations this second test revealed a rectangular
room which differed only in non-essentials from similar chambers
in present-day Hopi villages. We had no reason to expect a subterra-
nean room of this type even though there was some justification for
the belief that Kin Tiel was contemporaneous with prehistoric Hopi
settlements of the late Pueblo III and early Pueblo IV horizons. We
had noted no superficial evidence of Hopi culture, but Mindeleff had
remarked * that large circular depressions often revealed rectangular
rooms. This statement was based on the fact that he had found, only
a few feet from our second excavation, the walls of a rectangular
room which for lack of time he was unable completely to lay bare. We
finished the work he started in this particular chamber (our KT-II)
and the resultant floor plan closely resembles that of KT-I, the first
kiva we excavated at Kin Tiel.
For a better understanding of the Hopi type of kiva let us consider
this latter chamber (fig. 24). Its floor is divided into what we may
call the kiva room and the platform, or alcove. On the elevated plat-
form, spectators might gather to witness the rituals performed in
the larger space where the “ altar” appropriate to each ceremony was
arranged and the accompanying prayer dramatized.”
It is probable that this kiva, KT-I, is the oldest of the Hopi type
yet excavated, and while its shape seems to be a modification of the
earlier rectangular kiva, such as those at Betatakin, for example, in
this instance there appear to be two rooms combined and remodelled.
This is indicated by the difference in construction of the two divi-
sions: the walls of the platform are of masonry, whereas those of
the kiva room are merely the adobe plastered sides of a hole dug into
hard-packed sand. The kiva floor, or area devoted to ceremonial
purposes, is both wider and longer than that of the platform, though
both were under the same roof.
Other general characteristics of the special type noted in KT-I are:
offsets or jogs, in the side wall where the kiva room and platform
alcove meet; the deflector, or fire screen; the firepit; the ventilator,
in the lower face of the platform; the ventilator passageway beneath
*8th Ann. Rep., Bur. Amer. Ethnol., 1886-7, p. 92.
*8th Ann. Rep., Bur. Amer. Ethnol., 1886-7, p. 93.
* A detailed architectural description of the Hopi kiva of the nineteenth cen-
tury is given by Mindeleff in A study of Pueblo architecture, Tusayan and
Cibola. 8th Ann. Rep., Bur. Amer. Ethnol., 1886-7.
VOL. 82
SMITHSONIAN MISCELLANEOUS COLLECTIONS
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No. II PUEBLO RUINS IN ARIZONA—-HAURY AND HARGRAVE 85
the platform and outer wall; and the ventilator or air shaft connect-
ing with the passageway outside of the kiva wall. Banquettes and
pilasters are absent in both KT-I and II, and the presence of a
sipapu is questionable.
The ceiling height is not known, but in the northeast and south-
west corners, 6 feet 11 inches above the floor, three courses of
masonry remain as a possible support for the original roof timbers.
The roof was probably just above ground, as is the case with the
older historic Hopi kivas. The original ceiling height is estimated
to have been between 7 and 8 feet.
In determining the length of the chamber, the raised platform
alcove must be considered, for it is really a part of the same room.
In KT-I, the depth of the platform is 4 feet 1 inch; this, added to the,
length of the kiva room, makes a total length of about 17 feet. Over-
looking the principal floor area as it does, the platform provided
space for spectators just as we saw them during practice of the Bean
Dance at Oraibi in 1928."
In front of and on each side of the platform alcove is a jog or
offset (pl. 21, fig. 1). That on the west is 84 inches wide; that on the
east, 64 inches. Both offsets are of masonry abutting the cut banks
which form the kiva walls.
As previously stated, the side walls of the kiva are of adobe plaster
applied directly to the sandy face of the original excavation. I do
not recall any other pueblo ruin so late as Kin Tiel in which plastered
earth walls substituted for masonry. Only the firm condition of
the sand bed made this possible. But that the builders were taking a
chance is shown by the later collapse of the plastered banks.
The collapse of the roof was obviously caused by the caving of
the west wall. That this accident came suddenly and without warn-
ing is evidenced by the fact that five men were caught between the
settling ceiling and the floor. Two jumbled skeletons were on the
southwest end of the platform. A few feet northeast of the fire-
pit, and in a sprawled position with the head resting on a pile of
tumbled rocks, was found the third skeleton, which was partly
burned. The skull and upper part of the body were burned where
they came in contact with the burned part of a large juniper beam.
The lower part of the skeleton, lying free from the beam, was ex-
cellently preserved, being covered with wind-blown sand that had
sifted through the roof. Two tubular bone beads, several large circu-
* Nat. Geogr. Mag., Vol. 56, No. 6, p. 755. Washington, 19209.
86 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
lar beads of a white chalky material, and a large flint point were
found among the bones of the body. Fragments of both black-on-
orange and corrugated pottery were under this skeleton, and on the
floor nearby were sherds of black-on-white ware. The fourth skele-
ton was in the northeast corner, in a flexed position on the floor, with
face toward the firepit, and only partially destroyed by fire. Wind-
blown sand had sifted through the roof and covered the bones. Sherds
similar to those with skeleton no. 3 were found near skeleton no. 4.
Mixed with the charred bones of the latter were hundreds of blow-fly
pupae cases that had been carbonized by the fire when the roof was
burned. The fifth skeleton was found 3 feet 4 inches from the west
wall, and 1 foot 6 inches from the north wall. The jaw was that of a
youth whose burned bones were found 2 feet above the floor, resting
upon the collapsed wall.
The presence of blow-fly pupae cases seems to explain why the
kiva was burned, and from the facts found the story of the tragedy
may be partially reconstructed. In time the natural earth walls of
the kiva probably became so weakened by the weight of the roof,
and from the absorption of moisture, that the west wall caved in.
Five men were trapped and lost their lives. That the kiva and its
unfortunate occupants were not entirely burned at the time of the
accident is manifest by the occurrence of the carbonized pupae cases
of the blow-fly, and can only mean that the structure was fired after
the men had been dead for some time. From the proximity of dwelling
rooms to the scene of the disaster, we might infer that the ruined kiva
was intentionally fired for sanitary purposes.
Pueblo custom decrees that the kiva be partly if not wholly under-
ground. Even among the Hopi, whose villages stand on rocky buttes,
kivas are often built on a lower sandstone ledge. With at least one
side abutting the cliff face the prescribed subterranean, or semi-
subterranean, position was thus realized without excavation of solid
rock—a tedious task for folk not formerly possessed of metal tools.
Returning to Kin Tiel, the masonry walls at the platform end of
Kiva KT-I were excellently constructed of hard, fine-grained sand-
stone. This sandstone, possessing definite lines of cleavage, could
with little effort be split into blocks of fairly uniform size. The
average size is about 2 inches thick and 10 inches long. Breaking of
joints and dovetailing of corners was practiced but neither occurred
consistently. The mortar was fine, sandy and sparingly used because
the flat surfaces of the building stones fitted easily together.
NO. IL PUEBLO RUINS IN ARIZONA—HAURY AND HARGRAVE 87
The floor of KT-I, and that of the platform as well, is of smoothly
worn irregular sandstone slabs averaging I inch in thickness and about
24 by 3 feet square. In the stone floor, 2 feet 9 inches from the east
wall, are five holes in line. They average approximately 2? inches
in diameter and 19 inches from center to center. On the opposite
side of the kiva is a row of six similar holes, slightly closer together,
this second series being 14 inches from the base of the platform,
whereas the first hole in the other row lies 2 feet 10 inches from the
platform. Of six other holes, five lie in the main part of the floor;
one, in that of the platform. Since these latter six were either plugged
with clay or a sandstone stopper, it is believed that they had been
abandoned and superseded by the two series above described. It is
also a possibility that the slabs in which they occur may have been
salvaged from other rooms and re-used.
A possible explanation of this belief may be the clay-plugged
hole which lies 4 feet 6 inches from the north wall and 3 feet 3 inches
from the east. The position of this hole, primarily, prompts a sugges-
tion that it might be a sipapu, but it is not in line with the firepit,
deflector, and ventilator, and it is too far removed from the firepit
which, itself, is not carefully oriented (fig. 24). A careful exami-
nation of the earth beneath this hole revealed only undisturbed soil ;
not the clay-lined cylinder anticipated.
From our observations it would appear that only the two series
of five and six holes, respectively, were in use at the time Kiva KT-I
was abandoned. As to their functions, two theories have been ad-
vanced. The first is that such holes were used to anchor the lower
end of a loom, a theory to which the writer subscribes. It is well-
known that Hopi men have long woven blankets in their kivas.
The second and quite improbable explanation is advanced by a
Hopi who admitted that while such holes are often used in fastening
the looms to the floor their real purpose is ceremonial. According to
our informant, holes such as those under discussion were designed
to hold freshly grown plants. The writer has not observed flag-
stone floors in modern Hopi kivas nor has he seen this arrangement
of round holes. In Hopi ceremonial chambers a square log or plank
with a series of small rectangular holes is buried in the floor on
cither side.” These rectangular holes are definitely made for weaving.
They may lie not only in the floor proper, but at either side of the
platform, and even at the ends of the kiva, and cut as they are in
*Mindeleff, Victor, A study of Pueblo architecture, Tusayan and Cibola.
8th Ann. Rep., Bur. Amer. Ethnol., 1886-7, p. 132.
88 SMITHSONIAN MISCELLANEOUS, COLLECTIONS VoL. 82
wood, they are entirely unsuited for potting plants. And yet it
might be that, in prehistoric times, such floor holes served this dual
purpose; that some tradition of that dual function has survived
until the present.
The firepit, 3 feet 14 inches from the base of the platform, is in
line with the ventilator and deflector, though not parallel to the latter.
Though the length of the sides is not consistent, averaging 1 foot 44
inches, the firepit approximates a square. It is 7 inches deep and
lined with clay-coated slabs of sandstone, but no evidence of the
corners being rounded with clay was found. The top of the pit
is flush with the floor of the room.
Seventeen inches from the firepit, and between it and the ven-
tilator, was a deflector—a sandstone slab with rounded top and
corners. The deflector is 14 inches thick, 2 feet 2 inches wide and
2 feet 1 inch high. It was set several inches deep into the floor and
so firmly that it was broken off just above the floor by the falling
roof. Though cleanly broken when found, it was still in an upright
position supported by fallen débris. The narrow space between
the deflector and platform was the only part of the kiva floor not
paved with flagstones. Beneath the clay-surfaced area loose earth
and rocks were found.
At the south end of the kiva, 19 inches from the deflector, was
the platform (pl. 21, fig. 1) which was 2 feet 64 inches high and 4 feet
wide. The vertical face of this platform as well as the natural walls of
KT-I were coated with plaster. Upon this plaster was a coat of white-
wash. As is usual in kivas of this type, fresh air was drawn in through
a vertical shaft outside the walls of the chamber, through a passage-
way beneath the platform and thence by means of an opening in the
middle front. In KT-I, this opening measured 12 inches wide by
22 inches high; its lintel consisted of two superposed sandstone slabs,
6 or 7 inches wide and separated by adobe mortar, making a total
thickness of 6 inches. The total length of the passageway from the
entrance in the face of the platform to the back of the air shaft was
5 feet 64 inches. The passageway, or ventilator duct, was roofed with
small sticks covered by a 44-inch layer of adobe mortar as a support
for the flagstones of the platform. Most of these sticks had decayed
but their imprints remained where the passageway had not collapsed
under the weight of the falling roof. Like the sides of the ventilator
opening, the duct walls were of undisturbed earth, heavily plastered
with clay. Ata point 2 feet 5 inches from the entrance was a plas-
tered step 1 foot high. This level continued back to the base of the
shaft where the width narrowed to 11 inches. Between the step and
the shaft the height of the passageway remained 10 inches.
ISKOE UU PUEBLO RUINS IN ARIZONA—HAURY AND HARGRAVE 89
The ventilator shaft was built of sandstone blocks somewhat
smaller than those used in construction of the room walls. The base
of the shaft, where it opened to the passageway, was slightly D-shaped
and averaged 14 inches in diameter. A short distance from the
bottom, the shaft became circular; at its present top, 7 feet 6 inches
above the duct floor, it was octagonal and 11} inches in diameter.
The upper portion of the shaft had collapsed with the wall against
which it stood.
The roof construction of KT-I may be approximated by the burned
material found in the northwest end of the chamber. Beams 3 to
5 inches in diameter had spanned the room and supported the cus-
tomary series of ceiling poles. Upon these was a layer of brush
and grass overlaid by several inches of adobe mud. The thoroughly
charred timbers had been broken into fragments rarely more than a
foot in length. Among these we recognized only one pine beam; all the
others were either pifion or juniper. About the firepit and deflector
were a number of sandstone slabs that obviously had fallen with the
roof. Their positions suggest that they probably rimmed the kiva
hatchway and that the latter, as in modern Hopi kivas, were above
the firepit. If this supposition is correct, then the chamber was en-
tered by means of a ladder extending through the hatchway and rest-
ing on the platform floor.
On removing the wind-blown sand and fallen roof, midden débris
was found in quantity, and in such position that it was obviously
not thrown through the roof entrance but through the hole left by
the collapse of the west wall. At this point the top of the midden
was 4 feet 2 inches above the floor, which, however, was not the
greatest depth of débris, since the bottom of the midden rested upon
the fallen wall. Transversely, the midden extended from the face of
the south wall to the north edge of the firepit (see drawing). From
the firepit to the north wall the roof rested upon the floor.
The midden fill of KT-I was principally of wood ash, scattered
through which were discarded stone implements, bone awls, turkey
and small-mammal bones, and quantities of potsherds of black-on-
white, corrugated, and a ware of a peculiar shade of orange decorated
in black. Differences and even local characteristics have been noted,
though a more comprehensive study of the sherds collected will have
to be made before these differences can be adequately defined.
While excavating KT-I, a flexed burial was found, though no
relation to the conditions in the kiva was indicated. In the north-
east corner, on top of the fallen roof and 2 feet above the floor, the
go SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
body was found against the wall and curved around the corner with
the head against the east wall, though facing west. The bones were
so well preserved that the skeleton was removed for later study.
Between the skeleton and the roof were two sandstone slabs in a
vertical position, across which the skeleton lay. The slabs appeared
to be part of the fallen roof support, rather than those of a burial
cist. In a horizontal position on top of the skeleton lay a flat, oval
stone I foot wide, and 1 foot 8 inches long upon which was a crushed
corrugated olla. The skeleton was found 4 feet 5 inches below the
top of the excavation in alternating layers of sand and human excre-
ment, overlaid by about 2 feet 6 inches of adobe. The kiva at this
point was filled with rocks that were once a part of adjacent dwelling
rooms.
ARTIFACTS
With the exception of the beads and flint point found with burial
number 3, the only artifacts found came from the midden. Among
these discards were three types of stone axes; single and double
groove encircling the head, and three-quarter groove. Manos were
of two types: oval with two flat sides, and triangular. Similar grinding
stones are in use among the Hopi today, each type serving a different
purpose. Metates of fine-grained stone, of which three were found,
are used with the triangular manos to produce the finest meal. Ham-
merstones, pottery polishing stones, pot lids worked from thin sand-
stone tablets, a pestle, wedge, an “arrowshaft straightener,’ two
stone mauls, and a triangular piece of sandstone grooved at two
corners and weighing about four pounds, were also found.
KIVA KT-II
A depression just southeast of KT-I proved on excavation to be
a second kiva, which we designated KT-II (fig. 25 and pl. 21, fig. 2).
The two ceremonial chambers are not exactly in line, however, since
the southwest corner of the platform in KT-I is only 12 feet from the
northwest corner of KT-II, while the southeast corner of the platform
is 15 feet from the northeast corner of KT-IIJ. This difference in-
dicates that the kivas, though facing the southwest, varied as to the
number of degrees, a condition frequently noted in structures of
this type. The interval between the kivas was not examined, but
from its size and position it is thought to contain a room.
In comparing the kivas only minor differences were noted. They are
of the same type and approximately the same size, though KT-II
OI
HAURY AND HARGRAVE
PUEBLO RUINS IN ARIZONA
NO. II
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g2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
was not so deeply set into the ground. The masonry of KT-II is of
the same type and material, but the method of construction is slightly
different. The walls at the southeast and southwest corners of KT-II
are joined together, but abut at the northeast and northwest corners
where wedges were driven between to strengthen the walls (pl. 22,
fig. 1). The tying of the walls in the two corners noted is well done
and appears to have been intentional. All four walls are of masonry,
plastered, and firmly constructed, contrasting greatly with the in-
secure walls of KT-I. The walls were straight and firm, with the
exception of the north and south sides which were slightly bowed
inward from pressure of the material against them.
The room floor consists of sandstone slabs of the same material
and comparable in size to those in KT-I. On the east side of the floor
are five holes, in line and averaging 11 inches from center to center.
On the west side is another series of holes, averaging 15 inches apart.
All holes are approximately 2? inches in diameter. That they were
probably for the purpose of fastening loom frames to the floor is in-
dicated by the fact that both series were immediately beneath roof
beams running parallel with the series. The charred remains of these
beams may be seen in the accompanying photograph (pl. 22, fig. 2).
No evidence of a sipapu was found.
The roughly squared firepit is 3 feet 34 inches from the base of
the platform, and is in line with the ventilator and deflector. The
pit, 10 inches deep and lined with clay, is flush with the floor; its
corners are rounded.
The deflector, a sandstone slab 1 foot 1o$ inches from the ven-
tilator, is 1 foot 10 inches high, 2 feet 44 inches wide, and 13 inches
thick. The top is slightly rounded and the base is firmly set in the
floor.
As in KT-I, the ventilator entrance is in the front side of the
platform, which lies 2 feet 6 inches above the floor. Across the
bottom of this entrance, 18 inches wide by 23 inches high, is a sill
4 inches high; across the top, a sandstone lintel. The roof of the
passageway had been composed of small twigs, placed close together
and covered with adobe as in KT-I. Over this roof, which is 3 inches
thick, are the flagstones of the platform, From front to back, the
width of the platform is 3 feet 6 inches (fig. 25); the length of the
passageway, including the base of the air shaft, is 5 feet 6 inches.
The sides of the passageway are of masonry, and still show signs
of plaster. In the passageway, 20 inches from the entrance, is a clay
step 1 foot high. The floor is paved from the entrance to this step,
NO. II PUEBLO RUINS IN ARIZONA—HAURY AND HARGRAVE 93
the remaining portion being of unplastered earth. The masonry walls
follow the level of the floor, rising at the step and inclining slightly
to the base of the air shaft which is D-shaped, with a diameter of 15
inches. Just above the base, the shaft is circular and 15 inches in
diameter; its diameter decreases toward the top. No idea of the
original height of the shaft could be ascertained since the south
wall, against which the shaft was built, had collapsed, falling on the
platform and breaking through the passageway.
Like its neighbor, KT-II was also destroyed by fire. Charred roof
beams were on the floor and though they were for the most part
completely burned, their position gives an idea of the arrangement
of the roof. Three feet, one inch from the east wall was a pair of
beams, each about 3 inches in diameter. Fragments of this pair were
found at intervals for the entire north-south length of the kiva. Along
the west side, 2 feet 114 inches from the wall, was another pair of
beams, also averaging 3 inches in diameter. These two pairs extended
over the platform. In the middle of the kiva and parallel to those at
each side was a third pair of beams, of approximately the same
diameters as the others. Between these paired timbers were single
poles but so completely burned that it was impossible to determine
their spacing or diameter. Extending from east to west at intervals
and resting upon the principal beams, were smaller logs, in pairs.
One such pair lay against the north wall, while another pair crossed
the kiva between the deflector and the platform. A third pair crossed
the kiva about 2 feet north of the firepit. These last two pairs prob-
ably sustained the added weight around the entrance which, in kivas
of this type, is in the ceiling over the firepit.
Evidence of the use of grass or brush in roof construction was
found, but only in the northeast corner could the order be determined.
Here was charcoal where the east-west timbers crossed the north-
south beams; upon the latter was a charred mat of grass or twigs,
covered with several inches of adobe. The clay of the roof was
covered with wind-blown sand, through which were scattered a few
potsherds, but the greater part of the depression between the fallen
roof and ground level was filled with rocks from nearby walls.
Between the ventilator and the deflector, the only portion of the
floor not paved with flagstones, were found three smooth stones, a
large flint core, and a stone maul about 3 inches in diameter and 5
inches long with a groove encircling the middle. Between the de-
flector and the firepit were fragments of a corrugated olla, shattered
by the fallen roof. The base of this olla lay near the firepit and still
8
94 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
contained pieces of small bones while part of the top and side was
found in the pit. The only decorated pottery found on the floor were
a few black-on-white sherds but as no two belong to the same vessel
we assume they had weathered from the mortar of the walls. On
top of the fallen roof, however, black-on-white sherds and other
fragments of an unusual shade, somewhat between brown and orange,
with black decoration were found mixed with the rocks and sand.
Sherds from KT-II, in general, are the same types as those from
Gigi
DATING THE KIVAS
When selecting Kin Tiel for excavation it was hoped that a good
selection of datable pine timbers would be recovered. The surround-
ing forests today consist almost wholly of pion (Pinus edulis) and
juniper (Juniperus monosperma) with here and there a lonely yellow
pine (Pinus ponderosa). We may infer that comparable conditions
obtained during the occupancy of Kin Tiel for, in the two rooms we
cleared, only one pine timber was found and this, unfortunately,
proved too complacent for dating. We may rejoice, however, in the
fact that Doctor Douglass and Mr. Haury, concentrating upon the
most promising of the material in hand, finally succeeded in determin-
ing the cutting dates of 27 pifion beams used in the two kivas.
Of this number, 13 were from KT-I and of these six were cut
between A. D. 1264 and 1270; one in 1274; four in 1275; and two in
1276. The collecting of building material usually requires some time,
at least among the Hopi, for never are beams plentiful in this region
even since the advent of the telephone pole. My personal observations
confirm this fact. We believe, therefore, that the building date of
KT-I is 1276 rather than 1275 A. D., the year of preparation ; that the
earlier dates evidence re-use of timbers.
Of the 14 dated specimens from KT-II one was cut in 1266; one
in 1272; nine in 1275 (four of these are from the same tree) ; two
in 1276; and one in 1285. It is obvious from the grouping of these
dates that the accumulation period was during 1275, and that KT-II
was therefore probably constructed at about the same time as KT-I,
sometime during 1276 A. D. It is a significant fact that KT-I had no
timbers cut later than its supposed building date, which, in view of
the unexpected collapse of the walls, suggests that it was destroyed
soon after construction—probably when the summer rains saturated
the sand walls. In contrast, KT-II was in use for at least ten years
before its final abandonment, as is shown by the date 1285 which
No. II PUEBLO RUINS IN ARIZONA—HAURY AND HARGRAVE 95
doubtless marks a bit of repair. That the longer life of KT-II is
due to better workmanship than that of KT-I is obvious from the
manner in which the latter was destroyed. This, and the fact that both
kivas were constructed during the same year, suggests that KT-I]
was built shortly after the destruction of KT-1.
SUMMARY
In brief, the season’s work at Kin Tiel has contributed to south-
western archeology (1) by furnishing plans and details of construc-
tion of two kivas—the earliest of the Hopi type yet reported; (2) by
determining the building dates of these kivas; (3) by showing the
Hopi relationship of a pueblo that traditionally has been considered
as Zufi; and (4) by securing new pottery types definitely associated
with dated beams.
KOKOPNYAMA
The prehistoric Hopi Pueblo, Kokopnyama, lies 1 mile east of the
Jeddito Trading Post, on the north side of Jeddito Valley and a few
miles south of the Indian Agency at Keams Canyon, in central Navajo
County, Arizona.
The Hopi mesas, of sandstone underlayed with shale, serve as
natural water reservoirs while the broad valleys on either side are
filled with alluvial deposits. Since the dip of the rocks is toward the
south, numerous springs dot the south side of the mesas. In a desert
region where permanent springs are few, such favorable conditions
for permanent homes were eagerly sought by the aboriginal inhabi-
tants and remains of their habitations are found in great number under
the mesa rims. Though soil and water are the most important, other
factors aided in making Jeddito Valley an important culture center in
prehistoric times. The valley floor was green with herbs and grasses ;
the bordering mesas were covered with shrubs and timber (Jumni-
perus monosperma and Pinus edulis). A few miles to the north
and east were pine (Pinus ponderosa) and fir (Pseudotsuga taxifolia).
Sandstone was available for building material; and deposits of clay
and veins of coal were visible along the mesa slopes.
The earliest historical reference to the Jeddito Valley is included
with an account of the discovery of Awatobi, visited in 1540 by Tobar
and Cardenas with a small detachment from the Coronado Expedi-
tion.’ Later explorers to visit this valley were Espejo in 1583. Ojfate
in 1598, and De Vargas in 1692.” In the first half of the 17th century
* Winship, George P., The Coronado Expedition, 1540-42. 14th Ann. Rep.,
Bur. Amer. Ethnol., 1896.
* Bull. 30, Bur. Amer. Ethnol., 1912, pp. 560-61.
96 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
a Franciscan mission was established at Awatobi and this thrived
until its destruction during the Pueblo Revolt of 1680. In 1700 an
attempt to re-establish the mission failed and Awatobi was destroyed
by the irate inhabitants of some of the nearby Hopi villages.’
Our first archeological reference to Jeddito Valley is from Victor
Mindeleff,” who in 1882-83 devoted much time to mapping the larger
ruins, of which there are five. These are all situated on the north
side of the valley and are well known to all students of Pueblo ar-
cheology as Awatobi, Kawaioku, Chakpahu, Nesheptanga, and Kokop-
nyama. In 1892 limited excavations were made at Awatobi by the
late Dr. J. W. Fewkes of the Smithsonian Institution,” “ who was
followed, in 1907, by Dr. Frank Russell of Harvard.
With the exception of the survey made by Mindeleff, no archeo-
logical investigations were made at Kokopnyama until 1901 when
Dr. Walter Hough of the Museum-Gates Expedition spent several
weeks in the valley.” In 1917 Spier, of the American Museum of
Natural History, undertook a pottery survey,’ and in 1923 and 1926
Kidder made a stratigraphic test at Nesheptanga and surface exami-
nations at other ruins." These are the archeological investigations
made prior to the spring of 1928, when the writer determined a
pottery sequence for Pueblo IV ruins in the Hopi country.
GENERAL DESCRIPTION
A surface survey of Kokopnyama reveals a ruin about ten acres
in area with architectural features not unlike those of modern Hopi
pueblos, if recent influence in the latter is disregarded. The general
plan is essentially the same with house groups two or more stories
in height surrounding open courts. Middens often contain pottery
types of different periods. From this condition we surmised that new
structures were erected in unoccupied portions when a building had
become unsafe for living. Later investigations confirmed this sup-
position. This shifting back and forth of buildings as decay set in is
found at the older inhabited Hopi towns. Having previously deter-
* Bull. 30, Bur. Amer. Ethnol., 1912, p. 561.
*8th Ann. Rep., Bur. Amer. Ethnol., 1887
* Hough, Walter, Ann. Rep., U. S. Nat. Mus., 1901, p. 333-
*Fewkes, Jesse Walter, Expedition to Arizona in 1895. 17th Ann. Rep., Bur.
Amer. Ethnol., Pt. 2, p. 592.
*Ann. Rep., U. S. Nat. Mus., 1901, pp. 279-358.
° Spier, Leslie, An outline for a chronology of Zufi ruins. Anthrop. Papers,
Amer. Mus. Nat. Hist., Vol. XVIII, pt. 3, New York.
"Kidder, A. V., Southwestern archaeology.
NO LT PUEBLO RUINS IN ARIZONA—-HAURY AND HARGRAVE 97
mined the sequence of Hopi pottery types, it was easy to select that
part of the ruin where beam material of the desired age might be
found. Only the presence or absence of charred or decayed pine would
affect the success of our 1929 work.
The accuracy of Mindeleff’s survey of the site was confirmed
through several tests, and having definitely located a point in the ruin
corresponding to a known point on his map, we established a per-
manent benchmark. From this point a base line was run through the
ruin. At convenient points cement monuments were erected from
which all tests and excavations were accurately plotted.
