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SMITHSONIAN 
MISCELLANEOUS COLLECTIONS 


VOL. 85 


Ooo CCCong, 





“EVERY MAN IS A VALUABLE MEMBER OF SOCIETY WHO, BY HIS OBSERVATIONS, RESEARCHES, AND 
EXPERIMENTS, PROCURES KNOWLEDGE FOR MEN’’—SMITHSON 


(PUBLICATION 3175) 


CITY OF WASHINGTON 
PUBLISHED BY THE SMITHSONIAN INSTITUTION 
1933 


THE SCIENCE PRESS PRINTING COMPANY 
LANCASTER, PENNSYLVANIA 


ADVERTISEMENT 


The present series, entitled ‘‘ Smithsonian Miscellaneous Collec- 
tions ’’, is intended to embrace all the octavo publications of the In- 
stitution, 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, min- 
eralogy, 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 impor- 
tant additions to knowledge. 

C. G. ABzot, 
Secretary of the Smithsonian Institution. 


(iii ) 





LO; 


LE 


CONTENTS 


Aspot, C. G. Weather dominated by solar changes. 18 pp., 4 
text figs. Feb. 5, 1931. (Publ. 3114.) 

WETMoRE, ALEXANDER. The avifauna of the Pleistocene in 
Florida. 41 pp., 6 pls., 16 text figs. Apr. 13, 1931. (Publ. 
3115.) 

Watcott, CuarLes D. (With explanatory notes by Charles E. 
Resser.) Addenda to descriptions of Burgess shale fossils. 
40 pp., 23 pls., 11 text figs. June 29, 1931. (Publ. 3117.) 

Tuériot, I. Mexican mosses collected by Brother Arsene 
Brouard—III. 44 pp., 22 text figs. Aug. 25, 1931. (Publ. 
2122.) 

Brackett, F. S., and LippeLt, Urner. Infra-red absorption 
bands of hydrogen cyanide in gas and liquid. 8 pp., 5 text 
fies. Angry 5, tosis. ( Publ. 3123. ) 

Snopcrass, R. E. Morphology of the insect abdomen. Part I. 
General structure of the abdomen and its appendages. 128 
pp., 46 text figs. Nov. 6, 1931. (Publ. 3124.) 

McAtTer, W. L. Effectiveness in nature of the so-called pro- 
tective adaptations in the animal kingdom, chiefly as illus- 
trated by the food habits of Nearctic birds. 201 pp. Mar. 
15,1932. (Publ..3125.) 

SwanTon, JoHN R. Modern square grounds of the Creek In- 
dians. 46 pp., 5 pls., 15 text figs. Nov. 11, 1931. (Publ. 
3126.) 

WuLF, OLiver R. The determination of ozone by spectrobolo- 
metric measurements. 12 pp., 3 pls., 5 text figs. Nov. 30, 
1O¢1. (Publ 2127,) 

Micrer, Gerrit S., Jk. Human hair and primate patterning. 
12 pp, 5 pls..Wec. 19,1031. 0( Publ. 3130.) 

AvpricH, L. B. Supplementary notes on body radiation. 12 
pp., 5 text figs. Feb. 2, 1932. (Publ. 3131.) 














SMITHSONIAN MISCELLANEOUS COLLECTIONS 


VOLUME 85, NUMBER 1 


HHodgkins Fund 


and 


Roebling Fund 


WEATHER DOMINATED BY SOLAR 
CHANGES 


BY 
C. G. ABBOT 





(PUBLICATION 3114) 


CITY OF WASHINGTON 
PUBLISHED BY THE SMITHSONIAN INSTITUTION 
FEBRUARY 5, 1931 


The Lord Baltimore Preas 


BALTIMORE, MD., U. 8. A. 


Hovgkins Fund and iocbling Fund 


WEATHER DOMINATED BY SOLAR CHANGES 
By C..G. ABBOT 


My title suggests a radical change of view regarding weather and 
weather forecasting. Let us contrast, for a moment, weather and 
climate. All men realize that it 1s the sun which furnishes the heat 
which warms the earth, and that the regular motions of rotation of 
the earth upon its axis, and of its revolution in its orbit around the 
sun produce those periodic variations of the solar heating which 
govern climates. Differences in latitude and of proximity to oceans 
and to other great terrestrial features introduce alterations from place 
to place in these periodic changes of solar heating; thereby are pro- 
duced climatic differences. As regards weather, which consists in 
departures from regularity in climate, I suppose that practically all 
meteorologists have been holding hitherto that it depends principally 
on the complexities of the earth. According to that view, weather 
represents, as it were, the changing eddies and whirlpools in the 
Niagara of climate, due to the jutting rocks of local circumstances, 
and, owing to enormous complexities, 1s essentially unpredictable for 
any considerable time in advance. 

I shall present evidence to show that weather, on the contrary, is 
caused chiefly by the frequent interventions of actual changes of the 
emission of radiation within the sun itself. Local conditions, to be 
sure, alter the magnitudes and times of the effects of these interven- 
tions into terrestrial affairs by the variable sun, but in ways determin- 
able by statistical studies. Hopeful indications will be given that 
changes of the solar radiation and their weather-consequences may be 
predictable long in advance. 

Figure 1 shows the daily observations of the solar constant of radia- 
tion made at Montezuma, Chile, by the Astrophysical Observatory of 
the Smithsonian Institution since 1924. The values give the intensity 
of the sun’s radiation as it would be found by an observer in free 
space situated at the earth’s mean distance from the sun. As far as 
possible, they are independent of any effects of the varying trans- 


SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 85, No, 1 


1924 























































































































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Fic. 1.—Daily observations at Montezuma, Chile, of the “ Solar Constant of 
changing about the mean value, 1.94 calories. Circles, crosses, dots repr¢ 




























































































































































































SHOWS ASCENDING SEQUENCES 
SHOWS DESCENDING SEQUENCES 





* since 1924. Shows that the sun’s gift of rays to warm the earth is frequently 
ectively satisfactory, nearly satisfactory, and unsatisfactory observations. 


4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


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NO. I WEATHER DOMINATED BY SOLAR CHANGES—ABBOT 7. 


parency of our earth’s atmosphere. No appreciable 12-month peri- 
odicity appears in the results. This is a good sign of their indepen- 
dence of atmospheric influences. Full and dotted curves in Figure 1 
mark all the well-supported sequences of rising and of falling solar 
radiation. They occur in short intervals, averaging 5 days. All of 
those selected exceed 0.4 per cent in range, averaging 0.8 per cent. 
These rising and falling sequences are 111 and 106 in number, respec- 
tively. Many are lost because of unfavorable observing conditions. 

Figure 2 shows average changes in the mean temperature and the 
barometric pressure at Washington, D. C., associated with these rising 
and falling sequences of solar radiation, during the months of March, 
April, September, and October. These meteorological exhibits are 
average values representing the work of 7 years, and of about Io cases 
of each kind in each month. 

The method of computing the curves shown in figure 2 is illustrated 
in tables 1 and 2 as regards temperatures of March. The temperatures 
(which are the mean of maximum and minimum at Washington as 
published by the U. S. Weather Bureau) are arranged in consecutive 
series of 25 days each. In each series, the fifth day is that on which 
the solar change examined reached its culmination. Departures of 
temperatures are always computed from the first day of the series 
as the base. The mean values of all the departures occurring in 
March in the years 1924 to 1930 are given at the foot of the table. 
They are corrected to eliminate the secular rise of temperature which, 
of course, occurs during any 25-day interval at that season of the year. 
The final result is plotted in figure 2. The reader will see that in all 
cases there is a marked opposition between curves corresponding to 
rising and falling solar radiation, respectively. 

Eleven physicists to whom I have shown these results unanimously 
concur in advising me that the constant opposition of the weather 
effects following opposite solar causes demonstrates a physical con- 
nection between the weather of Washington and the changes in the 
solar constant of radiation as observed in Chile. Average changes 
of mean temperature of 5° Fahrenheit are found corresponding to 
solar changes averaging only 0.8 per cent. Hence we may suppose 
that on many occasions temperature effects caused by solar changes 
may reach 10°, and sometimes 15° or 20°. That is to say, major 
changes in weather are due to short period changes in the sun. So 
revolutionary is this conclusion for meteorology, that I hesitated to 


8 SMITHSONIAN MISCELLANEOUS COLLECTIONS vot. 85 


publish it until the unanimous approval of many competent critics 
encouraged me. I am further supported in this view by having found 
a similar opposition of relations prevailing not only at Washington 
but at Williston, North Dakota, and Yuma, Arizona, in all months 
of the year. 

By what physical connection are these surprising meteorological 
results produced by such small solar changes? We must discard at 
once, I think, the idea that changes of ground temperature, directly 
produced, communicate the effects to the surface air. For firstly, by 
Stefan’s law, in equilibrium conditions radiation varies as the fourth 
power of the absolute temperature. Hence a change of 1 per cent in 
radiation, if acting directly and in equilibrium conditions, should re- 
quire but + per cent change in the earth’s temperature. Actually the 
change of temperature observed exceeds 1 per cent, reckoned from 
the absolute zero. Secondly, in March and some other months, a tem- 
perature effect at Washington is found to be nearly simultaneous with 
the solar change. The solid earth has too large a capacity for heat to 
follow in temperature thus quickly. Thirdly, large effects occur at 
Washington 10 or 12 days, and sometimes 16 or 17 days, after the 
solar cause ceases. Not all of these effects can be direct. Fourthly, in 
September a reversal of sign is observed. 

Admitting that the meteorological effects are produced indirectly, 
let us recall: Firstly, that from 10 to 25 per cent of the solar radia- 
tion is primarily absorbed in the atmosphere itself, which has a very 
small capacity for heat. Secondly, that the atmosphere circulates in 
great cyclonic whirls. Thirdly, that the temperature of a station 
depends greatly on the prevailing wind direction. May it not be that 
the instantaneous changes of heat absorption in the atmosphere tend 
to displace centers of cyclones, and thereby to alter the wind direction 
at stations, thus altering their temperatures ? 

How shall we explain deferred effects occurring 10 or even 17 days 
after the culmination of solar sequences? May they not result from 
atmospheric waves drifting in a southeasterly direction from distant 
centers of action where primary effects are produced? If so, we 
must perceive that the average effects shown in tables I and 2 can 
form no trustworthy basis for forecasting individual cases. For 
primary and secondary effects, treading on each other’s heels, as it 
were, must often interfere, and either augment or reduce expected 
weather changes. 


NO. I WEATHER DOMINATED BY SOLAR CHANGES 





ABBOT 9 


SOLAR PERIODICITIES 


It would be encouraging from a forecaster’s standpoint if definite 
periodicities’ should be found in solar variations. In table 3 are given 
10-day mean values of solar radiation from 1918 to 1930." A ten- 
dency towards the recurrence of a certain form of 8 months’ period 
was discovered in the 10-day means. To evaluate this periodicity, 
the 10-day mean values were arranged in a table of 9 lines of 24 con- 
secutive values each, beginning with May, 1924. Mean values of the 
24 columns being computed, they resulted thus: 


8-month period 


Direct, (Mieanis@” wclsc eanelesie.es 40 41 42 41 44 41 41 #42 «41 43 «+42 «+40 41 42 
Smoothed Means ............ 40 41 42 42 43 #42 #42 «42 «42 «42 «#41 «4t)«64t) «(41 
Smoothed Departures ........ o +r +2 +2 +3 +2 +2 +2 +2 42 +1 +1 +1 GIL 
Direct. Means i. 2..0<50000<. 39 41 4o 38 41 4t 39 38 «37° «37 
Smoothed Means ............. 41 40 40 40 39 39 39 «638 © «603706037 
Smoocthed Departures ........ +1 0 oO o —I —I —I —2 —3 -—3 


« First two figures omitted. Thus for 1.940 calories, I substitute go. Departures are given 
from 1.940, omitting three figures. 


From these numbers a smoothed curve was drawn which gave the 
departures from 1.940 calories. Subtracting these departures, the 
original data were cleared of the 8-month periodicity from January, 
1924, to December, 1930. It was then perceived that another peri- 
odicity of 11 months seemed present. By a similar arrangement 
in lines of 33 consecutive revised 10-day means of solar constant 
numbers, the following values were computed, representing the 
11-month. periodicity : 


I1-month period 


Direct eiMeans! <24 sh eteasavie ce 4o 41 39 38 «638 «6936 «638 «30-335 37) 37s 34s 38s 400 
Smoothed Means ............ 41 40 39 38 38 37 37 37 36 360 «636637 «638 = 630 
Smoothed Departures ........ I 0 —I —2 —2 —3 —3 —3 —4 —4 —4 —3 —2 —I 
Direct Means ..........00000 40 41 43 44 41 40 38 42 42 40 42 #45 43 «46 
Smoothed Means ............ 40 41 42 42 41 40 40 41 4 42 42 «43 «45 ~«46 
Smoothed Departures ........ o I 2 2 I oO oO I I 2 2 RB 5 6 
DinectoiMieans cance secession 44 45 43 43° «41 
Smoothed Means ............ 45 44 43 #42 «4! 
Smoothed Departures ........ 5 4 2 2 I 


As these two periodicities had been evaluated solely from results 
of 1924 to 1930, I desired to see whether they were also in evidence 
from 1918 to 1923. For this purpose, I made templates fitting the 
smoothed-curve departures for both periodicities. These templates 
I traced again and again in their proper phases to fill the entire period 


* The best values are those obtained since January, 1924. Prior to August, 1920, 
all observations were made in the outskirts of the city of Calama, amid dust and 
smoke, and with less perfect equipment than subsequently. Prior to January, 1919, 
there was only one observation per day and by the “long” method. 


IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


from August, 1918, to December, 1930. I then added their amplitudes 
algebraically. This produced a curve which obviously bore a con- 
siderable resemblance to the curve A of figure 3 throughout its whole 
extent. This indicated that both 8- and 11-month periodicities have 
prevailed in solar radiation since 1918. 

I now desired to search for longer periodicities. It seemed better 
to use monthly mean values for this, as given in table 4 and figure 
3, A. Having read from the curve of combined departures of 8-month 
and r1-month periodicities the departures for the second decade of 
each month from 1918 to 1930, I subtracted these from curve A, 
figure 3, and replotted the again-revised data. This curve seemed to 
indicate the existence of a periodicity of 45 months. Arranging the 
corrected solar values in lines of consecutive 45’s, and proceeding as 
previously, the following result appeared: 


45-month period 


Direct) AMieans serxcis 2 <sieecrec ee 20) 32) “4135 37) 28) (4h) 633) at bag 47 37 edge AS 
Smoothed Means ............ 3333) 934) 1835) 590, 37 38.030) dOnt gen ageeasieraa 
Smoothed Departures ........ 7 —7 —-6 —-5 —4 —3 —2 —I o +1 +2 +3 +3 +4 
DinectedMieans; esjeissnes violetecerate 44-45 AY 38) 46-46: 44" 46" 4t 50)) 743) 437 az vad 
Smoothed! WMeans (.iiciices cece 44 45 45 45 46 46 46 46 46 46 45 45 45 45 
Smoothed Departures ........ +4 +5 +5 +5 +6 +6 +6 +6 +6 +6 +5 +5 +5 +5 
Direct” Means, aie .ciee sic.s.c.cteve oie 44 30 40 40 38 48 40 47 47 4I 4r 39 40 39 
Smoothed Means ............ 45 45 45 44 44 43 43 42 4 41 .40 39 37 36 
Smoothed Departures ........ See ci Saeed eet Ae eee ee eo o —I —3 —4 
Dinect je Mieans) 201 1e1a)s/</a1e1ers eres 36) 35, 83t 

Smoothed Means ............ 350 340633 

Smoothed Departures ........ —5 —6 —7 


After removing the 45-month periodicity as in former cases, there 
seemed to exist a periodicity of 25 months, which by similar treatment 
resulted as follows: 


25-month period 


Directs Means: i icewccs ceccecte 30!) © 33) 9530537 8 34 Bese AT aS sooo eso ues 740 Soma 
smoothed: Means ...10010...10cis- 32, 32 33, 34 34 35) 350.36. 37) 938) g05) 0 on 40 
Smoothed Departures ........ —8 -—8 —7 -6 -6 —5 —5 —4 —3 —2 —I —I o o 
Direct, IWeansi sacs cslesie esses 4%. 44 43 “40! 43 420 42 43) 42) 40n 33 
Smoothed Means ............ 40, 41 | 4t 42 42°) 42) a2 942° \4t 40. 735 
Smoothed Departures ........ o +1 +1 +2 +2 +2 +2 +2 +1 0 —5 


Removing the 25-month periodicity, as before, a nearly smooth curve 
resulted in which the 68-month period corresponding to a half sun-spot 
period was clearly seen. The coordinates of the five periods discovered 
are as follows: 


Coordinates of Periods 


Length Amplitude Date of Zero 

in Months in Calories Departure 
68 -O14 Dec. 15, 1929 
45 013 Sept. 15, 1930 
25 .O10 Nov. 15, 1929 
II .009 Dec. I, 1929 


8 .005 May I, 1930 


NO. I WEATHER DOMINATED BY SOLAR CHANGES——ABBOT ligt 














































































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1.938) 
1.936 


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1.94 1.93 
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Fic. 3.—Periodicities in solar radiation. 


TZ SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


I next made templates and traced the five periods hitherto described in 
a way to cover the entire interval 1918 to 1930. The total effect of the 


TABLE 3.—TZen-Day Solar Constant Values, 1918-1930 


Decade 1918 1919 1920 1921 1922 1923 1924 1925 1926 1927 1928 1929 1930 


Jan. I 1.943 1.968 1.956 1.924 1.946 1.037 1.945 1-944 1.9390 1.041 1.925 1.938 
2 1.048 1.967 1.053 1.946 ..... 1.943 1.939 1-943 1.938 1.931 1.941 1.937 
3 ieGkis Jeolte) Goode Te Q5 2s sterile 1.944 1.947 1.933 1-93I 1-942 1.939 1.929 
Feb. 1 T.OO2! LeO58. aicicie TeOll e934. W935) eciaccr 1.938 1.936 1.947 1.937 1.933 
Z 1.951 1.954 1-952 1.947 1-951 1-943 1-951 1.939 1.946 1.941 1.925 1.939 
3 1.930 1.956 1.958 1.948 1.923 1.938 1.938 1.929 ...-- 1.934 1.925 1.942 
Mar. 1 1.950 1.959 1-954 1-949 1.929 1.947 1.941 1.941 ..... 1.950 1.932 1.940 
2 1.942 1.948 1.940 1.939 1.036 1.944 1.036 1.048 1.044 1.945 1.931 1.937 
3 T2931  D2032" eee 1.932 1.931 1.942 1.941 1.932 1-941 1.945 1.932 1.941 
Apr 1 1.943 1-948 1.951 1.930 1-934 1-942 1.945 1.927 1.941 1.946 1.932 1-941 
2 1.957 1.956 1.941 1.937 1-928 1.948 1.950 1.937 1.945 1-940,1.942 1.938 
3 1.961 1.952 1.934 1-025 1-934 1-947 1.946 1.939 1.945 1.940 1.938 1.941 
May 1 1.953 1.950 1.946 1.924 1.934 1.944 1.946 1.937 1-947 1-943 1.936 1.045 
2 1.921 1.961 1.939 1.925 1.935 1.948 1.950 1.938 1.944 1.951 1.941 1.048 
3 TeQ45. TisQ50) VLQ4T) frais ae 1.937 1.950 1.954 1.942 1.944 1.949 1.937 1.942 
June 1 1.957 1.943 1.933 1.910 1.918 1.957 1.943 1.939 1.950 1.947 1.938 1.949 
2 1.938 1.034 1.936 1.913 1.934 1.956 1.943 1.946 1.943 1.948 1.932 1.944 
3 1.962 1.938 1.945 1.920 1.933 1-953 1.948 1.945 1.945 1.951 1.932 1.941 
July 1 1.951 1.945 1.960 1.904 1.034 1.946 1.052 1.942 1.9049 1.943 1.935 1-945 
1.961 1.940 1.957 1.913 1.928 1.951 1.954 1.949 1.942 1.942 1.931 1.949 
3. 1.921 1.950 1.951 1.953 1.918 1.944 1.942 1.047 1.944 1.946 1.940 1.935 1-947 
Aug. I 1.955 1.961 1.930 1.944 1.919 1.942 1.950 1.949 1.045 1.942 1.943 1.931 1.946 
2 L045, 13042 T2927 9s 5.0. 1.916 1.940 1.940 1.941 1.942 1.941 1.937 1.932 1.947 
3 1-950) 1-955) 1-932) asec I.92I 1.941 1.933 1-942 1.942 1.941 1.932 1.930 1.943 
Sept 042) 1.038) 1c0SI> ean uae 1.945 1.941 1.956 1.942 1.940 ..... 1.926 1.942 
2) 1.946 1.942" 1.044) Secu. 1.932 1.9044 1.950 1.946 1.940 1.942 1.938 1.928 1.929 
3 1.044 1.937 1.944 1.969 1.916 1.942 1.946 1.950 1.943 1.950 1.921 1.930 1.939 
Oct. IT 1.951 1-947 1.942 1.959 1.926 1.940 1.953 1.942 1.938 1.945 1.930 1.928 1.939 
2 1.930 1.949 1.950 1.969 1.929 1.942 1.949 1.949 1.937 1.044 1.935 1.933 1.041 
3. E933) 1.9601 “150431 1/0660... 1.938 1.048 1.046 1.929 1.943 1.927 1.926 1.939 
Nov. 1 1.928 1.958 1.951 1.953 1.929 1.934 1.948 1.944 1.931 1.945 1.924 1.932 1.942 
2 1.945 1-951 1.946 1.949 1.035 1.944 1.951 1.948 1.926 1.943 1.932 1.936 1.0943 
3 1.047 1.948 1.045 1.952 1.920 1.044 1.045 1.044 1.930 1.944 1.930 1.939 1.049 

Dec. 1 1.062 1.944 1.957 1.956 1.912 I 042 1.942 1.944 I 
3 : p 5 -935 1.949 1-930 1.941 1.94 
2 1.969 1.949 1.957 1.938 1.916 1.942 1.947 1.045 1.931 1.935 1.924 1.939 ean 
Zi I960" 2-058) I.056. a. 0 ee 1.912 1.921 1.939 1.046 £.935 1.939 1.027 1.940 1.951 


Taste 4.—Monthly Mean Solar Constant Values, 1918-1930 


Month 1018 1919 1920 1921 1922 1923 1924 1925 1926 1927 1928 1929 1930 


Jan. 1.043 1.064 1.955 1.048 1.946 1.942 1.943 1.941 1.938 1.940 1.938 1.9346 
Feb. 1.949 1.956 1.956 1.943 1.930 1.939 1-943 1.938 1.943 1.943 1.929 1.938 
Mar. 1.941 1.945 1-949 1.938 1.932 1.945 1.939 1.939 1-942 1.946 1.931 1.939 
Apr. 1.953 1-952 1.944 I.93I 1.932 1.946 1.947 1.934 1.944 1.942 1.937 1.940 
May 1.940 1.953 1.943 1.925 1.936 1.948 1.950 1.939 1.945 1-947 1.938 1.944 
June 1.955 1.939 1-939 1.914 1.928 1.955 1.945 1.944 1.946 1.948 1.934 1.943 
July I.92I 1.954 1.945 1.956 1.912 1.936 1.946 1.951 1.944 1.945 1.942 1.933 1.947 
Aug. 1.954 1.953 1.930 1.944 1.918 1.941 1.940 1.045 1.944 1.941 1.937 1.931 1.945 
Sept. 1.044 1.939 1.947 1.969 1.924 1.944 1.946 1.950 1.942 1.944 1.927 1.928 1.937 
Oct. 1.939 1.953 1.944 1.962 1.927 1.940 1.949 1.946 1.034 1.944 1.930 1.929 1.940 
Nov. 1.941 1.953 1.948 1.951 1.929 1.941 1.948 1.946 1.929 1.944 1.929 1.936 1.944 
ee 1.962 1.950 1.957 1.953 1.915 1.933 1-942 1.945 1.932 1.942 1.926 1.940 1.947 

early 

Mean .. 1.949 1.948 1.952 1.927 1.937 1.946 1.946 1.938 1.943 1.938 1.034 1.942 


five periodicities is summed up algebraically in curve B, which will 
be seen to represent the main features and even most details of 


Noe I WEATHER DOMINATED BY SOLAR CHANGES—ABBOT 13 


curve A of figure 3. Inasmuch as three of the five periodicities which, 
combined, yield curve B are determined entirely from the work of 
1924 to 1930, and the other two are to a large extent thus deter- 
mined, the part of curve B from 1918 to 1923 may be regarded as 
if it were a forecast. Its good fit* encourages us to expect to see 
these five periodicities continue to hold until 1933, producing the 
general march of solar variation forecasted in curve I of figure 3. 

In former publications dealing with possible solar periodicities, [ 
was indebted to Dr. D. C. Miller for the use of his harmonic analyzing 
machine. Two of the periods which I then thought real, namely of 
about 25 months and 11 months, are re-discovered by my present 
method. I feel better satisfied, however, this time, because there is 
nothing arbitrary about my present analysis. It does not assume 
periods not indicated by the observations as does the ordinary method 
of harmonic analysis, which deals with submultiples of some arbi- 
trarily assumed period. 

I propose soon to apply a similar method to the individual daily 
observations, in the hope of discovering shorter periodicities. Thus 
far I have not gone very far in this line, and will reserve it for a later 
paper. At present, I will only mention that in the year 1924 there 
appeared to be continuing periodicities of 45 days and of the eighth 
part thereof, 5.6 days. These are illustrated in curve H of figure 3. 
Other periodicities seemed to hold from 2 to 4 months and then 
disappear. 

So far, I have disclosed in solar radiation continuing periods of 
approximately $ and 4 of the 114-year sun-spot cycle, and of 1/16, 
1/36, and 1/50 of the Bruckner cycle of 33 years. Besides these there 
were periodicities approximating 45 and 5.6 days in the year 1924, of 
which it is uncertain whether they belong to these families, though 
they approximate to 1/90 and 1/720 of the 114-year cycle. 


WEATHER PERIODICITIES 


If, as suggested by the title, weather is governed by solar varia- 
tion, and if, as has just been shown, the solar variation from 1918 
to 1930 comprises five definite continuing periodicities, we should 
expect to find these same periodicities in the weather. 

For data to investigate this point, I took from ‘ World Weather 
Records ”’* the Washington monthly mean temperatures from 1918 


* Regarding discrepancies of 1918 to 1920, see footnote on page 9. 
* Smithsonian Misc. Coll., Vol. 79, 1927. 


T4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


to 1923. I supplemented them to 1930 by taking monthly mean values 
of “ Max.” plus “ Min.,” as given in the “ Climatological Data.’”’* In 
some previous work I had prepared a plot of the average yearly march 
of Washington mean temperatures. From this smoothed curve I took 
values corresponding to the 15th day of each month, and subtracted 
from my monthly mean data. Thus I obtained the temperature-depar- 
tures which constitute weather, as freed from the average march of 
events which constitutes climate. These results are plotted in curve A 
of figure 4 and given in column 9g of table 5. 

I then analyzed these temperature-departure data in the manner 
already explained regarding the solar data. I employed in my analysis 
the same periods of 68, 45, 25, 11, and 8 months used in the solar 
work. These were found to represent to a surprisingly close approxi- 
mation the variation of Washington temperature-departures since 
1918. The agreement with observed data was somewhat improved by 
adding a sixth period of 18 months. These six periodicities are shown 
graphically in curves C, D, E, F, G, H of figure 4, and their summa- 
tion in curve B. The actual data from which these curves are plotted 
are given in columns 1 to 8 of table 5. 

The reader, I think, will agree with me that the similarity between 
curves A and B of figure 4 is both close and significant. Not only are 
the main trends of the original observations fairly well reproduced in 
the periodic summation, but many of the details also. Discrepancies, 
indeed, occur at several times, and unfortunately a principal one is 
found in 1930. One, therefore, hesitates to predict that the tempera- 
ture departures of 1931 and following years will be defined by the 
same six periodicities without modifications of amplitudes or phases. 
Nevertheless the discrepancy of 1930 is not much more pronounced 
than several preceding ones, after which fair agreements returned. 

It may be objected that the five solar periodicities alone were in- 
sufficient to give the best representation, without adding a sixth of 
18 months not found conspicuously in solar variation. Is not this last 
periodicity possibly of terrestrial origin? May it not be due to some 
peculiarity of Washington surroundings which lends a predisposition 
to a periodicity of 18 months? For analogy, consider an automobile 
on a dirt road. It vibrates as the wheels strike the irregularities of 
the road, in a manner depending on these outside interferences. But 





“Issued monthly by the United States Weather Bureau, Washington, D. C. 


WEATHER DOMINATED BY SOLAR CHANGES—ABBOT 


I 


NO. 


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16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


TaBLe 5.—Periodic Analysis of Washington Temperature Departures 





Periodicities 
Degrees x 10 ahr. 
Original 
68 m 45m 25m 18 m 11m 8m Sum data 
TOTS VAN: ccisreremecis cre ceesocioss I 5 —7 14 —4 —16 —0.7 —9.7 
IED: warstersio siare crstateveiersie o 4 —5 19 —2 —3 1-3 0.8 
Mia ri reteicsis clcleists:-feleiciei 1 3 =—3 21 0 —I 1.9 5.1 
April, (seceerroe caistetsiors —2 3 —I 16 5 —7 1A) 0.2 
eliyammerctoforietereesctelslsrerereiers —3 3 3 a 8 —8 1.0 5-5 
AUN eee tere erorndeeciostelet —4 2 8 —4 10 —3 0.9 —2.7 
Ata liyjenetererereteveieceroteilereieiereie —s5 2 13 —I2 o 10 0.8 —2.9 
AIS ace ete —6 2 17 —I4 —2 6 0.3 2.5 
Sians “ddodeoouscadccdas —7 2 21 —14 5 —16 —0.9 —4.0 
(OYSEL. USaaoddecusocgncuuc —8 2 23 —9 12 —3 1.7 4.1 
ING Wapericrcticiemiaciiscerer —9 I 23 —4 10 —I 2.0 2.0 
WGC! (Sores tensa ccrecuereterete —10 I 23 oO —4 —7 0.3 5.2 
TOTO) JAN ~ siocetsisisissicistevclereiwtere @ —II oO 21 2 —2 —8s 0.2 4.5 
Ive Dae rarstetarsiels\eierarcrstonevercicte —12 —I 17 3 0 —3 0.4 1.2 
Mare 2 sisrcrnistelsl syeeyavsteisivele —13 —3 14 4 5 10 3.0 Bur 
April Rereewisissinclosisenee —14 —6 II 5 8 6 2.4 0.8 
DY same stavatels ctetevore tolerate everss —I4 —9 7. 7 10 —16 0.2 0.5 
AMUUTLS area rsioseul stele Sielarsteere.s —I5 —I0 5 10 oO —3 0.2 0.4 
Wivtlliy ‘etersieie oete'e «wis wioies assis —I15 —I0 2 14 —2 —I 0.3 —o0.I 
WAIST onccoorsisle slvewie lease —16 —Il oO 19 5 —7 0.6 —1.5 
Sept eaesciinccctlacser —I15 —Il —3 21 12 —8 Ter 1.2 
Oct ua ase eeteneeetece —15 —10 —s5 16 10 —3 0.8 6.8 
ING Wan icicteisiateveiniersiercioicteis —I5 —9 —7 7 —4 10 —0.3 2.5 
MD CCH erence sie ccleiete mle \eerere 14 7 8 4 2 6 —1.5 —4.2 
TOZOMPAI.. aiesicis crs sisiece crests (sieve —I4 —3 —9 —12 0 —16 —5.4 —4.7 
eb, atacistee 000 sine veces —13 8 —7 —14 5 —3 —2. —3.3 
Maar. oosies sis sienresisie nines —I3 29 —5 —I4 8 —I 0.4 2.2 
VA DTI Merete oens.s teers —I3 25 —3 —9 10 —7 0.3 —0.4 
IVa tavercicsareeceeisiase steve —I2 7 —I —4 oO —8 —1.8 —3.9 
MVIUATLE TS ease levefoxstarsfeyevorestorsrare —12 I 3} oO —2 —3 —1.3 —1.9 
Bia yr’ ers tstesale srororo tees ata; zisvere —ilI oO 8 2 5 10 1.4 —2.3 
PASM acisicisiversisleierscletele —I0 oO 13 3 12 6 2.4 —0.3 
Septat) sacle s:swie cavers ewleete —o oO 17 4 10 —16 0.6 0.6 
OGte antisense 5 ceeeitiecicle —8 I 21 5 —4 —3 12 4.8 
INGOV es, bsiaisiss2/oisisfevsverersieler svete —7 2 23 7 —2 —I zee 1.6 
ID) Oy epareressssis ote locetoisicin eters —6 3 23 10 ° —7 253 2.9 
TO2L Mat epercts: sive speresyavejayscctcine —4 5 23 14 5 —8 305 3-2 
EE Dee cretecsteisiaitts sisi ctcTovaisints —3 8 21 19 8 —3 5.0 3.0 
Mare suis wciciosceee oninele —I 10 17 21 10 ~ 10 6.7 12.2 
April, (adissisissilessiaiess oO 12 14 16 oO 6 4.8 5.8 
IMiaiy:, Sorretiscpiscieclouneete a 13 II 7 —2 —16 1.5 —1.8 
Viti Ctireteicreeiicrtelclesicters siete 4 13 7 —4 5 —3 2.2 0.7 
ALY ip, csserctore orvavetore era) erareiers 6 II 5 —12 12 —I 2.1 2.0 
AUD Sie cicccis sie saistecse 7 9 2 —I4 10 —7 0.7 —2.5 
SEpts sarapsisasine see@ecs 8 a o —I4 —4 —8 —I.1 —3.8 
OCEM Saiviea-sctsisistnners sas 9 5 —3 —9 —2 —3 —0.3 0.6 
INO Vatoe Section te cists 10 4 —5 —4 0 10 TS 3-2 
DD Cciataretstereietatteerisoore ter Il 3 —7 oO 5 6 1.8 1.5 
TO22EV AN ee Ncreentemics aes 12 3 —8s 2 8 —16 0.1 —1.4 
ebty were ciel setactctere cist atere 12 3 —9 3 10 —3 1.6 2.6 
Mar. Ce rverccccccccece I2 2 7 4 Oo —I TO iT 
SALDEAL™ (rete steno shies 13 2 —s5 Fi —2 —77 0.6 2.6 
AVM oeleleieyroboisteletctorsteistave 13 2 —3 a 5 —8 1.6 2.7 
JUNE > Ask misctwmerie tite 13 2 —I 10 12 —3 3:3 1.0 
Uitte avcrteerattcre tate cco or 12 2 3 14 10 10 5.1 —0.7 
ALi actararate stataverevelchavelane 12 I 8 19 —4 6 4.2 —2.0 
Sept cctsiec eee soe 12 I 13 21 —2 —16 2.9 1:7, 
OCt) soaienecasinesiioes II o 17 16 oO —3 4.1 3.0 
NOVA vostieiiteisetetlonnses II —I 21 7 5 —I 4.2 3.6 
DOCS sas wine rnfu te isyevereraynrs ore 10 —3 23 —4 8 —7 27, 1.2 
1923 Jan. siatololeterecerelerete Mijetrels 9 —6 23 —I2 10 —8 1.6 3-4 
ED: 2 ise tee ost tiins 8 —9 23 —14 oO —3 0.5 —3.4 
IM AtY Semitsreterteeintosite 7 —I10 21 —I4 —2 10 IZ 2.1 
PADIS sSevootante deminer 6 —10 17 —9 ig 6 1.5 0.6 
MIA hii sie crerocrtnseereies 5 —1I 14 —4 12 —16 0.0 —0.7 
JUMP o2jn(ato.ciessiveiareieielelie, ete 4 —i1 II ° 10 —3 Tn 21 
ANCA Ys sessreie: visite wivueie oveieierete 3 —10 7 2 —4 —f —0.3 —n6 
II yc teenie sete sin 2 —9 5 3 —2 —7 —o.8 —0.7 
ere Pincitidst ese I —7 2 4 0 —s —o.8 1.4 
nee Yeperera te rererorcierste eter ; 0 —3 oO 5 5 —3 0.4 —03 
OW scale cvstereainleheiertarsisiaye =I 8 —3 7 8 10 2.9 0.8 
DOCH tanioe ios ceieiteciee —2 29 —5 10 10 6 4.8 8.6 


NO. I WEATHER DOMINATED BY SOLAR CHANGES—ABBOT 17 


TABLE 5.—Periodic Analysis of Washington Temperature Departures—(cont’d) 





Periodicities 
Degrees x 10 Fahr. 
— = — Original 

68 m 45m 25m 18m I1m 8m Sum data 
EOPAMIAM SY yan alosarararcisrearors naerere —3 25 7 14 0 —16 Ta 1.8 
He botgencaceincceee etek —4 7 —8§8 19 —2 —3 0.9 — 7 
Mian, seajesciae slew nates —5 I —9o 21 FS —I 1.2 —o.6 
ADT lexckers sisiestaisresisistetere —6 0 —7 16 12 —7 0.8 —1.2 
May fies caiccscc aa neces —7 oO —5 7 10 —8 —0.3 —3.9 
WAIN Ce oercce vee. Sneek —8s oO —3 —-4 —4 —3 —2.2 —1.9 
uJintlivaiee steetnencca ceevstecte —9 I —I —12 —2 10 —1.4 —3.0 
PAC oO Saerneee ict eelomine vse —10 2 3 —I4 oO 6 —1.3 —2.0 
Septimecccic meres —II 3 8 —I4 cS —16 —2.5 —4.4 
GE serenrsieaeis area ances —I2 Ss 13 —9 8 —3 0.2 3-3 
NOV: odes ates cemcisiases —I13 8 17 —4 10 —I 17, Ley, 
IDECH pete ctcals eaterecoes —I4 10 21 oO oO —7 1.0 0.6 
TOZG lanl omer alee sees —I4 12 2 2 —2 —8 3 —0.4 
Feb. nevis sewelemactiocn —I5 13 23 3 5 3 2.6 7.3 
Miatig aescccarscdamatec —I5 II 23 4 12 10 4.5 4.9 
PADTIL, els iss wig soa va.ctore —16 9 21 5 10 6 3:5 3.8 
May? iets ioc aitsuisiiteles —I5 7 Te 7 —4 —16 —0.4 —3.4 
UNE cc ieivistc Seta iealeics se —15 5 14 10 —2 —3 0.9 4.2 
uty. Geyeteces care trernara, sarees —I5 4 II 14 oO —I se) O.1 
ATI oiioiee dgea aise wrelod —I4 a 7 19 5 —7 ta3 —1.6 
Septih casi aus aiyoertdeatas —I14 3 5 21 8 —8s 1.5 5.0 
OCEL eae ee ee mnateeses —13 3 2 16 10 —3 1.5 —3.9 
INO asthe aca ateecont acs —I3 2 0 7 o 10 0.6 —0.I 
IDECEM ais aontnco es ticles —13 2 —3 —4 —2 6 —I.4 1.0 
19026: Jans \eicdcsvesiewineiensiaser —I2 2 —5 —I2 5 —16 —I1.4 O.1 
Heb perish waicet weasels —I2 2 —7 —I4 12 3 —2.2 1.0 
Mars. Wevsiermayetastaaareraisielets —II 2 —8 —I14 10 —I —2.2 —3.2 
April eitceewicnatenaases —I10 I —o —9 —4 —7 —3.8 —I.2 
AYa Uasree sae tiereeiasee —9 I —7 —4 —2 —8 —2.9 2: 
Att Chyna cemetrcrecia st —8 0 —§5 oO oO —3 Ta —4.1 
Atl yx" Ta tesc Sviaranstnecs adversities —7 —I —3 ES 5 10 0.6 —0.2 
USS weiner saverstveraea ne —6 —3 —I 3 8 6 0.7 1.8 
Sept, 4tceaeehtemssan: —4 —6 3 4 10 —16 —0.9 TS 
Oct Sisasecs secoociows —3 —9 8 5 oO —3 —0.2 ie 
NOV: Giasiacere aie iesareediorarais —I —10 13 ii —2 —1 0.6 0.7 
DOCK sakace testes oes oO —10 r7 10 5 —7 r.5 0.4 
TOS TMUAT Veet ciee eis ences 2 —II 21 14 TZ —8 3-0 Lad 
IMGD:, ciewecieste veeereeaeais 4 —II 23 19 10 —3 2:2 Be, 
Mar, aeitadeiscuiseaeeids 6 —10 2 21 —4 10 4.6 4-7 
BATT sears orev state overatere<rarere 7 —9o 2 16 — 6 4.1 —1.2 
May. siwlasaeeéaiemanisions 8 —7 21 a 0 —16 1.3 TS 
MIN? Bs jac ccces ce eeccas 9 —3 17 —4 5 —3 2.1 —3.5 
Wutliye: vesssSec are setts ccetdicee 10 8 14 —12 8 —I a7 —0.4 
AUG, anjwaciecs creaccacae 11 20 II —I4 10 —7 4.0 —4.7 
Sept, seaiuscasneteacas 12 2 a —I4 o —k 2.2 —o.8 
OCtR canencce ceateenace 12 7 5 —9 —2 ag 1.0 4.4 
NOVA recess sciatic wey i 12 I 2 —4 5 10 2.6 6.6 

DEC. © ceed sie tects s anno 13 0 o o 12 6 aur 2) 
1928 Jan. Baverailefeexetere eee 13 oO —3 2 10 —i6 0.6 2.8 
BieD sc x stiercauicectrecesreacs 13 0 —5 3 —4 —3 0.4 1.9 
IMMrev ipa eceresste/e\sceresslaie)s fries 12 I —7 4 —2 —I 0.7 0.9 
NTA ee ass ocean at eeae ig 2 —8 5 oO —7 0.4 —I1.4 
ME AV "> Fetereiesotiarene/aceiststaverstare 12 3 —9 a 5 —8 1.0 —o.8 
WPM, ceyteasniieceen cee. 11 F —7 10 8 —3 2. —3.3 
AAU VP) Varescvetcvess cis cis icis ators II 8 —5 14 10 10 4.8 1.4 
PAN Getus gis heveawleieistoee 10 10 —3 19 oO 6 4.2 5.7 
DEP. adie cse ceewreaenc 9 12 —I 21 —2 —16 2.3 —2.9 
Oct? | Biisserodewe cecees 8 13 <4 16 5 —3 ae 4.6 
INOW severe aicrstevaxetenarsrsravere oe 7 II 8 Fi 12 —I 4.4 ie 
DG Ci: iperresinevas ars apererocnde 6 9 13 —4 10 —7 2.7 3.1 
1929 Jan. BiNes ovo) hayes a fore afatonets 5 7 17 —12 —4 — 0.5 1.2 
EDs | aacais nae aieelstvsitiasioc 4 5 21 —I14 —2 -3 1.1 —0.5 
SVT estes ciete seat eee B 4 ee —I4 oO 10 2.6 3-7 
ATL “GacvsAievatenies ac sear 2 3 23 —9 F 6 3-0 Ais 
LY cmerletee cio nieicisierersira I 3 23 —4 8 -16 1.5 0.4 
UMN, cave acmesneescinecce oO 3 21 oO 10 —3 3.1 —I1.0 
VE shen teers odeats ees —I 2 17 2 oO —I 1.9 0.0 
PANT G Ss sticraree ale loreranePer Nears —2 2 14 3 —2 —7 0.8 —0.2 
Epts a caseeeneeerete. —3 2 II 4 5 —8 Lo. —2.0 
OC. caeionenmenees ath —4 a F 5 12 —3 1.9 —0.4 
IN GV _ chor mianeectecrnsts —5 2 5 7 10 10 2.9 3.3 
DECs Sateen etae eae —6 I 2 10 —4 6 0.0 2. 


18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


Tas_e 5.—Periodic Analysis of Washington Temperature Departures—(cont’d) 


Periodicities 
Degrees x 10 Fahr. 


Original 
68 m 45m 25m 18 m Ir1m 8m Sum data 
TOZOWMAMG sete cidec ceva tcc —7 I ° 14 —2 —16 —1.0 3-5 
FEDS ateseciewisarisinaciers —8 oO —3 19 oO —3 0.5 6.9 
Mise tater ttises sosinewiew sie —9 —I —5 21 5 —I 1.0 2.0 
ADT! Feet cce.ciesscreraeinels —I10 —3 —7 16 8 —7 —0.3 0.0 
Miaiy? ~ cionicrosiacsiors eisvareloiers —II —6 —s 7 10 —8s —1.6 5.8 
ifitirl eM eecrerrstetecietelnee —I2 —9 —9 —4 oO —3 —3.7 1.9 
Italy? <cteletavscratevessvsterara sere ers —I3 —10 —7 —I12 —2 10 —3.4 1.9 
PAUSE) i srarasicreiocaeve ciate ohn elece —14 —10 —5 —I4 5 6 —3.2 2.1 
Septi! Si casiscmeerenrcsr —14 —II —3 —14 12 —16 —4.8 8.3 
OCH Meetnoteleryecteicieeiacit= —I5 == ed —I —9 10 —3 —2.9 
INO Vomanen seein meie —I15 —I0 3 —4 —4 —I —3.1 
ID GCN y cereinerereiatete sYerereincotet= 1s —9 8 oO —2 —7 —2.5 


at some special speeds, there are sometimes encountered “ sym- 
pathetic ” vibrations due to the make-up of the car itself. 

After all, the contribution of the 18-month periodicity to the fit 
between curves A and B is a minor feature. Is not their surprising 
agreement, which would still be striking if the 18-month curve F were 
omitted, significant because related to solar phenomena? Is it not 
indeed of promising import from the standpoint of long-range weather 
forecasting ? 


SUMMARY 


1. Contrary to the prevailing view, the weather appears to be gov- 
erned by variations in solar radiation. 

2. Long-continuing periodicities in solar variation are found which 
give promise of value for purposes of long-range weather forecasting. 
They appear to be submultiples of 114 and 33 years. 

3. All of these periodicities are found in Washington temperature- 
departures, and, combined, suffice to represent its main features. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS 
VOLUME 85, NUMBER 2 


THE AVIFAUNA OF THE PLEISTOCENE 
IN FLORIDA 


(WiTH S1x PLATEs) 


BY 
ALEXANDER WETMORE 


Assistant Secretary, Smithsonian Institution 





(PUBLICATION 3115) 


CITY OF WASHINGTON 
PUBLISHED BY THE SMITHSONIAN INSTITUTION 
APR 135 1934 


The Lord Baltimore Press 


BALTIMORE, MD., U. S. A. 


Tae AVIFAUNA.OF THE PLEISTOCENE IN FLORIDA 


By ALEXANDER WETMORE 
ASSISTANT SECRETARY, SMITHSONIAN INSTITUTION 


(WitH 6 PLATES) 


Pleistocene deposits of fossils containing numerous bones of birds 
have been known for years in the western part of the United States 
in the Fossil Lake area in Oregon, and in the asphalt beds and caverns 
of California, but such material in other sections of our country to 
date has been decidedly rare and of limited amount. It is of interest, 
therefore, to discuss recent discoveries of abundant avian remains in 
Pleistocene beds in several localities in Florida, with representation of 
a far larger number of species than has been found at any previous 
time in the East. 

Early report of birds in the Pleistocene in Florida came from the 
excavations at Vero on the east coast which initiated the argument re- 
garding the antiquity of man in that area (see fig. 1). There were 
found here remains of a jabiru described by E. H. Sellards, and later 
there came another collection from which Shufeldt named as new a 
gull, a teal, and a heron. More recent excavations by J. W. Gidley and 
by F. B. Loomis, and subsequent work by Doctor Gidley and C. P. 
Singleton near Melbourne, not far from Vero, have brought to light 
many bird bones, while investigations initiated by Walter Wetmore 
Holmes near St. Petersburg on the west coast, in what is known as 
the Seminole Field, have uncovered the most extensive series of fossil 
bird bones that have as yet been found in the eastern part of our 
country. This series is supplemented by bones collected in several 
localities in Manatee County by J. E. Moore, by a few bird bones 
secured by Mr. Holmes from a Pleistocene cave deposit near Lecanto 
in central Florida, and by specimens from several localities in the 
collections of the Florida State Geological Survey. 

The geologic conditions under which these fossils, other than those 
from the cave, are found are briefly as follows: At or below sea level 
on the east coast of Florida is a bed of cemented sand and broken 
marine shells that has been called the Anastasia formation, the Num- 
ber One stratum, or the Coquina layer. At the Seminole Field near 
St. Petersburg, the corresponding layer is of fine white sand con- 
taining many mollusks, less compact than the beds at Melbourne and 


SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 85, No. 2 


2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


Vero. This lower bed is overlaid by a stratum of fine white to light 
brown sand, from a few inches to several feet thick, containing occa- 
sional lenses or groups of marine shells and, locally, accumulations of 
fossilized bones. This is the Number Two layer, usually referred to 
as the bone bed, a deposit that is generally thicker on the east coast 
than on the west. From this bone bed have come fossil vertebrate 





Fic. 1.—Localities where collections of fossil birds have been 
made in Florida. 


remains. Above this bone bed appear Recent deposits of sand or 
humus that form the present surface, though in places the bone bed 
is exposed. 

The actual age of the specimens from the beds in question has been 
subject to some discussion. Dr. O. P. Hay” holds that the Number 


* Journ. Washington Acad. Sci., vol. 20, August 10, 1930, p. 335; and in earlier 
papers. 


NO. 2 AVIFAUNA OF PLEISTOCENE IN FLORIDA—WETMORE 3 


Two bed or stratum dates back to the early part of the Pleistocene, 
probably to the Aftonian period. In this he seems to be upheld by 
Dr. Wythe Cooke.’ Dr. G. G. Simpson’ considers it more probable 
that these deposits are of late Pleistocene age than that they date from 
the earlier part of that period. Doctor Simpson further concludes that 
the faunas from the Number Two bed of the east coast, from Saber- 
tooth Cave, and from the Seminole area “ represent a single phase of 
geologic time.” 

The writer cites these diverse opinions here without attempt to offer 
evidence from the bird material in favor of either one. 


DISCUSSION OF THE AVIFAUNA 


The five principal localities here considered with their fossils may 
be now treated briefly, but before taking these up in detail it is of 
interest to note that though grebes, cormorants, herons, ducks and 
geese, jabirus and other water loving birds are represented among the 
birds of these deposits, there have been found as yet no sandpipers, 
plovers, or other shorebirds, nor any terns or gulls (Larus vero of 
Shufeldt being the yellow-crowned night heron). The lack of gulls is 
of interest particularly since gulls are absent also from the Pleistocene 
of California, where only one bone of a gull has been identified in 
several hundred thousand specimens examined.’ 

In the present studies there have been identified 65 forms of birds 
from the Pleistocene of Florida. Of these three are fossil species of 
the Pleistocene, two of them, a teal, Querquedula foridana, and a 
turkey, Meleagris tridens, being known only from Florida, while the 
third, Teratornis merriami, was described originally from the deposits 
of Rancho La Brea in California. 

There are nine forms that have not been reported from modern 
Florida. Among these is a shearwater, Puffinus puffinus, a pelagic 
species of wide range in the Atlantic Ocean and the Mediterranean Sea 
that comes here in all probability merely as a casual straggler. The 
trumpeter swan, Cygnus buccinator, now nearly extinct, bred formerly 
in the interior of the continent, ranging south in migration to Texas. 
The whooping crane, Grus americana, a breeding form of the interior 
of North America, now nearly extinct, has been reported uncertainly 
from Florida. A small gray crane may be the Cuban bird or the little 
brown crane of western North America. The California vulture, 


* Amer. Journ. Sci., vol. 12, 1926, pp. 449-452. 
* Bull. Amer. Mus. Nat. Hist., vol. 54, February 19, 1920, p. 572. 
*See Miller, Loye, Condor, 1924, pp. 173-174, and 1930, p. I17. 


4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


Gymnogyps californianus, comes only from the West Coast in modern 
times, while an eagle, Geranoaétus sp., has a modern form in South 
America and fossils in the Pleistocene of California. The jabiru, 
Jabiru mycteria, Mexican turkey vulture, Cathartes aura aura, and 
wood-rail, Aramides cajanea are species known today from tropical 
America. 

The 53 forms remaining that are found:in modern Florida are listed 
below. Most of them are common today in the areas under study 
either as permanent residents or as migrants from the north during 
winter. 


Colymbus auritus 
Podilymbus podiceps 
Phalacrocorax auritus 
Anhinga anhinga 
Ardea herodias 
Casmerodius albus 
Egretta thula 
Hydranassa tricolor 
Florida caerulea 
Butorides virescens 
Nycticorax nycticorax 
Nyctanassa violacea 
Botaurus lentiginosus 
Plegadis sp. 

Guara alba 

Cygnus columbianus 
Branta canadensis 
Branta canadensis hutchinsi 
Anas platyrhynchos 
Anas rubripes 

Anas fulvigula 

Nettion carolinense 
Nyroca valisineria 
Nyroca affinis 
Erismatura jamaicensis 
Lophodytes cucullatus 


Cathartes aura septentrionalis 
Coragyps urubu 

Buteo jamaicensis 

Buteo lineatus 

Buteo platy pterus 
Haliaeetus leucocephalus 
Pandion haliaétus 
Polyborus cheriway 
Falco sparverius 
Colinus virginianus 
Meleagris gallopavo 
Grus canadensis (large form) 
Aramus pictus 

Rallus elegans 

Rallus longirostris 
Gallinula chloropus 
Fulica americana 
Zenaidura macroura 
Tyto alba 

Otus asio 

Strix varia 

Corvus brachyrhynchos 
Corvus ossifragus 
Agelaius phoeniceus 
Megaquiscalus major 
Ouiscalus quiscula 


Among these species there are 26 that have not before been recorded 
in the Pleistocene age, a considerable addition to the 114 modern 
species known previously from deposits of that period. 

The fact that at this writing 140 species of the birds found living 
today in that area of North America included in the limits of the 
official Check-list of the American Ornithologists’ Union are known 
as fossils in the Pleistocene illustrates clearly the stability in form of 
our existing species of birds, since this number is more than 15 per 
cent of the total living list (not counting subspecies) for the region 


NO. 2 AVIFAUNA OF PLEISTOCENE IN FLORIDA—WETMORE 5 


in question. Progress in our knowledge of these matters has been so 
rapid that it may be confidently predicted that eventually all of our 
existing species, except those of small size, will be identified in Pleis- 
tocene deposits. As conditions are seldom favorable for preservation 
of small bones of fragile texture, not much can be known of the 
smaller birds as fossils, for their preservation in that state is highly 
fortuitous. We may dream, however, of the discovery of ancient 
caves, inhabited long ago by Pleistocene owls, with great accumula- 
tions of bones of small birds from the pellets of these nocturnal 
predators—caves that have been hermetically sealed for tens of thou- 
sands of years that chance may bring to attention and so give us 
unexpected information on a fascinating subject. 

As his studies in avian paleontology have progressed the writer has 
become convinced that evolution of our existing birds so far as 
differentiation of species is concerned has taken place principally in 
the late Tertiary, and that variation since that time has been of slight 
degree, confined apparently to minor differences (in color and dimen- 
sion) such as are used in our modern studies to distinguish the less 
definitely marked of geographic races or subspecies. As our informa- 
tion increases it appears that some of the differences that we consider 
today as of subspecific value were in existence in birds of the Pleisto- 
cene, for example in the gray cranes and in the turkey vultures, and 
have persisted to the present without apparent change, a striking ex- 
ample of stability in these groups. 

The diversity in the bird life of North America at the time of the 
coming of the rigors of the Ice Age must have been truly remarkable 
since it would seem to have included most of our modern forms to- 
gether with a host of others now extinct that are slowly becoming 
known from the fossil record. The entire period since the opening 
of the Pleistocene has been one of extermination rather than of evolu- 
tion, a process that continued steadily until men appeared as the most 
active factor contributing to its progress. 


THE SEMINOLE AREA 


The region surrounding the small settlement of Seminole, not far 
from St. Petersburg, Pinellas County, Florida, has been designated as 
the Seminole area (see pl. 1). In 1924 Mr. Walter Wetmore Holmes 
discovered here a scute from the glyptodon Chlamytherium septen- 
trionale, and through continued search during the succeeding 5 years 
unearthed numerous other fossil bones including among them many 
remains of birds. It is the Holmes collection of fossil birds that 


initiated the writer’s present studies on the Pleistocene avifauna of 
Florida. 


6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 85 


According to data secured from Mr. Holmes and from a paper on 
the mammals of these beds by Doctor Simpson* of the American 
Museum of Natural History, the fossil bones come originally from 
one deposit in the area in question, many of them being obtained by 
excavating in the original deposit, some coming from erosional wash 
and redeposit by Joe’s Creek which runs through this area, and some 
from the dump along a drainage canal cut through this region. The 
bone-bearing layer is from 1 to 2 feet thick. According to Simpson 
“the lowest bed exposed is of white sand, with numerous marine 
shells, correlated by Cooke with the Anastasia formation of the east 
coast, and hence the equivalent of stratum No. 1 at Vero and Mel- 
bourne. Above this, sometimes with a barren sand layer intervening, 
is the bone bed, equivalent in age and character with stratum No. 2. 
This is generally overlain by a sandy soil, derived from it by weather- 
ing.” Gidley has distinguished above this in places deposits of sand 
and muck that he considers equivalent to stratum Number Three of 
the east coast. 

The list of mammals from this area as determined by Doctor Simp- 
son is extensive and includes among its 49 species a capybara, a bear, 
Arctodus floridanus, a saber-tooth tiger, two ground-sloths, two glypto- 
dons, tapirs, peculiar pigs, camelids, mastodon, and elephant that are 
considered typically Pleistocene species, in addition to opossums, 
moles, rabbits, rodents, skunks, minks, and deer of the same form 
as those occurring in the modern fauna. 

The bird remains collected by Mr. Holmes include 52 forms, this 
being the largest assemblage of fossil species secured to date at any 
point in the eastern portion of North America. The importance of the 
collection is very evident from examination of the list that follows. 
A teal, Querquedula floridana, a huge condor, Teratornis merriami, 
and a turkey, Meleagris tridens, are extinct species of the Pleistocene, 
the first and last being known only from Florida. The jabiru, Jabiru 
mycteria, the Mexican turkey vulture, Cathartes aura aura, and the 
wood-rail, Aramides cajanea, are forms that at the present time range 
in tropical America and are not now known in the present limits of 
the United States. An eagle, Geranoaétus sp., has its only living repre- 
sentative in South America though Pleistocene forms have been found 
in, California. Most remarkable are remains of the California condor, 
Gymnogyps californianus, and a larger condorlike vulture, Teratornis 


*Simpson, George Gaylord, Pleistocene mammalian fauna of the Seminole 
Field, Pinellas County, Florida, Bull. Amer. Mus. Nat. Hist., vol. 54, Febru- 
ary 19, 1929, pp. 501-500, 22 figs. 


NO. 2 AVIFAUNA OF PLEISTOCENE IN FLORIDA—-WETMORE 7 


merriamt. Bones of the former are abundant in the Pleistocene asphalt 
beds of California and the species is known today from northern 
Lower California north into California. It has never been recorded 
before in the eastern part of the country. Teratornis has been known 
previously only from the Pleistocene of California, its occurrence in 
beds of similar age in Florida indicating a former broad range that 
carried it clear across the continent. The occurrence of the whooping 
crane, Grus americana, and of two forms of the brown crane, one 
large and one small, is also of interest. There are 20 modern forms 
in the collection that have not previously been recorded as fossils. 
Following is the list of species from this area: 


Colymbus auritus 
Podilymbus podiceps 
Phalacrocorax auritus 
Anhinga anhinga 
Ardea herodias 
Casmerodius albus 
Hydranassa_ tricolor 
Florida caerulea 
Butorides virescens 
Nyctanassa violacea 
Botaurus lentiginosus 
Jabiru mycteria 
Plegadis sp. 

Guara alba 

Cygnus columbianus 
Branta canadensis 
Branta c. hutchinsi 
Anas fulvigula 

Anas sp. 

Nettion carolinense 
Ouerquedula floridana 
Nyroca affinis 
Nyroca sp. 

Cathartes aura aura 
Coragyps urubu 


Teratornis merriamt 
Buteo jamaicensis 

Buteo lineatus 

Buteo platypterus 
Geranoaétus sp. 
Haliaeetus leucocephalus 
Pandion haliaétus 
Polyborus cheriway 
Meleagris gallopavo 
Meleagris tridens 

Grus americanus 

Grus canadensis (large form) 
Grus canadensis (small form) 
Aramus pictus 

Rallus elegans 

Rallus longirostris 
Aramides cajanea 
Gallinula chloropus 
Fulica americana 
Zenaidura macroura 
Strix varia 

Corvus brachyrhynchos 
Corvus ossifragus 
Agelaius phoeniceus 
Megaqutscalus major 


Gymnogvyps californianus Ouiscalus quiscula 


MANATEE COUNTY 


From Mr. J. E. Moore of Sarasota, Florida, there have come three 
small collections of bones made at as many points in Manatee County. 
The first of these was forwarded to me through Dr. George Gaylord 
Simpson, and comes from deposits near the mouth of Hog Creek near 
Sarasota, Florida. These are said’ to have been found in a stratum 


* Simpson, G. G., Florida State Geol. Surv., 20th Ann. Rep., 1920, p. 274. 


8 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 85 


of blue clay 8 to 14 inches thick overlying limestone from 11 to 12 
feet below the surface. The birds are accompanied by remains of 
Smilodon floridanus, Megalonyx, Mylodon, Chlamytherium, Bore- 
ostracon, Tapirus, elephant, mastodon, and other Pleistocene mammals. 
The bird bones are dull black in color and are heavily fossilized. The 
presence of the California condor is notable. The following species 
are represented : 

Phalacrocorax auritus Gymnogyps californianus 

Botaurus lentiginosus Meleagris gallopavo 

A second collection forwarded by Mr. Moore was obtained from a 

canal within the city limits of Bradenton, the locality being known 
as the Florida Avenue pit. The type material of Parelephas floridanus 
Osborn* came from this point. Mr. Moore (in a letter) states that 
remains of Chlamytherium, Glyptodon and Bison latifrons were ob- 
tained here also. The bird bones examined vary from dull brown to 
dull black in color and are well fossilized. The night heron and ruddy 
duck are here first recorded from the Pleistocene of Florida and from 
these excavations come the best remains of Teratornis. The following 
species are represented : 


Ardea herodias Teratornis merriam 
Egretta thula Meleagris gallopavo 
Nycticorax nycticorax Grus canadensis 
Anas fulvigula Fulica americana 


The third collection was obtained by Mr. Moore at Venice Rocks, 
two miles south of Venice, Florida. The material is fragmentary and 
varies from light brown to black in color, some bones being more 
heavily mineralized than others. Following is the complete list of 
species : 


Buteo jamaicensis Ouerquedula floridana 
Buteo lineatus Nyroca affinis 
Haliaeetus leucocephalus Erismatura jamaicensis 
Casmerodius albus Jabiru mycteria 

Anas sp. 


SABER-TOOTH CAVE 


According to information supplied by Mr. W. W. Holmes, and a 
published account by Dr. George Gaylord Simpson * the sink known 
as Saber-tooth Cave (see pl. 2) is located in a bed of Ocala limestone 


* Amer. Mus. Nov., No. 393, December 24, 1920, p. 20. 
* Pleistocene mammals from a cave in Citrus County, Florida, Amer. Mus. 
Nov., No. 328, October 26, 1928, pp. 1-16, 11 figs. 


NO. 2 AVIFAUNA OF PLEISTOCENE IN FLORIDA—-WETMORE 9 


(Eocene) 1 mile northwest of Lecanto, Citrus County, Florida, on 
property belonging to Mr. D. J. Allen. Several years ago Murray 
Davis with some companions obtained there the canine tooth of a 
saber-tooth tiger (Similodon sp.) which was sent to the Florida State 
Geological Survey. Subsequently Mr. Holmes made arrangements 
for a thorough examination through the cooperation of Mr. Herman 
Gunter, Mr. J. E. King, and Mr. Allen. The work was done under 
Mr. Holmes’ direction in February and March, 1928. 

The entrance to this cave is described * as being “ through a broad 
sink terminating in two vertical shafts. Immediately under these the 
floor of the cave was from 25 to 40 feet below the shaft mouths and 
there apparently has never been an entrance practicable for large 
living mammals. On the floor below the sink and in pockets elsewhere 
was a deposit of red earth or clay in which were found numerous re- 
mains of Pleistocene animals, apparently representing a distinct unit 
fauna. There also occurred in the cave a younger bed of sand and 
humus containing no extinct mammals but with numerous remains 
of the recent white-tailed deer of the region.” 

The bird bones from these deposits are relatively few in number 
and are all fragmentary. They are light in color, somewhat stained 
by the reddish earth in which they were found, and present the chalky 
appearance usual in bones from limestone caves. 

The 10 species of birds identified offer little worthy of remark as 
all are found in the same area at the present time. The barn owl, 
Tyto alba, regularly inhabits caves. The two vultures, Cathartes and 
Coragyps, sometimes nest in or about caverns. Occurrence of the 
other species must be considered as due to chance except that possibly 
the screech owl and barred owl may have sought shelter in the cave. 

The bird bones are associated with remains of the saber-tooth tiger, 
a capybara, a fossil dog, ground sloth, horse, tapir, a camelid, and 
mastodon among Pleistocene species, together with a number of mam- 
mals that occur at the present time in this area. 

The list of species follows: 


Nyroca affinis Colinus virginianus 
Cathartes aura septentrionalis Meleagris gallopavo 
Coragyps urubu Tyto alba 
Haliaeetus leucocephalus Otus asio 

Falco sparverius Strix varia 


COLUMBIA COUNTY DEPOSITS 


From collections in the Florida State Geological Survey obtained in 
Columbia County about 3 miles northwest of Fort White, Mr. Her- 


Io SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


man Gunter has forwarded a number of bird bones for examination. 
Dr. G. G. Simpson, who has reported on the mammals from this 
deposit, states’ that part of this material was collected by J. Clarence 
Simpson of High Springs, with additional specimens obtained by L. G: 
Getzen and J. D. Lowe. The site is about a mile below the springs 
at the head of the Itchtucknee River. Following these earlier collec- 
tions a considerable number of bird bones were obtained by Mr. and 
Mrs. H. H. Simpson of High Springs during the latter months of 
1930. These latter specimens came to hand as the present report was 
being completed and have added decidedly to information concerning 
birds from this area. The bird bones are brown in color and are 
heavily fossilized. Associated mammalian remains include Hydro- 
choerus, Mylodon, Equus, Tapirus, Mylohyus, Mastodon, and Archi- 
diskidon columbi. 

Notable among the birds are the jabiru, the extinct teal, Querque- 
dula floridana, and the trumpeter swan. Curiously enough the collec- 
tion contains no remains of hawks or vultures. 

Following is the list of identified species : 


Colymbus auritus Querquedula floridana 
Podilymbus podiceps Nyroca valisineria 
Phalacrocorax auritus Nyroca affinis 

Ardea herodias Lophodytes cucullatus 
Nycticorax naevius Pandion haliaétus 
Jabiru mycteria Meleagris gallopavo 
Cygnus buccinator Grus americanus 
Branta canadensis Aramus pictus 
Branta canadensis hutchinsi Rallus elegans 

Anas platyrhynchos Gallinula chloropus 
Anas rubripes Fulica americana 


Anas fulvigula 


VERO AND MELBOURNE DEPOSITS 


The deposits at Vero have attracted the greatest public attention 
because remains of man were found there associated with bones of 
mammals currently considered of Pleistocene age. Among other speci- 
mens secured at this locality were a few bones of birds that were 
described by Shufeldt in 1916." In this account there are listed the 
turkey vulture, Cathartes aura, barn owl, Tyto alba, great blue heron, 
Ardea herodias, and several other species not certainly identified. 
Three forms were described as new, a teal, Querquedula floridana, a 


*Florida State Geol. Surv., 20th Ann. Rep., 1929, p. 270. 
* Florida State Geol. Surv., oth Ann. Rep., 1917, pp. 35-41. 


NO. 2 AVIFAUNA OF PLEISTOCENE IN FLORIDA—WETMORE Tet 


supposed heron, Ardea sellardsi, and a supposed gull, Larus vero, the 
two latter proving invalid (see beyond under Meleagris gallopavo 
and Nyctanassa violacea). 

Two bones forwarded by Mr. Gunter to the present writer include 
the cormorant, Phalacrocorax auritus, and turkey, Meleagris gallo- 
pavo. Mammal remains from this area include Hydrochoerus, Canis 
ayerst, Smilodon floridanus, Felis veronis, Megalonyx jeffersonii, 
Mylodon harlani, Chlamytherium septentrionale, Tatu bellus, three 
species of Equus, Tapirus veroensis, Mylohyus, a camelid, Archi- 
diskidon columbi, and Mastodon americanus. 

Near Melbourne (see pl. 3) in Brevard County, Dr. F. B. Loomis 
of Amherst College located a further deposit which was worked partly 
in cooperation with Dr. J. W. Gidley of the United States National 
Museum. Subsequently Doctor Gidley carried on extensive work in 
this general region during three winters, being assisted by Mr. C. P. 
Singleton of Melbourne. Later Mr. Singleton worked in these exca- 
vations for the Museum of Comparative Zoology. Bird material ob- 
tained has been scattering but has included some important finds. All 
of the specimens secured have been available for the present study. 
The specimens vary in state of preservation, some being heavily 
fossilized and others having a chalky texture. The latter are quite 
fragile. The bones vary from light to dark brown in color. 

The shearwater that comes first on the list may be a species of acci- 
dental occurrence as it ranges regularly at sea or about islands. The 
jabiru and the extinct teal, Querquedula floridana, are represented, 
as are a large and a small form of the gray crane, Grus canadensis, 
These birds accompanied species of mammals of supposed Pleistocene 
age. 

Following is a list of the birds that have been identified : 


Puffinus puffinus Buteo lineatus 
Phalacrocorax auritus Haliaeetus leucocephalus 
Ardea herodias Polyborus cheriway 
Casmerodius albus Colinus virginianus 
Nyctanassa violacea Meleagris gallopavo 

Jabiru mycteria Grus americanus 

Branta canadensis hutchinsi Grus canadensis (large and 
Ouerquedula floridana small forms ) 

Cathartes aura septentrionalis Strix varia 


Buteo jamaicensis 


ACKNOWLEDGMENTS 


In the identification of these specimens the writer is indebted to the 
American Museum of Natural History for the loan of a skeleton of 


I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


the jabiru, and to Dr. Hildegarde Howard, of the Los Angeles Mu- 
seum of History, Science and Art, for bones of Teratornis and 
certain other important material for comparison. 

Mr. W. W. Holmes, in addition to his specimens which he has 
most generously placed in the United States National Museum, has 
supplied much important information both in letters and in personal 
conversations. Ornithologists stand greatly indebted to him for the 
care and effort that he has given to the collection of his material which 
has caused such an increase in our knowledge of the fossil birds of 
this area. Photographs illustrating the Seminole area and Saber- 
tooth Cave were obtained from him. Thanks are due to Mr. J. E. 
Moore for important specimens that he has forwarded for the national 
collections. Dr. Thomas Barbour has kindly forwarded for study 
material collected by C. P. Singleton. Dr. J. W. Gidley has furnished 
data with regard to the deposits on the east coast and has supplied 
certain photographs. Drawings illustrating this report have been 
made by Mr. Sidney Prentice. 


ANNOTATED LIST 


Order COLYMBIFORMES 
Family COLYMBIDAE 
COLYMBUS AURITUS Linnaeus 
Horned grebe 
Colymbus auritus Linnaeus, Syst. Nat., ed. 10, vol. 1, 1758, p. 135. 


In the Holmes collection from near St. Petersburg there are one 
complete and several broken humeri. Part of another humerus is 
contained in collections from the Itchtucknee River, Columbia County, 
in the Florida State Geological Survey. All are similar to those of 
the modern birds. The species is known previously from the Pleisto- 
cene of Fossil Lake, Oregon, and from cavern deposits in Tennessee. 


PODILYMBUS PODICEPS (Linnaeus) 
Pied-billed grebe 
Colymbus podiceps Linnaeus, Syst. Nat., ed. 10, vol. 1, 1758, p. 136. 
In the Holmes collection from the Seminole Field there are limb 
bones of several individuals of this species which are similar to those 
of modern individuals. Other bones are found in the series from the 


Itchtucknee River, Columbia County, in the Florida State Geological 
Survey. This grebe today ranges throughout North and South 


NO. 2 AVIFAUNA OF PLEISTOCENE IN FLORIDA—-WETMORE 13 


America. As it has been recorded in Pleistocene beds in Oregon and 
California (McKittrick) its presence in deposits of that age in Florida 
indicates a similar wide distribution for North America during the 
ice Ace: 


Order PROCELLARITFORMES 
Family PROCELLARIIDAE 
PUFFINUS PUFFINUS (Briinnich) 
Manx shearwater 
Procellaria puffinus Brtinnich, Orn. Bor., 1764, p. 29. 


A left metacarpal with the fourth metacarpus missing was secured 
by J. W. Gidley near Melbourne, March 18, 1929. This shearwater 
as a species now ranges from Norway south into the Mediterranean, 
breeding in Iceland, the Azores and other islands, and at least casually 
on Bermuda. At the present time it occurs rarely along the coasts of 
North America. It has not been recorded previously as fossil nor has 
it been known before from Florida. 


Order PELECANIFORMES 
Family PHALACROCORACIDAE 
PHALACROCORAX AURITUS (Lesson) 
Double-crested cormorant 
Carbo auritus Lesson, Traité Orn., 1831, p. 605. 


Cormorants of this type apparently were as widely distributed in 
Florida during the Pleistocene as they are today, for in the collections 
here under review there are found the lower end of a tibio-tarsus and 
part of an ulna from stratum Number Two at Melbourne secured by 
J. W. Gidley, part of an ulna from Hog Creek, near Sarasota, obtained 
by J. E. Moore in 1928, and a sacrum and three fragments of humeri 
from the Holmes collections in the Seminole Field. In the collections 
of the Florida State Geological Survey there are a complete humerus, 
part of an ulna and other bones from the Itchtucknee River, Columbia 
County, another humerus, white in color, from Rock Springs in 
Orange County that is very doubtfully Pleistocene in age, and still 
another humerus from the north bank of the canal west of the rail- 
road bridge at Vero. 

The resident cormorant of this group found now in Florida, Phala- 
crocorax auritus foridanus, is smaller than the bird from farther north 
and west, Phalacrocorax auritus auritus, which comes to Florida as 


14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


a migrant during the winter season. It is interesting to observe that 
the distal ends of two humeri in the Holmes collection from the west 
coast differ decidedly in size, the transverse breadth across the troch- 
leae in one being 15.8 mm. and in the other 18.2 mm., thus exhibiting 
differences similar to those that mark the larger and smaller modern 
races. It appears possible that differentiation between these two f orms 
may have occurred in the Pleistocene, though on the other hand these 
two specimens may be merely extremes of individual variation existing 
at that time. On this scanty material the writer does not venture to 
identify the two as belonging certainly to distinct subspecies. 


Family ANHINGIDAE 
ANHINGA ANHINGA (Linnaeus) 
Snake-bird, water-turkey 
Plotus anhinga Linnaeus, Syst. Nat., ed. 12, vol. 1, 1766, p. 218. 


The snake-bird is represented by the distal end of a left humerus 
collected in the Number Two bed near Melbourne by Doctor Gidley 
on May 3, 1929. 

This species has not been recorded previously as a fossil. 


Family ARDEIDAE 


ARDEA HERODIAS Linnaeus 
Great blue heron 
Ardea herodias Linnaeus, Syst. Nat., ed. 10, vol. 1, 1758, p. 143. 


The genus Ardea is represented by four cervical vertebrae and 
two fragmentary metatarsi from Melbourne, collected by Gidley; by 
the lower end of a metatarsus and the upper part of a coracoid from 
the Seminole Field collected by Holmes; and by the lower end of a 
tibio-tarsus from Bradenton collected by J. E. Moore. The upper 
and lower ends of metatarsi and a broken tibio-tarsus are included in 
collections in the Florida State Geological Survey from the Itchtucknee 
River, Columbia County. All are referred here to the species herodias 
without consideration of the possible occurrence of the great white 
heron, Ardea occidentalis, confined today to southern Florida, since 
so far as present information goes these two supposed species are 
indistinguishable in their skeletons. The two specimens from the 
Seminole Field are larger than any modern bird seen, suggesting that 
possibly there was a larger heron of this type in existence in the 
Pleistocene. The differences are shown in the following measure- 


NO. 2 AVIFAUNA OF PLEISTOCENE IN FLORIDA—WETMORE 15 


ments: Modern Ardea herodias (seven specimens, including two from 
Florida) ; metatarsus, transverse breadth of trochleae 16.2-17.3 mm., 
smallest transverse breadth of shaft 5.9-7.0 mm.; coracoid, trans- 
verse breadth of head 13.8-15.7 mm. Fossils from Seminole Field: 
metatarsus, transverse breadth of trochleae 18.4 mm., smallest trans- 
verse breadth of shaft 8.2 mm.; coracoid, transverse breadth of head 
17.3 mm. The material is considered too fragmentary for further 
consideration at this time. 


CASMERODIUS ALBUS (Linnaeus) 
Egret 
Ardea alba Linnaeus, Syst. Nat., ed. 10, vol. 1, 1758, p. 144. 


The lower end of the left tibio-tarsus of a young individual is found 
in the collections made by W. W. Holmes in the Seminole Field. 
Another was identified in material collected near Venice by J. E. 
Moore. An ulna of an individual of large size was included in col- 
lections made near Melbourne by C. P. Singleton, in 1928, for the 
Museum of Comparative Zoology. 

This species has not been recorded previously as a fossil, the only 
other reference to its possible occurrence in the Pleistocene being that 
of Shufeldt,’ where a fragmentary metatarsus is listed as possibly 
from this species. 


EGRETTA THULA (Molina) 
Snowy heron 
Ardea thula Molina, Sagg. Stor. Nat. Chili, 1782, p. 235. 


A partly complete metatarsus collected at Bradenton by J. Ek. Moore 
is the first record of this species as a fossil. The specimen comes from 
a small individual. The snowy heron is fairly common in Florida at 
the present. time, and formerly existed there in large numbers. It 
has not been recorded previously as a fossil. 


HYDRANASSA TRICOLOR (Miiller) 
Louisiana heron 
Ardea tricolor Muller, Vollst. Naturs. Suppl., 1776, p. 111. 


The lower end of a right metatarsus comes from the Seminole Field 
near St. Petersburg. 
The present species is here first recorded as a fossil. 


* Florida Geol. Surv., Ninth Ann. Rep., 1917, pp. 40-41. 
2 


16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


FLORIDA CAERULEA (Linnaeus) 
Little blue heron 
Ardea caerulea Linnaeus, Syst. Nat., ed. 10, vol. 1, 1758, p. 143. 


This species, common in modern Florida, is represented by the 
lower end of a right tibio-tarsus and the distal portion of a right femur 
secured by W. W. Holmes in the Seminole Field. 

This heron is here first reported certainly in fossil deposits. 


BUTORIDES VIRESCENS (Linnaeus) 
Little green heron 
Ardea virescens Linnaeus, Syst. Nat., ed. 10, vol. 1, 1758, p. 144. 


A right metatarsus and the lower end of a left tibio-tarsus were 
collected in the Seminole Field deposits by W. W. Holmes. The 
metatarsus measures 51.7 mm. in length. 

The green heron is here found fossil for the first time. 


NYCTICORAX NYCTICORAX (Linnaeus) 
Black-crowned night heron 
Ardea nycticorax Linnaeus, Syst. Nat., ed. 10, vol. 1, 1758, p. 142. 


The lower end of a tibio-tarsus was obtained by J. E. Moore, 
at Bradenton, and a similar specimen was collected at the Itchtucknee 
deposit by Mr. and Mrs. H. H. Simpson. This heron, which is com- 
mon in Florida now, has been found previously in the Pleistocene 
of Rancho La Brea in California. 


NYCTANASSA VIOLACEA (Linnaeus) 
Yellow-crowned night heron 


Ardea violacea Linnaeus, Sys. Nat., ed. 10, vol. 1, 1758, p. 143. 
Larus vero Shufeldt, Journ. Geol., vol. 25, Jan.-Feb. (Jan.), 1917, p. 18; 
Florida State Geol. Surv., Ninth Ann. Rep., 1917, p. 40, pl. 2, fig. 21. 


The proximal ends of two left coracoids represent this heron in 
the material obtained by W. W. Holmes in the Seminole Field. 

The type of Larus vero Shufeldt, a left metacarpal secured at Vero, 
Florida (U.S. Nat. Mus. Div. Vert. Pal. No. 8832), on examination 
proves to be the yellow-crowned night heron. This species has not 
* been recorded before in the Pleistocene. 


NO. 2 AVIFAUNA OF PLEISTOCENE IN FLORIDA—-WETMORE 17 


BOTAURUS LENTIGINOSUS (Montagu) 
American bittern 
Ardea lentiginosa Montagu, Suppl. Orn. Dict., 1813, text and plate. 


arts of two humeri were obtained by W. W. Holmes in the 
Seminole area, and of another by J. E. Moore on Hog Creek near 
Sarasota, Florida. This inhabitant of marshes is widely distributed 
in Florida at the present time. 


Family CICONIIDAE 
JABIRU MYCTERIA (Lichtenstein) 
Jabiru 
Ciconta mycteria Lichtenstein, Abhandl. Kon. Akad. Wiss. Berlin (Phys. 
Klass.), for 1816-1817, 1819, p. 163. 
Jabiru? weillsi Sellards, Florida State Geol. Sury., 8th Ann. Rep., 1916, p. 146; 
pl. 26, figs. I-4, text-fig. 15. 

Apparently the great jabiru stork was common in Florida during 
the Pleistocene as it is represented in the present collections by many 
fragments of bones from a number of localities. In the Seminole 
Field near St. Petersburg W. W. Holmes obtained a number of frag- 
mentary specimens, including parts of the tibio-tarsus, coracoid, 
scapula, ulna, and metacarpus. Most of these are well fossilized 
though one fragment appears quite modern. A fragment from the 
head of a tibio-tarsus was obtained by J. E. Moore near Venice. A 
perfect metacarpal, a coracoid, and part of a metatarsus are found 
in collections from the Itchtucknee River, Columbia County, in the 
Florida State Geological Survey. At Melbourne in the excavations 
on the golf links J. W. Gidley secured the lower end of a right 
metatarsus, and parts of an ulna and a metacarpus from the Number 
Two stratum. In the collection made at Melbourne by C. P. Singleton 
for the Museum of Comparative Zoology there are parts of three 
right and one left tibio-tarsi and both extremities of a right meta- 
tarsus with the central part of the shaft gone. 

After careful comparison of the type»specimen of. Jabiru weillsi, 
a right humerus obtained at Vero, Florida, there is nothing evident to 
separate it from the modern Jabiru mycteria. It was differentiated 
in the original description principally on larger size, but, though large, 
it 1s equalled by modern birds in dimension, and is similar to them 
in its conformation. The original description gives the total length 
of the type humerus as 280 mm. Since then the bone has been broken 
and restored, in this process being lengthened until now it is 293 mm. 


18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


long, the extra length being obviously due to crushing of the shaft 
and the separation of these parts. In a previous publication* the 
present writer indicated that sellardsi was probably equivalent to 
modern mycteria, a belief that is now substantiated. 
The fossil material indicated above shows the same range in in- 
dividual variation in size as is found in the modern material examined. 
The jabiru is also known from the Pleistocene of Cuba.’ 


Family THRESKIORNITHIDAE 
PLEGADIS sp. 
Glossy ibis 
The lower end of a left tibio-tarsus from the Seminole Field col- 
lected by W. W. Holmes is from a juvenile bird and shows little 
evidence of fossilization. It is not practicable to determine whether 
it represents Plegadis falcinellus or P. guarauna, both of which occur 
in Florida. 
GUARA ALBA (Linnaeus) 
White ibis 
Scolopax alba Linnaeus, Syst. Nat., ed. 10, vol. 1, 1758, p. 145. 
The distal ends of right and left humeri, of two right ulnae, and 
the lower end of a left tibio-tarsus were collected by W. W. Holmes 
in the Seminole Field. The white ibis is locally common in Florida 


at the present time. 
This ibis is here first reported as a fossil. 


Order ANSERIFORMES 
Family ANATIDAE 
CYGNUS COLUMBIANUS (Ord) 
Whistling swan 

Anas columbianus Ord, in Guthrie’s Geogr., 2d Amer. ed., 1815, p. 319. 

The proximal ends of right and left coracoids were obtained by 
W. W. Holmes in collecting in the Seminole Field. In modern times 
this swan is found in winter occasionally in Florida, mainly along the 
Gulf Coast, its principal winter range in eastern North America being 
farther north. 

The whistling swan has been known previously as a fossil only from 
the Pleistocene beds at Fossil Lake, Oregon. 


1 Amer. Mus. Nov., No. 301, Feb. 29, 1928, pp. 2-3. 


NO. 2 AVIFAUNA OF PLEISTOCENE IN FLORIDA—WETMORE IQ 


CYGNUS BUCCINATOR Richardson 


Trumpeter swan 
Cygnus buccinator Richardson, Faun. Bor.-Amer., vol. 2, 1831, (1832), p. 464. 


One entire and three fragmentary humeri, a metacarpal, a coracoid, 
and a tibio-tarsus are included in the Florida State Geological Survey 
collections from near the head of the Itchtucknee River in Columbia 
County (Catalog nos. V-4576; V-4589; V-4599; V-4598, 2 specimens ; 
V-4599 and V-4826). These bones are in an excellent state of preser- 
vation, part being dark and part light in color. They agree perfectly 
with the modern bird, and are larger and stronger than the correspond- 
ing bones in the whistling swan. This fine bird, known previously from 
the Pleistocene of Fossil Lake, Oregon, formerly ranged widely 
through interior and western North America but at the present time 
is at so low an ebb of abundance as to be nearly extinct. It wintered 
formerly from southern Indiana and southern Illinois to Texas but 
has not been reported previously from Florida. 


BRANTA CANADENSIS (Linnaeus) 


Canada goose 
Anas canadensis Linnaeus, Syst. Nat., ed. 10, vol. 1, 1758, p. 123. 


From the Seminole Field this goose, a common species on the 
northern part of the Gulf Coast of modern Florida, is represented by 
the lower end of a left tibio-tarsus and the distal ends of right and 
left ulnae. Two ulnae and a broken metacarpal are found in collec- 
tions in the Florida State Geological Survey from near the head of 
the Itchtucknee River in Columbia County. 


BRANTA CANADENSIS HUTCHINSI (Richardson) 


Hutchins’ goose 
Anser hutchinsti Richardson, Faun. Bor.-Amer., vol. 2, 1831 (1832), p. 470. 


In the Holmes collection from the Seminole Field near St. Peters- 
burg there is a distal end of a left ulna of the Branta type that is a 
counterpart of B. canadensis except for its smaller size. Part of an 
ulna comes from the Itchtucknee River, and material secured near 
Melbourne for the Museum of Comparative Zoology by C. P. Single- 
ton includes a right humerus that also has the same characters. These 
are identified as from the Hutchins’ goose, a species rarely recorded 
from modern Florida. Current custom in recognizing this bird as a 
subspecies of the Canada goose is here followed though some doubt 
may be expressed as to whether the two are not specifically distinct. 


20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


This form is here identified certainly for the first time in a fossil state, 
the only previous records being open to question. Shufeldt* has re- 
corded it uncertainly from the Pleistocene of Fossil Lake, Oregon, 
listing it in his final table with a query.” 


ANAS PLATYRHYNCHOS Linnaeus 
Mallard 


Anas platyrhynchos Linnaeus, Syst. Nat., ed. 10, vol. 1, 1758, p. 125. 


Two humeri of the mallard, strong and robust bones, were collected 
in the Itchtucknee River deposits in Columbia County by Mr. and 
Mrs. H. H. Simpson. 

Remains of other ducks of the mallard type from the Seminole 
Field, collected by W. W. Holmes, include parts of humeri, an ulna, 
a metacarpal, and parts of several coracoids, all in such fragmentary 
form that it is not expedient to attempt to identify them specifically. 
Parts of two humeri of similar status were obtained by J. E. Moore 
near Venice. Probably the black duck and mallard are both repre- 
sented. 

The mallard is a regular migrant to Florida. 


ANAS RUBRIPES Brewster 
Black duck 


Anas obscura rubripes Brewster, Auk, 1902, p. 184. 


Collections from the Itchtucknee River, Columbia County, made by 
Mr. and Mrs. H. H. Simpson, include a humerus and a metatarsus of 
this species, which has not been recorded previously as a fossil. 

Like the mallard the black duck comes regularly to winter in Florida. 


ANAS FULVIGULA Ridgway 
Florida duck 


Anas obscura var. fulvigula Ridgway, Amer. Nat., vol. 8, February, 1874, 
Dako 
A metacarpal and the proximal and distal ends of two humeri are 
equal in size to a female of this species and are identified as this bird. 
These specimens come from the Holmes collection from the Seminole 
Iield. In material obtained by J. E. Moore at Bradenton there is part 
of another humerus. 





*Bull. Amer. Mus. Nat. Hist., vol. 32, July 9, 1913, pp. 147, 156, pl. 33, fig. 
414. 

* See also Shufeldt, Auk, 1913, p. 30, and Science, vol. 37, February 21, 1013, 
p. 307, where this same record is given as Branta canadensis hutchinsi (?). 


NO. 2 AVIFAUNA OF PLEISTOCENE IN FLORIDA—WETMORE 21 


Collections in the Florida State Geological Survey from near the 
head of the Itchtucknee River, Columbia County, contain two entire 
and four broken humeri, a coracoid, and two radii (the last being 
identified tentatively on basis of agreement in size and contour). 
Apparently the species was common in that area. These represent the 
first records for the Florida duck in a fossil state. This species breeds 
abundantly in Florida at the present time. 


NETTION CAROLINENSE (Gmelin) 
Green-winged teal 

Anas carolinensis Gmelin, Syst. Nat., vol. 1, pt. 2, 1780, p. 533. 

In material collected by W. W. Holmes in the Seminole Field there 
are parts of two right and two left humeri, of right and left ulnae, and 
a left coracoid. The humerus in this species is distinctly shorter than 
in the blue-winged teal. 

This duck has been reported previously from the Pleistocene of 
Oregon, and from several localities in California. 


QUERQUEDULA FLORIDANA Shufeldt 


Querquedula floridana Shufeldt, Florida State Geol. Surv., Ninth Ann. Rep., 
1917, p. 36, pl. 1, fig. 4, pl. II, fig. 25. 

The type specimen of this teal (figs. 2-3) was collected in stratum 

Number Two at Vero, Florida, and is now in the collections of the 












Tics. 2-3.—Type of Querquedula floridana 
Shufeldt (natural size). 


22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


United States National Museum. While closely similar to the humerus 
of the living blue-winged teal, Querquedula discors, the fossil is 
heavier and stronger throughout both in the shaft and in the proximal 
and distal ends. It thus bears out the characters assigned to it in the 
original description. 

In the Holmes collection from the Seminole Field there 1s one right 
and one left humerus nearly complete, and the fragments of three or 
more others that correspond very closely to the type specimen. With 
them are three broken metacarpals. A portion of a humerus was 
secured by J. E. Moore near Venice. The proximal half of a right 
humerus was collected by C. P. Singleton at Melbourne for the Mu- 
seum of Comparative Zoology. There are also a number of bones 
from the Itchtucknee River deposits, Columbia County, in the collec- 
tions of the Florida State Geological Survey. 

Following are measurements of the four complete humeri at hand. 

Type, Vero, Florida, total length 65.8, transverse diameter through 
trochleae 9.9, transverse diameter through head 16.1, transverse di- 
ameter of shaft at center 5.4 nim. 

Two specimens, Seminole Field, total length 65.7-70.2, transverse 
diameter through trochleae 10.3-10.5, transverse diameter through 
head 15.3-15.6, transverse diameter of shaft at center 5.I-5.3 mm. 

Three modern Querquedula discors, two males and one female, 
measure as follows: total length 65.4, 65.9, 61.8, transverse breadth 
through trochleae 9.7, 10.0, 9.2, transverse breadth through head 13.8, 
14.0, 13.1, transverse diameter of shaft at center 4.7, 4.8, 4.6 mm. 


NYROCA VALISINERIA (Wilson) 
Canvasback 
Anas valisineria Wilson, Amer. Orn., vol. 8, 1814, p. 103, pl. 70, fig. 5. 


A complete ulna is found in collections made by Mr. and Mrs. H. H. 
Simpson on the Itchtucknee River in Columbia County. 
This species in Florida is a winter migrant from the north. 


NYROCA AFFINIS (Eyton) 
Lesser scaup duck 
Fuligula affinis Eyton, Monogr. Anatidae, 1838, p. 157. 


A left humerus collected in the Number Two bed at Melbourne by 
J. W. Gidley in 1926, with a right metatarsus in the Holmes collection 
from the Seminole Field, part of an ulna obtained near Venice by 
J. E. Moore, and four complete and one fragmentary humeri, two 


NO; 2 AVIFAUNA OF PLEISTOCENE IN FLORIDA—-WETMORE 23 


ulnae, a metacarpal, and a tibio-tarsus from the head of the Itchtucknee 
River, Columbia County, constitute definite record of this duck. Some 
miscellaneous fragments from near St. Petersburg belong probably 
to this species but cannot be certainly identified. There is also a 
complete ulna from Saber-tooth Cave near Lecanto, collected in 1928 
by W. W. Holmes. These form the only certain Pleistocene records 
for this duck which is found in abundance in Florida during winter 
at the present time. 


ERISMATURA JAMAICENSIS (Gmelin) 
Ruddy duck 
Anas jamaicensis Gmelin, Syst. Nat., vol. 1, pt. 2, 1780, p. 510. 


The ruddy duck is represented by a metacarpal collected near 
Venice by J. E. Moore. This species is a common visitor to Florida 
during the winter. 


LOPHODYTES CUCULLATUS (Linnaeus) 
Hooded merganser 
Mergus cucullatus Linnaeus, Syst. Nat., ed. 10, vol. 1, 1758, p. 120. 


A humerus was obtained on the Itchtucknee River, Columbia 
County, by Mr. and Mrs. H. H. Simpson. 


Order FALCONIFORMES 
Family CATHARTIDAE 
CATHARTES AURA AURA (Linnaeus) 
Mexican turkey vulture 
Vultur aura Linnaeus, Syst. Nat., ed. 10, vol. 1, 1758, p. 86. 


In the Holmes collection from the Seminole Field there is the distal 
end of a right tibio-tarsus and the shaft of a left coracoid of a turkey 
vulture that are decidedly smaller than these bones in the modern 
bird of the United States, but that agree exactly in dimension with a 
specimen of the Mexican form from Matamoras. They are identified 
as typical aura which is here first recorded from within the limits of 
the United States, its modern range being from Mexico south to 
Panama and Columbia, including Cuba and Jamaica. The transverse 
breadth of the distal end of the tibio-tarsus in the Pleistocene speci- 
men is 11.0 mm. The modern specimen of aura from Matamoras 
(U. S. N. M. 1442) is exactly similar. In a series of eight modern 
birds of septentrionalis from Florida, Virginia, Maryland, and Penn- 


24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


sylvania this measurement ranges from 12.2 to 13.1 mm. with an 
average of 12.6 mm. The fossil bone in form is exactly like the 
modern aura. 

The existence of remains of two evidently distinct races of the 
turkey vulture in the Pleistocene deposits of Florida, of which one is 
now northern and the other southern in range is puzzling. Since the 
two forms are found in different deposits, septentrionalis coming 
from Vero and Melbourne on the east coast and Lecanto in the central 
part of the state, and aura from near St. Petersburg, if it is assumed 
that the record of aura is not due to a stray individual out of its 
normal range, it seems probable that the bone deposits in question 
were formed at different periods during the Ice Age. 


CATHARTES AURA SEPTENTRIONALIS Wied 
Turkey vulture 
Cathartes septentrionalis Wied, Reis. Nord-Amer., vol. 1, 1830, p. 162. 


The distal part of a left ulna obtained near Melbourne by C. P. 
Singleton in 1928 for the Museum of Comparative Zoology, and a 
fragment of a left metacarpal collected by W. W. Holmes in Saber- 
tooth Cave at Lecanto, Florida, are similar in size to the turkey 
vulture found today in Florida. Shufeldt* has reported this bird from 
Vero, Florida, and from his figured specimen it is evident that the 
large northern bird was, the one represented. 

This form ranges today throughout the greater part of the United 
States, being absent only in the north, and has had extended range for 
a long period of time, since it is known from the Pleistocene deposits 
of California. 


CORAGYPS URUBU (Vieillot) 
Black vulture 
Vultur urubu Vieillot, Ois. Amer. Sept., vol. 1, 1807, p. 23, pl. 2. 


Numerous fragments of bone from the Seminole Field include 
parts of metatarsus, tibio-tarsus, coracoid, humerus, and metacarpal, 
while from Saber-tooth Cave at Lecanto there are two bones, the 
distal part of a tibio-tarsus and the upper portion of a metatarsus, the 
latter from a juvenile individual. All this material was collected by 
W. W. Holmes. These remains are similar in size and form to those 
of modern individuals. 


‘Journ. Geol., 1917, p. 18; Florida State Geol. Surv., Ninth Ann. Rep.. 
TOL, p. 36, pl. 1, fie. 2: 


NO. 2 AVIFAUNA OF PLEISTOCENE IN FLORIDA—WETMORE 25 


The greater abundance of remains of the black vulture over those 
of the turkey vulture in these deposits is worthy of comment as indi- 
cating possibly the relative abundance of these two birds during the 
Pleistocene. The black vulture is first known as a fossil from this 
area. 


GYMNOGYPS CALIFORNIANUS (Shaw) 
California condor 
Vultur californianus Shaw, Nat. Misc., vol. 9, 1797, pl. 301. 


In the material secured on Hog Creek, near Sarasota, Florida, by 
J. E. Moore in 1928, there is part of the distal end of a left meta- 
tarsus (see pl. 4, and figs. 4-5) that agrees so exactly in form and 





Fics. 4-5.—Fragmentary metatarsus of Cali- 
fornia Condor (Gymnogyps californianus) from 
near Sarasota (natural size). 


dimension with two modern specimens of the California condor that 
there is no hesitancy in identifying it as that species. It may be 
remarked that Gymnogyps has the middle trochlea of the metatarsus 
decidedly smaller than the South American condor ultur, this serving 
to distinguish the metatarsus in these two genera without difficulty. 

In collecting in the Seminole area W. W. Holmes obtained a bit 
of a right humerus comprising the ulnar trochlea with the adjacent 
external parts, and the distal end of a right radius that are identified 
as remains of this species. 

The previously known range of the California condor has been 
entirely western as it has been found living in the coast ranges of 
California from Santa Clara County south into northern Lower Cali- 
fornia, ranging in earlier days north to the Columbia River. Though 


26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


it has been reported casually east to Arizona, according to Swarth* 
these records lack confirmation. Miller * has identified an ulna from 
cave deposits of uncertain age near Las Vegas, Nevada. The same 
author * in writing of the occurrence of this species in the Pleistocene 
asphalt beds of Rancho La Brea at Los Angeles, California, says that 
it is the most common of the American vultures in these deposits, its 
remains occurring in almost incredible abundance. With large series 
of Pleistocene material for examination he found remarkable uni- 
formity when compared with bones from modern specimens. 

The report of this species from the Pleistocene of Florida is the 
first note of the occurrence of a condor-like bird in eastern North 
America and gives an entirely unexpected extension of range for 
this species during the Ice Age. Discovery of further remains will be 
awaited with interest. 


Family TERATORNITHIDAE 
TERATORNIS MERRIAMI L. H. Miller 
Teratornis 


Teratornis merriamt L. H. Miller, Univ. California Publ. Geol., vol. 5, Sep- 
tember 10, 1900, p. 307, figs. I-09. 

Among fossils collected by W. W. Holmes in the Seminole area 
there were found a number of small fragments of bones from what 
was evidently a very large form of bird. After some study it was 
clear that these were from some large vulture so that on prelimi- 
nary examination they were placed among remains assigned to the 
condors. Critical study indicated certain puzzling differences from 
both the California and the South American condors and it was with 
much surprise that they were found to come from the great Tera- 
tornis known previously only from California where its remains have 
been found in the asphalt deposits at Rancho La Brea, McKittrick 
and Carpinteria. 

As stated above the remains from the Holmes collection are all 
highly fragmentary. The distal end of a left ulna (figs. 9-11), one 
of the most characteristic bits, agrees minutely in its somewhat intri- 
cate details with two specimens of Teratornis from California. Two 
bits of humeri include the articular surface of the head and the 
radial trochlea of a left humerus. There are further the distal ends 

* Pac. Coast Avif., No. 10, May 25, 1914, p. 83. 

* Condor, 1931, p. 32. 

“Carnegie Inst., Washington, Publ. No. 349, August, 1925, p. 81. 


NO. 2 AVIFAUNA OF PLEISTOCENE IN: FLORIDA—WETMORE 


bo 
Sy 


of left and right radii and the lateral facets from the head of a left 
coracoid. These likewise in size and detail are like the corresponding 
parts in Teratornis. In fact the agreement is so close that there is 
no basis for differentiating the Florida bird from that of California. 

















Fics. 6-8.—Metatarsus of Teratornis merriami from Bradenton 
(natural size). 


With the material described above at hand it has been highly grati- 
fying to find in specimens collected by J. 2. Moore at Bradenton a 
nearly complete metatarsus (pl. 5, and figs. 6-8) and a broken femur 
(fig. 12) that likewise agree in close detail with the bird of California. 






an ANI SS 





9 10 11 


Fics. 9-11.—Distal end of ulna of Teratornis merriami from the Seminole 
area (natural size). 





Fic. 12——Femur of Teratornis merriami from 
Bradenton (natural size). 


28 


NO. 2. AVIFAUNA OF PLEISTOCENE IN FLORIDA—WETMORE 29 
The femur was in a fragile condition and was received in several 
pieces. These have been so skillfully joined by N. H. Boss that they 
illustrate well the form of the bone. 

The identification of this form from Florida is one of the most 
unexpected results of this study of the Pleistocene from Florida and 
indicates a wide range in Pleistocene times for this peculiar bird. 


Bamily, ACGUIPT TRIDAI 
BUTEO JAMAICENSIS (Gmelin) 
Red-tailed hawk 
Falco jamaicensis Gmelin, Syst. Nat., vol. 1, pt. 1, 1788, p. 266. 


The red-tailed hawk is represented in the W. W. Holmes collection 
from the Seminole Field by the distal end of a left metatarsus, the 
lower ends of two left tibio-tarsi, and a left coracoid. J. IX. Moore 
secured part of an ulna near Venice. Gidley collected the lower por- 
tion of a left humerus in the lower part of the Number Two bed on 
the golf links at Melbourne. The species occurs today in Florida, and 
has been recorded previously as fossil in the Pleistocene of California. 

The red-tailed hawk has been known for many years as Buteo 
borealis, the original reference being Falco borealis Gmelin, Syst. 
Nat., vol. 1, pt. 1, 1788, p. 266, where it is species No. 75. The pre- 
ceding species, No. 74, Falco jamaicensis on the same page is based 
on the cream-colored buzzard of Latham’ described from a specimen 
from Jamaica, evidently an immature of the red-tailed hawk. As the 
name jamaicensis comes first on the page in question in Gmelin’s 
work it will replace the familiar borcalis as the specific name for 
this hawk. 

BUTEO LINEATUS (Gmelin) 
Red-shouldered hawk 


Falco lineatus Gmelin, Syst. Nat., vol. 1, pt. 1, 1788, p. 268. 


The red-shouldered hawk was apparently as common in Pleistocene 
times as today, for it is represented by a number of fragmentary 
bones. Holmes obtained a left humerus lacking the head and the 
distal ends of two tibio-tarsi from the Seminole area. J. E. Moore 
secured part of a metatarsus near Venice. In excavations on the golf 
links at Melbourne Gidley secured a nearly complete left metatarsus 
in 1926, a fragment of another in 1928, and a broken left femur 
in 1930. The red-shouldered hawk is represented in modern Florida 
by a resident form Buteo lineatus alleni that besides differing in color 


* Gen: Syn. Birds, vol. 1, pt. 1, 1781, p. 40: 


30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


from the northern race is slightly smaller, and by a migrant form 
Buteo lineatus lineatus that is present in winter and is slightly larger. 
The specimens from the west coast are from slightly smaller birds 
than those from Melbourne, suggesting that possibly two forms are 
presented in the fossil material. This cannot be definitely decided 
from the bones now at hand. 

The red-shouldered hawk is here first recorded in fossil form. 


BUTEO PLATYPTERUS (Vieillot) 
Broad-winged hawk 
Sparvius platypterus Vieillot, Tabl. Encycl. Meéth., vol. 3, 1823, p. 1273. 


In the Seminole Field in Pinellas County, W. W. Holmes obtained 
the distal end of a left humerus, and the proximal part of a left meta- 
carpus. This species has not been recorded previously as a fossil. 

The broad-winged hawk today is a winter visitor to Florida ar- 
riving from the north in October and departing in March. 


GERANOAETUS sp. 
Eagle 


The distal ends of three ulnae obtained by W. W. Holmes in the 
Seminole Field come from an eagle of this genus, which was repre- 
sented in the Pleistocene of California by two species G. fragilis and 
G. grinnelli and of which there is one living species G. melanoleucus 
in South America. The bones from Florida agree in size with the 
latter. The material is considered too fragmentary to warrant specific 
determination at the present time. The genus is here first recorded 
from eastern North America. 


HALIAEETUS LEUCOCEPHALUS (Linnaeus) 
Bald eagle 
Falco leucocephalus Linnaeus, Syst. Nat., ed. 12, vol. 1, 1766, p. 124. 


The bald eagle is represented by fragments from the Seminole 
Field, and by two broken radii from Saber-tooth Cave, near Lecanto, 
collected by W. W. Holmes, as well as by part of an ulna collected 
near Venice by J. E. Moore and a number of bones from near Mel- 
bourne, obtained by Gidley and Singleton. The collection made by 
Singleton for the Museum of Comparative Zodlogy contains part of 
a metacarpal. Several of the Melbourne specimens are practically 
complete, and show no differences from the modern bird which is 
common at present in Florida. 


NO. 2 AVIFAUNA OF PLEISTOCENE IN FLORIDA—WETMORE 31 


PANDION HALIAETUS (Linnaeus) 
Osprey 
Falco haliaétus Linnaeus, Syst. Nat., ed. 10, vol. 1, 1758, p. Ol. 


A nearly complete left metatarsus was obtained by J. W. Gidley on 
April 5, 1929, in the Number Two bed at Melbourne where it was 
associated with remains of a peculiar extinct bear, Arctodus flori- 
danus. In 1930 a femur was secured in the Itchtucknee deposits by 
Mr. and Mrs. H. H. Simpson. These are the first reports of the 
osprey in the Pleistocene of our continent. 


Family FALCONIDAE 
POLYBORUS CHERIWAY (Jacquin) 
Audubon’s caracara 
Falco (cheriway) Jacquin, Beytr. Gesch. Vogel, 1784, p. 17, pl. 4. 


On the golf course near Melbourne, Gidley collected an ulna that is 
identical with that of modern birds. On the west coast W. W. 
Holmes obtained numerous remains in the Seminole Field, these in- 
cluding parts of humeri, ulnae, a femur, a tibio-tarsus, and several 
metatarsi. Two fragmentary humeri and the lower end of a tibio- 
tarsus are similar in size to small modern specimens of the caracara 
from Florida. Parts of four metatarsi agree in having the distal troch- 
leae distinctly smaller than in any of the three modern birds seen. 
Ulnae and part of a femur also seem smaller than usual. It will be 
recalled that two subspecies of this caracara are now recognized, 
Polyborus cheriway cheriway of northern South America, and Poly- 
borus cheriway auduboni of Florida and the southwestern part of the 
United States south into Mexico, the former being smaller in size. 
The smaller fossil bones here under discussion seem to show approach 
to the modern race of South America. 

Remains of the caracara are common in the Pleistocene deposits 
of California but are here reported for the first time outside that State. 
In Florida the species at the present time is peculiar to the prairies of 
the Okeechobee and Kissimmee regions, where it is locally common. 


FALCO SPARVERIUS Linnaeus 
Sparrow hawk 
Falco sparverius Linnaeus, Syst. Nat., ed. 10, vol. 1, 1758, p. 90. 
Material collected in Saber-tooth Cave at Lecanto, Florida, in 1928 
by W. W. Holmes includes parts of right and left tibio-tarsi of this 


species, a common bird in this area at the present time. 
3 


32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


Order GALLIFORMES 
Family PERDICIDAE 
COLINUS VIRGINIANUS (Linnaeus) 
Bob-white 
Tetrao virginianus Linnaeus, Syst. Nat., ed. 10, vol. 1, 1758, p. 161. 


In material collected by W. W. Holmes in the Seminole area re- 
mains of the quail may be said to be common since this species is 
represented by six humeri more or less complete and part of an ulna. 
It is likewise common in the collection from Saber-tooth Cave near 
Lecanto where two humeri, a metatarsus and two femora were ob- 
tained. In excavations on the golf links at Melbourne in February, 
1928, Gidley secured two humeri at the line of contact between _ 
stratum Number One and stratum Number Two. 

These bones all appear similar to those of modern quail. The 
species is abundant in Florida, and has been previously reported as a 
fossil from Pleistocene cavern deposits in Tennessee. 


Family MELEAGRIDIDAE 
MELEAGRIS GALLOPAVO Linnaeus 
Turkey 


Meleagris gallopavo Linnaeus, Syst. Nat., ed. 10, vol. 1, 1758, p. 156. 
Ardea sellardsi Shufeldt, Journ. Geol., Jan—Feb. (publ. Jan.), 1917, p. 109. 


In the series of bird bones obtained in the Seminole area, Pinellas 
County, by W. W. Holmes, remains of the wild turkey far out- 
number those of any other species represented ; they include 98 frag- 
ments of metatarsi, tibio-tarsi, femora, coracoids, humeri, ulnae, and 
metacarpi. Most of these have been broken into small bits. The speci- 
mens exhibit the usual variation in size found in series of wild turkey 
bones, and do not differ from modern specimens. In the material 
collected on Hog Creek, near Sarasota, by J. E. Moore in 1928 there 
is included parts of a femur and a tibio-tarsus, the latter considerably 
worn. The proximal end of a metatarsus has been forwarded by 
Mr. Moore as taken at Bradenton. In Saber-tooth Cave near Lecanto 
Holmes obtained a single spur core from the metatarsus of a male 
individual. Collections in the Florida State Geological Survey from 
near the head of the Itchtucknee River, Columbia County, include 
metatarsi, femora, humerus, ulnae, and other bones, all more or less 


NO. 2 AVIFAUNA OF PLEISTOCENE IN FLORIDA—WETMORE 33 


fragmentary, and part of a sternum. There is part of a metatarsus 
in the same collections from the north bank of the canal between the 
Florida East Coast Railroad and the highway at Vero. Gidley col- 
lected a broken metatarsus near Melbourne March 1, 1928, and 
Singleton in the same year working in this same deposit for the 
Museum of Comparative Zoology secured parts of humerus, meta- 
tarsus, and tibio-tarsus. The wild turkey must have been abundant in 
Florida during the Ice Age. 

The type specimen of Ardea sellardsi Shufeldt,’ the distal end of 
a right tibio-tarsus, proves on examination to be from a wild turkey. 
The bone is from an individual apparently barely adult and of small 
size, possibly from a young female. The condyles are worn and 
abraded in such a manner as to mask their true form, leading to error 
in the earlier identification. The type in question is equalled in size 
by the smallest in a considerable series of modern wild turkey bones 
examined. Ardea sellardsi thus becomes a synonym of Meleagris 
gallopavo. The specimen was taken in Pleistocene deposits in stratum 
Number Three, near Vero, Florida. 


MELEAGRIS TRIDENS sp. nov. 


Characters—Metatarsus (pl. 6, and fig. 13) similar to that of 
Meleagris gallopavo Linnaeus * but male with three-pointed spur core. 

Description—Type, U.S. Nat. Mus. No. 12052. Central portion of 
shaft of right metatarsus, collected by W. W. Holmes, in the Semi- 
nole area, Pinellas County, Florida. Shaft strong, flattened antero- 
posteriorly below, and more rounded above ; anterior face with a wide, 
shallow groove that becomes obsolete at level of central spur; below 
this the anterior face is ridged and shallowly grooved by tendons 
leading to the toes; external side of shaft rounded; internal side more 
flattened, spurs rising from a common base in a broad buttress of 
bone projecting obliquely inward from the inner side of the posterior 
surface; central spur strong and heavy (tip partly broken away) ; 
with an accessory spur above and below of smaller size, the upper one 
slightly more acute than a right triangle in outline, relatively broad 
transversely, with the distal extremity widened laterally so that in 
form it is like a cog in a cogwheel; distal accessory spur longer, more 
slender, with a conical, rather sharp point; outer surface of buttress 
supporting spurs broadly grooved for the passage of tendons that in 
life passed down the back of the metatarsus ; a distinct, rather narrow, 


‘Journ. Geol., Jan-Feb. (publ. Jan.), 1917, p. 19. See also Florida State 
Geol. Surv., 1917, Ninth Ann. Rep., pp. 38-39, pl. 2, fig. 15. 
* Meleagris gallopavo Linnaeus, Syst. Nat., ed. 10, vol. 1, 1758, p. 156. 


34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


shallow groove across the base of the spur core buttress on the an- 
terior side, to allow passage of another tendon. Bone brown in color, 
well fossilized. 

Remarks.—In size and form the specimen here described is similar 
to the ordinary turkey, Meleagris gallopavo, except for the three 
points of the spur core. Description of it as a new species has been 
withheld for 2 years to allow careful consideration of its peculiarities. 
These are susceptible of three interpretations: (1) that the bone is 
pathological and therefore aberrant, (2) that it is simply an individual 
variant, and (3) that it represents a distinct species. 








Fic. 13.—Fragmentary metatarsus (type) of Meleagris 
tridens (natural size). 


With regard to the first it needs only casual inspection to determine 
that the entire bone, including the spur cores is entirely normal and 
without any indication of disease. The very regularity and symmetry 
of its development indicate that the increased number of spurs is not 
due to any injury. As for the second supposition, in the past two years 
the writer has examined critically all of the specimens of wild turkeys 
that have been available to him, has seen the tarsal bones of a con- 
siderable number, one hundred or more, that have come from Indian 
pueblos in the Southwest and elsewhere, has seen several hundreds of 
domestic turkeys, and has talked with persons who have reared do- 
mestic turkeys for years without learning of any instance where a 
male turkey had more than a single spur. Under these circumstances 


NO. 2 AVIFAUNA OF PLEISTOCENE IN FLORIDA—-WETMORE 35 


it has seemed entirely logical to consider this specimen as representa- 
tive of a peculiar species. Among numerous other tarsal bones in the 
material from the Florida Pleistocene it stands unique, indicating 
probable rarity. It is possible that some of the bones from other parts 
of the skeleton that have been identified as Meleagris gallopavo belong 
to M. tridens but this cannot be determined. 

It may be remarked that multiple spurs are not unusual among 
gallinaceous birds of the Old World, though hitherto unknown in any 
American species. The pheasants of the genus /thaginis regularly 
have two or more pairs of spurs in the male. The same is true of 
Polyplectron, while according to Ogilvie-Grant * duplicate spurs occur 
in Haematortyx, Caloperdix, and Galloperdix. The vulturine guinea 
fowl, Acryllium vulturinum, frequently has two to four lumpy spur- 
like processes on the tarsus. 

The type of Meleagris tridens is so fragmentary that it affords few 
measurements. The transverse diameter of the shaft just below the 
spurs is 9.0 mm. The buttress supporting the spurs is 30.6 mm. long. 
The form may be ascertained from the accompanying figure. 


Order GRUIFORMES 
Family GRUIDAE 
GRUS AMERICANA (Linnaeus) 
Whooping crane 
Ardea americana Linnaeus, Syst. Nat., ed. 10, vol. 1, 1758, p. 142. 


Remains of cranes obtained by W. W. Holmes in the Seminole 
Field, Pinellas County, include fragments of metacarpus, tibio-tarsus 
and femur, and of three ulnae. Two fragmentary ulnae and one 
radius are in the collections of the Florida State Geological Survey 
from the Itchtucknee River area in Columbia County. Gidley collected 
part of an ulna from stratum Number Two at Melbourne in 1930, and 
Singleton secured part of another ulna in the same beds in June, 1929, 
when collecting for the Museum of Comparative Zoology. All are 
easily distinguished from the bones of other cranes found with them 
by their much greater size. 

Though the whooping crane was recorded from Florida by early 
ornithologists, in recent years doubt has been cast upon these reports 
and the species seems not to have been certainly found in modern 


*Cat. Birds Brit. Mus., vol. 22, 1893, pp. 221, 222, 260. 


36 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


times south of Georgia. These records from the Pleistocene are 
therefore of exceptional interest. This species is now nearly extinct, 
only a few individuals being known to exist in the interior of our 
country. It has not been recorded previously as a fossil. 


GRUS CANADENSIS (Linnaeus) 


Gray crane 
Ardea canadensis Linnaeus, Syst. Nat., ed. 10, vol. 1, 1758, p. 141. 


Remains of gray cranes are common in the deposits at Melbourne 
and in the Seminole area, and contain representatives of two forms, 
one large in size and the other small. The large form has the dimen- 
sions of Grus canadensis pratensis Meyer, the Florida crane, which is 
resident in Florida today, and is supposed to be that race. There is 
now recognized however another race, Grus canadensis tabida Peters, 
of more northern and western range that resembles pratensis in size, 
but differs in coloration, so that there is no certainty as to the form 
that ranged in Florida during the Pleistocene. This larger bird is 
represented in the Pleistocene collections by a coracoid, a femur, and 
part of a metacarpal, all secured by Gidley near Melbourne, and the 
head of a metatarsus and the symphysis of a lower mandible collected 
by Holmes in the Seminole area, as well as by part of a tibio-tarsus 
secured by Moore at Bradenton. 

The smaller race from the Florida Pleistocene has the dimensions 
of the little brown crane, Grus canadensis canadensis, that now ranges 
in the western half of the United States, and might be supposed to 
be that form were it not that the Cuban crane, Grus canadensis 
nesiotes, is a bird of equally small dimension. In fact the differences 
between G. c. canadensis and G. c. nesiotes seem to rest on color 
characters that appear not to have been definitely worked out. The 
small form is represented in the Pleistocene collections at hand by the 
distal end of a humerus, parts of two radii, and two coracoids from 
Melbourne, obtained by Gidley, and the distal end of a humerus 
secured by Holmes in the Seminole area. 

The occurrence of these two races in the Pleistocene of Florida is. 
suggestive of the modern condition in the western part of the United 
States, where a large gray crane and a small one occur together during 
migration over a considerable area. 


NO. 2 AVIFAUNA OF PLEISTOCENE IN FLORIDA 





WETMORE 37 


Family ARAMIDAE 


ARAMUS PICTUS (Meyer) 
Limpkin 
Tantalus pictus Meyer, Zool. Ann., vol. 1, 1794, p. 287. 


The distal end of a left humerus was collected by W. W. Holmes 
in the Seminole area. Parts of five metatarsi and a broken tibio- 
tarsus are found in collections from the Itchtucknee River deposits 
in Columbia County, the specimens being in the Florida State Geo- 
logical Survey. All these are similar to the corresponding bones in 
modern birds. The species is of regular occurrence in Florida at the 
present time. It has not been recorded previously as a fossil. 


Family RALLIDAE 
RALLUS ELEGANS Audubon 


King rail 
Rallus elegans Audubon, Birds Amer. (folio), vol. 3, 1834, pl. 203. 


In the Seminole area W. W. Holmes secured a complete right 
femur, and Mr. and Mrs. H. H. Simpson obtained a humerus in the 
Itchtucknee beds in Columbia County. These bones of this species 
are distinguished from the clapper rail by larger size. 

This rail, common in Florida now, inhabits mainly fresh-water 
marshes. It has not been recorded previously as a fossil. 


RALLUS LONGIROSTRIS Boddaert 


Clapper rail 
Rallus longirostris Boddaert, Tabl. Planch. Enl., 1783, p. 52. 


The distal end of a humerus comes from the Seminole area, col- 
lected by W. W. Holmes. The clapper rail is a sedentary species 
inhabiting salt-water marshes that is common at the present time along 
the coast of Florida, where several subspecies, slightly differentiated 
from one another, occur in different geographic areas. 

It has not been reported previously as a fossil. 


ARAMIDES CAJANEA (Miiller) 


Wood rail 
Fulica cajanea Miller, Vollst. Nat. Suppl., 1776, p. 1109. 
The determination of two fragmentary metatarsi (see figs. 14-16) 
and a nearly complete femur collected by W. W. Holmes in the Semi- 


nole area as belonging to a form of wood rail, a group of birds com- 
prising several forms that range now from southeastern Mexico south 


38 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 85 


to Argentina, has been another of the unexpected finds in the present 
collection. The two metatarsal bones are well fossilized, one being 
black in color and the other brown. The femur contains somewhat 
less mineral. 

Identification to species of these bones has been difficult owing to 
lack of material for comparison. That they are not related to the 
large A. ypecaha and A. saracura of the area from southern Brazil 
southward is obvious. Likewise it is evident on close study that they 








Fics. 14-16.—Metatarsi1 of the wood rail Aramides cajanea from the Seminole 
area (natural size). 


are from a bird larger than A. axillaris and its allies, which are among 
the smallest forms of the group. They are smaller than albiventris, 
but agree with A. cajanea, which now ranges in two or more sub- 
species from Panama southward into Brazil, and are identified as of 
that group on this basis. The genus has not been previously recorded 
north of southeastern Mexico nor has it been previously encountered 
as a fossil. Its occurrence in the Pleistocene of Florida is quite in 
keeping with the various types of mammals of South American 
affinity that come from these same beds. 


NO. 2 AVIFAUNA OF PLEISTOCENE IN. FLORIDA—WETMORE 39 


GALLINULA CHLOROPUS (Linnaeus) 
Gallinule 

Fulica chloropus Linnaeus, Syst. Nat., ed. 10, vol. 1, 1758, p. 152. 

Parts of four humeri were collected by W. W. Holmes in the 
Seminole field on the west coast, and another humerus was secured 
on the Itchtucknee River, Columbia County, by Mr. and Mrs. H. H. 
Simpson. The species, abundant in present day Florida, has not been 
recorded previously as a fossil. 


FULICA AMERICANA Gmelin 
Coot 

Fulica americana Gmelin, Syst. Nat., vol. 1, pt. 2, 1780, p. 704. 

Bones of this species collected in the Seminole area by W. W. 
Holmes include one entire and two fragmentary humeri and the distal 
ends of two tibio-tarsi. Parts of a humerus and a coracoid were 
obtained by J. E. Moore at Bradenton. A number of other limb bones 
are found in the collections of the Florida State Geological Survey 
from the Itchtucknee River area in Columbia County. 

The coot is found now in abundance in Florida in winter and a few 
remain to nest during summer. The species has been reported pre- 
viously from the Pleistocene of Oregon. 


Order COLUMBIFORMES 
Family COLUMBIDAE 
ZENAIDURA MACROURA (Linnaeus) 
Mourning dove 
Columba macroura Linnaeus, Syst. Nat., ed. 10, vol. 1, 1758, p. 164. 


Two metacarpals and the proximal end of an ulna from the Semi- 
nole Field, obtained by W. W. Holmes, are in size similar to the 
modern forms of the mourning dove of North America, being larger 
than the bird of the West Indies. 


Order STRIGIFORMES 
Family TYTONIDAE 
TYTO ALBA (Scopoli) 
Barn owl 
Strix alba Scopoli, Annus I. Hist.-Nat., 1769, p. 21. 


In the collection obtained by W. W. Holmes in Saber-tooth Cave 
at Lecanto in 1928 there are a number of fragmentary bones of the 


40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


barn owl, including parts of the femur, tibio-tarsus, metatarsus, and 
ulna. The species is quite common in modern Florida. 


Family STRIGIDAE 
OTUS ASIO (Linnaeus) 
Screech owl 
Strix asio Linnaeus, Syst. Nat., ed. 10, vol. 1, 1758, p. 92. 
Parts of two humeri of the screech owl were collected by W. W. 


Holmes in Saber-tooth Cave near Lecanto. This bird is common and 
widely distributed in Florida. 


STRIX VARIA Barton 
Barred owl 

Strix varius Barton, Fragm. Nat. Hist. Penn., 17909, p. II. 

The barred owl, a common species in Florida at the present time, 
seems to have had equally wide distribution during the Pleistocene. 
In the Seminole area W. W. Holmes obtained a number of fragments 
including parts of the metatarsus, humerus, ulna, and metacarpus. 
In the excavation of Saber-tooth Cave at Lecanto Mr. Holmes further 
obtained a nearly complete femur. J. W. Gidley secured part of a 
metatarsus in the golf links area at Melbourne. The species has not 
been previously recorded as a fossil. 


Order PASSERIFORMES 
Family CORVIDAE 
CORVUS BRACHYRHYNCHOS Brehm 
Crow 
Corvus brachyrhynchos Brehm, Beitr. Vogelk., vol. 2, 1822, p. 56. 


Numerous remains of the common crow were secured by W. W. 
Holmes in the Seminole area, indicating that this species was as 
common during the Pleistocene as it is in Florida at the present time. 
Crows have been recorded previously from Ice Age deposits in Cali- 
fornia but not before from the Pleistocene of eastern North America. 


CORVUS OSSIFRAGUS Wilson 
Fish crow 
Corvus ossifragus Wilson, Amer. Orn., vol. 5, 1812, p. 27, pl. 37, fig. 2. 


A humerus, lacking the head, was obtained by W. W. Holmes in 
the Seminole area, this being the first report of this species for the 


NO. 2 AVIFAUNA OF PLEISTOCENE IN FLORIDA—-WETMORE 41 


Pleistocene. The bone is similar to that of the common crow but is 
decidedly smaller. The fish crow is widely distributed through the 
Florida Peninsula today. 


Family ICTERIDAE 
AGELAIUS PHOENICEUS (Linnaeus) 
Red-winged blackbird 
Oriolus phoeniceus Linnaeus, Syst. Nat., ed. 12, vol. I, 1766, p. 161. 


A right humerus lacking the distal end, and a left one with part of 
the head missing, were secured in the Seminole area by W. W. 
Holmes. This is a common resident of marshes throughout much of 
North America and abounds today in Florida. It has not been identi- 
fied certainly before from the Pleistocene. 


’ 


MEGAQUISCALUS MAJOR (Vieillot) 
Boat-tailed grackle 
Ouiscalus major Vieillot, Nouv. Dict. Hist. Nat., vol. 28, 1819, p. 487. 


The proximal portion of a right humerus found by W. W. Holmes 
in the Seminole area comes from an individual of small size. These 
grackles are common in Florida, ranging mainly about water. The 
species has not been recorded before from the Pleistocene. 


QUISCALUS QUISCULA (Linnaeus) 
Crow blackbird 
Gracula quiscula Linnaeus, Syst. Nat., ed. 10, vol. 1, 1758, p. 109. 


A nearly complete left humerus secured by W. W. Holmes in the 
Seminole Field comes from an individual of small size. This grackle, 
common in modern Florida, has not been recorded previously from 
the Pleistocene. 



















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SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL.“85)\NO. 2,. PL. 7 





1. General view of the Seminole area with excavation in foreground. 


Photograph by W. W. Holmes. 





~ 
i i 


25 


2. Stratification in excavation at Seminole with the bone bearing layer indi- 
cated by two trowels at center. Photograph by W. W. Holmes. 


4 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. (85, NO= 2), ‘RE. 2 





1. Entrance to Saber-tooth Cave near Lecanto, Fla. 





2. Excavations in Saber-tooth Cave. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85, NO. 2, PL. 3 


<y 





1. Stratification at Melbourne, the bone bearing layer 
being the dark band through the center. Photograph by 


J. W. Gidley. 





2. Excavations on the golf links at Melbourne, Fla. Photograph by 


J. W. Gidley. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. "85, NO. 2); IPL. 4: 





At left fragmentary metatarsus of California condor, collected by J. E. 
Moore at Venice, Fla., compared with modern specimen at right. (Natural size.) 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLS 855 NO. 2, PEL 5 


i. 

iM 

‘ 
Ry 





2 


J. E. Moore at Bradenton, Fla. 


1 
Metatarsus of Teratornis merriami obtained by | 
( Natural size. ) 


2. Femur of Teratornis merriami obtained by J. 
(Natural size. ) 


ie 
EE. Moore at Bradenton, Fla. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLE. 85; NO; 2) (PE. 6 





At left type of Meleagris tridens, compared with metatarsus of modern male 
Meleagris gallopavo merriami. (Natural size.) 





SMITHSONIAN MISCELLANEOUS COLLECTIONS 


VOLUME 85, NUMBER 3 


ADDENDA TO DESCRIPTIONS OF 
SUK Ghoo SotAle Osos 


(WiTH 23 PLATEs) 


BY 
CHARLES D. WALCOTT 


(With Explanatory Notes by Charles E. Resser) 





(PUBLICATION 3117) 


CITY OF WASHINGTON 
PUBLISHED BY THE SMITHSONIAN INSTITUTION 
JUNIE 295 1931 


Tbe Lord Waitimore Press 


BALTIMORE, MD,, U. 8 A. 


ADDENDA TO DESCRIPTIONS OF BURGESS 
SHALE. FOSSILS 


By CHARLES D. WALCOTT 
(Witn ExpLanatory Notes By CHARLES E. RESSER) 
(WitTH 23 PLATES) 
PREFATORY STATEMENT 


Shortly after his discovery of the remarkable Burgess shale fossils 
in British Columbia in 1910, Dr. Charles D. Walcott described the 
more striking species of the major classes of animals and plants 
represented, to bring to the attention of the scientific world their 
exceptionally well preserved anatomic details. Subsequent quarrying 
at the locality yielded many additional specimens of the described 
forms as well as examples of rarer species not secured in the first 
season’s work. 

During my 13 years’ association with Doctor Walcott he frequently 
dwelt upon the fact that he considered his papers on the Burgess 
shale forms rather in the nature of announcements than as completed 
studies of these wonderfully preserved fossils. He always intended 
to return to the study of the described species and to publish more 
detailed descriptions and interpretations of their form and structure. 
However, the stress of war times and advancing years prevented a 
realization of this hope. Nevertheless, from time to time, he had 
photographs prepared or made notes of his observations regarding 
structure, all of which were preserved with the collections. 

At the request of the National Museum authorities I have assembled 
these notes and illustrations for publication so that they may not be 
lost to science. It must be remembered that none of the statements, 
and particularly none of the interpretations, in the following pages 
should be regarded as Doctor Walcott’s final opinion, since he recog- 
nized many of them as tentative. He more than once stated that 
fully 15 years’ work remained to be done on the 35,000 Burgess shale 
specimens in the National Museum’s collections. 

All generic and specific names, having been created by Doctor 
Walcott, are, of course, to be credited to him, and not to us jointly. 

In order to show clearly exactly what Doctor Walcott wrote and, 
on the other hand, what I have added—chiefly by way of explana- 
tion—two type faces are used. Doctor Walcott’s manuscript is printed 
in 10-point type, while the explanations added by me appear in the 
aUatlSe DOME type: Cuartes E. Resser. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 85, No. 3 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


Nv 


INTRODUCTION 


It is obvious that one person cannot cover, even in a very cursory manner, 
the entire field of Cambrian stratigraphy and paleontology, especially with such 
rich faunas as the Burgess shale extant. Again, in the case of the Burgess 
shale faunas, none but a trained student in biology can do more than merely 
assign specific and generic names to the gross forms. Further, it is doubtful if 
any biologist, however versatile, could by long years of study perceive every- 
thing to be learned from these wonderful fossils. In view of these facts it has 
seemed advisable to encourage specialists to study the various classes repre- 
sented rather than attempt to monograph the subject as a whole. Accordingly, 
such a study by Dr. G. E. Hutchinson has recently been printed* and one by 
Dr. Rudolf Ruedemann has been submitted for publication.*? In this way we 
may hope to carry on the work suspended by the passing of the discoverer of 
these unique forms. 

The following descriptions either were prepared entirely by Doctor Walcott 
or are based on notes and illustrations left by him. 

Formation and locality—For every species described in this paper, the for- 
mation and locality are as follows: Middle Cambrian, Burgess shale: (Loc. 
35k) on the west slope of the ridge between Mount Field and Wapta Peak, 
1 mile (1.6 km.) northeast of Burgess Pass, near Field, British Columbia. 


DESCRIPTION OF GENERA AND SPECIES 
MARGARETIA, new genus 


The characters of this strange organism are presented in the specific descrip- 
tion or shown in the illustrations. 
Genotype.—M. dorus, new species. 


MARGARETIA DORUS, new species 
Plate 1, figs. 1-6 


More than 70 specimens of this peculiar organism have been assembled from 
the Burgess shale collections. In the following description comparisons are 
made with algae. Other notes by Doctor Walcott, apparently his latest, together 
with suggestions by Mr. A. H. Clark, and particularly the presence in the same 
drawer of specimens of Titanideum suberosum, indicate that Doctor Walcott’s 
latest opinion was that M. dorus might really be an Alcyonarian. 


Description——Mass forming a thin membranous perforated sheet, 
narrow at the base and expanding to a width of 1.5 cm. in 2 cm. 
distance ; length of narrow base about 1.5 cm, and of wider portion 
4 cm.; the perforations are elongate oval and apparently arranged on 
longitudinal and obliquely transverse lines; tegument presumably 


*Restudy of Some Burgess Shale Fossils. Proc. U. S. Nat. Mus., vol. 78, 


art. II, pp. I-24, pl. 1, year ? 
* Some New Middle Cambtian Fossils from British Columbia. To be printed 


in the Proc. U: S. Nat. Mus. 


NO. 3 BURGESS SHALE FOSSILS—-WALCOTT 3 


leathery as it is a black film with irregular, broken, longitudinal lines 
with more or less scaly edges. 


Microscopic structure undetermined. 


Observations.—The perforations in the tegument are not unlike 
those of the living alga Agarum turneri Post and Ruprecht, and one 
might imagine that a fragment of the strong frond of this species 
is similar to M. dorus, but the resemblance is only general ; the per- 
forations of M. dorus are more uniform than those of the beautifully 
perforate living alga Kallymenia perforata Agardh, which also has a 
far more delicate tegument. 


Holotype and paratypes —uvU. S. N. M., Nos. 83922, 83923a-e. 


REDOUBTIA Walcott 1918 
REDOUBTIA POLYPODIA Walcott 
Plate 2, figs. 2-3 


Redoubtia polypodia Walcott, 1918, Smithsonian Mise. Coll., vol. 68, no. 12, 
p. 5, fig. 5. 


The holotype is refigured since it was first published in the popular account 
of field explorations issued annually by the Smithsonian Institution, which does 
not reach paleontologists generally. 

Accompanying the original figure is the following statement, “An elongate 
creeping holothurian with numerous tube feet and tentacles.” 

Whether the second specimen really represents the same species appears some- 
what doubtful inasmuch as the tube feet are smaller and more numerous. The 
larger appendages above the specimen, as posed on the plate, are parts of 
another animal. 


Holotype and paratype. 





U.S. N. M., Nos. 83924 and 83925. 


PORTALIA Walcott 1918 
PORTALIA MIRA Walcott 
Plate 3, figs. 2-3 


Portalia mira Walcott, 1918, Smithsonian Misc. Coll., vol. 68, no. 12, p. 6, 
figs. 6, 7. 


Another holothurian first figured in the explorations account for 1918 is also 
refigured, to give it wider availability. This form differs from the preceding 
Redoubtia polypodia in having fewer and longer tube feet and in their apparently 
different grouping. 


Holotype-—U. S. N. M., No. 83027. 


4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


MISKOIA Walcott 1911’ 
MISKOIA PLACIDA, new species 
Plate 2, fig. I 


Comparing this form with M. preciosa Walcott, the genotype, the new species 
is stouter and shorter. As none of the specimens referred to the type species 
preserves the rear end, it is not possible to determine whether the lobate termi- 
nation of M. placida is also characteristic of the first species. The annulations 
of the body are clearly marked in the specimen illustrated, particularly on the 
counterpart, which is a mold of the exterior. Teeth are shown around the mouth 
as bright, shiny, curved, chitinous hooks. 


Holotype—U. S. N. M., No. 83028. 


CANADIA Walcott 1911 
CANADIA SPINOSA Walcott 


Plate 4; plate 5; plate 6, figs. 1-2 


Canadia spinosa Walcott, 1911, Smithsonian Misc. Coll., vol. 57, no. 5, p. 118, 
pl. 23, figs. 4-7. 

Original description — Body slender, formed of 20 to 21 segments 
that, when flattened on the shale, are a little longer than wide; each 
segment has a pair of parapodia with a dorsal and ventral bundle of 
strong non-jointed setae. The setae are finely illustrated by figures 4, 
6, and 7. Head minute, with a pair of large tentacles curving outward 
from the front anterior margins; a bundle of fine setae occurs on 
each side of the head back of the base of the large tentacles. A 
straight slender enteric canal is indicated on several specimens. Mouth 
and anus not seen, but probably at or near the end of the annelid. 

“ Dimensions.—The largest adult specimen has a length of 34 mm., 
with a width of the body at the seventh segment from the head 
of. 1.5 mm.” 


Fortunately many additional specimens of this interesting spiny worm were 
found after 1911. Those first illustrated give a fairly good conception of the 
general features; however, illustrations of additional specimens may show 
features particularly desired by the biological student at places where the other 
specimens are faulty. 


Plesiotypes—U. S. N. M., Nos. 83929a-e. 
Walcott, C. D., Smithsonian Misc. Coll., vol. 57, no. 5, p. 114, pl. 18, figs. 
1-5, IOI. 


NO. 3 BURGESS SHALE FOSSILS—WALCOTT 5 


CANADIA SETIGERA Walcott 
Plate 7, figs. 1, 4; plate 8, fig. 3 


Canadia setigera Walcott, 1911, Smithsonian Misc. Coll., vol. 57, p. 110, pl. 23, 
figs. 1-3. 

Canadia setigera Walcott, 1916, Ann. Rep. Smithsonian Inst., 1915, pl. 12, 
figs. I-3. 


The original description states that “this species differs from C. spinosa in 
being more elongate, slender, and with much smaller bundles of finer setae.” 

It is further stated that a series of 36 specimens shows gradation between the 
two types originally illustrated. It seems, however, that in reality several dis- 
tinct forms are included in the species as now constituted. 

Plesiotypes.——U. S. N. M., Nos. 83930a-c. 





CANADIA GRANDIS, new species 
Plate 0, fig. 10 


A single wide Canadia that shows the body annulations very well and that 
has numerous setae seems to differ from C. spinosa mainly in the larger bundles 
of setae. 

Holotype-——vU. S. N. M., No. 83932. 


CANADIA IRREGULARIS Walcott 
Plate 6, figs. 4-6; plate 7, fig. 3 
Canadia irregularis Walcott, 1911, Smithsonian Misc. Coll., vol. 57, p. 120. 


Original description.—“A slender species not over 20 mm. in length. 
The setae are irregular in size and appearance and suggest partially 
worn macerated specimens of the slender forms of C. setigera.” 

The specimens on which this description was based are now illustrated for 
the first time. A study of the figures, however, causes some doubt to arise 
regarding specific differentiation from C. grandis. 

Lectotype and paratypes—uU. S. N. M., Nos. 83933 and 83934a 
and b. 


CANADIA SPARSA Walcott 
Plate 6, fig. 3 
Canadia sparsa Walcott, 1911, Smithsonian Misc. Coll., vol. 57, p. 119. 


Original description.— A slender form with only two strong setae 
on each very short parapodia. Finer setae may occur but they are 
not shown in the one specimen.” 

This form is another that was not illustrated in 1911. In this case two ques- 


tions may be raised: First, the specific identity of all the specimens seems 
doubtful, and second, the generic reference to Canadia is also uncertain. 


Holotype —U. S. N. M., No. 83935. 


6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


CANADIA DUBIA Walcott 
Plate 7, fig. 2; plate 8, figs. 1-2; plate 0, fig. 8 
Canadia dubia Walcott, 1911, Smithsonian Misc. Coll., vol. 57, p. 110. 


Original description‘ This species is proposed to include a small 
chaetiferous annelid not over 10 mm. in length. One specimen shows 
a bundle of very fine setae on each side near the head.” 


Four of the original specimens are illustrated. 


Cotypes—U. S. N. M., Nos. 83936a-d. 


CANADIA SIMPLEX, new species 
Plate 0, fig. 9 
A tiny organism that appears as a slender tube with a termination surrounded 


by a ring of setae constitutes the material to which Doctor Walcott attached 


this name. 
Holotype —U. S. N. M., No. 83937. 


WIWAXIA Walcott 1911 
WIWAXIA CORRUGATA (Matthew) 
Plate 3, fig. 1 


Orthotheca corrugata Matthew, 1800, Trans. Roy. Soc. Canada, 2d ser., vol. 5, 
sec, 4; -p. 42) ple 1, lige 3: 

Orthotheca corrugata Walcott, 1908, Canadian Alpine Journ., vol. I, no. 2, 
p. 246; plat figs ti: 

Wiwaxia corrugata Walcott, 1911, Smithsonian Misc. Coll., vol. 57, no. 5, 


p. 123, pl. 21, figs. 1-4. 
A particularly fine example of this remarkable spined worm turned up in 
some of the more recent collections. Its picture is included since it may repre- 
sent a relatively undistorted specimen with most of the plates retained. 


Plesiotype.—U. S. N. M., No. 83038. 


OTTOIA Walcott 1911 
OTTOIA MINOR Walcott 
Plate 9, figs. 1-7 


Ottoia minor Walcott, 1911, Smithsonian Misc. Coll., vol. 57, p. 120, pl. 22, 
figs. 5-6. 
Original description—* This species differs from O. prolifica in 
its proportionally more slender form when elongated and straighter 
outline both when elongated and contracted. The hooks are also much 





NO 3 BURGESS SHALE FOSSILS WALCOTT 7 
finer and extend farther back on the anterior end. The annular lines 
and interspaces are also finer and more irregular.” 

Several specimens, among the many found subsequent to I91I, preserve some 
of the structure features very well, and illustrations were prepared by Doctor 


Walcott to show them. However, it is very doubtful whether the forms shown 
on plate 9, figures 2 and 4, belong to this species. 


Plesiotypes—U. S. N. M., Nos 83939a-g. 


PIKAIA Walcott 1911 
PIKAIA GRACILENS Walcott 
Plate 8, figs. 4-5 
Pikaia gracilens Walcott, 1911, Smithsonian Misc. Coll., vol. 57, p. 132, pl. 20, 
figs. 1-2. 

Original description— Body elongate, slender, and tapering at 
each end. It is formed of many segments that are defined by strong 
annular shiny lines. Head small with two large eyes and two tentacles 
as shown by figure 1. Back of the head the first five segments carry 
short parapodia that appear to be divided into two parts. 

“The enteric canal extends from end to end without change in 
character. It is relatively large along the central portions and tapering 
toward the ends. Judging from such specimens as the one illustrated 
by figure 2, its annulations correspond in size with those of the body. 

“Surface apparently smooth. Two entire adult specimens and 
several fragments of others indicate a length of about 5 cm.” 


Two additional figures are presented at this time. 


Plesiotypes-—U. S. N. M., Nos. 83940a-b. 


SELKIRKIA Walcott 1911 
SELKIRKIA MAJOR (Walcott) 
Plate 10 


Orthotheca major Walcott, 1908, Canadian Alpine Journ., vol. 1, p. 246, pl. 1, 
fig. II. 

Selkirkia major Walcott, 1911, Smithsonian Misc. Coll., vol. 57, p. 120, pl. 10, 
fig. 6. 


This species was first described from the Stephen formation on Mount Stephen. 
Later Doctor Walcott found apparently the same shell at the Burgess Pass 
quarry, but in this instance the soft body of the animal was preserved and there- 
fore, in the 1911 discussion, he removed it from the Hyolithidae to the poly- 
chaetous annelids. 

Photographs of two exceedingly well preserved individuals with the body 
extending beyond the shell are printed here for the first time. 

Plesiotypes.—U. S. N. M., No. 83041a-b. 





8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


AYSHEAIA Walcott 1911 
AYSHEAIA PEDUNCULATA Walcott 
Plate 11 


Aysheaia pedunculata Walcott, 1911, Smithsonian Misc. Coll., vol. 57, p. 117, 
pl. 23, figs. 8-0. 

Aysheaia pedunculata Hutchinson, 1930, Proc. U. S. Nat. Mus., vol. 78, art. 11, 
p. 14. 

This fossil has perhaps attracted wider attention than any other. Recently 
G. E. Hutchinson, of Yale University, studied this peculiar form and concluded 
that Aysheaia is an extinct Onychophora. Unfortunately, Mr. Hutchinson did not 
see the two individuals here illustrated, as they were buried away among 
numerous specimens of Ottoia. As the present assemblage of material progressed 
they came to light and are now illustrated, especially as they are perhaps the best 
preserved specimens available. 

At the time these photographs were found among the notes Doctor Walcott 
intended using in further publications relating to this animal, two letters were 
discovered, both suggesting that Aysheaia may be an Onychophora or a Perip- 
atus. The first letter, dated September 21, 1911, was written by Prof. W. M. 
Wheeler of Harvard University, and reads as follows: 

“T wish to thank you for your very interesting publications on the Middle 
Cambrian Annelids. On plate 23, I noticed two figures of Aysheaia pedunculata. 
This creature bears the most extraordinary resemblance to Peripatus, except for 
the head, and judging from the figures the ‘head’ may be something which 
does not belong to the fossil. I have just shown these figures to Mr. C. T. Brues, 
who has been working on Peripatus, and he also was struck with the remarkable 
resemblance. Is there any possibility that it might be Peripatus instead of an 
Annelid? If this should prove to be the case it would be a matter of the very 
greatest interest.” 

The second letter dated October 25, 1911, was written by Prof. Charles 
Schuchert, of Yale University, and contains the following : 

“The other point is one that Lull has called my attention to and refers to 
figures 8 and 9 of plate 23 which you call Aysheaia pedunculata. The question 
that I want to ask is, have you considered it as a possible Onychophora or 
related to Peripatus? Of course if one looks at your illustrations and compares 
them with the illustration of Peripatus given by Parker and Haswell in their 
Text-book of Zoology, page 607, in the edition of 1910, one can see considerable 
differences and yet there are in your figures several points in common to 
make one wonder whether you have not a marine ancestor of this land-living 
arthropod.” 