After selecting a section to be worked, it was found that Min-
deleff had considered only well-defined room outlines, probably con-
sidering the talus on the mesa slope to be fallen walls from rooms
above. Tests along the slope, however, revealed midden-filled rooms
with débris containing sherds of late Pueblo III and early Pueblo IV
types. The greater part of the pueblo occupied in late Pueblo IIT and
early Pueblo IV is then found to be outside of the ruin as mapped
by Mindeleff, the heavy lines outlined by him giving a fair idea of
the area occupied in late Pueblo IV (fig. 26).
DWELLING ROOMS
The distinctive characteristic of Kokopnyama masonry is the poor
quality of stone used. This material, Mesa Verde sandstone, is friable
and easily “ blocked,” but is unfit for “ dressing.” Also, because of its
softness, this stone is readily affected by weathering, which accounts
for the crumbled condition of exposed walls although Kokopnyama
is much more recent than many ruins of the Southwest whose walls,
excellently preserved, are constructed of more durable stone.’ Clay
used for mortar was dug from shale beds within the Mesa Verde
sandstone formation, and occurs in quantities under the mesa rim.
Of the rooms we opened all vary somewhat in size (fig. 27). The
normal thickness of walls is about 12 inches, though walls 24 or
more inches in thickness are not unusual where strengthening mea-
sures were found necessary. No orderly arrangement of blocks
was found, both large and small stones being used at random; nor
was any attempt at coursing apparent in those walls examined. The
granular surface of the walls was protected by coats of natural, yellow-
clay plaster—as is common today among the Hopi. Plastering seems
*22nd Ann. Rep., Bur. Amer. Ethnol., 1904, p. 134.
a “VY Of srt a}e
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VOL. 82
~
EVAR OL
SMITHSONIAN MISCELLANEOUS COLLECTIONS
No. ITI PUEBLO RUINS IN ARIZONA—HAURY AND HARGRAVE 99
to have been confined to the interiors of rooms at Kokopnyama, how-
ever, and though some Hopi dwellings with plastered exteriors are
found, they are few—the practice having been introduced into this
Size of Rooms
Mealing bin
Rooms | Length | Width | Depth | Door | Cache | Firepit
| eee ee ee
R-2 1102 Wy IE = Yes | |
R-3 6’ 9” ie Yes
R-6 aGe els) 2
R7 Ti oad NE
R-8 | 7'4"| 5/6”
Gr iG: | 5°40
R-9 SAS TM Dad OR: Sine
6’ 6”
R-1o | 10’ Grr: Yes
8” 6”
m-10a!| 8 6” | G 1” Yes
R-11 Bim0 |) 167 Was. |. Yes Ves Yes
R-13 ‘Cua Cia Me. 6 Yes
R-13a ? ? BG" Extends under R-11
im-i4 | 11’ 4” 9’ 2”
9” | 8
R-15 y aa intro 2
9
R-16 Geir) 319"
R-17 8 8
R-18 | 10’ Ge ; Mesa) Wes
R-g9a 8’ 8
R-21 3 Sp
= aa ie i Yes
R-25 | 14° Z 6” 6
| : |
BIG. 27%
region within recent years. Decoration of walls with painted designs,
or with incised drawings was not found, though incised stones were
found detached from the walls. Both methods of decoration have
been found at Kawaioku, a nearby ruin occupied at the same time as
I0O SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
Kokopnyama, where Hough records a wall elaborately decorated in
color, and the writer found geometric designs scratched upon the wall
plaster of a kiva.
DOORS
We found only seven doors, a surprising fact considering the
number of rooms opened. But when it is realized that until a few
years ago practically all entrances to first floor rooms of historic Hopi
pueblos were made through the roof, this condition will be understood.
Even today in Oraibi, the oldest inhabited Hopi village, many rooms
now partly buried by accumulations of sand and deébris are still en-
tered in this manner. There is hardly a room, either above or below
ground, in Oraibi or villages on the Second Mesa, that at some time
Doorways
Width | Height
f z 4 offsets offsets
Room Type Width |Height ee Lintel A Remarks
| Left Right) Left |Right|
R-3 | Rect. ete 44 2h oa tO aS ticles | |
R-1oa | Square | 1’ 9’ Toe — 6” | Stone
| / } Cy: | Base of door is
= ~ , 7 af 147 as =H |
R-11 Rect. | 6 2a | Sticks Acomleuel
Aen ae Th 4” 2! em 110" = | “eé
io | Partly de-
en |) op G hacer: 2! ? |
R-2 (aiae! Were WI SE hai : | | stroyed.
| | |
R-25 | T-shape | 2’ 2” 3/4” | 2’— | Stone | — 6” |— 43”| 1’ — | 1’ —
Fic. 28.
the writer has not been permitted to enter. It was observed that in
old dwellings on the ground floor which have for years been used
for storage, entrance was usually made through the ceiling. The
doorways at Kokopnyama were square to rectangular, or T-shaped.
Dimensions in all vary (fig. 28). Lintels were either of split juniper
sticks about 14 inches in diameter set in adobe mortar, or of sand-
stone slabs.
CACHES
Caches used both for domestic and ceremonial purposes were found
and the interiors of all were plastered. In some caches, corners were
rounded with plaster and sometimes emphasized until the cache was
* Archeological Field-Work in Northeastern Arizona. The Museum-Gates
Expedition of toot. Ann. Rep., U. S. Nat. Mus., 1901, pp. 270-358.
No. II PUEBLO RUINS IN ARIZONA—HAURY AND HARGRAVE IOI
oval or even circular while in others rounding was limited to the
upper rear corners, producing an oval top. More often the lintel
was a sandstone block of the wall masonry (fig. 29), but an excep-
tion is found in Room 11, where a cache is roofed with small sticks
set into adobe after the manner of door lintels. Time has proved the
strength of roofs made from adobe and wood, and it is not surprising
that this knowledge should be used where a substantial covering for
an opening is needed whether large or small.
Caches
| x
We ts S
- 3 :
= ° = = = S mS 8 Remarks
5 Sele ae: ee hs s = S
o | 6 = 3 © 8 & < 8
Ramee) Sd S a Q a, i) S ce
R-2 1’ 5” |— 6” |— 6” | — 6” | Yes | Stone | N. a’ 9’-N. E.| Circular
R-11 | 2’ 6” |— 8” | — 7” | 1’ 1” | Yes | Sticks | W. Near middle of room.
R-15 | 2’ 3” | —8” | —73”| — 8” | Yes | Stone | S. E. 7i”_S.W.| Slightly rounded.
R-15 | 2! 3” me 9” =o ae 8” Yes 66 S. E. xe GA W. “cc “
Rag |x? =| — 5’ |—4” |— 6” | Yes se N. W. 4’-N. No depth at top. Slopes
backward and down to
bottom.
R-23 | 2’ 1”|— 7” | — 6” |— 5”| Yes OU N. W. | 3’ 6’-N.W.| D-shape. In kiva.
R-23 ? ? ? ? Yes 2 N. W. ry No measurements.
R-24 | —— | 1’ 6” | 2’ 7” | 1’ 3” | Yes | Stone | N. W. 7”-N.W.| Top oval. Slopes down
ward.
R-24 | — 8” |— 6” |— 5” |— 8”! Yes wo N. W. |2’ 107-S.W.]| D-shape. Base flat. In
kiva.
Fic. 209.
FLOORS
Floors were of clay, with the exception of those in kivas and in
Room 10 which were of stone, and were found in all rooms, whether
on bedrock or midden fill. These floors were easily identified by their
smoothness, hardness, and thickness which varied from one to several
inches.
FIREPITS
Nothing characteristic was noted in clay floors, except the position
of the firepit, which was invariably in a corner, or against the wall
2 or 3 feet from the corner. This position does not apply to kiva
102 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
fireplaces, which will be discussed later. Firepits were made with
sandstone slabs set below the floor surface and plastered, or by
plastering the sides of a hole within the floor itself. Dimensions vary,
as shown in the accompanying table (fig. 30).
Fire pits
Room||Length| Width| Depth Sides Corners Remarks
R-1o 1’ — | —11”| — 8” 3 of Clay Slightly rounded
1 of Stone
R-18 Toe ten e ? Clay Rounded Against S. E. wall 4’ 7
from S. W. wall.
R-19 Ty 20a —or ore ? Clay Slightly rounded | In N. W. corner.
TGs 530:
MEALING BINS
Mealing bins were found in approximately the same position as
firepits, 7. e., against the walls, but confusion in identity is not likely
since firepits are usually lined with clay, contain ashes and extend
below the surface of the floor, whereas mealing bins are not lined
with clay, contain no ashes, and are built above the floor. In two
rooms bins were found in the corner, the walls serving for two sides
of the bin, while in one room a bin was constructed against the south-
west wall, 4 feet from the nearest corner. All mealing bins found were
larger than the firepits (fig. 31).
Mealing Bins
Room | Length | Width | Depth eee Ne er Remarks
R-13 i’ 10” mes 4g’ —1” 2 In N. W. corner, two walls serve
as sides,
R-19a gv | i — — 9” No 3 N. E. wall 1’ 6” from S. E. wall-
R-22 pi Rt Tes 1’ — No 3 Along S. W. wall about 4” from
S. E. wall.
R-11 2 At Tena — 11” —1” 2 In S. E. corner two walls serve
as sides.
EiGee sie
CEILINGS
Roof material was found in a number of rooms, but usually in
such a poor state of preservation and in such small quantities that
No. II PUEBLO RUINS IN ARIZONA—-HAURY AND HARGRAVE 103
only a faint idea of the ceiling arrangement could be inferred. Rooms
g and II are the exceptions. In Room 9g the ceiling was so perfectly
preserved that every step in construction was clearly defined. This
room is of average size, which increases the value of the illustration,
permitting the use of this roof as a typical example of those for
house dwellings. In a north-south direction were two large beams of
juniper and pifion of 6 and 4 inches diameter, respectively, support-
ing a framework of three small poles. These latter, averaging about 3
inches in diameter, were approximately 12 inches apart and were near
the middle of the room (fig. 32). Many pieces of split juniper were
placed between the east-west cross-poles, thus forming a ceiling upon
which brush or grass was laid. The roof was finished with a layer
of adobe. Beam holes in the walls naturally vary in size according
to the diameter of the beams used. Only the beams were imbedded
in adobe pockets in the walls, the split juniper sticks extending to
the wall surface. From the beams to the room floor was 4 feet 4
inches ; or, if one includes the diameter of the main beams, the ceiling
height would be 4 feet 10 inches. As there was no door through the
walls, the entrance must have been through the roof at the east end,
the only side not covered by the roof when excavated.
The ceiling of Room 11 was largely destroyed by fallen rocks, but
fully one-third remained in such excellent condition that comparison
with the roof in Room g revealed the same kinds of material and
same method of construction.
KIVA R-4
The best preserved kiva we found at Kokopnyama is indicated on
the map as R-4, and may be seen in plan on figure 33. The kiva is
built facing southeast on a sandstone ledge just below the mesa rim.
There was little or no soil on the ledge, but the section was once
covered by earlier house structures, the tumbled walls of which were
sufficiently deep to favor the building of a kiva. This location is
similar to that of many present-day Hopi kivas, and is in keeping
with the current Hopi custom of building their ceremonial chambers
on a lower ledge, if sufficient depth of soil cannot be found on the
ground level.
Three sides of kiva R-4 were covered either by abandoned rooms
or household débris but the fourth was exposed, being built on the
edge of the ledge. The greater part of the southeast wall had weathered
away but the remaining portion of the platform was easily recognized
and provided a starting point for excavating the kiva. Above its
fallen roof the chamber was filled with fallen masonry through which
were occasionally found potsherds of Jeddito black-on-yellow and
104 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
UCC CCC LU
Fic. 32.—Diagram of roof structure in R-9.
NO: IL PUEBLO RUINS IN ARIZONA—HAURY AND HARGRAVE 105
Sikyatki polychrome. There is no reason to believe that the kiva be-
came a refuse dump after it ceased to serve for ceremonial purposes
since the sherds were too few and scattered, and were not always
associated with ash deposits. It is believed these sherds were included
in the roofing clay.
That the kiva was used after it was abandoned for ceremonial pur-
poses, however, is clear, for the rear had been made into a small
dwelling or storage room, by building in a cross wall, partitioning an
area approximately 10 by 6 feet. The floor of this small room was 5
feet above that of the kiva and was made by leveling the accumulations
of fallen wall masonry that lay beneath it. The northwest and south-
west walls of the kiva were utilized in the smaller room, though an
auxiliary wall was built against and parallel to the northwest side.
These later walls were of inferior construction and were built of
irregular sandstone blocks, chinked with smaller pieces of the same
material set in thick clay mortar.
All walls of the kiva were of roughly shaped sandstone blocks
averaging about 4 by Io inches, and were set in thick mortar as in
the walls of dwelling rooms, which only differed from kiva walls in
that they were thinner and built of smaller stones. The kiva walls
were heavily plastered, there being 32 coats on the northeast wall,
the 29th of which was red. In the northwest wall, 4 feet 9 inches from
the northeast side and 3 feet 7 inches above the floor, was a badly
decayed wooden peg set in a hole approximately 24 inches in diameter.
In the southwest wall and near each end, had once been two more
pegs of about the same size and distance from the floor as that noted.
Only small pieces of the decayed wood were found in these holes. It
is presumed that these pegs were for hanging ceremonial paraphernalia
or other objects, as in modern Hopi kivas.
Originally, the kiva was much larger than at the time of abandon-
ment. On two separate occasions its dimensions were reduced by
strengthening walls (fig. 33). The original dimensions of the kiva
room, exclusive of the platform, were approximately 14 feet 6 inches,
by 114 feet. The back of the kiva, or the northwest wall, originally
was the plastered face of a midden, in which the kiva had been partly
excavated, and apparently served for several years since it was plas-
tered four times. That the “ midden wall” weakened is indicated by a
slight bow in the middle, which was strengthened by building against it
a masonry wall about 8 inches in thickness which was in turn further
strengthened by a second masonry wall constructed at a point 3 feet
5 inches from the “ midden wall,” and ingeniously arranged with a
106
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When first constructed, the rear wall of R-4 was the plastered face of a refuse heap,
like KT-I at Kin Tiel, and threatened another disaster which was timely averted by an
ingeniously arranged brace wall.
No datable beam material was collected.
No. It PUEBLO RUINS IN ARIZONA—HAURY AND HARGRAVE 107
brace wall of stone between it and the first masonry wall, the inter-
vening space being filled with loose rocks and earth (fig. 33 and pl. 23,
fig. 1). This was the northwest wall of the kiva when it was aban-
doned. Under the plaster coats, of which there were several, and im-
bedded in the mortar of the wall were two sherds of Jeddito yellow
ware. One of these was an undecorated surface, while the other
was an excellent example of Jeddito black-on-yellow. The northwest
wall showed no further evidence of weakening, but the southwest
wall eventually weakened, and was reinforced by building another
masonry wall against it. The new southwest wall is 20 inches thick
and wider than the offset on the southwest side by 5 inches.
A comparison of the various sizes of the kiva at different periods
of remodeling may be made from figure 33. On the northeast side the
offset is Ir inches. After the construction of the new wall on the
southwest side there remained only the offset on the northeast, but
by removing some of the masonry of the new wall the original size
of the offset on the southwest side was found to be 15 inches. Origi-
nally the platform was g feet 10 inches long, but this length was re-
duced 5 inches by extending the extra width of the auxiliary southwest
wall across the end. The platform is 3 feet 10 inches wide, and its
floor is 21 inches above the floor of the kiva room. Though the south-
east wall is missing to the level of the platform floor, the platform
walls and face are of similar material and construction to the walls
of the kiva room. The kiva, being on a ledge below the mesa top
and surrounded on three sides by tumbled walls, was constantly sub-
ject to erosion from the slope above. As a result of this action the
southeast ends of the kiva walls were reduced on the sides of the
platform to only a foot or two high, and the southeast wall was re-
duced to the top of the platform. At the top of the wall in the north-
west corner were two layers of masonry extending for several inches
over the plastered wall of the midden excavation, and since the depth
of the kiva at this point is 8 feet 4 inches, it is probable that this
represents the original height of the kiva. The height of the kiva at
the northwest corner of the present end wall is 7 feet 8 inches, but the
weathered slope of the walls from beyond this point to the southeast
wall indicates that the original height of the kiva was greater.
The floor was covered with flagstones of varying sizes, with the
exception of the space between the ventilator entrance and the de-
flector, which was of hard packed rock and earth. As noted in the
kivas at Kin Tiel, this space represents the hole dug in setting up
108 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
the deflector. About 6 inches below the level of the kiva floor in the
earth and rocks, was found a Jeddito black-on-yellow sherd. The
platform also was originally paved with flagstones but several were
removed by our Indian workmen before the error was discovered. The
only flagstones now remaining on the platform are at the southeast
end. On the floor of the kiva 2 feet from the northeast wall and 3
feet from the base of the platform, is a line of four holes with an
average diameter of 2 inches, which are 19 inches from center to
center. The belief that these holes were used in weaving is supported
by finding two “ loom blocks ” in the kiva (pl. 23, fig. 1). These blocks
are of sandstone and are about 114 inches long, 8 inches wide, and
634 inches thick. Sandstone blocks similar to these, but longer, and
with hand holds pecked in each end are found in modern Hopi kivas
where their uses also vary. At the present time so little weaving is
done that they are used primarily for seats. On the southwest side is
another line of four holes, 4 feet 1 inch from the platform, and 1 foot
from the southwest wall. These holes are slightly larger than those
on the northeast side, being 24 inches in diameter, and are 18 inches
from center to center. All holes on the southwest side were plugged
with clay, which may be accounted for by their nearness to the south-
west wall which was constructed after the kiva floor had been laid.
Two other holes on the same: side, and near the holes in line, were
also plugged with clay, indicating that when the kiva was abandoned
none of the holes on this side were being used. In the rear of the
kiva and parallel to the northwest wall were two more holes, 2 inches
in diameter, that were likewise plugged with clay. The position of
these holes—z2 feet from the northwest wall and almost in line with
the firepit and deflector—made it appear that one of them might have
been the sipapu, but this could not be definitely determined.
The deflector is a sandstone slab 14 inches thick, 20 inches wide,
and standing 16 inches above the floor. The top was rounded into
an arc. Seventeen inches from the ventilator entrance, the deflector
was set 9 inches into the floor. An interesting feature of the de-
flector is its relation to a firepit of which it was part. At each side
of the deflector was built an arm of sandstone and adobe that ex-
tended at right angles to the deflector, the whole resembling an arm-
chair without legs (pl. 23, fig. 2). Each arm is 12 inches long, 5 inches
wide, and 7 inches high. Between the arms is a peculiar firepit—
peculiar in that it has two floor levels, the first level with the floor,
the bottom being a sandstone slab, the two arms and the deflector
serving for three sides; the fourth side is open. Five inches from
NO. TI PUEBLO RUINS IN ARIZONA—HAURY AND HARGRAVE 109
the deflector and parallel with it is a pit 12 inches long, 74 inches
wide, and 10 inches deep, the bottom of which forms the second
level of the firepit. This deeper part is filled with ashes, but its
exact function is not known.
A second firepit 64 inches from the one described (pls. 23, fig. 2, and
24, fig. 1) is 12 inches square and 13 inches deep. There is nothing
unusual about this pit, which was clay lined with hand rounded cor-
ners, and in line with the first firepit, the deflector, and ventilator.
Like the first firepit it was fitted with a sandstone cover, and was
filled with wood ashes. The top is level with the floor.
Near the center of the platform is the entrance to the ventilator,
which is 14 inches square. Over the top, and sustaining the weight
of the platform floor is a lintel of sandstone slabs set in mortar. The
bottom of the lintel is a single slab, upon which are two shorter slabs,
placed end to end. From the platform to the bottom of the lintel is
7 inches. The length of the passageway is not known since it is partly
eroded near the southeast wall; it probably extended through the
platform as in the kivas at Kin Tiel. The length of the part remaining
is 3 feet 2 inches. For 7 inches back from the entrance, the floor of
the passageway is formed of a sandstone slab; but at a point 5 inches
from the entrance there is an upright slab 11 inches high, behind
which were found several lumps of red paint, two manos covered
with paint, two hammerstones, half of a Jeddito yellow bowl con-
taining fragments of small bones, and a piece of chert, all of which
were on the sandstone floor which extended about 6 inches beyond the
upright slab. Covering these artifacts and reaching to the top of the
upright slab was an unsmoothed floor of clay that gradually sloped
upward and back to the end of the passageway. Between the level of
the upright slab and the roof of the passageway was a distance of
6 inches. Though it could not be definitely determined that the up-
right slab with the clay behind it was a step reducing the height
of the passageway, as found in the kivas at Kin Tiel, still there is a
remarkable similarity both in principle and execution. The presence
of the artifacts behind the slab and beneath the clay floor might,
however, indicate that the slab or “step,” was at one time farther
back in the passageway—at the end of the sandstone floor possibly—
if there was an abrupt change in the floor level. Owing to erosion,
conditions near the rear of the passageway were unfavorable for ac-
curate notations. The floor of the platform was broken over the
passageway by the fallen roof of the kiva, but enough of the passage-
way roof remained intact to show the order of construction, which was
9
110 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
the same as that noted in the kivas at Kin Tiel, namely, a layer of
small sticks across the passageway covered by a thick layer of adobe.
In order to examine the interior of the passageway, however, it was
necessary to remove the broken floor, but after completing the investi-
gations the floor was replaced and the eroded section of the platform
rebuilt. In addition to this repair the walls were strengthened by re-
placing with cement part of the adobe mortar between the blocks.
Pressure from débris against the outside of the wall was partly
relieved by removing much of the material, and drainage was pro-
vided by trenching around the sides. No provision was made to drain
the interior of the kiva, since a crack in the bottom of the pit by
the deflector was thought to be large enough to carry off rainwater.
It is planned to make more permanent repairs at a later date.
A number of beam specimens were collected, but since the kiva
was not burned and conditions were unfavorable for preservation,
they were too badly decayed to give a comprehensive idea of the roof
construction. All specimens were either pifion, juniper, or cotton-
wood, and at the time of writing none of the specimens has been
dated.
KIVA R-27
The northeast wall of Kiva R-4 was the southwest wall of an-
other room, R-27, which when partly opened proved to be another
kiva. Due to lack of time this latter was not completely excavated,
though sufficient material was removed to reveal the deflector and
edge of the platform.
KIVA R-24
Previous to the discovery of Kiva R-27, another kiva, R-24, was
opened and studied (pl. 25, fig. 1). Both R-24 and R-27 are on the
sandstone ledge with R-4, and all face in the same general direction.
As was shown in the case of R-4, kivas on the ledge are most sus-
ceptible to erosion at the southeast end, so it was not surprising to
find that only the face of the platform remained intact. The weather-
ing of the slope had reduced the walls at the edge of the platform to
15 inches in the northeast corner, and 2 feet 6 inches in the south-
east. Though complete notes could not be taken at this end of the
kiva, the ventilator passageway remaining showed evidence of having
been roofed with sticks and adobe, while the entrance to the ven-
tilator was covered with a sandstone lintel.
Walls were of the same material and construction as those in R-4,
and were heavily coated with plaster, being about 14 inches thick.
NO: LE PUEBLO RUINS IN ARIZONA—HAURY AND HARGRAVE EEE
The northeast wall was 10 feet 4 inches long as contrasted to 11 feet
2 inches for the southwest wall. The same lack of consistency was
found in the length of the northwest wall and the corresponding width
of the kiva between the walls of the platform face. The height of
the kiva could not be determined, though those corners with less evi-
dence of erosion are the northwest and southwest, which are 6 feet 10
inches, and 6 feet 2 inches respectively from the kiva floor to the top
of the remaining walls. The vertical side of the platform is of
masonry and extends to the side walls, but on account of the weathered
condition it could not be determined whether there were offsets be-
tween the platform and walls.
In the northwest wall are two plaster-lined caches. The first of
these is 7 inches from the northwest corner, and on the same level
as the base of the cache and floor of the kiva. The height of the cache
from the floor to the middle of the top, which is an arc, is 2 feet 7
inches. The depths and widths at the top and bottom differ. The
bottom is 18 inches wide, the top 11 inches at the floor level, the depth
15 inches, while the top is oval and slopes downward toward the
rear and base of the cache. An unusual arrangement in this cache is
a shelf, 17 inches above the floor, composed of three cross sticks of
split juniper about I inch in diameter upon which rest two sandstone
slabs, one upon the other, the thickness of the shelf being about 3
inches. On the floor of the cache was found a large piece of gypsum.
The second cache is at the opposite end of the northwest wall, 2 feet
10 inches from the southwest corner. It is D-shaped with the flat side
at the bottom, which is 8 inches above the floor of the kiva. The cache
is 6 inches wide, 5 inches high, and 8 inches deep. Within the cache
was a quantity of red paint.
The floor of the kiva was paved with sandstone slabs, except be-
tween the deflector and the ventilator, where it was of clay ; many of
the flagstones were smeared with red paint. The writer has never
heard of kiva floors being painted, and from the quantity of red
paint found in the cache, he leans to the belief that possibly the sand-
stone flags of the floor were used for powdering the lump paint.
Along the northeast side of the floor, and 18 inches from the northeast
wall, was found the usual line of holes drilled into the stones of the
floor. These holes were four in number, each with a diameter of 24
inches, and averaging 17 inches from center to center. On the south-
west side was another line, also of four holes, 1 foot 3 inches from
the southwest wall; 1 foot 5 inches from center to center; and aver-
aging 2 inches in diameter. These holes on the southwest side were
I1I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
plugged with clay, as were also two other holes between the first hole
of the line and the firepit (fig. 34). The firepit might be said to be
in line with the deflector and ventilator, though it does not “ center ”
as in other kivas, the northeast side of the pit being almost in line
with the northeast edge of the deflector. The firepit is 4 feet from
the northeast wall and 18 inches from the deflector; is Io inches
square with slightly rounded corners, and 18 inches deep. It was
filled with wood ashes. Over the top was a sandstone cover 1 foot
square. The deflector was 18 inches from the ventilator entrance, or
midway between the ventilator and firepit. As in other kivas de-
scribed from this ruin, the deflector was a sandstone slab set firmly
into the floor. There was a decided lean toward the platform, how-
ever, which might have been caused by the falling roof. The de-
flector was 1 inch thick, 21 inches long, and 18 inches from the floor
to the top. The corners were rounded. On the northeast side of the
deflector was a loom block, while another was found near the south-
west corner of the room. Beneath the fallen roof and scattered on
the floor near the deflector and firepit were fragments of Jeddito
black-on-yellow and Jeddito corrugated ware.
In the southeast corner of the room and buried beneath the floor
was found a cache of two Jeddito black-on-yellow ollas. Both were
filled with a coarse sand identical to sand found on ant-hills of the
region. Inquiry among the Hopi, however, failed to verify the be-
lief that sand from ant-hills was used for ceremonial purposes, as
the discovery seemed to indicate. Both ollas were neatly protected by
a sandstone cover beneath the flagstones ‘of the floor.
KIVA R-23
It will be noted that all kivas heretofore described in this paper
have been of the Hopi type, and the consistency in plan and orienta-
tion would lead one to expect all other kivas of the same period and
region to be of the same arrangement. This was found not to be the
case, however, for on excavating the room northeast of R-24, a kiva
was disclosed that differed greatly in several details. This kiva, R-23
is rectangular (fig. 35), faces north, and has no platform, all of which
are radical changes ; other departures from the usual arrangement will
be noted in order. The material and construction of the walls is es-
sentially the same as others described and the floor was paved with
flagstones. This kiva, also located on the sandstone ledge below the
mesa top, is therefore subject to erosion, and on account of the orienta-
tion, the southeast side was affected more than that in the other two
NO. LT PUEBLO RUINS IN ARIZONA—HAURY AND HARGRAVE IT3
Os F 2 FEET
Fic. 34.—Plan of Kiva R-24.