This rather lengthy historical account is presented to show that at least three 
authorities arrived at the same conclusions independently. 

Plesiotypes—U. S. N. M., Nos. 83942a-b. 


LEANCHOILIA Walcott 1912 
LEANCHOILIA SUPERLATA Walcott 
Plate 12; plate 13, figs. 1-2; plate 14, figs. 4-5 


Leanchoilia superlata Walcott, 1912. Smithsonian Misc. Coll., vol. 57, p. 170, 
pl. 31, fig. 6. 


Original description.— Body elongate, with clearly defined head 
shield and nine strong body segments up to the point where the 


NO. 3 BURGESS SHALE FOSSILS—WALCOTT 9 


posterior part of the body is broken off. The anterior pointed end of 
the head is broken off in such a manner that the presence of a frontal 
appendage is suggested. The large opening on the side of the head 
indicates a large pedunculated eye comparable with that of Opabinia 
regalis (pl. 28, fig. 1). 

“ Appendages.—Of the head appendages, the antennae are the best 
preserved. These are large and composed of several strong joints, 
of which three now show from beneath the carapace; the second of 
these bears a long slender branch on its inner margin, and the third 
two branches, one of which is similar to that of the second joint. 
These two branches appear to be composed of one very long slender 
joint followed at the end by several short small joints that curve 
upward and presumably gave the branches flexible extremities ; the 
third and lower branch has a similar slender proximal joint that at 
its outer end has three slender, jointed branches. This structure makes 
a very effective clasper of each of the antennae. Back of the right 
antennae are two narrow appendages that may be the ends of one of 
the third and fourth pairs of head appendages. 

“The thoracic legs terminate in flat, elongate, broad, lanceolate 
joints. The terminal joint is about three-fifths the entire length of 
the leg, and has a fringe of strong setae on its outer and posterior 
margin. The condition of preservation is such that the details of struc- 
ture of the other portions of the leg cannot clearly be determined.” 


The illustrations presented herewith apparently were prepared by Doctor 
Walcott to exhibit the detailed structure of this interesting crustacean. 


Plesiotypes—U. S. N. M., Nos. 83943a-g. 


LEANCHOILIA MAJOR, new species 





Plate 13, fig. 3 


Several specimens, of which the best is illustrated, were labeled Leanchoilia 
major by Walcott. Just why he should have chosen this specific name is not 
readily apparent as these individuals are not sufficiently larger than L. superlata 
to warrant the designation. In fact there is but little difference between this 
form, which happens to be flattened out horizontally, and the specimen shown 
in figure 2, plate 13, referred to the genotype. 

Holotype-—U. S. N. M., No. 83044. 


NARAOIA Walcott 1912 
NARAOIA COMPACTA Walcott 
Plate 13, fig. 4; plate 14, figs. 1-3; plate 15, figs. 2-3 
Naraoia compacta Walcott, 1912, Smithsonian Misc. Coll., vol. 57, no. 6, p. 175, 
ple2s8y hesasea. 
Many specimens of this interesting form have been found since its 
preliminary description in 1912, but none shows the cephalic ap- 


IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


pendages in position or an uninjured trunk limb. The carapace is 
thicker than that of Burgessia bella and the body is firmly attached to 
the fused segments forming the posterior dorsal shield, and there is a 
close union between the body of the cephalic region and the carapace 


st 
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ewonee 





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H 


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a 
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i 
Alp 
aay 
i 
4} be 





Fic. 1.—Naraoia compacta Walcott. 


a, antennae; as, anal segment; ds, reflex margin of posterior carapace; en, 
endopodite; ex, exopodite; 7, intestine; m, mandible; m’, maxilla; mm”, maxillula; 
seg, segmented posterior carapace; st, stomach; tf, telson. 

(About 5.) Diagrammatic outline of ventral view of appendages, etc. 


that extends from the anterior ventral margin of the latter back to | 
the line of the anterior margin of the posterior dorsal shield. In the 
restoration (text figs. 1, 2) I have made an attempt to incorporate 
all information available. 


NO. 3 BURGESS SHALE FOSSILS—WALCOTT Tet 


Exoskeleton.—The dorsal exoskeleton as seen from its dorsal side 
is formed of a carapace and a posterior segmented shield. The true 
cephalic carapace or shell fold is attached to the cephalic somites near 
its anterior portion, probably as in the recent Apodidae or the asso- 
ciated Burgessia and also along the line of the body as far back as 
the anterior margin of the posterior dorsal carapace. The carapace 1s 
not known to have had a reflected anterior margin with a labrum at- 





hor eee |e —T=S-~.__ \\ WYS=-- 4. h 
WaT CRS i ! : 


tl, pa ae SEL. 


Fic. 2.—Naraoia compacta Walcott. 


c, Carapace; g, digestive glands; d.g., lateral digestive glands; c, eye; h.c., 
hepatic caeca; st, stomach; ¢./., thoracic limb. ; 
(About * 5.) Diagrammatic outline of the digestive organs. 


tached as in Burgessia, but it may have had, as none of the specimens 
shows the ventral side of this part in an uninjured condition. The 
carapace is broader than long, with a somewhat uniformly rounded 
outline except posteriorly where it is nearly transverse; it probably 
had a ventral as well as dorsal membrane between which the great 
hepatic caeca were located very much as are the shell glands in the 
Apodidae.”. For some unknown reason the anterior portion of the 
carapace is usually distorted by being crowded back so as to wrinkle 
and shorten it. 

The large shield of the posterior part of the exoskeleton is com- 
posed of 14 fused segments with a narrow border. It has the ap- 
pearance of the many-segmented pygidium of the trilobite belonging 


"In no other manner can I explain the wonderful preservation of the digestive 
tubes and caeca. 


I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


to the genera Ogygopsis or Orria,’ except that the median lobe is not 
as strongly marked. These two genera are mentioned as they occur in 
correlative Middle Cambrian formations and one of them in the 
Stephen formation which is nearly contemporaneous with the Burgess 
shale. The dorsal carapace and shield also appear similar in outline 
to the dorsal exoskeleton of the freshly hatched young of Limulus 
polyphemus as figured by Packard.* The posterior shield has each of 
the thoracic body segments, excepting the two posterior which pro- 
ject beyond it, attached directly to it, and it forms the dorsal side 
of the exoskeleton of the body segments. The exoskeleton of the free 
posterior segments and telson, of the cephalic segments, and of the 
ventral side of the trunk segments was so exceedingly delicate as to 
leave only a slight trace on the shale. Both the anterior carapace and 
posterior segmented shield were very thin and readily distorted. 

The anterior carapace slightly overlapped the posterior shield and 
the two were closely held together by the strong body as evidenced 
by their rarely being found separated. The telson is short and marked 
by fine short spines. 

Dimensions. —The largest specimen had a length of about 40 mm., 
the carapace being shortened by compression. A specimen that has 
been slightly narrowed by compression has the following dimensions : 


Menoth of carapace ccs. sere a sso cise ears pale ates 16 
Wadth oficarapacesiccasccme atene cities nite ae ile iol eters 15 
Length of posterior dorsal: shieldai..: ssc. a0 someteeien lens 17 
Width-of posterior dofsalishteldis: «sccm... soe ocrdeeie sen 14 


Eyes.—The eyes are represented by two small crescent-shaped 
bright spots a little in advance of the anterior end of the stomach as 
illustrated by the restoration (fig. 2). The position and form corre- 
spond quite closely to the paired eyes of the recent Apus lucasanus 
Packard.’ 

Cephalic appendages.—The antennae are uniramous, short jointed, 
and slender in their distal portion, and have a large proximal joint ; 
the intermediate joints are unknown. Of the cephalic limbs only slight 
indications were found of the proximal joints of three pairs, and a 
few terminal joints extending from beneath the carapace, nothing 
of their original form being preserved. All traces of cephalic ap- 


* Smithsonian Misc. Coll., vol. 64, no. 5, figs. 1 and 2, pl. 66, 1916. 

* Mem. Boston Soc. Nat. Hist., vol. 2, pl. 5, figs. 25, 25d, 1871. 

* Twelfth Ann. Rep., U. S. Geol. and Geog. Surv. Territories, Hayden, Pt. 1, 
pl. xvi, fig. 2, 1883. 


NO. 3 BURGESS SHALE FOSSILS—WALCOTT 13 


pendages are posterior to the hepato-pancreatic tubes passing from 
the stomach to the hepatic caeca. 

There is no clearly defined line between the cephalic and trunk 
limbs, but from the relations of the limbs in Burgessia and Marrella 
it is assumed that it is between the third pair of cephalic limbs and 
the supposed first pair of trunk limbs. The specimens are too much 
obscured by the compression they have undergone to permit of recog- 
nition of the detailed structure of the limbs. 

Thoracic limbs.—The specimen represented by figure 3, plate 14, 
has the distal ends of 17 thoracic or trunk limbs projecting beyond the 
left margin of the posterior shield; the shield in this specimen has 
not more than 14 fused segments outlined on it, so it is probable that 
the three anterior limbs belong with the body segments between the 
anterior segment of the posterior shield and the third pair of cephalic 
limbs. Another alternative is that the distal portion of the two anterior 
limbs extending beyond the margin of the shield belong to the maxilla 
and maxillula, which would leave only one pair of limbs from the seg- 
ment anterior to the posterior shield and posterior to the cephalic 
limbs. The limbs were so subject to displacement, however, that any 
deduction is very uncertain. The distal portion of the thoracic or 
trunk limbs shows an endopodite with a slightly curved terminal spine 
with a slender section back of it corresponding to the slender distal 
joint of the endopodite of Marrella and Burgessia; and then the joints 
broaden towards the coxopodite with slight indications of five joints 
between the distal joint and coxopodite. 

The exopodite is represented by many slender filaments that were 
attached to a multi-jointed arm or support similar in appearance to 
that of the exopodite of Marrella. The filaments are relatively broad, 
as they occur flattened on the shale. There are strong indications of 
large coxopodites, but none show their original form or the exact 
point of attachment of endopodite or exopodite, and the joints of 
the endopodites have been so crushed down as to be no longer definitely 
recognizable. The exopodites were nearly as long as the endopodites, 
and the filaments of the former are usually extended out to the end 
of the endopodites or beyond. 

Digestive organs—The exact location of the mouth is unknown, 
but from the apparent position of the antennae and proximal joints of 
the cephalic limbs, it was posterior to the point of entrance of the 
hepatic tubes, back of which the intestine was large with minor hepatic 
caeca opening into it through four small tubes, all of which are anterior 
to the posterior dorsal shield as indicated in the diagrammatic res- 
toration (fig. 2) ; beneath the posterior dorsal shield the intestine is 


14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


slightly constricted at the union of the trunk segments and extends 
back to the anal segment which protrudes posteriorly from beneath 
the shield ; the gullet connecting the mouth and the stomach must have 
extended forward and upward. 

The large hepatic glands and caeca are somewhat similar to those 
of Burgessia bella (text fig. 5); a short, strong tube with a well- 
marked anterior and posterior tube leads out from the stomach, and 
branching from these lateral tubes are a series of hepatic caeca. The 
small hepatic caeca are located between the long posterior tubes of the 
hepatic glands and the intestine and posterior to the main hepatic tube. 

Owing to the excellent state of preservation of many of the speci- 
mens showing the hepatic caeca it is probable that they were situated 
between the dorsal and ventral membrane of the carapace and thus 
held in position and protected from destruction; that they are pre- 
served at all is one of the wonders of this remarkable Burgess shale 
fauna. 

Functions of appendages——These were presumably the same as 
for similar organs in Marrella and Burgessia, and the mode of occur- 
rence is essentially the same. 

Diagrammatic restorations —The diagrammatic restoration (fig. 2) 
presents the outline of the carapace and posterior shield with the 
stomach, intestine, hepatic tubes and caeca of the digestive system 
outlined, also the thoracic trunk, the telson and the thoracic limbs 
as far as known and interpreted. The data for this diagram are very 
good except the jointing of the endopodites and the exact form of the 
coxopodites and proximal joints of the exopodites. In figure 1, the 
endopodites on each side have been omitted and the exopodites drawn 
in so as to show their structure and position above the endopodites 
and below the ventral membrane of the posterior dorsal shield. The 
sixth limb has both the endopodite and exopodite attached ; this should 
be compared with the thoracic limb of Marrella splendens (text fig. 9). 





We know so little of the cephalic limbs of Naraoia compacta that I 
hesitate to give a diagrammatic sketch of them, and it would not be of 
even tentative value if we did not have the cephalic limbs of Marrella 
and Burgessia for suggestion; from the latter and from the evidence 
afforded by a few specimens, the outline of figure I is drawn. 

Comparison with crustaceans—WNaraoia has many characters in 
common with the trilobite and some in common with Marrella, Bur- 
gessia, and Waptia, which will be spoken of in the discussion of this 
group of genera. 

Plesiotypes—U. S. N. M., Nos. 83945a-e. 


NOU 3 BURGESS SHALE FOSSILS—-WALCOTT 15 


NARAOIA SPINIFER, new species 
Plate 15, fig. 1 


Three specimens referred to this species are known, of which the 
one figured shows best the spines on the margin of the posterior 
dorsal shield; another preserves both the carapace and dorsal shield, 
the latter having marginal spines while the carapace has a smooth 
margin. On the third specimen the test of the dorsal shield is nearly 
all exfoliated on the left side so as to expose the body, several of the 
fringed exopodites, and coming from beneath them the distal portions 
of the endopodites. 

This species differs from Naraoia compacta in having eight short, 
small spines on the outer margin of the dorsal shield equally spaced 
between the anterior margin and a large posterior median spine ; all 
three of the known specimens of the posterior dorsal shield are also 
larger than those of N. compacta, as they average 25 mm. in length 
exclusive of the posterior median spine. The one specimen preserving 
the carapace indicates that it was similar to the carapace of N. com- 
pacta, the recognized differences between the two species being con- 
fined to the posterior dorsal shield. 

Holotype —U. S. N. M., No. 83946., 


BURGESSIA Walcott 1912 
BURGESSIA BELLA Walcott 
Plate 15, figs. 4-7; plate 16; plate 17; plate 18, fig. 1 


Burgessia bella Walcott, 1912, Smithsonian Misc. Coll., vol. 57, no. 6, p. 177, 
pl. 27, figs. 1-3; pl. 30, figs. 3-4. 

Since the publication of the original description of Burgessia bella 
Walcott, a large number of more or less well preserved specimens 
have been collected from the Burgess shale, a few of which preserve 
details of structure that make it possible to draw a diagrammatic resto- 
ration indicating the increase in our information of the cephalic and 
thoracic appendages (text figs. 3, 4). 

Exoskeleton—The exoskeleton is very delicate and the carapace 
is so thin as to be almost membranaceous. The segment to which the 
telson is attached appears to be partly covered ventrally by a heart- 
shaped plate that is attached to the anterior margin of the segment 
or to the posterior limb-bearing segment of the thorax; it suggests a 
supra-anal plate. 

There appear to be five cephalic, eight thoracic, and one abdominal 
segment, also a long, slender telson with numerous joints. One ex- 
ample 21 mm. in length has 30 joints. 


2 


16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 








o 
{\ 





1 


if 

| 
a 
a 
a 
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Fic. 3.—Burgessia bella Walcott. 


a, antennae; ap, anal plate; as, anal somite; c, carapace; d, doublure; en, 
endopodite; ex, exopodite; /b, labrum; m, mandibles; m’, maxillae; m”, maxil- 
lulae; ¢, telson. 

(About 7.) Diagrammatic outline of ventral side with appendages. Much 
of the data on which this figure is based are shown by the specimens illustrated 
on plates 16 and 17. The exact form and position of the proximal joints of the 
cephalic limbs is unknown, but their general outline and position are about as 
outlined. The thoracic limbs, especially the endopodites, are well preserved in 
several specimens, but the exopodites are rarely seen and then only as faint and 
delicate impressions on the shale. The protopodites are fairly well defined, also 
the trunk segments and telson. The posterior endopodite and exopodite both 
differ from those anterior to them. 


NOS BURGESS SHALE FOSSILS—WALCOTT 17 


Carapace.—Carapace semicircular, with a deep notch on the poste- 
rior side. It appears to have had an upper (dorsal) and lower (ven- 
tral) membrane between which the irregular ramifications of the 
hepatic caeca were located. The shell glands, so conspicuous in the 
Apodidae, have not been recognized in Burgessia. 

Labrum.—tThe labrum is attached to the reflected anterior rim 
(doublure) of the ventral side of the carapace and extends back 
nearly one-third its length ; the labrum is rounded posteriorly and has 
a shallow obliquely transverse furrow on each side just in advance of 
its posterior margin; it was thin, readily distorted by pressure and is 
rarely preserved; one of the best exantples is illustrated by figure 3, 
plate 17 ; it appears to have covered the anterior portion of the mouth. 

Eyes.—The eyes are indicated by a minute round spot on each side 
of the dorsal median axis of the carapace and a short distance within 
the anterior margin. 





Fic. 4.—Diagrammatic outline of a thoracic limb of Burgessia. 


pr, protopodite ; en, endopodite ; ex, exopodite. ; 
The outline is based on the examination of many specimens aided by the known 
form of the endopodite of Marrella which is somewhat similar. 


Dimensions.—The average length of the larger specimens is about 
1o mm. A few are 12 and many are 6 to 8 mm. in length exclusive 
of the long telson. The relative proportions of the carapace, thorax, 
and abdomen are indicated by the diagrammatic restoration (text 
fig. 3). 

Cephalic appendages.—These consist of well-marked antennae (figs. 
3, 4) and three pairs of limbs situated between the antennae and the 
hepato-pancreas tubes; there is evidence that the basal or proximal 
joints of the cephalic limbs are relatively large and the remaining 
joints slender, but their exact position in relation to the labrum and 
their details of form and structure are not determined. It is quite 
probable that they represent the mandibles, maxillulae, and maxillae 
very much as in Marrella, and I have so represented them in the 
diagrammatic restoration of the ventral view of the species (text 
fig. 3). 

Thoracic limbs.—The ten pairs of biramous thoracic limbs are uni- 
form in character with the exception of the posterior pair, which are 


18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


relatively smaller and more slender. Each limb has a strong proximal 
joint (coxopodite) to which the endopodite is attached; the latter is 
formed of four stout joints and two slender joints with two or three 
short spines at the end of the distal joints ; the four joints between the 
proximal and slender sixth joint may have a flattened extension on 
the ventral side as in the endopodite of Marrella that gives them a 
greater transverse diameter, and this may also occur in the sixth joint ; 
the distal joint is slender and probably cylindrical; the exopodite has 
not been seen attached to the protopodite, but from a number of speci- 
mens showing their position there is little doubt of their having been 
attached as on the thoracic limb of Marrella. The exopodite is an elon- 
gate oval, apparently unjointed lobe as seen in the specimen repre- 
sented by figure 4, plate 17; a fringe of fine, short filaments occurs 
on the ventral and outer margins; the delicate structure and small 
size makes it difficult to determine its exact nature, but as far as known 
it recalls the exopodite of Neolenus. One specimen indicates that there 
may have been an anterior support for the exopodite that extended 
beyond the flat filamentous lobe and terminated in two minute spines ; 
the proximal portion of the endopodites has been flaked off in this 
specimen so as to expose the exopodites ; the slender distal extensions 
may belong to the exopodites or they may be the ends of the endopo- 
dites of the opposite side flexed under. I am inclined, however, to 
think that they belong with the exopodites. What may be a modified 
exopodite has been seen in one specimen; it projects from beside the 
posterior thoracic endopodite and consists of a central axis with seven 
sharp spines projecting from its posterior side and a terminal spine ; 
or it may be an endopodite showing the edges of plate-like joints in the 
same manner as those of Marrella splendens (pl. 22, figs. 6, 7). 
Digestive organs ——The mouth was situated at the ventral side and 
probably bounded in front by the labrum and on the sides by the 
mandibles ; the mouth presumably opened into a gullet that passed 
into a large stomach apparently divided or forked anteriorly ; from 
the rear of the stomach a straight intestine extended back to the anus. 
A strong, relatively large tube is given off from each side of the 
stomach at about the fifth segment; these have strong branches at 
the proximal end, one extending forward and another backward, both 
of which have short bifurcating branches on both the outer and inner 
sides. In nearly all well-preserved specimens the large tube and often 
the large connecting tubes are rounded as though they were distended 
when buried in the sediment; this would accord with the view that 
these were large digestive glands that contained food in process of 


NOT 3 BURGESS SHALE FOSSILS—-WALCOTT 1g 


digestion, the ultimate or hepatic caeca secreting a digestive juice as 
in Lepidurus and other crustaceans having such glands.’ 


oat 


/\ 


/ 


ANN AS 


iW 
al sf 
a7 
) 
f 





Fic. 5.—Burgessia bella Walcott. 


ap, anal plate; as, anal somite; cs, central stomach; d.g., digestive glands; 
d.g'., lateral digestive glands; e, eye; h.c., hepatic caeca; 1, intestine; s’ and s”, 
anterior lobes of stomach; sc, anterior central lobe of stomach; f¢, telson. 

(About 7.) Diagrammatic outline of digestive organs. Most of the data 
on which this figure is based are shown by the specimens illustrated on plate 16, 
figures I, 3 and 4. The exact relations of the anterior central lobe of the stomach 
to the central stomach are unknown as there appears to be a line separating them. 


The anus is supposed to have been at the last thoracic 
beneath the platelike structure shown on figure 3, plate 17. 


segment 


Parker and Haswell, Text-book of Zool., vol. 1, p. 491, 1897. 


20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


Functions of appendages.—The functions of the cephalic and 
thoracic limbs were probably similar to the functions of those of 
Marrella splendens. 

Mode of occurrence.—This delicate and beautiful little crustacean 
occurs abundantly in association with Marrella splendens and Waptia 
fieldensis, and is quickly recognized, even when distorted and crushed, 
by its carapace and strongly marked hepatic caeca. The carapace is 
almost always outlined on the shale, as are frequently the body and 
telson; the large strong endopodites are usually more or less clearly 
marked, although their jointed structure has generally been lost in the 
flattening in the shale; the exopodites were so delicate that they are 
rarely preserved, and the same is true of the labrum and eyes. 

Comparison with crustaceans —Burgessia has certain characters in 
common with Marrella and Naraoia and belongs in a group with them 
which will be discussed later (p. 37). 

Diagrammatic restorations of ventral surface ——I have endeavored 
to present, in text figures 3, 4, and 5, interpretations of the structure 
of my Burgessia bella. 

Plesiotypes—U. S. N. M., Nos. 83947a-0. 


WAPTIA Walcott 1912 
WAPTIA FIELDENSIS Walcott 


Plate 18, figs. 2-5; plate 19; plate 20; plate 21, fig. 2 
Waptia fieldensis Walcott, 1912, Smithsonian Misc. Coll., vol. 57, no. 6, 1912, 
p. 181, pl. 27, figs. 4, 5. 

The general characters of this species were described in 1912, since 
when a large number of specimens have been collected from the Bur- 
gess shale, a few affording data from which a fairly accurate dia- 
grammatic restoration of the animal may be drawn (text fig. 6). 

Exoskeleton—The exoskeleton of the carapace, trunk, and caudal 
furca was very thin and readily distorted. The trunk consists of 5 to 7 
short cephalic segments ; 8 narrow thoracic segments, each bearing a 
pair of uniramous appendages; 6 long abdominal segments and 2 
broad lobelike terminal caudal furca or rami; the latter have three 
transverse lines indicating four fused segments. The posterior margin 
of the abdominal segments bears four or more strong spines with a 
fringe of small, short, sharp spines between them. The last or anal 
segment has a minute anal opening on a slightly rounded elevation 
near its posterior ventral margin. The abdominal segments have often 
been narrowed and lengthened, or broadened and shortened by dis- 
tortion in the shale. 


NO. 3 BURGESS SHALE FOSSILS—-WALCOTT 21 


Carapace.—The carapace when viewed from its dorsal side is elon- 
gate, narrowed anteriorly, expanded posteriorly, and has the outline 
of two broad lobes by the incurving of the rounded posterior margin 








































oS 
aS 
sa 


: rE 


BL eiae on 


Fic. 6.—Waptia fieldensis Walcott. 


a, antennae; a’, antennules; a./., abdominal limbs; a.o., anal opening; ¢, 
carapace; c.f., caudal furca; c./., cephalic limbs; e, eye; en, endopodite (??) ; 
ex, exopodite; h.c., hepatic caeca; i, intestine; 7. p., rostral plate; sh. gl., shell 
gland; st, stomach; th./., thoracic leg. 

(About X 3.5.) Diagrammatic outline of ventral view of appendages and 
digestive organs. Most of the data on which this figure is based are shown by 
specimens illustrated on plates 19 and 20. 


towards the median line; when folded over on its longitudinal axis 
each side is a long semi-oval with the narrow end in front; there is 


no evidence of the presence of a longitudinal median line or hinge 
in the many specimens collected from the Burgess shale. 


22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


Rostral plate-——A small triangular rostral plate with narrow, sharp, 
longitudinal median ridge has been seen in four specimens (see fig. 3, 
pl. 19; fig. 2, pl. 20); it is located in the median line between the 
antennae, 

Dimensions.—The average length of an entire adult specimen is 
from 40 to 50 mmn., the carapace being about 16 mm. in length with 
a width when flattened of 15 mm. The general proportions of the 
various parts are shown by figure 4, plate 27, of the 1912 paper. 

Eyes.—tThe eyes are relatively large and placed at the end of a 
stalk or peduncle that projects from beneath and on each side of the 
rostral plate as seen from above. The peduncles are slender at their 
proximal end and expanded in a broad oval outline on the distal third 
of their length, the expanded section carrying the elongate oval visual 
surface ; the peduncle appears to have had at least one joint at about 
the inner third of its length and to have been attached to a prostomium 
at its proximal end. 

Cephalic appendages——These consist of a pair of long jointed 
antennae that project forward beside and beneath the median rostral 
plate, and a pair of short lobelike antennules appear to be represented 
close to the eye and above the antennae (see fig. 3, pl. 19; fig. 2, pl. 20) 
in several specimens.. Traces of three pairs of cephalic limbs have 
been observed but their structure and form are unknown. 

Thoracic limbs ——A number of specimens have five strong thoracic 
limbs that extend from their union with the body trunk forward and 
outward beyond the edge of the carapace (see fig. 2, pl. 18) ; the distal 
joint is short and has three strong and two small curved spines project- 
ing from its outer end and fine spines along its margin; the three next 
joints are rather short and spiniferous, but the detailed character of 
the remaining joints is unknown. The limb observed is assumed to be 
the endopodite of a biramous limb, but the exopodite was not de- 
veloped or it was so small and delicate as not to be preserved in the 
fossil state. 

Abdominal limbs —Each of the abdominal limbs is represented by 
long, multi-jointed exopodites bearing long, slender filaments (see 
fig. 3, pl. 20). The proximal joint was probably short and without 
fringing filaments, but none of the specimens proves this to have been 
the case ; the exopodites are rather large at the proximal end, tapering 
gradually to a slender, flexible terminal section ; the filaments of the 
terminal section are sometimes gathered in tufts or bundles as shown 
by figure 3, plate 20. The filaments are usually flattened and matted 
together, but a few specimens show them to have been slender, 
cylindrical tubes similar to the filaments on the exopodite of Marrella 





= 


NO. 3 BURGESS SHALE FOSSILS—WALCOTT 23 


Splendens (see restorations). The presence of a rudimentary endop- 
odite is suggested on some specimens by an elongate, triangular, 
light-colored space on the proximal portion of the exopodites as shown 
by figure 3, plate 20; these light areas may be the outline of a space 
inside the broad arm of the exopodite, but they usually cross the axis 
of the arm diagonally ; if they do represent the endopodite they were 
exceedingly delicate and attached by a broad base beside the exopodite 
in such a manner as to be held almost rigidly in place, and they are 
always in the fossil state pressed against the proximal section of the 
exopodite, and they have a silvery sheen so characteristic of the con- 
tents of the inside of the limbs of all crustaceans of the Burgess shale 
preserving the limbs. I do not think that they represent the endop- 
odites, but they are the only suggestion of the latter thus far observed 
in connection with the abdominal limbs of Waptia. 






rp-- 





ie} 
i 
\\) 
| 


A 









\\ 
i 


\ 
\\\ 
LL 

i 


<I + 
ee 
\ ! 
[Ue 


“<i 


So 


I 
Is 











s 









Fic. 7.—IWaptia fieldensis Walcott. 


a, antennae; a.o., anal opening; c, carapace; c.f., caudal furca; e, eye; ex, 
exopodite; /i.c., hepatic caeca; 1, intestine; r./., rostral plate; sf, stomach; 
th. 1., thoracic leg. 

(X 3.) Diagrammatic side view of a section of the animal, illustrating the 
appendages, digestive tract, etc. 


Functions of appendages.—The functions of the antennules and 
antennae were presumably sensory as in recent Malacostracans, as 
they do not appear to have been modified for any other purpose, and 
the proximal joint, as far as known, did not function as a manducatory 
organ. 

The mandibles, maxillulae, and maxillae are unknown; the five 
pairs of thoracic limbs may have been used for crawling on the bottom, 
but with short joints and spinous distal joint they could not have been 
very effective; the exopodites of the eight pairs of abdominal limbs 
served as natatory organs and also as branchiae, the long delicate fila- 
ments presenting an extended surface area to the water. 

Digestive organs—What is known of the digestive system of 
Waptia indicates that it was somewhat similar to that of the living 


24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


Apus. The elongate globular stomach (st), with the small digestive 
or hepatic glands (d. gl.), long simple intestine (7) terminating on the 
last segment at the anus (am) all suggest corresponding organs in 
A pus, and it is highly probable that the mouth was ventral and com- 
municated with the stomach by a gullet extending upward and for- 
ward. The shell gland (sh. gl.) or renal organ is distinctly marked 
in several specimens and, as far as comparison is possible, is not 
unlike that of Apus. 

Observations —Waptia was a pelagic, free and active swimming 
animal using its abdominal limbs and the broad terminal rami for 
propulsion. The fact that it is found in association with algae and 
sponges is explained by the conclusion that the sessile forms of life 
were detached and drifted into the Burgess pool and deposited along 
with the pelagic forms that dropped to the bottom of the sea.’ 

The carapace of Waptia is much like that of Hymenocaris except 
that it is not separated into two equal parts by a median longitudinal 
hinge line, and there is no evidence of the presence of an adductor 
muscle scar on each side as in Hymenocarts. 


DIAGRAMMATIC RESTORATION OF VENTRAL VIEW OF THE 
BODY AND APPENDAGES, WITH OUTLINE OF 
DIGESTIVE ORGANS 


Most of the data on which the restoration is based is shown by 
the specimens illustrated on plates 18, 19 and 20. The form and 
position of the cephalic appendages are unknown with the exception 
of the antennae and possibly antennules; the proximal joints of the 
thoracic and abdominal limbs are outlined on the specimens though 
their form is not preserved, but otherwise the limbs are fairly well 
known. The body cavity is outlined by figure 3, plate 18, but it is 
not included in this diagrammatic figure. The shell glands (sh. gl.), 
stomach, intestine, and hepatic caeca are outlined, as they represent 
what is known of the digestive organs. 


Plesiotypes—U. S. N. M., Nos. 83948a-e. 


WAPTIA CIRCULARIS, new species 


Plate 21, hg. 3 


A single specimen with a short, rounded carapace was labeled W. circularis by 
Doctor Walcott. As far as the rather poor preservation permits a determination 
it would seem that otherwise it is similar to WW’. fieldensis. 

Holotype —vU. S. N. M., No. 83449. 


* Smithsonian Misc. Coll., vol. 67, no. 5, pp. 219, 220, 1919. Idem, no. 6, p. 265, 
under Habitat. 


NO. 3 BURGESS SHALE FOSSILS—-WALCOTT 2 


on 


SKANIA, new genus 


Description.—Dorsal shield thin, broadly rounded in front and 
tapering from the postero-lateral angles of the cephalic carapace to 
the posterior end of the shield. 

Cephalic carapace transverse with the postero-lateral angles ex- 
tended into spines ; posterior margin arched forward; frontal margin 
reflected to form a doublure to which a small elongate labrum is 
attached. Eyes unknown but indicated by a bright spot on the carapace 
a short distance outward from the side of the labrum. No traces of 
facial sutures. 

Posterior dorsal shield, elongate and formed of 14 or 15 fused 
segments with a more or less distinctly marked border. There is a 
short transverse segment or telson (pygidium) outlined, but whether 
it is free from the next anterior segment is unknown. 

Surface of test apparently smooth. 

Dimensions. —-This genus is based on a small animal, S. fragilis, 
5 to 17 mm. in length. 

Appendages.—There are indications of antennae, three pairs of 
cephalic limbs, and a pair of limbs for each segment of the posterior 
dorsal shield. 

Digestive organs.—An intestine extends from the posterior segment 
forward to the central part of the cephalic carapace where it widens 
out to form an elongate oval stomach. There are traces of hepatic 
caeca adjoining the stomach. 

Genotype—Skania fragilis Walcott. 

Stratigraphic range-—The stratigraphic range is limited to a band 
of dark siliceous shale about 4 feet in thickness forming a part of the 
Burgess shale member of the Stephen formation. 

Observations—The generic name is derived from Skana, the name 
of a glacier in the Mount Robson District, Alberta, Canada. 

The specimens representing the species of this genus are small and 
so thoroughly flattened in the shale that little more than a black film 
remains. This makes it very difficult to obtain details and also leaves 
some doubt as to whether the posterior dorsal shield is formed of 
fused or free segments.’ 


SKANIA FRAGILIS, new species 
Plate 21, fig. 1 


Description —General outline irregularly heart-shaped but subject 
to wide variation owing to distortion by compression. 





*At the head of these notes, Doctor Walcott later wrote, “A_ trilobite, 
CDs W.” 


26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


Dorsal shield Owing to its extreme tenuity there is usually little 
more than a dark film on the shale that has definite outlines, and shin- 
ing through it are traces of the digestive organs and the ventral limbs. 
The transverse cephalic carapace recalls that of Marrella without the 
great median spines; it is often incurved at the center of its anterior 
margin and laterally projects into long backward-curving, spine-like 
extensions that are so tenuous as to suggest that the cephalic carapace 
was formed of a delicate membrane. 





Fic. 8.— Skania fragilis Walcott. 


_ (X7.) Diagrammatic outline of ventral side showing the various parts as 
interpreted from several specimens. No details of the segments of the posterior 
dorsal shield are preserved, but the segments are clearly outlined. The intestine 
is quite definite, also the fact that it contracted at each segment and expanded 
into a stomach beneath the cephalic carapace. Only the proximal portion of 
the limbs is outlined, although fragments of the distal portion are preserved on 
one specimen. 

The thoracic portion of the dorsal shield is clearly segmented in two 
specimens, and traces of segmentation occur in others, but usually 
there is only a black smear on the shale with the outline of the intestine 
showing through it. There appear to be 14 or 15 fused segments and 
possibly a minute terminal segment or telson. None of the 29 speci- 
mens in the collection show the outlines of the median lobe, although 
one has a slight elevation along the line of the intestine. 

Labrum.—Traces of a narrow doublure and small labrum have been 
seen on two specimens; the labrum appears to have been elongate with 


an outline similar to that of the labrum of Burgessia. 


NO. 3 BURGESS SHALE FOSSILS—WALCOTT 27 


Dimensions.—The largest specimen has a length of 17 mm., but 
the average length is from 5 to 8 mm. 

Appendages—One specimen has several thoracic endopodites out 
of place on one side, and other specimens show the proximal joint 
obscurely but sufficiently well to recognize them; another specimen 
has what may have been slender antennae projecting from beneath 
the flattened labrum and posterior to it three pairs of slender ap- 
pendages in which all traces of joints have disappeared ; there are also 
on this specimen several threadlike, silvery lines extending from the 
central axis out to a margin which indicates that the limbs were long 
and slender ; none of the specimens clearly show the exopodites or any 
details of the limbs. Specimens of Marrella and Burgessia often 
have threadlike, silvery lines representing the limbs, these lines being 
the pyritized contents of the joints, the test having disappeared in the 
process of mineralization of the original specimen. 

Digestive organs.—The stomach is represented by an enlargement 
of the anterior portion of the intestine within the cephalic carapace, 
and the intestine extends back to the last segment; traces of hepatic 
caeca also occur beneath the cephalic carapace adjacent to the stomach. 

Observations —This very delicate form was placed, when sorting 
the collections, among specimens of the young of Marrella splendens, 
but it became evident upon close examination that they were quite 
distinct. They have a dorsal shield resembling that of Naraoia. I 
have examined all the specimens in hopes of finding free segments, 
but without results. There is no well-defined border about the pos- 
terior dorsal shield as in Naraoia, but there is a definite margin that 
is unbroken by the extension of the fused segments beyond it. 

The almost complete flattening of all the specimens prevents any 
comparison with the median lobe of the trilobites, and there is no 
indication of facial sutures although there are slight traces of eyes 
on the cephalic shield at about the same place as in Nathorstia.’ 

Holotype —U. S. N. M., No. 83950. 


MOLLISONIA Walcott 1912 
MOLLISONIA ? RARA Walcott 
Plate 21, fig. 4 
Mollisonia ? rara Walcott, 1912, Smithsonian Misc. Coll., vol. 57, no. 6, p. 198, 
pl. 24, figs. 6, 7. 
Original description.—* Of this species there are several fragmen- 
tary specimens. The species differs from M. gracilis, with which it is 


* Smithsonian Misc. Coll., vol. 57, no. 6, pl. 28, fig. 2, 1912. 


28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 85 


associated, in the character of the thoracic segments and pygidium ; 
also, so far as we can determine from this superficial study, there are 
seven segments and the pygidium shows distinct segmentation with 
a denticulated border.” 

A complete individual was found after the two fragments were described in 
1912. The angularity of the shield at the bottom of the specimen as mounted on 


the plate is characteristic as its essential angles and curves are repeated in all 
the specimens referred to the species. 


Plesiotype—U. S. N. M., No. 83951. 


MARRELLA SPLENDENS Walcott 
Plate 22, figs. 1-9 


Marrella splendens Walcott, 1912, Smithsonian Misc. Coll., vol. 57, no. 6, 
p. 193, pl. 25, figs. 1-6, pl. 26, figs. 1-6. 

In the preliminary note of 1912 the general form and appearance 
of the carapace and appendages of Marrella splendens were described 
and illustrated. Since then a large number of specimens have been 
collected, some of which have added to our information both of the 
carapace and ventral side. 

Exoskeleton—The exoskeleton with the exception of the cara- 
pace is very delicate and formed of a series of 31 segments or somites, 
to 24 of which a pair of biramous appendages are attached; also a 
terminal segment of the body forming a minute plate-like telson and 
five segments of the head indicated by the presence of four pairs of 
free appendages and one segment incorporated in the body of the 
carapace ; this is indicated by the anterior lateral free spines of the 
carapace with a pair of sessile eyes. As far as may be determined from 
the compressed fossil specimens the section of the body segments was 
broadly oval with a dorsal stergite and a ventral sternite section, the 
appendages being attached on the lower side on the ventral sternite 
below the margins of the dorsal stergite. 

This does not mean that the eyes necessarily represent the anterior 
segment but that they represent one segment whatever may have been 
its original position. 

Carapace.—Carapace strong, subquadrangular, and with two large 
dorsal postero-lateral, spinelike lobes (fig. 9) comparable with the 
postero-lateral lobes of the carapace of the Apodidae. At each antero- 
lateral angle a strong, backward-curving spine is attached by a close 
suture. These spines complement the great dorsal thoracic spines and 
may be compared with the movable or free cheeks of the trilobite. A 
narrow median carina or ridge extends the entire length of the lateral 
spines and the postero-lateral lobes. 


NO 3 BURGESS SHALE FOSSILS—WALCOTT 29 


Labrum.—The labrum is attached to the strong frontal rim 
(doublure) of the ventral side of the carapace (fig. 9), and extends 
back to within a short distance below the posterior median margin of 
the carapace ; its posterior lateral angles are extended into short, spine- 
like projections and the posterior margin appears to have been pro- 
vided with two short points. The labrum appears to have covered the 





Fic. 9.—Marrella splendens Walcott. 


Restoration as described in text: a, antennae; c.sp., carapace spine; d, doub- 
lure or reflex of anterior margin of carapace; e, eye; en, endopodite; er, 
exopodite ; m, mandible; mm’, maxillula; m”, maxilla; ¢, telson. 


anterior portion of the mouth ; this is indicated by the proximal end of 
the protopodite or basal joint of the large mandible and that of the 
antennae passing beneath it in many specimens (see pl. 22). 
Eyes—The eyes (fig. 9) have not been seen from the dorsal side 
of any of the several hundred specimens preserving the carapace, and 
rarely from the ventral side; this leads to the conclusions (a) that 
they were located on the lower anterior margin in such a position 
as to be concealed when the carapace was flattened by compression ; 


30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


(b) that they were of a very delicate structure and readily destroyed ; 
(c) that when preserved they were likely to be distorted and displaced 
by compression in the shale and could only be seen from the ventral 
side of the flattened carapace when they projected beyond the margin 
and were outlined on the shale. At first I thought that the eyes were 
situated on the carapace just within the line of its union with the 
large antero-lateral spines. Later I re-examined all the specimens 
showing the eyes, and found two that indicated that the visual surface 
of the eye was on one side of the suture separating the spines from 
the carapace, and the cap or palpebral lobe on the other, and one that 
quite clearly indicated that it was attached to the proximal end of the 
great spines, the latter being equivalent to the free cheeks of the 
trilobite. The interpretation of this is that the visual surface of the 
eye was attached to the great spine outside of the suture that outlined 
the spine from the carapace, and that the cap or palpebral lobe of the 
elevated visual surface of the eye was attached to the carapace as 
in the trilobites with elevated eyes and free cheeks. 

It is difficult to determine the extent of the elevation of the eye 
above the carapace, but from its inconspicuous position in the fossil 
state I strongly suspect that it was only slightly raised and that its 
field of vision was largely forward and downward; this would be in 
accord with the needs of a small, active, free-swimming animal that 
spent little time on the bottom. 

Among the trilobites the eye of Deiphon forbesi Barrande’* is at the 
proximal end of a large genal spine forming the free cheek, and the 
great eye of Bohemilla stupenda Barrande’* occupies nearly the entire 
width of the proximal end of the free cheek which is extended into a 
long strong spine. | 

Digestive organs.—The intestinal canal extends from the posterior 
margin of the labrum back to the small, platelike termination of the 
body ; it is contracted a little opposite the line of union of each of the 
segments (see figs. 6, 7, and 9, pl. 22) ; anteriorly the intestine widens 
out between the labrum and carapace to form what may have been 
the stomach; the narrow canals of the dorsal lobes passed into the 
space between the carapace and labrum and probably entered the en- 
larged intestinal canal as did the canals of the antero-lateral spines 
which appear to pass without interruptions through the close sutures 
that unite them with the carapace; the canals of the postero-dorsal 
lobes may represent the shell-glands or excretory organs of the recent 
Apodidae. 

* Syst. Silur. de Boheme, vol. 1, pl. 2B, fig. 4, 1852; suppl., pl. 2, fig. 19, 1872. 

“Idem, vol. 1, Suppl., pl. 14, fig. 3o. 


INKOG, 6) BURGESS SHALE FOSSILS—-WALCOTT al 


Cephalic appendages.—These consist of antennae, mandibles, simple 
slender maxillulae, and slender maxillae. The proximal joints of the 
cephalic appendages are so badly crushed and matted together beneath 
the labrum or just back of it that it has been very difficult to determine 
exactly their form and relations to each other, but it is highly probable 
that they were arranged as in the restoration (text fig. 9). 

Antennae.—The antennae are long, slender, and many-jointed, with 
fine spines at the distal end of each joint. As far as may be determined, 
the proximal joint was attached to the ventral surface beneath the 
postero-lateral angle of the labrum just in advance of the mandible. 
There is no evidence that it served as a jaw or manducatory organ 
except that in specimens preserving them their inner (proximal) end 
is in front of and adjoining the large proximal joint of the mandibles 
(see figs. 3, 6, and 7, pl. 22). 

Mandibles—The mandibles are formed of a strong proximal joint 
with four short, strong joints followed by five slender, elongate joints 
(see fig. 9), the latter being almost covered with very fine setae that 
give a plumose appearance to the appendage as it extends out beyond 
the great backward curving spines of the carapace. I examined hun- 
dreds of specimens before finding a proximal joint with its inner end 
sufficiently well preserved to suggest the character of its masticatory 
surface; two specimens indicate that it is as shown in figure 7, 
plate 22, and in the restoration. There is no evidence as to whether 
the proximal joint is composed of one long joint or two closely united 
short joints. The usual location of the mandibles in well preserved 
specimens is shown by figures 1 and 2, plate 22. 

Maxillulae—These are long, slender, and with about Io slender 
joints. They look like thoracic legs (endopodite) but their position 
and slender joints serve to distinguish them. Portions of them may be 
seen in figures I and 2, plate 22. 

Maxillae—As far as known the maxillae are formed of joints a 
little longer than those of the maxillula and about the same diameter ; 
both appear to have been slender, rather closely jointed, simple ap- 
pendages as far as the endopodite was concerned; there is strong 
evidence that an exopodite was present, similar to those of the exop- 
odites of the trunk appendages, but they have not been seen directly 
attached to the protopodite; where the parting of the shale is on the 
plane of the exopodites they are usually present next to the mandible 
and directly over the position of the maxillulae and maxillae, which 
suggests strongly that they were present. 

The maxillulae and maxillae were so slender that they are usually 
absent as the result of having been torn off or crushed between the 

3 


32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 85 
strong mandibles and the thoracic limbs. In figure 1, plate 22, their 
exopodites are shown on the left side, and on the right side the endop- 
odites of the maxilla with the exopodite of the maxillula, the endop- 
odite of the latter having apparently been pushed under and a little 
forward of the mandible. Sometimes the endopodite is present but 
the joints are indistinguishable or only a few can be seen. 

Thoracic limbs.—The biramous thoracic limbs appear to be uniform 
in character from the cephalon to the minute plate-like telson at the 
posterior end of the body. Each limb is formed of a protopodite, a 
jointed endopodite, and a jointed fringed exopodite. 

Protopodite.—The large protopodite is attached by its inner end 
to the lower side of the body segment about half way between the 


Vi © ---\---Sectron of Body 





Fic. 10—Diagrammatic outline of the posterior side of one of the anterior 
thoracic limbs: pr, protopodite; en, endopodite; e+, exopodite; f, filaments of 
exopodite ; 7, intestine. 

This figure indicates the point of attachment of the limb to the body, also 
approximate position of the intestine. 


ventral median line and the rounded outer side of the body apparently 
in the same manner as the trunk limbs of Apus, except that in the 
latter there is no evidence that the protopodite served as a gnathobase. 
The protopodite is elongate, apparently cylindrical at its inner end, 
and flattened somewhat at the distal end; it is strong, and supports an 
endopodite and an exopodite. It is usually flattened so as to appear 
of about the same width throughout its length; a few specimens in- 
dicate that it narrowed at its proximal end, essentially as shown in the 
restoration. 

Endopodites—The endopodite or leg is formed of six joints. The 
first five joints of the anterior limbs are rather flat and broad at the 





‘T find that at many places Doctor Walcott changed “ protopodite” to 
“coxopodite.” Whether this term was supposed to have been changed here also 
I was unable to ascertain —C. E. Resser. 


NO. 3 BURGESS SHALE FOSSILS—WALCOTT ns 


sides; narrow and slender on the dorsal and ventral view; short, very 
fine spines occur at their distal end and along the side of the joints. 
The slender distal joint is more nearly cylindrical and has a short, 
strong, slightly curved spine with one or two fine spines beside it ex- 
tending out from the end of the joint. The anterior legs appear to have 
been delicate and slender, but usually they have retained their natural 
position remarkably well. Usually the first joint of the endopodite 
of the fourth pair of limbs is slightly expanded, and the first and sec- 
ond joints of the fifth to seventh pairs of limbs, and the first five joints 
of the eighth to twentieth pairs of limbs. The expanded joints vary in 
degree of expansion from slight enlargement on the fourth limb to 
where the transverse diameter is considerably greater than the length 
of the joint. The latter recall the transverse flattened joints of the 
endopodite of the trilobite Triarthrus becki.’ 





Fic. 11.—Diagrammatic enlargement of a section of the exopodite showing the 
body and the attached cylindrical filaments. 


On some specimens showing the expanded joints the extended por- 
tion is very narrow from base to point, and gives the effect of a strong 
spine projecting from midway of the joint; in other specimens the 
base is as long as the joint and the apex is obtuse, which is the pre- 
vailing form. When in a natural condition the expansion of the joint 
was undoubtedly on the lower or the ventral side, and the fact that 
in the fossil state specimens occur with all the expansions pointing 
forward means only an accident of preservation; some occur with 
scarcely a trace of the enlarged joint, owing to the fact that the ven- 
tral side of the endopodite is buried in the shale, leaving the narrow 
dorsal side in view; in the restoration (fig. 9) I have outlined the flat, 
vertical posterior side of the endopodite. 

Exopodite—The exopodite is attached to the protopodite about 
midway of the length of the latter. It is formed of a long, strong 
proximal joint to which is attached a long, slender, multi-articulate 
appendage, each segment of which supports a long, slender, flat 


* Smithsonian Misc. Coll., vol. 57, p. 137, pl. 30, fig. 20, 1912. 


34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


(formerly round) filament, which is beautifully preserved in some 
specimens (fig. 8, pl. 22). The fringe of filaments often overlap 
those of the adjoining exopodite so as to form an imbricating series 
of fringes the entire length of the body. 

An exopodite 3 mm. in length has attached to it 42 filaments that 
average about 12 to 14 to the millimeter in the proximal portion, 
and 14 to 16 towards the distal end; they increase gradually in length 
from the proximal end until on the distal section they may be as long 
as the entire exopodite. A rusty specimen laid aside as of little value 
proved on cleaning to have the filaments preserved as long, slender 
cylinders or tubes (see fig. 8, pl. 22). 

Most of the fossil specimens. have the fringes extending forward 
and outward, but when the animal was living they undoubtedly ex- 
tended outward and backward so as not to impede its forward move- 
ment. 

As mentioned under Cephalic appendages, there is good reason to 
think that the two posterior oral appendages (maxillulae and maxillae) 
have in addition to the simple jointed endopodite an exopodite similar 
in structure to that of the thoracic limbs. 

E pipodite —A single specimen shows what I thought in rg12 to be 
a large epipodite or branchiae, but which I have now decided to be 
several of the fringed exopodites pressed down together and more or 
less macerated in the contents of the body which were squeezed out 
on that side. It was only after finding a number of examples showing 
the fringed exopodites arranged in this manner but not pressed into 
each other that I gave up the view that a large epipodite was present. 

Several specimens have been found since 1912 that indicate the 
presence of a small oval flattened lobe attached to the dorsal side of 
the protopodite, or it may be to the proximal joint of the exopodite, 
but it may be that this appearance is caused by the manner in which 
the protopodite and the segments of the body are matted down to- 
gether ; some of the thin oval bodies, however, are so clearly defined 
that they suggest the presence of a small epipodite, but I do not 
consider the evidence sufficient to warrant representing them on the 
restoration of the thoracic limb. 

Functions of appendages.—The proximal joints of the antennae 
may have assisted in mastication and may have had a sensory func- 
tion. The proximal joints of the mandibles undoubtedly served in 
mastication, but whether those of the slender leglike maxillulae and 
maxillae aided is undetermined, as nothing has been seen of either 


* Smithsonian Misc. Coll., vol. 67, pls. 20, 32, 1918. 


NO. 3 BURGESS SHALE FOSSILS—WALCOTT 35 


gnathobasic spines or lobes. It is probable, however, from their posi- 
tion and form they were of service in directing food to the mouth. 

The long, flat outer joints of the mandibles may have been used in 
swimming as an aid to the thoracic limbs. 

The endopodite of the biramous trunk appendages probably served 
both as a natatory and ambulatory leg, although from their delicate dis- 
tal joints they evidently were little used in walking. The fringed exop- 
odites may have assisted in swimming and they undoubtedly served as 
gills. The absence of a channel formed of gnathobases on the protop- 
odites of the trunk limbs such as occurs in the Apodidae and Trilobita, 
and also of any known organ for seizing food, leads to the conclusion 
that the exopodite may have served to direct a current of water bear- 
ing food towards the mouth. The long, strong filaments attached to 
the segments of the exopodite are comparable with the filaments of 
the exopodite of the trilobite Neolenus,’ and the remarks on the latter 
should be read in this connection. 

Diagrammatic restoration of the ventral appendages.—This is shown 
in text figure g and presents my interpretation of the arrangement of 
the appendages. The long body, with its numerous segments, is at- 
tached to the carapace in its cephalic region only. The antennae (a), 
mandibles (m), maxillulae (m’), and maxillae (1m”) are drawn in 
from the evidence given by many specimens, no one of which has 
all the limbs in place; several of the best specimens are illustrated on 
plate 22. The eyes (¢) are somewhat theoretically placed, but their 
general position is known despite the displacement most of them have 
been subjected to. On the right side the entire series of protopodites 
and endopodites forming the thoracic limb are represented ; the form 
of the inner end of the protopodite is based on indications afforded by 
several specimens, although usually the protopodite is crushed flat and 
appears to be of the same size throughout its length. On the left side 
the protopodite is cut away so as to show the approximate point of 
attachment of the proximal end of the exopodite. The latter are 
drawn from such specimens as are represented on figures 1-9, plate 22, 
and many others not illustrated. The relative position and form of 
the exopodite, endopodite and protopodite is shown by text figure 10. 

Mode of occurrence—Marrella splendens occurs abundantly in the 
compact, hard shale but there are few really fine specimens. This 
free-swimming, delicate little crustacean dropped down on the surface 
of the bottom and was speedily buried by fine mud settling over it; 
the mass of gradually hardening mud pressed the rounded body into a 








* Smithsonian Misc. Coll., vol. 67, no. 7, p. 370, 1920. 


30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


thin film and matted those parts resting on each other together unless 
there was a thin film of mud between them. When there was such a 
film of mud, it later hardened into shale and formed a plane of weak- 
ness along which the shale parted when, from the action of weathering 
or of force applied with hammer or chisel, the shale was split open. 
Sometimes the parting is between the matrix and the ventral or dorsal 
surface of the specimen, or it may be between the series of fringed 
exopodites and the endopodites ; a number of specimens in the col- 
lection show a part of the endopodites with the exopodites above or 
below them, and again the parting may have been above or below the 
exopodites on one side of the body and the reverse on the other side 
(see pl. 22). The structure of the body and the thoracic appendages is 
very clearly exhibited, but the cephalic appendages, labrum and cara- 
pace, are usually so matted together that it is difficult to distinguish 
the details of structure. 


OBSERVATIONS 


Marrella and the trilobite—Marrella has several characters in com- 
mon with the trilobite and others that are dissimilar. 


SIMILAR CHARACTERS 


A cephalic shield supporting a labrum. 

Sessile eyes on the proximal end of a great spine equivalent to 

the free cheek of the trilobite. 

3. A labrum (hypostoma) with the proximal joints of the cephalic 
limbs gathered at its posterior end in a manner comparable with 
that of the trilobite. 

4. A pair of biramous limbs for each trunk segment formed of a 
protopodite, jointed endopodite (leg), and a jointed exopodite, 
but without any known epipodite. 

5. Expansion of the joints of the endopodites on some of the thoracic 

limbs. 


Ne 


DISSIMILAR CHARACTERS 


1. Absence of a thoracic dorsal shield. 

2. Almost total absence of an abdominal section or pygidium. 

3. Position of proximal joint of antennae. 

4. A large third cephalic appendage (mandible). 

5. The manner of attachment of the coxopodite of each trunk limb 
directly by its proximal end to the side of the ventral surface 
of the body. 


ENO’ 3 BURGESS SHALE FOSSILS—-WALCOTT 3 


“I 


6. The coxopodite did not serve as a gnathobase. 
1 and 2 are considered to be more primitive characters. 
3, 4, 5, and 6 less primitive. 


My present conclusion is that Marrella is a less primitive form than 
the Apodidae, and while a more primitive form than the trilobite it is 
nearer the latter than the Apodidae, and should be grouped near it 
but not with the Trilobita. At the time of my preliminary examination 
of the crustaceans then known to me from the Burgess shale I placed 
Marrella and Nathorstia as progenitors of the trilobite,” but with our 
present information Marrella will be placed with Burgessia, Nathorstia 
being left under Trilobita. 


COMPARISON WITH CRUSTACEANS 


Marrella and the Branchiopoda.—Marrella, with its sessile eyes, 
carapace-like cephalic shield, labrum attached to the doublure, numer- 
ous trunk limbs, and the large mandible, suggests the Apodidae, but 
when we consider the well-developed antennae, large removable spine 
attached to the cephalic shield, biramous trunk limbs on each body 
segment consisting of a fully developed endopodite and exopodite, 
and the absence of caudal rami, the conclusion is that Marrella repre- 
sents a more advanced stage in the evolution of the Crustacea than 
Apus and its allies. The biramous limb of Marrella, like that of the 
trilobite, undoubtedly passed through the foliaceous or multiramous 
limb stage in its evolution, probably in pre-Cambrian time. 

Marrella differs from the Branchiopoda in: 


8 


Absence of lobed multiramous foliaceous trunk limbs with gnatho- 
bases and in the presence of biramous trunk limbs with protop- 
odite, jointed endopodite (leg), and jointed exopodite. 

b. Absence of furcal rami. 

c. Presence of a pair of biramous limbs on each trunk segment back 

to the telson. 


Marrella includes the following characters of the Branchiopoda: 


a. A true carapace arising from a fold of the integument. 


b. A labrum attached to the reflected margin or “ doublure ” of the 
carapace. 
c. A large mandible serving as a jaw in the process of mastication. 


Plesiotypes—U. S. N. M., Nos. 83486a-1. 


* Smithsonian Misc. Coll., vol..57, p. 161, 1912. 


38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


HELMETIA Walcott 1918 
HELMETIA EXPANSA Walcott 
Plate 27 


Helmetia expansa Walcott, 1918, Smithsonian Misc. Coll., vol. 68, no. 12, p. 7, 
fig. 8. 


This is another of the species first published in the Smithsonian explorations 
pamphlet for 1917, and for that reason is here reprinted. 
Holotype —U. S. N. M., No. 83952. 


EXPLANATION OF PLATES 


PLATE I 


Margaretta dorus new SpecieSeirmaccere ince oie ee eee eee 2 
Fic. 1. Impression of outer surface. 

2. A wider and less well defined specimen. Possibly a mutilated 
fragment. 

3. Impression of surface, on which the elevations apparently were 
more conical and more numerous, which may indicate another 
species. 

4. Unretouched photograph by reflected light of the typical form. 
This is again the impression of the outer surface. Compare 
the elongate elevations with the more conical ones in fig. 3. 


5. Unretouched photograph of the holotype. 
6. (Xx 2.) Another, less perfect specimen on which Doctor Walcott 
apparently laid stress. 
PLATE 2 
PAGE 
Miuskowe placida:, New \SPeCieS) Ss.cnoan cushion a sh ee ee ee 4 


Fic. 1. The holotype showing the annulations, the teeth around the 
mouth, and the manner in which the digestive tract seems to 
run along one side of the body, as it is compressed in the 
shale. 


Redoubtia poly podia: NV alcott s oco5 585s cias siace aig a 74 oie 02 © sure eleiieys ons reat 3 
Fic. 2. A specimen, possibly of a different species, since it has smaller 
appendages. Parts of a Hymenocaris? lie above it. 
3. (X1.5.) Holotype, as illustrated previously. 


Wawania corrugaia NVialCOttes case cal oli mie oreiocre) cist crcnevetors Lovelelstersicielere 6 
Fic. 1. Photograph of a very fine individual retouched to bring out the 
striae on the scales. 


PortalazmiracNValcott csc oon eee eins see cee AC Se SO OPE ie etree 3 
Fics. 2, 3. Counterparts of the holotype. A specimen of Miskota lies 
nearby. 


NO. 3 BURGESS SHALE FOSSILS—-WALCOTT 39 


PLATE 4 


Ht SPHOSE) NV AICOU soaccaisie grrr cin sails ane eels ala ea E Seaview Wins ss eve 4 
Fic. 1. (X 1.5.) Specimen having an unusually straight position, with 
all the spines turned backward and thus crowded together. 
2,3. Natural size and enlarged (3) views of a fine complete 
individual. The enlarged picture has been retouched. 


AREOLA ESP INOSO, NVAICOEE 5 2s cc ccrs Ss ecwie Soe a's « fiw sis sie aay oo mais Sie winigiaua ola Scie aie 4 
Fics. 1, 2. Natural size and enlarged (X 4) photographs. The larger one 
has been retouched. 


PLATE 6 


AAA SPinOSD NV AICO sacar Or cca e os cess ae tee We sate ss he dleore needs 4 
Fic. 1. (XX 2.) Photograph of a well preserved specimen. 
2. (X2.) Unretouched illustration of another specimen whose 
specific identity Doctor Walcott doubted somewhat. 


MPC PAT SAN ALCOULS oo. 313 vite Ad OS Sew Da eae Tailed Maw Raed Saws Qed ee 5 
Fic. 3. (X 2.) Unretouched photograph of this peculiar worm. 


CP AIUAIET COULATIS WV. ACO natin see utn Samir nd coos acosened beneege wanes 5 
Fic. 4. Small, somewhat broken specimen. (See pl. 9, fig. 3, for enlarge- 
ment. ) 


5. (X2.) An unretouched photograph of the lectotype. 
6. (2.) Coiled specimen doubtfully referable to the species. 


RPA CAS OTIC CT @- VV BICOUR <aue tc cfc oa s'dpars Sarena sea WG igs Se are e's cos ed Hel we ties 5 
Fic. 1. (2.) Unretouched photograph of a rather complete curled 
specimen. 


4. (X 2.) Unretouched photograph showing the attachment of the 
setae. 


(DOM EATOMO UO 2Oe WVIAICOtE 5 sa. ible ec nectaya ever sin reais ars ciai ers: fede gene S aie ele Sochareiar es 6 
Fic. 2. (X 3.) A good illustration of this small form. 


pata regulars. WalCOtts: 0... ics cwcs cds cadawnecensn bes sume cael secs 5 
Fic. 3. (X3.) Enlargement of specimen illustrated as fig. 4 on preced- 
ing plate. 


40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


PLaTE 8 
PAGE 
Canadia dubtas Walcott) ccc.226 6 oe ne eoreke Cee nae | eR eee 6 
Fic. 1. A fairly large specimen. 
2. (X2.) <A specimen showing the intestinal tract. 
Canadiavsetiger am NVialcotta- see tee ee ee ee EEC COE Eee 5 


Fic. 3. (X3.) Retouched figure of a specimen showing the bundles of 
spines particularly well. 


Pikaia.gracilens. Walcottsaacnisnane ee ee eee Cee eee 7 
Fic. 4. (X 2.) A wide specimen. The illustration is slightly retouched. 
5. (X2.) Another rather well preserved example. 
PLATE 9 
PAGE 
Ottoialmmor Walcottic: cca oacon See ce leeks viele cone Ie Oe EEE 6 
Fic. t. (X2.) <A short form that may have its shape because the mouth 
parts are retracted. 
2. (2.) Small specimen, showing intestinal tract, doubtfully 
referred to the species. 
3. (X2.) Apparently the anterior end of a poorly preserved indi- 
vidual. 
4. (x 2.) Another form like fig. 2. 
5. (X2.) <A specimen with a peculiar restriction that may be 
accidental. 
6,7. Retouched photographs of a particularly fine individual. The 
whole specimen is enlarged somewhat (1.5) and the an- 
terior end considerably (X 4). 
GanadiavdubiaaNWVialcottaces eee cee eee eee 6 
Fic. 8. (x 2.) Unretouched photograph of a small individual. 
Canadia simples, mew SpeCieS.\seieiee dae ellie ete teed eis helt eee eee 6 
Fic. 9. ( 3.) Retouched photograph of the holotype. 
Ganadial grandis; new species sans: eee ee te ene ae eee 5 
Fic.10. (X 2.) Retouched figure of the holotype. 
PLATE 10 
PAGE 
Selkirkia: major \@Wialcott)) 2 sckc se cor cctetie seit etna ee eae ae 7 


Fic. 1. (circa X 3.5.) A large individual, possibly preserving the origi- 
nal proportions of the shell. 

2. (circa X 3.5.) Another specimen whose shell is more or less 

crushed, but with even better preservation of the soft parts. 


NO. 3 BURGESS SHALE FOSSILS—W ALCOTT 41 
PLATE. 


PASEO a PEGUNCUIUTO VWalCOtl oss aAnie cack dais senda 4 kaka lndn bos 3480.0 dels 8 
Fic. 1. (x 2.) An unusually perfect specimen. 
2. Large coiled specimen that Doctor Walcott regarded as possibly 
representing a different animal, but which may appear odd 
only because of its unusual attitude. 


PLATE 12 
PAGE 
ECONGHOIMNG ASUPCTIALde Wi alCOtt jc cero oie aicie suvlei o-oo ei aicl da, nese Ss aw ade on ss 8 
Fic. 1. (X 2.) Retouched figure of a fragmentary specimen. 
2. Natural size unretouched photograph of a well preserved indi- 
vidual. 
3. Enlarged, retouched figure of the counterpart of the preceding 
figure. 
PLATE 13 
PAGE 
Leanchoilia superlata Walcott.............0. cee cee cc cecsenvececcsecees 8 


Fic. 1. (3°). An appendage apparently belonging to this species. 
2. An individual flattened out in the horizontal plane, thus giving it 
an unusual aspect. 


NECOMGHOUIGE MAT ONss NEW {SPCCIES ayes crest ie Seuss bc Gries ea siseree ieee ices sme es 10 
Fic. 3. Illustration of the holotype. 


WWaraoia compacta, Walcott. ... 62c<.0c ss ce cscs ec wes reece secs severe ees 10 
Fic. 4. (X 3.) An unretouched photograph of a good specimen. The 
appendages are only faintly shown extending beyond the 
carapace at the rear of the specimen. (See pl. 15, fig. 3.) 


PLATE 14 
PAGE 
ON COMPACTION VV ALCOLGS 5 cena sn bere oo gag adie ood eb ennes pagans dosages 10 
Fic. 1. (X 2.) Dorsal view of a compressed and somewhat distorted 
specimen, on which the outline of the body trunk is well 
shown and, on the left side, the ends of 16 exopodites pre- 
serving their fine filaments. 
2. (xX 2.) Dorsal view of a specimen with the carapace shortened 
nearly one-half by compression. The central portion showing 
the hepatic glands or caeca is unusually well preserved. The 
posterior dorsal shield has the outlines of the fused segments 
and the outer border clearly indicated, and on the left side, 
where a piece is broken out, two of the exopodites are seen, 
and at the posterior margin, traces of endopodites. 
3. (X 3.) Enlarged view of a specimen with peculiar features 
along the axis. 


Eeanchowha superlata Walcott... 6s <siecs0ccscevecoevacccvsngusboeccecese 8 
Fic. 4. ( 2.) Details of a separate appendage. 
5. (x 2.) Another appendage of a different type. 


42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


PLATE 15 


PAGE 


Naraota spintioronew iSpeCieS.2 achaiiack sae se See ee ee eee ee 
Fic. 1. (X2.) Posterior dorsal shield with traces of segmented body 
and thoracic limbs; a dorsal view of another specimen shows 
clearly defined exopodites with the slender, distal ends of 

endopodites projecting from beneath them. 


15 


Naraoia compacta. Walcotts.. isos naciciecinee ties Sei ae ree eee 10 
Fic. 2. (> 2.) Smaller, retouched picture of the specimen shown in 
fig. 4, pl. 13. 
3. (X 1.5.) Incomplete individual clearly showing the axis. 
Burgéessia bella Walcott): 47s seine & see eee so oes eee 1 
Fic. 4. (XX 3.) Specimen showing the outlines of 10 thoracic legs 
formed of the six joints of the endopodite and the large 
protopodite, also the annulated intestine and fragments of the 
crushed carapace. 

5. (x 4.) A dorsal view of a specimen in which the stomach and 
portion of the intestine as well as the large hepatic tubes are 
distended so as to be moderately convex. The specimen also 
shows the position of the antennae and portions of the hepatic 
caeca. 

6. (xX 4.) A macerated specimen indicating the position of the 
eyes, the antennae, a portion of the labrum, the mandibles, 
the maxillulae, and the maxillae; also the proximal portion 
of seven pairs of the thoracic limbs, with a fair indication of 
the point of attachment of the limbs to the body. 

7. (X3.) Another very fine individual. 

PLATE 16 
PAGE 
Bungessiaivelia, Wialcottaana tes oe oe eee ee eee eer IS 


Fic. 1. ( 2.) Fragmentary specimen indicating structure of limbs. 
(Possibly oriented incorrectly.) 

2. (x4.) A specimen preserving 10 pairs of thoracic limbs, show- 
ing their approximate place of attachment to the body, and 
also their expanded joints and general form. 

The carapace has been crowded back and crushed, but the 
antennae project from its anterior side in an almost natural 
position. 

3. (X2.) <A badly decayed specimen that shows the manner of 
attachment of the limbs. 

4. (x 4.) Dorsal view of a specimen with the digestive organs 
beautifully preserved. These include the intestine posterior to 
the large hepatic tubes, the stomach anterior to the latter, 
and also the anterior and posterior branches of the main 
hepatic tubes and the numerous finer hepatic caeca, which 
occur on both the outer and inner sides of the main branches. 

This specimen gives most of the data for the restoration 
of the digestive organs as shown by text fig. 5. 


NO: 3 BURGESS SHALE FOSSILS—WALCOTT 43 


5. (XX 4.) Specimen showing the expanded joints of the endopo- 
dites of the cephalic limbs. 

6. (> 3.) Partly side view of a crushed and distorted specimen 
illustrated for the purpose of showing the exopodites that 
occur near the body, the proximal part of the endopodites 
with the protopodites having been flaked off from above the 
exopodites in the specimen; in their natural position the 
exopodites were probably above and between the endopodites. 
The outer ends of the long, strong endopodites are well 
shown in this specimen, although the joints have been ob- 
scured. The slender jointed leglike structure associated with 
the exopodites may be the distal part of the endopodites of the 
right side. 


PLATE 17 


TaRP a SEL OCIA NN AICOUE erate cise ee viahaye 6.6: o'ale cisiqin Wikre pantie doh Side oyna d arden wren 15 
Fic. 1. (X3.) Another specimen preserving mainly the * skeleton.” 

2. ( 2.) A small individual crushed obliquely. 

3. (X3.) Ventral view of a specimen preserving the reflected 
anterior margin and labrum, also the antennae, outlines of 
the inner portions of the cephalic limbs, and more or less 
distinctly outlined thoracic endopodites with transversely ex- 
panded joints. The anal plate is clearly indicated, also the 
anterior end of the telson. 

4. (4.) Side view of a crushed specimen preserving on the left 
lower side the outlines of 10 thoracic legs, on the right side 
the outlines of four entire lobelike exopodites and the distal 
end of four posterior to them. The exopodites still show 
slight traces of fine filaments (?) along the posterior and 
outer margins. 


PLATE 18 
PAGE 
PSSA Dela WV BICOtts ma. v a othe soe ete SAG A PUES POSNMA <ribak dias wa 15 
Fic. 1. (X2.) An incomplete specimen showing the manner in which 
the limbs project beyond the carapace. 


Vie iICMn GLA GTISUS NV a lCOLtc. qatsicncse aie vinic etkde-oies citi. sii oh sate asi eieie easels 6 20 
Fic. 2. (X 4.) Side view of a flattened specimen preserving six or more 
cephalic limbs, the limbs from both sides being more or less 
crushed down together. 
3. (X2.) Specimen showing on the right side a flattened uninjured 
antennae in its natural form, and on the left side only the out- 
line of the interior of the joints. The latter mode of occur- 
rence is quite common for the antennae and other appendages. 
This specimen also has the stalked eye preserved on the 
right side. 


44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


4. (x 2.) A few thoracic exopodites in which the flattened fila- 
ments are unusually well preserved. 

5. (x 4.) Side view of a flattened specimen showing the outlines 
of body through the carapace, the stalked eye, antennae, four 
cephalic limbs, and several thoracic limbs. 

This specimen is particularly instructive, as it shows the 
outline of the thoracic limbs from the body to their distal end. 


PLATE 19 
PAGE 
Waepna fieldensis Walcotts. 2.2 seoe onto oe oe Co ero ene Oe 3 20 
Fic. 1. (X 2.) Anterior portion of an individual showing the limbs. 

2. (Xx 2.) Unretouched photograph of a distorted specimen show- 
ing the eye particularly well. 

3. (X 2.) Dorsal view of the carapace of a specimen showing the 
outline of the body, also the small rostrum between the 
the antennae and the eyes. 

4. (X 3.) Specimen with details of eyes and antennae. 


PLATE 20 


Wapiti feldensis: Walcott s,. css sacar cee oe iets ee ao eee eee 20 
Fic. 1. (xX 2.) Posterior portion of specimen preserving the three seg- 
ments, the anal opening, and the lobed, segmented caudal 
furca. 

2. (3 3.) Anterior portion of a carapace, median rostral plate, 
antennae, stalked eyes, and a palp on the left side. 

3. (X< 2.) Side view of a flattened specimen in which a number of 
the exopodites retain the fringing filaments, some of the latter 
being gathered in a cluster at the distal ends. This specimen 
also shows the outlines of elongate triangular, light-colored, 
shiny places that may indicate the interior of the exopodite 
or possibly a rudimentary endopodite. 


PLATE 21 
PAGE 
SRaniaafragtliss Walcott: couccsyoccleoc tine oe ees oo ae oe eos 2 
Fic. 1. (X 4.) Specimen showing the flattened intestine and outline 
of thoracic segments. 


Weaptia: feldensts. NValcotty ..s.a5 ws iesinage © ehcdetavers arp arncio esis te ene eis telerayaieret 20 
Fic. 2. (> 2.) Dorsal view of a flattened specimen in which the cara- 
pace has been removed from over the cephalic area and por- 
tion of the thoracic region. The proximal portion of eight 
thoracic limbs is clearly shown, also of three pairs of cephalic 
limbs. 


NOD 3 BURGESS SHALE FOSSILS—WALCOTT 45 


Te MICN GIT EUIGLIS TOW ‘SPECIES aise sch aie mans «cis ould a diaibe's sie ee secre ue ss 4 24 
Fic. 3. ( 2.) Side view of a flattened specimen illustrating the form 
of the carapace, one side of which has been crushed upward 
and backward. The exopodites of the thoracic limbs appear 
to have the same structure as those of Waptia fieldensis. 


MUNG MUSONTOL SONG. NVAlCOUE sera.) aelees.ere cyst cis 0 2 4S sid orsiaiwg so G's aiesdieibicieie’ere auc 27 


Fic. 4. (X 2.) <A retouched figure of a most excellent specimen. 


PLATE, 22 


ARRCG SPIEndenS WalCOtl a. isc ccscuwicrs sos cdsisies de gees ee oe esa cee tes 28 
Fic. 1. (X 4.) Dorsal view of a specimen showing the mandibles, the 
antenna on the left side extending down beside the mandible, 
and the exopodites referred to the maxillae and maxillulae ; 
what may be the endopodite of the maxillulae is shown on the 
right side below the fringing filaments of the exopodite of the 
maxilla. 

(x 4.) In this ventral view the mandibles have been pushed 
forward so that the mandible and antenna on the right side 
are in a vertical position and side by side, and the endopodites 
of the maxilla and maxillula are in advance of their natural 
position; on the left side two endopodites of thoracic limbs 
are faintly outlined. 

3. (xX 4.) Ventral view of a specimen in which the joints of the 
mandibles have been crowded together and somewhat short- 
ened; the proximal joint on the right side is well exposed by 
the labrum having been pushed forward; the inner margin 
of the joint is serrated but not quite as clearly shown as on 
the specimen represented by fig. 0. 

4. (XX 4.) The dorsal view of this specimen is illustrated to show 
the position of the antenna on the right side which is the 
same as the antenna on the left side in fig. I, and on both 
sides in fig. 5; the position of the mandibles is also well 
shown. 

5. (X4.) Ventral view showing the position of the mandibles 
beside the labrum, also the exopodites of the maxillae and 
maxillulae posterior to them; the antennae appear to have 
been torn away in the crowding forward of the mandibles. 

6. (x 4.) Ventral view of a specimen showing the mandibles, the 
one on the right side preserving the proximal joint with a 
serrated inner margin. This mandible has been colored white 
in order to bring it out more clearly in the reproduction; the 
thoracic exopodites are very clearly shown on both sides, the 
endopodites having been largely exfoliated. 

(X< 4.) Ventral view of a specimen preserving a complete man- 
dible on the left side, also several endopodites of the thoracic 
limbs, and on the right side six joints of the mandible and a 
few imperfect thoracic endopodites, which are exposed on both 
sides by the exfoliation of the exopodites. 


ty 


NI 
. 


46 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


8 (x 6.) Enlargement of the filaments of one of the thoracic 
exopodites in which the filaments retain their original round, 
slender tubular form. This is the only specimen among 
several hundred that I have examined which has escaped 
flattening by compression. This is due to the pyritization of 
the filaments of this particular specimen. 

9. (X4.) Ventral view of a specimen illustrated to show what 
appears to be a small oval flattened lobe attached to the 
dorsal side of a protopodite or it may be to the proximal 
joint of the exopodite; a number of the thoracic exopodites 
with the filaments projecting forward occur on both sides, 
also fragments of the large postero-lateral spines of the 
carapace which lie above the exopodite of the thoracic limbs. 


PLATE 23 


Helmetia expansa Walcott. cnass 2258) so -0s Joe oe ene ee eee 38 
Photograph of the holotype. 


1 


VOL. 85, NO. 3, PL. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS 





BURGESS SHALE FOSSILS 


FOR EXPLANATION SEE PAGE 38. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOEI"85),, INO: 3; (PES 





BURGESS SHALE FOSSILS 


FOR EXPLANATION SEE PAGE 38. 





SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85, NO. 3, PL. 3 


5 ae Pact i 
— 
Se on a 





BURGESS SHALE FOSSILS 


FOR EXPLANATION SEE PAGE 38. 


VOL. 85, NO. 3, PL. 4 


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FOR EXPLANATION SEE PAGE 39 


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FOR EXPLANATION SEE PAGE 39, 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85, NO. 3, PL. 6 





BURGESS SHALE FOSSILS 


FOR EXPLANATION SEE PAGE 39. 


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BURGESS SHALE FOSSILS 


FOR EXPLANATION SEE PAGE 39. 


VOL.85, (NO-°3), PES 8 


SMITHSONIAN MISCELLANEOUS COLLECTIONS 





BURGESS SHALE FOSSILS 


FOR EXPLANATION SEE PAGE 40. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85, NO. 3, PL. 9 


cy * S73] ores 
SRS 4 is ; 





BURGESS SHALE FOSSILS 


FOR EXPLANATION SEE PAGE 40. 


on 


VOL. 85, NO. 3, PL. 70 


SMITHSONIAN MISCELLANEOUS COLLECTIONS 





BURGESS SHALE FOSSILS 


FOR EXPLANATION SEE PAGE 40. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS 


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a 


BURGESS SHALE FOSSILS 


FOR EXPLANATION SEE PAGE 41. 


VOE. 85, 7NO. 3, PES 14 





SMITHSONIAN MISCELLANEOUS COLLECTIONS VOES85, NO 3) PEaat2 





BURGESS SHALE FOSSILS 


FOR EXPLANATION SEE PAGE 41. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL 85, NO. .3;, PE. 13 





BURGESS SHALE FOSSILS 


FOR EXPLANATION SEE PAGE 41. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85, NO. 3, PL. 14 





BURGESS SHALE FOSSILS 


FOR EXPLANATION SEE PAGE 41. 


NO*. 3; PEE 15 


VOL. 85; 


SMITHSONIAN MISCELLANEOUS COLLECTIONS 








BURGESS SHALE FOSSILS 


FOR EXPLANATION SEE PAGE 42. 


VOES 85) \NO- 3; (REZaG 


SMITHSONIAN MISCELLANEOUS COLLECTIONS 





BURGESS SHALE FOSSILS 


FOR EXPLANATION SEE PAGE 42. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS 


BURGESS SHALE FOSSILS 


FOR EXPLANATION SEE PAGE 43. 


VOL. 85, 


NO. 3, PL. 


17 





VOL. 85, NO. 3, PL. 18 


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BURGESS SHALE FOSSILS 


FOR EXPLANATION SEE PAGE 43. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85, NO. 3, PL. 19 





BURGESS SHALE FOSSILS 


FOR EXPLANATION SEE PAGE 44. 


VOL. 85, NO. 3, PL. 2 


SMITHSONIAN MISCELLANEOUS COLLECTIONS 





BURGESS SHALE FOSSILS 


FOR EXPLANATION SEE PAGE 44. 


| SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85, (NO. 3); PL. 21 





BURGESS SHALE FOSSILS 


FOR EXPLANATION SEE PAGE 44. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85, NO. 3), PE, 22 





BURGESS SHALE FOSSILS 


FOR EXPLANATION SEE PAGE 45. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85, NO. 3,\RL. 23 





BURGESS SHALE FOSSILS 


FOR EXPLANATION SEE PAGE 46. 





SMITHSONIAN MISCELLANEOUS COLLECTIONS 


VOLUME 8&5, NUMBER 4 


MEXICAN MOSSES COLLECTED BY 
BROTHER ARSENE BROUARD — II] 


BY 
I. THERIOT 
Fontaine la Mallet, France 








S 
' % he 
es) 





(PUBLICATION 3122) 


CITY OF WASHINGTON 
PUBLISHED BY THE SMITHSONIAN INSTITUTION 
AUGUST 25, 1931 





The Lord Baltimore Presa 


BALTIMORE, MD., U. S. A. 


oe 


MEXICAN MOSSES COLLECTED "BY BROTHER 
ARSENE BROUARD—III 


By I. THERIOT 
FONTAINE LA MALLET, FRANCE 


With the present paper the study of Brother Arscne’s mosses comes 
to an end. In addition to this collection a rather large number of 
species collected by Brother Amable in 1926-27 are included. These 
last are from the Valley of Mexico and adjacent regions. Together 
the two collections comprise about 300 species, among which 18 had 
not been found previously in Mexico, and 44 are entirely new. Brother 
Arséne’s and Brother Amable’s explorations have thus enriched the 
bryological flora of Mexico by more than 60 species, These valiant 
botanists are deserving of great praise and also of thanks from all 
those who are interested in bryology. Referring to the figures given 
by Cardot* the Mexican flora now includes, with the Arséne and 
Amable contributions, about 700 species of mosses. We are therefore 
well above the 400 species enumerated in the Prodrome of Bescherelle. 

It remains now for me to fulfill the very agreeable duty of ex- 
pressing my gratitude to Brother Arsene and Edwin b. Bartram, 
who have assumed with such kindness the task of translating my 
French text into English, and to the Smithsonian Institution for 
publishing my studies. 

The two previous parts were published in the Smithsonian Miscel- 
laneous Collections as follows: Vol. 78, no. 2, pp. 1-29, June 15, 1926; 
vol. 81, no. I, pp. 1-26, August 15, 1928. In the introduction to the 
first of these there will be found a list of localities in the states of 
Michoacan and Puebla from which specimens are cited; this will 
prove useful also for the present and final contribution. 


DITRICHACEAE (continued) 
DISTICHIUM CAPILLACEUM (Sw.) Bry. Eur. 


Valle de México: Desierto (Bro. Amable 1500). 
Very probably new to Mexico. 


eikey: Bryol. 38:97. 1011; ‘40: 33. 1013. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 85, No. 4 


2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


BRYOXIPHIACEAE 
BRYOXIPHIUM MEXICANUM Besch. Journ. de Bot. 6: 180. 1892 
Eustichium norvegicum Schimp. in Besch. Prodr. Bryol. Mex. 29. 1871. 


Valle de México: Desierto (Bro. Amable 1629). 


DICRANACEAE (continued) 
CAMPYLOPUS (Eucampylopus) ANGUSTI-ALATUS Thér., sp. nov. 


Distr. Federal: Rio Frio, alt. 3,000 meters (Bro. Amable 1724). 

Habitu, colore, rete foliorum C. Pittiert R. S. W. sat similis, sed 
differt : caulibus elongatis usque 6 cm., foliis magis concavis, angustis 
(basi 0.5 mm.), acumine angustiore, cellulis laminae minus incrassatis, 
costa angustiore (0.36 mm.). 





| 2 3 
\ 
ae 
KS 
| C 
x 12 


Fic. 1—Campylopus angusti-alatus Thér. 1, leaf; 2, acumen; 3, lamina ceils 
in the upper third; 4, alar and suprabasal cells; 5, cross-section of a leaf near 
the base; 6, cross-section of costa. 


In size this species suggests C. A pollinairei Ther.,’ from Colombia, 
but may be distinguished from it by the leaves, which are erect- 
appressed when dry, little or not flexuous, shorter and only half 
as wide, very narrow toward the base (8 to to rows of cells), and 
by the non-incrassate cells of the lamina. 


CAMPYLOPUS DESTRUCTILIS (C. M.) Jaeg. 


Dicranum destructile C. M. Bot. Zeit. 17: 220. 1850. 
Campylopus Arsenei Thér. Smithsonian Misc. Coll. 787: 5. 1926. 


Having seen from the Berlin Museum the types of C. Chrismari 
(C. M.) and C. destructilis (C. M.), both Mexican species, I have 


* Archiv. Bot. 2: 187. 1928. 


NO. 4 MEXICAN MOSSES—THERIOT 3 


come to the conclusion that they are distinct, the first belonging to 
the subgenus Eucampylopus, the second to the subgenus Pseudocampy- 
lopus; furthermore, that C. Arsene: Thér, does not differ from the 
latter. 


CAMPYLOPUS (Eucampylopus) SAINT-PIERREI Theér., sp. nov. 


Hidalgo: El Chico, on rocks, alt. 2,600 meters, leg. Marius Saint- 
Pierre (Bro. Amable 1589). 

Sterilis. Caespites densi. Caulis 2-3 cm. altus, simplex vel parce 
ramosus, inferne radiculosus, basi terra obrutus, dense foliosus. Folia 
sicca erecta, parum flexuosa, humida erecto-patula, lanceolata, longis- 
sime subulata, canaliculata, marginibus integris, apice denticulatis, 





Fic, 2.—Campylopus Saint-Pierrei Thér. 1, leaf; 2, acumen; 3, upper cells 
at a; 4, suprabasal cells toward b; 5, alar cells; 6, cross-section of leaf near 
the base; 7, section from the acumen; 8, cross-section of costa near the base. 


5-6 mm, longa, 0.7 mm. lata; costa basi 0.3 mm., breviter excurrente, 
dorso superne sulcata, haud lamellosa ; auriculis distinctis, valde exca- 
vatis, cellulis alaribus numerosis, minutis, vesiculosis, rete supra- 
basilari chlorophylloso, cellulis marginalibus (3-4) linearibus, hyalinis, 
sequentibus quadratis, internis rectangularibus, parietibus incrassatis, 
sinuatis; cellulis laminae rhomboidalibus valde incrassatis, juxta- 
marginalibus minutis, juxtacostalibus sensim majoribus. 

At first sight a cross-section of the costa seems to indicate the 
subgenus Palinocraspis. Under a moderate magnification this section 


4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


appears as a thin opaque slice, in which is seen only the median arc 
of eurocysts covered on both sides by cells with a small lumen. 

But, using a higher magnification for the same section, one notices 
that the ventral surface is formed of a unique layer of cells with 
very thick walls showing a lumen like a “ cat-eye”’ (not punctiform). 
These cells, which are not stereids, remind one exactly of those found 
in C. matarensis Besch., from La Réunion Island. Like the last named 
species, C. Saint-Pierrei belongs to the group “ C.b.”* 

Another peculiarity is that the comal leaves often end in a rather 
long hair point, which is discolored or subhyaline, recalling to mind 
that of the group Trichophylli. 


POTTIACEAE 
MOLENDOA OBTUSIFOLIA Broth. & Par. in Card. Rev. Bryol. 40: 36. 1913 


Puebla: Hacienda. Alamos (4631, 4697); Rio Alseseca (574). 
Vera Cruz; Cordoba (10978). Distr. Federal: Mixcoac (9432, 9434, 
9462, 9465, 9466, 9468). Valle de México (Bro. Amable): Desierto 
(1244) ; San Juanico (1261). 

The leaves of this species show a peculiarity which has not yet 
been pointed out: the margins of the acumen are a little thickened 
by the bistratose marginal cells, such as may be seen in some other 
species of the family Pottiaceae. 


MOLENDOA OBTUSIFOLIA Broth. & Par. var. INCRASSATA Thér., var. nov. 

Puebla: Hacienda Batan (5007 p. p.), associated with Trichosto- 
mopsis crisptfolia Card. 

Caulis 1 cm. altus. Folia breviora latioraque, I mm. X 0.25 mm.; 
rete opaco, cellulis incrassatis, costa 50 p. 

Here the thickening of the cell walls is more accentuated than 
-in the type form: toward the apex the lamina is composed of 2 layers 
of cells throughout, while in the middle the thickening is confined 
to 2 or 3 rows of marginal cells. 


ANOECTANGIUM LIEBMANNI Schimp. in Besch. Prodr. Bryol. Mex. 15. 1871 


Morelia: Jesus del Monte (7965). 


ANOECTANGIUM COMPACTUM Schwaegr. Suppl. 1: 36, pl. rr. 1811 


Puebla: (4800). 


*E. & P. Nat. Pflanzenfam. ed. 2, 10: 187. 1924. 


NO. 4 MEXICAN MOSSES—THERIOT 5 


ANOECTANGIUM APICULATUM Schimp. in Besch. Prodr. Bryol. Mex. 16. 
1871 

Puebla: Hacienda Alamos (4770). Morelia: Cerro San Miguel 

(5042). Valle de México: Puente de la Venta (Bro. Amable 1388). 


ANOECTANGIUM EUCHLORON (Schwaegr.) Mitt. Musc. Austr. Amer. 176. 
1869 


Morelia: Loma del Zapote (7508). 


ANOECTANGIUM CONDENSATUM Schimp. in Besch. Prodr. Bryol. Mex. 16. 
1871 
Valle de México (Bro. Amable): Contreras (1483); Desierto 
(11627 )'. 


HYMENOSTOMUM (Kleioweisia) SEMIDIAPHANUM Thér., sp. nov. 


Morelia: Cerro San Miguel, on earth (5040). 

Dioicum? fl. masc. haud vidi. Tenellum ; caulis 1 mm. altus, pauci- 
foliatus. Folia sicca crispula, madida patula, inferiora minuta, caetera 
sensim majora, oblonga-lanceolata, breviter acuminata, acuta, concava, 





Fic. 3.—Hymenostomum semidiaphanum Thér. 1, entire plant, dry; 2, moist 
plant; 3, 4, 5, leaves; 6, acumen; 7, basal cells; 8, cross-section of leaf toward 
the base; 9, cross-section of acumen; 10, fragment of cross-section from acumen. 


superne canaliculata, marginibus integerrimis, valde involutis, folia 
media 1 mm. longa, 0.3 mm. lata; costa basi 40 », percurrente vel 
breviter excurrente ; rete basilari in dimidio inferiore hyalino, cellulis 
rectangularibus, parietibus tenuibus, cellulis laminae opacis, densissime 


6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 85 


papillosis, papillis elatis, diam. 6 ». Pedicellus 3 mm. longus, pallidus, 
siccus valde arcuatus, humidus erectus; calyptra cucullata, dimidium 
partem capsulae obtegens; capsula oblonga, clausa; operculum haud 
secedens, e basi conicum oblique rostratum ; sporae papillosae, 18-20 wm. 

Easily distinguished from the other species of the subgenus Kleio- 
weisia by the length of the pedicels. 


HYMENOSTOMUM MEXICANUM Card. Rev. Bryol. 36: 70. 1909 


Puebla: Road to Cholula. (4849). Valle de México: Tenayuca 
(Bro. Amable 1380). 

These plants conform closely to Cardot’s type except in the in- 
florescence. The type specimen has a paroicous inflorescence (teste 
Cardot) ; here it is autoicous (male flower on a short pedicel under 
the female flower). However, I do not think it is wise to base a 
new species on this difference alone; it seems more logical, in my 
opinion, to conclude that in H. mexicanum the inflorescence is variable, 
occasionally paroicous but more often autoicous, as is normal among 
the species of the genus Hymenostomum. 


GYMNOSTOMUM CALCAREUM Bryol. Germ. 1: 153, pl. 10, f. 15. 1823 


Valle de México: Morales (Bro. Amable 1595), c. fr. 
This species is new to Mexico. 


GYROWEISIA OBTUSIFOLIA (Hampe) Broth. in E. & P., Nat. Pflanzenfam. 
E3389. 1902 

Puebla: Cerro Guadalupe (666); Hacienda Alamos (4634). 
Morelia: Loma del Zapote (7507). 

The leaves of this species are identical as to form and areolation 
with those of Husnoticlla revoluta. But this is the only likeness, the 
leaves of G. obtusifolia being exactly plane on the margin and having 
the costa half as narrow, not dilated at the apex. 


GYROWEISIA PAPILLOSA Ther., sp. nov. 


Morelia: (4927) ; Loma Santa Maria (7887). 

Pusilla, caulis vix 1 mm. Folia erecto-appressa, parum crispula, 
difficile emollita, oblonga vel oblongo-lanceolata, summo late obtusa, 
valde concava, integerrima, marginibus planis, costa basi 30 », superne 
dilatata, sub apicem evanescente, cellulis basilaribus hyalinis, rectangu- 
laribus, cellulis mediis et superioribus quadrato-hexagonis, chloro- 
phyllosis, papillosis, 8-10 », parietibus tenuibus. Pedicellus pallidus, 


NO. 4 MEXICAN MOSSES—THERIOT 


ae 


erectus, 8-10 mm. longus; capsula oblonga, erecta, basi attenuata ; 
operculum conico-rostratum ; annulus latus, peristomium sub orificio 
insertum, dentes papillosi. 











Py 
3 
EL) 

Eby 


at 
ed 






CET) 
EL) 
Ey 





PE 
)=] 
Cr 






amas 
[ze 
yy, 


(a) 
ss 





Fic. 4.—Gyroweisia papillosa Thér. 1, 2, leaves; 3, cells in upper part at a; 
4, middle cells near ); 5, basal cells; 6, cross-section of a leaf; 7, 8, perichaetial 
leaves; 9, capsule; 10, fragment of peristome; 11, annulus. 


Differs from G. obtusifolia by the very unequal, narrower leaves, 
whose cells are more chlorophyllose and papillose, also by the capsule 
with a broader annulus. 


HUSNOTIELLA REVOLUTA Card. Rev. Bryol. 36: 71. 1909 


Puebla: Guadalupe (662, 671, 800); Hacienda Santa Barbara 
(4519, 4520, 4595 p. p.). Morelia: (4922, 7653) ; Andameo (4829, 
4841, 4842, 4845). Distr. Federal: Mixcoac (9441, 9447, 9475 P. P., 
9483). Valle de México (Bro. Amable): Desierto (1249); San 
Juanico (1227) ; Penon (1255) ; San Borja (1259). 

Forma ELATA Ther., form. nov.—Stems longer, up to 10 to I5 
mm., green above, discolored below. Leaves dimorphous, the lower 
short, oval as in the type, those of the young shoots elongated, almost 
twice as long as the lower stem leaves. Valle de México: Contreras 
(Bro. Amable 1333). 


HUSNOTIELLA REVOLUTA Card. var. PALMERI (Card.) Thér., comb. nov. 
Husnotieclla Palntert Card. Rev. Bryol. 37: 121. 1910. 


Puebla: (698) ; Rancho Santa Barbara (4594b). Morelia: Loma 
Santa Maria (7880, 4902). Distr. Federal: Mixcoac (9471, 9475 
Pp. p., 9490). 


8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 85 


In reducing H. Palmeri Card. to a mere variety, following my own 
observations, I feel in perfect agreement with Mr. R. S. Williams 
and also with Cardot himself.’ 


HUSNOTIELLA TORQUESCENS (Card.) Bartr. Bryologist 29: 45. 1926 

Didymodon torquescens Card. Rey. Bryol. 36: 83. 1900. 

Morelia: (7652); Bosque San Pedro (4577, 4583); La Huerta 
(7967) ; Loma Santa Maria (4913, 7882, 7886). 

These specimens differ slightly from the type in being green, not 
glaucous, above ; but the agreeinent is complete in so far as the fruit 
and the form and structure of the leaves are concerned. 


HYMENOSTYLIUM CURVIROSTRE (Ehrh.) Lindb. 


Puebla: Boca del Monte (4737) ; Hacienda Batan, with var. sca- 
brum Lindb. (4960) ; Esperanza (7976). 


HYMENOSTYLIUM INCURVANS (Schimp.) Broth. in E. & P. Nat. 
Pflanzenfam. 1°: 389. 1902 


Gymnostomum incurvans Schimp. in Besch. Prodr. Bryol. Mex. 15. 1871. 

Puebla: Esperanza (4740). Valle de México: Desierto (Bro. 
Amable 1500 p. p.). 

For want of good capsules the determination remains uncertain. 
The male and female flowers are lateral, an abnormal character for the 
genus Hymenostylium. This fact is, however, not unique; I have 
previously mentioned a similar case in connection with H/. curvirostre.’ 


TRICHOSTOMUM CLINTONI C. M. Linnaea 38: 636. 1874 


Morelia: Jesus del Monte (7613 p. p.) ; Loma Santa Maria (7860, 
7874) ; Campanario (7643). 


TRICHOSTOMUM INVOLVENS Card. Rev. Bryol. 40: 34. 1913 


Puebla: Cerro Guadalupe (674, 802) ; Rancho Guadalupe (4612). 

E. G. Paris distributed as T. lamprothecium C. M. a moss collected 
by Bro. Arséne at Puebla which is very different from this species 
and should be referred to T. involvens. 


TRICHOSTOMUM CHLOROPHYLLUM C. M. var. BREVIFOLIUM Thér., 
var. nov. 

Puebla: (4994, 4995). 

Folia breviora latioraque. 

* Rev. Bryol. 40: 34. I913. 

* Bull. Acad. Internat. Geogr. Bot. 11: 319. 1902. 


NO. 4 MEXICAN MOSSES—THERIOT 9 


TIMMIELLA SUBANOMALA (Besch.) Broth. in E. & P. Nat. Pflanzenfam. 


I’: 392. 1902 


Trichostomum subanomalunm Besch. Prod. Bryol. Mex. 33. 1871. 


Puebla: Hacienda Alamos (582, 4513, 4635, 4769) ; Finca Guada- 
lupe (730). Morelia: Campanario (4773, 7523) ; Loma Santa Maria 
(4907). Distr. Federal: Mixcoac (9448). Valle de México (Bro. 
Amable): Desierto (1215, 1251, 1271) ; Rio Frio (1711). Hidalgo: 
EL Chico (1579). 


TIMMIELLA ANOMALA (Bry. Eur.) Limpr. Laubm. Deutschl. £: 590. 1888 


Valle de México: Tenango (Bro. Amable 1686). 

This species, which grows also in California, Arizona, Louisiana, 
and Florida, has not previously been recorded from Mexico. The 
specimens are identical with the European moss, 

T. subanomala is very close to T. anomala, and to my mind seems 
to be only a subspecies or a species of secondary rank. It differs 
from JT. anomala by the position of the male flowers, the form of the 
acumen (which is more gradually narrowed and more acute), and by 
the much broader costa, up to two and a half times as wide. 


TRICHOSTOMOPSIS CRISPIFOLIA Card. Rev. Bryol. 36: 74. 1900 


Puebla: Rancho Santa Barbara (4593, 4594a, 4595, 4599) ; Rancho 
Posadas (4807). Morelia: Cerro Azul (4934, 4936). Valle de 
Mexico (Bro. Amable): San Jeronimo (1593); Tenango (1688). 

Forma CRASSIRETIS Thér., form. nov.—Cellulae laminae valde 
incrassatae——Puebla: Hacienda Batan (5007 p. p.). 


TRIQUETRELLA FERRUGINEA (Schimp.) Thér., comb. nov. 
Barbula ferruginea Schimp. in Besch. Prodr. Bryol. Mex. 37. 1871. 


San Cristobal (F. Muiiller), original specimen, Puebla: Boca del 
Monte (Purpus, in hb. Cardot), (Arséne 4726) ; Esperanza (4742). 
Valle de México: Desierto (Bro. Amable 1419). 

The leaves are triquetrous, squarrose-spreading when moist; the 
areolation and papillae suggest the genus Leptodontium. I found in 
no. 4726 a single capsule. The fruit of this species not being known, 
I describe it: 

Folia perichaetialia erecta, vaginantia; pedicellus flavidus, 1 cm. 
altus ; capsula parum inclinata, oblongo-cylindrica ; operculum oblique 
rostratum. 


Io SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL: 85 


LEPTODONTIUM ANGUSTINERVE Thér., sp. nov. 


Puebla: Esperanza (4741 ; 4743). 

Sterile. Caespites sat densi, intense virides. Caulis 4-5 cm. longus, 
simplex vel parce ramosus, laxe foliosus. Folia sicca erecta, parum 
patula, flexuosa, madida patula, oblongo-lanceolata, sensim acuminata, 
acuta, carinata, decurrentia, marginibus integerrimis, in medio reflexis, 
1.2 mm. longa, 0.3 mm. lata; costa basi 45 », percurrente, dorso papil- 





Fic. 5.—Leptodontium angustinerve Thér. 1, 2, cauline leaves; 3, acumen; 
4, middle cells; 5, basal cells; 6, papillae from the dorsal side; 7, cross-section 
near the lower third of leaf; 8, 9, fragments of 7. 


losa; cellulis laminae quadrato-hexagonis, incrassatis, 12X10 p, pa- 
pillosis, papillis 1-2, elatis, obtusis, cellulis apicalibus laevibus, cellulis 
basilaribus rectangularibus, incrassatis, laevibus. 

This species is rather close to L. filescens Hampe, but is distin- 
guished by the narrow acute leaves, the narrow costa, the incrassate 
cells with only 1 or 2 large papillae, the smooth apical cells, and the 
more compact basal areolation. 


LEPTODONTIUM ARSENEI Ther., sp. nov. 


Morelia: Cerro San Miguel (5073). 

Sterile. Caespites sat densi, compacti, fusco-virides. Caulis 4-5 cm. 
altus, dense radiculosus, regulariter foliosus. Folia sicca crispula, 
humida patulo-squarrosa, vaginantia, e basi erecta, breviter oblonga, 
superne dilatata, sat abrupte in acumen angustum, canaliculatum, sub- 
obtusum contracta, marginibus integris, in medio foli revolutis, 
1.3 mm. longa, 0.5 mm. lata; costa basi 70 m, percurrente, dorso pa- 


INOS MEXICAN MOSSES—THERIOT Il 


pillosa, cellulis laminae opacis, vix distinctis, quadratis, 7-8 », dense 
et tenuiter papillosis, cellulis basilaribus juxtamarginalibus linearibus, 
laevibus, hyalinis, internis rectangularibus, papillosis, paulum chloro- 


phyllosis. 





Fic. 6.—Leptodontium Arsenci Thér. 1, 2, 3, cauline leaves; 4, cross-section 
in the acumen; 5, fragment of 4; 6, upper cells; 7, cells in the upper part of 
leaf base toward a; 8, marginal cells of leaf base at b. 


This species also belongs to the group of L. filescens. It may be 
separated from it by the more elongated stems, by the leaves with a 
short sheathing base, by the narrower acumen (subobtuse and not 
apiculate), and the smaller median cells. 


LEPTODONTIUM FILESCENS (Hampe) Mitt. Musc. Austr. Amer. 50. 1869 
Valle de México ; Desierto, upon bark associated with Rozea stricta 
Besch. (Bro. Amable 1444). 
A Colombian species, new to Mexico. 


LEPTODONTIUM SQUARROSUM (Hook.) Par. Ind. Bryol. 732. 1896 
Holomitrium serratum (Schimp.) C. M. Syn. 2: 587. 1851. 
Valle de México (Bro. Amable): Desierto (1431); Rio Frio 


(1396). 
The last plant is a form with strongly undulate leaves. 


LEPTODONTIUM EXASPERATUM Card. Rev. Bryol. 36: 74. 1909 
Valle de México: Rio Frio (Bro. Amable 1690). 


LEPTODONTIUM sp. 


Valle de México (Bro. Amable): Puenta de la Venta (1391): 
Desierto (1439 p. p.). 

The present species, which unquestionably is new, was recognized 
as such independently by my friend Edwin B. Bartram, who will 
shortly publish a description. 


I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


TORTULA SUBNIGRA Mitt. Musc. Austr. Amer. 164. 1869 


The position of this moss has remained uncertain until now. Jaeger 
classified it as a Barbula. Brotherus* thought that perhaps it was a 
Didymodon. Lastly, J. Cardot, who saw its capsules in good condition, 
verified the absence of a peristome and asked~ if it might not be 
convenient to separate Mitten’s species from true Tortula to constitute 
the type of a new subgenus. 

Mr. Bartram and myself think that we must go further. Trans- 
verse sections of the leaves show that the costa possesses the structure 
of the Trichostomoideae and not of the Pottioideae. Another peculiar 
fact is that the lamina is composed of two layers of cells, except at 
the margin. These noteworthy characters, in addition to that presented 
by the sporophyte, justify the creation of a new genus. We are 
happy to dedicate this Mexican genus to our eminent friend J. Cardot, 
whose last and precious works contributed so much to a better under- 
standing of the bryological flora of Mexico, and who has enriched 
it with such a large number of new species. 


NEOCARDOTIA Thér. & Bartr., gen. nov. 


Caulis erectus. Folia sicca crispata, humore patulo-squarrosa, cari- 
nata, acuta, marginibus inferne revolutis, superne serratis ; rete opaco, 
bistratoso ; cellulis basilaribus linearibus, laevibus, hyalinis, caeteris 
minutis, dense papillosis ; costa breviter excurrente, in sectione trans- 
versali e strato medio cellulis eurycystis et stereidis ventralibus et 
dorsalibus composita. Folia perichaetialia perlonga, erecta, vaginantia ; 
capsula erecta, cylindrica, symmetrica, gymnostoma, annulata ; opercu- 
lum breviter conicum, cellulis recte seriatis ; sporae laeves. 

The affinities of the genus Neocardotia are with the genus Lepto- 
dontium, as much through the habit of the plant as by its serrated 
cauline leaves and strongly sheathing perichaetial leaves, 2 to 3 times 
larger than the stem leaves. 

A single species : 


NEOCARDOTIA SUBNIGRA (Mitt.) Thér. & Bartr., comb. nov. 
Tortula subnigra Mitt. Muse. Austr. Amer. 104. 1860. 


Caespites nigrescenti-virides ; caulis 2-4 cm. altus. Folia 3.5-4 mm. 
longa, 1 mm. lata; costa basi 60-80 », dorso rugulosa ; cellulis laminae 
6-7 », paulum incrassatis, papillis minutis, obtusis; pedicellus 1 cm. 


a ap & Pe Nat. Pflanzentam. Tee OF; L902'5 ed. 2 10. 2/735 1024. 
S / 
* Rev. 3ryol. 38: IOI. TOLL. 


NO. 4 MEXICAN MOSSES—THERIOT L3 


longus, sporae diam. 12-18 ».—Often mixed with other mosses, and 
almost always with Braunia secunda. 

Mexico: Loc. class. (Humboldt) ; La Cima (Barnes & Land 373 
p. p.). Valle de México (Bro. Amable): Salazar (1294, 1716) ; 
Contreras (1470). Hidalgo: Mineral del Chico (Orcutt 6841). 
Arizona: Santa Rita Mts. (Bartram 813). 

The plant from Arizona shows some differences as compared with 
the Mexican plant, slight differences it is true, but worthy of mention; 
it is a little more siender, the leaves are more shortly acuminate, and 
their cells less regularly bistratose. 


HYOPHILA DENTATA Card. Rev. Bryol. 40: 36. 1913 
Morelia: (7896). 


HYOPHILA MEXICANA Thér., sp. nov. 


Valle de México: Tizapan, on earth (Bro. Amable 1613 p. p.). 
Sterilis. Caulis perbrevis ; folia 2 mm. longa, 0.6 mm, lata, margini- 
bus integerrimis, planis, superne paulum involutis; cellulis laminae 
> 7, ’ 
hexagonis, papillosis, parietibus tenuibus, diam. 6-7 ,, costa latissima, 
S Tat 
go-100 pw, in mucronem brevem acutum excurrente. 





Fic. 7.—Hyophila mexicana Thér. 1, leaf; 2, acumen; 3, cross-section toward 
the middle; 4, median cells; 5, basal cells. Hyophila subangustifolia Thér. 
6, 7, cauline leaves: 8, median cells; 9, marginal cells toward a; 10, basal cells; 
II, perichaetial leaf; 12, 13, capsules. 


In form, size, and areolation of the leaves this species may be com- 
pared with H. Bescherellei C. M. It differs in the very short stems 
and perfectly entire leaves, and in the hyaline basal cells, with thin, 
soft walls, These last characters distinguish our species from H. frag- 
ilis Card. 


14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


HYOPHILA SUBANGUSTIFOLIA Thér., sp. nov. 


Valle de México: Tizapan, on earth, associated with the preceding 
species and other mosses (Bro. Amable 1613 p. p.). 

Dioica. Caespites incohaerentes laxiusculi, virides. Caulis perbrevis, 
2-3 mm. altus. Folia sicca crispula, humida erecto-patentia, 1.3-1.6 
mm. longa, 0.40—0.45 mm. lata, oblongo-lanceolata, obtusa, breviter 
mucronata, marginibus planis, integris, superne parum involutis ; costa 
60 p» lata, breviter excedente, dorso laevi; cellulis basilaribus hyalinis, 
ad costam elongate rectangulis, ad marginem brevioribus, superiori- 
bus minutis, quadratis vel hexagonis, papillosis, obscuris, diam. 7-8 p. 
Pedicellus erectus, 4 mm, longus ; capsula oblonga, gymnostoma, annu- 
lata; operculum oblique et longe rostratum, capsulam subaequans ; 
sporae papillosae, diam. 18 ». Flos masculus ignotus. 

I can find no better comparison for this than H. angustifolia Par. & 
Ren., from Madagascar. It differs from the latter in its shorter stems, 
oblong-lanceolate leaves (wider, scarcely involute above, and not 
cucullate at the apex), more compact areolation, and longer opercu- 
lum. 

WEISIOPSIS STENOCARPA Thér., sp. nov. 

Valle de México: Desierto (Bro. Amable 1205 p. p.). Growing 
as isolated stems among other mosses, such as Didymodon oeneus and 
Campylium hispidulum var. Sommerfeltii. 


Jl 


Fic. 8.—Weisiopsis stenocarpa Thér. 1, 2, cauline leaves; 3, acumen; 4, me- 
dian cells; 5, basal cells; 6, antheridial bud; 7, perichaetial leaf; 8, capsule; 
g, annulus; 10, fragment of peristome. 





Autoica, pusilla. Caulis ascendens, 2-5 mm. altus. Folia sicca cri- 
spula, humore patula, lanceolato-acuminata, acuta, basi plicatula, 
marginibus planis, integris, 2-2.2 mm. longa, 0.3-0.4 mm. lata, costa 


NO. 4 MEXICAN MOSSES—THERIOT I 


an 


basi 60 », percurrente vel breviter excedente; rete opaco, papilloso, 
papillis densis, minutis, cellulis quadrato-hexagonis, 8-9 p, cellulis 
basilaribus laxioribus, hyalinis, rectangularibus. Folia perichaetialia 
longiora (3 mm.) ; pedicellus erectus, pallidus, 5-6 mm. longus; cap- 
sula angustissime cylindrica 1.5 mm. longa, 0.26 mm. crassa, annulata, 
peristomium sub ore insertum, dentes lineares, integri, tenuiter pa- 
pillosi; sporae laeves, 12 » crassae. Calyptra? Operculum? 

Comparable to WW. stomatodonta (Card.) Broth. in form and size 
of capsule, but very different in its autoicous inflorescence, longer, 
more narrowly acuminate leaves (with margins not involute), looser 
basal areolation, and entire peristome teeth. 

Furthermore, does Cardot’s species, which has the peristome teeth 
divided into two branches, really belong to the genus Weisiopsis? 


DIDYMODON (Erythrophyllum) PATENTIFOLIUS Thér., sp. nov. 


Valle de México: Xoquiapan (Bro. Amable 1676); Mixcoac 
(Arséne 9442). 

Dioicus, tenellus, obscure viridis. Caulis erectus, simplex, vix 
2 mm. altus. Folia sicca crispula-patula, humida patentia, carinato- 





Fic. 9.—Didymodon patentifolius Thér. 1, 2, cauline leaves; 3, median cells: 


x 


4, basal cells; 5, 6, cross-sections in acumen; 7, cross-section of costa near 
base; 8, perichaetial leaf; 9, dry capsule; 10, moist capsule; 11, peristome; 12, 
fragment of peristome. 


concava, lanceolato-ligulata, acuta, marginibus anguste revolutis, in- 

tegris, 1.5-1.6 mm. longa, 0.3 mm. lata; costa papillosa, basi 60 

crassa, percurrente ; cellulis laminae hexagonis, chlorophyllosis, obscu- 

ris, papillosis, 8-9 », basilaribus laxis, hyalinis, teneris, oblongo-hexa- 

gonis, vel rectangularibus. Folia perichaetialia similia sed majora, 
2 


16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


haud vaginata ; pedicellus purpureus, erectus, 10-12 mm, longus ; cap- 
sula oblonga (2 mm. longa c. operculo), annulata, peristomii dentes 
e€ membrana basilari humillima erecti (0.6 mm. alti), fere usque ad 
basin in 2 crura filiformia papillosa divisi; sporae laeves, 12-15 p; 
operculum conico-rostratum. 

Distinguished at a glance from D. oeneus by its small size. The leaf 
margin is so closely revolute that at first sight the border seems to be 
thickened and formed of two layers of cells. 


DIDYMODON OENEUS (C. M.) Schimp. in Besch. Prodr. Bryol. Mex. 28. 1871 
Trichostomum Oeneum C. M. Syn. 2: 628. 1851. 


Puebla: Esperanza (4802). Valle de México: Desierto (Bro. 
Amable 1205 p. p., 1417). 


DIDYMODON CAMPYLOCARPUS (C. M.) Broth. in E. & P. Nat. Pflanzenfam. 
I°: 405. 1902 


Trichostomum campylocarpum C. M. Syn. 2: 628. 1851. 
Valle de México: Desierto (Bro. Amable 1248). 


DIDYMODON INCRASSATO-LIMBATUS Card. Rev. Bryol. 36: 81. 1909 


Morelia: (7914); Rincon (4567); Loma Santa Maria (4886) ; 
Cerro San Miguel (5085). Valle de México: Tlalpan (Bro. Amable 
1348, 1349). 

According to these specimens the peristome teeth are a little twisted, 
and not exactly straight. Furthermore, the cells of the operculum are 
arranged in oblique rows, a character which is also found in Pringle’s 
no. 10588. No. 4567, from Rincon, is a form with acute leaves whose 
margins are less strongly thickened. 


DIDYMODON FUSCO-VIRIDIS Card. Rev. Bryol. 36: 83. 1909 
Valle de México: San Juanico (Bro. Amable 1264). 


DIDYMODON PUSILLUS Card. Rev. Bryol. 36: 82. 1909 
Valle de México (Bro. Amable): Desierto (1207 p. p., 13am 
1615); Rio Frio (1746) ; Xoquiapan (1752). 
A robust form, the stems tall, up to 2 cm., leaves wider at the base, 
inargins plane or slightly reflexed. These plants seem to mark a 
transition toward the last preceding species. 


DIDYMODON DIAPHANOBASIS Card. Rev. Bryol. 37: 125. 1910 
Valle de México (Bro. Amable): Contadero (1363); Rio Frio 
(1704). 


NO. 4 MEXICAN MOSSES—THERIOT Dy, 


DIDYMODON MEXICANUS Besch. Prodr. Bryol. Mex. 28. 1871 


Puebla: Hacienda Alamos (4764). Valle de México: San Juanico 
(Bro. Amable 1322, 1332). 


DACTYLHYMENIUM PRINGLEI (E. G. Britt.) Card. Rev. Bryol. 36: 72. 1909 


Puebla: (9493); Rancho Santa Barbara (4598, 4811). 
These specimens represent a form with less papillose leaves and 
a nearly smooth costa. 


BARBULA BESCHERELLEI Sauerb. 


Puebla: Cerro Guadalupe (686a, 799). Morelia: Punguato 
(5048) ; Campanario (7564) ; Cerro San Miguel (5087). Tlaxcala: 
(613). Valle de México: Desierto, Contadero (Bro, Amable). 


BARBULA BESCHERELLEI Sauerb. var. CRASSINERVIA Theér., var. nov. 


Distr. Federal: Mixcoac (9470, 9473). 

Folia basi cordata, sat abrupte contracta, costa lata, 90 »; folia 
perichaetialia late ovata vel oblonga, breviter acuminata; capsula 
anguste cylindrica. 


BARBULA BESCHERELLEI Sauerb. var. STENOCARPA Card. 


Valle de México: Xoquiapan (Bro. Amable 1749). 


BARBULA ALTISETA Card. Rev. Bryol. 36: 83. 1909 


Tlaxcala: (621, 720). 
A robust form, the stems longer and the leaves less strongly revo- 
lute than usual. 


BARBULA GRACILIFORMIS Schimp. in Besch. Prodr. Bryol. Mex. 35. 1871 
Puebla: Cerro Guadalupe (668, 669, 680, 801). Distr. Federal: 


Mixcoac (9460, 9461). Tlaxcala: Acuitlapilco (725). 
Nos. 680 and 8o1 are more robust forms. 


BARBULA GRACILESCENS Schimp. in Besch. Prodr. Bryol. Mex. 34. 1871 


Puebla: (600). Morelia: (7946, 7948) ; Bosque San Pedro (4577 
P- p., 4578, 4581, 4925, 4926) ; Loma Santa Maria (4888, 4904) ; 
Jests del Monte (7622). Distr. Federal: Mixcoac (9431, 9435, 9454, 


9456, 9458, 9469, 9480). 


18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


Barbula altiseta Card., B. graciliformus Schimp., and B. gracilescens 
Schimp. are very closely allied species, especially the last two. 

According to authentic specimens 8. gracilescens is distinguished 
from B. graciliformis by its flexuose leaves when dry (not stiff and 
subimbricated), often narrower, with the acumen a little longer and 
thinner. I have not detected any differences in areolation, costa, or 
recurvature of the leaf margins. And inasmuch as I have found forms 
that could not be definitely connected with either species, | am much 
inclined to believe that these so-called species are in reality but forms 
of a single one. 


BARBULA TERETIUSCULA Schimp. in C. M. Syn. 1: 614. 1849 


Puebla: (g07); Fort de Lorette (4622). Morelia: Loma Santa 
Maria (7864 p. p.). 


BARBULA SUBTERETIUSCULA Card. Rev. Bryol. 36: 85. 1909 


Puebla: Rancho Posadas (4808). 


BARBULA BOURGEANA Besch. Rev. Bryol. 36: 35. 1909 


Puebla: (4996); Rio San Francisco (4999); Hacienda Alamos 
(4637). 


BARBULA ORIZABENSIS C. M. Linnaea 40: 638. 1876 


Puebla: Hacienda Alamos (578); Cerro Guadalupe (4616) ; 
Rancho Guadalupe (4590 p. p.). Distr. Federal: Muixcoac (9464, 
9467 ). 

All of these plants are sterile but are identical with the type, which 
I have been able to examine. 

The author compares this species with B. spiralis Schimp. It is 
distinguished, he says, by its less twisted leaves and its cylindrical 
capsule. These characters are rather intangible and valueless to one 
who has seen a series of specimens of B. spiralis, Happily other 
important and obvious characters are available: In B. orizabenstis the 
margins of the leaves are merely reflexed and not revolute; they are 
plane at the apex, the costa is thinner and not widened in the acumen, 
and finally the areolation is chlorophyllose and papillose almost to 
the base. These characters establish the true position of B. orizabensis 
by the side of B. unguiculata and not of B. spiralis. 

| have noticed in all my specimens, as well as in Pringle’s no. 10574, 
the presence of abundant moniliform propagula in the leaf axils. 


NO. 4 MEXICAN MOSSES—THERIOT 19 


BARBULA SPIRALIS Schimp. in C. M. Syn. 1: 622. 1849 
Puebla: (892) ; Mayorazgo (4673, 5975) ; Cerro Guadalupe (655, 
663, 675, 681, 690, 691, 692, 693, 4619, 4620) ; Rancho Guadalupe 
(728, 4591, 4602 p. p., 4604 p. p., 4607, 4609 p. p.); Rio Alseseca 
(7o1); Malinche (6003). Distr. Federal: Tlaquecomeca (9478) ; 
Mixcoac (9446, 9447, 9449, 9482). Tlaxcala: Acuitlapilco (741, 
742). Morelia: Campanario (7922) ; Andameo (4830, 4843, 4844) ; 
Cuincho (5082) ; Cerro Azul (5053) ; Loma del Zapote (7509). Valle 
de México (Bro. Amable): San Juanico (1231, 1262); Texcoco 
(1287) ; Guadalupe (1228) ; Pefion de los Bafios (1256). 
Of all the mosses known to Mexico this species seems to be the 
most common. 


BARBULA EHRENBERGII (Lor.) Fleisch. var. MEXICANA Thér., var. nov. 


Nuevo Leon: Monterrey (Bro. Abbon 10969). 
A forma typica differt foliis valde revolutis, 


BARBULA (Hydrogonium) RUBRICAULIS Thér., sp. nov. 
Nuevo Leon: Monterrey (Bro. Abbon 10968). 
Sterilis. Caespites densi, glauco-virides. Caulis erectus, simplex, 
ruber, 1.5-2 cm. altus, basi terra obrutus, laxe foliosus. I[*olia sicca 





Fic. 10.—Barbula rubricaulis Thér. 1, 2, leaves; 3, acumen; 4, median and 
marginal cells; 5, basal cells. Barbula Abbonti Thér. 6, 7, leaves; 8, cross- 
section of leaf; 9, acumen; 10, median cells; 11, basal cells. 


erecto-flexuosa, madida erecta, paulum patula, oblongo-ligulata, sub- 
obtusa, concava, decurrentia, marginibus integris, planis vel uno 
latere parce reflexis ; costa basi 60-70 p, apicem attingente, rete chlo 


20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


rophylloso, cellulis subquadratis, 10 x 8 p, laevibus, parietibus incrassa- 
tis, rete basilari laxiore, cellulis juxtacostalibus rectangularibus, hya- 
linis, margines versus quadratis vel breviter rectangularibus. Caetera 
desunt. 

In color of the stems and in form, size, and areolation of the 
leaves B. rubricaulis is very close to B, dialytrichoides Thér., from 
China, differing only in its nearly smooth areolation. I have not seen 
the fruit. 


BARBULA (Hydrogonium) ABBONII Thér., sp. nov. 


Nuevo Leon: Monterrey (Bro. Abbon 10970). 

A B. rubricauli proximo differt: caule viride, foliis siccis valde 
patulis, humidis subsquarrosis, brevioribus, 1.4 mm. longis, 0.60-0.65 
mm. latis, costa latiora, go-100 p crassa, cellulis mediis majoribus, 
15-20 wX 10-15 p. 

The leaves show the median areolation of B. Ehrenbergiana var. 
mexicana and the basal areolation of B. rubricaulis; the leaf margins 
are narrowly revolute three-fourths of the way up from the base. 


BARBULA (Streblotrichum) CALCAREA Thér., sp. nov. 


Morelia: Loma Santa Maria, on calcareous rocks (4891). Valle 
de México: Desierto (Bro. Amable 1620). 





Fic. 11.—Barbula calcarea Thér. 1, 2, 3, cauline leaves; 4, median cells; 
5, basal cells; 6, 7, perichaetial leaves; 8, deoperculate capsule. 


Pusilla. Caulis gracilis, simplex, 3-5 cm. altus, laxe foliosus. Folia 
mollia, sicca appressa, humida patula, elliptica vel oblonga, late ro- 
tundata, decurrentia, marginibus integerrimis, inferne planis, superne 


NO. 4 MEXICAN MOSSES—THERIOT 21 


valde revolutis, 0.9 mm. longa, 0.4 mm. lata; costa basi 60 », continua, 
dorso papillosa; cellulis mediis opacis, indistinctis, dense papillosis, 
diam. 10-12 p, superioribus minoribus, rete basilari laxiore, pellucido, 
cellulis rectangularibus, chlorophyllosis, plus minus papillosis, infimis 
laevibus. Folia perichaetialia pauca, intima 2 duplo longiora, convo- 
iuta, longe vaginantia, apice lingulata, obtusa; pedicellus tenuis, palli- 
do-luteus, 7-8 mm, longus ; capsula (immatura) minuta, anguste-cylin- 
drica ; operculum rostratum, Caetera ignota. 

By its slender habit, its loosely foliate stems, and leaves revolute 
in the upper two-thirds, the present species is immediately distin- 
guished from B. hypselostegia Card. and B. Muenchii Card., both of 
which also have obtuse leaves. 


BARBULA (Streblotrichum) STENOTHECA Thér., sp. nov. 


Valle de México; Rio Frio, on earth (Bro. Amable 1726). 
Dioica. Caespites sat densi, obscuro-virides. Caulis erectus, flexu- 
osus, gracilis, remote foliosus, 10-15 mm. altus. Folia sicca incurvato- 


, (i 


i. 


Hy, 
A 





Fic. 12—Barbula stenotheca Thér. 1, cauline leaf; 2, acumen; 3, median 
leaf cells; 4, basal cells; 5, 6, cross-sections of leaf; 7, cross-section of costa 
near base; 8, 9, perichaetial leaves; 10, dry capsule; 11, calyptra; 12, peristome. 


crispata, madida patula, oblongo-lanceolata, subobtusa, breviter apicu- 
lata, marginibus integris, usque ad apicem revolutis, 2 mm. longa, 
0.6 mm. lata, costa valida, basi 60 » crassa, dorso laevi, breviter ex- 


22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


currente; rete opaco, cellulis hexagonis, 10 pm, parietibus tenuibus, 
dense et minute papillosis, cellulis inferioribus rectangularibus, pellu- 
cidis, laevibus, parietibus firmis. Folia perichaetialia numerosa, ex- 
terna patulo-subsquarrosa, intima convoluta, longe vaginantia, decolo- 
1ata, longissima, 4 mm. longa, obtusiuscula, apiculata ; pedicellus ruber, 
15 mm, longus; capsula erecta, angustissime cylindrica vel arcuato- 
cylindrica, 4 mm. longa, 0.4 mm. crassa; calyptra } partem capsulae 
obtegens ; operculum longe conicum, 1.3 mm. longum; annulus sim- 
plex; peristomium elatum, 1 mm. altum, dentibus valde contortis, 
membrana basilari brevi; sporae laeves, 8-9 vp. 

Differs widely from the other Mexican species of the section Stre- 
blotrichum in size and habit, and especially in the dimensions of the 
capsule. 


MORINIA EHRENBERGIANA (C. M.) Thér., comb. nov. 


Barbula Ehrenbergiana C. M. Syn. 1: 636. 1849. 
Barbula trichostomoides Besch. Prodr. Bryol. Mex. 38. 1871. 
Morinia trichostomoides Card. Rev. Bryol. 37: 124. I910. 


Valle de México: Desierto, on earth (Bro. Amable 1240). 

While studying this specimen I recognized, by a happy chance, its 
identity with Barbula Ehrenbergiana C. M. and with Morinia tri- 
chostomoides (Besch.) Card. The name established by Muller has 
priority, hence the above new combination. 


ALOINA CALCEOLIFOLIA (Spruce) Broth. in E. & P. Nat. Pflanzenfam. 
I°: 428. 1902 


Puebla: (704) ; Mayorazgo (4672). 


ALOINELLA CATENULA Card. Rev. Bryol. 36: 76. 1909 


Valle de México (Bro. Amable) : Desierto (1207 p. p., 1217 p. p.) ; 
Salazar (1296 p. p.). 
Terrestrial, in isolated bits, always associated with other mosses. 


TORTULA PARVA Card. var. LATIFOLIA Thér., var. nov. 


Puebla: (4509); Rancho Santa Barbara (4593, 4600) ; Hacienda 
Alamos (4720). Morelia: Cerro Azul (4933). Valle de México: 
Cartridge Factory (Bro. Amable 1459 p. p.). 

A forma typica differt statura robustiore, foliis longioribus et duplo 
latioribus, 1.2-1.8 mm. X 0.6-0.8 mm. 


NO. 4 MEXICAN MOSSES—THERIOT 23 


‘“TORTULA AMPHIDIACEA (C. M.) Broth. in E. & P. Nat. Pflanzenfam. 
I°: 424. 1902 


Barbula? amphidiacea C. M. Linnaea 38: 639. 1874. 


Puebla: Rancho Santa Barbara (4810 p. p.). Morelia: Pare San 
Pedro, c. fr. (4587) ; Cerro Azul (4932). Valle de México: Conta- 
dero (Bro. Amable 1300, 1308 p. p., 1360). 

The plant from San Pedro bears capsules. The fruit being un- 
known, I describe it: Pedicel short, 6-7 mm., almost hidden by the 


6 
iS 'O BD ww og OOSQYT 
CX OEY FS ESSeE 


2 ie cy Y (s @ \ ) 
<90 an , OOOGS& 


: 0 
ib OO S 9090 
', . A ay ee Oo 
2OQ*gOO0G 


x 200 a 


& 





Fic. 13.—Tortula amphidiacea. 1, 2, stem leaves; 3, cross-section of a leaf; 
4, the same, costa; 5, upper marginal cells toward a; 6, median cells; 7, basal 
cells; 8, propagulae. 


long and numerous innovations ; capsule oblong-cylindrical. All the 
capsules are old and have neither operculum nor peristome. 

The species is well characterized in other particulars: The leaf 
is acute and entire, more or less marginate at the base with several 
rows of narrow cells, the lamina with differentiated, incrassate and 
smooth cells ; the costa is percurrent or short-excurrent ; the stems bear 
oblong propagula. 


TORTULA RIPICOLA Ther., sp. nov. 


Valle de México: Morales, on earth, bank of a small stream, asso- 
ciated with Fissidens tortilis Hampe & C. M. (Bro. Amable 1596 
Das) 

Sterilis, pusilla. Caulis inferne denudatus, superne rosulato-folio- 
sus, 0.5-I cm. altus, in axillis foliorum propagula numerosa, fusca, 
sphaerica gerens. Folia sicca paulum crispula, humore patula, ovato- 
lanceolata, obtusa vel raro subacuta, breviter mucronata, marginibus 


24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


integerrimis, toto ambitu revolutis, 1.5 mm. longa, 0.6-0.7 mm. lata; 
rete opaco, dense et minute papilloso; cellulis quadrato-hexagonis, 
chlorophyllosis, haud incrassatis, diam. 6 p, basilaribus paucis, hyalinis, 
laevibus, breviter rectangularibus, costa basi 60 », dorso minute pa- 





Fic. 14.—Tortula ripicola Thér. 1, 2, 3, leaves; 4, 5, acumens; 6, median leaf 
cells; 7, basal cells; 8, 9, 10, 11, cross-sections of a leaf; 12, propagulae. 


pillosa, in mucronem brevem excedente, structura in sectione trans- 
versali ut in genere. 

Group of T. papillosa Wils. Well characterized by its small size, its 
leaves broadened at base, revolute all around and short-mucronate, 
and its very compact areolation. 


TORTULA FRAGILIS (Tayl.) Mitt. Musc. Austr. Amer. 172. 1869 
Tortula confusa Card. Rey. Bryol. 36: 87. 1900. 
Tortula Pringlet Card. Rev. Bryol. 36: 87. 1909. 
Puebla: (4510); Esperanza (4940). Distr. Federal: Tlalpam 
(9494). 


For the synonymy of this species the reader is referred to a recent 
note by Mr. E. B. Bartram.’ While studying Bro. Arséne’s collection 
I also formed a clear idea that Cardot’s two species could not be main- 
tained, as the author himself apparently suspected. 


TORTULA OBTUSISSIMA (C. M.) Mitt. Musc. Austr. Amer. 174. 1869 


Puebla: Cerro Guadalupe (667, 673). Tlaxcala: (606). Valle de 
México (Bro. Amable): San Juanico (1232) ; Tenayuca (1376) ; 
Xoquiapan (1675). 


‘ Bryologist 29: 53. 1926, 


NO. 4 MEXICAN MOSSES—THERIOT 


to 


cn 


TORTULA OBTUSISSIMA (C. M.) Mitt. var. CONNECTENS (Card.) Thér., 
comb. nov. 


Tortula connectens Card. Rey. Bryol. 36: 87. 1900. 


Puebla: Fort Guadalupe (4621). Morelia: Cerro Azul (4531). 
Distr. Federal: Mixcoac (9485). 

The characters indicated to separate T. connectens from T. obtu- 
sissima do not appear constant, and I combine the two species. 


GRIMMIACEAE (continued) 
GRIMMIA INVOLUCRATA Card. Rev. Bryol. 36: 105. 1909 


Valle de México (Bro. Amable): Tlalpam, c. fr. (1448) ; Zaca- 
tenco, ster. (1352). 


GRIMMIA PRAETERMISSA Card. Rev. Bryol. 36: 105. 1909 


Valle de México (Bro. Amable): Rio Frio, on rocks (1401 p. p., 
1681). 

The capsule is sometimes pale and scarcely exserted, sometimes 
brown, longer-pedicellate, and well exserted. 


GRIMMIA CALIFORNICA Sull. in U. S. Rep. Expl. Miss. Pacif. 4: 187, pl. 4. 
1856 

Valle de México (Bro. Amable): Rio Frio, intimately mixed with 

the preceding species (1401 p. p.) ; Llano Grande, alt. 3,700 meters 


(1724, 1734). 


GRIMMIA PULLA Card. Rev. Bryol. 36: 106. 1909 


Valle de México: Contreras, on rocks (Bro. Amable). 

The plants are fruited, but the over-ripe capsules have lost their 
peristomes. Sporophyte pseudo-lateral, because of the 1-3 rather 
elongated innovations borne under the male flower ; pedicel 2-2.5 mm. 
long, suberect when dry, arcuate when moist ; capsule oblong, strongly 
furrowed. 


SPLACHNACEAE 
TAYLORIA (Eutayloria) TORTELLOIDES Thér., sp. nov. 


Hidalgo: El Chico, 2,600 meters (Bro. Amable 1587 p. p.). Grow- 
ing as isolated stems among other mosses, especially with Bryuwi 
Ehrenbergianum., 


26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


Dioica? Flos masculus ignotus. Caulis brevis, vix 1 cm. altus, 
simplex vel parce ramosus, inferne denudatus, radiculosus, pauci- 
foliatus, apice rosulato-foliosus. Folia sicca valde crispata, nitida, 
humore patentia, oblongo-spathulata, e basi contracta, decurrentia, 
apice rotundata, apiculata, apiculo brevi, obliquo, marginibus planis, 
inferne paulum reflexis, integris vel remote et obtuse denticulatis, 3 
mm. longa, 2 mm. lata; costa basi 120 p, raptim attenuata, sub apicem 
evanescente, in sectione transversali ut in genere; cellulis mediis 
hexagonis, chlorophyllosis, parietibus tenuibus, 60 x 30 », marginalibus 





Fic. 15.—Tayloria tortelloides Thér. 1, lower leaf; 2, comal leaf; 3, upper 
cells at a; 4, median cells at Db; 5, marginal cells toward c; 6, basal cells; 7, 
young dry capsule; 8, moist capsule; 9, wall of capsule orifice; 10, fragment ot 
peristome. 


(2-3 ser.) elongatis, inanis, cellulis basilaribus rectangularibus, parce 
chlorophyllosis. Folia perichaetialia similia, intima minora; pedicellus 
erectus, perbrevis, 1.5 mm. longus, laevis, pallidus; capsula sub- 
cylindrica, brevicollis, 2 mm. longa; operculum obtuse conicum, colu- 
mella inclusa, peristomii 16 dentes liberi, opaci, dense papillosi, 0.4 
mm, longi; sporae laeves, 15-18 » crassae. Calyptra? 

The extremely short pedicel and the entire leaves, rounded apicu- 
late, broadly spatulate, and shrivelled when dry (like some Tortula), 
readily distinguish this plant from the other species of the subgenus 
Eutayloria. 





NO. 4 MEXICAN MOSSES—THERIOT 


No 
N 


BRYACEAE (continued ) 
MIELICHHOFERIA SAINT-PIERREI Thér., sp. nov. 


Valle de México: Lerma; leg. Marius Saint Pierre (Bro. Amable 
1685). 

Paroica, laxiuscula caespitosa, tenella, viridis. Caulis julaceus, 
2-3 mm. altus, ramis erectis, vix 5 mm. longis. Folia caulina conferta, 
imbricata, ovato vel oblongo-lanceolata, acuminata, acuta, 0.8-1.2 mm. 
longa, 0.4-0.5 mm. lata, marginibus parce et anguste reflexis, superne 
remote denticulatis vel sinuolatis, costa basi 30-36 » crassa, subpercur- 





Fic. 16.—Mielichhoferia Saint-Pierrei Thér. 1, 2, 3, stem leaves; 4, leaf from 
an innovation; 5, median cells; 6, apical cells; 7, basal cells; 8, moist capsule; 
9, fragment of peristome and annulus. 


rente, rete membranaceo, cellulis elongate rhomboideis, 60-70 px 12 p, 
marginibus angustioribus, basilaribus rectangularibus; folia ramea 
angustiora, marginibus erectis. Folia perichaetialia caulinis similia ; 
pedicellus erectus, 8-12 mm. longus; capsula inclinata vel subhorizon- 
talis, symmetrica, oblongo-cylindrica, collo attenuato instructa ; annu- 
lus latus ; peristomium simplex, membrana basilari subnulla, processus 
angusti, 10 p lati, granulosi haud appendiculati; sporae sublaeves, 
12-15 »; operculum convexum, mamillatum. 


LEPTOBRYUM PYRIFORME (L.) Wils. 


Valle de México: Tlalpam (Bro. Amable 1246 p. p.). 


EPIPTERYGIUM MEXICANUM (Besch.) Broth. 


Valle de México (Bro. Amable): Puente de la Venta (1400) : 
Santa Rosa (1513); Desierto (1642). 


28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 85 


EPIPTERYGIUM MEXICANUM (Besch.) Broth. var. ANGUSTIRETE Thér., 
var. nov. 
Valle de México: Contreras (Bro. Amable 1659). 
Folia angustiora, cellulis chlorophyllosis, angustioribus, 


MNIOBRYUM INTEGRUM (Card.) Broth. in E. & P. Nat. Pflanzenfam. 
ed. 2, 10: 363. 1924 
Webera integra Card. Rey. Bryol. 40: II. 1913. 
Valle de México: Contreras (Bro. Amable 1478). 


WEBERA SPECTABILIS (C. M.) Jaeg. 

Webera cylindrica (Mont.) Schimp. in Besch. Prodr. Bryol. Mex. 52. 1871. 

I have received from Bro. Amable rather numerous collections 
of Webera of the present relationship, but frankly, I have not suc- 
ceeded in distinguishing W. cylindrica from W. spectabilis. The 
characters I had considered distinctive are rarely combined on the same 
plant and all of them show a rather wide variability, as, for instance, 
in the width of the leaf, the recurvature of the margin, the width 
of the costa and of the cells, and the length of the capsule. My con- 
‘clusion is that we must unite the two species. Webera spectabilis has 
priority. 

WEBERA PSEUDO-BARBULA Thér., sp. nov. 

Valle de Mexico (Bro. Amable): Desierto (1630, 1643) ; Con- 
treras (1658) ; Lerma (1684).—In all these localities the plants grow 
in company with Anomobryum filiforme var. mexicanum, a remarkable 
fact of association. 





Fic. 17.—Webera pseudo-Barbula Thér. 1, stem leaf; 2, marginal and median 
cells; 3, apical cells; 4, leaves of innovations; 5, apical cells of 4; 6, marginal 
and median cells of 4; 7, propagula; 8, perichaetial leaf; 9, dry capsule; 10, 
moist capsule; 11, peristome (fragment) ; 12, fragment of annulus. 


NO. 4 MEXICAN MOSSES—-THERIOT 29 


Dioica. Caespites laxi, virides. Caulis brevis, 5 mm. longus, superne 
innovationibus elongatis, 10-15 mm. longis, in axillis foliorum superi- 
orum propagula fusca, numerosa, subglobosa gerens. Folia sicca 
erecta, parum flexuosa, humore erecto-patula, ovato-acuminata, 1.2-1.5 
mm. longa, 0.5 mm. lata, marginibus planis, interdum anguste revo- 
lutis, elimbatis, integerrimis, apice denticulatis ; costa basi 60 p, sen- 
sim attenuata, ante apicem evanescente ; cellulis anguste rhomboideis, 
chlorophyllosis, 70-90 » x 8-9 pw, ad marginem angustioribus ; folia in- 
novationis similia sed minora. Folia perichaetialia longiora, ovato- 
lanceolata, acuminata, intima anguste lanceolata, tenui-acuminata, 
marginibus revolutis, costa percurrente; pedicellus flexuosus, 20-25 
“mm. altus; capsula suberecta vel inclinata, oblonga collo breviore 
attenuata; operculum convexum, mamillatum; annulus latus; exo- 
stomii dentes pallidi, haud marginati, dorso inferne laeves, supernc 
papillosi, 0.27 mm. alti, membrana ad 4 dentium producta, processus 
lineares, fugaces, cilia rudimentaria; sporae diam. 12-15 p. 

Very close to W. didymodontia (Mitt.) Broth., which is distin- 
guished at a glance by its globular capsule. 


BRACHYMENIUM (Dicranobryum) SAINT-PIERREI Thér., sp. nov. 


Valle de México: Contreras, on earth; leg. Marius Saint-Pierre 
(Bro. Amable 1338 p. p.). 

Dioicum. Caulis brevis, 2-3 mm. altus, inferne denudatus, inno- 
vationibus numerosis, clavatis. Folia sicca appressa, oblonga, breviter 





x 120 





Fic. 18.—Brachymenium Saint-Pierret Thér. 1, plant at natural size; 2, stem 
leaf; 3, median cells; 4, lower leaf of innovation; 5, upper leaf of innovation; 
6, median cells; 7, marginal cells; 8, basal cells; 9, moist capsule; 10, oper- 
culum; 11, fragment of annulus, 


30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


acuminata, mucronata, elimbata, marginibus integris, planis; costa 
flexuosa, basi 40 p, breviter excurrente, cellulis longe hexagonis, 
36-60 »X10p, basilaribus quadratis; folia innovationis inferiora 
minuta, caetera sensim majora, valde concava, apice congesta, Folia 
perichaetialia majora, deltoidea ; pedicellus pertenuis, flexuosus, 12-15 
mm. longus ; capsula suberecta vel horizontalis, microstoma, oblongo- 
cylindrica, collo longo attenuata; annulus latus; operculum conico- 
convexum, mamillatum; peristomium externum normale, 0.32 mm. 
altum, internum? (capsulae immaturae) ; sporae laeves, 18  crassae. 

In the size and form of its capsule this species recalls B. rubricarpum 
(Besch.). It differs in its leaves, which are of another form, short- 
mucronate, with the hair-point not spreading when dry, in its areola- 
tion, the median cells of which are a little shorter and the marginal 
cells not differentiated, and in its paler capsule. 


BRYUM BOTTERII C. M. Linnaea 38: 622. 1874 


Valle de México (Bro. Amable): Desierto (1619, 1633, 1637) ; 
Rio Frio (1709); Salazar (1714); Llano Grande, 3,700 meters 


(1736). 
BARTRAMIACEAE (continued) 
BARTRAMIA ITHYPHYLLA (Hall.) Brid. 


Valle de México: Rio Frio (Bro. Amable 1405, 1407 p. p.). 
This species is new to Mexico. 


BARTRAMIA THRAUSTA Schimp.; C. M., Nuov. Giorn. Bot. Ital. 4: 41. 1897 


Valle de México: On rocks (Bro. Amable 1502 p. p., 1503 Pp. p.)- 

An interesting discovery. This species belongs to the South Amer- 
ican flora and was known previously only from Bolivia and Argen- 
tina. Unfortunately I found only two specimens, these among tufts 
of Anacolia intertexta. 


BARTRAMIDULA MEXICANA Schimp. in Besch. Prodr. Bryol. Mex. 58. 1871 
Valle de México: Desierto (Bro. Amable 1624). 


PTY CHOMITRIACEAE 


PTYCHOMITRIUM LEPIDOMITRIUM Schimp. in Besch. Prodr. Bryol. 
Mex. 41. 1871 


Valle de México (Bro. Amable) : Contreras (1443, 1469) ; Salazar 
(1719). 


NO. 4 MEXICAN MOSSES—THERIOT 31 


HEDWIGIACEAE (continued) 
HEDWIGIDIUM IMBERBE (Sm.) Bry. Eur. 


Valle de México: Salazar, on trunks of trees (Bro. Amable 1294 
p.p:). 

A new genus for Mexico, also, I believe, for all of North America. 
This number (1294) was made up of a close intermixture of four 
species, two of them predominant: Hedwigidium imberbe and Nco- 
cardotia subnigra; the two others, Hedwigia albicans var. viridis and 
Braunia secunda, were represented by a few plants only. 

It was an easy matter to separate Neocardotia subnigra and Hed- 
wigia albicans, but quite another thing with regard to Braunia secunda, 
whose presence I did not even suspect. If some fruiting plants had 
not been present this species would have been overlooked, its size and 
appearance being so similar to those of Hedwigidium imberbe. 

It is rather unusual to find, associated in such a manner, two species 
that are indistinguishable either to the naked eye or by the use of a 
hand lens except by their fruit; and it is even more unusual to be 
unable to find morphological and anatomical characters by which to 
separate them. The form and size of the leaves, recurvature of the 
borders, plication of the lamina, and areolation, all are identical. I do 
not know of another example of such an association and such a 
similarity. 


AMBLYSTEGIACEAE 
CAMPYLIUM CHRYSOPHYLLUM (Brid.) Bryhn, Explor. 61. 1893 


Hypnum chrysophyllum Brid. Musc. Rec. 2°: 84. pl. 2. 1801. 


Morelia: Loma Santa Maria (7870). 


CAMPYLIUM HISPIDULUM (Brid.) Mitt. var. SOMMERFELTII (Myrin) 
Lindb. Musc. Scand. 38. 1879 
Puebla: Rancho Guadalupe (4602). Morelia: Cerro Azul (4561) ; 
Loma Santa Maria (5103, 5105, 7859 p. p.); Campanario (7462 
p.p.). Valle de México (Bro. Amable): Desierto (1222 p. p., 1238, 
1432) ; Contreras (1462, 1490). 


AMBLYSTEGIUM SERPENS (L.) Bry. Eur. 


Distr. Federal: Tlalpam (9493). 
This species seems to be new to Mexico. 
3 


32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


AMBLYSTEGIUM VARIUM (Hedw.) Lindb. var. ARSENEI (Par. & Broth.) 
Thér., var. nov. 


Amblystegium Arsenci Par. & Broth., Ms. 


Puebla: Rio San Francisco (5004). Valle de México: Tlalpam 
(Bro. Amable 1453). 

I had previously received this plant from E. G. Paris under the 
name A. Arsenei Par. & Broth., sp. nov., likewise from Rio San 
Francisco. Indeed, at first sight it would appear different from 
A. varium in several respects: (1) Its narrower leaves; (2) its 
greatly developed perichaetium, the perichaetial leaves being almost 
4 times longer than the cauline leaves; (3) the capsule not arcuate 
when dry. 

These characters, however, fade out to some extent upon close 
examination: (1) In A. varium the form of the leaves is very vari- 
able ; (2) if specimens of A. varium are found with an inconspicuous 
perichaetium and short perichaetial leaves, there are others whose 
perichaetium is as well developed as in A. Arsenei; (3) there remains 
only the character afforded by the form of the capsule. This last is 
not sufficient to justify the recognition of a species. 


AMBLYSTEGIUM ORTHOCLADUM (Beauv.) Jaeg. 


Puebla: Finca Guadalupe (737) ; Hacienda Alamos (4723, 4725). 
Morelia: Bosque San Pedro (4569). 

The last plant, probably half submerged, is a form with greatly 
elongated stems and branches and a thicker nerve (60 4). 


AMBLYSTEGIUM RADICALE (Beauv.) Mitt. Musc. Austr. Amer. 569. 1869 


Puebla: ‘Hacienda Batan (934); Rio San Francisco (5000). 

Plants sterile, the determination only probable. The plant from 
Rio San Francisco has the stems and branches laxly foliate and 
the larger leaves widely spreading, either dry or moist. 


AMBLYSTEGIUM JURATZKANUM Schimp. 
Valle de México: Tlalpam (Bro. Amable 1346 p. p.). 


The nerve extends well into the apex of the leaf. This is almost 
the only difference I could find, as compared with the preceding 
plants identified as A. radicale. 


AMBLYSTEGIUM HYGROPHILUM (Jur.) Schimp. 


Puebla: Hacienda Batan (5008). 
A new species for Mexico. 


NO. 4 MEXICAN MOSSES—THERIOT 33 


HYGROHYPNUM PALUSTRE (Huds.) Loesk. 


Puebla: San Felipe (4504, 4505). 
This species was not known previously from Mexico. 


DREPANOCLADUS EXANNULATUS (Giimb.) Warnst. var. MEXICANUS 
(Mitt.) Card. Rev. Bryol. 37: 54. 1910 

Puebla: Hacienda Batan (4961). Querétaro: Cienaga de la 
Canada (11002). 

The var. mexicanus seems close to var. pinnatus (Boul.), from 
which it may be distinguished by its almost entire leaves, with thinner 
costae. The most conspicuous character of this variety consists in the 
very marked apical prominence of the cells. It is, perhaps, the first 
time this peculiarity has been noted in connection with D. exannulatus. 


PLATYHYPNIDIUM SUBRUSCIFORME (C. M.) Fleisch. Laubm. Fl. Jav. 4: 
1537- 1922 

Hypnum subrusciforme C. M. Linnaea 38: 658. 1874. 

Rhynchostegium malacocladum Card. Rev. Bryol. 37: 71. 1910. 

Puebla: (699); banks of Alseseca (700); Cerro Guadalupe 
(676); Hacienda Alamos (4626, 4629, 4761). Valle de México 
(Bro. Amable): Morales (1597) ; Tenango (1687). 

I have noticed the variability of this species with regard to the 
form of the acumen and of the capsule. 


PLATYHYPNIDIUM PRINGLEI (Card.) Broth. in E. & P. Nat. Pflanzenfam. 
ed. 2, II: 347. 1925 


Rhynchostegium Pringlei Card. Rev. Bryol. 37: 70. 1910. 

Puebla: Hacienda Alamos (4628, 4769 p. p.); Hacienda Batan 
(5006). Morelia: Pare San Pedro (4589) ; Andameo (4822) ; Cam- 
panario (7534) ; Loma Santa Maria (4908, 4910). 


PLATYHYPNIDIUM OBTUSIFOLIUM (Besch.) Broth. in E. & P. Nat. 
Pflanzenfam. ed. 2, II: 347. 1925. 

Rhynchostegium obtusifolium Besch. in Card. Rev. Bryol. 37: 71. 1910. 

Morelia: Cerro San Miguel (4870, 5041, 5071); Campanario 
(7631); Loma Santa Maria (4890, 4917). Distr. Federal: Tlal- 
pam (9492). 
PLATYHYPNIDIUM OBTUSIFOLIUM (Besch.) Broth. var. SUBACUTUM 

Thér., var. nov. 


Leaves subacute and contracted at the apex. 
Valle de México: Tlalpam, in water (Bro. Amable 1450, 1452). 


34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


BRACHY THECIACEAE 


PLEUROPUS BONPLANDII (Hook.) Broth. in E. & P. Nat. Planzenfam. 1’: 
1136. 1908 


Leskea Bonplandtu Hook. in Kunth Syn. Pl. Aequin. 1: 61. 1822-28. 


Puebla: Esperanza (4745, 4754). Valle de México (Bro. Ama- 
ble): Desierto (1438); Santa Rosa (1504). Hidalgo: El Chico 
(1580). 

BRACHYTHECIUM 


I must confess that the study of the Mexican specimens belonging 
to the genus Brachythecium has been an extremely laborious task : 
The sterility of many of them on the one hand and, on the other, 
the difficulty if not the impossibility of obtaining, for the sake of com- 
parison, good and complete specimens of the types, are among the 
more important contributory causes. 

I studied nearly 60 numbered specimens and drew almost all of 
them. They belong, excepting five or six, to the sections Acuminata 
and Salebrosa. Now except for B. salebrosum (Hoffm.) and B. lai- 
reticulatum Card., the Mexican species of this group are very diffi- 
cult to identify. For one specimen that agrees with the type there are 
many others which combine characters common to several species and 
which one hesitates to attribute to one rather than the other, Hence 
I gave three different names in succession to the same specimen with- 
out being entirely satisfied with any of them. My conclusions are as 
follows : 

(1) Several of my determinations remain uncertain; they are 
merely probable. 

(2) Some of the Mexican species are very polymorphous, like 
our B. rutabulum, and their forms have been taken for new species. 
Therefore one must not be surprised to find indeterminable specimens 
which in turn seem like new species. 

It will be a task for future bryologists, those who will have the 
privilege of studying the flora im situ, to weigh these variable species 
and to make, with a thorough knowledge of the facts, whatever re- 
ductions are necessary. 


BRACHYTHECIUM TENUINERVE Card. Rev. Bryol. 37: 65. 1g10 


Puebla: Xuehitl, near Esperanza (7988). A form which by its 
laxer areolation marks a tendency toward B. lanceolifolium Card. 

Valle de México (Bro. Amable): Contreras (1221, 1483); De- 
sierto (1618) ; Salazar (1717); Llano Grande (1738). This is a 
form with very elongate branches. 


NO. 4 MEXICAN MOSSES—THERIOT 35 


BRACHYTHECIUM ALBULUM Besch. in Card. Rev. Bryol. 37: 66. 1910 

Morelia: Bosque San Pedro (4582). 

I have seen only a very incomplete specimen of the type. The 
present specimen seems to differ from it by the longer and more 
slender acumen of the leaves. 


BRACHYTHECIUM LANCEOLIFOLIUM Card. Rev. Bryol. 37: 66. 1910 

Puebla: Cerro Guadalupe (796) ; Hacienda Alamos (4760) ; Rio 
San Francisco (5003). Morelia: Loma Santa Maria (5089). Distr. 
Federal: Mixcoac (9453). Valle de México (Bro. Amable): Santa 
Teresa (1339) ; Contadero (1364) ; Tizapan (1612). 


BRACHYTHECIUM LANCEOLIFOLIUM Card. var. GRACILE Card. Rev. 
Bryol. 37: 66. 1910 
Puebla: Hacienda Batan (935) ; Hacienda Alamos (4799). More- 
lia: Campanario (7452). 


BRACHYTHECIUM CLADONEURON (C. M.) Par. Ind. Bryol. 132. 1894 
Hypnum cladoneuron C. M. Linnaea 38: 652. 1874. 
Puebla: Hacienda Alamos (4696). 


BRACHYTHECIUM COMTIFOLIUM (C. M.) Jaeg. 
Hypnum comtifolium C. M. Linnaea 38: 653. 1874. 
Valle de México: Desierto (Bro. Amable 1238). 


BRACHYTHECIUM TROCHALOBASIS C. M. Bull. Herb. Boiss. 5: 238. 1897 


Puebla: Esperanza (4729). Morelia: Cascade de Coincho (4713) ; 
Carindapaz (7951) ; Santa Clara (4886). 


BRACHYTHECIUM FLEXIVENTROSUM (C. M.) Jaeg. 

Hypnum flexiventrosum C. M. Linnaea 38: 653. 1874. 

Morelia: Cerro San Miguel (7546) ; Campanario (7940) ; Cerro 
Azul (4532, 4541, 4554, 4788). Distr. Federal: Tlalpam (9498). 
Valle de México (Bro. Amable) : Desierto (1222 p. p.) ; San Juanico 
(1261).. 

Several of these specimens oscillate between this species and the 
preceding one. In their long and slender acumen and flexuose costa 
they tend toward B. flexiventrosum; but the habit, the short nerve, and 
the short pedicel (1 cm. or less) bring them nearer to B. trochalo- 
basis. I am not far from believing that these two species should 
be united into one. 


36 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


BRACHYTHECIUM SERICEOLUM Card. Rev. Bryol. 37: 66. 1910 


Puebla: (4997) ; Hacienda Batan (4975). 


BRACHYTHECIUM FLEXINERVE Card. Rev. Bryol. 37: 67. 1910 


Puebla: (4862); Hacienda Santa Barbara (740). Tlaxcala: 
(4855). 


BRACHYTHECIUM ALBO-FLAVENS Card. Rev. Bryol. 37: 68. 1910 


Puebla: Rancho Guadalupe (4614) ; Esperanza (4666). Morelia: 
Campanario (7537, 7506); Cerro Azul (4530). 


BRACHYTHECIUM ALBO-VIRIDE Besch. in Card. Rev. Bryol. 37: 69. 1910 


Puebla: Boca del Monte (4674). Morelia: Campanario (7539). 

I recognize in these specimens most of the characters attributed 
to B. albo-viride: the green color of the tufts, the laxly foliate 
branches, the lanceolate leaves strongly excavate at the base, long- 
acuminate, with a costa reaching to two-thirds and even three-fourths 
of the leaf ; but I have not noticed that the stems are more slender and 
the branches more tenuous than in B. albo-favens. 


BRACHYTHECIUM LAXIRETICULATUM Card. Rev. Bryol. 37: 67. 1910 


Valle de México: Desierto (Bro. Amable 1412). 


BRACHYTHECIUM ACUTUM (Mitt.) Sull. Icon. Musc. Suppl. 99. p/. 75. 1874 


Pueblo: Rio San Francisco (5001 p. p.) ; sterile plant. 


BRACHYTHECIUM SALEBROSUM (Hofim.) Bry. Eur. 


Puebla: Esperanza (4515, 46064, 4690). 


BRACHYTHECIUM SALEBROSUM var. POLYOICUM Thér., var. nov. 


Synoicous and unisexual flowers, male and female, on the same 
stem. 

Puebla: Hacienda Batan (4937). 

I combine this curious form with B. salebrosum on account of its 
characters as a whole. It is much nearer to it than to the known 
synoicous or polyoicous species B. acutum, B. conostomum, and 


B. Mildeanum, 


NO. 4 MEXICAN MOSSES—THERIOT 37 


BRACHYTHECIUM INTEGRIFOLIUM Thér., sp. nov. 
Distr. Federal: Tlalpam (9499). 


Sterile. Caulis repens, radiculosus, sat regulariter pinnatus, ramis 
inaequalibus, 3-4 mm., usque 10 mm. longis, patulis, attenuatis. Folia 
caulina sicca et humida erecto-appressa, e basi decurrente latissime 
cordato-ovalia in acumen longiusculum, patulum, acutum sat subito 
constricta, haud plicata, marginibus planis, integerrimis, 1.6-1.7 mm. 


Tey iC - 


, ? 





Fic. 19.—Brachythecium integrifolium Ther. 1, 2, 3, stem leaves; 4, median 
cells; 5, basal and alar cells; 6, branch leaves; 7, acumen. 


longa, 0.8-0.9 mm. lata, costa ad 3 evanida; rete pellucido, chloro- 
phylloso, cellulis linearibus, par ean tenuibus, 35-45 «Xx 5-6», cellu- 
lis basilaribus et alaribus laxioribus, breviter rectangularibus. Folia 
ramea minora, secunda, subfalcata, 1 mm.xo.5 mm. Caetera ignota. 

In size and habit like B. hylocomioides Card., but that species has 
denticulate non-falciform leaves and looser areolation. It suggests 
also B. refexum Starke, but is easily distinguished by its entire leaves, 
with the nerve reaching only to the base of the acumen. 


BRACHYTHECIUM CORBIEREI Card. Rev. Bryol. 38: 42. 1911 


Valle de México (Bro, Amable) : Desierto (1222 p. p., 1441) ; Rio 
Frio (1692, 1703, 1706) ; Contadero (1307). 
The last number (1307) represents a form with long, flexuose 


stems, irregularly ramose, with long, slender, almost flagelliform 
branches. 


BRACHYTHECIUM PLUMOSUM (Sw.) Bry. Eur. 
Puebla: Huejotzingo (4856). Morelia: Loma Santa Maria 
(4896). Valle de México (Bro. Amable): Desierto (1245, 1616, 
1640) ; Contreras (1668). Hidalgo: El Chico (1586). 


38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


Of the material listed no. 1668 may be classified as var. sublaevipes 
Card.,’ because the pedicel is scarcely papillose at the top; under a 
strong magnification one can see only separated, depressed, low pa- 
pillae. In this specimen a single costa is the exception; most of the 
leaves have a double nerve of very variable length, sometimes very 
short. The Bryologia Europaea indicates that this case is not of rare 
occurrence. 


BRACHYTHECIUM HASTIFOLIUM Card. Rev. Bryol. 37: 69. 1910 


This species is not mentioned by Brotherus in his treatment of 
Brachythecium in the second edition of Die naturlichen Pflanzen- 
familien, but I am inclined to think that it is the one cited in the genus 
Heterophyllum under the combination H. hastifoliwm (Card.) Fleisch. 

Cardot says, “ Costa ad % evanida.’’ How can this character agree 
with the genus Heterophyllum, which has “ Rippe sehr kurz oder 
fehlend?”’ How could a moss which a bryologist of the standing of 
Cardot affirms to belong to the genus Brachythecium have at the same 
time the characters of the family Brachytheciaceae and those common 
to the genus Heterophyllum of the family Sematophyllaceae ? 

I have endeavored to solve this puzzle.. An examination of no. 
10474 of Pringle’s exsiccata brought the answer to me. The speci- 
men in my collection labelled Brachythecium hastifoliwm Card. 1s 
not this species, but Heterophyllum affine (Hook.) Fleisch. Now if 
one turns to the original description, where Cardot discusses Prin- 
gle’s no. 10474, which he considers as a form of his Brachythecium 
hastifolium, the inference is clear that Pringle distributed under 
this same number (10474) two different species—the form just men- 
tioned and Heterophyllum affine. 1 take no pride in this discovery, 
but I cannot understand why such an expert and conscientious bryolo- 
gist as Fleischer failed to find the clue and thus allowed himself to be 
misled into giving full confidence to a specimen which did not agree 
with the original description and was distributed by a collector who 
was not a bryologist. 

My conclusions are: First, that Heterophyllum hastifolium 
(Card.) Fleisch. is a myth, and that this combination must disappear 
from nomenclature ; secondly, that the binomial, Brachythecium hasti- 
folium Card., which applies to one of the best characterized species 
of the subgenus Salebrosium, ought to take again its place. 





*Rev. Bryol. 37: 70. I9gT10. 


NO. 4 MEXICAN MOSSES—THERIOT 39 


RHYNCHOSTEGIUM SAINT-PIERREI Thér., sp. nov. 


Valle de México: Contadero, on bark; leg. Marius Saint-Pierre 
(Bro. Amable 1298). 


6 


x 200 





| 
\4 


! 
h 
Fie, 20.—Rhynchostegium Saint-Pierrei Thér. 1, stem leaves; 2, apical cells; 


3, upper cells; 4, median cells; 5, branch leaf; 6, marginal cells; 7, 8, perichaetial 
leaves; 9, moist capsule. 


j,? 
| | 


* 200 


es /| / Va ; 
PL) 





Rh. leptomerocarpo (C. M.) sat simile, sed differt colore smaragdo- 
viridi, caulibus gracilibus, laxe foliosis, foliis siccis valde patulis, duplo 
angustioribus (1.7 mm.X0.5 mm.), tenuiter acuminatis, paulum de- 
currentibus, rete densiore (cellulis mediis go-120 » x6»), foliis peri- 
chaetialibus duplo majoribus, pedicellis longioribus (2 cm. longis). 


RHYNCHOSTEGIUM HUITOMALCONUM (C. M.) Besch. Prodr. Bryol. Mex. 
107. 1871 


Hypnum huitomalconum C. M. Syn. 2: 248. 1850. 


Morelia: Cascade de Coincho (4712a) ; Andameo (4827). Valle de 
Mexico: Tlalpam (Bro. Amable). 


RHYNCHOSTEGIUM LEPTOMEROCARPUM (C. M.) Besch. Prodr. Bryol. 
Mex. 107. 1871 


Hypnum leptomerocarpum C. M. Syn. 2: 354. 1850. 


Puebla: Hacienda Alamos (586). Morelia: Loma Santa Maria 
(4868, 4894, 5062). Distr. Federal: Tlalpam (9430a) ; Cuajimalpa 
(9487, 9489). Valle de México (Bro. Amable): Santa Rosa (1515) ; 
Contadero (1315). 


40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


RHYNCHOSTEGIELLA ARSENEI Thér., sp. nov. 


Puebla: Hacienda Santa Barbara, on sandy ground (739). 

Sterile. Caespites lutescenti-virides, nitidi, Caulis repens, gracilis, 
laxe foliosus, ramosus, parce radiculosus, paraphyllis raris; ramis 
erectis, brevibus, 2-3 mm. longis, sat confertis, sat dense foliosis. 
Folia caulina et ramea sicca erecto-patula, humida patentia, anguste 
lanceolata-acuminata, acuta, decurrentia, marginibus planis, toto am- 


3 2 i 
x 200 
x 200 


x 200 





Fic. 21—Rhynchostegiella Arsenei Thér. 1, stem leaves; 2, apical cells; 
3, median cells; 4, marginal cells; 5, basal cells. 


bitu minute denticulatis, 0.6-0.7 mm. x 0.2 mm. ; costa basi 30 m percur- 
rente; rete opaco, cellulis linearibus, 36-4046 p, basilaribus sat 
distinctis, marginalibus subquadratis (2-3 ser.), subhyalinis. 

This species can be compared only with FR. Jacquini (Garov.) 
Limpr. and Rk. Teesdalu (Sm.) Limpr. It is distinct from both by its 
leaves denticulate all around and from the first species also by its 
larger branch leaves, differentiated basal areolation, and more densely 
foliate branches ; from the second species by its acute, decurrent leaves. 


EURHYNCHIUM SUBSTRIATUM Thér., sp. nov. 


Valle de México: Llano Grande, alt. 3,700 meters, on rocks (Bro. 
Amable 1735). 

I, striato (Schreb.) simillimum differt: statura graciliore, ramis 
brevioribus, foliis minus profunde sulcatis, caulinis angustius de- 
currentibus, rete basilari praecipue ad angulos densiore, foliis rameis 
margine dentibus brevioribus, 





NO. 4 MEXICAN MOSSES—THERIOT 4I 


I segregate this plant from £. striatum, because, in addition to the 
characters enumerated above, the European species 1s absolutely un- 





Fic. 22.—Eurhynchium substriatum Thér. 1, stem leaf; 2, median and mar- 
ginal cells; 3, basal cells; 4, 5, branch leaves; 6, cross-section of branch leaf; 
7, apical cells; 8, median cells; 9, perichaetial leaf. 
known throughout the American continent and it is therefore hardly 
possible to consider the moss from Llano Grande as a local form. 


EURHYNCHIUM STOKESII (Turn.) Bry. Eur. 


Puebla: Boca del Monte (4738) ; a form with stems less densely 
branched, elongate, and laxly foliate. Valle de México: Contreras 
(Bro. Amable 1518). 


ENTODONTACEAE (continued) 


PTERIGYNANDRUM FILIFORME (Timm.) Hedw. var. MEXICANUM Ther., 
var. nov. 


Folia valde secunda, latiora (0.5 mm.), cellulis apice parum pro- 
minulis, costa gemella usque ad 4 folii producta. 

Valle de México: Santa Rosa (Bro. Amable 1503). Hidalgo: 
Mineral del Chico (Orcutt 6649). 

The typical form of this species has not yet been found in Mexico. 


ROZEA STRICTA Besch. Prodr. Bryol. Mex. too. 1871 


Valle de México (Bro. Amable): Desierto (1418, 1425, 1444) ; 
Llano Grande (1731). 


ENTODON JAMESONII (Tayl.) Mitt. Musc. Austr. Amer. 525. 1869 


Morelia: Cerro Azul (4779). Valle de México (Bro. Amable) : 
Desierto (1245) ; Contadero (1311). 


42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. &5 


ENTODON ABBREVIATUS (Bry. Eur.) Jaeg. 


Valle de México (Bro. Amable): Contadero (1302, 1305) ; San 
Rafael (1280). 

The pedicel is very variable in length. In the same tuft I have seen 
pedicels 3 mm. long and others up to g mm. long. 


ENTODON ABBREVIATUS (Bry. Eur.) Jaeg. var. TURGESCENS Thér., 
var. nov. 


Caules et rami turgidi, folia 2.2 mm.xX1.3 mm., valde concava, 
cochleariformia, rete laxiore, cellulis mediis 70-go px 9 p. 

Valle de México: Contadero (Bro. Amable 1362). 

The facies of this variety is very different from the usual forms 
of E. abbreviatus. In its leaves and their areolation it comes close to 
Pringle’s no. 15226, identified by Cardot as E. brevipes (Schimp.). 


SEMATOPHYLEACE AE 


RHAPHIDORRHYNCHIUM OBLIQUEROSTRATUM (Mitt.) Broth. in E. & P. 
Nat. Pflanzenfam. ed. 2, 11: 428. 1925 


Sematophyllum obliquerostratum Mitt. Musc. Austr. Amer. 472. 1860. 
Morelia: Campanario (7926, 7941). 
RHAPHIDORRHYNCHIUM DECUMBENS (Wils.) Broth. in E. & P. Nat. 
Pflanzenfam. ed. 2, 11: 427. 1925 


Hypnum decumbens Wils. (Ms.); Sematophyllum decumbens Mitt. Muse. 
Austr. Amer. 488. 1860. 


Valle de México: Desierto (Bro. Amable 1439). 


SEMATOPHYLLUM CAESPITOSUM (Sw.) Mitt. Musc. Austr. Amer. 479. 
1869 


Morelia: (7893, 7909, 7912); Cerro Azul (4784) ; Campanario 
(7515, 7516, 7520, 7537, 7559 7552; 7557, 7935)- 
SEMATOPHYLLUM CAESPITOSUM (Sw.) Mitt. var. LATICUSPIDATUM 

(Card.) Thér., comb. nov. 

Rhaphidostegium caespitosum var. laticuspidatum Card. Rev. Bryol. 40: 39. 
1913. 

Morelia: (7890, 7916) ; Campanario (7642). 

SEMATOPHYLLUM HAMPEI (Besch.) Broth. in E. & P. Nat. Pflanzenfam. 
ed. 2, IL: 433. 1925 
Rhynchostegium Hampei Besch. Prodr. Bryol, Mex. 105. 1871. 


Morelia: Campanario (7518). 





NO. 4 MEXICAN MOSSES—THERIOT 43 


HY PNACEAE (continued) 


STEREODON FALCATUS (Schimp.) Fleisch. in E. & P. Nat. Pflanzenfam. 
ed. 2, I: 452. 1925 

Stereodon subfalcatus (Schimp.) Fleisch. in E. & P. Nat. Pflanzenfam. ed. 2, 
IIs 452. 1925. 

Further observations have convinced me that in these two species 
of Schimper’s there is only a single specific type.’ 

New localities: Valle de México (Bro. Amable): Desierto (1210, 
1221) ; Acopilco (1201) ; Salazar (1236). 
HYPNUM AMABILE (Mitt.) Broth. in E. & P. Nat. Pflanzenfam. ed. 2, 11: 

454. 1925 
Ectropothecium amabile Mitt. Musc. Austr. Amer. 513. 1860. 


Puebla: (4945, 4947, 4948, 4949, 4951, 4953, 4956, 4959). Distr. 
Federal: San Angel (9479). 


ISOPTERYGIUM CYLINDRICARPUM Card. Rev. Bryol. 37: 56. 1910 
Valle de México: Desierto (Bro. Amable 1247, 1623). 
TAXIPHYLLUM PLANISSIMUM (Mitt.) Broth. in E. & P. Nat. Pflanzenfam. 
ed. 2, 11: 462. 1925 
Tsopterygium planissimum Mitt. Musc. Austr. Amer. 408. 1860. 
Puebla: Hacienda Alamos (584). Distr. Federal: Tlalpam 
(10999). 


ISOPTERYGIUM PLANISSIMUM Mitt. var. LAXIRETE Thér., var. nov. 
A forma typica differt: rete laxiore, cellulis diam. 8-9 p. 


Morelia: Loma Santa Maria (4877). 


VESICULARIA VESICULARIS (Schwaegr.) Broth. in E. & P. Nat. 
Pflanzenfam, 1°: 1094. 1908 


Hypnum vesiculare Schwaegr. Suppl. 2°: pl. 199. 1827. 
Nuevo Leon: Monterrey (Bro. Abbon 10969). 


MICROTHAMNIUM THELISTEGUM (C. M.) Mitt. Musc. Austr. Amer. 504. 
1869 


Hypnum thelistegum C. M. Syn. 2: 269. 1850. 

Morelia: Campanario (7924). 

Sterile, the determination uncertain. The cauline leaves are sharply 
dentate and the branch leaves secund. 


*See, Smithsonian Misc. Coll. 787: 28. 1926. 


AA SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 
MICROTHAMNIUM SUBTHELISTEGUM (Card.) Broth. in E. & P. Nat. 
Pflanzenfam. ed. 2, IL: 471. 1925 

Mittenothamnium subthelistegum Card. Rev. Bryol. 37:55. I9Io. 


Morelia: Jesus del Monte (7608a). 


HYLOCOMIACEAE 
LEPTOHYMENIUM EHRENBERGIANUM (C. M.) Fleisch. in sched. 


Hypnum Ehrenbergianum C. M. Bot. Zeit. 14: 408. 1856. 
Hylocomium Ehrenbergianum Besch. Prodr. Bryol. Mex. 111. 1871. 


Vera Cruz: Jalapa (7998). 


POLYTRICHACEAE, (continued) 
POGONATUM BESCHERELLEI Hampe in Besch. Prodr. Bryol. Mex. 63. 1871 


Valle de México: Salazar, alt. 3,100 meters (Bro. Amable 1715). 


POLYTRICHUM ALPINIFORME Card. Rev. Bryol. 37: 6. 1910 


Valle de México (Bro. Amable): Contreras (1667) ; Xoquiapan 
(1750). 

The last plant, which is in fruit, affords an opportunity to complete 
the description: 

Folia perichaetialia numerosa (12-15), remota, longe et late vagi- 
nantia (vagina 4-5 mm. longa, 0.2 mm. lata), in acumen angustum 
abrupte contracta, humida patulo-squarrosa. Pedicellus 20 mm. altus ; 
capsula minuta, oblonga, laevis ; calyptra angusta, elongata, 6-7 mm., 
parce pilosa. Caetera ignota (capsulae immaturae). 








SMITHSONIAN MISCELLANEOUS COLLECTIONS 
VOLUME 85, NUMBER 4 (ADDENDUM) 


INDEX TO PAPERS BY I. THERIOT ON MEXICAN MOSSES COL- 
LECTED BY BROTHER ARSENE BROUARD PUBLISHED 
BY THE SMITHSONIAN INSTITUTION’ 


INDEX 


[Synonyms in italic. Page numbers of principal entries in heavy-faced type. ] 


PNOMiaRGA CEO IT TO] atten. setae 8 staves ete Pes ettad. crcuoveteia ties dy crenata alep bie aly rage Siders III, 22 
EMOTE Cll awe CALCU awe srottna Setey natetee Sica ahe oettrotssay2i-dcnye slate lari des, dvs ios exaudio iterates III, 22 
EMD SteP Lat APSCHEL cid cel os eles ala dn es ein arn Go eth eda Boog Dele ders i iieee2 
Frny sare Op Eye LUiTtameece says coe ha catvinis lol atbitouay shave, airi-d delat datleneusl ch actieie onavanees duct oe ee 
Siuratzikan tists. sje rete eee A fete eal mies cereals deere eae el 2 

COTgt nO Cle Cla trey ance aes vam aoa Bete o ees Sean hi Sh dias oh Stare Sh Sr toude coud Avoute ce, SM ottowarrd 6. LIT 32 
HAG T Gell CMMs RVs 2 ae bis orate BL SEN RA NSB LA asl at sae afove cusriipleiasine dea tes Iii s2 
Gla) GILG) aeteberene once oraheret oi seeney crore eterna cise Ga aaersie ice cree entice ere oie eee Lilien 
SATU TTUN cc Geese hte treats ay af Shoes aya Mearns nseste APM G asnsnd sl Mau D aed adialaioea: Tht, 32 

PN TES ETE Lessa ries cathe SVS AS re Ica oP Ra ns en eesine sone a BaP Ditne2 

PMC OT AMITIRE ECGS alm ceeraics a eee a emis is csne Sieressiieseuese aie sucieloreus sioveen overstate eise oo aera I, 18 
UTS CUO l Tel ot emer crerensaareueheee nen caaer erences ace teen oie a rene ore eure 1, 18 

SELL lei reese acy enemas lesa rete teens ec ieiet reich at canter Oe ahs ets 
SMDSESSil I Sme@merer araecre oat ieee ee er earn I, 19 
BMIOECtalcilmy APICWAtUIM v.44p6s0 4 spanwise dine esoend che sd bdevar baeee ha Deas 
HeCHINUp) A CULIT II sepemeraye ex aievensyevenstene lem gererevetciaie eaxceierstere reve ate ie cies re Sieiava chees eA 
BONG elMSa tlimtin pantera ies sete eiyeee tre eusee wieresie oraniesic ese eae eidigen ale Ill, 5 
SULCIN ORO acters teers ks sis iene iees ice inion Sernt neers eee es 
mesa ATE AMAT Ie? "eres seas ay sco wiane eaicinta ae 4'a int eave Sc clase une cae are a seetesst aie tet Ill, 4 
EMIGINODE YM ALiTOrmMe MEXICAN. .<.s 0.40 esgee vesee ven ceusveeeeseaes II, 1x 
DISA hitin Meee eae eiiere reste Ticet erie een eee oe eee eee Le ex 
VAMOMVOGOW TOCCOGE 5. 6.606scccecescarsncwscecoesacccancececceateae iW, 2 
PES EEOC IA (DVECIPES 6 cuando G4 a4 velwn 658 Saareidae siuee a vas eck eaccead ac 3 
Piet UCLA Came Senet ere Ine eaten rere neTeTE ere Te eee rere Setar ole oteretea sien seis ee 
PRPIEUNCCTISIS: Sen enrebice ence tee See sie Sone Ao sve Sees ent ches Wanda nd Ds 

Hilt cA CMe arty acea aS erEa Toya icteetsa tee terete a eee Ie ey ice eiecele he tue Sle Oalbaans's a I, 4 

TST MA CL See sveneniercececircesee aes HTT eee oh ee aca altace nn i 
SUMS 1 cals ewe sy aeryoe cece eee vercw ys reestal susyerau wer state vane Tete den Goae ear nci six eter cne ore aanns | oe 
ESL ETE eS TS SOT Te 2 
Atrichum contermanum ....00. ccc cece cece ccc eens cnecceeseevecevcccuves [ea2o 
Mrrelleri *contermintm +..4-s00-.0e6.0. 4 ase ce lt cent ecole c I, 20; II, 26 

TON GUY el UAT pee eee eae eee reo y Seay areyen UIE aT, Th ire MOREE Aedes sem vas 1, I, 20 


* Part I, Smithsonian Misc. Coll., vol. 78, no. 2, June 15, 1926. 
Part II, idem, vol. 81, no. 1, August 15, 1928. 
Part III, idem, vol. 85, no. 4, August 25, 1931. 


45 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


Barbula Abbomity cekcccnea dees oe ere oe ees SHS Hee TREE RRS III, 20 
AIEISStaY etic eis cvs ei Gla reo ees oe ICE aren eR or snepe ERE eienetonoeyetnenots ii rzeere 

GIN PUD LAC CLL ” si esha cet ees Roe ecto hae Sao eS et RR Nach ntoeS coheed eo teceg= Ill, 23 
Bescherelletie sian sacicearscryett tree torpeletay toecuntenteleterer ene ee oer Tench meene rate Tag, 
CLASGCIMERV IQA oh SO rene ee ae ERR ee einer ere mien Ee taker Mia 

SEEN OCAL PA % dra A syd sie ate a eee repairer ee ete eaten tere Lae 
IByoyoirereaehich nogueongsoco pec Fi Naa PI Nis ry aN ch leg Say a OR III, 18 

Cal Careat ecto 8 Sie cect ee ie AES Seo Bars ies rated eM Cen nee Iieezo 
dialivérichoides s 4.2 o/s oe wre oictaieicverse terete eheicte-s etetehanchete RoteverrNeveie raha tate fetes fete III, 20 
FEV ORD CH GONG Cie ules i wtarter a Sister tee aye ae eRoeneh A avete Mel) ieee toa eatche ez 
Hhrenberoii mexicana meorcaerie seein irr reer III, 19, 20 
POFVUGINED. waned na She ice hata nie hom ee ae elas eto De Seer III, 9 
PEACUESCEMS. 5. in. aero Syapkl es a ese Ste ers o elas hac ree OS arson austohal pape Lil. 27s 
PHACHIOLMIS: eee ciae oes siekaals arcisieshe aes chiara ora marae rereraie erie tiie Li r7ers 
hypselostepia. esyris seawnrste aracee Sonia oa amin siete se anemia he aera eee III, 21 
NATE TE CHALE: Ue ave ce tev mesos cotskg oua Soe aiie oe toce SRS Sea Sad Say oSne EN nen sone eters Eisen 
OVIZADEMSTS:  eacrex ees cisy so Soe cue iereter oh eelallele taverethlsh Pepateheto AMONG Caevoroen chee note treks Ill, 18 
PUbTiCA ys, =z serstersafoide koe Shee Gua ae earch oleae te Lh nos Zo 
SPIT ALES <a caiee cea wicca setae) te wicteadev ona loreal She area terete fole sfeomicpatater te aa nero III, 18, 19 
SEETLOCH EGA ikon odie, Sua cra eee cls cnc aiwtal ai cn Sotelo ouanigee Sic re Mauey ores ehorenenterete Li om 
Stubteretiuscula) sc.ccts 6 <a5 os cris a sroisin o-o,50e aucun uanvis es elsievn lorie euerepehtele III, 18 

TO Caahbk cele len mena ao Samad neo mop ccaccuscmacnamdos odoudlD ono tc Ill, 18 
ERICH O STOIMOLD ES ia ace, Hevseiere hia are ele Saree aie el Oe vole Oba cversy el srepeenstert Likes 
Wasi ttl atten: say Metin ce wi atey hatese aie aeons oats oustay tetrsalct woke tcust ss Sistine tae IL a8 
Bantramua thy plivila, eeeecaais oe emiayae ac rst xstoieerela cinecne ie Cera et eerste iene III, 30 
En aan tL S Eats Ae here eee Te IN sr tie Rot aeceeteys ie Ore coke cen accuse tees chai fame Miteso 
Bartramidulaimexicatias s.ceis/s osu siete eveuetteors seta erasers & alee siokeie rete) aerarsiotaeenenens III, 30 
Brachymentunme barbacaimOmtise 2 7.) scien crcieneie syste eerie fae ako eee eter redone Dis 
Capilla res nc elaine seeranai shane here snes aie tusr teva orate tay tsrover cite le Siatadl her sfegeenste a aaa II, 9 
Chlorocanpitiniwe: mercies cee eset eborere ster usta tortor stoner keekersracnerater II, 10 
COndensatutn sae ie Ce erste Oo eel cielo eit rer okey tel Revs atreionens II, 10 
EMIGUUT cca as cone cle cae Se whe aise ctoyeilafe efeharon eighore oo eteyaoa ap Severe teats cher «tee keine II, 8 

it 1CAtUtM Vora cys ses cies ei cis ciate ellevere esc letancnere siiaeye ei x steeere Totcoter Miah Melsnetelts Herel II, 9 
TE ZAM ONT heel ee ees ieee oe a eas OOOO eros mete eae ll xo 
AEC OLE Fars Penske Siete abe are Os, Oe ROIS RTE ECT CUAL Crate MeN We 
TERI CATIUTI wea icts ctoetereceieere econ tees renewals listeners) sionerensyeneterelorekoteliheKcntonetare II, ro 
INAitreric bitte eae ye ves rere rerericiess ace icioveie rene rcret ete telereree Velev eterno tener Melt oherehs tate II, 9 
TEQUET LSS weoretcs csc eta ahaiieTes Sat eNes ay cod aT aOT ST Certo PMC H Ns cl oy sroueteowolotoitevo mete stsonerch seks II, 8 
PLLVIGUITINT soleete OTR esr oe ene veme ere H A ene ic ieee ooh nereleers ter tenn ekeroRs II, ro 
Saints PLERT OL sera ceercvsie Siero ero cree A ore a we ve re ferew clemency Rolcey eichaeoners Ill, 29 
SQUATVUVOSUIN. Gcraratetciaveraidta ws 01h tie sieie) siels a'0 area lelavara/=imennve l= Waele sitet ere Lie 

SYS tylitaini: shce ciara inet eons = ctas narrctors ees eiaiensile ete erate II, 10 
Brachythecitum sActtuim' casa sos oo te ee cce esie eens cin inyeg eee eters Til, 36 
BIBO= Aa eNSS Asc sc ae ae haste INOS OHS ww Blcveke ene cct ake Greteko MRCS emreene MI 36 
ABO | VL a Rc ascaeca nese toler ee ca ol ele acre aes tone Renae ee poeeeemenontere III, 36 
SUBS Ga rea 2.8, eae each oor tesco bate oro mene cas Pane SE ce eR ete RcTened emetoree III, 35 
CLACOM EE OL rise een eae eel oe ae Coste elas eae cI tar ton ener Ill, 35 
eres at Colt Lr Rae ee ee Morir ayteye cM Tale iss 
GOTO SEGAL NN ee re ster cscs css ee edie ete eae ai ca NT Len tate oot one Iis6 
COTBISL EL: ccs d eelece iG ecw ev ree eto eore lis Fe raed Pe ess ae aU redone oc Eos erst aes TIT 937 


HEINER VE alts aise Dia teleoc rh oe eet TST Ue UT a SEE ere eerie hereon III, 36 





NO. 4 MEXICAN MOSSES—THERIOT A7 
Brachythecium—Continued. 

HE MAVEMEUOSUIII Veeteve skate er, eee eit Crete cd Eile cvsy ot Aud Seuss Sane Meal aiacete. Seveienae tii 35 

avs Gr T UTI Sees os Scent stent ater tse shayie Oat ksysiDaNe WOR ate «cies. III, 38 

IyglG COMO LES dere, «5 fe se Be oa acoleia.c MEI SiS So aegietoy cs aie 37, 

ATCC Casi LT CITT Weageu ee dyah Pat Sea sears Sait =. oa SA tpayednne els ee eeheews stl III, 37 

HAM CEO LATO LIUITIN fis sce va. asroeseaklo ears aves ay laye 6 vnsieye O aya ers avdrmvagens ts yet Pisa ss 

ated Gl Cig rer eave tey tio eka ve ee ome awe goatee he ak os hes cat yayzropenuseans sWevatene tev Ill, 35 

Paes e tal CLUE CA EELEYN, 07 6: sy sear tevctiateuetete teeuscs Stat ye ralsl 6,6 Soren aie. a, cveNape, exact utbanecorvensare 66 III, 36 

ANT Cea ttt rn aetna oo eon ¥ke Mey Sibs Schr os a Absyen saSs vale Gecitsou Sus teeeyee Spo ysebyES 8 aus III, 36 

PUTA S CHT AMineyaysy nese ods ekavayev or Cayeyaranaietaparexe cieiers we scree eager whey aa ce eeeeye eter Les 7, 

TET CORUII YM Mera consis cite oye abst av S00 oe os SV Syaos oelints 2) ote oe Be thysceis ) outyleee yeas Inn 37 

Gall GPO Street tees es once ether ocr oee eres eretectere crereimesea cuentas: te Lie s6 

TOO OL CUM area stun ceeveie ys ore reser ons ere eeserr eo eaele ete ercre eee ee ee lees 

SETHI E OMT Avec ucita cre eecnau sun oe tap ve erteciete nia cies ei arauerseeravansrentiase erensuntelerc ran III, 36 

PEIN UIULD © eV Coae wavereumasyfeusyeeey sxe apsye stores tetereietaney ery eerie ieicreeccinienriooeyes oheroreneictre I, 34 

OG Hell OM asts ae seree peer enas ene eeray. eens aereeaiererrm everson III, 35 

FEMA ZA MUP TEU 25 2 Bleisie cuscnisueia Se een e wo 6lm 6 Gyayts Suess Flom aed aloes Ge oe lvsd 

Ce Tot | Tc we eeeyege eee een etane omens use Uae cereceretayeteticna e¥euole a cieieye cheretameices tei: Lez 

lehiiantiiania ese eee eee cite. Sates ieee las 

PUL Gevtay emaersrcmra ss ncpe mus eentriseisrers eee cise stele met eee naar eens den may ence ae Le2e3 

GANIESCEM Sumer tae Ee oe erro eee l323 

SE COUITI Cl cll ya cence emetic ee V Sec eer aaa ere iise ee tees fo Sotto nc ens teers e235) Dil ar 

ENTICLGICLIXI Mee een acer ene eh tee nenereer ene raeye cities ees et ans I, 24 

aSSINGL Siamese meister teeter eee ones ete ren Tiane Sie Rare cwi te salen enavoee 3 24 

DEG Ga ee eee: Sicreee te rg ce crete Tee ON enc ohereecce ez 

GLEN FIAGI (OS a eat ao ne Ea oe eee ie oto CL ee RU eC OE eee Le22 

HSRC UILE Ice raTItGITMC Ici ercics iets scereieasie a tenoceneeo eens o.srei.s.s 61 are eecerctouc the sestha eo t.a? I-20 

EC) TTT Call CCS ct ee a ee ON Petey ce ME eT, Non ain nye cert erny Ns ts cata) I, 20 

Hes qy, OI 1) LI VUATIT THE CATIVITI eye eiene areve eterna ete ates slate core. 0s fay oi adavte neste e cue. .e- cyst Tez 

FeTTo ATI DIWZOLE DIS: wereateasers.cevatst sects aise eneses cSt wo cua Shaye shu ay bvela toe ash nee Avesis. I, 16 

UTC (|< eee a RENE STEM cia Al a oan. Br caste Ar Dens ya 

AIGS GLUE TUTTI mmeeve grote naan aate terete her cieta ema ES viata lola days ay Stapnsnata eeen erst aie eet 7ien i eaeer 

DEachy can pun Mea are teeta tees eae Ae ae rte Nace eR eae II, 11 

EMMOPOCAU MIT amare ee kn eee lee «ced  a tudes heen ar whales rt ae 

COSLAICEIS Cua erent oe ee cerca cutee hoes oa eran fete Peete =e eer 

LL CCITT penne a OIE OS 3, nes gee oes eM ane mc goaeg er re RNS fe cies tae Mey, 

EEL cUY (1S gaa stn gntl eee Pha wed catoR ee a dhe ee, PINNARAL RAI ola @ NEE wih Sovh i 

PAGS C11 GH wmeese cme te arch sapee eae T SOI cee ie Sete io ak arse Seat cathe etaeier eset et I, 16 

IS OLLG Lol imenemenscerep siete eiisenk oo esae eas Geers es Sees Gee sys mh tube ionvha cae ayre sills ayes ure es III, 30 

FS OUUG CCA) U1 1y Meee tee cee mee U Pre -cce ee tcataienns Meese tae ete oe aie vince tiers 

Glia SEU reenere ene crsea te nmemetet ae cient tote, Meas. 6 stenc ste oos oavintye eve. eeie oxtail ee II, 12 

SASS LITIMM ere ee Te enete ecient asics slates aca AR Pog ork toot ane ee oiSuatin'nva geod ROH I, 15 

OU UL NMR esc emen ToMen tener swe wae tess cue o1act ee aye ou auevel Re) ae ire 

EOD CITT meter neyetete tener ees iets ots ai ee cneccono iets Bo fe az. dod deeetvbs Benehtees.« II, 15 

GidyMoOdontiunt oo... ccc cece ccc ec cc cecucuceencaceveusctecevevevees he 

Ilirenbercianiiin eae. secs ses oe sda. oes ws aveeves sleces dccecsece- tet 

DMO ICOT OUI bn aie ep ateseciniese SR so Seales Salad Sas Babs Dak ak he Shas II, 9 

Ue eM UMN eae eee ee ne tye eerie Sec vs Sek asa tk cee eau oisee ae ist anche oe I, 18 

POMCeGLOliuti ea ane cian eee ee ie Ue a Minow eden eda enki tl, 311 

Weutalinn DAVCLiTmereatere ene ei eet te ier Sia eae sc ens TR eraios ad li rs 

PB eraRMIITENIATACIINN 1p vseezssseiase ay Gun Cs eee eae erect Sewn arte ae aR wood tea ears I rE 


48 SMITHSONIAN MISCELLANEOUS COLLECTIONS vor. 85 


Bryum—Continued. 


VUES OIA nese Gealecon Gt oh opehihe Teele) Sear aacuahis oeseed Neue AINE oper Veh enes Chern eeeNIS oe Seamer TT K@ 
{TAG HO DALAT heaters ee Slee cies acs ioeh eearaene Che eee Mee reReRe te eer ater Tule eres! 
FULATDAE TAL TaLy eerie ce eh aes cycthe cot suntan lata cgRateG Neat aele eaten aaetace date ee ei epee eters I, 18 
COSA tear eee Sac Bien a sie Wac ettre censuewonauuee eaiictlens Gieictiens euemaccieas oheeeme uel eee Das 
SPECLUDALEN, Fixo% bynpec edn aioe oleae ert Ieee Maloferalete srelei 6 epee arene ILG 
SCMAT EU OSIM ie-ciereys owicsonvs Ghee histones Bevae eben eee eImeIs s apaert en eee eres 
ISD CHUN SEMA racine oe etic lehclciete ete ciel enir clever elei ces) cro iovectorehs ef rei he tetetcvers Ir 
Stubelimbattn, 225. cs wih wons.cror a once ge deledvcistos sete tiersyele elo.cis na seattle Liang 
Campy lium *chiry, Sop lay lenin sa. oo 2st tie es elaiwte eye elevelaeteie: = lle olereieve «re reretete er -tote ines 
hispidulum: Sommienteltti cjasmeec sacise sale stlenclelecias eccleeteeieee 1. ann 
Gampylopius, ame mstisal ters; ye vererssccchansvevoi crete stele evel ieleuate ecetyael iene cel eaerer erates hi 
AOlMait els seiccdes ssi ers @ alo cepts oreyeoveen ieee bneeral iterate ctegen aslo Sperone Lie 
DASES OTE: Sets a ceee ca lana Ors ah ece Ua oie tek UMN rakesta roe eeee ole TeL tote ouslanat Wnenatenner eer 125 
AVS CEI: Face als eis is ote LAGS o Meu OLS OHS Osea NO BARS ENCES eue Te oMersy eed styebehel cre Reto 2 
Ghristiarty scecetoats Gok cAea Ocean Oe eee EOUor er ierat re latene Teo ese 
GESEUCENUS. 2:5, & cece. adverse ds aevcvoraherave esersie, OWS eLel eras Ss Gis mussel aera) w ahateeeheaiterens Te 
LITER OPE SUIS! cis th srs shu Ses Sede ae ealsie iat che tla Paral custo ay witsbegetciren ou Stevslist so aoneriete tat eel ees 
TTIATATENISIS® ances deo oh ctNauan avon cols de a tara ete ctioye Ye el cvaLeliellatansr-e/orelleyawe lavabeeroderamer acre III, 4 
{TREK CALITIS * Vorsieearsuee devas, Sateeshraaay casa be tcue ate (itnavahorete Waa cs a ateieeoley sxe se toearaee Lone I,7 

BP TEELE LE as gr tesco vere roar een oe tep a ra tad at ole Late aver™m ACR NSs eyelet Suche ere Tae s7eh a eee aenee Tit 
PUUSHLTTTS, so, orace, scislels/ aoe are crercva cei emma aver Haver oye! oye oTnlare ede) oilers alaTehe te etetenate rs ttctode IG 

TR OGUTT cctv Sea carey cies eR: ar ciate nat olerete ei Gie ee I ere ee ere I, 8 

G aint PV eREEI/ lees eonapaeete i toera Davegoc deresaistua te (orators chee ol ote aveweeetro mipeirenotey sreyeenes hens eas 
SLID ELIT RACES ees aces tie cecc hovel Neferieyc ellos fo onstel oy exerci oesicvatatsloretedaneuct foley eketareg rey tonne Te, 
Geratodomm put purettse teres crests elevererarsje.ollelet\ste sie) eo iehlcie Mietelounverelobevovetey<l-tey fereere Ae 
SEEMO CAT PUSi un. esl care eneyelcsterehastnt = spel aae elle el ctetetnt tetiar silage aiedctsy eneractors I, 23 tie 
GOscinodomeATSEMe|ty ste crcicre see erete aravere le tersovellelajotals ietaTers aieraial"s\n/ote’= @' my efatel atenetetele II, 4 
WYieiciintai. savarse erase seciapom Soule celatsvorney cates or Ate intel sisioueye tava eke arneorsy Retainers Li 
Gry phaea apiculatar caterers ets eles oss te aaueyeray stene exayaneteteeetoransi ye 1 rareer eats II, 16 
ACHE AL A eer ene ere eee ey aren eLeR STDC Te rere seN Ae raledeisteliosevedele oncaclcleveret te aane teen kenors II, 16 
fiLT RO TALS doc eee ence rs ese arm soco ape oie lO aie daceeanoteoueyol over ote otulstaerch dereinnsecteterotaeke II, 16 
OLIZAD AS cere, aaiete cere ee ee a rat eae one Ge eatin wteieicnan wie oe eos Meyer aoe ae Il, 26 
PAESHISE carshate gale sncteere ciere'at'e whet ieicr oshoueler stale) she illest ler sYat=ioats) a avelietr icy e baleNeketaxel Thay 
CE@CUIT CTS a nssseie se el cece oo chen eveves seh aiene ee relat orsiistel clays (eleietonevouapenerensires cciet TS ey, 

SF LOLIE EL eet ere ere SUS ike FIST ae ret Terre vNaVON odsue ol taeelelsToreterhstchea cesta 1, 07, 
Cyclodictyon! albicans © 27 ji. si. 20 -lefersye eee aie no eieos nee eterna ene ores lize 
INTSCHOIME Ronee het ert ie Soe miei tehe rior eek recht er kre II, 20 
Etre Cha Currant tetoes avo ae avee eG OGLE asie erceselchsi hate tototay Repenel ster Wen-tone II, 20 
Tete lyrraatririd eco rhe avec etal ees Sector Tote feted clo shehoton oneal nel eel heNeomeretenexe ie ett: Tl ven 
Dactylhymenium Pringlei .............- eee e eee e cece center renee eee Ill, 17 
Dendropogonella rufescens ........-..ee eset cette tet teen ents TL a7, 
Bicratellas varicie cceccassctaretetere eee tera toro 01c1 ole ite elereleley sxe 6 he Mics stiles rebalecete oOo rete I, 
DicranOdOntuint COSEGTICENSE ccc do. ocise wi ciniern = wie nics oisles acleibia sosisie'e wicinielare sie ieee 
DiCrAanUm Ce SETUCTLC ec eater oe eros) ores ie teree tect oink oustiotcks = erevalieder si(el skoteksi afer orerst ere talents iia 
EFT OALII cece oc «5 oor eiacos eco rene Chetatae's tale #1 Serna te aie (otere inieiansls antes eee ES 
Didymodon campylocarpus ...----- +e see e eee e eee e eee teense eee IT, x6 
Giciplanmobasise ve 2s ict kpc certs asrelielt ate tal eee yarornwserelervtnsees orenaletacs memes III, 16 

CAA COS VAITICIS eet aA rere tortie ler are eye cue a iekeiakey atc tenctebey clove eet shoberei we III, 16 
INCraSSAtOlitmbatts) cic ce mteis uae oe ces oshetelere ce shencterehelesoleeltererenotonenstentorare III, 16 


ATLCUCATIUIG esta ey ehe Ser een aria e feveenae ef okeneis eactey eno PeceteneNc totsielrerleiem eer choc ronan OT a7, 





NO. 4 MEXICAN MOSSES—THERIOT AQ 


Didymodon—C ontinued. 


OEIC CLS ee ere rete eNc Pare re ever ce eeiers Petes stereos oss. eter oreeeh rete sae nse Suess III, 16 

DS UCTUCUNO INTIS eee arava sesvevensleievcteierelcrciciets ole nic ieaeis ocho ora eee oedema acoso: Li x5 
PUTS TT Seemann erecstet oemie reece renee tere arere etc nerc ek ere tomer siete eects cates oe III, 16 
LOU MAS CON Steet aerer cere) otet cet ereh crates etsee tet cutee zea eee or aie aoe ae Cave oss Tis 
MDSti cline Gaia Cetin merece ceeveretec stone cremate. cee Dee crete s eile ae orsr oe wht cueue ake Ill, x 
Drepanocladus exannulatus meXiCanus ....... 6.60. e eee sense reese Mies 
PSM ct Gee aargerieie, aes Greta ater 6 aesrcee cols geal ena Grd owe Ghai oe Seka Piss 
PECIGOPOLNE CUMIN: AMADUE oo 5% slc.sis.cicceie:o.co.0 48 mcileseo asr010 acvle. ore alevniste ele shvie oes Ill, 43 
TMUOC OM, SAD DGEVIALUS) se1ec susedsun cients od a sutsevalee viele ole rere [27 see S55) ci, Az 
ELIF S'S COM SUMP ra eters etna riay ows enlarges tran aianstoten caine Re eee i as 

BRC VAD CSM me ene ls eset tena Saat cccihh ua odie tended cee ahy aie, die tie III, 42 

Clay LEO USI DLE VISCUUIS Et sir. tete ete cstte lore 6 ced sa've.cs saleuens a caee'e I, 26; Il, 2 
AIEEE CT US ween tier Ae eee sates oa yeah vans Shay Ses ante tog acces tend i's audattsveees she I, 26 
TILE XU CALAIS Meee vee te ey trey ester te yaa allay du ah ust ive tease n Gata feces eaves: os ezOse Lila 

VitiTem Chitmmerrete oo ee eee ic teas ee nee GM CRS bin bad. odeottne bus’ Attonte a I, 26 
AIFASTTT GS Oni eens sie Bh ya oe te cesta oteee e saceieé vaievs ccs parte at failed ops a Rpepees ee 1, Ax 
PoMplery Sil MEKICAMUM |. hes vies dba w csaveulacaowes ers le Sis tile 27, 
BOP IEC Ie Co Raters Seaeae ee na Nias. air o.< eats plate detiek OG aati O LR iiss 
Brytnnrodontinm Cylndricaule’ ccc casas sees en ee eae see vs ce eee 45/5 IN 26) 
METS (itrtmen ar teem tee ieee repre ona as Tacs apne Gye Re Varad eae esterase ern caine 27, 
rEeysito WITT sya se easier tenner Maye clean noet ne aie sister eure een eer 

CULE ENG ULE EEL! | Rae oP aE Pee Lee27 
MOT AS CULT Ik mee Aedes erie Nee ena corse irne ria cian ere re aesie ie emiee Resta II, 26 
ret] Oo Ot Mente yateet rarer atinceeveyeerneuenee eeatinicuss Airlie lenin sre aisieierere esos oe 2, 
EOS Rca AR My irene Eales Choestatcj sits ey aveisae stage el usunra Sec verctey cera na eee ties 
FetieMyMehi tim StOKeSIl . as agaidc me aienie sense ners aes dec cues un eee on TA 
EMILE ahha lta c eed maha Pod Rhijsed beat G eee oon a estat oe ate III, 4o 
ETUCLISS tilele UUTM Iaeyeuereisyanereremaceeacncrene ee meitiie oence eee eae Tee III, 40 
EUSTICHUUNG MOTUCOICUM. .05ba<B5 es evncdlew asides cieeascuceecductsseesees 2 
ao Orice CENtata panto oais.cise- vaineutrs Ware Avcusislaie crsisfs ee csess deci nev ermece ernest Jel sear 
HA ATICTAVA'S oaks eas eyace a eetnera seenayeeseicrere siete sitesi srare serene ciate eee ez re lier 
CGtO DIE Diarisen eae eae ia tee einai eee. ear 
ALTE TIGA ae Crane pay ere oye yoo ian Meaney Te Ri tvante) oeroet eee earns eer 
TENE Nel LNT cla eee eas eee ee Rea er eee een II, 21 
PVECIIUIN OA ae ye ontva is Pe nace cesriciomers oi ecise titra emies ones 28 ssl 27 
BIER ICAU «ate achat Aion ig Ae acre sonnei aee, sho aC eins RGAE les Gee Es I, 28 
ISSTC EMS, ACCUALISS o o.c..0.5 bee trove oes stein oo olan aie ste oe eld) sisie a cle eas eecedaediees I, 10, Ir 
PMTISEMEl y ereacis ain oncieeriesenToeieiciciel steerer riers MI ee ile dns Sam bles I, 8 

AAS PUCIMLOLC ES o,e00 crc eens eieyeuste «ere eve 4, Atal e nEeoe te prepaa ite hudba sealed I, 14 

ENT COUAT Cl lens, Sratstele vere emeiere tekinete cuss stir orerene series Smee oneal hicre sews irr; 13 
Nha yas stays re vi Byer ee Seg gees cies wieises yas) yield Ges ARS he eoald b: aigetT ey 
ARNO STG 5 gees Ne wre Gane cer stieae Geer eae or ortega os ade aie ln I, 14 
emICraspedOnhy tS) dat acess veces es oases demas Save de beeioeead I, 12 
Eeiat ya tcl las pe creepenetrcuae seen ee eee tee eeoreaeieass ete ee eee eee ee ce 4 I, 8 
NOMCTCECUTHENS 4 asc esc ececisiee so secant eee ee feed ein eb dece ee I, 10 
MINEO ACAI S: aewey vee ereinmeetere crtiseaiclevcle rey siecaiarercistans meee eras ees reed cncvy? aus Lar 
PCA IOL SOU warraine ke cts eae aged chee ee ateicuveis eee once Sea ok ete cod aevie aloe ek [12 
PR OPROVIMES) ec agate oes Se0 Sem sth aigte saree oe one a4 a oa cen ioe 6 cee I, 9 


Fg MAGS Rares cee negara. n caies Chatee a Gyn coy be rerareton enue. ses Vn ted aie whine tend T8210 


50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


Fissidens—Continued. 


pseudo-exctlis y ona vsunc ote se Tatne conte Se ee Oe Ee eee 7 Hg 

Ra vene hits xin ay-tarescsraiclene.« oc Seti ore ei eee eke eerealerenetiie hey chon etna p eters enya 
POCIIMALUIUS.  dcicitccrsiee «CeO Oe Aer steeO oo CNG OUT ETOC TC een eR Ctra I9 
DHCD UROLIWS™ 2 een ne teak aaa neti tare te Reon ee secon eee cee cea ee Tee eerae I, 9 

PORCUMTS eats Saye beceeietdon anavdlra atte Bioncrac cle eueeet ae, Neagle potate pees steno Met eee I, 9 
brevifolivs. io. U eons e sociswe ine hab an oye see Moose eee aan 1) 
Funatia. annulatay i.) siSoucaie ne iennin a nee etOn oon oe eS Ee Sn eT ae Re Ie 
apiculatipilOSat Ach. sche eiows foe Seloers Os see om Ele seta ectae Ooi reece II, 5 

Cal VESCEMS) 4<.34¥ tee aeyexeit © oe sae alee soul “tones apse eral ake easier o: se 

CON VOliutay jeter veel soy cen See eee ee OE ener ee TG 
EPID SH OSTE RIA. ro. ccacacee oss orise okt BaSrlA ole cut CO DLE MTR RTO ECT Veen reer ae ees 

hy. orometriCay® os 4.<ccgioke s Ses Se eis a aloe eee oto a een ee sis eet ee ea LG 
Calvescens: Owes veal ecient: Cele ear tae ee ne ee eee Il, 6 

OLEH OPOMay us Hae Ree eseerese rele Te Sieees 1s a OT ee IE eee ees 
SABEODIL vonseaveote cee ert yore ta tolerates eae Ronse tated ST OKIE CEE Ee ee TS 
GrimmiaoArsenel:, octet scien saeiaeten ae Sues oo eine ciete s Seles ceo eee arenes 1 
CalifOrnical We. tieo Reach oie Oe ere Oe aS Oiler eee I 53) Wiis 
IMVOlUICEAtA 4 J. ter | .\besm re verucmaetare kore ereuntols te tale er eielonays orcreieds entrees erent Ill, 25 
Oval Late sie k eae sega sci oiecss ae eiails ec arere ea eyetay age siete osevoteneteraisnater stern eee II, 5 
practermissay 2: cei see yrmanue: mh rreseek Se eetotea ee aoe eae stoic ere oe tik 2s 

PU Ba Gee oh cect See wis Soe aren Saver SR wit abet eel ora oe auc orca 2 ee 2s 
Gymnostomunm calcareuniaa. amis ee erratic) ieieistoe eterna III, 6 
INCUL TONS etre LOS OS TTA ee NRE eo ae Ti, 8 
Gyroweisia- Obtusitolia® searentse cancaikrane os ener ten noe epee erator ere LLG 
pPapillOSa Wassaeecie dels | fe eekti hi, Sreilor are ater ace “arte ouaredets avelene tego se Panes a beporeyev oncom Ill, 6 
lekehoereleetebhonnal sonedojolanAlibhigls soqueccoq 4oancronaddgudondaccedacacsobagor II, 24 
ledwiciaalbicansp swt cire act koe ore ete ee eee eee eee I, 21 
Hedwisidiummeimberbes ameiccem peer comin ocean octet ism 
ELerpetine wrong WOCCOA!! s.screc «)aciciayorel tone etic cetera store ouedetelotodtvey uals logerele einen 24 
Eleterophyllumiiafiiie, Wace sere niente caterer reise sictel eo eus Late mora ene III, 38 
Pasta Ole scalars See hse es Oe ROS OOO Sa ELSI a a Ill, 38 
HolomitrivinisarbOreume versie fence or oe etek ere sella terse tear cn eee eA 
PLOLMPEH UM, <hy.c 5% wsita eedeasls ewes Sees ee, Beate SO one heme eased Ne eee ees 
SOUT CEMIN Mite cs ois caer ers ROBLES pe Bee SECC eT ee ee Le sim 
Hiusnotiella: -Palamert ccs. cis crsmue see oo eie oe oeust rosie everapale eo slete Glebesneneicrs eraeyars Te; 
TEVOUIEAL cy vileiahine totttare Moe, Mop Set olucic le euler a onan caay.c) MORI SEERA RE TSTE ee RCW LE Ora 
CLE ee areas heehee a Bee eNOS IEG Pie TE Per ceed oreo eis roae Ee, 
Peal rv Or thy cos oeesy fel denies tase creer ecole alan et ctereh telcnel Mepetetee re Ralerteret one Meare Lies 
TOPQUESCENS? Cote fit 2 eel tin tesian ee Seer ear eee el eaten ae caer eats, eeestie seer aence ae petoae TES 
Eby orohypntimispalusthe /sapre vtemicicctetor Srertestine se oeiedtetne eter ra ere iiss 
Hylocomum Ehrenberguamum soa... otic os st cece essa sess cys se ne III, 44 
HiymenostomumsimexXiCanuin > sic mitoras ssetvaseia terol ie onatotede ts toe cil atotescoiste eee 1,16 
SMIUCIaphanutms co se cava sete ret ais iaclehe veel seer alee olteneor onto oes eetenew ene Ill, 5 

Eby menostyliumCurgyinoStre tacsin clin rrr eke ce sitet re y-rteter rer eres Ill, 8 
orb} 650) eee NS CI ico acaciocn cir cine arc omiGo ce atta salted on III, 8 
INICUEVANS awe) racic oe Ge a Oe ee De ener es 
Hyophilavangustiiolia ..s aay ce natant s n(n also cats ele esye anchotelala Simei a a oro ae III, 14 
Bescherellei® ..dns4 Siesc.s dere oe Nee TO Ie Oo ee eee Tike 


CLE MEAEA  ieccar ag Saensna tA BzeiaSere fe ota mioueleie lee teceatiahe oi holes te ate oie Rete PT, 13 








NO. 4 MEXICAN MOSSES—THERIOT 51 
Hyophila—Continued. 
Hats) all Sem Sees eevee gene ov aren tanetee tee veyetay cueite ome evens fe cosa romtinroe er coc tayias ah cerians wuioiay ait irs 
AT Csxal (Uk) Asura Pe en eear Gee ee enN SUE err tee Neier err Gcn eect se @adale) stnkege keieuckarete ic Pil. rs 
SUDAN PUIStINO lias epumeyse meeieiclterele meoteecaierei asics rere dee ees tee enol III, 14 
elaygi OTL U (Tle CEL UG Cet Semleneperene nV oteaetene ei enehcter et seve eier dee: cls sk ciniaterst ores: siti ayn. ti eudssarbusicce Te20 
ANNI <LI Caen ees utente tee were Sen aye I ICT rates ote oni pohyan ares ene e hd dh 4 III, 43 
CHEN SO PI LLUGIN sarorerace oie cotnersiascstiavecs Saathsnscere © lelse ete rar'sroialete: olsnselavelle aie. axaltore at 
GLOLOUEIL GOVE ET eS Tee TA Se Mises Bod ee eee IBA ei 
GOPENO LMETH rete ere ae Tate RSet us Cites Ries asta leah Sas EERO ae LT 35 
LECTIN CHS mre Ree tan oo eee skier aes nGBn Ca SHEN oto ANNA foe eee IAD 
PAN CIUD CONAN UIM ct se specdeT lesbo lec ss fo se, Bae oetS Seta eoAeeT tse Misiare ub ae Oaoei III, 44 
LGATUIGTILLOSWU THEM metanees orem coreMe dee aPSEE tego asl Seon a fe sT ea. wee Sakae Petes ectieha a 35 
WarbOmalcOnutt oo. 0c cece cece ccccrsenscce tee bevettuteecccersess Les) 
VOD EOUUCH OG OM DUT aeetee ee srasstden totisacusteies oid ates sors sie) sais Gis Sees F saNeNS 68s III, 30 
FLOVGH.O PHL LLU MMER et siete peee soit eRe ele a MS ett coco fs Ge evaded ce alps < S aence SeETERENE sore II, 24 
SLLIES GUO TTILC Met eadare, cia SaoN aa esr ashes eerdees SRS 4a. 16ad) Me dh aubys eis seners vised wee yea tees DU ee 
LIUCUUSLG CULTIUMMMPRCCMNET terest AS sae fei ioca ashe "as areas QIAKOE ints ciate. ote oer III, 43 
EISUCILLC CRM RR eg TES ARs 0 oh Peg cake Hata =, 6 oe eter tg PE Oh cea As 
lsopterycium) cylindnicarpim, 2. ).2.-62 0s. csc eens eee meee ne es ss US 
LCVOUSST VLU TN metero coe otk tee asta aL oc Sette nied ules chstens sess, Goede aciotewste teen Teas 
ADA TEE CMM escalate coreg ses Iw avec he GO Ate Sire WA sass 
WWeptobiry um) PInMrOnmNe e244) .ccuce ease sss cee eae tere -ses+ scans euessuas I, 15 
PD ViLgLAO INIT Cemetery sacar rt ete ors Be ee aN anaes Caceres teeire siesee n sunray ene to 27 
Leptodontium angustinerve ......... 0... cc ce cee cece ee cee eee eaes III, ro 
PENSTSS TL Clie Wades osha Font Le PaMCNe UES A Cay cfs Sees IES cs ates ous, Span cea Tee a we cigtanenecensie os III, ro 
NaS Cll Ul] fleets reenter emer reeteeteiterc terete eae ciercier aereeae eas Lenn 
LCS CEN Summer aera emt c a tetor sere ier ce ec III, 10, 14 
MelICOIdeSp eee ee eee eet eee eee eee I, 15 
SU Llatgs@ SULIMLp ae ueneners sr tererrelcren terse ecineaey eee sitemeter recs acleereeentioreeiconen ie Ill, 11 
TAL Gea liver eaten ep oy aueee tute cent a avr cue enene cies eierece etree COs ores a eaare eee eas 
Leptohymenium Ehrenbergianum .............00ecee eee eens bees III, 44 
Leptotrichii leptocarpum .....0ccc veces cccce seme ne ccsta nt seessessuvavs I, 8 
BASIC OP ON PIQWOUL. oss ters aie sae etsiciers oe noe'e ere, sate ene, dae) sieve) renee G¥ op Biss bie scien Ill, 34 
PILEALUG OGY pA eINrye erat ie rete ee eee aoe Ore ee oe rio II, 22 
Et COUMOMMCHYPtOteGaman ame serie tee rieereeicieteieer te ere leis ere ee tie awe rz. 
Say EUS US tnnmenam haute Rian wma es ee Geis a Patna s ewes sees «es. oe 24 
Bima pencias mexicana, 2. ev. saueis situate ele aisieiei sis aie0sg slesscosu rece ere echomte, 2 ly 225,23 
AGU IM Ate eke mrorneee erat te iceiceereeecee ieee eee hceiopeie es Il, 23 
Opal el roe ver ee oie ce yey cn eyeraeueue ere ser ecticee aieneleetene teen enn Ie Tr el enti II, 23 
Macromitrium flexuosum ......00c ccc vec cec cc cevcuceececcsesevevcettes II, 16 
Rete SECO TIM secsyn as yeriiute eee sy asitee so ae en are orien tetas dla xe Ck es ans 
IDO Cte paca erers ayaa riciraeee en erciere emitter Tae eiass cry, Sere omhectae thar Its 
TIN Gel CEUNACT TTD shay chase eer ey onsts veeRye eee Vn ene eee ie ones ohare tan ce ex aiaes. Occasions sansa ohne II, 16 
PYGMOp iy ltt rege seeeste ey epee trace nies sete eter ere noe a Mieneye ce cart. II, 16 
ee ei os ee eres rena aera we eee are ree cash Sletee ssh es sha Seba MB yn Po II, 16 
LOMEMOS tle meaennencvercrsverenerevensusinicrerceneearie areca cae eine ere ee tenet Il, 16 
Miefeoriumillecebriutm <a cs 4 cferere so ee ee cel scle ae oa. conelewies soocusse. 6 II, 18 
HELE LILO EI Cie serve ageiaiets epee earch crarde viele cise Moers eect ain Sraneoe cat II, 18 
Wietzlerella scoStaniGensis: ¢..00c0.ces- cave suiees seecneecewsemicevivudecten Li 2 
Neotel T3102 ary oy apse tan oreoueh i oensienea ay sreconrereetye hole terest che einer rere saaeie even eee 


52 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 





Microthammnium. subthelistectuml \..oq-celece eo ceiice sceictcneelstenerecteuets Ill, 44 
tel TSC meta a, < chs laleas specie Gi ck Metio-a) seca ayes emeenerone eiigatana itesar alters tag rewee ten TA 
IMB Sa noone Seimei a goocokoonsoedonuiscn cu dasoddubooopoeDec TY 25 
Mittenothamnium subthelistegum 22.0... ce cece c sees scons seeesencsees III, 44 
Mintobrytim albicans.) oo c.csraie ici ios ceteis te baie icles sc orera ie terete 
AMES SU Uta, Geog pises oes Chews ease eseacacel ett renee pedals ee ane astetene chase hatchet coher eee III, 28 
Menivim walbrcans\ a: Aeacaditeersthee ree oe ee teas CIS: eect cine noe ene Toe ae 
POS EATUITINE meters epee lars iavekakerenelayererelsiiev er cuoh aN ehencietetenaneyre ovetarel loch anc tator Nene Me ice i estetens 
Milendoa -obtusifiolias .a2.. i422 eaten cos ocesee bate orc ee eer Ill 4 
{NGLASSAtA. sais ool ere ers alee Rie aie che Oe Nee chen ere ee) olein: rete eee Lie | 
Moriniasl hrenberoitamea cfc cti ste = oro crctars close segalotetaget ete fareleee nel teterctet Aletta Like | 
PUIG SEOULOLLES™ xcs eed eadavs HORT INE eets to ore weotatoney aI ehedhe Ree tO ere Ti 22 | 
Niecleeran chiloracatslis: caged snc cates sana ae crscn nore crolenciay Sstepene eicte layer sieyeinre eaehe II, 19 | 
lornschtichtata <5 seis ces wisterstete etsy arenas = eielolsierietetenerereie omic overs lot teters II, 19 | 
MIZE SBI eee oe Rae A Te ENS ele ote STG ales tel aoa etan eee hechctes Naan II, 18 | 
LONGUS CEG he oa oxciereishe ote artes ie niet Meroe eoesn tat te sense Nel elo Coteaeh ee II, 26 
OPDIGhVGNG 2 oda cine co aveees visinns Stine tle efaleler= @ wale le) m (ress olsie sey iav aioe II, 19 
BETES AS eee OS TAS SR eee NaNO OTe CO aires Cee ae es Il, 25 
FUP GOSCOMS © esc wichosaisisnyeverar8/ ey shalin ae) ese ei ese tele tie) «i ein ofore! oats tol aelaera esac TL 
Nieocardotia cubmigra sc cco ace scrim telole + alaynininlajesat= alot siete Giot-poteregeuelers Lire 
Oreas yy Niartiamayecs cicero cetecreie oral evar elciote lor crch atete clteneieneraeite te enn Narcrerse Jules 
TOK ICAO. &cvecciecdasoarecrw arti ity lads ooied eps sien oiera mis oe emer eel oO stave oteuctotie as Ronstore ie 
Oxrthodicranumpetlawellare: 426.4 weirs selore wlete ee ue cyelete, ow errpelene Miel=ts ache or eres ee eA 
Orthotrichum’ diamham tne aco e cess eee iy lee eis UR eye exe recente Tos 
TSO Z ATOM ere EAST onion le Facet on eveneNo rsa Pech ane DT, a6 
Taal ACOp Mayline tere eters ete tetacad eos) ona ls oiete sellin over ceed ace ep eyetaelenece vel ero kenele Th oes 
pYCTOp liyelltatray ertstee mre tsre wie rere vsl ee cae Osa er ole lye mileysi ofebereteranio tele a lotesenteenere ers 
RECUEVAIISE Meee rae EE CI eon oe oe oreo Ncrenlotcycketeere Liens 
Papiblaria appressa qo toca ce eeierede cic fete es crensh vue wuctate fafa tone orckaienecels terse a tate II, 18 
DG ppeiy woot eae erate ess eas eer eto tatete io ence tncetote cleioteaeae ate a iea ven she II, 18 
EAH Tite ree, on cetera Se ee a Te eee Nee ee toeneetet II, 18 
TIUSTESCEMS. Ses eteieieiel cies cease Ma etesehae renr fe Reve reat eda eke: her teeter eee II, 18 
STAD Ade OL teh er ee ae ee oe NI SNS Ios ed eiict nce g heey arsenic Cet Reena ee II, 18 
Philonotis: amblyoblastas  aajace otis otk liek trope Voteloo s oveneovercteren ete ole otenaTerayre rere I, £9 
CUT Vat annie neice ote te ate aC ere ceener tenet eto choice tol atone on ean retest neni Remade I, 19 
Clesamtuilgy ccs: erect ci aienneere teenie oparean ome erorare Gieleter ete ccefete lazer tier aero I, 19 
STAMUMICOIA a aynins <a a e Cee ee tenet ane ke wld teaters oe telene oka teter ies tele [, ro 
NA TTe CONS1S enone ork ae eee abayete eater a eee ens reteset NOI acto ceee sees a ctemeretee I, 20 
TURE CEAL CA otitis erccvag cs ote ere eet eielcw oN oneTe oreo eC ekcpet eels orelionever cde tarsi ere eeey I, 19 
TAAL CATS. Sea etars spac pense erate Oe Meira erate edsiote lene Leiria oe eens Lao 
WETAGLALISY aves se iechresstc oe caverec Deeg aay eon leet oapst econ eko ears tees Steer tel eae feet cnet I, 21g 
Pilotrichella cflescillish eesmersow ecient ioeioeteys aero terieter eet ey rneisr terete II, 18 
FUP GOST OMS iid aa Shae ROTA Ia einer ee See Oe eee 118 
Pylothichumutasciculatumls acre rie etote tere ren ri ereicne(naretereretsens Il 2 
ATIC ICANT TI day ae rese ts tate ott enasa eleeie cea sien ese ORO ee OT Ieee II, 19 
Platyeyriella shelicodontoidesie tac ccrec ers creates tcc stoke relator poleteheteoltehter eae 


imbricatitolia neta eee eee CeCe eee Se ele 


NO. 4 MEXICAN MOSSES—THERIOT 53 
iatynypnidiain ODEUSHOMUAD 66s. cc cuss ose ve Go ccs o's « seeid vas eearw ates ie ss 
SUG UAL MEET reso ere rere Srecetaraie cite nici east es ico non ears ace hor eau oes lees 
Pettt1 es Ud eer tere che ae RII eo a aches acted Aes aise tetas Mood Lit, 33 
SSNS LU SESLA COE Ig 1Ot els taper a Cyne ese e's a ee GE hes doar are es Wc a a sivas iiiess 
lenrochaete Mitel vaca aac ein ssc fe oo bee arcla lees or Gtelecece so2ene'eereealy ecets'avv eave I, 14 
TILELGUG OT] CL RAINE RO TO rhe tect dae T cA A a ent Be beh a Ne I,.14 
SCL Tg OS mercenaria ie See Mvag e Mee S PNAS cas cake auction spt cesblac aye CeMaNen ioe feta ta ate este Pers 
Pets HOM DIA] Se ae ean eeie aha deus Sia ais abiac ateiads enlerdiata comes ae hes DM 3% 
Reo OMIALLITY yA ESI Oasis Gos, susiera tate a, aisye ¥ ACh 6: ee wd ase had aoasecdee oS wy Ctedordcg a weet ian 
BS ESC HEE CIEL OMe prem cyan ls 2c: Mhe. eR Saha.) Spain So ese Una. oO a III, 44 
ILL CED CULS Camm ee Re Sen ela, MEER tay GR py ae ar 5s / USE yesh nds PSUS HERS tae gous) oft enc et Mons 2 
OVILO SULIT UMMC RSM Pepe eNe ary nes as Us ake honey cus ae eee dh diavel at TE Se tO ise eaeeS I, 20 

GUS PIC Atlin cet oer mas Ooms ieee: Naked sei erSue Oo an eatin evateicey ores Ti 26 

Gy ci CUrtip 2 cPepeicteke orcie oe shoe bes Ge leersiers e506 Hg ssi to sso wiscatee nadine oe aye arsé ane I, 20 

Efe Sot SLO TTT alee reyes sae oud toe eaiyore oso da Payee thes fon sesh avandia chaser sc eerdite Bek Rdae tues I, 20 

WO ZA OUR heer tin ok eto ae Sean ores ASHER Nee rae hs Gath atnad II, 26 
IROZATIO Ihrer Ma tere tens, Aerie oer creme ie cians ciate Sieve eicleie e tiriaiere memes cre teeter: I, 20 
EOIMICSTS Camere eter tena areyare acetone ots mucky ase vey cre ietgis ee vere fee ae RG osialareis ler 

Fad (E10) 5) 01) <a ee aE ge ee ae 

PTR AUCH LE CONG o5:a¥ ate his aihtiniatdigin as Bina eae GG dd idseee: $614 49 Hiaserecdeea me done i 
olytrichtim calpintiorin€ ..c6--e..e00c oss sees cs senes sense we. le Tae Dl maea 
ATLUTl eUTstd te eueverete eyelets siete arte ei sicimantens seco er oieiiel aie ierelcneretereieror tea reer eke J, 21 
MLITIME TI MUTIN, eyetes aerpteeises eis ere esters sce erereree ie ves eretereuetere siaeiaiere [een leas G 

He TeOTNG Gl OLY NeESETI Clap foycusaaieveveroievene sierevcte site sities errs crekaycas stein sir ietel es kare oterocicde I, 25 
GiliciteiSumeterasta ier aceemerstierey cient icietete ec Steise eerie ciate ets ecto: I, 26 
GIs Smee ge erer tay recreate cietceorsiseticn sae tare athe eee ees I, 24 
VAEVAVISCUL ti Speers reeneien eres reterweneiene siete tstei eaters oy ctescu ec is 5 Ween Gee 
LCG DOC IITs neem eros erence rece Skate ce eh dare nt I, 26 
MITE CAMS, caper ee saetere einie eo etoncnaiers terete ears si aehae Cote eee ce witiccs oem 4 
site a CHanererearsiscis reise cise teri neers earn Mea ee, 2 len 
TASCUUOLES CMe SUUCULCHUIGLO: sere aero accins 2 ee Seine pede © veel Sieicus ie ieeteros « II, 24 
Eterigynandrum filifiorme mexicanum ...... 555+ ieee .e0b bees e de dew'ws Ill, 41 
EAeCLODIGVONSIS Illex! CANcdap ere emirates aiieiere seeinciione aarti ea esiare esc Ik 17 
IESstelTd Oi tomes erases sialekeresversuetarersbens or ieierers camisetas ei eet sxere on ar hee Is Beal TE 

Gy GHOM Ute pI OMiititItieerel ei olor wits eestor eet aecters-ceie <6 e e selsha (ous ocae x Ill, 30 
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ITILEL ICC 1d Men ee eee ReCn ee Reena tii ter ieercrcien tennessee eleva sities c10 oe 1,28 

SC IIMAMCEL. sae pad acabaw sede toe sajaietviadls oho Ladi eacteteie 2 OF Decachydatiaie I, 28 

Sil biiell Gata mane Na ory airara iter aes tesie amen eet aeons usd. 6 Se tea wa nies I, 28 
RATES GC ACCII 111 <1 Cl alee teee Neer terror areustecvereate te ise kaltc1cue sites giendveusetle aka ee cs I 24 
erigcl dia eGchleatitGlia: S222 5.01 «oer gos suse ig Save acaete burg aeiwralaSated'eed II, 17 
Maco pilin latistipulacetitmess smc desis cf ecin« eae = wesc «vice ase ev cerecfs one I, 29 
EOUMELITOSTILIIN exeressererere ers cae re eee eter tere ee Caio ree ttrede cee St ed Sse chictits afisk Su, S88 I, 29 
napoidoprhynchitim dectimbens ..2ecae. 2% oes sla sca ors weiss woe ses os lil 42 
GME OScratlim meena eter evertererstere Corre ims rchecces ae hae ous Malo cgeiayaee < iia 
Rhaphidostegium caespitosum laticuspidatum ..........: Me ers eee ITT, 42 


incom focommtilt roLmlsieemeaeiersetneaicia sities cts ecco ecteietnctenreenie ook oc. e Li 22 


54 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 
RihynehostesiellasArsener ataeecas eck roe eer eee eee ee cere eee lil, 40 
VACQUI to widatee.c Aer lot oe Sel eae Oe SOE Ee OU eee III, 40 
Meesdaliil ..cheseecsdeacvctas ie MESO AL ACEC CR eer eer Cio eer III, 40 
Rhynchostesium selamper accep acc cee acie eit heir LL 42 
ttitOmalComumay %.,., cise dea apoiice Gavtustsnn athens Wave ooo eaters eee eae III, 39 
LEPLOMEROCAN DUM sce niekia cree serse eae Seas Ie OO eee eee ili 349 
WUGLACOCTO DUAN. totat cis diac syaraue sd eustions ot aticone ash ekepeen ene EINE eR PISCE ieee 
OOUSUFOLININ, oaioraierecdte sin Nafta eae ache Rese ee a oo Oe COO eer TIT; 38 
PUG Le be ateieiaavateres 0 suai cD apeua tro el aeee a oe SO IO aoe ee Ss 
Sammt=Prerneiy. Bais cs oye oss ecco aco Sis ere dora Olena tone tea ee It, 39 
ROZea+ BOUTS Gata ver mec e icccuelsike ete tae tonee Ra eee ee i228 
SELICEAL .< svsdete « Sesis vos Bicvoder grsveucutivel omairoraley sicko wesee the eral Shek teers cla alc ce ae ene LIT, 4a 
Sematophyllumecaespitosum seer ceeeiece aie ein ee neiaron ner ane ne Ill, 42 
laticuspidaturnin tose remot teen eee ee oe ee ee Ta yA2 

DE CUMDEINS: ae Sika tks yd cn eee se NOE le OO ae Ee III, 42 
Elaimpei” «i csuaeseie cer Gee eee COC ee Oe OER ECE eee Ere Dies 
ODI GWer OSIM ATU 6 See Sacco Coe AES ee eee I 42 
Stereodon ataleatusir meme sco oer eee eee aT ee ee IT, “43 
SILO OLCOBUS) tevin fersjee See NI Cane ASIA oe Pics er ee 43 
Syvanblephanise melicophylilagerecicsspencettscietien eile creer ie enteric eer ee ie} 
siaxaphyllcimysplaniSsimumnenrsee cm cece iter ies teenie Ill 4s 
NARIRETE Vis ak Ne eye SPR Cea OOO hae oO De eee LS 5 
‘ayloriawtontelloidess saa. crac en eo eae Sore ete see eee III, 25 
ihuicditimaimest Canim ete eee orem eerie Enmore re I; 24 
MiTAC OFICIAL eee reer One cae Ion CCE 25 

OV EOC OV PUNY racks seas SRST Es OO Eee Ee ees 
HODUS HIT er epsesecs, ace sesso aioe Ble aie Sia outa orn) ene cars Maree Si eieraisie- oleae ee 25 

S Chiluimmbe ne erie varsectssecalee Meer ee meee GP one ae ERG eee is 
LOMLOVUSCUNALHE MINE UC ONIN ecotetsee leis eietele le eeclclel sire cieleieiolelcieieleiel = tle eter ti2s 
Aber Clee limite epee eras ie arrest eee ic IS EN ree NIP ae ee ee ee Tia 
AMA USTALAT 1. caters ress sreeyavoReuelfdeo tier c Suet govehceelece ions Revel re: cLevoten one aoe Ths 
‘Mimmuicllapanomalauers erect eee eee eee ee III, 9 
Sua mOrmal aye secceve wacko epane sooeease chan ere a areata ake ous see aa teen nese eve esis Recor Reger Ill, 9 
Tortulacamphidiacea® 22. cere ee eee en er ee ee ence Ill, 23 
GCOMPUSD (Foch x eee cs ge 6 oe ato ogee se Naialalat co Pete rotates aoe a oh ed eae oer Iii24 
COMME CECWS Scotian Guntakomes atene WO, = taueilees Bic Mt ales uereua cle) element tae Mesa reHST LCOS TT, 2s 

AA OAT Se mraye cre cote ei kiclyase Greteae koe ceo me Sierra ic a OTC IO CEs EMG eee Wi; 24 
OBUASTSS ITA Lee. acd cct oe eacre see eesnssct ocevs eae cnt rtNCes ean us ete cio ea ee Ill, 24 
COMME CHENS: sears Sac er oe roeice eon RCT eter ere tiers iis 
Papillosae wroevsiec pees nets eae ale este ee aero nreme pero III, 24 
parva latitoliay. 3,2 ot S-gcew ote eree was ste yciinve tacos sasere co eranctvarer tcccaansttoun epee Ill, “22 
PAG LOU, sports in tnt site, een Lagan oo bog aa vers) eS. tere Oe ays Say ey syatte Po ararenaaa eee Vere eee gees III, 24 
TUPI COlA Mra gy creuslshenead ues ens «, sto bateucopetsunyehtc vole mravel cleyaacin epee teen eee Til, 23 
SUDO NUGT OL, oo escih abNe ooeie oi ste von eeUeiee SS Re anne TET ShG Creuse ere ae Saeeerrovorst eer ayers TIT. 2 
TrichostomopsissCrispitolia, eee aaa ae ie oe Oe ee eee ee To 
GCUASSIFE LISI. ccm aieclereies avs aston cor caens sence nite tk euchaiionemennee caste cocker ney areere III, 9 
Mir1chostOmuUummncantpylOCA PUI) avers eysreleie aicratetcelotele) stoyeletaiere alehctenstieleraiereheoeers III, 16 
chiorophytlum (brevitoliumy @.er o. nsqsacr seer cies oer TIES 
(Nt mite head Sore etucts olga tower corsa ter cuvligc es sho cle oer Ne ere Tara hen eta Ill, 8 


TELVO VIET Suc: tse ds ratece se eee eee EEE Ue ALLS Site BSE TOT tee en ene III, 8 


NO. 4 MEXICAN MOSSES—THERIOT 55 


Trichostomum—C ontinued. 


DeMMDO UNE CAUITIUm teres is, eeeiere rs lercuer ster eset eye vs. cos airs csc. Meenas teatpmeseushara ese III, 8 
LLCO UU amarante charsieiens 6 iskcierNaielis cist ele ren cael arc fe Aleianc ye ae eke eieal $s, OG couse ts I, 14 
© ICEL ITEM OT ec earch lal oct ce oan ocoheaty setuthe dere cote III, 16 
ee ECUMOOL ROPE os cease 1c eo eja eee pO nGeNee esd oY Sie EER eR Ee hea De III, 9 
MeriG etre lar herEiUSiNed: Sarsac. aie creas ccs aidle lade .c0es sued as ee shece sundae Siwy vis III, 9 
Bose Cll cata leey C1 Cillatise Perenere se cvaretars Geese sres oy5s Fin ivan sicaiebe ot oadecd ke tcmttetiay nical home III, 43 
vee [espe COMTI TINA TicUt cameo ays eee: cece eee roe ose tee sage atau adestslay sea pe veh te sh bs taste Moma aie drese eee 
fava lAT1G ta] Gaeta reais reret eSNG ls Miele eiaxs yo aera Aver hata eyaa ee CoA a haat ses Shoe Tea 
RMLLIUCL IGG CE tare Sets Lopes Sie Sie RS RATE Silo BO yep eG Bu eeay Nea BUS esac take ce oe hoch ITI, 28 
CGV IN OM ON tla weaes shstlerss es ca qe sb ohans 21, Sa as ivciole-wagtna ahaha ee eA ies le2o 
RAVAGE mersrtepere eee etc ahs eat see uch ey eae RR Ay aith BAF ald Siobndystal ste vadiarolas Gleneoks III, 28 
ISGUIG Om sail lei ie. ct tateeeterercics everarey cleus rose: s/S)ace-aaevens wieie colores aieleue arvane eee /ee III, 28 
TAC GUL 01) LSM Ie yh Pearc AROA Se ssh esse OA aes) ata) a ss yauera syseynre es Tiegs Tih 28 
ZAC AMECCAT IA alahetcs cc aes! a. oiee take staie' aft ate ere endls: se tsabehs-ocdia 0nd ape. ected aeons os II, 7 
BMersIONSIs. SUCHOCAT DA, </s\os esis wis si eae oe ea cig ged ened syns ree wee See 10 ee 
SHO MAO OM bal meverts sin aeevaieiarel tues a eusiek cis wisielsny ysneye 6 Goad’ gee esis maner swve.g es IIT, 15 
PY COMON ODEUSITO]US Qencsscice nce sassw oe sass e gems sens eweessmaecvenes II, 14 
PUM OI isis feta st Ase hr iStr een ne etotvig hts ave ipa ear bb aot onayeriyseeatease ee 2 II, 14 
Seta OHO LITT Stata: systah are ores sHesayselateo es totais exaea le asec a1 sncweicl <n «, wuaregpnarecche were Il, 14 
ERRATUM. 





Part III, page 43 (Smiths. Misc. Coll. 85, no. 4)—For Isopterygium planis- 
simum Mitt. var. laxirete Thér., var. nov., substitute Taxiphyllum planissimum 
(Mitt.) Broth. var. laxirete Thér., var. nov. 








SMITHSONIAN MISCELLANEOUS COLLECTIONS 


VOLUME 85, NUMBER 5 


INFRA-RED ABSORPTION BANDS OF 
HYDROGEN CYANIDE IN GAS 
AND LIQUID 


BY 
F. S. BRACKETT 
Chief, Division of Radiation and Organisms, Smithsonian Institution 
AND 
URNER LIDDEL 
Fertilizer and Fixed Nitrogen Investigations 


Bureau of Chemistry and Soils, U.S. Department of Agriculture 





(PUBLICATION 3123) 


CITY OF WASHINGTON 
PUBLISHED BY THE SMITHSONIAN INSTITUTION 
AUGUST 5, 1931 





The Lord Baltimore Press 


BALTIMORE, MD., U. 8. A. 


INFRA-RED ABSORPTION BANDS OF HYDROGEN 
CYANIDE IN GAS AND LIQUID 


ByiS. BRACKET, 


CHIEF, DIVISION OF RADIATION AND ORGANISMS, SMITHSONIAN INSTITUTION, 


AND 
URNER LIDDEL, 


FERTILIZER AND FIXED NITROGEN INVESTIGATIONS 


BUREAU OF CHEMISTRY AND SOILS, U. S. DEPARTMENT OF AGRICULTURE 


INTRODUCTION 


The absorption spectrum of hydrogen cyanide in gas phase in the 
region from 3u to 15m was investigated originally by W. Burmeister,’ 
and more recently at higher resolution by E. F. Barker.” These investi- 
gations have shown the presence of a strong doublet at 14, weaker 
bands at 7p, 4.74, and 3.64, with another.very strong band at 3.04p. 
Whereas the frequency relations supported the view earlier held that 
the 7u, 4.74, and 3.6u bands were respectively second, third, and 
fourth harmonics of a fundamental at 14, the intensity of the 4.7y 
band led Barker to question this interpretation and to suggest that 
very likely a new fundamental was present at approximately the same 
wave-length as the third harmonic. The 3.6” band is thus more 
likely to be a combination of the new fundamental at 4.7, with the 
lower frequency vibration. 

The band occurring at 3.04 is recognized as another fundamental. 
The bands at 14m and 7p are clearly of the doublet character. Mol- 
ecular moments of inertia are readily calculated from these data. From 
Burmeister’s curves for the 14 band, yielding an apparent separation 
of maxima of 37.5 cm.}, one calculates a moment of inertia of 
33 X 10° *° g. cm.? If, however, the relatively large slit-width at which 
this work was carried out is taken into account, estimates may be 
made as to the degree of overlapping, and more probable positions of 
the two components of the doublets may be plotted from the com- 
posite curve. On this basis a larger separation is obtained, of the 
order of 50 cm.'. The 7 band was investigated by Barker at suf- 
ficiently high resolution so that no such correction needs to be made. 





* Verh. Deutsche Phys. Ges., vol. 15, p. 589, 1913. 
* Phys. Rev., vol. 23, p. 200, 1924. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 85, No. 5 


2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


Because of some uncertainty as to the exact intensity values, how- 
ever, the form of the curve introduces some uncertainty in. the 
determination of the separation of the maxima. In view of this, and 
of the inadequate resolution of Burmeister’s apparatus, Barker’s cal- 
culation of 13.2x10°*° g. cm.” for the moment of inertia, based on 
the separation of 58 cm., is perhaps in as good agreement as could 
be expected with the corrected value of Burmeister. As the band 
occurring at 4.7m is certainly composite, no great significance can be 
attached to calculations of moments of inertia involving data on this 
band. 

Recently, R. M. Badger and J. L. Binder* have carried out a 
photographic investigation of the absorption spectrum in the near 
infra-red of hydrogen cyanide vapor in a 280 cm. absorption cell. 
They have observed two bands in this region occurring at A7QI2 
and A8563, respectively. In these bands they have been able to resolve 
the fine structure attributed to rotation in the molecules. On the 
basis of their measurements, they interpret each of the bands as com- 
posed of a P and & branch. From this rotational structure they are 
able to compute an accurate moment of inertia of 18.79 x 10°*° g. cm.? 
This calculation is in a reasonable agreement with the values based on 
doublet separation for the 14m and 7p bands. The absence of a Q 
branch is in harmony with the observations of a clearly doublet 
character of both the latter bands. In regard to the apparent central 
maxima in the bands occurring at 4.74 and 3.6u the question naturally 
arises as to the possible presence of a Q branch. It should be borne 
in mind, however, that these may readily be explained as due to 
overlapping. 

Assuming three fundamental frequencies corresponding to the 
bands at I4pu, 4.74, and 3.04», which have been designated respec- 
tively as 6, v2 and 1, Badger and Binder have interpreted the near 
infra-red bands as 3v,+v. for the band at A8563 and as 4v, for the 
band at A79g12. Because of the absence of a Q branch, they have 
assumed a linear arrangement of atoms, and on the basis of three 
fundamental frequencies, offered an interpretation of the three funda- 
mental modes of vibration corresponding to these fundamental fre- 
quencies. From an analysis of probable atomic distances of separation, 
they have come to the conclusion that the molecule must be hydrogen 
cyanide rather than hydrogen isonitrile (HNC). 


* Phys. Rev., vol. 37, p. 800, 1931. 








NO. 5 HYDROGEN CYANIDE—-BRACKETT AND LIDDEL 3 


EXPERIMENTAL RESULTS 


The results to be presented here were obtained with an automatic 
recording apparatus yielding high resolution and possessing certain 
novel features. An earlier self-recording instrument of high resolution 
was set up at the University of California by F. S. Brackett, yieid- 
ing on effective slit-width of 10 A. A similar instrument, but one 
with considerably greater aperture, was constructed by E. D. McAlis- 
ter at the University of Oregon, yielding an effective slit-width of 
6 A. The instrument used in the present investigation at the Fixed 
Nitrogen Laboratory is of approximately the same aperture, though 
of considerably greater focal length, and yields the same effective 
resolution. 






Prisms & Litfrow Mirror 


Collimating Mirror 


=a bes en rae TES sega 
Bie Sakae ei ee ee — = ri ource 


Fic. 1—Diagrammatic sketch of spectrograph. 


The instrument is of the Littrow form, wherein the light twice 
traverses two 60° prisms. The aperture is limited by the smaller of 
the two prisms, whose face is 20 cm. high and 15 cm. basal width. 
The instrument is used at a focal length of 2 meters. The body, 
a large casting, may be sufficiently evacuated to remove atmospheric 
absorption. Wave length variation is accomplished by rotation of 
the plane mirror, which is coupled with the motion of the photographic 
plate by a lever system. The use of a mechanical lever system with a 
variable pivot permits a wide range of variation of relative motion of 
the plate carriage to the angular rotation of the mirror, giving prac- 
tically any desired spread of spectrum. This improved mechanical 
system, together with the use of photographic plates instead of film 
or paper, gives a much greater reproducibility of spectrum and 
accuracy of wave length than heretofore obtained. The calibration 
was effected with mercury arc spectra and water vapor bands, the 


4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


observed wave lengths being consistent with grating measurements 
within +2 A. 

The thermocouple used is a modification of the type of single 
junction vacuum thermocouple described by Brackett and McAlister.’ 
The source of continuous radiation is a tungsten ribbon filament, 
using 16 amperes current at 6 volts and working at an approximate 
temperature of 2900° K. 


INVESTIGATION OF LIQUID HYDROGEN CYANIDE 


In the present investigation, the absorption spectrum of hydrogen 
cyanide in liquid phase has been studied with cell thicknesses of I mm., 
Icm., 5 cm., and 30 cm. 

The liquid hydrogen cyanide was obtained through the courtesy of 
W. B. Wood of the Plant Quarantine and Control Administration of 
the Department of Agriculture. This product had quite a perceptible 
odor of hydrogen sulfide. The original sample containing about 
1500 cc. was distilled over P,O;, primarily to remove any water, but 
a considerable quantity of sulfur was precipitated, as was expected 
from the presence of hydrogen sulfide. The second 500 cc. fraction 
was taken as an experimental sample. A drop of it did not affect 
lead acetate paper. This purification was made possible through the 
courtesy of Drs. G. E. Hilbert and L. B. Howard of the Fixed 
Nitrogen Research Laboratory. , 


aN H 
\ 
| 
<p aon ae Oe Oe ee aE SL 


Fig. 2.—Energy transmission curve showing absorption spectrum of 5 cm. 
cell of liquid HCN at low dispersion. Slit width approx. 40A. 


Figure 2 shows the absorption of a 5 cm. cell in the region from 
7uto 2u. The steadiness of the thermocouple will be apparent from the 
smoothness of the record. This illustration shows the instrument set 
for a relatively narrow spread, covering the entire region, and gives 
a general idea of the relative intensities. Actual observations of wave 


Rev: Sei. Instr} vol; 1, p: 181, 1930. 











NO. 5 HYDROGEN CYANIDE—BRACKETT AND LIDDEL 5 
lengths, however, were made mostly on a much wider spread, in- 
cluding simply the region from Ip to 2n. A typical plate at this spread 
of the same cell length is shown in figure 3. Figure 4 shows an 


! 1 : Ny — ~~ eee 


10 Ll Le i 14 WED, 16 Ly 18 


Fic, 3—Energy transmission curve showing absorption spectrum of 5 cm. 
cell of liquid HCN at high dispersion. Slit width approx. 9A. 


analysis of the bands in this region, the frequencies of the maxima 
being plotted against percentage absorption. Table 1 gives the sum- 
mary of the data obtained. The values of the fundamentals in vapor 
are inserted for comparison since no liquid values have been obtained 





SY+d, 


2),+20, 
t l 
5000 6000 7000 8000 9000 10000 11000 12000 cm‘ 


Fre, 4—Diagrammatic representation of absorption maxima observed with 
assignments of designations. Broken lines show vapor absorption. Fre- 
quency is plotted against per cent absorption. 





in that region. Intensity values, both as to percentage absorption and 
absorption coefficient are only approximate. Generally the estimated 
uncertainties in the frequency values indicated arise from the diffi- 
culty of setting upon broad absorption maxima like those shown in 
figure 3. Still less favorable are the conditions of the values for 41 
and 371+ 12, which were obtained on a low spread plate such as that 
shown in figure 1. In other cases the larger uncertainties indicated 
arise from the proximity of strong absorption bands. It will be seen 
that the observed values of Av lie well within the probable values to 
be expected, taking into account the normal variation to be expected 
in the successive differences, together with the probable uncertainty 
of measurement. The agreement certainly excludes any uncertainty 
as to identification. Not only do the wave lengths lead definitely to 


6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 85- 


the identification indicated, but the approximate intensities are con- 
sistent with such an interpretation. Of the entire 15 bands observed, 
only one has not been identified. This is almost immeasurably weak, 
and occurs in a position slightly displaced from the frequency where 
we should expect 3v,+6. This excellent agreement throughout leaves 
little doubt as to the correctness of the choice of fundamentals pro- 
posed by Badger and Binder. 


TABLE I1—HCWN Bands in Liquid 
Abs. 


Notation Abs. coeff. r v Ap 
% k A em.) cm.-1 
4” 6.6 O14 8000 12500 + 30 
3000 
3" 57. U7, 10527 9500 = 10 
3090 
24 96.0 64 15600 6410 = 10 
(1) vapor 3200 
217+ 22 * 2. -004 9500 10527 = 20 
2040 
211 + 2 47. 213 11787 8487 += 10 
2077 
2” 6410 + 10 
4v2 12. .024 12540 7974 = 10 
IQIO 
Bye (50) oe 16490 6064 + 10 
(2v2) 
(v2) vapor 2090 
2r, + 26 14. .020 12760 7837 + 10 
719 
21+ 6 8&8. 42 14050 71S == TO 
: 708 
2V4 6410 + 10 
(6) vapor 710 
3r1 + v2 207) 008 8650 TTSOI ==\25 
2061 
3r1 g500 = 10 
713 
31 — 6 10. .O17 11380 8787 = 10 
3r2 + 5 42. Ll 14690 6804 = I0 
740 
3¥2 6064 + 10 
2r2 + 26 (20) ays 17900 5587 +15 


vy + re 78. -30 19000 5203 22 15 

















NO. 5 HYDROGEN CYANIDE—BRACKETT AND LIDDEL 7 


INVESTIGATION OF VAPOR 


The absorption spectrum has been obtained of saturated vapor at 
22.5°C., with a 30 cm. length of cell. The three bands observed in 
the gas absorption, interpreted as 2v,, 3v1, and 2v;+ 2 all show clearly 
a doublet structure. In the stronger bands, 2v; and 3, shown in 
figure 5, separations of maxima are obtained of 47+2 cm. and 
50+2 cm.“, which yield moments of inertia 21+2x10-* g. cm.’ 
and 18+2x107*° g. cm.” respectively. This is consistent, within the 
order of the accuracy of the work, with the more accurate value ob- 
tained by Badger. The combination band is too weak to obtain 
separation values of significance. On the basis of this conclusive 
identification of fundamentals, the clearly doublet character of the 
gas absorption implying the absence of a Q branch, and the approxi- 


Oil —~— 


| 


| 
| 
1034 tre 1.1340 


ype 


1.5280 BO ST 





Fic. 5.—Energy curves showing absorption spectrum of 30 cm. Saturated 
HCN vapor. Slit width approx. 9A. 


mate values of moments of inertia, we had independently come to the 
same conclusions regarding the arrangement of atoms, the approxi- 
mate separations, and the probable modes of vibration before the 
publication of the work of Badger and Binder. 

The position of the 2y, and 37; bands, however, is not consistent 
with the formula 


Vn = 3333-70 — 43.70" 
Assuming the formula 
vV=Nwo — N-wo¥ 
our values indicate a variation in +. This is evident from table 2, 
where the values of Av. or 2w)x have successive values 59, 107, 


and 133, indicating values of wo, varying from 30 to 67, as against 
Badger’s value of 43.7 based only upon v; and 41,. Third differences 


8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


suggest that a constant value of )1 may be approached equal to or 
slightly greater than 67. On the basis of this value we may compute 
a heat of dissociation corresponding to an absolute electron voltage 
of 5.5 volts. This is in much better agreement with the value com- 


TABLE 2.—HCN Bands in Vapor 


Notation Abs. A v Ava I v(aver.) Av Av, yy va 
% Mh cm.-—! cm.-1 Tot? cms! cmt cm: cm.-1 cm,.-t 
(41) 12636 12500 + 30 136 
2901 


3n sSot2 18 9645 133 950010 145 


3124 
80 1.5280 6544 +2 
Qn A 2 OST 107. 6410+10 III 
75 1.5391 6497 +2 
3231 
(11) 3.04 3290 59 
3200 


bo 


1.1610 8613 £6 
2. + 44 10 8501 8487 =10 104 
1.1670 8569 +6 


bo 


2070 
(211) 6521 
(72) 4.7 2090 


puted from chemical data, 4.2 volts, than would be obtained from 
Badger’s constant value of 43.7. 

The displacements to lower frequencies in passing from vapor to 
liquid show a marked increase for the higher harmonics, with the 
exception of the 4v, value, for which it must be remembered that the 
liquid value is only approximate. 








SMITHSONIAN MISCELLANEOUS COLLECTIONS 
VOLUME 85, NUMBER 6 


MORPHOLOGY OF THE INSECT ABDOMEN 


PART J. GENERAL STRUCTURE OF THE ABDOMEN 
AND ITS APPENDAGES 


BY 
R. E. SNODGRASS 


Bureau of Entomology, 
U. S. Department of Agriculture 


qoneeeeseg 





(PuBLICATION 3124) 


CITY OF WASHINGTON 
PUBLISHED BY THE SMITHSONIAN INSTITUTION 
NOVEMBER 6, 1931 





Te Lord Baltimore Press 


BALTIMORE, MD., U. S. A. 


MORPHOLOGY OF THE INSECT ABDOMEN 


PART I. GENERAL STRUCTURE OF THE ABDOMEN AND ITS 


APPENDAGES 


By R. E. SNODGRASS 
BuREAU OF ENTOMOLOGY 
U. S. DEPARTMENT OF AGRICULTURE 


CONTENTS 


PAGE 
MTVETO@ CHUA CELO tlle sep eee epee tanta eee ie: < cas Meee dene renee tere ter eas, dakeke Susieus, sue ereie I 

lee he -abdominallsclenOtizatiome. sriecee iciec se sic leper cosiesertsreiss arenaicn Slouaueveus oasis 6 
Mipm ne mab comaitialieSeett ett aeeewtere irate cette <- c's lereter state ss « ies sielels otals cnusials 14 
Abi eaviscetal USCESTMENtSs aa suse ete e a .o.8 eo oecaie eieiale aielwlaie @ sctiefere a cls.cesvaiave 16 
iiveecenital mse otnenihs amentyem eres ated eerrtecrolcwet tore: o ersuebsrSoer ale cceloee 17 
Mhemposteenitall Seo ments aes «= nels eto yetie cic lee ap vsiensie ecte.0 ale 19 

ieee ire aba ominal emusctlatuce ers acai sessilis ts erete winters ear ceieis 28 
General plan of the abdominal musculature....................... 31 

The abdominal musculature of adult Pterygota.................4. 42 

The abdominal musculature of endopterygote larvae............... 48 

The abdominal musculature of Apterygota................ eee eee 56 
INereliies abdominals appemdac ese m.aebimer es vrais sierra ene seit se naire 62 
Bodyeappendacesmon a@hilopodasi sts arc ecta lee cus srnicerere sete ise cosine 5 
Abdominal appendaresvor | Grustacediws... sauce ae eases cies ee tere 68 

The abdominal appendages of Protura................eeeeeeecees 70 
General structure of the abdominal appendages of insects........... 71 

The abdominal appendages of Collembola................. 2.000. 2 

The abdominal appendages of Thysanura.................--00000 74 

The abdominal gills of ephemerid larvae..................0000005 77 
Lateral abdominal appendages of sialid and coleopterous larvae.... 70 

The abdominal legs of lepidopterous larvae..................... 83 

“WER ONO DOC Sema ees se eeeke ness rer eis od So ais SoSNERAG ust onal ues esos Gadbenevditeos 88 

PR ERCERCIE SCUITOWOUSI ie eeetrc srerhe enece ists ee ainieis acein vie ® eesteie te ciceterete is 2 

The terminal appendages of endopterygote larvae................. 06 
Terminal lobes of the paraprocts...0<52- leas sesso wise seecwe cess 107 
Morphology of the abdominal appendages................ 0020000 108 
Abbreviations used on the figures.............c.ceeccceeccccceuees 122 
Tere RGU Semon eee acon lant eee Seti Nag ar nals Govan ye dart ais kek 641d act seens 123 


INTRODUCTION 


The incision of the insect into head, thorax, and abdomen is in 
general more evident in the cervical region than at the thoracico- 
abdominal line ; but anatomically the insect is more profoundly divided 
between the thorax and the abdomen than it is between the head and 


SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 85, NO. 6 


2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


the thorax. The series of segmental appendages is continuous, and 
uniform except for minor adaptational modifications, from the head 
to the metathorax, inclusive ; on the abdomen the appendages abruptly 
cease, or are greatly altered in form. The musculature of the abdomen, 
consequently, is sharply differentiated from that of the thorax. Even 
the respiratory mechanism attests that the thoracico-abdominal bound- 
ary marks some deep-seated change in the body organization, since 
the closing apparatus of the spiracles is almost invariably different 
in one way or another before and behind the intersegmental fold 
separating the thorax from the abdomen. The definitive head contains 
three segments that have been, comparatively speaking, but recently 
transferred from the body to the cephalic region; the waist line of 
the insect has long been definitely established, and only in a single 
order has an abdominal segment been given over to the thorax. The 
abdomen is distinctly the visceral region of the body, and its major 
active functions in adult insects are those of respiration, copulation, 
and oviposition. 

Yet, notwithstanding the functional and structural differences that 
have come to separate the insect body into cephalothoracic and ab- 
dominal regions, we can not avoid the assumption that modern in- 
sects are derived from polypod ancestors, and that the abdominal 
segments at some time in the past history of the Hexapoda had the 
same essential structure as that of the primitive thoracic and gnathal 
segments. In studying the morphology of the abdomen and its ap- 
pendicular organs, therefore, we must attempt to find in the modern 
structure a basic plan of organization the same as that we are led to 
believe exists in the cephalic and thoracic regions from a’study of the 
segments, the appendages, and the musculature in the preabdominal 
parts of the body. Considering, then, the nature of the task, it is not 
surprising that students of insect morphology find in the abdomen 
problems far more difficult of solution than are those encountered in 
the head or thorax, and that there are many fundamental problems 
in the abdomen which are still unsolved. 

To the systematist in entomology the study of the abdomen, or 
particularly of the genital appendages, is becoming of ever increasing 
importance, and specialists are coming to feel acutely the need of a 
fundamental understanding of the organs that have been found in so 
many cases to give the best characters by which species may be dis- 
tinguished. Unfortunately, however, no investigator has yet discovered 
a means for determining with certainty the homologies of the organs 
most useful for diagnostic purposes. In truth, we may say that the 


NO. 6 INSECT ABDOMEN—SNODGRASS a 
morphology of the insect abdomen is a field in which no angel yet 
has trod, and is, therefore, one in which the fool unhindered may 
achieve his destiny. However, it is reputed that there is some merit 
in knowing oneself to be a fool, and, if it is the wisdom of the wise 
to enter only where the foolish have sprung the trap, the fool at least 
has a mission to perform. Hence, the writer offers the generalizations 
contained in this and a second paper to follow, not with the conviction 
that they will prove infallible, but in the hope that others will try 
to disprove them—and thereby establish their value. 

The principal conclusions derived from the study of the abdomen 
and its appendages to be given later in detail may be itemized in 
advance as follows: 

1. The insect abdomen consists primarily of 12 segments, the first 
11 of which are true somites, while the last is the periproct, or telson. 

2. The twelfth segment is practically obliterated in all the true 
Insecta, except for possible remnants in larval Odonata. 

3. The eleventh somite becomes the functional anal segment with 

- the suppression of the twelfth segment. Its tergum forms the epiproct. 
The venter of the eleventh segment is distinct in some Thysanura, 
but it is usually reduced or obliterated, except for two terminal lobes, 
which are the paraprocts. The appendages of the eleventh segment 
pie tie Cercl, 

4. The tenth segment is usually distinct in generalized insects, but 
it is often combined with the eleventh to form a composite terminal 
segment. The embryonic appendages of the tenth segment are sup- 
pressed in postembryonic stages of the more generalized insects ; they 
form the postpedes of holometabolous larvae, and perhaps the ap- 
pendicular processes of the proctiger, or tenth segment, of some adult 
Holometabola. 

5. The eighth and ninth somites are the genital segments, which 
bear the gonopods. The median gonopore of the female is located 
typically behind the eighth sternum, that of the male behind the ninth. 
Deviations from these positions are secondary; but the opening of 
the paired oviducts of Ephemerida between the seventh and eighth 
abdominal sterna probably represents a primitive condition, exhibited 
also by the embryos of certain insects (see Heymons, 1892, and 
Wheeler, 1893). 

6. The abdominal appendages of adult and larval insects are serially 
homologous with the legs and mouth parts. Each consists of a basis, 
and of one or two movable appendicular processes. The basis appears 
to comprise the coxal and subcoxal regions of a typical appendage, 
the two parts being either distinct or united. The coxal part often 


4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


bears an eversible or retractile sac having various functions, repre- 
sented by the vesiculae of Thysanura, the gill-bearing lobes of certain 
neuropterous larvae, and the plantar lobes of caterpillars and sawfly 
larvae. The basis of an abdominal limb usually takes the form of a 
lobe or plate of the body wall, and in the pregenital segments the 
limb bases are generally united with the sterna in adult insects. The 
appendicular process of the basis more commonly present is the 
stylus of the lower insects, or its derivatives, including the clasper- 
like organs borne by the male gonopods of the higher insects. The 
other limb process is the gonapophysis, which occurs only on the 
gonopods. Both the styli and the gonapophyses may be movable by 
muscles arising within the supporting basal lobes or plates, or on 
segmental areas derived from the latter. 

7. No positive evidence can be adduced from the known facts of 
anatomy or embryology to establish the homology of either the stylus 
or the gonapophysis. Many structural interrelations, however, sug- 
gest that the stylus is the telopodite of the appendage, and that the 
gonapophysis is an endite process of the basis. 

8. The genital appendages, or gonopods, have the same essential 
structure as the appendages of the pregenital segments. Their dis- 
tinguishing feature is the presence of the gonapophyses arising 
mesally from the bases. In the female, the gonapophyses of the two 
pairs of gonopods form the first and second pairs of valvulae of the 
ovipositor ; in the male the gonapophyses of the ninth segment be- 
come the parameres. The styli of the gonopods are usually suppressed 
in the female of pterygote insects; those of the ninth segment of 
the male form the movable claspers, or harpes, of the copulatory 
apparatus in the Endopterygota. 

g. The bases of the gonopods in adult female insects become plates 
supporting the first and second valvulae; those of the second genital 
segment may form a third pair of valvulae. In the male the bases of 
the single pair of gonopods often form distinct pleural plates of the 
ninth segmental wall between the tergum and the sternum, or they 
may fuse with either the tergum or the sternum, or with both; again 
they may unite with each other to form a plate either coalesced with 
the sternum or free and independently movable behind the latter. 

10. The parameres of the male are associated with the median 
penis in the lower insects, generally uniting with the latter except 
in Thysanura; but the penis may be suppressed, and the parameres 
then unite with each other+and inclose the terminal part of the 
ejaculatory duct to form the more complex copulatory organ known 
as the aedeagus. The parameres are to be identified throughout the 


no. 6 INSECT ABDOMEN—SNODGRASS 5 


orders by the muscles inserted on them, which take their origins in 
the supporting basal plates. 

11. The styli of the male gonopods become the movable claspers 
known as the harpes in the copulatory apparatus of holometabolous 
insects. They are to be identified by their muscles which arise in the 
supporting basal plates. The harpes may be divided each into a pair 
of movable claspers. 

12. Numerous accessory appendicular lobes and processes may be 
developed on all parts of the male genital segment and on segments 
associated with it. These organs are secondary and are not necessarily 
homologous in the several orders ; they are often flexible at their bases, 
but are to be distinguished from the true harpes and from the para- 
meres, with which they are associated, by their lack of muscles. 

13. The postpedes, present in holometabolous larvae of several 
orders, are the pygopods, or appendages of the tenth somite. The 
postpedes are probably transformed into the appendicular processes 
of adult males known as socii, found in adult Trichoptera and 
Lepidoptera, or into the cercus-like appendages of adult chalasto- 
gastrous Hymenoptera. 

14. The cerci are the uropods, or the appendages of the eleventh 
somite. Typically each is situated in a membranous area laterad otf the 
base of the epiproct, and above the paraproct. Muscles that move 
the cercus arise on the tenth tergum, or also on the epiproct, but these 
muscles are not necessarily primitive muscles of the cercal appendages. 
There is no intrinsic evidence that the cerci have any genetic relation 
with the paraprocts. It is doubtful if true cerci occur in any holo- 
metabolous insects, except possibly in females of Mecoptera. 

It will be evident from statements given above, and more flagrantly 
apparent in discussion to follow, that the writer still gives much 
importance to the value of muscles as determinants of skeletal homol- 
ogies—and this in the face of the edict against such practices recently 
put forth by H. J. Hansen (1930). However, there surely can be 
no question that in studying the insect skeleton we are dealing with 
the passive elements of mechanisms, in which the active parts are the 
muscles. The principal sclerotic areas of the body segments, and of 
the limb segments, are always directly or indirectly associated with 
muscle attachments or with mechanical strains resulting from muscle 
actions, and there is every reason for believing that scierites have 
been correlated with muscles in their evolution, if not necessarily in 
their origin. It is, of course, true that, just as some sclerites are 
secondary productions, so undoubtedly are some muscles. We must 
admit that all kinds of deviations from a rule are possible ; but a few 


6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 85 


exceptions do not discredit evidence supported by a long series of 
uniform interrelations between muscles and skeletal parts. However, 
a mere discussion of the matter is useless, and in a final analysis the 
identities of muscles must be established by a study of the muscle 
innervation. But, in the meantime, practical results may serve as a 
basis of judgment. The results of the studies to be given in the fol- 
lowing pages will appear principally in Part II of this paper, which 
will attempt to analyze the organs of oviposition, and especially the 
male organs of copulation, according to the light thrown on the homol- 
ogies of their parts by an examination of their musculature. The 
muscles furnish a means as yet but little used for identifying corre- 
sponding structures in the male genital apparatus of the various orders, 
and it will be found that they at least give something definite as a 
working basis in a comparative study of the genitalia. 

For most of the identified material on which the present paper is 
based the writer is indebted to specialists in the United States Na- 
tional Museum, including the entomologists of the Bureau of En- 
tomology, Department of Agriculture, and Dr. Waldo L. Schmitt 
and his associates, of the Museum’s Division of Marine Invertebrates. 
Specimens of Heterojapyx and Nesomachilis, however, were obtained 
through the interest of Dr. R. J. Tillyard of Australia. Furthermore, 
much valuable criticism and information has been contributed by 
Dr. A. G. Boving, Mr. Carl Heinrich, Dr. H. E. Ewing, and other 
Museum entomologists of the Bureau of Entomology. 


I. THE ABDOMINAL SCLEROTIZATION 


For purposes of morphological description it is necessary to dis- 
tinguish regions of the body wall from the sclerites that may partly 
or entirely occupy the regional areas. Considering the body as a whoie, 
there are two principal surface regions, one including the back and 
sides above the limb bases, the other the under surface between the 
limb bases. The first is the dorsum (fig. 1 A, D) ; the second is the 
venter (V). Then, in a metameric animal, each somite is likewise 
divided into a segmental dorsum and a segmental venter. Separating 
the dorsum and venter of each segment are the latero-ventral limb 
bases (LB, LB). The regions of the limb bases may be termed the 
pleural areas of the segments. The free distal part of any limb, mova- 
ble in a vertical plane on the basis, is the telopodite (Tlpd). 

It is now well understood that the hardened areas, or sclerites, of the 
body wall of insects, as well as of other arthropods, are not “ chitin- 
ized” or “strongly chitinized” regions, but that they are areas of 


0 ee ee 


No. 6 INSECT ABDOMEN—SNODGRASS 7 


the cuticula in which other substances than chitin predominate. It 
has been shown by Campbell (1929), for example, that the exocuticula 
of Periplaneta contains only about 22 per cent of chitin, while the 
soft endocuticula contains about 60 per cent; and according to 
Kunike (1926) the wing covers of a May beetle contain by weight 
75 per cent of nonchitinous substances, and those of a grasshopper 
as much as 80 per cent. The writer, therefore, follows the suggestions 
of Ferris and Chamberlin (1928) in designating the sclerites as areas 
of sclerotization rather than of “ chitinization.” 

Sclerotization of the body wall usually produces definite piates in 
the several segmental regions. According to the scheme of nomen- 
clature adopted in this paper, a major segmental plate of the dorsum 





Fic. 1—Diagrams illustrating the theoretical primitive structure of the ap- 
pendages and their relation to the body wall. 


A, cross section of a segment, showing the basis of each appendage (LB) 
movable antero-posteriorly on the body segment by an axis (a-b) in a vertical 
plane, and the telopodite (7/pd) movable dorso-ventrally at the coxo-trochan- 
teral joint (ct). 

B, the basis of each appendage subdivided into a subcoxa (Scx) and coxa 
(Cx) by a secondary joint with a vertical axis (c-d); the upper part of the 
subcoxa forming the “pleuron” of the body segment. 


is a tergum (fig. 1B, T), a corresponding plate of the venter is a 
sternum (Stn), and a single plate or group of plates in the pleural 
region is a pleuron (Pl). Subdivisions or component elements of 
these principal segmental plates then become tergites, sternites, and 
pleurites, respectively, since the suffix ite grammatically can mean 
only “a part of ” some larger unit designated by the stem of the word 
to which it is appended. 

The plan for distinguishing and naming the segmental regions, the 
major sclerites, and the subdivisions of major sclerites given above 
is not generally followed; but the writer has not found any nomen- 
clatural scheme for these parts that is consistently applied, or that 
adequately meets the situation. Some writers define the terms “ terga” 
and “‘sterna”’ as the segmental dorsal and ventral regions, and then 
designate the principal plates of these regions the “ tergites”’ and 


8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 85 


“sternites.”’ This usage is without other objections than that it leaves 
us no specific names for the subdivisions or component minor sclerites 
of major areas of sclerotization. Writers that adopt it seldom follow 
it consistently. On the other hand, many entomologists find it con- 
venient for descriptive purposes to distinguish the segmental plates of 
the abdomen from those of the thorax as “ tergites ”’ and “ sternites.” 
The use of these terms in this manner, however, is not only arbitrary, 
but it is ungrammatical, since it is clear that the terms ending in 
ite may be applied to metameric units of any particular group of 
somites only if the names ‘‘ tergum” and “sternum” are defined as 
the entire dorsum and entire venter of this limited group of somites— 
a usage which no one pretends to adopt. In general, nomenclatural 
troubles arise not so much from a scarcity of suitable terms as from 
a lack of consistent application of words in common use. The writer, 
therefore, employs the terminology recommended above as the one 
most adaptable to the needs of anatomical description. And yet, it will 
be found that absolute consistency is not possible ; the insects are sure 
to present some anatomical conditions that can not be made to fit 
with any nomenclatural scheme that can be devised. Consistency 
is said to be a jewel, but an excess of jewelry may become a burden. 

Little is known concerning the nature of the sclerotic substances 
in the cuticula of insects, or of the procedure by which a specific area 
of the body wall becomes continuously sclerotized. We may believe, 
however, that minor sclerites may have been produced phylogenetic- 
ally by the secondary subdivision of major sclerites, though in the 
ontogeny of the individual they may proceed from separate centers 
of sclerotization. On the other hand, it is unquestionably true that 
primarily distinct areas of sclerotization may unite, and give no trace 
of their independent origin in the development of the embryo or pupa. 
In the abdomen of most adult insects, for example, the pleural 
sclerotizations derived from the limb bases are fused with the primi- 
tive sterna, and each definitive “ sternal ” plate in such cases is a triple 
structure, though it may lose all trace of its composite origin. 

The spiracles are important landmarks in the study of the abdom- 
inal segments. They never exceed eight pairs in number, and while 
one or more of the posterior pairs may be absent, the presence of a 
pair of spiracles is often better proof of the site of a primitive seg- 
ment than is evidence derived from the sclerites. The primary posi- 
tion of the spiracles is a matter on which opinion differs. There can 
be no doubt that the spiracles are subject to migration, and that in 
certain insects they have undergone an extreme displacement; but 
in the more generalized segments of most insects the spiracles lie in 


No. 6 INSECT ABDOMEN—SNODGRASS 9 


the sides of the segments above the regions of the limb bases, and 
therefore in the lateral parts of the dorsum (figs. 1, 2A, Sp). 

the tergal sclerotization of a segment does not invade the spiracular 
areas, the spiracles commonly lie in membranous lower parts of the 
dorsum between the tergum and the limb bases (fig. 2 A), or between 
the tergum and the definitive sternum, which has absorbed the limb 
bases (B). The tergal plates of the abdomen, however, often extend 


Stn 






Fic. 2—Diagrams showing the sclerotization and the retractor mechanism of 
the abdomen. 


A-E, variations of sclerotization above and below the dorso-pleural line (a-a) 
separating the dorsum from the region of the limb bases (LB). F, the retractor 
mechanism as seen in vertical section, resulting from secondary segmentation 
produced by the usual type of sclerotization in adult insects. G, two consecutive 
segmental plates and their connecting muscles. 

a-a, dorso-pleural fold; Ac, antecosta; acs, antecostal suture; DMcl, dorsal 
longitudinal muscles; /sg, primary intersegmental fold; LB, limb basis; Mb, 
secondary intersegmental membrane; MWcl, “longitudinal muscle; patg, parater- 
gite; Pc, precosta; PJ, pleural plate formed of the limb basis ; cg definitive ster- 
num including areas of limb bases; Sp, spiracle; Stn, primary sternal plate; 
T, tergum. 


so far downward on the sides of the dorsum as to include the spiracles 
in their lateral parts (C). In some cases the spiracles occur in inde- 
pendent lateral, or paratergal, sclerites of the dorsum (D). Finally, 
the definitive ventral sclerotization is sometimes produced upward 
on the sides of the abdominal segments, and the spiracles may then 
be included in the lateral parts of the sternal plates (FE) ; but in such 
cases it is to be suspected that the apparently sternal areas containing 


IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


the spiracles are really paratergal sclerotizations that have secondarily 
united with the sterna. 

In the larvae of holometabolous insects there is usually a more 
or less distinct groove extending along each side of the abdomen 
below the line of the spiracles (fig. 3 A, B, C, a-a), which is continued 
upon the thoracic region above the regions of the pleural, or subcoxal, 
sclerites, when the latter are present (B, C, Scx;). This groove, 
therefore, evidently marks the division between the dorsal and pleural 
areas of the abdominal segments, and may be termed the dorso- 





Fic. 3—External structure of the abdomen in holometabolous larvae. 


A, larva of Carpocapsa pomonella, showing the dorso-pleural fold (a-a) sepa- 
rating the dorsum from the subcoxal areas in both the thorax and the abdomen. 
B, larva of Silpha obscura, with series of paratergal sclerites (patg) above 
dorso-pleural fold, and series of pleural subcoxal sclerites (Scx) below fold. 
C, larva of Pteronidea ribesti, showing same structure as in B. 


pleural groove. Sclerites or lobes of the body wall lying immediately 
above this groove, then, belong to the dorsum, and may be called 
paratergites (fig. 3 B, patg), but not “ pleurites”’ or “ epipleurites.” 
Latero-ventral plates or lobes lying below the tergo-pleural groove 
are properly termed abdominal pleura (B, Scx) if it is clear by their 
position or by the presence of appendicular organs upon them that 
they are the equivalents of the thoracic subcoxal pleura (Scx;). It 
is convenient, however, to extend the term “ pleurites ” to any lateral 
sclerites situated ventrad of the dorso-pleural groove, or below the 
line of the spiracles, though such sclerites may be evidently secondary 
sclerotizations of this region (fig. 24 D, rpl, 2pl, etc.). But if there is 





No. 6 INSECT ABDOMEN—SNODGRASS TT 


reason to believe that lateral sclerites are secondary subdivisions of 
earlier formed pleurosternal plates, they may be given the non- 
committal name of parasternites. When true pleural plates or lobes 
of the abdomen are subdivided longitudinally, the upper and lower 
parts may appropriately be termed epipleurites and hypopleurites, 
respectively ; but such a division seldom occurs in the abdominal 
pleura, and the term “ epipleurite ” is commonly misapplied by students 
of insect larvae to paratergal lobes, or sclerites of the dorsum. 
The fact that the dorso-pleural groove forms a conspicuous line 
of infolding along the side of the abdomen in many insect larvae 
(fig. 3, a-a) is probably the reason for its frequently having been 
termed the “ pleural suture.” Hopkins (1909) thus named it in his 
study of the larva of Dendroctonus, and he designated the iateral 
lobes above the groove “ epipleural ” and those below it “‘ hypopleural.” 
The former he believed represented the epimeron of a thoracic 
pleuron, and the latter the episternum. No such homology as this, 
however, is possible, since the pleural suture of a thoracic segment 
is morphologically a vertical groove in the subcoxal sclerotization cf 
the leg bases, taking only secondarily a horizontal position in the meta- 
thorax of adult beetles. The so-called ‘‘ pleural suture ”’ of the larval 
abdomen, moreover, as we have seen (fig. 3 A, B, C, a-a), extends 
into the thorax above the subcoxal sclerotizations (Sc), and thus 
throughout the body separates the dorsum from the true pleural 
region. Lateral lobes or sclerites of the abdomen lying above the 
dorso-pleural groove are, therefore, paratergal (fig. 3 B, patg), and 
not “epipleural.’”’ Only the so-called “hypopleural”’ areas lying 
ventrad of the dorso-pleural groove, that is, between it and the true 
sternal region, are properly pleural in the sense that they correspond 
with the subcoxal areas of the thorax (B, Scx;) containing the scler- 
ites of the thoracic pleura. The ventro-lateral lobes or plates of the 
larval abdomen may, then, be termed the abdominal pleura inasmuch 
as they appear to represent the subcoxae of the thorax. The abdominal 
pleura are never divided vertically in a way to suggest a true homology 
with the division of a thoracic pleuron into episternum and epimeron. 
The relation of the muscles to the lateral lobes of the abdomen 
in the larvae of Coleoptera has been studied by Boving (1914) and 
by Craighead (1916). Boving, here following Hopkins (1909), calls 
the lateral groove of the abdomen the “ pleural suture,” but in all his 
subsequent work he terms it the “ ventro-lateral suture.” Craighead 
identifies the lateral areas of the abdomen with the corresponding 


’ 


areas of the thorax in cerambycid larvae, but since he regards the 


12 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 
jateral groove of the abdomen as the “ sternopleural suture,” he 
takes the paratergal areas to be the abdominal pleura. 

The abdominal terga.—The dorsal sclerotizations of the abdomen 
in general take the form of simple tergal plates characteristic of any 
region of the body in which a secondary segmentation has been 
established. Each tergum presents anteriorly a submarginal or often 
marginal internal ridge, the antecosta (fig. 2F, G, Ac), on which 
the principal longitudinal muscles usually have their attachments. 
The antecostal suture (F, acs) is generally but faintly marked, and 
the precosta (F, G, Pc) varies from a scarcely perceptible marginal 
rim to a wide flange extending a considerable distance anterior to the 
muscle attachments (G, Pc). Apodemal processes are frequently ex- 
tended from the anterior margins of the terga. From the antecosta 
of the first abdominal tergum there is commonly developed a pair of 
phragmatal lobes, and the precostal part of this tergum, together 
with the antecosta and the phragma, may be separated from the rest 
of the tergal plate to form a so-called postnotal or postscutellar plate 
of the metathorax. Otherwise the abdominal terga usually preserve 
their structural unity. The postcostal areas of the abdominal terga are 
seldom marked by sutural lines in adult or nymphal insects, and where 
such lines do occur they can not be supposed to have any homology 
with the sutures of a wing-bearing thoracic tergum, which adapt the 
latter to its function in the wing mechanism. 

The dorsal regions of the abdominal segments of soft-bodied holo- 
metabolous larvae are usually divided transversely by impressed lines 
or by strongly pronounced topographical features. The dorsal areas 
thus formed are evidently mere adaptations to the contractile move- 
ments of the larvae and have no morphological significance. That 
the external body features of eruciform and vermiform larvae are 
secondary larval characters is evident from the structure of the head, 
which shows that such larvae are lateral derivatives from highly 
evolved adult forms representing the immediate ancestors of the 
order. 

The abdominal sterna.—The definitive sternal plates of the abdomen 
are in general similar to the tergal plates, each being a continuously 
sclerotized area of the ventral integument of its segment, always 
including the primary intersegmental area anterior to the somite, 
corresponding with the intersternites, or spinisternites, of the thoracic 
region. The antecostae may be coincident with the anterior margins 
of the sternal plates, or set well back from the margins (fig. 2 F, Ac) 
with distinct precostal regions before them. In the Japygidae a short 
anterior division of each abdominal sternum is separated by a mem- 





NO. 6 INSECT ABDOMEN—SNODGRASS 13 


branous fold from the rest of the plate, forming a distinct sternal 
apotome (fig. 24 D, Apt). Apodemal processes to give more effective 
action to protractor and dilator muscles are commonly developed from 
the anterior and lateral margins of the abdominal sterna. 
Notwithstanding the apparent unity of structure in the abdominal 
sterna, it is probable, as already stated, that the ventral plates of the 
pregenital segments in most adult insects are triple structures, each 
including in its composition the area of the true sternal sclerotization 
of its segment, and the areas of the limb bases of the corresponding 





Fic. 4—Ventral plates of abdomen of Nesomachilis maoricus. 


I, II, VI, VIII, ventral plates of segments one, two, six, and eight in male. 
LB, limb basis, or basal plate of appendage: rvs, muscles of retractile vesicle; 
Stn, primitive sternal sclerite; Sty, stylus; ]"s, eversible and retractile vesicle. 


segmental appendages. A comparatively generalized condition ‘is to be 
seen in larvae of Ephemerida, in which lateral lobes of the abdomen 
supporting the gills (fig. 34 A, B, LB), though forming a part of the 
ventral wall of each segment, are distinct from the areas of the primary 
sterna (Stn), and occupy the primitive position of limb bases on the 
sides of the segments between the tergal and sternal sclerotizations. 
A similar but less primitive condition is that occurring in some of the 
Thysanura, as in the Machilidae (fig. 4), where each of the definitive 
sterna in the pregenital region of the abdomen consists of a small, 
median, true sternal sclerite (Stn) and of two, large, lateral stylus- 
bearing plates (LB, LB) clearly representing the limb bases. 


14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


The frequent occurrence of styli on abdominal segments of insects 
in which the sterna are undivided plates leaves little doubt that the 
definitive abdominal sterna of insects in general are composite plates 
including the limb bases as integral parts of their areas. Evidence 
of the inclusion of the limb bases in the adult abdominal sterna might 
be derived also from other sources, as in the Lepidoptera, where 
the abdominal appendages of the larva at the time of pupation are 
flattened out in the form of discs, and merge into the ventral areas 
from which later the adult sterna are produced. 

A definitive sternal plate that includes the primitive sternum and 
the bases of the adjacent limbs is sometimes called a “ coxosternum,” 
but, as will be shown later, there is a question as to whether the 
abdominal limb bases represent the coxae or the subcoxae, or include 
both of these usual basal elements of the appendages. A composite 
sternal plate, therefore, is more appropriately distinguished from a 
primitive sternal plate by the term sygosternum proposed by Prell 
(1913). For the same reason the name “ coxite,’ often given to the 
limb base element of the zygosternum, is objectionable as being more 
specific in its meaning than is warranted by the known facts of the 
origin of the part in question. Besides this, the suffix ite implies that 
a structure so-named is “a part of” a coxa, and this implication 1s 
clearly not intended. 


II. THE ABDOMINAL SEGMENTS 


Entomologists sometimes nominally distinguish the segments of 
the insect abdomen frdm those of the thorax as “ urites,’ a term 
perhaps recommended by its brevity, but one which, by inference, 
reduces the entire abdomen to the status of a “ tail.”’, Consistent with 
this usage, the abdominal appendages would all be “ uropods,” but 
the custom of carcinologists in applying the latter term only to the 
terminal pair of appendages has better anatomical sanction. (Lan- 
kester, 1909; Sedgwick, 1909. ) 

From embryological evidence there appears to be little doubt that 
the primitive number of abdominal segments in typical insects is at 
least 12 (fig. 5 A). Twelve segments are actually present in adult 
Protura (B), each having distinct tergal and sternal plates, but the 
tenth and eleventh are said to be added by “ epigenesis,” that is, they 
are developed during postembryonic growth. In many of the Aptery- 
gota and in the lower Pterygota, 11 segments are present without 
question, while in some forms there are possible rudiments of a 
‘twelfth segment. The twelfth or primitive terminal segment is the 


No. 6 INSECT ABDOMEN—SNODGRASS 15 


periproct (fig. 5 A, Prpt), which carries the anus, but does not have 
appendages. It is the terminal piece of the body anterior to which 
the true somites are formed, segmentation of arthropods being intra- 
somatic and not a process of budding. Some investigators claim that 
a pair of coelomic sacs is formed in the periproct. The presence of not 
more than six segments in the abdomen of Collembola is usually 
taken to be the result of reductive specialization, or “ degeneration,” 
in these insects, considering that the existence of even six segments 
is obscure in some forms by a loss of intersegmental lines. Tillyard 
(1930), however, sees in the Collembola a primitive “ protomorphic ” 
condition in which only nine postcephalic somites have ever been de- 
veloped. He points out that segmentation in the collembolan embryo 
produces six abdominal segments and no more, and that we have, 


fC aE ea IXApd ae 







fe XApd 


5 XApd 
(Cer) 





~Prpt 
XI ae 
ee ari xis B 

Fic. 5.—Examples of the presence of twelve segments in the hexapod abdomen. 


A, posterior end of abdomen of young embryo of Gryllotalpa (from Heymons, 
1895), with large periproct (Prpt), or twelfth segment, behind the last true 
somite (X/) bearing the cerci (Cer). B, end of abdomen of adult proturon, 
Acerentulus confinis (from Berlese, 1910), showing twelve distinct segments. 


therefore, no evidence of a greater number of somites ever having 
been present in this group of arthropods, which he would not ally 
closely with the other insects. A reduction in the number of abdominal 
segments is the rule in both immature and adult insects generally. 
As just stated, evident remnants of the periproct are rare except in 
the Protura. While 11 segments are distinct in many of the more 
generalized insects, in the higher orders, especially in the Holome- 
tabola, not more than 10 segments are usually present, and sometimes 
only nine are evident. In the more specialized insects there is a 
tendency toward elimination of the first abdominal segment, but 
generally reduction takes place at the posterior end of the body. 
Since the periproct is commonly lacking, or reduced to a circumanal 
membrane, the eleventh somite, which carries the last pair of seg- 
mental appendages, becomes the definitive anal segment. The tenth 


16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


segment is sometimes more or less united with the eleventh in orthop- 
teroid insects; but in the Holometabola the eleventh segment 1s 
generally suppressed, and the body ends with the tenth segment, 
though possibly in certain holometabolous larvae the terminal segment 
contains a remnant of the eleventh somite, while in coleopterous larvae 
the abdomen ends in a distinct anal lobe, which appears to represent 
the eleventh segment. 

The association of the organs of copulation and oviposition with 
the eighth and ninth segments of the abdomen is usually accompanied 
by adaptive structural modifications in these segments that conspicu- 
ously differentiate the latter as the genital segments, and separate the 
rest of the abdomen into a pregenital region and a postgenital region. 
Since the pregenital region contains most of the internal abdominal 
organs, its segments may be termed the visceral segments. The seg- 
ments beyond the ninth, which are usually more or less reduced and 
united with each other, constitute the postgenital segments. It is not 
possible, of course, in all cases to divide the abdomen consistently into 
visceral and genital regions, since modifications adaptive to the major 
functions of the eighth and ninth segments often affect one or several 
of the preceding segments, but yet, for general descriptive purposes, 
the term “ genital segments ” will have a specific meaning. 


THE VISCERAL SEGMENTS 


To describe here in full the visceral region of the abdomen would 
be to repeat many well known facts without adding anything of 1m- 
portance. The seven segments of this region are usually of simple 
structure and differ but little from one another. In adult pterygote 
insects they lack appendicular organs, and the definitive sterna prob- 
ably always include the areas of the primitive limb bases. The first 
segment is more subject to modifications than are any of the others. 
In winged insects the antecosta of its tergal plate bears the third pair 
of phragmatal lobes, and the precosta is usually much enlarged, form- 
ing the so-called postnotal, or postscutellar, plate of the metathorax 
(fig. 16, PN;), which, together with the base of the phragma, is 
frequently removed from the tergal region of the first abdominal seg- 
ment and more closely associated with that of the metathorax. The 
rest of the first segment is often reduced, or fused with the second, 
and the sternal sclerotization is sometimes obliterated. The first pair 
of spiracles, however, are nearly always retained, and the spiracles 
will generally furnish a key to the basal segmentation of the abdo- 
men where the segmental limits are obscured. In the aculeate Hy- 





No. 6 INSECT ABDOMEN—SNODGRASS 17 
menoptera the entire first abdominal segment is so intimately united 
with the metathorax that it forms anatomically a part of the thoracic 
region of the body. Modification of the posterior visceral segments 
will be noted in connection with a study of the genitalia. In females 
of higher Diptera the functional visceral region is reduced to five 
segments by the conversion of the posterior segments into a tubular 
organ of oviposition. 


THE GENITAL SEGMENTS 


The lateral ducts of the reproductive organs in the majority of 
the Hexapoda open into a common, median outlet tube. [Exceptions 
to this rule occur in the males of Protura, in both males and females 
of Ephemerida, and in males of some Dermaptera, the two lateral 
ducts in these cases opening separately to the exterior. The position 
of the genital apertures varies within the Hexapoda through an 
extreme of six segments. In the Collembola the gonopores of botit 
sexes are situated on the fifth abdominal segment, while in the Pro- 
tura they occur between the eleventh and twelfth segments. In the 
Ephemerida the paired oviducts open between the seventh and eighth 
abdominal segments, and the vasa deferentia open on the penes be- 
tween the ninth and the tenth segments. In all other Pterygota, except 
Dermaptera, and in Thysanura and Dicellura, the single female aper- 
ture lies between the eighth and ninth segments, and the male aperture 
between the ninth and tenth segments. Apparent exceptions to this 
rule occur where some of the terminal segments are fused, where 
one or more of the pregenital segments have been obliterated, or 
where, as in female Lepidoptera, the gonopore has evidently under- 
gone a secondary change in position. 