Beneath the floor of R-24 were found two Jeddito black-on-yellow jars filled with
coarse sand which was probably reserved for ceremonial purposes. In the rear wall were
two caches which are said by the Hopi to have been used in the Bean Ceremony. Kiva
R-24 was constructed in 1380 A. D,
II4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
kivas, resulting in the complete destruction of the southeast wall and
part of the flooring on that side. The northeast and southwest walls
were also partly destroyed at the southeast ends, so that complete
measurements could not be taken. The remaining portions of these
two walls are 8 feet 2 inches, and 7 feet 10 inches respectively. The
northwest is complete, though badly bowed from the weight of the
débris on the outside, and is 10 feet 3 inches long. With the tops of
the walls weathered, the ceiling height of the kiva could not be de-
termined, though it is known to have been more than 4 feet 9 inches
as determined from the present height of the walls in the northwest
corner. In the northwest wall, 3 feet 6 inches from the northwest
corner of the room, is a D-shaped cache 2 feet high, 1 inch above the
floor. The cache is 7 inches wide, 6 inches high, and 5 inches deep.
In the floor of the kiva, 3 feet 6 inches from the northeast wall, 1 foot
3 inches from, and parallel to the northeast wall, are four clay-plugged
holes, 2 inches in diameter, and averaging I foot 5 inches from center
to center. Three feet 9 inches from the northwest wall, and 3 feet 8
inches from the northeast wall is the sandstone deflector, 14 inches
thick, and 18 inches wide. The height of the deflector is not known
since the deflector was broken off 9 inches above the floor, nor could
the missing top be found.
One foot 7 inches from the deflector and near the center of the
kiva is a firepit of plastered sandstone slabs, with slightly rounded
corners (pl. 26, fig. 1). The northwest side of the pit is 10 inches long,
the northeast side is 12 inches; the other two sides correspond in
length to their opposites. The depth is 21 inches. Between the deflec-
tor and the northeast wall is another firepit which lacks the regularity
of the former. The southeast corner of the pit is 6 inches from the de-
flector as contrasted with 3 inches from the southwest. A better idea
of the shape and position may be gained by referring to figure 35.
The dimensions are 17 inches for the southwest side, 11 inches for the
northeast, 12 inches for the northwest, and 10 inches for the south-
east. The sides are of sandstone slabs, plastered and with slightly
rounded corners. For a depth of 6 inches from the top of the pits,
and level with the floor of the kiva, was wind-blown sand under which
were ashes. Lying on top of the wind-blown sand in the second fire-
pit were Jeddito black-on-yellow and Sikyatki polychrome sherds.
The ventilator is not as truly aligned with the deflector or firepit
as in R-4 and R-24, nor does it conform specifically with other ven-
tilators examined in Hopi type kivas. The entrance to the ventilator
is in the face of the northeast wall 3 feet 6 inches from the northwest
wall, and was originally 8 inches wide but has been reduced in width
by partly filling the opening with clay. It is 9 inches high, 5 inches
NOS EE PUEBLO RUINS IN ARIZONA—HAURY AND HARGRAVE 115
deep, and is roofed with sticks covered with clay. The base of the
entrance and passageway is level with the kiva floor, and is of clay
(pl. 26). This short passageway—if it may be so termed—is built into
the masonry wall of the kiva and connects with a circular airshaft 6
inches in diameter that extends upward at right angles to the passage-
way. The airshaft is a hole running vertically through the middle of
the wall from the ventilator passageway to the remaining top of the
wall, which is 3 feet above the floor of the kiva. The airshaft is strik-
SCALE
| oamanal hanna acs ae
om wae | 2 FEET
Fic. 35.—Plan of Kiva R-23.
_ Kiva R-23 is the only kiva excavated in which the ventilator is on the north side. In
this instance the entire passageway and airshaft are within the north wall. From among
the coals in the firepit a datable specimen gives the year 1416 A.
ingly like an ordinary flue. In a small midden-fill on top of the roof
clay were sherds of Jeddito black-on-yellow and Sikyatki polychrome.
Further investigations of conditions concerning this kiva might
determine it to be the rear of an abandoned kiva remodeled for cere-
monial purposes, or another type of kiva used contemporaneously
with the Hopi type. It is possible that the northeast and southwest
walls can be followed along their bases for a sufficient distance to
settle this question. It should be remembered, however, that the
principal object of our limited excavations was search for beam speci-
116 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
mens, so the writer could not always investigate conditions as thor-
oughly as he would have liked.
BEAM SPECIMENS
As has been previously stated, the primary objective of the expe-
dition was to secure datable beam material. With this in mind Kokop-
nyama was examined in 1928 and, though the site proved prolific in
the desired pottery types, little hope was felt for the discovery of pine
timbers, since the predominating woods today are juniper and pinyon.
Our activities were, therefore, directed to Showlow where pine trees
were still growing within a short distance of the ruin, and where pre-
vious examination had revealed that the ruin was destroyed by fire
and pine charcoal was abundant.
After the memorable discovery of HH-39, which “ bridged the
gap,’ my attention was directed to the less favorable Hopi country in
hope of substantiating the Showlow results from a widely separated
district. The chance was admittedly slim since the mesas are devoid
of pine, but having previously discovered several datable specimens
from Kawaioku, a pueblo of later date, some slight encouragement
was gained, and it was hoped that the earlier pueblo of Kokopnyama
might fulfill our needs. At this time Doctor Douglass had not recog-
nized pinyon as reliable material, and it was not until much laboratory
work had been done by Mr. Haury that Doctor Douglass realized the
value of pinyon, and that once familiar with its characters, absolute
dates could be determined. This discovery was joyful news to me,
for I had begun to feel that my efforts at Kin Tiel and Kokopnyama
were wasted, since only a few unfavorable specimens of pine had been
found. I had collected a number of pinyon specimens, both of wood
and charcoal, and with this latest development, I knew that not only
could Kin Tiel and Kokopnyama be dated, but that countless other
ruins situated in similar desert regions would eventually give up their
SECTetS:
Unlike the Showlow ruin, Kokopnyama was not burned, which
further lessened our chances, since the preservation of wood is de-
pendent upon many factors that are rarely encountered in the proper
combination. To date 42 specimens from Kokopnyama have been
dated, 20 of which are charcoal, among which was only one pine speci-
men. The remaining 22 wood specimens were all from dwelling or
storage rooms and gave the following dates: 1380, 1430, 1383, and
1389 ; two of 1369, four of 1370, and one of 1399; three of 1400, and
one of 1416; one of 1400; one of 1255; one of 1371; and two of 1269.
These dates are grouped according to their association.
NO. II PUEBLO RUINS IN ARIZONA—HAURY AND HARGRAVE DL,
With the exception of one specimen of charcoal found in a refuse
heap and which was dated in 1928° all others were from kivas. Kiva
R-24 was the only one in which dated roof beams were found. Seven-
teen of these dated 1380, two others giving the years 1362 and 1368
respectively. The great number of specimens dating 1380 A. D.
would indicate that the kiva was either constructed in that or the
following year. The remaining charcoal specimen was found in the
firepit of Kiva R-23 and dated 1416, proving that the kiva was in
use after that year. Many specimens from Kokopnyama still remain
to be dated.
POTTERY CHRONOLOGY
Though several expeditions were made to the Hopi country in the
latter part of the 19th century, it was not until 1917 that an effort was
made through correlation, and stratigraphic and statistical methods
to work out a chronological sequence of pottery development for
the Little Colorado area, which includes the Hopi country. This work
was undertaken by Spier, of the American Museum of Natural His-
tory, whose principal object was a determination of the Zufi series.
The occurrence of Hopi pottery was lumped under the term “ Buff-
ware,” which term was applied to Hopi as well as Zufti wares. Thus,
the Hopi sequence remained unsolved, and it was Kidder in 1923
and 1926 who first threw light upon the subject. As a result of a
survey of a number of ruins, from surface examination and strati-
graphic investigation he was able to determine that a yellow ware
with black decoration preceded the Sikyatki polychrome which was
in use at the Hopi pueblos when the Spaniards arrived in 1540.
This was the condition of affairs in the spring and summer of 1928
when the writer made a survey for Dr. A. E. Douglass in the interest
of the Second National Geographic Society Beam Expedition, in an
effort to determine those sites occupied or abandoned during the
period known in the Douglass tree-ring chronology as the “ Gap,”
a period which covered the time between late black-on-white wares and
the development of early historic pottery types.
Briefly, the result of the 1928 study of Hopi pottery was the es-
tablishment of a pottery sequence for Pueblo IV in the Hopi country,
* Douglass, Andrew Ellicott, The secret of the Southwest solved by talkative
tree rings. Nat. Geogr. Mag., Dec., 1929.
* An outline for a chronology of Zufi ruins. Anthrop. Papers, Amer. Mus.
Nat. Hist., Vol. XVIII, pt. 3, New York, 1917.
118 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
which substantiated the order suspected by Dr. Kidder. This sequence
has withstood the test of stratigraphy, and has been verified by dates
from beams found at Kawaioku in close association with pottery
specimens representing the different periods. This development is not
clearly understood as yet, and will be presented at some future time
after the study of pottery specimens collected has been completed.
The tentative classification used in field research is based upon the
order in which characteristic pottery types first appear, and though
the order of development has been verified, the periods as designated
may later be revised. In the following classification only the type
specimen for each division will be given. The classification is as
follows:
Pueblo III (Basic complex): Kayenta black-on-white and Kayenta
polychrome.
Pueblo: 1V, Period: A : Jeddito black-on-orange
Pueblo IV, Period B : Jeddito black-on-yellow
Pueblo IV, Period C : Sikyatki polychrome
In an effort to help clarify the matter of Southwestern terminology,
names of ceramic types follow the recommendations of the 1927
Pecos Conference. They are binomial—a geographic locality followed
by a descriptive term. Generally speaking the type specimens of
Pueblo III and Pueblo IV of the Hopi region are easily described ;
specifically, they are difficult of description. Pueblo III at the present
time embraces the types found at Betatakin and Keet Seel, as de-
scribed by Kidder,’ and others. Though it is reasonable to expect
further divisions of Pueblo III in the Hopi country, our knowledge
is not yet sufficient to designate any one type as representative of a
definite division, so we are necessarily forced to classify the period
by its ceramic development. Period A of Pueblo IV is different,
however ; it is characterized by a ware of a peculiar shade of orange,
sometimes ranging from almost red to a near-brown, which is dec-
orated with black paint and has sherd temper. This is Jeddito black-on-
orange and is found consistently associated either with late black-on-
white, an earlier developed ware, or with Jeddito black-on-yellow, the
succeeding type. Color alone is not sufficient for the identification of
these types, for specimens in peripheral sites often are similar in
color though they differ in other details which are recognized by one
familiar with either type. Frequently the difference cannot be fully
* An introduction to the study of Southwestern Archaeology. A. V. Kidder,
Yale Univ. Press, New Haven, 1924.
NOs LT PUEBLO RUINS IN ARIZONA—-HAURY AND HARGRAVE 12 fe)
explained, and is recognized by the “ feel” of the ware—a fact noted
by all familiar with Pueblo pottery. Sherds of the period types dealt
with in this paper are common in ruins belonging to the same cultural
development, the difficulty in identification being limited only to periph-
eral finds. Jeddito black-on-yellow is characterized by the smooth-
ness and quality of the ware, partly depending upon the fineness of
the tempering material, which is seldom obvious to the naked eye.
The basic color shades vary from cream to rich orange, while the
decoration may be from light brown to black. The appearance of this
type is sudden, suggesting a radical change in technique or materials—
possibly both. A paper dealing with important differences in decora-
tive technique and correlation with dated beams is in preparation.
In addition to normal pottery types as given in the preceding para-
graph, occasional intrusive sherds from other regions have been found
on the surface of the pueblo and in refuse heaps. Most notable of
these are Zufi glaze-paint specimens as noted by Hodge.’ A sherd of
“ green glaze on white or creamy slip ” (Zufi Chronology, Period C)
was found associated with a few Jeddito black-on-yellow sherds mixed
with the fallen walls of Kiva R-4. The relation of these sherds to the
occupation of the kiva is rather indefinite, though it is felt that they
were included in the wall masonry. No specific importance, other
than the occurrence of this Zufi type during Pueblo IV can therefore
be given. Four-mile polychrome has also been found on the ground
surface, but is less common than the Zufii wares. It occurs commonly
in Pueblo IV ruins along the Little Colorado River and at Chevelon
to the south. Rarely a sherd of Middle Gila pottery is also found.
Though it is not intended to comment upon the various forms of
pottery found at Kokopnyama since this subject is worthy of separate
note, it is of interest to remark upon the discovery of a problematical
form found in a test hole. The vessel is a plate-like object with per-
forations around the edge. This form is described by Mr. Haury.
The Kokopnyama specimen is smaller than that from the White
Mountains. It is 7 inches in diameter, has one row of perforations
around the edge, and four rows quartering the vessel. Perforations
are made from the inside and average 4 inch from center to center.
Like the Pinedale specimen, clay was adhering to the inside when
discovered. This would strengthen the belief that it was made for a
pot rest to sustain the weight of large jars in the making, though no
plausible explanation has been given for the perforations. Hopi
workmen suggested that it might have been used in an obsolete cere-
* Kidder, A. V., Southwestern archaeology, p. 91, 1924.
120 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
mony in which vessels of this description were used. It is reported
that the perforations were to hold flowers as in a modern flower
holder.
Kokopnyama would give little encouragement to a pot hunter, and
the paucity of complete or restorable specimens was a blow to the
investigators since we had expected a nice collection of these excel-
lent wares as a by-product. Only one room, R-11, contained un-
broken vessels, practically all other rooms either being filled with
refuse or containing nothing at all. Room R-14 will prove of value,
however, since it was filled with refuse containing a great quantity of
broken pottery vessels, the majority of which were Jeddito black-on-
orange. Associated with these specimens were a few late black-on-
white vessels, but none of the later Jeddito black-on-yellow ware was
found. This is the only occurrence of the orange ware found at
Kokopnyama in which it was not also associated with the yellow ware.
Many of these specimens are restorable, and it is expected that much
will be learned from a study of them.
SUMMARY
Briefly, the results of the 1929 work at Kokopnyama have con-
tributed to our knowledge of prehistoric Hopi culture, in that:
(1) Details of dwelling rooms have been found similar to those of
modern Hopi pueblos, if Spanish and American influence is dis-
regarded ;
(2) A knowledge of ceiling. construction is gained and found to be
essentially the same as in historic Hopi structures ;
(3) Three kivas, two completely and one partially excavated, fur-
nished data for comparison with ancestral (at Kin Tiel) and modern
Hopi types; thus making it possible to have the evolutionary de-
velopment from 1276 A. D. (see Kin Tiel) to the present time ;
(4) Many beam specimens were acquired which, when dated, will
give approximate years for important stages of kiva development,
and associated pottery types;
(5) A determination was made of the definite time of decadence of
old and rise of new pottery types;
(6) Confirmation was obtained from stratigraphic evidence of pre-
historic Hopi pottery sequence as previously determined by other
means; and
(7) Much cultural material was acquired.
SMITHSONIAN MISCELLANEOUS COLLECTIONS Woes eG INO Wale delbe 25)
1. Deflector and platform, Kiva KT-I, Kin Tiel.
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2. Note shadow of airshaft, Kiva KT-II, Kin Tiel.
2, MOA
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SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82, NO2 11> PL. 23
1. Kiva R-4, Kokopnyama, showing brace wall between rear wall of kiva and
first masonry wall.
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2. Kiva R-4, Kokopnyama, showing combined deflector and _ firepit.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82, NO. 11, PL. 24
1. Kiva R-4, Kokopnyama, with platform restored.
2. Kiva R-4, Kokopnyama, showing covers to firepits.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOE R82 NO eagle 25
1. Kiva R-24, Kokopnyama; general view. Note ollas buried in upper right
corner Ot room.
iS)
Room 11, Kokopnyama; part of fallen roof removed showing buried olla
in corner. Note beam hole (top) and cache.
SMITHSONIAN MISCELLANEOUS COLLECTIONS WADE 325 INO Waly AL. 2s
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2. Room 10, Kokopnyama; firepit behind upright slab in corner.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82, NO. 11, PL. 27
1. Jeddito black-on-yellow bowls, Kokopnyama.
2. Late black-on-white ware ollas from Kin Tiel.
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SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 82, NUMBER 12
THE FIVE MONACAN TOWNS IN’
VIRGINIA, 1607
(WITH 14 PLATEs)
BY
DAVID I. BUSHNELL, JR.
(PUBLICATION 3070)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
NOVEMBER 18, 1930
The Lord Gattimore (Press
BALTIMORE, MD., U. S&S A.
THE FIVE MONACAN TOWNS IN VIRGINIA, 1607
By DAVID L BUSHNELL, JR:
(WitTH 14 PLATEs)
Late in April, 1607, three small ships sent out from England passed
the Virginia Capes, entered Chesapeake Bay, and soon sailed a short
distance up the James. These were the Godspeed, Discovery, and
Sarah Constant, bearing colonists destined to form the first permanent
English settlement in the New World. On May 14, they had come to
anchor off a chosen site where Jamestown was soon to be established.
The spirit of adventure which had inspired many to undertake the
long and tedious voyage across the Atlantic continued to dominate
their acts, and within the following year they had reconnoitered much
of the surrounding region, then a vast wilderness claimed and occupied
by native tribes.
So venturesome were the colonists, and so desirous were they of
becoming acquainted with the country and its primitive inhabitants
that on May 21, one week after landing, a party of 23, one of them
Captain John Smith, left Jamestown to explore the river. They con-
tinued up the stream, made rapid progress, and two days later arrived
near Powhatan’s village, not far from the eastern bounds of the
present city of Richmond. The English desired to continue up the
valley beyond the falls, and endeavored to persuade Powhatan to
furnish guides to accompany them, but in this they were not success-
ful. To quote from a narrative prepared by a member of the party:
‘“Dynner Done we entred into Discourse of the Ryver how far it
might be to the head therof, where they gat their Copper, and their
Iron, and how many dayes loryne it was to Monanacah, Rahowacah
and the Mountains Quirank: requesting him to have guydes with us
also in our intended March; for our Captaine Determyned to have
travelled two or 3 dayes lornye a foote up the Ryver: but without
gyving any answer to our Demaundes, he shewde he would meete
us himselfe at the overfall and so we parted. This Nauiraus ac-
companyed us still in the boate. According to his promyse he
(Pawatah) mett us; where the fellow whome I have called our kinde
Consort, he that followed us from Turkey Ile, at the Coming of
Pawatah made signe to us we must make a shoute, which we Dyd.
SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 82, No, 12
2 SMITHSONIAN MISCELI.ANEOUS COLLECTIONS voL. 82
“Now sitting upon the banck by the overfall beholding the same,
he began to tell us of the tedyous travell we should have if wee pro-
ceeded any further, that it was a Daye and a halfe Iorney to Mona-
nacah, and if we went to Quiranck, we should get no vittailes and
be tyred, and sought by all means to Disswade our Captayne from
going any further: Also he tolde us that the Monanacah was his
Enmye, and that he came Downe at the fall of the leafe and invaded
his Countrye . . . . But our Captayne out of his Discreyton (though
we would faine have seene further, yea and himselfe as desirous
also) Checkt his intentyon and retorned to his boate.”* The party
returned safely to Jamestown where they arrived June 22.
On this, the first journey to the falls, the inhabitants of all the
many native villages encountered were allies, but the colonists heard
of others beyond, to the westward, who were enemies of the allied
tribes. This is the earliest known reference to the Monacan.
Early in the spring of 1608 another expedition was projected “ to
discover and search the commodities of Monacans countrie beyound
the Falles,” but the exploration was not attempted as events at James-
town made it necessary for all to remain near by. However, during
the autumn of the same year Captain Newport, having just returned ~
from England, “ with 120 chasen men . . . . set forward for the dis-
covery of Monacan.” The expedition appears to have been quite
successful, although scant records of the happenings have been pre-
served. One narrative tells how “ Arriving at the Falles, we marched
by land some fortie myles in two dayes and a halfe; and so returned
downe the same path we went. Two townes we discovered of the
Monacans, called Massinacak and Mowhemenchouch; the people
neither used us well nor ill, yet for our securitie we tooke one of
their petty Kings, and led him bound to conduct us the way.’* This
is the earliest known record of the entering of the Monacan territory
by Europeans.
The region beyond the bounds of the Powhatan confederacy, up
the valley of the James from the falls, was regarded by Indian and
colonist alike as a separate and distinct land. This is suggested by a
statement contained in a letter written by George Yardly to Sir Henry
Payton in London, dated ‘“ James town, this XVIII of November,
1610,” in which he referred to an expedition planned by the Governor,
who intended going “up unto a famous fall or cataract of waters,
where leaving his pinnasses & Boates safe riding, so purposely to
loade up go into the Land called the Monscane.” *
* Archer Relation. Jn Smith, Arber ed., pp. xlv-xlvi.
* Smith, Arber ed., p. 438.
* Bodleian Library, Oxford. MSS. Eng. Hist., C. 4, fol. 3.
Lee es
NO: 12 _ MONACAN TOWNS IN VIRGINIA—BUSHNELL 3
Strachey writing a few years later told what was then known of
the country beyond the falls: “ Concerning the high-land little can
we say as yet, because thereof little have we discovered: only some
Indians’ relations and some fewe daies’ marches into the Monocan
country of our owne, have instructed us thus far.
“This high land, or Britannia, then say we, is the mayne and firme
continent, which extendeth, we wot not how far, beyond that cataract
or fall of water, which the Indians call Paquachowng, from whence
one daies’ jorney into the Monocan country. Our elder planters (at
their first comyng) proclaymed His Majestie King of the country at
Mohominge (a neighbour village), and sett up a crosse there with
His Majestie’s name inscribed thereon . . . . From the falls our men
have heretofore marched (as the river led them) about forty or fifty
miles, and fownd a high land woody, little champion, with rising hills,
rocky and mountanous ... .”"
Continuing Strachey wrote (p. 131): “For mineralls we will
promise nothing ; but the hope of which, seeing the low grownd, yields
manie faire shewes; the mountaines cannot be doubted but that in
them manie sortes will be found: and our people, in their first dis-
covery into the Monocan country discovered two mynes, the one
within six miles of the head of the falls, which takes the name of
Namantack, the fynder of yt: which is conceaved wilbe worth the
exploring, and with little charge; the other lyes in the myd-waie be-
tweene twoo townes of Monocan, the nearest called Mowhemincke,
the furthest, Massinnacock, distant one from another fourteen miles.”
The preceding references must necessarily apply to discoveries
made by Newport during the autumn of 1608.
A map of Virginia, usually attributed to Captain John Smith, was
presented in his “ Generall Historie of Virginia,” 1624. It shows
the course of the James far above the falls and many miles beyond
the spot reached by Newport in 1608. A cross appears on the map
at the beginning of the falls, the meaning of which may be understood
by quoting from Strachey’s reference to the map. He gave this
quaintly worded explanation (p. 42): “In which mappe, observe
this, that, as far as you see the little crosses either rivers, mountaines,
or other places, have discovered ; the rest was had by informacion of
the salvadges, and are set downe accordinge to their instruccions.”
In this instance it is evident the cross marks the farthest point reached
by Smith in May or June, 1607. He had never gone beyond the falls
* Strachey, William, The Historie of Travile into Virginia Britannia, p. 25.
Hakluyt Society, London, 1840.
4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL, 82
and wrote, when describing the James then “ called Powhatan, ac-
cording to the name of a principall country that lyeth upon it... . .
It falleth from Rockes farr west in a Country inhabited by a nation
they call Monacans. But where it commeth into our discovery it is
Powhatan. In the farthest place that was diligently observed, are
falles, rockes, shoules, &c. which makes it past navigation any higher.” *
And it is believed the dotted line following the left bank of the river
indicates the trail traversed by Newport and his party in the autumn of
1608. The region beyond the end of the trail, as shown by the broken
line, was described by the Indians. The Monacan with whom Newport
came in contact would undoubtedly have been well acquainted with
the country as far westward as the mountains and even beyond, and
it may have been from “ one of their pettie Werowances ”’ that were
learned the description of the course of the river, the position of its
principal tributary beyond the falls, and the locations of the large
towns. But whatever may have been the source of the information,
the map made more than three centuries ago was remarkably accurate,
as may be seen by comparing it with a very recent plan of the same
region. The one tributary shown on the Smith map was the present
Rivanna, but its name, by which it was known to the Indians, has not
been preserved.
Five villages, or rather centers of population, are indicated on that
part of the old map designated as the territory of the Monacans.
These are:
a. Mowhemcho. On the right bank of the James.
b. Massinacack. On the right bank of the James beyond Mo-
whemcho.
c. Rassawek. At the junction of the James and the Rivanna.
d. Monahassanugh. On the James beyond the mouth of the Rivanna.
e. Monasukapanough. On the Rivanna.
The spelling of the names differs in the text where they are often
given as:
a. Monacans. d. Monahassanuggs.
b. Massinnacacks. e. Mouhemenchughes.
c. Russawmeake.
The name Monacan was first applied to the territory occupied by
the five tribes as well as to the confederacy which they composed.
Later the first town entered by the English—Mowhemcho—became
known as Monacan Town. This was the village of the Monakins of
Op. cit, p: 346:
NO: D2 MONACAN TOWNS IN VIRGINIA—BUSHNELL 5
Lederer, and Manakin Town or Maningkinton of Michel. The mean-
ing of the word is not known.
The spelling of the names as given on the Smith map will be fol-
lowed when referring to the sites in the present sketch, although it
is not possible to discover which form is the more nearly correct. A
section of the map showing the five Monacan towns, the course of
the James and of its principal tributary, the Rivanna, is reproduced
in figure 1. A second map, figure 2, is presented for comparison with
the preceding. The base is traced from a recent map issued by the
United States Geological Survey, and on this have been indicated
the sites of the five Monacan towns mentioned by the early writers.
They will be described separately in the following pages.
How long the country had been occupied by the Siouan tribes can
never be determined. Others had preceded them, but who they were
or whence they came may ever remain unknown. The earlier habitat
of the Siouan tribes, to which stock the Monacan belonged, is believed
to have been in the valley of the Ohio, from which region they crossed
the mountains to the eastward and later occupied the lands where
they were encountered by the Virginia colonists early in the 17th
century. A comparison of the material to be recovered from sites
eastward from the Ohio may make it possible to trace the line of
migration of these tribes; this would require much time and careful
study, but if successful would prove of the greatest interest.
CAE OPALBIES
The Falls of the James, which tended to separate the regions
occupied by the two groups of tribes, may be more correctly described
as a series of rapids extending several miles. It was an important
place for fishing, and was frequented by the Indians whose camps
would have been found scattered along the wooded banks of the
stream, a very rough and broken bit of country.
The last village of the Algonquian tribes up the river was a mile
or more below the foot of the rapids. It is not improbable that the
upper part of the falls, some miles distant from the Algonquian
village, was often visited by the inhabitants of the nearest Monacan
town, Mowhemcho. Powhatan’s statement to Captain Newport at
the time of their first visit in 1608 that the Monacan “ came Downe
at the fall of the leafe and invaded his Countrye,” would indicate that
the Monacan rather than the Algonquian dominated the region and
did not fear the latter.
There was formerly in “The Byrd Title Book ”—a manuscript
volume belonging to the Virginia Historical Society—a drawing or
6
—
—
h
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
a Mafsinaca C
ti
Ni Git
Wee
>
Y }
LALA QD Cattachiptre
Fic. 1—Section of the Smith map, 1624, showing the country occupied by
the Monacan.
se
NO. I2 MONACAN TOWNS IN VIRGINIA—BUSHNELL
NVONIJACANS
MASSINACACK
Ie
MOWHEMCHO ie = 2
Ik
a
The //ralls Seale of Miles 4
® RICHMOND ie
Fic. 2—Detail of a recent survey, with sites of Indian towns added
For comparison with map of 1624, figure I.