The genital apertures are described by some writers as being situated 
on the segments, while others state that they occur between segments. 
The gonopores, in truth, are probably located on the posterior parts of 
the ventral surfaces of primary segmental areas, but since these parts 
of the primitive somites become the intersegmental membranes of the 
definitive segments, the gonopores of adult insects are anatomically 
intersegmental. They lie behind the primary sterna of the segments 
on which they are situated, and only rarely is a secondary sclerotization 
formed behind them (male Odonata). The male gonopore is usually 
carried outward on an evagination of the body wall forming a simple 
penis, or it is situated on a more complex copulatory organ composed 
of the penis and the parameres, or of the parameres alone, known 
as the aedeagus. 


18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


Owing to the uniformity in the relation of the genital apertures to 
the eighth and ninth abdominal segments, these segments in the 
majority of insects become specifically the genital segments. Their 
appendages form the principal organs of egg-laying and copulation, 
and may therefore be designated gonopods. In some of the simpler 
insects the gonopods are lacking and the genital segments have no 
distinctive external features; but usually the segments show some 
conspicuous structural adaptation to the functions of copulation or 
oviposition. 

The eighth segment——Modifications of the eighth segment (fig. 6, 
VIII) occur principally in the female, since it is on the ventral part 


VII VIII oe x XM Cer MiiGgey) 
ppt 





Fic. 6—Diagram illustrating the concept of the structure of the abdomen 
adopted in this paper. 


a-a, dorso-pleural line separating tergal region from pleuro-sternal region; 
Cer, cercus; Eppt, epiproct; 1Gon, 2Gon, first and second gonapophyses; /a, 
lamina subanalis; LB, limb basis; Papt, paraproct (lobe of eleventh sternum) ; 
sa, lamina supra-analis; Sp, spiracle; Stn, primary sternum; Sty, stylus; T, 
tergum; Tcl, telson (twelfth segment, greatly reduced or obliterated in insects). 


of this segment that the first gonapophyses (1Gon), or genital proc- 
esses of the eighth gonopods, are developed, and become the ventral 
blades, or first valvulae, of the ovipositor in all species provided with 
an ovipositor. The female genital opening is normally situated be- 
tween the bases of the first gonapophyses in the membrane behind 
the primitive eighth sternal plate, but the latter is frequently prolonged 
beneath the base of the ovipositor, forming the subgenital plate of the 
female. The bases of the gonopods of the eighth segment are never 
united with the eighth sternum in female insects having an ovipositor. 
In the Thysanura they are large, stylus-bearing plates or lobes which 
retain the normal position of limb bases, but in pterygote insects they 


No. 6 INSECT ABDOMEN 





SNODGRASS 19 
appear to form small suspensional sclerites of the first valvulae, known 
as the valvifers, which always lack styli. Gonapophyses of the eighth 
segment are known to be present in male insects only in some species 
of Machilis, but the eighth segment of the male is frequently more or 
less modified when associated with the ninth in the copulatory 
mechanism. 

The ninth segment.—The second genital segment (fig. 6, 7X) usu- 
ally has less of the typical form than does the first. It is the somite of 
the second gonapophyses (2Gon), or genital processes of the ninth 
gonopods, which form the second valvulae of the ovipositor in the 
female, and the usual parameres in the male. The sternum of the 
ninth segment is generally reduced or rudimentary in the female, but 
the bases of the gonopods are commonly retained, either in the form 
of lobes, or as blade-like pieces of the ovipositor, the third valvulae. 
In the male the ninth segment retains a generalized structure in the 
Thysanura (fig. 33, B, C), but in the pterygote insects it is subject 
to many modifications and takes on a great variety of forms. The 
bases of the gonopods in male Pterygota either remain as distinct 
lobes of the segment, or they unite with each other, or with the 
sternum, or with both the sternum and the tergum. The styli, if re- 
tained, usually take the form of movable clasping organs. Various im- 
movable lobes also may be developed from the ninth segment of the 
male, and sometimes from the eighth, which serve as accessory organs 
in copulation. 

The intromittent organ of the male arises in the Thysanura behind 
the region of the ninth sternum and between the bases of the gono- 
pods (fig. 33 B, C, Pen) ; but in insects having the gonopod bases 
united with the sternum, it arises posterior to, or usually above, the 
limb base area of the composite sternum. The membranous area 
from which the organ arises is, furthermore, generally more or less 
inflected between the ninth and tenth segments, forming a genital 
chamber above the ninth sternum, and the ninth sternum is often 
extended posteriorly as the male subgenital plate, or hypandrium. 
The intromittent organ has the form of a simple, tubular penis in the 
Thysanura, but in most pterygote insects it is a more complex struc- 
ture, called the aedeagus, formed of the penis and parameres, or 
of the parameres alone. The external genitalia will be fully described 
in Part II of the present paper. 


THE POSTGENITAL SEGMENTS 


Beyond the second genital segment there are never more than three 
segments represented in the hexapod abdomen (fig. 6, X, XJ, X//), 


2 


20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


and it is only in the Protura that the last of these segments is well 
developed (fig. 5 B, X//). In some of the lower insects apparent 
traces of the terminal segment are to be found, but in most of the 
true Insecta there are only two postgenital segments present. A still 
further reduction, however, has taken place in many insects, with the 
result that but one segment is to be recognized beyond the ninth. 
Where two postgenital segments are present there is little doubt that 
they are the tenth and the eleventh; and where the number of post- 
genital segments is reduced to one, it is usually to be assumed that the 





Fic. 7—Terminal abdominal structures of Thysanura. 


A, end of abdomen of male Nesomachilis maoricus. B, caudal filament and 
cercus of same removed, showing their origin from small eleventh segment nor- 
mally retracted into the tenth. C, dorsal view of same. D, ventral view of 
eleventh segment, with bases of caudal filament and cerci. E, dorsal view of 
eleventh segment and terminal appendages of Thermobia. F, ventral view of 
same. 

An, anus; Cer, cercus; cf, caudal filament; Eppt, epiproct; LB, limb basis; 
Papt, paraproct; sa, possible rudiment of lamina supra-analis; Stn, primary 
sternum; Sty, stylus. 


single end-segment is the tenth somite, and that the eleventh has dis- 
appeared, though it is probable that the tenth and the eleventh somites 
are in some cases combined in the definitive terminal segment. 

The tenth segment.—The tenth somite of the abdomen is clearly 
retained as the numerical tenth segment in insects in which there are 
two distinct postgenital segments. It is a complete annulus in the 
Thysanura (fig. 7A, X), quite distinct from the eleventh segment 
(B, X/), which is mostly concealed within it. In the Odonata the 
tenth segment is a continuously sclerotized ring (fig. 12B, C, X), 


NO. 6 INSECT ABDOMEN—SNODGRASS Zi 


beyond which are the parts of the eleventh segment, and apparent 
remnants of the twelfth (A). Likewise, in larvae of Plecoptera the 
tenth segment 1s cylindrical in form (fig. 8 A, X), and the parts of the 
eleventh segment (Eppt, Papt) are quite distinct from it. The struc- 
ture in an adult plecopteron is essentially the same as that of the 
larva, but the tenth segment is smaller. In the Ephemerida, both 
larval and adult forms, the tenth somite is a well-developed segment, 
which, because of the reduction of the eleventh somite, forms the 
terminal segment of the body and appears to carry the long cerci. 
Its tergal region is produced posteriorly in a median lobe, and thus 
resembles the supra-anal plate of other insects, but the small true 
epiproct of the eleventh segment lies beneath the lobe of the tenth 
tergum and carries the median caudal filament. The venter of the 
ephemerid tenth segment appears to contain the anal opening, but 
it is evident that the anus is drawn forward and that the paraprocts are 
united with the bases of the cerci. 

Among orthopteroid insects the tenth segment is variable ; its ven- 
tral region is usually membranous, and in some families its tergal plate 
is fused with the eleventh tergum, or epiproct. In the Phasmidae, 
however, the tenth segment is large and normally developed (fig. 
8 E,G, X). In Diapheromera it has distinct tergal and sternal plates, 
the tergum overlapping the edges of the sternum in the female (F), 
though the two plates are ankylosed in the male to form a strong 
support for the clasper-like cerci (2). The paraprocts are united 
ventrally with the tenth sternum (F, Papt) and appear to be lobes of 
the latter. The ventral region of the tenth segment 1s membranous 
in most other Orthoptera (fig. 8 D), though the dorsum usually 
contains a distinct plate (V7). In Acrididae the tenth tergum is a 
narrow transverse sclerite fused laterally with the ninth tergum, 
but it is separated from the epiproct by a complete suture. In Blat- 
tidae (fig. 40 A), Tettigoniidae, and Gryllidae (fig. 8 B), however, 
the tergum of the tenth segment (X), is generally fused with the 
epiproct (Eppt), and since the paraprocts become continuous with 
the membranous ventral wall of the segment, the tenth somite in 
these families loses the status of an independent body segment. 

It must be noted here that the principal muscles of the cerci arise 
on the tergum of the tenth segment. The size of the tenth segment, 
therefore, generally varies according to the size of the cercal muscles, 
the segment being large in insects having long, mobile cerci (fig. 8 A, 
XS), and strongly developed in insects in which the cerci have 
a grasping function, as in Japyx (fig. 40 C),and in Diapheromera (fig. 
8E, X). When the cercal musculature is weak the tenth segment 


22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


is usually reduced, and its tergal plate becomes small, or unites with 
the epiproct. In certain cases, however, the tenth segment is developed 
quite independently of any relation to the cerci, as in some of the 
Homoptera, in which the cerci are rudimentary or absent. In the 
cicada (fig. 8 C) the tenth tergum is a strong plate produced down- 
ward on the sides into a pair of hooked lobes (/) embracing the distal 
end of the aedeagus. 


XT Cer Eppt 
4 eae 


2 ae 







~ 
Cc WUIS= 
Fic. 8—Terminal abdominal structures of various insects. 


A, larva of plecopteron, ventral view, showing cerci, paraprocts, and epiproct 
as parts of eleventh segment. B, Gryllus assimilis, dorsal view, showing union 
of epiproct with tenth tergum. C, Magicicada septendecim, with eleventh seg- 
ment distinct from tenth. D, Scapteriscus didactylus, female. E, Diapheromera 
femorata, male. F, female of same, showing paraprocts fused with tenth ster- 
num. G, lateral view of genital and postgenital segments of female Diapheromera 
femorata, showing ovipositor, and subgenital plate produced from eighth ster- 
num. H, end of abdomen of female Panorpa consuctudinis. 


True appendages are absent from the tenth segment in postem- 
bryonic stages of all Apterygota and hemimetabolous Pterygota. 
Rudiments of appendages, however, are well known to be present 
on the tenth segment of many insect embryos (figs. 5 A,g A, XApd). 
The idea that these appendages are developed in the female into the 
third pair of valvulae of the ovipositor is now generally regarded as 
erroneous, since it is clear that both the second and the third pairs 
of valvulae are parts of the gonopods of the ninth segment. Berlese 
(1906) records an anomaly found in an adult female of Locusta 


No. 6 INSECT ABDOMEN—-SNODGRASS 23 


viridissima, consisting of two pairs of appendicular processes on the 
tenth abdominal segment closely resembling the two pairs of valvulae 
on the ninth segment, the outer pair corresponding with the valvulae 
formed of the basal plates, the inner pair with those formed of the 
gonapophyses. It is scarcely to be supposed, however, that an abnor- 
mality of this kind is a “‘ reversion” to an ancestral normality. The 
embryonic limb rudiments of the tenth abdominal segment in all the 
more generalized insects are normally suppressed before hatching. 
In the Holometabola, on the other hand, appendicular structures fre- 
quently occur on the tenth segment in postembryonic stages, and there 
is little doubt that such organs on the larva, typically represented 
by the postpedes of caterpillars, are true limb structures; in adults 
they include the socii of Lepidoptera and Trichoptera, and the cercus- 
like processes of Tenthredinidae, which appear to be derived from 
the larval postpedes. The appendages of the tenth, or pygidial, seg- 
ment of the abdomen may be generally designated the pygopods. 

The eleventh segment (uro-segment).—The eleventh abdominal 
segment represents the last true somite of the body, and is present in 
the embryos of many insects as a well-developed ring bearing the 
rudiments of the terminal pair of appendages (fig. 5 A, Cer). The 
segment is present in adult Protura as a fully formed annulus with 
tergal and sternal plates (B, X/), and in some of the lower Insecta 
having 11 distinct segments in the abdomen the eleventh segment 
is retained likewise as a complete annulus. This condition is well 
shown in Nesomachilis (fig. 7B) where the eleventh segment, though 
mostly concealed within the tenth (A), consists of continuous tergal 
and sternal regions (C, D), and bears laterally the long, filamentous 
cerci (Cer). The tergal region is produced into the median caudal 
filament (cf), and the sternal bridge supports a pair of broad subanal 
lobes, the paraprocts (D, Papt), separated by a median cleft. In 
Thermobia the eleventh segment has a distinct tergal plate, or epiproct 
(E, F, Eppt), but the sternal bridge is lost, and the sternal region 
of the segment is represented only by the paraprocts (F, Papt), upon 
which are borne the cerci (Cer). The median ventral region of the 
eleventh segment is generally obliterated in pterygote insects that 
have a well-developed tenth segment, but in some of the Homoptera, 
as in the cicada (fig. 8 C), the venter of the eleventh segment is not 
only present but it contains a distinct sclerotic sternal remnant (X/S). 

The adult abdomen of most of the lower Pterygota ends with a 
supra-anal plate (fig. 6, Eppt) which is in every way suggestive that it 
corresponds with the epiproct, or tergum of the eleventh segment, in 
the Thysanura. Some entomologists, however, basing their opinion on 


24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


Heymons’ (1895) assertion that the eleventh segment of the embryo 
in Gryllotalpa and other Orthoptera is lost during development, regard 
the supra-anal plate of pterygote insects as belonging to the twelfth 
segment. On the other hand, Wheeler’s (1893) observations on the 
development of Niphidium are fully convincing that the cercus-bear- 
ing eleventh segment persists in the Orthoptera, and, though it be- 
comes reduced, forms the terminal parts of the adult abdomen asso- 
ciated with the cerci. Wheeler’s idea that the rudimentary appendages 
of the tenth segment become the inner valvulae of the ovipositor 1n 
the female does not conform with the evident facts of comparative 
anatomy, but this detail of interpretation does not affect his exposition 
of the segmentation. 

When we compare the usual circumanal structures of pterygote in- 
sects with the parts of the eleventh segment in the Odonata (fig. 12 A, 





Aa 
An JX 
Fic, 9.—Posterior segments and appendage rudiments of embryos of N7phi- 


dium. (Outlines from drawings by Wheeler, 1893, showing segmentation and 
appendages, but with other details omitted. ) 


A, male embryo. B, female embryo, each with cercal appendages (Cer) on 
eleventh segment. C, female embryo in later stage, showing retention of eleventh 
segment structures (NX/). 


Eppt, Cer, Papt), in which Heymons himself has shown that the 
twelfth segment (Prpt) is present though rudimentary, we can 
scarcely question the identity of the parts in all cases. In other words, 
the epiproct, the cerci, and the paraprocts, which in larval Odonata 
clearly belong to the eleventh segment, must be eleventh segment 
structures in all Pterygota, as they are in Thysanura. Heymons’ 
(1904) claim that the appendages of the eleventh segment in the 
Odonata are not true cerci, and that the latter are represented in the 
apparent paraprocts finds no support in comparative anatomy, and 
has been generally rejected on the ground that it would set the Odonata 
apart from all other insects. 

The writer would, therefore, agree with Crampton (1918) that the 
epiproct is in all insects the tergum of the eleventh segment (fig. 6, 


NO. 6 INSECT ABDOMEN—SNODGRASS ‘25 


Eppt), but would dissent from Crampton’s opinion that the cerci 
and paraprocts pertain to the tenth segment, since in such forms 
as Nesomachilis (fig. 7 B) they clearly belong to the eleventh segment, 
and embryologists agree that the cerci are appendages of the eleventh 
segment (fig. 5 A). That the paraprocts at least belong to the same 
segment as the epiproct is evident from their usual anatomical rela- 
tions with the latter. Ford (1923), in her review of the musculature 
of orthopteroid insects, says, “‘ from the musculature we find that the 
supra-anal plate and paraprocts form a symmetrical group,” and fur- 
ther she observes that “the transverse muscles between the supra- 
anal plate and paraprocts show that all three belong to the same seg- 
ment.” The segment represented by the epiproct and paraprocts Ford 
believes is the twelfth, but she bases her opinion largely on Heymons’ 
statement that.the eleventh segment is suppressed in the adult. 

Furthermore, the anatomical relations between the paraprocts and 
the cerci do not support Crampton’s (1920, 1921) contention that 
the paraprocts are the bases of the cercal appendages. The cerci may 
be united with the paraprocts (figs. 7 F, 8A), but generally they 
arise independently in membranous areas behind the tenth tergum 
between the epiproct and paraprocts. The cerci never have muscles 
arising in the paraprocts, and the ventral musculature of the para- 
procts indicates that these plates are merely lobes of the eleventh 
sternum (fig. 6, Papt), as they are actually in Nesomachilis (fig. 
7D). With the usual suppression of the eleventh sternal area, 
however, the paraprocts may appear to arise from the posterior margin 
of the tenth sternum, and they are sometimes continuous with the 
latter (fig. 8 F, Papt). 

The cerci, as shown by their development, are the entire appendages 
of the eleventh segment. Their primitive bases may be represented 
by a small, ring-like segment at the root of each organ (Heymons, 
1896, Walker, 1922), and, as noted above, they are sometimes united 
with the sternal paraprocts, but the muscles of the cerci always have 
a tergal origin. As already observed, most of the cercal muscles arise 
on the tergum of the tenth segment. These anterior muscles of the 
cerci, however, appear to be derived from the intersegmental, longi- 
tudinal muscle fibers primitively extending between the tenth and 
eleventh terga, which have secondarily become motors of the cerct. 
In some Orthoptera, each cercus has a single muscle arising on the 
epiproct. 

The terminal structure of the generalized insect abdomen has a 
certain resemblance to that of a malacostracan crustacean (fig. 10 A). 
The twentieth body segment of the crustacean represents the eleventh 


26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


segment of the insect abdomen, and its appendages, the uropods 
(20Apd), evidently correspond with the cerci. The telson (Tel) 
being suppressed in insects, the tergum of the eleventh abdominal 
segment (twentieth somite) becomes the supra-anal plate, or epiproct. 
The cerci, therefore, may be regarded as the uropods of insects. 
The twelfth segment—Among adult Hexapoda a twelfth segment 
of the abdomen is developed as a complete annulus with tergal and 
sternal plates only in Protura (fig. 5 B, X/7). In the arthropods gen- 
erally the terminal segment is the periproct, or end piece of the body 
containing the anus, anterior to which the true appendage-bearing 
somites are formed. In the malacostracan Crustacea the periproct 





FrG. 10.—Terminal abdominal structures of a crustacean, Anaspides tasmaniae. 


A, posterior part of abdomen, showing the uropods (20Apd) as terminal 
appendages of penultimate segment. B, ventral view of telson, showing lamina 
supra-analis (sa) and laminae sub-anales (/a) surrounding anus (An). 


forms the telson (fig. 10 A, Tel), typically a broad terminal lobe of the 
abdomen, having the anus situated in the basal part of its ventral 
surface (B, An) between two lateral valve-like flaps (/a). A distinct, 
anus-bearing, terminal lobe of the body is said to be present in the 
embryos of some insects (fig. 5 A, Prpt), but in adult insects there 
is never more than a vestige of the periproct, or rudiment of a seg- 
ment beyond the cercus-bearing eleventh somite (fig. 6, XJ/). 

The best example of the retention of a twelfth segment in insects 
is furnished by the larvae of some Odonata, in which the anus is con- 
tained in a small circular fold (fig. 12 A, Prpt) ordinarily concealed 
between the bases of the epiproct (Eppt) and the paraprocts (Papt). 
In the walls of this cirecumanal fold, as Heymons (1904) has shown, 


NO. 6 INSECT ABDOMEN—SNODGRASS 


to 
N 








/ \ Y = = ¥ Z x \ 
va ei / 
Hypt Eppt Papt (© Hypt D 
Fic. 11.—Posterior segments of a noctuid caterpillar. 


A, dorsal view. B, lateral. C, ventral. D, posterior. 
An, anus; Eppt, epiproct; Hypt, hypoproct; Papt, paraproct; Pp, postpedes 
(pygopods ). 


1Gor’ J 
IXLB Sty 





2Gon Papt paptl 
Fic. 12.—Terminal abdominal structures of odonate larvae. 


A, aeschnid larva, posterior view of end of abdomen, with epiproct and para- 
procts spread out, exposing the periproct (Prpt) containing lamina supra-analis 
(sa) and laminae sub-anales (/a) surrounding the anus (An). B, lateral view 
of same with parts in normal position. C, end of abdomen of larva of Archilestes 
grandis, showing gill plates formed of caudal filament (cf) of epiproct (Eppt), 
and of lobes (paptl) of paraprocts (Papt). 


28 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 85 


there is a small dorsal sclerite, or lamina supra-analis (sa), and two 
lateroventral sclerites, or laminae infra-anales (la). These sclerites 
are lost in adult Odonata, but a small supra-anal lobe, possibly a rem- 
nant of the lamina supra-analis, projects from beneath the epiproct. 
A similar lobe occurs in Nesomachilis (fig. 7 D, sa), as well as in some 
other Thysanura, and in larvae of Ephemerida. The supra-anal lobe 
of these insects might be regarded, therefore, as a dorsal remnant of 
the telson (fig. 6, Tel). In most insects, however, no trace of a twelfth 
segment is to be found, and the periproct must be supposed to have 
been reduced to the membranous area at the end of the eleventh 
segment in which the anus is situated. 


Ill. THE ABDOMINAL MUSCULATURE 


We do not have as yet a sufficient knowledge of the comparative 
myology of Arthropoda to furnish a basis for any theory as to the 
nature of the primitive body musculature in this group of animals, 
in which mobility of the body is a characteristic feature. Widely 
different patterns of muscle arrangement are encountered 1n the several 
arthropod classes, and even within a single class, while, among the 
insects, extraordinary differences occur often between larval and 
adult stages of the same species. 

In the Insecta the abdominal musculature consists typically of dorsal 
and ventral longitudinal fibers, dorsal and ventral transverse fibers, 
and lateral dorsoventral fibers; but in none of these muscle groups 
do all the fibers often retain their characteristic positions. 

The development of the body muscles has been described by Cholod- 
kowsky (1891), Heymons (1895), and Nelson (1915). The dorsal 
and lateral muscles are formed from the lateral somatic plates of the 
mesoderm; the ventral muscles arise from the median ventral parts 
of the mesoderm where the somatic and splanchnic layers are united. 
The muscle rudiments, or anlagen, according to Heymons, in insects 
having open coelomic sacs (Blattidae, Gryllus, Acrididae), are formed 
from sac-like evaginations of the mesodermal walls of the seg- 
mental cavities, which are at first tubular, but sooner or later become 
solid strands of cells. In the higher insects, however, in which the 
coelomic sacs are small or but little developed, the muscles either are 
formed by the proliferation of cells from the mesoderm segments, or 
they arise directly from mesenchyme tissue at points corresponding 
with the position of the coelomic sacs of lower insects. 

Since the muscles are derived from the walls of the embryonic 
coelomic sacs, or from the metameric divisions of the mesoderm, we 


ee 


No. 6 INSECT ABDOMEN—SNODGRASS 29 


may assume that the primitive somatic fibers were all intra-segmental 
in arrangement, as they are in the Annelida. With the acquisition 
of secondary segmentation in arthropods, however, consequent upon 
the development of sclerotic plates in the body wall, the principal 
longitudinal fibers became functionally intersegmental (fig. 2 F). The 
body of the animal can thus be shortened by a telescoping of its seg- 
ments brought about by contraction of the longitudinal muscles, and 
it can be compressed by contraction of the lateral dorsoventral muscies. 
In most cases the opposite movements result either from the elasticity 
of the body wall, or from pressure generated by contraction in one 
part of the body transmitted to another through the medium of the 
body liquid and the visceral organs; but in many insects a dilator ap- 
paratus is developed in which certain muscles in both the longitudinal 
and dorsoventral systems become antagonistic to the retractors and 
compressors. 

The abdominal musculature of adult insects is simpler than the 
thoracic musculature because of the absence of leg muscles. There is 
no evidence that the definitive lateral muscles of the abdomen have 
been derived from the body muscles of the limbs. Muscles of the 
movable parts of the abdominal appendages, as will be shown in the 
next section, arise generally within areas of the body wall that may 
be attributed to the limb bases (figs. 32 B, C, 34 B, 36D), except 
the muscles of eversible or retractile sacs which in some cases have 
evidently extended to the dorsum. The general segmental plan of 
the abdominal musculature is usually repeated with only minor varia- 
tions in each of the visceral segments ; in the genital and postgenital 
segments it is more or less obscured by special modifications. 

A rather simple scheme of abdominal muscle arrangement prevails 
throughout all adult pterygote insects; but in the Apterygota and in 
larval forms of holometabolous insects the musculature may be highly 
complex. Some students regard the complex types of musculature as 
representative of a primitive condition from which the simpler types 
have been derived by elimination. There are reasons, however, for 
taking the opposite view, as will later be shown. 

Something is known of the abdominal musculature in most of the 
principal orders of insects ; but the Odonata, Orthoptera, Coleoptera, 
Hymenoptera, and the larvae of Lepidoptera and Diptera have re- 
ceived special attention. Trichoptera and Neuroptera, on the other 
hand, have been particularly neglected, and little has been done on 
the abdominal musculature of Hemiptera, and of adult Lepidoptera 


30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


and Diptera. The following literature contains the principal descrip- 
tions of the abdominal muscles of the various orders of insects known 
to the writer. 

PROTURA: Berlese (1910). COLLEMBOLA : Lubbock (1873). 
THYSANURA: Grassi (1887), giving the characteristics of the 
abdominal musculature of Campodea, Japyx, Machilis, and Lepisma. 
ODONATA: Wallengren (1914), Whedon (1919), Ford (1923), 
Steiner (1929), with descriptions of muscles of first three segments 
in adult by Backhoff (1910) and Schmidt (1915), figures showing 
larval musculature by Calvert (1911, 1915), and a tabulation of the 
muscles by Tillyard (1917). EPHEMERIDA: Ditrken (1907). 
ORTHOPTERA: Gryllus, Voss (1905), DuPorte (1920) ; Divip- 
pus, first three abdominal segments, Jeziorski (1918) ; general com- 
parative study of abdominal muscles of orthopteroid insects, Ford 
(1923). HEMIPTERA: Aphis fabae, Weber (1928). ANO- 
PLEURA: Haematopinus suis, Florence (1921). COLEOPTERA: 
Melolontha, Straus-Dirckheim (1828); Hydrophilus, first three 
abdominal segments, Berlese (1909) ; Dytiscus, Bauer (1910), Speyer 
(1922), Korschelt (1924) ; larval musculature of other species, Ber- 
lese (1909), Boving (1914), Craighead (1916), Paterson (1930). 
LEPIDOPTERA: larval musculature, Lyonet (1762), Lubbock 
(1859), Berlese (1909), Forbes (1914, 1916). HYMENOPTERA: 
larval muscles of a chalcid, Tiegs (1922), of the honeybee, Nelson 
(1924) ; adult musculature of pregenital segments of Vespa, Berlese 
(1909), of Apis, Carlet (1890), Betts (1923), Snodgrass (1925) ; 
full account of abdominal musculature of Apis, Morison (1927). 
DIPTERA: larval musculature partly or briefly described or figured 
in Syrphidae, Ktinckel d’Herculais (1875), in Chironomus, Miall 
and Hammond (1900), in Anopheles, Imms (1908), in Musca, 
Hewitt (1910, 1914), in Khagoletis, Snodgrass (1924) ; full account 
of larval muscles of Psychoda, Dirkes (1928), of Culicidae, Samt- 
leben (1929). 

Less appears to have been done on the body musculature of other 
Arthropoda than on that of insects. Descriptions of the abdominal 
muscles of Crustacea will be found in the paper on the musculature 
of Astacus fluviatilis by Schmidt (1915), in that on Pandalus danae 
by Berkeley (1928), and in that on Copilia dana by Riester (1931). 
A paper by Becker (1926) describes the dorsal body musculature 
of Chilopoda. 


ieee 


eee 


NO. 6 INSECT ABDOMEN—SNODGRASS 51 


GENERAL PLAN OF THE ABDOMINAL MUSCULATURE 


A review of the literature cited above gives a fairly comprehensive 
survey of the abdominal musculature of insects in most of the principal 
orders. There are notable blanks, however, since such important 
orders as Neuroptera and Trichoptera are omitted entirely, and 
adult Lepidoptera and Diptera have been given scant attention. On 
the other hand it is gratifying to find that we have, as a basis for 
a comparative study of insect myology, very full accounts of the body 
musculature of the Odonata, Ephemerida, and Orthoptera. In the 
Apterygota, we are indebted to Berlese for an excellent study of the 
muscles in Protura, to Lubbock for a description of the collemboian 
musculature, and to Grassi for brief descriptions of the characteristic 
differences in the musculature of representative genera of Dicellura 
and Thysanura, to which is added in this paper an account of the 
abdominal muscles of Heterojapyx ; but a more complete study of the 
musculature of Machilidae and Lepismatidae, and perhaps of Cam- 
podea, is still to be desired. When we look to the papers treating 
of holometabolous larvae, we find again satisfactory and in some cases 
complete accounts of the body musculature in Coleoptera, Lepidoptera, 
Hymenoptera, and Diptera, but note with regret a lack of information 
on Neuroptera and Trichoptera. 

To present here even a summary of the details known concern- 
ing the abdominal muscles of insects would occupy an unwarranted 
amount of space. A careful review of the facts to be obtained from 
the works above listed, however, shows that we may with confidence 
make certain broad generalizations concerning the fundamental plan 
of the abdominal musculature of adult pterygote insects. The basic 
plan is found to be simple; but, as so often occurs in insect mor- 
phology, more difficulties are encountered in finding suitable terms to 
express the facts than in discovering the facts themselves. 

Voss (1905) classified the abdominal muscles as longitudinal mus- 
cles, transverse muscles, and lateral muscles (Flankenmuskeln). This 
classification is logical inasmuch as it probably conforms with the 
primitive arrangement of the fibers. The muscles of the so-called 
longitudinal groups, however, do not always preserve a lengthwise 
arrangement; they are often strongly oblique, and some of them 
frequently take a transverse position. The lateral muscles are desig- 
nated “ dorsoventral’? muscles by many writers, but, though their 
attachments are usually dorsal and ventral, some of their fibers com- 
monly run in an oblique direction, The lateral muscles have also 
been termed “ transverse’ muscles, but, as Samtleben (1929) points 


32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


out, only the crosswise dorsal and ventral muscles are literally trans- 
verse in position. Again, the body muscles are sometimes classed 
as dorsal, ventral, and lateral muscles, the dorsals and ventrals com- 
prising longitudinal, oblique, and transverse fibers, and the laterals 
including dorsoventral and oblique fibers. This classification is evi- 
dently the most nearly consistent one that can be devised, and it has 
the added merit of being in conformity with the embryonic develop- 
ment of the muscles. Unfortunately, however, in naming the secon- 
dary muscle groups or individual muscles according to it, the plan 
brings out such terms as “‘ median longitudinal dorsal muscles,” ‘‘ ex- 
ternal median longitudinal dorsal muscles,” or “* second internal median 





Fic. 13.—Diagrams of the more simple types of segmental musculature. 


A, simple condition of musculature in right half of a segment, with dorsal 
(d) and ventral (wv) fibers attached intersegmentally, external laterals (le) 
intrasegmentally, and internal laterals (/i) on intersegmental folds. 

B, upper ends of internal lateral muscles (/i) migrated posteriorly, separating 
a paratergal muscle band (pf) from the rest of the dorsal muscles (d). 

C, cross section of segment, showing relations of principal groups of muscles. 
dl, lateral dorsal muscles; dm, median dorsals; Je, external laterals; li, internal 
laterals ; p, paratergal muscle; td, dorsal transverse muscles; ftv, ventral trans- 
verse muscles; v, ventrals. 


longitudinal dorsal muscle.” Still more unwieldy do such terms be- 
come when put into Latin form. 

It is evident that strict anatomical and nomenclatural consistency in 
dealing with the body musculature leads to impractical results. The 
writer, therefore, has adopted a classification and nomenclature that 
recognizes the anatomical arrangement of the muscles, but which, 
in order to shorten the names, errs somewhat on the side of specificity 
in terminology. Five principal groups of muscles are distinguished 
and designated as follows: I. porsAL mMuscLes (fig. 13 A, d), the 
fibers of which are typically longitudinal and primarily intersegmental 
in their attachments. II. veNnrRAL MUSCLES (v), resembling the 
dorsals in that their fibers are typically longitudinal and attached 
primarily on the intersegmental lines. III. LATERAL mMuscLes (J), 











NO. 6 INSECT ABDOMEN—SNODGRASS a3 


typically dorso-ventral, their fibers both intersegmental and intraseg- 
mental. IV. TRANSVERSE MUSCLEs (C, ft), lying internal to the longi- 
tudinals, including dorsal transverse muscles (td), and ventral trans- 
verse muscles (tv). V.SPIRACULAR MUSCLES (s), generally not more 
than two connected with each spiracle, one an occlusor (os), the other 
a dilator (dls). 

All the body muscles are bilateral in their origin, and all of them 
except the ventral transverse muscles remain so in the definitive state. 
The fibers of the ventral transverse muscles, which primarily are 
intersegmental, Heymons (1895) says are at first attached mesally on 
a fold of the body wall between the nerve cords. Later the fold is sup- 
pressed and the fibers from opposite sides become continuous across 
the sternal region. The fibers of the longitudinal dorsal and ventral 
muscles are always separated into symmetrical lateral groups along 
the midline of the dorsum and venter, but the lateral sets of dorsal 
transverse muscles come together on the ventral wall of the heart. 

Each primary group of muscles may undergo an endless diversifica- 
tion resulting both from a multiplication of fibers in the group, and 
from a rearrangement of the fibers brought about by changes in the 
points of attachment. With respect to the dorsal and ventral muscles, 
the most general departure from the simple plan, in which the fibers 
all lie in a single plane, consists of a differentiation of the fibers 
in each group into internal muscles and external muscles. Thus it 1s 
found in nearly all insects that the dorsal and ventral muscles com- 
prise each two layers of fibers, namely, imternal dorsals (di) and 
external dorsals (de), and internal ventrals (vi) and external ventrals 
(ve). A second form of diversification affecting the dorsal and ven- 
tral muscles is a more or less distinct grouping of the fibers into 
median and lateral muscles. In most insects, therefore, we may distin- 
guish four sets of dorsal fibers, and correspondingly four sets of 
ventral fibers. The several resulting muscles may then be distinguished 
as median and lateral internal dorsals (fig. 14 A, B, dim, dil), median 
and lateral external dorsals (dem, del), median and lateral internal 
ventrals (vim, vil), and median and lateral external ventrals (vei, 
vel). 

The lateral muscles are more subject to irregularities than are the 
dorsal and ventral muscles, but they likewise are often divided into 
internal laterals (fig. 13 B, C, lv) and external laterals (le). 

Associated with the dorsoventral lateral muscles there is sometimes 
present a longitudinal muscle or group of longitudinal fibers lying 
external to the upper ends of the internal laterals (fig. 13, p). This 
muscle is called the ‘“ epipleural”” muscle by Ford (1923), but since 


34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


it occurs on the region of the dorsum, being situated above the line of 
the spiracles, and therefore not on the true pleural region, the writer 
would term it the paratergal muscle (p). Since this muscle belongs 
to the dorsum it should be classed as a dorsal muscle. 

The fibers of the transverse muscles are never differentiated into dis- 
tinct layers, but they may be variously grouped in both the dorsal 
system (fig. 13 C, td) and the ventral (tv). 

The spiracular muscles comprise usually not more than two mus- 
cles associated with each spiracle. One is an occlusor of the spiracle, 
the other a dilator. The occlusor is seldom lacking ; the dilator is less 
constant. 

To express more concisely the principal groups of abdominal mus- 
cles and their subdivisions, we may tabulate the muscles in the follow- 
ing manner : 

I. Muscutti porsaLes (d). 

1. M. dorsales interni (dz). 
a. M. dorsales interni mediales (dim). 
b. M. dorsales interni laterales (dil). 
2. M. dorsales externi (de). 
a. M. dorsales externi mediales (dem). 
b. M. dorsales externi laterales (del). 
3. M. paratergales (p). 
II. MuscuLt VENTRALES (v). 
I. M. ventrales interni (v1). 
a. M. ventrales interni mediales (vim). 
b. M. ventrales interni laterales (vil). 
2. M. ventrales externi (ve). 
a. M. ventrales externi mediales (vem). 
b. M. ventrales externi laterales (vel). 
III. MuscuLt LaTERALEs (J). 
1. M. laterales interni (Jz). 
2. M. laterales externi (Je). 
IV. MuSCULI TRANSVERSALES (ft). 
1. M. transversales dorsales (td). 
2. M. transversales ventrales (tv). 
V. MUSCULI SPIRACULORUM (s). 
1. M. occlusores spiraculorum (os). 
2. M. dilatores spiraculorum (dls). 

Each secondary group of muscles is often again subdivided into 
several bundles of fibers. These ultimate individual muscles may be 
given numerical designations, beginning medially in the case of the 
longitudinal muscles and anteriorly with the lateral muscles. Thus 


No. 6 INSECT ABDOMEN—SNODGRASS 35 


the individual muscles of the median internal dorsals may be 
specifically indicated rdim, 2dim, 3dim, etc., the external laterals rle, 
2le, 3le, etc., and the muscles of the other groups in like manner (fig. 
15 A). If it is desired to show that a muscle belongs to a particular 
segment, this may be expressed by adding to its symbol a Roman 
numeral designating the number of the segment, thus rdimI1, 3vimlV, 
aleV I, etc. In describing the complete musculature of a species, how- 
ever, the writer has found it more practical to number the muscles 
with Arabic numerals, rather than to attempt to follow any system 
of lettering that pretends to identify homologous muscles in con- 
secutive segments. 

The dorsal muscles—The muscles of the dorsum are primarily 
composed of longitudinal fibers of segmental length attached on the 
intersegmental folds ; in many larvae the principal dorsal fibers retain 
this primitive condition. Wherever the dorsum, however, contains 
fully-developed sclerotic terga, a secondary segmentation is estab- 
lished, and the folds on which the dorsal muscles are attached become 
the antecostae of the definitive tergal plates (fig. 14C, Ac). The 
longitudinal, primitively intrasegmental muscles thus become func- 
tionally intersegmental, and serve to contract the abdomen in a length- 
wise direction by retracting each tergum into the posterior end of 
the segment preceding, as far as the intersegmental membrane will 
allow. The anterior end of a longitudinal abdominal muscle, there- 
fore, may be termed the origin, and the posterior end its isertion. 

The differentiation of the dorsal fibers into internal and external 
muscles is the rule in both adult and larval stages of pterygote in- 
sects. The internal dorsals commonly retain their longitudinal posi- 
tions, their segmental lengths, and their attachments on the antecostae ; 
but there are many departures from this generalized condition. Fre- 
quently the fibers take an oblique position, and sometimes they become 
shorter than segmental length by a migration of their origins to the 
postcostal area of the tergum, or of their insertions to the precostal 
area, The external dorsals seldom retain a segmental length ; typically 
they are short muscles lying in the posterior parts of the segments 
(fig. 14 C, de), and often they become strongly oblique, sometimes 
actually transverse, giving a movement of torsion between the two 
segments they connect. Finally, the external dorsals may become 
completely reversed in position (D, de), their origins being so far 
back on each tergum that they lie posterior to the points of insertion 
on the anterior edge of the precostal rim of the following tergum. 
In such cases, the external dorsals become antagonistic to the internal 


3 


30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


dorsals (di), and function as abdominal protractors, since their con- 
traction lengthens the abdomen by decreasing the overlap of the 
segments. 


dem 





Se 





Fic. 14.—Diagrams illustrating more evolved types of musculature, and seg- 
mental mechanisms. 


A, dorsal muscles differentiated into internal and external median dorsals 
(dim, dem), and internal and external lateral dorsals (dil, del) ; lateral muscles 
(1) differentiated into tergo-sternal (t-s), tergo-pleural (t-p), and sterno- 
pleural (s-p) groups; ventral muscles differentiated into internal and external 
median ventrals (vim, vem), and into internal and external lateral ventrals 
(vil, vel). 

B, illustrating the compressor and dilator mechanism of an abdominal seg- 
ment, in which some of the lateral muscles function as compressors (cp), and 
others, attached ventrally on edge of tergum, become dilators (dlr), 

C, usual arrangement of dorsal muscles as seen in longitudinal section, with 
internal dorsals (di) attached intersegmentally on antecostae (4c), and external 
dorsals (de) arising on posterior parts of terga, both sets acting as retractors 
of the terga. 

D, modification by which external dorsals (de), attached in posterior folds 
of terga, become antagonistic to internal dorsals (di) and act as protractors of 
the terga. 


The division of the dorsal muscles into median and lateral groups 
of fibers affects both the inner and the outer layers (fig. 14 A, B, dim, 
dil, and dem, del), but it is not always apparent, and the lateral dorsals 
are sometimes absent. The line of division of the inner dorsals into 


No. 6 INSECT ABDOMEN—SNODGRASS By 


median and lateral muscles is sometimes marked by the points of 
origin of the dorsal transverse muscles on the tergal wall (figs. 13 C, 
15 B, td). 

The paratergal muscle of the dorsum (figs. 13 B, C, 14 B, p) is not 
commonly present in adult insects, or, at least, its fibers are not 
generally separated from those of the other lateral dorsal muscles. It 
is well developed in the Acrididae (fig. 15 B, 169), where it is repre- 
sented in each of the pregenital segments but the first (fig. 16) by 
a band of intertergal fibers lying above the line of the spiracles ex- 
ternal to the upper ends of the internal lateral muscles (fig. 15 B, 
175, 176). According to Ford (1923) it is also present in the 
Plecoptera (“‘epipleural muscle’’). The paratergal muscle occurs 
more commonly in larval insects. In the abdomen of the larva of 
Dytiscus it is represented by a lateral band of fibers (fig. 18, p) cut 
off from the other dorsal muscles (di/, dill) by the upper ends of the 
internal lateral muscles (JiJJ). An apparently corresponding mus- 
cle, consisting of a pair of parallel fibers extending lengthwise on 
each side of the body above the line of the spiracles, is characteristic 
of the caterpillars (figs. 20, 21, p). In the larva of Tipula there is a 
broad band of paratergal fibers on each side of the body attached on 
the middle of the laterodorsal areas of successive segments (fig. 
22,\p). 

The ventral muscles——TVhe ventral abdominal muscles undergo an 
evolution parallel in most respects with that of the dorsal muscles. 
Their fibers are nearly always differentiated into internal and external 
layers, and those of both groups are commonly separated into 
median and lateral groups. The fibers of the internal layer are typically 
intersegmental wherever complete sternal plates are developed, and 
serve as retractors of the ventral arcs of the segments. The external 
ventrals are usually short and take their origins on the posterior parts 
of the sterna. Frequently they become sternal protractors by a re- 
versal of their position, owing to the carrying forward of their points 
of insertion on apodemal arms of the anterior margins of the sterna 
until their morphologically posterior ends lie anterior to their points 
of origin on the preceding sterna. The length of the sternal apodemes 
commonly gives to the ventral protractor mechanism of the abdomen 
a greater effectiveness than has that of the dorsum. 

The lateral muscles—It.is difficult to make satisfactory general- 
izations concerning the lateral muscles of the abdomen, because these 
muscles are subject to more variations in position and attachments 
than are either the dorsals or the ventrals. Most commonly the lateral 
muscles are tergo-sternal in their attachments (fig. 13 C, li, le), and 


38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 85 


typically they are vertical in position; but they may comprise tergo- 
pleural and sterno-pleural fibers (fig. 14 A, t-p, s-p), as well as 
tergo-sternal fibers (t-s), and generally some of them are oblique. 
A division of the lateral muscles into internal laterals and external 
laterals (fig. 13 B, C, li, le) is not always apparent, often because 
of the absence of the internal group, but it is of common occurrence. 

The internal lateral muscles, when present, are longer than the 
external laterals because their upper attachments are at a higher 
level on the dorsum than are those of the external muscles (figs. 
13. C, li, 15 B, 175, 176). The position of the internal laterals along 
the sides of a segment is variable. The muscles are usually situated 
in the middle or anterior parts of the segments (fig. 16), but in 
some cases they are limited to the extreme anterior regions, and in 
certain holometabolous larvae they lie on the intersegmental folds. 
The internal lateral muscles, however, do not in all cases constitute 
a homogeneous group of muscles ; one or more sets of anterior fibers, 
such as those forming the first internal lateral muscle of Dissosteira 
(fig. 15 B, 175), lie internal to the lateral tracheal trunk, while the 
more posterior fibers, as the second internal lateral of Dissosteira 
(176), may lie external to the tracheal trunk. In some insects, on 
the other hand, the entire series of internal lateral fibers are internal 
to the lateral tracheal trunk (fig. 22). 

An example of the limitation of the internal lateral muscles to the 
intersegmental regions is well shown in the larva of Rhagoletis po- 
monella (fig. 23), a trypetid fly, in which the muscles consist of 
slender hands of fibers (7) lying laterally on the intersegmental folds 
in both the abdomen and the thorax. Similar intersegmental muscles, 
comprising each three groups of fibers, are described by Samtleben 
(1929) in the larvae of Culicidae as “‘ musculi dorsoventrales medi- 
ales,” the upper attachments of which are between the ends of the 
dorsal longitudinal muscles, and the lower attachments between the 
ends of the ventral muscles. In the larva of Tipula (fig. 22) the 
internal lateral muscles consist of a series of approximately vertical 
fibers (li) occupying the anterior half of the lateral wall of each seg- 
ment, but the anterior fibers in each segmental group are attached 
on, or close to, the intersegmental fold. All of these fibers lie internal 
to the ventrolateral tracheal trunk (L7ra) and a broad band of lon- 
gitudinal paratergal fibers (p). In the caterpillars a group of several 
internal lateral fibers (figs. 20, 21, li) arise from the lateral extremity 
of each ventral intersegmental fold and diverge posteriorly to the 
dorsum, going internal to the lateral tracheal trunk (Tra) and the 
paratergal muscles (p/p). 


No. 6 INSECT ABDOMEN—SNODGRASS 39 


Similar groups of internal lateral muscles occur in both the abdomen 
and the thorax of the larva of Dytiscus. According to Speyer (1922), 
a two-branched internal lateral muscle, ‘‘ musculus dorsoventralis 
abdominis a,” occurs in the anterior part of each of the first five seg- 
ments of the abdomen in the larva of Dytiscus marginalis (fg. 18, 
lill). The lower ends of these muscles are inserted on the interseg- 
mental folds, but their upper ends are attached in the anterior parts of 
the segments following on the tergal plates between median and 
lateral groups of fibers of the dorsal longitudinal muscles (di, /). 
In the thorax, the upper ends of the corresponding muscles (/i) are 
more nearly intersegmental ; their lower attachments are on the inter- 
segmental “ furcillae ’’ and on the sternal apophyses. These muscles 
of the larval thorax possibly correspond with muscles of the adult 
described by Bauer (1910), “ musculi levatores prothoracis and meso- 
thoracis,” extending from the sternal apophysis of the prothorax 
and mesothorax to the first and second phragmata, respectively. Sim- 
ilar muscles are sometimes present in the thorax of other adult in. 
sects ; one such occurs in Acrididae attached ventrally on the prosternal 
apophysis and dorsally on the intersegmental fold in front of the 
mesepisternum (see Snodgrass, 1920, figs. 32, 34, 50). 

The fragmentary review of the position of the internal lateral mus- 
cles given above suggests that the anterior fibers at least of each 
segmental group represent lateral dorsoventral muscles that are 
primarily intersegmental (fig. 13 A, li). The fibers have a tendency 
in the abdomen to migrate posteriorly, especially on the dorsum (B), 
though they may extend backward along the lateral edges of the 
sternum also (fig. 22). Their upper ends thus cut off a lateral group 
of fibers (fig. 13 B, fp) from the longitudinal dorsals, which become 
the paratergal muscles. These primarily intersegmental internal lateral 
muscles run mesad to the lateral tracheal trunks in some insects 
(Acrididae, lepidopterous larvae, tipulid larvae), and their homologues 
presumably should do so in all insects, but this point has not been 
determined. Other internal lateral muscles of the abdomen, lying 
external to the lateral tracheae, are probably intrasegmental in their 
origin. 

The external lateral muscles are typically dorsoventral and intra- 
segmental (fig. 13 A, B, le). Some of them, however, are frequently 
oblique (fig. 15 B, 775, 770), and the latter may include an interseg- 
mental muscle (fig. 15 A, 4/1). The dorsoventral fibers are sometimes 
attached on the pleural membrane or on “ pleural’’ sclerites, form- 
ing thus tergo-pleural and sterno-pleural muscles (fig. 14 A, t-/~, s-p). 


40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


While most of the lateral muscles are compressors of the abdomen 
(fig. 14. B, cp), since they serve to approximate the sternum to the 
tergum in each segment, some of them, at least in insects that make 
active respiratory movements, serve as dilators (dir). The lateral 
dilators become mechanically antagonistic to the compressors by rea- 
son of the fact that their points of origin are on the lower edges 
of the terga ventral to their insertions on the overlapped edges of 
the sterna. As in the case of the sternal protractor muscles of the 
abdomen, the effectiveness of the dilators is commonly increased 
by the dorsal extension of their points of insertion on apodemes of the 
sterna. 

There is no evidence to suggest that any of the lateral abdominal 
muscles of adult pterygote insects are derived from the primitive body 
muscles of the lost appendages. In larval forms that retain appendage 
rudiments on the abdomen, the lateral muscles lie mesad of the limb 
bases (figs. 34 A, 36 C, 1), attached above on the tergum and below 
on the sternum. The persisting muscles of the abdominal appendages 
pertain to the distal movable parts of the organs, and these muscles 
take their origins within the limb bases (figs. 34 A, 36D). Exceptions 
to this occur in the case of the muscles of retractile vesicles of holo- 
metabolous larvae (fig. 36 C, D, rvs), which take their origin on the 
dorsum, but these muscles are not retained in the adult.. The branchial 
muscles of ephemerid larvae (fig. 15 A, bmcls) are said to persist in 
the adult stage, but they do not appear to correspond with any of 
the lateral muscles in other pterygote insects. 

The transverse muscles——The transverse muscles of the abdomen 
are best known as the muscles of the dorsal and ventral diaphragms 
(fig. 13 C, td, tv). It seems probable that primitively these muscles 
were intersegmental in position, their fibers being attached on the 
intersegmental folds, one set being dorsal, the other ventral. 

The fibers of the dorsal transverse muscles arise typically in groups 
on the anterior edges of the lateral parts of the abdominal terga, and 
spread mesally to their insertions along the ventral wall of the heart. 
Only in a few insects are they evenly distributed along the entire 
length of the tergum, or collected into anterior and posterior groups. 
The usual anterior origin of the fibers, therefore, suggests that the 
dorsal transverse muscles are primarily intersegmental. In the cater- 
pillars (fig. 21 A, td) they practically have this position, except that the 
diverging inner ends of the fibers spread into the anterior and posterior 
parts of the adjoining segments. Usually the muscles of the dorsal 
diaphragm extend from the second to the eighth or ninth abdominal 
segment, but in the Blattidae they are said to occur not only in the 


NO. 6 INSECT ABDOMEN—SNODGRASS 41 


first abdominal segment, but also in the mesothorax and the meta- 
thorax (Brocher, 1922). In the larvae of anisopterous Odonata, 
according to Whedon (1919), a muscular dorsal diaphragm is present 
only in the fourth or the fifth segment of the abdomen. 

The ventral transverse muscles in some of the Orthoptera (‘Tet- 
tigoniidae and most of the Gryllidae) take the form of widely sepa- 
rated compact bundles of fibers crossing the anterior parts of the 
segmental sterna. In others, as in the Acrididae, the origins of the 
fibers are distributed along the sides of the sterna, and the muscles 
form a typical ventral diaphragm occupying most of the length of the 
abdomen. Ford (1923) thinks that the compact type, that is, the one 
in which the fibers form individual transverse muscles segmentally 
arranged, is the primitive type, and that it has been derived from a 
diffuse or web-like type. The writer, however, believes that the 
relations may be the reverse, especially considering Heymons’ 
(1895) statement that the transverse muscles of Orthoptera are 
formed in the embryo along the intersegmental folds. In the larvae 
of anisopterous Odonata, according to Whedon (1919), there is 
in the abdomen only a single, large, spindle-shaped, somewhat flat- 
tened ventral transverse muscle lying in the extreme anterior part 
of the sixth segment, attached laterally on the intersegmental fold. 

In the higher insects in which ventral transverse muscles are present 
in the adult, as in Hymenoptera, the muscles form a continuous sheet 
of tissue over the ventral sinus. Ventral transverse muscles are usu- 
ally absent in holometabolous larvae. In the honeybee larva, Nelson 
(1924) says, there is present in the newly hatched larva a well- 
developed ventral diaphragm consisting of a continuous sheet of trans- 
verse fibers, but in older larvae it becomes a vestigial structure formed 
of more or less isolated fibers entirely too few in numbers to con- 
stitute more than a loose and insignificant meshwork. 

The spiracular muscles—The musculature of the abdominal spira- 
cles includes one or two muscles associated with each spiracle. The 
muscle most generally present is an occlusor. This is a short muscle 
usually attached at both ends on the base of the spiracular atrium, 
where its contraction compresses the inner end of the atrium and so 
closes the entrance to the trachea. In the Acrididae the occlusor 
muscle arises dorsally on the tergal wall close behind the spiracle. A 
dilator, or opening muscle of the spiracle, antagonistic to the occlusor, 
occurs at least in most of the Orthoptera, Lepidoptera, and Hymenop- 
tera, but it is absent in Odonata, some Orthoptera, and Coleoptera. 
The dilator commonly takes its origin on the tergum or on the lateral 
margin of the sternum of the segment in which the spiracle is situated. 


42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


The musculature of the thoracic spiracles is usually different from 
that of the abdominal spiracles, as is the structure of the spiracles 
themselves. The spiracles of Apterygota and Ephemerida are said 
to have no musculature. 


THE ABDOMINAL MUSCULATURE OF ADULT PTERYGOTA 


The musculature of the visceral segments of the abdomen in 
pterygote insects adheres closely to the generalized plan of structure, 
though there are usually slight aberrations in the first segment or 
first and second segments. The musculature of the genital and post- 
genital segments is often highly specialized or reduced, but it is un- 
doubtedly derived from the same muscle pattern as that prevailing 
in the less modified segments. The usual departures from the gener- 
alized musculature in the visceral region of the abdomen consist 
principally of a reduction in the number of muscles, a shortening in 
the length of some of them, and a shifting of the points of attach- 
ment, bringing about simple changes in the position of certain mus- 
cles. A brief examination of the orders in which the abdominal mus- 
culature is best known will serve to show the extent and nature of the 
modifications that take place in the visceral segments. The more ex- 
tensive modifications in the specialized genital and postgenital seg- 
ments need not concern us here. 

E phemerida.—The most generalized abdominal musculature of the 
adult pterygote type occurs in the Ephemerida, and the muscle pattern 
is here essentially the same in both adult and larval stages. We may, 
therefore, follow Dutrken’s account of the larval musculature of 
Ephemerella ignita, which can easily be verified in any ephemerid 
species. Most of the abdominal muscles (fig.15 A), except those 1n- 
serted on the gill bases (bmcls), lie in a single plane against the body 
wall, and are comprised in dorsal (d), lateral (/), and ventral (v) 
groups. The first two dorsals (rd, 2d), counting outward from the 
median line, and the second and third ventrals (2v, 3v) in most of 
the segments are typical intersegmental, longitudinal muscles attached 
on the anterior margins of successive segmental plates. The third 
dorsal (3d), however, is atypical in that most of its fibers take their 
origin on the middle of the tergum and cross the following segment 
to be inserted on the anterior margin of the second tergum follow- 
ing. This muscle Durken calls a “ compound intersegmental ” mus- 
cle. The fourth dorsal (4d) and the first and fourth ventrals (7v, 
jv) are short muscles arising on the posterior parts of the segmental 
plates before those of their insertions. These muscles thus appear 





NO. 6 INSECT ABDOMEN—SNODGRASS 43 


to correspond with the external dorsal and ventral muscles usually 
more definitely differentiated from the internal muscles in the majority 
of the Pterygota. 

The true lateral body muscles of Ephemerella include only the 
intrasegmental, vertical, tergo-sternal muscles (fig. 15 A, zl, 2l, 31), 
and the intersegmental, oblique tergo-sternal muscle (4/). The 
groups of branchial muscles (bimcls), inserted in the larva on the 
bases of the gills, are described and figured by Durken as arising 
on the lateral parts of the sterna. The areas on which these muscles 


NIT 
/ 












\ / 
Ac ms RB Ivs 


Fic. 15—Examples of abdominal musculature. 


A, musculature of left half of seventh and eighth segments (seen externally) 
of abdomen of larva of Ephemerella ignita (from Diurken, 1907) ; bmcls, bran- 
chial muscles, shown here as originating on lateral part of sternum, which is 
probably the ventral area of limb basis (see fig. 34 B). 

B, muscles of right half of third abdominal segment (seen internally) of an 
acridid, Dissosteira carolina, 167, internal median dorsals; 168, internal lateral 
dorsal; 769, paratergal muscle; 170, external median dorsal; 177, external 
lateral dorsal; 172, median ventral; 173, internal lateral ventral; 774, external 
lateral ventral; 775, 176, internal laterals; 177, 178, 179, external laterals; td, 
insertion point of dorsal transverse (cardiac) muscles. 


arise, however, are clearly distinct from the true sterna (fig. 34 A, 
B, Stn), and very evidently represent the bases of the abdominal 
limbs (LB), of which the gills (Brn) are the distal movable parts. 
The branchial muscles, therefore, are not body muscles, but are in- 
trinsic muscles of the appendages, and take their origins within the 
limb bases. The true lateral body muscles (fig. 34 A, J) are tergo- 
sternal in their attachments and lie mesad of the lobes (LB, LB) 
supporting the gills. The gill muscles of the ephemerid larva, Diirken 
says, are retained without change in the adult. They do not appear 


44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


to have representatives in the pregenital segments of any other adult 
pterygote insect. 

Odonata.—The muscles of the first three abdominal segments of 
adult Odonata are described by Backhoff (1910) and by Schmidt 
(1915) in connection with a study of the male genital organs, and 
some of the abdominal muscles of odonate larvae are figured by Cal- 
vert (I9gII, 1915). A more complete description of the larval muscu- 
lature as a part of the respiratory mechanism is given by Wallengren 
(1914), and of that of the adult by Steiner (1929), while a full 
account of both the adult and larval muscles in Zygoptera and Anisop- 
tera will be found in the paper by Whedon (1919) on the morphology 
of the odonate abdomen, a few errors in which are corrected by 
Steiner (1929). 

In the abdominal musculature of the Odonata there is nothing 
to suggest a type of structure more primitive than that of other 
Pterygota ; the fundamental plan of the muscle arrangement is that of 
pterygote insects in general, and has little to distinguish it from the 
muscle pattern of orthopteroid insects. The generalized plan of 
musculature is best retained in the larvae of Zygoptera. The muscles 
here comprise internal and external longitudinal dorsals, internal and 
external longitudinal ventrals, and dorsoventral and oblique lateral 
muscles. The internal dorsal and ventral fibers are of segmental 
length ; but the externals in each set are short, taking their origins 
on the posterior parts of the segments. In the Anisoptera the larval 
muscles are more strongly developed than in the Zygoptera, evidently 
as an accommodation to the respiratory and locomotor functions 
of the rectum, and the broad internal dorsal and ventral bands of 
fibers take on oblique direction. The adult musculature is much re- 
duced in the abdomen, and most of the muscles are very short, but 
the arrangement of the muscles shows no radical departure from 
the fundamental pterygote pattern better preserved in the larva. 

Orthoptera—The comparative myology of the abdomen is better 
known in the Orthoptera than in any other of the larger orders of 
insects owing to the comprehensive review by Ford (1923) of the 
abdominal musculature of orthopteroid insects. Then, too, Voss 
(1905) in his thorough study of the thorax of Gryllus includes an 
account of the muscles of the anterior abdominal segments, and 
Du Porte (1920) describes the entire musculature of the abdomen 
in the same genus. 

The abdominal musculature of the Orthoptera and related orders 
shows in all groups a differentiation of the dorsal and ventral muscles 


NO. 6 INSECT ABDOMEN—SNODGRASS 45 
into internal and external layers of fibers, and in most cases a well 
marked separation between median and lateral fibers in each of these 
groups. In a general statement on the abdominal musculature of 
orthopteroid insects, Ford (1923) says: ‘In the common ancestors 
of the orthopteroid insects the tergal musculature probably consisted 
of two broad layers, an internal longitudinal and an external oblique, 
with the inner layer approximately equaling the length of the tergum, 
and the outer layer much shorter. Of the present-day orders the 
Blattaria approach closely this hypothetical type.” Of the ventral 












PNz I 140 14:9 
! Il/ 
197 
ae va | 
lay — Vil 
—— font 
rs = AA 
= 
= LAA 
i a Lares 
LS] Sls & | i 
Se = 
DEEL [7A 














7. Yi [aN \ \ \ 
Pe 145 143 IS 154 ie 187 202 
Fic. 16.—Musculature of the right half of the first five segments of the ab- 
domen of Dissosteira carolina, together with dorsal muscles (7112) of meta- 
thorax. (Compare with fig. 15 B.) 


140-197, median internal dorsals; 150-198, lateral internal dorsals; 143-202, 
median ventrals; 744, first lateral internal ventral; 145, first lateral external 


ventral. 


musculature Ford says: “The hypothetical type of sternal muscu- 
lature is similar to the tergal, having the two-layered arrangement 
of longitudinal ental and oblique ectal muscles. The Blattaria again, 
resemble the hypothetical type.” 

Since the external muscles of the dorsum and venter are not always 
oblique, being often parallel with the internal muscles, and since, 
furthermore, the dorsal and ventral muscles are not always differ- 
entiated into external and internal layers (fig. 15 A), it would seem to 
the writer more probable that all the dorsal and ventral fibers were 


46 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


primarily longitudinal and of segmental length, and that they had 
this arrangement in pre-orthopteroid insects. The internal dorsals and 
ventrals are likewise often oblique. Obliquity, therefore, would ap- 
pear to be secondary also in the external muscles, in which it may 
be so accentuated that the muscles lie in a transverse direction, or are 
even reversed in position, 

The lateral muscles of the Orthoptera are variable in their positions 
and in their attachments. They include typical vertical and oblique 
intrasegmental tergo-sternal muscles, oblique intersegmental tergo- 
sternal muscles, and in some cases muscles that may be termed “‘ tergo- 
pleural” and “ sterno-pleural,” since they are inserted on the lateral 
membranes or on sclerites below the line of the spiracles. The so- 
called pleural areas on which these last named muscles are attached, 
however, probably really belong either to the dorsum or to the venter 
of the segment, and, if so, none of the lateral muscles is properly a 
* pleural” muscle. 

In the Acrididae the internal dorsal muscles are distinctly separated 
into median and lateral groups of fibers (figs. 15 B, 16, 167, 168) 
by the points of attachment of the dorsal transverse fibers on the 
tergum (fig. 15 B, td). The external dorsals assume very oblique 
or transverse positions (170, 171). The ventral muscles are well 
differentiated into median and lateral groups of internal fibers (172, 
173) and into lateral external muscles (174). The external ventrals 
(174) are sternal protractors by a complete reversal in the relation 
between their points of attachment. The lateral muscles in the third 
and succeeding segments (figs. 15 B, 16) comprise two internal dor- 
soventral laterals (fig. 15 B, 175, 176), and three external laterals 
(177, 178, 179), of which the first (777) is an abdominal dilator by 
reason of its sternal attachment being on the upper end of a large 
lateral sternal apodeme (/A4p). The upper ends of the internal 
laterals (175, 176) are attached on the tergum between the lateral 
internal dorsals (168) and a broad paratergal dorsal muscle (1769). 
This last muscle is the “epipleural’’ muscle of Ford (1923), who 
says a similar muscle also is present in the Plecoptera. 

Coleoptera——The abdominal musculature of adult Coleoptera is 
known principally from the description of Melolontha vulgaris by 
Straus-Durckheim (1828), and of Dytiscus marginalis by Bauer 
(1910) and Korschelt (1924). 

The adult musculature of the abdomen of Dytiscus is relatively sim- 
ple. As described by Bauer (1910) it consists of dorsal longitudinal 
muscles, ventral longitudinal muscles, and lateral muscles, to which 
list should be added the transverse muscles of the dorsal diaphragm. 


No. 6 INSECT ABDOMEN—SNODGRASS 47 


Bauer terms the lateral muscles ‘“‘ musculi transversales abdom- 
inis,” but, as pointed out by Samtleben (1929), the lateral muscles are 
dorsoventral and should not be termed “ transverse.” The largest mus- 
cles of the Dytiscus abdomen are the dorsal muscles. These consist of 
broad bands of fibers forming a wide sheet of muscles against the ter- 
gal region in each of the first six segments. The ventral muscles are 
present only in segments III, 1V, and V. They include large median 
ventrals and small lateral ventrals. The median ventrals form three 
pairs of muscle sheets occupying the median sternal region of the seg- 
ments, the fibers of the opposite groups in each pair converging pos- 
teriorly. The lateral ventrals (“ musculi ventrales externi’”’ of Bauer) 
are very small, each arising on the posterior lateral angle of the 
sternum of its segment, and being inserted on the anterior margin of 
the sternum following. The lateral muscles (‘‘ musculi transversales ” 
of Bauer) comprise a pair of small, oblique tergo-sternal muscles 
crossing each other in the form of an X in each side of segments I] 
to V inclusive, and a single oblique muscle in segment VI. 
Hymenoptera—The honeybee furnishes the principal information 
that we have on the abdominal musculature of Hymenoptera. The 
muscles of a typical abdominal segment of the honeybee have been 
described by Carlet (1890), Betts (1923), and Snodgrass (1925) ; 
the complete abdominal musculature is given by Morison (1927). 
The muscles characteristic of the part of the abdomen involved in 
respiration are well shown in the third and fourth segments (fig. 17). 
The dorsal muscles consist of three sets of fibers in each half of the 
seginent, two of which are internal and one external. The internals 
form a broad median band of fibers (dim) slanting ‘from in front 
posteriorly and medially, and a slenderer lateral muscle (dil) ex- 
tending from in front posteriorly and laterally. The external dorsal 
isa short muscle (del) arising laterally on the posterior margin of the 
tergum and extending forward to its insertion on the tip of a lateral 
tergal apodeme of the following segment. The two sets of dorsals 
are thus antagonistic, the internal fibers being tergal retractors, and 
the external fibers tergal protractors. The ventral musculature com- 
prises internal and external muscles, which are likewise antagonistic. 
The internal fibers form an oblique median internal ventral (wi) 
on each side of the sternum, the two converging mesally in the form of 
a V, anda slenderer lateral muscle (vil) oblique in the opposite direc- 
tion. The external ventrals consist of a single small, fan-shaped lateral 
muscle on each side (vel), arising laterally on the posterior part of 
the sternum and inserted anteriorly on the lateral anterior apodeme 


48 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL 85 


of the following sternum. The lateral muscles comprise three tergo- — 


sternal muscles in each side of the segment. .The first (71) is a dilator 
of the abdomen, since it arises ventrally on the lateral part of the ter- 
gum and is inserted dorsally on the tip of the lateral apodeme of 
the sternum; the second and third laterals (2/, 31) are oblique tergo- 
sternal compressors of the abdomen. 






i i 
If 


li 





Fic. 17—Musculature of two consecutive abdominal segments of adult Apis 
mellifica, showing the muscles on the left side. 


Ac, antecosta; del, lateral external dorsal muscle; dil, lateral internal dorsal; 
dim, median internal dorsal; zl, 2/, 3/1, first, second, and third laterals; dls, 
dilator muscle of spiracle; Sp, spiracle; vel, lateral external ventral muscle; 
vil, lateral internal ventral; vim, median internal ventrals. 

The first lateral muscles (z/) are dilators of the abdomen, the second and 
third laterals (2/, 31) are compressors; the internal dorsal and ventral muscles 
(dim, dil, vim, vil) are retractors of the segments, the external dorsals and ven- 
trals (del, vel) are protractors. 


THE ABDOMINAL MUSCULATURE OF ENDOPTERYGOTE LARVAE 


The body musculature of endopterygote, or holometabolous, larvae, 
in its*higher forms of development, attains an extreme degree of com- 
plexity ; in its simpler forms it differs but little from the body muscu- 
lature typical of all adult Pterygota. It appears, therefore, that the 
complex types of larval musculature represent specialized conditions 
adapting the larvae to their individual ways of living, and are not to 
be interpreted as meaning that insects are derived from ancestral 
worm-like forms having an intricate body musculature. The changes 
in the musculature that occur during the pupal metamorphosis are to 
be regarded as alterations necessitated by the restoration of the 
normal adult musculature, which involve varying degrees of destruc- 
tion or reconstruction in the special, temporary larval musculature. 

Coleoptera.—The larval musculature of Trichoptera and Neurop- 
tera has not been fully studied, nor do we have any comparative work 


a 





No. 6 INSECT ABDOMEN—SNODGRASS 49 


on the larval muscles of the Coleoptera. The complete account of the 
muscles of the Dytiscus larva given by Speyer (1922) and by Kor- 
schelt (1924), however, furnishes a basis for an understanding of 
the relation between the larval musculature and the musculature of 
adult insects. The structural changes which take place in the trans- 
formation from the larval to the imaginal musculature have been 
described by Breed (1903) in a trogositid, Thymalus marginicoliis. 

The abdominal musculature of the larva of Dytiscus marginalis, 
as described by Speyer, consists of four primary groups of fiber 
bundles, namely, dorsal muscles, ventral muscles, and, on each side, 
a set of lateral (dorsoventral) muscles. In the region of the first seven 
abdominal segments, the dorsal muscles comprise an internal set of - 





Fic. 18.—Body muscles in right half of thorax and first two abdominal seg- 
ments of Dytiscus marginalis larva. (Outline from figure by Speyer, 1922, re- 
lettered in accord with muscle nomenclature adopted in this paper.) 


di, internal dorsal muscles; /’, internal lateral muscles, upper ends of which 
cut off paratergal muscles (p) from the other dorsals; wi, internal ventrals. 


intersegmental longitudinal fibers of segmental length (fig. 18, dil, 
dill), and outer sets of short fibers extending from the posterior 
parts of the terga to the following intersegmental folds. The ventral 
muscles consist likewise of internal (vi/, vill) and external sets of 
intersegmental fibers. In each side of the first five abdominal segments 
Speyer distinguishes six lateral (dorsoventral) muscles. Five of 
these are external laterals, three of which are tergo-sternal and two 
tergo-pleural in their attachments. The other lateral muscle is an 
internal lateral and consists of two bundles of fibers (liJJ), which 
arise ventrally by a common base on the intersegmental fold. Dorsally 
the two branches are inserted on the anterior lateral part of the 
tergum between median and lateral sets of the dorsal longitudinal 
fibers (di and p). In the thorax the muscles (li) corresponding with 


50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


the internal laterals of the abdomen occur between the prothorax 
and mesothorax, between the mesothorax and metathorax, and be- 
tween the metathorax and first abdominal segment. They are attached 
above more nearly on the anterior margins of the terga, and are in- 
serted ventrally on the intersegmental “ furcillae ”’ and on the sternal 
apophyses. The posterior migration of the upper ends of the internal 
laterals in the metathorax and abdomen cuts off a lateral group of 
fibers (fp) from the longitudinal dorsals that evidently corresponds 
with the paratergal muscle of Dissosteira (fig.-15 B, 160). 

It is clear that the abdominal musculature of the Dytiscus larva 
differs in no essential respect from that characteristic of adult insects 
generally. It presents a more primitive condition in that the internal 
lateral muscles retain ventrally their intersegmental attachments, 
whereas in most adult insects, when present, their ventral ends have 
migrated posteriorly along the edges of the sterna (fig. 15 B, 175, 
170, tie. iO): | 

The figure given by Berlese (1909) of the muscles in the first three 
abdominal segments of the larva of Pentodon, and the studies of 
Boving (1914) and of Craighead (1916) on the abdominal muscula- 
ture of coleopterous larvae, including species of Cleridae, Trogositidae, 
Elateridae, and Scarabaeidae, suggest that the chief deviation’ from the 
Dytiscus larval muscle pattern consists only of a greater diversification 
in the position of the muscles, and of an increase in the number of 
muscles or individual fibers in each group. In any case it is clear that 
the larval musculature in the Coleoptera presents at most but a small 
increase in complexity beyond the minimum characteristic of adult 
pterygote insects. Proceeding from this condition found in the 
Coleoptera, therefore, we may expect to find that the more complex 
musculature of other holometabolous larvae represents only a more 
highly specialized condition. 

Hymenoptera.—In the larvae of Hymenoptera the body muscula- 
ture also retains a relative simplicity. The pattern of the abdominal 
muscles of the honeybee larva (fig. 19), as described by Nelson 
(1924), departs but little from the basic plan of the general adult 
pterygote musculature, though it is somewhat more complex than the 
abdominal musculature of the adult honeybee (fig. 17), and is not at all 
like the latter in detail. The dorsal muscles of a typical abdominal 
segment of the larva (fig. 19) consist of broad bands of internal 
longitudinal fibers (di) of segmental length, and of shorter, oblique 
external fibers (de). Some of the external fibers, by a transposition 
of their posterior attachments on the intersegmental fold, have come 


no. 6 INSECT ABDOMEN—SNODGRASS 51 


to overlap internally the internal dorsals. The ventral muscles include 
internal oblique (vi) and external oblique muscles (ve), all of seg- 
mental length. The lateral muscles comprise dorsoventral and oblique 
external laterals (/e), and a strong, oblique, internal sterno-tergal 
muscle (Ji) attached on the consecutive intersegmental folds. It may 
be questioned whether this last muscle represents the internal laterals 
of the Dytiscus larva (fig. 18, li), but the only difference between 
the two is that the upper end of the muscle in the bee larva is attached 
on the intersegmental fold following that of its ventral attachment, 
a change that might have come about by a posterior migration of its 
dorsal end. 





wit\ 


WF. 
Wf 





Fic. 19—Musculature of right half of two consecutive abdominal segments 
of honeybee larva. (Figure from Nelson, 1924, but relettered in accord with 
muscle nomenclature adopted in this paper.) 


Con, ganglionic connectives; de, external dorsal muscles, some of them secon- 
darily internal at posterior ends; di, internal dorsals; Gung, segmental ganglion; 
Ht, heart; le, external lateral muscles; //, internal lateral muscle; Sp, spiracle; 
ve, external ventral muscles; vi, internal ventrals. 


Lepidoptera.—the larvae of Lepidoptera have long been noted for 
the great number of muscles that lie against the body wall, and for the 
extreme complexity in the arrangement of the fibers. Fully 150 mus- 
cles, mostly individual fibers, may be counted in a typical abdominal 
segment of any caterpillar (figs. 20, 21). The principal muscles of 
the innermost layer (figs. 20, 21 A) are definite bands of parallel 
longitudinal fibers having segmental lengths and attached on the inter- 
segmental folds. In the ventral region there are also strong external 
muscles of segmental length having an oblique position. Most of 
the external fibers, however, are of various lengths and are disposed 
in all directions against the body wall (fig. 21 B). On each side of 
the body, between the principal dorsal and ventral groups of muscles 


4 


52 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


there is a pair of slender, longitudinal paratergal muscles (figs. 20, 
21 A, B, p) lying just above the line of the spiracles and the lateral 
tracheal trunks (7va). Anteriorly in each segment the paratergal 
muscles are crossed internally by a group of internal lateral fibers 
(li) arising ventrally on the intersegmental fold, and diverging dor- 
sally and posteriorly to their attachments on the dorsum. These in- 
ternal lateral muscles of the caterpillar lie internal to the lateral tra- 
cheal trunks. The dorsal transverse muscles of the caterpillar (figs. 
20, 21 A, td) arise in groups immediately dorsad of the paratergal 
muscles from the posterior margins of the intersegmental folds. 





P Tra 2S et ee VNC vi Tra Zp 





Fic. 20.—Ventral musculature of fourth abdominal segment of a caterpillar, 
Estigmene acraea. 


li, internal lateral muscles; p, paratergal muscle; fd, origins of dorsal trans- 
verse (cardiac) muscles; Tyra, lateral tracheal trunk; vi, internal ventral mus- 
cles; NC, ventral nerve cord. 


The complexity of the body musculature of the caterpillar appears 
to demonstrate that the muscle system of insects has no limits imposed 
on its possibility of diversification both by multiplication and by re- 
arrangement of its fibers, since there is no reason to believe that the 
intricate pattern of the caterpillar muscles represents in any way the 
primitive plan of insect musculature. In the other organization of the 
lepidopterous larva there is little to suggest a primitive condition. The 
head and mouth parts present the typical fundamental structure of 
these organs that has been developed in adult Pterygota, and on this 
basic structure have been built up the many special features of the 


NO. 6 INSECT ABDOMEN—SNODGRASS 53 


caterpillar head and mouth parts adapted to the needs of the larva. 
The alimentary canal of the caterpillar is highly specialized in its 
musculature. The ae of the nervous and tracheal systems 








eS 


CLL 


Fic. 21.—Abdominal muscles of a caterpillar, Estigmene acraca, seen by re- 
moval of inner muscles shown in figure 20. 

A, muscles of right half of fourth segment (//”) after removal of internal 
ventrals (vi). B, outermost muscles in right half of third segment, showing 
particularly the leg muscles; cross-hatched areas represent bases of hair-bearing 
tubercles. 

Cx, basal rim of leg; di, internal dorsal muscles; Ht, heart; /i, internal lateral 
muscles; p, paratergal muscles; td, dorsal transverse (cardiac) muscles ; Tra, 
lateral tracheal trunk; vi, internal ventral muscles. 


is not necessarily an indication of a primitive state; it is merely the 
retention of a generalized structure in these organs accompanying ¢ 
high specialization in others. The presence of appendages on the pre- 


54 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


genital segments of the abdomen likewise signifies nothing more than 
the retention of organs useful in the larval stage. In short, the worm- 
like form of the caterpillar and of other holometabolous larvae has 
no phylogenetic significance. It is a secondary adaptation, derived 
from the normal adult pterygote structure, accompanied by numerous 
specializations peculiar to the larva, and later discarded. The com- 
plex musculature of the caterpillar is only one of the features in the 
larval organization that have been specially evolved from the gener- 
alized adult structures of the immediate ancestors of the Lepidoptera 
to enable the caterpillar to perform more efficiently the duties that have 
devolved upon it from the apportionment of the life processes between 
the immature and adult stages of the individual. 

Diptera——The musculature of the larvae of Diptera shows a 
unique type of specialization in its highest development, but at the 
other extreme it has a pattern corresponding entirely with that of the 
generalized plan of abdominal musculature in adult Pterygota. 

The simpler forms of dipterous larval musculature, known in the 
Tipulidae, Psychodidae, Chironomidae, Culicidae, and Tabanidae, con- 
sist of dorsal and ventral bands of longitudinal fibers, and of lateral 
dorsoventral muscies. A primitive type of musculature occurs in the 
Psychodidae, where, as described by Dirkes (1928) for Psychoda 
alternata, the dorsal and ventral muscles are mostly longitudinal and 
attached on the intersegmental folds, though a few in each set are 
shorter than segmental length. In the first abdominal segment there 
are five dorsoventral laterals and two oblique laterals on each side. 
The first of the dorsoventral muscles is attached on the intersegmental 
fold between metathorax and abdomen, the others follow along the 
side of the segment. A similar condition exists in the Culicidae, as 
described by Samtleben (1929), except that here some of the inner 
muscles of the dorsal and ventral series in each segment cross ob- 
liquely over the outer muscles, and the inner lateral muscles are con- 
fined to the anterior parts of the segments, where they are attached on 
the intersegmental folds between the ends of the dorsal and ventral 
muscles. In the larva of Tipula (fig. 22) the musculature is compli- 
cated by a great increase in the number of fibers in all the principal 
groups, and by a diversification in their points of attachment, but 
there are few fibers taking an oblique course. In both the dorsal and 
ventral groups certain sets of fibers are attached regularly on the 
intrasegmental transverse folds of the body wall, and some of the 
median ventral fibers form somewhat oblique interlacing bundles. The 
internal lateral muscles (Ji) comprise a series of dorsoventral fibers 





5 


Cn 


No. 6 INSECT ABDOMEN—SNODGRASS 


lying in the anterior half of each segment internal to the laterai 
tracheal trunk (LTra). Outside of these muscles, and external to the 
tracheal trunk, is a wide band of longitudinal paratergal fibers (/) of 

























































































i 


ABLE 
SEW ue Bl 


mM Ho 
FY best 
J TG 
(ES 


= 
Fy 
| 
H 
a 
HI 


































































































Fic. 22.—Musculature of right half of two consecutive abdominal segments 
of Tipula abdominalis larva. 

di, internal dorsal muscles; /i, internal lateral muscles distributed in anterior 
half of segment internal to lateral tracheal trunk (LT7ra) ; p, band of paratergal 
fibers; vi, internal ventral muscles. 





Fic. 23.—Musculature of the body wall of the larva of a cyclorrhaphous 
dipteron, Rhagoletis pomonella. 

A, musculature of the thoracic and first abdominal segments, seen from above, 
with pharynx and connected parts turned forward. B, musculature of right 


half of two consecutive abdominal segments. 
_ASp, anterior spiracle; fs, frontal sacs; Ht, heart; /e, external lateral muscles ; 
li, internal lateral muscles; Phy, pharynx, turned forward; wi, longitudinal 


bands of internal ventral muscles. 


segmental length, but attached on the median intrasegmental folds of 
the dorsum. The numerous external laterals lie against the body wall 
external to the paratergal fibers. 


56 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


In the larvae of the higher Diptera the dorsal and ventral muscu- 
lature appears to be merged into a double series of oblique muscles 
regularly crossing one another to form a network pattern repeated 
throughout the thorax and the abdomen (fig. 23). The only muscles 
that preserve a longitudinal direction are two bands of ventral fibers 
continued from the prothorax to the end of the abdomen (v7). 
There can be little doubt that this type of musculature represents a 
highly specialized condition, correlated with the great specialization 
which the maggot shows in nearly all other parts of its body organ- 
ization. The lateral muscles, on the other hand, appear to retain 
a primitive condition. The internal laterals consist of slender fibers 
lying on the intersegmental folds (Ji) along the sides of the body. 
The external laterals (Je) comprise a small group of fibers in the 
side of each segment against the body wall external to the network 
of oblique muscles. 


THE ABDOMINAL MUSCULATURE OF APTERYGOTA 


The body musculature of apterygote hexapods is not well known 
in all the major apterygote groups; it has been carefully studied in 
representatives of Protura, Collembola, and Dicellura, but only casu- 
ally examined in Thysanura. Particularly desirable, therefore, would 
be a complete account of the body muscles of Machilidae and Le- 
pismatidae. 

Protura—The abdominal musculature of the Protura is fully 
described by Berlese (1910) in his monograph on the “ Myriento- 
mata.” In this group of hexapods, Berlese says, “ the musculature 
is extraordinarily complex by reason of the great multiplicity of 
fibers extending in all directions, very much as in the larvae of meta- 
bolic insects.” The muscle pattern of the proturan abdomen as shown 
by Berlese, however, is not complex by comparison with that of a 
caterpillar or of a muscoid maggot, and the proturan body muscles 
clearly fall into the three usual categories of insect muscles, namely, 
dorsal muscles, ventral muscles, and lateral muscles, to which are to 
be added the body muscles of the appendages. 

The dorsal abdominal muscles of Protura are divided into external 
dorsals and internal dorsals. The internal dorsals (muscles of the 
second stratum of Berlese) consist of broad bands of fibers in the 
Acerentomidae attached on the successive tergal antecostae. In the 
FEosentomidae they are differentiated into median and lateral groups 
of fibers. The external dorsals (muscles of the third stratum of 

serlese) include two large oblique muscles on each side of each seg- 


NO. O INSECT ABDOMEN—SNODGRASS 57 


ment; one (the intersegmental tergal muscle of Lerlese) arising 
medially on the anterior part of the tergum and inserted laterally on 
the antecosta of the following tergum, the other (the intersegmental 
tergopleural muscle of Berlese) arising anteriorly on the tergal ante- 
costa and inserted posteriorly on the “pleuron” of the following 
segment. The so-called ‘ pleuron,” however, is probably to be re- 
garded as a paratergal sclerite. 

The ventral muscles of the abdomen include likewise external 
ventrals and internal ventrals. The internal ventrals consist of paired 
bands of longitudinal fibers extending throughout the length of the 
abdomen. Each muscle band is divided into a median group of fibers 
attached on the sternal antecosta of each segment, and into longer 
lateral muscles in the first five segments attached on alternate sterna. 
The external ventrals occur only in the first three abdominal seg- 
ments of Acerentomidae. Those of the first segment extend from the 
center of the sternum laterally to the bases of the appendages of 
this segment, and serve as adductors of the appendages. In the sec- 
ond and third segments corresponding pairs of muscles arising on 
the anterior median part of each sternum diverge posteriorly to the 
antecosta of the following sternum. 

The lateral musculature of the abdomen in Protura has a very sim- 
ple pattern. The lateral muscles comprise intrasegmental vertical 
tergo-sternal muscles, and intersegmental oblique tergo-sternal mus- 
cles. The intrasegmental laterals include in each of the first three 
abdominal segments of EKosentomidae, and in the first segment only of 
Acerentomidae, a pair of tergal muscles inserted ventrally on the base 
of the appendage, and in the following segments of Acerentomidae 
a single lateral tergo-sternal muscle. The intersegmental laterals con- 
sist of two slender muscles arising laterally at each end of the tergal 
antecosta in each segment, one of which goes to the anterior margin 
of the sternum of the preceding segment, the other to a corresponding 
point on the sternum of the following segment. From the first ab- 
dominal tergum a muscle extends downward to the posterior edge of 
the metathoracic sternum, and another goes forward to the posterior 
edge of the mesothoracic sternum. 

An analysis of the proturan musculature, as described by Berlese, 
thus shows that the Protura suggest nothing different as to the pattern 
of the primitive body musculature of the Hexapoda from the idea 
to be derived from a study of the muscles of adult Pterygota. 
Since the lateral musculature in the Protura does not match with that 
of any pterygote insect, it does not appear to be the prototype of 
the lateral musculature characteristic of the Pterygota, and therefore, 


58 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 85 


probably represents a special development. The Protura have no 
transverse muscles. Berlese describes a dorsal septum above the 
alimentary canal, but, he says, it is composed entirely of a connective 
tissue membrane and contains no muscle fibers. A closed dorsal vessel 
is likewise absent. 

Collembola.—The account of the body musculature of the Collem- 
bola given by Lubbock (1873) is so complete and so convincing in 
its detail that no doubt can be entertained of its accuracy, though 
apparently no subsequent investigator has verified it, or given any 
attention to the musculature, other than that of the appendages, in 
this interesting group of insects. Lubbock describes the muscles of 
Lomocerus as an example of the musculature of a “ linear ” species, 
and those of Smynthurus to illustrate the musculature of a “ globular ” 
species. It is clear that the muscle pattern in the abdomen of the 
former is more generalized than in that of the latter, but that in both 
forms the musculature is modified in adaptation to the specialized 
functions of the abdominal appendages. 

The abdominal musculature of Tomocerus is highly developed, 
consisting of strong bands of longitudinal dorsal and ventral muscles 
differentiated into internal and external groups of fibers, and of 
vertical and oblique dorsoventral lateral muscles. In the first segment 
two strong muscles arising on the tergum are inserted on the eversible 
vesicle of the collophore. In the third segment groups of lateral 
muscles are attached ventrally on the sternal region in the neighbor- 
hood of the tenaculum, but they do not appear necessarily to be 
primarily muscles of the pair of appendages presumably combined 
in this organ. The muscles of the furcula, or spring supported on the 
fifth segment, take their origins in the fourth and third segments, 
but they appear to be parts of the system of longitudinal body muscles 
rather than specific muscles of the leaping appendage. In Smynthurus 
the abdominal musculature is highly modified. The longitudinal mus- 
cles appear to be reduced and are mostly absent in typical form. On 
the other hand, there is a great development of vertical and oblique 
dorsoventral muscles associated with the base of the furcula, taking 
their origins in the posterior and middle parts of the abdomen. 

In no respect can the collembolan musculature be said to be primi- 
tive ; but it is evident that it may be derived from the same generalized 
plan of muscle arrangement that underlies the abdominal musculature 
of adult pterygote insects. 

Dicellura.—lIt is most interesting to aad in Grassi’s description of 
the muscles of Campodea that the pattern of the abdominal muscula- 
ture of this primitive apterygote insect conforms closely with the 


No. 6 INSECT ABDOMEN—SNODGRASS 59 


fundamental plan of the abdominal musculature of pterygote insects. 
According to Grassi, the musculature of an abdominal segment of 
Campodea comprises longitudinal dorsal and ventral muscles, oblique 
dorsal and ventral muscles, and dorsoventral lateral muscles. The 
longitudinal muscles are clearly the internal dorsals, and internal 
ventrals. The oblique dorsal muscles are the external dorsals. The 
external ventrals are represented by a pair of muscles convergent from 
the posterior margin of the segment to the mid-sternal region below 
the ganglion. These muscles Grassi terms musculi subganglionares. 
In addition to these there are also small oblique and transverse lateral 
ventral muscles. The true lateral muscles include several small tergo- 
sternal fibers on the sides of each segment. Finally there are the 
muscles of the styli and eversible vesicles. 

By comparison with Campodea, or with almost any other insect, 
the body musculature in the Japygidae is extremely intricate, being 
highly complicated by the presence of numerous muscles that appear 
to have no relation to muscles in a simple type of musculature. The 
following account of the abdominal musculature of a member of this 
group is based on a study of specimens of the Australian Heterojapyx 
gallardi, for which the writer is indebted to Dr. R. J. Tillyard. 
Females of this huge japygid reach a length of 40 millimeters, and 
a dissection of the muscles in well preserved specimens is not a 
particularly difficult task. 

The entire body musculature of Heterojapyx anterior to the ninth 
abdominal segment is highly complex, there being in each of the first 
eight segments of the abdomen at least 4o pairs of muscles, the 
arrangement of which makes a most intricate pattern against the body 
wall (fig. 24). In the mesothorax and metathorax the musculature is 
quite as complex, and in many details quite different from that of the 
abdomen, and is more diversified by the presence of the leg muscles. 
In the ninth abdominal segment the musculature is simplified. In the 
tenth it consists of a single pair of fiber bundles, but these constitute 
two great lateral muscles, almost completely occupying the segment, 
which act as adductors of the cercal forceps. 

The 40 muscles in either half of a typical abdominal segment of 
Heterojapyx, shown in figure 24 representing segment VI, are com- 
prised in the following groups: 

I. Dorsat MuscLes.—A median band of inner longitudinal inter- 
segmental dorsals (A, ta, 1b, Ic); two medio-lateral oblique inter- 
segmental dorsals (A, B, 2, 3) ; two latero-median oblique interseg- 
mental dorsals (B, 4, 5); and an outer longitudinal intersegmental 
dorsal (B, 6). 


60 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 





| / 
/ 7), 
VUS B 
Fic. 24 A, B.—Musculature of sixth abdominal segment of Heterojapy.+ 
gallardi. 
A, muscles of ventral region and right half of sixth segment, and anterior 


part of seventh segment. B, same view with muscles 7, 8, 9, 17, 10, 20, 21 partly 
or entirely removed. 








No. 6 INSECT ABDOMEN—SNODGRASS 61 


Fic, 24 C, D.—Outermost muscles of right half of sixth abdominal segment 
of Heterojapyx gallardi. 


C, posterior part of right half of sixth segment and anterior part of seventh 
sternum. D, right half of sixth segment, showing segmental sclerites. 

Ac, antecosta; Apt, sternal apotome; /t, laterotergal lobe; Wb, intersegmental 
membrane; Pc, precosta; rpl, 2pl, 3pl, 4pl, pleural sclerites; Sp, spiracle; Sty, 
stylus; V/S, VIS, sixth and seventh abdominal sterna; ’ JT, V/IT, sixth and 
seventh abdominal terga; 7-40, muscles of sixth segment (see text, pages 59, 62). 


62 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL: 85 


IJ. VENTRAL MUSCLES.—A group of internal transverse interseg- 
mental sterno-pleural fibers (A, 7) ; an oblique intersegmental sterno- 
sternal muscle (A, 8); two imner longitudinal intersegmental ven- 
trals (A, 9, 10); five outer oblique intersegmental ventrals, four 
of which are median (B, 77, 72, 13, 14), and one lateral (75). 

II. LaTerRAL MUSCLES.—An anterior intrasegmental tergo-pleural 
muscle (A, B, 16); an oblique intersegmental sterno-tergal muscle 
(A, 17) ; an oblique intrasegmental tergo-sternal muscle (A, B, 18) ; 
a series of five intrasegmental transverse tergo-sternal muscles (A, 
19, 20, 21, 22, 23); two short lateral intra-tergal muscles (D, 24, 
25); two small anterior intrasegmental tergo-pleural muscles (D, 
26, 27); two median intrasegmental pleuro-sternal muscles (C, D, 
28, 29) ; an oblique intersegmental pleuro-sternal muscle (A, B, C, 
30) ; a group of small external posterior lateral intrasegmental mus- 
cles (C, 32, 32, 33, 34, 35; D, 38) ; and two small posterior interseg- 
mental muscles, one sterno-pleural (C, 36), the other pleuro-sternal 
(vas 

IV. Muscles OF THE STYLUS.—Two small muscles (D, 39, 40) 
arising in the posterior lateral lobe of the sternum, inserted on the base 
of the stylus (Sty). 

This complex and strongly developed musculature of Heterojapy., 
which presumably is characteristic at least of the Japygidae, contains 
nothing to suggest that it represents the primitive plan of the body 
‘musculature of insects. It indicates, on the other hand, a highly 
specialized condition giving to these very small creatures a strength 
out of proportion to their size, which might enable them to burrow 
into hard soil or to insinuate their bodies into minute irregular spaces. 
In the multiplicity of individual muscles and in the diversity of 
their attachments, the body musculature of H eterojapyx resembles 
that of a caterpillar, but there is not the remotest likeness in detail, 
showing that the complexity of the muscle pattern in each case is but 
the result of a high degree of specialization adaptive to demands for 
dexterity of body movements. Both the caterpillar and Heterojapyx 
demonstrate the limitless potentiality of the insect muscular system, 
and make it all the more surprising that there are so few departures 
from the fundamental plan of muscle arrangement. 


IV. THE ABDOMINAL APPENDAGES 


There is no more vexing subject in the whole field of insect mor- 
phology than that of the homologies of the appendicular organs of 
the abdomen. Embryology shows at most that these organs are de- 


No. 6 INSECT ABDOMEN—SNODGRASS 63 


rivatives of the segmental appendages ; it gives no positive evidence as 
to what part of a primitive limb may be preserved in the definitive 
rudiment, since the latter, whatever it may be, develops directly from 
the embryonic rudiment, instead of following what we should suppose 
would be the course of the phylogenetic evolution of the organ. 
Comparative anatomy is more likely to foster illusions than to lead 
to definite results, for while certain categories of facts may seem 
to align themselves satisfactorily in some limited scheme of suggested 
homology, the plan invariably breaks down when wider generaliza- 
tions are attempted. The writer, therefore, can offer nothing new on 
the fundamental morphology of the abdominal appendages of insects 
that is likely to be generally accepted. Even so, however, it will 
be sufficiently worth while to bring together the principal facts at 
present known concerning the anatomy of the various appendicular 
structures. 

Though the appendages of the insect abdomen are rudimentary in 
the sense that they do not in any case represent a fully-developed 
limb, they are in all cases specialized by a structural adaptation to 
some particular use. The abdominal appendages of most interest to 
entomologists are those of the genital segments, and if we can dis- 
cover a means of identifying these organs in the various insect orders, 
this discovery alone will be of much practical value, and it then be- 
comes a less consequential matter if we can not fully decide the exact 
morphological nature of the organs themselves. 

It is not possible, however, to study with profit any modified or 
specialized appendicular organ without having some concept of the 
nature of the primitive limb structure from which it has been derived. 
Since there are current several different ideas concerning the funda- 
mental structure of a primitive arthropod limb, it is therefore neces- 
sary for a writer to make clear at the outset of a discussion the par- 
ticular theory from which he proceeds. The following sketch will 
give briefly the view on this subject here taken, and a more extensive 
discussion at the conclusion of this section will examine the possibil- 
ities of interpreting the structures of the abdominal appendages of 
insects according to the terms of the theory adopted, which is essen- 
tially that of Borner (1921), though with differences in special appli- 
cations. 

A comparative study of arthropod appendages soon shows that the 
number of segments in the limbs, the relative size of the segments, 
and even the segmental musculature are so variable in different 
arthropod groups that none of these features can be used as a guide 


64 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


for establishing the homologies of the segments or parts of the limb 
in any specific case. There are two joints of the limb, however, that 
recur in the same form in such a large number of appendages in 
the various arthropod groups as to suggest that they represent two 
primary points of flexure in the primitive ambulatory appendages, 
and that they may, therefore, be accepted as “‘ constants ” in the limb 
structure. These joints in a thoracic leg of an insect are the coxo- 
trochanteral joint, and the femoro-tibial joint (the Hiiftgelenk and 
the Kniegelenk of Borner, 1921). The first (fig. 25 A, f-g) divides 
the appendages into a basal region, or limb basis (LB), and a distal 
shaft, or telopodite (Tlpd), which is movable on the basis in a vertical 
plane by a horizontal, dicondylic hinge (f-g). The limb basis, in the 





b B Pa. \Ptar 


Fic, 25.—Diagrams showing the relation of the base of a tee te to the body, and 
the theoretical progress of segmentation in the limb. 


A, theoretically primitive appendage divided into a basis (LB) and a telopodite 
(Tlpd); the first movable antero-posteriorly on the body by a vertical axis 
(a-b) between tergum and sternum, the second movable on the basis in a ver- 
tical plane by a dicondylic, horizontal hinge (f-g) with levator and depressor 
muscles (O, Q) arising in the basis. 

B, the fully segmented appendage: the basis divided into coxa (Cx) and sub- 
coxa (Scv), the latter becoming the pleuron; the telopodite divided into the 
usual segments of an arthropod leg beyond the coxa. 


sense here understood, includes the potential coxa and subcoxa, which 
in some arthropods are differentiated as distinct parts of the basis (B, 
Cx, Scx), the coxa then becoming the functional or movable base of 
the appendage, while the subcoxa becomes a part of the lateral and 
ventral walls of the supporting body segment. The second funda- 
mental joint of the limb forms the “knee” (A, ft), and divides the 
telopodite into a proximal trochantero-femoral piece, and a distal 
tibio-tarso-praetarsal piece, the two movable on each other in a vertical 
plane by an articulation which is either monocondylic, or dicondylic. 

If we conceive, thus, that the primitive arthropod limb is divided 
primarily into a basis and a telopodite, we should expect the baso- 
telopodite joint to be the point of flexure most generally preserved, 


No. 6 INSECT ABDOMEN—SNODGRASS 65 


and, as above noted, a joint does occur in the proximal part of 
practically all fully-developed arthropod appendages that is evidently, 
from its structure and musculature, to be identified as the joint be- 
tween the primitive basis and the telopodite. This joint is the coxo- 
trochanteral joint of an insect’s leg. It is then reasonable to assume 
that the same joint 1s retained in reduced appendages, and that, finally, 
in an unsegmented limb rudiment it is the telopodite that has been 
lost, and that the part which remains is the basis. 

The appendages of arthropods are prone to develop appendicular 
processes on the limb segments. Such processes may be either endites 
or exites, or both forms may occur on the same segment. [Endites are 
developed particularly on the basis, serving as masticatory lobes on 
the gnathal appendages. In the Crustacea, exites of the basis are often 
gill-bearing organs, and an exite of the proximal segment of the 
telopodite commonly forms an outer branch of the appendage known 
as the exopodite. The study of rudimentary appendages becomes 
complicated by the fact that it is often difficult or impossible to 
determine whether a persisting part represents the main shaft of the 
limb, or an appendicular process of the latter. 


BODY APPENDAGES OF CHILOPODA 


The centipedes furnish a good example of arthropods that have 
retained a long series of body appendages preserving the form and 





Ftc. 26.—Somewhat diagrammatic cross section of a body segment of Litho- 
bius, showing the relation of the subcoxa (Scr) and coxa (Ca) to the body 
and to the telopodite (7/pd). 


function of ambulatory limbs. Each appendage of the pregenital re- 
gion of the body is implanted in a membranous pleural area of its 
supporting body segment between distinct tergal and sternal plates 
(figs. 26, 27 A). The movable basal piece of a typical chilopod leg is 
a small segment generally termed the coxa, or coxopodite (Cx). The 


66 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


coxa supports the six-segmented telopodite (fig. 26, T/pd), the proxi- 
mal segment of which, or first trochanter (177), is articulated to the 
coxa by a typical coxo-trochanteral hinge (fig. 25 B, f-g). Surround- 
ing or partly surrounding the base of the coxa, in most of the pre- 
genital segments but the last, is an area of the body wall containing 
one or several small sclerites (fig. 27 A, Scx). These sclerites appear 
to belong to the subcoxal region of the primitive limb basis, since 
upon this region are inserted the tergal muscles of the appendage, and 
within it arise muscles of the coxa. The coxa turns antero-posteriorly 
upon the subcoxa by an approximately dorso-ventral axis (fig. 25 B, 
c-d). 

The large terminal pair of legs of a chilopod borne by the last pre- 
genital segment (fig. 27 A, T/pd) are supported each upon a single 
large plate in the lateral segmental wall (LB). The basal joint of each 
of these legs clearly corresponds with the coxo-trochanteral joints of 
the preceding appendages, and a comparison of the leg-bearing plate 
of this segment (LB) with the coxal and subcoxal sclerites of the 
segments immediately anterior to it leaves little doubt that the single 
‘“pleural”’ plate of the last segment represents both the coxa and the 
subcoxa of the preceding segments (Ca, Scv). In other words, the 
large pleural plates supporting the legs of the last pregenital segment 
are the limb bases (LB) undivided into coxal and subcoxal parts as in 
the other segments. The condition here, of course, may be the result 
of a secondary union of the subcoxal sclerotizations with the coxa, 
but it gives a convincing demonstration of the potential unity of the 
coxal and subcoxal regions of the limb basis, and at least suggests 
a primitive condition in which the limb basis occupied the lateral walls 
of the body segment between the tergal and sternal plates (A, C, IT, 
IStn). The levator and depressor muscles of the telopodite of the 
last pair of legs arise on the plate of the limb base and on the 
sternum (D, O, Q), and have their insertions on the first trochanter 
(rey 

The basal structure of the last pair of legs in the chilopoda is 
paralleled exactly in that of the legs of more generalized Arachnida 
as in the Phalangidae (fig. 46 A), in which the free part of each leg 
is supported on a large basal plate (1B) implanted in the lateral wall 
of the body. Borner (1904) regards the single basal plate of the 
terminal pair of chilopod legs as the united coxa and subcoxa; but in 
the Arachnida, he concludes (1921) that subcoxae are absent and that 
the plates supporting the telopodites are the coxae alone. It is not 
clear why structures so evidently similar should be differently inter- 
preted. 


No. 6 INSECT ABDOMEN—SNODGRASS 67 


In the Diplopoda the free basal segment of the leg, judging from 
its structure and the nature of its articulation with the next segment, 
would appear to be the coxa, and since the sternal plates of the Diplop- 
oda surround the bases of the legs, we may conclude with Borner 
(1921) that the definitive sterna include the subcoxae. Silvestri 
(1903), however, regards the free basal segment of the diplopod leg 
as the subcoxa, and the next segment as the coxa, though the latter 





Fic. 27.—Appendages and terminal body structures of Chilopoda. 

A, posterior part of body of Scolopocryptops sexspinosa, telopodites removed 
except from last segment, showing union of subcoxal sclerites with coxa in large 
pleural plate (LB) on last pregenital segment. B, ventral view of genital and 
pregenital segments of Lithobius. C, lateral view of same. D, base of pregenital 
appendage of Lithobius. E, telopodite of right gonopod of Lithobius, mesal 
surface. 

An, anus; Cx, coxa; Crpd, coxopodite; GO, gonopore; Gp, gonopod; gS, 
sternum of genital segment; GSeg, genital segment; /st, intersternal sclerite; 
LB, limb basis; ]Stn, sternum of pregenital segment; /7, tergum of pregenital 
segment; O, levator of telopodite; Prpt, periproct; Q, depressor of telopodite; 
Scx, subcoxa; Stn, segmental sternal plate; T/pd, telopodite; rT yr, first trochan- 
ter; 277, second trochanter (praefemur ). 


segment has all the structural features and usual relations of a 
first trochanter. 

Following the last leg-bearing segment in the Chilopoda comes 
the definitive genital segment (fig. 27 A, GSeg), beyond which is the 
periproct (Prpt), or anal segment. According to Heymons (1gor) the 
genital region of the body contains two small somites in the embryo, 
parts of both of which are sometimes retained in the adult stage. The 


5 


68 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


apparently single genital segment of the adult may be a mere mem- 
branous ring (A, GSeg), or it may be a well-developed segment with 
dorsal and ventral plates (C, gT, gS). In Lithobius the ventral plate 
of the genital segment bears laterally on its posterior margin a pair of 
small, three-segmented appendages, the gonopods (B, C, Gp, E), the 
basal muscles of which (E, O, Q) arise on the sternal plate of the 
segment. The definitive sternum of the genital segment, thegefore, is 
clearly a composite plate which includes the true bases of the genital 
appendages united with the primitive segmental sternum. The free 
genital appendages, then, are not the entire gonopods, but are the 
telopodites of the latter, and their muscles (E, O, Q) are the levators 
and depressors of the first trochanter (Tr). 

In the males of many insects of the higher orders the structure of 
the second genital segment and its clasperlike appendages (the harpes) 
closely resembles the condition in Lithobius. Though the claspers are 
but one-segmented, they are movable by muscles arising in the gonopod 
bases, and the latter are generally more or less united with the sternum. 
In the insects, however, it is not so clear that the claspers are the true 
telopodites of the gonopods, since there is evidence to suggest that 
they may be other appendicular processes of the bases of the genital 
appendages. 


ABDOMINAL APPENDAGES OF CRUSTACEA 


All the body segments of the Crustacea anterior to the telson are 
usually provided with well-developed appendages. In the lower crus- 
tacean groups, the appendages of the entire body series, as in Apus 
(fig. 28 A), are fundamentally uniramous in form, though the various 
segments may be provided with endite and exite lobes. Each limb 
consists of a basis (LB), called the coxopodite, and of a telopodite 
(Tlpd). The frequent biramous form of crustacean appendages (C) 
is evidently the result of the hyper-development of an exite of the 
basal segment of the telopodite (the first trochanter, or basipodite, 
Bspd). The shaft of the telopodite beyond the basipodite then be- 
comes the endopodite (Endpd). The exite lobes are movable by mus- 
cles arising in the limb segments that support them. 

The abdominal appendages of the Malacostraca are typically bira- 
mous limbs (fig. 28 C) in which the endopodite (Endpd) is usually 
reduced to the size of the exopodite (Expd). The basis, or coxopodite 
(Capd), and the basipodite (Bspd) may be distinct segments, but in 
some forms (B) they are united in a single protopodite (Prtpd). 
In certain cases the abdominal appendages become practically unira- 


No. 6 INSECT ABDOMEN—SNODGRASS 69 


mous by a suppression of the endopodite (B, Endpd), or by its con- 
version into a genital process. In such cases the functional or loco- 
motory shaft of the appendage is the exopodite (Ewpd). A crustacean 
limb of this type of structure furnishes an analogy with the abdominal 
limbs of Thysanura on the assumption that the stylus of the latter 
(fig. 31 A, Sty) is the exopodite, and that the endopodite has been 
entirely suppressed, or preserved only in the gonapophyses of the 
genital appendages (B, Gon). 





Fic. 28.—Abdominal appendages of Crustacea. 


A, Apus longicaudata, appendage from posterior part of body, left, anterior 
surface. B, Anaspides tasmaniae, third abdominal appendage of left side, anterior 
surface. C, Spirontocaris groenlandicus, abdominal appendage of left side, 
anterior view. 

Bnd, basendite; Bspd, basipodite; Capd, coxopodite, or limb basis; Endpd, 
endopodite; Expd, exopodite; LB, limb basis, or coxopodite; Prtpd, protopodite 
(united basis and basipodite) ; 7/pd, telopodite. 


The genital claspers of the higher insects, which are clearly homo- 
logues of the abdominal styli of the Thysanura, may thus be 
likened either to the gonopods of the Chilopoda (fig. 27 C, Gp), if we 
assume that they are the main shafts of the telopodites, or to the ab- 
dominal appendages of such crustaceans as Anaspides (fig. 28B), 
if we assume that they are exites of the appendages. As will later be 
shown, however, it is difficult to obtain positive evidence as to the 
nature of the insect abdominal styli; whether they are likened to the 
main shaft of the telopodite or to an exopodite branch depends largely 
on the student’s bias toward a myriapodan or a crustacean ancestry 
for the insects. 


70 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 85 


THE ABDOMINAL APPENDAGES OF PROTURA 


A pair of short, cylindrical appendages is present on each of the 
first three abdominal segments of all adult Protura. These appendages 
arise from the membranous parts of these segments between the pos- 
terior angles of the tergal and sternal plates. They are best developed 
in Eosentomidae, where the three pairs are alike in size and structure, 
and each organ (fig. 29 B) consists of two segments and a small 
terminal vesicle (v) which is eversible and retractile. In Acerento- 
midae (A) the appendages of the first pair are like those of the 
Eosentomidae, but the second and third pairs are simple, tuberculiform 


2 oex 


—Cx 








Fic. 29.—Abdominal appendages of Protura. 


A, Eosentomon germanicum, abdominal leg (from Prell, 1913). B, Aceren- 
tomon doderoi, first abdominal leg (from Berlese, 1910). 

Cx, coxa; I, promotor muscle of limb base; J, remotor muscle; rv, retractor 
of vesicle; Scr, subcoxa; T/pd, rudiment of telopodite; v, terminal vesicle. 


protuberances, unsegmented and lacking the terminal vesicle. Each 
appendage of the larger type in the two families, as described by 
Berlese (1910), is movable by two tergal muscles (B, /, J) inserted 
on the basal segment, one anteriorly, the other posteriorly. The sec- 
ond segment is provided likewise with two muscles, one arising 
anteriorly, the other posteriorly in the proximal segment, the two 
crossing each other axially to be inserted on opposite sides of the base 
of the distal segment. The terminal vesicle is retracted by a single 
large muscle (rv), which takes its origin mesally on the base of the 
first segment of the appendage, and is inserted on a central depression 
of the ventral face of the vesicle. The extrusion of the vesicle is 
evidently brought about by blood pressure from within the body. 


No. 6 INSECT ABDOMEN—SNODGRASS 71 


There appears to be no reason to doubt that these abdominal 
appendages of the Protura are remnants of true post-thoracic limbs. 
They have, as Berlese points out, a certain resemblance to the ab- 
dominal legs of lepidopterous larvae; but a closer comparison shows 
differences in the segmentation and musculature which makes it seem 
probable that there is no close genetic relation between the two sets of 
organs. Prell (1913), in his study of Eosentomon germanicum, finds 
at the base of each abdominal leg two small sclerotizations which 
he regards as remnants of the subcoxa (fig. 29 A, Scv). The large 
basal segment he believes is the coxa (C2) and the smaller distal seg- 
ment the rudimentary telopodite (7/pd). The homology of the 
terminal vesicle (v) is doubtful. The organ does not appear to 
represent the eversible sacs of Thysanura, since the latter are borne 
by the limb bases (fig. 4, Vs) ; it might be, however, as Prell sug- 
gests, the praetarsus, since it has a certain resemblance to the vesicular 
praetarsus of Thysanoptera. The most likely homologue of the 
proturan leg vesicles is to be found in the eversible sac on the 
collophore of Collembola (fig. 30 B, v), which probably represents the 
united vesicles of a pair of fused appendages. 


GENERAL STRUCTURE OF THE ABDOMINAL APPENDAGES OF INSECTS 


Most of the appendicular organs found on the abdominal region of 
insects fall into two quite distinct categories distinguished by the 
insertion points of their muscles. In those of one group the muscles 
are inserted on the base of the organ; in those of the other the 
muscles traverse the organ and are inserted within its distal extremity. 
Appendicular structures of the first class are typically stylus-like in 
form, though they take on various other shapes. They include such 
organs as the abdominal styli of the Thysanura and the more general- 
ized Pterygota, the furcula of Collembola, the gills of ephemerid 
larvae, the terminal claws of trichopterous larvae, the lateral abdominal 
appendages of larvae of Sialidae, the gonapophyses, the movable 
claspers of male pterygote insects, and the cerci. Organs of the sec- 
ond class are sac-like or tubular in form, and are usually retractile and 


-eversible. They include the collophore of Collembola, the eversible 


vesicles of Thysanura, the gill-bearing tubercles of some sialid larvae, 
and the plantar lobes of the abdominal legs of larvae of Lepidoptera 
and chalastogastrous Hymenoptera. 

If we could accept the two categories of abdominal appendicular 
structures, distinguished by the muscle insertions, as morphological 
groups of organs, the study of the abdominal appendages of insects 


72 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


would be much simplified. But, unfortunately, there are other fea- 
tures than the muscle insertions to be considered, such as the origins 
of the muscles and the position of the organs on the body, that make 
it doubtful if either constitutes a homogeneous group. Furthermore, 
organs of each type frequently occur together supported on a common 
basal structure, and the latter must then be reckoned as an essential 
part of the primitive appendage, of which the free, movable parts are 
but accessory structures of one kind or another. Any attempt to make 
theoretical generalizations on the primitive form or on the homologies 
of the abdominal appendages of insects will be premature until we have 
more closely examined the structure of the principal types of such 
organs as occur in both larval and adult stages of the various insect 
orders. 

The abdominal appendages of insects fall into three groups corre- 
sponding with the subdivision of the abdomen into pregenital, genital, 
and postgenital regions. The appendages of the pregenital segments 
may be designated the pregenital appendages; those of the genital 
segment are the gonopods; those of the first and second postgenital 
segments are conveniently termed the pygopods and uropods, respec- 
tively. The pygopods are the anal legs, or postpedes, of holometab- 
olous larvae, and probably the socii and so-called “cerci” of holo- 
metabolous adults. The uropods are the true cerci. 


THE ABDOMINAL APPENDAGES OF COLLEMBOLA 


The well known appendicular organs on the abdomen of Collembola 
include organs of each type of structure as defined above according 
to the muscle insertions. 

The collophore (fig. 30 A, Col) is a large, thick, tubular pouch of 
the body wall projecting from the sternal region of the first abdominal 
segment. In most species it ends in a bilobed terminal vesicle (B, v), 
which is ordinarily retracted but is capable of being protruded by 
blood pressure. The entire collophore is traversed by a pair of large 
muscles (rv) arising on the tergal region of the body and inserted on 
the lobes of the terminal vesicle. The structure of the collembolan 
collophore thus suggests that it is formed by the union of a pair 
of abdominal appendages resembling those of the Protura (fig. 29), 
though in the latter the retractor muscle (B, rv) arises in the base 
of the appendage, and the appendage itself is movable by two muscles 
(J, J) arising in the body and inserted on its base. In some of the 
Collembola each lobe of the vesicle is produced into a long eversible 
tube. 





No. 6 INSECT ABDOMEN—SNODGRASS 73 


The typical spring, or leaping organ of the Collembola, known as 
the furcula (fig. 30 D), has quite a different type of structure from 
that of the collophore. It consists of a large median base, the manu- 
brium (mn), and of two slender arms, each of which is subdivided 
into a long proximal segment, the dens (d), and a short terminal 
segment, or mucro (m). On the base of the manubrium are inserted 
flexor and extensor muscles arising in the fourth and third abdominal 
segments, but, as already observed, these muscles apparently belong 
to the system of longitudinal dorsal and ventral body muscles, and 
are not specifically muscles of the spring. In Tomocerus vulgaris 





Fic. 30.—Abdominal appendages of Collembola. Tomocerus vulgaris. 


A, lateral view of insect. B, anterior view of collophore. C, tenaculum. D 
furcula. 

a, aperture between bases of furcular arms receiving prongs of tenaculum; 
ab, abductor muscle; ad, adductor muscle; Col, collophore; d, dens; Fur, fur- 
cula; m, mucro; mn, manubrium; rv, retractor muscles of vesicle; v, terminal 
vesicle of collophore. 


(fig. 30) each of the arms of the furcula is provided with an abductor 
muscle (D, ab) and an adductor muscle (ad) having their origins in 
the manubrium. According to Quiel (1915) adductor muscles are 
absent in Orchesella cincta, though he says a few obliquely transverse 
fibers are present in the manubrium. It is possible that Quiel did 
not observe in studying sections that these transverse fibers are 
attached on each side to a slender adductor tendon of the dens. The 
structure of the furcula readily suggests that it is composed of a 
pair of segmental appendages united by a fusion of the coxae, which 
become the manubrium, while the reduced telopodites become the 
arms. 


74 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


The minute tenaculum of the third abdominal segment (fig. 30 C) 
looks like a miniature furcula, and likewise suggests that it has been 
produced in the same manner by the union of the bases of a pair of 
appendages. 


THE ABDOMINAL APPENDAGES OF THYSANURA 


It is in the Thysanura that the abdominal appendages best preserve 
the fundamental structure characteristic of the abdominal appendages 
of adult Pterygota, as shown in the gonopods of the latter group ; but, 
as will be seen later, it appears that the abdominal appendages have 
a more primitive form in the larvae of Ephemerida and in the larvae 





Fic. 31.—Diagrams of structure of abdominal appendages of Thysanura and 
Pterygota. 


A, a typical pregenital appendage. B, a gonopod, or genital appendage. 

gmcls, muscles of gonapophysis; Gon, gonapophysis; LB, limb basis, usually 
a lobe or plate of body wall; rvs, retractor muscles of eversible vesicle; smcls, 
muscles of stylus; Stn, sternum; Sty, stylus; /’s, eversible and retractile vesicle. 


of certain holometabolous insects: The thysanuran appendages retain 
most completely their independence in the Machilidae. 

The appendages of the pregenital region of the abdomen are 
typically developed on each of the pregenital segments except 
the first in Machilidae (fig. 4, II, VI), and those of each pair are 
distinct from the small median sternal plate (Sin), though their 
bases (LB, LB) are ankylosed with the latter, and are united medi- 
ally with each other. Each appendage consists of a broad basal plaie 
(fig. 31 A, LB), of a stylus (Sty) borne by the distal free margin of 
the basal plate, and of an eversible sac, or vesicle (V's), lying mesad of 
the stylus and retractile into the basal plate (fig. 4, II, Vs). The 
posterior part of the basal plate projects from the ventral wall of the 


NO. 6 INSECT ABDOMEN—SNODGRASS 75 


abdomen as a free flap with a membranous dorsal wall (fig. 32 B). 
Both the stylus and the eversible sac are provided with muscles arising 
within the basal plate (figs. 31 A, 32 B, smcls, rvs). The muscles of 
the stylus are inserted on the base of the stylus; the muscles of the 
vesicle traverse the latter, when the vesicle is everted, to be inserted 
within its distal extremity. 

The appendages of the pregenital segments are never developed 
into any other form in the Thysanura than that which they have 
in the Machilidae, but they may be variously reduced, or united with 
the sternum. The styli and eversible sacs are sometimes absent, or 
either organ may occur alone (fig. 4, I, VIII). A pair of vesicles 
is frequently present on each basal plate, but the stylus never occurs 





Fic. 32.—Appendages of Thysanura and Dicellura. 


A, base of metathoracic leg of Nesomachilis maoricus, showing styliform spur 
on coxa. B, typical structure of a pregenital appendage of same, dorsal view. 
C, posterior lateral part of sternum of Heterojapyx gallardi with united limb 
basis bearing the stylus. 


in duplicate. In Lepismatidae the basal plates of the appendages in 
each segment are fused with the primary sternum to form a large 
zygosternum, which is the definitive sternal plate of the segment. 
The same is true in the Dicellura, though the regions of the limb 
bases may remain partially separated from the region of the primary 
sternum (fig. 32 C). 

The basal plates of the thysanuran appendages are commonly called 
“coxae”’ (or “coxites”) by American entomologists, while certain 
European entomologists call them “ subcoxae.’’ The idea that the 
plates are coxae is based chiefly on the fact that in the Machilidae 
stylus-like spurs occur on the coxae of the second and third pairs of 
thoracic legs (fig. 32 A, Sty?), which appear to be homologues of 
the abdominal styli. The question of the possible identity of the 


76 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 85 


thoracic and abdominal styli will be discussed in the concluding part 
of this Section, but in any case the term “ coxite” should not be used 
to designate the stylus-bearing plates, because the word can properly 
mean only “a part of a coxa.” Since the writer is inclined to believe 
that the abdominal plates in question represent both the coxae and 
the subcoxae of the thoracic appendages, they are here termed simply 
the limb bases, or basal plates. 

The gonopods, or appendages of the genital segments, have the 
same structure as the pregenital limbs in Thysanura, with the excep- 
tions that they always lack eversible vesicles, and that typically each 
bears a median genital process, or gonapophysis (fig. 31 B, Gon). 
Two pairs of gonapophyses are regularly present in the females of 





Fic. 33.—Genital and postgenital segments of Machilidae. 


A, Machilis variabilis, male, ventral plates of eee abdominal segment, dor- 
sal view, with small first gonapophyses (1Gon). B, ventral plates of ninth seg- 
ment of same, dorsal view, showing second pair of gonapophyses (2Gon) at 
sides of median penis (Pen). CG, Nesomachilis maoricus, male, posterior part of 
abdomen, ventral view, showing absence of gonapophyses, and ventral structure 
a taleil segment, bearing paraprocts (Papt), caudal filament (cf), and cerci 
(Cer 


Thysanura, one pair borne by the gonopods of the eighth segment, the 
other by the gonopods of the ninth segment. In the male, gonapophy- 
ses are known to occur on the gonopods of the eighth segment only 
in certain forms of Machilis (fig. 33 A, 1Gon); they are usually 
present on the gonopods of the ninth segment (B, 2Gon), but they 
may be absent from both genital segments (C). Each gonapophysis, 
when present, arises from the median, basal angle of the free dorsal 
surface of the stylus-bearing plate, and is provided with short muscles 
(fig. 31 B, gmcls) arising within the supporting plate and inserted 
on its base. 

Between the bases of the gonopods of the ninth segment in the male 
is a short membranous penis (fig. 33 B, C, Pen), a tubular evagination 





No. 6 INSECT ABDOMEN—SNODGRASS 77 


of the body wall from behind the region of the ninth sternum, having 
the opening of the ejaculatory duct at its extremity. 

The thysanuran gonopods contain, in their simplest form, the 
fundamental elements of the organs of copulation and oviposition of 
pterygote insects. In the male the gonapophyses of the second gono- 
pods become the so-called parameres of the copulatory apparatus ; in 
the female, the first and second gonapophyses become the first and 
second valvulae of the ovipositor, and the basal plates of the second 
gonopods form the third valvulae, when the last are present. 

The uropods, or cerci, of the Thysanura are typically long, multi- 
articulate filaments (fig. 7 A, Cer) borne by the eleventh segment 
(B, XJ). In the Dicellura the abdomen contains only 10 segments, 
and the uropods, therefore, in this group appear to belong to the 
tenth segment. They are filamentous in Campodea, styliform in 
Projapygidae, and take the shape of large pinchers in Japygidae (fig. 
40 C, Cer). The uropods differ from the preceding appendages in 
that they are not differentiated into a basal plate and a stylus, and 
they bear neither eversible sacs nor processes corresponding with 
the gonapophyses. 


THE ABDOMINAL GILLS OF EPHEMERID LARVAE 


The abdominal gills of ephemerid larvae, together with the lateral 
lobes of the body wall supporting them (fig. 34 A), appear to be ap- 
pendages of a more primitive form than the abdominal appendages 
of the Thysanura. Their structure, with certain modifications, is 
repeated in the abdominal appendages of several groups of holometab- 
olous larvae; but it does not furnish the basis of the structure of 
the gonopods in adult pterygote insects, which, as already stated, 
is to be derived from that of the gonopods of the Thysanura. 

The ephemerid larval gills have various shapes, some being taper- 
ing stalks, either single or double, fringed with filaments (fig. 34 A, B, 
Brn), while others are expanded into broad plates ; but, whatever the 
form, each organ is movably attached by its base to a large lateral 
lobe of the body wall (LB). The gill is movable by muscles arising 
in the ventral part of the supporting lobe (A, B, bmcls). 

The gill-bearing lobes are not movable, since there are no body 
muscles inserted upon them. The vertical lateral muscles of the 
abdominal segments extend from the tergum to the edge of the ster- 
num mesad of the gill lobes (fig. 15 A, 7/, 21, 3l, fig. 34 A, 1). The gill- 
bearing lobes therefore have the character of limb bases implanted 
in the pleural areas of the segments between the tergal and sternal 


78 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


plates (compare LB of fig. 34 A with LB of fig. 1 A). The lateral 
and ventral surfaces of each lobe are often separated by a sharp fold 
or ridge (fig. 34 B, LB), and the ventral part of the lobe may be 
more or less united with the sternum of its segment (Stn). The gill 
(Brn) arises from the posterior end of the lateral surface of the 
supporting lobe, and is usually provided with four muscles inserted 
on its base (C). The branchial muscles, as above noted, take their 
origins on the ventral plate of the supporting lobe (B), and therefore 





Fic. 34—The abdominal gill-bearing appendages of ephemerid larvae. 


A, diagrammatic cross section of abdominal segment, showing limb base lobes 
(LB) bearing gill appendages (Brn), which are evidently the styli, each mov- 
able by muscles (bmcls) arising in the bases; the lateral body muscles (/) 
extend from tergum to sternum. 

B, external view of a gill and its supporting lobe (LB), in which arise the 
gill muscles (bmcls). 

C, musculature and tracheation of a gill, lateral view. 


not on the sternum, as stated by Dtirken (1907), who did not dis- 
tinguish the ventral plate of the gill basis from the true segmental 
sternum. 

The old idea that the ephemerid larval gills are appendages of the 
dorsum is no longer tenable. The organs are very evidently rudiments 
of segmental appendages, as claimed by Heymons (18 96a, 1896) 
and by Borner (1909), each being composed of a basis (LB), and 
of a terminal appendicular part (Brn) movable on the basis by mus- 


NO. 6 INSECT ABDOMEN—SNODGRASS 79 


cles arising in the latter. The gill bases are interpreted by Borner as 
the subcoxae of the abdominal appendages, since they follow exactly 
in line with the subcoxal, or “ pleural,” plates of the thorax. The 
gills, therefore, Borner contends, are the equivalents of the legs, 
and the proximal end of each represents the coxa. On the other 
hand, we might assume that the gill basis includes the equivalents 
of both the subcoxa and the coxa of a thoracic leg, in which case the 
gill shaft or plate might be supposed to be the telopodite with its 
proximal end representing the first trochanter. The presence of the gill 
muscles arising in the gill basis suggests this homology, since the 
muscles are comparable with the trochanteral muscles of a leg (fig. 
26, O, Q). But again, the gill resembles the stylus of a thysanuran 
appendage, and there is doubt as to whether the abdominal styli 
are true telopodites or secondary appendicular processes of the coxae. 

Leaving aside, for the present, the question of homologies between 
the parts of the abdominal appendages and those of the thoracic 
appendages, a comparison of the abdominal appendages of the ephem- 
erid larva with the abdominal appendages of Thysanura leaves 
little doubt that the gill-supporting lobes of the former are the 
equivalents of the stylus-bearing plates of the latter, and that the 
gill stalks or plates are the homologues of the styli. Neither the 
abdominal gills of ephemerid larvae nor the styli of Lepisma are 
present on the early postembryonic stage of the insect. In the newly 
hatched larva of Ephemerella vulgata, according to Heymons (1896), 
there are lateral protuberances of the abdominal segments, especially 
prominent on segments II and VII, which are derived during em- 
bryonic development from the embryonic limb rudiments. The giils 
first appear as outgrowths from these abdominal lobes about four 
days after hatching. The styli of Lepisma saccharina, Heymons 
(1897) says, appear likewise a considerable time after hatching, and 
arise from the parts of the ventral plates of the eighth and ninth 
abdominal segments that are derived from the “ Anlagen” of the 
embryonic appendages. 


LATERAL ABDOMINAL APPENDAGES OF SIALID AND COLEOPTEROUS 
LARVAE 


The larva of Sialis (fig. 35 A) presents at least an excellent imita- 
tion of an insect that has carried the primitive polypod condition 
into a postembryonic stage. The long, tapering, segmented, ap- 
pendicular organs, usually termed “ gills,’ projecting from the sides 
of the first seven abdominal segments have a striking resemblance to 


8o SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


legs. Each appendage (C) is a hollow process of the body wall, dis- 
tinctly jointed, and composed of six segments, of which the proximal 
three are relatively thick, while the region of the distal three is slender 
and rapidly tapering to the apex. 

Each appendage is supported on a lateral lobe of the body segment 
(fig. 35 C, LB). The series of lobes appears at first glance to belong 
to the dorsum of the abdomen, but each one, though so closely amal- 
gamated with the tergum that the spiracle appears to be situated on 
its dorsal part, really occupies a pleural position between the tergum 





Fic. 35——Abdominal appendages of sialid larvae. 


A, larva of Sialis, showing jointed appendages of abdomen. B, metathoracic 
leg of Sialis larva, with subcoxal sclerites at its base. C, abdominal appendage of 
Sialis larva, showing division into basis (LB) and jointed stylus (Sty). D, 
posterior end of larva of Chauliodes, dorsal view, showing pygopods (Pp) of 
terminal segment. E, the same, ventral view. 


and the sternum of its segment. The lumen of each lobe is separated 
from the general body cavity by a vertical sheet of tergo-sternal 
lateral muscles, and within the lobe arise anteriorly and posteriorly 
muscles inserted on the base of the movable shaft of the appendage. 
It is clear, therefore, that we have here an organ corresponding 
in every respect with a gill-bearing appendage of an ephemerid larva 
(fig. 34 B), and that in both of these structures the basal lobe repre- 
sents the stylus-bearing plate of Machilis, and the movable distal 
appendage the stylus. 





no. 6 INSECT ABDOMEN—SNODGRASS 81 


The distinct segmentation of the abdominal appendages of the 
Sialis larva almost unavoidably gives the impression that these organs 
are the true telopodites of the abdominal limbs. The impression, 
moreover, comes close to a conviction when it is discovered that each 
appendage is provided with internal muscles in addition to those in- 
serted on its base. The presence of these muscles was first mentioned 
by Heymons (1896a) ; and it can be demonstrated by dissection and 
staining of specimens preserved in alcohol that bundles of muscle 
fibers are present in at least each of the first three segments, inserted 
on the bases of the second, third, and fourth segments, but it is diffi- 
cult to make an exact study of them without properly prepared ma- 
terial. The slender distal part of the shaft beyond the third segment 
is penetrated by a branched trachea, and this part of the appendage 
might serve as a tracheal gill ; but the strong musculature of the proxi- 
mal part of the organ, and the long hairs that fringe the segments, 
suggest that the abdominal appendages of the Sialis larva have an 
important locomotory function. 

In the sialid genera Chauliodes and Corydalus the larvae are like- 
wise provided with long, lateral abdominal appendages, a pair being 
present on each of the first eight segments, and a terminal pair on the 
last segment (fig. 36B). In these genera, however, the appendages 
are simple, tapering, hollow processes of the integument, unsegmented, 
and containing no muscles. Each is supported on a lateral lobe of the 
body wall (LB). 

The basal lobes of the lateral appendages of the Corydalus larva 
are large and prominent as seen in a transverse section of an abdominal 
segment (fig. 30C, LB, LB). Each projects laterally beyond the at- 
tachments of a set of strong tergo-sternal lateral body muscles (/) ; and 
on the inner margin of the ventral wall of the lobe arise three muscles 
(D, smcls), one anteriorly and two posteriorly, which are inserted on 
the base of the distal appendicular process. Here again, therefore, we 
find repeated the same structures that occur in the ephemerid larval 
gills and in the thysanuran abdominal appendages. In the Corydalus 
larva the appendage-bearing lobes of the abdomen fall in line with 
the subcoxal lobes of the thoracic segments (fig. 36 A, Scv), rather 
than with the long coxae (Cx); but the muscles of the abdominal 
appendages (D, smecls), taking their origins in the supporting lobes, 
can be compared only with the basal muscles of the leg telopodite 
(A, O, Q) inserted on the trochanter, The abdominal lobes, there- 
fore, would appear to contain both the subcoxal and the coxal parts 
of the limb bases. 


82 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


The Corydalus larva differs from the Sialis and Chauliodes larvae 
in that the basal lobes of the first seven abdominal appendages bear 
each a large ventral tubercle supporting a circle of respiratory fila- 
ments (fig. 36 B, Vs). When the filaments are removed it is seen that 
each tubercle is subdivided distally into three terminal lobes (C, Vs), 
and dissection reveals the fact that the tubercle is provided with a large 


Sty 





Frc. 36.—Thoracic and abdominal appendages of the larva of Corydalus 
cornutus. 


A, transverse section through anterior end of metathorax, showing subcoxal 
lobes (Scx) above bases of coxae (Cx), and levator and depressor muscles of 
trochanter (O, Q) arising in coxa. 

B, last four segments of abdomen, showing segmental appendages. 

C, transverse section through posterior part of an abdominal segment, seen 
from behind, showing limb base lobes (LB) supporting each a stylus (Sty), 
and a retractile gill-bearing tubercle (I’s). 

D, section through base of right abdominal appendage and gill tubercle, seen 
from behind, showing muscles of stylus and gill tubercle. 

FE, antero-mesal view of right appendage (pygopod) of tenth abdominal 
segment. 

F, section of terminal appendage, showing insertion of retractor muscles 
behind bases of claws. 

Cx, coxa; d, claws (crochets); e¢, insertion point of retractor muscles of 
claw-bearing tubercle (planta); /, lateral body muscles; LB, limb basis; O, 
QO, muscles of trochanter; rvs, retractor muscle of gill-bearing or claw- 
bearing lobe of appendage; Sc, subcoxa; smcls, muscles of stylus; Sp, spiracle; 
Stn, sternum; Sty, stylus; 7, tergum; /’s, retractile lobe of appendage (retrac- 
tile vesicle). 


retractor muscle (C, D, rvs) arising on the dorsum of the segment and 
inserted by three diverging branches on the distal surfaces of the three 
terminal lobes of the tubercle. The gill tubercles are thus highly 
suggestive of the eversible sacs of the thysanuran appendages, except 
for the difference that their retractor muscles arise on the tergal 





CO 


no. 6 INSECT ABDOMEN—SNODGRASS 3 
region of the body segment, instead of in the bases of the appendages, 
as in the Thysanura (fig. 32 B, rvs). On the other hand, as we shail 
presently see, the gill tubercles of the Corydalus larvae are almost 
identical in structure with the abdominal feet of lepidopterous larvae. 

The appendages of the last abdominal segment in the larva of 
Chauliodes and Corydalus are remarkable structures in that they com- 
bine the characters of the preceding appendages of Corydalus with the 
features of an abdominal leg of a caterpillar. Each of these terminal 
appendages (figs. 35 D, Pp, 36 B, E) consists of a large, hollow, 
somewhat cylindrical lobe of the body wall. The basal part of the 
organ (fig. 30B, E, LB) bears laterally a tapering process (Sty) 
similar to the lateral processes of the preceding appendages, and ends 
distally in a thick tubercle (Vs), which lacks gill filaments, but is 
armed on its terminal surface with a pair of large, curved claws (d). 
The appendage is traversed by a strong retractor muscle (F, rvs) 
taking its origin on the dorsum of the tenth segment, and having 
its insertion on the distal wall of the tubercle at the posterior ends 
of the long bases of the claws (e). The resemblance in structure 
and mechanism of these appendages to the “ anal” legs of caterpillars 
is so striking that it is difficult to believe the likeness is fortuitous. 
The terminal body segment is better developed in the Corydalus larva 
(fig. 36B, X) than in Chauliodes (fig. 35 E), in which its dorsal 
part is rudimentary. 

Lateral appendicular processes of the abdomen, similar in every 
respect to those of the sialid larvae, are present also on certain aquatic 
coleopterous larvae, especially in the families Dytiscidae and Gyrin- 
idae. In the gyrinid Dineutes, for example, the larva is provided with 
long, tapering processes arising from lateral lobes of the body wali 
on each side of each of the first eight segments, and with a pair of 
two-branched processes on the ninth segment. Each process is pene- 
trated by a trachea from the lateral respiratory trunk, and is fur- 
nished with two short, antagonistic muscles arising in the supporting 
lobe of the body and inserted on its base. The larva of the dytiscid 
Coptotomus has the same equipment of lateral processes, but the writer 
did not find muscles connected with them in a specimen examined. 


THE ABDOMINAL LEGS OF LEPIDOPTEROUS LARVAE 


A typical abdominal leg of a caterpillar consists of three parts 
(fig. 37 A). At the base is a ring of flexible integument (mb) ; be- 
yond this is a longer, cylindrical section (Cx) forming the greater part 
of the appendage, and frequently having a sclerotic plate in its outer 

6 


7 


84 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 85 


wall often marked by distinctive groups of setae (fig. 3 A) ; distally 
the leg ends in a retractile lobe (Vs), called the planta, which bears 
the claws, or crochets (d). 

Functionally the planta is the most important part of the leg, and 
structurally it is the most variable. The proximal parts of the ap- 
pendage differ principally in relative size in different species. The 
planta in its more generalized condition is a short cylindrical pad with 
a circular distal surface, on the center of which is inserted a group 





Fic. 37.—Structure of an abdominal leg of a caterpillar. Malacosoma 
americana. 


A, posterior view of a pair of abdominal legs. B, ventral view of a left ab- 
dominal leg, showing crochets (d) turned outward, and insertion of retractor 
muscles (¢) at inner margin of plantar lobe (Vs). C, dorsal view of leg mus- 
culature, right side. D, posterior view of musculature of a right leg. 

a, dorso-pleural groove; Cx, coxa; d, crochets; e, insertion point of retractor 
muscles of planta; mb, membranous area between subcoxa and coxa; rvs, 
retractor muscles of planta; Scx, subcoxa; Vs, planta (retractile vesicle). 


of retractor muscle fibers. In such cases the crochets may be arranged 
in a complete circle around the distal plantar surface, with their re- 
curved points turned peripherally and upward. With most cater- 
pillars, however, the claws are limited to a semicircle or a small arc 
usually on the inner margin of the planta (fig. 37 B), and in such cases 
the planta itself (Vs) generally becomes asymmetrical by a reduction 
or obliteration of its outer half. The planta then assumes the form 
of a lobe projecting to the mesal side of the axis of the limb, marked 
by the insertion point of the retractor muscle (e), and its claws (d) 


No. 6 INSECT ABDOMEN—SNODGRASS 85 


curve mesally and upward when the planta is protracted in the usual 
position. The various types of foot structures resulting from modi- 
fications in the form of the planta and in the arrangement of tie 
crochets characteristic of the different groups of caterpillars have 
been described by Fracker (1915). 

Immediately above each abdominal leg there is usually a prominent 
lobe or swelling of the body wall (fig. 37 A, Scv), separated from the 
latero-dorsal area of the segment by a distinct groove (a). Corre- 
sponding lobes are present also on the legless abdominal segments, and 
likewise on the metathorax and mesothorax (fig. 3 A, Scr). The 
serial identity of these suprapedal lobes of the abdomen and thorax 
is demonstrated by their uniform position relative to the appendages, 
and by the fact that in many species they bear similar or identical 
marks or groups of setae. In the anatomy of the caterpillar, therefore, 
the abdominal and thoracic appendages appear to be homodynamous 
structures. Eastham (1930), in his study of the embryology of 
Pieris rapae, says: “The prolegs which are retained on their seg- 
ments must be regarded as true appendages. They develop in the same 
manner as those of the head and thorax, have the same relation to their 
own somites, and a musculature develops in connection with each com- 
parable to that of the thoracic limbs though of a weaker order.” 

The suprapedal lobes of the caterpillar (fig. 3 A, Scx) are clearly 
the subcoxal areas of the appendages, since those of the thorax are 
identical with the areas which in certain other holometabolous larvae 
contain the pleural sclerites of the thorax (B, C, Scv3). The free 
part of the abdominal appendage in the caterpillar is, therefore, ap- 
parently the coxa (fig. 37 A, Cx). The planta (Vs), then, is either 
a rudiment of the telopodite, or a highly specialized retractile vesicle 
of the coxa. Further light on the morphology of the caterpillar proleg 
may be obtained from a study of the musculature. 

The musculature of an abdominal leg of a caterpillar is compara- 
tively simple. It comprises two sets of muscles, those of one set 
being inserted on the base of the principal part of the leg (fig. 37 D, 
Cx), those of the other on the planta (Vs). The muscles inserted on 
the proximal rim of the leg include three groups of fibers represented 
in Malacosoma americana and Estigmene acrea as follows: (1) a 
series of median fibers (fig. 21 B, 37 C, D, r) arising on the midline 
of the venter, or also on the mesal parts of the anterior and posterior 
intersegmental folds, and converging to the mesal rim of the base of 
the principal segment (Cx) of the leg; (2) a group of two fibers (fig. 
37 C, D, 2) arising on the groove (a) above the suprapedal lobe of 
the body wall, and inserted on the mesal rim of the leg base posteriorly 


86 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


just ventrad of the insertions of the median muscles; (3) a group 
of two or three fibers (C, D, 3) taking their origins on the middle of 
the lateral wall of the segment posterior to and a little above the level 
of the spiracle (fig. 21 B), and having their insertions on the outer side 
of the proximal rim of the leg (figs. 21 B, 37 C, D, 3). 

The muscles of the planta take their origins outside the leg from 
two widely separated parts of the body wall. The plantar muscles 
of Malacosoma americana (fig. 37 C, D) comprise four fibers. The 
principal one is a long fiber (4) arising on the middle of the lateral 
wall of the segment, close to the origins of the lateral muscles of 
the leg (3), from which point it curves downward into the leg to be 
inserted on the inner face of the planta. The other plantar muscles 
arise on the dorso-pleural groove (a) above the suprapedal lobe (fig. 
37 D, Scx). One consists of a single fiber (C, D, 5) arising pos- 
teriorly just before 2, and entering the leg with 4. The other in- 
cludes two fibers (6) in Malacosoma, represented by a single fiber 
in Estigmene (fig. 21 B, 6), arising anteriorly on the dorso-pleural 
groove, and curving posteriorly and downward into the leg to join 
with 4 and 5. In the distal part of the leg (fig. 37 D) all the fibers 
of the plantar group unite to form a common stalk which is inserted 
on the inner surface of the ventral wall of the planta. In caterpillars 
having a disk-shaped planta, the muscle insertion is at or near the 
center of the latter, but with species in which the planta has the form 
of a mesal lobe, the muscle attachment is at the outer side of the 
plantar lobe (fig. 37 B, D, e). 

On comparing, in the caterpillar, the musculature of an abdominal 
leg with that of a thoracic leg, it is found that though there is ne 
exact correspondence in the number and arrangement of the fibers, 
there is a general similarity in the disposition of the muscles sufficient 
to suggest a derivation of the muscles in the two cases from one 
fundamental plan of musculature. Thus, in the metathorax of Mala- 
cosoma (fig. 38 A, B) there is a set of sternal fibers (a) arising an- 
teriorly on the intersegmental fold, and inserted mesally on the rim of 
the coxa (Cx), which correspond with the median muscles of an ab- 
dominal leg (fig. 37 C, D, 1). Likewise, there are muscles from the | 
lateral wall of the thoracic segment inserted on the outer rim of the | 
coxa (fig. 38 B, C, b), having thus the same relation to the appendage | 
as the fibers of muscle 3 in the abdomen (fig. 37C, D). In the | 
thorax there are several subcoxo-coxal muscles (fig. 38 B, c) which | 
have no exact counterparts in the abdomen, though in the latter there | 
is a muscle from the groove above the subcoxal lobe (fig. 37 C, D, 2) 
to the inner margin of the apparent coxal segment of the leg. The — 





SNODGRASS 87 


No. 6 INSECT ABDOMEN 7 


other muscles in the base of a thoracic leg (fig. 38 B, C, d, e, f, g) 
are coxo-trochanteral and coxo-femoral muscles, representatives 0} 
which are entirely absent in the abdominal legs. On the other hand, 
the plantar muscles of the abdomen have no evident counterparts in 
the thorax. 

The general parallelism between the muscles of the abdominal ap- 
pendages and those of the thoracic legs shows that the musculature 
of the prolegs in the mature caterpillar is, as Eastham says of the 
comparable to that of the thoracic limbs 
Moreover, if the musculature has any 


ce 


musculature in the embrvo, 


though of a weaker order.” 


c 


Wy 


1 CW ee ace 
¢ SS” mM i : 
; ; a G/, 


} 
[ 
ot — 


q caine % | Sap 





Fic. 38.—Body muscles of a thoracic leg of a caterpillar. 


A, external body muscles and leg muscles of ventral area and right half of 
metathorax of Malacosoma americana. B, basal muscles of right metathoracic 
leg of same. C, coxal and coxo-trochanteral muscles of right mesothoracic leg 
of Estigmene acraea. 


bearing on homologies in the segments of the appendages, it shows 
that the suprapedal lobes of the body wall above the abdominal limbs 
are the subcoxae (fig. 37 A, D, Scv), and that the principal parts 
of the legs are the coxae (Cx). The plantae of the abdominal ap- 
pendages, however, have no evident homologues in the thoracic legs, 
nor do their muscles correspond with any of the muscles of the legs 
in the thorax. 

The nature of the planta becomes clear when we compare an ab- 
dominal limb of a caterpillar (fig. 37 D) with an abdominal appendage 
of the Corydalus larva (fig. 36C, D). A striking resemblance is 


88 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 85 





seen at once between the retractile planta of the former and the gill- 
bearing tubercle (Vs) of the latter. Moreover, the likeness amounts 
almost to a structural identity when the planta is compared with the 
claw-bearing lobe of one of the terminal appendages of the Cory- 
dalus larva (fig. 36 E, Vs). In each case the organ is provided with 
strong retractor muscles arising on the dorsum of the body segment. 
It is a mere detail that the muscles in Corydalus (fig. 36 D, rvs) branch 
to the several lobes of the gill tubercle; in the terminal appendages 
the bundle of retractor fibers (F, rvs) tapers to a narrow stalk inserted 
at the bases of the claws exactly as in the foot of the caterpillar (fig. 
37 D). The Corydalus larva lacks only the accessory muscles of the 
planta arising in the base of the subcoxa. We can, therefore, scarcely 
avoid the conclusion that the planta of the caterpillar’s abdominal 
leg is an eversible vesicle of the limb basis, here borne by the coxa, 
which is not consolidated with the subcoxa. The representatives of 
the styli, preserved in the movable distal processes of the sialid ap- 
pendages (fig. 36 C, D, E, Sty), have been lost from the abdominal 
limbs of all lepidopterous larvae. 

The anal legs, or appendages of the last abdominal segment of the 
caterpillar, differ from the legs of the preceding abdominal segments 
only in details of their musculature. Their structure will be described _ 
in the subsequent discussion of the terminal appendages of holometab- 
olous larvae. 

The abdominal appendages of chalastogastrous hymenopterous 
larvae have the same essential structure as the abdominal limbs of 
caterpillars, though they are not so highly organized, and the plantar 
lobes are reduced to terminal disks of the coxal segments (fig. 3 C). 


THE GONOPODS 


The appendages of the eighth and ninth segments of the abdomen 
are potentially gonopods because of the association of the openings 
of the genital ducts with these segments. They are, however, not 
necessarily modified for reproductive purposes, as in the males of cer- 
tain Thysanura in which the appendages of the eighth and ninth 
segments (fig. 33 C) do not differ structurally from those of the 
preceding segments. 

Typically a gonopod is distinguished from the pregenital and post- 
genital appendages by the development of a median process from the 
proximal part of its base. A complete gonopod, therefore, consists 
of a basal lobe or plate (fig. 31 B, LB), of a lateral, distal stylus (Sty), 
and of a median, proximal gonapophysis (Gon). Both the stylus and 


no. 6 INSECT ABDOMEN—SNODGRASS 89 
the gonapophysis may be movable on the basis by muscles arising in 
the latter and inserted on their bases (smcls, gmcls). So far as has 
been ascertained neither of the appendicular parts of a gonopod ever 
contains intrinsic muscles. In the males of some Ephemerida the 
genital styli are distinctly jointed and thus separated into apparent 
segments, though the latter are not independently movable. In the 
Thysanura the gonapophyses are marked by circular constrictions, 
varying from a few to many according to the length of the organs 
(fig. 33 A, B), but the resulting subdivisions have none of the charac- 
ters of true segments, and are entirely comparable with the annulations 
of the caudal filament and cerci (C; cf, Cer). 

In female insects the gonopods form the ovipositor, when this organ 
is present, and both pairs of appendages enter into its composition. 
In the Thysanura the gonapophyses only are involved in the ovipositor, 
the basal plates and the styli retaining the structure typical of these 
parts in the pregenital segments of Machilidae. Evidently the condi- 
tion here represents a primitive stage, in which two pairs of median 
apophyses of the appendages of the eighth and ninth abdominal seg- 
ments form a simple egg-laying organ. In female Pterygota the styli 
of the gonopods are usually lost, those of the first pair being always 
absent, and the basal plates are transformed into a suspensory appara- 
tus for the gonapophyses. The basal plates of the first gonopods (fig. 
39 A) evidently become the small sclerites known as the valvifers 
(B, Vif), which support the first gonapophyses (1//1), though there 
is a difference of opinion on this point. Those of the second gonopods 
form lobes (JX LB) supporting the second gonapophyses (2V1), or 
they are drawn out into long processes that become a third pair of 
blades in the ovipositor (C, 37). The component blades of the adult 
ovipositor are commonly called valvulae. It is to be observed that only 
the first and second pairs of valvulae (B, C, rVl, 2V1) represent the 
gonapophyses (A, 1Gon, 2Gon), those of the third pair (C, 3/1) 
being derived directly from the basal plates of the second gonopods, 
the styli of which are apparently lost. 

The ovipositor is absent or rudimentary in many groups of insects, 
but its wide distribution throughout the orders leaves little doubt 
of its being a primitive structure of the Insecta. It has no homo- 
logue in other Arthropoda, and it is doubtful if the rami of the gono- 
pods in the Crustacea are homodynamous with the gonapophyses 
of insects. The various theories concerning the possible homologies 
of the genital processes of the gonopods in insects will be considered 
in the closing discussion of this paper. 


F 


go SMITHSONIAN MISCELLANEOUS COLLECTIONS — VOL. 85 


In male insects the history of the gonopods is much more involved 
than in the female, and the evolution of the genital appendages into 
organs of copulation has produced many different kinds of struc- 
tures. It is only in certain species of Machilis that both pairs of gono- 





VIII LB (VIF) XLB Sty D 
\ 


Vig. 





N \ 
G iVI 2V1 3V1 


Fic. 39.—Diagrams showing the morphology of the ovipositor, and the exter- 
nal genitalia of the male. 


A, female genital segments and generalized structure of the gonopods. B, an 
ovipositor with two pairs of valvulae formed of the gonapophyses. C, an ovi- 
positor with three pairs of valvulae, the third pair (3/1) formed of the basal 
plates of second pair of gonopods (A, LB). 

D, generalized structure of ventral parts of ninth abdominal segment of male, 
showing sternum (Stn), gonopods (Gp, Gp), and penis (Pen). E, specialized 
structure of male genitalia; basal plates of gonopods (LB, LB) united by pons 
basalis (pnb), parameres united with penis to form an aedeagus (ded), and 
styli (Sty) transformed into clasping lobes. 

Aed, aedeagus; Gon, gonapophysis; 1Gon, 2Gon, gonapophyses of first and 
second gonopods; LB, basal plate of gonopod; Ovd, oviduct; Pm, parameres 
(male gonapophyses) ; pub, pons basalis; smcls, muscles of stylus; Stn, ster- 
num; Sty, stylus; 7, tergum, rV1, 2V1, 3V1, first, second and third valvulae; 
Vif, valvifer. 


pods bear gonapophyses (fig. 33 A, B, 1Gon, 2Gon). In all male 
pterygote insects gonapophyses are lacking on the eighth segment, and 
the gonopods of the ninth segment only enter into the copulatory 
apparatus, though accessory structures of the eighth and the tenth 
segment may be included. 





No. 6 INSECT ABDOMEN—SNODGRASS gI 


The basal plates of the second gonopods in the male may unite with 
each other and with the primary sternum of their segment, as do those 
of the preceding segments, to form a composite zygosternum, and in 
such cases the styli either retain the typical shape of styliform organs, 
or they are lost. The gonapophyses, usually termed the parameres, 
however, in the more generalized insects, are associated with a median 
intromittent organ, or penis (figs. 33 B, 39 D, Pen), which is a tubular 
evagination of the segmental wall behind the ninth sternum, bearing 
the opening of the ejaculatory duct at its extremity. In the higher 
insects the primitive penis becomes partly or entirely suppressed, and 
the parameres unite with it or with each other to form the secondary 
and often more complicated intromittent organ usually termed the 
aedeagus (fig. 39 E, Aed), which incloses the ejaculatory duct and 
bears the gonopore. 

The basal plates of the gonopods of the ninth segment in the male, 
if not completely amalgamated with the sternum, may form free lobes 
of the ninth segment, or they may unite with each other, with the 
sternum, or with the sternum and the tergum of the ninth segment. 
In this way the genital segment of the male, especially in holometa- 
bolous insects, assumes a great diversity of structure, and it is often 
reduced to a simple continuously sclerotized annulus. The ninth seg- 
ment, however, regardless of its form, always bears the aedeagus, 
which may be partly or wholly concealed in a genital chamber of its 
ventral part, and it generally carries clasping organs of various forms 
on its posterior margin. Usually, among the clasping organs of the 
ninth segment, or often the only structures having a clasping func- 
tion, is a pair of lobes flexible at their bases and independently mov- 
able by muscles taking their origins in the basal plates of the gonopods, 
or in the regions of the ninth annulus derived from the gonopod bases. 
These movable claspers, designated the harpes by students of Lepi- 
doptera, are evidently the homologues of the styli of the more general- 
ized insects (fig. 39 D, E, Sty). 

It is most important, now, to observe that in the fundamental or- 
ganization of the gonopods there are only two sets of appendicular 
structures that are independently movable by muscles inserted directly 
on their bases. These structures are the styli and the gonapophyses. 
Therefore, in the ninth segment complex of the male genitalia, there 
will generally be two sets of appendicular structures, the harpes and 
the parameres, provided with muscles arising in the basal plates of 
the gonopods, or in the parts of the ninth segmental ring derived 
from the latter. By a study of the genital musculature, then, thesc 
two structures can be identified with certainty in almost all cases 


92 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


throughout the orders, unless one or the other or both are absent. 
In addition to these fundamental, musculated processes, however, 
there are innumerable other secondary genital processes having no 
necessary homology in the different orders, which may be developed on 
the ninth segment, on the aedeagus, or on the segments preceding 
and following the ninth. These structures, except in rare cases, have 
no muscles of their own, and are not independently movable, though 
some of them may be moved incidentally by the usual segmental mus- 
cles attached at their bases. The movable claspers derived from the 
styli, however, are sometimes divided, and each may be separated into 
two quite distinct parts provided individually with groups of muscle 
fibers. In such cases there will appear to be, as in some of the Hy- 
menoptera, a pair of movable lobes on each side of the genital 
apparatus. 

A more detailed analysis of the structure of the organs of ovi- 
position and copulation, as shown in the principal orders of insects, 
will form the subject matter of Part II of this paper, wherein will be 
presented also a larger body of evidence in support of some of the 
statements that seem arbitrary in the brief discussion given above. 


THE CERCI (UROPODS) 


The prevalence of cerci in so many orders of insects, and the almost 
universal occurrence of the organs in the more generalized groups 
leave little doubt that the cercal appendages are primitive structures, 
and that, in some form, they must have been characteristic features 
of the early insect ancestors. The anatomy and structural variations 
of the cerci are well known; functionally the appendages are in most 
cases sensory organs, though they are frequently modified in form 
to serve mechanical purposes; morphologically they are subjects of 
diverse opinion among speculative entomologists. The essential facts 
known concerning the cerci can be briefly stated. 

In the Thysanura the cerci evidently belong to the eleventh ab- 
dominal segment. The last typical segment of the body in such forms 
as Nesomachilis (fig. 7A) is the tenth (X), which is a complete 
annulus. From within the posterior margin of this segment there 
project the three terminal filaments, of which the lateral pair are 
the cerci (Cer). If the group of filaments is pulled out of the tenth 
segment, it is seen that the three of them arise from a common basal 
ring (B, XJ), which has all the aspects of a reduced segment, in this 
case the eleventh, normally concealed within the tenth. The eleventh 
annulus presents a wide dorsal region (C, XJT) prolonged into the 





No. 6 INSECT ABDOMEN—SNODGRASS 93 


median caudal filament (cf), a narrow ventral region (D, X/Stn) 
bearing a pair of broad posterior flaps (Papt), and two prominent 
lateral lobes supporting the cerci (Cer). There can be little doubt, 
therefore, that the caudal filament and the cerci here belong to the 
eleventh abdominal segment. The lobes of the eleventh sternum are 
evidently the paraprocts (D, Papt). Projecting from beneath the base 
of the caudal filament (D, cf) is a small median lobe (sa), possibiy 
a remnant of the true telson, represented by the lamina supra-analis of 
the twelfth segment, better developed in odonate larvae (fig. 12 A, sa). 

The terminal parts of the abdomen are less simple in some other 
thysanurans than they are in Nesomachilis. In Thermobia (fig. 7 E, 
F), for example, the eleventh segment is largely obliterated except 
for a distinct tergal plate, or epiproct (E, Eppt), which is connected 





Fic. 40.—The cerci and associated parts. 


A, posterior segments of Periplaneta orientalis, dorsal view, showing union of 
tenth and eleventh abdominal terga. B, ventral view of terminal segment of 
same. C, cerci of Heterojapyx gallardi, and muscles of tenth segment that move 
them. 


laterally by a pair of small sclerites with the bases of the cerci (Cer). 
The paraprocts of Thermobia are large sclerotic plates (F, Papt) 
supporting the cerci. They would thus appear to correspond with 
the lateral lobes of the eleventh segment and the median paraproct 
plates of Nesomachilis (D). A sternal region of the eleventh segment 
distinct from the paraprocts is not evident in Thermobia. 

In the Dicellura (Campodeidae and Japygidae) the abdominal 
segments beyond the tenth are obliterated, and the cerci are sup- 
ported directly by the end of the tenth segment (fig. 40 C, Cer). 

The cerci of the Pterygota most commonly appear to belong to the 
tenth abdominal segment, since they arise at the posterior edge of 
the latter, usually from membranous areas inclosed by the adjacent 
angles of the tenth tergum, the epiproct, and the paraprocts (fig. 12 A, 
B, Cer). In Periplaneta the cerci of the adult insect arise from be- 


94 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


tween the lateral angles of the last tergal plate and the upper 
angles of the paraprocts (fig. 40 A, B), but the terminal plate 
of the dorsum is here clearly a composite sclerite formed of the united 
tenth tergum (A, X) and the epiproct (XJ). Frequently the cerci 
are more closely associated with the paraprocts than with the tergal 
plates. In any case, however, the intermediate position of the cerci 
in adult Pterygota gives no positive evidence of the segmental rela- 
tions of these appendages in this group of insects. 

On the other hand, the ontogenetic evidence of the nature of the 
pterygote cerci seems to be quite definite, for it is stated by Ayers 
(1884), Cholodkowsky (1891), Wheeler (1893), and Heymons 
(1896a) that the cerci in the embryos of Orthoptera are formed 
directly from the appendages of the eleventh abdominal segment (figs. 
5 A, 9A, B, Cer). Heymons claims that the eleventh segment itself 
disappears from the adult abdomen, and that the cerci thus come 
to have an apparent intersegmental position between the tenth and 
the twelfth segments. As already shown, however, it appears more 
probable that the eleventh segment is usually represented in the 
adult by the epiproct and the paraprocts, and that it is the twelfth 
segment which is lacking, or reduced to a circumanal fold (fig. 12 A, 
Prpt). 

The association of the cerci with the upper basal angles of the 
paraprocts, or their actual connection with these plates in some cases, 
as in Thermobia (fig. 7 F), has given rise to the idea that the cerci 
and the paraprocts have a genetic relation to each other. Thus, 
Crampton (1920, 1921) contends that the paraproct is the base of a 
segmental appendage of which the cercus is the distal part. Accord- 
ing to Heymons (1896), on the other hand, the embryonic cercus 
represents the entire appendage of the eleventh segment, including the 
basis, which in the pregenital segments unites with the primary seg- 
mental sternum to become a lateral part of the definitive sternal plate. 
In the adult insect, Heymons says, the cercal base usually disappears 
as an evident lobe, though a rudiment of it is retained in young 
nymphs of Gryllus and Decticus as a small basal ring supporting the 
free part of the organ (fig. 8 B). 

The musculature of the cerci, so far as it is known, is always dorsal, 
there being no muscles from the sternal region of the abdomen or from 
the paraprocts in any way associated with the cerci. The origins of the 
muscles present, however, give no clew to the segmental relations of 
the cerci, since the muscles arise either on both the tenth and eleventh 
terga, or on the tenth tergum alone. In her study of the abdominal 
muscles of Orthoptera, Ford (1923) finds that each cercus is typically 





\ 
| 
j 
} 
f 
t 
| 
( 
\ 


no. 6 INSECT ABDOMEN—SNODGRASS 95 


provided with four muscles. Three of these muscles, distinguished 
as an abductor, a depressor, and an elevator muscle, take their origins 
on the tenth abdominal tergum; the fourth arises on the supra-anal 
plate or epiproct. The muscle from the epiproct, Ford says, is absent 
in Gryllotalpa, but it is present in Gryllus, Neoconocephalus, Ceutho- 
philus, and Melanoplus. In Gryllus, because of the union of the tenth 
tergum and the epiproct, this muscle, however, has shifted forward to 
the area of the tenth tergum. The writer has found only three mus- 
cles in Dissosteira inserted directly on the base of the cercus, two 
arising on the tenth tergum, and one on the epiproct. 

The origin of the anterior muscles of the cerci on the tenth segment 
might be construed as evidence in favor of the view that the cercal 
appendages belong to the tenth segment; or, on the other hand, it 
might be taken as favoring Heymons’ claim that the true eleventh 
segment has been obliterated. However, it is not necessary to assume 
that the muscles associated with the cerci are primarily muscles of 
these appendages. The great bundles of fibers that operate the 
pincer-like cerci of Japyx almost fill the large tenth abdominal seg- 
ment (fig. 40 C, mcl), but they appear to be the longitudinal dorsal 
muscles normal to this segment, which secondarily function as cercal 
muscles by reason of their posterior attachments at the bases of the 
cerci. Ford (1923), observing that most of the cercal muscles in 
Orthoptera arise from the tenth tergum, asserts that these muscles are 
““intersegmental muscles between the tenth and eleventh segments,” 
while the muscles from the epiproct, she says, represent “ the inter- 
segmental muscles between the eleventh and twelfth terga.” (Her 
reference of the posterior muscles to the twelfth tergum is in accord 
with her acceptance of Heymons’ claim that the eleventh segment 
has disappeared in the adult.) 

Whatever may be the nature of the dorsal muscles of the cerci, 
the fact is significant that the organs have no ventral musculature— 
in this respect cerci differ from styli and gonapophyses. The absence 
of muscles from the paraprocts to the cerci, moreover, weakens the 
comparison between the paraprocts and the stylus-bearing plates of 
the preceding abdominal segments, since the stylus muscles always 
take their origin in these plates. The termination of the ventral mus- 
culature of the abdomen in the paraprocts, on the other hand, makes 
it almost certain that the paraprocts are terminal lobes of the eleventh 
sternum. 

Cerci are usually absent in holometabolous insects, but cercus- 
like appendages occur on the eleventh abdominal segment in females 
of Panorpa (fig. 8H, Cer?), and on the terminal segment of adult 


96 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


Tenthredinidae, which is numerically the tenth abdominal segment. 
‘The panorpid appendages may be true cerci. Appendages occurring 
on an apparent tenth segment might be suspected of being cerci if 
there is evidence that this segment is composed of the tenth and 
eleventh somites, a condition which frequently occurs in orthopteroid 
insects, where there is no doubt that the terminal appendages are 
the cerci. In the Tenthredinidae, however, there is reason to believe, 
as will be shown later, that the terminal appendages of the adult are 
not the cerci, but are appendicular organs of the tenth segment cor- 
responding with the socii of adult Lepidoptera, and that they are de- 
rived from the postpedes of the tenth somite of the larva. 


THE TERMINAL APPENDAGES OF ENDOPTERYGOTE LARVAE 


Appendicular organs representative of abdominal limbs are present 
on the last abdominal segment in some or most of the larvae of 
Neuroptera, Trichoptera, Lepidoptera, and chalastogastrous Hymen- 
optera. These larval appendages of the terminal segment have a 
lateral or latero-ventral position, and are movable by muscles in some 
cases attached on their bases, but more generally inserted within 
their distal parts. The appendages most resemble jointed limbs in 
the Trichoptera. In Neuroptera, Lepidoptera, and Hymenoptera they 
commonly have the structure typical of the abdominal legs of the 
caterpillars. Since the terminal segment in these larvae is evidently 
the true tenth somite of the abdomen, or the tenth and the eleventh 
somites combined, there is little doubt that the terminal appendages 
are the pygopods. The Endopterygota differ thus from the more gen- 
eralized Exopterygota in that some of them retain the tenth segment 
appendages in postembryonic stages. 

In addition to the true appendicular organs, there may be in endop- 
terygote larvae also processes developed from the dorsum of the last 
segment, and lobes of various forms associated with the anal opening, 
or protruded from within the rectum. Processes resembling cerci 
occur in some coleopterous larvae, but their morphology is uncertain. 

It seems probable that the pygopods of endopterygote larvae are, 
in certain orders, caried over to the adult stage as processes which 
sometimes occur on the proctiger, or terminal segment of the imago. 
These processes have various forms in the Trichoptera and Lepidop- 
tera, and are termed the soci by students of the latter; in chalasto- 
gastrous Hymenoptera they resemble cerci, and are frequently called 
“cerci.”” Busck and Heinrich (1922) have observed that in the micro- 
lepidopteron Ethmia machelhosiella the anal prolegs, with their 





no. 6 INSECT ABDOMEN—SNODGRASS 97 


crochets, are retained in the pupa, and Bottimer (1926, fig. 3 A) 
shows a similar retention of the anal legs in the pupa of Chaetocampa 
crotonella. It is unfortunate that socii are absent in these species, 
for we might expect to find the socii developed within the anal pro- 
legs of the pupae. More positive evidence of the identity of the termi- 
nal larval appendages with the terminal appendages of the adult is pre- 
sented by Middleton (1921) in a study of the chalastogastrous Hy- 
menoptera. Middleton claims that the anal prolegs, or postpedes, of the 
larva of Pteronidea ribesti are transformed during metamorphosis into 
anal lobes of the pupa, and that within these lobes are developed the 
so-called ‘‘ cerci” of the adult sawfly. These appendages of the adult 
insect, he points out, are not borne by the tergum of the anal segment, 
but arise from a lateral membranous area of the venter of this seg- 
ment, and thus morphologically have the same position as have the 
postpedes of the larva. The anal segment in both the larval and the 
adult sawfly is numerically the tenth abdominal segment, and its ap- 
pendages are therefore not the true cerci in either case. 

Trichoptera—The abdomen of trichopterous larvae lacks appen- 
dages except on the terminal segment. In some forms, as in Platyphy- 
lax designatus, a fringe of slender setae along each side of the abdomen 
from the second to the end of the eighth segment (fig. 41 A, B, a) 
evidently marks the dorso-pleural line separating the dorsum of the 
abdomen from the region of the limb bases (LB), since, if the line of 
this fringe were carried into the thorax, it would run dorsad of the 
thoracic subcoxae (A, Scv;). On the ninth segment there is nothing 
to mark the dorso-pleural boundary; but on the terminal segment 
the base of the appendage (B, Pf) has a lateral position corresponding 
with the limb base areas of the segments preceding the ninth. The 
terminal segment of Platyphylax is a hemispherical lobe with a long, 
median anal cleft on the ventral part of its distal surface (An). It is 
evidently the tenth somite, or pygidial segment. 

The pygopods of trichopterous larvae differ considerably in different 
families and genera. Their principal variations have been described by 
Ulmer (1903) and by Krafka (1924). According to Ulmer there 
are two principal types of these appendages. Those of one type are 
short ; those of the other, characteristic of Hydropsychidae and Rhya- 
cophilinae, are long and leg-like. In both types the limb terminates in 
a hook-like claw. The structure of the two forms of appendages 
is here illustrated from Platyphylax designatus (fig. 41 B, Pp) and 
an unidentified species of Hydropsyche (F, Pp). 

In Platyphylax designatus each larval pygopod together with its 
supporting structure (fig. 41 B, Pp) consists of a large basal plate (C, 


98 SMITHSONIAN MISCELLANEOUS COLLECTIONS ~— VOL. 85_ 


b), a smaller intermediate plate (c), and a free apical claw (d) hav- 
ing its point turned downward and outward. The apical claw has a 
movement of adduction on the middle plate, and the latter is movable 
on its hinge with the basal plate. The muscles of this appendage may 
be studied by cutting the terminal body segment into lateral halves. 
It is first seen that the base of the appendage is crossed internally by 
several slender transverse muscles attached on the segmental walls. 
The muscles inserted on the appendage consist of three flat groups of 





Fic. 41.—Structural details of trichopterous larvae. 


A, metathorax and anterior part of abdomen of Platyphylax designatus. 
B, posterior abdominal segments of same. C, appendage of tenth segment of 
same. D, mesal view of right appendage, showing muscles. E, mesal view of 
terminal appendage of Hydropsyche sp., showing muscles. F, posterior end of 
abdomen of same, with intestinal processes (ip) extruded from anus. 

a, line of dorso-pleural groove; An, anus; b, proximal sclerite of appendage; 
Cx, coxa; d, claw of appendage; e, f, intersegmental folds between eighth and 
ninth, and ninth and tenth abdominal segments ; 7, intestinal processes; LB, area 
of abdominal limb base; Pp, pygopod; Scx, subcoxa. 


fibers (fig. 41 D), all arising on the anterior margin of the basal 
plate (b). Two of these muscles (za, rb) converge downward and 
posteriorly to a common point of insertion on the inner margin of 
the base of the apical claw (d), and thus evidently function as ad- 
ductors of the latter. The third muscle is a broad sheet of fibers 
(2) lying external to the others, and inserted on the dorsal margin of 
the intermediate plate (c). Associated with the appendage is a pair of 
slender, vertical fibers (5) crossing the inner face of the basal plate 
and attached above and below it to the membranous walls of the 





No. 6 INSECT ABDOMEN—SNODGRASS 99 


terminal body segment. The ventral muscles of the ninth body seg- 
ment (C, 4) are attached posteriorly on the intersegmental fold (f) 
just before the lower part of the basal plate of the appendage, but 
no muscles are inserted directly on the latter. 

The terminal appendages of the larva of Hydropsyche (fig. 41, 
F, Pp) represent quite a different type of appendicular organ. Each 
pygopod here is a large, two-segmented, leglike structure projecting 
from the ventro-lateral part of the terminal body segment. The 
proximal segment of the appendage (E, b) is elongate and cylindrical ; 
the distal segment (d) is a strong, decurved claw with a high, nar- 
row base articulated by its dorsal end with the dorsal extremity of 


_the basal segment. Ventrally, just before the base of the claw, there 


is a small triangular sclerite (c) in the lower, membranous wall of 
the proximal segment. 

The musculature of the terminal appendage of Hydropsyche (fig. 
41 E) differs in several respects from that of Platyphylax (D). As 
in the latter, however, there are no muscles inserted on the base of 
the organ, but there are an adductor muscle (E, 1) inserted on the 
base of the apical claw (d), and two flexor muscles (2, 3) inserted 
on the small ventral sclerite (c). The adductor (1) arises, not in 
the basal segment of the appendage as in Platyphylax (D, 1a, 1b), 
but in the proximal part of the last body segment. One of the flexor 
muscles (E, 2) is a broad fan of fibers arising on the dorsal wall of the 
basal segment, and thus suggests an identity with muscle 2 of Platy- 
phylax (D) arising on the basal plate of the appendage. The other 
flexor muscle is a long bundle of fibers (E, 3) arising on the interseg- 
mental fold (e) between the eighth and ninth abdominal segments 
along with the fibers of the ventral longitudinal muscles of the ninth 
segment (4), but separating from the latter posteriorly to enter the 
appendage. This muscle apparently has no representative in Platy- 
bhylax (D). 

The relation between the two types of appendages described above 
is obscure. The only suggestion that can be made is that the basal 
plate of the Platyphylax appendage (fig. 41 D, b) corresponds with 
the basal segment of the Hydropsyche appendage (FE, b) and that the 
intermediate plate of the former (c) is represented by the small ven- 
tral sclerite (c) of the latter. The apical claw (d) is evidently the 
same in both. Still more difficult is it to find possible homologies 
between either of these two types of trichopterous appendages and 
the terminal appendages of the sialid larvae, Chauliodes and Corydalus 
(figs. 35 D, E, 36 B, E), or the abdominal legs of lepidopterous larvae 
(figs. 37, 42 C). The basal plate or basal segment (0) in the tri- 

7 


100 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


chopterous appendage, however, may be the limb basis of a more 
typical abdominal appendage, while the apical claw (d) and the asso- 
ciated middle plate (c) may possibly represent the stylus. Judging 
from the structure there is little probability that the trichopterous 
claw is homologous with the claws of Corydalus or Chauliodes, or 
with the crochets of lepidopterous larvae. The lack of similarity 
in the larval abdominal appendages of Trichoptera and Lepidoptera 
is somewhat surprising, considering the many other structural like- 
nesses between these two orders. 

Many trichopterous larvae are provided with a group of slender 
processes protractile through the anus (fig. 41 F, ip), which arise 
from the intestinal wall. In the species of Hydropsyche figured there 
are from four to six of these processes. The structures are hollow, 
thin-walled tubules containing no tracheae, and are therefore usually 
termed “blood gills”; but the idea of their respiratory nature is 
based on their structure and on the fact that they can be entirely 
exposed by protraction through the anus, for no one apparently 
has made any physiological experiments on their function. According 
to Branch (1922) the organs arise as diverticula of the intestinal 
wall produced posteriorly from the six folds of the pre-rectal part of 
the proctodeum, and each is provided with a three-branched muscle 
taking its origin on the intersegmental membrane between segments 
VIII and IX of the abdomen, and extending through the lumen to 
the tip of the processes. When retracted the processes lie in the rec- 
tum with only their extremities exposed in the anal aperture.. Pro- 
traction evidently is accomplished by internal pressure resulting from 
a contraction of the abdominal walls. 

Neuroptera——tThe pygidial appendages in the larvae of the stalid 
genera Chauliodes and Corydalus, as already noted, are long, thick 
structures projecting posteriorly and ventrally from the terminal 


segment of the abdomen (fig. 35 D, E, Pp, fig. 36 B, E). Each organ ~ 


consists of a large basis (fig. 36 E, LB), supporting laterally a flex- 
ible, tapering process representing the stylus (Sty), and bearing dis- 
tally a short, cylindrical lobe (V's) armed with two strong claws (d). 
The appendage has a striking resemblance to the typical abdominal 
leg of a caterpillar, except that the latter has no representative of the 
stylus. The distal lobe, which clearly is serially homologous with 
the gill-bearing tubercles of the preceding appendages in Corydalus 
(fig. 36 B, C, D, Vs), is remarkably like the planta of the caterpillar’s 
leg (fig. 37 D, Vs). It is retracted by a strong muscle (fig. 36 F, rvs) 
arising on the dorsal wall of the terminal body segment, and inserted 
on the distal wall of the lobe at the posterior or upper end of the long 


no. 6 INSECT ABDOMEN—SNODGRASS IOI 


bases of the claws. The claws themselves are in every way suggestive 
of the crochets of the caterpillar’s foot. 

Notwithstanding the general structural resemblance between the 
terminal appendages of Chauliodes or Corydalus and the abdominal 
legs of caterpillars, it can scarcely be supposed that the foot struc- 
ture in either case has been derived immediately from that of the 
other, since the two-clawed condition would be a highly specialized 
one in the Lepidoptera. All that may be claimed is that the funda- 
mental structure of the larval abdominal limbs is the same in both 
the Neuroptera and the Lepidoptera. The neuropterous appendage 
is the more primitive in that it retains the stylus, which has the form of 
a segmented appendage in Sialis (fig. 35 A, C). We may assume that 
the gill-bearing tubercles of the Corydalus larva have been formed 
secondarily from the foot lobes, or retractile vesicles, as a better adap- 
tation to aquatic life in this genus, and that the vesicles have been 
lost from all the appendages in Sialis, and from all but the terminal 
appendages in Chauliodes. 

Lepidoptera.—The so-called anal legs, or postpedes, of caterpillars 
are so similar to the legs of the preceding abdominal segments as 
scarcely to need a separate description. The musculature of the two 
sets of appendages, however, differs in some respects. The plantar 
lobe of each anal leg is retracted by a large dorsal muscle (fig. 42 A, 
B, C, rvsd) and a small ventral muscle (rvsv), both arising from the 
intersegmental fold (f) before the ninth abdominal segment. The 
lateral muscles of the leg are reduced to a few fibers (D, b) lying 
external to the large dorsal retractor of the planta. Between the bases 
of the legs there is a sheet of transverse ventral muscles (B, tv), 
which appear to belong to the wall of the last body segment rather than 
to the appendages. 

The great development of the dorsal retractor muscles of the anal 
legs, the reduction of the lateral muscles, and the presence of the 
ventral retractors of the plantae are all features correlated with the 
function of the postpedes in the caterpillar, which usually have a 
stronger independent forward movement than do the legs of the pre- 
ceding segments. 

The large terminal segment of lepidopterous larvae appears to be 
a compound segment composed of the tenth abdominal somite, with 
its appendages, the pygopods, and the reduced eleventh segment, bear- 
ing the anus, but lacking cerci. Figures of the embryo of Pieris rapac 
given by Eastham (1930) show clearly a well-developed tenth abdom- 
inal segment bearing the last pair of appendages, and beyond it a 
large terminal lobe, containing the anus, which is evidently the 


102 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


eleventh somite. In the caterpillar (fig. 11) the eleventh somite is 
apparently represented in the terminal segment by the four postpedal 
lobes surrounding the anus (C, D), which may be supposed to be the 
epiproct (Eppt), the paraprocts (Papt), and a hypoproct (Hypt). 
There is, however, no evidence of a dual composition of the terminal 
segment of the caterpillar furnished by the musculature of this seg- 
ment. Both the dorsal and the ventral internal longitudinal muscles 





Fic. 42—Musculature of the terminal segment and pygopods of a noctuid 
caterpillar. 


A, dorsal muscles and leg muscles seen from below in terminal segment cut 
open along mid-ventral line and spread out. B, ventral musculature of same 
segment, seen from above. C, posterior end of abdomen, showing origins of leg 
muscles. D, basal rim of left leg (L) and group of small muscles (b) lying 
beneath dorsal retractors of planta (rvsd). 

An, anus; b, group of small antero-lateral lez muscles; de, external dorsal 
muscles; di, internal dorsal muscles; Eppt, epiproct; f, intersegmental fold 
between ninth and tenth segments; Hypt, hypoproct; L, basal rim of leg; J, 
lateral muscles; Papt, paraproct; rvsd, dorsal retractor of planta; rvsv, ventral 
retractor of planta; tv, transverse ventral muscles; vi, internal ventral muscles; 
Vs, planta. 


(fig. 42 A, B, di, vi) extend continuously from the anterior inter- 
segmental fold (f) to the epiproct and the paraprocts. Two large 
sets of internal dorsal fibers (A, de) arise in the notches between 
the epiproct and paraprocts and are inserted on the dorsal plate of the 
segment. The eleventh segment, therefore, if represented here at all, 
is reduced to the circumanal lobes; and the fibers of the longitudinal 
muscles of the tenth and eleventh segments have become continuous. 











No. 6 INSECT ABDOMEN—SNODGRASS 103 


Continuity of muscle fibers is of frequent occurrence wherever the 
intersegmental connections are lost, or where segmental boundaries 
are obliterated. 

Chalastogastrous Hymenoptera.—The larvae of the sawflies and 
horntails are provided with terminal appendicular organs of several 
varieties, all borne by the last abdominal segment, some arising from 
the dorsum, others from the venter. 

On the dorsum there is in some forms, as in Pteronidea ribesii 
(fig. 43 A), a pair of small, immovable lateral processes (a) arising 
from the end of the last segment above the anal opening (An). These 
processes have been regarded as rudimentary cerci (Crampton, 1919), 
but they are more evidently mere cuticular outgrowths, as claimed 
by Middleton (1921), analogous with the urogomphi of coleopterous 
larvae (fig. 44 C, ug), since they have none of the characteristics of 





Fic. 43.—Terminal appendicular structures of larvae of chalastogastrous 
Hymenoptera. 


A, Pteronidea ribesti (Tenthredinidae). B, Cimbex americana (Cimbicidae). 
C, Cephaleis sp. (Pamphiliidae). 

a, paired processes of dorsum of terminal segment; An, anus; b, median proc- 
ess of dorsum of terminal segment; Pp, postpedes, or pygopods; rvs, retractor 
muscle of plantar lobe; ls, planta. 


true cerci, and, according to Middleton, take no part in the formation 
of the lateral, cercus-like appendages of adult Tenthredinidae. In 
certain other chalastogastrous larvae a median process, or postcornu 
(fig. 43 C, b), is borne on the end of the tenth abdominal segment. 
This process varies characteristically in shape and size in different 
families, as tabulated by Middleton (1921). It occurs in larvae that 
bore into stems or that live in silk-spun tents or within the hollow of 
curled leaves. 

The ventral appendages of the pygidial segment of chalastogastrous 
larvae likewise vary in form in different groups. Those of species 
that live in the open closely resemble the anal legs of lepidopterous 
larvae. In Pteronidea ribesii, for example (fig. 43 A), a ventral pro- 
jection of the tenth segment beneath the anus, bearing two small 
terminal lobes (ls), is clearly the homologue of a pair of appendages 





104 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


such as occur on the more anterior segments of the abdomen. The 
free lobes (V's) are evidently comparable with the plantae of the ab- 
dominal legs of caterpillars (fig. 37 D, Vs), though the retractor mus- 
cles of the lobes in the sawfly larva take their origins from a point 
on the side of the segment just anterior to the cleft of the anal open- 
ing (fig. 43 A, rvs). In other forms, as in Cimbex americana (B), 
the ventral appendages (Pp) consist apparently of the plantar lobes 
only, which arise directly from the flattened venter of the pygidial 
segment. Again, as in the Cephidae and Pamphiliidae, typical “ pro- 
legs ” are replaced on the tenth segment by slender jointed appendages 
(C, Pp); but these “ arthrostyli,” as they have been called (Cramp- 
ton, 1919), are evidently alternative forms of the anal “ prolegs,” or 
postpedes, since they arise at approximately the same points as do 
the typical postpedes in other forms (A, Pp), and do not occur con- 
jointly with the latter. They are not, however, provided with muscles, 


so far as the writer could discover; but each is penetrated by a large 


nerve, and bears sense organs on the distal segment having the form 
of minute disks, in addition to setae on the proximal and middle 
segments. The jointed form of the terminal appendage is, therefore, 
evidently an adaptation to a sensory function instead of a locomotory 
one. 

Crampton (1919) suggests that the jointed appendages, or “ arthro- 
styli,’ of chalastogastrous larvae do not represent the “ prolegs ” 
directly, but that each has the relation to the latter of a stylus, that 
is, it is an appendicular part of the true appendage. Middleton (1921), 
on the other hand, thinks that the jointed appendages are direct repre- 
sentatives of the unjointed postpedes, because the two organs have 
identical positions on the tenth segment, and do not occur together. 
He would attribute the difference in form to the different habits of the 
larvae, since those species having typical, fleshy postpedes feed in the 
open on leaves and grasp the edges of the latter with the terminal 
appendages, while those having slender, jointed postpedes, bore into 
the stems of plants, or live in the protection of web nests or curled 
leaves. To the writer it appears most probable that the two forms 
of appendages are identical organs, and that the jointed variety is a 
secondary modification of the typical postpedes for a sensory function, 
the jointing being a mere subdivision of the appendage and not a true 
segmentation. 

The observation made by Middleton (1921) that the postpedes 
of the larva become the cercus-like appendages of the adult sawfly has 
been discussed in the introductory part of this section, and need be 
given no further attention here, except to point out its importance, 


y 


; 
. 


a 


No. 6 INSECT ABDOMEN—SNODGRASS 105 


if true, in establishing an identity between the terminal appendages 
of larval insects and the appendicular processes of the tenth segment 
in certain adult holometabolous insects. 

Coleoptera.—In many families of the Coleoptera the abdomen of 
the larva is provided with a pair of appendicular processes arising 
from the dorsum of the ninth segment. These structures have been 
variously termed styli, cerci, pseudocerci, corniculi, and urogomphi. 

The abdomen of the larva of Dytiscus ends with a transverse pos- 
terior surface of the narrow eighth segment (fig. 44 A), in the upper 
part of which the last pair of spiracles open through a median, vertical 
slit (VIIISp). From the membrane below the spiracular area of the 
eighth segment there arise laterally two slender, tapering processes 
(ug?) fringed with long hairs. Between the bases of these organs 





Vill NIISp an 


r An 
/ el ect? \G i 





Fic. 44.—Terminal appendages of coleopterous larvae. 


A, posterior end of abdomen of Dytiscus circumcinctus larva. B, end of ab- 
domen of a silphid larva, Thanatophilus sp., with exserted anal lobes (from 
Kemner, 1918). C, end of abdomen of carabid larva, Oodes helopioides, with 
exserted anal lobes (from Kemner, 1018). 

An, anus; anl, anal lobes; ug, urogomph. 


is a small, median plate (JX7?), which appears to be a remnant of 
the ninth abdominal tergum. Ventral to it is the anal opening (An). 
According to Speyer (1922, Korschelt, 1924), each of the terminal 
appendages of the larva of Dytiscus marginalis is provided with three 
muscles, two inserted dorsally on its base, and one ventrally, all of 
which arise on the tergum of the eighth segment. 

The morphology of the larval appendages of Dytiscus is difficult 
to determine. The apparent position of the organs on the rudimentary 
ninth segment makes it doubtful that they are true cerci, and the dorsal 
origins of their muscles is not in accord with the musculature of 
styli. It is claimed by Blunck (1918, Korschelt, 1924) that in all 
Coleoptera the first two primitive somites of the abdomen are united 
in the first definitive segment, and that, therefore, the segment bear- 
ing the terminal appendages is really the tenth. In this case we might 


106 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 85 


regard the terminal appendages of the Dytiscus larva as the cerci. 
However, the evidence presented by Blunck of the fusion of the first 
and second segments of the abdomen is not generally accepted by 
students of Coleoptera, and is not convincing, while the fact that the 
segment preceding the appendicular processes bears the eighth pair of 
abdominal spiracles would ordinarily be taken as conclusive evidence 
that this latter segment is the true eighth somite of the abdomen, 
especially since these respiratory apertures, as described by Blunck, 
have the structure typical of lateral abdominal spiracles. 

For the present, therefore, the nature of the terminal appendages of 
the Dytiscus larva must be left in doubt, but if the organs do not 
belong to the series of lateral, stylus-like appendages, fully repre- 
sented in Coptotomus and Gyrinidae, it is possible that they are struc- 
tures homologous with those dorsal processes often developed on the 
ninth abdominal segment of other coleopterous larvae, which appear 
to be mere cuticular outgrowths, though they may become movable 
at their bases. The muscles inserted on the processes in Dytiscus are 
evidently not specific muscles of the appendages, but groups of seg- 
mental or intersegmental fibers that, by reason of their attachments 
at the bases of the processes, serve to move the latter. 

The abdomen of most other coleopterous larvae consists of 10 
distinct segments (fig. 44 B, C). The ninth segment is usually well 
developed and frequently supports a pair of processes (B, C, ug) 
arising from its dorsum. These processes are termed urogomphi by 
Boving, since from their segmental position it is clear that they can 
not be cerci. In some species the urogomphi are fixed outgrowths 
of the posterior end of the ninth tergum (C) ; in others they arise 
from the membrane behind the ninth tergal plate (B), and are then 
flexible at their bases. They vary much in size and shape from 
short, spine-like points to long, thick processes or multiarticulate fila- 
ments, and they are sometimes distinctly jointed. The urogomphi 
appear to be equivalent structures in all cases, and are probably but 
cuticular outgrowths of the ninth abdominal tergum, which in 
certain species become movable by a membranization of the tergal 
wall at their bases. The mobile appendages of the Dytiscus larva, 
therefore, are perhaps urogomphi of the rudimentary ninth segment, 
secondarily movable by some of the intersegmental muscles normally 
extending between the eighth and ninth segments. 

The tenth abdominal segment in many coleopterous larvae is a 
short, often tubular structure (fig. 44B, C, X), at the extremity 
of which is a small retractile and eversible membranous pad sur- 
rounding the anus, known as the “ pygopodium ” (C, XJ), which may 





No. 6 INSECT ABDOMEN—SNODGRASS 107 


be produced into two or more soft lobes (anl). It is claimed by Kem- 
ner (1918), from embryological evidence, that the so-called pygo- 
podium is the rudimentary eleventh abdominal somite. The eversible 
pygopodial lobes, however, in no way represent appendicular organs, 
Kemner asserts, nor are they evaginations of the wall of the rectum, 
as they have been supposed to be; they are merely productions of the 
cercumanal area on which the longitudinal muscles from the tenth 
to the eleventh segment are attached. If the organs in question, there- 
fore, are not of an appendicular nature, the term “ pygopodia ”’ should 
not be given to them, since it is convenient to apply this name spe- 
cifically to the true appendages of the tenth, or pygidial, segment. 
The presence of a distinct though rudimentary eleventh segment 
in larvae of Coleoptera is of interest because of the general suppres- 
sion of this segment in holometabolous insects. 


TERMINAL LOBES OF THE PARAPROCTS 


In a few of the lower Pterygota an appendicular lobe is borne by 
each of the paraprocts. These processes have been termed “ para- 
processi”’ by Crampton (1920). Examples of paraproct lobes occur 
in the Odonata and in the tridactylid Orthoptera. 

The paraproct processes of Odonata occur in adult Anisoptera in 
the form of small, seta-bearing lobes projecting posteriorly from the 
ends of the paraprocts. Corresponding lobes are not present in the 
larvae of this group of dragonflies, in which the paraprocts, together 
with the elongate epiproct, form the valves that close the anal open- 
ing (fig. 12A, B). In the larvae of Zygoptera, however, paraproct 
lobes are highly developed as the large, flat, tracheated plates that form 
the lateral caudal gills (C, paptl). The median gill (cf) is a similar 
lobe of the epiproct (Eppt), and is evidently comparable with the 
median caudal filament of Thysanura (fig. 7 A, B, C, cf). 

In the Orthoptera paraproct lobes are well developed in the Tri- 
dactylidae, where they have the form of long processes resembling 
the cerci (fig. 45 A, B, paptl). In some species of Ripipteryx those of 
the male are incurved at the ends and are said to be used as claspers 
during copulation. In Ellipes (fig. 45 A, B) each process is borne 
on a membranous area at the end of the short paraproct (Papf). 
Crampton (1918, 1921) has given considerable significance to the 
“ paraprocessi ”’ of the Tridactylidae, which he regards as homologous 
with the styli of the preceding segments, and as representatives of 
the exopodites of crustacean appendages (fig. 45 C, D, expd). Most 
other writers, including Walker (1919), regard the processes as sec- 


7 


108 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 





ondary lobes of the paraprocts. It is highly unlikely that the struc- 
tures in question are styli, since, as already shown, the paraprocts are 
not limb bases. Crampton’s view regarding the relation of the para- 
proct process to crustacean exopodites will be discussed in the fol- 
lowing Section of this paper. 


Milly "IX xX. Ger 





Fic. 45.—Terminal appendages of a tridactylid insect and an isopod crustacean. 


A, posterior part of abdomen of Ellipes minuta (tridactylid orthopteron). 
B, the same, dorsal view. C, end of abdomen of Porcellio (isopod crustacean), 
ventral view. D, the same, dorsal view. 

An, anus; Cer, cercus; Endpd, endopodite; Eppt, epiproct; Expd, exopodite; 
la, lamina sub-analis; Papt, paraproct; paptl, lobe of paraproct; Prtpd, protop- 
odite; sa, lamina supra-analis; Tel, telsqgn (fused with tergum of twentieth 
segment). 


MORPHOLOGY OF THE ABDOMINAL APPENDAGES 


There is no need at present to offer proof of the serial identity 
of the appendicular organs of the insect abdomen with the segmental 
appendages of the thoracic and cephalic regions of the body. Embry- 
ologists have amply demonstrated the continuity of appendage rudi- 
ments on the entire series of primitive somites, at least 20 in all, leav- 
ing only a prostomial lobe and a periproctial region devoid of true 
limb structures. To determine the homologies of the parts of the 
abdominal appendages with the parts of the better developed append- 





No. 6 INSECT ABDOMEN—SNODGRASS 109 


ages on the anterior regions of the body, or with those of theoretically 
more generalized appendages, is quite another matter. 

Theoretical Structure of Arthropod Appendages.—At the outset of 
an attempt to study the morphology of the rudimentary abdominal 
appendages of insects a difficulty is encountered arising from the 
lack of uniform opinion as to the structure of a generalized but 
fully segmented arthropod limb. Particularly is it necessary in a 
study of rudimentary appendages to know the structure of the basal 
part of a primitive appendage. In the thorax of modern insects 
the functional base of a leg is the coxa, and yet, it seems almost 
certain that at an earlier stage the true basis of the limb must have 
included the subcoxal region now forming the so-called pleuron and in 
some cases a lateral part of the definitive sternal plate of the support- 
ing body segment. Then, the further question arises as to whether the 
subcoxa was once a free segment of the appendage, or whether it has 
been evolved secondarily by a differentiation of the primitive limb 
basis into subcoxal and coxal parts. Differences of opinion on such 
questions as these have led immediately to different interpretations of 
the basal parts of the abdominal appendages, and consequently to 
different views concerning the nature of the distal parts. 

A simple condition of the limb base occurs throughout the Arach- 
nida, which is well shown in any one of the legs of a phalangid 
(fig. 46 A). It is to be seen here that the leg is supported on a large 
basal segment (LB) that occupies the lateral wall of a segmental area 
of the body, and that it turns slightly forward and rearward on a 
dorsoventral axis (a-b) extending from the tergum above to the 
sternum below. To this large basal segment the telopodite is articu- 
lated by a dicondylic hinge on a horizontal axis (f-g). The first seg- 
ment of the telopodite is a trochanter (z77r). 

In the majority of the Crustacea the proximal region of the leg has 
the same structure as in the Arachnida, there being a single basal 
segment, the coxopodite (fig. 46 B, C, LB), implanted directly in the 
ventro-lateral wall of the supporting body segment, and often articu- 
lated dorsally (a) with the tergal plate (B, 7). The basis, however, 
is not prolonged ventrally as in the phalangid (A), and is inclined to 
be cylindrical (C). The telopodite articulates with the basis by its 
proximal segment (the first trochanter, or basipodite, B, C, rT7r) 
on a horizontal, dicondylic hinge (f-g) having the same type of struc- 
ture as that in the arachnid (A). 

If, now, we look at a typical thoracic leg of an insect (fig. 46D), 
it is seen that the proximal joint in the appendage corresponding struc- 
turally and functionally with the joint between the basis and the telop- 


Ilo SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


odite in the phalangid or crustacean leg (A, B, C, f-g) is that be- 
tween the coxa and the trochanter (D, f-g). The coxa (Cx), how- 
ever, is separated from the tergum (7), and often from the sternum 
(Stn), by a sclerotization, known as the “ pleuron ” (Scx), occupying 
the position in the lateral wall of the supporting body segment that is 
occupied by the limb basis in the phalangid and crustacean (A, B, C, 
LB). In other words, the basal region of the appendage in the case 
of the insect leg (D, LB) is composed of two parts (Scx, Cx) cor- 


/ 
1Tr+2Tr 





Fic. 46.—Basal parts of arthropod legs. 


A, third right leg of a phalangid, Liobunum, showing limb basis (coxopodite) 
elongate ventrally (LB), turning antero-posteriorly on dorsoventral axis (a-b), 
with telopodite movable dorsoventrally on horizontal axis (f-g). 

B, last right ambulatory leg of a crustacean, Anaspides tasmaniae, with small 
basis, or coxopodite (LB), on which the telopodite turns dorsoventrally on hori- 
zontal axis (f-g) of first trochanter, or basipodite (177). 

C, leg of an isopod crustacean, Porcellio, with elongate basis, or coxopodite 
(E 2 } Ae which the telopodite is articulated by a horizontal hinge (f-g) as in 
A and B. 

D, mesothorax and middle leg of a young acridid nymph, Melanoplus, showing 
region of limb basis (LB), between the horizontal baso-telopodite hinge (f-g) 
and the tergum (7), subdivided into subcoxa (Sc) and coxa (Cr). 

a, dorsal articulation of limb basis with body; a-b, axis of articulation of 
limb basis with body; c, dorsal subcoxo-coxal articulation; Cx, coxa; Cxrpd, 
coxopodite (limb basis) ; f-g, axis of baso-telopodite hinge; Fm, femur (merop- 
odite) ; h-i, axis of hinge between first and second trochanters; 7-k, axis of 
trochantero-femoral hinge; LB, limb basis (coxopodite); Scr, subcoxa; Stn, 
sternum; 7, tergum; 17,, first trochanter (basipodite) ; 27r, second trochanter 
(ischiopodite). 


responding with the single plate or segment (coxopodite) forming the 
basis of the arachnid and crustacean appendages illustrated (A, B, C). 
A condition similar to that in the insect thorax is found, however, 
in the thoracic region of the decapod crustaceans, where a large 
pleural plate occurs on each side of the body intervening between 
the coxae and the tergum. This plate is clearly a composite structure, 
the segmental areas of which are evidently derived from the sub- 
coxal parts of the leg bases, because they support the gills of the 





no. 6 INSECT ABDOMEN—SNODGRASS III 


branchial chamber. Subcoxal sclerites occupying the lateral walls 
of the body segments are again found in the Chilopoda (fig. 27 A, 
Scx), and in the last leg-bearing segment they are here united with 
the coxa to form a single basal plate (LB) of the appendage, occupy- 
ing the pleural region of the segment. 

From comparative studies such as those just cited, the writer has 
come to regard the pleural subcoxa, or sclerotic area of the lateral 
body wall supporting the free part of an appendage, as the proximal 
part of the primary limb basis secondarily separated, wherever it 
occurs, from the distal part of the basis, which becomes the movable 
coxa, or functional base of the limb. On the other hand, many 
students of Arthropoda regard the subcoxa as a primitive limb seg- 
ment, which has become suppressed or fused with the coxa wherever 
traces of it are not to be found in modern forms. The entire absence 
of a subcoxa in all Arachnida and in the majority of Crustacea, how- 
ever, is against this view; and the lack of uniformity in the subcoxo- 
coxal musculature, when a subcoxa is present, suggests that the sub- 
coxo-coxal joint is a recent division of the limb basis that has occurred 
principally in the Chilopoda and the Hexapoda. If the subcoxa is a 
secondary formation, then it must be assumed that the subcoxo-coxal 
muscles likewise are secondary, and that, as the subcoxa became 
differentiated from the coxa, most of the primitive basal muscles of 
the appendage were transferred to the coxa. 

The subcoxa in its more primitive condition is best seen in the 
Chilopoda and in the thoracic segments of apterygote Hexapoda. It 
here consists of a circular fold or slight elevation of the body wall 
supporting the leg, containing one or more small sclerites, particularly 
in the region above the coxa. The large “pleura” of the thoracic 
region of pterygote insects, or the pleural plates of the branchial cham- 
bers of decapod crustaceans, undoubtedly represent highly specialized 
developments of the subcoxae, adapting the latter to uses quite inde- 
pendent of any function connected with the legs. The subcoxal plates 
in the thorax of holometabolous insect larvae, however, are relatively 
small and are closely associated with the coxae (figs. 3 B, C, 41 A, 
Scx;). The region of the subcoxa surrounds the base of the coxa, 
but its ventral arc is reduced to a fold, which generally in the thorax 
of adult insects unites with the sternum. The sclerotized area of 
the subcoxa may be broken up into several small sclerites; in the 
thorax of pterygote insects there is typically a large supracoxal plate 
known as the ‘‘ pleuron.” 

In a former paper the writer (1928) has given reasons for believing 
that the body of a gnathal appendage represents the basis of a leg. It 


I1I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


is similarly divided secondarily into two parts (cardo and stipes), 
though it is not clear that the two parts exactly correspond with the 
subcoxa and coxa of a thoracic leg. The palpi, however, are thus 
seen to be the telopodites of the head appendages. If the interpretation 
concerning the primary segmentation of a gnathal or a thoracic 
appendage into basis and telopodite is now carried to the appendages 
of the abdomen, the basal plates or basal lobes of the latter become 
the true limb bases, and the telopodites should be freely movable ap- 
pendicular processes of the bases. 

In general, then, it appears that the arthropod limb is divided by a 
joint near its base into a proximal part, the primary limb basis, and 
into a distal part, or telopodite. The baso-telopodite joint is the coxo- 
trochanteral joint of a fully segmented limb, which is the joint be- 
tween the coxopodite and the basipodite in terms usually employed 
by carcinologists. The use of the term “basipodite” by Borner 
(1904, 1921) to designate the subcoxo-coxal base of the limb creates 
a duplication in nomenclature that is likely to be confusing. The 
movement of the telopodite on the basis is typically in a vertical 
plane, produced by levator and depressor muscles arising in the basis 
(hee 25205) O78). 

On the assumption that the basal mechanism of all the limbs is 
fundamentally the same in all groups of arthropods, we can imagine 
a simple primitive condition in the arthropodan ancestors in which 
the entire series of appendages had a uniform line of flexure near 
the body, along which the distal parts of the limbs, or telopodites, 
were movable in a vertical plane on their bases. The bases, on the 
other hand, turned forward and rearward on the axes of their attach- 
ments on the body. Wherever the basis is differentiated into a coxa 
and a subcoxa, the primitive basal movement of the appendage on the 
body is lost, but is replaced by a vertical axis of promotion and remo- 
tion between the subcoxa and the coxa, as the latter becomes sec- 
ondarily the functional base of the limb. Finally, if the limb becomes 
rudimentary and loses its basal musculature, the basis might become 
transformed to a simple immovable lobe or plate of the wall of 
the supporting body segment, with the telopodite reduced to an ap- 
pendicular process movable by muscles arising in the basis. 

Other Theories on the Morphology of the Abdominal Appen- 
dages.—The principal problem encountered in a study of the abdom- 
inal appendages of insects is that of determining the homologies of the 
parts of the appendages with those of a generalized limb. The basis, 
the stylus, the gonapophysis, the eversible vesicle, each raise ques- 
tions as to its nature and derivation. 


’ 





no. 6 INSECT ABDOMEN—SNODGRASS 113 


Discussions on the morphology of the abdominal appendages of 
insects, and speculations on their possible homologies with the limbs 


of Crustacea have continued for half a century without leading to 


definite conclusions. They began at. least with Wood-Mason (1879), 
who, in a paper on the origin of insects, interpreted the stylus-bear- 
ing plates of the abdomen of Machilis as the protopodites of primi- 
tively biramous appendages, of which the endopodites are represented 
by the gonapophyses, and the exopodites by the styli. The eversible 
vesicles, Wood-Mason suggested, may be homologues of nephridia, 
those of the eighth and ninth segments being converted into the 
genital ducts. 

Diversities of opinion soon followed the publication of a more 
widely read paper by Haase (1889) on the morphology of the seg- 
mental appendages, containing not only a clear exposition of the 
appendicular nature of the stylus-bearing plates in the abdomen of 
Thysanura, but also a demonstration of the triple origin of the de- 
finitive abdominal sterna of insects in general from the union of the 
rudimentary embryonic limbs with the median sternal area in each 
segment. The styli, Haase claimed, are not the leg rudiments, but 
secondary structures of the nature of hairs, which have been con- 
verted into locomotory organs from sensory organs. The eversible 
vesicles he believed function as blood gills, but are to be traced back 
in all cases to coxal glands. 

Wheeler (1893) and earlier students of the embryology of insects 
regarded the gonapophyses of the genital segments as the direct 
representatives of the appendages of these segments. Considering 
the late development of the gonapophyses, however, and their in- 
variable median position on the body of the insect, Heymons (1896a) 
contended that the genital processes are secondary integumental out- 
growths having no relation to the appendages, and that the latter 
are preserved in the styli and cerci. Heymons’ heterodox opinion 
brought a severe criticism from Verhoeff (1896), who defended the 
limb nature of the gonapophyses as an established fact, and main- 
tained that the identity of the abdominal styli with the thoracic styli 
of Machilis could not be disputed, and that therefore both are merely 
appendicular processes of the legs. In reply to Verhoeff, Heymons 
(1896b) emphasized his former statements in evidence of his view 
concerning the nature of the appendicular parts of the abdomen, 
pointing out that, during embryonic development of the Orthoptera, 
the abdominal appendages disappear, and the gonapophyses are later 
formed quite independent of the limbs. He argued that if the 
gonapophyses are the limbs, intermediate stages should be found some- 


y 


114 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 85 


where between gonapophyses and legs. The styli, Heymons reasserted, 
are direct derivatives of the abdominal appendages, and occur in 
primitive forms such as Campodea and Japyx in which gonapophyses 
are lacking. From the abdominal limb rudiments of the embryo, Hey- 
mons showed, are produced not only the typical styli, but also the 
cerci, and the lateral gills of the larvae of Ephemerida and Svalis. 

Since Heymons’ views are based on embryological studies they 
deserve more attention than purely theoretical considerations. In his 
studies of the development of the appendages of Periplaneta, Ectobia, 
and Mantis, Heymons (1896a) observes that each limb rudiment 
of the ninth segment is early marked by a circular constriction, which 
divides the appendage into a broader proximal part and a slenderer 
distal part. The proximal part flattens out and finally is incorporated 
in the definitive sternal plate of the segment, while the distal part 
becomes longer and slenderer and develops directly into the stylus. 
The gonapophyses, on the other hand, Heymons claims, are secondary 
outgrowths of the sterna in the Orthoptera. Concerning them he 
says: “In Gryllus there can be no doubt of the nature of the 
gonapophyses. Abdominal extremities are present in the embryo on 
the eighth and ninth segments, but they later degenerate, and in old 
embryos as well as in young larvae leave not the slightest rudiments. 
It is only later, in older larvae, that the gonapophyses appear, and 
they are therefore undoubtedly to be regarded as secondary integu- 
mental outgrowths.” Thus, according to Heymons, the stylus and 
not the gonapophysis is the representative of the telopodite in an 
abdominal appendage. 

It must be conceded that the facts of embryonic development do not 
necessarily recapitulate phylogenetic evolution, since we can never 
be sure that the early stage of an organ reproduces the primitive form 
of that organ, and this must be particularly true of a rudimentary 
structure. Thus, if the telopodite of a limb bearing a basal exite 
process has long been lost, the limb rudiment in the embryo might be 
supposed to consist of the limb basis and the accessory process, and 
to lack the telopodite element entirely. Hence, while Heymons’ 
evidence of the nature of the styli is highly suggestive that the styli 
are the rudiments of the telopodites, it does not demonstrate the 
point. On the other hand, Heymons’ line of reasoning concerning 
the gonapophyses makes it seem almost certain that the genital proc- 
esses are not the telopodites of the gonopods, but the facts of develop- 
ment can scarcely be taken as evidence that the gonapophyses do 
not belong to the genital appendages. It is amply proven in the Thy- 
sanura that the gonapophyses are processes of the gonopods, and in 


/ 


| No. 6 INSECT ABDOMEN—SNODGRASS It5 


ee 


the Orthoptera they are undoubtedly outgrowths of the parts of the 
definitive sterna derived from the bases of these appendages. 
Verhoeff (1903), retaining his former views on the homologies 


between the abdominal and thoracic appendages, but going more into 


detail, proposed, on theoretical grounds, that the stylus-bearing plates 
of the insect abdomen represent the coxae, which in the abdomen 
he distinguished as “ coxites,” or in the genital segments as “ gono- 
coxites.” According to Verhoeff’s theory, the telopodites are lost 
from all the abdominal appendages except those of the eighth and 
ninth segments, where they become the gonapophyses; the styli are 
secondary lateral outgrowths of the coxae, equivalent to the coxal 
spurs of the thoracic legs of Machilis; and the eversible vesicles are 
median coxal structures comparable with the coxal glands of Dip- 
lopoda. 

Borner, though at first taking Heymons’ view of the nature of the 
gonapophyses, later (1904) agrees with Verhoeff that the genital 
processes represent the telopodites of the abdominal appendages, pre- 
served only on the gonopods. Regarding the supporting plates, how- 
ever, Borner differs from Verhoeff in that he identifies them as 
“basipodites,” meaning by this term that each plate is the equivalent 
of the coxa and subcoxa of a thoracic leg. (The same idea concerning 
the nature of the basal plates is followed in the present paper, but 
Borner’s term “ basipodite” is replaced with “limb basis” to avoid 
confusion with the more usual application of the other word to the 
first trochanter. ) 

Silvestri (1903, 1905) regards the basal segment of an arthropod 
limb as being in all cases a subcoxa (including the so-called coxop- 
odite of Crustacea), and he would divide the appendage into a basis 
(subcoxa) and a telopodite at the subcoxo-coxal joint. Applying this 
interpretation of the basal structure of the limb to the abdominal 
appendages of insects, Silvestri (1905) identifies the stylus-bearing 
plates of the Thysanura with the subcoxae. The styli he regards as 
the rudiments of the telopodites, with their bases representing the 
coxae. Silvestri, therefore, admits no homology between the abdominal 
styli and those of the thorax in Machilis; the leg styli he claims are 
secondary outgrowths of the coxal integument. Verhoeff (1903) had 
figured a coxal muscle attached to the leg stylus of Machilis, but 
this supposed muscle Silvestri shows does not exist—an observation 
in accord with statements by earlier as well as by subsequent writers, 
and one easily verified. 

The most interesting feature in Silvestri’s interpretation of the 
morphology of the abdominal limbs is his proposal that the genital 

8 


‘ 





116 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


apophyses are serially homologous with the eversible sacs of the pre- 
genital segments. Silvestri argues as follows: On the first abdominal 
segment of Projapyx and Anajapyx there are borne on each sub- 
coxa (stylus-bearing plate) a typical stylus and, mesad of it, a 
cylindrical or conical process. In Machilis and Nicoletia each sub- 
coxa of the first segment has a retractile vesicle, and in Campodea 
only a cylindrical process. The following six segments of Machilidae 
bear on the subcoxae both styli and vesicles, but on the next two, the 
genital segments, each subcoxa has a stylus and, in some genera, mesad 
to it a genital process. Thus Silvestri contends that the gonapophysis 
is evidently an eversible sac permanently everted. Muscles he ob- 
serves are attached to each appendage, though he does not point out 
that those of the gonapophysis are inserted on the base of the process, 
while those of the vesicles traverse the organ to be inserted in its 
extremity. 

The abdominal appendages of insects have not lacked attention 
from students of arthropod phylogeny, because their several parts 
make up a composite limb pattern that may be supposed to conform 
with the biramous structure of crustacean appendages, and thus 
indicate either that insects are closely related to the Crustacea, or that 
the primitive arthropod limb was a biramous structure. Writers who 
espouse the idea of a crustacean ancestry for insects, following 
Wood-Mason (1879), interpret the stylus as the exopodite, and find 
the homologue of the endopodite in the gonapophyses of the genital 
appendages. The theory must assume that the endopodites have been 
suppressed on the pregenital segments, since the eversible vesicles 
are eliminated as possible telopodite homologues by the fact that 
they sometimes occur in duplicate. 

The theoretical possibilities of aligning the appendages of insects 
with those of Crustacea have been exhaustively searched by Cramp- 
ton. In a study of the terminal appendages of the tridactylid orthop- 
teron, Ellipes, Crampton (1921) adduces evidence that he takes 
to be conclusive of the biramous nature of insect appendages. The 
dorsal pair of terminal appendicular processes in the Tridactylidae 
are undoubtedly the cerci (fig. 45 A, B, Cer) ; the ventral pair (paptl) 
are the lobes of the paraprocts (‘‘ paraprocessi”). After removing 
the end of the abdomen and spreading the parts out from below until 
they lie in one plane, Crampton makes a comparison of the tridactylid 
appendages in this position with the uropods of an isopod crustacean 
in the normal position (fig. 45 C, D), and arrives at the conclusion 
that the cerci of the former correspond with the endopodites of the 
latter, and that the paraproct lobes of Ellipes represent the exopodites 





no. 6 INSECT ABDOMEN—SNODGRASS 117 


of the isopod. The paraprocts themselves Crampton regards as the 
protopodites (i. e., combined coxopodites and basipodites) of the 
terminal appendages. The interpretation of the paraproct lobes as 
exopodites is consistent with other evidence that the styli are exop- 
odites, for the paraproct lobes fall in line with the styli, or would 
do so if styli were present in the tridactylids, and the abdominal styli 
are serially continuous with the thoracic styli of Machilis, which 
appear to have an exopodite status, and therefore suggest that the 
entire series of styliform organs are exopodites. 

There are several weak places in the above line of reasoning. In 
the first place, the writer fails entirely to get Crampton’s view from 
the comparison between the tridactylid and the isopod, since, with 
the terminal parts of both in the normal condition (fig: 45 B, D), 
the cercus of the insect (Cer) surely has the position of the exopodite 
(Expd) of the crustacean uropod, while the paraproct lobe (papil) 
corresponds in position with the small endopodite (Endpd) borne 
by the basal plate of the uropod (C). In the second place, a more 
careful examination of details shows that the cercus (B, Cer) has 
no anatomical relation with the paraproct (Papt), being situated 
dorsad of the latter in a position corresponding with that of the base 
of the crustacean uropod (D). Furthermore, as has already been 
shown, there is no evidence whatever to support the idea that the 


-paraprocts of insects are parts of the appendages. Their musculature 


indicates that they are mere lateroventral, subanal lobes of the eleventh 
abdominal sternum. The cerci have no muscles arising in the para- 
procts. Finally, the embryological evidence concerning the nature 
of the cerci appears to show definitely that the cerci are the entire 
appendages of the eleventh segment, and that their bases, if present 
at all, are retained in a basal ring of each organ. Hence, until some 
radically new information comes to light concerning the cerci, there 
is no question of exopodite or endopodite connected with them. Our 
present information is to the effect simply that the cerci are the appen- 
dages of the eleventh abdominal segment. 

The lobes of the paraprocts, whether the ‘“ paraprocessi”’ of the 
tridactylids, the small lobate ends of the paraprocts of the Anisoptera, 
or the lateral gill plates of the Zygoptera, have no validated claim 
to an appendicular origin. They must, then, for the present be re- 
garded as secondary outgrowths of the subanal lobes of the sternum 
of the eleventh abdominal segment, comparable to the various median 
outgrowths of the supra-anal plate of the same segment. 

The most nearly convincing evidence of the biramous nature of 
insect appendages is, admittedly, the presence of styliform processes 


‘ ’ 


118 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


on the mesothoracic and metathoracic legs of Machilis closely resem- 
bling the styli of the abdomen. Without this coincidence, or if 
Machilis and its coxal spurs had not survived to modern times, it is 
doubtful if entomologists would ever have thought of regarding the 
abdominal styli or the cerci as other than direct rudiments of legs. 

Conclusions —The abdominal appendages of insects are rudimen- 
tary limbs. Each consists of a basis and usually one or more distal 
appendicular parts, including a stylus, an eversible or retractile vesicle, 
and a gonapophysis. From the facts known of the comparative struc- 
ture of the abdominal appendages, and from theoretical considera- 
tions we may draw the following tentative conclusions relative to the 
homologies of the parts of the appendages, but it must be admitted 
that the evidence at hand is not sufficiently definite to establish any 
particular view concerning them. 

The limb bases of the abdominal appendages are the lobes or plates 
of the walls of the body segments that support the appendicular proc- 
esses. They are usually well developed in larval insects, but in most 
adults they are partially or wholly united or blended with the sternum, 
or in the male genital segment fused also with the tergum; in the 
eleventh segment they are reduced to small basal rings of the cerci, 
or entirely obliterated. Generally there is no distinction between coxal 
and subcoxal regions in the limb bases of the abdomen. In position 
on the body the abdominal limb bases usually fall in line with the 
subcoxae or pleural areas of the thorax. But since there is no 
apparent reason for the development of large subcoxal plates on 
the abdominal segments, such as those of the pterygote thorax, 
it is not to be supposed that the limb bases of the abdomen repre- 
sent the subcoxae alone. It is more probable that their principal parts 
are derived from the flattened coxae, or that the structures in most 
cases may represent primitive limb bases undifferentiated into coxae 
and subcoxae. In the caterpillars and sawfly larvae, however, the 
principal segment of each abdominal leg appears to be the coxa, which 
is quite distinct from a subcoxal lobe of the wall of the body segment 
to which it is attached. The abdominal limb bases are seldom movable 
on the body, since they commonly lack muscles corresponding with 
the basal muscles of the gnathal and thoracic appendages arising on the 
body wall. Exceptions to this rule are found in the male genital seg- 
ment, where the bases of the gonopods are occasionally provided with 
muscles and are independently movable ; but in such cases it is to be 
suspected, at least, that the muscles are secondary developments. 

The styli, or other homologous appendicular processes of the ab- 
domen, are of wide occurrence in insects, and serve a variety of 


¥ 






no. 6 INSECT ABDOMEN—SNODGRASS 11g 


functions, to which they are adapted by structural modifications. The 
styliform type is not necessarily the primitive form of the organs. 
The abdominal styli may be serially homologous with the thoracic 
coxal styli of Machilis, or the latter may be merely large spurs re- 
sembling the abdominal styli, from which they differ in lacking mus- 
cles. The abdominal styli are individually movable on the limb bases 
by muscles arising in the latter. If they are not the rudimentary main 
shafts of the abdominal limbs, they are exite lobes of the coxae analo- 
gous with the epipodites of crustacean appendages. They are not 
comparable with the exopodites of Crustacea, because the exopodite 
branch of a typical biramous limb is an exite of the first trochanter, 
or basipodite. 

There are many reasons for regarding the abdominal styli or their 
derivatives in insects as the rudimentary telopodites of the abdominal 
appendages. The styli seldom lose their muscles, except when they 
are immovably united with the bases; in some insects they take an 
active part in locomotion; they may be jointed in a manner sug- 
gesting at least a true segmentation, and in the larvae of Sialis they 
have intrinsic muscles in their basal segments. The styli of the gono- 
pods in male pterygote insects, especially in the holometabolous orders, 
are commonly modified to serve as grasping or clasping organs during 
copulation. The styli are the most generally persistent of the distal 
parts of the abdominal appendages. If it were not for their likeness 
in apterygote and orthopteroid insects to the coxal spurs of Machilis, 
it seems doubtful if the abdominal styli would ever have been regarded 
as anything else than the rudimentary telopodites of the abdominal 
appendages, represented in a similar form by the cerci on the eleventh 
segment. 

The vesicles of the abdominal appendages of Apterygota, the gill 
tubercles of the larva of Corydalus, and the plantar lobes of the larval 
abdominal legs of Lepidoptera and chalastogastrous Hymenoptera are 
all organs of a similar and unique type of structure. They are essen- 
tially exserted or invaginated lobes of the coxal areas of the limb bases 
lying mesad of the bases of the styli, and are retractile by muscies 
inserted within their distal parts. In the case of the Apterygota the 
muscles arise in the limb bases; in the others they arise from the 
lateral walls of the body. We might, with Verhoeff, regard these 
sacs as derivatives of coxal glands, since integumentary glands some- 
times take the form of eversible and retractile pouches. The coxal 
vesicles, however, serve a variety of purposes, and they are more 
simply explained as endite lobes of the coxae, which in some cases 
have become normally invaginated. They may thus be likened to the 


¥ 


I20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 





endite lobes of the gnathal appendages, and, as the latter, they some- 
times occur in duplicate on each appendage. 

The gonapophyses are hollow, median outgrowths of the bases of 
the gonopods, developed in both male and female insects during 
postembryonic stages. They are movable by muscles arising in the 
limb bases, or in areas of the genital segments derived from the latter, 
except where they are operated by the segmental muscles of the body. 
Theoretically the gonapophyses may be supposed to be either the 
telopodites (endopodites) of the gonopods, or endite lobes of the 
bases of these appendages. There are several objections to the first | 
view. The gonapophyses, for example, are never truly segmented, and 
never have a form suggestive of a leg structure; they occur only 
on the appendages of the genital segments, unless the cylindrical : 
processes of the first abdominal segment in certain Dicellura are | 
homologous structures; and finally, they serve only in a particular | 
capacity in connection with oviposition and copulation, except in | 
insects where they are secondarily adapted to form a stinging organ. 
These facts, together with the invariable median position of the 
gonapophyses on the gonopod bases, suggest that the gonapophyses 
are basal endites of the gonopods, movable by muscles arising in the 
basis, as such endites usually are, and specially adapted to the repro- 
ductive functions. The genital apophyses might then be regarded as_ | 
serially homologous with the eversible or retractile vesicles of the 
pregenital appendages and certain larval pygopods, in which case 
Silvestri’s idea that they represent “ permanently everted eversible 
vesicles” is better stated in the reverse, namely, that the eversible 
vesicles are inverted gonapophyses. However, the vesicles do not 
occur at points on the gonopod bases corresponding with the origins _ 
of the gonapophyses, and the musculature of the two sets of organs 
is characteristically different. It is perhaps possible that the gona- 
pophyses are subcoxal endites, and the vesicles coxal endites. 

If none of the appendicular processes of the abdominal limps can _ 
be satisfactorily identified with the telopodite of a primitive appendage, 
we must conclude that the abdominal telopodites have been lost from 
all but the terminal segment, where they form the cerci, and that 
the various persisting appendicular structures are accessory processes 
of the limb bases. Otherwise, we must choose between the gona- 
pophyses and the styli as possible representatives of the telopodites. 
Of the two, the styli certainly present better credentials, considering 
their occurrence on many segments of the abdomen as contrasted 
with the segmental localization of the gonapophyses, their structural 
and functional versatility by comparison with the limitations of the 


a 


a a i ny 


no. 6 INSECT ABDOMEN—SNODGRASS 121 


genital apophyses in form and use, and their leglike relations to the 
limb bases as opposed to the median, proximal origins of the gona- 
pophyses. All these points qualify the abdominal styli for true limb 
rudiments, and give the genital processes the status of basendites 
specially developed for reproductive purposes. The pretensions of the 
abdominal styli to telopodite origin are opposed only by their similar- 
ity to the thoracic styli of Machilis; but there is nothing to show 
that these leg structures are not mere coxal spurs resembling in form 
but fundamentally unlike the musculated appendages of the abdomen, 
the styliform shape of which is but one of their many structural 
adaptations. 

A discussion of the phylogeny of insects, or particularly of the 
possible origin of insects from any other group of existing arthropods 
is beyond the intended scope of the present paper. A recent work by 
Tillyard (1930) on the evolution of the Insecta, though somewhat 
partisan in favor of myriapodan descent, gives many reasons for 
believing that insects are not directly related to the Crustacea. And 
yet, the weight of evidence, whether put forth by claimants of a 
myriapodan or a crustacean ancestry for insects, seems to depend 
largely on minimizing or disqualifying the evidence on the other side. 
However, if we were to give equal weight to arguments on both sides 
of the question, the insects would be cut off from all ancestral ties, 
and thereby deprived of a respectable pedigree—unless they are able 
to take care of themselves through all the unknown ages of time 
before they are first known to us as fully-winged hexapods in the 
Carboniferous deposits. To the writer it appears that all the principal 
arthropod groups must represent independent lines of descent from 
some remote ancestral forms embodying the potentialities of a spider, 
a crab, a centipede, or an insect. It has recently been emphasized 
by Clark (1930) that the chronic inability of the evolutionary theory 
in its usual form to explain the lack of intermediates between the 
major groups of animals constitutes a real weakness of the theory, 
which calls for a new concept of the method by which distinct types 
of organisms have been produced. The condition to which Clark 
refers is well exemplified within the Arthropoda, where connective 
forms between the classes are unknown. Moreover, it is impossible 
to construct imaginary arthropods that will fill the blanks, as, for 
example, the three-cornered gap between the crustaceans, the myria- 
pods, and the insects. Considering that embryos develop before 
our eyes by ways that are still inscrutable, it takes a strong faith in 
established ideas to believe that organic evolution has proceeded 
entirely by the means we have furnished for its guidance. 


122 


ABBREVIATIONS USED ON 


a-a, dorso-pleural groove. 

a-b, axis of articulation of limb base 
on body. 

ab, abductor muscle. 

Ac, antecosta. 

acs, antecostal suture. 

ad, adductor muscle. 

Aed, aedeagus. 

An, anus, 

Ap, apodeme. 

Apd, segmental appendage. 

Apt, sternal apotome. 


Bnd, basendite. 

Brn, branchia (gill). 

brncls, branchial muscles. 

Bspd, basipodite (first trochanter ). 


c-d, axis of subcoxo-coxal joint, 
Cer, cercus (uropod). 

cf, caudal filament. 

Col, collophore. 

Con, nerve connective. 

ct, coxo-trochanteral joint. 

Cx, coxa. 
Cxpd, coxopodite. 


D, dorsum. 

d, dorsal muscles. 

del, lateral external dorsal muscles. 

dem, median external dorsal muscles. 

dil, lateral internal dorsal muscles. 

dim, median internal dorsal muscles. 

dl, dilator muscle. 

dm, median dorsal muscles. 

DMcl, longitudinal dorsal muscles, 

DTra, dorsal longitudinal tracheal 
trunk. 


Endpd, endopodite. 
Eppt, epiproct. 
Expd, exopodite. 


f-g, axis of baso-telopodite joint. 
Fm, femur (meropodite). 

ft, femoro-tibial joint. 

Fur, furcula, 


SMITHSONIAN MISCELLANEOUS COLLECTIONS 





VoL. 85 


THE FIGURES 


gmcls, muscles of gonapophysis. 
Gng, ganglion. 

Gon, gonapophysis. 

Gp, gonopod. 

GSeg, genital segment. 


Hpn, hypandrium. 
Ht, heart. 
Flypt, hypoproct, 


/, tergal promotor muscle of appen- 
dage. 

ip, intestinal process. 

/sg, intersegmental fold. 

/st, intersternite. 


J, tergal remotor muscle of appendage. 
k, interfurcal sternal ridge. 


L, leg. 

!, lateral body muscles, 

Ja, lamina subanalis. 

LB, limb basis. 

le, external lateral muscles. 

li, internal lateral muscles. 

lStn, last pregenital sternum. 

IT, last pregenital tergum. 

LTra, lateral “longitudinal 
trunk, 


tracheal 


Mb, intersegmental membrane (con- 
junctiva). 

mb, membrane. 

Mcl, longitudinal muscles. 

mcls, muscles. 

mn, manubrium. 


NIG, neural groove. 


O, levator muscle of first trochanter. 
Ovd, oviduct. 


p, paratergal muscle. 
Papt, paraproct. 

paptl, lobe of paraproct. 
patg, paratergite. 

Pc, precosta. 





NO. 6 


Pen, penis. 

Ph, phragma. 

Pl, pleuron. 

pl, pleurite. 

Pm, paramere (male gonapophysis of 

ninth segment). 

meta-postnotum (precosta of 

first abdominal segment). 

pnb, pons basalis. 

Pp, pygopod. 

Prpt, periproct. 

Prtpd, protopodite (united coxopodite 
and basipodite). 

Ptar, praetarsus (dactylopodite). 


PNs, 


O, depressor muscle of first trochanter, 


rvs, retractor muscles of vesicle of leg 
base, or of its derivatives. 

rvsd, dorsal retractor of planta. 

rusv, ventral retractor of planta. 


S, definitive sternum (usually includ- 
ing limb bases). 

SAsz, metasternal apophysis. 

sa, lamina supra-analis. 

Sex, subcoxa. 

Sp, spiracle, 

s-p, sterno-pleural muscle. 

smcls, muscles of stylus, or of its de- 
rivatives. 

Stn, primary segmental sternum. 


INSECT ABDOMEN—SNODGRASS 123 


Sty, stylus of leg base, or its deriva- 
tives. 


T, tergum. 

Tar, tarsus (propodite). 

Tb, tibia (carpopodite). 

td, dorsal transverse muscles, 

Tel, telson. 

Tlpd, telopodite. 

t-p, tergo-pleural muscle. 

Tr, trochanter. 

1Tr, first trochanter (basipodite). 
2Ty, second trochanter (ischiopodite). 
Tra, trachea. 

t-s, tergo-sternal muscle, 

tv, ventral transverse muscles. 


ug, urogomph. 


I’, venter. 

v, ventral muscles. 

vel, lateral external ventral muscles. 

vem, median external ventral muscles. 

vil, lateral internal ventral muscles. 

zim, median internal ventral muscles. 

V1, valvula. 

Vif, valvifer. 

V Mel, ventral longitudinal muscles. 

V NC, ventral nerve cord. 

I’s, vesicle of limb base, or its deriva- 
tives. 


1-3, thoracic segments. 
I-XIT, abdominal segments. 


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} 














a a 





or 
a 





SMITHSONIAN MISCELLANEOUS COLLECTIONS 
VOLUME 85, NUMBER 7 


EFFECTIVENESS IN NATURE OF THE SO-CALLED 
PROTECTIVE ADAPTATIONS IN THE ANIMAL 
KINGDOM, CHIEFLY AS ILLUSTRATED BY 
THE FOOD HABITS OF NEARCTIC BIRDS 


BY 
W. L. McATEE 
Bureau of Biological Survey 
5. 


Wis epartment of Agriculture 





(PUBLICATION 3125) 


GITY OF WASHINGTON 
PUBLISHED BY THE SMITHSONIAN INSTITUTION 
MARGH 15, 1932 





SMITHSONIAN MISCELLANEOUS COLLECTIONS 


ViGl..85; INO:\7 


EFFECTIVENESS IN NATURE OF THE SO-CALLED PROTECTIVE 
ADAPTATIONS IN THE ANIMAL KINGDOM, CHIEFLY 
AS ILLUSTRATED BY THE FOOD HABITS OF 
NEARCTIC BIRDS 


By W. L. McATEE 
ERRATA 


Page 56. Inthe table Identifications of Lepidoptera, the middle column is a relic 
from a set of calculations of the percentages of identifications among 
all insects. The appended figures are to be substituted as represent- 
ing the percentages of identifications among all Lepidoptera. In 
explanation of the third column in this table, it may be said that it 
differs from others given further on in the work by omission of 
figures for families not represented among the food identifications. 


Percentage of 
identifications 
among all 
Lepidoptera 
2.6992 
.0270 
.0270 
8060 
.OO19 
.2055 
.O108 
.0270 
0108 
.0108 
-4700 
.0324 
.O412 
.0378 
6383 
6.1015 
.0216 
3078 
-3570 
1508 
8438 
11.4458 
25.3300 
.0487 

0054 
GLOL 
.0054 
-O108 
2218 
0112 
7248 
68.5670 
1.2279 
3.5376 


Page 86. The figure 1 before the decimal in the entry for the family Diopsidae 
should be deleted. 

Pages 102-105. Insert the word “aquatic ” after the word “all” in the heading 
for the middle column on each of these pages, with the exception of 
that at the bottom of page 105. 





mePrPECTIVENESS IN NATURE OF -THE SO-CALLED 
BROLteGhivi ADAPTATIONS IN THE ANIMAL 
RINGDOM, CHIEFLY AS ILLUSTRATED BY THE 
BOOD HABITS OF NEARCTIC BIRDS 


By W. L. McATEE 
BUREAU OF BIOLOGICAL SURVEY, 
U. S. DEPARTMENT OF AGRICULTURE 


CONTENTS chee 
BMG TIGL ONY mhrae arse citer enn are a csiske ie DEE weed Shiale mn sey Ea em edvelee eS 
WMBrotective adaptations ............ecececececcccccccccececuceeevuccuces 4 
Banimalsneaten by, WNearctie birds. ccs. acces cee vw sce es setae oe ccn ae es 6 
Datarcited andehow. Obtained... 2 ccs sees. oes tee seems sessed cones 6 
IG@entinicatlons Ol vanimeals 100d es. ec Oleg. fee os eo eee cise esse cer at mance 8 
Protozoa (one-celled animals)................. cece ee cece cease 9 
I GRIeT deve OMONEES ) a atrsc wise eS aes aks ¥ <8 dime bate au she wae neg 10 
Coelenterata (hydras, jellyfishes, sea-anemones)................. II 
Platyhelminthes (flatworms, flukes).............. 0000 c cece eeees 13 
Nemathelminthes (threadworms, roundworms).................. 13 
aliochelminthes u(rotikers)) se s.40s «ese cece sions sea cece nee ae 13 
Molluscoida (corallines, lampshells)................ 000 cece eee 13 
Echinodermata (sea-cucumbers, sea urchins, starfishes)........... 14 
PTV ia Cea tcta am (GVO LTIIS)) 8 repehey er eyete eye arenes sierra onsies eevee Sines Se ra ce Anesth Rhy 15 
Arthropoda’ (jointed “animals)..22 2.2.22. 0.0<s..02 ste. st eeene 16 
@rustaceas(crabs) shrimps, SOWwbUeS )\: g0. 42.2 ce cee bone ees i 
Myriapoda (thousandlegs, centipedes)....................5- 22 
SEC tame CITISECESD) pameittepiere etek niekcieds aitieys chs. dle'csurare 2% 0 o.shae. ote 2 
EN Puc hate (AWA C1 eSS)ettISECES!) penton n oss sce cratic: <. osia ce aieuccelanci eBs 2 
Odonata (dragonflies, damselflies)...............0000005 28 
Nomeathiaa (may ilies) et wtatetaee suse a eae cess eect eee 20 
Plecoptera i(stonefiies)) 20.220... cae ce ce aecdssesss 30 
NSO teraen CLEtIILeS) geese wyatt ea yeneete stn daeserd,= a/steptonk 31 
Orthopteroidea (embracing the following 5 groups)...... 32 
WW ermaptena | Canwiesi ae aie ses ner ae © icacere esau eceeie oe BD 
Cheleutoptera (walleinestichks i. 22.siideawesiuws teen cae: 33 
Saltatoria (grasshoppers, locusts, crickets).............. 34 
| Paleo pretramn( HOACIES )maremitts suis acts octete otis sees eaaieea 38 
Dictyopteral (meantids)): sees s «sees wes sees eee ease oe 39 
| (Worrodentiay GpsocidS)) en. cec feeds ce es see cease. e eu: 30 
| Matlophaga (bilinear Hee) 22 cc diese rent debe encatoten ot 40 
Sipnoneaptencaem heas\mawera teas ese ceciaeete seen see eta: 40 
MDVSANOp Grae AiTipS) | etandhoes vnc Wie seated siete gas rw eee d 4o 
Rhynchota (bugs, cicadas, leafhoppers, scale insects)..... 4! 

Neuropteroidea (dobsonflies, snakeflies, scorpionflies, ant- 
Lions Caddishies i) a emisscee ees asic ead aage cence de cece 49 
Lepidoptera (moths, butterflies)................0..00 00 52 


SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 85, NO. 7. 


2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


PAGE 

Coleoptera’ (beetles): Seeion- tis 2m: steceitercls.- eee 63 

Meeaptera (scorpiontlies)) 2: asics sors 21 aleve elem ellen 84 

Diptera, (ilies) joceicaastpeisoed sto nie ta cee eee 84 

Hymenoptera’ (ants, ‘bees, wasps)!:.----.- ce «ee 88 

Arachnida (scorpions, spiders, ticks; ‘etC.)).. 2.0.2.2... 9. seeee 07 

Mollusca (snails, slags, mussels, limpets).....2 00-9. ----- eee IOI 
Chordata (lancelets, tunicates, vertebrates).............-+- scare 110 

Pisces: (fishes )* 5.02). ae staternctteten net aos ok on ee aie te eee 112 

Amphibia (salamanders, toads, frogs) .............+-sssesme 120 

Reptilia (turtles, lizards, snakes).........:.2.. ..-++s09ame 123 

Aves iCbitds))! (fs tadten 2nGoe wee rte ictus 6 Soe oni eee 126 

Mammalia’ (mamimeals)! 2.55 2c clele es = aleiele 2 oye eves ee 131 

IDF orci SREP ASAeUOM cH oddone dnadinenoccubsono aos moggDoowonedooococs 135 
Indiscriminancy of predators other than birds.................se.m0e 136 
More theoretical aspects of indiscriminancy by predators............. 140 
Indiscriminancy of normal checks other than predators.............+. 141 
Relative importance of natural’ checks.2..0c 2. s.ieeas + ee eee 141 
Summary —acax ews ore Sos o cies etolsias slnmtare\ercrers seta alee ia tt te 143 
Bibliographiy: << woes cis niet mocteraieve, ote olen tet sieve piers ieee) wate eva eta ea 145 


INTRODUCTION 


In a previous paper’ the writer set forth reasons for believing 
that the results of experimental tests of the effectiveness of the so- 
called protective adaptations in protecting animals from their enemies 
are not trustworthy indications of what occurs under natural condi- 
tions. In the present contribution he proposes to show just what 
insects and other animals are actually preyed upon by wild birds 
of the United States, Canada, and Alaska, giving also incidental 
notes on other enemies. This evidence reflecting food habits under 
natural conditions goes far to show how little the alleged protective 
devices have to do with choice of food by vertebrates. 

Judging from the literature of the subject since 1912, the con- 
tentions of the article on the experimental study of the food habits 
of animals seem to have been generally admitted, or at least regarded 
as too well supported to be lightly attacked. Only one essay has been 
seen by the writer, that seems in any way a reply, namely an account 
of “ Experiments and Observations Bearing on the Explanation of 
Form and Colouring,” * by C. F. M. Swynnerton, who refers to my 
criticism of the experimental method as “ rather over-vigorous.” The 
vigor of the criticism is admitted but in view of the absurdity of the 
arguments against which it was directed, it can hardly be considered 


‘The experimental method of testing the efficiency of warning and cryptic 
coloration in protecting animals from their enemies. Proc. Acad. Nat. Sci. 
Philadelphia, June 1912, pp. 281-364 (Sept. 6, 1912). 

? Journ. Linn. Soc., Zool., vol. 33, pp. 203-385, London, June 30, 1919. 








NO. 7 PROTECTIVE ADAPTATIONS—McATEE 3 


excessive. Undeniably selectionists have been absurd in their dis- 
quisitions on adaptations ; for instance “ eye-spots ” on a butterfly’s 
wings are to direct the attack of enemies to a nonvital spot, while 
“eye-spots”” on a caterpillar are “terrifying” and prevent even 
a touch where merely a touch would be fatal; in numerous species 
of birds the male is colored red and black or orange and black, 
characteristics that selectionists say have been developed by sexual 
selection as an attraction to the opposite sex, yet the females of 
these birds are supposed to be repelled by the same colors in possible 
insect prey ; red insects are said to be warningly, red fruits invitingly 
colored, and so on. A popular foible of similar type is that of sports- 
men who hold up to admiration the marvelous protective coloration 
of game birds, and in the next breath complain of severe depredations 
on these birds by “‘ vermin.” 

But this is digressing and the writer is glad to acknowledge that 
if all of the experimenters had been as critical of their methods and 
conclusions as Mr. Swynnerton, the tone of his former paper would 
have been quite different. For instance Mr. Swynnerton carried 
on more experiments than any of the authors reviewed in the previous 
communication, before he, according to his own confession, learned 
how to experiment. This in itself confirms the writer’s charges that 
the experiments he reviewed were both inadequate and misinterpreted. 
It may further be stated that the principal conclusions Mr. Swynner- 
ton draws from his experiments and observations would have been 
agreed to in advance by anyone experienced in the study of bird food. 
Thus he concludes that birds show preferences among the food items 
_ available to them, and that predatory animals of various groups show 
more or less agreement in preferences. From his general experience 
with birds he decides also that ‘‘ Unless through sheer impossible 
hardness, size, etc., there is practically no such thing as ‘ inedibility,’”’’ 
and he appreciates that a group of insects, limited in numbers as are 
butterflies, will not be taken by insectivorous birds out of proportion 
to their abundance as compared to all insects available. 

These things did not require experimental test for they are cor- 
roborated in every thorough report on the natural feeding habits 
of birds. What can not be admitted, however, is that preferences 
of birds learned by feeding them upon some certain group of insects 
to an extent far greater than the birds ever prey upon them in nature, 


‘ 


reflect normal feeding habits, nor that there is evidence of intensive 
enough feeding by discriminating enemies upon any group of insects 


*A brief preliminary statement of a few of the results of five years’ special 
testing of the theories of mimicry. Proc. Ent. Soc. Lond., 1915, pp. xxxii-xliii. 


4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


to meet the requirements of the selection theories. We further can 
not admit what the experimenters imply, namely, that the analyses 
of the stomach contents of birds fail to reveal the approximate num- 
bers present of certain insects (such as butterflies) which they believe 
are eaten to a considerable extent. This point will be discussed later, 

So much for what has happened between the previous paper and 
the present, which as stated, will be devoted chiefly to an exposition 
of the animal food of nearctic birds, with special reference to the 
so-called protective adaptations. 


PROTECTIVE ADAPTATIONS 


The characteristics of animals that are usually classed as protective 
adaptations include resemblance to generalities or details of the en- 


vironment, whether through color or other modification of the animal 


itself or utilization by it of materials from the environment for 
concealment, the possession of protective bristles, spines, hard in- 
teguments, stings, poisonous bites, and the like, and nauseous or 
irritating odors or tastes. There are animals with actually poisonous 
properties among many of the phyla including species with poison 
glands and special organs for using the poison in offense or defense, 
among Coelenterata, Echinodermata, Arachnida, Insecta, and Pisces; 
others with poison glands connected with the mouth organs among 
worms, spiders, other arachnids, mites, myriapods, chilopods, insects, 
fishes, and reptiles; animals with unarmed poison glands among 
coelenterates, echinoderms, myriapods, insects, mollusks, amphibians ; 
and others poisonous in a variety of ways so that practically all phyla 
are represented. The colors of the animals possessing dangerous 
qualities in many cases are said to be warning in nature, and the 
colors of animals which resemble them but lack the disagreeable 
qualities are termed mimetic. The subject of protective adaptations 
has very largely become one of coloration especially as associated with 
the qualities of animals from the supposed point of view of possible 
predators. 

A statement of the various classes of color adaptations is here 
quoted from Prof. E. B. Poulton, the leading advocate of the view 
that these adaptations are really protective and that they have been 
developed by natural selection. 

Protective and Aggressive Resemblance—By far the most widespread use 
of colour is to assist an animal in escaping from its enemies or in capturing its 
prey; the former is Protective, the latter Aggressive. It is probable that these 
were the first uses to which non-significant colours were put. The resemblances 


are of various kinds; the commonest cases are those of simple concealment. 
The animal passes undetected by resembling some common object which is of 











NO. 7 PROTECTIVE ADAPTATIONS—McATEE 5 


no interest to its enemies or prey respectively, or by harmonising with the 
general effect of its surroundings; the former is Special, the latter General 
Resemblance, and both may be Protective or Aggressive. Among the most 
interesting Special Aggressive Resemblances are the cases of Alluring Colour- 
ing, in which the animal, or some part of it, resembles an object which is 
attractive to its prey.’ 

Protective and Aggressive Mimicry.—Mimicry is in reality a very important 
section of Special Resemblance. The animal gains advantage by a superficial 
resemblance to some other, and generally very different, species which is well 
known and dreaded because of some unpleasant quality, such as a sting or an 
offensive taste or smell, &c., or it may even be protected from the animal it 
resembles: this is Protective Mimicry. When, however, the animal resembles 
another so as to be able to injure the latter or some other form which accom- 
panies it or is not afraid of it, the Mimicry is Aggressive? 

Warning Colours——When an animal possesses an unpleasant attribute, it is 
often to its advantage to advertise the fact as publicly as possible. In this way 
it escapes a great deal of experimental “tasting.” The conspicuous patterns 
and strongly contrasted colours which serve as the signal of danger or inedibility 
are known as Warning Colours. In other cases such colours or markings enable 
individuals of the same species easily to follow those in front to a place of safety, 
or assist them in keeping together when safety depends upon numbers. It is these 
Warning Colours which are nearly always the objects of Protective Mimicry.’ 


Following is a copy of Poulton’s table ‘ classifying color adaptations : 











I. Apatetic colours.— Il. Sematic Ill. Epigamic 

Colours resembling some part of the | colouwrs—Warning colours——Colours 
environment or the appearance of an- | and signalling col- displayed in court- 
other species. ours. ' ship. 

A. Cryptic col- | B. Pseudo- | 
ours.—Protective | sematic colours.— | 
and Aggressive False warning and 
Resemblances. signalling colours. 

1. Procryptic 1. Pseudapose- 1. Aposematic | 
colours——Protec- | matic colours.— colours—Warning | 
tive Resem- | Protective Mimi- | colours. | 
blances. | cry. 

2. Anticryptic 2. Pseudepise- 2. Episematic 
colours —Ag- | matic colours.— colours.—Recog- 
gressive Resem- Ageressive Mimi- | nition Markings. 
blances. ‘ery and Alluring 

Colouration. | | 





Having presented the foregoing outline of protective color and 
other adaptations, references to them in succeeding pages will be made 
without further explanation of the terms involved. 


* The colours of animals, pages 19-20, 1890. 
* Idem, p. 20. *Tdem, p. 21. *Tdem, p. 338. 


¥ 


6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 











ANIMALS EATEN BY NEARCTIC BIRDS 


Data CITED AND How OBTAINED ! 


The main body of data used herein consists of the records of 
animals identified in the contents of the stomachs of about 80,000 
nearctic birds examined in the United States Biological Survey since 
1885. These stomachs represent a wide range of species of all of 
the families of birds occurring in the region; the birds were collected 
at all seasons and in practically all parts of nearctic America. While 
not evenly distributed in any of these senses, the collection is very 
satisfactory for the purpose in hand, and yields a mass of precise 
information on bird food that far surpasses anything of the kind 
available elsewhere. 

A word about the methods of investigating bird food may be 
desirable. The gizzards of birds, together with the gullets or crops 
when they contain food, are received chiefly from persons collecting 
birds for some other scientific purpose, although in some cases they 
are especially obtained to throw light on the relations of birds to some 
crop, or useful or injurious animal. They are preserved usually with 
formalin in the field and in alcohol after receipt at the laboratory. 
Contents of a stomach being examined are removed either wet or dry 
as best fits the particular case and transferred to watch glasses or 
small white blotters for sorting and identification of the material 
under compound binocular dissecting microscopes. A great deal of 
the analysis is done at a magnification of 8 diameters but special 
study of difficult subjects is continued when necessary under higher 
powers. 

At this point it may be well to comment on the popular misconcep- 
tion that anything found in a bird’s stomach necessarily is ground 
up and in all but unrecognizable state. As a matter of fact the reverse 
is true. Most birds swallow their food whole ; consequently in any col- 
lection of birds a certain proportion will have swallowed some food 
items just before death. These things often are in perfect condition ; 
they may be, and sometimes are, used for cabinet specimens. The 
nearly or quite whole objects usually furnish clues to the fragmentary 
material, and in the great majority of cases it is possible to sort out 
completely all components of the food. It is the exception when the 
finely ground food remains defy separation and identification. De- 
terminations are carried as far as practicable; each member of the 
staff of analysts is a specialist in some line and they cooperate freely ; 
specimens defying their combined efforts, if in fair or better condi- 
tion, are submitted to advanced investigators elsewhere. The records 








NO. 7 PROTECTIVE ADAPTATIONS—McATEE 7 
quoted in the following pages include more or less of the handiwork 
of practically every prominent American systematic zoologist of the 
period. Nevertheless everything is not identified, far from it ; expert 
assistance has not been available in some cases when needed, too far 
digested residues sometimes occur, and rarely we find also well-pre- 
served but puzzling objects that indefinitely defy classification. 

These, however, are but minor flaws in the system; the ground- 
work of our faith in the results of stomach analysis is the law of 
averages. Given good distribution geographically and seasonally, 
which necessarily follows from miscellaneous collecting carried on 
for so many years, the reliability of results varies directly with the 
number of stomachs. The collection (80,000) here reported upon 
is believed sufficient to furnish fairly dependable data, although addi- 
tions are made almost daily to the list of animals identified from bird 
stomachs. 

The total number of identifications of animals from these stomachs, 
counting those of whatever degree, once for each time identified 
irrespective of the number of individual specimens concerned, is 
237,399. 

It was impracticable to compute the total number of individual ani- 
mals concerned for the reason that these were not counted in all 
cases. Moreover this figure would not have been especially useful 
in the absence of estimates for comparison of the actual animal popu- 
lation of significant areas. In casting about for a standard which 
would afford some idea of the frequency of occurrence of animals of 
various groups, the estimated number of species therein proved to be 
the only one available for the whole range of the animal kingdom. 
That the number of species in taxonomic groups bears a general re- 
lation to the number of individuals can not be questioned. It is easy 
to point out exceptions, but remember we can only deal with this 
problem in an approximate way, and it goes without saying that on 
the average a group more numerous in individuals will have devel- 
oped more species than one less numerous. The correspondence is 
not exact, but it is sufficient to give a fair working idea of the position 
of the various groups in the scale of frequency of occurrence, the 


* The tabulation necessary to yield this figure was an immense one (covering 
nearly a thousand typewritten pages) and has been found, it is not surprising, 
to contain some errors. These are so small, however, that rectification of them 
would not cause changes of more than a fraction of one per cent in any part 
of the results, except in the table for Coleoptera, pp. 65-67. Hence they do not 
invalidate the figures at all for the purpose here used of showing in a general 
way the tendencies exhibited by our birds in their choice of animal food. 


v 


8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


very standard we wish for comparison with frequency of identification 
in the stomachs of collected wild birds. The general correlation of 
these factors to be noted in the tables presented in subsequent pages 
increases confidence in the value of the method.’ 


IDENTIFICATIONS OF ANIMAL Foop 


To illustrate the way in which data was assembled, that for the 
phyla may be given in rather more detail than is planned for the 
balance of the report. The figures for number of species in the various 
phyla are compiled from various estimates * of this nature; the facts 
that these are not strictly up-to-date nor anything like exact are of 
no consequence in a field where only approximations may be hoped for. 

The subjoined table shows the estimates used for the number of 
described species in each phylum and the percentage that figure bears 
to the total number of animals known. 


Phyla of Animals and the Number and Percentage of Species in Each 


Percentage 


of species 
Estimated in this 
number of phylum 

species among the 

Phylum known whole number 

ProtozOals manna asia OEE ome arose 8,000 1.4272 
PORUEET areeva shy vee ease cree rece aero tebe: 2,500 .4400 
Coelenterata: 225 sce ake oto aero 4,500 8028 
Platyhelminthes werner ose aes 5,000 .8920 
INemathelminthesy saeeciaenineseeee rec 1,500 .2076 
Erochel minthes sy ere lee caress ake 500 0892 
Moliitiscoidameereesce teeter eon: 1,700 3032 
Echinodermata seni. de cies acer 4,000 -7136 
Aninitilatay 632 Asem een Sct cee Ghinee 4,000 7136 
ATT OPOddimn ean clten cone or nee 418,250 74.6188 
Molluscapce ence satya ec eer 61,000 10.8828 
Chordata? sa. .ures sehen en ae eee eee 49,505 8.8427 
AROtallsingersetn eos ee oe Cite acer 560,515 99.9995 


“Here may be mentioned the law demonstrated by Olaf Arrhenius (Journ. 
Ecol., vol. 9, no. 1, p. 99, Sept., 1921) that among plants ‘ The number of species 
increases continuously as the area increases.” Since as a rule the number of 
individuals also increases with the area, the parallelism between the number 
of individuals and that of species is further confirmed. 

* Pratt, H. S., On the number of known species of animals. Science, vol. 35, 
pp. 467-468, March 22, 1912. 

Henshaw, H. W., Number of species of living vertebrates. Science, vol. 36, 
pp. 317-318, Sept. 6, 1912. 

Handlirsch, A., Die fossilen Insekten, pp. 1182-1188, 1908. 





NO. 7 PROTECTIVE ADAPTATIONS—McATEE 9 


Classifying the 237,399 identifications of the animal food of 
nearctic birds and calculating the percentage of the determinations, by 
phyla, we reach the results shown in the next table, the percentage 
of species in each phylum among the whole number of known species 
being repeated for ease in comparison. 


Identifications of Animal Food by Phyla 


Percentage 





of species 

in this 

phylum 
* among the 

Percentage of whole 
identifications number of 

among those animal 

Number of of all species 

Phylum identifications animals known 

PeVOCOZOA: ss cess cece set seee I2 .0050 1.4272 

IROTITEra 26. sees. sss ene os 2 .0008 .4460 
@oclenterata (2.04.42... 122 .O514 8028 
Nemathelminthes ......... 24 .OIOL 2676 
Molluscoida ~ ...6...ssse0% 134 0564 3032 
Echinodermata ........... 125 .0526 -7136 
Aminulata ..sccsiecsescses 1,131 4764 7136 
mrthropoda .........+..--. 210,752 88.7751 74.6188 
IMMOINISCA, .0 5c. oe cece eee 1770 4.9583 10.8828 
| Ghordata ce. hedics ceca von 13,326 Biobes 8.8427 


Without going into details, it is apparent that the percentage of 
| identifications preserves very well a relative ratio to that of the num- 
_ ber of species and presumably, therefore, to the abundance of indi- 
viduals in the phyla. What variations there are seem obviously due 
_ to differences in the availability to birds of the differing types of 

animals. 

Taking up the phyla in order, we begin with the 


PROTOZOA (ONE-CELLED ANIMALS) 


Protective adaptations —Judging from what is asserted about other 
phyla, phosphorescence and the possession of bright colors in some 
groups and of silicious or calcareous, often tuberculate or spinose, 
tests or shells or of exoskeletons formed of foreign bodies in others, 
are characters that would be deemed of protective significance in 
Protozoa. 

Bird enemies.—Protozoa are too small to engage the attention ot 
birds, those found in stomachs being Foraminifera strained from 
water or mud, or picked up as gravel by ducks. It is probable also 
that protozoa are consumed, along with the stems and leaves of 
aquatic plants upon which they often are abundant, by wild ducks 


® 


1D 


IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


which feed upon such vegetation. Stomach analysis, however, has 
not been directed toward the recognition of such minute material. 

Number of identifications, 12; percentage of identifications among 
those of all animals, .o050 ; percentage of species in this phylum among 
the whole number of animal species known, 1.4272. 

Other enemies.—Protozoa are the prey of others of their kind— 
of bacteria, of rotifers, of flatworms, of amphipods and other small 
crustacea, and of mollusks; they are eaten also by the young of 
numerous species of fishes, by the adults of specialized forms (men- 
haden, gizzard shad), and by the larvae of batrachians. 

Discussion.—Protozoa, because of their minute size and general 
inaccessibility to birds, would not be expected to enter largely into 
the food of this class. The forms eaten by birds are among the best 
“protected” protozoa, but the possession of shells can hardly be 
considered as an adaptation for protection from enemies in the case 
of animals so small as to be easily devoured by almost any carnivorous 
animals encountering them and which exist in such enormous numbers 
that vast areas of sea bottom are covered with remains of their 
shells. In this case as in many others, numbers are so large and re- 
production so great that the inroads of all enemies are fully dis- 
counted. Losses to predatory enemies are only a fraction of the total 
death rate. 


PORIFERA (SPONGES) 


Many sponges are pervaded by calcareous or silicious spicules 
which may render them more or less undesirable as food for pre- 
datory animals. Some are brightly colored and some phosphorescent. 
‘Sponges do not appear to be edible by Fishes or even the higher 
Crustaceans or Molluscs. Countless lower animal forms, however, 
burrow in their substance, if not for food, at least for shelter, and 
the interior of a sponge is frequently found to be teeming with 
small Crustaceans, Annelids, Molluscs and other Invertebrates.” * 

Sponges have been identified from only 2 stomachs of nearctic 
birds (Canada goose and lesser scaup) and from their low degree 
of accessibility to birds, not many cases of feeding upon them would 
be expected. 


1Entries under this head for the various groups treated are intended as 
suggestive rather than as exhaustive. A list of papers from which much of 
this information has been gleaned forms the special bibliography on pp. 145-201. 
Notes on the food of reptiles, amphibians, and mammals are mostly from 
analyses of stomach contents in the Biological Survey. 

? Parker, T. J., and Haswell, W. A., A text-book of zoology, vol. I, p. 126, 
IQI0. 





NO. 7 PROTECTIVE ADAPTATIONS—McATEE Ve 


Number of identifications, 2; percentage of identifications among 
those of all animals, .oo08; percentage of species in this phylum 
among the whole number of animal species known, .8028. 

Discussion.—Information at hand seems to indicate that sponges 
are used very little as food by other animals; sea-urchins, marine 
worms, amphipods, and mollusks, however, are recorded as predators. 
Fresh water sponges are eaten to some extent by fishes. Whether 
this is credited to their “ protective adaptations” is of little moment 
as the fact remains that sponges do not multiply excessively nor 
overrun the earth as forms that lack enemies are in theory supposed 
to do. 

Sponges have bright and varied colors and one case of mimicry has 
been pointed out.’ If it be true as apparent from observations thus 
far, that they have few or no enemies, natural selection can not be 
advanced as an explanation of their color phenomena. If sponges 
without enemies have adaptations of the same character as other 
groups with numerous enemies, it would seem evident that selection 
by predatory animals has no necessary connection with the adaptations. 


COELENTERATA (HYDRAS, JELLYFISHES, SEA-ANEMONES) 


Protective adaptations-—Some coelenterates have a chitinous cuti- 
cle, others have a calcareous skeleton, and many of them have nema- 
tocysts or stinging cells. Numbers of them are brilliantly colored or 
phosphorescent but it must be noted also that many are transparent 
or nearly so, showing that possession of protective devices (as the 
nematocysts) is not always accompanied by the development of 
“warning colors.” 

Bird enenties.—The Coelenterata most often found in bird stomachs 
are the Hydrozoa (such as Abictinaria, Sertularclla, and Thuiaria). 
They have been identified 113 times from the stomachs of 13 species 
of ducks, 2 of gulls, and one each of murre, murrelet, and shearwater. 
Sea-anemones (Anthopleura, Aulactinia) have been identified four 
times from stomachs of a scoter, an eider, an oyster-catcher, and a 
gull, Aleyonaria from two ducks, and coral from one. 

Number of identifications, 122; percentage of identifications among 
those of all animals, .o514 ; percentage of species in this phylum among 
the whole number of animal species known, .8028. 

Other enemies—Hydroids are eaten by marine worms, by sea- 
urchins and sea-anemones, and also by fishes, as the cod, haddock, 

*McIntosh, W. C., The coloration of marine animals. Ann. Mag. Nat. Hist. 
7th ser., vol. 7, p. 223, Mar., 1901. 


' 
12 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


sand launce, lump sucker, cunner, scup, filefish, and flatfishes ; cteno- 
phores are eaten by the spiny dogfish, flatfishes, whiting, and cod; 
sea-anemones are eaten by cod,’ haddock, tilefish, flatfishes, the sun- 
fish Mola, spiny dogfish, whiting, and by the so-called jellyfishes and 
by whales. Holothurians and some fishes (Scarus) feed on corals. 

McIntosh notes that the brightly colored jellyfishes “ have precisely 
the same habits as the uncoloured and transparent,” which raises 
doubt as to the validity of the selectionist interpretation of the facts. 
The brightly hued and the translucent forms are equally palatable 
to whales and other animals using jellyfishes as food. He adds with 
regard to sea-anemones that “the view that the gaudy colors .... 
act as a warning is not borne out by the eagerness with which the 
cod swallows the brightest, such as Stomphia, while the smaller flat- 
fishes fill their stomachs with Edwardsiae.’” (Ann. Mag. Nat. Hist. 
7th ser., vol. 7, pp. 2242225, 1901.) 

Discussion.—Coelenterates are another group of animals but 
slightly available to birds and seem to be taken in full proportion to 
the degree of availability. The nematocysts seem a futile defense 
against animals of the groups here mentioned as coelenterate enemies, 
and must be also in the case of the myriads of crustacea (possible 
enemies) all of which have a chitinous exterior and which more- 
over manipulate their food in the chelae before chewing it, a process 
that would result in the harmless discharge of the stinging cells. 
It is alleged that hermit crabs have a commensal relation with certain 
hydroids which grow upon the shells they inhabit and that they are 
protected from their enemies by the presence of the inedible stinging 
hydroids.” This is not the case where the bird enemies are concerned, 
as the sea ducks which are the principal bird enemies of hydroids, 
often swallow the hermit crab, shell, hydroids and all. Many of the 
examples identified from bird stomachs came from precisely this 
source. With respect to the practical aspect of the case, it would 
appear that in its shell retreat and its own strong claws the hermit 
crab has much more efficient defenses than the nettlelike hydroids. 
It seems more likely that the latter merely grow on mollusk shells as 
a convenient substratum. From the habit some hermit crabs have of 
frequently changing their abode, the advantage held by a “com- 
mensal”’ hydroid may be lost at any moment. 


"McIntosh notes that sea-anemones are a valued bait for cod. 
* Parker and Haswell, Zoology, vol. 1, p. 144, 1910. 


Ss 


NO. 7 PROTECTIVE ADAPTATIONS—McATEE 13 


PLATYHELMINTHES (FLATWORMS, FLUKES ) 


The majority of organisms of this phylum are parasitic and there- 
fore not available to predatory animals. Perhaps some of the fresh- 
water planaria and the marine nemerteans have been found but not 
identified in the stomachs of shore birds, but so far we have no 
positive determination of a worm of this phylum as bird food. Forbes 
reports a small catfish (Noturus) feeding on fresh-water planaria. 
McIntosh says that marine planarians of both plainly and brightly 
colored forms are eaten by sea-anemones and fishes. Fresh-water 
planarians also are eaten by fishes. Stiles intimates that carp destroy 
large numbers of the liver fluke (Fasciola hepatica) in the cercaria 
stage. 


NEMATHELMINTHES (THREADWORMS, ROUNDWORMS) 


Again a vast number of worms of this phylum are parasites, abun- 
dantly so, in fact, of birds themselves. In order to reckon as food 
only those so taken, all nematodes other than Gordiidae have been kept 
out of the computations. The records for Gordiidae number 24, the 
percentage of these identifications among those of all animals is 
oIoI, and the percentage of known species’ of Nemathelminthes 
among all animals according to the estimates used in the present 
paper, .2676. The nematodes have a tough cuticle but no special 
defenses ; nevertheless they certainly are not eaten out of proportion 
to their numbers, but considering availability to birds, they may 
possibly be eaten somewhat in ratio to the frequency with which they 
are encountered. They are eaten also by flatworms and by various 


fishes. 


TROCHELMINTHES (ROTIFERS) 


None of these have yet been identified as food of nearctic birds, 
though possibly rotifers taken in with foliage of aquatic plants may 
have been overlooked. Rotifers are eaten by the young of a number 
of fishes. 


MOLLUSCOIDA (CORALLINES, LAMPSHELLS) 





Protective adaptations —Except for the shells of the brachiopods, 
and cuticular walls of some bryozoa, no special protective features 
have been developed by the Molluscoida. 

Bird enemies.—Only three brachiopods have as yet been identified 
from the stomachs of nearctic birds—not a matter for surprise in 


* An enormous number of Nematodes await description. 


14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


view of the small number and marine habitat of the species of these 
animals. The other Molluscoida that have been found in bird stomachs 
include Cyclostomata (having calcareous zooecia), Cheilostomata 
(with calcareous or chitinous zooecia), and Phylactolaemata. Large 
numbers of the statoblasts of the latter group, the fresh-water bryozoa, 
have been disclosed in the stomachs of wild ducks. 

Number of identifications of Molluscoida, 134; percentage of 
identifications among those of all animals, .0564; percentage of species 
in this phylum among the whole number of animal species known, 
.3032. 

Discussion.—Considering their low availability to birds, animals of 
this phylum probably are taken in due proportion. Fresh-water bryo- 
zoans have been recorded from stomachs of many species of fishes, 
and the marine forms from a smaller number. Marine bryozoans 
are preyed upon by worms, amphipods, decapods, and starfishes. 


ECHINODERMATA (SEA-CUCUMBERS, SEA-URCHINS, STARFISHES ) 


Protective adaptattons—All of the echinoderms have a calcareous 
exoskeleton and in many the surface is beset with tubercles or spines. 
The starfishes and sea-urchins are armed also with pedicellariae or 
grasping organs, which in some cases in the latter group are said to 
be poisonous. Some sea-cucumbers have the “ Cuvierian organs” 
which throw out long viscid filaments. Starfishes, especially the 
brittlestars and many crinoids, have the supposedly protective faculty 
of snapping off their arms or portions thereof. The colors of echino- 
derms are often conspicuous and in certain cases have been termed 
warning. 

Bird enemies.—Starfishes have been identified 28 times in the 
stomachs of nearctic birds here reported upon; sea-urchins (Strongyl- 
ocentrus, Echinarachnius) 92 times; and sea-cucumbers, 3 times. 
The birds (19 species) eating them were chiefly ducks collected im 
northern seas. 

Number of identifications, 125 ; percentage of identifications among 
those of all animals, .0526; percentage of species in this phylum among 
the whole number of animal species known, .7136. 

Other enemics.—Starfishes and sea-urchins prey upon one another, 
and are very commonly eaten by cod, haddock and other species of 
Gadus, by argentines, dragonets, rocklings, wolffishes, rays, sharks, 
tautog, scup, smelt, flatfishes, and others. Sea-cucumbers are less 
commonly taken by the same predators. Blue foxes on the Pribilof 
Islands feed on sea-urchins in winter. Sea-urchins and starfishes are 
consumed also by crabs, sea-anemones, and marine worms. 





NO. 7 PROTECTIVE ADAPTATIONS—McATEE 15 


McIntosh comments interestingly on the enemies of echinoderms: 
“The colours of Echinoderms are often most conspicuously bright,” 
but Hippasterias, which is brilliant orange-red, is eaten by gulls, 
cod, catfishes and by other starfishes. “ The sand-stars (e. g. Ophiura 
lacertosa) are often tinted of a hue resembling their surroundings, 
yet they and the more brightly tinted forms are common in the 
stomachs of fishes and are eagerly devoured by gulls when stranded 
on the beach.” “ The brown and purple hues of sea-cucumbers may 
in some way stubserve protection... . yet both they and the trans- 
parent forms are found in the stomachs of fishes.” (Ann. Mag. Nat. 
Hist. 7th ser., vol. 7, pp. 225-226, 1901.) 

Discussion——Echinoderms have a number of protective devices 
but also it would appear, numerous and effective enemies. Birds prey 
upon this group to fully as large an extent as could be expected, con- 
sidering the slight degree to which they come in contact with echino- 

derms. 
| It should be noted that while practically all starfishes and sea- 
urchins have similar protective adaptations, some are very gaudily, 
others modestly colored; in one case or the other, the natural selec- 
| tion theory as to the connection between special defenses and color- 
ing is untenable. The sea-urchins with calcareous tests, abundant 
spines, and pedicellariae seem unusually well defended, but that this 
does not mean freedom from enemies is shown by the great fecundity 
| of sea-urchins, individuals of some species, e. g., Echinus esculentus, 
yielding 20,000,000 eggs per season. 


ANNULATA (WORMS) 


Protective adaptations —The chaetopods including the most com- 
mon marine worms and the majority of earthworms have chitinous 
setae on all segments of the body. The earthworms are habitual bur- 
rowers, and some of both fresh- and salt-water annelids live in tubes. 
A few in each group are phosphorescent, and many of the marine 
worms are highly colored. A. R. Wallace says" “Among the crea- 
tures which probably have warning colors as a sign of inedibility are 
_. . . those curious annelids the Nereis and the Aphrodite or sea- 
mouse.” 

It should be noted however that many of the brightly colored forms 
live in burrows or tubes, thus taking care not to advertise their 
“inedibility.” _Leeches sometimes have strongly contrasting color, as 
for example greenish with red and black spots. 


‘Darwinism, p. 266, 1896. 
2 





¥ 


16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


Bird enemies.—The identifications of Annulata from nearctic birds 
are: 690 for the Polychaeta, chiefly Nereidae, which have been found 
in the stomachs of more than 70 species of birds and in numbers up 
to 500 in a single stomach; 428 for the Oligochaeta or earthworms 
from 44 species of birds; and 11 of Hirudinea or leeches from tro 
species. The robin (Planesticus migratorius) feeds habitually and 
voraciously upon earthworms and the woodcock (Philohela minor) 
makes about half of its diet of these annelids. 

Number of identifications, 1131; percentage of identifications 
among those of all animals, .4764 ; percentage of species in this phylum 
among the whole number of animal species known, .7136. 

Other enemies.—Studies that have been made of the food of fishes 
indicate that a very large number of marine fishes prey upon the 
Nereidae and. other annelids. They are eaten also by other worms, 
starfishes, sea-urchins, sea-anemones, gastropods, and crabs. Fresh- 
water oligochaetes form a steady contribution to the diet of the fishes 
of their environment. Earthworms are eaten by predacious beetles, 
by most batrachians, by some turtles, snakes and by various mam- 
mals including shrews, skunks, and the armadillo, but especially by 
the moles (Parascalops brewert, 26 per cent of the food; Scalopus 
aquaticus, 31 per cent; Scapanus townsendi, 40 per cent; Condylura 
cristata, 50 per cent.) Leeches are eaten by a variety of mammals, 
birds, reptiles, amphibians, fishes, crustaceans, snails, and insects. 

Discussion.—Both of the annelids (Nereis and Aphrodite) Wallace 
mentions as being warningly colored are eaten by birds and fishes, 
Nereis frequently and in large numbers. Considering the aquatic 
habits of most of the annelids it would appear that they are taken 
by birds as often as could be expected. It is evident furthermore 
that they have numerous other predatory foes and that they probably 
contribute their full quota of food toward the dietary requirements 
of the animal kingdom. 


ARTHROPODA (JOINTED ANIMALS) 


As recorded previously in the table of phyla, the Arthropoda, in- 
cluding the exceedingly numerous class of insects, furnish, as would 
be expected, a very large preponderance of the animals eaten by birds. 

Number of identifications, 210,752; percentage of identifications 
among those of all animals, 88.7551; percentage of species in this 
phylum among the whole number of animal species known, 74.6188. 
The disproportion of the percentage of capture to that of frequency 
reflects the relatively greater availability to birds of arthropods over 
the other phyla. 








NO. 7 PROTECTIVE ADAPTATIONS—McATEE 17 

Since arthropods are by far the most important phylum of animals 
as bird food and since it is with reference to the included class of 
insects that the theories of protective adaptations have been most 
highly elaborated, it is desirable to subdivide the phylum for the pur- 
poses of the present discussion. 

The tabulation below shows the number of identifications and their 
percentages for the four classes of Arthropoda that are available for 
food to nearctic birds. 


Arthropoda 
Percentage 
of species 
Percentage of in this class 
identifications among the 
among those whole number 
Number of of all of arthropod 
Class identifications arthropods species known 
(nustacea: 2s... .2 os skeeacies 6,086 2.8877 3.8254 
IMvGIA POG ceecce. cece. 2,862 1.3530 4781 
ISG Ctae aaron is at e-wieaiiare we hiee 190,919 90.5801 91.8589 
PNG ACINITC aw sed els soins swe aoe 10,885 5.1648 3.8254 


It is evident that the percentage of identifications for each class 
corresponds very well with the frequency of such animals as indicated 
by the number of species. The validity of such comparisons used 
throughout this paper thus receives further corroboration. 


CruSsTACEA (CRABS, SHRIMPS, SOWBUGS) 


Protective adaptations ——The exoskeleton of crustacea is either 
chitinous or much thickened or calcified. Several groups have bivalved 
carapaces in which the animal is nearly or entirely enclosed. The 
barnacles have a more complicated covering of calcareous plates, some- 
times thick and hard. Most of the decapods have strong grasping fore- 
legs, and have furthermore the faculty of snapping these off when 
properly stimulated. Some crustacea have burrowing habits (Hip- 
pidae) and others (hermit crabs) use the shells of univalves for 
shelter, while the terrestrial sowbugs roll themselves up into a ball 
when disturbed. 

As to color, some of the smaller aquatic forms are translucent or 
transparent; the ostracods are said to assimilate with the general 


color of their environment, while some copepods and decapods are 


brilliantly colored. As to form, Mortensen says: “‘ The typical cases 
of adaptation to life among algae are especially found among the 
Caprellids ; they might be said to represent the Phasmids and Geo- 
metrid larvae, among marine animals ” (p. 77). “ Zdothea marina and 


18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


related species . . . . bear a striking resemblance to the plants (es- 
pecially Zostera leaves) on which they are found” (p. 78).’ 

“Many Crustacea,’ according to Parker and Haswell, “ present 
instances of protective and aggressive characters, i. e., modifications 
in form, colour, etc., which serve to conceal them from their enemies 
or from their prey. Probably the most striking example is that of 
certain crabs (Paramithrax) which deliberately plant Sea-weeds, 
Sponges, Alcyonarians, Zoophytes, etc., all over the carapace, and 
are thus perfectly concealed except when in motion.” * 

Poulton expresses the same view in the following language: “ Cer- 
tain palatable animals make use of the Special Defence and Warning 
Colours of other forms. Thus, the common English hermit-crab, 
Pagurus bernhardus, commonly carries on its borrowed shell the con- 
spicuous stinging sea-anemone Sagartia parasitica; while another 
English species, Pagurus prideauxit, inhabits a shell which is invari- 
ably clothed by the flattened Adamsia palliata. Two crabs (Polydectus 
cupulifer and Melia tessellata), from Mauritius, described by Mobius, 
invariably held a sea-anemone in each claw. Two other groups of 
animals, sponges and Ascidians, in addition to sea-anemones, are 
avoided by enemies of the Crustacea, and these are also employed 
by the latter. Thus the British hermit-crab Pagurus cuanensis is 
found in shells which are covered with a (generally) brightly-coloured 
sponge (Suberites domuncula). Mobius also describes a Mauritian 
hermit-crab (Ascidiophilus caphyraeformis) which lives in a case 
formed by an Ascidian.” ® 

Bird enenues—Most of the leading subdivisions of the Crustacea 
contribute to the food of birds, apparently about in proportion to their 
accessibility. The tabulation of numbers of species of Crustacea used 
for comparison with those of percentages of identifications was made 
from “A list of the Crustacea of New Jersey including the adjacent 
region or that of the Middle Atlantic States,’ * the only check list 
available for any considerable area of our region. 

The Anostraca (fairy shrimps) are locally eaten more extensively 
than indicated above and the fact is an illustration of the principle 


* Mortensen, T. H., Observations on protective adaptations and habits, mainly 
in marine animals. Vidensk. Meddel. fra Dansk. naturh. For. Kjob., bd. 69, 1920. 

* Zoology, vol. 1, p. 601, 1910. 

* Poulton, E. B., Essays on evolution, pp. 356-357, 1908. 

Thomas Scott records a copepod (Acidicola rosea) which lives within the 
branchial sac of an ascidian as having been eaten by a sole Pleuronectes micro- 
cephalus. (20th Ann. Rep. Fishery Bd. Scotland, p. 525, (1901) 1902.) 

“Fowler, H. W., Ann. Rep. N. J. State Mus., pp. 463-5908 (1911), 1912. 





NO. 7 PROTECTIVE ADAPTATIONS 





McATEE 19 


that in natural history as elsewhere sweeping statements based on 
partial or negative evidence are dangerous. No exception may be 
noted for one which claims that “enemies play no part in keeping 
down the numbers of Artemia (brine shrimps), or of Ephydra 
(alkali flies) in the larval stage.” * Dr. Alexander Wetmore, of the 
Smithsonian Institution, who has had considerable experience about 
Great Salt Lake to which locality the quoted assertion relates, has 
pointed out * that Artemia and Ephydra are by no means free from 
enemies. Shovellers, lesser scaups, golden-eyes, green-winged teal, 
Wilson’s and northern phalaropes, avocets, and black-necked stilts all 
feed extensively upon both of these animals. But for the fact that 


Identifications of Crustacea 


Percentage 
of species 
Percentage of in this group 
identifications among those 
among those of all Crustacea 
Number of of all of the Middle 
Group identifications Crustacea Atlantic States 
RIMIGOHtIAEE — 6s: sess ids oso 573 9.4150 ee 
PNTIOStEACA-speeyeiae sav se eee ee 28 .4601 1.3667 
ladOCera: Wesrs oH ey ss sieie 90 1.4788 4.1002 
PU STLACOU A. “org Qe site van ts See os 207 3.4012 3.8724 
COPEPOdS -o.0.5 60d von sane ceess 13 2136 18.2232 
@irripedia: .sse..0..2se4 00 4o1 6.5888 5.9225 
MSOPOMA Ss aces cecacmencs ses 385 6.3259 14.3508 
Amphipoda ..6..i26....0800. 986 16.2010 9.3394 
OTMACEA is. oie 20 ean ess be dun t as ee 3.1891 
ptomatopoda: Fss.i0<cssecs0s een seeee Q112 
CMIZOPOddy s5..06 eee. se cia ne 48 .7887 1.1389 
Wecapodare sce saeeeeee ee 3355 55.1260 37.5853 


stomach analyses have not been made of other birds collected at the 
same place, it would undoubtedly be possible to add the names of 
a number of species to this list. Doctor Wetmore states that “ the 
toll taken by birds from the brine shrimp and alkali fly larvae and 
pupae during the course of a season constitutes a mass of individuals 


almost beyond comprehension . . . . The immense numbers of these 
creatures . . . . must be attributed to the large number of offspring 


produced rather than to an absence of enemies.” The number of 
records for the minute Cladocera is fully up to expectations. Among 
other items of this group the egg capsules or ephippia of Daphnia 
have been found in numerous stomachs of grebes and wild ducks, and 
in number up to 250 in a single stomach. 


*Vorhies, Charles T., Notes on the fauna of Great Salt Lake. Amer. Nat., 


vol. 51, pp. 494-499, Aug., 1917. 
” Amer. Nat., vol. 51, pp. 753-755, Dec., 1917. 


20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


The ostracods have been identified chiefly from the stomachs of 
15 species of wild ducks, and no fewer than 1,200 have in two in- 
stances been taken from a single stomach. Most of the barnacles 
were found in the stomachs of 22 species of ducks, gulls, and shore- 
birds from northern waters. Amphipods were eaten by more than 80 
species of birds, largely shorebirds, ducks, and other waterfowl ; nearly 
70 species of amphipods were identified and the number of individuals 
taken by single birds ran up as high as 2,500. The isopods were con- 
sumed by more than 75 species of birds, the land-forms or pillbugs 
alone by about half that many; the greatest number of individuals 
of terrestrial sowbugs found in a single stomach was 60, of aquatic 
forms, 256. In the Decapoda it may be of interest to note that 392 
of the identifications were of shrimps, hundreds of individuals being 
present in some stomachs ; 1,592 of Astacidae (crawfishes), the great- 
est number taken by one bird being 49; and 794 of crabs of various 
kinds. Among groups of crabs represented, there were the following 
numbers of captures, the figures in parentheses in each case denoting 
the largest number of individual crabs found in a single stomach: 
Sand crabs or sandbugs, Hippidae, 61 (14) ; stone crabs, Lithodidae, 
go (16) ; hermit crabs, Paguridae, 35 (40) ; mud crabs, Pilumnidae, 
186 (36) ; swimming crabs, Portunidae, 39 (16) ; edible crabs, Can- 
cridae, 41 (18) ; shore crabs, Grapsidae, 180 (26) ; and fiddler crabs, 
Ocypodidae, 272 (19). 

Number of identifications 6,086; percentage of identifications 
among those of all arthopods 2.8877; percentage of species in this 
group among the whole number of arthropod species known, 3.8254. 

Other enemies.—Crabs of various genera (including hermits) are 
a staple item of food for many fishes, such as the dogfish, rays, eels, 
sea bass, squeteague, scup, tautog, swellfish, toadfish, tilefish, hake, cod, 
haddock, sculpins, and flounders. Crawfishes are relished by fresh- 
water fishes and are eaten also by snakes, turtles, and various mam- 
mals such as the muskrat, raccoon, skunks, mink, and otter. Such large 
and powerful forms as lobsters are eaten by sea bass, rockfish, tautog, 
sharks, dogfish, rays, and skates. Amphipods are captured by the 
plant Utricularia, by insects, hydras, sea-anemones, and starfishes. 
Practically all adult fresh-water fishes eat amphipods and isopods, 
and when young prey upon Cladocera, Copepoda and Ostracoda. 
Starfishes and bony fishes such as Coregonus, Salvelinus, Alosa, her- 
ring, sticklebacks, and roaches continue feeding on these small forms 
when adult. Marine fishes take similar crustacea available to them, 
particularly the abundant shrimps and Mysidacea. Whales and seals 
consume enormous quantities of isopods and Euphasiacea. Caprellids 





NO. 7 PROTECTIVE ADAPTATIONS—McATEE Zi 


are eaten by fishes as well as by birds. Cladocera and Copepoda are 
eaten freely by larval salamanders and to a lesser extent by recently 
transformed frogs. They and all other small fresh-water crustaceans 
fall a prey to Hydra and aquatic insects. Small marine forms are 
engulfed even by protozoans. Barnacles are eaten by the tautog, and 
by sea-anemones and sea-urchins. More than 80 kinds of crustacea 
have been identified from stomachs of haddock taken in waters about 
Scotland (Thomas Scott). Crustacea have parasites from among 
their own ranks, and from among the worms. 

Discussion—Most of the small crustacea are translucent or trans- 
parent but this does not save them from their foes. Practically all 
aquatic animals “ get their start’ by feeding on these crustacea, the 
list including a great variety of insects, fishes, and batrachians. Many 
of them continue feeding upon crustacea when adult, and so com- 
mon is this habit that in many cases small crustacea are the animal 
basis of the food for the entire fauna of certain waters. This is true 
of the Artemia of Great Salt Lake, previously discussed, and con- 
spicuously so of the Mysidacea, Amphipoda, Isopoda, Euphasiacea 
and Macrura of northern oceans, where everything from other 
crustacea, through fishes and birds up to whales preys incessantly upon 
them. There is no question of special protection here but solely of 
numbers and fecundity. 

The protection that crustacea might be supposed to derive from their 
more or less indurated exoskeleton is discounted by the fact that in 
most cases there are plenty of enemies large enough to swallow them 
whole. Of what avail for instance is the bivalved shell of the almost 
microscopic Ostracoda? The same principle applies all along the line 
up to and including the crabs, for most crab-eaters swallow their 
prey entire ; however there are some crabs that grow so large they are 
possibly almost free from enemies when adult. 

The claws of the large decapods naturally are of little avail against 
enemies so voracious as to swallow the crustaceans whole and there 
is no evidence known to the writer that the self-mutilation practiced 
by decapods results in enemies swallowing the claw and letting its 
owner escape. 

Birds find the Hippidae or sandbugs, despite their burrowing habits, 
and hermit crabs, adopted shell and all, are freely eaten by birds and 
fishes. In numerous cases the hermit crabs found in bird stomachs 
were those with hydroids and bryozoa growing on their carapaces 
or shelters. Why should it even be supposed that such combinations 
of animals are protective when the enemies of one of them are in 
most cases enemies of all? For instance the diving ducks and fishes 


7 


22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


which relish crabs, including hermits, also eat mollusks, bryozoa, and 
hydroids. What difference is it to them that a mollusk shell con- 
tains a hermit crab rather than its original occupant, or that hydroids 
are growing on it when these animals are browsed from rocks, etc., 
elsewhere? McIntosh has brought up this same point with regard to 
species of Hyas which become covered with a growth of algae and 
invertebrates, yet covered with parasites as they are, abound in 
stomachs of the cod.’ They are eaten by other fishes and by birds 
also. Conclusions of a similar nature undoubtedly must be drawn in 
the case of those crabs associating sea-anemones and ascidians with 
themselves. Both of these classes of animals have their enemies 
which probably would engulf crab and all in cases where the animals 
were together. 

The caprellids noted by Mortensen as resembling algae and by 
Parker and Haswell as so closely assimilated in form and color to 
Hydrozoa and Polyzoa as to be difficult of detection nevertheless are 
detected and eaten by some birds and by numerous fishes, and the 
protectively formed and colored isopods of the genus /dothea are 
represented by 51 records for 6 species in stomachs of 18 kinds of 
birds. 

The fiddler crabs (Uca), so abundant and conspicuous on the mud 
flats of the southeastern coast of the United States, have one claw 
enormously developed, thus having the principal characteristics of the 
so-called protected species, a special mode of defense, and living ex- 
posed and conspicuously in large numbers. They are freely eaten by 
birds however and for this single genus of a few species, we have 
271 records from 24 species of birds. 


Myriapopa (THOUSANDLEGS, CENTIPEDS ) 


Protective adaptations.—The centipeds and millipeds exhibit differ- 
ences that would warrant their being treated as separate classes; 
customarily, however, they are considered together. The following re- 
marks on their protective adaptations are quoted from F. G, Sinclair.’ 


The means of defence possessed by these animals ... . differ very much 
in the different species of Myriapods. In the Centipedes the animals are 
provided with a powerful weapon in the great poison claws which lie just 
beneath the mouth, and which are provided with large poison glands, which 
supply a fluid which runs through a canal in the hard substance of the claw 
and passes into the wound made by the latter. The effect of this fluid is 


Ann. Mag. Nat. Hist. 7th ser., vol. 7, p. 229, Mar., 1901. 
* Cambridge Nat. Hist., vol. 5, pp. 36-37, 1910. 


NO. 7 PROTECTIVE ADAPTATIONS—McATEE 23 


instantaneous on the small animals which form the food of the Centipedes. I 
have myself watched Lithobius in this country creep up to a blue-bottle fly 
and seize it between the poison claws. One powerful nip and the blue-bottle 
was dead, as if struck by lightning. I have also seen them kill worms and also 
other Lithobius in the same way. When another Lithobius is wounded by the 
poison claws it seems to be paralyzed behind the wound. The Millepedes, on 
the other hand, have no such offensive and defensive weapon. They rely for 
protection on the fluid secreted by the stigmata repugnatoria (or glandulae 
odoriferae) mentioned before. This fluid has been shown to contain prussic 
acid, and has a very unpleasant odour. Most of the Millepedes are provided 
with these glands; but in the cave Myriapods mentioned before, the animals 
have not to contend against so many adversaries, and these glands almost 
disappear. Other Myriapods defend themselves by means of the long and stiff 
bristles with which they are provided, e. g., the little Polyxvenus. 


Bird enemics—Centipeds have been identified 236 times from the 
stomachs of 65 species of nearctic birds, and millipeds 2,598 times 
from 98 species. The latter were identified more than 50 times in the 
case of each of 12 species of birds. The highest number of millipeds 
found in a single stomach—that of a starling—was 40. More than a 
tenth of the starling’s annual food in the United States consists of 
millipeds. 

Number of identifications, 2,862; percentage of identifications 
among those of all Arthropoda, 1.3580; percentage of species in this 
class among the whole number of arthropod species known, .4781. 

Other enemies.—A. H. Wirkland in his report on the ‘‘ Usefulness 
of the American toad”’’ states that Io per cent of the food of 149 
individuals examined consisted of millipeds and that as many as 77 
were found in a single stomach. Myriapods are eaten also by frogs, 
salamanders, lizards, snakes and turtles. Among mammals the com- 
mon mole (Scalopus), armadillo (Tatu) and civetcat (Bassariscus ) 
(and the mongoose as introduced into Trinidad) are known to feed 
on centipeds, and Brewer’s mole (Parascalops) and the armadillo on 
millipeds; shrews prey upon both groups. Centipeds are eaten by 
predacious beetles, frequently prey upon each other, and it appears 
that often the male is consumed by the female following pairing. 
Millipeds are the chief food also of certain Lampyrid larvae, are 
eaten by ground beetles, and are parasitized by phorid flies. 

Discussion.—Vhere is a more or less prevalent belief that myriapods 
are “ specially protected” animals. This idea is reflected in an article 
on “The hothouse milliped as a new genus,” in which the author, 


‘Farmers’ Bull. 196, U. S. Dep. Agr., 16 pp., 1904. 


24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


O. F. Cook, says:* “ Prussic acid and other corrosive secretions 

. .render . . . . the millipeds distasteful to birds and other animals 
that might prey upon them.” This statement implies that millipeds 
have no natural enemies, an Utopian condition that probably no 
animal enjoys. In fact the evidence here adduced shows that milli- 
peds and centipeds as well, have numerous effective bird enemies, 
which together with special enemies in other groups, no doubt prey 
upon them about in proportion to their availability. From the com- 
paratively small numbers of myriapods and their secretive habits, it 
could not be expected that they form a very high percentage of the 
food of carnivorous animals. This reasonable expectation certainly 
is fully satisfied by the showing here made of the activities of their 
natural enemies. 


InsEctTA (INSECTS) 


From tabulations appearing earlier in this article, it will have been 
noted that arthropods contribute more than 88 per cent of all the 
records of the animal food of nearctic birds and insects more than 
go per cent of the arthropods. To repeat the figures for the latter 
group, insects furnish 190,919 identifications, which is 90.5891 per 
cent of those of all arthropods. The percentage of species of the class 
Insecta among the whole number of arthropod species known is 
91.8589. 

Not only are insects the most numerous class of jointed animals, 
and the most important item of the animal food of birds, but they are 
also the group about which most has been written in a theoretical way 
as to protective adaptations (especially color) and as to the relation of 
these adaptations to predatory foes. On all these accounts it is de- 
sirable to discuss the insects in greater detail, certainly in most cases 
by orders and in some instances by families. Tabulations have been 
prepared, therefore, showing numbers of identifications by orders and 
families, with their relative percentages. The first of these is a 
distribution of the total number of identifications by orders. 

The reader may have wondered why some of the tabulations as 
to relative numbers of insects have not been based on the inventories 
of some of the larger museums. However, this matter has been con- 
sidered and the invalidating factor in such statistics is that such col- 
lections are always more or less specialized either as a result of 
policies of the museum or of the receipt of collections from special- 
ists. Thus among insects, such favorite groups of the amateur as 


* Proc. U. S. Nat. Mus., vol. 40, p. 625, IQII. 


















| 
Tr | 
Order pe | Beene 
hysantira ......... | * 100 | * 700 
@dondta ........:; : 705 16,642 
MSOPteTa ss csc. ebsss 173 / 1+ 100,000 
| Ephemerida ........ | 
| Plecoptera ......... | 
Gorrodentia ........ | ve | . 
Mecoptera ......... a aan 
| Trichoptera ........ | 
Neuroptera ......... | 
| Mallophaga ........ 472 | '1,250 
Dermaptera ........ 180 1,098 
Orthoptera .......... 2,556 25,988 
Hemiptera ......... | * 3876 244,637 
Lepidoptera ........ 30,653 275,920 
MUIPUEPA, occ cess eas 10,253 210,880 
Siphonaptera ....... *130 * 432 
Coleoptera ......... * 32,500 * 738,000 
Hymenoptera ....... | 17,638 493,757 
Thysanoptera ....... 200 750 
Strepsiptera ........ 159 414 
| 
BPOLAI! fees 4.0 serdns | 98,925 2,125,189 


1 Estimated. 


NO. 7 PROTECTIVE ADAPTATIONS—McATEE 





Percentage 
of species 
in this 
group 


1010 
.7120 
1747 


6534 


.1262 
1818 
2.5915 
3.9147 
30.9595 
10.3555 
fists 
32.8250 
17.8143 
.2020 
1605 


25 


Lepidoptera and Coleoptera are always copiously represented, while 
such orders as Thysanura and Thysanoptera in most cases are 
obviously neglected. However, one museum tabulation with percent- 
age designations added, is herewith appended as a further demon- 
stration (notwithstanding the defects just pointed out and the ap- 
proximate nature of some of its figures) that multiplicity of species is 
more or less closely correlated with abundance of individuals. This 
table is adapted from one presented by Dr. J. M. Aldrich in the 
Smithsonian Report for 1g19 (1921), p. 373. 