8 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
rather plan of the rapids and islands in the James which was made
before the year 1700. At a point between an island and the right
bank of the river was indicated the position of the “ Indian Fish
Traps.” This was evidently just within the western limits of the
present City of Richmond where ancient fish traps may still be seen
in the river. They are clearly defined when the water in the James
is low, and under such favorable conditions they were photographed
by the writer in October, 1926. The view is reproduced as the frontis-
piece, plate I.
Beverley * described several ways of fishing, and had undoubtedly
witnessed all being practiced by Indians with whom he had come in
contact. He wrote in part: “At the falls of the Rivers, where the
Water is shallow, and the Current strong, the /ndians use another
kind of Weir, thus made: They make a Dam of loose Stones, whereof
there is plenty at hand, quite across the River, leaving One, Two, or
more Spaces or Trunnels, for the Water to pass thro’; at the Mouth
of which they set a Pot of Reeds, wove in the Form of a Cone, whose
Base is about Three Foot, and perpendicular Ten, into which the
Swiftness of the Current carries the Fish, and wedges them so fast,
that they cannot possibly return.”
The preceding description applies perfectly to the fish trap in the
James, where three “ Spaces or Trunnels ” may be distinctly traced
pointing down the stream. Beverley may have had this exact site in
mind when he wrote his account so many years ago.
White oak splints, similar to those used in making baskets, were
formerly “ wove in the Form of a Cone ”’ to serve as fish traps. They
were used extensively in this part of Virginia as elsewhere. The
maximum diameter of the large end was usually about one-third the
length of the finished trap. At this end the weaving returned inward
for a short distance, the opening becoming smaller so as to prevent
the escape of the fish. At the opposite end the warp elements extended
some inches after the woof was discontinued, coming closer together
and finally touching, thus serving to close the end of the trap. They
were placed where the current was strong, with the small end pointing
down stream.
MOW HEMCHO
This village, later to be known as Monacan Town, was the first
encountered by Newport in 1608 in passing up the valley of the James
from the falls. It stood on the right or south bank of the river and
* Beverley, Robert, The history and present state of Virginia, Book 2, p. 33.
London, 1705.
NO. I2 MONACAN TOWNS IN VIRGINIA—BUSHNELL 9
probably covered some of the level area bordering the stream in the
extreme eastern part of the present Powhatan County, between
Bernards Creek on the east and Jones Creek on the west. It does not
appear to have been a palisaded village but rather an open settlement.
Gardens were probably near the scattered habitations. It was a beauti-
ful site for a native village. On the north it was protected by cliffs
rising abruptly from the left bank of the river, on the south it
was bounded by high, rolling land from which issued springs of clear
water. Game was abundant throughout the region.
Three centuries ago Mowhemcho was an important center, probably
the home of some hundreds of individuals who lived in a land of
plenty, where food was easily obtained. How long the site may have
been occupied will never be known. By the close of the 17th century
few Indians remained in the vicinity, and during the year 1699 a
Huguenot colony took possession of the land and there established
a settlement which continued for some years. Huguenot, near the
middle of the tract, and Huguenot Springs on Bernards Creek, about
2 miles from the bank of the James, are names which tend to identify
the site. At the present time few traces of a native settlement can be
discovered on the surface, which has been subject to overflow during
the past centuries, but much may be hidden beneath deposits of sand
and alluvium.
The Huguenots settled part of the area in 1699, but it is quite
evident that Indians continued to occupy a portion of the site. Three
years later they were briefly mentioned by a Swiss traveler.’ This
was in April, 1702, when Michel stopped at “ Manakin town,’ and
later wrote in his journal (II, p. 123): “ The Indians often visit
there, bringing game, rum and other small things. There is a good
opportunity to trade with skins. They (the Indians) often bring
pottery and when desired fill’d with corn.” How corn was prepared
in Virginia at that time is not revealed. He continued and again
mentioned the Indians (p. 132): “ In their homes they are naked, as
I have seen one at Maningkinton, who came back from hunting. He
had nothing but his gun, knife and powder horn, except a linen rag
which covered his sexual parts a little, and a deer skin protecting
his feet, that the thorns might not hurt him. He had also a tuft of
feathers behind his ear.”” Such was the appearance of a Monacan in
the early Spring of 1702. Several drawings made by Michel and
* Michel, Francis Louis, Journey from Berne, Switzerland, to Virginia—
October 2, 1701 to December 1, 1702. The Virginia Magazine, Virginia Hist.
Soc., Richmond. January and April, 1916.
Io SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
reproduced in the articles cited, are believed to represent Monacan
Indians as described by him, however the sketches are not mentioned
in the journal and for that reason are not positively identified.
The Indians of the village, or those who frequented the settlement,
had been in possession of guns for many years. When Lederer’ and
his party reached the village of the “ Monakins,” May 22, 1670, they
were, so he wrote: “ welcomed by them with volleys of shot.” But
the village at that time was already much reduced and undoubtedly
many of the people had wandered away to seek new homes elsewhere.
In 1669, the year before Lederer’s visit, the Monacan could bring
together only 30 warriors.
MASSINACACK
Massinacack was the second native village reached by Newport
in 1608 after having entered the Monacan territory. It is shown
on the Smith map of 1624 as situated south of the James. Strachey
mentioned it being some 14 miles distant from Mowhemcho, or
Monacan Town, which would place it on the right bank of the river
about the mouth of the present Mohawk Creek, a mile or more south
of the town of Goochland, on the opposite side of the James.
Lederer was at the Monacan village late in May, 1670. A crude
map, part of which is now reproduced in figure 3, accompanied the
narrative of his journey and shows the James, or Pawathan fl. from
near the falls westward. A dotted line passes up the right bank of
the river and is evidently intended to represent the trail which he
followed. This reached Monakin and. continued to Mahock. The
former was the Mowhemcho of Smith, the latter was undoubtedly
another form of Massinacack, from which the present name of the
creek has been derived. Two streams are represented coming together
at Mahock and at once suggest the confluence of the James and the
Rivanna, but it probably indicates the creek now bearing the name
Mohawk flowing into the James. This was,- without doubt, the site
of the ancient village of Massinacack.
Lederer’s narrative is vague and uncertain, the distances given
are not reliable, but the names of streams and of places which appear
on the map and are repeated in the text may be quite accurate. For
this reason the present Mohawk Creek is believed to have been the
Mahock of Lederer. By continuing due westward from this creek
the party would have reached the James flowing in a northerly
*Lederer, John, The discoveries of .... Begun in March 1669, and ended
in September 1670. London, 1672. Reprint, Rochester, 1902.
NO. 12 MONACAN TOWNS IN VIRGINIA—BUSHNELL
2;
S
Fic. 3.—Section of the Lederer map, 1670, showing the trail up the James to
Monakin-and Mahock, with Sapon Nahissan far to the southwest.
12 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL, 82
direction ; this was probably what they considered to be “‘ the south-
branch of James river, which Major Harris observing to run north-
ward, vainly imagined to be an arm of the lake of Canada.”
During the 18th century there was an important ferry across
the James at, or very near, the site of this early native settlement.
Leading southward from the ferry was a road over which colonists
went to Carolina. As many of the roads developed by the first
settlers followed the lines of ancient trails it is within reason to
believe the same route had been traversed by Indians through
generations.
RASSAWEK
No account has been discovered of a European having visited
the village of Rassawek, although early writers referred to it as the
principal town of the Monacan confederacy. Smith wrote in 1612:
“ Upon the head of the Powhatans are the Monacans, whose chiefe
habitation is at Russawmeake,” but all his knowledge of the place
had been derived from Indians. It stood evidently at the confluence
of the James and Rivanna, some miles beyond the point where
Newport turned to retrace his way to Jamestown in the autumn of
1608. The site had been abandoned before white settlers entered the
region and consequently its exact position may never be known.
Viewed at the present time the most desirable and logical location
for an extensive village would have been on the right bank of the
Rivanna, within the angle formed by the two streams. Here is a
wide bottom with high, rolling land a short distance from the James
touching the Rivanna. When timbered and in its natural condition
this would have been a beautiful site for a native settlement. The
proximity of the two streams would have afforded some protection.
The wooded hills to the north would have sheltered the frail habi-
tations from the winter winds. Fish and wild game, ever plentiful,
could have been easily taken for food. This was probably the site
of the village of three centuries and more ago.
Many traces of Indian occupancy have been discovered within
a radius of a few miles of the mouth of the Rivanna, but all should
not be attributed to the Monacan. The junction of two streams
always attracted the Indian and it is evident others had lived there
before the coming of the Siouan tribes to the valley of the James.
It is believed that the members of the five tribes or groups men-
tioned in the present sketch had similar manners and customs, and
that all disposed of their dead as did the people of Monasukapanough,
whose village stood on the banks of the Rivanna far above Rassawek.
NO. 12 MONACAN TOWNS IN VIRGINIA—BUSHNELL 1s
Consequently the discovery of other sites along the course of the
James, where the dead had been buried in shallow pits scattered
through the village, suggests that some other tribe or tribes may have
preceded the Monacan. Numerous signs of Indian occupancy have
been encountered on Elk Island, a large island in the James a short
distance below the mouth of the Rivanna, but there is no reason to
believe it was ever occupied by the important village of Rassawek.
The burials discovered on the island do not appear to have been of
Monacan origin; however, related Siouan tribes could have occupied
this and other sites in the valley of the James.
MONAHASSANUGH
As Mooney has so clearly shown, the Monahassanugh of Smith
were the Tutelo of later narratives. To quote from his interesting
work’ (p. 37): ‘“‘ The Tutelo and Saponi tribes must be considered
together. Their history under either name begins in 1670. ....
Monahassanugh and Nahyssan are other forms of Yesa", the name
given themselves by the last surviving Tutelo, and which seems to
have been the generic term used by all the tribes of this connection
to designate them as a people.” And again (p. 31): “ In Nahyssan
we have the Monahassanugh of Smith, the Hanohaskie of Batts, and
the Yesang of Hale. The last is evidently the generic root word, the
prefix Mo, Mona, or Na in the other forms probably giving a specific
local application to the common term. Thus from Lederer’s statement
that Sapon was a Nahyssan town we understand that the Saponi were
a subtribe or division of the people who knew themselves as Yesang.”
The ancient village of Monahassanugh is believed to have stood
on the left bank of the James, about 14 miles up the stream from
Wingina, in Nelson County. The river is here bordered on the
north, or left bank, by broad fertile bottom lands which extend for
some miles above and below the site; while on the opposite side
cliffs rise abruptly, steep, rugged and broken. The site resembles
that of Mowhemcho or Monacan Town, although the relative position
of the cliffs and low ground is reversed, the former being on the left
bank of the river and the latter, which was occupied by the village,
on the right.
Stone implements have been found scattered over much of the
low ground, arrowheads of white quartz and of brownish quartzite
have been recovered in vast quantities. Numerous fragments of
*Mooney, James, The Siouan tribes of the East. Bull. 22, Bur, Amer. Eth-
nol., Washington, 1894.
If SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
pottery bearing the imprint of textiles, stone gorgets, pipes, etc.,
have been collected on the site—all proving the former existence here
of an extensive, permanent village.
The site was visited by Fowke about the year 1892. He wrote:’
“ The Indian trail from the Shenandoah valley, through Rockfish gap,
crossed James river at an island near Norwood. For 5 miles below
in the river there is a succession of pools and rapids, with many
large rocks in the channel which are covered only in time of high
water. The hills on the south with scarcely an exception reach to
the water, there being only a few narrow strips of level ground. On
the north the bottom lands are wide and continuous.
“The only indications of Indian occupancy on the southern side
in this vicinity are opposite the island. On the northern side, however,
aboriginal remains may be found on every farm. They are most
abundant on the lands .... three miles below Norwood.
“The floods of 1870 and 1877 disclosed numerous small deposits,
probably more than 200 in all, containing burned stones, pieces of
pottery, arrowheads, and great quantities of quartz chips. They are
in nearly straight rows, from 25 to 50 feet apart, and extend for
several hundred yards along the river.” Many stone implements were
discovered, and “all these things point to a village of considerable
size, but a most careful search of the whole area, especially along
the river bank and in the numerous gullies, failed to reveal a bone
of any description.”
The material recovered was similar to that found on the site of
Monasukapanough, on the banks of the Rivanna, to be described in
the following section. The chipped ax- or celt-like implements found
on both sites are the most characteristic of all the objects recovered.
Three typical examples from the James River site are shown in
figure 4, for comparison with others found on the banks of the
Rivanna, plate 5.
Fowke’s failure to discover a cemetery, or to find any traces of
human remains, tends to strengthen the belief that this was a Siouan
village where the burial customs were the same as those of the
people of Monasukapanough, related tribes having the same customs
and ways of life. Undoubtedly a large burial mound, or possibly
several, once stood on the low grounds bordering the left bank of the
James. These, the ‘“‘ Indian Graves ” of early records, were probably
to have been encountered in many localities, but were destined to be
destroyed when the land was cleared and cultivated ; soon all traces
1 Fowke, Gerard, Archeologic investigations in James and Potomac Valleys.
3ull. 23, Bur. Amer. Ethnol., Washington, 1894,
—-
ke
MONACAN TOWNS IN VIRGINIA—BUSHNELL I5
12
NO.
Size,
natural
1
Fic. 4——Implements found on the site of Monahassanugh.
(U. S. Nat. Mus. Cat. No. 136175)
16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
of the mounds disappeared and their existence was forgotten. How-
ever the mound which stood on the bank of the Rivanna was to live
in history, and the careful manner in which it was examined by
Jefferson will ever be of interest.
About the year 1654 many Indians came from a distance and “ sett
downe near the falls of James river, to the number of six or seaven
hundred.” * It appears they came to seek a new home, in no manner
antagonistic to the colonists and desiring peace. However, after they
had been there some months the English endeavored to expel them and
this resulted in one of the greatest and most disastrous battles ever
fought by early settlers and Indians. Totopotomi and his Pamunkey
warriors had become allies of the English, but he and the majority
of his followers fell when the entire force was routed and defeated.
The identity of the Indians who had thus come to the region of the
falls to seek a peaceful home, and who proved themselves such
worthy warriors, has never been determined. The name Rechahecrian
or Rickohockan has been applied to them, believed by some to have
been the Cherokee, although it was Mooney’s later belief that they
were. Erie who had come southward. However, a statement by
Lederer makes it appear they were the people of two Monacan groups,
the Massinacack and Monahassanugh, who may have come from
farther up the James to settle a new home more protected from the
war parties of the Iroquois.
As has already been explained the Mahocks and Nahyssans of
Lederer were probably the Massinacack and Monahassanugh of
Smith and other early writers. Thus when Lederer mentioned the
great encounter and said: “a great Indian king called Tottopottoma
was heretofore slain in battle, fighting for the Christians against the
Mahocks and Nahyssans.”’* he did not doubt the identity of the people
against whom the English and the Pamunkey allies fought. Lederer,
on his map, gave the name Rickohockans to a tribe then living far to
the westward beyond the mountains. The name or term has never
been clearly understood or translated, and with slight variation of
spelling has been used to designate several tribes in widely separated
parts of the country. But the word may have been a term applied
under certain conditions and not the definite name of any tribe or
group of tribes. If this belief is correct it could have been applied
to Siouan as well as to Iroquoian or other tribes.
The Rickohockans, so-called, were to Lederer a vague group,
evidently known to him only during his travels away from the English
1 Hening, I, p. 402.
Ope cite sp. 10.
NO. I2 MONACAN TOWNS IN VIRGINIA—-BUSHNELL 17
settlements. Had he associated them in any way with the great
battle he would undoubtedly have mentioned them in that connection,
but this he failed to do. The Algonquian and Monacan tribes had
ever been enemies, it was known as early as 1608, and this fact may
explain the willingness with which Totopotomi and the Pamunkey
warriors joined the English in attacking their ancient tribal enemies.
MONASUKAPANOUGH
This name, as it appears on the Smith map, corresponds with the
position of an extensive village site on the banks of the Rivanna, in
Albemarle County, directly north of the University of Virginia and
about one-half mile up the river from the bridge of the Southern
Railway. At that point the Rivanna makes a wide bend, flowing from
the west, then turning and continuing in a southeastwardly direction.
On the right or south bank there is a wide, fertile bottom, bounded
on the north by the river and on the south by cliffs sloping to the
low grounds. On the left or north bank of the stream the bottom is
far less extensive than on the opposite side, but it is rather higher
and less liable to be overflowed, and the cliffs are nearer the river.
This is believed to have been the site of the ancient settlement of
Monasukapanough. The village appears to have occupied both sides
of the river, with a ford that made it possible to pass from one side
to the other, although canoes were probably in constant use on the
stream. A plan of the region is shown in figure 5.
The translation of the name of the village has not been determined,
nor has that of the name of the related settlement which stood on
the bank of the James. As mentioned there is a ford across the
Rivanna at this place—-shallow water—which may have to do with the
first part of the name. This is suggested by statements by William
Byrd, in the year 1728, during the running of the line between Virginia
and North Carolina,’ when he had an old Saponi Indian acting as
guide. To quote from the remarkable narrative (p. 42): On Septem-
ber 28 “ We proceeded to the canoe landing on Roanoke, where we
passed the river with the baggage. But the horses were directed to
a ford about a mile higher, called by the Indians Moni-seep, which
signifies, in their jargon, shallow-water. This is the Ford where the
Indian traders used to cross with their horses, in their way to the
Catawba nation.” And on October 2 they crossed a large creek “ which
the Indians called Massa-moni, signifying, in their language, Paint
* Byrd, William, The Westover manuscripts: containing the history of the
dividing line. .... Petersburg, 1841.
18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
creek, because of the great quantity of red ochre found in its banks.”
Later on the same day they crossed another creek called “in the
Saponi language, Ohimpa-moni, signifying jumping creek, from the
frequent jumping of fish during the spring season.” It would now
be interesting to know if the name Jumping Branch, applied at the
present time to a branch of Hardware River, in Albemarle County,
perpetuates an ancient Siouan name.
Mooney was of the belief that Monasukapanough was _ possibly
“the original of Saponi.”’ There is little reason to doubt the correct-
7 2 \
}
NN . a
Peo \ / \
2 aa te x 4 ys
—o ie 1 . o PD)
GD =)
River
1. GARNETT SITE
2. MONASUKAPANOUGH (
3. MOON SITE
4. CAIRN
Scale of Miles ff
fo) —(] 2 t(
Fic. 5.—A section of the Rivanna Valley, in Albemarle County, Virginia,
showing the position of Monasukapanough and lesser sites.
Zz
ness of this belief. Lederer stated that he “ arrived at Sapon, a village
of the Nahyssans.” The latter, as previously shown, were the Mona-
hassanugh whose name appears on the map of 1624. Therefore it is
quite evident that at the time of the settlement of Jamestown, 1607,
the site on the banks of the Rivanna was occupied by the Saponi,
closely allied with the Monahassanugh or Tutelo, whose village stood
on the bank of the James some miles away in a southwesterly direction.
Had it not been for the work and interest of Jefferson, no account
of the great burial mound which once stood at the ancient village of
Monasukapanough would now be available. It would have disappeared
NO. I2 MONACAN TOWNS IN VIRGINIA—BUSHNELL 19
as have the burial places once belonging to other villages of the Siouan
tribes and no reference to it would have been preserved. The site of
the Indian town was visible from Monticello, and the burial mound
stood near the south, or right bank of the Rivanna within the area
shown in plate 2. Jefferson desired to know the nature of the contents
of the work and, so he wrote* (p. 139): “ For this purpose I de-
termined to open and examine it thoroughly. It was situated on the
low grounds of the Rivanna, about two miles above its principal fork,
and opposite to some hills, on which had been an Indian town. It was
of spheroidical form, of about 40 feet diameter at the base, and had
been of about twelve feet altitude, though now reduced by the plough
to seven and a half, having been under cultivation about a dozen years.
Before this it was covered with trees of 12 inches diameter, and round
the base was an excavation of five feet depth and width, from whence
the earth had been taken of which the hillock was formed. I first dug
superficially in several parts of it, and came to collections of human
bones, at different depths, from six inches to three feet below the
surface. These were lying in the utmost confusion, some vertical,
some oblique, some horizontal, and directed to every point of the
compass, entangled, and held together in clusters by the earth. Bones
of the most distant parts were found together, as, for instance, the
small bones of the foot in the hollow of a scull, many sculls would
sometimes be in contact, lying on the face, on the side, on the back,
top or bottom, so as, on the whole, to give the idea of bones emptied
promiscuously from a bag or basket, and covered over with earth,
without any attention to their order.” And to continue: “I proceeded
then to make a perpendicular cut through the body of the barrow,
that I might examine its internal structure. This passed about three
feet from its center, was opened to the former surface of the earth,
and was wide enough for a man to walk through and examine its
sides. At the bottom, that is, on the level of the circumjacent plain, I
found bones; above these a few stones, brought from a cliff a quarter
of a mile off, and from the river one-eighth of a mile off; then a
large interval of earth, then a stratum of bones, and so on. At one
end of the section were four strata of bones plainly distinguishable ;
at the other, three; the strata in one part not ranging with those in
another. The bones nearest the surface were least decayed . . . . Ap-
pearances certainly indicate that it has derived both origin and growth
from the accustomary collection of bones, and deposition of them
together ; that the first collection had been deposited on the common
1 Jefferson, Thomas, Notes on the State of Virginia. Philadelphia, 1794.
2
20 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
surface of the earth, a few stones put over it, and then a covering of
earth, that the second had been laid on this, had covered more or less
of it in proportion to the number of bones, and was then also covered |
with earth; and so on.”
There is reason to believe some Indians continued to occupy the
site until after the beginning of the 18th century. They may have
been few in number, but among the number must have been some
who were descendants of others who had lived there when Monasu-
kapanough was a large village. As late as the middle of the century
some were living who knew of the burial place of their dead. Jeffer-
son, referring to the mound which he had examined, told how “a party
passing, about thirty years ago, through the part of the country where
this barrow is, went through the woods directly to it, without any
instructions or enquiry, and having staid about it some time, with
expressions which were construed to be those of sorrow, then returned
to the high road, which they had left about half a dozen miles to pay
this visit, and pursued their journey.” Only those who had retained
a memory of the burial place could, or would, have made such a
pilgrimage.
The exact position of the mound may never be determined, but it
certainly stood on the low ground, on the right bank of the Rivanna,
evidently nearer the river than the cliffs, and it may have been some
distance above the ford.
During the month of June, 1911, I examined part of the low ground
in the endeavor to find some trace of the native village to which the
burial mound had belonged. Nothing was discovered on the surface;
all had been covered in the past years. Nine excavations were made
about 50 yards from the river bank, and beginning about that same
distance west of the road leading to the ford. One excavation was
30 feet in length, others were 5 or 6 feet square, all were 2 feet or
more in depth. In seven of the nine excavations small fragments of
pottery were encountered at an average depth of about 20 inches, bits
of quartz and quartzite, and pieces of charcoal were also met with in
some excavations. No traces of bones of any sort were found. The
superstratum, some 20 inches in thickness, represents the alluvium
deposited by the river since the village was occupied, and may have
resulted from one or more freshets during the past century. The
greatest freshet known was in 1877, at which time, so it is said, most
of the low ground was overflowed to a great depth. When the waters
receded some parts of the area were covered with a thick deposit of
sand while on other sections the soil had been washed away and the
surface lowered. Many stone objects of Indian origin were exposed.
‘syueq S}I SULIops0q
Soot} 94} Aq Uspply St Weaijs oy “eUURAY dq} JO YOU sy oy} Woy YNOS Suryooy ysnourdeynseuoyy jo ops ayy
@ “Id ‘Gt “ON ‘28 “10A SNOILOA1100 SNOANVITISOSIN NVINOSHLIWS
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82, NO. 12, PL. 3
Object found on the site of Monasukapanough, suggesting a human head.
ixacte sizes
(U. S. Nat. Mus. Cat. No. 350136)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOC S82 INO ed 25 ele 4:
Objects found on the surface, Monasukapanough. +4 natural size.
> 2
(U. S. Nat. Mus. Cat. Nos.: a, 350101; b, 350102; c, 350107; d, 350109; e€, 350110)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL 82); NO: 1/2) Pia5
Objects found on the surface, Monasukapanough. 4 natural size.
@G.S: Nats Mus: (Gat) Noss: (a) 350114 bs s50mnss) ce 3500103) d.6350nL 75) eps sOunLe))
i"
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLE 82) NOn 2) PL 6
ei:
: = Se gts {
Objects found on the surface, Monasukapanough. 4 natural size.
(U. S. Nat. Mus. Cat. Nos.: a, 350120; b, 350121; c, 350122; d, 350123)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82, NO. 12, PL.
ie rit ak ‘
aye
Objects found on the surface, Monasukapanough. 4 natural size.
(U. S. Nat. Mus. Cat. Nos.: a, 350124; b, 350139; c, 350125; d, 350126)
NO. 12 MONACAN TOWNS IN VIRGINIA—BUSHNELL 21
Axes, discoidal stones, and numerous chipped implements are men-
tioned as having been discovered, but now all are scattered and lost.
Undoubtedly a great number of interesting specimens could have been
collected at that time, proving it to have been the site of an extensive
native village. Evidently Jefferson did not suspect the existence of
part of the great village on’ the side of the river on which the mound
stood. He mentioned the hills on the opposite side “on which had
been an Indian town,” which may have been the more important part
of the settlement, as it has now become the more interesting.
On THE Lert BANK
Much of that which precedes refers to conditions on the right
bank of the Rivanna, but the great village also occupied some ground
on the opposite side of the stream. The land on the north or left
bank rises rather abruptly from the water, continues quite level for
100 yards or more and then becomes much higher. This compara-
tively level area of some 20 acres or more is thus bounded on one
side by the Rivanna and on the east and north by rising ground which
in some places is quite steep. On the west the cliffs approach the
river. Several large springs issue from the surface on the site of the
village. Before the land was cleared of timber the ground was
necessarily irregular and broken, and was traversed by several gullies
extending from the bordering cliffs to the river, worn deep by the
waters flowing from the springs which would have supplied the wants
of the settlement. The area has now been cultivated for many years,
the surface leveled and worn down by the plow, but while it remained
in its natural condition surrounded as it was by wooded cliffs, it
would have appeared hilly and broken; these were the hills on which
Jefferson said “ had been an Indian town.”
The central portion of the level area is the more elevated and
slopes gradually to the west and east. It is believed this part has
never been covered by the waters of the Rivanna although the lower
ground has been overflowed several times within recent years, always
leaving deposits of sand and alluvium on the surface.
A general view of the site is reproduced in plate 2. This was taken
from the high land on the north. In the foreground is the section
north of the river; the course of the stream is indicated by the line
of trees bordering its banks. Beyond is the low ground on the right
bank of the river, with the cliffs rising in the distance.
Many stone objects have been discovered scattered over the surface
of the higher part of the level ground where they may never have
been covered by water, even at the times of great freshets. The
22 -SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
specimens have thus remained since they were lost or abandoned by
the last inhabitants of the village—believed to have been the Saponi,
who left the site some time before the year 1670, although some
may have lingered behind. About 70 years later colonists entered the
valley of the Rivanna. The ground has now been cultivated for many
years and, undoubtedly, numerous objects both large and small have
been broken by the plow, but some of unusual interest have been
discovered within the past few years.
The material collected on the surface consists of objects of stone,
both chipped and polished, and numerous fragments of pottery, many
of which bear the imprint of textiles. No specimens made of shell,
bone, or metal have been discovered, and nothing of European origin
to suggest contact with the colonists has been encountered on the site.
Many of the stone implements, or weapons, are crudely made, but
with edges worn and polished as a result of much use. These are
seldom broken or incomplete although a number of fragments of well
made polished celts have been found, as well as more perfect speci-
mens with only the cutting edge battered or fractured, suggesting
rough usage. Perfect or complete objects of the finer workmanship
are not found. This fact is difficult to explain unless the better pieces
were carried away when, as it is believed, the majority of the people
of the village moved to another locality during the latter part of the
17th century. The crudely chipped implements may have been made
by the last native inhabitants of the site, thus representing the close
of the stone age in this part of Virginia.
The material collected on this very interesting site is now in the
United States National Museum and will be briefly described.
A very unusual specimen is shown in plate 3, the photograph being
exact size; material, greenish gray chlorite schist. It was found by
the writer on the surface near the center of the site, October, 1928.
This suggests a human head with a pointed base; extreme height
34 inches, width 12 inches, thickness from front to back 14 inches.
The material is comparatively soft and the surface of the stone has
probably become smooth and somewhat worn away during the years
since it was made, thus losing some of its sharpness and detail which
it might otherwise have possessed. The true meaning or purpose
of the object is not known, but it at once suggests Beverley’s
reference to an idol which he discovered in a temple belonging to one
of the Algonquian tribes of Virginia, probably about the year 1700.
The various parts of the idol were found with the exception of the
head, which had been removed and secreted. This small stone head,
although found on the site of a Siouan village, may have been some-
NO, I2 MONACAN TOWNS IN VIRGINIA—BUSHNELL 23
thing of the same sort. The pointed projection was undoubtedly
fashioned to be inserted in a base or body to hold it in an upright
position. Unfortunately very little is known of the customs and
beliefs of the people who once occupied the ancient village.
Chipped objects, usually with ground edges and showing evidence
of much use, are quite numerous on the site and many have been
discovered during the past few years. All are crudely made—rough
but apparently well suited for some definite purpose. The actual use
of these specimens is not known although the majority may have been
implements used in and about the village. Possibly some were hafted
to serve as hoes in the gardens, others may have been inserted in
wooden handles forming serviceable weapons. Four distinct forms
are recognizable but no attempt will be made to distinguish them by
name. They will be described and referred to as types A, B, C, and D.
Examples of the four types, belonging to the collection, are illustrated
in the plates.
Type A, plate 4. This is the simplest form. The great majority
are made of greenstone, which occurs on the site. The two specimens
at the top of the plate are exceptionally well made—far superior to
the average. The cutting edge is sharp and well ground. The three
examples below are rather more characteristic and many similar pieces
have been recovered. The cutting edge of the largest of the three
has been battered and worn away, causing it to become concave, as is
shown in the photograph. However, the extremities of the edge, both
above and below, are smoothed from use.
Type B, plate 5. These are the most interesting and distinctive
objects found on the site. The flaring blade is quite unusual, and the
narrow base suggests the use of a wooden handle in which the stone
may have been inserted. Similar specimens have been recovered from
the site of Monahassanugh, on the bank of the James. (Compare
fig. 4.)
Type C, plate 6. Examples of this type are widely distributed
throughout piedmont Virginia and eastward. All are recognized by
the slight grooves on the narrow, opposite sides. They vary somewhat
in detail, but are usually very rough, crudely made, and very often
with the surface greatly weathered. These are far more numerous
than the better made specimens with ground or polished surfaces
and clearly defined grooves. Two of the latter are shown in plate 7.
Type D, plate 7. A type specimen is shown at the bottom of the
plate, the distinguishing feature being the two cutting edges at opposite
ends. A number of examples of this type have been discovered on
the site but the majority are crude and roughly made, some being
3
24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
fashioned from natural pebbles, and evidently intended for hard usage.
However, two other specimens, quite similar to the type specimen
which is illustrated, were found. The three were discovered within
a very small area. All are rather massive and approximately the same
size. The hafting evidently passed across the middle and rested in
the two slight grooves. The fourth specimen shown in this plate is a
fragment of a well finished implement which, after having been
broken, was used as a hammer. The two ends show the effect of long
use, being much battered and worn. In this condition it may or may
not have been attached to a handle.
With very few exceptions all specimens illustrated in the four
plates are made of greenstone or a related rock. Some are more
weathered than others, but whether this condition should be attributed
to greater age of the object or to the material of which it is made has
not been determined.
Small stone objects, in addition to the numerous arrowheads and
related forms, are found scattered over the surface of the site of the
village. Thus far very few have been recovered, and although they
are often fractured, they tend nevertheless to reveal some part of the
art of the inhabitants of the ancient settlement. Examples are illus-
trated in the upper part of plate 8. Top row: a is a curious object
made of soapstone. It appears to be complete and suggests, in form,
the claw of a bear. It is believed to have been a fetish rather than an
implement of any sort. Next, b is a small stone disk, maximum
diameter 13%; inches, thickness 7; inch. It is made of an igneous
rock, and the surface is now brownish and greatly weathered. There
appear to have been two small perforations on the edge less than
4 inch apart, but this part of the original surface has been broken
away, allowing only a section of the perforations to remain. The third
specimen on this row, c, is a fragment of a well made, polished tablet
which had probably been perforated. Its greatest thickness is about
zs inch; material, reddish brown slate.
Below the three pieces just described are four objects which may
be termed tools. The use of d and ¢ is not known, but the two chipped
specimens, f and g, show evidence of having been used as scrapers.
Both are made of chert.
On the lower part of plate 8 are shown typical examples of
projectile points, i, together with some larger pieces which may have
served as knives. All are made of grayish quartzite. There is also an
excellent example of a disk or blade, i, being one of two similar speci-
mens discovered on the surface of the slope of the hill rising just
east of the site. The greatest thickness of this piece is less than one
inch; material, grayish quartzite.
NWO: I2 MONACAN TOWNS IN VIRGINIA—BUSHNELL 25
Fragments of two banner stones were found. Both are made of the
same material, a light greenish-gray talc schist. The larger piece repre-
sents about one-half of the entire specimen. As restored it is shown in
figure 6, full size.
The majority of the arrowheads recovered from the surface of
the site are made of the white quartz which is so plentiful throughout
the region. And of these more than one-half are of the simple tri-
angular type, without notches. Many are roughly made. Examples
are presented in plate 9, figure 2.
Two specimens are illustrated on this plate in addition to the quartz
points. Plate 9, figure I, a, is a pitted hammer stone made of quartzite
greatly weathered. On it are two pits, on opposite sides. The second
, i
ee re ee ee ee ee ee ee ee ee ee
oa
3 4
OO he ee ee re ee a ee ee Le
Fic. 6.—Banner stone as restored. Found on the site of Monasukapanough.
Natural size.
(U. S. Nat. Mus. Cat. Nag. 350135)
specimen, b, is smooth and worn; material, quartzite; thickness 12
inches. This may have been a chukey stone, used in the game, although
its shape suggests a muller, or mano stone, which could have been used
in conjunction with a mortar of the form shown in plate 10, figure 2.
This mortar was discovered near the center of the site; material,
coarse sandstone which occurs on the hill just east of the low ground ;
extreme length of slab of stone 154 inches ; depth of depression 1 inch.
USE OF SOAPSTONE
A small object made of soapstone has already been described. Three
fragments of vessels made of the same material have been found on
the surface, but no trace of a tobacco pipe or of an ornament of any
sort has been encountered. However, there is proof that small pieces
of the stone were worked into shape. It is evident bits of the stone
26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, 82
were carried from the quarries to the village, there to be cut and made
into finished products, but what they were has not been discovered.
Four small pieces of soapstone have recently been found on the
site, not far from the left bank of the Rivanna and all close together.
These show the effect of having been sawed on opposite sides, then
broken apart. Three of the specimens are figured in plate 10, figure 1,
and reveal clearly the shallow grooves worn in the soft stone during
the process of cutting. A piece of stone had probably been used in
sawing the comparatively soft material, and this had evidently been
quite thick as is indicated by the angle of the cut surface of the re-
maining portion.
Another specimen of soapstone, showing the effect of sawing and
breaking, is in the collection of the National Museum (U. S. Nat.
Mus. Cat. No. 170257). This was found at the junction of the Dan
and Staunton Rivers, near Clarksville, Mecklenburg County, where
the two streams unite to form the Roanoke. Eastward, in the adjoin-
ing county of Brunswick, stood Fort Christanna. The region was
much ‘frequented by Indians, and the Saponi and Tutelo—the ancient
inhabitants of Monasukapanough and Monahassanugh—are known to
have occupied islands in the Roanoke for a short period during their
southern movement, before turning northward to settle at Fort
Christanna.
_ CORDS, TEXTILES, AND BASKETRY AS REVEALED BY
IMPRESSIONS ON POTTERY
The journal written by Col. William Byrd while engaged in running
the dividing line between Virginia and Carolina contains much inter-
esting information. In it are many references to the beliefs and
customs of the Indians, and much of this is believed to have been
related by Bearskin, the old Saponi hunter and guide from the village
near Fort Christanna, who accompanied the expedition and served it
so faithfully.
On November 10, 1728, Byrd wrote in part:* “One of the men,
who had been an old Indian trader, brought me a stem of silk grass,
which was about as big as my little finger. But, being so late in the
year that the leaf was fallen off, I am not able to describe the plant.
The Indians use it in all their little manufactures, twisting a thread
of it that is prodigiously strong. Of this they make their baskets
and the aprons which their women wear about their middles, for
decency’s sake. These are long enough to wrap quite round them and
1 Byrd, William, The Westover manuscripts: containing the history of the
dividing line. .... Petersburg, 1841.
NO, 12 MONACAN TOWNS IN VIRGINIA—BUSHNELL 27
reach down to their knees, with a fringe on the under part by way
of ornament.”
The plant to which Byrd referred was undoubtedly a milkweed,
probably Asclepias pulchra, often termed Indian hemp, and known to
many persons in this part of Virginia as silk weed. The long fiber
is easily detached from the stalk, and when twisted forms a very
tough and strong cord.
Milkweed is plentiful throughout the region, and the fiber derived
from the stem was undoubtedly used by the people who occupied the
ancient village which stood on the banks of the Rivanna. Innumerable
small fragments of pottery have been recovered from the surface of
the site, many of which bear the imprint of cords or of basketry.
These fragments have been found on both sides of the river, but all ex-
amples to be considered at this time have been collected on the left
bank and, with few exceptions, have been found within 50 yards of
the water. The land has been plowed and harrowed for many years
and as a consequence it is seldom that a piece of pottery more than
an inch in length can be discovered. Much of it may never have been
very hard and the bits have now worn away until the impressions on
the surface of many have become faint and scarcely discernible. Casts
of nine small pieces are shown natural size in plate 11, figure 2. These
are the clearest impressions selected from several hundred.
The specimen in the middle of the top row is unusually hard and
compact, and for that reason the impression of cords has remained
very clear and distinct. It is an interesting fragment, but whether it
represents basketry or cloth has not been determined. On either side
of this are examples of coarser cords. The small triangular fragment
in the middle of the bottom row, and likewise the specimen on the
left, is believed to represent a form of basketry. No piece yet found
bears the imprint of a coarse, net-like material.
White quartz, pulverized or reduced to very small pieces, was mixed
with the clay of which the vessels were made.
As mentioned, the great majority of fragments of pottery have
been found near the left bank of the river, where the ground is rela-
tively high and has seldom been overflowed. A few pieces of rock
showing the effect of fire together with small bits of charcoal have
been revealed by the plow near the highest point of the bank, some
20 yards from the water. Quantities of flakes of quartz, quartzite,
chert, and some of jasper occur on this part of the site, and near by
were found the several pieces of worked soapstone already described.
This may have been a much frequented section of the village and
possibly sweat houses once occupied the high ground overlooking
28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
the water, with the mat- or bark-covered habitations scattered through
the surrounding wooded area—now cultivated lands where the crude
stone implements are found.
Many objects have already been recovered from the site but it is
expected that others, of equal or even greater interest, may be dis-
covered during the continued examination of the area, to be described
at another time.
THE. ~SAPPONEY, INDIAN TOWN”
There is no known record of a white man having visited Mona-
sukapanough, the ancient Saponi village on the banks of the Rivanna,
and consequently no description of the settlement has been preserved.
It was probably an extensive and important center. It is believed
that some time before the year 1670 the people, or at least the greater
part of them, moved from the valley of the Rivanna and went south-
ward to establish a new village which, according to Mooney, “‘ was
probably on Otter river, a northern tributary of the Roanoke, in what
is now Campbell county, Virginia, nearly south of Lynchburg.” *
Here they were visited by Lederer in 1670, and by the Batts party
during the following year, but these explorers failed to describe the
settlement. Soon the movement was resumed; they wandered far,
nearly reaching the center of North Carolina, later returning to
Virginia.
A generation after their first contact with Europeans, through
the influence of Governor Spotswood, the Saponi and remnants of
other tribes became established in the vicinity of Fort Christanna,
about 10 miles north of Roanoke River, in the present Brunswick
County, Virginia.
Although the Saponi had undoubtedly changed greatly from their
primitive state, yet they must have retained many of the manners and
ways of life practiced in earlier years at their ancient home on the
Rivanna. An interesting and at this time most valuable account of
the people as they appeared in the spring of 1716 is to be found in
the journal of one who visited them at that time.” The journal is a
record of a journey made by Fontaine and Governor Spotswood from
Williamsburg to Fort Christanna and return during the first ten days
of April, 1716. To quote from the journal: “ The 5th day—After
breakfast, I went down to the Sapponey Indian town, which is about
a musket-shot from the fort. I walked round to view it. It lieth in
NOp.icits p. 34:
*Journal of John Fontaine. In Memoirs of a Huguenot family, by Ann
Maury. New York, 1853.
NO: 52 MONACAN TOWNS IN VIRGINIA—BUSHNELL 29
a plain by the river-side, the houses join all the one to the other,
and altogether make a circle; the walls are large pieces of timber
which are squared, and being sharpened at the lower end, are put
down two feet in the ground, and stand about seven feet above the
ground. These posts are laid as close as possible the one to the
other, and when they are all fixed after this manner, they make a
roof with rafters, and cover the house with oak or hickory bark,
which they strip off in great flakes, and lay it so closely that no
rain can come in. Some Indian houses are covered in a circular
manner, which they do by getting long saplings, sticking each end
in the ground, and so covering them with bark; but there are none
of the houses in this town so covered. There are three ways for
entering into this town or circle of houses, which are passages of
about six feet wide, between two of the houses. All the doors are on
the inside of the ring, and the ground is very level withinside, which
is in common between all the people to divert themselves. There is
in the centre of the circle a great stump of a tree; I asked the reason ~
they left that standing, and they informed me it was for one of their
head men to stand upon when he had anything of consequence to
relate to them, so that being raised, might the better be heard.” Con-
tinuing he described briefly the interior of the structures: ‘‘ Their
houses are pretty large, they have no garrets, and no other light than
the door, and that which comes from the hole in the top of the house
which is to let out the smoke. They make their fires always in the
middle of the house; the chief of their household goods is a pot and
some wooden dishes and trays, which they make themselves; they
seldom have any thing to sit upon, but squat upon the ground ; they
have small divisions in their houses to sleep in, which they make of
mats made of bullrushes ; they have bedsteads, raised about two feet
from) the ground, upon which they lay bear and deer skins, and all
the covering they have is a blanket. These people have no sort of
tame creatures, but live entirely upon. their hunting and the corn
which their wives cultivate. They live as lazily and miserably as any
people in the world.
“ Between the town and the river, upon the river side, there are
several little huts built with wattles, in the form of an oven, with
a small door in one end of it; these wattles are plaistered without
side very closely with clay, they are big enough to hold a man, and are
called sweating-houses. When they have any sickness, they get ten
or twelve pebble stones which they heat in the fire, and when they
are red-hot they carry them into these huts, and the sick man or
woman goes in naked, only a blanket with him, and they shut the
30 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
door upon them, and there they sit and sweat until they are no more
able to support it, and then they go out naked and immediately jump
into the water over head and ears, and this is the remedy they have
for all distempers.”
The appearance of the Indians, and the manner in which the
Governor was received the day of his arrival at the fort, was told
by Fontaine. He wrote in his journal: ‘About three of the clock,
came sixty of the young men with feathers in their hair and run
through their ears, their faces painted with blue and vermilion,
their hair cut in many forms, some on one side of the head, and
some on both, and others on the upper part of the head, making it
stand like a cock’s-comb, and they had blue and red blankets wrapped
about them. They dress themselves after this maner when they go
to war the one with the other, so they call it their war dress, and it
really is very terrible, and makes them look like so many furies.
These young men made no speeches, they only walked up and down,
" seeming to be very proud of their most abominable dress.
“After this came the young women; they all have long straight
black hair, which comes down to the waist, they had each of them a
blanket tied round the waist, and hanging down about the legs like
a petticoat. They have no shifts, and most of them nothing to cover
them from the waist upwards ; others of them there were that had two
deer skins sewed together and thrown over their shoulders like a
mantle. They all of them grease their bodies and heads with bear’s
oil, which, with the smoke of their cabins, gives them an ugly smell.
They are very modest and very true to their husbands. They are
straight and well limbed, good shape, and extraordinary good features,
as well the men as the women. They look wild, and are mighty shy
of an Englishman, and will not let you touch them. The men marry
but one wife, and cannot marry any more until she die, or grow so
old that she cannot bear any more children; then the man may take
another wife, but is obliged.to keep them both and maintain them.
They take one another without ceremony.”
The children were bound to boards that were “ cut after the shape
of the child,” with two pieces at the bottom to which the child’s legs
were tied. A cord passed through a hole in the top of the board with
which it could be attached to a limb of a tree, ‘‘ or to a pin in a post
for that purpose, and there the children swing about and divert
themselves, out of the reach of any thing that may hurt them.” They
were kept in this manner until they were about two years of age.
Governor Spotswood was evidently greatly interested in the
Indians and on April 6, 1716, “ asked the boys to dance a war dance,
SMITHSONIAN MISCELLANEOUS COLLECTIONS Wales S372 INO, G25 Lats}
Objects found on the surface, Monasukapanough. 4 natural size.
(U. S. Nat. Mus, Cat. Nos.: a, 350130; b, 3501333 c, 350134; d, 350131; e, 350132;
f, 9, 350145; h, 350150; 1, 350148)
4
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLE 825 INO a2t eile)
I. a, pitted stone; b, mano stone, Monasukapanough. 4 natural size.
(U. S. Nat. Mus. Cat. Nos:: a, 350140; b, 350141)
2. Objects found on the surface, Monasukapanough. 4 natural size.
(U. S. Nat. Mus. Cat. No. 350151)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLES 82.) NO. 12, PES Oo
1. Three pieces of soapstone showing the effect of sawing, Monasukapanough.
Natural size.
(U. S. Nat. Mus. Cat. No. 350137)
1
2. Sandstone mortar, Monasukapanough. About } natural size.
(U. S. Nat. Mus. Cat. No. 350142)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOE 825 NO a h2) Please
2)
2. Casts of impressions on fragments of pottery, Monasukapanough.
Natural size.
(U. S. Nat. Mus. Cat. No. 350154)
WO; E2 MONACAN TOWNS IN VIRGINIA—BUSHNELL 31
so they all prepared for it, and made a great ring; the musician
being come, he sat himself in the middle of the ring; all the instrument
he had was a piece of board and two small sticks; the board he set
upon his lap, and began to sing a doleful tune, and by striking on the
board with his sticks, he accompanied his voice; he made several
antic motions, and sometimes shrieked hideously, which was answered
by the-boys. As the men sung, so the boys danced all round, endeav-
oring who could outdo the other in antic motions and hideous cries,
the movements answering in some way to the time of the music.
All that I could remark by their actions was, that they were repre-
senting how they attacked their enemies, and relating one to the
other how many of the other Indians they had killed, and how they
did it making all the motions in this dance as if they were actually
in the action.”
It is within reason to believe that among the Indians who gathered
at the Saponi village early in April, 1716, to greet Governor Spots-
wood were some who, as children, had lived at Monasukapanough on
the banks of the Rivanna. But during that period of wandering,
changes had taken place in the habits and ways of life of the people;
nevertheless much that was witnessed and recorded by Fontaine had
probably been similarly enacted long before at the older town. The
earthen pots and wooden dishes and trays, the mats made of bul-
rushes, the mantles made of two deer skins sewed together, and the
small sweat houses standing near the river bank—all were details
that would have been seen at the more primitive town on the Rivanna.
In 1728, 12 years after Governor Spotswood’s visit to Fort Chris-
tanna, the line between Virginia and North Carolina was being run
westward from the coast. On September 29 Col. William Byrd, of
the Virginia commission, secured a Saponi Indian from the village
near the fort to serve as guide and hunter for the party during the
latter part of their journey through the wilderness. To the English
he was known as Bearskin. He was worthy and capable, and kept the
camp well supplied with game. On Sunday, October 13, he explained
the religion of the Saponi which was recorded at length by Byrd—’*
a remarkable account of the primitive beliefs of a tribe of which
so little is known.
The habitations at the Saponi village near Fort Christanna, as
described by Fontaine in 1716, were quite unusual and are believed
to have been of English conception. It is doubtful if any structure
of a similar nature ever stood at Monasukapanough, where the small
Of. c., B: 51.
32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, 82
habitations were undoubtedly the typical mat- or bark-covered lodges.
But strongly made log structures were to have been encountered else-
where in Virginia, and their occurrence was likewise recorded by
Fontaine. In the month of June, the year before he visited the Saponi
settlement with Governor Spotswood, Fontaine made a journey from
Williamsburg to the German colony on the Rappahannock. He had
crossed the Mattaponi and was in King William County when at
some point on the left or north bank of the river, possibly about due
north of the present Pamunkey Indian Reservation, he encountered
a single Indian habitation. This was June 12, 1715, and he wrote in
his journal that day:* “ The day very windy. We see by the side of
the road an Indian cabin, which was built with posts put into the
ground, the one by the other as close as they could stand, and about
seven feet high, all of an equal length. It was built four-square, and
a sort of a roof upon it, covered with the bark of trees. They say it
keeps out the rain very well. The Indian women were all naked, only
a girdle they had tied round the waist, and about a yard of blanketing
put between their legs, and fastened one end under the fore-part of
the girdle, and the other behind. Their beds were mats made of
bulrushes, upon which they lie, and have one blanket to cover them.
All the household goods was a pot.” Unfortunately Fontaine failed
to record the name of the tribe to which this family belonged, but the
lodge, its surroundings and the condition of its occupants, were
probably characteristic of the time and country and were in no way
exceptional. Indian families such as this, living off and apart from
others, would undoubtedly have been found in many parts of tide-
water and piedmont Virginia. And after the towns of Mowhemcho,
Massinacack, and Rassawek ceased to be important settlements, as
they were during the early years of the 17th century, many isolated
cabins would have been encountered within “ The Land called the
Monscane.”
The five Monacan towns, as they are known to have stood early in
the 17th century, have now been mentioned. It is believed the identi-
fication of the sites is accurate and conclusive. And although the
exact position of Rassawek has not been determined, it must be
agreed that the settlement was situated somewhere within a rather
restricted area between the James and Rivanna, on or near the right
bank of the latter stream and not far from its mouth. No other
Monacan villages were referred to by the early writers and if any
existed they have been lost to history. But throughout the entire
* Op. cit., p. 264.
i
NO; 12 MONACAN TOWNS IN VIRGINIA—BUSHNELL 33
region once dominated by the Siouan tribes—a region embracing the
valley of the James from the falls to the mountains, where wild game
was abundant and food easily procured—are to be found traces of the
period of Indian occupancy ; village and camp sites on the banks of
streams, quarries where soapstone was obtained, and innumerable
stone implements scattered over the surface. However, as said else-
where, all must not be attributed to the Monacan tribes.
Scattered sites have been visited and examined and it is planned
to describe them, together with the material collected, in a subsequent
paper. Two localities will now be mentioned, both on the banks of
the Rivanna, one above, the other below the site of Monasukapanough,
and both believed to have been in some manner related to the great
village.
GARNETT SITE
The site of the large village, believed to have been Monasuka-
panough, is on the banks of the South Branch of the Rivanna, the
Fic, 7—Double edged instrument found on the Garnett site. Natural size.
: (U. S. Nat. Mus. Cat. No. 350156)
lesser North Branch joining it about 2 miles below. Some 7 miles
above this site, and 1 mile or more below the junction of Moormans
River and Mechum River—two streams which unite to form the
South Branch of the Rivanna—is one of the best places for fishing
on the entire river. On the right bank are great masses of rock against
which the water flows to a depth of 10 feet or more. Cliffs reach near
the bank on the opposite side. The water is usually clear and cold,
34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
and shaded along the banks by overhanging trees. Just below this
point a small creek flows into the Rivanna from the right or south.
Evidences of a camp have been discovered near the creek, but whether
it was large or small cannot be determined as the land has been culti-
vated for many years and the surface has often been covered by the
waters of the river. Many objects of stone and some fragments of
pottery have been found. The material is identical with that collected
from the surface at Monasukapanough, which suggests that this may
have been a fishing camp belonging to the people of the large village.
The characteristic chipped ax- or celt-like implements are found here
and the arrowheads are typical. One specimen from the site is quite
rare. A small double edged chisel or celt, made of greenstone, 32
inches in length. It is shown exact size in figure 7.
This is a beautiful spot, secluded and well protected, and could have
been reached by canoe from the large settlement down the river.
MOON SITE
Jefferson described the mound which he examined as being on the
low grounds of the Rivanna “ about two miles above its principal fork.”
The “ principal fork” is now known as the North Branch. Where
the two branches unite the main stream bends and flows in a more
southerly course. A wide bottom on the right extends far above and
below the mouth of the North Branch, on the opposite side, but this
is quite low and consequently often overflowed. Traces of an encamp-
ment were discovered near the south end of the low ground, back from
the river where the surface begins to rise. Several stone implements,
bits of pottery, and a single fragment of a well-finished soapstone
vessel were found scattered over the surface. Also many chips of
white quartz, and larger masses of the same from which smaller
pieces had been broken. Two small, crudely made, grooved imple-
ments were found on the higher ground not far from the residence.
They are typical examples of the widely distributed form shown in
plate 6.
High rugged cliffs rise a short distance from the left bank of the
river, opposite the southern part of the low grounds on the Moon
plantation. Quantities of white quartz encountered here may have
been one source of material for the making of arrowheads, and the
pieces discovered on the Moon site had probably been carried across
the stream from this outcropping. The cliffs and low grounds on the
left side of the river are heavily timbered and with much undergrowth,
and as a consequence it is very difficult to discover any traces of
Indian occupancy.
NO, 12 MONACAN TOWNS IN VIRGINIA—BUSHNELL 35
A cairn formed of rather small stones, shown in plate 12, figure 2,
about 3 feet in height and 15 feet in diameter, stands on the summit
of the cliff overlooking this part of the low grounds. The cliff is here
a narrow ridge and the cairn commands an extensive view of the
county westward to the Blue Ridge and in the opposite direction across
the intervening valley to the South Western Mountain. This is about
1 mile below the mouth of the North Branch.
The map of Virginia and Maryland, drawn by Augustin Herrman,
was completed in 1670 but not engraved and issued until three years
later. No Monacan towns are indicated on the banks of the James or
of its tributaries. Lederer made his journey up the valley of the
James during the months of May and June of that year, but evidently
his travels were not known to Herrman until the appearance of his
brief volume of “ Discoveries ” which was printed in London in 1672.
The engraving of the map may already have been completed by
William Faithorne in England and consequently it would have been
too late for Herrman to have added the newly acquired data.
The Herrman map shows the course of the James, with one large
tributary far to the westward. This was probably the Rivanna, known
from the earliest days of the colony. A legend on the map, placed
north of the latter stream, reads thus:
Mount Edlo This name derives from a Person that was in his
Infancy taken Prisoner in the last Massacra over Virginia,
And carried amongst others to this Mount, by the Indians,
which was their watch Hill, the country there about being
Champion and not much Hilly.
Mr. Fairfax Harrison has suggested to the writer the possibility
of Mount Edlo being some point in Albemarle County west of where
the Rivanna passes the South Western Mountain. If this is correct
the high narrow ridge, surmounted by the cairn, may have been the
spot indicated. The great village, Monasukapanough, was less than
3 miles distant, and the entire region shows evidence of Indian
occupancy. The country had probably not been seen by a European
and the description of the land, as recorded in the legend, was neces-
sarily vague and uncertain.