Sununary of U. S. National Museum Collection, June 1919 


Percentage 
of specimens 
in this 
group 


.0329 
7821 
4.7000 


.0587 
.0510 
1.2214 
11.4979 
12.9082 
9.9113 
.0203 
34.6860 
23.2005 
.0352 
0194 





SMITHSONIAN MISCELLANEOUS 





COLLECTIONS — VOL. 85 


Identifications of Insects 


Number of 


Order 1 identifications 
APC A aera crn cxetorere trees 5 
Odonata (further unidenti- 

{LCC Been iare Cicer ee 2,082 
LVS Optenas sence reece 245 
Anisopteramereiecie noe 707 
YAW (OWA) Soa cacnoccost 3,03 
Agnathalarpciceriereincicie ai 484 
Plecopteras bination tant 80 
ESoptetar: tyes asi aie 129 
Dermaptera eee ae oe 18 
Cheleutoptera ............ 26 
Diphtheropterawaesetoe ces 5,005 
Orthoptera (Sens. str.)... 6,280 
Raleoptera wee: ere ce 117 
Dicty,optenape amram cter 58 
Saltatoria (further uniden- 

LIMEUM eects cee errors 6,450 
Orthopteroidea (further 

tinidentifled)\ meses 3590 
All Orthopteroidea ...... 109,003 
Corrodentiaiessericiceeaee 17, 
Moallophagarere 2 sacs ciel 6 
SIPHONAPtehAagecdectome sh I 
Fleteropterae sacra bose 11,530 
Elomopteram: qaee ree creme 5,215 
Hemiptera (further uni- 

dented) hit, sade cneineni 5,650 
Allg ICHMCHOLC ae ects 22,395 
Neuroptera (Sens. lat.)... 119 
Megalopteray sh aecmeece es 167 
Rhaphidioideai vs. csc -m92 «ae 54 
Neuroptera (Sens. str.)... 108 
Phryganoideay aeatraseccee 866 
All Neuropteroidea ...... 1,314 
Wepidopteta meres sect 18,487 
Coleopteramaacas coer 85,322 
iMecapteramemeenercenecte. 5 
Da pteramtetca te svar metres 10,836 
Fiymienopterarja-tees. «cette 27,02 
Unidentified) i 52 sc0.0.4 00 2,676 


Percentage of 
identifications 
among those 
of all 
insects 


.0026 


1.0905 
.1283 
-3703 

1.5891 
-2535 
.O419 
.0077 
0094 
.0136 

2.9829 

3.2893 
.0613 
.0304 


3.3784 


.1880 
9.9534 
.0089 
0031 
.0005 
6.0392 
2.7315 


2.9594 
11.7300 
.0623 
.0875 
.0283 
.0506 
-4536 
6882 
9.0831 
44.6899 
.0026 
5.6757 
14.1551 
1.4016 





Percentage 
of species 
in this group 
among the 
whole number 
of insect 
species known 


1691 


2003 
.3383 
5986 
1041 
.0780 
.OOII 
.1301 
.6507 
.7809 
.8589 
3123 
.2082 


2.9410 
.0780 
-3383 
.0130 

4.9457 

3.6442 


8.5899 
.0156 
.O104 
.3383 
.3044 
7187 

15.6180 
40.2032 
.0260 


11.4432 
17.1798 


‘The arrangement of the orders of insects in this tabulation is a compromise 
among several systems. 


There are no accounts given of Platyptera (embiids), Zoraptera, Notoptera 
(grylloblattids), Siphunculata (body lice), Apocoleoptera (beaver beetles), 
and Suctoria (fleas) because we have no records of bird enemies. 


NO. 7 PROTECTIVE ADAPTATIONS—McATEE 27 


Total number of identifications, 190,919; percentage of identifi- 
cations among those of all arthropods, 90.5891 ; percentage of species 
in this class among the whole number of arthropod species known, 


91.8580. 
APTERA (WINGLESS INSECTS) 


Protective adaptations —The springtails and their allies appear to 
have few adaptations such as are commonly called protective, their 
defense being agility in some cases, and secretive habits in others. 
Some species have coxal glands supposed to be repugnatorial. 

Bird enemies.—While only five records of Thysanura are included 
in the tabulations here reported upon others have been made since 
and it seems probable that birds which feed about small pools, on 
the quiet surface of which Collembola sometimes abound, or on the 
edges of snowfields, will be found to pay due attention to thysanurans. 

Total number of identifications, 5; percentage of identifications 
among those of all insects, .0026; percentage of species in this 
group among the whole number of insect species known, .1691. 

Other enemies.—In reports of the Pennsylvania Department of 
Agriculture and others treating the same groups of animals, seven 
species of salamanders, four of frogs, and one toad are recorded as 
feeding on Thysanura. Hamilton, reporting on 400 stomach con- 
tents of young toads, says: “ Collembola comprised 6.2 per cent of 
the diet. The springtails sometimes occurred in large numbers in 
the stomachs examined, and together with thrips appeared to be 
an important food of all small anurans” (Copeia, 1930, p. 45). 
Forbes reports them being eaten by a Coccinellid beetle’ and a 
small fish* (Labidesthes sicculus), Needham, by the brook trout,’ 
and Pearce by two species of fishes, a killifish and the mudminnow.* 
They are known to be preyed upon also by aquatic hemiptera, and 
are cannibalistic. 

Discussion —Thysanura are chiefly minute insects, many of which 
spend their whole lives in well-concealed places. The forms which 
live more or less exposed appear to have enemies among animals 
interested in such small morsels of food. However information on 
the subject thus far is inadequate and no doubt will be increased by 
more intensive investigation of potential predators. 





* Bull. Ill. State Lab. Nat. Hist., vol. 1, no. 6, p. 52, May, 1883. 
7 Op. cit, vol. 2, p. 525, 1888. 

* Bull. 68, N. Y. State Mus., p. 205, 1903. 

“Bull. U. S. Bur. Fisheries, vol. 35, p. 285 (1915-16), 1918. 


28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


ODONATA (DRAGONFLIES, DAMSELFLIES ) 


Protective adaptations.—Dragonflies are fairly large, powerful, 
predacious insects with remarkable ability for flight. They are 
held in fear by illiterate people, a feeling possibly inspired by the 
large mobile head consisting chiefly of eyes, and the strikingly con- 
trasted color-pattern of many of them. A dark ground color with 
vivid spots of green or yellow, answering to the description of warning 
color, is common among dragonflies; some also have brilliant red, 
blue, and metallic colors. 

On the other hand, members of the order known as damselflies in 
general are weak on the wing and of slighter and more delicate 
structure. Some of them also are brightly colored but many are 
dull. The immature stages of both dragonflies and damselflies’ are 
aquatic, and predacious, and invariably inconspicuously colored. 

Bird enemies.—It might perhaps be expected that damselflies would 
be more frequently captured by birds than dragonflies, but this does 
not seem to be the case, the determinations for these groups so far 
standing at 245 damselflies and 707 dragonflies. However, 2,082 
identifications do not indicate which suborder is concerned. About 
200 species of birds are known to eat Odonata, and nymphs as well as 
adults are freely taken. No fewer than 100-125 nymphs have been 
taken from the gullet and gizzard of individual ducks, yellow-legs, and 
magpies. Regarding the adults, Needham says: “It is doubtful 
whether anything that flies is able to capture in flight one of the 
swiftest dragon flies.” " However, we have records of birds eating 
Epiaeschna heros, one of the largest and swiftest of the dragonflies of 
the United States, and Anax junius, another of the giant species, is 
commonly eaten by the pigeon hawk. No fewer than 28 individuals 
of Anax were found in a single stomach of this falcon, and adult 
dragonflies, mostly Anax junius, were found in 120 out of 181 
stomachs of the species. In a lot of dragonfly wings, picked up under 
the home of a colony of purple martins at West Chester, Pa., were 
represented about 63 individual dragonflies, largely Epiaeschna heros, 
but including also, Anax junius, Libellula pulchella, and Anax 
longipes. 

Number of identifications, 3,034; percentage of identifications 
among those of all insects, 1.5891 ; percentage of species in this group 
among the whole number of insect species known, .5986. 

Other enemies.—Odonata are notoriously cannibalistic both in the 
nymphal and adult stages. Diving beetles, water scorpions and other 





“In Ward and Whipple, Fresh-water biology, p. 890, 1918. 


* 











Ty 


NO. 7 PROTECTIVE ADAPTATIONS—McATEE 29 


aquatic hemiptera, salamanders, frogs, turtles, and many kinds of 
fishes prey upon the nymphs. Ants, spiders, robber flies, chipmunks, 
snakes, frogs, toads, and fishes feed to some extent also on adult 
dragonflies, obtaining most of them no doubt when teneral. They are 
parasitized by nematodes, mites, and flies.’ 

Discussion.—Odonata, both immature and adult, are freely preyed 
upon by a variety of enemies and no special defense can be assumed 
except the great expertness in flight of some of the dragonflies. This 
we have seen does not foil various birds nor of course predators from 
their own ranks. All in all it would seem that Odonata are preyed 
upon fully in proportion to their abundance. 


AGNATHA (MAYFLIES ) 


Protective adaptations-—The nymphs that live in water are plainly 
colored; some cling closely to various objects in their environment, 
while others swim in a rapid darting manner. The adults also are 
usually inconspicuously colored. 

Bird enemies.—Our tabulation shows mayflies to have been identi- 
fied from the stomachs of 108 species of nearctic birds. A nighthawk 
has been known to contain 400 adults at one time or many thousands 
of eggs, the remains of the digestion of adults. As many as 250 
nymphs have been found in a godwit’s stomach. Mayflies periodically 
are exceedingly abundant and then are preyed upon by practically all 
kinds of insectivorous birds. An interesting account of the behavior 
of birds in the presence of a swarm of ephemerids is given by Dr. S. D. 
Judd in his “ Birds of a Maryland Farm” ;* on this occasion 40 
species of birds were observed eating mayflies. This list adds nine 
to the species of birds known from stomach examination to feed 
upon mayflies. 

Number of identifications, 484; percentage of identifications among 
those of all insects, .2535; percentage of species in this group among 
the whole number of insect species known, .1041. 

Other enemies.—Mayfly nymphs are eaten by the nymphs of stone- 
flies and dragonflies, by water bugs, most fresh-water fishes, and to 
some extent by salamanders and turtles; the adults are preyed upon 
by fishes and adult dragonflies, spiders, toads, and bats. 

Discussion—Mayflies are good food for predacious animals and are 
eaten freely, so much so as to cause David Sharp to remark:* ‘‘ That 


*For full discussion of dragonfly enemies, see Bull. U. S. Bur. Fisheries, 
vol. 36, pp. 209-211 and 222-232 (1917-18), 1920. 

* Bull. 17, U. S. Biol. Survey, pp. 22-24, 1902. 

*Cambridge Nat. Hist., vol. 5, p. 442, 1910. 





30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, 85 


insects so fragile, so highly organized, with a host of powerful 
enemies, but themselves destitute of means of attack or defense, 
should contrive to exist at all, is remarkable.” Doctor Sharp here 
falls into the same error the selectionists often do, namely, of taking 
the struggle for existence too seriously. While mayflies are a favorite 
food of many predators, the evidence does not indicate that they are 
eaten out of proportion to their numbers. They are also very fecund, 
practically the whole body content of a female mayfly consisting of 
the two egg masses. The annual occurrence of swarms covering the 
foliage along streams (dating back as far as such things were re- 
corded) is proof enough that enemies do not permanently reduce the 
numbers of mayflies, and furthermore that the so-called defenses or 
protective adaptations, of which mayflies are so nearly destitute, are 
not essential to the maintenance of species in large, even overwhelm- 
ing numbers. 


PLECOPTERA (STONEFLIES ) 


Protective adaptations—tThe stoneflies are mostly plainly colored 
but some are rather bright yellow; they are poor fliers but some of 
them are said to emit a liquid from the basal articulations of the legs, 
a performance usually classed as protective. The nymphs are aquatic 
in habit, good swimmers, and obscure in color. 

Bird enemies.—Stoneflies have been identified in the stomachs of 
AI species of nearctic birds, usually in no very large numbers. The 
total number of identifications is 80; the percentage of identifications 
among those of all insects, .0419; and the percentage of species in 
this order among those of all insect species known, .0780. 

Other enemies ——Dragonfly nymphs prey upon those of stoneflies, 
and a few fishes, salamanders, frogs, and turtles feed upon these 
insects, either in the immature or mature condition according to 
availability. Needham says: “‘ Hudson has demonstrated the im- 
portance of stoneflies as fish food in the mountain streams of New 
Zealand”? (Fresh-water Biology, 1918, p. 884), and Muttkowski 
reports that 90 per cent of the food of trout in Yellowstone National 
Park consists of them. 

Discussion——The Plecoptera are a small group of insects of re- 
stricted habitat, one we should therefore not expect to find preyed 
upon extensively. They are eaten by various enemies, however, more 
or less in proportion to their abundance, and the evidence does not 
seem to indicate that special defenses of any kind enter into the 
equation. 


| 
| 
| 
| 
| 
| 
| 
| 


NO. 7 PROTECTIVE ADAPTATIONS—McATEE 31 


ISOPTERA (TERMITES ) 






Protective adaptations —Termites pass most of their lives concealed 
in galleries in wood or underground or in well-built nests. They have 
strong jaws, a caste of soldiers especially well-armed in this respect, 
and they emit a corrosive secretion. The color is usually yellowish 
to brownish, but some species have the body reddish and the wings 
dark, nearly black, thus having a coloration approaching that termed 
warning. 

Bird enemies.—Stomach examination has revealed termites in the 
dietary of 38 species of nearctic birds. The occasions when termites 
are available to most birds are infrequent, but when they come, the 
insects usually are in great abundance. Accordingly large numbers 
are eaten, and single stomachs have yielded as many as 215 termites 
in the case of a nighthawk, 400 in that of a pileated woodpecker, and 
| 1,100 in that of a flicker. The writer has twice observed numbers of 
English sparrows gobbling up termites upon emergence and Hagen 
_ has recorded’ a case in which 15 species of birds were in attendance 
ona swarm of white ants, the robins among them so gorging them- 
selves that their bills stood open. 
| Number of identifications, 129; percentage of identifications among 

those of all insects, .0677; percentage of species in this group among 
| the whole number of insect species known, .ogIT. 

Other enemics—Termites are as much sought after by some other 
| animals as they are by birds and even are eaten by man. It has been 
said that in the Tropics “ The flight of the winged termites is a great 

event in the animal year.”’* In India cockroaches, frogs, lizards, rats, 
bats, jackals, mongooses, jungle cats, and dogs have been observed * 
preying upon them. In the United States, besides wild birds and 
domestic fowls, salamanders, frogs, toads, lizards, spiders, centipeds, 
crickets, robberflies, ants, and beetle larvae prey upon termites. The 
insects have parasites also among the fungi, protozoa, nematodes, 
and mites." 

Discussion—The enemies of termites are comparatively well- 
known, not wholly because they are numerous or active, but also 
because termites are ‘‘ economic ” insects and have therefore been the 
subject of considerable study from many points of view. Although 


* Hagen, H., Proc. Boston Soc. Nat. Hist., vol. 20, p. 118, (1878-1880) 1881. 

*Longstaff, G. B., in Shelford, R., Naturalist in Borneo, p. 37, 1916. 

*Rothney, G. A. J., Proc. Ent. Soc. London, 1918, pp. Ixiv-lxvi. 

*For an account of these miscellaneous enemies, see Snyder, T. E., U. S. Nat. 
Mus. Bull. 108, pp. 116-118, 1920. 


3 





32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


they have protective adaptations of various kinds, termites are eaten 
freely by numerous animals. Birds prey upon them eagerly when 
occasion offers, but on the whole not out of proportion to the abun- 
dance of the insects. 


ORTHOPTEROIDEA 
Percentage 
of species 
Percentage of in this group 
identifications among the 
among those whole number 
Number of of all of nearctic 
Group identifications Orthopteriodea Orthopteroidea 1 | 
Winidentitiediee a seeeaeoe 6,800 35.8310 wae 
Dermapteram ease eee 18 0047 1.8404 
Paleopterayeacsaceeeoe ace 117 .6157 4.41904 
Dicky Optenaauasmecteemietiac 58 3052 2.1136 
@heleuitoptera We sa-n esos 26 .1368 1.4531 
Diphtheroptera ss... 04 5,605 29.9688 69.3525 
@xnthopterayyees sere 6,280 33.0472 20.7307 


DERMAPTERA (EARWIGS) 4 


All earwigs have pincer-like appendages at the end of the abdomen, 
the function of which is little understood. One suggestion is that they 
are for defense, but in what way they might serve for this purpose 
is not clear. Many earwigs have glands producing a fetid secretion. 
These insects in general are inconspicuous but a few have brilliant 
colors. Earwigs are seldom met with in the United States and the 
record of their bird enemies is short—18 identifications in the 
stomachs of 15 species of birds. Percentage of identifications among 
those of all insects, .o094; percentage of species in this group among 
the whole number of insect species known, .1301. That this result is 
merely a reflection of the infrequency of earwigs is indicated by the | 
fact that in Great Britain where these insects are much more common, 
the records of birds eating them are proportionately higher. Thus 
Robert Newstead, treating of a mere fraction of our number of | 
stomach examinations gives records for seven species of birds and | 
notes that 23 earwigs were found in the stomach of a green wood- 
pecker and 4o in that of a whimbrel. F. V. Theobald and William 
McGowan in their report * on ‘' The Food of the Rook, Starling and 
Chaffinch,” note that each of these birds prey upon earwigs, and 





‘Computed from Scudder, S. H., Catalogue of the described Orthoptera of 
the United States and Canada, Proc. Davenport (Iowa) Acad. Nat. Sci., vol. 8, 
IOl pp., 3 pls., 1990. | 

* Suppl. Journ. Board Agr. [London], vol. 15, no. 9, Dec., 1908. | 

* Suppl. Journ. Board Agr. [London], vol. 15, no. 15, May, 1916. 





A A SL OE gS est 


nO. 7 PROTECTIVE ADAPTATIONS—McATEE 33 


W. E. Collinge records* five species of birds as feeding on these 
insects. Among other enemies of earwigs are batrachians, of which 
6 species of salamanders, and 16 of frogs have been recorded in 
the United States as feeding on Dermaptera. The earwigs are neither 
an extensive nor an abundant group of insects and we should not 
expect to find them preyed upon by insectivorous animals to any 
marked degree. 


CHELEUTOPTERA (WALKINGSTICKS ) 


Protective adaptations —As their vernacular names, stick and leaf 
insects imply, these insects bear resemblances to objects in the vege- 
table kingdom that have caused them to be considered as having 
reached the very acme of protective adaptation. “Some,” says David 
Sharp, “look like sticks, or stems of grass; some have a moss-like 
appearance, while others resemble pieces of lichen-covered bark. The 
members of the tribe Phyllides are leaf-like. A certain number 
are covered with strong spines, like thorns. Some, if not all, of the 
Phasmidae,” he adds, “ have the habit of ejecting a stinking fluid that 
is said to be very acrid” (264). The eggs of walkingsticks are 


peculiar in shape and sculpturing and many of them resemble seeds. 


Bird enemics.—Records of walkingsticks in the identifications of 
bird food here discussed total 26, and pertain to 18 species of birds. 
The crow blackbird heads the list with seven captures. 

Percentage of identifications among those of all insects, .0136; 
percentage of species in this group among the whole number of insect 
species known, .6507. 

Other enemics.—Predacious hemiptera, mantids, lizards, and sper- 
mophiles may be mentioned among the enemies of stick-insects, and 
ichneumon flies are said to parasitize both adults and eggs. 

Discussion.—The apparent discrepancy in the indices of frequency 
of occurrence of stick-insects in bird food and in nature is to be 
explained by the relatively poor representation of this group in the 
United States, we having but 11 species. If we grant that the form, 
color, and sluggishness of these insects has a protective value in 
relation to predators, we must admit that these qualities facilitate also 
the destruction of the walkingsticks by grazing animals, which engulf 
indiscriminately huge mouthfuls of browse together with any insects 
thereon that are not agile enough to beat an instantaneous retreat. 
In the same way if the resemblance of the eggs to seeds is to be 

*The food of some British wild birds, 1913. 

* Cambridge Nat. Hist., vol. 5, p. 260, 1910. 


"" 


34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


regarded as significant, it would appear to put these objects in 
jeopardy, as the proportion of the food of birds and other animals 
made up of seeds is immensely greater than that composed of insect 
eggs. Judged as a protective adaptation, therefore, this case would 
seem to fit the old adage of “out of the frying-pan into the fire.” 
Two authorities who have paid special attention to the subject, how- 
ever, conclude that the resemblance to seeds of these eggs has no 
bionomic importance.’ 


SALTATORIA (GRASSHOPPERS, LOCUSTS, CRICKETS ) 


Owing to the facts that identifications in hundreds of cases were 
not carried as far as they might have been, and that it is impracticable 
to tabulate them by families, we put all the leaping orthoptera together, 
rather than consider separately the two orders into which these forms 
are usually grouped. 

For convenience part of the tabulation of identifications is here 
repeated in revised form. 


Identifications of Saltatoria 


Percentage of 

identifications 

among those 
Number of of all 


Group identifications Saltatoria 

Diphtheroptera 

(Grasshoppers, locusts) <.gct js cieels ease ore 5,005 30.3185 
Orthoptera (Sens. str.) 

(Green grasshoppers, katydids, crickets). 6,280 33.4328 
Saltatoria 

(Burther unidentified) ) os...sc.6e64- ee 6,450 34.3378 
Orthopteroidea 

(Further unidentified,no doubt Saltatoria) 359 T.QII2 
AllSaltatoriarse crear er eo tree cen orris 18,784 


Protective adaptations —A. R. Wallace says:* ‘‘ The whole order 
of Orthoptera, grasshoppers, locusts, crickets, etc., are protected by 
their colours harmonising with that of the vegetation or the soil on 
which they live, and in no other group have we such striking examples 
of special resemblance.” 

With special reference to American insects, A. P. Morse makes the 
following statement:* The coloration 


is, with few exceptions, highly sympathetic in character, harmonizing with or 
resembling very closely, often to a marvelous degree, the background of the 


* Sharp, D., Willey Zool. Results, Cambridge, 1808, p. 75-04. 
Severin, H. H. P., Ann. Ent. Soc. Amer., vol. 3, pp. 83-92, 1910. 
* Natural selection and Tropical nature, p. 46, 1891. 

“Proc. Boston Soc. Nat. Hist., vol. 35, no. 6, p. 244, 1920. 








INO. 7 PROTECTIVE ADAPTATIONS—McATEE 35 


insect’s environment. Earth tints, rock and sand textures, the infinitely varied 
browns, greens, and grays of living and dead vegetation, yellow, orange, rose, 
and silvery white are all represented in spots and streaks, the effect being to 
merge the insect indistinguishably into its background while at rest, thus shield- 
ing it in a very high degree from the observation of its foes. These colors 
are of great protective value at the present time, natural selection continually 
acting to preserve and perfect them, but though highly protective in character, 
they are without doubt primarily due to physiological processes and influences 
as yet imperfectly understood. 

This type of coloration is admirably illustrated among New England species 
by the Seaside Locust and Sand Locust which live on sandy backgrounds, the 
Snapping and the Ledge-loving Locusts on rock habitats, the Coral-winged 
and the Clear-winged Locusts in fields; and in the plant-perching species the 
Pine-tree Locust with its background of lichened pine bark, the Red-legged 
and the Two-striped Locusts among the yellowish green of herbage, and other 
species of Melanoplus—M. mancus, M. fasciatus, etc..—whose darker tints 
resemble those of fallen leaves from the Vaccinium thickets amid which they 
live. 

One who has not watched these creatures out of doors can appreciate to 
but a slight degree the effectiveness of sympathetic coloring as a means of 
concealment. Let him but try to pick out from its background immobile grass- 
green Cone-head, leaf-brown Shield-backed Grasshopper, or any of the Locusts 
just mentioned, and he will realize as never before the importance to the 
defenceless insect of Mother Nature’s protective mantle of invisibility. 


The wing-covers of certain katydids and allied forms are very 
leaflike, the resemblance being carried so far in certain cases, it 1s 
said, that the spots like those due to fungi and the tracks of leaf- 
mining insects are closely imitated. 

The leaping powers of the Saltatoria, remarkably developed in some 
forms, must be classed as defensive; most of these insects have 
powerful mandibles also, a few of them indeed being markedly 
carnivorous. 

Locusts of the subfamily Oedipodinae, especially, have another 
adaptation some consider protective. For instance E. B. Poulton 
says:* “ The brightly coloured hind wings of many moths (Catocala, 
Tryphaena, etc.) and grasshoppers (Oedipoda, etc.) which flash out 
conspicuously when the insect becomes active, and disappear equally 
suddenly when it alights, probably serve, as Lord Walshingham has 
suggested [Proc. Ent. Soc. Lond. 1890, pp. l-liii], to confuse a 
pursuing enemy.” It may be noted that Morse considers these colors 
as recognition markings. 

Finally, among protective adaptations, certain Orthoptera are said 
to mimic other insects, as for instance Membracidae, Phasmidae, ants, 


‘Essays on evolution, p. 303, 1908. 





30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


and beetles, though we have none of these forms in the United States ; 
the mole crickets and a few other forms have special fetid secretions, 
and the brown drop that so many orthoptera exude from the mouth 
when captured is said to be a protective device. 

Bird enemies.—Nearly a tenth of all the identifications of insects 
in bird stomachs are of leaping orthoptera. To name the birds that 
eat grasshoppers is to name all birds not strict vegetarians. When 
these insects are abundant, birds of all sizes turn their attention to 
the Orthoptera and for the time being make them a staple food. Asa 
constant article of diet also, they are important to many birds. The 
number of identifications of Saltatoria from stomach contents was 
50 or more in the case of over 20 species of birds, more than 100 in 
22 additional species, more than 200 in Io other species, in excess of 
T,000 in two cases, namely, of the common crow, and the meadowlark, 
and more than 1,500 for the starling and crow blackbird. Expressed 
in proportions of the annual subsistence of certain birds most fond 
of the insects, we find according to Biological Survey records that 
Saltatoria compose 21.29 per cent of the food of the western bluebird 
(based on the examination of 217 stomachs), 22.01 per cent for the 
eastern bluebird (855) ; grasshopper sparrow (170), 23 per cent; the 
eastern and western meadowlarks combined (1,514), 26.08 per cent; 
the Arkansas kingbird (109), 27.76 per cent; Franklin’s gull (93), 
43.43 per cent; and the scissor-tailed flycatcher (129), 46.07 per cent. 

These are illustrations of the relations of birds to leaping orthoptera 
under normal conditions. When species of these insects become exces- 
sively abundant as they frequently do, the gathering of the bird clans 
to feed upon them is proverbial. No instance is more celebrated than 
that studied by Prof. Samuel Aughey during an invasion of the Rocky 
Mountain locust in Nebraska. He found locusts in the stomachs of 
no fewer than 172 species of birds varying in size from the tiny 
hummingbirds up to the largest hawks, and including such usually 
exclusively vegetarian birds as the passenger pigeon and mourning 
dove. Professor Aughey was eye-witness also to 33 additional species 
of birds preying upon the locusts.’ 

For a modern illustration of the same phenomenon, we may cite a 
brief investigation made by the Biological Survey during a grasshopper 
outbreak in South Dakota in 1920. Out of the 26 species of birds 
collected, representatives of 24 had been eating the hoppers; of 19 
species every bird collected had taken grasshoppers, and for the 


4 Notes on the nature of the food of the birds of Nebraska. First Ann. Rep. 
U. S. Ent. Comm. (1877), App. I], pp. [13]-[62], 1878. 











NO. 7 PROTECTIVE ADAPTATIONS—McATEE 37 


species eating them, the insects composed from 40 per cent to 90 per 
cent of the total food. 

In another study of the effect of birds upon a severe irruption of 
grasshoppers in California, H. C. Bryant estimated that birds were 
destroying daily more than 120,000 grasshoppers per square mile in 
the infested area.’ 

In countries, notably Africa, where migrations of large numbers 
of locusts are of regular occurrence, various species of birds have more 
or less specialized in following these flights and feeding on the mi- 
grants, so much so, in fact, as to earn for themselves the name of 
locust birds.” 

Number of identifications of Saltatoria, 18,784; percentage of 
identifications among those of all insects, 9.6506; percentage of 
species in this group aniong all insect species known, 1.6308. 

Other enemies —All stages of the Saltatoria are much sought for 
by various animals. The larvae of Cantharid beetles and of bee flies 
(Bombyliidae) subsist upon the eggs, as do also certain mites, and 
the egg masses of some species are dug up and devoured by various 
mammals, as moles, mice, spermophiles, and skunks. The nymphs 
and adults fall a prey to vertebrates of nearly all sizes and descrip- 
tions, ranging from bears, through coyotes, foxes, badgers, skunks, 
civetcats, weasels, wood rats, squirrels, spermophiles, moles, shrews, 
and mice, to lizards, tortoises, snakes, salamanders, frogs, and toads. 
If any seek to escape their land enemies by jumping into the water, 
they are snapped up by fishes. The adults are destroyed in large 
numbers by parasitic diptera and hymenoptera. Most of the predatory 
invertebrates are fond of grasshoppers, this being particularly true of 
dragonflies, tiger beetles, ground beetles, robber flies, digger wasps, 
and spiders. In the case of the latter, S. W. Bilsing found grass- 
hoppers in 20 per cent of the webs of Epeira trifolium, in 35 per cent 
of those of Argiope riparia, 44 per cent of those of A. trifasciata, and 
in 53 per cent of those of Agalena naevia. Grasshoppers are parasit- 
ized by nematodes and protozoa and are subject to bacterial and fungal 
diseases, which last are said sometimes to destroy them “ in myriads.” 

Discussion—tThe Saltatoria or leaping Orthopteroidea are promi- 
nent in the insect world more through average large size and the 





*Univ. California Publ. Zool., vol. 11, no. 1, pp. 16-17, Nov. 1, 1912. 

* See Badenoch, L. N., True tales of the insects, pp. 127-128, 1899; La Baume, 
W., Beihefte z. Tropenpflanzer, vol. 11, no. 2, pp. 65-128, Apr., 1910; Agr. Journ. 
Cape of Good Hope, vol. 18, pp. 820-833, 1901; vol. 19, pp. 99-106, 165-171, 
248-262, 1901 ; vol. 28, pp. 364-366, 1906; Trans. South African Phil. Soc., vol. 1, 
p. 103, 1880. 





38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


abundance of individuals rather than through abundance of species. 
Almost everywhere in the United States that herbage is plentiful, 
grasshoppers in the late summer rattle away from the approaching 
pedestrian in such numbers as to form a veritable rolling barrage of 
insect projectiles. No insects are more conspicuous in action, yet on 
close examination the observer finds that the individual hopper is dull 
and obscure in color. The point is worthy of attention because it proves 
that the formula that abundant and conspicuous insects tend to be 
warningly colored and inedible has numerous exceptions. None of 
our grasshoppers of the northeastern United States are warningly 
colored, unless the Oedipodinae with brightly and contrastingly colored 
hind-wings, and in many instances a loudly rattling flight, may be so 
considered. Whatever their status in adaptation theories such genera 
as Arphia, Dissosteira, and Hippiscus seem to supply their full quota 
to the food of birds and other predatory enemies. On the other hand 
some of the “sympathetically” colored species mentioned in the 
remarks on adaptations quoted from Morse are the very bread of 
avian diet. Grasshoppers of the genus Melanoplus for instance were 
identified 543 times among the records here considered, and were 
found in the stomachs of more than 85 species of birds. These and 
other Acridids are taken not only frequently but often in quantity, for 
instance, the remains of no fewer than 123 specimens were found at 
one time in the stomach of a common crow and 340 in that of a 
Franklin’s gull. Judging from the records, the green grasshoppers or 
Locustidae and the crickets also bear their appropriate burden of 
predatory attack. 

The imposing total of 17,641 identifications of Saltatoria, more than 
a tenth of all insect determinations, shows what an important staple 
for the birds these creatures are, and how poorly their prevailing 
elaborately cryptic coloration succeeds in foiling their enemies. They 
are preyed upon voraciously not only by birds but by a host of other 
animals, but the effect of the attacks of predators, parasites, and 
diseases together in no way suggests that the Saltatoria are a dis- 
appearing race. Despite persecution, these insects abound and the 
reasons are high fecundity and the great surplus of food available to 
them ; these are substantial realities and outweigh immeasurably those 
airy intangibilities classed as protective adaptations. 


PALEOPTERA (ROACHES ) 


Protective adaptations——The comparatively few native species of 
roaches in the United States are secretive and nocturnal in habit but 
appear to have no other special protective adaptations. The introduced 


NO. 7 PROTECTIVE ADAPTATIONS—McATEE 39 


species live chiefly in structures of man hence have little relation to 
the indigenous fauna. 

Bird enemies—Thirty-six species of birds share the 117 identifica- 
tions of roaches in the food of nearctic birds. The number of records 
was 10 or more in the case of four species of birds, and the number 
of specimens eaten was as high as 10 in two instances but usually 
was less. 

Percentage of identifications among those of all insects, .0613; 
percentage of species in this group among the whole number of insect 
species known, .3123. 

Other enemies.—Roaches seem to be more or less regularly eaten 
by toads, frogs, the armadillo, spiders, rats, scorpions, and wasps. 
They have specific parasites among the Evaniidae. 

Discussion.—Owing to the poverty of the roach fauna of the United 
States, research here is not likely to throw much light on relations of 
these insects and their adaptations to predators. Tropical species are 
said to resemble various other organisms, including isopods, myrio- 
pods, longicorn, and coccinellid beetles, and hemiptera of the family 

| Miridae. But since all of these models themselves are freely eaten 
| by predators, the significance of the resemblances is hardly that usually 
| attributed. In the United States natural enemies would seem to be 
| proportional to the scanty population of native roaches. 

} 

| 

| 


DICTYOPTERA (MANTIDS) 


Like the roaches, the mantids of the United States are few in 
number and do not exhibit the unusual modifications displayed by 
some of the tropical representatives of the group. The principal 
defenses of our species must be their comparatively large size among 
insects and their highly predatory nature. However, these character- 
istics are of little avail against still larger predators and we find these 
insects taken by birds in numbers probably bearing no distant relation 
to the frequency of mantids in the country. Number of identifica- 
tions, 58; percentage of identifications among those of all insects, 
.0304; percentage of species in this group among all insect species, 
.2082 (for the world, of course). The number of species of birds 
~ concerned in the records here cited is 21. Mantids are eaten also 

by lizards. 


CORRODENTIA (PSOCIDS ) 
The Psocidae, which include the booklice seen in houses, are 


delicate and minute insects. Many of the out-of-door species are 
winged and the wings bear color patterns which may assimilate the 


7 


40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


insects more or less to the bark surface upon which many of them 
dwell. These insects mostly below the size of food objects ordinarily 
taken by birds were identified 17 times in the stomachs of nine species 
of birds. In one case, that of a chimney swift the stomach contained 
hundreds of specimens, gleaned no doubt from a swarm on the wing. 

Percentage of identifications among those of all insects, .0089 ; per- 
centage of species in this group among the whole number of insect 
species, .0780. 

MALLOPHAGA (BITING LICE) 


The only opportunity birds have to get these usually minute insects 
is to capture those parasitic on their own bodies, or in the case of 
raptorial birds to engulf some with their prey. Apparently either of 
these occurrences is rare; six records for as many species of birds 
being all included in the present tabulation. Percentage of identifica- 
tions among those of all insects, .0031; percentage of species in this 
group among all insect species, .3383. 


SIPHONAPTERA (FLEAS) 


Only a single instance of a flea being eaten by a bird has thus far 
come to light ; the opportunities for getting these small agile insects 
must be very few since our native birds are parasitized by fleas to only 
a very slight extent. That fleas are in no way distasteful (as food) 
to some of their hosts is evident to anyone who has observed dogs, 
monkeys, and other animals in their persistent and often successful 
search for these pests. 

Percentage of identifications among those of all insects, .0005 ; per- 
centage of species in this group among all insect species, .0130. 


THYSANOPTERA (THRIPS) 


Protective adaptations—Some are contrastingly black and white 
colored and the immature stages of many are red. It is doubtful 
however if these colors have any warning significance. ‘The small size 
and secretive habits of these insects doubtless are the most effective 
factors in restricting predation upon them. 

Bird enemies.—No identifications of thrips appear in the analyses 
of the stomach contents of nearctic birds here reported upon. 
Wetmore reports a thrips from the stomach of a hummingbird 
(Anthracothorax viridis) from Porto Rico. (Bull. 326. U. S. Dep. 
Agr. p:°73) 1916:) 

Other enemies.—Thrips are eaten by small predacious hemiptera, 
especially Anthocoridae, and egg parasites are known. Hamilton 








i Ny ‘ ss 
a ys 


NO. 7 PROTECTIVE ADAPTATIONS—McATEE 4! 


(Copeia, 1930, p. 45) says of 400 young toads examined, “ Thrips 
formed 10.1 per cent of the food, but were found in all but a few 
stomachs. These small insects appear to be a staple article of diet 
for young Bufo.” 

Discussion——Thrips are too small for most birds to notice, but 
considering our ignorance of the subject, the notes on enemies given 
indicate that they have foes, the character and number of which, 
probably as in other cases, are regulated largely by the factor of 
availability. 


RHYNCHOTA (BUGS, CICADAS, LEAFHOPPERS, SCALE INSECTS) 


For the reason that the term Hemiptera in a broad sense was used 
for about one-fourth of all the identifications of Rhynchota, it is not 
practicable entirely to separate Heteroptera and Homoptera. However 
the identifications of these groups are distributed as far as possible to 
families in the tables presented. In using these tables, it should be 
kept in mind that could the incomplete determinations have been dis- 
tributed, the figures would average about a fourth higher throughout. 

Protective adaptations—The popular expression ‘a nasty bug’ 
undoubtedly has reference, in most instances, to insects of this order, 
many of which produce scents disagreeable to human senses. Theorists 
have assumed these must also be repulsive to animal predators, a 
doctrine briefly stated in the following quotation from E. B. Poulton: 
“The Heteroptera (Hemiptera) are obviously, as a whole, a specially 
protected group, commonly defended by taste or smell from large 
numbers of insect-eating animals.” * 

A great series of Heteroptera are more or less aquatic in habit and 
thus are screened from the attacks of purely terrestrial enemies. 
Some are very active, as the Saldidae and many Miridae; some are 
said to be “ mimics,” as for example immature Nabidae resembling 
ants and certain Reduviidae resembling wasps. 

Mimicry, so-called, is exemplified among the Homoptera, also, as 
some Fulgoridae are considered to resemble Lepidoptera in appear- 
ance. The Membracidae with a host of bizarre forms, are thought to 
present cases of mimicry to ants, and of resemblance to thorns and 
seed pods of plants. One author further remarks: “ Evidently the 
strong pronotal processes, which are often sharp and hard enough to 
pierce the skin if the insect is seized suddenly, are unpalatable and 
irritating.” “ Quoting Poulton again (op. cit., p. 4): ‘Allusion must 





‘In Buckton, G. B., A monograph of the Membracidae, separate, p. 3, 1903. 
*Funkhouser, W. D., Mem. 11, Cornell Univ. Agr. Exp. Sta., p. 417, June, 
IQI7. 





42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. Ser 


be made to the special and curious defence by a waxy secretion which 
is common in the Homoptera. The method may be compared to the 
defensive silken walls of the cocoon in other insects, while the long 
trailing filaments of wax borne by certain species of Homoptera may 
play the same part as the ‘ tails’ on the hind wings of many Lepidop- 
tera, or the ‘tussocks’ of hair on some of their larvae—all these 
probably acting as directive structures which divert the attention of 
an enemy from the vital parts.” 

Many plant lice have the waxy filaments alluded to by Poulton, 
while most of them exude special secretions from the cornicles, sup- 
posed to be protective. Leafhoppers of various groups have been 
thought to resemble color or structural details of plants they frequent, 
and as for scale insects, their small size, waxy secretions and great 
resemblance to the bark upon which they rest, have given them high 
rank among the theoretically protected insects. Indeed they have been 
thought well-nigh immune to attack and one author has intimated that 
birds never eat scale insects. (Smith, J. B., Proc. State Hort. Soc. 
N.. J: vol. 20; p: 90; 1904.) 

Bird enemies.—Below are tabulations of the identifications of 
Hemiptera in the stomach contents of nearctic birds followed by sup- 
plemental comment. Comparative percentages are not given for the 
plant lice, scale insects, and mealybugs as these have not been cata- 
logued with the same degree of thoroughness as the other groups. 

Total number of identifications of Hemiptera, 22,395; percentage 
of identifications among those of all insects, 11.7301 ; percentage of 
species in this order among those of all insect species known, 8.5899. 

The Corixidae, although they spend practically all of their existence 
in water and usually on the bottom, do not thereby secure immunity 
from bird enemies. 

Like other hemiptera, however, they are supposed to be specially 
protected, one author saying: 


As to the function of the stink-apparatus in the adult Corixa, there is no 
need to look beyond defence. The insect frequently leaves the water, and it is 
then exposed to all the dangers met with by the land Heteroptera. Also there 
is no reason to doubt that the odoriferous secretion is equally efficacious against 
enemies in water.’ 


Results indicate that this efficacy is nothing remarkable; indeed it 
is a fallacy to suppose that so abundant and accessible a group does 
not pay due toll to predators. The number of species of birds that 


‘Brindley, Maud D. Haviland, On the repugnatorial glands of Corixa, 
Trans. Ent. Soc. London, vol. 77, p. 13, 1929. 


PROTECTIVE ADAPTATIONS—McATEE 43 


Identifications of Rhynchota 





| 


Percentage 
of species 
Percentage of in this group 
identifications among the 
among those whole number 
Number of of all of nearctic 
Group identifications Rhynchota Rhynchota 1 
Unidentified .............. 5,050 27.5325 
Heteroptera (further uni- 
dentified)) to..c 00.00.0608 389 1.8956 Sete 
Scutelleridae ............. 187 -QII2 8673 
MOGIGACE eis sock e cate wcs 232 1.1305 1.5077 
Rentatomidae ........+0:. 5,582 27.2011 5.5037 
Coreidae: 26-566. ssc caren c 305 1.9248 4.1361 
PNTadidae® Gi hocaasdetu selene 15 .0731 2.0014 
Neididae .2.........05-55. 7 0828 2668 
VY SACIGAG “oc sc 5k. se esses 524 2.5334 2.8353 
Pyrrhocoridae ............ 18 .0877 7338 
Mingitidae .............:. 66 3216 2.3349 
Enicocephalidae .......... ee ee 0667 
Phymatidae .............. 19 1.0926 .4003 
IREGUVIIdAG .oecce.. es. sss 633 3.4846 3.7692 
BICDLIMAe 2.6.6. nsec nan ee 2 .0097 1334 
| Mesoveliidae ............. 16 0780 0334 
INADTGAG acces ai hoa 4 ne oes 163 7943 7005 
| Gimicidae’ eae ces ats one 2 .0097 1334 
| Amnthocoridae ............ 3 .0146 1.1007 
Termatophylidae ......... a eee .0334 
IMMTIGAE. isco oes. 0d ae oa. 518 2.5242 14.8101 
Isometopidae ............. 1334 
Dipsocoridae ............. .I001 
Schizopteridae ............ or wie 0334 
Hydrometridae ........... 22 .1072 0067 
PREETIGdE 629% ocees ce ga ds. 228 I.11T0 6671 
Veliidae ..............000. 32 -1559 6004 
Saldidaes so.a. 6+ cece ciee se 74 3606 1.0674 
Notonectidae ............. 327 1.5935 6004 
Naucoridae .............. 306 1.4911 .4336 
INGDICAE: chs-0. 0:0 0s o ecioe secs 40 .1949 2668 
Belostomidae ............. 326 1.5886 .6671 
| Gelastocoridae ............ 3 £0146 2001 
| Ochteridae ............... a ae 1001 
| Worixidae. 225... .02..--:- 4391 6.7783 2.0347 
| Homoptera (further uni- 
Gentine )...asescess can 107 5214 we 
Oi rahdae..aiaisais ficewiete way 556 2.7004 2.7685 
| Gercopidae ..scs..s.-0..0 6 102 .4970 8673 
| Membracidae ............. 960 4.6781 6.2042 
| rcadellidae’ 3... ... 22.6521 1,435 6.9927 25.0837 
| Fulgoridae ............... 590 2875 IIl.9414. 
| PS VMNIGAC 23. a)3.e 00,644 cece ma's-s 122 5945 4.7365 
| 
| 
; 


P 


California Publ. Ent., vol. 2, 902 pp., 1917. 


*Computed from Van Duzee, E. P., Catalogue of the Hemiptera of America 
north of Mexico, excepting the Aphididae, Coccidae and Aleurodidae. Univ. 





44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


feed upon them shown by the present tabulation is 85 and the number 
of specimens taken at a meal ran over 200 in several cases, and up to. 
1,300 in one instance (eared grebe). The Belostomidae or giant water 
bugs, including the largest of North American Heteroptera, have 
strong, grasping forelegs and a stout beak which readily pierces the 
skin of man, making an aching, evidently envenomed wound. Not- 
withstanding these characteristics they do not escape the birds. Fifty- 
three species are on our list of captors, ducks, herons, and the like 
preponderating ; in two cases, both herons, as many as IO specimens 
were found in a stomach at one time. The Notonectidae again are 
sharply biting and exceedingly active under-water bugs ; but the larger 
types are eaten by no fewer than 44 species of birds, sometimes in 
considerable number (30-57), while the little crawling and obscure 
Plea were identified in 60 stomachs of 12 species of birds, in numbers 
up to 40 in a single instance. The Gerridae, so very active on the 
water surface, fall a prey to at least 49 kinds of birds, sometimes 
being taken in considerable numbers (20-40). The Miridae or plant 
bugs are agile and rapid in their movements and of great variety in 
form and color, but corresponding with their abundance and wide 
distribution, we find them preyed upon by 108 species of birds. The 
Anthocoridae and Cimicidae, both odoriferous families, seem poorly 
represented in our tables, but from their habits we should hardly 
expect the latter to be found at all, while most Anthocoridae also live 
largely hidden lives. We have found Nabidae in the stomachs of 
52 species of birds, Emesidae in 10, and Reduviidae in 115; these 
highly predatory forms therefore seem to have bird enemies about in 
proportion to their abundance. 

There are only a few species of Pyrrhocoridae in the United States 
and none of them are abundant ; hence the 18 captures by nine species 
of birds are perhaps not below proportion ; while the relations of birds 
to the Lygaeidae shows again that an abundant and diversified group 
is sure to be frequently taken by a large variety of birds. In this 
family may be specially mentioned Myodocha serripes, a bug with an 
extraordinarily long neck, for what purpose is unknown; at any rate 
it is one of the most bizarre of the group in our area, but it is eaten 
by more than 20 species of birds and no fewer than 27 specimens have 
been found in a single stomach (purple martin). The large red and 
black Lygaeus species were taken by 14 species of birds, and the 
superabundant chinch bug (Blissus leucopterus), frequently observed 
in prodigious numbers, by 29. Three species of birds, the bobwhite, 
meadowlark, and brown thrasher, had records of a hundred or more 
chinch bugs at a meal. These facts contrast strongly with the state- 





NO. 7 PROTECTIVE ADAPTATIONS—McATEE 45 


ment that ‘‘ Very few birds prey upon it because of its repulsive smell 
and taste. It is questionable whether any of them are fond of it.” 
(Garman, H., Bull. 74, Ky. Agr. Exp. Sta., p. 56, May, 1898.) In the 
series of Heteroptera composing the Coreidae and the groups aggre- 
gated as the Pentatomidae or Pentatomoidea, we have the typically 
stinking bugs. Practically all of them have powerfully scented secre- 
tions usually of a character obnoxious to man, but it is not evident 
that they are equally so to birds. Some of our Coreids (Thasus) are 
too large for most of our birds to prey upon, but those of the next 
rank in size are more or less freely taken, as Acanthocephala by 12 
species, in numbers as high as 14-22 by Franklin’s gull; and the 
various species of Leptoglossus by 16, 10-15 individuals at a meal by 
the same gull; Alydinae, nearly as large and equally smelly, are preyed 
upon by 21 kinds of birds. All Pentatomids are eaten so freely that 
it is difficult to pick the most representative examples. However, to 
begin, let us consider Podisus, a predacious, but nevertheless highly 
scented genus; it has been found in the stomachs of 29 species of 
birds, the most remarkable record being for a bird not included in 
these tabulations, namely a black duck collected in Maine, which had 
in its gullet alone 525 specimens of Podisus sereiventris. 

One of the largest and most highly scented stink bugs of our fauna 
(Acrosternum hilaris) was found in the stomachs of 37 species of 
birds, in number up to 26 in one instance (purple martin), while for 
our typical and most abundant genus ( Fuschistus) 62 avian predators 
are known. The number of specimens found in a stomach exceeded 
10 in a number of cases, and in one, that of a Franklin’s gull, 
reached 175. The little Thyreocoridae, polished black with touches 
of yellow on the costa, were found in the stomachs of 65 kinds of 
birds, and the Scutelleridae in 60. 

FE. A. D’Abreu in his report on “ Some insect prey of birds in the 
(Rep. Proc. Third Int. Meeting, Pusa, 





Central Provinces ” [of India 
1919, Vol. iti, p. 866, 1920) says “ Pentatomids seem a favorite diet 
with birds.” He gives notes also on bird enemies of 16 other families 
of Rhynchota. 

The only report on the food habits of birds in the American Tropics, 
namely, the “ Birds of Porto Rico” (Bull. 326, U. S. Dep. Agr., 
1916), by Alexander Wetmore, in the accounts of the species through- 
out shows Hemiptera to be taken in due proportion. 

The Cicadidae are chiefly large insects, a factor which to some 
extent must limit the number of their bird enemies ; however the list 
here drawn upon shows 87 species and there are four records for one 
of our smallest birds, the house wren. Some of the larger birds 





40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


devour considerable numbers of the smaller cicadas, for example, 
30 Okanagana rimosa were found in a nighthawk’s stomach and from 
19 to 41 Tibicina septendecim in each of several crow stomachs. It is 
of interest in this connection that adult as well as immature domestic 
fowls have been killed by crop-binding due to eating too many cicadas. 
(Weekly News Letter, U. S. Dep. Agr., vol. 6, no. 46, p. 14, June 18, 
1919.) Wild birds, however, not only take large numbers of cicadas, 
but feed on them steadily day after day when the chance comes. The 
English sparrow and the crow blackbird are notable examples of this 
and it has been concluded by entomologists that broods of the 
periodical cicada issuing in parks and other places, where exposed to 
concentrated attacks of these species, are doomed to extinction. 
(Smith, J. B., Economic entomology, pp. 142-143, [1896]; Mar- 
latt, C. L.,. Bull. 90, U.S. Dep: Ags. py 10, 1804.) 

Our records do not show whether any immature Cercopidae 
(spittle insects) are eaten by birds, but the adults are taken by 41 
species. One chimney swift had eaten about 100 cercopids of the 
genus Clastoptera. Despite the numerous defenses they are said to 
have, Membracidae were eaten by no fewer than 136 species of birds 
represented in the present tabulation and in numbers up to 26 indi- 
viduals in a single stomach. They have been found in 15 or more 
stomachs of each of the following species: Least, great-crested and 
ash-throated flycatchers, wood pewee, meadowlark, Brewer’s black- 
bird, Bullock’s oriole, English sparrow, cliff swallow, red-eyed, soli- 
tary, and warbling vireos, bush-tit, and ruby-crowned kinglet. The 
tree hoppers identified belong to 21 different genera, indicating that no 
partiality is shown. Membracids with the most prominent horns and 
spines of any in our fauna, such as those of the genera Campylenchia, 
Platycotis, Thelia, Ceresa, and Platycentrus, are taken with the rest. 
During stomach examinations 175 kinds of nearctic birds have yielded 
leafhoppers (Jassidae sens. lat.) and 10 or more stomachs of no 
fewer than 35 species have disclosed them. In a number of cases 
from 20 to 50 leafhoppers were found in single stomachs and in one 
case (barn swallow) a thousand. 

The fulgorid fauna of the United States is scanty and our records 
of birds feeding on these insects correspond. Beyond the fact that 
they are well distributed through the various groups of the family 
and pertain to 18 species of birds, there is little of special interest 
concerning them. Some lesser yellow-legs had eaten from 50 to 
400 each. The Psyllidae were found in the stomachs of 46 kinds 
of birds and the Aphididae in 86. Cases are known in which the 


* 


NO. 7 PROTECTIVE ADAPTATIONS—McATEE 47 


former have been devoured very extensively by birds, an entire 
orchard having been cleared of the pear psylla by nuthatches. (Zool. 
Bull. Pennsylvania Dep. Agr., vol. 3, p. 79, July, 1907.) Plant lice 
were found in large numbers in the stomachs of some birds, up to 
200 or more in each of five species of the finch family, 300 or more 
in three of them (pine siskin and two goldfinches), about 650 in the 
stomach of a nighthawk and 1,600 in that of a wood duck. On a 
200-acre farm in North Carolina, birds were found to be destroying 
more than a million grain aphids daily. (McAtee, W. L., Yearbook, 
U. S. Dep. Agr. (1912), pp. 397-404, 1913.) Aleurodidae have not 
as yet been identified from stomachs of nearctic birds ; possibly some 
may have been confused with scale-insects. The latter, notwith- 
standing deprecatory statements that have been made relative to birds 
as predators upon them, have been found in the stomachs of 88 species 
of nearctic birds. No fewer than 100 Eulecanium cerasifex were 
found in the stomach of a rose-breasted grosbeak, 300 Margarodes 
in one of a scaled quail, 304 Saissetia oleae in that of a black-headed 
grosbeak and 200, 700, and 800 of the same scale, respectively, in 
three stomachs of the pine siskin. 

Other enemies.—Salamanders, toads, and frogs are recorded in the 
| Pennsylvania reports as feeding upon both Heteroptera and Homop- 
' tera, as are also the common swift lizard (Sceloporus undulatus) and 
_ the copperhead (Agkistrodon contortrix) and the hog-nosed snake 
(Heterodon platirhinos). The same source credits five species of 
| turtles with eating Heteroptera and one with devouring Homoptera. 

Munz found that all the common frogs feed on Hemiptera about as 

freely as upon any other insects, and Garman found bugs in 6 out of 
20 stomachs of the common toad. Winton reports the Texas horned 
| 
| 
| 
. 





lizard (Phrynosoma cornutum) as eating stink bugs. 

Aquatic hemiptera, particularly Corixidae, are eaten by most fresh- 
water fishes, while scattering representatives of the terrestrial families 
are taken now and then as opportunities occur. Forbes records from 
fish stomachs representatives of 14 families of Heteroptera and three 
of Homoptera. Among mammals, the common mole is known to take 
leafhoppers, chinch bugs, and other species; shrews do not entirely 
neglect Hemiptera; the nine-banded armadillo devours Cydnidae and 
Pentatomidae. 

Insect enemies of Hemiptera include both nymphal and adult 
dragonflies, the former getting considerable numbers of Corixidae and 
the latter representatives of various families. Robber flies feed freely 
upon Hemiptera, ground beetles and ladybirds devour them, and the 

4 


ai 





48 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


Nyssonidae, Mimesidae, and Crabronidae, among predacious hymen- 
optera, prey more or less selectively upon Homoptera; in the eastern 
States a large Sphegid wasp is a special foe of cicadas. Other enemies 
of cicadas include dragonflies, wasps, predatory beetles and bugs, 
mantids, spiders, mites, hymenopterous and dipterous parasites, 
fishes, snakes, turtles, squirrels, badgers, armadillos, skunks, moles, 
and fungi. Spiders consume many Hemiptera of a wide variety and 
are credited with being among the most important natural enemies of 
leafhoppers. The latter insects are heavily parasitized by the Dry- 
inidae, and by at least five other families of Hymenoptera, by Pipun- 
culidae, and Strepsiptera; and are preyed upon by larvae of Chryso- 
pidae, and by Coccinellidae, Reduviidae, and certain other insects. 
The Pyrrhocoridae said to be specially protected are preyed upon 
by spiders, pseudoscorpions, thrips (the eggs), tachinid flies, reduviid 
bugs, and lizards. The Coreidae have special parasitic foes among the 
Tachinidae; while the order of Rhynchota in general is subject to 
hymenopterous parasites, the abundance of plant lice and scale insects 
in particular depending to a large degree in many cases upon the 
relative numbers of these destructive foes. Lycaenid caterpillars feed 
upon aphids, coccids, jassids, and membracids. A page would scarcely 
suffice to list the numerous enemies of plant lice which include, besides 
parasites, coccinellid, lampyrid, syrphid, hemerobiid, and chrysopid 
larvae, in addition to adult ladybird beetles, assassin bugs, and other 
insects, mites, and spiders. Fungi are known to destroy, at times, 
large numbers of hemiptera, among which may be mentioned plant 
lice, scale insects, mealybugs, and the chinch bug. 
Discussion.—Despite their malodorous secretions and other “ pro- 
tective devices ’’ there can be no doubt that Rhynchota are taken fully 
in proportion to their abundance by nearctic birds, and the evidence 
is that their other enemies are numerous and effective. If we consider 
the most pronouncedly repugnant species found in the United States, 
such as the harlequin bug (Murgantia histrionica) and the squash bug 
(Anasa tristis),’ we find that severe infestations of the former have 
been kept in check by English sparrows (Sherman, F., Bull. North 
Carolina Dep. Agr., vol. 32, no. 7; p. 21, July, 1911) and that the 
squash bug has a number of deadly enemies. It has been shown that 
the volatilized secretions of squash bugs if confined in a glass con- 
tainer are capable of killing toads (Weed and Conradi, Bull. 89, New 


‘It is worth noting that of these two common and exceedingly malodorous 
bugs, one is warningly, one obscurely colored. Where is the correlation that 
theories as to warning colors demand ? 





———— 


NO. 7 PROTECTIVE ADAPTATIONS—McATEE 49 


Hampshire Agr. Exp. Sta., pp. 21-23, Feb., 1902), and the conclusion 
was drawn that “ toads do not ordinarily devour many of these pests.” 
Perhaps they do not “devour many ” of them, nor, with the whole 
insect world available for them to prey upon, should they be expected 
to specialize upon squash bugs, but they do eat them, as found by 
Kirkland (Bull. 46, Hatch Exp. Sta., p. 26, 1897) and also by 
Biological Survey investigators. Bird enemies also are not lacking, 
present records showing six species of birds known to feed upon 
Anasa tristis and four upon other species of the genus. The harlequin 
bug is sometimes heavily parasitized also, while the squash bug has 
both tachinid and hymenopterous parasites and is subject to a bacterial 
disease. 

Disregarding the “ protective adaptations”’ and reasoning alone 
from the prevalence of hemiptera, there would be no presumption 
that these insects would constitute a tenth of the food of any species 
of birds, yet they actually do contribute 10 per cent or more of the 
subsistence of the following 12 species in the United States: Nuttall 
woodpecker (number of stomachs examined 53), percentage of 
Rhynchota in the food, 14.76 per cent ; Scissor-tailed flycatcher (129), 
10.17 per cent; eastern phoebe (370), 10.38 per cent; black phoebe 
(344), 10.56 per cent; crested flycatcher (265), 14.26 per cent; least 
mycatcher (177), 11.12 per cent; Bullock’s oriole (162), 10 per cent; 
sharp-tailed sparrow (51), 12 per cent ; spotted towhee (139), 14 per 
cent; purple martin (205), 14.58 per cent; barn swallow (467), 
15.1 per cent ; and rough-winged swallow (136), 14.9 per cent ; more 
than 20 per cent of the food of two birds, namely the black-headed 
grosbeak (225), 21 per cent, and cliff swallow (375), 26.32 per cent, 
and more than 30 per cent of the total diet of the violet-green swallow 
(110), 35.96 per cent. 

If the glandular secretions of hemiptera had the repugnatorial, not 
to say dangerous, qualities attributed to them, there would be no such 
wholesale preying upon them as is shown in the foregoing data. 
Descending to milder forms of “ protection” as afforded by pointed 
protuberances and secretions of wax, we find that the “ hardihood ” 
of birds (from the selectionist point of view), or in other words their 
tendency not to be bound by human criteria, is so great that such 
devices simply do not count. 


NEUROPTEROIDEA (DOBSONFLIES, SNAKEFLIES, SCORPIONFLIES, ANT-LIONS, 
CADDISFLIES ) 
In the period during which the records of bird food here discussed 
were obtained, the conception of the group of insects broadly termed 
Neuroptera has gradually evolved from that of a catch-all for net- 





50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


veined insects to a restricted group of very few families. Hence 
many of the determinations (in want of re-examination) cannot be 
definitely correlated with modern classification. With little doubt, 
therefore, there has been confusion of records between Neuropter- 
oidea (in the present sense) and some of the orders elsewhere discussed 
as Agnatha, Mecaptera, and especially Plecoptera. Hence the tabula- 
tion figures given are only approximations, and a better conception 
of the relations of birds to the insects would be obtained by lumping 
all of the so-called net-veined insects together. 

Protective adaptations—The Neuropteroidea have not been given 
so much attention by adaptationists as some other groups of insects, 
but certain supposedly protective features have been pointed out or are 
suggested by analogy with other described cases. Dobsonflies are 
large, the larvae and females have powerful biting-jaws, while in 
some cases the males have enormously developed mandibles of less 
sturdy construction, and the coloration of the wings of some presents 
strong contrasts. The latter characteristic is possessed by the Myrme- 
leonidae also, while their larvae, the ant-lions, have large, strong jaws 
and an antlike odor. Some Ascalaphidae are said to resemble dragon- 
flies both in appearance and habits; Chrysopidae have vile smells 
earning them the name of stink flies; and Mantispidae not only have 
predatory forelegs but are said to be protected by their resemblance 
to Hymenoptera. (Poulton, E. B., Trans. Ent. Soc. London, 1902, 
p- 536.) “ The well-known cases of Caddice-worms (Trichoptera) 
are partly for concealment and partly for defence.” (Poulton, Colours 
of Animals, p. 77, 1890.) Some of them resemble snail-shells. 

Bird enemies.—While Neuroptera (sens. lat.) have been identified 
from the stomachs of 56 species of birds there is little object in dis- 
cussing further this heterogenous assemblage. Sialidae (dobsonflies 
etc.), despite their average large size and biting powers, were taken 
by 38 kinds of birds; 58 specimens were found in the stomach of a 
Bonaparte’s gull and from 55 to 93 larvae in three stomachs of lesser 
scaups and 192 in one of a canvas-back. Snakeflies, of bizarre ap- 
pearance, and of limited distribution in the United States were 
identified in the food of 22 species of birds; and Mantispidae, “ pro- 
tected by their resemblance to Hymenoptera,” and also by considerable 
rarity in our fauna, were found in the stomach of 11 species. Stink 
flies (Chrysopidae) were eaten by 18 kinds of birds, and Myrme- 
leonidae by 20. Ascalaphus was identified but once, quite in keeping 
with its extreme rarity, and Hemerobiidae 11 times. The figures for 
identifications are low for scarce or locally distributed groups, but 


4 
| 
| 
| 
4 
| 





NO. 7 PROTECTIVE ADAPTATIONS—McATEE SI 


when we come to one of common and general occurrence, the corre- 
sponding rise in frequency of capture by birds is apparent at once. 
The caddisflies, more numerous in species and individuals than all our 
other Neuropteroidea together, appropriately contribute nearly two- 
thirds of the total number of records for the group. The number of 
species of birds feeding upon them is 113, and of these 45 or more had 
taken the “ specially protected’ larvae. The number of records of 
caddisflies determined was 10 or more for 23 species of birds, and 
more than 20, 30, and 40 in the case of four, three, and three species 
respectively. The number of specimens taken by single birds exceeded 
30 of larvae in a number of instances and ran as high as 207 (in a 
scaup duck), and of adults reached such figures as 280 and 400 in 
the case of the nighthawk. 


Identifications of Neuropteroidea 


Percentage 
of species 


Percentage of in this group 
identifications among the 
among those whole number 
Number of of all of nearctic 
Group identifications Neuropteroidea Neuropteroidea + 
Neuroptera (sens. lat.).... 119 9.0504 sha 
Megaloptera ....5...6.00% 167 12.7094 3.5648 
Rhapidioidea ............. 54 4.1096 1.6886 
Neuroptera (sens. str.).... 108 8.2192 32.2700 
IPhryeanoidea ......+s0+5% 866 65.9061 62.4774 


All Neuropteroidea ....... 1,314 


Other enemies —Forbes reports that neuropteroid larvae compose 
about 10 per cent of the food of the sucker and catfish families in 
Illinois ; he found caddis larvae in the stomachs of 17 species of fishes. 
According to various authors, these larvae are an important element 
in the food of most kinds of trout. Salamanders, frogs, larvae of 
stoneflies, and parasitic hymenoptera also are enumerated among the 
enemies of caddis larvae. Forbes found larvae of Sialidae in seven 
species of fishes, and these are known to be eaten also by frogs and 
turtles. Chrysopidae have been seen to be eaten by frogs, salamanders, 
and ants, and they have numerous hymenopterous parasites sometimes 
destroying inmates of about half of the cocoons. (McGregor, E. A., 
Can. Ent., vol. 46, pp. 306-308, 1914.) Frogs are recorded also as 
capturing Mecaptera, as are also lizards and larvae of ant-lions. 
Robber flies and dragonflies apparently devour any Neuroptera chance 
throws their way. 





*Computed from Banks, Nathan, Catalogue of neuropteroid insects (except 
Odonata) of the United States, 53 pp., 1907. 





52 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


Discussion.—lIt is obvious from the available data on enemies of 


Neuropteroidea that the small or rare groups have few, the large and 
abundant families many foes, the result that would be predicted with 
‘protective adaptations” discounted. The group most numerous in 
species and individuals, namely the caddisflies, has the most enemies, 
and their larvae, said to be well defended from enemies, form one of 
the staple elements of the food of fresh-water fishes all over the globe, 
as well as a favorite prey of aquatic birds. 


LEPIDOPTERA (MOTHS, BUTTERFLIES ) 


Protective adaptations—In the space that can be devoted in this 
paper to protective adaptations of Lepidoptera it is impossible to do 
more than call attention to general aspects of theoretical considera- 
tions, since what has been written on the subject would fill many 
volumes. This flood of literature is due principally to the fact that 
Lepidoptera have been regarded as the chief examples of the phe- 
nomena of warning colors and of mimicry, subjects that have been 
expounded and discussed at great length. 

Warning coloration, it need hardly be stated, designates the con- 
spicuous, often highly contrasted, patterns, which it is held may be 
assumed with relative impunity by tough, distasteful, or dangerous 
species. Batesian mimicry is the more or less pronounced resemblance 
to these species by others supposedly less qualified to cope with the 
struggle for existence, while Mullerian mimicry is mutual approach 
in appearance by species all of which belong to “ specially protected ” 
groups. As remarked in my 1912 paper, these theories were chiefly 
built up at a time when there was almost complete ignorance of the 
actual feeding habits of predacious animals, and attempts to secure 
evidence on the subject by experiment were in most cases characterized 
by a singular lack of appreciation of the vital factors involved and of 
realities in nature. 

The following statement by Alfred Russell Wallace gives the gist 
of the principal nearctic instances of mimicry among Lepidoptera: 
“In North America, the large and handsome Danais archippus with 
rich reddish-brown wings is very common, and it is closely imitated 
by Limenitis misippus, a butterfly . . . . which has acquired a color 
quite distinct from that of the great bulk of its allies. In the same 
country there is a more interesting case. The beautiful dark bronzy- 
green butterfly, Papilio philenor, is inedible both in larva and perfect 
insect, and it is mimicked by the equally dark Limenitis ursula. There 
is also in the Southern and Western States a dark female form of the 





NO. 7 PROTECTIVE ADAPTATIONS—-McATEE 53 


yellow Papilio turnus, which in all probability obtains protection from 
its general resemblance to P. philenor.” (Darwinism, p. 248, 1896.) 

Mimicry of another order of insects, the Hymenoptera, is shown 
by many of the clear-winged moths (Syntomidae and Sesiidae) as 
adults; and of black-and-yellow ringed larvae, it is said they gain 
great advantages from resemblance to the justly respected appearance 
of hornets and wasps. 

The majority of adults of Lepidoptera, especially the moths, exhibit 
in greater or less perfection what is called cryptic coloration, that is 
resemblance to details of the environment, exemplified by the species 
that are inconspicuous on bark, old leaves and the like. This style of 
protective adaptation also is attributed to many larvae and pupae. On 
this topic Poulton says: “ There is no better instance of special pro- 
tective resemblance than that afforded by the larvae of Geometrae, 
‘stick caterpillars’ or loopers as they are often called. These cater- 
pillars are extremely common and between two and three hundred 
species are found in this country [Great Britain]; but the great 
majority are rarely seen because of their perfect resemblance to the 
twigs of the plants upon which they feed.” (Poulton, E. B., The 
colours of animals, p. 26, 1890.) 

This idea is pushed to an extreme by another author as shown by 
the following quotation relating to the caterpillar of “a geometrid 
moth. In the larval state the insect bears a very close resemblance 
toa twig. Its habit of clinging to a real twig with its posterior ‘legs’ 
and allowing the body to swing out, adds to the illusion. The head 
of the caterpillar resembles a leaf bud, while in color the entire 
creature is an exact counterpart of a rough apple twig, the plant upon 
which it naturally feeds. Thus complete immunity is secured from the 
attacks of birds and all enemies which depend chiefly upon sight.” * 
(Howes, Paul Griswold, Insect behavior, pp. 164-165, 1919.) 

Adaptations of caterpillars supposed to repel enemies, which have 
received the most attention from writers on the subject, include: 
armatures of hairs or spines, repugnant odors, warning colors, and 
terrifying attitudes, in addition to various special resemblances. 
Among the latter, Howes considers especially remarkable those that 
“rely for their protection upon their mimicry of the excreta of birds. 
I have been completely fooled by these larvae on more than one 
occasion. They frequently rest in the center of a green leaf and while 
conspicuous, never suggest a living insect to the uninitiated. In color, 
the upper and lower portions of the body are dark chocolate brown, 


*See pages 56 and 85 for the facts as to immunity of loopers. 


7 





54 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


banded through the center with pure white, which suggests the lime 
so often seen in the excreta of birds. The entire creature is highly 
glossed, which gives a fresh and moist appearance to the object, which 
makes no attempt to conceal itself, depending entirely upon its 
strangely camouflaged body for protection.” (Insect behavior, p. 165, 
TOLOS) 9 

Poulton summarizes the purpose of caterpillar adaptations as fol- 
lows: “In the remarkable. abundance and variety of methods by 
which concealment is effected in Lepidopterous larvae, we probably 
see a result of their peculiarly defenceless condition. .... Hence 
larvae are so colored as to avoid detection or to warn of some un- 
pleasant attribute, the object in both cases being the same—to leave 
the larva untouched, a touch being practically fatal.’ (The colours of 
animals, p. 51, 1890.) 

On the concealment of lepidopterous pupae, the same author says: 
‘Protective Resemblance, either Special or General, is seen in nearly 
all exposed pupae, but most chrysalides are buried in the earth or 
protected by cocoons. The cocoons are often sufficient defense, 
because the silk is very unpleasant in the mouth; but such protection 
only applies in the warmer weather when there is an abundance of 
insect food. In the winter, insectivorous animals are pinched by 
hunger, and would devour the pupa in spite of the cocoon. We there- 
fore find that all cocoons which contain pupae during the winter are | 
well concealed, either spun between leaves which fall off and become 
brown, or hidden under bark or moss, or constructed on the surface 
of bark with a color and texture which renders them extremely diffi- 
cult to detect.” (Ops cit; pp. S1-52-) 

Pausing only long enough to note the incorrectness of the statement 
‘all cocoons which contain pupae during the winter are well con- | 
cealed ” (witness those of Saturniidae, not to speak of the cases of 
many Tineidae), we may pass to Howes’ more imaginative account. 


We find, for instance, the chrysalis of a butterfly, a species of Vanessa. It | 
hangs by a tiny silk-fastened stem under a protecting fence rail. Within the 
shell of the chrysalis, there is nothing but a mass of disintegrating tissues, a 
thick fluid, studded with globules of fat. It is neither caterpillar nor butterfly. 
It cannot thrash about from side to side or make a demonstration, there are no_ 
spines to pierce a would-be enemy, no wings by which the creature might take 
flight. It is as helpless now as so much custard, for the insect is in the process 
of change from one form to another. 


*This comment ignores the fact that a great many birds habitually devour | 
the excreta of their young, even returning to it when accidentally dropped, and 
this nestling excreta is exactly of the luscious appearance described by Howes. 


a 


NO: 7 PROTECTIVE ADAPTATIONS—McATEE 55 


Such is the actual condition of the pupal butterfly, but let us examine its 
outer covering. It is a frightful-looking object, armored, and covered with 
sharp spikes between which beady false eyes peer out. It is absolutely harmless 
but appears otherwise. To birds it is doubtless a thing to beware of, yet one 
tiny puncture of its brittle covering would reveal a delicious feast within. 

Many insects are thus protected, ones that could not compete in any form 
of battle. They are given immunity from attack because they could not ward 
it off themselves. In the case of the transforming pupa, some such form of 
protection becomes a necessity. A butterfly in the making is as helpless as the 
egg from which it sprung, so Nature resorts to camouflage to terrorize the 
destroyers of her children. (Insect behavior, p. 168, 1910.) 


Aside from the fact that Vanessa pupae do not enjoy immunity 
(see p. 62), we may well inquire whether birds are not Nature’s 
children just as much as the butterflies, and just as fully entitled to 
be her beneficiaries ? 

Bird enemies.—Identification to species especially has lagged more 
in the case of lepidopterous items of food, than in those of any of the 
other larger orders of insects, due chiefly to poor condition of the 
remains of adults, and to lack of knowledge of larvae. Unidentified 
Lepidoptera exceed 2.85 times those in some degree identified, and in 
considering the relation of the percentages of identifications to those 
of the number of species of various groups, the former figures should 
be multiplied by 2.85. 

In view of the very unsatisfactory distribution of identifications of 
_ Lepidoptera to families (over 70 per cent of the whole number being 
| merely as Lepidoptera), it would be of little avail to discuss the 

relative importance of family groups as bird food. Rather it will be 
better to treat the subject along lines of general interest already 
developed, as the preference between larval and adult Lepidoptera, 
the extent to which hairy caterpillars are eaten, and the relation of 
birds to butterflies, the chief illustrations of mimicry theories. 

The question as to which is eaten most extensively, adult or larval 
Lepidoptera, is easily answered in favor of the latter. As the table 
shows 68 per cent of all records of Lepidoptera are for larvae, further 
unidentified ; moreover, it is certain that the great bulk of specimens 
identified to families also were larvae. Thus the Noctuidae deter- 

-mined were chiefly cutworms, the Geometridae were mostly loopers, 
the Tineidae principally case-bearers, and so on. Caterpillars, not 
further identified, were found from 50 to 100 times in the stomachs 
of 23 species of birds; from 100 to 200 times in 21 species ; from 200 
to 300 times in eight species (downy woodpecker, blue jay, red-winged 
and Brewer’s blackbirds, warbling vireo, black-capped chickadee, 
hermit thrush, and bluebird) ; from 300 to 400 times in two species 








: 


SMITHSONIAN MISCELLANEOUS COLLECTIONS 


Identifications of Lepidoptera 


Gelechtidde 445-6 elena 
drortricidacwencts scien 
Grambidaeweeeee neers 
Pyralidae: Fess ccm ere 
Sesiidaet tans hum. c oven 
(Cossidaemeprer cee een ete 


Psychiddensstraaus. ete ees 
Geometridae ee ne 
Bombycidae en) snes nee 
[Easicampidae mse 2 hate: 
[iparidae f.hcoraen eat 
Notodontidae 4 2i2¢25..20. 
INoctuidae tne ae iae ens 
INGarIStidacumaiee recite 
AT ctiidaeweree eee eee ene 
Ceratocampidae- 5-45-42 - 
Satunntidaeweee see eee eer 
Sphingidae haere erates 
Moths (further unidenti- 

Hed) L aie wmech seen ors 
ALLE VOLS oe eer eee 
Elespeniidaem amen 
Teyana ci daeaeemreertiee tree 
Nymphalidae) .:2...-....+- 
Pieridaes sce sce ae sien Oe 
Bapilionidae seer mere 
Butterflies (further  un- 

Ad entitled) eee reer 
AU DULLCRTUVE St rtaiacioieis eee’ 
Lepidopterous eggs ...... 
Lepidopterous larvae ..... 
Lepidopterous cocoons and 

chysalidestys ca -peeiies 
Lepidopterous adults ..... 


Number of 
identifications 


499 


149 


38 


oO 
AN NH MN ND 


174 


118 
1,128 


68 


29 
156 


2,116 
4,083 


Percentage of 
identifications 
among all 
Lepidoptera 


2008 
0021 
0021 
.0626 
.0071 
.O160 
.0008 
.0021 
.0008 
.0008 
.0306 
" 0025 
.0731 
.0029 
.0496 
4742 
.OO17 
0286 
.0277 
.O122 
.0056 


.8896 
1.9085 
.0038 
0004 
.0248 
.0004 
.0008 


0172 
.0474 
4233 
5.3201 


0954 
2749 


Percentage 
of species 
in this family 
among 
described 
nearctic 
species of 
Lepidoptera 1 


6.3575 
2.9296 
4.4548 
7.2183 
10.0573 
1.5252 
3I70 
4983 
-1963 
12.2318 
JOISI 
3624 
.2265 
1.2533 
32.1049 
.2265 
1.7668 
1812 
4681 
1.2533 


90.1541 
2.0447 
-0453 
2.5218 
9664 
Bit 


9.8459 


*Computed from Dyar, H. G., A list of North American Lepidoptera, etc., 
U.S. Nat. Mus. Bull. 52, 723 pp., 1902. 


VOL. 85 





| 
t 
( 
{ 
| 








NO. 7 PROTECTIVE ADAPTATIONS—McATEE 57 


(red-eyed vireo and robin) ; and more than 400 times in the following 
five species: crow (438), starling (727), meadowlark (474), crow 
blackbird (600), and English sparrow (466). One hundred or more 
caterpillars further unidentified were found in single stomach con- 
tents of each of the following birds: sparrow hawk, downy wood- 
pecker, hairy woodpecker, black-billed cuckoo, yellow-billed cuckoo, 
crow, starling, crow blackbird, hermit thrush, wood thrush, and robin. 
A very characteristic phase of the destruction of caterpillars by birds 
is their use as a special food for the young ; numerous species of birds 
make a practice of feeding the young a very much higher proportion 
of caterpillars than is taken by the adults. 

It has often been asserted that hairs and spines are very effective 
in protecting certain caterpillars from birds." Bastin for instance, 
states that “stinging hairs defend their possessors from almost all 
birds except the cuckoos.” (Insects, their life-histories and habits, 
p. 168, 1913.) These claims ignore the fact that birds are véry well 
equipped with relatively insensitive bills and feet for removing spines 
and hairs from larvae if they choose. Some birds do this, others 
actually dissect caterpillars, eating parts they want from the inside, 
piecemeal. Hairy and spiny armature is no bar to birds with such 
feeding habits, and, furthermore, do not seem to be of any great 
service in relation to numerous birds which swallow entire larvae thus 
defended. A characteristic statement about hairy caterpillars is: 
“Tent caterpillars have few enemies..... Our two species of 
Cuckoos make it a regular business to feed upon these worms which 
no other birds will eat.” (Lugger, Otto, Fourth Ann. Rep. Ent. Minn. 
(1898), p. 142, 1899.) 

Seventeen of the species of birds included in the tabulations on 
which this paper is based had eaten tent caterpillars or the eggs from 
which they hatch; numbers of larvae taken at a meal ran up as high 
as 200 in case of the black-billed cuckoo, and of eggs as high as 1,047 
in that of a blue jay. Compiling records from the reports of ento- 
mologists and others who have found birds feeding upon tent cater- 
pillars, we get a list of 43 species of bird predators upon the so-called 
“Orchard” species (Malacosoma americana) and 32 upon the 
“Forest” species (IM. disstria). Caterpillars even more offensively 


*In the case of this as in other similar claims, we may, well ask why such 
theoretically effective defenses have not been developed by a larger proportion or 
in fact by all larvae? The most cursory consideration of the subject shows 
that hairiness of caterpillars is in the main a phyletic character. A few related 
families include the great bulk of the hairy larvae. 


* 


58 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 





hairy than these are those of the gipsy and the brown-tailed moths ; 
the hairs of the latter species especially cause a troublesome and 
painful rash upon the skin of man. Nevertheless 46 kinds of birds 
are known to eat caterpillars of the former species and 31 those of the 
latter. (For a valuable article on bird enemies of these and other hairy 
caterpillars, see Forbush, E. H., Bull. 20, n. s., U. S. Div. Enum 
pp- 85-93, 1899. ) | 
The larvae of the tussock moth (Orgyia leucostigma) are supposed 
to be especially distasteful to birds, but Forbush records (Mass. Crop 
Rep., July, 1900, p. 36) nine species of birds as feeding upon them. 
The writer*has observed English sparrows and robins eating them; | 
in the spring of 1921 in Washington, D. C., the larvae were quite 
common and robins were feeding freely upon them, carrying them 
to their young, I believe, as it was a common sight to see the birds | 
flying with the white tufts showing at the tips of their bills. | 
Again records of birds feeding on fall webworms (Hyphantria 

| 

\ 





fextor) are relatively scanty, only six species being named, yet careful 
observation in the field has proved one of them to be a very effective 
foe. Dr. C. Gordon Hewitt informs us that of the various factors 
operating in the reduction of this insect in Nova Scotia in 1916, the 
red-eyed vireo was most important and “it was estimated that about 
40 per cent of the larvae had been destroyed in the webs by this bird.” 
(Rep. Dominion Ent. 1917, p. 8.) A later report shows an average | 
destruction of 68 per cent. | 

It would not be necessary to refer to the preying of birds upon 
smooth caterpillars, a thing universally done, except for theoretical dis- 
quisitions as to the “protected nature” of certain groups. The 
Geometridae, loopers or measuring worms, are said to be protected by 
resemblance to twigs, etc., a statement made without giving due weight 
to the fact that such a defense depends upon immobility whereas these 
caterpillars must be in motion the greater part of the time while 
searching for and devouring food. Forty-four species of birds are 
recorded as feeding upon Geometridae in our tabulation and numbers 
of specimens as high as 20 were taken at one meal by the starling and 
go by the robin. 

Larvae of the Sphingidae are said to be protected by their “ horns ” 
and by “ terrifying attitudes,” but 44 species of birds covered by the 
present investigation do not seem to agree with the theorists on these 
points. Ten species are known to prey upon Dielephila lineata alone, 
and in field observation the crow has been known to clear tomato 
patches of the hornworm (Phlegethontius sexta). 





NO. 7 PROTECTIVE ADAPTATIONS—McATEE 59 


Many printed pages have been devoted to discussion of the question: 
“To birds eat butterflies? ’”’ but the natural answer: “ Certainly, but 
probably not out of proportion to their abundance,” seems not to have 
occurred to the disputants. 

At this point it will be well to say a word about the alleged difficulty 
of identifying adult Lepidoptera, especially butterflies in the stomach 
contents of birds. But for this, some argue, the number of records 
of butterflies eaten would be much larger. The assumption is made 
that the scales are necessary to identification, and since they are so 
easily rubbed off, determination will usually be impossible. This ob- 
jection serves mainly to exhibit the ignorance of its proposers relative 
to the analysis of the contents of bird stomachs. In the first place 
when adult Lepidoptera have been eaten at all recently, that fact is 
evident to the practiced eye, even unaided, on first glance at the 
stomach contents. A characteristic fuzzy, felted appearance, due to 
the distribution of the hundreds of scales throughout the mass, tells 
the tale at once. Even after digestion is far advanced the scales do 
not disappear because they are so numerous and stick to everything, 
and they are evident under magnifications used in the analysis of 
practically every stomach contents. Moreover were all scales absent, 
it would be possible to unroll the wing membrane, if swallowed, and 
examine the venation; the antennae also would usually be present ; 
and the form of the head, thorax, and body, which are characteristic, 
could be made out. 

In addition we would remind the reader that all things found in 
birds’ stomachs are not ground to a powder. Just the reverse in fact 
is true; birds feed more or less constantly, and whenever shot they 
will as a rule just have swallowed some article of food which, of 
course, will be in good condition for study. In the long run all con- 
stituents of the food will be found nearly or quite intact in the 
stomachs in proportion to the frequency in which they are taken. 

However it is unnecessary to discuss the matter further. One need 
only consider the extent to which we have identified certain insects 
far more fragile than butterflies, as mayflies (Ephemeridae) 484 
records, midges (Chironomidae) 1,003 records, and crane flies (T1- 
pulidae) 1,565 records, to be assured that there is no likelihood what- 
ever of a butterfly being overlooked during careful stomach analysis. 

Of the 113 records of birds eating Rhopalocera included in the 
present tabulations, 24 refer to larvae and two to chrysalides. It is 
worth noting that one of the larval records was for Anosia plexip pus, 
two for Papilio species, and six for l’anessa species, supposedly the 


60 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


best protected forms. Nymphalidae and Hesperiidae are most numer- 
ously represented among the adults taken. 

The 87 records of imago butterflies are distributed among T5 species 
of birds, but all save 18 of them pertain to a single bird, the pigeon 
hawk. The specimens of this hawk examined were taken on their 
southward migration at a point that is in the migration path of butter- 
flies also, so that opportunities for catching these insects were at the 
best. (It is worth noting here that dragonflies, swallows, swifts, and 
bats also using this same migration track were preyed upon by the 
pigeon hawk.) In this case as in many others the abundance and 
availability of prey is shown to have great influence upon the choice 
of food by birds. Amid the butterflies, this hawk preyed upon them; 
elsewhere we have no record of its doing so. Clearly the other birds 
(14 species) in whose stomachs butterflies have been found (18 
records). are only occasional predators upon them. This is only what 
would be expected, for ordinarily butterflies, numerically, are no 
considerable part of the insect fauna; when under extraordinary cir- 
cumstances they do become over-abundant they are more frequently 
devoured by birds. Thus Bryant found Brewer’s blackbird eating 
large numbers, and three other species of birds smaller numbers, of 
Eugonia californica during an unusual outbreak of the species. (Con- 
dor, vol. 13, pp. 195-208, Nov. 1911.) 

Summary of identifications of Lepidoptera: Total number 18,487; 
percentage of identifications among those of all insects, 9.6831 ; per- 
centage of species in this group among the whole number of insect 
species, 15.6180. 

Other enenues.—For the most part fishes are only casual devourers 
of Lepidoptera, getting chiefly larvae which fall into the water, most 
of which would perish anyway. However, gamy fishes such as trout 
snap up adults that incautiously fly near the surface of the water. 

Bullfrogs have been observed feeding freely on Papilio turnus 
adults (Mallonee, Science, 1916, pp. 386-387) and half a dozen leopard 
frogs have been noted as eating 500 Argynnis aphrodite in a week 
(Shiras, Nat. Geogr. Mag., 1921, p. 174). Kirkland found cutworms, 
tent and other caterpillars to compose 28 per cent of the total food of 
149 toad stomachs examined by him, and Munz found lepidopterous 
larvae in stomachs of four species of frogs. In 209 leopard frogs, 
Drake found one imago, one chrysalid, and 121 larvae of Lepidoptera. 
Surface reports remains of Lepidoptera in stomachs of eight species 
of salamanders, one toad, and nine frogs. In the Tropics lizards are 
said to be important enemies of adults of this order and our lizards 





- a 2 amen ieaeredeiedeenetie 
$$ c rere 


NO. 7 PROTECTIVE ADAPTATIONS—McATEE 61 


are known to eat both larvae and imagos. Surface records Lepidop- 
tera from the stomachs of five species each of snakes and turtles. 

Cutworms are commonly taken and other caterpillars and chrysa- 
lides are devoured to a smaller extent by moles. A number of small 
mammals, such as opossums, spermophiles, ground squirrels, tree 
squirrels, prairiedogs, grasshopper mice, skunks, raccoons, shrews, 
armadillos, the mongoose, and Nasua feed more or less regularly on 
caterpillars, and take an occasional pupa or imago. Bird has observed 
that field mice and skunks are effective enemies of the gall-making 
larvae of Papaipema. (Can. Ent., vol. 41, no. 2, pp. 67-68, Feb., 1909. ) 
Haskin has reported squirrels devouring large numbers of Aelitaea 
chalcedon adults (Ent. News, vol. 27, no. 8, p. 370, Oct., 1916), and 
Attwater found wings of several hundred Danais archippus that had 
been eaten by the Texas grasshopper mouse (Bull. Amer. Mus. Nat. 
Hist., vol. 6, p. r81, 1894). Bats catch moths, and monkeys also have 
been reported as eating butterflies commonly. (Trans. Ent. Soc. 
London, 1912, p. iv, XVii-Xviil.) 

The insect enemies of Lepidoptera also are numerous and some ot 
them are exceedingly destructive. Robber flies and dragonflies are 
frequently observed devouring adult Lepidoptera, and a Natal col- 
lector considers Mantidae the chief enemies of butterflies. (Proc. Ent. 
Soc. London, 1906, p. lii.) Spiders catch them directly or trap them 
in their webs, Phymatidae lie in wait for them, and predacious beetles 
sometimes capture them. However, the latter predators are more 
serious foes of caterpillars, in the pursuit of which they have as 
fellows numerous wasps. Ants, chrysopid larvae, and other insects 
and mites feed upon the eggs; and parasites often destroy large pro- 
portions of the eggs laid. Parasites of lepidopterous larvae also are 
legion, including numerous species of Hymenoptera and Diptera, and 
they take a large toll from every generation of the insects. Exceed- 
ingly high percentages of parasitism have frequently been observed, 
reaching locally in a few cases even to 75 and 100 per cent. It has 
been found in one case at least that no fewer than 63 species of 
hymenopterous parasites attack a single species of moth. (Cambridge 
Nat. Hist., vol. 5, p. 521, 1910.) 

Bacterial diseases frequently kill large numbers of caterpillars and 
sometimes locally extirpate certain species. 

Discussion.—It is one thing to record a proved fact, but quite 
another to assert that a certain thing does not occur in nature. Our 
stock of verified data stands as an imperishable record and addition 
to it, not subtraction, is the rule. Let none be tempted therefore to add 





62 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


to the vast body of speculation that has proceeded from slight premises 
by data in a preceding paragraph apparently indicating that birds do 
not eat many chrysalides of butterflies. For in that case we must refer 
him to Bryant’s statement that about 15 per cent of the pupae of 
Eugonia californica at a time when they were very abundant showed 
evidences of attack by birds (Condor, vol. 13, p. 200, Nov., 1g1r), 
and to Chittenden’s that “in one case it was found that during the 
winter the number of pupae of the cabbage butterflies was reduced 
more than go per cent by birds feeding upon them.” ( Farmers’ 
Bull. 766;,°Ur S: Dep. Ars p: ©; TOr6s) 

In this paper we cannot possibly discuss all of the data relating to 
predators upon insects and other animals, but the evidence we present 
in our tabulations surely goes far to prove that no groups are neglected 
by predators (except as availability or sheer size dictates) and that 
the various groups are preyed upon more or less in proportion to their 
numbers. As applied to Lepidoptera this rule is apparent in the 
greater number of records for such large families as the Noctuidae, 
Tineidae, and Tortricidae for instance as contrasted to such smaller 
ones as the Sphingidae, Arctiidae, and Bombycidae or of the more 
numerous Nymphalidae to the less numerous Papilionidae. Due to 
the high proportion of unidentified Lepidoptera, our tables are not as 
complete and informing as could be desired, but where there are ap- 
parent exceptions to the rule of proportional loss to predators, data 
from other sources usually indicates unreliability of the apparently 
negative evidence. For instance the records of Geometridae in our 
table seem too low for this rather important family which is un- 
doubtedly numerous in individuals. But that this is due solely to the 
make-up of our material is proved at once by reference to the litera- 
ture; no fewer than 73 species of nearctic birds have been observed 
feeding on cankerworms (Paleacrita and Alsophila) for instance. 
Wellhouse, who reports finding cankerworms in 98 of 100 stomachs 
of birds (36 species) collected near Lawrence, Kans., in 34 of which 
they composed the total food, says: “ Probably no insect is a favorite 
food of more species of birds than the cankerworm larva.” (Bull. 
Univ. Kansas, vol. 18, no. 1, p. 301, Oct. 1917.) In a study of birds 
in relation to cankerworms in Illinois, Forbes found these larvae to 
compose 45 per cent of the food of a collection of 55 birds (15 species) 
and that one species, the cedarbird, was destroying them at the rate 
of at least 90,000 per month. (Forbes, S. A., Trans. Illinois State 
Hort. Soc. (1881), pp. 123-130, 1882.) 





NO. 7 PROTECTIVE ADAPTATIONS—McATEE 63 





The snow-white linden moth (Ennomos subsignarius) has a typical 
twiglike caterpillar, but several entomologists have testified that it was 
practically exterminated in cities by the English sparrow. (See 
Merrick, G. W., Bull. 286, Cornell Univ. Agr. Exp. Sta., p. 62, 
Nov., 1910.) 

The only other important family of moths which our tabulations 
might indicate to be neglected by birds is the Pyralidae. With little 
doubt this condition is due either to the larvae not being recognized 
er to our stomach material not being fully representative. Certainly 
birds are known to be enemies of our pyralid larvae, a little search 
revealing records of avian predators upon Lowvostege similalis, L. 
sticticalis, Pilocrocis tripunctata, Pinipestis simmermanm, Diatraea 


-saccharalis, Acrobasis nebulella, and Pyrausta nubilalis. Five species 


are known to feed on the last-named, the corn rootborer, while of 
Loxostege sticticalis, the beet webworm, it is recorded that: “‘ Insect- 
eating birds devour the worms in large quantities. Where the worms 
were abundant [in Colorado] ... . blackbirds were attracted in 
flocks of thousands and in several instances . . . . the worms were all 
cleaned out of fields by them in the course of two or three days.”’ (Gil- 
lette, C. P., Bull. 98, Colorado Agr. Exp. Sta., p. 10, Mar., 1905.) 

These instances emphasize the universal scope of the predatory 
activities of birds; in general the enemies of economic species of 
insects are better known, and fully discounting the fact that they are 
most studied, this is only another way of saying that the most abun- 
dant species have the most numerous enemies. 


COLEOPTERA (BEETLES ) 


Protective adapiations —While more pages have been written about 
warning colors, mimicry and the like in Lepidoptera, which insects 
furnished the inspiration for this line of speculation, the important 
and extensive order of Coleoptera has been far from neglected and 
perhaps the most positive statements of all have been made regarding 
the “protected”? status of some of its members. In conclusions 
derived from G. A. K. Marshall’s data on ‘“ The Bionomics of South 
African Insects” (Trans. Ent. Soc. London, 1902, pp. 393-584), 
Prof. E. B. Poulton in discussing the chief specially defended 
Coleoptera mentions: “ The groups about which there seems to be no 
doubt at all—conspicuous, constantly refused by insect-eaters, and 
hable to be mimicked by other Coleoptera are the following: FEroty- 
lidae, Coccinellidae, Malacodermidae, including the Lycinae, Lam- 
pyrinae and Telephorinae, Melyridae, Cantharidae, Chrysomelidae, 


5 





64 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 85 


Endomychidae, and Pyrochroidae.”” The Cleridae are cited as a family 
that while undoubtedly distasteful, in forming color associations take 
the colors and patterns of other insects “rather than impress the 
’ The following four 
‘at any rate partially distasteful ” : Scarabae- 
idae, Cetoniidae, Tenebrionidae, and Lagriidae. The longicorns are 
thought to include a few distasteful species in addition to many that 


stamp of their own likeness on the assemblage.’ 
families are said to be ‘ 


mimic aculeate Hymenoptera and other specially defended insects. 
Cicindelidae are said by Wallace to be protected by cryptic coloration, 
the refuge of the weak, while Poulton and Shelford have recorded 
them as models mimicked by species less prepared for the struggle 
for existence—a tribute to the strong. 

“The Carabidae are a powerful specially defended group,” writes 
Poulton (op. cit., pp. 513-514) “and it is of advantage to be recognized 
as belonging to the group, even though it is no doubt of still greater 
advantage to be mistaken, as may happen at a distance, or on a super- 
ficial view, or during rapid movement, for the still more formidable 
Mutillidae and ants”’..... “ Dr. A. R. Wallace has always thought 
that the extreme hardness of the mimicked Curculionidae and An- 
thribidae is the character which protects them.” (Poulton, op. cit., 
PP. 522-523.) 

Comment of this kind could be cited indefinitely, for something or 
other has been claimed to be “ special protection ”’ for practically every 
group of beetles. It is undesirable and unnecessary to cite this matter 
in detail, but some attention should be given to the subject of repug- 
natorial secretions which has figured considerably in accounts of 
protective adaptations of beetles. For convenience, a summary of the 
occurrence of such secretions is quoted from a recent article on the 
topic: 

“It has been well understood that the presence of defensive or 
repugnatorial scent glands in certain insects exists in direct adaptation 
to the needs and habits of their owners and in close response to their 
environment ; also that such glands are of very frequent occurrence 
and with much variation as to position, form, and function; and that 
their presence is of value to the insect for repellent, defensive and 
warning purposes. .... Biologically speaking, the principle involved 
in such cases, though often modified, is practically identical with that 
of the mephitic, sulphuretted, oil-like fluid ejected by the skunks. Thus 
far anal glands are known to be present in the following families of 
Coleoptera: Cicindelidae, Carabidae, Dytiscidae, Gyrinidae, Staphy- 
linidae, Silphidae, and Tenebrionidae. The blood itself serves as a 





NO. 7 PROTECTIVE ADAPTATIONS—McATEE 65 


repellent fluid in the Meloidae, and in the Coccinellidae and Lam- 
pyridae, and it issues from a pore at the end of the femur as a yellowish 
fluid. The cantharidin in the blood of some species of Meloidae, 
commonly known as ‘ Spanish Fly,’ forms an especially caustic pro- 
tection against birds, predacious insects and reptiles.” (Wade, J. S., 
Notes on defensive scent glands of certain Coleoptera, Psyche, vol. 28, 
nos. 5-6, p. 146, Oct.-Dec., 1921.) 

Bird enemies.—It is worth pointing out that about 15 per cent of 
all the determinations of beetles were not carried to the family, and 
consequently that the percentages for the various families should be, 
on the average, about a seventh larger than shown in the tabulation. 


Identifications of Coleoptera 


Percentage 
of species 
in this family 
Percentage of aeomaued 
identifications nearctic 
Number of among all species of 
Family identifications ! Coleoptera Coleoptera 2 
Gicindelidae ......-....... 649 -7606 6146 
Carabidae Josie cs nds eeas 15,887 18.6200 11.6730 
| EATMPMIZOIdAG 23.5 0e0es208 jes ae Aono ens 
| Omophronidae ........... 16 (Olay) 0808 (15) 
| Ip Idae.* <akeswecweseas’ 363 4254 .2210 
ID tISCIMAC. ey cic cess 4 cowie es 1,729 2.0264 1.79054 
MV PAMIGAC een oaig waieie Sa eas 64 .0750 .0808 
Hydrophilidae ............ 2,418 2.8340 1.0244 
Platypsyllidae .......00.. eee Shi .0053 (1) 
Brathinidae’ .....0%. 6. << ae tnd .O16I (3) 
IBeptinidae o5...4 samesuecaes ee ree O16 (3) 
Silphidae ...........2000: 409 4794 .7386 
lambidae ..2.:...-...0.- wir ah: 0323 (6) 
Scydmaenidae ............ vee a .9381 (174) 
Orthoperidae ............. 4 .0047 .3073 (57) 
Stapnylinidae .../....e2.60% 1,605 1.8811 14.8163 
Pselaphidae .............. 3 .0035 1.9140 
Clavigeridae ...........0. aoe ae 0161 (3) 
IM OAC acc ecw haan tana ds ae on 4690 (87) 
Sphaeritidae ............. nae ae 0053 (1) 
Colydiidae ............... 5 .0059 .4520 
Murmidiidae ............. ee Ka 0269 (5) 
Monoedidae .............. ee oie 0053 (1) 





* There is an omission of 737 records of Carabidae and 574 of Chrysomelidae, 
enough to make more than 1.5 per cent of the total of beetle records. 

*Computed from Leng, C. W., Catalogue of the Coleoptera of America 
north of Mexico, 470 pp., 1920. 

*The number of nearctic species in the family. 


66 


SMITHSONIAN MISCELLANEOUS COLLECTIONS 


Identifications of Coleoptera—Continued 


Family 
athridudae hence ee oe 
Mycetaeidae 
See 
Bhalacridacmns-eeeicree cent 
Coccinellidae’ =. 22... eee ees 
Allecnlidaeteeor.t os eee 
Menebrionidaew mse eeaeceo 
IDSKepaRGRYS sono pone boceoer 
Mionomidaea- eee eee 
Melandryidae.-.)ae cn eee 
Ptinidae 
Anobiidae } 
Bostruchidaceesasaceeee cee 
ley ctidacmene ec eirraee 
Sphindidaemeass see seers 
Gisidae, sence oe a aaeienee 


Lucanidae 
ee 
Cerambycidae {eases aoe 
Chrrysomelidae) 22..-...02- 
Mylabridaey. scone cmisen ces 
Scaphididae 2 see aac e ae 
listeridae’ sana seenem eer ite 
Lycidae 

Lam.pyridae 

Bheieodiinan ere 
Telephoridae 
Melyridaeleensscce seers 
Cleridae i 
Corynetidae | 
@rthniidaewee ence cree 
levinexylidaemeatcse eee ee 
Telegeusidae ............. 
Micromalthidae .......... 
Cupedidae’ sac. ss cece. 
Cephaloidae. =. ss... ee ae 
@edemeridae ~22ssss- 02.6 
Mordellidaeweccae een 
Rhiphiphoridae ........... 
IMeloidaemsnermetiosane once 
Wevialitidae secmeise ecco 
Pythidaewarerscts nie cere 
Pyrochroidaec,cm.c tk ser ¢ 


Number of 
identifications 


16 


879 


38 


ONS: 


34 


279 
10 


Percentage of 
identifications 
among all 
Coleoptera 


.0187 
.OO12 


.0316 
1.7053 
.0398 
2.5749 
.0094 


0457 
£0508 


.0281 


.0047 
15.5317 
1418 


1.8577 
6.6407 
.O551 
0094 
1.2459 


1.0302 


0445 


0410 


.OOI2 
.0023 
.O105 
0398 
.0023 
.3270 
.O1I7 
.0070 
-0047 





VOL. 85 


Percentage 
of species 
in this family 
among 
described 
nearctic 
species of 
Coleoptera 


.5007 
.2048 


.6308 
1.9517 
6685 
6.1411 
0916 
.0323 (6) 
4367 


1.4550 


3288 
.0862 (16) 
-0323 (6) 
-4582 
5.3701 


1724 


6.0548 
5-2515 
5014 


2.0704 
1.5095 


1.7307 
1.1806 


.0269 (5) 
.0107 (2) 
0053 (1) 
.0053 (1) 
0215 
0431 
2641 
7656 
-1347 
1.2239 
O61 
.0916 
-0593 


NO. 7 PROTECTIVE ADAPTATIONS—McATEE 


Identifications of Coleoptera—Continued 


Family 


Pedilidae | 
Anthicidae 
Euglenidae | 
Cerophytidae 
Ceb.ionidae 
Plastoceridae 
Rhipiceridae 
Bilateridae .4......secea0 303 
IMielasidae: sic ..cnise sa dec 0s 
Throscidae ............... 
Buprestidae .............. 
Psephenidae 
PDEVOPIdaG ees fake oan 
Helmidae 
Heteroceridae ............ 
(GeOmysSIdae: cn. aaiewevete si oer 
Dascillidae ) 
Riedie ere 
Chelonariidae ............ 
Dermestidae ............. 
Biyiethidaes —scialeleiie esses. 
PROV SSOCICAG 6 ag a5 <i cewe cs 
MSiOmiddae..<aisaeces daecucs 
Nitidulidae ............... 
Rhizophagidae ............ 
Monotomidae ............. 
GiGi idaes ens sa- 2 eee 2 
HeGOLY IGA: civ..me,2 <6 « sores 
Derodontidae ............ 
Cryptophagidae ........... 
Mycetophagidae .......... 
Brentidae ...............- 
Platystomidae ...:....... 
Belidae 2... cc ccc ccc ee cess 
@urcilionidae s.... 8.0... 
Platypodidae ............. 
Scolytidae ............... 
Water beetles (further uni- 
dentified) .............. 
Rhynchophora (further 
unidentified) ........... 
Beetles (further unidenti- 
HEC) We tive fake arco See es yarts 
Beetle larvae (further uni- 
dentified) .............. 


Number of 


identifications 


NI 


143 
17 


190 
B12 


Percentage of 
identifications 
among all 
Coleoptera 


0716 


.0082 


5.2612 
.0047 
.0070 
7759 


.0387 


.1676 


67 


Percentage 
of species 
in this family 
among 
described 
nearctic 
species of 
Coleoptera 


T53L1 


1939 


3.1056 
-3073 
-1347 

2.0434 


.3072 


-0593 
.0107 (2) 


3288 


.0053 (1) 
.6955 
.5220 
.0215 (4) 
.3450 
LL, 
-0754 (14) 
.1941 
.4582 
3828 
.0269 (5) 
.7278 
1725 
.0323 
3342 
.0053 
9.9153 
0215 
2.0488 





68 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


The tiger beetles “are avoided on account of their ferocity ” 
(Bastin, Insects, their life-histories and habits, p. 151, 1913), and have 
been referred to as “ dreaded insects ” (Poulton, Colours of animals, 
p. 252, 1890), but what creatures capable of feeling dread so regard 
these beetles is unexplained ; certainly the facts indicate they are not 
birds. The 649 records included in the present tabulation are dis- 
tributed among 99 species of birds. Eight species have 10 or more 
records each, two others, the eastern meadowlark and eastern kingbird, 
over 20, the crow more than 60, and the crow blackbird 94. No fewer 
than 25 larvae of tiger beetles were found in a single stomach of an 
eastern bluebird, and 156 adults in that of a sparrow hawk and 164 in 
that of a long-billed curlew. If tiger beetles ever evade attacks by 
birds it is by celerity of motion rather than by any special defenses. 

With respect to Carabidae or ground beetles, Forbes in his report 
on the food of thrushes may have given some comfort to protective 
adaptation theorists when he said: ‘‘ We note, however, a remarkable 
deficiency of the highly colored genera—such as Galerita, Brachynus, 
Lebia, Platynus, Chlaenius, etc., which are either absent, or found but 
rarely in these birds’ (thrushes, bluebird) food. Evidently these more 
showy beetles are protected by some more effective means than ob- 
scurity of color.” (Forbes, S. A., Bull. Illinois State Lab. Nat. Hist., 
vol..t; no. 6, p. 57, May, 1883:)) 

However, this statement is but another instance of the danger of 
generalizing from insufficient data. In the study of the food of birds 
and other animals we are always adding to the list of species eaten 
and to the number of times they are taken ; the movement is never in 
the contrary direction. We are constantly finding enemies of forms 
previously held to be more or less exempt, and usually to an extent 


which more than compensates for previous lack of knowledge on the . 


subject. 

In the present instance such progress in knowledge since Forbes’ 
study is indicated by 535 records of the capture of Chlaenius by 
41 species of birds, 254 for Platynus by 55 species, 44 records for 
Galerita by 13 species, 39 for Lebia by 21 species, and eight for 
Brachynus by seven species ; figures more or less in harmony with the 
relative abundance in individuals of these groups. In this connection 
it may be well to note also F. H. Chittenden’s remark that Lebia 
grandis “is protected by its warning color from rapacious birds.” 
(Farmers’ Bull. 1020, U. S. Dep. Agr., p. 16, 1919.) Six of the 39 
Lebia records here cited are for grandis, and the writer submits that 
six records for this single species out of a total of 85,322 for all 





NO. 7 PROTECTIVE ADAPTATIONS—McATEE 69 


beetles (18,548 nearctic species) fully satisfies expectations based on 
the relative availability of the species to birds. 

Species of Agonoderus, much more common than Lebia but just as 
contrastingly colored, contribute 188 records to our tabulations and 
were eaten by no fewer than 57 species of birds. From ro to as many 
as 50 specimens had in several instances been taken at a meal. There 
are 57 records for Casnonia, a small genus of “long-necked” dis- 
tinctly “ warningly-colored ” beetles in stomachs of 14 species of birds. 

Even black, alone, the predominant color among Carabidae has been 
held to have a warning value, but Amara, Anisodactylus, Harpalus, 
and Pterostichus, chiefly typically black species, are eaten by the 
hundreds. There are 445 records for the powerful Pasimachus (80 
individuals in one crow stomach), and 497 for the species of Calosoma 
which are not only large, but some of which have contrasting blue 
margins, others fiery spots, and all powerful, ill-scented excretions. 
In fact, it is everywhere evident that the special defenses alleged for 
the Carabidae are more in the nature of pleasing fictions for theorists 
to speculate upon than practical reliances for the beetles concerned. 
Eloquent is the fact that between a sixth and a fifth of all determina- 
tions of beetles in the stomachs of nearctic birds are of Carabidae. 

The Haliplidae, all of which have “ warning colors,’ and the 
Dytiscidae and Gyrinidae, said to be protected by anal glands, all seem 
to be preyed upon in proportion to their abundance. The Silphidae 
quite generally have nauseous excretions and include numerous species 
with distinct warning colors, but it is the latter forms such as Necro- 
phorus with 102 records and Silpha with 213 that most evidently are 
eaten in due proportion. The apparent falling of records of this 
family below the index of frequency must be attributed to the smaller 
and rarer species with more concealed habits being overlooked, rather 
than to the larger familiar ones enjoying immunity on account of 
alleged special defenses which they possess in the highest degree. 

That the Staphylinidae is the family most numerous in species, and 
probably therefore of individuals, among all Coleoptera is a fact not 
realized by the average collector. It has been brought out only by the 
accumulated research of generations of coleopterists, and its lack of 
obviousness must be attributed to the secretive habits of so many of 
these small or even minute beetles. Most of them spend their lives 
chiefly under cover of various kinds, for example, in fungi, in leaf- 
mold, under bark, in old logs, and in ant nests, and it must be on this 
account that the records of birds capturing them are not very much 
more numerous, rather than that they are disliked. In fact the 1,605 

- determinations for them proves they are not disliked, and these 


70 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


records are shared by more than 160 species of birds. Fifteen of these 
kinds of birds had more than to records each, nine others more 
than 20, six others from 30 to 60 records, one other, the chimney 
swift, 76, the crow 190, and another, the starling, more than 200. 
In several instances the number of specimens found in a stomach was 
as many as from 20 to 50 and larger numbers were 85 for the bald- 
pate and 150 for the dowitcher. Such data certainly do not indicate 
distaste for Staphylinidae, hence the failure of the total number of 
captures to come up to theoretical expectations must be due to some 
other factor, presumably the small size and concealed habit of living 
characteristic of so large a proportion of the beetles of this family. 

The same causes also serve to explain why a number of the minor 
families of beetles have not yet been identified in bird stomachs ; the 
Platypsyllidae, and Leptinidae are parasitic upon mammals, the Scyd- 
maenidae, and Clavigeridae mostly live in ant nests, the Ptiliidae are 
minute, while the others most of which have five or fewer species in 
our region owe their degree of immunity to their very rarity. 

Passing now to one of the larger groups of beetles about whose 
protected status “there seems to be no doubt,” namely the malaco- 
derms, variously regarded as forming from one to four families, we 
find that they are devoured in no mincing way by nearctic birds. 
While various authors refer to these beetles (generally known as 
Lampyridae in the United States) in terms varying from distasteful 
to inedible or immune, our records show 879 determinations of them 
from bird stomachs. All of the groups were preyed upon, the Lycinae 
and Phengodinae least, however, because they are scantily represented 
in our fauna. The adult lampyrids identified were eaten by no fewer 
than 108 species of birds and the larvae by 25. Larvae in number up 
to 50 were found in a bluebird’s stomach, and in three instances as 
many as 100 were taken from a single stomach of the robin. Our 
most common lampyrids are Chauliognathus and Telephorus. The 
former genus was identified 179 times in the stomachs of 34 species 
of birds. Three of these had from 30 to 38 records each and the 
number of individual beetles eaten ran as high as 30 in a single in- 
stance. Telephorus (Cantharis) were determined 274 times in the 
stomachs of 35 species of birds; the number of imagines in a stomach 
ran as high as 16 and of larvae, 100. If the Lampyridae fail in any 
degree to attain proportional representation among the food items 
taken by nearctic birds it is due to the nocturnal habits of a large 
number of the species. The diurnal species seem to be captured as 
frequently as would be expected. 








| 





NO. 7 PROTECTIVE ADAPTATIONS—McATEE Fu 


The Melyridae (Malachiidae) are poorly represented compared to 
the Lampyridae, yet upon inspection of the records it does not seem 
that they are really avoided. Six genera and at least 10 species of 
these beetles were identified ; 21 species of birds had eaten them, and 
for one of these birds, Say’s phoebe, there were eight records of 
feeding on Collops. Identifications of the Cleridae again include 
numerous (21) species distributed among an equal number of species 
of birds. One of these birds, the red-eyed vireo, had eight of the 
records. In our experience Cleridae occur chiefly scattered and in 
small numbers, a type of distribution with which the records of birds 
preying upon them seem to harmonize. 

Of the Histeridae, Donisthorpe says: “All the species of this 
family are protected by their oval shape and hardness. They also 
‘feign death.’ ” . the “species which are spotted with red, are 
probably protected by their resemblance to Coccinellidae.” (Trans. 
Ent. Soc. London, 1901, p. 354.) The prevailing color in this family, 
1. e. black, has also been said to have a warning significance. Our 
records show 1,063 identifications of Histeridae representing 116 
species of birds; they are very freely eaten by some of these birds, 
the number of records per species exceeding 20 in the case of at least 
12 kinds, and the number of specimens eaten at a meal running up 
to as high as 200 as a maximum. 

The family of blister beetles ( Mylabridae, Cantharidae, or Meloidae 
as it is variously known) is especially noted for the presence in the 
bodies of its members of a vesicant poison, cantharidin, of which as 
small a quantity as one grain has proved a fatal dose for a human 
being. Bastin says of them “the blood contains cantharidin, an 
extremely caustic substance, which is an almost perfect protection 
against birds, reptiles, and predacious insects.” (Insects, their life- 
histories and habits, p. 167, 1913.) While these beetles are supposed 
to enjoy a very high degree of protection from natural enemies, 47 
species of birds included in the tabulations here discussed had fed 
upon them. Seven of the species had 10 or more records apiece of 
preying upon blister beetles, the eastern kingbird having no fewer 
than 77. In some cases from 12 to 16 specimens of cantharids were 
found in single stomachs and a maximum of 31 in the case of a 
magpie; more than 30 species in all of these beetles were identified. 

Pyrochroidae are said to be another specially defended group of the 
first order, but in view of the fact that there are only 11 nearctic 
species of the family and they usually rare, we believe that the four 
records of our birds capturing them are as many as could be expected. 
One of the birds eating Pyrochroidae, namely a hairy woodpecker, 





Jz SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


must have had unusual luck in order to obtain the 12 specimens it 
contained. 

Donisthorpe says “ The Elaters ‘ feign death’ and their ability to 
‘skip’... . is no doubt of great use to them. Mr. Holland points 
out that many of them possess a colour and shape suggesting the ap- 
pearance of bits of dry brown stick.” (Trans. Ent. Soc. London, 1901, 
p. 360.) Over four thousand (4,489) records of these beetles being 
eaten by nearctic birds show that the protective devices mentioned are 
of no particular account. There would appear to be no doubt whatever 
that birds feed upon Elateridae whenever available to them. 

The larvae of Buprestidae live in wood, and the adults have hard 
chitin and metallic or other brilliant coloration, but since there are 
more than 650 records of their occurrence in the stomach contents of 
nearctic birds, it is certain that concealment of the larvae rather than 
color protection is their main defense. Heteroceridae or mud beetles 
certainly seem well concealed to the human eye but the records indi- 
cate they are taken fully in proportion to their abundance. Dermes- 
tidae, said to be protected because they are carrion-feeders, are taken 
freely considering their availability in nature. Byrrhidae are thought 
to be excellent examples of cryptically defended insects. “ The legs 
and antennae are packed close to the body, fitting into cavities for 
their reception and the beetles then represent rabbits’ dung, or little 
lumps of earth; they in no way suggest the appearance of living 
beetles.” (Donisthorpe, Trans. Ent. Soc. London, 1901, p. 357.) 
However 312 records for them show American birds are not especially 
deceived by the alleged protective devices. 

It is unnecessary to comment on every family, but coming to the 
Erotylidae we have a group which though small in numbers is said 
to be one of the most highly protected groups. However, in the United 
States, insects of this family in general do not have the bold habits 
supposed to be associated with warning colors; in fact most of them 
feed concealed in fleshy fungi. Correspondingly most of the determi- 
nations of beetles of this family are for the species which live exposed 
as Languria, for which there are 10 records, probably all that should 
be expected for a single small genus. Similarly the Endomychidae 
are protected by feeding inside of fungi or on fungi growing on the 
under side of logs rather than by their “ warning colors.” It should 
puzzle selectionists to explain why these and other brightly colored, 
supposedly distasteful insects have such retiring habits that their 
“ warning coloration ” is seldom displayed. 

Contrasting these elusive beetles with another brightly colored but 
decidedly not secretive group they are supposed to mimic, the Coc- 








NO. 7 PROTECTIVE ADAPTATIONS—McATEE 73 


cinellidae, it is easy to see what factor makes for greater depredations 
by birds ; it is none other than the frequently mentioned “ availability.” 
Endomychids and Erotylids are red and black or yellow and black 
beetles, less abundant and much more retiring in habits; while coc- 
cinellids with the same colors are more common and live unconcealed. 
The former are relatively seldom captured, the latter are freely eaten. 
No better example of the influence of availability in guiding choice of 
food by birds could be desired. This despite universal acclaim of 
coccinellids as specially defended insects. “All the lady-birds are very 
gaily colored” says Donisthorpe. “They boldly walk about with- 
out any attempt at concealment, as do also their larvae. Both their 
larvae and pupae are very brightly spotted. The distastefulness of the 
perfect insects was proved* by Jenner Weir, and has since been con- 
firmed by both Poulton and Wallace.” (Trans. Ent. Soc. London, 
1901, p. 354.) 

Packard states that “ The Coccinellidae are . . . . protected by a 
yellow mucilaginous disagreeable fluid oozing out of the sides of the 
thorax,” (Journ. N. Y. Ent. Soc., vol. 3, p. 116, 1895), and Wallace 
says: “ The Coccinellidae or lady-birds are another uneatable group.” 
(Darwinism, p. 234, 1896.) Let us see. The total number of records 
of coccinellids in the food of nearctic birds is 1,455 and these are 
shared by 127 species. Twenty-seven kinds of birds had to or more 
records each, nine of which ran over 50, and three over 100. Not only 
is the effect of availability noted in birds eating more coccinellids than 
other similar but less abundant and conspicuous beetles, but its 
influence is evident in at least two other ways, namely that leaf-feeding 
birds, as warblers and vireos, get the most ladybird beetles, and that 
in California where coccinellids are notably more abundant than they 
are in the eastern States, a larger number of birds feed upon them and 
they get a great many more of the beetles. The largest numbers of 
coccinellids found in individual stomachs were 12 and 18 taken by 
English sparrows, 13 by the summer warbler, 14 by the warbling vireo. 
and 15 by the valley quail. 

We now come to the consideration of three families (the Scara- 
baeidae and Cetoniidae being reckoned as one) which Poulton says are 
“at any rate partially distasteful.” Regarding one of these fami- 
lies, the Lagriidae, which has only 17 species in the nearctic fauna, it 


* The “ proof” was experimental, of course; for the value of this proof see 
my I9I2 paper. Also note that Meisner’s results on the poisonous effect of 
Coccinellid juices (Ent. Bl. Nurnberg, vol. 5, no. 9, pp. 180-182, Sept. 20, 1909) 
are controverted by a repetition of his experiments by Heikertinger (Wien. Ent. 
Zeit., vol. 38, Heft 4-8, pp. 109-113, June 15, 1921). 





74 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


may be said that almost any small number of captures by our birds 
would satisfy expectations. There are eight records distributed among 
six species of birds, about all that probabilities demand. As to the 
Tenebrionidae so many species of which have secretions nauseous to 
man, the nearly 2,200 records are eloquent of the fact that these 
beetles are not disliked by birds. If they do enjoy any degree of 
immunity from bird attacks, it is probably on account of their char- 


acteristic nocturnal or otherwise seclusive habits. The number of — 


species of birds known to prey upon Tenebrionids is in excess of 175; 
nine birds had over 20 records each, five others more than 40, one 
additional over 50, and two others more than 100. The number of 
specimens taken at a meal ran up to 44 in two cases and to 46 in 
another and 53 in still another. The number of species of Teneb- 
rionidae identified was over 100, including 12 of Eleodes, the largest 
and most potently odoriferous of the family. 

Of these a recent article says: “ It was interesting to note that the 
quantity of the secretion voided varies noticeably with the different 
species under observation, both under field and under laboratory con- 
ditions, and some of the species, notably Eleodes tricostata Say, 
undoubtedly have the habit of erecting the abdomen in a threatening 
manner when approached, though no secretion may be voided. Such 
species undoubtedly find protection through imitation of the threaten- 
ing movements of their more formidable associates. Two of the 
characteristics of the Eleodes are their slowness of movement, and 
their habit of coming out of their hiding places about sunset for 
feeding purposes, and their presence is readily noticed on the bare 
sandy plains by birds, skunks, and other enemies, hence their protec- 
tive secretion, or, in the absence of this, their threatening maneuvers 
are no doubt of highest value to them.” (Wade, J. S., Notes on 
defensive scent glands of certain Coleoptera, Psyche, vol. 28, nos. 5-6, 
p. 148, Oct-Dec, 1921.) 

In this connection it may be said that our tabulations show 51 
records of birds feeding on Eleodes tricostata and tog for the other 
species. Other large Tenebrionids as Asida and Nyctobates are well 
represented in the table of determinations as are also the metallic 
torms such as Helops, Meracantha, and [:pitragus. Blapstinus with 
286 records for 11 species is the favorite genus, and the reason is 
what ?—simply that it is the most widely distributed and the most 
numerous in individuals. 

With respect to the other “‘ partially distasteful” family of beetles, 
the Scarabaeidae (sens. lat.), the more than 13,000 records in our 
tabulations speak for themselves. The selectionist protectionists have 


NO. 7 PROTECTIVE ADAPTATIONS—McATEE 75 


simply made a very bad guess. Consider for instance the Cetoniini, 
the best ‘“‘ protected’ tribe, some of which are said to mimic bees in 
flight. Our most numerously represented genus, Euphoria, has 445 
records, of which 148 are for the most beelike species of all, E. inda. 
Cotinis, very poorly represented in our fauna, has 156 records, and 
Cremastochilus, noted for their association with ants, 77. 

Species of Onthophagus “live in and about dung and are of a 
colour which conceals them well in those surroundings.” (Donis- 
thorpe, Trans. Ent. Soc. London, 1901, p. 358.) However, they were 
preyed upon 642 times by the birds included in the present tabulations. 
The species of Aphodius also are dung-feeders and said to be protected 
The number of determinations of this genus is approximately 3,565 ; 
in numerous cases 100 of these beetles were found in single stomachs 
and in one instance no fewer than goo. A warningly colored species, 
A. fimetarius (with the thorax black and elytra red), was identified 
in 913 stomachs. Consider the entirely different case of a beetle, the 
rose chafer (Macrodactylus), known to be actually poisonous (see 
Science, n. s., vol. 43, pp. 138-139, Jan. 28, 1916) besides having 
protective (cryptic) coloration and long spiny legs: although there 
are but two species in the country, we have 52 records in our tabula- 
tions representing 15 species of birds. The larger numbers of speci- 
mens taken were: nine by a crow, 12 by a road-runner, and from 
15 to 40 in five instances by the eastern kingbird. These records show 
that the most potent protective adaptations possible do not necessarily 
protect. The highly significant fact about the case is that predators 
do not seem to recognize the dangerous qualities of the rose chafer ; 
every generation of young chicks and pheasants will pay a heavy 
death toll if permitted to stuff themselves with these beetles. Even 
trout kill themselves in the same way. But what advantage is all this 
to the beetle? Those that cause the death of some predators, them- 
selves lose their lives, that is, all of those actually proved “ fit” in 
this respect are eliminated ; the only effective poisonous action is upon 
young (among birds)—adults can and do eat them freely. No con- 
siderable body of predators has ever been killed, and ‘“ warning color ” 
has not been acquired (the rose chafer is a uniform and inconspicuous 
brownish-yellow). Theories as to protective adaptations seem to suffer 
from every angle of this case. (For fuller discussion of the subject 
see Lamson, Geo. H., Journ. Econ. Ent., vol. 8, no. 6, pp. 547-548, 
Dec., 1915; Bates, J. M., Science, n. s., vol. 43, pp. 209-210, Feb. 11, 
1916; and McAtee, W. L., The Auk, vol. 33, no. 2, pp. 205-206, 
Apr., 1916.) 





¥ 


76 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


Some curiosity may be felt as to the relations of birds to the large | 
Scarabaeidae and Lucanidae with thoracic horns and especially strong 
or greatly enlarged mandibles. In nearctic America we have few | 
species in any of these groups; another limiting factor important in 
relation to bird predators is the large size of these beetles. Never- | 
theless all of the genera are represented in the food of birds, Passalus | 
by 32 records, Platycercus by 19, Lucanus by 25, Ceruchus by three, | 
Dorcus by six, Sinodendron by seven, and lucanids further unidentified 
by 28 determinations ; our largest scarabaeid, Dynastes is represented | 
by five identifications, Strategus by 27, Xyloryctes by seven, Copris | 
by 62, and Phanaeus by 252. The latter genus, besides being | 
“horned,” has brilliant metallic colors. | 

The long-horned beetles or Cerambycidae include many species | 
with showy colors, but selectionists as a rule have not attributed dis- | 
tasteful qualities to the group; rather they have considered these | 
beetles mimics of various more strongly “ protected” insects such as | 
wasps and other Hymenoptera and weevils. Numerous longicorns | 
have cryptic coloration also, but their chief defense must be residence 
of the long larval stages in wood where they can be reached only by 
a small proportion of insect predators. The imago state, only in which 
the colors theorized about are displayed, is of relatively short duration. 
Bearing these facts in mind we believe the records show that longi- 
corns are fed upon to such an extent as to indicate that in proportion 
to availability they contribute their due share to the subsistence of 
birds. 

The total number of determinations in the present tabulations is 
1,585, shared by 162 species of birds. Twenty-one kinds of birds have 
from 10 to 19 records each; six additional species from 20 to 29; six 
others from 30 to 39; one other 42; still another 53; and two as many 
as 169 and 173 respectively. The woodpeckers, on account of their 
peculiar qualifications for obtaining the larvae, naturally are the chief 
enemies of Cerambycidae. Several of the species prey upon these — 
beetles to the extent of from ro to 50 per cent of their total food. The 
number of adult beetles taken at a meal by these or other birds ex- 
ceeded 30 in a number of cases and in four ran as high as 83, 100, 
102, and 168. The last named figure is for one of our most wasplike 
species, Xylotrechus colonus, in the stomach of a raven. There are 
10 identifications of Nylotrechus; of the other wasp-colored long- 
horns, we have the following numbers of determinations: Cyllene, 10 
(28 specimens of C. robiniae in the stomach of a magpie) ; Calloides 
nobilis 1, Neoclytus 11, Clytanthus 12, Clytes 5, Strangalia 6, Typo- 
cerus 16, and Leptura 39. It is noticeable that the numbers appended 


bad 


ee — eeeeeeeeeeeeeeeEOEOEOEeeEeEeEeEeEEeEEeEEeeEeEEE—EeEeEeEeEeEeeEEEE_E_E_E_______E EO _—_—_—_V—_—_—_ 


eo = - —_ A A A A LLL LLL LLL LLL LLL RL LLL LD | 
pa ee eee — 


NO. 7 PROTECTIVE ADAPTATIONS—McATEE 77 


correspond very closely to the relative abundance in individuals of 
these genera. Of the ant-suggesting genera, we have for Euderces 
five records and for Cyrtophorus one; and for the ichneumonid- 
mimic Neolorchus three identifications. 

The figures for the distinctly warningly colored genera are 
Acmaeops seven, Desmocerus two, Gaurotes two, Tetraopes nine, and 
Oberea five. Such a catalogue shows that all the forms, whatever their 
alleged “‘ protection” are eaten more or less, and there is no other 
explanation of the comparative extents to which they are taken so 
reasonable and satisfying as that it probably depends almost entirely 
on their relative abundance and availability to birds. 

The Chrysomelidae or leaf beetles are classed by Professor Poulton 
as undoubtedly specially protected, and Donisthorpe writing of them 
under another name says: “‘ The Phytophaga are considered to be all 
more or less distasteful, and no doubt justly so. Many species have 
been proved to be so, and the group is mimicked by various orders of 
beetles throughout the world.” (Trans. Ent. Soc. London, Igor, 
p. 367.) Selectionists should have been somewhat restrained in their 
theorizing by the very name Phytophaga, for the leaf beetles and their 
allies being groups that subsist directly upon vegetation, must ac- 
cording to inevitable law in the organic world form the base of a 
column of predacious life more or less exclusively dependent upon 
them. Like the grazing mammals, all plant-feeding insects, no doubt, 
have their lions, wolves, and eagles, their hyaenas, jackals, and 
vultures. 

No reason appears from the records of bird food here discussed to 
warrant doubt that the leaf beetles do in fact contribute their full 
quota toward the subsistence of predatory animals. The total number 
of identifications of Chrysomelidae is 5,666, and these are shared by 
well over 200 species of birds, so it is certain that practically all of our 
birds feed more or less upon these beetles. More than 230 species of 
Chrysomelidae are represented in the determinations, this in turn 
indicating that all tribes of the family are preyed upon. The Cassidini, 
on account of their bright colors and specialized larvae, receive fre- 
quent mention as a specially protected group but our scant representa- 
tion of this tribe seems to bear its share of bird predation; Cassida 
3 records, Physonota 2, Coptocycla 48, and Chelymorpha 48. Again 
correspondence of the number of identifications with observed fre- 
quency of the insects is quite-evident. 

Resemblance to caterpillar droppings always is spoken of by selec- 
tionists as a prime defense, and one tribe of our leaf beetles, the 
Chlamydini, exhibits this to a high degree. When feigning death, as 


78 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


they do when disturbed, they “resemble the excrement of certain | 
caterpillars so closely as to render their eee dificult . . . . and | 
it is said that birds will not pick them up.” (Blatchley, Coleoptera 
of Indiana, p. 1,114, 1910.) Two genera represent this tribe in our | 
fauna and of these, Chlamys has been found 56 times in bird stomachs | 
and Exema 17 times; 10 of the latter beetles were contained ina | 
single stomach of a Bewick’s wren. The number of determinations | 
| 
! 
| 
| 
| 
| 
| 
| 
| 
| 
| 





cited, in view of the few species we have of this tribe, fully satis 
the probabilities. 

The genus Diabrotica, chiefly yellow and black species, has received 
special attention from the standpoint of protective adaptations. ‘“] 
believe,’ says C. J. Gahan, “that the species of Diabrotica are pro- 
tected, and that the species of Lema derive advantage by mimicking 
them.” (Trans. Ent. Soc. London, 1891, p. 369.) The tabulations 
of bird food here discussed show 41 records of Diabrotica vittata 
distributed among 17 species of birds; 107 of D. 12-punctata for 42 
species of birds (18 specimens being found in a stomach of a cliff 
swallow) ; and 194 records of D. soror for 22 kinds of birds (a 
black-headed grosbeak had eaten 21 of these beetles). There are also 
34 other records for scattering and unidentified species of the genus. 
Thus there is no evidence of special protection for Diabrotica; as for 
Lema the species are much less numerous in individuals, and that is 
the real reason they are captured less frequently by birds; we have | 
22 identifications shared by 14 species of birds. 

One other Chrysomelid, the Colorado potato beetle (Leptinotarsa 
z0-lineata) , has had its protective adaptations pointed out on numerous 
occasions, and like the rose chafer, among the Scarabaeidae, seems to 
be actually poisonous. (See Riley, Seventh Missouri Rep., 1875, 
pp. 6-7.) However, our records show that 23 species of birds devour 
the insect and r1 others are added by the literature of the subject. 
One hundred and eighteen identifications of this pest are included in 
our tabulations; the larger number of specimens found in single 
stomachs are 8 in that of a starling, 10 in a sharp-tailed grouse, 
12 in a black-headed grosbeak, and 14 in a rose-breasted grosbeak. 
Birds such as the bob-white, crow, and rose-breasted grosbeak are 
recorded as having cleared fields of these pests. 

Before leaving the Phytophaga or Chrysomelidae it may be well to 
cite certain records of large numbers of individuals being taken at a 
meal by birds, since they show not only-that there is no restriction of 
bird attack to certain tribes of the family but also that there is no 
restriction of the more important avian predators to certain groups 
of birds. Some of the larger records are: 36 specimens of Micro- 


NO. 7 PROTECTIVE ADAPTATIONS—McATEE 79 


rhopala vittata taken by a starling ; 47 Donacia by a red-winged black- 
bird; 50 of Systena sp. by a Baird’s sparrow; 50 of Disonycha 
caroliniana by a horned lark; 58 of Myochrous denticollis by a house 
wren; 212 Donacia subtilis by a Franklin’s gull; 250 Colaspis 
brunnea by a nighthawk ; and about 300 FE pitrix cucumeris by each of 
five individual tree swallows. 

Bruchidae spend so much of their lives within seeds that they are 
little exposed to attack by birds; an advantage which probably is 
compensated for by their being devoured with the seeds by some birds 
and other seed-eating animals. However, this is a subject that has 
scarcely been investigated. Our 47 records represent nine or more 
species of bruchids and were distributed among 29 species of birds. 
Expectations based on availability of free bruchids probably are 
satisfied. 

The great series or suborder of beetles known as the weevils or 
Rhynchophora, for the most part, are said to be cryptically colored, 
resembling seeds, buds, bark, bits of earth, bird droppings, etc. 
Wallace adds: ‘One of the characters by which some beetles are 
protected is excessive hardness of the elytra and integuments. Several 
genera of weevils (Curculionidae) are thus saved from attack and 
these are often mimicked by species of softer and more eatable 
groups.” (Darwinism, p. 260, 1896.) However, it should be pointed 
out at once that those who predicate hardness as a defense against 
predators do so without due reflection upon the digestive powers of 
animals. 

Recall the fragmentation and gulping down of bones by dogs; the 
swallowing of snails, shells and all, by squirrels; while reptiles, am- 
phibians, birds of prey, and predatory mammals either swallow their 
vertebrate prey whole or in large pieces, the bones included ; waterfowl 
and shorebirds habitually take shellfish entire, including such hard- 
shelled forms as clams and oysters; gallinaceous birds are provided 
with gizzards which grind up the hardest seeds; and finches and 
numerous other birds are just as effectively equipped if on a smaller 
scale. Not only do birds with gizzards grind up their food materials, 
but the grit and pebbles they swallow are in most cases gradually 
ground down and pass out through the intestines in the form of fine 
sand. Most predators, in fact, have either a powerful mechanical or 
a resistless chemical digestion that as a rule is fully competent to dis- 
pose of anything entrusted to it. With such digestive powers at the 
service of predators, it is extremely unlikely that hardness in the 
degree possessed by weevils is any bar to their being eaten ; moreover 
being jointed, weevils are thoroughly susceptible to chemical digestion. 

6 


8o SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 85 


In illustration of the factor of hardness as related to bird food the 
three genera Lixus, Thecesternus, and Sphenophorus, representing 
as many families of weevils, may be discussed. Lixus is so hard that 
the entomologist usually finds drilling a necessary preliminary to 
pinning ; moreover the species are full of vitality, sometimes living 
through 24 to 36 hours in the cyanide bottle. Records of this genus 
in our tabulations total 102, distributed among 27 species of birds. 
No fewer than 18 specimens were found in a single stomach of a 
nighthawk. Thecesternus, a weevil with unusually thick and hard 
integument, was identified 151 times in the stomachs of 22 species of 
birds. Twelve specimens were taken by a meadowlark at a meal and 
15 by arobin. The billbugs of the genus Sphenophorus not only are 
hard, but like Lixus have much ability to resist the fumes of cyanide 
and prolonged submersion in water. However 1,397 determinations 
were made representing 34 species of these weevils. They were found 
in the stomachs of no fewer than 11o species of birds. Some of the 
larger numbers taken from single stomachs were: 10 in the cases of 
the upland plover, clapper rail, and yellow-headed blackbird; 11 in a 
robin ; 12 in an avocet ; 17 in a crow blackbird ; 20 in a killdeer ; 33 in 
a crow; and 34 in a magpie. 

Hardness thus appears to be of no consequence as a defense. Brief 
attention may be paid to a few other of the so-called protective devices 
of weevils. One of the obscurely colored genera, with the habit of 
dropping to the ground and feigning death, is Rhinoncus ; such weevils 
are said to resemble seeds, but what good this would do, since most 
birds eat seeds, theorists have left unexplained. Rhinoncus has been 
identified 73 times from the stomachs of 30 species of birds, of one 
of which, the olive-backed thrush, an individual had eaten 20 of these 
weevils. Rhodobaenus, our only conspicuous red and black weevil, 
was identified 14 times in-the stomachs of Io species of birds, and 
Tyloderma, black weevils with whitish or yellowish markings, 133 
times in 48 species of birds. Fifteen specimens of Tyloderma were 
taken from the stomach of a meadowlark. 

To mention the relations of birds to a few representative genera of 
weevils, we may record that the rare Otidocephalus were identified 
six times in the stomachs of an equal number of species of birds ; that 
the minute Apion were taken 91 times by 36 species; the nut weevils 
(Balaninus) 380 times by 85; the cotton-boll weevil (Anthonomus 
grandis) 348 times by 43 species of birds (23 other species are re- 
corded as enemies in the literature); the alfalfa weevil (Hypera 
murinus) 2,222 times by 50; the clover root weevils (Sitona) 1,611 





NO. 7 PROTECTIVE ADAPTATIONS—McATEE 81 


times by 94; the engraver beetles (ps [Tomicus|) 120 times by 24; 
and the Anthribidae 29 times by 21 species. 

A few of the larger numbers of weevils found in single stomachs 
also may be cited; thus 109 Dorytomus mucidus were found in one 
stomach of a downy woodpecker; 153 Calandra oryzae in a barn 
swallow ; 167 Barypithes pellucidalis in a starling ; 282 Hyperodes sp. 
in an eared grebe; and 281 larval and adult alfalfa weevils in a 
Brewer’s blackbird, 300 in a killdeer, and 317 in a valley quail. 

The nearly 20,000 identifications of weevils in birds’ stomachs attest 
to the frequency of their capture, and records such as those just cited 
to the relish with which they are eaten. 

Though to all weevils are attributed various protective adaptations, 
weevils of all sorts are preyed upon; the secret of the whole relation- 
ship between prey and predator in this as in other cases is distribution 
of the attack. All available food supplies are sought by predators and 
the amount of attention they receive is in direct proportion to their 
availability. 

Total number of identifications, of Coleoptera 85,322; percentage 
of identifications among those of all insects, 44.6899; percentage of 
species in this order among those of all insect species known, 46.2032. 

Other enenues.—lIt is difficult to summarize what is known regard- 
ing the predatory foes of so extensive an order as the Coleoptera. 
Fresh-water fishes prey systematically upon both larvae and adults 
of the aquatic beetles but secure other forms only incidentally. How- 
ever it appears that falling into the water or otherwise becoming 
available as prey for fishes is a more or less frequent happening to 
terrestrial beetles, since most of the families are represented in the 
food of these animals. (See especially Forbes, papers, bibliography 
p. 188.) 

Kirkland reports Coleoptera as making up the following percentages 
of the food of 149 common toads: ground beetles and their allies, 8 per 
cent ; May beetles and allies, 6 per cent ; wireworms and allies, 5 per 
cent ; weevils, 5 per cent; potato beetles and allies, 1 per cent ; carrion 
beetles, 1 per cent ; and miscellaneous beetles, 1 per cent. Drake found 
Coleoptera to constitute 33 per cent of the whole number of animals 
consumed by 209 leopard frogs and 54 per cent of the insects. The 
number of specimens of various families identified was: Carabidae 
176, Cicindelidae 44, Hydrophilidae 1, Staphylinidae 12, Coccinellidae 
13, Erotylidae 1, Elateridae 1, Spondylidae 2, Cerambycidae 4, Chryso- 
melidae 2, Tenebrionidae 1, and Rhynchophora 146. It is worth noting 
that this author says of weevils: ‘‘ The habit of these insects of drop- 
ping to the ground when disturbed gives the frog a chance to capture 


82 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


them.” This is just the habit the selectionists have declared protective. 
Klugh reports finding 35 Colorado potato beetles and five other 
Coleoptera in 25 stomachs of the leopard frog. Surface’s report shows 
that other frogs and toads feed extensively upon beetles, the larger 
families contributing most heavily ; the salamanders also eat a great 
many beetles, especially aquatic forms. Lizards, snakes, and turtles 
also feed upon beetles, some of the smaller terrestrial snakes taking a 
great many of them. Pack reports lizards feeding on beetles of such 
“protected” groups as Coccinellidae, Erotylidae, Meloidae, and 
Chrysomelidae. 

Among mammals, moles and shrews prey freely upon beetles, 
taking Scarabaeidae and their larvae more and wireworms and ground 
beetles less often. Spermophiles, prairiedogs, chipmunks, squirrels, 
grasshopper mice, and other rodents as well as raccoons, foxes, and 
coyotes prey upon beetles occasionally, and such animals as bats, 
skunks, and armadillos depend upon them to a much larger extent. 
There is no reason to believe that the ‘“‘ protected” groups of beetles 
fare any better with mammalian than with avian predators. However 
citation of a few instances of the capture of such beetles may be 
advisable. A series of three armadillo (Tatu novemcinctum) stomachs 
from Texas contained Carabidae and Scarabaeidae in profusion, also 
weevils, Histeridae, Lampyridae, Staphylinidae and Tenebrionidae 
(including Eleodes). The stomach of a skunk (Mephitis occidentalis) 
collected at Nelson, Calif., held 60 per cent of pupae of the Colorado 
potato beetle; two shrews (Sorex vagrans amoenus) from Crater 
Lake, Ore., had fed on Silphidae, one to the extent of 50 per cent, the 
other to 100 per cent of the total food. A prairiedog (Cynomuys 
gunnisoni) from Magdalena, N. Mex., had nothing but remains of 
Calosoma triste in its stomach, and a badger (Taxidea taxus) from 
Ash Meadows, Nev., had eaten no fewer than 150 Calosoma 
prominens. 

Passing to the enemies of beetles in the insect kingdom, it is well 
known that the various predatory tribes make no exception of beetles 
even though their generally hard integuments would seem to be a bar. 
Mantids, chrysopids, robber flies, predacious bugs and beetles, wasps, 
ants, dragonflies, and spiders all feed upon beetles, and every tabula- 
tion of the species eaten by them shows “ protected ” forms liberally 
represented. Beetles are subject to numerous parasites which attack 
them in all stages from egg to imago, and like most insects, they at 
times are decimated by fungal or bacterial intruders. 

While it has been impossible in the limits of this paper to discuss 
fully the enemies of beetles other than birds, a few cases may be cited 









NO. 7 PROTECTIVE ADAPTATIONS—McATEE 83 


in more detail as showing how the activities of such foes supplement 
the predatory activities of birds. Take for example the wood-boring 
beetles, which although they are eaten by birds to an extent that 
indicates that no special protective adaptation operates in their favor, 
yet are shielded from most birds during the greater part of their lives 
by living under bark or even within solid wood. However this habit 
does not put them out of the reach of predatory and parasitic insects. 
Thus Kleine records 159 hymenopterous parasites of Cerambycidae. 
and 136 beetle predators and 157 hymenopterous parasites of Sco- 
lytidae in Europe. (Ent. Bl. Nurnberg, vols. 4-5, 1908-1909.) The 
cotton-boll weevil (Anthonomus grandis) again passes the larval and 
pupal stages apparently well hidden from most enemies, yet some of 
the 66 kinds of birds known to prey upon it remove the immature 
stages from the cotton bolls, and in addition the weevil has 54 insect 
enemies, about half of which attack the concealed stages. (Bull. 100, 
iS. Bur. Ent. 1912.) 

So it is with the mechanically protected species; all have in the 
chains of their life histories weak links, of which hungry predators 
and assiduous parasites are not slow to take advantage. For contrast, 
consider the case of the Colorado potato beetle, an insect exposed 
almost throughout its life history, and with all of the attributes— 
color, reflex bleeding, nastiness, even poisonous qualities—of a most 
highly “ protected”’ insect. Besides the 27 species of wild birds 
known to feed upon this insect, ducks, chickens, guinea fowls, skunks, 
snakes, frogs, toads, at least eight species of Pentatomidae, two of 
Reduviidae among bugs, and eight of Coccinellidae and seven of 
Carabidae among beetles, besides robber flies, wasps, spiders, pha- 
langids, and mites prey upon the various stages. Despite all of its 
protective adaptations, the Colorado potato beetle undoubtedly has its 
full quota of foes; its rapid increase and spread over the United 
States was due to enormous increase by cultivation of a favored food 
plant and not to lack of enemies. Dr. F. H. Chittenden remarks: 
“Few, if any, noxious insects have so many recorded natural enemies 
as the Colorado potato beetle.” (Bull. 82, pt. 7, U. S. Bur. Ent., p. 85, 
Feb., 1911.) In other words, the potato beetle, being an important 
economic insect, has been much studied, and among other things we 
have learned that it has numerous enemies. If less were known about 
the species it would be hailed as a marvelous instance of protective 
adaptation ; facts are a terrible handicap to theorizing. 