SOAPSTONE
Steatite, or soapstone, is found in several localities in the region
between the Rivanna and the James. It was quarried and used by the
Indians, and what is believed to be the most extensive quarry worked
by them within these bounds was discovered in 1926, on a high ridge
30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
a short distance south of Damon, Albemarle County. It is about 2
miles in a direct line north of east from Schuyler, and between 5 and
6 miles northwest from the nearest point on the left bank of the James.
Schuyler is on the banks of Rockfish River, some 6 miles above its
mouth, in Nelson County. Since 1926 the surface has been stripped
and quarries have been opened on the site, thus destroying traces of
the work done by Indians before the coming of Europeans to this
part of Virginia.
When visited in 1926* great masses of soapstone outcropped on
the surface. These followed a general direction from southwest to
northeast and had a dip of about 60° to the southeast. The broken,
irregular surface was heavily timbered, and evidently the entire region
had changed little in appearance during the past centuries. Near by
Fic. 8.—Fragment of an unfinished flat-bottomed soapstone vessel. 4 natural size.
(U. S. Nat. Mus. Cat. No. 332025)
were several large springs. For 1,000 feet or more along the ridge it
was possible to trace the pits, dug by Indians generations ago, from
which quantities of soapstone had been removed. Twenty or more
excavations were thus discovered and probably others were so filled
with the accumulated mold and moss as not to have been distinguish-
able. The pits varied from 10 to 30 feet in diameter and appeared
to have been from 2 to 4 feet in depth. Some were quite distinct ;
others may have been joined beneath the mass of mold and thus in
reality have been parts of a large excavation.
The surface surrounding the pits was covered with pieces of the
stone, some large, others small, which had been removed from the
quarries and evidently rejected as being unfit for use. But only a
Bushnell, David I., Jr., Ancient soapstone quarry in Albemarle County,
Virginia. Journ. Washington Acad. Sci., Vol. 16, No, 19, November 18, 1926.
NO. I2 MONACAN TOWNS IN VIRGINIA—BUSHNELL 37
small amount of the stone was visible, or projected through the thick
vegetal mold which had formed since the quarries were last used by
the Indians (pl. 13).
Many broken, unfinished vessels were discovered in the vicinity of
the pits. These had been fractured in the process of making and
abandoned as useless. Consequently this was not only a quarry but
a workshop, where the vessel was fashioned in the rough, later to be
smoothed and polished.
That two types of bowls were made here is indicated by the many
fragments recovered from the surface in the vicinity of the pits. One
type had a flat bottom; in the second type it was more rounded. An
admirable example of the first form, found near one of the pits on
the northern part of the ridge, is shown in plate 14, figure 1; length
Fic. 9.—Fragment of an unfinished, round-bottomed soapstone vessel.
+ natural size.
(U. S. Nat. Mus. Cat. No. 332024)
about 17 inches. It is very rough and unfinished, and the surface
reveals distinctly the irregular cuttings made by crude stone imple-
ments. One side had been broken, probably while it was being hol-
lowed out, and the vessel discarded. No attempt had been made to
smooth or finish the surface, either within or without. The knobs to
serve as handles project from the upper edge, at ends.
The second type, with rounded bottom, usually has knobs extending
from the ends an inch or more below the upper rim to serve for
handles. The two forms shown in figures 8 and 9 are examples found
on the surface near the pits. Of the latter type are the two fractured
specimens said to have been discovered in a cultivated field near the
foot of Buck Mountain, in Buckingham County. This would be a
short distance south of the right bank of the James, and a few miles
from the extensive village site, believed to have been Monahassanugh,
between Norwood and Wingina, in Nelson County on the opposite
side of the river. The two specimens, as restored, are shown in plate
38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
14, figure 2. The smaller is in the collection of the United States
National Museum. The inside diameters are 13% inches and 102
inches ; depth 53 inches; thickness from } to 3 inch. The larger speci-
men, not in the Museum collection, is of the same proportions but
with a maximum diameter, inside, of 16% inches.
Innumerable vessels and smaller objects were undoubtedly made
in the vicinity of the quarries to be carried away to distant villages,
as well as for use in nearby camps; nevertheless very few pieces of
soapstone vessels are found. Several small fragments have been dis-
covered on the site of Monasukapanough, together with four pieces
of the material that had been sawed. This latter may have been ob-
tained from an outcropping a few miles north of the ancient site.
Another small fragment of a well finished vessel was encountered
on the Moon site, already mentioned. Two tobacco pipes made of
soapstone are said to have been found some years ago on a ridge
just east of the Southern Railway at Arrowhead, about 8 miles south
of the University of Virginia. They were not seen by the writer but
were described as being quite small and very well made. The lack of
more traces of soapstone on the village and camp sites is difficult to
explain. Many vessels were made and used, as shown by evidence at
the quarries, but all have disappeared.
The quarries south of Damon may be the most extensive group
worked by Indians in this immediate section of Virginia, but there
are several other localities where soapstone was obtained and vessels
made. Extensive quarries, possibly worked by the people of the
ancient village of Monahassanugh, are situated midway between Nor-
wood, on the left bank of the James at the mouth of Buffalo River,
and Arrington, a small station on the Southern Railway in Nelson
County. Many broken, unfinished vessels have been discovered here,
and the two specimens shown in plate 14, figure 2, may have been
made at this quarry. Another quarry, not visited by the writer, is
in Goochland County about 8} miles in a direct line northeast of the
site of Rassawek, at the mouth of the Rivanna, and 14 miles due
south of the present village of Tabscott.
There are other outcroppings of soapstone in the region, in addition
to those already mentioned, which show evidence of having been
worked by Indians. And although all are well within the bounds of
the territory occupied by the Monacan tribes, the last of the native
tribes to claim this part of Virginia, the soft stone may have been
discovered and worked by others who had preceded them. Nor is
the scarcity of objects made of soapstone less mysterious than the
identity of the tribes by whom the quarries were opened and the
utensils and ornaments fashioned.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLS 2 NOW 12) Pema2
2. Cairn on summit of cliff below mouth of the North Fork.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82, NO. 12, PL. 13
eos Raat
SN adh
2. One of the larger pits worked by the Indians,
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82, NO. 12, PL. 14
1. Unfinished soapstone vessel from quarry south of Damon.
About } natural size.
(U. S. Nat. Mus. Cat. No. 332023)
2. Two soapstone vessels, restored, from near foot of Buck Mountain,
Buckingham County, Virginia.
(Upper, U. S. Nat. Mus. Cat. No. 342083; lower, owned by Mrs. Wirt Robinson)
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 82, NUMBER 13
A NOTE ON THE SKELETONS OF TWO
ALASKAN PORPOISES
(WiTH ONE PLATE)
BY
GERRI Ss VIGEERT Sik:
Curator, Division of Mammals, U. S. National Museum
(PUBLICATION 3107)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
DECEMBER 23, 1936
The Lord Gaftimore Press
BALTIMORE, MD., U. S A.
A NOTE ON THE SKELETONS OF TWO ALASKAN
PORPOISES
By GERRIT SS: MILLER; JR.
CURATOR, DIVISION OF MAMMALS, U. S. NATIONAL MUSEUM
(WitH ONE PLaTeE)
Skeletons of two Alaskan porpoises, Phocena phocena (Linnzus)
and Phocenoides dalli (True), have recently been mounted in the |
United States National Museum. Before these skeletons were placed
on exhibition the photographs were made that are reproduced on the
plate accompanying this article.
In his original account of Phocenoides (Bull. Amer. Mus. Nat.
Hist., Vol. 30, pp. 31-50, May 16, 1911) Andrews figured the
skull, jaw, hyoids, sternum, scapula, flipper, and several vertebrae;
but these individual parts fail to give an adequate idea of the striking
peculiarities of the genus as compared with its relative Phocena.
Chief among these peculiarities are the length and slenderness of the
chevron bones, of the ribs (length of longest rib contained about
44 times in length of vertebral series, as compared with 54 times in
Phocena), and of the dorsal spines (longest spines contained about
104 instead of 174 times in length of vertebral series) ; the shortness
(or compression) of the vertebral centra by virtue of which the 45
centra of the dorso-lumbar series are crowded into appreciably less
space than that occupied by only 29 dorso-lumbar centra in Phocena.
Particularly noticeable also are the differences in form of the scapula,
arm and hand.
In the great height of the dorsal spines and the compression of the
centra Phocenoides appears to be unique among living members of
the family Delphinide. There is no approach to these conditions in
the genus Neomeris.
The two skeletons, as mounted, are of the same length, 1,760 cm.
In the Phocena the depth of chest, including vertebral spines, is about
30 cm., while in the Phocenoides it is about 44 cm.
Considering the peculiarities of the skeletons, the skulls of Phocena
and Phocenoides are remarkably alike. In the teeth and gums, how-
ever, the differences are pronounced.
*See Miller, Proc. U. S. Nat. Mus., Vol. 74, Art. 26, pp. 1-4, pls. 1-4,
January 26, 1920.
SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 82, No. 13.
to
SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
EXPLANATION OF PLATE
Both figures greatly reduced to approximately uniform scale
Upper Fic. Phocena. No. 218737, U. S. Nat. Mus. Collected at St. George
Island, Alaska, by C. D. Hanna.
Lower Fic. Phocanoides. No. 219334, U. S. Nat. Mus. Collected near
Wrangell, Alaska, by E. P. Walker. (The skull is tilted slightly to the right.
This exaggerates the flatness of the rostrum as compared with the rostrum
of Phocena.)
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ba cil ae Le FP: PRL ee | he oo
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, a
L “Id ‘€L “ON 28 “100A SNOILO311090 SNOANVIISOSIN NVINOSHLIWS
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 82, NUMBER 14
THE SUPPOSED OCCURRENCE OF AN ASIATIC
GOAT-ANTELOPE IN THE PLEISTOCENE
OF COLORADO
(WitTH Two PLaTEs)
BY
GERRIT S. MILLER, JR.
Curator, Division of Mammals, U. S. National Museum
(PUBLICATION 3108)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
DECEMBER 22, 1930
The Lord Baltimore Press
BALTIMORE, MD., U. S. A.
THE SUPPOSED OCCURRENCE OF AN ASIATIC GOAT-
AINTEEOPE IN THE PLEISTOCENE: OF
COLORADO
By GERRIT S. MILLER, JR.
CURATOR, DIVISION OF MAMMALS, U. S. NATIONAL MUSEUM
(WitH Two Pirates)
In the Bulletin of the Geological Society of America, Vol. II, pp.
610-612, pl. 57, August 10, 1900, Mr. F. W. Cragin described and
figured the right humerus and right metacarpal of a “ Goat-antelope
from the Cave Fauna of Pike’s Peak Region.” He proposed for this
animal the new name Nemorhedus palmeri, and concluded that the
discovery of the two leg bones extended the former range of the
Asiatic genus Nemorhedus to Colorado. Furthermore he reasoned
that :
Tf the range of the Pike’s Peak capricorn corresponded nearly with that of the
Himalayan, and the cave of the capricorn-eating carnivore was conveniently
located within the zone of the greatest abundance of the quarry—s5,000 to 6,000
feet above sea level—the Rocky Mountain plateau must have stood something
like one or two thousand feet lower in its capricorn epoch than today, as the
present elevation of the cave approaches 7,000 feet; and as the two conditions
above predicated are those most likely to have prevailed, it seems quite probable
that Nemorhadus, as an element of the North American fauna, belonged to
the Champlain phase of the Glacial epoch.
Apparently no one has yet submitted these findings to critical ex-
amination. Hay (The Pleistocene of the Middle Region of North
America and Its Vertebrated Animals, 1924, pp. 144, 273, 275) ac-
cepts the species as a genuine member of the Pleistocene fauna of
Colorado.
The specimens of Nemorhadus palmeri are now in the United
States National Museum. They were originally entered under the
number 8042, Division of Vertebrate Paleontology, but they have now
received the number 255680 in the Division of [recent] Mammals.
A few months ago, at the request of Mr. E. R. Warren, these speci-
mens were examined by Dr. J. W. Gidley, who concluded that they are
geologically recent in origin and not of Pleistocene age. He therefore
asked me to compare them with the corresponding parts of such un-
gulates as now occur in the Rocky Mountain region. On doing this
SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 82, No. 14
2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
I am unable to find any characters by which they can be distinguished
either generically or specifically from existing American sheep, not-
withstanding Mr. Cragin’s belief (Colorado College Studies, Vol. 8,
p. 23, April, 1900) that they “ differed widely ” from the correspond-
ing bones of these animals.
The similarity in both size and form between the type specimen,
the humerus, of “ Nemorhedus palmeri” and the humeri of three
bighorns is made sufficiently evident by the photographs reproduced
in plates 1 and 2. Individual peculiarities can be seen in the cave bone
as in each of the others; but the specimen appears to be characterized
by nothing more important.
Finally, the condition of the femur is such as to lend no support to ,
the idea that the bone pertained to a member of the Pleistocene fauna.
There is no indication of mineralization. On protected parts there are
thin deposits of a fine reddish dust that is readily removed with water,
leaving the surface clean and fresh in appearance. When charred a
small fragment gives off the characteristic odor of burned bone.
In the absence of evidence to the contrary, I therefore have no hesi-
tation in regarding the name Nemorhedus palmeri Cragin as a
synonym:of Ovis canadensis Shaw. Mr. Warren writes me under
date of August 18, 1930, that the type locality is “ Glen Eyrie,” the
former home of General Palmer, for whom the supposed new species
was named. This place is in a valley or canyon about five miles north-
west of Colorado Springs, across the “ Mesa.”
EXPLANATION OF PLATES 1 AND 2
All figures about two-thirds natural size
Fic. 1. Ovts dalli Nelson. Female. Lapierre House, Yukon, Canada. No. 20963,
U. S. Nat. Mus.
Type of Nemorhedus palmeri Cragin. Near Colorado Springs, El Paso
County, Colorado. No. 255680, U. S. Nat. Mus.
Fic. 3. Ovis sp. Male. Exact locality unknown’ (received alive from D. E.
Wintermute, Gila, Arizona). No. 49777, U. S. Nat. Mus.
Fic. 4. Ovis canadensis Shaw. Male. Delta County, Colorado. No. 49704,
U. S. Nat. Mus.
Fic.
Ny
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOR S2 INO 4 erie
Right humerus of four specimens of Rocky Mountain sheep (Fig. 2,
type of Nemorhadus palmerit Cragin).
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82, NO. 14, PL.
Right humerus of four specimens of Rocky Mountain sheep (Fig. 2
type of Nemorhadus palmeri Cragin).
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 82, NUMBER 15
THREE SMALL COLLECTIONS OF MAMMALS
FROM HISPANIOLA
(WitH Two PLatTEs)
BY
GERRIT S. MILLER, JR.
Curator, Division of Mammals, U. S. National Museum
(PUBLICATION 3109)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
DECEMBER 24, 1930
The Lord Baftimore Press
BALTIMORE, MD., U. 8. A.
THREE SMALL COLLECTIONS OF MAMMALS FROM
HISPANIOLA
By GERRIT S. MILLER, JR:
CURATOR, DIVISION OF MAMMALS, U. S. NATIONAL MUSEUM
(WitH Two PLatEs)
The United States National Museum has received from Haiti and
the Dominican Republic three small collections of mammals that have
not yet been reported on.
Of these, the first was made in a sheltered side crevice, probably
once the nesting place of the giant Haitian barn owl, near the bottom
of a deep sink hole called the Trujin, on the massif of La Selle, Haiti.
How he explored this cavity by means of a tall pine, felled and lowered
into the hole to serve‘as a ladder, has been told by Dr. Alexander Wet-
more in “ Explorations and Field-Work of the Smithsonian Institu-
tion in 1927,” p. 36. The bones from this source are particularly in-
teresting because they represent an almost “ pure culture” of the
native mammal fauna, nearly uncontaminated by introduced European
rodents. Among the Trujin remains is the most nearly complete skull
of Brotomys yet collected, and a series of Nesophontes skulls that
indicates the presence of well-defined sexual characters independent
of size.
The second collection was made during March, 1929, in caves near
En Café on Gonave Island, Haiti, by Arthur J. Poole, who has de-
scribed his experiences in “ Explorations and Field-Work of the
Smithsonian Institution in 1929,” pp. 71-73. It shows that most of
the genera of extinct mammals found in the caves of the Haitian
mainland were also represented in the fauna of Gonave.
The third collection is from the neighborhood of Constanza, in
the mountainous interior of the Dominican Republic (altitude about
4,000 feet), a region that Herbert W. Krieger visited during the
spring of 1930. It consists of two lots of bones. One of these was
dug from an Indian refuse deposit, about 25 feet deep, on the valley
floor at Cerro de Monte, 6 km. east of Constanza. The other was con-
tained in owl pellets found in a shelter under an overhanging ledge
about 100 feet up the northern flank of Monte Culo de Maco above
the Arroyo Limoncillo at a point some 10 km. southwest of Constanza.
The mass of about half a peck of dry, partly disintegrated pellets was
SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 82, No. 15
bo
SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
brought entire to Washington. In it I found, for the first time, re-
mains of Nesophontes, Brotomys, and Isolobodon in dejecta that
appear to be those of the small living barn owl. As usual with this
bird the ubiquitous roof rats made up the bulk of the food, as indi-
cated by the undigested remains. The Brotomys must have been about
the same size as one of these rats, while the /solobodon, a very young
individual, could not have been much larger. The finding of these
remains in recent owl pellets, and particularly in pellets from the rain
forest region where there is no reason to suppose that disintegration
is likely to be very long delayed, as it might be in the dry parts of the
island, is extremely important. It points to the probability that mem-
bers of three of the supposedly extinct genera of Hispaniolan mam-
mals may, in reality, be still alive.
(1) TRUJIN COLLECTION
NESOPHONTES HYPOMICRUS Miller
Anterior part of skull, 5; palate, 1 ; right maxilla, 3 ; left maxilla, 5;
right mandible, 20; left mandible, 20; periotic bones, 30; tympanic
rings, 10; sternal manubria, 7; scapulae, 2; humeri, 43; ulnae, 27;
radii, 4; innominates, 21; femora, 52; tibiae, 25; astragali, 35; cal-
canea, 42.
Two of these skulls first called my attention to the fact, afterward
verified in many specimens from other localities, that individuals of
Nesophontes hypomicrus (pl. 2, figs. 1 and 1a) and N. paramicrus
(pl. 2, figs. 2 and 2a) possess upper canine teeth that are either high
and strong or low and weak. The conditions in N. zamicrus are not
yet known. That the weak teeth are not deciduous canines is shown
by their presence in old individuals with much worn molars. It seems
plain, therefore, that the differences should be attributed to sexual
dimorphism and that the individuals with weak canines should be
regarded as females, those with strong canines as males. While this
conclusion seems necessary it must be admitted that such sexual
differences in the size and form of the canine are unusual if not
unique among the insectivores. I have been unable to find an approach
to it in any of the insectivore genera represented in the National
Museum collection.
EPTESICUS HISPANIOLZ Miller
Skull, nearly perfect, 1; broken rostra, 3; fragments of palate, 37 ;
mandibles, 89. Also many periotics, finger bones, and loose teeth.
The preponderance of remains of this species in the food of an
owl that had a varied bat fauna to select from is rather remarkable.
NO: 15 MAMMALS FROM HISPANIOLA—MILLER
W
TADARIDA sp.
Mandibles, 2.
RATTUS RATTUS subsp.
Mandible, 1 ; auditory bulla, 1 ; femur, 1; tibia, 1; heads of tibiae, 3 ;
astragali, 2; calcaneum, 1; ulna, I.
These few rat bones were the only remains of introduced European
rodents found in the deposit. Bones of the house mouse were entirely
absent.
BROTOMYS VORATUS Miller
Imperfect skull, 1; palate and rostrum, 1; right maxillae, 2; left
maxilla, 1; left nasal, 1 ; fragmentary palates, 5; loose cheekteeth, 27 ;
pieces of frontal, 2; pieces of braincase, 2; occipitals, 1; auditory
bullae, 2; mandibles, right, 5; left, 5; humeri, 2; ulnae, 4; radius, 1 ;
innominates, I2; sacra, 4; femora, 9; tibiae, 7; heads of tibia, 4;
astragalus, 1; calcaneum, I.
The skull (pl. 1, fig. 2), though imperfect, is the most nearly com-
plete specimen that has yet been found. It permits, for the first time,
a rather full comparison with the skull of Proechimys (pl. 1, fig. 1),
and it shows conclusively that, so far as cranial characters are con-
cerned, the differences between the two animals are of no more than
generic importance. Chief among the characters in which the island
genus differs from its mainland relative are the broader, deeper, less
downward bent rostrum, deeper zygoma (in a perfect specimen this
would be even more evident than it is in the slightly injured zygoma
of the Trujin skull), less developed supraorbital bead (not well shown
in the Trujin skull but obvious in other specimens), and less contrast
in size and form between the alveolar pit made by the inner root of
each cheektooth and the two pits made by the outer roots. Apparently
there is less space between the paroccipital process and the posterior
margin of the glenoid fossa, but this may be partly or wholly due to
incorrect reconstruction of this part of the skull. The auditory bulla
in the Trujin specimen is not exactly in place; it should lie about 1.5
mm. farther back. As compared with the mandible of Proechimys
semispinosus that of Brotomys voratus (see Smithsonian Misc. Coll.,
Vol. 82, No. 5, pl. 1, fig. 3, December 11, 1929) is more robust. In
particular the depth at the sigmoid flexure is noticeably greater in
proportion to the total length of the mandible. The coronoid process
is larger and the angular process is longer.
In its dental characters Brotomys likewise differs no more from
Proechimys than the latter differs from some of its living South
x.
4 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
American relatives. The upper incisors are more abruptly curved and
their course is slightly if at all evident on the side of the rostrum or
in the antorbital foramen. Similarly the shaft of the lower incisor is
less conspicuous on the under side of the mandible; its base does not
extend quite so far beyond the root of m,; as in the case of Proechimys.
All the maxillary cheekteeth are 3-rooted as in Proechimys, but the
inner root is merely larger than either of the others and not specialized
in form as it is in the South American animal. In both genera the
mandibular teeth are 3-rooted with the exception of the 2-rooted
premolar. The large root is placed behind and the two small roots
in front.
The enamel pattern is less complicated than that of Proechimys
semispinosus. In all of the teeth except pm, the pattern is unmodified
pentamerous with the two reentrant folds of the paramere slightly
deeper than the single reentrant of the protomere. In the lower pre-
molar the protomere bears a shallow second reentrant fold near the
front of the crown. This fold is so shallow that it is soon cut off as an
enamel lake. No species of Proechimys that I have seen is as simple
as this in its enamel folding. The fundamental conditions. present
in Brotomys are, however, exactly reproduced in Cercomys, though in
this South American genus there is a slight specialization in the form
of a peripheral narrowing of the reentrant folds of the metamere
which causes these folds to become early isolated as lakes.
PLAGIODONTIA HYLAUM Miller
Right mandible (no teeth), 1; left auditory bulla, 1; fragment of
ulna, 1; fragments of innominates, 2; femur, 1; tibia, 1 ; epiphysis
from head of tibia, 1; astragalus, 1; calcaneum, 1.
PLAGIODONTIA sp. ?
A first or second upper molar, a first or second lower molar, and
three separate plates of a larger tooth, apparently m,; or mz, suggest
the occurrence of a Plagiodontia with teeth more compressed antero-
posteriorly than they are in any of the three species now recognized.
Two auditory bullae slightly different from those of P. hyleum may
pertain to this animal.
ISOLOBODON LEVIR Miller
Probably two individuals, both immature, represented by the left
half of a palate and three loose teeth.
INOS LS MAMMALS FROM HISPANIOLA——MILLER 5
(2) GONAVE COLLECTION
NESOPHONTES HYPOMICRUS Miller
Two skulls lacking most of the braincase. One retains the right
canine; this tooth has the form and size that appears to indicate the
female sex.
BROTOMYS VORATUS Miller
Fragment of palate, 1; mandibles, 35.
These specimens appear to agree in all respects with those col-
lected on the Haitian mainland.
ISOLOBODON PORTORICENSIS Allen
Skull, lacking rostrum and teeth, 1; palate with three teeth, 1;
right upper premolar, 1; mandibles, 23 ; humerus, 1.
All of this material is referable to the large Jsolobodon. The tooth-
rows of the 10 largest mandibles range from 19 to 20 mm. in alveolar
length, thus closely agreeing with the measurements of I1 jaws of
I. portoricensis from the mouth of San Juan River, Dominican Re-
public (19 to 20.8 mm.), and decidedly exceeding those of 15 jaws
of /. levir picked for their large size from a series of 281 collected in
the vicinity of Monte Cristi (16 to 17.6 mm.).
APHATREUS MONTANUS Miller
Six mandibles, all from immature individuals.
HEXOLOBODON PHENAX Miller
Imperfect mandible of a young individual, with pm, in place.
Two bullae, an atlas, and the lower extremity of a femur, all of
which appear to be too large to have pertained to an /solobodon, may
represent this animal.
ACRATOCNUS COMES Miller
Upper canine, 1; imperfect molariform teeth, 2; left fourth meta-
carpal, 1; phalanges, 2; centrum of vetebra, 1.
(3) CONSTANZA COLLECTION
(A) OWL DEPOSIT
NESOPHONTES PARAMICRUS Miller
Anterior part of skull, 1; braincase, 1; mandibles, 11; humeri, 4;
femora, Io.
6 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
All of these bones were found in the mass of partly disintegrated
owl pellets. Many of them have the appearance of such freshness that
it is easy to believe that they were dropped by the owls within a period
not greater than a year or two before the time of Mr. Krieger’s work.
One of the femurs, for instance, retains a patch of dried tissue on the
anterior basal portion of the greater trochanter and a loose web of
hairs in the digital fossa and concave inner aspect of the greater
trochanter. The braincase (pl. 2, fig. 3) is packed full of hair by the
action of the owl’s stomach, and the broken anterior part of a skull
(pl. 2, fig. 3a) gives similar evidence of recent submission to digestive
action. Most of the jaws have hairs adhering to the teeth or in the
spaces between the roots. One has a felt-like mass plastered against
the inner side of the ascending ramus.
As regards specific characters this material appears to be in per-
fect accord with the original material from St. Michel, Haiti.
NESOPHONTES HYPOMICRUS Miller
Imperfect rostra, 2; mandibles, 14; humerus, 1 ; femora, 3; tibiae, 4.
In one of the imperfect rostra the antorbital canal is packed with
hair and there are tufts of hair in the spaces between the teeth. Most
of the jaws have hairs adhering to the teeth:
NESOPHONTES ZAMICRUS Miller
A mandible, an ulna and a tibia represent this species. All three
bones have traces of hair and other organic matter adhering to their
surfaces. The mandible measures: total length, about 13 mm.; depth
through coronoid process, 4.0; distance from articular process to
anterior border of first molar, 8.4; combined four molariform teeth
(alveoli), 5.2.
PHYLLOPS HAITIENSIS (J. A. Allen)
Broken skull of one individual and mandible of another.
EPTESICUS HISPANIOLZ Miller
Broken skull and nearly perfect jaw, both apparently parts of
one animal. The scarcity of bat remains is a peculiar feature of this
owl deposit.
BROTOMYS VORATUS Miller
A mandible and femur. The femur is smeared with hair and half
digested organic material. Its digital fossa and concave inner surface
NO. 15 MAMMALS FROM HISPANIOLA—MILLER a
of the greater trochanter are packed with a felt-like mass of fine hair
(pl. 2, fig. 5), in the same condition as the femur of a roof rat (pl. 2,
fig. 4) near which it was found.
ISOLOBODON sp.
Two halves of the palate from a young individual. On the left side
the teeth have fallen out and the alveoli are packed with hair (pl. 2,
fig. 6a). The femur of a young hystricoid rodent (pl. 1, fig. 6) is
probably a part of the same animal. It is shorter and decidedly more
robust than the femur that I refer to Brotomys.