Discussion—In general we have seen that whatever the beetle, 
something in the way of protective adaptation has been claimed for it, 
yet practically all are eaten. On the other hand we have also seen that 


84 SMITHSONIAN MISCELLANEOUS COLLECTIONS ~— VOL. 85 


the large families of Coleoptera, those abundant in individuals, are 
most freely eaten by birds, while the small families with few species 
escape with small losses. It is the old story over again of food sup- 
plies (beetles in the present consideration) being drawn upon in pro- 
portion to their abundance and availability. 


MECAPTERA (SCORPIONFLIES ) 


Protective adaptations —The scorpionflies are predacious ; those of 
the genus Panorpa commonly have yellow bodies and black markings 
in the wings ; and the males have enlarged genitalia carried aloft some- 
what like the stings of scorpions. The species of Bittacus resemble 
crane flies. 

Bird enemies.—We have only five records of scorpionflies being 
eaten by nearctic birds, these being distributed among four species. 

Number of identifications, 5; percentage of identifications among 
those of all insects, .0026; percentage of species in this group among 
the whole number of insect species, .0260. 

Other enemies.—There seem to be no records of such. 

Discussion.—Poverty of data is the chief characteristic of the record 
for scorpionflies. These insects are not an obtrusive part of the insect 
fauna and have been little studied. The question of the efficiency of 
their protective adaptations in relation to predators can hardly be 
intelligently discussed at present. 


DIPTERA (FLIES) 


Protective adaptations——Not much has been written about the pro- 
tective adaptations of diptera, the suggestion most often made being 
that a considerable number of them “ closely resemble wasps, and bees, 
and no doubt derive much benefit from the wholesome dread which 
those insects excite.” (Wallace, Natural selection, p. 69, 1891.) The 
families that have the most numerous species supposed to resemble 
Hymenoptera are the Stratiomyidae, Bombyliidae, Asilidae, Co- 
nopidae, and Syrphidae. Many flies have metallic colors, which are 
alleged to be warning; such insects are common among the Stratio- 
myidae, Dolichopodidae, Tachinidae, and Muscidae. A large number 
of Diptera pass the greater part of their lives in the larval stage and 
many of these larvae are more or less protected from birds by their 
habitat, as the Cecidomyidae in galls, the Mycetophilidae and others in 
fungi, the Culicidae, Chironomidae, many Tipulidae, the Simuliidae, 
Stratiomyidae, Tabanidae, and Ephydridae in mud or water; and 
various others in excrement and other decaying organic matter. Of 
course this sort of protection is of no avail in the case respectively of 





~ 


NO. 7 PROTECTIVE ADAPTATIONS—McATEE 85 


birds which eat galls or fungi or which obtain their food chiefly in or 
about water, or which feed directly upon or by tearing up carrion and 
the like. 

Bird enemies.—Until comparatively recently it was very difficult to 
get identifications of flies found in bird stomachs, and even more so 
of their larvae, hence nearly half of the identifications of Diptera 
were not carried further than to the order. 

The large proportion of the unidentified to the total number of 
records of Diptera has one advantage, namely that it is distributed 
probably among nearly as great a number of species of birds as are 
the records for all flies. Since that number exceeds 250, we may be 
sure that there is no group of birds that habitually avoids Diptera. 
Among these records are numerous instances of from 50 to 500 
specimens of flies or their larvae being taken at a single meal, good 
evidence that the flies concerned were not at all distasteful. These 
data are sufficient commentary also on the state of determinations of 
Diptera, scores of specimens being present and not being named even 
to the family. The material will be re-examined in the future to 
obtain more satisfactory results, but there has been no time for that 
in connection with the present paper, which is wholly a by-product. 

A satisfactory discussion of the relations of birds to Diptera is 
hardly possible therefore, and the best that can be done is to present 
the fragmentary data available and to make allowances for deficiencies. 





Identifications of Diptera 


Percentage 
of species 
Percentage of in this family 
identifications among the 
among those whole number 
Number of of all of nearctic 
Family identifications Diptera Diptera 1 
Tipulidae ................ 1,505 14.4426 5.7309 
Did ACS cass eseiaysier oo ere etna he 3 LO27y, 0922 
Psychodidae ............. 5 0461 .3920 
Chironomidae ............ 1,003 9.2562 3.0903 
Gilicidae | Seve .e sc ce sneie ne 112 1.0336 1.9257 
Mycetophilidae ........... 53 4891 2.9058 
Cecidomyidae ............ 15 1384 1.6835 
Bibionidae ............... 140 1.2920 8648 
STII AG: cis-sparsisiesaa'sieis hare 8 .0738 3173 
Blepharoceridae .......... I .0092 1384 
ny pada: Yates cise sete oe ere I .0092 .0807 
Orphnephilidae ........... 08 Ra .OITS 
Stratiomyidae ............ Vee 6.7553 3.4362 
Wabanidae- :.....<-.cce~0es 336 3.1008 3.5285 
Acanthomeridae .......... Lee aie S153 


“Computed from Aldrich, J. M., A catalogue of North American Diptera, 


etc., Smithsonian Misc. Coll., vol. 46, pp. 1-680, 1905. 


86 





SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


Identifications of Diptera—Continued 


Number of 


Family identifications 
Meptidacmeracesen = sore 42 
Nemestrinidae "22.2... Js: eae 
Cyrtidaernavesaset chee I 
Bombyliidae) 2asseesssee- 8 
Wherevidae wary: sss cence 5 
Scenopinidael wanes se. soe I 
My daidaey ess eene nee 
A proceridaemer aaa eee nae vine 
Asiltdaet tration eae 170 
Dolichopodidae 232 soe 86 
Hmpididaes joann eee 48 
onchopteridaess.2.0 en ae 
Bhoridae: t5ccque ntsc ) 
latypezidae (ye sa. ones 
Pipunculidae” Wem ose ee - sie 
Synphidaemancecceeeeaee 259 
Conopidael ae neemagce nek ae 
Oestridaey tases eee 4 
Machinidae jcc eecs ass 54 
Dextidaen perenne oe es 
Sarcopnagidaey (sen ay-ue. 102 
Muscidae (sens. lat.)...... 512 
“Anthomyiidae: ..5...2...:. 109 
Scatophagidae: Aasnace nee 79 
Heteroneuridae ........... 7 
EHelomyzidae sesnee oe II 
Borboridae) saceere eee 2 
Phycodromidae 22... 5.22 20 
Sciomyzidaene seaen sarees 13 
Sapromyzidael sas... o6 oo" - 24 
Ontalidae 2a csacnas asta 18 
Rhopalomeridae .......... one 
dinypetidae mace aa.t elena 10 
Mucropezidae sa.s-- 42... ae 
Sepsidacmusccr acetic ene 8 
Rsilidaepuncsetcsc sore ate 
Diopsidacmnn ceils ere ate 
E phy dridae sas acs eee 305 
©scinidde meer nea 20 
Drosophiliddes a... - 24-5. 2 
Geomyzidae ssn ce ccc es I 
Agromyzidae’ (22... e-- ose 2 
Elippoboscidae 2.....-. 4. -1- I 
Nycteribiidaer cms. ee eee es 
Wnidentiied 44-02 ose 4,904 


Percentage of 
identifications 
among those 
of all 
Diptera 
387 6 


.0092 
.0738 


0461 
.0092 


1.5688 
.7936 
-4430 


.0830 
2.3902 
.0369 
4983 
9413 
4.7250 
1.0059 
.7290 
.0646 
1015 
.26076 
.1846 
.1200 


2215 
1661 


-0923 


.0738 


to 


8147 
.1846 
10184 
.0092 
.0184 
0092 


45.2566 


Percentage 
of species 
in this family 
among the 
whole number 
of nearctic 
Diptera 


1.4644 
0092 
-4843 

5.2581 
.8187 
.1268 
-5074 
.0807 

6.3536 

6.2959 

5.4196 
-0346 
.7380 
2998 
.3229 

9.3401 

1.0378 
-3459 

12.6841 

1.9833 

1.4183 


3.4478 
9917 
-1614 
.4728 
.2707 
.0231 
.7380 

1.2338 

1.7181 
.0346 

2.3984 
-7380 
.3344 
.2883 

I.O1I5 

1.6720 

1.5451 
8763 
.1730 

1.0054 
5304 
.0576 











NO. 7 PROTECTIVE ADAPTATIONS—McATEE 87 


From the foregoing table it is evident that crane flies (Tipulidae), 
midges (Chironomidae) and mosquitos (Culicidae) are adequately 
represented, and it is fair to say that an important reason for this 
showing is that the groups of birds eating most of these flies and their 
larvae have been examined rather recently and that in consequence 
closer identification of their food items has been made. This would 
indicate that records for the other families will be similarly increased 
by future studies. It is worth noting that most of the larvae of 
Chironomidae which are so commonly eaten by birds are red (a 
warning color) so much so as to be popularly called “ bloodworms.” 
There are numerous instances of hundreds of these larvae being taken 
at a single meal. 

The more than 10,000 records cf Diptera mark these insects as a 
valuable bird food; as in other cases certain birds prey to a greater 
extent upon the group than others ; of these may be cited seven species 
of swallows which make 13 per cent to 40 per cent of their total food 
of flies and an equal number of flycatchers consuming them to the 
extent of from II per cent to 44 per cent of their entire subsistence. 

Total number of identifications, 10,836; percentage of identifica- 
tions among those of all insects, 5.6757; percentage of species in this 
order among the whole number of insect species known, 11.4432. 

Other enemies—Fishes are among the most important enemies of 
flies having aquatic immature stages. Pearse, writing of the food of 
33 species of fishes in Wisconsin lakes, reports 20 per cent of their 
food to consist of flies and their larvae, chiefly the latter. Marine 
Chironomidae are eaten by shrimps and sea-anemones. A variety of 
fishes, the top minnows and killifish in particular, are such efficient 
enemies of mosquito larvae that they have been widely used in 
mosquito-control campaigns. Diptera are eaten quite freely by frogs 
and toads and to a lesser extent by lizards, snakes, and turtles. Among 
mammals, shrews, moles, and bats feed regularly and extensively upon 
Diptera; other mammals that get at least some Diptera are mice, 
squirrels, foxes, and armadillos. 

Among their own kind, 7. e., insects, about all the predacious kinds 
feed freely on flies. The latter are soft-bodied insects easily pierced 
by the sucking predators or chewed up by the biting kinds. Tiger 
beetles, assassin bugs, mantids, ants, panorpids, dragonflies, and robber 
flies and other predacious members of their own order habitually feed 
upon flies. A number of families of wasps, such as the Nyssonidae, 
Bembecidae, Crabronidae, and Vespidae, prey freely upon Diptera, 
and spiders gain from their ranks a considerable share of their sub- 


88 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


sistence. Flies appear to be subject to parasitism only to a conpara- 
tively slight extent but some of them are decimated by fungal diseases, 

Discussion—While flies in the adult stage appear to have a degree 
of freedom from predators, it is evident that the immature stages of 
many groups of them pay a heavy toll; the chief food of predacious 
fly and beetle larvae that live under bark, in decaying fungi or carrion 
are the fly larvae they find there; the chief food especially of the 
young of a great many fresh-water fishes again are fly larvae and 
pupae; and a very important element of the food of the mold- and 
earth-traversing shrews and moles are the larvae of flies. Fly larvae 
perish in large numbers also because of the drying up or exhaustion 
of their breeding nidus. Possibly some relative good fortune for the 
adults may be only compensatory, but so little is known about the 
subject that discussion is not on a very firm basis. Regarding the fate 
of adults it is worth while recalling the all but universal destruction at 
times wrought among the ranks of its hosts by the fungus Empusa 
muscae. 

Evidence showing the importance of availability as regulating the 
consumption of dipterous food is presented in testimony of an Alaskan 
correspondent about birds feeding on mosquitos. These insects, so 
much more prominent an element of the insect fauna of that territory 
than they are in the United States, apparently are fed upon much 
more freely by birds. This correspondent, A. H. Twitchell, a reindeer 
breeder, reports all small*birds frequenting the vicinity of his camp, 
as myrtle, blackpoll, and Wilson’s warblers, Gambel’s sparrow, and 
Alice’s thrushes preying regularly on mosquitos and feeding them 
extensively to their young. 


HYMENOPTERA (ANTS, BEES, WASPS) 


Protective adaptations——In selectionist writings, Hymenoptera 
usually are classed as the very acme of protected insects, and pro- 
tective qualifications are broadly assigned to the whole group. Poulton 
says: “Ants and wasps are known to be aggressive dominant insects 
avoided by the majority of insect-eating animals.” (Essays on evolu- 
tion, p. 281, 1908.) Drummond, in similar vein, declares that “ well- 
armed or stinging insects are always conspicuously ornamented with 
warning colours. The expense of eating a wasp, for instance, is too 
great to lead to a second investment in the same insect, and wasps 
therefore have been rendered as showy as possible so that they may 
be at once seen and as carefully avoided. The same law applies to 
bees, dragonflies, and other gaudy forms; and it may be taken as a 
rule that all gaily-coloured insects belong to one or other of these two 


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NO. 7 PROTECTIVE ADAPTATIONS—McATEE 89 


classes; that is, that they are either bad eating or bad stingers.” 
(Tropical Africa, p. 163.) 

Mimicry of a group is supposed to be a tribute to its specially 
defended character and it is said that: “ The hymenoptera including 
the formidable hornets, wasps, bees and ants are more frequently 
mimicked than any other order.” (Poulton, The colours of animals, 
p. 245, 1890.) “‘ Stinging hymenoptera . . . . are sedulously avoided 
by insectivorous creatures in general.” (Bastin, Insects, their life- 
histories and habits, p. 247, 1913.) Numerous Hymenoptera which 
do not possess stings are said to mimic those that do have them and 
species of one non-stinging group, the sawflies, are alleged to be 
protected in the larval stage by distasteful or disagreeable internal or 
external secretions. 

Bird enemies.—For many years difficulty was experienced in ob- 
taining identifications of hymenoptera and the following table plainly 
shows the effect of this situation, more than a third of the determina- 
tions being to the order only. 


Identifications of Hymenoptera 


Percentage 
of species 
of this family 
among the 
whole number 
Percentage of of species 
identifications of these 
among those groups in 
; Number of of all New York 
Family identifications Hymenoptera State 1 
BOVE CAG vhs sus Ssg'PA Kaw eee 2 .0074 .4640 
Pamphiliidaé ........«.<.- 5 0185 1.2529 
Tenthredinidae ........... 263 .9732 17.5407 
PIpMVATUdae: 55. ovis caweled 5 0185 .3248 
BUIICIGAC) ules Sawer xed «eins 16 .0592 3719 
MPEDIIGAe: Co se.t sic bec wed 2 .0074 2320 
O©ryssidae) ce sicc ew oct see I .0037 .1392 
Tenthredinoidea (further 
unidentified) .......... 85 3145 a 
Mipionidae % i.<ihdwescans 2 2604 9745 
PRAY SUGAG <5, fap cwlsaiseoox 13 .0481 .2320 
Capitoniidae ...........0.5 I .0037 nuk 
BTACONICAG: 2s sacs Saceae 28 .1036 9.0488 
Fvaniidae ................ 5 0185 3719 
Trigonalidae ............. I .0037 .6496 
Ichneumonidae ........... 1,003 4.1184 25.6614 
Ichneumonoidea (further 
unidentified) ........... 13 .0481 


*Computed from Bradley, J. Chester, Hymenoptera, in A list of the insects of 
New York, etc., Mem. 101, Cornell Univ. Agric. Exp. Sta., pp. 870-1,033, 1926, 
the most comprehensive checklist of nearctic forms available. 


go 


SMITHSONIAN MISCELLANEOUS COLLECTIONS 


Identifications of Hymenoptera—Continued 


Family 
Hicitidaeteerer reise cenit: 
Cynipidaewnanyae eee 
Pteromalidae 7... acces ee 
Eupelmidaer.... ees ccmiee 
Callinomidaess. ea coe: 
Eanytomidaes sees cere 
Berilampidaeva. casa ede e 
Ghalcididaew ater aac tr 
IWeucospidael semen see er 
Chalcidoidea (further uni- 

denititied|)imesmirineeierrats 
Platyeastnidaceen-as eerie 
Scelionidaes-aeee eres 
Geraphronidacesa..e acne 
Diapriidae’ vere cis estan 
Belytidae, facemas acca ausee 
Serphidae a2c; ance ue oeice 
Pelecinidae nc. aese ocr 
Serphoidea (further  uni- 

Centitied)) eee 
Hormicidacma.cneea ieee. 
Miyrinicidaes.ccc.cr eee 
Formicoidea (further uni- 

dentitred) meee eeeericee 
Ghrysididaeien een eee 
Bethylidaeseamecescicmaeee. 
Diryinidaes eae acter 
Soolhittkeseoopedeumoddacdse 
IMAGINES Gaagecoocnnnoe 
Mintillidae Mearsctcetse eee 
Psammocharidae .......... 
Mumenidae jae seyta acre te: 
Wespidacue ener erate: 
Vespoidea (further uniden- 

titted) Me ercacsterserta cers 
Sphecidacwarecieermerce cre 
Bembecidae ...5......:... 
Sphecoidea (further uni- 

Gentitied)) erence 
Flalictidaeyciwoncicscto oe 
Andrenidacmescnrcseiecicent 


Number of 
identifications 


14 


= 


_ 
SOM OOUNT IDE RO BS ON eS 


144 


23 
134 
92 


Percentage of 
identifications 
among those 


Hymenoptera 


of all 


.0518 
.1400 
0814 
OTT 


0444 


.0185 


OIII 
2701 
.0037 


.0518 
.0259 
0148 
.0074 
.0444 
.0259 
0296 
0333 


.0666 


7.7410 
4.4404 


34-9715 


_ 


2257 
-0333 
0148 
.0540 
OLLI 
0851 
.1332 
1295 
5328 


8215 
.2812 
0148 


.O851 
.4958 
.3404 


Percentage 
of species 
of this family 
among the 
whole number 
of species 
of these 
groups in 
New York 
State 
0464 

6.7286 
.9280 
.1392 
.6032 

1.5313 
.0464 

3.1091 
.0464 


.7889 
1.1137 
.4640 
5508 
.4176 
0464 


1.1137 
1.2529 


1.0673 
1392 
3248 
9281 
.0928 
9745 

3-0195 

1.8562 
5104 


3.2019 


2.5980 


9745 
2.4504 


VOL. 85, | 


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7 PROTECTIVE ADAPTATIONS—McATEE gl 


Identifications of Hymenoptera—Continued 
Percentage 
of species 
of this family 
among the 
whole number 


Percentage of of species 
identifications of these 
among those groups in 
Number of of all New York 
Family identifications Hymenoptera State 
IPantireidae@ .2:....0..<s. ; 2 .0074 3248 
Nomadidae ............... 12 .O444 8817 
Buceridae’ .. 0.2.00 00660: 10 .0370 shan 
Anthophoridae ........... 4 0148 5568 
ely lACId ae! fs 2 sects cee nes I .0037 3248 
@olletidaes.. 2. s.c6in.- 8 0296 3248 
Megachilidae ............. 18 0666 1.6241 
Geratinidae 22.2.2. 05.0..-6.- 2 .0074 .0464 
SELIG Ce eee ieiec cate farats odoinn ate I .0037 .0928 
Gy OCOpid ae ecm atsatie = wale 2 .0074 .0464 
END a Cra tact teuay ote cit Serer ose 139 5143 8353 
Apoidea (further unidenti- 
CCR Maret scn ees cre eet 3 B72 1.3705 
Unidentified .............. 10,682 39.5266 


Examination of the preceding tabulation shows again the influence 
of availability upon choice of food. It is at once evident that the 
groups more numerous in species and individuals are taken most often 
by birds. Whether all are taken in sufficient number to satisfy expec- 
tations is subject to discussion but the relativity of capture to abun- 
dance is unmistakable. Superfamilies such as the Cynipoidea and 
Chalcidoidea, owing to the minute size of most of their species, could 
not be expected to figure largely in the diet of birds, and the same is 
true for most Serphoidea. These are just the groups and the only 
ones in the table except the Sphecoidea that seem obviously to be 
inadequately represented. The Sphecoidea perhaps verge toward the 
opposite limit of size for bird food. 

Since so many Hymenoptera were determined no further than to 
the order, the number of species (over 300) of birds eating these 
unidentified forms may be taken as an approximation to the entire 
number of bird species consuming Hymenoptera. It is enough at any 
rate to indicate that Hymenoptera are eaten by birds of all groups 
studied, just as the total number of records (over 27,000) of Hymen- 
optera clearly shows that these insects are one of the most important 
elements of bird food. 

Beginning our consideration of the Hymenoptera with the sawflies, 
it may be said that some of these insects are alleged to obtain pro- 
tection from their resemblance to stinging members of the order. As 





Q2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


to the larvae, Poulton says: ‘‘ Numerous experiments have convinced 
me that the latter are almost invariably distasteful.” (Essays on 
evolution, p. 238, 1908.) However, the present tabulations reveal more 
than 30 species of birds as predators upon sawfly larvae and no fewer 
than 50 to 100 specimens of these larvae have been found in single 
stomachs of the mockingbird and from Io to 25 in those of other 
species. Hewitt records seven species of British birds as feeding upon 
larvae of the large larch sawfly (Nematus erichsonii) (Bull. 10, Div. 
Ent. Dom. Can. Dep. Agr., p. 22, 1912), and attributes to them great 
destruction of the larvae. The 380 records of Tenthredinoidea in 
our table are distributed among 99 species of birds and such wasp- 
suggesting forms as Cimbex, and the horntails of various kinds, with 
24. records, seem to be proportionally represented. 


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Most of the Ichneumonoidea are not credited with any special 
defenses besides their resemblance to stinging hymenoptera, and the 
more than 1,200 records of their being eaten would seem to indicate 
that this means of protection is more imaginary than real. Some of 
the more interesting records may be cited as indicating the extent to 
which birds eat these insects: Protapanteles: 50 specimens in the 
stomach of an English sparrow (one of a series of 12 containing 10 or 
more each), and 120 in one stomach each of a Brewer’s blackbird and 
an Aleutian sandpiper ; Ichneumonidae, further unidentified: 68 speci- 
mens in a sanderling’s stomach (19 birds have from 10 to 42 records 
each) ; Ichneumon sp.: 37 specimens in the stomach of a burrowing 
owl; Glypta tuberculifrons: 44 individuals taken at a meal by a 
yellow-throated vireo; Ophion spp.: 54 records for these large 
ichneumons which can sting. 

Most ants, their size considered, can bite severely ; their body fluids 
contain formic acid and other pungent substances ; and many of them 
also can sting. As further tribute to their prowess the reference of 
Poulton may be quoted to the “ numerous mimetic resemblances to the 
aggressive, abundant, and well-defended ants.”’ (Essays on evolution, 
p. 252, 1908.) Badenoch says that ant-models “as a rule are exempt 
from persecution.” (Romance of the insect world, p. 300, 1893). 
The confidence of selectionists in the protective nature of ant mimicry 
is further shown in the following statement by Donisthorpe on Nabis 
lativentris: “I consider this insect to be an ant mimic in its earlier 
stages, when it is usually found in the company of ants. From this 
mimicry it obtains protection from outside enemies, both as much 
when away from ants as when with them.” (Ent. Monthly Mag. 
ard seri, vol.7, pp; 137-138, 1921.) 





NO. 7 PROTECTIVE ADAPTATIONS—McATEE 93 


But why this conclusion? The more than 12,000 records of ant- 
eating by the birds represented in our tabulations certainly indicate 
no sort of immunity on the part of ants from the attacks of birds. 
These records are shared by well over 300 species of birds which, 
practically speaking, means that all birds eat ants. Ninety-three of 
the species of birds represented in our tabulations have from 10 to 
4g records of ant-eating each, 18 others from 50 to 99, 17 others 
from 100 to 199, four additional over 300, and one additional species, 
the eastern flicker, in excess of 500. All these records are among the 
Formicoidea further unidentified, three-fourths of the total for all 
ants. From 200 to 300 ants at a meal is a common number; the 
swallows often get 800 or more; the nighthawk 1,000; and wood- 
peckers 2,000 or more. In two cases stomachs of flickers yielded more 
than 3,000 ants each, and in one case more than 5,000. Out of 684 
stomachs of this last named species, 524 contained ants. 

In this connection the extent to which ants enter into the diet of 
certain birds is of considerable interest ; our five species of thrushes 
of the genus Hylocichla consume ants to an average of 12.65 per cent 
of their total food, while 16 species of woodpeckers, the food of 
which was tabulated by Prof. F. E. L. Beal, ate ants to extents vary- 
ing from 5 to 85 per cent of their entire subsistence, the average for 
the 16 species being 28.49 per cent. 

The stinging ants, of course, are the most highly “ protected” of 
all and it is unfortunate for our discussion that the group is so 
poorly represented in the United States. Myrmicidae, including 
Ponerinae and Dorylinae, are more or less generally provided with 
stings, which however in the most of our species are too small to 
inflict damage on a human subject. Our tabulations show 1,200 
records for Myrmicidae, and they are eaten in just as large numbers 
as are other ants. The harvester ants of the genus Pogonomyrmex 
are larger and equipped with stings which can painfully wound a 
human being. We have 66 records of these ants being taken by 25 
species of birds; no fewer than 200 and 400 individuals were taken 
from the stomachs of two Texan nighthawks. Mitchell and Pierce 
write of birds feeding freely on Pogonomyrmex and note a case of a 
group of nesting jackdaws (Megaquiscalus) cleaning up a colony in 
a short time. (Proc. Ent. Soc. Washington, vol. 14, no. 2, p. 72, 
June, 1912.) 

Among the remaining, mostly stinging, Hymenoptera are the 
Chrysididae, supposed to be protected by their hardness, abililty to 
roll into a ball, and by metallic colors. We have 61 records of these 
being eaten, shared by 37 species of birds. The Vespoidea or wasps 


‘ 


QO4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


as a whole have 822 records representing 140 species of birds. Some 
of the larger numbers of wasps consumed at a meal were 10 Vespula 
germanica by a wild turkey, 10 Polistes sp. by a yellow-billed magpie, 
and the following numbers of unidentified wasps by the birds men- 
tioned: purple martin 17, olive-sided flycatcher 24, and kingbird 30. 

The 103 records of Sphecoidea are distributed among 43 species of 
birds, none of which took any notable number of these large insects. 
Bees all sting, and the 797 records of their being eaten by the birds 
examined by us would seem to indicate considerable disregard for 
the stings on the part of birds. The number of species of birds repre- 
sented in these bee-eating records is 144. Thirty-two species of birds 
took honey bees (Apis mellifera) on a total of 118 occasions and nine 
species of birds ate bumble bees a total of 18 times. These numbers 
of determinations seem in fair proportion to the availability of the 
bees concerned. The largest numbers of bees taken at a meal were 
26 Andrenidae by a rose-breasted grosbeak, 34 honey bees by a cactus 
wren, and 106 of the domestic species by a road-runner. 

It is of interest to note that besides the thrushes and woodpeckers 
previously mentioned, two other groups of birds are very notable 
consumers of Hymenoptera. Thus the seven species of swallows make 
an average of 24.9 per cent of their diet of these insects, and 14 
species of flycatchers average 33 per cent. 

Summary.—Number of identifications of Hymenoptera 27,025; 
percentage of identifications among those of all insects, 14.1551; per- 
centage of species in this group among all insect species, 17.1798. 

Other enemies—Hymenoptera, having so few aquatic representa- 
tives, do not figure in the diet of fishes as anything but an incidental 
item, consisting of specimens, a considerable proportion of them ants, 
that have approached too near or have fallen upon the surface of 
the water. 

Passing to batrachian enemies of Hymenoptera we may note that 
Kirkland found ants to compose 19 per cent of the contents of 
149 toad stomachs, and that he had evidence also of their feeding ex- 
tensively upon honey bees. Garman also found not only the common 
toad (Bufo lentiginosus) but also the pigmy toad (Bufo quercicus) 
to be very fond of ants. Toads have been observed to feed freely 
upon the larger stinging insects also, such as yellowjackets and wasps. 
Drake found 25 ants and 21 other Hymenoptera in 209 stomachs of 
the leopard frog. Insects of this order, especially ants, are eaten by 
all frogs and toads and to a considerable extent by salamanders also. 
Most lizards feed freely on ants, bees, and wasps. Winton found 
agricultural ants (Pogonomyrme.x) in 80 per cent of the horned-toad 








NO. 7 PROTECTIVE ADAPTATIONS—MCcATEE 95 


stomachs (485) examined by him, and Mitchell and Pierce record 
the extermination of a colony of these ants by horned-toads. Several 
species of snakes and a few turtles feed to a slight extent upon 
Hymenoptera. 

Among mammals, moles prey extensively upon ants, and shrews 


-and bats by no means avoid them. In our country armadillos are 


destructive ant eaters and in other continents various mammals 
specialize upon ants. Spermophiles and other slightly insectivorous 
rodents include ants and other Hymenoptera in their bill-of-fare. 
Skunks are assiduous in digging out the nests of yellowjackets (lVes- 
pula), the comb, its contents and active inhabitants of the nest all 
being devoured. Mice, weasels, foxes, and especially badgers simi- 
larly ravage the nests of bumble bees, while bears plunder not only 
these insects but also honey bees and hornets. Meadow mice and 
shrews have been found to be among the most effective enemies of 
sawflies, extracting the larvae from the cocoons, and these and deer 
mice take a heavy toll of the Hessian fly, nibbling the stem-galls and 
devouring their inmates. Squirrels feed freely upon galls produced 
by Hymenoptera. 

The insect enemies of Hymenoptera are numerous and effective and 
strangely enough many of them are within the ranks of the order. 
Philanthidae use Aculeates for food, many bees, cuckoo wasps, and 
the like live parasitically in the nests of other Hymenoptera; the 
surprising phenomena of hyper-parasitism reveal numerous serious 
enemies of Hymenoptera among their own kin; and a number of 
dipterous parasites of sawflies, bees, and wasps are known. The 
so-called guests in the nests of bees and ants destroy many of the 
larvae of their hosts. Predacious insects such as assassin bugs, Phy- 
matidae, dragonflies, and robber flies feed freely upon Hymenoptera, 
the last-named foes almost appearing to have a preference for the 
larger and better armed sorts of stinging Hymenoptera. Spiders of 
certain species entrap and devour large numbers of Hymenoptera. 
Nematode and protozoan parasites exist and some Hymenoptera have 
important fungal and bacterial diseases. 

Discussion.—According to selectionists, Hymenoptera are the most 
highly protected insects and the so-called mimicry of examples of this 
order, such as the ants, by numerous spiders, long-horned beetles and 
rove beetles, plant-bugs, and other insects is regarded as strong 
evidence for the truth of the claim. Let the case be presented in the 
words of an advocate (Poulton, Essays on evolution, p. 260-261, 
1908) : “ The means by which the resemblance to ants is brought about 
are diverse, the end—the resemblance itself—is uniform. Further- 

7 





96 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


more the likeness is almost always detailed and remarkable, however 
it is attained, while the methods made use of differ absolutely. . . . . 
We are compelled to believe that there is something advantageous in 
the resemblance to an ant, and that Natural Selection has been at 
work. The phenomena do not merely disprove all other suggested 
causes of change ; they constitute the most powerful indirect proof of 
the operation of Natural Selection.” 

If the above reasoning has any application so far as the attacks of 
predators upon ants are concerned, we should expect some evidence 
that ants are relatively free from such attacks. Let us see what is the 
case. Beginning in the very homes of the ants we find, among crea- 
tures habitually living in ant colonies, that numerous Staphylinid 
beetles devour the brood, besides crippled and even normal ants; the 
larvae of Clythrinae (Chrysomelidae) feed on the eggs; lycaenid 
caterpillars and paussid beetles eat the eggs and larvae. Numerous 
ectoparasitic mites and some chalcidids also attack the ants in their 
domiciles, as well as entoparasites among the Strepsiptera, Phoridae, 
Conopidae, Braconidae, Chalcididae, Proctrotrupidae, and Nematoda. 
Ants have very important predatory enemies in their own ranks, 
namely the doryline and slave-making ants. Ant-lions of the family 
Myrmeleonidae, Diptera with similar habits, predacious wasps, es- 
pecially the Crabronidae, assassin bugs, ground and tiger beetles, and 
spiders are serious invertebrate enemies of ants. Most toads, frogs, 
and lizards, the amphisbaenids, and certain snakes feed upon ants; 
fishes take them when opportunity affords; practically all birds eat 
ants, several groups as the song thrushes, ant-thrushes (Formi- 
cariidae), and woodpeckers depending upon them for a large part 
of their food; in the same way most of the insectivorous mammals 
are fond of ants and several groups of this phylum are specialized 
ant eaters, namely Echidnidae (spiny anteaters) among the Mono- 
tremata, the banded anteater (Myrmecobius) among the Marsupialia, 
and nearly the whole order of Edentata (antbears, pangolins, and 
armadillos ).” 

In fact it would be difficult to name a group of insects that is so 
thoroughly preyed upon as the ants, and impossible to name one that 
has so many specialized foes scattered through the various animal 
phyla. So far as predatory attack is concerned, it would seem that 
ant-mimics court rather than avoid danger. To recapitulate: if there 
is any virtue in the protective adaptations of the “aggressive, abun- 


*For a comprehensive account of “The predacious enemies of ants,” see 
Bequaert, Bull. Amer. Mus. Nat. Hist., vol. 45, pp. 271-331, 1922. 





NO. 7 PROTECTIVE ADAPTATIONS—McATEE 97 


dant, and well-defended ants,” it should be apparent in some lessening 
of predatory attacks upon them. However the very opposite is true 
and the case affords the best sort of proof of the thesis of this paper, 
namely that the number of enemies, or perhaps better stated, the total 
losses to enemies, is in direct proportion to abundance of the group 
concerned. 

Selectionists regard bees as very highly protected insects, but taking 
the honey bee as type of the group because more is known about the 
species than any other, we find that bee-keepers complain bitterly of 
the numerous enemies of the inmates of their hives. Wasps, velvet- 
ants, robber flies, Phymatid bugs, mantids, and dragonflies are serious 
insect enemies of honey bees; spiders, toads, lizards, rats, mice, and 
skunks prey upon them; numerous wild birds join in the attack (32 
nearctic species according to our tabulations), and domestic ducks are 
said to be insatiable in devouring bees. A mite is the primary cause of 
the so-called Isle-of-Wight disease among hive-bees; ants and wax 
moths destroy the comb ; and there are at least two serious infectious 
diseases. At times bees rob other colonies, the rifling being accom- 
plished however only after great slaughter. In the case of the honey 
bee, much study has been devoted to the insect and we know consider- 
able about its enemies, but the ruling principle is as clear in this case 
as in that of the ants, namely, that common species have numerous 
enemies. 

Since ants and the honey bee fairly exemplify two of the main 
phases of protective adaptation in Hymenoptera, despite which these 
species clearly have their full quota of enemies, we cannot doubt that 
other species of the order, when they are as well known, will prove to 
have predatory foes fully in proportion to their relative abundance. 

In fact the 27,000 records of Hymenoptera now available are suf- 
ficient indication that the order contributes its due toll to the subsis- 
tence of one of the chief groups of its enemies—the birds. 


ARACHNIDA (SCORPIONS, SPIDERS, Ticks, Erc.) 


Protective adaptations—Most arachnids possess venom of suf- 
ficient strength, and means of injecting it into other creatures, to 
enable them to overcome the animals upon which they prey. Numbers 
of them have chelicerae, which in a few cases are rather powerful. 
The poisonous nature of many of the species has been greatly exag- 
gerated especially by primitive races of man so that they are held 
in extreme dread. 





98 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


Besides these direct means of defense, ticks and spiders exhibit in 
a high state of development that class of protective adaptations known 
as cryptic coloration (both defensive and aggressive). Certain groups, 
however, are brilliantly colored; some spiders also have the body 
integuments hardened and produced in the form of angles or spines, 
and numerous spiders resemble ants. Among the forms of this class 
ordinarily coming to the attention of man, spiders are by far the most 
numerous, but the small often minute mites prove, when the care 
necessary to their study is bestowed, to be exceedingly abundant. 
However, these small forms are beneath the notice of most birds. 

The following table shows the relation of the records of the various 
orders to their approximate representation by species in the nearctic 
region. 


Identifications of Arachnida 


Percentage 
Percentage of of species 
identifications in this order 
among those among all 
Number of of all nearctic 
Order identifications Arachnida Arachnida 1 
Wnidentitied ence or eae 26 2386 oe 
Maphostta sj sacae seas 20 .1836 .0513 
Microthelyphonida ....... ae ee 0513 
Sconpionidaweeeeeeeeenios 18 .1652 1.1801 
Pseudoscorpionida ........ 80 7343 2.4115 
Redipalpida wrmse ease: 4 .0367 .3078 
Solpucidateee: sete eee oe 2 .2203 6157 
Phalancidam eter eres 478 4.3873 3.4376 
Araneidauicjcs «rete esuciee 9,966 91.4729 66.7044 
IN CAtitlaencetoe aa eee 258 2.3681 23.0886 
Pycnogonidare ss see cciae rd 1010 2.1549 


Bird enemies.—Birds certainly specialize upon the group of arach- 
nids—spiders—that to man seems most abundant and easily available, 
over OI per cent of their arachnid captures coming from this order. 
We have records of more than 300 species of nearctic birds feeding 
on spiders. The freedom with which they take these creatures is 
illustrated by the following citations of records; of those identified 
to the order alone or about 92 per cent of the total, 81 birds have 
from 10 to 49 captures each; 28 birds from 50 to 99; 15 from 100 


*Computed from Comstock, J. H., The spider book, etc., 721 pp., 1912, with 
numbers of Araneida and Acarina approximated from the following works, 
respectively: Banks, N., Catalogue of nearctic spiders, U. S. Nat. Mus. Bull. 
72, 80 pp., 1910, and Banks, N., A catalogue of the Acarina, or mites, of the 
United States, Proc. U. S. Nat. Mus., vol. 32, pp. 505-625, 1907. 


af 


NO. 7 PROTECTIVE ADAPTATIONS—McATEE 99 


to 199; two additional above 200; one more above 300; besides the 
following with greater numbers of records; English sparrow, 420; 
Eastern meadowlark, 425; crow blackbird, 621; starling, 631; and 
crow, 722. Some of the larger counts of spiders taken at a meal 
were 25 by a Say’s phoebe, 33 by a greater yellow-legs, 46 by a 
wood duck, 58 by a Louisiana heron, 187 by a starling, and 300 by 
a hairy woodpecker. 

A rather small proportion (less than 8 per cent) of spiders found 
in bird stomachs were identified, but results obtained along this 
line show the determinations are distributed to cryptically colored 
groups as the Epeiridae (30 records) and Thomisidae (28) ; more 
brilliant forms as the Attidae (158) and to the formidable Ly- 
cosidae (370), in a way that would indicate availability to be the 
principal factor in choice. There are two records of Synemosina 
formica, the most antlike of our spiders, a small and rather un- 
common form that one would expect no more frequently regardless 
Or its “ 
resemblance to an ant is no protection whatever against predators. 


protected”’ status. As noted in the last section, however, 


There are 134 records of the cocoons or egg-cases of spiders being 
eaten showing that even these quiescent stages do not escape the birds. 
In bulk spiders do not ordinarily form any considerable percentage 
of the total food of birds, but the proportion runs as high as 6 per 
cent and 8 per cent of the annual diet in the case of certain song 
thrushes and petty flycatchers. 

There are 34 records in our tabulations of ticks being eaten, and 
224 of mites. Of interest in connection with the latter are the finding 
of 100 Parasitidae in the stomach of a red-eyed vireo; 320 mites 
further unidentified in the stomach of a pipit ; 535 water mites in the 
gizzard of a green-winged teal, and 594 of the same group in the 
stomach of a pied-billed grebe. 

Such geographically restricted and relatively uncommon forms as 
the scorpions, pedipalps, and solpugids, even though having only a 
small number of records each, would seem, nevertheless, to be amply 
represented, considering their availability. Pseudoscorpions are 
present throughout our area but lead chiefly concealed lives; the 80 
records are distributed among 22 species of birds. 

The daddy-long-legs, or Phalangida, with 478 determinations cer- 
tainly have not been slighted; 10 of the birds taking them have Io to 
1g records each; two others over 20 records ; and one each additional, 
the yellow-billed cuckoo, 34; and crow blackbird, 60. Large numbers 








100 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


of the arachnids captured for one meal were 20 by a yellow-billed 
cuckoo and 77 by an ovenbird. 

Number of identifications, 10,885; percentage of identifications 
among those of all arthropods, 5.1648; percentage of species in this 
class among those of all arthropods, 3.8254. 

Other enenues.—Spiders are frequently snapped up by fresh-water 
fishes, and mites often, especially the water mites. Pycnogonids, or 
sea-spiders, have occasionally been found in the stomachs of marine 
fishes and are consumed also by sea-anemones. Kirkland found 
spiders to compose 2 per cent of the food of 149 common toads 
examined by him, and Drake found them to constitute about 27 per 
cent of the entire number of animals found by him in the stomachs 
of 209 leopard frogs. However, practically all frogs, toads, and sala- 
manders that have been studied have been found to feed upon spiders, 
often extensively, and mites, pseudoscorpions, and phalangids are not 
neglected. Lizards commonly prey upon spiders, while snakes and 
turtles so far have rarely been observed to do so. 

Spiders appear to constitute an important element of the food of 
our shrews, and a lesser, though frequently taken, item in the diet of 
the moles. We have records of the wood rat and armadillo preying 
upon spiders, and evidence that the badger at times is a destructive 
enemy of scorpions. Monkeys and marmosets are said to be very fond 
of spiders, and anteaters also are listed among their foes. 

Of invertebrate enemies various wasps (Pompilidae, Sphegidae, and 
Trypoxylonidae) are among the most effective destroyers of spiders, 
some species preying exclusively upon them, temporarily at least, even 
upon single species. The writer has found the cells of Pelopaeus 
cementarius filled entirely with flower spiders, Misumena americana, 
supposedly one of the most perfectly protected (cryptically colored) 
species. Dragonflies prey upon spiders. Tiger beetles and ants eat 
spiders and mites, ground beetles and ladybirds also figure as enemies 
of mites and spiders. Water mites are preyed upon by dragonfly 
nymphs and aquatic Hemiptera. There are a number of kinds of 
spiders that habitually prey upon their fellows when adult, and canni- 
balism among young spiders is the rule. Mantispidae and other 
spiders eat the eggs and young of spiders, and there are many proc- 
trotrupid and ichneumonid parasites of the eggs. Parasitic nematodes 
also afflict the adults. Scorpions are notoriously cannibalistic, so much 
so in fact that it is said in some cases that whenever two of them are 
found together, one is eating the other. 





Ss es ee 


0 OO EE EEO ns 


McATEE IOI 





NO. 7 PROTECTIVE ADAPTATIONS 


Discussion.—All spiders have venom and some of them are large 
and venomous enough to be able to kill birds. The case would seem 
to be crucial for the usefulness of this direct means of defense, but 
we may well say, in the light of the evidence, that the defense is 
entirely disregarded by birds. Not only do our records show more 
than 10,000 records of spiders having been eaten by more than 300 
species of birds, but the birds emphasize their disregard for the 
dangerous qualities of spiders by making them in many cases the 
staple food for their callow young. Such minor protective adaptations 
as those of color and form necessarily fall with the greater, and there 
is no evidence whatever but that birds eat spiders under any and all 
conditions as freely as they choose. The nearly 1,000 records of 
arachnids other than spiders seem to be distributed among the orders 
in very just proportion to the extent these creatures are available to 
birds. No evidence of “ special protectedness ” obtrudes itself. 


MOLLUSCA (SNAILS, SLUGS, MUSSELS, LIMPETS) 


Protective adaptations The great majority of mollusks are 
equipped with calcareous shells into which they can entirely withdraw. 
Besides this protection more than half of the species are aquatic and 
hence are more or less out of reach of many birds. Many land snails 
have the apertures of their shells furnished with processes or teeth 
which partly bar these openings and operculi to close them. Snails 
and especially slugs secrete mucus freely ; a habit thought by some to 
be protective. Numerous mollusks are colored more or less in harmony 
with their environments, this being especially noted of marine forms 
living on seaweeds, gorgonians, etc. The nudibranch mollusks are 
characteristically brightly colored and have been said to be distasteful. 
Of shelled mollusks, Wallace remarks: “ The brilliant colors of the 
scallops (Pecten) and some other bivalve shells are perhaps an indica- 
tion of their hardness and consequent inedibility.” (Darwinism, 
p. 266-267, 1896. ) 

Bird enemies.—The tabulation of identifications herewith presented 
is the best that could be made so far as comparative records is con- 
cerned; these had to be gleaned from two sources as noted, which 
between them do not include all of the families, nor, because of 
disparity of data, do they give even the grand divisions comparable 
treatment. 


102 SMITHSONIAN MISCELLANEOUS COLLECTIONS 


Identifications of Mollusca 
Aquatic shells 


Percentage of 


identifications 
among those 
Number of of all 
Group identifications Mollusca 
Wmidentitved! 225.055 saree ee 1,032 8.7673 
Pelecypoda (further  uni- 

Gentine)! sc. 2 scene 513 4.3582 
Ostreidae) Hee kee. oswieae ten 552 4.6895 
Atomitdaeanse ere teu eer 
Dimyidaeteene ences 
Spondylrdaey Geese sees ots ifr 
Pectinidaes.mne cent 193 1.6306 
TSIM a Cae Met ee ee 
INWACBIINGEYS Geo ducuuabecece re Sate 
IMBWGINGEI® ooncacoasoducgne 674. 5.7260 
Wnioniddew eee 8 .0679 
NTCIdAC aeee Gee near ere 73 .6201 
Niiculidaetnaseen nee ore 45 3823 
Medidaewe can eee re I £0085 
Solenomyidae .....:.....%. I .0085 
@anditidae@ ses. erie oer aces theta 
Astantidacus.aje- caer cn 20 .1699 
Grassatellidacmetsa- ease cer aide 
Egycinidae, snrdas sate 2 0170 
Ungulinidae nae. c25 sem: 

Cyrenellidaci ses asese nee 

leucinidae wasnt core ce 

Diplodontidacwar.cem seme or 

@hamidaey seaport cee ee oor Bre 
Cardidaewesseeeeeeeeee 20 .1609 
Wenilidaemeecemsseacterice: 2 .O170 
lsocarditdaes we siteseiees aee <a ae 
Wenenidaerts se oasecon sso 131 1.1129 
Corbiculidacwemereee ees ere fo) 
Betricolidae ss senterecrs cis 28 2378 
Donacidaeueeemmne ar eee 122 1.0364 
Psammobidae -....526 4. he Sie 
drelilimidaeecarasoee ces oe 324 27525 
Semelidacs ache cane ae 
Gnathodontidae ........... 4 .0340 
Mactnidaew-n-rrasoe cans aoe 21 1784 
Anatinidae wan cmniy. ceils =i 


‘Compiled from Dall, W. H., A preliminary catalogue of the shell-bearing 
marine mollusks and brachiopods of the southeastern coast of the United 
States, with illustrations of many of the species, U. S. Nat. Mus. Bull. 37, 


221 pp., 74 pls., 1880. 







Percentage 
_ of species 
in this group 
among marine 
mollusks of the 
Southeast 
Coast 1 


2516 
.1887 
.0629 
1887 
1.8868 
6918 
.5031 
1.6353 


2.0755 
6289 
1.9497 
.1887 
.4402 
6918 
.1887 
.5031 
.6289 
.0629 
1.4465 
2516 
3145 
817 
0620 
.1887 
1.9497 
1258 
.4402 
3773 
-3773 
2.3270 
6280 
.1258 
3145 
8176 


VOL. 85, 





PROTECTIVE ADAPTATIONS—McATEE 


N 


Identifications of Mollusca—Continued 


Aquatic shells 


Percentage of 
identifications 
among those 
Number of of all 
identifications Mollusca 


Group 

HEVONSiGaewees cree aoc cies 
Verticordiidae ............ 
Cuspidariidae ............. 
Poromyidae .............. 
IPandondae ......--6.s +-- “0 wae 
@orbulidae........ 0.05200 2 7716 
WWitvidaeg were cares ee wae ace 10 .0849 
DHIGAVIdAG oe ae tds iietbaiets 25 2124 
pap hitdacweniericls + cees sien 400 3.3982 
Solentdae’ star. sacciscasinee 4 .0340 
Gastrochaenidae .......... 

tO ladidacmenee sans. aeee sale sie oes 
sNereCIdae muerte aeons aseetesie 5 .0425 
Gastropoda (further  uni- 

Gentitied Jee. ecco ane: 3,421 29.0631 
Wentalitdaeweccs. as ose I 0085 
Iimacimidaes 2 42.42.6605 0m. 

Gavolinidae asses aan eae 

Gyimluliidaes enw. «ese ce 

G@loOnidaetses, se eee 

@liopsidaesseeess sone n aes 

Pneumodermatidae ....... 

Actaeonidae .............. 

PIMP iCuidae eaters nae oe rye 
sornatinidacwes ee eee 77 6541 
Scaphandridae ............ 
PNDIUStrIidde? Giese: css cee 
Tat aen etesyererte tere erecta 
Philinidae) foe. ae. see eee: 
Gastropteridae ............ 
Umbraculidae ............ 
INDIVSUGACG Gane cess eens - 
Pleurobranchidae ......... 
@nchidiidae 1s) eo. 
Veronicellidae ........+... ee 
Auriculidae .............. 158 
Siphonariidae ............ 
(Gacimiidace eine eee so Die 
Terebridae ............... I .0085 
(Conidaer scsi steer Sear ae ee 
Pleurotomidae ............ 43 3653 
@ancellariidde 2. ....0:22: ae Sak 
Olividaes: 5 sécsscscsswitecs It 0934 


1.3423 


103 


Percentage 
of species 
in this group 
among marine 
mollusks of the 
Southeast 
Coast 


.4402 
.6289 
1.1321 
.5031 
-3145 
6918 
£0629 
.1887 
3145 
2516 
7547 
5031 


to 


.7673 
.6289 
1.1950 

.0629 

.0029 

1258 

.0629 

.6289 

1258 
1.2207 
.5060 
1258 
.6918 
“3073 
.0629 
1258 
1258 
.1887 
.0629 
0629 
.00603 
.1887 
.0629 
.6289 
.6918 
8.3019 
5031 
5031 


+ 


Marginellidae 


Turbinellidae 


Columbellidae 
Pyramidellidae 
Amphiperasidae 
Cerithiopsidae 
Trichotropidae 
Seguenzidae ......s55-64< 
Murritellidae S.ese weae cs 


Philomycidae 
Ampullariidae 
Assimineidae 


Aquatic shells 


Number of 
identifications 


LS 


176 


SMITHSONIAN MISCELLANEOUS COLLECTIONS 


Identifications of Mollusca—Continued 


Percentage of 
identifications 
among those 


of all 


Mollusca 


0595 


.1529 
0525 
3.6021 
2.0394 
9430 


.O169 
1.4952 


1444 
03.40 
1.8520 
.O169 
1104 
0595 


VOL. Sm 


Percentage 
of species 
in this group 
among marine 
mollusks of the 
Southeast 


Coast 
2.9560 
3145 
1258 
1.0692 
1.6981 
1.9407 
5031 
1.6352 
2.6415 
3.3962 
2516 
1.0692 
3-3333 
1.0063 
.1887 
5031 
3145 
3773 
.6918 
5031 
3145 
1.0063 
6918 
1.0063 
1258 
.0629 
.1887 
.8176 
.3145 
5031 
3145 
3145 
6918 
2516 
.1887 
8176 
1.5004 
3145 


1258 
.1887 





NO. 7 PROTECTIVE ADAPTATIONS—McATEE 
Identifications of Mollusca—Continued 
Aquatic shells 
Percentage of 
identifications 
among those 
Number of of all 
Group identifications Mollusca 
Wruneatellidae’ wa... 5.006. 
@horistidadl 2... cc0re ee ae se 
Galyptracidae .... 1. assess > 135 1.1469 
Gali emer tier eer Doe 
Amaltheidae ............. 
Xenophoridae ............ anes Sais 
INicit1 C1C Gamera en eteteme ta ie areee ie ere 47 3993 
Wamieliaritdae: sees. scte we es ae Ae 
Ncimaecidaey ame. aclele sane ers 43 3053 
Ee petidae meee so ecees cares 
Setivel limidaere. tae. cre acusc = aise 
Addisonudae 5.2.00 .s.s.26: 
@occulinidaes... cca ese 
Phasianellidae ............ 
eiturbimidaew.. .oe4 ven <+0s on otis ee 
MTOChIdae any saesia see eee 18 1529 
Delphinulidae ............ 
Cyclostrematidae ......... ee ayes 
Nf tetitst cl aaa ee ieee onsen tee ene 374 2773 
Stomatiidae: wee qantas 
Telia TOT AG a. ss eres weer 
Scissunellidaé: ..4.40%a0 
Pleurotomariidae ......... 
Taissurellidae 2.2...:2..... ee ae 
@hitonidae =.c 226s s2e oo. 26 2200 
Cephalopoda (further uni- 
Gentified)s ves cn ere te 86 7306 
Wolieinidae .22...02665+24- 3 .0254 
Land shells 
Percentage of 
identifications 
in this group 
among those 
of all land 
Mollusca 
Number of (except the 
Family identifications unidentified ) 
Cyclostomatidae .......... 
diruncatellidaey ass. .saeiee wee Bere 
lelicinidde: ..22...0.+..-+ 14 7.9090 
Flelicidae 2..6..e00.2.0205 70 39.5479 


105 


Percentage 
of species 
in this group 
among marine 
mollusks of the 
Southeast 
Coast 


.2516 
.0629 
.5031 
.1887 
.1887 
.1258 
1.5004 
.1887 
3145 
.2516 
.0629 
.0629 
3773 
.1887 
8176 
3.9622 
6289 
8805 
.5031 
.0629 
.0629 
.1887 
1258 
2.2012 
1.4465 


1258 


Percentage 
of species 
in this group 
among all 
nearctic land 
Mollusca + 


5277 
1.0554 
1.0554 

37-7305 


*Compiled from Pilsbry, H. A., and Johnson, C. W., A classified catalogue 
with localities of the land shells of America north of Mexico. Reprinted from 


The Nautilus, 1897-1898, Philadelphia, 35 pp., 1808. 


106 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 . 


Identifications of Mollusca—Continued 
Land shells 


Percentage of 
identifications 


in this group Percentage 

among those of species 

of all land in this group 

Mollusca among all 
. Number of (except the nearctic land 
Family identifications unidentified ) Mollusca 

Bulimultdae ess eee ae mie 2.9023 
Wiocoptidaess yee: Natt sive 4.2216 
Papidae: 23.53 tees ae cranes 42 23.7287 13.1925 
Nchaniti deus I .5049 1.5831 
Glandinidae 25... - ses see- Ae ae 1.0554 
sRestaceliltda@ems. se tase ee ee .2038 
Circinantidacene eee neces Ae 1.8469 | 
Lonitidacweyeencee eee 27 15.2542 17.6779 
Mimacidaciee eee nee eree 2 1.1299 1.5831 
ANOLOMGLKS Goaneaoceaccega6 ae es 3-6939 
Pinlomycidaes ya. I 5649 1.3192 
ndodontidae --4-24442400% 6 3.2808 4.2216 
Succinetdae eee I4 7.9090 5.8047 
Vasinulidae’ 3s. 520.4. see ; an 2638 


The outstanding impression given by the foregoing table is that 
notwithstanding their relatively low availability to birds, mollusks of 
practically all kinds are eaten. In general it is also true that the large 
groups more numerous in species and individuals contribute most 
heavily to avian subsistence, while the small groups of less abundance 
get off with a light toll. 

Let us see what are the relations of birds to some of the protected 
mollusks. In a paper “On the Adaptive Coloration of Mollusca” 
(Proc. Boston Soc. Nat. Hist., vol. 14, pp. 141-145 (1871) 1872), 
Edward S. Morse alludes to protective coloration of several species. 
His remarks with comment deduced from our tabulations are here- 
with presented. “Among the marine forms we notice the adaptive 
coloration of certain species very well marked. The common Littorina 
of the coast swarms on the bladder weed, the bulbous portions of 
which are olive brown in color or yellowish according to age. The 
shells of the Littorina found upon it, present in their varieties these 
two colors and are limited to these colors ” (p. 143). Our tabulations 
show 503 records of capture of Littorina, 11 species of which were 
identified from the stomachs of 46 kinds of birds. These shells are 
freely eaten as the following instances of large numbers taken by 
single birds testify: Pacific eider 110, surf scoter 120, black duck 150, 
purple sandpiper 205, and greater scaup duck 350. 








i 





NO. 7 PROTECTIVE ADAPTATIONS—McATEE 107 

“The few species common to the mud flats exposed by the retreat- 
ing tide are colored black or dark olive.’ Examples: //yanassa 
obsoleta, Nassa trivittata, Rissoa minuta (p. 143). There are 78 
records of Jlyanassa obsoleta distributed among 18 species of birds. 
Thirty-two of these shells were taken at a meal by a greater scaup 
duck, and from 42 to 62 by each of six knots. There are 98 records 
of Nassa trivittata, two of them being 275 and 285 specimens in the 
stomachs of greater scaups; and there are three determinations of 
Rissoa minuta. 

Lacuna vincta: The colors ‘‘ quite match the Laminarian upon 
which they are found ” (p. 143). This species was identified 39 times 
in nine species of birds in which numbers from 32 to 75 were found, 
and in one case, that of a golden-eye, no fewer than 116. 

“ Margarites helicina | have found in numbers on the large Lami- 
narian and on seaweed at low water mark and its color is decidedly 
protective’ (p. 144). Our tabulation shows 10 records of this species 


‘ 


of shell, distributed among five kinds of birds. 

“A very evident case of protective coloring is seen in the three 
species of Crepidula found on our coast. Crepidula fornicata is drab, 
variously rayed and mottled with brown, and it lives attached to 
stones near the roots of the large Laminarian or upon stones clothed 
with algae of similar colors, or attached to the large Mytilus. Crepi- 
— dula convexa, a much smaller species, lives on the roots of seaweed. 

Professor Perkins records its occurrence on the black shell of 
_ Ilyanassa obsoleta, This Crepidula has a very dark brown shell, ac- 
cording well with the dark color of its various places of lodgement. 
Crepidula plana or unguifornus lives within the apertures of the shells 
of larger species of Gasteropods, as Buccinum, Natica, Busycon and 
others. The shell of this Crepidula is absolutely white ” (pp. 144-145). 

All of the limpets named in the foregoing quotation have been 
identified from the stomachs of nearctic birds, and the total number 
of records for species of Crepidula is 135. Fifty specimens of 
C. glauca were found in one gizzard of a greater scaup duck and 60 in 
another. The “ protection” of C. plana is very undependable since 
all of the mollusks named as its hosts are swallowed whole by birds 
and other predatory enemies of shellfish. With reference to a special 
enemy of limpets “ it has been calculated that a single flock of oyster- 
catchers, frequenting a small Scotch Loch, must consume hundreds 
of thousands of limpets in the course of a single year.” (Cooke, 
Cambridge Nat. Hist., vol. 3, pp. 56-57, 1895.) 

As an example of the land snails thought to be defended from some 
enemies by the toothed apertures of their shells, the genus Polygyra 








108 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


may be mentioned. Twelve species have been identified from the birds 
represented in our tabulations, the total number of determinations 
being 42. Slugs were identified three times, but our findings in this 
respect probably are not representative since in Great Britain it is 
said that: “ Every kind of slug and snail is eaten greedily by black- 
birds, thrushes, chaffinches, and in fact by many species of birds.” 
(Cooke, Cambridge Nat. Hist., vol. 3, p. 58, 1895.) With regard to 
highly colored shells, such as Pecten, conjectured by Wallace to be 
protected, it may be said that our table shows 193 records for Pecten 
and the Biological Survey has been called upon to make a special study 
of damage to the scallop industry by wild fowl in the vicinity of 
Marthas Vineyard, Mass. Teredos were identified from the stomachs 
of four Bachman oyster-catchers and of one egret. 

It should be remarked that the very large number of records of 
Ostreidae in the tabulations is due also to a special investigation of 
the bird enemies of Ostrea lurida. The high records for Tellinidae 
(Macoma, especially) and Paphiidae (Paphia staminea) are chiefly 
by-products of this same study. The large numbers of identifications 
for such dominant families as the Mytilidae, Nassidae, Columbellidae, 
and Litorinidae among marine shells and Amnicolidae and Lymnae- 
idae among fresh-water ones, need no explanation. 

Number of identifications, 11,771; percentage of identifications 
among those of all animals, 4.9583; percentage of species in this 
phylum among the whole number of animal species known, 10.8828. 

Other enenies——Mollusks are preyed upon to an important extent 
by very many marine fishes, as well as by most of the rays and sharks ; 
among these being numerous forms specialized (as by possession of 
the pavement-like pharyngeal dentition) to feed upon shell fish. For 
such fishes as the haddock, cod, wolffish, and flounders they are a 
staple food. Field found razor-clams in 4 per cent of 388 stomachs 
of the smooth dogfish (Mustelus canis); and in 3.68 per cent of 
516 stomachs of the summer skate (Raja erinacea). The same author 
found mollusks of various kinds in 17.64 per cent of the stomachs of 
306 cunners (Tautogolabris adspersus), and in 27.2 per cent of those 
of 33 toadfish (Opsanus taw). Some of the marine fishes are known 
to be enemies of certain specially defended mollusks ; as predators on 
Crepidula, the scup, tautog, swellfish and toadfish may be mentioned ; 
upon chitons, the haddock and flounders (Pleuronectes) ; and upon 
Eolis and other nudibranchs, the cods, gurnards and_ flounders. 
Cephalopods, especially squids, are a favorite food of many of the 
highly predacious fishes as the sharks, rays, bonito, swordfish, blue- 
fish, mackerels, pollock, and haddock. It would be easy to compile a 


Sena er 
9 


NO. 7 PROTECTIVE ADAPTATIONS—McATEE LOg 


very long list of squid-eaters. As to the extent to which these cephalo- 
pods are taken, Field reports squid from the following percentages 
of the stomachs examined by him: Summer skate 6.39 per cent, 
smooth dogfish nearly 10 per cent, and goosefish 17.39 per cent. 
Cuttlefishes are known to be eaten by the bonito, cod, whiting, and 
gurnard, and octopods by the ling, haddock, and conger eel. 

Turning to the fresh-water mollusks, we find that they are equally 
beset by enemies. Pearse reports 2 per cent of the total food of 
32 species of fishes in Wisconsin lakes: consists of these animals, and 
from Forbes we learn that mollusks make up about one-fourth of 
the food of the dogfish (Amia) and a sheepshead (Aplodinotus), 
about half that of the suckers (Catostomus), rising to 60 per cent in 
the case of the red-horse (Neorostoma), and a considerable propor- 
tion (14 to 16 per cent) of the food of the perch (Perca flavescens), 
catfishes, sunfishes, top minnows, and shiner (Abramis). Almost all 
fishes eat mollusks to some extent and practically all groups of 
mollusks suffer from these predatory attentions. 

Taking up the relations of amphibians to mollusks, it may be noted 
that Kirkland found 1 per cent of the food of 149 toads to consist of 
snails and slugs, and Drake found 29 of these mollusks in 209 
stomachs of the leopard frog. In general it may be said that most 
frogs consume aquatic snails when in the larval state and land snails 
when adult. With reference to European conditions, Cooke adds: 
“Frogs and toads are very partial to land mollusca. A garden at- 
tached to the Laboratory of Agricultural Chemistry at Rouen had been 
abandoned for three years to weeds and slugs. The director intro- 
duced 100 toads and go frogs, and in less than a month all the slugs 
were destroyed.” (Cambridge Nat. Hist., vol. 3, p. 58, 1895.) Snails 
are eaten by most salamanders, the kind, whether water or land, 
depending on the habits of the salamanders concerned ; small mussels 
even are consumed by some of the thoroughly aquatic forms. 

Reptiles do not prey very extensively upon mollusks, yet snails are 
frequently eaten by lizards; slugs and snails are eaten by several 
species of snakes and by most turtles, the aquatic forms of the latter 
group consuming some bivalves. 

Among mammals we find that some of the land forms consume 
mollusks to a slight extent ; shrews, rats, white-footed mice, squirrels, 
and chipmunks may be mentioned as examples; a specimen of the 
eastern chipmunk (Futamias striatus) taken near Fairfax, Va., had 
packed in its cheek-pouches or swallowed more than 47 Pomatiopsis 
lapidana. It is well known that the muskrat preys extensively upon 
fresh-water mussels, and the mink and otter must be listed as foes of 


1 } 


Ilo SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


fresh-water mollusks. The food of the walrus consists mainly of 
shellfish. Dyche reports that the California sea-lion feeds very largely 
upon squids and octopods and it is known that squids form a con- 
siderable proportion of the diet of sperm and other whales. 

The enemies of marine mollusks include also sea-anemones,. star- 
fishes, and boring univalves of the genera Purpura, Polynices, Thais, 
Lunatia, Natica, Cycotypus, Fulgur, and Urosalpinx. Fresh-water 
mollusks form an important element of the food of dragonfly nymphs, 
and a lesser one of horse fly larvae, water beetle larvae, water bugs, 
leeches, and crayfishes. JIand snails are attacked by predacious 
beetles and fly larvae. Mollusks also have enemies among such para- 
sitic groups as mites, nematodes, and trematodes. 

Discussion—From the abundance of their enemies and from the 
extent to which these predators feed upon mollusks (more than 8,000 
records for birds in our tabulations), it is evident that the possession 
of a shell as a means of defense has been entirely discounted so far 
as predators of any size are concerned. The relations of birds to the 
protectively colored forms show that some of these (Litorinidae) are 
freely eaten; the brightly colored shells (Pectenidae) also are freely 
taken, as well as the very hard and thick-shelled ones (Ostreidae). 
Slugs, snails, limpets, teredos, chitons, and cephalopods pay their toll 
also, testimony to the all-pervading search for food by birds. In fact 
the evidence is that birds feed more or less indiscriminately upon all 
mollusks of suitable size that are available to them. Other enemies 
follow mollusks, especially the marine forms, where most birds can 
not, and it would seem that the whole molluskan world is exploited 
as a source of food to as large an extent as could be expected. 


CHORDATA (LANCELETS, TUNICATES, VERTEBRATES) 


While the Chordata with 13,326 identifications contribute only 
5.6133 per cent of the total determinations of the animal food of 
birds, yet the phylum comprises so many familiar animals that it 
probably will be best to treat it more in detail as was done in the case 
of insects. 

Number of identifications, 13,326; percentage of identifications 
among those of all animals, 5.6133; percentage of species in this 
phylum among the whole number of animal species known, 8.8427. 

A tabulation of the records of Chordata with frequency indices de- 
rived from estimates for the world fauna gives the following results: 









NO. PROTECTIVE ADAPTATIONS—McATEE Gt 


N 


Identifications of Chordata 


Percentage 
of species 
in this class 
among those 


Percentage of of the whole 
identifications number of 
among those Chordate 
Number of of all species 
Class identifications Chordates known 
Wirochorda. .. ..c0. sea. e=s 5 .0375 2.6228 
HEATS COSiersers eretaercistavers cao © 4,92 36.9427 26.2281 
Xm piibia #e .2ss0 see eso 097 7.4816 4.4386 
TED tll ice eerste teeters aiskeiseae ior 6905 5.2153 12.1053 
PNGVGS: a Hisseses tei) atl eis Hit als S05 3,555 20.6771 40.3510 
Mammalian er emcccys s)..<.<iek 3,151 23.6454 14.1228 


The urochordates listed are ascidians, in three cases being identified 
as Boltenia ovifera. While the identifications of urochords is far 
from proportional to the frequency of these animals, the result is 
only what would be expected in view of their strictly marine and 
chiefly submerged habitat in which they are exposed to the attacks of 
only a very small proportion of our birds. It may be noted here that 
tunicates have numerous enemies, however, among fishes which take 
the pelagic and both simple and colonial fixed ascidians. It is on 
record that these animals are not uncommon in the stomachs of cod 
and haddock, and they have been found also in herring, flatfishes, 
tilefish, cunner, scup, the great sunfish (Mola), and a number of 
other fishes. They are taken also by sea-anemones and sea-urchins. 

Omitting the Urochorda and figuring frequency indices from the 
fairly well known numbers * of North American species in the various 
classes we obtain the following table: ° 


Identifications of Vertebrata 


Percentage 
of species 

Percentage of Number of in this class 

a identifications nearctic among those 

among those species of all 

Number of of all in this nearctic 

Class identifications Vertebrata class Vertebrata 
Pisces ....2 4,023 36,0565 3,054 61.3253 
Amphibia .. 907 7.4844 141 2.8313 
Reptilia ... 605 52073 308 6.1847 
PN CS Sted seks! BISGs 26.6870 Sol 16.0843 
Mammalia . 3,151 23.6542 676 13.5743 


In this as in other cases we clearly observe the tendency for losses 
to predators to correspond to the extent and abundance of a group. 
In fishes the largest class are preyed upon the most, but less than their 





‘Counts derived from standard works on the various classes, as noted later in 
connection with the tabulations by classes. 


8 


II2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 





relative abundance would seem to warrant, for the reason that, as a 
group, they are relatively inaccessible to birds, many of the deeper- 
water forms being entirely so. 


Pisces (FIsHEs) 


| 
| 


Protective adaptations.—It has been held that the great group of 
spiny-rayed fishes is protected from enemies to a greater degree than 
the soft-finned families, and in general harsh scales and spines are 
deemed protective. Some fishes have poison glands connected with 
certain specialized spines. Some of the species with disagreeable 
qualities have colors that are said to be warning, while the great | 
majority of fishes exhibit varying degrees of cryptic coloration, many | 
of them having more or less ability to change in color in response to 
that of their environment. Such in brief are some of the more or | 
less theoretical defenses of fishes; as to actual physical protection, it | 
may be said that fishes are shielded from most birds by their aquatic 
habits and many of them even from water birds by their living at | 
considerable depths. 

Bird enemies.—It is well known that whole families of the so-called 
lower orders of birds are specialized to prey upon fishes, for example 
the loons, terns, cormorants, anhingas, pelicans, mergansers, herons, 
and kingfishers. There are special fish eaters in other groups, and 
many birds not at all specialized to prey upon fishes nevertheless con- 
sume them to some extent more or less habitually. Nearctic birds 
which subsist almost exclusively upon fishes include: the western 
grebe, Caspian, royal and Cabot terns, black skimmer, anhinga, double- 
crested cormorant, brown and white pelicans, man-o’-war bird, and 
osprey. Others making fishes from 50 to go per cent of their diet are: 
the common loon, Holboell’s grebe, black, Mandt’s and pigeon guille- 
mots, common and Brunnich’s murre, kittiwake, glaucous-winged, 
herring, and ring-billed gulls, gannet, violet-green cormorant, Ameri- 
can and red-breasted mergansers, bald eagle, and belted kingfisher. 

Nearly 5,000 records of fishes being eaten are contained in our 
tabulations of the food of nearctic birds, and of these nearly half were 
identified no further than the class. The remaining determinations 
grouped by families are listed herewith: 


7 





NO. 7 PROTECTIVE ADAPTATIONS—McATEE 113 


Identifications of Pisces 





Percentage 
of species 

Percentage of of this group 
identifications among 
among those North 

Number of of all American 
Group identifications fishes fishes 1 
Wtdentified ...0606..000 0% 2,253 45.7052 err 
Branchiostomidae ......... : : .I310 
Heptatremidae ........... mie ae .0327 
IMiyxinidaege. ......ce0s6css trys cine 0327 
Petromyzonidae .......... se ae 3274 
Chlamydoselachidae ....... or se .0327 
Elexanchidae ......:.6.0.«- er Ss 0982 
Heterodontidae ........... Sars aes 0055 
Scylliorhinidae ........... toe ae 1965 
Ginglymostomidae ........ ake ave .0327 
Pseudotriakidae .......... Aer ae .0327 
SLEICAG: (on ger avare Miata’ ent aves .9823 
Syria! aa..sc sos siete xneee — Sa .0982 
PUOPNGAS facveee die desde ne na .0327 
amchiaridae <.sck..deeaeales ‘hig we .0327 
IBamnidae wie lace cree se «le spots <a .1637 
Cetorhinidae ............. mar Sears .0327 
Rhinodontidae ..c......: ee oak .0327 
Squalidae’...............- nets ee .1637 
Walatidaee ascent es ecer Sieg ee 0827, 
Echinorhinidae ........... ae aes .0327 
Squatinidae .............. oe Sate .0327 
PP TISHOAC Is. daa tee ate acnaas + ore aa 0655 
Reino DatiGae’ a). siete ee ee ote .2047 
Rand aes a wets See.ccuncweuie rere ae .6221 
WNarcobatidae <...........- or Ae .1310 
Dasyatidae o....... 08000 sn dds a 5239 
Myliobatidae ............. we 2 26019 
Miaritrd aes es. a s00 se00 2s sae aes .0055 
Chimaeridae ............. ere rts .1310 
Polyodontidae ............ mE ee .0327 
Acipenseridae ............ ae .1965 
| Lepisosteidae ............. 5 1016 1310 
| ATHLIGAC! Bias es su vie asso uses I .0203 .0327 
| Siluridae ............0.0.. 100 2.0313 3.1107 
Koricaritdae ac. sse sees ee te 3274 
Catostomidae ............. 62 1.2504 2.1284 
@yprinidaeess.. ssm.< eee 482 9.7909 7.3346 
Prythrinidae. ....3<6.00s.. aes ae .0327 
Characinidae ............. ae at 6221 
RpOdes: id vase ceie vdeo « 2 .0406 


*Computed from Jordan, D. S., and Evermann, B. E., The fishes of North 
and Middle America, etc., U. S. Nat. Mus. Bull. 47, 4 vols., 1896-1900. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS 


Identifications of Pisces—Continued 


Percentage of 
identifications 
among those 
Number of of all 
Group identifications fishes 


Gymnotidae =. -se ee ooueee 

Symbranchidae :...:...... 

Denichtyidaey aa.sscgene eer cae oe 
Anenillidae 252.425. cene. 16 -3250 
Simenchelyidae ........... 
ivophidacmernn cen see ri ee 
Synaphobranchidae ....... 
Leptocephalidae .......... 
Muraenesocidae .......... 
Nettastomidae ...:.....0+. 
INemichthyidde (254. e0..6- 
IW ARCELOE pers otrs Bo auuoeee 
Ophichthyidaewsasssse. eee I .0203 
Miitraentdaer as eeimeeite rier 

Saccopharyngidae ......... 

Eurypharyngidae ......... 

Blopidacey eee scene: 

Aibulidaeieen see eer 

Ehodontidacwessrceee eer 

Ghanidaca.-s see eee 

Dorosomidae 

Glugeidae I Areca eee 249 5.0579 
Bneraulididae a: . 4s. esse SI 1.0360 
Alepocephalidae .......... aan ers 
Salmonidae, 2. sss ne ee cs 198 4.0220 
ehymallida cesar eer rst Se nits 
Argentinidae ............. 30 .6004 
Microstomidae ........... 
Synodentidae jcc ce see 
Ailopidaeiecrsiic seraeree eter 
Benthosauridae ........... 
Bathypteroidae ........... 
Ipnopidacue enone eee 
Rondeletiidae ............. 
@etomimidae’ 3.2 )..2.56..- 
Myctophidaes 2.5 reer: <- 
Maunolicidae, Jc. asses 


to 
° 
HR - 
One 
aN 


.0406 


bo: 


Astronesthidde ........... 
Stomiatidae <5. 2s seeeesee 
Malacosteidae ............ 
Allepisaunidae’ s.0ce sce... 
Odontostomidae .......... 
Paralepididae s/.4.5.42702% 






VOL. 85 


Percentage 
of species 
of this group 
among 
North 
American 
fishes 


0655 
.0327 
.0327 
.0327 
0327 
.0327 
.0982 
.2619 
-4584 
0655 
2292 
1310 
.9496 
.9405 
.0327 
.0327 
£0655 | 
0327 
.0982 
.0327 


1.3097 


8841 
3602 
1.0478 
0055 
-3929 
1310 | 
‘sr 
.1310 | 
0655 | 
"0655 

.0327 

.0327 

0655 

.4080 | 
.0655 
-2047 
0982 
.1965 
.0327 
.1637 
.0327 
.1965 | 





- 





py 


NO. 7 PROTECTIVE ADAPTATIONS—McATEE 


Identifications of Pisces—Continued 


Percentage of 


identifications 
among those 
Number of of all 

Group identifications fishes 
Sternoptychidae .......... 
Idiacanthidae . 
falosauridae ..24...0...+. 
Notacanthidae .......... 
iEmpogenyidae 2.2.00 .56 2.6% 
Byam it aes Acres ziwc els ee ds a8 ee a 
lWimbridae .aacase.s Ss vena: 9 .1828 
NE CIT AC Uet nnd sa Ne etacsniseetasns Be .4460 
IPoeciliidae 2... ..2.2.0..4-- 530 10.7659 
Amblyopsidae ............ ae aoe 
ESO CIA aeemne cuter earsieiiny yas 8 162 
Hemiramphidae ........... 4 O82 
Scombresocidae ...... av a ae 
TX OCOCHGAG kavisisa acs coe se I .0203 
Gasterosteidae ............ ToT 2.2547 
Aulorhynchidae .......... 
Aulostomidae ............. 
astulariidae™ ..25.....220s. 
Macrorhamphosidae..... ss fae 
Syngnathidae ............ 6 1210 
Percopsidae oi... ...6.< . — 
Aphredoderidae .......... I .0203 
Atherinidae .............. 38 .7719 
Mugilidae ................ 18 3656 
Sphyraenidae ............ 
Polynemidae ............. are et 
Ammodytidae ............ 25 .5078 
Bathyclupeidae ........... 
Stephanoberycidae ........ 
Trachichthyidae .......... 
ELV CIAAE! 2. one es: Stasi 
Holocentridae ............ 
OLVEMINIGAC seen. yee sions 
Mullidae ................. Sede 
Scombridae .............. I .0203 
Gempylidae ............... 
Lepidopidae .............. 
Trichiuridae ............. 
Istiophoridae ............. 
Xela t1daes yaa vate es ays senate 
Nematistiidae ............ ae : 
Carangidae ............... 7 .1422 
Pomatomidae ............ 4 0812 
Rachycentridae ........... I .0203 


Percentage 
of species 
of this group 
among 
North 
American 
fishes 


.0982 
.0055 
2202 
-1905 
.0327 
.0327 
.0055 
.1965 
3.8638 
.1310 
.6221 
2292 
.0055 
.6221 
.2292 
0327 
.0327 
0982 
.0327 
1.1460 
.0055 
.0327 
151033 
.5566 
.1637 
.1637 
.1310 
0327 
.0055 
.0327 
-3274 
-4584 
.0327 
2619 
4912 
2202 
.1310 
.0327 
0982 
0327 
0327 
1.0074 
0327 
0327 


FT 


116 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


Identifications of Pisces—Continued 








Percentage 
of species 
Percentage of of this group 
identifications among 
among those North 
Number of of all American 
Group identifications fishes fishes 
Nomeidaer saan leciyeeieesiee see id .1637 
Coryphaenidae -.-.-....-. I .0203 .0055 
ampridae) -... 2d: sls steve sare syle .0327 
Pterachidaey soe ..c cern a sae .0327 
Branndaea aan ss others siete Rahs .1310 
Stemeseniidae ta. =. she. ne cial 0327 | 
Centrolophidae” Gros... 5s. ste oes 0055 
Stromaterdaes meee tiels me siels 1965 
Icosteidacynemsee cere ee wee ne .1310 
Grammicolepididae ........ ee tre 10327 
Metragonunidae te. oe eee e a. oe .0327 | 
Pempheridae’ (:4.2..-21--- oe ae .1310 
Bilassomidacwnssriece: tenes aes Bae .0055 
Centrarchidaeia sacs eens 164 23313 1.0478 
KGQhIMIMGEYS) bossastendoodacs nee ees .0055 
Percidacsn eee eee 171 3.4735 2.8160 | 
Cheilodipteridae .......... : ae .4Q12 | 
Centropomidac) -y...-4-cr ae ae 4257 | 
Serrantdae a. wa. ee eerie = 5 .1016 3.0779 
Ioloyolobi Vansengepoooudoc ae acy. .0327 
Priacanthidae cease eee aoe a .1310 
Mutianidae 4 asec eaene se ne 1.1133 
Elaemtlidae neces sree 2 .0406 1.8009 
Sparidae Ae oanc cee acer 5 1016 7858 
Miaenidae meee cae cien nh See 0655 
Gerridaceae eee eee ter 3 .0609 5500 
Keyphosidae saeeees see ae Rte 3929 
Sciaenidae) Go seeee ace. sees 14 2844 3.5036 
@innhitidae sea-ice eee ae sas .0655 
Embiotocidae =. 2..:.000 ee 7 .1422 5804 
Gichlidaeyeeane eer eee er here er toe 1.8337 
Pomacentridae (2... cee ie ware .9823 
aloridaer ae ds sess eer ne aleve II 2234 1.6044 
Scanidageae sacnensoo eee ste Pre 1.4407 
LADAC asics oc hs a ae ciclo ane ee .0982 
(GEROMEV Goo ogbaccsoomucs ete nee .0055 
Epliippidaein -siierieee crete nrae ee .0g82 
Chaetodontidae ........... a ner 6549 
Lanchdacee sere sys Seis .0327 
Meuthididaeyy.c)s cess ee nes Seay .29047 
Mrtacanthidae: saeco aa ae .0327 
Balistidacmey.arcae eee ; mere 5239 
Monacanthidae ........... 3 .0609 3002 


@stractidaeweawac aciierle ter ae ord: .1310 


nee. PROTECTIVE ADAPTATIONS—McATEE 7 


Identifications of Pisces—Continued 


Percentage 
of species 





Percentage of of this group 
identifications among 
among those North 
Number of of all American 
Group identifications fishes fishes 
Tetraodontidae ........... ee aes 6221 
Canthigasteridae ......... Su ea .0055 
Diodontidae .........6.0.. I 0203 3602 
IMIG OGG oo nec cds kaye em i bes Sar 00655 
SCOMpaenidae sa..s%0. od es aoe Save 2.6523 
Anoplopomatidae ......... Lee .0055 
Hexagrammidae .......... 4 .O812 3274 
(Botticla G see vis eters ensiete cadver ees 188 3.8188 4.2567 
Ramphocottidae .......... on eats .0327 
ENOOM1 AG maeicrser eis ct asei Se eee 2 .0406 1.1788 
Gy clopteridde, mera. ce sm ane Sav. 2047 
{ NEiparididaes.. ase« vol. <6 sive sas 1.2443 
Merrmlidae ec ectes «00 aces oa I .0203 8186 
| Peristediidae ............. Meee oe .1310 
Cephalacanthidae ......... ater oe .0327 
\ Callionymidae ............ Sash ae 1310 
GOBNGAC: .uSee2.u.dneccawe a 10 .2031 2.8487 
Echeneididae ............. ee . aes .2292 
i Malacanthidae ........... See ae 51637, 
| Opisthognathidae ......... setae any 3602 
Bathymasteridae .......... ate Aa 0982 
Chiasmodontidae .......... oe ae .0055 
Chaenichthyidae .......... ayes ne .0327 
| Mrichodontidae®..3.....0.«: ae ae 0055 
Dactyloscopidae .......... ean A327. 
Uranoscopidae ............ 2 .0400 .1637 
Batrachoididae ........... 17 3453 327d 
Gobiesocidae ............. I .0203 S841 
Blenniidae ................ 160 3250 4.5187 
Cryptacanthodidae ........ Boe a .0982 
Anarhichadidae ........... I 0203 1905 
@erdalidac ee eee ae iene .0982 
Ptilichthyidae ............ Bes ane .0327 
Scytalinidae .............. one ms .0327 
| OATCICAG ee acne eee ae ae 1.0805 
Derepodichthyidae ........ oe siipt .0327 
Ophidiidae ............... ae wos 5506 
Lycodapodidae ........... oe 2 are .1310 
BPierasteridae-. 0 < nie seas wees me 0982 
| Brotwhdae’ cb ccusice sha idee ee tie 8841 
| Bregmacerotidae .......... — eas .0055 
Merlucciidae ............. ee ee .0982 
| (Graditdac eee weiter 31 .6207 1.1788 
| Macrounidae ....e-0eee os: ee 3 1.0805 
| 
| 


118 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


Identifications of Pisces—Continued 





Percentage 
of species 
Percentage of of this group 
identifications among 
among those North 
Number of of all American 
Group identifications fishes fishes 
INesalecidacwer ea eo eer ae wa .0327 
rachypteridae s2.4.ceeee Bee ah £0982 
Stylephonidaceen seer ae aC .0327 
Pleuronectidae i 
TER INGEST ay aes 0 60 798 
Soleidae if : 2 3/2 
Lophidaes ee naa oeeae ake ae .0327 
Antennariidae, sas250 4.666) ae a: .4Q12 | 
Ceratiidacnckh sana eceoues tae ne 3274 
Ogcocephalidae ........... ae eee 3274 


Total number of identifications of fishes, 4,923; percentage of iden- | 
tifications among those of all vertebrates, 36.9565; percentage of 
species in this class among those of all nearctic vertebrates, 61.3253. 

Commenting on this table it is obvious that a wide range of fishes | 
is preyed upon, and that the families known to be most abundant in | 
individuals almost invariably are those most extensively consumed by 
the birds. As to the bearing of this data on protective adaptations, we 
see the spined catfishes well represented, more so in fact than the 
equally abundant and only negatively if at all defended suckers. No 
fewer than 36 small catfishes were found in the stomach of a single 
belted kingfisher. The very spiny sticklebacks are eaten enough to 
show that their spines are no deterrent to the attacks of birds; no 
fewer than 150 of these little fishes have been taken from the stomach 
of a great blue heron. Advancing to the true prickly-scaled and spiny- 
finned fishes, we note that Centrarchids (sunfishes, bass, etc.) and 
perches are freely taken. High counts of sunfishes in stomachs are | 
12 in that of a least bittern, 14 in an anhinga, and 18 in a little green 
heron. Twenty yellow perch have been eaten at a meal by the least 
bittern and the great blue heron and no fewer than 25 darters by the 
little green heron. The Cottidae or sculpins often have a highly 
developed armature of spines about the head, but there is no evidence 
that it protects them from birds. Flatfishes (Pleuronectidae) repre- 
sent almost the acme of protective coloration, especially of power to 
simulate the background, but they seem to be proportionally repre- 
sented in our table. One double-crested cormorant had eaten 16 
Symphurus plagiusa. 

The unidentified fishes were distributed among approximately 165 
species of birds to which a considerable number would have to be 
added to give the total number of fish-consuming species. A family 





a 


NO. 7 PROTECTIVE ADAPTATIONS—MCcATEE 119 
of fishes that almost everything “ picks on,’ such as the minnows 
(Cyprinidae), was represented in the stomachs of 44 species of birds. 
Numbers up to 50 of these little fishes were found in stomachs of the 
belted kingfisher and the hooded merganser, and in the case of young 
of the common carp a high count of 106 was made from the stomach 
contents of a glossy ibis. Thirty-nine species of birds are known to 
prey upon the common killifishes and their allies ; and numbers were 
taken from many stomachs, the maximum being 526 from a little 
blue heron. 

Other enemies.—Fishes have no more destructive enemies than the 
predacious element among their own kind. Among the highly preda- 
tory marine forms may be mentioned the dogfishes and other sharks, 
swordfish, bluefish, squeateague, conger eel, and the angler, and among 
fresh-water fishes, the gars, sculpins, trout, amia, pikes, and bass. 
In a study of the fisheries of Buzzard’s Bay, Mass., Field estimated 
that two species of sharks destroy more than 500,000 fishes annually 
in that body of water. Pearse found fishes to compose 12.3 per cent 
of the food of 32 species of their class in Wisconsin lakes. Forbes 
notes that the principal piscivorous fishes of Illinois, those which 
obtain three-fourths or more of their total subsistence from their 
fellow fin-bearers, are Lota, Stizostedion, Esox, Micropterus, Icta- 
lurus, Leptops, and Lepidosteus. Six other species are listed as taking 
from 25 per cent to 65 per cent of fish food. 

Predators devour fishes in all stages, and there are numerous 
special enemies of fish spawn; worst among these are other fishes 
such as the suckers, sculpins, minnows, sticklebacks, killifishes, top 
minnows, and trout. 

Not many enemies of fishes are numbered among our batrachians 
and reptiles, those worthy of note including only the bullfrog, Nec- 
turus and Cryptobranchus of amphibians; the king, garter, and water 
snakes, copperhead, rattlesnake, and cottonmouth moccasin among 
snakes; and the painted terrapin, and snapping and _ soft-shelled 
turtles. 

Some mammals are important enemies of fishes but the number is 
not large; we may mention the raccoon, mink, otter, seals, sea-lions, 
porpoises, and whales. 

The young of fishes especially fall a prey to a variety of insects, as 
the larvae of aquatic beetles and of dragonflies, and to several kinds 
of water bugs and to hydras. Insects also, and crawfishes and leeches 
prey upon the eggs of fishes, and squids are said to be among the most 
destructive foes of adult fishes. Parasites of fishes abound and are 
recruited from the ranks of such diverse groups as bacteria, proto- 


120 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


zoans, cestodes, trematodes, and crustaceans. Fishes are destroyed in 
large numbers sometimes by fungoid diseases ; and enormous numbers 
of them perish by being stranded in pools, overflowed by high tides or 
freshets, which later dry up. 

Discussion.—Forbes in his discussion of the ‘‘ Food Relations of 
Fresh-water Fishes” notes that: “ The soft-finned fishes were not 
very much more abundant, on the whole, in the stomachs of other 
species than were those with ctenoid scales, spiny fins, and other de- 
fensive structures,—an unexpected circumstance which I cannot at 
present explain” (p. 479). The natural comment upon this remark 
is that the fact detailed does not need to be explained, only accepted, 
theoretical bias being cast aside. He goes on to say: “ Only the 
catfishes seem to have acquired defensive structures equal to their 
protection, the predatory apparatus of the carnivorous fishes having 
elsewhere outrun in development the protective equipment of the best- 
defended species” (p. 480). Examining the basis for this statement 
we find that Forbes examined the stomachs of about goo adult or 
nearly adult fishes, and that he found catfishes in five of these 
stomachs; darters were identified only four times, whitefish only 
twice, and round suckers only three times, yet all of these are groups 
which equal or exceed catfishes in abundance. There is no reason 
therefore for saying their defenses are unusually efficient ; from the 
table on p. 113 we see that birds take catfishes in due proportion. 

Some kind of protectedness is claimed for practically every kind 
of fish, yet we see that all groups of them are devoured by natural 
enemies, and where data is available, predation seems to be very much 
in proportion to abundance. This principle is especially evident in 
depredations upon fishes if carried back through the life history of 
these animals; young fishes are more abundant than adults and they 
are greedily devoured by many piscivorous animals; while fish eggs, 
most abundant of all, are sought by a perfect swarm of predators. 
The grand principle of predation proportional to population is well 
supported by the known relations of fishes and their foes. 


AMPHIBIA (SALAMANDERS, Toaps, FroGs) 


Protective adaptations ——All amphibians have skin glands that se- 
crete a slime which some have thought to function partly as a defense. 
Toads in particular, frogs to a lesser extent, possess poison glands 
also, and “‘ experiments have proved that toad poison injected into the 
system will kill any vertebrate, the dose being proportionate to the 
size of the animal.’’ (Dickerson, Mary C., The frog book, p. 17, 


- 





NO. 7 PROTECTIVE ADAPTATIONS—McATEE I21I 


1906.) Some amphibians have warning colors, but it is noticeable that 
the nearctic species having such coloration (certain Ambystoma) do 
not possess especially noxious secretions, while our toads are not at 
all warningly colored. The real defense of most amphibians lies in 
their habits, such as aquatic life, nocturnal activity, and lying in 
seclusion in burrows or under logs and rocks. Most of the species are 
very fecund also. 

Bird enemies.—The extent to which the various families of Am- 
phibia have been identified from the stomachs of nearctic birds is 
shown in the subjoined table. 


Identifications of Amphibia 


Percentage 
of species 
Percentage of of this group 
identifications among 
among those North 
Number of of all American 
Group identifications amphibians amphibians ? 
Unidentified .............. 132 13.2396 
Urodela (further unidenti- 

TEC) oer eat cont aayreroainees esac 124 12.4372 ees 
iINe@cturidae=ssa0.5. sents 4 .4012 1.4184 
Typhlomoigidae .......... sists oe .0709 
Amphuimidae ............ oe oe 1.4184 
Cryptobranchidae ......... — ee £0709 
Salamandridae ............ I 1003 227 
Ambystomidae ........... 16 1.6048 14.8932 
Plethodontidae ........... 8 .802 31.9140 
DSIFEMICAC ssc aiele cesses - a Fett 1.4184 
Anura (further unidenti- 

LIEGE) 5 Maes sts teyercee tks ce sae, 3 4o 4.0120 ae 
Discoglossidae ............ ae oan: .0709 
Scaphiopodidae ........... wae Bie. 2.8368 
Bttonmidaey sees eels ee 60 6.0180 9.9288 
Hylidae ................-- 77 7.7231 14.1840 
Leptodactylidae ........... mene ae 2.8368 
INatidae eee ees esis 535 53.6605 12.0564 
Brevyicipitidae, .... seen cr ie 2.8368 


While the Ranidae are more abundant and accessible to birds than 
most of the other amphibians, even so they seem considerably over- 
represented in the preceding tabulation, a circumstance that is ex- 
plained in part by the fact that greater numbers proportionally of 
the stomachs of aquatic birds have been examined than of any other 


group. 


* Computed from Stejneger, L., and Barbour, T., A check list of North Ameri- 
can amphibians and reptiles, pp. 5-40, 1917. 





I22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 ; 


Fifty-three species of birds are recorded as preying on Ranidae, of 
which the common crow is the most voracious (numbers as high as 
24 and 29 individual frogs being counted in stomachs of this species), 
and has the largest number of records (197). Thirty-four frogs were 
found in the stomach of one little blue heron. 

Among 14 species of toad eaters, the following are most important : 
common crow 16 records, red-tailed hawk 10, red-shouldered hawk 
9g, and broad-winged hawk 9. The most frequent consumer of 
salamanders also is the common crow with 83 records. 

Total number of identifications of amphibians, 997 ; percentage of 
identifications among those of all vertebrates, 7.4844; percentage of 
species in this class among those of all nearctic vertebrates, 2.8313. 

Other enemics.—Fishes occasionally eat the eggs of toads and 
frequently devour tadpoles of both toads and frogs, and the larger 
predacious fresh-water fishes are fond of frogs. The bullfrog 
especially preys upon other frogs and the gopher frog is a special 
enemy of toads. The Anura more or less frequently are cannibalistic 
upon the young of their kind, while larvae of salamanders regularly 
devour their brethren. Aquatic salamanders also eat the eggs and 
larvae of frogs. Snapping turtles, soft-shelled turtles, and alligators 
prey upon frogs, but it is particularly among snakes that the most 
deadly enemies of the Anura occur. The garter snakes and hog-nosed 
snake are especially fond of toads, while snakes in general eat frogs 
and also salamanders. In their account of the “* Snakes of Okefinokee 
Swamp,” Wright and Bishop report that: ‘‘ With the larger snakes, 
the food most generally sought is Anura or Amphibia in general. 
It is par excellence the food of the aquatic snakes, and with these four 
or five species it is usually some species of Rana, though Acris, Choro- 
philus or Hyla may rarely appear as their prey. [qually important 
are frogs in the food of the larger land snakes, five species being 
addicted to them. With these the southern and oak toads (Bufo) are 
easily of first importance, with the tree frogs (//yla) and the narrow- 
mouthed frog (Engystoma) occupying second and third places. In 
fact, these 10 snakes prefer the soft-bodied frogs and toads to any 
other food of the swamp (reptilian eggs not considered).” (Proc. 
Acad. Nat. Sci. Philadelphia, vol. 67, p. 147, Apr. 1915.) 

Among mammals the skunk is known to be fond of toads, and 
coyotes, skunks, weasels, minks, otters, wildcats, and the brown rat 
feed upon frogs. No doubt most of these animals will take sala- 
manders also when the opportunity occurs; the little spotted skunk 
and coyote are definitely known to do so, one stomach of the latter 
animal yielding 15 Ambystoma. The mongoose was found to feed 








NO. 7 PROTECTIVE ADAPTATIONS—McATEE 123 
commonly on toads and frogs in three separate investigations of its 
food habits in Trinidad. 

Leeches prey upon both eggs and young of amphibians and there 
are numerous insects which destroy tadpoles. Such are the giant water 
bugs, backswimmers, water scorpions, predacious diving beetles, and 
their larvae, and the nymphs of dragonflies. Finally, it should be 
mentioned that myriads of amphibian eggs and young perish because 
of the unwise choice by their parents of too temporary bodies of 
water for their egg-laying. 

Discussion—The relations of predators to amphibians throw an 
interesting light on the efficiency of protective adaptations in averting 
the attacks of foes. Clearly the Ranidae or frogs are more preyed 
upon than any other group, certainly much more so than the toads. 
The theorist on adaptations attributes this to the superior special 
defenses of toads, but with no doubt whatever the difference in 
amount of predation on these two groups is a direct reflection of their 
relative abundance. 

If toads really were specially protected, if their so-called defenses 
actually saved them from a certain proportion of predatory attacks, 
should they not increase continually relative to the Ranidae? The fact 
that they do not is the best proof that could be asked that their 
“ special defenses ” do not actually function in nature. In short there 
is no reason to believe in the case of amphibians but that the attacks 
of predatory enemies bear a close relation to abundance and availa- 
bility of the various orders and families. Where a certain group 
appears to have an advantage in escaping certain foes, to a degree, it 
invariably proves that it suffers extraordinarily from attacks of other 
enemies. 


Repritia (TurtLes, LizArps, SNAKES) 


Protective adaptations —Although turtles have the direct defenses 
of their shells, jaws and claws, several of the species have also a 
strong musky odor, and some exhibit warning colors. Numerous 
lizards have cryptic coloration; one of our species is poisonous and 
one has the faculty of changing its color considerably. Many lizards 
drop their tails easily, a device said to aid them in eluding enemies. 
The horned-toads besides their protective coloration have more or 
less prominent spines on the back of the head. Many snakes exhibit 
cryptic coloration and a number of them have offensively odorous 
secretions. Certain serpents practice intimidatory actions and a con- 
siderable number of our species are dangerously venomous. 


124 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


Bird enemies.—Following is a tabulation of the records of Reptilia 
found in the stomachs of nearctic birds. The total number 695 seems 
proportional to the abundance of animals of this group in the United 
States. 


Identifications of Reptilia 


Percentage 
of species 
Percentage of of this group 
identifications among 
among those North 
Number of of all American 
Group identifications reptiles reptiles 1 
Winidentittedit ere oe 21 3.0215 aah 
Crocodylidae .. 5.06. e ee I .1438 .6404 
Lacertilia (further uniden- 

tied) eer anc ee 140 20.1432 Soe 
Gekkontdaemeons mene ee ae Ae 9741 
Kublepharidae) s+. cm. eee war Sais -6494 
lewanidaeweeeateteeeeee 47 6.7624 22.0790 
PMGTAUIOAS san goadacasaeoct 2 2878 2.9223 
Anntellidae Maecenas Pies ie 6494 
Helodermatidae ........... ee Weg 0325 
Nantusiidacw eerie aa ae 1.2988 
Mendae qoecc vse eeertor: 5 7194 4.8705 
Scincidaes cates emceicrr 26 3.7409 4.8705 
Ophidia (further unidenti- 

HEM IES te Meer te Cy aone 250 35.9700 et 
Bipedidaehns tasers ae ees .0325 
Léeposternidae ...........- oe ie .0325 
Leptotyphlopidae ......... Bich ae .6404 
Boidaeenyoascaren ere Bre Mee .O741 
Golubuidacie enone III 15.9707 35.0076 
Elapidae ..... sect tree hecreeaee Stas ait .6404 
Grotalidace sseeseee eee I .1438 5.8446 
Chelonia (further unidenti- 

TEC rack isrs ciseicree exaiere toate 84 12.0859 vie 
Kanosternidae) =e eee oe 2 2878 2.2729 
Chelydridaemeen. sade I 1438 6404 
Mestudinidde=.eesdae sce: 3 4316 0.7410 
Ghelonudae......0.-.40...: ae ae 2.2729 
Dermochelidae ............ Ban we 6494 
eri ony Chtdae. .jesew oitearsl ee I .1438 1.2988 


In commenting on the foregoing table the obvious fact is recalled 
that our birds can hardly assume the role of predators upon turtles 
except in the case of rather small young of these animals. This 
limitation considered, g1 records seems fully as many as could be 


*Computed from Stejneger, L., and Barbour, T., A check list of North 
American amphibians and reptiles, pp. 41-125, 1917. 





ee 








NO. 7 PROTECTIVE ADAPTATIONS—MCcATEE 125 
expected. Forty-five species of birds participated in the lizard-eating, 
including some surprisingly diminutive ones such as the canyon and 
Carolina wrens and the white-eyed vireo. The road-runner, crows, 
jays, butcherbirds, and the Carolina wren took lizards most frequently. 
The chameleon, despite its powers of color change, was identified more 
often than any other species of lizard, namely, 24 times in the 
stomachs of 10 species of birds. One swallow-tailed kite had eaten 
seven specimens at a meal. Horned-toads and swifts, notwithstanding 
their defenses, which as it happens are diametrically opposed in 
character, were “among those present ”’ in the stomachs. 

In contrast to the comparatively wide distribution of the lizard 
determinations, those of snakes were shared by only 26 species of 
birds. Crows, hawks, and owls were the most important of these 
predators; and it is worth mentioning that the little Carolina wren 
again unexpectedly appears in the list. The superlatively cryptic 
green snake (Cyclophis aestivus) was eaten by red-shouldered and 
broad-winged hawks ; the swift racers (Bascanion) by five species of 
hawks and the crow; the desperately bluffing hog-nosed snakes by the 
red-tailed and Swainson’s hawks; the stinking garter and water 
snakes by several kinds of birds; and the redoubtable and warningly 
colored king snakes by the red-shouldered hawk. A great blue heron 
had swallowed a water snake (Natrix fasciatus) slightly over 25 
inches long. The only venomous snake identified in the stomachs was 
Crotalus confluentus from a great horned owl but field observers 
credit another of our birds, the road-runner, with occasional depreda- 
tions on rattlesnakes. 

Total number of identifications of reptiles, 695; percentage of 
identifications among those of all vertebrates, 5.2173; percentage of 
species in this class among those of all nearctic vertebrates, 6.1847. 

Other enemies—While some of the larger predatory fresh-water 
fishes may occasionally devour a young turtle or small snake, actual 
records of the occurrence have not come to hand. The only one of 
our amphibians known to be a reptile eater 1s the bullfrog, which has 
been observed to eat snakes and newly-hatched turtles and alligators. 
Reptiles have numerous destructive enemies among their own ranks. 
Snapping turtles eat snakes; several kinds of snakes eat turtle eggs 
and a few the young; a few species of lizards prey upon other lizards, 
and a number of snakes devour both these animals and their eggs. 
Snakes are the worst enemies of snakes, such species as the racers, 
king snakes, ring-necked snakes, coral snakes, water moccasin, and 
copperhead being conspicuous in this respect. The king snakes are 


126 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


immune to the poison of the venomous serpents and kill them when- 
ever they run across them. Among mammals, skunks, raccoons, and 
bears dig up and devour the eggs of turtles; skunks, foxes, and wild- 
cats eat snakes and lizards ; the badger is known to feed upon tortoises 
and snakes, the coyotes on horned-toads and garter snakes, the opos- 
sum on horned-toads, and ground squirrels and grasshopper mice upon 
lizards. 

Discussion—The reptiles are not a very numerous group in our 
fauna and it would appear that they have natural enemies in due 
proportion. While some of the turtles are monarchs of the waters 
they inhabit when adult, yet their young must run the gauntlet of 
numerous enemies which cut the number down so that there are no 
indications whatever of an increase in the number of these species. 
So it is apparently with all the forms that when adult seem too large 
to have many enemies to fear; they are small and relatively helpless 
in the earlier stages of their life, and it is then that predators do 
great execution. In the class of reptiles, fratricide in almost every 
direction seems to be one of the most important elements of natural 
control. That such control is effectively exercised, the relatively 
stationary character of the reptile population sufficiently attests. 


Aves (Brrps) 


Protective adaptations—Much has been written about protective 
coloration in the bird world, including the nests, the eggs, the sitting 
bird upon the nest, and later the nestlings, the fledglings with their 
special plumages, and extending to the adults of hundreds of species, 
some of which (Anatidae) have a special protective dress, the eclipse 
plumage, during the season when the flight feathers are moulted. The 
ringed plovers of numerous species are said to have ruptive color 
patterns tending to break up the outline of the birds and render them 
inconspicuous. (The phylogenetic significance of this group character 
apparently is ignored. ) 

Crests of birds in some cases are said to be used to frighten their 
enemies, as are various sudden displays of contrastingly colored 
feathers elsewhere. Boldly marked birds of colors held to be warning 
in other classes of animals are numerous and the unusual often in- 
tense and striking coloration of the lining of the mouth of certain 
nestlings is held to be warning in effect. It has even been claimed that 
the color of some bird eggs advertises their low digestibility and that 
they are therefore avoided by all but ravenously hungry predators. 

Bird enemies.—Birds, not content with preying upon animals of 
every class from protozoans to mammals, also draw upon their own 











NO. 7 PROTECTIVE ADAPTATIONS—McATEE 127 


kind to the extent of a fourth (26.6 per cent numerically) of all their 
vertebrate food. The following table shows the distribution to families 
and two more inclusive groups of the determinations that have thus 


Identifications of Aves 


Percentage 


| far been made of birds in the stomachs of nearctic birds. 
Percentage of of species 
identifications in this group 
among those among 
Number of of all nearctic 
Group identifications birds birds 1 
Birds (further  unidenti- 
LOC) ercpasha tie ero is us bees 301 8.4669 
Egg-shell .............0.. 463 13.0327 avr 
Golymbidae sas sos seean ee 6 .1688 -7490 
Gaviidaeime nese ecierciess < sete Live 6242 
PRICIOAE + fta earn an am pias aener4 3 .0844 2.7465 
SLEECOLATNGAC aa 6 specs ee eT .4994 
Waridae: Gaeecmncds asiesne 3 0844 5.2433 
ReynchoOpidaes a. ser. cee eet res ee 1248 
Diomedeidae .........-:.. Ge vee 6242 
Procellaridae .........-.. is are 3.8700 
Phaethontidae ............ ars ae 3745 
Stlidae ieyesec. ate ase) cena aca tendes aol Se -7490 
PNTININO 1CAC cele a saws iets eee ead Se .1248 
Phalacrocoracidae: . oss. ne ses -7490 
elG@CANIGAG tas stc% ce araialenee oe Bae .2407 
Fregatidae ............... oe wa 1248 
PNTIALICAGH wreccisseese-a susteserse stees 26 7314 7.2407 
Phoenicopteridae ......... si ate .1248 
Plataleidae 2... 2ae de.ees te ele .1248 
rdid ae wate twice cause stare Rate joer .4004 
W@icomidde Viste .cms noes ele — .2407 
INT CEIMAE fos csi sie ew es ook 2 10563 1.7478 
Gri da eins Soyo sie Cate ee aor ee 3745 
PXGAIM GAG? ec taciscsaiaesee ee ate 1248 
wall Cla Ge eres creye: aiefevetensue areieiss 19 25345 1.9974 
Phalaropodidae ........... a 0844 3745 
Recurvirostridae .......... 2 .0563 2497 
Scolopacidae ............. 53 1.4908 5.2433 
Gharadnidae 35.....-6.-- 5 .1406 1.7478 
IADMGIZACACS es0cse -s eee oe ce ere 49904 
Haematopodidae .......... eure nets -4994 
Wacanidae . caes cc. 2s css eee ae ae 1248 
Odontophoridae .......... 2 6470 .8739 
Tetraonidae ............:- 26 FATA. 1.7478 
yieisiairil Gale eanerasere eaters ates 265 7.4542 
Meleagridae .............. ne ; ane 





*Computed from Check list of North American birds, prepared by a Com- 
mittee of the American Ornithologists’ Union, 3rd ed. (rev.), 430 pp., 1910. 
9 






128 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


Identifications of Aves—Continued 


Percentage 

Percentage of of species 

identifications in this group 

among those among 

Number of of all nearctic 
Group identifications birds birds 

Gracid tet res ate prss nochanaiters bate beta 1248 
Columbidaeweacc. eee II 3094 1.6229 | 
Cathartidae: .c.-.0< ceoeemiee aia 3745 \ 
Buteomidae, so. dav eoveees ae a 2.8713 
Bal conidaewy sero aaeeiae 14 3938 1.4980 | 
Pandionidae’ 2.52.25 02000: oe ae .1248 if 
Alticonidden®scas2 css satrns: sé eats .1248 | 
Steisidaetsc.asncseos gos 15 4219. 2.3720 | | 
Psittacidaencrcstr. Acts notin tea aes .1248 
Guculidies 0 ee: 6 1688 8730 | 
‘Arogonidae mass. ae we .1248 | 
Alcedinidae 22.044 a et 3745 ‘| 
Picidaenstsat kee SI 1.4346 2.9962 
Caprimulsidae (2..-22..4.. I .0281 -7400 
Micropodidaei-.. 2.402. . 4. 17 .4782 .4004 | 
irochilidac.. 2400.64 eee ae 2.2471 
Cotingidae 2. ke" ee ae sine eats .1248 
Pyraniidae. si... 10 2813 3.8700 
Ataudidae. 32. 1.:b ase 12 3375 2497 | 
Cotvidaeatrss. ere eee 20 5626 2.8713 
Sturnidae ye a eee eee 2 .0563 .1248 
heteridae 22k aber III 3.1223 2.3720 | 
Fringillidae J... 006.602. .2. 992 27.9040 11.6101 
anpaniddes.ete yee ae Il 3004 4904 
Hirundinidae ............. 76 2.1378 1.6229 
Bombyeillidae -.23..5..5 -- 34 .9564 24907 
Ptilogonatidae a. -..0. hs sai nets .1248 
Keanidael es. coseGcs oe I 0281 .2407 
Wareonidaen -. eee 88 2.4754 1.4980 | 
Goerebidae 4. 25). es. ee ust .1248 
Mniotiitidae’ 22 ee. a. 2 488 13.7270 6.8662 
Miotacililidaes {2 -eni eee 7 .1969 8739 
Cinclidach eee eee Soe can .1248 
Mimidae. ta. cc. seve oes oe 45 1.2658 1.3732 
Miroclodytidaes tees 17 .4782 1.7478 
@erthndde ait. Sakae. 4 .II25 1248 
Sittidaenneeoa ees 15 .4219 .4004 
Pakidacmeemnne ea 38 1.0689 1.8726 
Chamacidacen eee I .0281 1248 
Sylvildacwy i eee 25 -7032 -7490 
durdidacaeeces eee eee 243 6.8353 1.8726 


About one-eighth (13.02 per cent) of all the records are for bird 
eggs, and the number of species feeding upon eggs is so considerable 





NO. 7 PROTECTIVE ADAPTATIONS—McATEE 129 


and represents so great a variety of birds (55 species) that the late 
Prof. F. E. L. Beal, taking these facts in connection with his field 
observations, was constrained to express the belief that scarcely a 
species of bird exists that upon good opportunity, can resist the 
temptation to eat another bird’s eggs. Numbers of identifications 
such as 6 for the yellow-billed cuckoo, 10 for the brown towhee, 
12 for the Baltimore oriole, 10 for the California towhee, and 11 for 
the bank swallow prove that egg or at least egg-shell eating is not 
confined to birds of the recognized predatory groups. Probably a 
number of the records are due to birds swallowing bits of their own 
egg-shells. On the other hand eggs may be punctured as by the house 
wren, or eaten without swallowing any of the shell, occurrences not 
likely to be registered in the evidence brought to light by stomach 
examination. 

Of the records for predation upon the various families of birds, it 
may be said that the high number for Phasianidae represents domestic 
poultry almost entirely, and that of the other families, the two— 
sparrows and warblers—undoubtedly most numerous in individuals 
are those which bear the brunt of predatory attack. The rather high 
number of determinations of Turdidae reflect the abundance of the 
robin which contributed nearly 45 (43.6 per cent) of the total. The 
Icteridae, next in line, are birds of great abundance, which might be 
expected to rank still higher among the avian contributors to the 
subsistence of their predatory relatives. However, there is no evidence 
that they are at all immune to attack, as the great flocks of blackbirds 
wintering in our southern latitudes are constantly harried by pre- 
dacious birds in variety and force. 

The sparrows, most persecuted of all, because most available, repre- 
sent almost the acme of protectively colored birds; the bob-whites 
(16 records), ruffed grouse (11), and their allies, also cited, as 
marvels of cryptic coloration are certainly eaten freely considering 
their relative numbers. It is of interest that birds of prey by no 
means spare each other, and it seems that a slight advantage in size 
is all that is needed to induce this strained predation; indeed there 
are records of intra-specific cannibalism. The pugnacious kingbird 
and other members of the family of tyrant flycatchers do not escape ; 
the aerially expert swifts and swallows pay their due toll; and the 
green-coated vireos, best blended with foliage of any of our birds, 
are freely eaten. 

Birds “ warningly colored” that are represented in the dietary of 
other birds as illustrated by our tabulations include the bobolink 
(19 records), Baltimore oriole 4, orchard oriole 3, lark bunting 6, 


130 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85. 


cardinal 3, rose-breasted grosbeak 1, black-headed grosbeak 3, scarlet 
tanager 11, Blackburnian warbler 5, bay-breasted warbler 12, myrtle 
warbler 16, magnolia warbler 16, Canadian warbler 13, Wilson’s 
warbler 10, hooded warbler 1, and the robin 106. These birds cer- 
tainly have the colors and arrangement of colors said to be warn- 
ing in the case of other animals, but brought home in the instances 
of these familiar and practically defenseless species, for none of 
which can any degree of inedibility be assumed, and in the light 
of the fact that all are eaten, some freely, some less so, in relation to 
their numbers, the theory of warning coloration becomes a wraith 
of the imagination so tenuous that one cannot understand why it ever 
received serious consideration. 

Total number of identifications of birds, 3,555; percentage of 
identifications among those of all vertebrates, 26.6870 ; percentage of 
species in this class among those of all nearctic vertebrates, 16.0843. 

Other enemies.—Fishes are not recorded as serious enemies of 
birds, but it is probable that sharks and some other highly predacious 
forms take some toll of birds that rest on the surface of the ocean. 
The goosefish is known to have eaten seven wild ducks at a meal and 
to have attacked such large birds as geese and loons. In fresh-water, 
bass have been observed to capture swallows. (Fins, feathers, and 
fur, p. 8, Dec. 1921.) The bullfrog is the only one of our amphibians 
known to eat birds, but records of its so doing are fairly numerous 
and some of the birds taken are surprisingly large (e. g. woodcock). 

Among the snakes we find very serious enemies of birds, some of 
the expert climbing species especially, making birds, their eggs and 
young a considerable part of their diet. Most noteworthy in this 
respect are the pilot snake and black snake. Other bird-eaters are the 
garter, house, hog-nosed, king, and all of the Crotaline snakes. 

The larger predacious mammals are very fond of birds and must 
be numbered among their worst enemies. Such are the opossum, 
wild cats, foxes, coyotes, raccoon, badger, and skunks. Smaller species 
as the weasels and mink are no less destructive and even the highly 
vegetarian squirrels never lose an opportunity to devour the eggs and 
young of birds. The red or pine squirrels are universally acknowl- 
edged to be among the most destructive foes of birds. The domestic 
cat, large numbers of which lead a more or less feral life, possibly is 
the most deadly single enemy of birds. 

Recently much evidence has been gathered showing that the larvae 
of certain flesh flies (family Muscidae, sens. lat.) parasitize the nest- 
lings of various birds, this activity resulting in the destruction of 


EEE EE LLL eS 


INO. 7 PROTECTIVE ADAPTATIONS—McATEE I31 


numerous broods. Birds have other external as well as internal para- 
sites also, the relation of which to mortality is not well known. An 
occasional bird falls a victim to mussels or other bivalves, to cray- 
fishes, and to mantids and spiders. 

Discussion —“ Warningly colored ” nearctic birds, according to our 
tabulations, are eaten along with the others, the common ones fre- 
quently, the rarer ones to a lesser extent. Our most extensive family 
and the one most numerous in individuals, occupies the logical, if 
unenviable niche, as the most important contributor to the subsistence 
of predatory species. This family, the finches, includes many of the 
most “ protectively colored” species. Fortunately there is other direct 
evidence of the way in which nearctic predators react to protective 
coloration. I refer to Dr. Raymond Pearl’s paper on the “ Relative 
Conspicuousness of Barred and Self-colored Fowls” (Amer. Nat., 
vol. 45, pp. 107-117, Feb., 1911). Natural enemies captured in one 
year 325 individuals out of a total of 3,443, a flock which contained 
both barred and solid-colored fowls. By all theories of protective 
coloration, the latter are the more conspicuous and should pay a 
higher toll to predatory enemies. Of the total number of birds 10.05 
per cent were self-colored and of all the eliminated birds 10.77 per 
cent were self-colored. Thus these monochrome birds were taken 
almost exactly in proportion to their numbers in the whole flock. This 
is precisely the result that would be expected by those who have 
learned by study of the subject that availability is the one strongest 
factor in choice of food by predators. With availability as the con- 
trolling factor it follows that in the long run, and on the average, 
losses to predators will be very closely in proportion to the relative 
abundance of the group concerned. 


MamMMatiA (MAMMALS) 


Protective adaptations.—Many of the mammals are conceived to be 
very perfect exemplifications of protective or cryptic coloration. A 
few are credited with noxious qualities, accompanied in the case of 
the skunks only, in our fauna, by warning coloration. The short dense 
fur of moles and shrews is said to be a deterrent to predators and 
these animals are thought to be protected by a strong musky secretion 
also; shrews have even been credited with poisonous bites. However 
the most potent defenses of mammals in general against birds are 
their large size, and their teeth and claws. 

Bird enemies.—Despite the size and direct means of defense of 
many species, mammals pay a heavy toll to bird predators. In our 





132 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


complete tables, the group without doubt is over-represented, owing 
to the fact that stomachs of the hawks and owls have been kept 
examined practically up to date. However this fact probably does not 
materially affect the relative numbers of identifications for the differ- 
ent families, as shown in the following table for mammals alone. 


Identifications of Mammalia 


(Land mammals only) 


Percentage of Percentage 
identifications of nearctic 
among those species 
Number of of all in this 
Group identifications mammals group 1 
Further unidentified (in 

many cases carrion).... 331 10.5046 
Carrion (identified to spe- 

GIES) is ites, Serle eee oes 18 5712 nee 
Didelphtidae” fn aca. eee a 2 0635 2058 
Malpidaeden wi cen ee eee 22 .6982 1.4790 
Sortcidaee eee eee ee 274 8.6957 6.5076 
Phyllostomidae ........... no os 2958 
Vespertilionidae .......... 19 .6030 3.5490 
Molossidaesnaatceie eee aM ae 1479 
Wirsiddeern a ume roe Ot: 3.2538 
(Grint Saunoweansecodsnds I 0317 5.6202 
iRrocyonidaers: = sania. s aa: wee ae 5916 
Miistelidaew esas. cre eee I 0317 8.5782 
Heldaents tose sa eee eer I 10317 f 
Rodentia (further unidenti- 

fied)\\ Ge aatcnen inti cr 86 2.7203 See 
IMiuridae) ses assent eect 1,816 57.6326 24.1077 
Geomvyidaeesc cee mates 31 .9838 7.9866 
Heteromyidae ............ 16 .5078 8.5782 
Aapodidaewierrieen rece I4 4443 1.7748 
Enithizontidae: 2.54... « ee a 2058 
Aplodontidaey 22.5. ae saves I .0317 7395 
Sciunidaean nena ee ec ote 173 5.4903 14.6421 
Petauristidae secs 15 .4760 -7395 
@astoridae iyce. scat ene oe oe 2958 
©chotonidaen... sense sone Sie seus 1.7748 
ILNSOGES Ao oadodasoocoone 330 10.4729 2.9580 
Dasypodidacueias alos: age vide 1479 
Mayassuidae .n.a7ceceeen ee are Sree .1479 
Gerviddewt yt. sate cere aye aaah 3.9933 
Antilocapridae ............ ae Ate .1479 
Boviddersueerni-cscrcseue te ae esr 


*Compiled from Miller, Gerrit S., Jr., List of North American land mammals 
in the U. S. National Museum, 1911, U. S. Nat. Mus. Bull. 79, 455 pp., 1912. 








NO. 7 PROTECTIVE ADAPTATIONS—MCcATEE 133 


Let us now take up some of the groups of interest in the order of 
their appearance in the tabulation. From the large number (274) 
of records for shrews it would appear certain that the alleged special 
defenses of these animals are no protection against birds. Thirteen 
species of shrews were identified in the stomachs ; 27 species of birds 
are known to prey upon our common short-tailed shrew and 23 upon 
unidentified species of Sorex. Shrews are by no means gregarious, 
nevertheless five specimens of Sorex personatus were taken at a meal 
by a great gray owl. Considering their almost exclusively under- 
ground life, moles were captured fully as often as would be expected ; 
the number of species of birds preying upon them is 12. 

Bats, again on account of their nocturnal activity, are not greatly 
exposed to the attacks of birds. Six predators upon them are recorded 
in our tabulation with a total of 19 identifications. While the Mus- 
telidae are provided with unusually strong musky scents, they are also 
rather above the size for many birds to attack. The single determina- 
tion in our table, attributed to a crow, might perhaps be more correctly 
added to the records of carrion. Skunks, of this family, customarily 
cited as examples of animals having noxious qualities and warning 
coloration to advertise them certainly are too large for all except a 
very few species of our raptors to conquer. However there are a 
number of published and other records of the great horned owl 
preying upon skunks. 

Muridae (mice and rats) are secretive, elusive animals with what 
would be called highly protective coloration, but this does not prevent 
their being the staple mammal food of birds. Meadow mice, perhaps 
our most ubiquitous rodents, are eaten by the largest number of 
species of birds, namely 44. Twenty-six species of birds are known 
to feed on the house mouse and 35 upon deer mice (Peromyscus). 
We have records of five species of birds preying upon our largest 
member of this family, the muskrat, and eight upon the smallest 
(Reithrodontomys ). 

Pocket gophers, like the moles, spend most of their lives under- 
ground and this fact limits the opportunities of birds for capturing 
them, yet there are 31 records for 11 species of birds; nocturnal and 
burrowing habits shield also the pocket mice and kangaroo-rats. 
Captures in these groups probably are in proportion to their reduced 
availability. Jumping mice (14 records), a more diurnal group, seem 
to be proportionately represented. 

Erithizontidae (porcupines) are entirely beyond the size of prey 
practicable for birds, though possibly some of them are captured 


134 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


when young. Mountain-beavers also are rather large and are inacces- 
sible to any but owls, of which the great horned owl contributed our 
only record of their being eaten. 

The large number of determinations of members of the squirrel 
family, cover, it must be recalled, such diverse groups as the spermo- 
philes, prairiedogs, groundhogs, tree squirrels, and chipmunks. There 
are only two records of the groundhog, a very large rodent, one being 
captured by a goshawk and the other by a golden eagle. The number 
of identifications (15) of the chiefly nocturnal flying squirrels seems 
as large as could be expected. The cryptically colored rabbits are 
exceedingly common and live fully exposed to predacious birds, fac- 
tors which go far toward accounting for the very large number of 
records of their being eaten. The remaining families in the list all 
consist of animals so large that only a few of the most formidable 
birds can prey upon them, and then only upon the young. There are 
observations of such occurrences, but it so happens that our records 
of stomach contents do not include any of them. 

Total number of identifications of mammals, 3,151 ; percentage of 
identifications among those of all vertebrates, 23.6542; percentage of 
species in this class among those of all nearctic vertebrates, 13.5743. 

Other enemies——Fishes have few opportunities to capture mam- 
mals, but trout have been known to feed upon meadow mice and 
lemmings, and it is probable that other highly predacious fresh-water 
fishes occasionally get small mammals that venture near or in the 
water. The bullfrog is the only one of our amphibians known to eat 
mammals, an occasional mouse falling to its lot. The snapping turtles 
also get some mice and sometimes even capture animals as large as 
rabbits. Among snakes we find many habitual predators upon mice 
and other small mammals. Some results of studies of the food of 
snakes by the Pennsylvania Division of Zoology may be briefly cited: 
Pilot snake, mice 22 per cent of the diet, squirrels 11 per cent, weasels 
4 per cent; black snake, mice 26 per cent, rabbits 4 per cent, other 
mammals 7 per cent; milk snake, mice 71 per cent, other mammals 
II per cent; copperhead, mice 41 per cent, shrews 4 per cent, other 
mammals 8 per cent. In the case of the timber rattlesnake, mice, rats, 
and rabbits composed nearly the whole diet. This is known to be true 
also of most of our venomous snakes, 

The worst foes of mammals, however, are their own kind, and the 
diversity of their predatory habits may be indicated by brief refer- 
ences to their mammal prey. Opossums take limited numbers of small 
mammals, while raccoons and skunks prey more extensively upon 








yy 


NO. 7 PROTECTIVE ADAPTATIONS—McATEE 135 


them, especially upon mice and ground squirrels. The bob cats or 
lynxes are fond of mice, ground squirrels, rabbits, and other rodents, 
occasionally prey upon small domestic stock, and are known to eat 
skunks and porcupines. The mountain-lion specializes upon deer, but 
eats a variety of wild mammals, including foxes, skunks, coons, porcu- 
pines, and bob cats. House cats take mice, rats, moles, shrews, and 
rabbits. Coyotes and wolves prey upon the young of deer and 
domestic stock, and upon prairie dogs, spermophiles, and other small 
rodents. On the bill-of-fare of our various species of foxes are shrews, 
mice, ground squirrels, pocket gophers, kangaroo-rats, and rabbits. 
Badgers also take all of these mammals and in addition, prairiedogs 
and mountain-beaver. The black-footed ferret is a special enemy of 
the prairiedog, and relishes rabbits also. Weasels are ferocious 
enemies of small mammals in general, and for their size, shrews are 
fiendish predators. They commonly overpower and devour other 
shrews and mice of their own or even of slightly greater bulk. The 
polar bear preys especially upon seals, and the killer whales also 
destroy these animals, as well as wearing down and devouring the 
largest of all mammals, whales. 

Discussion.—Limitations due to relative sizes allowed for, we see 
the same phenomenon in the case of mammals as in those of other 
elements of bird food, namely that the more available (this usually 
meaning abundant) groups are preyed upon most extensively, while 
those which are less abundant or whose habitat is somewhat out of 
the domain of birds are not so often captured. We see that the 
burrowing moles and pocket gophers escape with moderate losses, but 
that the abundant mice, and the both common and relatively easier 
found rabbits suffer severely. It is evident also that the mammals 
outside the range of prey of birds have serious enemies, chiefly other 
mammals; and it is further evident that, taking all mammal enemies 
into consideration, they are most numerous in the case of so abundant 
and ubiquitous a group as the mice, and proportionally less numerous 
for other less abundant families. 


DISCUSSION 


Availability is a mighty factor in the choice of food by birds. 
Within the limits imposed by special habitats, bodily modifications, 
and the relative sizes of predator and prey, birds are prone to feed 
upon what is abundant and easily obtained. Not only is this very 
natural procedure the everyday order, but it is conspicuously exempli- 


136 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


fied by the characteristic flocking * of birds to the scene of insect out- 
breaks or of other occurrences of unusual abundance of food. 

Constant seeking of the available leads to a wide distribution of 
predatory attack because of seasonally or otherwise variable abundance 
or availability of many of the food organisms, further on account of 
the greater or lesser restriction of predators to specific habitats in 
each of which the range of food items is different, and because of the 
specialization of various predators in methods of seeking food. 

That the predatory attacks of birds are amazingly distributed over 
the entire animal kingdom, preceding pages bear witness. If it be 
asked whether birds eat bats or moles, flyingfishes or hermit crabs, 
dragonflies or mole crickets, sea-urchins or bryozoans, the ariswer is 
ever in the affirmative. Given an animal group comprising only a 
small number of species we find that there are only a few records of 
birds preying upon it. Given one of large numbers of species we 
invariably find it is an important item of bird food. If the validity 
of depending upon the number of species as an index of frequency 
be questioned, no matter. The tendency for feeding to be distributed 
over the whole range of the available food organisms and in at least 
rough proportion to the known abundance of the various groups, is 
beyond dispute. 

This principle, predation in proportion to population, stands out 
clearly in the tabulations of the animal food of nearctic birds here 
presented and discussed. Compared to it the effect of the so-called 
protective adaptations on character of food is negligible. If these 
adaptations controlled choice of bird food to a significant extent, 
discrimination would everywhere be evident ; finding indiscriminancy, 
on the contrary, we must conclude that the ruling criterion in choice 
of food is availability. 


INDISCRIMINANCY OF PREDATORS OTHER THAN Brrps 


Nearctic birds, as a group, are little influenced by the protective 
adaptations of available prey. Let us see what can be said of other 
classes of predators. 

Odonata.—In a general article on “‘ Predacious Insects and their 
Prey,” Prof. E. B. Poulton says of a tabulation of dragonfly victims : 
“ Short as it is, the list is extremely interesting, and raises the expec- 
tation that dragonflies will be found to prey rather largely upon 


* American instances are summarized in the following paper: McAtee, W. L., 
The rdle of vertebrates in the control of insect pests, Smithsonian Rep. 1925, 
PP. 415-437, 7 pls., 1926. 





NO. 7 PROTECTIVE ADAPTATIONS—McATEE 137 


specially defended groups of insects.” (Trans. Ent. Soc. London 
1906, p. 401, 1907.) 

Agnatha, Plecoptera, et al—In a report which deals with the food 
of stoneflies, mayflies, caddisflies, and Diptera in trout streams, Mutt- 
kowski and Smith state that “Aquatic insects in rapid streams are 
opportunists as regards food and eat whatever becomes available.” * 

Orthoptera—Professor Poulton in the paper referred to says of 
the prey of these insects: “ The proportion of specially protected 
forms was very high.” (Op. cit., p. 408.) 

Rhynchota—Quoting from Poulton again, he says of bugs, “ So 
far as it is possible to judge from the ... . table it appears that 
Hemiptera will prove to be extremely dangerous foes to the specially 
protected groups.” (Op. cit., p. 403.) 

Diptera.—Writing of the food of the larvae of aquatic midges, 
A. L. Leathers says:* “ The organisms found were so similar, both 
in number and variety, to those available in a given locality that there 
seemed to be little or no sorting in their method of feeding.” 

Professor Poulton remarks on robber flies that “A study of the 
table at once shows that the Asilidae are most indiscriminate in their 
attacks. The stings of the Aculeates, the distasteful qualities of 
Danainae and Acraeinae, and of the odoriferous Lagria, the hard 
chitinous covering of Coleoptera, the aggressive powers of Odonata, 
are alike insufficient protection against these active and voracious 
flies.” (Trans. Ent. Soc. Lond. 1902, p. 336.) 

Parasites —“ Certain species and groups of species [of insects | 
.... have, as far as we know, relatively few parasites in any 
ReSiON: « . . « This is sometimes considered to be due to the posses- 
sion of protective devices of certain kinds, but the explanation is not 
satisfactory. Neither systems of colorations, nor nettling hairs, nor 
an armour of chitinised plates, nor rapidity of movement, nor the 
existence of toxic principles in the blood prevent insects from being 
decimated by parasites.” * 

Miscellaneous insects —‘* Many groups of predacious insects also 
appear especially to attack the conspicuous, easily-captured prey pro- 


vided by the groups with warning colours. This has been observed in 


*Muttkowski, R. A., and Smith, G. M., The food of trout stream insects in 
Yellowstone National Park, Roosevelt Wild Life Ann., vol. 2, no. 2, p. 261, Oct., 
1929. 

* Bull. U. S. Bur. Fisheries, vol. 38, Doc. no. 915, p. 3, 1922. 

*Thompson, W. R., On natural control, Parasitology, vol. 21, no. 3, p. 279, 
Sept., 1929. 


138 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 = 


the case of the predacious asilid flies, Dragonflies, Hemiptera, Man- 
tidae, and Locustidae.” (Poulton, E. B., Essays on evolution, p. 318, 
1908. ) 

Arachnida.—* Spiders are for the most part not very particular as 
to the insects they catch.” (Bristowe, W. S., Proc. Zool. Soc. Lond., 
1929, p. 643.) 

“Tt is quite probable . . . . that certain species of spiders, together 
with Mantides and other predacious insects, will be found to be among 
the chief, perhaps the chief non-parasitic enemies of aposematic 
insects.” (Poulton, E; B:, Trans. Ent: Soc: Lond: 1902) ps 3278) 

Pisces —‘ In general, fish are opportunists as far as their food 
is concerned. They eat what animal food is available, regardless of 
the origin.” * 

The closeness with which the brook trout is guided by availability | 
in its choice of food is indicated in the following table by Dr. P. R. 
Needham based on studies near Ithaca, N. Y.: 





Comparison of Available Aquatic Fish Foods in Stream Bottoms and Aquatic 
Foods Consumed by Trout * | 











Available aquatic | Consumed aquatic 

foods oods | 

Order | 

Number Per cent Number Per cent | 

| 

Mavyitlya nymphs) asec oct | e236 30.90 | 350 30.12 | 
Caddisfly larvae and pupae....... 1,335 Pieyp | 528 44.07 
Stonefly, nymMps gos. seen osureeeel 921 14.67 4I 3.47 
Fly jarvae, and! pupae.-secececsse | 869 13.84 | 187 15.82 
Beetleslarvacecm-nce usta aac | 476 TES all 33 2.79 
Crayfish and ‘shrimps... «220.0% «1. | 235 20 Aas 14 1.18 
Miscellameousys-nicntce se nese ere | 125 1.99 23 1.04 
MOtalstase rreneesa.giinaca tomes fates 6,277 99.98 1,182 99.99 

















1 Quantitative studies of the fish food supply in selected areas, Suppl. 18th Ann. Rep. 
New York Conserv. Dep. 1928, p. 227, 1929. 

Further testimony to the effect of availability on the food of fishes 
is contained in Muttkowski’s study * of “ The Fauna of Lake Men- 
dota” in which he shows that insects form about 60 per cent by bulk 


*Muttkowski, R. A., The ecology of trout streams in Yellowstone National 
Park, Roosevelt Wild Life Ann., vol. 2, no. 2, p. 229, Oct., 1929. 

*Muttkowski, R. A., Trans. Wisconsin Acad. Sci. Arts and Letters, vol. 19, 
pp. 374-482, 1918. 





NO. 7 PROTECTIVE ADAPTATIONS—McATEE 139 


of the macrofauna of the lake and about that proportion of the total 
diet of the fishes. 

Some of the above remarks tending to emphasize feeding on pro- 
tected forms are special pleading because their authors felt under the 
necessity of proving “ protected” insects do have enemies. The 
various groups of predators thus referred to, however, prey upon 
other than the specially protected insects, just as birds do, and ex- 
amined in that light, the comment “ indiscriminate”? would in most 
cases fit their food habits. An adaptation of Poulton’s tabulation for 
robber flies illustrates the point. 


Percentage 
of species 
in this group 
among the 
whole number 


Number Percentage of insect 
Name of prey of records of records species known 
@rthopteratesss- ees ese 13 5.70 2.04 
ny tehOta yaa cere eee ace ee 12 5.26 8.58 
Neuropteroidea .......... 7 3.07 eit 
Iepidoptera: s..c.s. cess 32 14.03 15.61 
Coleoptera “i.e. .ceee ess os 40 17.54 46.20 
Dipteraerccmt cme eemeetes 57 25.00 11.44 
Hymenoptera ............ 67 29.34 17.17 


This does not look very different from tabulations for birds, and 
clearly illustrates the same propensity demonstrated for that class, 
namely of preying largely upon the groups most numerous in species, 
and presumably therefore in individuals—in other words upon what 
is most available. 

That availability does largely govern choice of food is the very 
thing that creates problems in wild life economics. When man invades 
the domain of wild life and in the various phases of his husbandry 
makes available large supplies of new foods, they are immediately 
attacked and up to a certain limit the enemies steadily increase in 
variety and abundance. It is needless to cite examples of this uni- 
versal phenomenon from the vegetable kingdom. All may not realize, 
however, that it prevails also in the animal world. The temerity of 
the pioneer establishing an orchard in a clearing in the foothills where 
the crop is largely harvested by wild life, is paralleled by that of the 
sheep raiser who grazes his herds in mountain meadows where they 
are attacked at once by wolves, coyotes, wild cats, bears, and other 
predators. Man’s taking the domestic fowl wherever he goes furnishes 
material for further demonstration of the supreme influence of 
availability. The chicken, a native of Asia should have no * natural 


140 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 
* in other parts of the world, nevertheless in America for 
instance, a large number of predators, including hawks, owls, crows, 
jays, skunks, weasels, and foxes eagerly welcomed the new food. 


enemies ’ 


More THEORETICAL ASPECTS OF INDISCRIMINANCY BY PREDATORS 


The experience we have when we place inviting food supplies in 
abundance before the birds indicates what must happen in nature 
under similar circumstances. If we imagine a world of food available 
to predators we must realize that the elements composing it will be 
utilized very much in proportion to their abundance. This is only 
what would be expected if there is or ever was such a thing as the 
oft-mentioned “ balance of nature.” To preserve a balance, natural 
checks must be in proportion to population. If they were not appar- 
ently so, no balance would have been observed and the term balance 
of nature would never have been invented. 

Distribution of predation in proportion to population also is what 
we should expect if the theory of adaptive radiation, or the occupation 
of every possible ecologic niche is correct. Given the world of prey 
to exploit it is inevitable that predation will extend in all possible 
directions. No source of food will be left untouched if by any possi- 
bility it can be drawn upon. Under so searching a campaign for food 
each inevitably will be utilized in proportion to its abundance. 

That this principle actually is at work is well shown by a series * of 
studies by Harry B. Weiss which indicate that regardless of locality 
there is a more or less fixed set of ratios between types of food habits 
of insects. Thus from several widely separated areas the insect popu- 
lation groups into from 45 to 55 per cent of phytophagous species, 
from 15 to 27 per cent of saprophagous, from 14 to Ig per cent 
harpactophagous, from 10 to 12 per cent parasitic, and from 1 to 
4 per cent of species of miscellaneous feeding habits. 


“Insect food habits and vegetation. Ohio Journ. Sci., vol. 24, no. 2, pp. 100- 
106, Mar., 1924. 

Ratios between the food habits of insects. Ent. News., vol. 35, no. 10, pp. 
362-364, Dec., 1924. 

Notes on the ratios of insect food habits. Proc. Biol. Soc. Wash., vol. 38, 
pp. 1-4, Jan., 1925. 

Insect food habit ratios on Quelpart Island. Psyche, vol. 32, no. 2, pp. 92-94, 
Apr., 1925. 

The similarity of insect food habit types on the Atlantic and Western Arctic 
Coasts of America. Amer. Nat., vol. 60, no, 1, pp. 102-104, Jan.Feb., 1926. 








INO: 7 PROTECTIVE ADAPTATIONS—McATEE I4I 


Weiss’ table summarizing these interesting data is substantially 
quoted as follows: 











| I eee Har- | | Patten 
Nai ae pee pacto- | Para- feeders, 
| her | Paag phag phag- | sitic. | misc. 
| ber of | ous. | ous. ous. Per | species. 
| species Per | Per Dep cent | Per 
cent cent | cent cent 
| | | | 
Quelpart Island .............. S771” se. Oe oes 19 10 | I 
Western Arctic Coast of N. A.| 400} 47 27 14 10 2 
SSrateror IN. Dione seen. eiisiielen | 10,500! 49 | I9 16 12 4 
Dtatevor Connl........s..000%5 6,781 | 52 19 16 10 3 











The similarity of the figures whether for the States of New Jersey 
or Connecticut, for the Pacific Coast of Arctic America, or from far 
flung Quelpart Island (Corea) shows that there is at work some 
principle controlling choice of food that overrides whatever effect the 
so-called protective adaptations may have. 

It is almost certain that the constancy of the ratios is due to a 
tendency (one might well say a compulsion) toward distribution of 
predatory and parasitic attack. This distribution is one that lays every 
group under tribute, that takes toll from each so long as the tax is 
more easily collected there than elsewhere, but when that condition 
fails turns toward more easily available supplies. 

Predation is thus kept proportional to population and_ practical 
indiscriminancy as to factors other than availability must result. 


INDISCRIMINANCY OF NATURAL CHECKS OTHER THAN PREDATORS 


It will not be questioned, we believe, that from the standpoint of 
protective adaptations such checks as parasites, bacterial and fungal 
diseases, heat, cold, and other climatic factors, are indiscriminate in 
action. 


RELATIVE IMPORTANCE OF NATURAL CHECKS 


Years ago Chittenden writing of “Insects and the Weather ”' 
stated: “ It also appears to me what has been observed by Mr. Marlatt 
in the case of scale insects . . . . is true in general, viz., that favor- 
able or unfavorable climatic conditions are of greater importance in 


* Bull. 22, n. s., U. S. Div. Ent., pp. 51-64, 1900. 





142 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


determining the abundance or scarcity of insects as a whole than are 
other natural checks such as parasitic and other enemies, or even 
fungous or bacterial diseases ’’’ (p. 63). 

Recent studies have only crystallized long-held views to the effect 
that the grand overwhelming factors of insect control are climatic.’ 
Thus Uvarov in discussing “ Weather and Climate in their Relation 


2 


to Insects,” * says: 


Apart from the seasonal rhythm in the appearance and activities of insects, 
there is a more or less strongly marked periodic fluctuation of a species from 
year to year. Only relatively few insect pests are equally numerous and injurious 
every year, while most of them are practically negligible, except in certain 
years, when mass outbreaks occur. It would be out of place to discuss here 
all the causes for these periodic fluctuations, but I would like to point out that 
recent researches in this direction tend to throw some doubt on the commonly 
accepted idea that the chief controlling factor is the parasites, since a number 
of cases have become known in which the factors normally keeping an insect 
species down are almost entirely of meteorological order. This has been ad- 
mitted for the cotton boll weevil in America (Hunter and Pierce, 1912), for 
the corn-borer in Europe (Thompson and Parker, 1928), for the almond sawfly 
in Palestine (Bodenheimer, 1928), for the cotton seed bug in Egypt (Kirk- 
patrick, 1923), for plague fleas in India (Hirst, Rogers), for vine-moths in 
Europe (Stellwaag, 1925), and for some other notorious pests. 


Again Bodenheimer in answering * the question “ Welche Faktoren 
regulieren die Individuenzahl einer Insektenart in der Natur?” states 
that parasites, predators, and scarcity of food, are rarely or only 
secondarily of regulatory significance, but that climatic factors are 
the real controlling influences. 

Accepting the great superiority of meteorological phenomena as 
regulative factors we may make some inquiry as to the relative im- 
portance of other controlling agencies. Diseases sometimes are 
dramatically destructive, but they rarely have a steady regulatory 
influence. 

Among parasitic and predacious organisms it must be presumed, 
except for specific limiting factors, that their effectiveness as control 
agencies will be more or less in keeping with their total numbers. 
Thus we can deduce from a table such as that on page 9 that most 


*This statement has general validity, for insects are nine-tenths of the 
terrestrial animals above the size of nematodes, and probably a large proportion 
of the smaller animals, as well as part of the tenth of larger size are subject to 
similar checks. 

* Uvarov, B. P., Conference of [British] Empire meteorologists, 1929, Agri- 
cultural Section, pp. 17-18. 

* Bodenheimer, F. S., Biol. Zentralbl., vol. 48, pp. 714-739, 1028. 








NO. 7 PROTECTIVE ADAPTATIONS—McATEE 143 


of the groups can play only minor roles in the whole drama of pre- 
dation, and that insects must occupy the center of the stage, regardless 
even of the superior individual size of the chordates. 

To put the case in other language we may quote from David Sharp,’ 
“Tnsects form by far the larger part of the land animals of the 
world; they outnumber in species all the other terrestrial animals 
together, while compared with the vertebrates their numbers are 
simply enormous ” (p. 83). 

“Insects derive their sustenance primarily from the vegetable king- 
dom. So great and rapid are the powers of assimilation of the Insect, 
so prodigious its capacity for multiplication, that the mammal would 
not be able to compete with it were it not that the great horde of six- 
legged creatures has divided itself into two great armies, one of 
which destroys the other” (p. 521). 


SUMMARY 


The hypotheses about protective and warning colors and mimicry 
are part of the Natural Selection group of theories. These coloration 
phenomena and other protective adaptations are supposed to have been 
developed and perpetuated by the selective value they had in shielding 
their possessors from attack by predators. 

Preceding sections of this discussion call attention to the evidence 
that one group of predators after another is known either to prey 
habitually upon “specially protected” groups, or to be so largely 
guided in choice of food by availability as practically to ignore pro- 
tective adaptations. 

The former is admitted to be true of dragonflies, robber flies, 
mantids, predacious locustids and Hemiptera, parasitic insects, and 
of spiders, while the latter is stated to be characteristic of the aquatic 
immature forms of mayflies, stoneflies, caddisflies, and two-winged 
flies, and of fishes. Data cited throughout the main body of the 
present paper show a high degree of indiscriminancy also on the part 
of amphibians and reptiles. 

In fact this general indiscriminancy on the part of predators is so 
evident that even ardent advocates of the selection theories have been 
impressed by it and one of them, G. A. K. Marshall, in a paper on 
the “‘ Bionomics of South African Insects ” says: 

If the view advocated by many, that birds cannot be reckoned among the 
principal enemies of butterflies in the imago state, be true, then I consider that 


we may practically abandon the whole theory of mimicry as at present applied 
to the Acraeinae and Danainae of South Africa at all events, for from what I 


* Cambridge Nat. Hist., vol. 5, 1910. 
* Trans. Ent. Soc. London, 1902, p. 356. 
10 


7" 


144 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


have observed of these insects, J am convinced that their warning coloration 
cannot have reference to either Mantises, Asilidae, or lizards, which are prac- 
tically the only other enemies that can be taken into account. .... That 
they [birds] have been the chief, if not the only agents in the production of 
mimicry whether Batesian or Miillerian I have little doubt. 