(8) INDIAN DEPOSIT
Mr. Krieger informs me that he found numerous kitchenmiddens
on the valley floor near Constanza. In most of those that he examined
there were few bones, and these few were fragmentary. The midden
at Cerro de Monte, from which he took numerous mammalian remains
in fairly good condition was not more than 20 feet long, 6 feet wide
and 24 feet deep. In it he found no artifacts of Spanish origin; but
the collection includes an atlas, calcaneum, astragalus, phalanx, and
six teeth of the domestic pig (pertaining to at least two individuals )
an upper molar of a colt, and five mandibles, two femurs and a
humerus of Rattus rattus subsp. All of these remains of European
mammals, together with a few human bones, are in exactly the same
condition as the bones and teeth of the extinct rodents with which
they were associated.
NESOPHONTES PARAMICRUS Miller
A femur 18 mm. in length unquestionably pertains to this species.
Its presence in an open kitchenmidden like the one at Cerro de Monte
rather strongly suggests that the Indians may have used the small
insectivores, as well as the large Solenodon, for food. The humerus
of Nesophontes that I found in the kitchenmidden in a cave on San
Lorenzo Bay might easily have been dropped there by an owl (Smith-
sonian Misc. Coll., Vol. 82, No. 5, p. 4, December 11, 1929). Such
an origin for the femur at Cerro de Monte seems unlikely.
BROTOMYS VORATUS Miller
Mandible, 1; femur, I.
8 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
PLAGIODONTIA ADIUM F. Cuvier
Palates, 2; mandible, 1; odd cheekteeth, 4. One of the palates is
represented on plate 2 (fig. 7).
PLAGIODONTIA HYLEZUM Miller
Palate with full set of teeth, 1 (pl. 2, fig. 8); premaxillae with
incisors, 2; occipitals, 2; mandibles, 6.
ISOLOBODON LEVIR (Miller)
Palates, 4 (one with full set of teeth) ; premaxillae with incisors,
7; mandibles, 34 (18 left, 16 right). Also numerous small bones and
fragments of skulls that appear to have pertained to this animal.
CAVIA PORCELLUS (Linnzus)
(Plate 2, figs. 9, 9a)
Fragment of zygoma and palate with pm* in place, 1 ; mandibles, 4
(2 left, 2 right) ; femora, 2. One of the femora pertained to an adult,
the other to a young animal. The mandibles (two of which are figured
on pl. 2, figs. 6 and 6a) represent four individuals. They are similar
to those that Mr. Krieger unearthed at Anadel in 1929 (see Muller,
Smithsonian Misc. Coll., Vol. 82, No. 5, p. 11, December 11, 1929),
and I am unable to detect any characters by which they can be dis-
tinguished from jaws of the ordinary domestic animal. The same is
true of the right half of a palate with all four teeth in place that
Mr. Krieger collected at Boca Chica, on the coast about 20 miles east
of Santo Domingo City in 1930.
From Oviedo’s account of the “ cori” it seems evident that guinea-
pigs were well known to the Spaniards at Santo Domingo City during
the first half of the sixteenth century. Whether they brought the
animals from South America or found them already in the possession
of the natives at the time when Hispaniola was discovered is a ques-
tion that may never be answered. I was originally inclined to suppose
that the Spaniards themselves were responsible for the occurrence of
guineapigs on the island (Smithsonian Misc. Coll., Vol. 82, No. 5,
p. 14, December 11, 1929). Oviedo’s record for Santo Domingo City
and Mr. Krieger’s discovery of remains at Boca Chica and on the
shore of Samana Bay are in accord with this supposition, as both
localities were settled by the Spaniards. In fact, some remnants of a
Spanish house can still be seen at Anadel in the same field with the
kitchenmidden. But it is less easy to harmonize the belief in Spanish
NO. 15 MAMMALS FROM HISPANIOLA—MILLER 9
introduction of the guineapig with the finding of the animal’s remains
at such a remote and inaccessible locality as Constanza. This may
point, like the occurrence of a South American monkey’s teeth in a
Precolumbian grave in Cuba (Miller, Smithsonian Misc. Coll., Vol. 66,
No. 13, December 8, 1916), to early native trade between South
America and the Greater Antilles; but it must be admitted that the
presence of remains of pig and horse in the midden near Constanza
counts against such a view.
10 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
EXPLANATION OF PLATES
PLATE I
All figures natural size
Fics. 1, 1a, 1b. Proechimys semispinosus Tomes. No. 113273, U. S. Nat. Mus.
San Javier, Ecuador.
Fics. 2, 2a, 2b. Brotomys voratus Miller. No. 255696, U. S. Nat. Mus. Trujin,
Massif de La Selle, Haiti. Altitude 6,500 feet.
PLATE 2
Figs. 1-3 enlarged 4, other figures natural size
Fic. 1. Nesophontes hypomicrus Miller, male. No. 255697, U. S. Nat. Mus.
Trujin, Massif de La Selle, Haiti.
Fic. ta. Nesophontes hypomicrus Miller, female. No. 255698, U. S. Nat. Mus.
Same locality.
Fic. 2. Nesophontes paramicrus Miller, male. No. 255699, U. S. Nat. Mus.
St. Michel, Haiti. :
Fic. 2a. Nesophontes paramicrus Miller, female. No. 255700, U. S. Nat. Mus.
St. Michel, Haiti.
Fics. 3 and 3a. Nesophontes paramicrus Miller. Nos. 255301 and 255300, U. S.
Nat. Mus. Near Constanza, Dominican Republic. From owl pellets.
Fic. 4. Rattus rattus subsp. No. 255701, U. S. Nat. Mus. Near Constanza,
Dominican Republic. From owl pellet. Mass of hair in digital fossa.
Fic. 5. Brotomys voratus Miller. No. 255702, U. S. Nat. Mus. Same locality
and condition as fig. 4.
Fics. 6 and 6a. Isolobodon sp. No. 255703, U. S. Nat. Mus. Same locality and
condition as figs. 4 and 5. Hair in digital fossa of femur and in
two alveoli of palate.
Fic. 7. Plagiodontia edium F. Cuvier. No. 255704, U. S. Nat. Mus. Near
Constanza, Dominican Republic. From Indian deposit.
Fic. 8. Plagiodontia hyleum Miller. No. 255283, U. S. Nat. Mus. Same locality
BIS) anlar, of,
Fics. 9 and oa. Cavia porcellus. Nos. 2552099 and 255296, U. S. Nat. Mus. Same
locality as figs. 7 and 8.
“‘stmojo1g ‘@ So Sy > a2 a
Pu cSieeia om Sed rs v FS
bos ose os VS2 5S 3 5
= a a ra Z a Z
Unsprayed (control).. TAU 2 3 Te = (O83 5) OO
Calcium arsenate and
SUPAII fs ston tiraeioes TOO. S290. = 0.31 | 2.43
Magnesium arsenate 18 Sept. 5 32
ATIGMSU Sa Layecieeteie 6 ae Ne | oye Os Sieh (il oye
Lead arsenate and
SUT stevseatausecrel craic USM jl ALPES ONO) iets:
Unsprayed (control)..| 131 |4.10+0.38| 1.00
Magnesium arsenate
and table molasses..| 194 |6.06+0.40| 1.48
Magnesium arsenate
and corn sirup...... B57 4260 = 0.40;|) 1.20
Magnesium arsenate
and black-strap mo-
IASSES* hr eSieniacemise es TIES Ses = 0440) osso
22 Sept. 25 32
(4) Beetles can distinguish differences between water and sweet
liquids——To determine whether the above is correct, 24 series of
tests were conducted by using six sweet solutions, each of five of them
being prepared at the rate of 1 gm. or I cc. of granulated cane sugar,
grape sugar (dextrose), a high quality table molasses, corn sirup, or
black-strap molasses to 25 cc. of water; and the sixth at the rate of
20 mg. of saccharine to 25 cc. of water (table 5).
From the information given in tables 2 to 5, with additional notes,
it may be concluded that Mexican bean beetles exhibit “ likes ” and
“dislikes” when fed substances which produce the four human at-
38 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
tributes of taste. They “disliked” water containing salts, acids,
bitter materials, and saccharine, but “liked” the other sweet sub-
stances, including cane sugar, grape sugar, table molasses, corn
sirup, and black-strap molasses, and even showed preference between
them. To the writer the saccharine solution was sweetest, but dis-
tasteful; the cane sugar, less sweet, and tasteful; and the grape sugar,
least sweet, and less tasteful. The beetles showed “ dislikes’ and
TABLE 7.—Tests to determine whether bean foliage sprayed with arsenicals,
is repellent to Mexican bean beetles
x 2 vs ra s
3s vé ena Rees :
ee Bn ee adi :
aoe Hog ee S3e| 88 3
ae B25 a2 3.8 | Bus 2 EB
% ae eos ge yes Zo A =
Unsprayed (control) .| 191 |5.97 0.49] I.00
Calcium arsenate ....| 160 |5.00+0.62/ 0.84 |
: 17 Sept. 3 32
Magnesium arsenate..| 177 |5.53 0.56] 0.93
ead arsenate’ 2.55... III |3.47+0.40| 0.58 |
Unsprayed (control)..| 275 |8.59=0.67| 1.00
eat arsenate) sien 727) a Se 54 220250) mOnod! 7 Sone as
agnesium arsenate..| 122 | 3.81 0.42] 0.44
Kead@arsenatey ssc. 122 |3.81+0.48| 0.44
Unsprayed (control)..| 172 |5.38=0.46/ 1.00
Calemmyarsenates: |) £59) 4507, == 0.35) ono2
Magnesium arsenate..| I04 | 3.25 +0.25/ 0.60 24 Sent 27 32
Wead-arsenate: ... <1. LOL ||) 3et6) = 0422) 0559
SuMMARY of above:
Leaves, unsprayed
(controls) 1... 638 | 6.64 + 0.33| 1.00
Calcium arsenate ....| 406 |5.170.29| 0.78 a enie pce above
Magnesium arsenate..| 403 | 4.20+0.26) 0.63 | | O68
ead@arsenate mace cer Be || Jodi} 22 O22 |) Ook
“likes”? in somewhat the same order. To the writer the picric-acid
solution was most bitter, the quinine less bitter, and the strychnine
least bitter. The insects “ disliked ” these solutions in about the same
order. To the writer the solutions containing acetic acid and hy-
drochloric acid had practically the same degree of sourness, while the
diluted lemon juice was sourer. The beetles also showed only slight
differences between them. In regard to the salty solutions, the writer
disliked only the magnesium sulphate solution, but the beetles pre-
ferred it to the other two.
No. 18 SENSE ORGANS OF COLEOPTERA—McINDOO 39
(5) Bean foliage sprayed with sweetened arsenicals is more at-
tractive than unspraycd foliage—To ascertain whether the above is
correct, 16 series of tests were conducted. The arsenicals were pre-
pared as stated on the following page, then 1 gm. of granulated cane
TABLE 8.—Tests to determine whether bean foliage sprayed with sweetened
magnesium arsenate is more attractive to Mexican bean beetles than
foliage sprayed with non-sweetened magnesium arsenate
ico] o > n
o Le a Zs % 2
a 0° = oO a4
i a3) (Fire > 2 Sie ae
a Ev Ze s ov ro)
oHM tr owv Sao oT
8 eee ae Z52| 28 2
Ae S38 Sa aioe cs ES be} E
os o6o8 a] Caer Se) a =
4 aS = foo Zi fa) Z
Magnesium arsenate
(control) eee ete oe Wh | ApzASS OAH) 100
Magnesium arsenate
and table molasses
(C2BtOF5O) ccc at: 192 |6.00+0.61| 2.70
Magnesium arsenate 23 Sept. 26 32
and corn sirup (2 to
5/0) aie a eee atte PEA | 3. 50'2=.0.30) 1.60
Magnesium aresnate
and black-strap mo-
IeSSESS (2 \fO450):..52 | 122) 3.88 10.43)| 1:72
Magnesium arsenate |
| |
(control) snes see $2) | 2.5@== 0.28) 1.00
Magnesium arsenate
and black-strap mo-
lases, (2).t0°-50) ¢o42 | 132/140 02 0.36) 126%
Magnesium arsenate |
and black-strap mo- |
IESE (Mie FO) sooo |) wits) Weiogk{o)==(0),4%))| Waele)
Magnesium arsenate
and black-strap mo-
lasses (4 to 50)....| 134 | 4.19 =0.32| 1.63
25 | Sept. 3o 32
sugar or I cc. of a high quality table molasses, corn sirup, and black-
strap molasses was added to 25 cc. of the spray mixture (table 6).
According to the results given in table 6, it is again shown that
these beetles like their food sweetened.
(6) Bean foliage sprayed with arsenicals is repellent—To deter-
mine whether bean foliage sprayed with arsenicals is eaten as readily
as are unsprayed bean leaves, 24 series of tests were conducted. The
4
40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
leaves were sprayed with calcium arsenate, magnesium arsenate, and
lead arsenate at the rate of I pound of powder to 50 gallons of water.
The calcium-arsenate mixture also contained lime at the rate of 1.5
pounds to 50 gallons of water (table 7).
The results given in table 7 clearly show that arsenicals are repel-
lent, but not sufficiently so to prevent the foliage from being eaten.
Lead arsenate was most repellent, magnesium arsenate was less so,
and calcium arsenate was least repellent. The word “ deterrent” is
probably the better expression in this case.
TABLE 9.—Tests to determine whether water extract and steam distillate of
bean leaves are attractive to Mexican bean beetles
g 3 Ee ak: 2
> BES 2 age es g
ww og 3 5 2 are is nw coin is
ae a2 5 = E is 5 Ls =
pee Bee Sey geet ee |) eae 3
Bae $85 he Sa AEIS 2 gE
ae8 Sie ie Peers eo é y
Leaves, unsprayed
(controler 190 | 5.04 0.39] 1.00
Water on cotton.... 280 ORO 71 ==004)) NOR mS
Water extract on cot-
21 Sept. 18 BD
COMM Garerrciccoteisieere seca 167 | 5.220.40/] 0.88
Water extract and
cane sugar (1 to
25) TON mGOLtOne ser | 677: 20. 16e= 0.82) 3.56) 4)
Water on cotton | | |
|
(control) mane Ars lO. 48420741 |) bL00 21 Sept. 19 64
Distillate on cotton.. | 436 ONO TH==10843) MOS [
(7) Bean foliage sprayed with sweetened magnesium arsenate is
more attractive than foliage sprayed with non-sweetened magnesium
arsenate —To determine whether the above is true 16 series of tests
were conducted by using magnesium arsenate (I pound to 50 gallons
water) with molasses added at the rate of 2, 1, and 4 gallons to 50
gallons of the spray mixture (table 8).
The results given in table 8 once more show that sweetened food is
preferred to non-sweetened food.
On September 30 four small bean plants, each bearing six leaves,
were sprayed. Two of these were sprayed with magnesium arsenate
alone and the other two with a mixture of magnesium arsenate and
black-strap molasses (1 to 50). One plant sprayed with the non-
sweetened mixture and one with the sweetened mixture were put to-
No. 18 SENSE ORGANS OF COLEOPTERA—McINDOO 4!I
gether in one end of a cage, and the other two sprayed plants were
arranged likewise at the other end of the cage. Soon after placing
100 beetles in the cage the insects climbed upon the sprayed foliage,
paying apparently no more “attention ” to the sweetened leaves than
to the non-sweetened ones, but after a few hours and thereafter until
October 3, when the experiment was ended, the sweetened leaves
bore the more beetles and were the more eaten. The final result
TABLE 10.—Tests to determine whether chemotaxis or phototaxis is more
infortant in the finding of food by Mexican bean beetles
n ! Os ze a
ae BES 2 gee aalii Ea
bane 22s 28 25 | 8 82
aE ES See plese, thay g*
23-5 Gas a2 ial Se 2 EY
sks Coes ga Bets) 23 a 25
A
Bean leaves, not
sprayed (control).| 68 | 2.12+0.31| 1.00
Apple leaves not 32 tests.
SVG. Beside sia o's A | 10012 == )0.02' |, 0106 ba Sepeas Beetles
Green water on cot- in direct
OME, ce saheeetarats S04 196 | 6,120.76) 2.88 sunshine.
Green sugar water (1
to 25) on cotton..| 635 | 19.84+0.99| 9.34
Bean leaves, not
sprayed (control).| 116 | 3.62+0.24| 1.00
Mulberry leaves not |
SPLAVER! f.5 eters. LO) || O250 == 0.00) On56
Mulberry leaves,
; 22 Sept. 24 32
sprayed with sugar
water (I to 25)... 73 2:28 '0.30') 0:63
Bean leaves, sprayed
with sugar water
LO 25))icc skagen 221 6.91 +0.42| 1.90
a Be I
showed that the sweetened leaves bore 69.7 per cent of all the beetles
counted on the four sprayed plants.
(8) Water extract and steam distillate of bean leaves tested —To
test the diluted juice of bean foliage, a water extract was prepared by
adding 50 cc. of water to 10 gm. of leaves, cut into small pieces. After
macerating the pieces and decanting the liquid through cheesecloth,
50 cc. of a greenish liquid was secured. To test the steam distillate
of bean foliage, 100 cc. of water was added to 30 gm. of leaves, cut
into small pieces, and then 50 cc. of a clear and odorous distillate was
collected (table g).
42 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
The results given in table 9 show that when water extract of bean
leaves was compared to unsprayed leaves it was about equally attrac-
tive, while sweetened water extract was about 3.56 times as attractive.
Steam distillate from bean leaves was not attractive, but gave practi-
cally the same result as did water, indicating that its faint odor had
no attractive influence.
(9) Chemotaxis more important than phototaxis in the finding of
food. Sixteen series of tests were conducted to ascertain whether
phototaxis or chemotaxis, or possibly thigmotaxis, is the more im-
portant in the finding of food. Squares of bean leaves, apple leaves,
TABLE 11.—Tests to determine whether repellents would protect beans from the
Mexican bean beetle
i a ge 3 E
BEE gE B Be | se FE
"BE zee Es ee, | se
ao ees Ss g ore 2 3} é S
Unsprayed (control)..| 226 | 7.06 +0.49| 1.00 |
iargrand winner rece 216 |6.75 £0.53| 0.96 |
Nicotine sulphate and | | +19 Sept. 9 32
litteeene ree ene | 106 16.12 0.47| 0.87 |
Derris product ...... 120 | 3.78 SOA 7A Oss3 |
Unsprayed (control)..| 256 | 8.00 0.64} 1.00
Cresol iets ane Ra eee eRe | 251 |7.84+0.49 | 0.08 “6 Sephira fe
Nicotine sulphate ....| 219 6.840.560) 0.85 |
Betaynaphtholiss5-er 142 |4.44%0.47| 0.55 |
and mulberry leaves, all of the same shape and size (1 in. square),
but some bearing a film of cane sugar, represented practically the
same color, form, and texture, but differed chemotactically. Cotton
wet with green water and with green sugar water (a green dye being
used) also somewhat resembled the leaves in color but differed in
other respects (table 10).
The results given in table 10 show that sweetened food is preferred
to non-sweetened food and that chemotaxis is more important than
phototaxis in bringing about the results obtained.
(10) Repellents would probably not protect beans ——To ascertain
whether certain substances, usually known as repellents, would keep
the beetles away from the treated leaves in the four small cages,
bean foliage was sprayed with the following: Tar and lime, combined
as a dust; nicotine sulphate and lime, combined as a dust, a com-
No. 18 SENSE ORGANS OF COLEOPTERA—McINDOO 43
mercial Derris product, consisting mostly of pyridine; cresol, U. S.P.
(1 cc. shaken in 400 cc. water); 40 per cent nicotine sulphate in
water (I to 400) ; and beta naphthol (1 gm. powder in 400 cc. water ;
powder not all in suspension) (table 11).
The results given in table 11 show that the repellents more or less
protected the leaves, but not sufficiently so to prevent them from
being eaten. The Derris product and beta naphthol were the only ones
which might be considered promising, yet their protective value was
about equal to that of lead arsenate, as already shown in table 7.
Ill. THERMOTAXIS
After having searched the literature for references on other
tropisms not yet discussed, the writer found a few more concerning
Coleoptera, but only two of these references pertain to the orientation
of beetles to temperature. Much experimental work on various tem-
peratures, particularly as control measures, has been done, but very
little of it can be discussed from the tropic point of view.
I. REVIEW OF LITERATURE
Fulton (17) devised a crude temperature gradient with which he de-
termined that the choice of temperature of adult click beetles is much
below the usual maximum temperature in open fields during summer.
He also says that negative phototaxis causes the beetles to seek dark
hiding places during the day. Wireworms, or the larvae of these
beetles, were found more resistant to heat than were the adults, but
they did not voluntarily seek higher temperatures. Seasonal move-
ments of the larvae may be closely correlated with changes in soil
temperature.
Grossman (21) tried three methods to determine the orientation of
cotton boll weevils to heat stimuli, but decided that only the results
obtained by using a new apparatus were reliable. This apparatus was
constructed by using 16 copper bars 4 inch wide and 5 inch thick, with
gz inch insulating space between each two bars. Using only two
variables, temperature and light, 126 boll weevils were tested 1,993
times. The average temperatures to which they reacted definitely were
130° F. at the hot end of the apparatus and 26° F. at the cold end.
B. SENsory RECEPTORS
Since tropic responses are brought about largely by external stimuli
affecting either‘the special sense organs or others not definitely known
and localized, called the general sense organs, it is only natural to
discuss the tropic responses and sensory receptors in the same paper.
44 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
i; PHOTORECEPTORS
According to the phototactic responses of the Mexican bean beetle
and its larva, already discused, the compound eyes and ocelli in this
species are normally developed and seem to function adequately, so
far as beetles are concerned. It is recalled that the adults are always
photopositive and that the larvae up to the time of pupation are photo-
positive, too, but when ready to pupate they become photonegative.
Whether the negative reaction is caused by a change in the structure
of the ocelli is not known.
Since the morphology of insects eyes has often been discussed and
as the writer (46, 47) has recently cited reviews on this subject, no
further discussion is needed here. Also, the other sense organs and
senses of beetles will be discussed only briefly.
ITI. CHEMORECEPTORS
Chemoreceptors include both olfactory and gustatory organs, but
we are not absolutely sure that insects have true chemoreceptors, al-
though their organs certainly belong to the same category.
I. SO-CALLED OLFACTORY ORGANS
(A) ANTENNAL ORGANS
The organs on the antennae of the Mexican bean beetle are com-
paratively few; that is, these antennae are nearly bare in comparison
to most antennae (fig. 6). Only four types of sense organs were
found on them. They are as follows: (1) Two groups of tiny hairs
(St) ; and (2) three or four pores (P), called olfactory by the writer,
lie on the base of the first antennal segment; (3) the Johnston organ
(J) lies at the distal end of the second segment; and (4) five areas
of thin-walled hairs (OHr) were found on the distal ends of the ninth,
tenth, and eleventh segments. All of these structures are sense organs,
because sense cells were found connected with them, while the larger
hairs (Hr), usually called sense bristles, were found to be non-
innervated.
Of these four types of sense organs only the olfactory pores and
thin-walled hairs may be regarded as so-called olfactory organs. The
thin-walled hairs are numerous and most of them lie on the dorsal
surface of the antennae (fig. 6, OHr). Under a high-power lens they
appear long and slender, have thin, almost transparent walls (C, OHr),
and are connected with sense cells.
From the preceding it is evident that pore plates, found only on
the antennae of aphids, bees, wasps, and on some beetles, are totally
absent on the antennae of the Mexican bean beetle. The pore plates,
——————— —————— ee Oe
No. 18 SENSE ORGANS OF COLEOPTERA—McINDOO 45
when present, are considered the olfactory organs by most writers.
Figure 7 illustrates the antennal organs of a water beetle, copied from
Hochreuther (25). The pore plates (PP), hollow pit pegs (HPPq),
and massive pit pegs (MPPg) might be called olfactory organs, but
Hochreuther regarded only the hollow pit pegs as probably olfactory
in function. If they really act as olfactory organs, then the mouth
parts, thorax, legs, and sexual organs must aid in receiving odor
stimuli, because Hochreuther found them also on these parts of the
anatomy.
Fic. 6.—Drawings of antennae of adult Mexican bean beetle, showing organs
on them. A, ventral surface, and B, dorsal surface, showing location of follow-
ing: Hr, noninnervated hairs; J, Johnston organs; P, pores called olfactory
by the writer; OHr, so-called olfactory hairs; and St, tactile hairs. C, a semi-
diagrammatic drawing from a section through tenth segment, showing structure
of so-called olfactory hairs (OHr) and their sense cells (SC). A and B, X 53;
zhatal (Ga Se Covey
(B) OLFACTORY PORES
The writer (46, p. 1105) in 1926 cited references pertaining to
these organs in beetles and in 1929 he (47, p. 27) stated why they
were called “ olfactory pores.” In 1915 (41) he made a comparative
study of them in 50 species of beetles belonging to 47 genera and rep-
resenting 34 families. In that study only the legs, elytra, and wings
were examined for these pores. A group of pores (fig. 8, A and B, r)
was always found on the peduncle of each elytron. The number of
pores in it ranged from 12 to 310, and the more pores in the group
the smaller they were and the closer they were together. Of the 47
46 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
B
Fic, 7.—Antennal organs of a water beetle, Dytiscus marginalis, copied from
Hochreuther (25). A, longitudinal section through a hollow pit peg (HPPg);
B, longitudinal section through a small massive pit peg (MPPg) and 2 pore
plates (PP). This drawing is a combination of Figs. 32 and 58 from Hochreuther,
slightly modified. C, a small tactile hair from first segment, total preparation;
and D, portion of Fig. 12 from Hochreuther, showing 4 small sense bristles
from second segment. NF, nerve fiber; SC, sense cell; SCG, sense cell group;
and SF, sense fiber.
Fic. 8.—Portion of left elytron (A and B) and left wing (C) of Mexican
bean beetle, showing position of olfactory pores as indicated by numbers Z to 4
and letters a to d on dorsal surfaces. A shows relative sizes of peduncle of
elytron and group rt when compared with size of basal margin (BM) of elytron;
A and C, X 12; and B, X 67. The lower side of A and B is the outer margin
of the elytron. C, costa; MD, muscle disk; Me, media; R, radius; RP, radial
plate; Sc, subcosta; ScH, subcostal head.
No. 18 SENSE ORGANS OF COLEOPTERA—McINDOO 47
winged species examined, 11 had only one group of pores on each
wing, 21 had two groups on each wing, 12 had three groups on each
wing (C, 2, 3, and 4), and 3 had four groups on each wing. The
Fic. 9.—Position of sense organs on legs of adult beetles; dots, marked 5
and 6, and e and f, being olfactory pores; and St, tactile hairs. A to F, legs of
Mexican bean beetle, X 12; and G, distal end of tibia from front leg of Epicauta
marginata, showing 5 olfactory pores on tibial spine. The drawing of each leg
in which the tarsus is shown represents the outer surface and the portion of leg
without tarsus represents the inner surface of the same leg. A, right front
leg; B, left front leg; C, right middle leg; D, left middle leg; E, right hind
leg; and F, left hind leg.
number of pores on a pair of wings ranged from 130 to 982. The
number of pores counted on all six legs of an individual ranged from
49 to 341. There were usually two groups of pores on each trochanter
(fig. 9, 5 and 6). Sometimes a pore was found at the proximal end of
48 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
the femur. A few pores were always found at the proximal end of
each tibia (e and f), and sometimes pores were found in the tibial
spines (G) and on the tarsi.
In regard to water beetles, the better the legs are adapted for loco-
motion in water, the fewer pores they have. The smallest winged
species examined had 273 pores, which is the smallest number counted
of all the species, and the largest species had 1,268 pores which is the
largest number of all the species examined. The wingless species had
more pores on the legs than usual. As a rule, the smaller the species,
the fewer its pores and the larger they are, comparatively speaking.
As a rule, no generic and specific differences were found, except vari-
ations in number of pores, the amount of variation depending on the
sizes of the individuals compared. There were no individual and
sexual differences other than slight variations in number of pores.
The pore apertures or pits are round, oblong, slitlike, or club-
shaped. On the elytra and wings (fig. 10, A and B, Ap) they are al-
ways round or oblong. On the legs (C) they have all four of the
enumerated shapes.
The spindle-shaped sense cells (fig. 10, C, SC) of most beetles lie
in the lumens of the appendages outside the pore cavities. A small
chitinous cone (Co) is always present. It is formed by the hypoder-
mal cell at the mouth of the pore after the insect has emerged from
the last pupal instar, and at the same time when the chitinous integu-
ment is being considerably thickened. The sense cells are fully devel-
oped when the insect emerges into the imago. The sense fiber pierces
the cone, and comes in direct contact with the outside air. This state-
ment is denied by other writers. In the legs of the lady-beetle Epi-
lachna borealis the pore apertures lie in the center of domes (fig.
10, C) above the general surface of the legs.
A large nerve and a large trachea run into each elytron (fig. Io,
A, N and Tr) and wing. In the peduncle of the elytron they run
through the radial plate just beneath the group of olfactory pores.
Branches from the nerve are given off which connect with the sense
cells. The large nerve and trachea passing into the wing soon divide
so that a smaller nerve and a smaller trachea (B, N and Tr) run
through each main vein. The largest trachea passes through the sub-
costa, and the largest nerves pass through the veins bearing the ol-
factory pores. These nerves give off branches which connect with
the sense cells. The sense cells (C, SC), wherever found, are always
surrounded by blood (B/).
In a study of the sense organs of the cotton boll weevil, the writer
(46) found the olfactory pores common to both the adult and larva;
ee
No. 18 | SENSE ORGANS OF COLEOPTERA—McINDOO 49
Fic. 10.—Diagrams showing portions of elytron, wing, and leg of adult beetles,
to illustrate internal and external anatomy of these appendages and of olfactory
pores and hypodermal glands. A, oblique transverse-longitudinal view of portion
of peduncle of Epilachna borealis. The transverse portion passes through group
1 of the olfactory pores and radial plate (AP) in the direction of the line marked
X in figure 8 B. B, transverse-longitudinal view of portion of wing of Orthosoma,
passing through pores on radius () and media (Me). C, transverse-longitu-
dinal view of proximal end of trochanter belonging to right hind leg of E.
borealis, passing through group 6 of olfactory pores (4 pores on right) and
group 5 (3 pores at left). Ap, pore aperture, B/, Blood; Co, chitinous cone;
GC, hypodermal gland cell; Hr, noninnervated hair; N, nerve; Po, pore of
hypodermal gland; SC, sense cell; SV’, sense fiber; and 77, trachea.
ie) SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
but the other so-called olfactory organs, which are nothing more
than ordinary innervated hairs, are common only to the antennae of
the adult, although similar innervated hairs are also found on other
parts of both adult and larva. In the adult the olfactory pores were
found on the head capsules, legs, elytra, wings, and mouth parts, and
at the base of the antennae; in the larva, on the head capsule, base
of antennae, mouth parts, clypeus, and second thoracic segment. The
individual and sexual variations found in the pores were small, al-
though the females have 13.7 per cent more pores than have the
males.
Fic. 11.—External view of single olfactory organs and noninnervated hairs
on larva of Cotinis nitida, X 320. A, 4 organs from trochanter, showing pore
border (PB), pore aperture (Ap), and pore wall (PIV). B, 2 organs and a hair
from hypopleural region; C, 2 organs from maxilla; D, 2 organs and a hair
from labrum; FE, 2 organs from labium; F, 2 organs and a hair from epicranium ;
and G, a hair from first antennal segment.
On the larva of the green June beetle (Continis nitida L.) the writer
(44) found the olfactory pores unusually numerous and consisting of
two types. The single olfactory organs are isolated pores, not arranged
in groups. They were found on the antennae, all mouth parts, head
capsule, thorax, and legs, and average 1,359 pores per individual
larva. This number is slightly more than the total number of pores
found on the elytra, wings, and legs of an adult of the same species.
Their external anatomy is unusual in that the pore border (fig. 11,
PB) is radially striated, while the border around the hairs never
shows striae. The compound olfactory organs (fig. 12) are variously
shaped plates, each of which bears many apertures. They were found
only on the distal halves of the last antennal segments. Figure 13
illustrates the internal anatomy of the single and compound organs.
No. 18 SENSE ORGANS OF COLEOPTERA—McINDOO 5!
Fic. 12.—External view of compound and single olfactory organs and olfactory
pegs on distal half of last antennal segment of larva of Cotinis nitida. A, 9 com-
pound organs and 3 single ones on ventral side of antenna, viewed from a flat
surface, X 100; 2 of the compound organs at extreme tip are not shown. B, 6
compound organs, 4 single ones, and 1 group of olfactory pegs (Pg) on dorsal
side of antenna, viewed from a flat surface, X 100. C, external view of a com-
pound and a single organ, 320; the small circles represent pore apertures.
Fic. 13.—Internal anatomy of sense organs on antenna of larva of Cotinis
nitida. A, longitudinal section through tip of antenna, showing innervation of
compound (P/) and single olfactory organs (P), and tactile hair (St); two-
thirds diagrammatic, 100. (At this magnification the pore apertures are never
discernible). B, cross section through single olfactory organ from antenna,
< 500. Ap, pore aperture; Hyp, hypodermis; NB, nerve branch; NF, nerve
fiber; SC, sense cell; SF, sense fiber; and Tr, trachea.
52 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
The olfactory pores on several males and females of the Mexican
bean beetle were examined, but they were actually counted on only
one female, and these are illustrated in figures 6, 8, 9, 14, and 15. In
this study only one totally new fact was learned. In all the previous
studies on beetles, no olfactory pores were seen on the ventral side of
the peduncles of the elytra, but in this position on the bean beetle 7
pores were seen on one elytron and 6 pores on the other. The groups
Fic. 14.—External view of chemoreceptors of adult Mexican bean beetle. A,
noninnervated hairs and group 5 of olfactory pores on trochanter of front leg,
x 500. B, ventral surface of distal end of maxillary palpus, showing numerous
noninnervated hairs (Hr) with their gland pores, isolated olfactory pores (P),
group 7 of olfactory pores (7), and a plate bearing numerous so-called taste
hairs (THr), X 100. C, group 7 of olfactory pores, markings on chitin, hairs,
and gland pores, < 500.
of pores are numbered, as usual, from I to 7, and small letters are
used to indicate the position of some of the isolated pores. Group I
on the elytra (fig. 8, A) contains 58 pores on the left peduncle and
65 on the right one. Groups 2, 3, and 4 on the dorsal surface of the
wings (C) have as follows: Group 2, 58 and 64 pores; group 3, 43
and 38 pores; and group 4, 70 and 64 pores. Isolated pores on the
wings are as follows: At a, 2 on the ventral side; at b, 8 on the dorsal
side; at c, § on the dorsal side; and at d, 12 on the dorsal side and 8
on the ventral side. The number of pores in groups 5 and 6 and at e
No. 18 SENSE ORGANS OF COLEOPTERA—McINDOO 53
and f on the legs (fig. 9) can be counted by inspection. Group 7, con-
sisting of 8 slit-shaped pores, lies on the ventral surface of the ter-
minal segment of the maxillary palpus. All the remaining pores
counted are isolated ones found on the antennae and mouth parts.
The total number of pores on all appendages of the same bean beetle
are as follows: Wings 397, elytra 136, legs 95, maxillae 32, labium
14, antennae 8, mandibles 6, and labrum 4, making 692 in all. The
Fic. 15.—Internal structure of sense organs on adult Mexican bean beetle,
500. A, an olfactory pore, gland cell (GC), and sense hair from dorsal surface
of labrum. B to G, olfactory pores; B, from mandible, C, from maxilla; D,
from elytron; E, from trochanter; F, from tibia; and G, from wing. H, sense
hair from trochanter. I, drawing, two-thirds diagrammatic, from longitudinal
sections of distal end of labial palpus, showing innervation of so-called taste
hairs (THr), tactile hairs (St), and olfactory pore (P). J, semidiagrammatic
drawing from, 3 cross sections through base of first antennal segment, showing
tactile hairs (St), gland pore (Po), olfactory pore (P), trachea (Tr), and
nerve (NV).
fact that this number is small for an adult insect might be correlated
with the fact that the bean beetle is “ stupid ’’ when the olfactory re-
sponses are considered.
The olfactory pores on several individuals of all four instars of
bean-beetle larvae were examined. Since no differences in number
and position were observed, the pores were carefully studied on only
individuals of the fourth instar. They are illustrated in figure 16.
The total number of pores on all appendages and the head are as
follows: Legs 30, maxillae 12, head capsule 6, antennae 4, labrum 4,
54 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 82
labium 4, and mandibles 2, making 62 pores in all. The fact that this
number is extremely low for any insect may help to explain why these
larvae did not respond readily to odor stimuli.
is e
LG; D
Fic. 16.—Position of olfactory pores (dots), 12 to 15 so-called taste hairs
(Thr) at tip of each maxillary palpus, 8 tactile or so-called olfactory hairs
(St) at tip of each antenna, and ocelli (Oc) on larva of bean beetle, * 32.
A and B, inner and outer sides respectively of right front leg. The number of
pores on the legs is nearly constant, and they shift only slightly in position.
C and D, dorsal and ventral surfaces respectively of the head and head appen-
dages. On the base of each antenna one pore is on the dorsal side and one on
the ventral side. On each terminal maxillary segment there is a slit-shaped pore.
2. SO-CALLED TASTE ORGANS
Several writers, particularly Nagel (58), have described certain
tiny peglike hairs on the mouth parts of insects as taste organs, but
no one has ever demonstrated that they perform such a function.
Hochreuther (25) found many “ Tast- und Geschmackszapfchen ”’
on the maxillary and labial palpi of the water beetle Dytiscus margi-
No. 18 SENSE ORGANS OF COLEOPTERA—McINDOO 55
nalis. The earlier papers concerning the chemoreceptors of Coleoptera
are reviewed by Deegener (see Schroder (72, pp. 150-151). Since
Minnich’s papers on the taste organs of butterflies and flies have
recently been reviewed by the writer (47, pp. 36-39), they will not be
discussed here. The reader should know, however, that according to
the experiments conducted by Minnich certain butterflies bear so-
called taste organs in their tarsi, and certain hairs on the proboscis of
the blowfly serve as gustatory organs. The most recent paper by
Minnich (55) discusses the chemical sensitivity of the legs of a
blowfly.
The writer (46) described and Tice many tiny peglike hairs
found on the cotton boll weevil, but did not attribute a gustatory
THr
a
ih, ith bi
Yy/ Wy
i /
Fic. 17.—Internal anatomy through tip of maxillary palpus of adult Mexican
bean beetle, showing following: Hyp, thick hypodermis; PH, pseudohairs; PI,
soft plate; Po, gland pore connected with gland cell, which lies some distance
from pore; SC, sense cells; St, tactile hairs; and THr, so-called taste hairs. A,
drawing, two-thirds diagrammatic, & 300; and B, semidiagrammatic, 500.
function to any of them. According to position, and possibly to struc-
ture, the ones on the tips of the labial and maxillary palpi are best
suited to be taste organs. The same type of hairs was also found at
the same place on both adult and larva of the Mexican bean beetle
(figs. 14-17, THr). The ones at the tip of the maxillary palpus of
the adult (fig. 14, B) are the most numerous and most conspicuous
of any yet observed by the writer, and consequently they would appear
to have some function otber than that of touch. These tiny, thin-
walled, and transparent hairs arise from a slightly convex plate, which
is soft, flexible, and transparent. The number of hairs on the organ
illustrated in figure 17 is about 447. An oblique cross section through
the fourth or terminal maxillary segment is represented by figure 17,
5
56 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
A. The transparent plate (P/) is bordered by tiny pseudo-hairs (PH)
and the hypodermis (B, Hyp) just beneath the plate is very thick.
Each hair is connected with a sense cell (SC) and these cells almost
fill the lumen of the segment. The sense cells are very long and
slender and have conspicuous nuclei.
Now, if aqueous solutions can pass quickly through the walls of these
sense hairs in order to stimulate the nerves inside, these structures
would be excellent taste organs. Or, if air can pass quickly to the
nerves, they would then be olfactory organs. The fact that the bean
beetle possesses two of these highly developed sense organs helps
to explain how these insects were able to distinguish so readily between
the various aqueous solutions and insecticides fed to them.
Ill. AUDIRECEPTORS
Since the writer (46, p. 1119; 47, p. 39) has already reviewed the
literature on the sense of hearing in insects, no further review is
necessary here, other than to cite the recent book by Eggers(14).
I. JOHNSTON ORGANS
In caustic-potash preparations of the antennae of the adult bean
beetle the location of the Johnston organs may be determined by
focusing downward with the microscope when looking at the distal
end of the second antennal segment. A serrated structure (fig. 6, A
and B, J) will be observed to encircle the segment. The distal ends
of the sense cells are attached to this structure. In longitudinal sec-
tions the Johnston organs appear about as shown in figure 18, A. At
the base of the second segment the nerve divides into two branches,
which run directly to the sense cells (SC). Formerly the Johnston
organs were assumed to be auditory in function, but more recently
they have been called muscular receptors or statical-dynamic organs
to register the movements of the antennae.
2. CHORDOTONAL ORGANS
Chordotonal organs very often accompany the Johnston organs,
as illustrated by the writer in the cotton boll weevil; but none was
found in the Mexican bean beetle. Many sections through the larvae
were also made and studied, but no chordotonal or Johnston organs
were found.
Since the writer has never reviewed the literature on the so-called
auditory organs in larvae, the reader is referred to the paper by Hess
(24) who gives a brief history of the chordotonal organs and de-
No. 18 SENSE ORGANS OF COLEOPTERA—McINDOO 57
scribes them in cerambycid larvae. Hess determined that the pleural
discs in these larvae are the points of attachment of abdominal
chordotonal organs. Two of Hess’s drawings (fig. 18, B and C) were
copied to illustrate the internal structure of these organs in coleopter-
ous larvae.
Fic. 18.—Internal anatomy of so-called auditory organs of beetles. A, draw-
ing, two-thirds diagrammatic, from longitudinal sections through second an-
tennal segment of an adult Mexican bean beetle, showing Johnston organs con-
sisting of groups of sense cells (SC), X 500. Band C (after Hess), longitudinal-
vertical section of the pleural zone and chordotonal ligament from a larva of
Ergates spiculatus, showing following: AF, axis fiber; C, chitinous cap; CC,
cap cell; EC, enveloping cell; EK, end knob; F, fibrils of cap cell; N, chordo-
tonal nerve; Sc, scolopale; SC, sense cell; TL, terminal ligament; and lV’, vacuole.
IV. THIGMORECEPTORS
I. TACTILE HAIRS
Hochreuther (25) made a thorough study of the sense hairs on
a water beetle (Dytiscus marginalis). On the basis of external
structure, he separated them into five divisions. Vom Rath (63, 64)
found sense cells connected with all the small hairs on the maxillary
palpi of Coccinella septempunctata, Melolontha vulgaris, and Tenebrio
58 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
molitor, and also with all the small hairs on the labial palpi of the
last species. The present writer (46) found tactile hairs on the cotton
boll weevil as follows: Sense hairs (Sensilla trichodea) on the head
capsule, antennae, mouth parts, thorax, legs, wings and abdomen;
sense bristles (S. chaetica) on nearly the same parts; and sense pegs
(S. basiconica) on the head capsule, mouth parts, and genitalia.
In regard to tactile hairs on the Mexican bean beetles, all parts of
the integument were not searched for them, but practically all the
innervated hairs already discussed might be considered as touch
hairs; however, the sense hairs (Sensilla trichodea) are considered
to have no function other than that of touch. On the base of each
antenna lie two groups of these hairs (figs. 6, B, and 15, J, St) and
each trochanter bears one or two groups (fig. 9, C, and 15, H, St).
They were also found on the maxillary and labial palpi (fig. 15, I, St)
of the adult and on the head (fig. 16, C, St) of the larva.
’
C. Scent-PRoDUCING ORGANS AND REFLEX “ BLEEDING’
The study of scent-producing organs follows as a corollary to that
of tropisms and sensory receptors, and reflex “ bleeding” is closely
related to them. Since the sense of smell is such an important means
of communication among insects, it is probably true that all insects
have structures for producing odors. In fact these structures have
already been described for most insect orders, and particularly for
Coleoptera.
The writer (43) in 1917 reviewed the literature on this subject.
A brief summary of that review concerning beetles follows: The
simplest type of a scent-producing organ in beetles is composed of
unicellular glands distributed over the entire body surface. In some
beetles these unicellular glands are grouped and thus form glands
varying considerably in complexity. In Malachius two pairs of car-
uncles serve as the scent-producing organs; unicellular glands lie in
the walls of these structures. In Dytiscus, Gyrinus, and Acilius two
different kinds of liquids issue from unicellular glands situated in the
articular membranes between the thoracic segments. The liquid
emitted at the femoro-tibial articulation during reflex “ bleeding ” of
certain beetles seems to be secreted by two types of unicellular glands
at this location. The highest type of scent-producing organ among
insects is the anal glands of beetles. These have been found in several
families.
In regard to the Mexican bean beetle, no careful search was made
for the purpose of finding scent-producing organs other than the
No. 18 SENSE ORGANS OF COLEOPTERA—McINDOO 59
unicellular glands distributed over the entire body surface. In fact
this type of scent organ is the only one in lady-beetles known to the
writer. The bean beetle, like other coccinellids, is well supplied with
these glands. All parts of the body surface are covered with com-
paratively large hairs. Near the base of each hair there is usually one
and sometimes two gland pores (figs. 6, 10, 14, 15, 17-19, Po). The
large gland cells (figs. 10, C, and 15, A, GC) are variously con-
structed, but are always connected with reservoirs lying in the integu-
ment. In some of the smaller appendages, for example the maxillary
palpi, where the available space is limited, the gland cells lie some
distance from their pores and often nearly fill the lumen of the
appendage.
The writer (42) in 1916 reviewed the literature on reflex “ bleed-
ing’ in beetles and added further information by using the squash
lady-beetle, Epilachna borealis. When disturbed certain coccinellid
and meloid beetles fold the antennae and legs against the body, eject
small drops of liquid from the femoro-tibial articulations, and feign
death. There has been a controversy as to how the liquid is expelled
so quickly and as to whether the liquid is blood or is a glandular se-
cretion. The writer has now shown that in regard to the squash lady-
beetle and the Mexican bean beetle (/. corrupta) the phenomenon is
a true reflex, but that instead of the liquid being blood, it is a secretion
from two types of hypodermal glands and that it passes to the exterior
through innumerable tubes opening near and in the articular mem-
brane. The gland pores of the first type, with reservoirs, lie in groups
on the tarsi and around the femoro-tibial articulations. Two groups
of these are located at the extreme proximal end of the tibia and two
at the distal end of the femur around the articular membrane (fig. 19,
A, Po). The gland pores of the second type, without reservoirs, lie
in the articular membrane, marked a in figure 19. The discharge of
the amber-colored secretion is accomplished by putting the gland cells
under a high blood pressure. This is made possible by a muscular con-
traction in the femur whereby the blood is forced into a specially de-
vised chamber containing the gland cells which belong to the pores in
and near the femoro-tibial articulation. The glandular secretion is
bitter and has an offensive odor. Its chief purpose is that of pro-
tection, but it probably also aids the beetles in recognizing the different
individuals and sexes of the same species.
Hollande (26) in 1911 wrote a large paper in which he reviewed
the literature on the phenomenon of discharging “ blood ” in insects
and on the toxicity of this substance. He also added new information
on these subjects. He reports that self-bleeding has been found in
60 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
Orthoptera, Hemiptera, Coleoptera, Diptera, Lepidoptera, and Hy-
menoptera. Under his general conclusions he states that some authors
believe that the discharged liquid is blood, while others think it is a
glandular secretion. The manner in which the liquid is discharged
is little known, except in a few cases. In general it is admitted that
the blood is discharged by a reflex action, being a means of defense.
He discusses four methods in which the blood is discharged and gives
ni = a MWvrece
nee NH
m AN Le : : ;
C
Fic. 19.—Drawings, illustrating reflex “bleeding” in lady-beetles. A, diagram
of a section through femoro-tibial articulation of Epilachna borealis, showing fol-
lowing: a, pores of gland cells without reservoirs; BS, blood sinus; H, mem-
brane dividing lumen of leg into two chambers; Po, pores of gland cells ‘having
reservoirs; and St, sense hairs. B and C, portion of tubercle on larva of bean
beetle; B, distal end of tubercle having 6 branches (Br), each of which is
terminated with a hair (Hr), 32; and C, distal end of a branch, showing hair
arising from a socket, which is surrounded by 5 processes (Pr), only 2 being
shown, X 320.
examples of insects for each method. He further remarks that the
ejected blood is usually very toxic.
While discussing coleopterous larvae, Hollande shows how coc-
cinellid larvae protect themselves by discharging blood. As an example
he used Epilachna argus, whose body is covered with chitinous tu-
bercles, which in turn bear many smaller branches, each of which
is terminated by a hair. The discharged blood is accomplished by a
rupture of the chitin near the base of the hair. When one seizes the
No. 18 SENSE ORGANS OF COLEOPTERA—McINDOO 61
larva with the fingers, the hairs pierce the epidermis on the fingers
and are then broken off, causing the blood of the larva to exude as
small drops.
The larvae of the bean beetle are likewise covered with hairlike
tubercles (fig. 19, B), which bear many branches (Br), each of which
is terminated by a hair (Hr). While picking up the larvae the writer
observed a yellowish liquid on his fingers. This liquid was bitter and
very distasteful. After carefully examining the larvae under a bi-.
nocular, it was learned that the bitter liquid came from the tips of the
branches (C, Br). Using a needle it was possible to touch the hairs
(Hr) lightly, so that they broke at their weakest point; that is, at the
socket which is surrounded by five processes (Pr).
SUMMARY
This paper is written as a complement to the writer’s (47) former
one entitled “ Tropisms and Sense Organs of Lepidoptera,” and con-
tains information of a similar nature, but dealing with Coleoptera
alone. A large mass of literature on the sense organs and tropisms
of beetles, including papers on light traps, attractive baits, and repel-
lents, has been consulted; but only the more important information
found has been briefly summarized.
The Mexican bean beetle was selected to represent the Coleoptera.
When tested to odor stimuli alone this beetle was found to be an un-
favorable insect ; but when the adults were allowed to come in contact
with the substances to be tested as foods, the beetles clearly demon-
strated their “likes” and “ dislikes”’; and when tested to light and
gravity in a dark-room, the adults proved to be almost ideal for this
purpose. In order to obtain comparative results which could be treated
statistically, new technique and apparatus were devised, and the more
important experiments were repeated many times under controlled
conditions. The more important results obtained are as follows:
When tested in a phototactic box, which lay on a table by a south
window in bright light, although not in direct sunshine, larvae of the
first and second instars were weakly photopositive or indifferent to
light. Most of the larvae of the third instar and the more active ones
of the fourth instar were strongly photopositive. As a rule, the larvae
up to the time of pupation were found to be photopositive, but when
ready to pupate they became photonegative. Whether the negative
reaction is caused by a change in the structure of the ocelli is not
known. Hundreds of adult bean beetles were also tested and all
proved to be photopositive, most of them being strongly so.
62 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 82
In a dark-room in which the temperature and relative humidity
were fairly constant many tests were conducted to determine the differ-
ence between the phototactic and geotactic responses of adult bean
beetles and their larvae, with and without the use of light. The in-
sects were confined in a photo-geotactic box, just above or below
which lay a water screen to prevent the infra-red or heat rays from
reaching the beetles. Under these conditions the following results
were obtained. For active, overwintering adult beetles the geonega-
tive or upward response, when light was used, was 25.60.20 per cent
stronger than the geopositive or downward one; but when no light
was used, it was 54.6+0.17 per cent stronger, indicating that when
the beetles were forced downward by the light this stimulus overcame
about one-half of the geotactic one. Old beetles of the second brood
did not respond so readily, yet their geonegative responses were
stronger than their geopositive ones. Larvae of the third instar did
not respond readily and they went up only slightly more than down.
When light was used, active larvae of the fourth instar reacted as
readily as did the overwintering adults; but when no light was used,
they did not respond so readily, although they went up more than
down. |
While searching for attractants and repellents an improved feeding
method was devised. The adult bean beetles were confined in four
small wire-screen cages, each of which contained a row of the same
four foods, but differently arranged. This series of tests, with the
foods differently arranged each time, was then repeated three times in
the forenoon, and usually the four series were again repeated in the
afternoon. Each individual food used was therefore tested 16 times
in the forenoon and usually 16 times in the afternoon. According to
the arrangement of food, no two rows in the same cage were exactly
alike ; likewise, no two rows of all 16 rows were identical, although
the distribution of food was not so complete. This plan was adopted
in order to equalize the number of beetles counted on the same food
which lay in all four positions during any one series of tests; and
furthermore, everything possible was done to obtain reliable data
which could be treated statistically. Using this plan the following
results were obtained.
To determine whether bean beetles “like” or “ dislike’ the four
classes of substances which produce the four human attributes of
taste, many series of tests were conducted. It was soon learned that
they have “likes”? and “ dislikes”’ in regard to food. They “ dis-
liked” water containing salts, acids, bitter materials, and saccharine;
but “liked” the other sweet substances tested, including cane sugar,
No. I8 SENSE ORGANS OF COLEOPTERA—McINDOO 63
grape sugar, table molasses, corn sirup, and black-strap molasses, and
even showed preferences between them.
Bean foliage, sprayed with arsenicals, was repellent, but not suffi-
ciently so to prevent the leaves from being eaten. Lead arsenate
was most repellent; magnesium arsenate was less so; and calcium
arsenate was least repellent. Bean foliage sprayed with sweetened
arsenicals was more attractive than unsprayed foliage. Bean foliage
sprayed with sweetened magnesium arsenate was more attractive than
foliage sprayed with nonsweetened magnesium arsenate. This would
indicate that it might be of economic importance to use sweetened
arsenicals in control measures, particularly to poison the overwinter-
ing beetles early in the season.
In regard to the tropic receptors of the bean beetle, the following
may be stated. The structure of the compound eyes and ocelli was
not studied, but these organs are normally developed and seem to
function adequately, so far as beetles are concerned.
Two kinds of so-called smelling organs—certain hairs on the an-
tennae, and pores, called olfactory by the writer—are fully described.
These hairs appear long and slender, have thin, almost transparent
walls, and are connected with the sense cells. They are numerous and
lie in five groups on the distal ends of the ninth, tenth, and eleventh
segments. The olfactory pores on the adult beetle were found as
usual on the elytra, wings, legs, mouth parts, and antennae. The total
number counted was only 692. The pores on the larva lie on the head,
antennae, legs, and mouth parts. The total number found was only
62. The fact that the total number of pores on both adult and larva
is comparatively small might be correlated with the fact that this
species is ‘‘ stupid” when olfactory responses are considered.
A so-called taste organ was found at the tip of the maxillary palpus
of the adult. It is a soft plate which bears about 447 tiny, thin-walled
sense hairs. The fact that the bean beetle possesses two of these
highly developed sense organs helps to explain how these insects were
able to distinguish so readily between the various aqueous solutions
and insecticides fed to them.
The only so-called auditory organ found in the bean beetle lies in
the second antennal segment. These structures, called Johnston or-
gans, were formerly assumed to be auditory in function, but now are
believed to be muscular receptors to register the movements of the
antennae.
The remaining receptors described are the tactile hairs, which are
widely distributed over the surface of the beetle.
64 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 82
In connection with the receptors the scent-producing organs and the
phenomenon of reflex “bleeding”? were studied. The only scent-
producing organ found was the unicellular glands, which are dis-
tributed over the entire body surface. The bean beetle, like other
coccinellids, is well supplied with these hypodermal glands. The chief
purpose of the secretion is that of protection, but it probably also aids
the beetles in recognizing the different individuals and sexes of the
same species. When disturbed the adults eject small drops of a
glandular secretion from the femoro-tibial articulations, This is called
reflex “bleeding.” The larvae of the bean beetle also protect them-
selves in a similar manner. When they are handled or even touched
the yellowish and bitter “ blood’ exudes from ruptures at the bases
of the hairs, which terminate the branches on the tubercles.
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