In other words selectionists practically rest their case on the re- 
actions of birds to protective adaptations. The principal object of 
the present paper has been to show what those reactions are so far as 
nearctic birds are concerned, and there is no reason to suspect that the 
results are otherwise than typical for birds of the world. | 

The most outstanding feature of the records of the animal food of 
nearctic birds undoubtedly is the marvellous distribution of them 
through the phyla, orders, and subordinate systematic groups. Within | 
size limits, animals of practically every kind accessible to birds are 
preyed upon, and as we consider the records for group after group a 
tendency for the number of captures to be in proportion to the abun- 
dance of the animals concerned is unmistakable. Availability un- 
doubtedly is the chief factor involved in the choice of food, and pre- 
dation therefore tends to be in proportion to population. 

Considering bird predation alone this principle leads to a high de- 
gree of indiscriminancy in attack upon the whole kingdom of animal 
life. The combined attack of birds plus all other predators still more 
closely approaches complete indiscriminancy. In other words there is 
utilization of animals of practically every kind for food approximately 
in proportion to their numbers. This means that predation takes place 
much the same as if there were no such thing as protective adaptations. 
And this is only another way of saying that the phenomena classed by 
theorists as protective adaptations have little or no effectiveness. | 

Natural Selection theories assume discrimination in the choice of | 
prey. The principle of proportional predation so obvious from the 
data contained in this paper vitiates those theories for it denotes | 
indiscrimination, the very antithesis of selection. | 

Finally so far as the types of adaptations discussed in this paper are | 
concerned the influence of such factors as disease and climatic factors, 
the last the most important of all in reducing animal populations, is 
completely indiscriminate. 








The total mortality of animal groups is known normally to be in 
strict proportion to their numbers, 7. e., a pair of the new generation 
remains, to replace a pair of the old and it is apparent elimination of 
all but that pair is very largely due to agencies indiscriminate in their 
action. There would seem, therefore, to be no discriminative eliminat- 
ing forces of sufficient strength to bear the very great burden put 
upon them by natural selection theories. 





NO. 7 PROTECTIVE ADAPTATIONS—McATEE 145 


BIBLIOGRAPHY 


A complete bibliography of the subject matter of this paper would fill vol- 
umes and nothing like it is attempted here. Articles on food habits cited 
are chiefly those from which notes supplemental to our tabulations were gleaned ; 
few of foreign origin are included. The citations relating to enemies of various 
groups are illustrative only, not exhaustive, and some of exotic origin are 
used where helpful in indicating the wide distribution of predatory attack. 

The bibliography is primarily one of predation and so far as possible entries 
are distributed according to the thing eaten. When the feeding habits are 
varied, entries are filed according to the group the diet of which is reported 
upon. Titles classifiable under either of these criteria are arranged according 
to the phyla or orders to correspond with divisions of the text. Those unclassi- | 
fiable are grouped at the beginning of the bibliography as Miscellaneous. 


MISCELLANEOUS 


Buiecvap, H. 

1915. Food and conditions of nourishment among the communities of in- 
vertebrate animals found on or in the sea bottom in Danish waters. 
Rep. Danish Biol. Sta. to the Board Agr., vol. 22, 1914, pp. 41-78. 
Notes on food and enemy relations of many groups. 

Ciark, A. H. 

1925. Life in the ocean. Smithsonian Rep. 1923, pp. 309-304. Chiefly a 
discussion of food relations, many notes from which are used in 
this paper. 

Ewine, H. E. 

1929. A manual of external parasites. 225 pp., 96 figs. Includes informa- 
tion on the hosts attacked by mites, ticks, Mallophaga, Anoplura, 
and fleas. 

Force, Epitu R. 

1925. Notes on reptiles and amphibians of Okmulgee County, Oklahoma. 
Copeia, no. 141, p. 26, Apr. Notes on examinations of the stomachs 
of 6 species. 

KyerSKOG-AGERSBORG, H. P. 

1920. The utilization of echinoderms and of gasteropod mollusks. Amer. 
Nat., vol. 54, pp. 414-426, Sept.-Oct. Notes on these forms as food 
of fishes; short bibliography. 

McATEE, W. L. 

1918. Bird enemies of brine shrimps and alkali flies. The Auk, vol. 35, 
no. 3, p- 372, July. Eight species of birds mentioned as feeding 
extensively on both. Doctor Wetmore states: “The toll taken by 
birds from the brine shrimp and alkali fly larvae and pupae during 
the course of a season constitutes a mass of individuals almost 
beyond comprehension.” 

McIntosuH, W. C. 

1899. The resources of the sea as shown in the scientific experiments to 
test the effects of trawling and of the closure of certain areas off 
the Scottish shores. 248 pp., 32 tables, many pls. Notes on food 
of fishes, and other marine animals. 


146 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


Pratt, Henry S. 
1923. Preliminary report on the parasitic worms of Oneida Lake, New 
York. Roosevelt Wild Life Bull., vol. 2, no. I, pp. 55-71, Oct. 
Parasites of fishes, birds, reptiles, frogs, and mollusks. 
WEED, C. M. 
1884. The food relations of birds, frogs, and toads. Ent. Lab. Mich. Agr. 
Coll., pp. 20-20. Catbird 3 stomachs, robin 6, bluebird 2, crow 
blackbird 2, spotted frog 8, green frog 4, and toad 7. 
YonGE, C. M. 
1928. Feeding mechanisms in the invertebrates. Biol. Rev., vol. 3, no. 1, 
pp. 21-76, Jan. Notes on numerous food and enemy relationships. 
Full bibliography. 


PROTOZOA 


See entries under Miscellaneous, Blegvad, McIntosh, Yonge. 


PORIFERA 


See entries under Miscellaneous, Blegvad, McIntosh, Yonge. 


COELENTERATA 


See entries under Miscellaneous, Blegvad, McIntosh, Yonge. 


PLATYHELMINTHES 
GAMBLE, F. W. 
1910. Platyhelminthes and Mesozoa, Cambridge Nat. Hist., vol. 2, pp. 1-120. 
“ Turbellaria are carnivorous. .... Land Planarians feed on earth- 
worms, molluscs and wood-lice; fresh-water Planarians on Oligo- 
chaet worms, water-snails, and water-beetles; marine forms devour 
Polychaet worms and molluscs... .. Certain Rhabdocoelida are 
mess-mates of Molluscs and Echinoderms, and a few others are 
truly parasitic—a mode of life adopted by all Trematodes save 
Temnocephala” (p. 4). 
Stites, Cu. WARDELL. 
1902. Frogs, toads, and carp (Cyprinus carpio) as eradicators of fluke 
disease. 18th Ann. Rep. U. S. Bur. Animal Industry, 1901, pp. 
220-222, figs. 197-203. Carp apparently destroying large numbers 
of Fasciola hepatica. 


NEMATHELMINTHES 


VAN ZwWALUWENBURG, R. H. 
1928. The interrelationships of insects and roundworms. Bull. Exp. Sta. 
Hawaiian Sugar Planters’ Assoc., Ent. Ser. vol. 20, 68 pp., Jan. 
Some are primary parasites of orthoptera and bumble bees. 


TROCHELMINTHES 
Harroc, M. 
1910. Rotifera, Gastrotricha, and Kinorhyncha. Cambridge Nat. Hist., 
vol. 2, pp. 197-238. They devour algae, Infusoria, and other rotifers 
Cp: 212): 








NO. 7 PROTECTIVE ADAPTATIONS—McATEE 147 


MOLLUSCOIDA 
OssurN, RayMmonp C. 
1921. Bryozoa as food for other animals. Science, n. s., vol. 53, pp. 451-453, 
May 13. Two species of ducks and 10 of fishes noted as predators 
upon bryozoa. 


ECHINODERMATA 
CLARK, Hupert LYMAN. 
1920. Echinoderms in birds’ stomachs. Science, n. s., vol. 51, pp. 594-595, 
June 11. Ducks and gulls feeding on holothurians and brittlestars. 
Dawson, J. W. 
1867. The food of the common sea urchin. Amer. Nat., vol. 1, no. 3, pp. 
124-125, May. Minute seaweeds mixed with diatoms and remains 
of small sponges. 


ANNULATA 
BENHAM, W. B. 

1910. Chaetognatha. Cambridge Nat. Hist., vol. 2, pp. 186-194. “ The food 
of the Chaetognatha consists of floating diatoms, Infusoria, small 
larvae, and . . . . Copepods, small Amphipods, larval fishes”; they 
are also cannibalistic (p. 190). 

1910. Polychaeta. Cambridge Nat. Hist., vol. 2, pp. 245-344. “ The Nereidi- 
formia are mostly carnivorous, and feed on small Crustacea, Mol- 
lusca, sponges, and other animals; and Polynoids are even said to 
eat one another.” The Terebellids and Cryptocephala feed on minute 
organisms strained from water; the deep sea forms feed on Radio- 
laria and Foraminifera (p. 206). 

Buiarir, W. N. 

1927. Notes on Hirudo medicinalis, the medicinal leech, as a British Species. 
Proc. Zool. Soc. London 1927, pp. 999-1,002. The larvae live on 
frog-tadpoles and fish, and the adults on horses and cattle. 

Miter, JoHN A. 

1929. The leeches of Ohio. Distribution of the species together with what 
is known of their occurrence, food, and habitat. Ohio State Univ. 
Contr. vol. 2, Stone Labr., 38 pp. Feed on turtles, fishes, frogs, 
mammals, snails, worms, and insect larvae. 

Moore, J. PERcy. 

1923. The control of blood-sucking leeches, with an account of the leeches 
of Palisades Interstate Park. Roosevelt Wild Life Bull., vol. 2, 
no. I, pp. 9-53, I pl. 17 figs., Oct. Natural enemies of leeches 
include domestic and wild ducks, herons, kingfishers, crows, rats, 
minks, turtles, snakes, frogs, newts, fishes, crayfishes, dragonflies, 
and other predacious insects and leeches (pp. 20-30, 36). 


CRUSTACEA 
Bonnot, PAUL. 

1930. Crayfish. California Fish and Game, vol. 16, no. 3, pp. 212-216, 
figs. 65-67, July. Scavengers, will eat anything organic either alive 
or dead; destroy fish spawn. Are preyed upon by many fishes, 
other crayfishes, salamanders, snakes, turtles, kingfishers, raccoons, 
and man. 


148 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


Breper, C. M., Jr. 

1922. Notes on the summer food of Chilomycterus schoepfi Walbaum. 
Copeia, no. 104, pp. 18-19, Mar. 20. Analysis of contents of 26 
stomachs; chiefly crabs. 

Cott, HucuH B. 

1929. The Zoological Society’s Expedition to the Zambesi, 1927: No. 2, 
Observations on the natural history of the land-crab Sesarma 
meinerti, from Beira, with special reference to the theory of warn- 
ing colours. Proc. Zool. Soc. London, 1929, pp. 679-6092, pl. 1, 
figs. 1-4. “Crabs have numerous enemies; they are preyed upon 
by many small mammals, such as jackals, civets, and mongooses, 
but more especially by birds.” Pelicans, secretary-birds, herons, 
ibises, storks, owls, hawks, gulls, and waders (p. 689). 

Empopy, GEORGE C. 


1910. A new fresh-water amphipod from Virginia, with some notes on its - 


biology. Proc. U. S. Nat. Mus., vol. 38, pp. 299-305, 17 figs., 
June 18. Notes on 3 species of fish eating this Eucrangonyx, p. 305. 
Forses, S. A. 

1880. On the food of young fishes. Bull. Ill. State Lab. Nat. Hist., vol. 1, 
no. 3, pp. 66-79, Nov. Notes on stomach examinations; entomo- 
straca the most important food. 

1883. The first food of the common white fish. Bull. Ill. State Lab. Nat. 
Hist., vol. I, no. 6, pp. 95-109, May. Entomostraca. 

HANKINSON, T. L. 

1914. Young whitefish in Lake Superior. Science, n. s., vol. 40, pp. 239-240, 

Aug. 14. Notes on food, chiefly entomostraca. 
KENDALL, W. C. 

1923. Fresh-water Crustacea as food for young fishes. Rep. U. S. Comm. 
Fisheries, 1922, app. I, 32 pp., 10 figs. Copepods and ostracods in 
part carnivorous; Malacostraca and isopods, and amphipods, scaven- 
gers; crayfishes, carnivorous. Enemies of amphipods include fishes, 
birds, insects, Hydra, and the plant Utricularia; crustaceans im- 
portant food for young fishes. Bibliography. 

Kuiueu, A. B. 

1927. The ecology, food-relations, and culture of fresh-water Entomostraca. 
Trans. Royal Can. Inst., vol. 16, pt. I, pp. 15-98, May. The chief 
enemies of entomostracans are fish, dragonfly nymphs, and Hydra. 
They are eaten also by Corethra, young larvae of Dytiscus, tadpoles 
of Rana sylvatica, and by the entomostracans Leptodora kindtu and 
Cyclops fuscus. Their chief food is planktonic Chlorophyceae. A 
long bibliography. 

McATEE, W. L. 

1913. Some bird enemies of amphipods. The Auk, vol. 30, no. I, pp. 136- 
137, Jan. Amphipods preyed upon by 30 species of birds, including 
6 species of shorebirds and 14 ducks. 


MYRIAPODA 
Sincrarr, F. G. 

1910. Myriapoda. Cambridge Nat. Hist., vol. 5, pp. 20-80. Food of milli- 
peds, vegetable, of centipeds, animal, including diptera, other insects, 
worms, other centipeds. Centipeds eaten by South American 
Indians. 





NO.. 7 PROTECTIVE ADAPTATIONS—McATEE I49 


MISCELLANEOUS INSECTS 
Bairp, A. B. 
1923. Some notes on the natural control of the larch sawfly and larch case 
bearer in New Brunswick in 1922. Proc. Acadian Ent. Soc., vol. 8, 
1922, pp. 158-171. Lygaeonematus erichsonii. Birds ate about 10 
per cent and tachinid flies parasitized about 15 per cent of the 
larvae; a pentatomid, ants, and coccinellids were minor predators; 
hymenopterous parasites of the cocoons were scarce, but shrews 
consume about 40 per cent of them; natural enemies account for 
about 75 per cent of each brood. 
Coleophora laricella. Birds sometimes consume 75 per cent but on 
the average about 25 per cent of the larvae; ants and pentatomids 
take a few; hymenopterous parasites of the pupal stage were of 
slight importance. 
Beat, F. E. L., McAtee, W. L., and Katmsacu, E. R. 
1927. Common birds of Southeastern United States in relation to agricul- 
ture. Farmers’ Bull. 755, U. S. Dep. Agr. (rev.), 43 pp. 22 figs. 
In the introduction are statements about the bird enemies of certain 
groups of insects as 66 species vs. the cotton-boll weevil, 41 for 
the cottonworm, corn ear worm 12, white grubs 57, wireworms 128, 
billbugs 55, armyworm 43, cutworms 88, chinch bug 24, corn leaf 
beetle 22, corn root worm 26, leafhoppers 100, clover leaf weevil 
25, clover-root borers 74, cucumber beetle 19, sweet-potato flea 
beetle 28, grapevine flea beetle 23, bean leaf beetle 109, rice weevil 20, 
potato beetle 26, periodical cicada 33, and horse flies 40. 
CLARINVALL, AM. 
1928. De la disparition brusque des invasions d’ insects. Bull. Soc. Centr. 
Forest. Belgique, vol. 31, pp. 266-278, 316-335, and 378-302, 9 figs. 
Lack of food, climatic factors, natural enemies, and disease; the 
last three causes are given special attention; parasitic Hymenoptera 
and Diptera receive much comment and predators of the following 
groups are discussed: mammals, birds, Neuroptera, Coleoptera, 
Diptera, and Hymenoptera. 
Forsusy, E. H. 
1900. Birds as protectors of woodlands. Massachusetts Crop Rep., July, 
1900, pp. 26-39. Contains lists of birds feeding on gipsy moth, 
46 species; brown-tail moth, 29; forest tent caterpillar, 25; orchard 
tent caterpillar, 32; cankerworms, 51; tussock moth, 9; may 
beetles, 8; and plant lice, 34. 
GrIRAULT, A. A. 
1907. Hosts of insect egg-parasites in North and South America. Psyche, 
vol. 14, pp. 27-39, Apr. Coleoptera, 9 species, 10 egg parasites; 
Diptera, I species, I egg parasite; Hemiptera, 28 species, 43 egg 
parasites; Hymenoptera, 9 species, II egg parasites; Lepidoptera, 
51 species, 68 egg parasites; Neuroptera, I species, 2 egg parasites ; 
Orthoptera, 26 species, 47 egg parasites. 
1o1t. Hosts of insect egg-parasites in North and South America, II. 
Psyche, vol. 18, no. 4, pp. 146-153, Aug. Coleoptera, 6 species, 
6 parasites; Hemiptera, 10 species, 16 parasites; Hymenoptera, 
2 species, 4 parasites; Lepidoptera, 15 species, 22 parasites; 


150 





SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


Neuroptera, I species, I parasite; Odonata, 2 species, 5 parasites; 
Orthoptera, 4 species, 4 parasites; Platyptera, 1 species, I parasite. 


McATEE, W. L. 


1911. Economic ornithology in recent entomological publications. The Auk, 


vol. 28, no. I, pp. 138-142, Jan. Clover root curculios (Sitones 
hispidulus) are recorded as being taken by 24 species of birds. 
Sorghum midge (Contarina sorghicola) apparently eaten by hum- 
mingbirds. New Mexico range caterpillar (Hemileuca oliviae)— 
preyed upon by robins. Crane flies (Tipulidae)—86 species of 
nearctic birds are known to feed upon tipulids and their eggs. 
Pentatomidae are eaten freely by a great variety of birds. Yellow 
bear caterpillar (Diacrisia virginica)—two species of birds, the 
black-billed cuckoo and the bob-white, are recorded as enemies. 
Mosquitos are preyed upon by more than 20 species of North 
American birds. 


1911. Economic ornithology in recent entomological publications. The 


Auk, vol. 28, no. 2, pp. 282-287, Apr. Oakpruner (Elaphidion 
villosum) is recorded as preyed upon by four species of birds. 
Potato beetle (Leptinotarsa decemlineata)—21 species of birds 
are recorded as enemies of this pest. Hop flea beetles are eaten 
by killdeers and cliff swallows. Gipsy moths are recorded as being 
taken by 46 species of birds and brown-tail moths by 31. Alfalfa 
leaf weevil (Phytonomus murinus)—the English sparrow and the 
black-headed grosbeak are recorded as feeding on this weevil. 


1911. Economic ornithology in recent entomological publications. The 


Auk, vol. 28, no. 4, pp. 505-500, Oct. Muiullipeds are recorded as 
being eaten by 83 species of birds, certain beetle larvae, toads, 
armadillos and skunks. Cutworms are eaten on sight by prac- 
tically all birds that glean their food from the ground or from low 
vegetation. Flea beetles are eaten by various species of birds, in 
the case of Crepidodera by 26 species. Harlequin bug (Murgantia 
histrionica) is kept in check locally by the English sparrow. 
Cabbageworm (Pontia rapae) eaten by English sparrow. Codling 
moth—mortality during the winter as high as 90 per cent caused 
chiefly by birds, the beetle larvae, Tenebrioides sp., and diseases. 
Larch sawfly (Nematus erichsoni)—red-eyed vireos and cuckoos 
reported as feeding upon larvae of this insect. 


1912. Economic ornithology in recent entomological publications. The 


Auk, vol. 29, no. 3, pp. 416-417, July. Billbug (Sphenophorus 
callosus) recorded as taken by nighthawks. False wireworms 
(Eleodes) preyed upon by 13 species of birds. Agricultural ant 
(Pogonomyrmex barbatus molefaciens)—eight species of birds 
recorded as foes. Alfalfa weevil recorded as taken by 31 species 
of birds, notable mention being made of the English sparrow. 


1913. Economic ornithology in recent entomological publications. The 


Auk, vol. 30, no. I, pp. 128-132, Jan. Boll weevils preyed upon 
by 53 species of birds. A rice weevil (Lissorhoptrus simplex)— 
the only natural enemies recorded are two species of birds. Plum 
curculio—seven species of birds recorded as enemies. Leafhoppers 





NO. 7 PROTECTIVE ADAPTATIONS—McATEE 151 


known to be preyed upon by more than 120 kinds of birds, numerous 
species taking them in abundance. Nabidae, Lygaeidae, and spiders 
also mentioned as enemies. 

1913. Economic ornithology in recent entomological publications. The 
Auk, vol. 30, no. 4, p. 602, Oct. Eight species of birds observed 
feeding on the larvae of the fruit tree leafroller (Archips argyros- 
pila). May beetles and their larvae (Lachnosterna) preyed upon 
by 60 species of birds, the crow and crow blackbird probably being 
the most important enemies. 

1914. Economic ornithology in recent entomological publications. The 
Auk, vol. 31, no. 3, pp. 421-422, July. Sugar-beet wireworm 
(Limonius californicus )—the California shrike an important enemy. 
Crambus calignosellus—the quail and the kingbird noted feeding 
on this species. Crambus laqueatellus—the wood pewee observed 
taking large numbers. Rose aphid (Macrosiphum rosae)—house 
finch and white-crowned sparrow feeding on these aphids. Chinch 
bug—17 species of birds recorded as foes. 

1915. Economic ornithology in recent entomological publications. The 
Auk, vol. 32, no. 2, pp. 253-254, Apr. Wireworms (Elateridae) are 
recorded as being taken by 90 species of birds. Grasshoppers— 
upward of 100 species of birds are known to feed upon these insects. 
Alfalfahopper (Stictocephala festina)—four species of birds re- 
corded as enemies. Midges (Chironomus) are recorded as preyed 
upon by six species of birds. 

1915. Bird enemies of forest insects. Amer. Forestry, vol. 21, no. 6, pp. 681- 
691, June. Bark beetles are preyed upon by more than 45 species 
of birds. Round-headed and flat-headed wood borers—the larvae 
of these insects are recorded to be eaten by all kinds of wood- 
peckers. Flat-headed apple tree borers are recorded as taken by 
the downy woodpecker. Carpenter ants—fully 50 species of birds 
are known to eat these insects. An average of nearly 30 per cent 
of the food of woodpeckers is recorded as being ants. No fewer 
than 46 kinds of birds are known to feed upon the gipsy moth in 
one or another of its stages. Thirty-one species of birds are re- 
corded as enemies of the brown-tail moth. Orchard tent cater- 
pillars are preyed upon by 43 species of birds, forest tent cater- 
pillars by 32 and cankerworms by more than 50. Snow-white 
linden moth—the English sparrow is recorded as an important 
check on this insect. Plant lice are preyed upon by most small 
birds. Scale insects are known to be taken by more than 60 species 
of birds. Cicada—fishes and tortoises when opportunity presents, 
frogs, toads, lizards, squirrels, and a multitude of birds prey upon 
these insects. 

1915. Economic ornithology in recent entomological publications. The 
Auk, vol. 32, no. 4, pp. 520-521, Oct. Katydids—birds recorded as 
important foes, special mention being made of chipping sparrows. 
Calosoma sycophanta—crows and hairy woodpeckers recorded as 
enemies of this beetle. Armyworm—more than 20 species of birds 
recorded as foes. 





15 





SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


1915. birds that feed upon pecan insects. Proc. Nat. Nut Growers’ Assoc., 


pp. 40-41, Dec. 1. Pecan leaf caterpillar—birds known to feed 
upon pests of this genus (Datana) are robins, starlings, and two 
species of cuckoos. Fall webworm—four species of birds noted as 
enemies of this pest. Pecan weevil—64 kinds of birds are known 
to feed upon these beetles and congeners. White ants—of the 
27 species of birds feeding on white ants, a flicker is recorded to 
have taken 1,100. Oakpruners are known to be preyed upon by 
four species of birds. Cyllene—five species of birds are recorded 
as enemies. Bark beetles are devoured by more than 45 kinds of 
birds. Plant lice and scale insects each are taken by 60 or more 
species of birds. 


1916. Economic ornithology in recent entomological publications. The 


Auk, vol. 33, no. 2, pp. 216-217, Apr. Twelve-spotted cucumber 
beetle—27 species of birds recorded as enemies. Grasshoppers—six 
species of birds feeding upon them during an outbreak in New 
Mexico. Pine moth—the hairy woodpecker recorded as the most 
efficient natural force in restraining the Zimmerman pine moth. 


1916. Economic ornithology in recent entomological publications. The 


Auk, vol. 33, no. 4, pp. 448-450, Oct. Armyworm (Cirphis uni- 
puncta)—crows recorded as great destroyers of this pest; cow- 
birds and grackles also reported doing good work. Meadowlarks 
and robins observed eating the larvae. The armyworm has many 
natural enemies, among which are insects, reptiles, birds, and 
mammals. Skunks and toads undoubtedly eat thousands both of 
caterpillars and pupae. Clover leafhoppers are recorded taken by 
nine species of birds. Corn and cotton wireworm (Horistonotus 
uhleri)—birds are the only enemies of this pest recorded. Velvet- 
bean caterpillar—the “ricebird” and the mockingbird eat many 
of these. California green lacewing flies (Chrysopa californica) 
are recorded as taken by two species of birds, the western wood 
pewee and the nighthawk. 


1917. Bird enemies of a few insect pests. The Auk, vol. 34, no. 2, pp. 230- 


231, Apr. Grasshoppers are eaten by practically all birds, excep- 
tions being the strictly vegetarian doves and pigeons. Fall army- 
worm—several common wild birds recorded as important enemies. 
Cabbageworm—tbe English sparrow, chipping sparrow and house 
wren recorded as enemies of this pest. Velvetbean caterpillars 
are preyed upon by the red-winged blackbird, the mockingbird, 
and the field sparrow. 


1918. Economic ornithology in recent entomological publications. The 


Auk, vol. 35, no. 2, pp. 251-253. Apr. Potato aphid (Macrosiphwmn 
solanifolii)—chipping sparrows, quail, and English sparrows ob- 
served feeding on this pest. Sweet-potato leaf folder (Pilocrocis 
tripunctata) reported taken by the boat-tailed grackle. Cabbage 
looper (Autographa brassicae)—boat-tailed grackle observed feed- 
ing on adults and larvae. Pecan-leaf casebearer (Acrobasis nebu- 
lella) larvae taken by three species of birds. Fall webworm 
(Hyphantria textor )—red-eyed vireos recorded as destroying about 
40 per cent of the larvae in Nova Scotia in 1916; other bird foes 








NO. 7 PROTECTIVE ADAPTATIONS—McATEE 153 


noted are yellow-billed cuckoos and Baltimore orioles. Emperor 
moth (Samia cecropia)—cocoons destroyed by woodpeckers. 

1918. Economic ornithology in recent entomological publications. The 
Auk, vol. 35, no. 4, pp. 493-405, Oct. Round-headed apple tree borer 
(Saperda candida)—entomologists record birds as enemies; the 
present note names five species as feeding upon the adults. Root- 
worms—37 species of birds recorded as enemies of Diabrotica 
duodecimpunctata, and 23 species as enemies of Diabrotica soror. 
Green plant-bugs (Nezara spp.) identified in stomachs of 31 kinds 
of birds, 100 individuals being found in the stomach of a Franklin’s 
gull. Whitegrubs (Lachnosterna spp.) taken by 78 species of 
birds and 2 of toads. 

1919. Economic ornithology in recent entomological publications. The 
Auk, vol. 36, no. 2, pp. 305-307, Apr. Woodpeckers noted as prey- 
ing extensively upon larch bark beetles and borers. Grape root 
borer (Memythrus polistiformis)—the crested flycatcher observed 
feeding on the adults. Peach-tree borers (Sanninoidea exitiosa and 
S. pictipes)—two species of birds recorded as foes. Cankerworms 
preyed upon by 75 species of birds. Whitegrubs—several groups 
of birds named as enemies. 

1920. Economic ornithology in recent entomological publications. The 
Auk, vol. 37, no. 2, pp. 322-325, Apr. False wireworms (Eleodes)— 
24 species of birds recorded as enemies. Lotus borer (Pyrausta 
penitalis )—red-winged blackbirds noted as foes of this pest. Round- 
headed apple tree borer (Saperda candida)—‘‘ Woodpeckers de- 
stroy great numbers of the borers by removing them from their 
burrows. .... In some cases from 50 to 75 per cent.” Ten kinds 
of birds recorded as enemies. Flat-headed apple tree borer—12 
species of birds recorded as preying upon the adults of Chryso- 
bothris. Striped cucumber beetle (Diabrotica vittata)—17 species 
of birds noted as foes. Grainbugs (Chlorochroa spp.) recorded as 
taken by eight species of birds. Whitegrubs (Phyllophaga) preyed 
upon by 81 species of birds, the common crow being the most 
important enemy of both adults and larvae. 





1921. Economic ornithology in recent entomological publications. The 
Auk, vol. 38, no. 2, pp. 302-304, Apr. Spotted apple tree borer 
(Saperda cretata)—“ by far the most effective natural check to the 
increase of this borer seems to be the woodpeckers.” Clover stem 
borer (Languria mozardi)—hymenopterous parasites, toads, and 
five kinds of birds recorded as enemies. Beet leaf beetle (Mono-ia 
puncticollis)—enemies recorded are ladybird beetles, a stink bug, 
parasites, toads, and birds. Cabbage flea beetles (Phyllotreta spp.) 
12 kinds of birds noted as foes. Grapevine flea beetle (Altica chaly- 
bea)—eight species of birds recorded as feeding upon this insect. 
Clover leaf weevil (Hypera punctata) preyed upon by 42 species 
of birds. 

1922. Local suppression of agricultural pests by birds. Smithsonian Rep. 
1920, pp. 411-438, pls. 1-3. In more than 70 cases birds apparently 
exterminated one or another of 32 insect pests locally. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


1923. Economic ornithology in recent entomological publications. The 


Auk, vol. 40, no. I, pp. 161-162, Jan. Pale western cutworm 
(Porosagrotis orthogonia)—records the western grasshopper spar- 
row, horned larks, and possibly other wild birds as enemies. Green 
June beetle (Cotinis nitida)—19 kinds of birds recorded as foes; 
southern corn root worm (Diabrotica 12-punctata) taken by 4c 
species of birds. Potato beetle (Leptinotarsa decemlineata)—fed 
upon by 25 species of birds. 


1923. Economic ornithology in recent entomological publications. The 


Auk, vol. 40, no. 3, pp. 557-559, July. Corn ear worm (Heliothis 
obsoleta)—17 species of birds feed on this pest; more than 50 
larvae were found in a single stomach of the boat-tailed grackle. 
Cloverleaf weevil (Hypera punctata)—records 43 species of birds 
preying upon this weevil. European corn borer (Pyrausta 
nubilalis)—five species of birds recorded preying upon larvae and 
three species observed catching the moths. Tussock moth (Hemero- 
campa leucostigma)—-12 species of birds known to feed upon this 
insect in one stage or another. Lacewing flies (Chrysopidae)—17 
species of birds recorded as predators, most of them taking the 
adults, but five known to eat the larvae. 


1924. Economic ornithology in recent entomological publications. The 


Auk, vol. 41, no. I, pp. 191-193, Jan. American silkworm (Samia 
cecropia)—Dr. John Tothill concludes from his observations in 
Nova Scotia that nearly three-fourths of the caterpillars are eaten 
by birds (orioles, robins, etc.), and about 85 per cent of the pupae 
are destroyed by woodpeckers. Apple leaf skeletonizer (Hemero- 
phila pariana)—chipping sparrow observed feeding on the larvae. 
Mormon cricket (Anabrus simplex )—three species of birds men- 
tioned as enemies, but birds said to be important factors in the 
control of these insects. 


1924. Economic ornithology in recent entomological publications. The 


Auk, vol. 41, no. 4, pp. 629-632, Oct. False wireworms (Eleodes 
spp.)—13 kinds of birds recorded as enemies. Argus tortoise beetle 
(Chelymorpha cassidea)—identified in the stomachs of 14 species 
of birds, most often in those of the starling and kingbird. Codling 
moth (Carpocapsa pomonella)—woodpeckers recorded as important 
enemies, special mention being made of the red-bellied. Oak sap- 
ling borer (Goes tesselatus)—woodpeckers noted as destroying 
many larvae and pupae. Larch sawfly (Lygaconematus erich- 
soni )—four species of birds recorded as feeding upon the larvae, 
consuming about Io per cent of them in New Brunswick. Larch 
casebearer (Coleophora laricella)—4 species of birds recorded as 
enemies. Spruce budworm (Tortrix fumiferana)—several species 
of birds and insect parasites noted as foes. 


1926. Economic ornithology in recent entomological publications. The 


Auk, vol. 43, no. 3, pp. 396-308, July. Most common birds are 
enemies of the Japanese beetle (Popillia japonica). Green June 
beetle (Cotinis nitida)—observations show that starlings feed on 
the larvae and cardinals on the adults; in addition to these two 
birds, 22 other species are known to prey upon this pest. Striped 
cucumber beetle (Diabrotica vittata)—17 species of birds known 





NO. 7 PROTECTIVE ADAPTATIONS—McATEE 155 


to feed on this beetle. The larvae or pupae of cattle grubs (Hypo- 
derma) were found in stomachs of four species of birds; the robin 
also observed feeding on the larvae. Cankerworms (Alsophila 
pometaria and Paleacrita vernata)—76 species of birds listed as 
predators. Cabbageworm (Pieris rapae)—‘“‘ Birds which are known 
to feed upon cabbage worms are the chipping sparrow, English 
sparrow, and house wren.” 

1926. Relation of birds to woodlots. Roosevelt Wild Life Bull., vol. 4, 
no. I, 152 pp., 22 pls., Oct. Contains a section (pp. I0I-136) on 
forest insect pests and their bird enemies discussed under the 
following heads: Plant lice (Aphididae), scale insects (Coccidae), 
cicadas (Cicadidae), walkingsticks (Phasmidae), flat-headed wood 
borers (Buprestidae), leaf chafers (Scarabaeidae), leaf beetles 
(Chrysomelidae), round-headed wood borers (Cerambycidae), bark 
beetles (Scolytidae), caterpillars (Lepidoptera), and sawflies, borer 
wasps, and ants (Hymenoptera). 

1926. The role of vertebrates in the control of insect pests. Smithsonian 
Rep. 1925, pp. 415-437, 7 pls. General notes on amphibians, reptiles, 
and mammals as enemies of insects. Summarizes 109 cases of con- 
trol and 88 of local suppression of insects by birds. 

1928. Economic ornithology in recent entomological publications. The 
Auk, vol. 45, no. 4, pp. 526-528, Oct. Satin moth (Stilpnotia salicis) 
—five species of birds noted as enemies in Massachusetts. Western 
robins and bats reported as feeding on it in British Columbia. 
Lygus elisus—26 species of birds known to feed upon plant bugs 
of this genus. Cotton-stainers (Dysdercus spp.)—record of nine 
kinds of birds preying upon cotton stainers, and three species 
feeding upon other bugs of the same family. Fall armyworm 
(Laphygma frugiperda)—lists 13 species of birds as enemies, and 
notes that English sparrows have several times been observed to 
eradicate local infestations. Pale western cutworm (Porosagrotis 
orthogonia)—horned larks observed doing effective work against 
this pest. 

MurtrxKowskI, R. A., AND SMITH, G. M. 

1929. The food of trout stream insects in Yellowstone National Park. 
Roosevelt Wild Life Ann., vol. 2, no. 2, pp. 241-263, Oct. Stone- 
flies, carnivorous forms prey chiefly on larvae and pupae of 
mayflies, caddisflies, midges; mayflies are chiefly scavengers; 
caddisflies: the carnivorous species are inclined to be cannibalistic, 
but they take also rotifers, midge larvae and pupae, and dead 
insects. Bibliography. 

Patcu, E. M. 

1906. White grubs and June beetles. [In circulars, finance, meteorology, 
and index.] Bull. 137, Maine Exp. Sta., pp. 286-287. Enemies 
noted: Skunks, moles, and ground squirrels in addition to a large 
number of birds prey on the grubs. Besides toads and frogs and 
possibly insectivorous snakes, a large number of birds feed on the 
adult beetles. Cecropia moth, p. 294: enemies noted are chickens, 
turkeys, and swine. The tent caterpillar, p. 206: natural enemies 
of this caterpillar include birds and parasitic insects; it is also 
susceptible to attack by bacterial and fungus diseases. 








156 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


TuHompson, W. R. 


1929. On the relative value of parasites and predators in the biological 
control of insect pests. Bull. Ent. Research, vol. 19, pt. 4, pp. 343- 
350, Mar. Mentions parasitic habits for 3 families of Coleoptera, 
the Strepsiptera, 2 families of Neuroptera, 2 of Lepidoptera, 8 of 
Diptera, and 19 of Hymenoptera, and predatory habits in 3 families 
of Orthoptera, 9 of Neuroptera, the Odonata, some Corrodentia 
and Thysanoptera, in 8 families of Hemiptera, the Dermaptera, 19 
families of Coleoptera, the Mecoptera, 2 families of Lepidoptera, 15 
families of Diptera, and 6 of Hymenoptera. Thinks value of preda- 
tors has been underestimated. 

Uvarov, B. P. 

1928. Insect nutrition and metabolism. A summary of the literature. Trans. 
Ent. Soc. London, pp. 255-343. Largely technical on metabolism, 
but some details of food are given. There is a section on carnivo- 
rous insects, pp. 269-270. Bibliography. 


APTERA 
MACNAMARA, CHARLES. 
1924. The food of Collembola. Can. Ent., vol. 56, no. 5, pp. 99-105, May. 
Feed on sap, pollen, diatoms, algae, carrion, and Collembola. 


ODONATA 
Burnuam, Epwarp J. 
1899. Preliminary catalogue of the Anisoptera in the vicinity of Manchester, 
N. H. Proc. Manchester Inst. Arts and Sci., vol. 1, pp. 32-34. 
Certain birds appear to feed exclusively upon these insects while 
they last. The dragonflies mentioned are Macromia illinoensis and 
Tetragoneuria semiaquea. 
CALVERT, PHILtp P. 
1893. Catalogue of the Odonata (dragonflies) of the vicinity of Phila- 
delphia, with an introduction to the study of this group of insects. 
Trans. Amer. Ent. Soc., vol. 20, no. 3, pp. 205-206, July-Sept. 
Notes on numerous bird enemies of dragonflies. 
LAMBOoRN, Rost. H. 
1890. Dragonflies vs. mosquitoes, 202 pp., 9 pls. Nymphs feed on mosquito 
larvae, upon each other, upon water beetles, bugs, and small fishes. 
Not worth encouraging as mosquito enemies; fish and waterfowl 
also are foes of mosquitoes. 


IEWCAS SVs 
1908. Foe of dragonfly nymphs. Entomologist, vol. 41, p. 16. Notonecta 
glauca. 


Lyon, Mary B. 

1915. The ecology of the dragonfly nymphs of Cascadilla Creek (Odon.). 
Ent. News, vol. 26, no. 1, pp. 1-15, Jan. Notes on stomach contents 
of 36 specimens, midge larvae the most prominent item of food, 
but mayflies, Corixids, dytiscids, amphipods, cladocera, ostracods, 
hydrachnids, and snails were eaten. 

Moorr, J. PEercy. 

1900. Kingbirds eating dragonflies. Ent. News, vol. 11, p. 340. Epiaeschna 

heros; habitually captures them. 


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NO. 7 PROTECTIVE ADAPTATIONS—McATEE 157 


NEEDHAM, JAMES G. 

1808. Birds vs. dragonflies. Osprey, vol. 2, nos. 6-7, pp. 85-86, Feb.-Mar. 
Review of notes by Rene Martin on European hobby and swift 
as enemies; Merops persicus lines its nest with their wings; when 
teneral, chipmunks, frogs, toads, snakes, ants, and birds eat them. 
Various birds eat nymphs. 

1918. [Food of Odonata.] In Fresh-water biology, by Ward and Whipple, 
p. 890. Diptera and other dragonflies. 

NEEDHAM, JAS. G., AND HEywoop, Hortense B. 

1929. A handbook of the dragonflies of North America, 378 pp., illus. Food, 
flies, mosquitoes, honey bees; enemies, birds, frogs, fishes, water 
snakes, spiders, other dragonflies; large numbers sometimes de- 
stroyed by storms. 

SuaArp, Davin. 

1910. [Enemies of Odonata.] Cambridge Nat. Hist., vol. 5, pp. 424-425. 
Hawks, bee-eaters, other birds, fishes, snakes, newts, aquatic Cole- 
optera, Hemiptera, and other Odonata. 

WatkKer, E. M. 

1924. The Odonata of the Thunder Bay District, Ontario. Can. Ent., vol. 
56, no. 7, pp. 170-176, July; no. 8, pp. 182-189, Aug. Dragonflies 
found in stomachs of sucker, whitefish, sturgeon, and golden-eye 
ducks ; a dragonfly nymph observed eating an adult of same species. 

Witson, CHARLES BRANCH. 

1920. Dragonflies and damselflies in relation to pondfish culture, with a 
list of those found near Fairport, Iowa. Bull. 36, U. S. Bur. 
Fisheries, pp. 182-264, pls. 67-60, figs. 1-63, Aug. Notes on con- 
tents of alimentary canals of 250 nymphal and many adult Odonata; 
citation of previous studies; full bibliography. Odonate nymphs, 
diving beetles, water-scorpions, other aquatic Hemiptera, Hydra, 
nematodes, fungi, birds, fishes, reptiles and amphibians prey on 
nymphs; Diptera and Hymenoptera parasitize the eggs; and birds, 
other dragonflies, ants, spiders, robber flies, frogs, and fishes prey 
upon the adults, which also have both external and internal 
parasites. 


AGNATHA 


See various entries under Pisces; also Muttkowski and Smith under Miscel- 
laneous Insects. 
NEEDHAM, JAMES G. 

1920. Burrowing mayflies of our larger lakes and streams. Bull. U. S. 
Bur. Fisheries, vol. 36 (1917-1918), pp. 269-292, pls. 70-92. May- 
flies of prime importance as food of fishes; quotations from Forbes, 
Waener, and Pearse, as to their value (pp. 270-271). 


PLECOPTERA 


See various entries under Pisces; also Muttkowski and Smith under Miscel- 
laneous Insects. 


158 SMITHSONIAN MISCELLANEOUS -COLLECTIONS VOL. 85 


ISOPTERA 
[HacEN, H.] 

1881. [Letter on birds vs. termites.] Proc. Boston Soc. Nat. Hist., vol. 20, 
1878-1880, p. 118. Record of 15 species of birds following an 
emigration of white ants, robins, bluebirds, and sparrows being 
mentioned. 

[LonestaFF, G. B.] 

1918. A flight of winged termites at Barrackpore. Trans. Ent. Soc. London, 
1918, pp. Ixiv-lxvi. Lizards, bullfrogs, rats, cats, dogs, jackals, 
mongoose, crows, Indian mynah, bats and cockroaches observed 
eating white ants. 

Snyper, T. E. 

1920. [Nearctic Termites.] Notes on biology and geographic distribution. 
U. S. Nat. Mus. Bull. 108, pp. 87-211. Termite checks include 
parasitic fungi, protozoans, nematodes, mites, and predacious ants, 
robber flies, beetle larvae, crickets, spiders, centipeds, lizards, and 
domestic and wild birds (pp. 116-118). 

1924. New termites and hitherto unknown castes from the Canal Zone, 
Panama. Journ. Agr. Research, vol. 29, no. 4, p. 182, Aug. I5. 
Ants and anteaters as foes. 


DERMAPTERA 
BRINDLEY, H. H. 

1920. Notes on certain parasites, food, and capture by birds of the common 
earwig (Forficula auricularia). Proc. Cambridge Phil. Soc., vol. 
19, (1916-1919) pp. 167-177. Fourteen species of British birds 
known to eat earwigs; also domestic fowls. 

Morcan, W. P. 

1924. Notes on the function of the forceps in earwigs. Proc. Indiana 
Acad. Sci., vol. 33, (1923), pp. 303-306, 7 figs. Earwigs are preda- 
tory and cannibalistic, use forceps in capturing and holding prey. 

Suarp, Davin. 

1910. Forficulidae—earwigs. Cambridge Nat. Hist., vol. 5, pp. 202-216. 

Eat larvae, snails, flowers, vegetables. 


CHELEUTOPTERA 
Bapenocy, L. N. 
1899. [Enemies of Phasmidae.] True tales of the insects, p. 48. Birds, 
lizards, mantids, bugs; eggs parasitized. 
Suarp, DAvIp. 
1910. Phasmidae—stick and leaf insects. Cambridge Nat. Hist., vol. 5, 
pp. 260-278. Vegetarian, but sometimes cannibalistic. Enemies in- 
clude birds, Hemiptera, ichneumon flies. 


SALTATORIA 
AUGHEY, SAMUEL. 

1878. Notes on the nature of the food of the birds of Nebraska. 1st Ann. 
Rep. U. S. Ent. Comm. (1877), Appendix II, pp. [13-62.] Records 
migratory locusts from the stomachs of 172 species of birds and 
field observations on 33 other species eating them. 





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INOS 7 PROTECTIVE ADAPTATIONS—McATEE 159 


BapenocH, L. N. 

1899. [Enemies of locusts.] True tales of the insects, pp. 127-128. Especially 
“locust birds”; but bears, skunks, squirrels, mice, frogs, and lizards 
are mentioned. 

BRUNER, LAWRENCE. 

1902. Grasshopper notes for 1901. Bull. 38, Div. Ent., pp. 39-49. Chickens, 
turkeys, blackbirds, sage grouse and sharp-tail grouse mentioned 
as natural enemies. 

1905. Grasshopper conditions in Nebraska, Northeastern Colorado, Wyom- 
ing, Montana, and Western Kansas during the summer of 1904. 
Bull. 54, U. S. Bur. Ent., pp. 60-64. ‘“ Magnificent examples” of 
the usefulness of gulls as grasshopper destroyers; turkeys used to 
combat the insects. 

Bryant, H. C. 

1912. Birds in relation to a grasshopper outbreak in California. Univ. 
California Publ. in Zool., vol. 11, no. 1, pp. 1-20, Nov. Los Banos; 
15 species of land birds were found to eat the insects. Tame ducks 
also important. The native birds were calculated to be destroying 
daily 120,445 grasshoppers per square mile. 

Burritu, A. C. 

1920. Meadowlarks control cricket pest. California Fish and Game, vol. 6, 
no. I, p. 38, Jan. Meadowlarks recorded as important enemies of 
the coulee cricket. 

CrippLe, NorMAN. 

1920. Birds in relation to insect control. Can. Field-Nat., vol. 34, no. 8, 
pp. 152-153, Nov. Crows, gulls, black terns, blackbirds, and grouse 
recorded as destroying large numbers of grasshoppers. 

1922. [Enemies of grasshoppers.] Can. Field-Nat., vol. 36, no. 4, pp. 66-68, 
Apr. Diptera, Hymenoptera, Coleoptera, birds, mammals (skunks, 
badgers, weasels, pocket mice, shrews, gophers), snakes, toads, and 
frogs. 

GILLETTE, C. P. 

1905. The western cricket. Colorado Agr. Exp. Sta., Bull. 101, 16 pp., 
Apr. Anabrus simplex. Bears and coyotes feed upon this pest but 
birds destroy them in greatest numbers; hawks, sage grouse and 
blackbirds noted (p. 7). 

Grasse, P. 

1924. Les ennemis des Acridiens ravageurs francais. Rev. Zool. Agr. Appl., 
Bull. Soc. Zool. Agr., vol. 23, no. I, pp. 1-14, pl. 1, figs. 1-4, Jan. 
Mammals, birds, reptiles, spiders, mites, wasps, beetles, flies, nema- 
todes and Protozoa. 

LuGGeErR, OTTOo. 

1889. Notes on the Rocky Mountain locust in Otter Tail County, Minne- 
sota, in 1888. 5th Bienn. Rep. Dep. Agr. Minn., Suppl. 1, pp. 
305-343, 22 figs. Nematodes, mites, tachina flies, bee flies, blister 
beetles, ground beetles, soldier beetles, robber flies, digger wasps, 
dragonflies, birds, skunks, shrews, toads, snakes, and_ turtles 
mentioned as enemies. 

rT 





160 SMITHSONIAN MISCELLANEOUS COLLECTIONS — VOL. 85 


McAtTeEg, W. L. 

1913. Economic ornithology in California. The Auk, vol. 30, no. I, pp. 
132-136, Jan. H. C. Bryant records 22 species of water and shore 
birds and 40 species of land birds as enemies of grasshoppers. 

1917. Economic ornithology in recent entomological publications. The 
Auk, vol. 34, no. 4, pp. 497-498, Oct. Grasshoppers are found on the 
bill-of-fare of practically all wild birds; freely eaten also by 
chickens and turkeys. 

Merritt, D. E. 

1916. [Enemies of grasshoppers.] Bull. 102, New Mexico Agr. Exp. Sta, 
pp. 15-16, Apr. Birds; fields near breeding grounds of the black- 
birds are free from grasshopper damage; poultry; skunks; mites; 
parasitic flies; ground beetles; blister beetles; bee flies. 

MorseE, ALBERT P. 

1920. [Enemies of Orthoptera.] Proc. Boston Soc. Nat. Hist., vol. 35, 
p. 271. Frogs, toads, salamanders, snakes, lizards, birds, mice, 
moles, shrews, skunk, and fox. 

SANDERSON, E. DwIGHT. 

1906. The differential locust. Bull. 57, U. S. Bur. Ent., pp. 19-26, figs. 
g-11. Melanoplus differentialis. Blackbirds and bobolinks sup- 
pressing an infestation; a conopid fly parasite also mentioned. 

1906. Report on miscellaneous cotton insects in Texas. Bull. 57, U. S. 
Bur. Ent., p. 22. Blackbirds and bobolinks checking an outbreak 
of Melanoplus differentialts. 

SHARP, Davin. 

1910. [Enemies of Orthoptera.] Cambridge Nat. Hist., vol. 5, p. 201. 
Cantharidae, Bombyliidae, and mites destroy eggs; birds and mam- 
mals eat adults. 

SmitH, Harrison E. 

1915. The grasshopper outbreak in New Mexico during the summer of 
1913. Bull. 293, U. S. Dep. Agr., 12 pp. 2 figs. Six species va 
birds, several species of lizards, prairiedogs, a sarcophagid parasite, 
and a wasp observed doing notable execution. 

TREHERNE, R. C. AND BUCKELL, E. R. 

1924. The grasshoppers of British Columbia. Bull. 39, Dominion of Canada 
Dep. Agr., pp. 29-35, Oct. Enemies include: nematodes, Diptera, 
Hymenoptera, Coleoptera, fungi and bacteria. 

U. S. ENToMoLocicaAL CoMMISSION. 

1878. First annual report .... for the year 1877 relating to the Rocky 
Mountain locust, etc., pp. 477-++-[204], 111 figs., 5 pls. Invertebrate 
enemies (pp. 284-334) include larvae of anthomyiid and sarcophagid 
flies, ground beetles, blister beetles, click beetles, soldier beetles, 
robber flies, and mites all attacking the eggs; and the following 
preying upon the locusts after birth: mites, ground beetles, tiger 
beetles, robber flies, wasps, tachinid and sarcophagid flies, ichneu- 
monids and nematodes. The vertebrate enemies (pp. 334-350) 
include birds, hogs, skunks, prairie squirrels, mice, and toads. 
Appendix II [pp. 13-62], is devoted to an account of the food of 
birds especially in relation to the locust. 





NO. PROTECTIVE ADAPTATIONS—McATEE 161 


N 


PALEOPTERA 


SHARP, DAVID. 

1910. Blattidae—cockroaches. Cambridge Nat. Hist., vol. 5, pp. 220-241. 
Food chiefly dead animal matter, but a great variety of refuse 
also taken. Enemies include birds, rats, scorpions, spiders, and 
wasps (Ampulicides). 


DICTYOPTERA 


SHaArp, DAvID. 
1910. Mantidae—soothsayers or praying insects. Cambridge Nat. Hist., 
vol. 5, pp. 242-259. Voracious, eating insects of all kinds including 
their own, and even small birds. 


CORRODENTIA 


Food animal and vegetable refuse, and fungi. 


MALLOPHAGA 


Externally parasitic on birds and mammals. 





SIPHONAPTERA 


External parasites on birds and mammals. 


RHYNCHOTA 
CiarK, L. B. 

1928. Seasonal distribution and life history ot Notonecta undulata in the 
Winnipeg Region, Canada. Ecology, vol. 9, no. 4, pp. 383-403, pl. 
20, I fig., Oct. Summary of literature as to food and enemies, pp. 
395-399. Food: eggs of giant water bug, water-boatman, eggs and 
nymphs of dragonflies, ostracods, copepods, newly hatched fishes. 
Enemies: giant water bug, water-scorpion, waterstrider, dragonfly 
nymphs, fishes, and birds. 

Curran, C. Howarp. 

1920. Observations on the more common aphidophagous syrphid flies 
(Dipt.). Can. Ent., vol. 52, no. 3, pp. 53-55, Mar. Larvae of five 
species consumed on the average from 15 to 47 plant lice daily. 

Distant, W. L. 

1892. A monograph of the Oriental Cicadidae, pp. vii-vill. “ The Cicadidae 
appear to be one of the most non-protected families of insects and 
are the victims of most predacious creatures.” Mentions birds, 
mantids, spiders, dragonflies, wasps, hymenopterous parasites and 
fungi. 

BEUKE, C,. 15, 

1929. The known predacious and parasitic enemies of the pea aphid in 
North America. Research Bull. 93, Wisconsin Agr. Exp. Sta., 
47 pp., 3 pls., 32 figs. Mites, spiders, crickets, lacewing flies, 
Hemiptera, Coleoptera, Diptera, Hymenoptera, and birds. Lace- 
wings, ladybirds, and Syrphidae appear to be most important. 


162 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


FuLitaway, Davin T. 

1920. Natural control of scale insects in Hawaii. Proc. Hawaiian Ent. 
Soc. (1919), vol. 4, no. 2, pp. 237-246, June. Forty-four species 
of scale insects, 87 parasites, 20 predators; varying up to 3 preda- 
tors and 7 parasites to a species. 

GarMAN, H. 

1898. The chinch bug. Bull. 74, Kentucky Agr. Exp. Sta., pp. 45-70, 
figs. 1-10, May. Coccinellids, toads, quail, and meadowlarks prey 
upon it (p. 51); great fluctuations in abundance caused by disease. 

GIBSON, EpmunpD H. 

1916. The clover leafhopper and its control in the central States. Farmers’ 
Bull. 737, U. S. Dep. Agr., 8 pp., 5 figs., June. Agallia sanguino- 
lenta. More than 100 species of birds, chickens, turkeys, and guinea 
fowl prey upon leafhoppers (p. 5). 

Hanrt, H. 

1916. Beitrage zur Kenntniss der Cicadinenfeinde. Zeitschr. wiss. Insekten- 
biol., vol. 12, pp. 200-204, 217-223, 274-279, figs. Stresiptera, 
Dryinidae, Serphoidea, Pipunculidae, Neuroptera, Nematoda, mites, 
fungi. Bibliography. 

Huncerrorp, H. B. 

1919. The biology and ecology of aquatic and semi-aquatic Hemiptera. 
Kansas Univ. Bull., vol. 11, 265 pp., 30 pls. Gelastocoridae—grass- 
hoppers, lace bugs, beetle larvae, capsids. Ochteridae—tabanid 
larvae. Saldidae—drowned flies, etc. Hydrometra—Ostracoda, 
culicid larvae and pupae. Mesovelia—springtail, crambus, chalcid, 
Hydrometra, Ostracoda. Gerris remigis—midges, notonectid 
nymphs; jassids, etc., falling in water, snails. Rheumatobates— 
Ostracoda and fallen insects. Microvelia—Ostracoda and fallen 
insects, waterfleas. Belostoma—fish, snails. Lethocerus—tfrog, fish. 
Nepa—mayfly nymphs, Gyrinidae, Daphnia, Cyclops, fish eggs, fish, 
tadpoles. anatra—ostracod, fish, mayfly nymphs. PlJea—Ostra- 
coda and other small Crustacea. Notonecta—cannibalistic, Ostra- 
coda and other small Crustacea, corixids. Buenoa—Entomostraca, 
corixids. Corixid nymphs cannibalistic. 

JENSEN-Haarup, A. C. 

1924. Wasps preying on cicadas, Ent. Meddel, vol. 14, pp. 323-324. Also 

birds, spiders, mantids, and dragonflies noted as their enemies. 
JoHNSoN, Rosweti H. 

1907. Economic notes on aphids and coccinellids. Ent. News, vol. 18, no. 
5, pp. 171-174, May. Coccinellids, syrphids, spiders, and fungi as 
aphid destroyers. 

KirKaLpy, G. W. 

1907. [Enemies of Aleyrodidae.] Bull. 2, Board Agr. and For. Terr. 
Hawaii, pp. 80-84. Three species of flies, 4 of beetles, 1 of hemip- 
tera, 22 of hymenoptera, I neuropteron, I thysanopteron, I mite, and 
2 fungi. 

Loranpo, N. T. 

1929. A biological method for destroying bedbugs. Sci. Monthly, pp. 265- 

268, Sept. Spiders, reduviid bugs, cockroaches, and ants as enemies. 

















NO: 7 PROTECTIVE ADAPTATIONS—McATEE 163 


LuccER, OTTo. 

1895. [Enemies of the chinch bug.] Bull. 37, Minnesota Agr. Exp. Sta., 

pp. 178-179. Birds, reptiles, frogs, toads; some specified. 
MacAnprews, A. H. 

1923. Some notes on the natural control of the pine bark aphid (Chermes 
pinicorticis Fitch) in New Brunswick, 1922. Proc. Acadian Ent. 
Soc., vol. 8, 1922, pp. 52-56. A coccinellid exerted from 75 to 90 per 
cent of the natural control, and a syrphid fly and ant-lion the 
remainder. 

Martartt, C. L. 

1907. The periodical cicada. Bull. 71, U. S. Bur. Ent., 181 pp., 6 pls., 
68 figs. Natural enemies include dipterous, hymenopterous, and 
mite egg parasites, tachinid parasites of the adult, wasps, birds, 
squirrels, fishes; in some cases birds ate the insects as fast as they 
emerged. 

McAtTEE, W. L. 

1907. Birds that eat scale insects. U. S. Dep. Agr. Yearbook, 1906, pp. 
189-108. Fifty-seven species of birds are recorded as feeding upon 
scale insects. 

1913. Relation of birds to [an outbreak of] grain aphides. U. S. Dep. 
Agr., Yearbook 1912, pp. 397-404. Spring migrant birds on about 
100 acres of grainfields in North Carolina destroyed about 1,000,000 
grain aphids daily. 

1918. Bird enemies of tree hoppers (Membracidae). The Auk, vol. 35, 
no. 3, pp. 373-374, July. Treehoppers identified in the stomachs of 
more than 120 species of birds, as many as 26 individuals being 
found in a single stomach. 

McGrecor, E. A. 

1927. Lygus clisus: a pest of the cotton regions in Arizona and California. 
Techn. Bull. 4, U. S. Dep. Agr., 14 pp., 7 figs., July. Bugs of genus 
Lygus have been found in stomachs of 26 species of birds; Re- 
duviidae and spiders also noted as enemies (p. 8). 

MontTIzAMBERT, ERIC. 

1908. Lampyrids and aphids. Can. Ent., vol. 40, no. I, p. 36, Jan. Tele- 
phorus carolinus extirpating colonies of Siphonophora rudbeckiae 
(a red aphis). 

Moznette, Geo. F. 

1915. Notes on the brown lace-wing (Hemerobius pacificus Bks.). Journ. 
Econ. Ent., vol. 8, no. 3, pp. 350-354, pl. 15, June. Number of 
aphids devoured daily by each of five jarvae varied from 24 to 27. 
Captive. 

Myers, J. G. 

1927. The natural enemies of Dysdercus. Ann. Ent. Soc. Amer., vol. 20, 
no. 3, pp. 290-204, Sept. In an article entitled ‘ Ethnological Obser- 
vations on some Pyrrhocoridae of Cuba,’ the author reports on 
observations, published records, and experiments with natural 
enemies of Dysdercus. The actually observed enemies are spiders, 
pseudoscorpions, thrips (the eggs), tachinid flies, reduviid and 
other bugs, lizards, and birds. 





164 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


REINHARD, EDWARD G. 

1925. The wasp Hoplisus costalis, a hunter of tree-hoppers. Journ. Wash- 
ington Acad. Sci., vol. 15, no. 5, pp. 107-110, Mar. 4. An exclusive 
enemy of Membracidae; 12 species identified from nests. 

SHERMAN, FRANKLIN, JR. 

The harlequin cabbage bug. Bull. North Carolina Dep. Agr., vol. 32, no. 7, 
pp. 17-24, July. Murgantia histrionca. The English sparrow is 
quite an efficient aid in keeping this pest in check (p. 21). 

SmitH, Harry S. 

1917. Insect parasites and predators as adjuncts in the control of mealybugs. 
Monthly Bull. California Comm. Hort., vol. 61, nos. 3-4, pp. 108- 
114, Mar.-Apr. One chrysopid, seven coccinellids, one agromyzid, 
and one syrphid as predators, and six Hymenoptera as parasites 
upon species of Pseudococcus. 

Surrace, H. A. 

1907. [Enemies of plant lice.] Zool. Bull. Pennsylvania Dep. Agr., vol. 5, 
no. 3, pp. 81-82, July. Warblers, wrens, titmice, kinglets, chickadees. 

1907. [Enemies of the periodical cicada.] Zool. Bull. Pennsylvania Dep. 
Agr., vol. 5, no. 3, p. 74, July. Skunks, squirrels, moles, chipmunks, 
pigs, poultry, most birds, snakes (four species mentioned) and 
turtles listed. 

1907. Psyllidae. The jumping plant lice. Zool. Bull. Pennsylvania Dep. 
Agr., vol. 5, no. 3, pp. 78-79, July. White-breasted nuthatches 
practically freed an orchard from pear psyllids (Psylla pyricola). 

THompson, W. L. 

1928. The seasonal and ecological distribution of the common aphid pre- 
dators of central Florida. Florida Ent., vol. 11, no. 4, pp. 49-52, 
Feb. Cycloneda feeding on seven species of aphids; Hippodamia 
on 5, the larvae averaging 56 and the adults 87 bean aphids per 
day; Scymnus on 5; a few other species briefly mentioned, of 
which Coccinella, Rodolia, and Chilocorus are said to be primarily 
scale devourers. 

Wesster, F. M. 

1897. [Enemies of the periodical cicada.] Bull. 87, Ohio Agr. Exp. Sta., 
pp. 61-63, Noy. Birds, especially the English sparrow, parasitic 
flies, dragonflies, soldier-bugs, ground beetles, hogs, and poultry. 

1907. [Enemies of the chinch bug.] Bull. 60, U. S. Bur. Ent., pp. 58-59. 
Birds, frogs, nematodes, ants, ladybeetles, predacious Hemiptera, 
ground beetles, lacewing flies, spiders, and parasitic fungi. 

1909. The Chinchbug (Blissus leucopterus Say.). Circ. 113, U. S. Bur. 
Ent,, 27 pp., 8 figs., Nov. Natural enemies, bobwhite (100-400 
eaten at a meal), 15 other birds, frogs, ants, ladybirds, anthocorids, 
carabids, chrysopids, spiders, and diseases. 

WILDERMUTH, V. L. 

1915. Three-cornered alfalfa hopper. Journ. Agr. Research, vol. 3, no. 4, 
PP. 343-362, pl. 43, fig. 1, Jan. Stictocephala festina. Spiders, ants, 
mites, egg parasites, birds, and toads recorded as enemies (pp. 
359-360). 


NO. 7) PROTECTIVE ADAPTATIONS—McATEE 165 


WIittiaMs, C. B. 

1921. Report on the froghopper-blight of sugar-cane in Trinidad. Mem. 1, 
Dep. Agr. Trinidad and Tobago, Jan. Tomaspis saccharina Dist- 
ant. Enemies listed are: 2 species of hymenopterous parasites 
and 2 species of thrips predatory upon the eggs; 3 kinds of birds, 
1 syrphid larva, and 1 nematode upon the nymphs; and 17 species 
of birds, 2 of grasshoppers, 7 of ants, 1 of Lampyridae and 5 of 
Reduviidae predacious on the adults, besides sundry lizards, toads, 
frogs, spiders, mites, and two fungoid diseases. 


NEUROPTEROIDEA 


Insects chiefly predacious in the larval state and often also as imagos. For note 
on Trichoptera, see Muttkowski and Smith under Miscellaneous Insects. 


McGrecor, E. A. 

1914. Some notes on parasitism oi chrysopids in South Carolina. Can. 
Ent, vol. 46, pp. 306-308, fig 1. Forty-eight out of 99 cocoons were 
destroyed by hymenopterous parasites. 

SMITH, Rocer C. 

1922. The biology of the Chrysopidae. Mem. 58, Cornell Univ. Agr. Exp. 
Sta., pp. 1287-1372, pls. 75-88, figs. 154-163, June. Parasitized in 
all stages; ladybirds eat the eggs. Certain birds feed on adults. 
Robber flies noted catching adults and some Hemiptera prey on 
the larvae. 


LEPIDOPTERA 
AINSLIE, C. N. 

1910. The New Mexico range caterpillar. Bull. 85, U. S. Dep. Agr., pt. 5, 
pp. 59-96, figs. 1-53, June. HWemileuca oliviae—insect parasites, 
mites, robber flies, and robins recorded as enemies (pp. 88-93). 

ALLEN, J. A. 

1894. On the mammals of Aransas County, Texas, with descriptions of 
new forms of Lepus and Oryzomys. Bull. Amer. Mus. Nat. Hist., 
vol. 6, pp. 165-198. Onychomys longipes. Quotes a note from 
H. P. A[ttwater] to the effect that he found several hundred 
wings of Danais archippus, the bodies of which had been eaten 
by the Onychomys. Allen adds “ This observation is of special 
interest from the fact that this butterfly is supposed to be ‘pro- 
tected’ by a nauseous odor or taste that renders it unpalatable 
to animals” (p. 181). 

BabEeNocH, L. N. 

1899. [Enemies of the lictor moth.] True tales of the insects, p. 215. Notes 
on hymenopterous and dipterous parasites. The caterpillars of a 
species may be collected persistently for years for breeding and, 
because of parasites, none of them reach the perfect stage. 

Batt, E. D. 

1904. The codling moth. Bull. 87, Utah Exp. Sta., pp. 119-120. Birds, 
particularly the downy woodpecker and the chickadee, bats, ants, 
spiders, and predacious insects, recorded as enemies of this pest. 






166 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


BARBER, G. W. 

1925. The efficiency of birds in destroying overwintering larvae of the 
European corn borer in New England. Psyche, vol. 32, no. 1, 
pp. 30-46. Birds credited with destroying an average of 61 per cent 
of the larvae in test cases. 

BEEBE, WM. 

1924. Notes on Galapagos Lepidoptera. Zoologica, vol. 5, no. 3, pp. 51-50, 
pl. A., Jan. 11. Birds feeding on butterflies (Callidryas eubule 
and Agraulis vanillac) and moths. “ The relation between birds 
and butterflies is quite a negligible factor in any lepidopterous theory 
of evolution of pattern, color, form, or activity” (p. 57). 

BELE Eee 

1924. Notes on Asilus sericeus Say (Diptera, Asilidae). Journ. New York 
Ent. Soc., vol. 32, no. 4, p. 219, Dec. Capturing Hesperiidae, in- 
cluding Epargyreus tityrus. 

BERGER, E. W. 

1920. The semitropical army worm. Quart. Bull. Florida State Plant 
Board, vol. 4, no. 2, pp. 17-34, figs. 4, Jan. Xylomyges eridania. 
Birds, the spined soldier bug, a wasp, tiger beetles, ground beetles, 
and parasitic insects recorded as enemies (pp. 26-28). 

Birp, HENRY. 

1909. [Enemies of Papaipema maritima.] Can. Ent., vol. 41, no. 2, pp. 67-8, 
Feb. Field mice and skunks. “As very few pupae escape in any 
locality these animals go over, they become an important factor 
in the economy of the species.” 

BrEAKEY, E. P. 

1929. Notes on the natural enemies of the iris borer, Macronoctua onusia 
Grote (Lepidoptera). Ann. Ent. Soc. Amer., vol. 22, no. 3, pp. 
459-464, Sept. Six species of Diptera, two of Hymenoptera, one 
beetle, one bird, and rodents. 

Britton, W. E. 

1906. The gypsy moth and the brown-tail moth. Bull. 153, Connecticut 
Exp. Sta., p. 7. Several species of parasitic Hymenoptera, Diptera, 
and predacious insects attack both the gipsy moth and brown-tail 
moths in Massachusetts ; they are also devoured by birds, toads, and 
other insectivorous animals. 

Brooks, Frep E. 

1907. The grapevine root-borer. Bull. 110, West Virginia Agr. Exp. Sta., 
pp. 19-30, 5 pls., Nov. Memythrus polistiformis family Sesiidae. 
Great crested flycatcher feeding upon it (p. 28). 

BrYANT, Harorp C. 

1911. The relation of birds to an insect outbreak in northern California 
during the spring and summer of 1911. The Condor, vol. 13, 
pp. 195-208, Nov. Stomach examination revealed that four species 
of birds fed upon the butterflies (Eugonia californica), which 
formed an average of 32.8 per cent of their food. In addition a 
western flycatcher was observed feeding upon them. Chickens and 
ducks also reported as taking numbers of these insects. 





NO. 7 PROTECTIVE ADAPTATIONS—McATEE 167 


Burcess, A. F., AND CrossMAN, S. S. 

1929. Imported insect enemies of the gipsy moth and the brown-tail moth. 
U. S. Dep. Agr. Techn. Bull. 86, 147 pp., 6 pls., 55 figs. More 
than 93 million parasites and predators liberated and a high degree 
of control was obtained by 10924; parasite population fluctuates. 
Heteroptera, dermestids, spiders, nematodes, mites, birds, and fungi 
contribute to the mortality, as do starvation and severe weather. 

CHITTENDEN, F. H. 

1916. The common cabbage worm. Farmers’ Bull. 766, U. S. Dep. Agr., 
p. 9, Nov. The English sparrow, chipping sparrow, and house 
wren are known to feed upon cabbageworms. It is certain that 
other species eat them—one instance noted where the pupae were 
reduced more than 90 per cent by birds. 

1922, List of natural enemies of the celery leaf-tyer (Phlyctaenia rubigalis 
Guen.). Can. Ent., vol. 64, no. 8, p. 174, Aug. Twelve hymenop- 
terous parasites. 

Crark, A. H. 

1926. Carnivorous butterflies. Smithsonian Rep. 1025, pp. 439-508, figs. 
1-5. Lycaenidae. feeding on ants or upon ant-tended insects, as 
aphids, coccids, jassids, and membracids. 

CocKERELL, T. D. A. 

1898. Preliminary notes on the codling moth. Bull. 25, New Mexico Agr. 
Exp. Sta., pp. 55-58. Woodpeckers, and quite likely the kinglet, 
bats, toads, hymenopterous parasites, clerid beetle larvae, and 
parasitic fungi are recorded as enemies of the codling moth. Men- 
tion is also made of the house mouse. 

Cottins, C. W. 

1926. Observations on a recurring outbreak of Heterocampa guttivitta 
Walker and natural enemies controlling it. Journ. Agr. Research, 
vol. 32, no. 7, pp. 689-6900, Apr. 1. List of 15 hymenopterous, 
dipterous, and nematode parasites, and 8 coleopterous, hemipterous, 
and mammalian predators. Bibliography. 

Comstock, J. H. 

1879. Report upon cotton insects. U. S. Agr. Comm. 1879, 511 pp., 3 pls., 
77 figs. Enemies of the cottonworm (pp. 138-214) include hogs, 
dogs, cats, raccoons, bats, wild birds, poultry, spiders, Chrysopa, 
dragonflies, mantis, Hemiptera, robber flies, tiger beetles, ground 
beetles, soldier beetles, ladybirds, boll worms, wasps, ants, chalcid, 
ichneumonid, and tachinid parasites, flesh flies and phorids. 

CooLey, R. A. 

1908. An army cutworm (Chorizagrotis auxiliaris). Bull. 71, Montana 
Exp. Sta. pp. 146-147. Several species of wild birds, domestic 
fowls, parasitic flies and wasps, besides beetles, are named as foes. 

1930. The codling moth. Bull. 42, Montana Exp. Sta., p. 7. Birds con- 
sidered to be great destroyers of this insect. 

CrIDDLE, NORMAN. 

1920. Fragments in the life-habits of Manitoba insects—II. Can. Ent., 
vol. 52, no. 6, pp. 121-125, July. Proteopteryx oregonana checked 
by parasites, Calosoma, and birds. 





168 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


Crug, S. E. 

1926. Tobacco cutworms and their control. Farmers’ Bull. 1494, U. S. 
Dep. Agr., 13 pp., 11 figs, Aug. Among the agencies of natural 
control are mentioned spiders, ground beetles, birds, and toads. 
Birds play an important part. 

[Davis, W. T.] 

1914. [Dragonflies eating butterflies.] Ent. News, vol. 25, no. 4, p. 191, 
Apr. Mr. W. T. Davis said “ That the dragonflies, especially on 
the west coast of Florida, were quite a nuisance to collectors on 
account of their catching many of the smaller butterflies that were 
disturbed.” 

Dustan, ALAN G. 

1923. The natural control of the white-marked tussock moth under city 
and forest conditions. Proc. Acadian Ent. Soc., vol. 8, 1922, 
pp. 109-126, pls. 15-16. In the city the principal enemies are para- 
sites of the eggs and larvae, while in the forest, birds, ants, and 
spiders assume that role. - 

FELT, He ee 

1912. [Green maple worm (Xylina antennata Walk.)]. 27th Rep. State 
Entomologist, 1911, New York State Mus. Bull. 155, pp. 50-51, 
Jan. Nine species of birds actually observed eating or carrying 
away caterpillars, and nine others apparently associated in the work. 

FLOERSHEIM, C. 

1906. On some enemies of the diurnal Lepidoptera. Ent. Rec. vol. 18, 
no. 2, pp. 36-39, Feb. Two cases of birds; predacious beetles very 
abundant and get many sleeping butterflies. Spiders and coccinellid 
larvae eat butterfly larvae; eggs destroyed by Hemiptera. 

Forses, S. A. 

1883. The regulative action of birds upon insect oscillations. Bull. Illinois 
State Lab. Nat. Hist., vol. 1, no. 6, pp. 1-32, May. Chiefly on 
birds in relation to an outbreak of cankerworms which made up 
35 per cent of food of all birds in the affected orchard. Schedule 
of all food items identified. 

ForsusH, E. H. 

1899. The destruction of hairy caterpillars by birds. Bull. 20, Div. Ent., 
U. S. Dep. Agr., pp. 85-93. List of 46 species with detailed notes 
on feeding habits of some of them, especially in relation to gipsy 
moth and brown-tail moth larvae, and tent caterpillars. 

GarMAN, H. 

1895. [Cutworm enemies]. Bull. 58, Kentucky Exp. Sta., p. 106, Nov. 

Birds, chickens, turkeys, and pigs, besides insect parasites. 
Gipson, ARTHUR. 

1915. The army-worm. Dominion of Canada Dep. Agr., Ent. Branch, 
Bull. 9, 34 pp., 19 figs. Cirphis unipuncta. Many species of wild 
birds, large numbers of parasitic and predacious insects, domestic 
poultry, toads, and skunks recorded as enemies. Bacterial and 
fungous diseases also recorded attacking this worm (pp. 13-17). 





NO. 7 PROTECTIVE ADAPTATIONS—McATEE 169 


GILL, JouHN B. 

1913. The fruit-tree leaf roller. U. S. Dep. Agr. Bull. 116, pt. 5, pp. 91-110, 
pls. 12-16, Mar. Archips argyrospila—eight species of birds, para- 
sitic Hymenoptera and Diptera and predacious beetles and ants 
recorded as enemies; a small mite also noted feeding upon the 
eggs of the leaf-roller (p. 102). 

GILLETTE, CLARENCE P, 

1905. The beet webworm. Colorado Agr. Exp. Sta., Bull. 98, pp. 3-12, 2 pls., 
Mar. Lo-vostege sticticalis. Records of insect-eating birds devour- 
ing these in quantities, mention being made of large flocks oi 
blackbirds. Parasitic Hymenoptera also noted (pp. I0-I1). 

HARDENBERG, C. B. 

1912. The willow tree caterpillar (Angelica tyrrhea, Cramer). Agr. 
Journ. South Africa, vol. 4, no. 3, pp. 397-418, Sept. Parasitic 
flies and wasps attack the caterpillar and eggs. The larvae are 
reported to be distasteful to birds although they are said to be 
eaten by some tribes of Kaffirs. Guinea fowls and meerkats feed 
upon the pupae, and moles probably do so. They are also attacked 
by a fungus (pp. 412-416). 

Haskin, J. R. 

1916. Butterflies as food for squirrels. Ent. News, vol. 27, no. 8, p. 370, Oct. 
Melitaea chalcedon, evidence of destruction of 25 or more by gray 
squirrels; California. 

Herrick, GLENN W. 

1910. The snow-white linden moth. Bull. 286, Cornell Agr. Exp. Sta., pp. 
51-64, figs. 54-58, Nov. English sparrow freed cities of this pest— 
Ennomos subsignarius (p. 62). 

Horton, J. R. 

1922. A swallow-tail butterfly injurious to California orange trees (Papilio 
zolicaon Boisd.). Monthly Bull. Dep. Agr. California, vol. 11, 
no. 4, pp. 377-387, Apr. Larvae of Chrysopa californica sometimes 
destroy 80-90 per cent of the eggs. Young caterpillars eaten by 
same foe, also by Zelus renardi, and a hymenopterous parasite; 
the pupa by Chalcis ovata (p. 385). 

Howarp, L. O. 

1904. The insect book. The Nature Library, vol. 8, pp. 56-57. Parasitic 
Hymenoptera attacking the cotton caterpillar; an instance of de- 
struction of 95 per cent of the eggs. 

Howarp, L. O. AND CHITTENDEN, F. H. 

1907. The catalpa sphinx (Ceratomia catalpae Bdy.). Cire. 96, U. S. Bur. 
Ent., p. 6. Cuckoos, the catbird and the Baltimore oriole recorded 
as enemies. 

1909. The green-striped maple worm (Anisota rubicunda Fab.). Circ. 110, 
U. S. Bur. Ent., p. 5. Domestic fowls and nine species of wild 
birds recorded as enemies. 

Jounson, E. E. 

1926. Birds eating butterflies. The Field, London, vol. 147, p. 658, Apr. 15. 

Observations made of a stonechat taking butterflies. 


170 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


KERSHAW, JOHN C. W. 
1905. Butterfly-destroyers in Southern China. Trans. Ent. Soc. London, 


1905, pp. 5-8. Has seen only a dozen attacks by birds in five years; | 


lizards destroy most adults. Notches in wings made by striking 
twigs, etc., a cuckoo the worst enemy of larvae, taking even the 
hairy and most conspicuous kinds. Also ants fastening on butterfly 
tongues. 

KirKLAND, A. H. 


1896. The army worm. Massachusetts Crop Rep. 1896, pp. 34-36. Birds 
are recognized as being the most important enemies of the army 
worm. Toads, parasitic flies, and beetles also are recorded as being 
enemies of this pest. 

LamBorn, W. A. 


1912. Butterflies a natural food of monkeys. Trans. Ent. Soc. London, 

1912, p. iv. Mangabeys eating butterflies at mudholes. 
Lawson, G. 

1888. Insect injuries to field and orchard crops. Provincial Crop Rep. 
Nova Scotia, 1888, p. 29. American and forest tent caterpillars 
are eaten by the crow and cuckoo and to a lesser extent by the 
linnet and swallow; cankerworms by the linnet (purple finch?). 

LINTNER, J. A. 

1888. Cutworms. Bull. 6, New York State Mus., 36 pp., 28 figs. Natural 
enemies include wild birds, poultry, toads, ground beetles, preda- 
cious bugs, mites, spiders, parasitic Diptera and Hymenoptera 
(pp. 23-28). 

LounssBury, C. P. 

1895. Canker worms, army worms, etc. Bull. 28, Hatch Exp. Sta., p. 15. 

Poultry, birds, frogs, toads, beetles, and parasitic flies listed as foes. 
LucceEr, OTTOo. 

1892. Tent-caterpillars. Ann. Rep. Minnesota Hort. Soc., p. 372. Cuckoos, 

skunks, and Calosoma are enemies. 
MaLioneg, A. M. 


1916. Frogs catching butterflies. Science, n. s., vol. 43, pp. 386-387, Mar. 17. 

Rana catesbiana eating a number of Papilio turnus. 
MaALtty, F. W. 

1893. Report on the boll worm of cotton (Heliothis armiger Hubn.). Bull. 
29, U. S. Div. Ent., p. 26. Woodpeckers and sparrows reported as 
enemies. 

MAnopers, N. 

1912. [Enemies of Danais chrysippus.] Trans. Ent. Soc. London, p. 446. 
Trichogramma evanescens parasitizes large numbers of eggs; ants 
eat the eggs; larvae are commonly parasitized; spiders and ants 
eat them; they are cannibalistic; adults eaten by lizards, and 
sometimes by birds. 

Martatt, C. L. AND Orton, W. A. 

1906. The control of the codling moth and apple scab. Farmers’ Bull. 247, 
U. S. Dep. Agr., p. 9. Woodpeckers noted as preying upon the 
codling moth. 









INO. 7 PROTECTIVE ADAPTATIONS—McATEE 7 


McAtTEE, W. L. 

1912. Bird enemies of the codling moth. Yearbook U. S. Dep. Agr. 1911, 
pp. 237-246. Birds recognized as most effective natural enemies— 
from 66 to 85 per cent of the hibernating larvae recorded as being 
destroyed. 36 species of birds known to prey upon this moth. 

1923. Another insect birds should not eat. The Auk, vol. 40, no. 3, p. 560, 
July. Red-humped apple caterpillar (Schizura concinna)—recorded 
as preyed upon by six species of birds. 

1924. Birds as factors in the control of the fall webworm. The Auk, vol. 
41, no. 2, p. 372, Apr. Results of eight years’ study in Canada, 
red-eyed vireo destroys 11.4 to 80.5 per cent of the broods, averag- 
ing more than 68 per cent; birds “tremendously important” 
in control of the insect. 

1925. Economic ornithology. The Auk, vol. 42, no. 3, pp. 464-465, July. 
European corn borer—This pest is recorded as preyed upon by 
four species of birds. 

1926. Birds feeding on the European corn borer. The Auk, vol. 43, no. 3, 
p. 395, July. Red-wing blackbirds and downy woodpeckers re- 
corded as feeding on the larvae. 

1927. Economic ornithology in recent entomological publications. The 
Auk, vol. 44, no. 3, pp. 458-450, July. European corn borer re- 
corded as taken by six species of birds. 

NEAL, H. V. 

1912. Monkeys eating butterflies. Trans. Ent. Soc. London, 1912, pp. xvii- 

xviii. Commonly do so in Lagos. 
OTANES, F. O. 

1925. The rice stem borer (Schoenobius incertellus Walker). Philippine 
Agr. Rev., vol. 18, no. I, pp. 81-82. “The adult moths are said 
to be preyed upon by birds, mudfish (dalag), spiders, frogs, and 
mantids” (p. 82). 

BACK, Hi. J. 

1922. Toads in regulating insect outbreaks. Copeia, no. 107, pp. 46-47, 
June 20. Feeding exclusively on sugar-beet webworms and taking 
from 20 to 40 each. 

PatcH, EpitH M. 

1908. The saddled prominent, Hfeterocampa guttivilta (Walker). Maine 
Agr. Exp. Sta., Bull. 161, pp. 311-350, figs. 14-40, Nov. Predacious 
bugs and beetles, hymenopterous parasites, a fungus, skunks, 
domestic fowls, and wild birds recorded as enemies. (Pp. 340-348.) 

1921. A meadow caterpillar. Bull. 302, Maine Agr. Exp. Sta., pp. 300-320, 
2 pls., Dec. Ctenucha virginica. One dipterous and 5 hymenopter- 
ous parasites. “ Though covered with stiff hairs, the over-worked 
theory that such caterpillars are thereby immune from birds’ attacks 
cannot stand up against the testimony of my pet thrush which 
whips these caterpillars vigorously against the floor of the cage 
until, in a surprisingly short time, their bodies are beaten limp 
and naked, whereupon they are swallowed in one gulp.” 

berrit, I. EL. 

1904. The codling moth in Michigan. Bull. 222, Michigan Agr. Exp. Sta., 
pp. 78 and 89-90. Birds most important; shrews, parasitic Hymen- 
optera and fungi also mentioned. 

12 


172 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


Puitiips, W. J. and Ktnc, Kennetu M. 
1923. The corn earworm. Farmers’ Bull. 1310, U. S. Dep. Agr., 17 pp., 
18 figs., Jan. Heliothis obsolcta. Seventeen species of birds, egg 
parasites, and other parasitic insects, certain ants, and spiders 
recorded as foes. The earworms’ cannibalistic habit is the most 
important factor in reducing its attacks on corn (pp. 10-12). 
Purpps, C. R. 
1927. The black army cutworm. Maine Agr..Exp. Sta., Bull. 340, pp. 201- 
216, figs. 29-30, May. Agrotis fennica. Three kinds of flies and 
one of wasp parasites, predators including bugs, beetles, wasps, 


and birds recorded as enemies (pp. 212-213). 
PRANK) Elke 


1929. Natural enemies of the sugar cane moth stalkborer in Cuba. Ann. 
Ent. Soc. Amer., vol. 22, pp. 621-640, 7 figs. Its burrowing habit 
is “an admirable protection against direct, or artificial, but does 
not seem to afford any great amount of protection against the 
attack of a rather formidable array of natural enemies.” Tachinids, 
sarcophagids, and Hymenoptera recorded as parasites, and earwigs, 
beetles, and ants as predators. Bibliography. 

Poos, F. W. 

1928. An annotated list of some parasitic insects. Proc. Ent. Soc. 
Washington, vol. 30, no. 8, pp. 145-150, Nov. Parasites varying 
from r to 14 in number bred from each of 19 hosts, mostly 
Lepidoptera. 

Poutton, E. B. 

1911. The attacks of tachinid flies upon the African Danaine genus Amauris. 
Trans. Ent. Soc. London, tort, p. xcix. Twenty out of 25 pupae 
of Amauris psyttalea parasitized; another lot all parasitized. This 


a good indication enemies of adults are scarce; otherwise species 
would be rare. 


QUAINTANCE, A. L. 


1908. The apple-tree tent caterpillar (J/alacosoma americana). Circ. 08, 
U. S. Bur. Ent., p. 6. Black-billed and yellow-billed cuckoos, blue- 
jay, crow, chickadee, Baltimore oriole, red-eyed vireo, chipping 
sparrow, and yellow warbler mentioned as enemies; also the com- 
mon toad. 


SANDERSON, E. D. 


1903. The codling moth. Bull. 59, Delaware Agr. Exp. Sta., pp. 7-8. 
Parasitic worms and insects, soldier beetles, named as enemies as 
well as 10 or more species of birds which are the most efficient. 

1905. The gipsy moth in New Hampshire. Bull. 121, New Hampshire 
Agr. Exp. Sta. p. 99. Ground beetles, parasitic insects, and 
several species of birds are recorded as preying upon the gipsy 
moth. 

1906. The brown-tailed moth in New Hampshire. Second report. Bull. 
122, New Hampshire Agr. Exp. Sta., p. 127. Parasitic insects, 
predacious bugs, toads, bats, and several species of birds, particu- 
larly the English sparrow, recorded as enemies. 





ae SSS 








INO©:, 7 PROTECTIVE ADAPTATIONS—McATEE 173 


1909. The codling moth and how to control it by spraying. Bull. 143, 
New Hampshire Agr. Exp. Sta., pp. 64-82. Downy woodpeckers 
and nuthatches are recorded as being the most important enemies 
of the codling moth. 

SANDERSON, E. D., HEApLEE, T. J., AND Brooks, CHARLES. 

1907. Spraying the apple orchard. Bull. 131, New Hampshire Agr. Exp. 
Sta., pp. 18-19 and 35. Woodpeckers and nuthatches are recorded as 
feeding extensively on the codling moth. 

SAUNDERS, ARETAS A. 

1916. A note on the food of the western robin. The Condor, vol. 18, no. 9, 
p. 81, Mar.-Apr. Robin feeding on Papilio rutulus, and a chipmunk 
feeding upon the same species and also on P. eurymedon. 

SHARP, D. 

1910. [Parasites of winter moth.] Cambridge Nat. Hist., vol. 5, p. 521. 
“The destructive winter moth—Chetmatobia brumata—is known 
to be subject to the attacks of 63 species of Hymenopterous para- 
sites. So abundant are these latter that late in the autumn it is 
not infrequently the case that the majority of caterpillars contain 
these destroyers.” 

SHERMAN, F. 

1921. Observations of natural enemies of the fall cankerworm (Alsophila 
pometaria Peck) in forests of southern Alleghany Mountains in 
1920. Journ. Econ. Ent., vol. 14, no. 6, pp. 478-481, Dec. Names 
15 species of birds presumably of material help, five predacious 
insects, and 3 parasites of which one destroyed from 25 to 4o per 
cent of the eggs. 

SHIRAS, GEO., 3RD. 

1921. Frogs eating butterflies. Nat. Geogr. Mag., vol. 40, no. 2, p. 174, 
Aug. Leopard frogs catching about 500 “blues” in a week; also 
eating many Argynnis aphrodite. 

SxkalrFe, S. H. 

1921. Some factors in the natural control of the wattle bagworm. South 
African Journ. Sci., vol. 17, nos. 3-4, pp. 291-301, July. Acan- 
thopsyche junodi Heylaerts. Out of a total of 50,687 examined, 
just over one per cent were destroyed by birds and rats, 19 per 
cent by insect parasites, 16 per cent by fungous disease, and 17 
per cent by other diseases. Only one-quarter of one per cent 
survive the early perils of their life. 

SmirTH, J. B. 

1895. [Zeusera pyrina.] 15th Ann. Rep. New Jersey Exp. Sta. 1894, pp. 
531-532. Almost all insectivorous birds, especially woodpeckers and 
the sparrow, in addition to bats, and predacious insects, are re- 
corded as enemies of this insect. 

SPENCER, G. J. AND H. G. Crawrorp. 

1923. The European corn borer in Ontario. Ontario Dep. Agr. Bull. 2095, 
II pp., 10 figs., Mar. Ants, aphis-lions, ladybird beetles, ground 
beetles, crickets, a parasitic fly, and several species of birds noted 
as foes. One instance recorded of downy and hairy woodpeckers 
destroying 60 per cent of the borers (pp. 7-8). 






174. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


ToTHILL, JoHn D. 


1922. The natural control of the fall webworm [Hyphantria cunea Drury] 
in Canada together with an account of its several parasites. Dep. 
Agr. Dominion of Canada, Bull. 3, n. s. (techn.), 107 pp., 6 pls., 
99 figs. Tabulations of the destruction by various enemies in differ- 
ent localities and years; birds average most important, parasites 
sometimes important, sometimes not. 

1923. Notes on the outbreaks of spruce budworm, forest tent caterpillar, 
and larch sawfly in New Brunswick. Proc. Acadian Ent. Soc., 
vol. 8, 1922, pp. 172-182. Spruce budworm. Natural checks effective 
in New Brunswick were eggs, larvae, and pupal parasites, spiders, 
and birds; and in British Columbia, parasites and birds. Nothing 
of importance on enemies of the other forms. 

TROUVELOT, LEOPOLD. 

1868. The American silkworm. Amer. Nat., vol. 1, pp. 30-38, 85-94, 145- 
149. Telea polyphemus. Thrushes, catbirds, orioles; 95 out of a 
hundred worms become the prey of these feathered insect hunters. 

U. S. ENtTomMoLocIcAL CoMMISSION. 


1883. 3rd Rep. U. S. Ent. Comm., p. 125. All insectivorous birds, hogs, 
chickens, turkeys, toads, and frogs prey upon the armyworm. 
“The worms themselves, when hard pushed, will even devour each 
other.” 

1883. 3rd Rep. U. S. Ent. Comm., pp. 175-178. Forty or more species of 
wild birds, notable mention being made of bluebirds, cedarbirds 
and butcherbirds, and parasitic and predacious insects, also hogs, 
are recorded as enemies of the cankerworm. 

1885. 4th Rep. U. S. Ent. Comm., pp. 87-90. More than 20 species of wild 
birds, poultry, hogs, raccoons, skunks, opossums, bats, tree frogs, 
lizards, spiders, and numerous kinds of predacious insects are 
recorded as preying upon the cottonworm. 

Vickery, R. A. : 

1929. Studies on the fall army worm in the Gulf Coast District of Texas. 
Techn. Bull. 138, U. S. Dep. Agr., 63 pp. Numerous hymenopterous 
parasites sometimes destroy 40-50 per cent of the caterpillars. 

Warren, B. H. 

1897. The army worm. Ann. Rep. Pennsylvania State Coll. 1806, pp. 164- 
220, 16 pls. Much on natural enemies including tachinids and 
ichneumonids, ground beetles, birds, mammals, and toads. 

Wesster, R. L. 

1909. The lesser apple leaf-folder. Iowa State Coll. Exp. Sta. Bull. 102, 
pp. 181-212, figs. 1-13, Mar. Peronea minuta—tachinid and hymen- 
opterous parasites recorded as the most important natural enemies 
although birds and diseases are also important factors (pp. 
206-211). 

WEED, C. M. 

1809. The forest tent caterpillar. Bull. 64, New Hampshire Agr. Exp. 
Sta., pp. 77-98, figs. 20-33, Apr. Clisiocampa disstria—preyed upon 
by insects, spiders, toads, and birds. Ten kinds of birds feeding 
on larvae, one on the cocoons and four on the adults. 





NO. 7 PROTECTIVE ADAPTATIONS—McATEE 175 


1899. The spiny elm caterpillar. Bull. 67, New Hampshire Agr. Exp. Sta., 
pp. 125-141, figs. 40-51, Oct. Vanessa antiopa—egg parasites 
chalcid, ichneumonid and tachinid flies on caterpillars; Calosoma, 
wasps, cuckoos, and toads named as enemies (pp. 138-140). 

1900. Insect record for 1899. Bull. 72, New Hampshire Exp. Sta., pp. 

64-65. The Baltimore oriole and the cuckoos are especially noted 

as enemies of tent caterpillars. 

1900. The forest tent caterpillar. Second report. Bull. 75, New Hampshire 
Exp. Sta., pp. 120-121. Eighteen or more species of birds are 
recorded as preying upon these caterpillars. 

1902. [Enemies of cankerworms.] Bull. 90, New Hampshire Agr. Exp. 
Sta., 1902, p. 35, Mar. Robins, bluebirds, cedarbirds, and many 
others feed freely upon the pests. 

West, L. S. 

1923. Immunity to parasitism in Samia cecropia Linn. (Lep.: Saturntidae; 
Dip.: Tachinidae.) Ent. News, vol. 34, no. 1, pp. 23-25, Jan. Ineffec- 
tiveness of attack of 35-40 tachinid larvae; nevertheless lists two 
Tachinidae and eight species of Hymenoptera that do successfully 
parasitize this host. 

YorHers, M. A. 

1913. Eugonia californica Bdy. in the Pacific Northwest. Can. Ent., vol. 
45, no. 12, pp. 421-422, Dec. “I think that the total disappearance 
of these caterpillars and chrysalids was no doubt due to birds” 
(p. 422). 

Youn, R. A. 

1907. Insects affecting the poplar. Proc. Columbus Hort. Soc. 1906, pp. 
68-82. Birds constitute an important agency in keeping the 
Hemerocampa leucostigma in check (p. 74). 


COLEOPTERA 


Acassiz, L., AND Casot, J. ELtior. 

1850. Lake Superior, etc., p. 72. Monochamus scutellaris preyed upon by 

Canada jay and two species of Picoides. 
BiackMaNn, M. W. 

1915. Observations on the life history and habits of Pityogenes hopkinsi 
Swaine. Techn. Publ. no. 2, New York State Coll. Forestry, pp. 
11-66, 6 pls., Nov. Natural enemies include beetles, mites, and 
parasitic Hymenoptera (pp. 53-560). 

Britten, H. 

1926. A pentatomid bug preying on beetle larvae. North Western Nat., 
vol. 1, p. 38. Rhacognathus punctatus found sucking larvae of 
beetle (Hydrothassa marginella). 

1927. Red ants and beetles. North Western Nat., vol. 2, p. 256. Myrmica 
ruginodis killing beetles (Melandrya caraboides). 

Brooks, F. E. 

1919. The flat-headed apple-tree borer. Farmers’ Bull. 1065, U. S. Dep. 
Agr., 12 pp., 13 figs. Chrysobothris femorata. Woodpeckers, and 
other birds, ants, and six species of hymenopterous parasites re- 
corded as enemies (p. 9). 






176 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 85 


Burcess, A. F. Anp CoLitns, C. W. 

1917. The genus Calosoma, including studies of seasonal histories, habits, 
and economic importance of American species north of Mexico and 
of several introduced species. Bull. 417, U. S. Dep. Agr., 124 pp., 
19 pls. 5 figs. Caterpillars the principal food of these beetles; 
cannibalism, and attacks of toads, skunks, and birds the chief 
organic checks; predatory bugs, and parasitic flies also noted 
(pp. 10-13). 

CARTWRIGHT, OSCAR L. 

1929. The maize billbug in South Carolina. Bull. 257, South Carolina 
Agr. Exp. Sta., 35 pp., 13 figs. May. Natural enemies (p. 31) 
include egg parasite, predacious beetles, ants, and wasps. 

CHAMBERLIN, F. S., AND TENHET, J. N. 

1923. The tobacco flea-beetle in the southern cigar-wrapper district. 
Farmers’ Bull. 1352, U. S. Dep. Agr., 9 pp., 8 figs. Epitrix parvula. 
A spider, lygaeid bug, and birds noted as enemies (p. 5). 

CHITTENDEN, F. H. 

1911. Notes on various truck-crop insects. Bull. 82, pt. 7, U. S. Bur. Ent., 
pp. 85-93, fig. 24. Natural enemies of Leptinotarsa decemlineata, 
pp. 85-88; 1 beetle, 3 hemiptera, 16 wild birds, and guinea fowls. 

1926. Notes on the behavior of Cotinis nitida L. and its bird enemies. Proc. 
Biol. Soc. Washington, vol. 39, pp. 15-17, Feb. Starling and 
cardinal eat it. 

CHITTENDEN, F. H., anp Fink, D. E. 

1922. The green June beetle. Bull. 891, U. S. Dep. Agr., 52 pp., Io pls., 
7 figs. Cotinis nitida. Natural enemies (pp. 31-37) include para- 
sitic flies, digger wasps, ground beetles, mites, various mammals, 
and birds; fungal and bacterial diseases also noted. 

CHITTENDEN, F. H., anp Marsa, H. O. 

1920. The western cabbage flea-beetle. U. S. Dep. Agr. Bull. 902, 21 pp., 
4 figs., 1 pl., Oct. Phyllotreta pusilla. Hymenopterous and worm 
parasites recorded as well as 12 species of birds feeding on beetles 
of this genus. 

Davis, JoHN J. 

1913. Common white grubs. U. S. Farmers’ Bull. 543, 20 pp., 12 figs., 
July. More than 60 species of birds, domestic fowls, skunks, a 
number of predacious and parasitic insects recorded preying upon 
white grubs at one stage or another (pp. 13-15). 

FENTON, F. A., AND DunHAM, E. W. 

1929. Biology of the cotton boll weevil at Florence, S. C. Techn. Bull. 
112, U. S. Dep. Agr., 75 pp., 35 figs. Considerable variation exists 
in mortality rate from parasitism (by three species of Hymen- 
optera) from as low as 2.37 to as high as 51.52 per cent; predators, 
heat, proliferation by the plant, disease and unknown causes take 
their toll, the average total from all these causes being about 40 
per cent; then from the number that go into hibernation only an 
average Of 3.27 per cent survive. 


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NO. 7 PROTECTIVE ADAPTATIONS—McATEE 177 


FrEYTAUD, JEAN. 

1922. Le Doryphore, Chrysomcle nuisible a la pomme de terre (Leptino- 
tarsa decemlineata Say). Rev. Zool. Agr. Appl., vol. 21, Numero 
special, 48 pp., 13 figs., Aug. Natural checks include skunks, birds, 
snakes, frogs, spiders, phalangids, mites, beetles, bugs, wasps, robber 
flies, and parasitic flies (pp. 14-17). 

Forses, S. A. 

1880. Notes upon the food of predacious beetles. Bull. Illinois State Lab. 
Nat. Hist., vol. I, no. 3, pp. 149-152, Nov. Both vegetable and 
animal, the latter including beetles, larvae, and plant lice. 

1880. Notes on insectivorous Coleoptera. Bull. Illinois State Lab. Nat. 
Hist., vol. 1, no. 3, pp. 153-160, Nov. Carabidae, Lampyridae, 
Coccinellidae, from stomach examination. Animal food, mites and 
their eggs, ants, caterpillars, beetles and their larvae, plant lice, 
and centipeds. 

1883. The food relations of the Carabidae and Coccinellidae. Bull. Illinois 
State Lab. Nat. Hist., vol. 1, no. 6, pp. 33-64, May. Report on 
dissections of 175 Carabidae and 39 Coccinellidae. Animal food 
included Hymenoptera, Lepidoptera, Diptera, Neuroptera, and 
Coleoptera, spiders, mites, myriapods, mollusks. Notes on birds 
as enemies of Cicindelidae and Carabidae. 

1907. On the life history, habits, and economic relations of the white-grubs 
and may beetles. Bull. 116, Illinois Agr. Exp. Sta., pp. 447-480, 
Aug. Principal enemies, swine, crows, blackbirds, and Tiphia; 
other parasites Macrophthalma, Sparnopolius, Pyrgota, and Ophion 
(pp. 468-475). 

ForsusH, E. H. 

1912. 4th Ann. Rep. State Ornithologist Mass., 1911, 32 pp., 4 pls., 9 figs. 
Galerucella luteola. Cedar waxwing clearing trees of infestations 
of the elm leaf beetle (pp. 19-20). 

Hess, WALTER N. 

1920. The ribbed pine borer. Mem. 33, Cornell Agr. Exp. Sta., pp. 367- 
381, pl. 8, figs. 61-66. Rhagium lineatum. Woodpeckers, most im- 
portant; a parasite reared (pp. 378-379). 

Hopkins, A. D. 

1896. The relation of insects and birds to present forest conditions. Proc. 
Amer. Forestry Assoc., vol. 11, pp. 175-176. Woodpeckers recorded 
as enemies of bark and clerid beetles. 

Hystop, JAMES A. 

1912. The false wireworms of the Pacific Northwest. Bull. 95, pt. 5, U. S. 
Bur. Ent., pp. 73-87, figs. 22-27. Numerous species of birds, 
horned-toads, garden toads, skunks, parasites, and disease re- 
corded as enemies (pp. 84-86). 

1915. Wireworms attacking cereal and forage crops. Bull. 156, U. S. 
Dep. Agr., 34 pp., 8 figs. Elateridae—a long list of bird enemies 
given; horned-toads, mites, predacious flies, hymenopterous para- 
sites, nematodes, fungi (pp. 25-20). 

IncRAM, J. W. 

1927. The striped blister beetle on soy beans. U. S. Dep. Agr. Leafl. 12, 
5 pp., 3 figs. Epicauta lemniscata—three species of birds and a 
robber fly named as enemies. 


178 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


KALMBACH, E. R. 


1914. Birds in relation to the alfalfa weevil. Bull. 107, U. S. Dep. Agr., 
64 pp., 5 pls. Forty-five species of birds found to feed on this 
comparatively recently introduced pest; also domestic fowls, toads, 
frogs, salamanders, horned-toads, snakes, and shrews. 

McAteer, W. L. 

1914. Bird enemies of Diabroticas. The Auk, vol. 31, no. 1, p. 120, Jan. 
Southern corn root worm (Diabrotica duodecim-punctata) recorded 
as preyed upon by 24 species of birds; western corn root worm 
(Diabrotica longicornis) taken by the nighthawk and the wood 
pewee. 

1915. Bird enemies of two beetle pests. The Auk, vol. 32, no. 3, pp. 377- 
378, July. Oncideres putator—it is believed that the southern downy 
woodpecker and the Texas woodpecker attack the larvae of this 
pest. Monocrepidius vespertinus—the records show that these 
beetles are devoured by eight species of birds. 

MaiL, G. ALLEN. 

1930. Winter soil temperatures and their relation to subterranean insect 
survival. Journ. Agr. Research, vol. 41, no. 8, pp. 572-592, Oct. 15. 
Few parasites; mites, birds, fungal and bacterial diseases reduce 
them, but climate a control factor of much importance. 





Murr, F. 


1917. The introduction of Scolia manilae Ashm. into the Hawaiian Islands. 
Ann. Ent. Soc. Amer., vol. 10, no. 2, pp. 207-210, June. A parasite 
of the beetles Anomala orientalis and Adoretus tenuimaculatus. 

[NeEtson, E. W.] 

1921. Report of chief of Bureau of Biological Survey, 34 pp. Bird enemies 
of the Japanese beetle (Popillia japonica) mentioned (p. 14); five 
species of birds and the toad listed. 

QUAINTANCE, A. L., AND JENNE, E. L. 

1912. The plum curculio. Bull. 103, U. S. Bur. Ent., 250 pp., 20 pls., 33 
figs. Natural enemies (pp. 139-154) include an egg parasite, 
hymenopterous and dipterous parasites of later stages, ants, chry- 
sopids, carabids, lampyrids, fowls and wild birds; also the toad. 

SATTERTHWAIT, A. F. 

1919. How to control billbugs destructive to cereal and forage crops. 
Farmers’ Bull. 1003, U. S. Dep. Agr., 23 pp., 24 figs. Insect, 
worm, and fungus parasites, toad and bird predators mentioned, 
the birds apparently most important (pp. 19-20). 

SCHUSTER, W. 

1909. [Beetles and their enemies in the bird world.] Ent. Blatt. Nurnberg, 
vol. 5, no. 7, pp. 142-144, July 15. Birds the principal enemies of 
beetles; notes on European bird foes of various families of beetles ; 
similar notes for Lepidoptera. 

SLINGERLAND, M. V. 

1906. The bronze birch borer: an insect destroying the white birch. Bull. 
234, Cornell Agr. Exp. Sta., pp. 65-78, figs. 31-38. Agrilus anxius. 
Woodpeckers and chalcid parasites mentioned as foes. 





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PROTECTIVE ADAPTATIONS—McATEE 179 


se 


1931. The predatory enemies of Elateridae (Coleoptera). Ent. News, vol. 


Wesp, J. L. 


42, no. 5, pp. 137-140, May; no. 6, pp. 158-167, June. Mites, 
pseudoscorpions, spiders, hemiptera, beetles, flies, hymenoptera, 
amphibians, reptiles, birds, and mammals; birds most important ; 
predators more effective than parasites. Bibliography. 


1906. The western pine-destroying bark beetle. Bull. 58, pt. 2, U. S. Bur. 


Wesster, F. 


Ent., pp. 17-30, pls. 2-3, figs. 7-12. Dendroctonus brevicorms. 
Woodpeckers destroyed large percentage in some trees (p. 27). 
M. 


1880. Notes upon the food of predacious beetles. Bull. Illinois State Lab. 


Nat. Hist., vol. 1, no. 3, pp. 149-152, Nov. Chiefly on vegetarian 
Carabidae but notes on carnivorous species of Carabidae, Staphy- 
linidae, and Lampyridae; the prey mentioned includes plant lice, 
cricket, grasshopper, and beetles. 


1892. Underground insect destroyers of the wheat plant. Bull. Ohio Agr. 


Exp. Sta., vol. 5, no. 9, pp. 221-247, 8 figs., Dec. Wireworms— 
crows, thrushes, robins, blackbirds (p. 228). Whitegrubs—poultry, 
crows, jays, nighthawks, robin, catbird, brown thrasher, wood 
thrush, red-headed woodpecker ; swine, bats, badger, weasel, martin, 
rat, skunk, raccoon, fox, mole, frogs, digger wasps, robber flies, 
and fungi (pp. 236-237). 


1913. The southern corn rootworm, or budworm. U. S. Dep. Agr. Bull. 5, 


II pp. 2 figs., Sept. Diabrotica 12-punctata—12 species of birds 
and parasitic flies (pp. 9-10). 


1913. The western corn rootworm. Bull. 8, U. S. Dep. Agr., 8 pp., 5 figs., 


Sept. Diabrotica longicornis preyed upon by nighthawks, wood 
pewees, a parasitic fly, and chinch bug fungus (p. 6). 


WILDERMUTH, V. L. 
1910. The clover-root curculio. Bull. 85, pt. 3, U. S. Bur. Ent., pp. 29-38, 


WILson, C. 


figs. 15-19. Sitones hispidulus—14 species of birds recorded as 
enemies (p. 37). ; 
B. 


1923-1924. Life history of the scavenger water beetle Hydrous (Hydro- 


philus) triangularis, and its economic relations to fish breeding. 
Bull. Bur. Fisheries, vol. 39, pp. 9-38, 22 figs. Food of larvae, 
snails, midge larvae, fishes, other water beetle larvae, tadpoles, and 
several groups of insects and crustaceans in smaller quantity. Food 
of adults, vegetable matter, fishes. Enemies of Hydrophilus in- 
clude cannibalistic larvae, dragonfly nymphs, frogs, fishes, birds. 
Bibliography. 


1923-1924. Water beetles in relation to pondfish culture, with life histories 


of those found in fishponds at Fairport, Iowa. Bull. U. S. Bur. 
Fisheries, vol. 30, pp. 231-345, figs. 1-143. Larvae highly cannibal- 
istic, dragonfly nymphs are enemies, as are also, mites, hydra, 
ants, fishes, turtles, frogs, and snails; foes of pupae include hymen- 
opterous parasites, horse fly larvae, and ants; of adults, turtles, 
fishes, birds, toads, and frogs. Notes are given on the feeding 


180 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


habits of the larvae and adults of a number of water beetles; fish 
destruction not so apparent as would have been inferred from 
previous literature. Bibliography. 


MECAPTERA 





Predacious. 


DIPTERA 
ALEXANDER, CHARLES P. 

1929. The crane-flies of New York. Mem. 38, Cornell Univ. Agr. Exp. 
Sta., pp. 699-1132, pls. 12-97, June. Ninety-one species of birds, 
besides foxes, mice, shrews, moles, amphibians, fishes, mites, spiders, | 
dragonflies, Diptera, Coleoptera, Hymenoptera, Protozoa recorded | 
as enemies of crane flies in one stage or another (pp. 721-734). 

BroMtey, S. W. | 

1923. Observations on the feeding habits of robber flies. Part I. Psyche, | 
vol. 30, no. 2, pp. 41-45, Apr. Tabulation of the prey of 26 Procta- 
canthus rufus, all Hymenoptera and in 14 cases honey bees. Six 
records for P. brevipennis include three of beetles, one ant, one | 
blow fly, and one assassin bug. 

1930. Bee-killing robber flies. Journ. New York Ent. Soc., vol. 38, no. 2, 
pp. 159-176, pl. 10, June. Especially the honey bee; review of | 
records from various countries; discussion of the U. S. species, 
with notes on other kinds of prey taken by some of them. The 
Dasypogoninae tend to favor Hymenoptera, the Laphriinae beetles, 
while the Asilinae are more general feeders. 

Borris, ALG: | 

1913. Economic and biologic notes on the giant midge: Chironomus 
(Tendipes) plumosus Meigen. Bull. Wisconsin Nat. Hist. Soc., | 
vol. 10, nos. 3-4, pp. 124-163, Apr. Swallows, red-winged black- 
birds as enemies (p. 146); other notes in annotated bibliography 
refer to fishes, birds, Utricularia, and fungi as natural checks. 

1913. Notes on Lake Michigan swarms of chironomids; quantitative notes 
on spring insects. Bull. Wisconsin Nat. Hist. Soc., vol. 11, nos. 1-2, 
pp. 52-69, June. Enemies of adults include mites, spiders, ants, and 
birds (pp. 66-67). 

CocKErRELL, T. D. A. 

1894. On the habits of some Asilidae. Ent. News, vol. 5, no. 6, pp. 173- 
174, June. Mallophora fautrix eating Odynerus sp.; Mallophora 
sp. eating honey bee; Proctacanthus philadelphicus preying upon 
Erax dubius, and butterfly, Synchloe lacinia var. crocale; Procta- 
canthus milberti preying upon Bembex sp. and honey bee; Pro- 
machus princeps preying upon Odynerus annulatus. 

CUTHBERTSON, ALEXANDER. 

1926. The trout as a natural enemy of crane-flies. Scottish Nat., 1926, pp. 

85-88. Salmo fario an important consumer of crane flies in all 
stages; earthworms, phalangids, and click beetles also in the 
stomachs. 

1926. Spiders as enemies of crane-flies. Scottish Nat., 1926, pp. 127-129. 

List of species that eat crane flies, of which the names are given. 
Special study of the prey found in webs of the wood spiders Zilla 
atrica and Z. #-notata. 


NO. 7 PROTECTIVE ADAPTATIONS—McATEE 181 


HARSHBARGER, W. A. 

1894. The bold robber-fly and the mantis. Ent. News, vol. 5, no. 6, p. 169, 
June. Asilid attacked mantis (Stagmomantis carolina) but was itself 
captured and partly eaten. 

HILDEBRAND, S. F. 

1919. Fishes in relation to mosquito control in ponds. Rep. U. S. Comm. 

Fisheries 1918, App. 9, 15 pp., 18 figs. 
Hine, JAMEs S. 

1906. Habits and life histories of some flies of the family Tabanidae. Bur. 
Ent., U. S. Dep. Agr., Techn. Bull., no. 12, pt. 2, pp. 10-38, 12 
figs., Aug. Birds, hornets, and spiders noted as enemies. 

Howarp, L. O. 

1904. The insect book. The Nature Library, vol. 8, pp. 158-159. Outbreaks 
of the armyworm sometimes completely controlled by tachina flies. 
They also attack grasshoppers, bugs, and beetles, sawflies and saw- 
fly larvae and bumble bees and wasps. 

1910. Preventive and remedial work against mosquitoes. Bull. 88, U. S. 
Bur. Ent., 126 pp., June 20. Use of natural enemies, salamanders, 
dragonflies, predacious mosquitoes and fish (pp. 62-72). 

Hystop, J. A. 

1910. The smoky crane-fly. Bull. 85, pt. 7, U. S. Bur. Ent., pp. 119-132, 
figs. 60-66. Tipula infuscata. Natural enemies include a tachinid 
parasite, ground beetles, ants, mites, and birds; a long list is given 
of birds that feed on crane flies; fungi also kill the insects. 

LEATHERS, A. L. 

1922. Ecological study of aquatic midges and some related insects with 
special reference to feeding habits. Bull. U. S. Bur. Fisheries, 
38, Doc. no. 915, 61 pp., 44 figs., May. Food includes Protozoa, 
small Crustacea, diatoms, algae, and other vegetation. 

MatTueson, Roperr. 

1929. A handbook of the mosquitoes of North America. 268 pp., 25 pls. 
Food; suck blood of mammals, birds, amphibians, and snakes (pp. 
39-41); enemies, birds, bats, fishes, tadpoles, salamanders, and 
insects (pp. 71-72). 

SmitH, K. M. 

1927. A study of Hylemyia (Chortophila) brassicae Bouche, the cabbage 
root fly, and its parasites. With notes on some other dipterous 
pests of cruciferous plants. Ann. Appl. Biol., vol. 14, pp. 312-330. 
Description of life-history, enemies, and parasites. The larvae of 
a small beetle (Aleochara bilineata) destroy the pupae of the fly ; 
while a cynipid and a braconid parasitize the larvae, which are also 
eaten by the carnivorous larva of an anthomyid fly. The larva 
of the beetle is itself parasitized by a proctotrupid. 

Twinn, C. R. 

1931. Observations on some aquatic animal and plant enemies of mosquitoes. 
Can. Ent. 63, no. 3, pp. 51-61, Mar. Other mosquito larvae, water 
beetle larvae, dragonfly and damselfly nymphs, backswimmers, 
water-scorpions, caddis larvae, salamanders, fishes, hydras, and 
bladderworts. Bibliography. 


182 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


Van Dine, D. L. 
1907. The introduction of top-minnows (natural enemies of mosquitos) 
into the Hawaiian Islands. Press Bull. 20, Hawaiian Agr. Exp. 
Stas) lO pps 3 tigss, July 2s: 
WEED, C. M. 
1902. [Enemies of Bibio albipennis.] Bull. 900, New Hampshire Agr. Exp. 


Sta., pp. 32-33, Mar. Fishes eating those falling in lake; chief 
food of robin in early spring. 


HYMENOPTERA 
BerLtawsky, A. G. 

1927. [Enemies of bees.] Vragi Pchet, Leningrad, 204 pp., 2 pls., 148 figs. 
Mammals, birds, reptiles, amphibians, insects, arachnids, worms, 
and protozoans. 

BEQUAERT, J. 

1922. The predacious enemies of ants. Bull. Amer. Mus. Nat. Hist., vol. 
45, Pp. 271-331, pls. 24-25, Oct. Spiders, beetles, ant-lions, Diptera, 
Hymenoptera, amphibians, lizards, birds and mammals, including 
man, discussed at length. ‘“ There is certainly little or no evidence 
to show that, as the theory is often expressed, ants are unpalatable 
to most insectivorous animals” (p. 271). 

Bicetow, N. K. 


1922. Insect food of the black bear (Ursus americanus.) Can. Ent., vol. 
54, no. 3, pp. 40-50, Mar. Vespula diabolica, V. consobrina, and 
ants; notes on observations of others. 
Davis, Wo. T. 


191g, A remarkable nest of Vespa maculata, with notes on some other 
wasps’ nests. Bull. Brooklyn Ent. Soc., vol. 14, nos. 4-5, pp. 119- 
123, Oct.-Dec. Notes on food habits of Vespa spp., cannibalistic, 
eat flies and damselflies; robber flies are their enemies. 

1924. Oak apple galls destroyed by gray squirrels. Bull. Brooklyn Ent. 
Soc., vol. 19, no. 3, pp. 91-93, I fig., June. Amphibolips confluens 
freely eaten. 

GRAHAM, S. A. 


1928. The influence of small mammals and other factors upon the larch 
sawfly survival. Journ. Econ. Ent., vol. 21, no. 2, pp. 301-310, Apr. 
Lygaeonematus erichsoni. Small mammals, probably Microtus 
chiefly, destroy from 50 to 80 per cent of the hibernating cocoons; 
parasites and fungi about Io per cent. 

GRONEMAN, CArL F. 

1923. Birds as destroyers of gall insects. Audubon Bull. (Illinois Audubon 
Soc.), pp. 13-15, 6 figs., Fall issue. Birds and squirrels recorded 
as enemies. 

HEIKERTINGER, FRANZ. 

1919. Die metoke Myrmekodie. Tatsachenmaterial zur Losung des Mimi- 
kryproblems. Biol. Zentralbl., vol. 30, no. 2, pp. 65-102, Feb. Enemies 
of ants (pp. 81-100), insects, spiders, amphibians, reptiles, mam- 
mals, birds. 





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NO. 7 PROTECTIVE ADAPTATIONS—McATEE 183 


Hersey, J. L. 

1873. Bees and kingbirds. Can. Ent., vol. 5, pp. 159-160. Kingbirds and 
purple martins feed on honey bees, mostly drones; kingbirds feed 
freely on dragonflies also. 

Howarp, L. O. 

1904. [Prey of Proctotrypoidea.] The insect book, p. 51. Gall flies, gall 
gnats, butterflies, moths, beetles, and the eggs of spiders, bugs, 
butterflies, and moths. 

Hunter, W. D. 

1912. Two destructive Texas ants. U. S. Dep. Agr., Bur. Ent. Circe. 148, 
6 pp., Apr. Pogonomyrmex barbatus molefaciens. Eight species of 
birds and the horned lizard recorded as enemies. 

IseLy, Dwicut. 

1913. The biology of some Kansas Eumenidae. Kansas Univ. Sci. Bull., 
vol. 8, no. 7, pp. 235-3090, pls. 34-37, July. Prey includes larvae of 
several families of Lepidoptera, of two of beetles, and of sawflies. 
Bibliography. 

Puitiirs, E. F. 

1917. Beekeeping. Chap. 22, Bee diseases and enemies, pp. 397-416. Three 
diseases of the brood, two of adults; enemies include two wax 
moths, toads, birds, mice, rats, and other small mammals, certain 
spiders and mites, dragonflies, various Hemiptera, the death’s head 
moth, Mediterranean flour moth, a dipterous parasite (Braula 
caeca), blister beetle (Meloe) and other beetles, wasps, hornets, 
and ants. ‘‘ Dragonflies are so destructive to queens as to make 
queen-rearing unprofitable in some places.” 

Suarp, D. 

1910. [Summary of the prey of Fossores.] Cambridge Nat. Hist., vol. 6, 
pp. 92-93. General notes on prey of 16 families of wasps. 

1910. [Prey of Ichneumonidae.] Cambridge Nat. Hist., vol. 5, p. 551. 
“Most of the species, in the larval state, live inside the larvae of 
Lepidoptera, and they thus keep the myriads of caterpillars within 
bounds, the number of these destroyed by ichneumons being pro- 
digious. Some of the family are, however, external parasites, and 
some are known to attack spiders and insects of other Orders than 
Lepidoptera.” 

Swenk, M. H. 

1910. A new sawfly enemy of the bull pine in Nebraska. Rep. Nebraska 
Agr. Exp. Sta., pp. 3-33, 18 figs. Diprion n. sp.—natural checks 
include ichneumonids, tachinids, chipmunks, birds, and a bacterial 
disease. 

Wit.iaMs, F. X. 

1913. Monograph of the Larridae of Kansas. Kansas Univ. Sci. Bull. 
vol. 8, no. 4, pp. 121-213, pls. 22-30, July. Prey includes Orthoptera 
chiefly, but also Hemiptera, and spiders. Bibliography. 

1913. Notes on the habits of some wasps that occur in Kansas. Kansas 
Univ. Sci. Bull., vol. 8, no. 6, pp. 223-230, pl. 33, fig. 1, July. 
Harpactus preying upon Gypona cinerea, Mimesa upon Athysanus 
exitiosus and other jassids; Prionyx upon locusts. 








184. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


ARACHNIDA 
Bitsine, S. W. 

1920. Quantitative studies in the food of spiders. Ohio Journ. Sci., vol. 20, 
no. 7, pp. 215-260, May. Summarizes a large number of observa- 
tions on prey actually seen eaten by spiders, and upon insects 
found in their webs; gives also some experimental results. 

CALveErRT, Puiip P. 

1923. Studies on Costa Rican Odonata. XY. Megaloprepus, its distribution, 
variation, habits, and food. Ent. News, vol. 34, no. 6, (Food), 
pp. 171-174, June. Feeds on spiders. 

Lincecum, G. 

1867. The tarantula killers of Texas. Amer. Nat., vol. 1, no. 3, pp. 137-141, 
May. Pompilus formosus Say feeds on Mygale hentzii and other 
large spiders. 

(COVELL) oat 

1915. Insects captured by the Thomisidae. Can. Ent., vol. 47, no. 4, pp. 115- 
116, pl. 2, Apr. Crab spiders prey upon butterflies, dragonflies, 
wasps, bumble bees, honey bees, and large flies. 

McAtTekr, W. L. 

1911. Bird enemies of the Texas-fever tick and other ticks. The Auk, vol. 
28, no. I, pp. 136-138, Jan. A résumé of seven publications on the 
subject; of the birds mentioned, 12 species are inhabitants of the 
United States. 

Savory, THEO. H. 

1928. The biology of spiders. 376 pp., 16 pls., 121 figs., London. Food 
(pp. 116-125), flies, wasps, bees, ants, beetles, earwigs, butterflies, 
moths, harvestmen, woodlice, and other spiders; more rarely cater- 
pillars, worms, fish, birds. “‘ They show no trace of discrimination.” 
Enemies (pp. 176-179) include birds, toads, lizards, mammals, 
harvestmen, spiders, wasps, and ichneumon flies, and other parasites. 


MOLLUSCA 
Baker, F. C. 

1916. The relations of mollusks to fish in Oneida Lake. Techn. Publ. 
no. 4, New York State Coll. Forestry, 366 pp., 50 figs., one table, 
one map, July. On pp. 154-218 is summarized information on food 
of 54 species of fresh-water fishes, especially in relation to mollusks. 

1918. The relation of shellfish to fish in Oneida Lake, New York. Cire. 21, 
New York State Coll. Forestry, pp. 11-33, figs. 1-16, Aug. Some 
snails carnivorous, eating other snails, leeches, and small fish; 
shellfish form a large part of the food of many species of fishes; 
other enemies of shellfish include flukes, dragonfly nymphs, horse 
fly larvae, water bugs, water beetle larvae, leeches, crawfishes, frogs, 
salamanders, turtles, ducks, other water birds, muskrats, mink and 
otter. 

SEQUAERT, J. 

1925. The arthropod enemies of mollusks, with description of a new dip- 
terous parasite from Brazil. Journ. Parasitol., vol. 11, pp. 201-212, 
fig. 1. Carnivorous snails probably the most important predatory 


a 





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NO. 7 PROTECTIVE ADAPTATIONS—McATEE 185 


enemies; predacious beetles, mites, and dipterous parasites also 
numbered among their foes. Bibliography which is abstracted in 
the paper. 

BisHop, SHERMAN C, 

1921. The map turtle, Graptemys geographica (Le Sueur) in New York. 
Copeia, no. 100, pp. 80-81, Nov. 15. Feeding on Unio complanatus. 

CHURCHILL, E. P., AND LEwis, SARA I. 

1924. Food and feeding in fresh-water mussels. Bull. U. S. Bur. Fisheries, 
vol. 39, 1923-1924, pp. 439-471, figs. I-26. Protozoa, diatoms, other 
algae, organic detritus. Bibliography on the food of fresh-water 
mussels, and upon that of lamellibranchs in general. 

Cooke, A. H. 

1895. [Enemies of mollusca.] Cambridge Nat. Hist., vol. 3, pp. 56-62. 
Birds, rats, frogs, toads, beetles, mongooses, monkeys, walruses, 
whales, fishes, other mollusks, trematodes, nematodes, and mites. 

Dyeue, L. L. 

1903. Notes on the food habits of California sea-lions (Zalophus californi- 
anus Lesson.) Trans. Acad. Sci. Kansas, 1901-1902, pp. 179-182. 
Food found in numerous stomachs, chiefly squids. 

FEDERIGHI, HENry. 

1930. Control of the common oyster drill. Econ. Circ. 70, U. S. Bur. Fish- 
eries, 7 pp., 5 figs. Urosalpinx cinerea “destroys oysters to the 
value of several million dollars annually in the United States” 
(p. FE). 

FIELp, IrvinG A. 

1911. The food value of sea mussels. Bull. U. S. Bur. Fisheries, vol. 20, 
1909, pp. 85-128, pls. 18-25. Food (pp. 92-95), mostly diatoms 
and Protozoa. Enemies (pp. 95-97), are numerous, fishes, mollusks, 
sea-stars, crows, rats, parasitic crabs. 

Forrest, H. E. 

1927. Fishes, Caradoc and Severn Valley Field Club, record of bare facts 
for the year 1926, p. 19. Stomach of an eel (Anguilla vulgaris) 
from the Severn was full of small bivalves (Sphaerium corneum). 

HERRINGTON, Wo. C. 

1930. The Pismo clam. Fish. Bull. 18, California Div. Fish and Game, 
69 pp. 16 figs. Tivela stultorum—birds, rays, starfish, and marine 
snails are enemies (pp. 52-54). 

Moore, H. F. 

1908. Volumetric studies of the food and feeding of oysters. Bull. U. S. 
Bur. Fisheries, 28, pp. 1297-1308, pl. 125, 6 figs. Ninety-five per 
cent diatoms; remainder of equally minute plants and animals. 

Ritcutr, J. 

1927. A remarkable whale invasion. Scottish Nat. 1927, pp. 161-163. A 
school of false killers (Pseudorca crassidens) visited the Dornoch 
Firth in October, 1927, and some of them ran aground there. 
This whale is very rare, and had hardly been seen anywhere for 
80 years (a few appeared off Western Europe in 1861 and 1862, 
and it was also seen in Tasmania). Examination of these whales 
showed they feed mainly on large cuttlefish. 


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186 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 85 


STEVENSON, CHARLES H. 
1892. A bibliography of publications in the English language relative to 
oysters and the oyster industries. Extract from Rep. U. S. Comm. 
Fish and Fisheries for 1892, art. 3, pp. 305-359. Some of the papers 
referred to deal with the food and enemies of oysters. 
StiIves, CH. WARDELL. 
1902. Frogs, toads, and carp (Cyprinus carpio) as eradicators of fluke 
disease. Ann. Rep. U. S. Bur. Animal Industry 1go1, pp. 220- 
222, figs. 197-203. By feeding on snails the intermediate hosts. 


PISCES 


Apams, CHAs. C., AND HANKINSoN, T. L. 

1928. The ecology and economics of Oneida Lake fish. Roosevelt Wild 
Life Ann., vol. 1, nos. 3-4, pp. 242-548, I pl., figs. 179-244, Nov. 
Notes on food and enemies of most of the species; full bibliography. 

Auiin, A. E. . 

1929. Seining records and food of the intermediate stages of Lake Erie 
fishes. Suppl. 18th Ann. Rep. New York Conserv. Dep. 1928, pp. 
95-106. Cyprinidae and Catostomidae feed on algae and diatoms; 
the smaller Percidae on crustaceans and insect larvae, and the 
larger Percidae, Esocidae, and Catostomidae (fish eggs) on smaller 
fishes and fish eggs. 

ANNIN, J. 

1902. In Rhead, Louis, The speckled brook trout, pp. 129-140. Winged 
enemies include night heron, kingfisher, ducks, loons, grebes, fish 
hawk, bald eagle and barred owl. 

Baker, F. C. 

1918. The productivity of invertebrate fish food on the bottom of Oneida 
Lake, with special reference to mollusks. Techn. Publ. no. 9, New 
York State Coll. Forestry, 264 pp. Notes on food of five species 
of fishes (pp. 214-216). Bibliography. 

Bargour, T. 

1921. Spiders feeding on small cyprinodonts. Psyche, vol. 28, no. 4, pp. 

131-132, Aug. Dolomedes tenebrosus. 
BiceLow, N. K. 

1924. The food of young suckers (Catostomus commersonu) in Lake 
Nipigon. Univ. Toronto Studies, no. 24, Publ. Ontario Fish Res. 
Lab., no. 21, pp. 81-115. Rotifers, Cladocera, insects. 

Breper, C. M., Jr. 

1921. The food of Mustelus canis (Mitchill) in mid-summer. Copeia, no. 
101, pp. 85-86, Dec. 20. Notes on contents of 102 stomachs. (Fish 
in 10, crabs 44, Nereis sp. 1, univalves and vegetable matter 3.) 

1922. Observations on young bluefish. Copeia, no. 106, pp. 34-36, May 20. 
Contents of 15 stomachs listed; 86 per cent fishes. 

Breper, C. M., Jr., AND CrAwrorp, D. R. 

1922. The food of certain minnows. A study of the seasonal dietary cycle 
of six cyprinoids with especial reference to fish culture. Zoologica, 
vol. 2, no. 14, pp. 287-327, figs. 1ro1-128, Aug. Semotilus bullaris, 
87 per cent insects, including larvae and adults of several orders 
plus some worms, millipeds, crayfish; Leuciscus vandoisulus, 98 


7" 


NO. 7 PROTECTIVE ADAPTATIONS—McATEE 187 


per cent insects together with worms, spiders, crayfish and slug; 
Notropis procne, 36 per cent insects and in addition some worms 
and water mites; Notropis cornutus and Rhinichthys atronasus, 57 
per cent insects plus same additional items as in last; Exoglossum 
maxillingua, 35 per cent insects, plus worms and fish eggs. Most 
of the insects were terrestrial species. 

Carr, A. M. 

1908. Food of fishes. Rep. Sci. Invest. Northumberland Sea Fisheries 
Comm. 10907, pp. 68-71. Reports on stomach examinations of 10 
species. 

1909. The food and condition of fish obtained from the North-east coast. 
Rep. Sci. Invest. Northumberland Sea Fisheries Comm. 1908-1909, 
pp. 41-50. Stomach analyses of seven species of fishes (pp. 43-46). 

CHAMBERLAIN, F. M. 

1907. Some observations on salmon and trout in Alaska. U. S. Bur. Fisher- 
ies, Doc. 627, 112 pp., 5 pls. Enemies (pp. 107-109) include trout, 
sculpins, mergansers, golden-eyes, mallards. The trout feed on 
other fishes, insects and their larvae, snails, and bivalves. 

Cote, Leon J. 

1905. The German carp in the United States. Rep. U. S. Comm. Fisheries 
1904, pp. 523-641, pls. 1-3. Considerable on food and economic 
relations. Bibliography. 

DERYKE, WILLIS. 

1922. The food of the fishes of Winona Lake. Indiana Dep. Conserv., 47 
pp., 1 pl, 1 map. Notes on 17 species, 6 of which are treated 
in some detail; yellow perch: young, midge larvae, Entomostraca, 
amphipods; adults, chiefly fish; bluegill: young, chiefly midge 
larvae and Entomostraca; older, the same plus caddis larvae, 
insects, snails; large-mouth black bass: young, amphipods, Clado- 
cera, mayfly and midge larvae; larger, chiefly fish; log perch: 
amphipods, Cladocera, midge, caddis, and mayfly larvae, snails; 
skipjack: chiefly nonaquatic insects; sunfish: snails, midge larvae, 
insects; hogmolly: midge, and mayfly larvae, oligochaetes, Bibli- 
ography. 

Eaton, E. H. 

1928, The Finger Lakes fish problem. Suppl. 17th Ann. Rep. New York 
Conserv. Dep. 1927, pp. 40-46. Tabulation of food of some 30 
species, 3 of which are almost exclusive fish-eaters, 7 others 
largely so, 8 feed chiefly on larval, and 5 on flying insects. Six 
species eat many scuds, and 4 even as adults, subsist more or less 
on plankton Crustacea. Enemies of fish, besides their own kind, 
include lampreys, turtles, snakes, loons, grebes, and mergansers. 

Etmurirst, RICHARD. 

1926. Notes on fishes from the Firth of Clyde. Scottish Nat., pp. 151-158, 
and 179-186. Full notes on food of cod and briefer reference to 
that of some other species. Nine kinds of fishes (including itself) 
listed as predatory on herring. 

13 





188 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


Exrop, M. J. 

1929. The fishes of Flathead Lake. Montana Wild Life, vol. 2, no. 1, pp. 
6-9, June. Notes on food of: Catostomus spp.: Insects, Ento- 
mostraca; Ptychocheilus oregonensis: Mainly insects such as may- 
fly and caddisfly larvae, grasshoppers, some fish and shrimps; 
Mylocheilus caurinus: FEntomostraca and insects; Leuciscus gillu: 
Entomostraca and insects; Salmo clarkti: Beetles, mayflies, grass- 
hoppers; Salvelinus malma: Fishes including Coregonus and 
Ptychocheilus; Coregonus williamsoni: Larvae of Tipulidae, 
Simuliidae, Planorbis, Physa; Micropterus salmoides: Fish, insects. 

FIELD, Irvine A. 

1907. Unutilized fishes and their relation to the fishing industry. U. S. 
Bur. Fisheries Doc. no. 622, 50 pp. Notes on the food of eight 
species. 

Forses, S. A. 

1880. The food of fishes. Bull. Illinois State Lab. Nat. Hist., vol. 1, no. 3, 
pp. 18-65, Nov. Notes on stomach examinations for numerous 
species. 

1883. The food of the smaller fresh-water fishes. Bull. Illinois State Lab. 
Nat. Hist., vol. 1, no. 6, pp. 65-94, May. Examination of 319 
stomachs representing 25 species; food chiefly neuropteroid and 
chironomid larvae, and Entomostraca; other animal items, fishes, 
mollusks, Hymenoptera, Diptera, Coleoptera, Hemiptera, Thysan- 
ura, Arachnida, amphipods, isopods, worms, and protozoans. 

1890. Studies of the food of fresh-water fishes. Bull. Illinois State Lab. 
Nat. Hist. vol. 2, pp. 433-473. Many stomach examinations of 28 
species; tabulation of items and percentages. 

1890. On the food relations of fresh-water fishes. Bull. Illinois State Lab. 
Nat. Hist., vol. 2, pp. 475-538. Summary of the preceding papers, 
discussion of fishes as predators on other fishes, on mollusks, 
insects, crustaceans, worms, fresh-water sponges, and protozoans. 
Schedule of food items and the species taking them. 

Futon, T. WEMyYSS. 

1903. The distribution, growth, and food of the angler (Lophius pisca- 
torius.) 21st Ann. Rep. Fishery Board Scotland 1902, pp. 186-217. 
Analyses of 280 stomach contents; 269 containing fishes, 10 squids, 
and I a crab. 

Gupcer, E. W. 

1927. Hydras as enemies of young fishes. Nat. Hist., vol. 27, pp. 270-274, 
3 figs. 

1929. Wide-Gab, the angler fish. Nat. Hist., vol. 20, no. 2, pp. 155-159, 
illus., Mar.-Apr. Case of attempt to swallow a gull; review of 
literature, showing that birds up to the size of the loon are eaten; 
seven wild ducks from one stomach; the principal food, however, 
is fishes. 

HANKINSON, THomAsS L. 

1908. A biological survey of Walnut Lake, Michigan. Rep. Biol. Surv. 
Michigan Geol. Surv. 1907, pp. 158-288, pls. 13-75. Food of several 
species of fishes noted from examination of stomachs (pp. 200-216). 








NO. 7 PROTECTIVE ADAPTATIONS—McATEE 189 


HarNELL, J.. AND Nayupo, M. R. 

1924. A contribution to the life history of the Indian sardine, with notes 
on the plankton of the Malabar Coast. Madras Fisheries Bull. 17, 
pp. 129-197, 10 pls. Food extensively treated; consists of diatoms, 
peridineans, infusorians, Heliozoa, larval bivalves, and copepods. 

HILDEBRAND, S. F., AND Towers, I. L. 

1927. Food of trout in Fish Lake, Utah. Ecology, vol. 8, no. 4, pp. 389-397, 
Oct. Contents of 181 stomachs tabulated, the more important items 
being Daphnia, Gammarus, midges and vegetation; leeches, snails, 
dragonfly nymphs, and fishes and their eggs are other items of 
the food. 

JOHANSEN, FRITs. 

1912. The fishes of the Datimark Expedition. Danmark-Ekspeditionen 
Gronl. Nordostkyst 1906-1908, vol. 5, no. 12, pp. 633-675, 5 figs., 
pls. 44-46. Notes on the food of Gadus and Salmo. 

JouHNson, Ropert S., AND Stapleton, M. F. 

1917. Fish ponds on farms. App. 2, Rep. U. S. Comm. Fisheries 1915, 
29 pp. Cannibalistic and other predacious fishes, turtles, snakes, 
birds, and minks are the principal foes. 

Jupay, CHANCEY. 

1906. A study of Twin Lakes, Colorado, with especial consideration of 
the food of the trouts. Bull. U. S. Bur. Fisheries, vol. 26, pp. 147- 
178, pl. 3. In addition to notes on contents of 370 trout stomachs 
of six species this publication contains a good bibliography and a 
digest of papers relating to Entomostraca as food of fishes. 

KENDALL, WILLIAM C., 

1897. Notes on the food of four species of the cod family. Rep. U. S. 
Fish Comm. 1896, App. 3, pp. 177-186. A long list of food items. 
“Protective mimicry seems of little avail against these fishes.” 

KENDALL, WILLIAM C., AND DENcE, W. A. 

1927. A trout survey of the Allegany State Park in 1922. Roosevelt Wild 
Life Bull., vol. 4, no. 3, pp. 291-482, figs. 54-86, tables, July. Notes 
on 112 stomach contents (pp. 472-474, table 27): Midges, caddis- 
flies, beetles, ants and other Hymenoptera, Diptera, grasshoppers, 
plant lice, lacewing flies, stoneflies, mayflies, spiders, crustaceans, 
and fish. Bibliography. 

Knicut, A. P. 

1927. Losses in speckled trout fry after distribution. Science, n. s., vol. 65, 
pp. 525-526, Aug. Losses 71-08 per cent, mostly to natural enemies, 
birds, trout and other fishes. 

KrAAtz, WALTER C. 

1923. A study of the food of the minnow Campostoma anomalum. Ohio 

Journ. Sci., vol. 33, pp. 265-283. Diatoms, algae, etc. 
Lezour, Marie V. 

1924. The food of young herring. Journ. Marine Biol, Assoc. United 
Kingdom, n. s., vol. 13, pp. 325-330. Among animal items, Infusoria, 
larval mollusks, copepods. 


190 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


Lewis, RALPH C. 

1929. The food habits of the California sardine in relation to the seasonal 
distribution of microplankton. Bull. Scripps Inst. Oceanography, 
Techn. Ser. 2, pp. 155-180, 2 figs. Items of animal food are schizo- 
pods and copepods. 

Linton, EpwIn. 

1901. Fish parasites collected at Woods Hole in 1808. Bull. U. S. Fish 
Comm., vol. 19, 1899, pp. 267-304, pls. 33-43. Notes on fish food, 
Pp. 270-284. 

1901. Parasites of fishes of the Woods Hole region. Bull. U. S. Fish 
Comm., vol. 19, 1890, pp. 405-492, pls. 1-34. Summary of parasites, 
pp. 425-488, contains many references to food of fishes. 

1921. Food of young winter flounders. Rep. U. S. Comm. Fisheries, 
App. 4, 14 pp. Pseudopleuronectes americanus, food of young 
principally amphipods, other small Crustacea, and annelids; food 
of adults, annelids, Crustacea, ascidians, fish, mollusks. Almost 
as much on parasites (Sporozoa, trematodes, nematodes, and 
Acanthocephala) as on food. 

MacCoy, CLinTon V. 

1929. The mackerel in New England. Bull. Boston Soc. Nat. Hist., vol. 
53, PP. 3-7, Oct. Food, small fish, squids, pteropods, amphipods, 
copepods. Enemies, whales, porpoises, sharks, dogfish, bluefish, 
gannets, parasitic worms. 

MarsHALL, W. S., AND GILBER, N. C. 

1905. Notes on the food and parasites of some fresh-water fishes from 
the lakes at Madison, Wis. Rep. U. S. Comm. Fisheries 1904, 
App., pp. 513-522. Incidental notes on food. 

Moore, J. Percy. 

1922. Use of fishes for control of mosquitoes in northern fresh waters of 
the United States. Rep. U. S. Comm. Fisheries, App. 4, 60 pp., 
7 pls. Food of roach: Entomostraca, insects, mites, Protozoa; 
mudminnow: Insects, Crustacea, mollusks, Protozoa; killifish: 
Oligochaetes, mollusks, Entomostraca; top minnow: cannibalistic; 
blue-spotted sunfish: Midge larvae, Entomostraca, amphipods; 
long-eared sunfish: Midge larvae, Entomostraca, oligochaetes; 
common sunfish: Midge larvae, Entomostraca, snails, mites, tad- 
poles. All eat mosquito larvae. Bibliography. 

MurrKkowskI, RicHArpD A. 

1925. The food of trout in Yellowstone National Park. Roosevelt Wild 
Life Bull., vol. 2, no. 4, pp. 471-497, figs. 114-133, Feb. Stoneflies, 
90 per cent; mayflies, caddisflies, adults and young of all; and 
water-trapped land insects. : 

1929. The ecology of trout streams in Yellowstone National Park. Roose- 
velt Wild Life Ann., vol. 2, no. 2, pp. 155-240, figs. 53-116, Oct. 
Food of trout, pp. 222-230, conclusions as in his 1925 paper on the 
subject. Food of insects, pp. 230-233; see under Muttkowski and 
Smith. 

NEEDHAM, JAMES G. 

1903. Food of brook trout in Bone Pond. Bull. 68, New York State Mus., 

pp. 204-217. Contents of 25 stomachs tabulated. 








NO. 7 PROTECTIVE ADAPTATIONS—McATEE IQI 


NEEDHAM, JAMES G.; JupAY, CHANCEY; Moore, EMMELINE; SIBLE, CuAs. K.; 
AND TiTcoms, JOHN W. 

1922. A biological survey of Lake George, N. Y. N. Y. State Conserv. 
Comm., 78 pp., 27 figs. Much on the food of fishes; the staples of 
the diet of carnivorous fry are waterfleas, midges (all stages), 
other insects, scuds (amphipods), and crayfishes; cannibalism 
prevalent (p. 63); food of adults of eight species outlined on pp. 
65-68. Lake trout: Principal food, lake smelt, other items yellow 
perch, and caddisflies; black bass: Perch, crawfish, grasshoppers, 
scuds; pike: Other fishes; yellow perch: Staples, midge larvae, 
mayfly nymphs, scuds, snails, secondary, caddisworms and water- 
fleas; bullhead: Scuds, midge larvae, mayfly nymphs, snails; long- 
eared sunfish: Mayfly nymphs, midge larvae, ants, scuds, water- 
fleas, miscellaneous insects, and crayfishes ; common sunfish: Snails, 
mayfly nymphs, caddisworms, beetles, midge larvae, various insects ; 
rock bass: Crayfish, fishes, insects. 

NEEDHAM, P. R. 

1929. Quantitative studies of the fish food supply in selected areas. Suppl. 
18th Ann. Rep. New York Conserv. Dep. 1928, pp. 220-232. Ithaca, 
N. Y., Erie-Niagara watershed. Foods consumed by trout in com- 
parison with available foods; in the case of aquatic foods the re- 
lation of consumption to availability is very clear. This is a 
reworking of a similar paper in the 17th Ann. Rep. (1927) 1928, 
Pp. 192-206. 

New York CoNnsERVATION DEPARTMENT. 

1928. A biological survey of the Oswego River System. Suppl. 17th Ann. 
Rep. New York Conserv. Dep. 1927, 248 pp., 12 col. pls., text figs., 
maps. Much on fish food; in a tabulation of food items of adults 
of 31 species, midges, mayflies, and minnows seem to be most 
commonly used; and of young of eight species, copepods, Cladocera, 
and midges. 

Pace, Wo. F. 

1895. Feeding and rearing fishes, particularly trout, under domestication. 
Bull. U. S. Fish Comm., 1894, pp. 280-314. Some notes on natural 
food, and an indexed bibliography. 

Patterson, A. H. 

1926-1927. Food of the Sturgeon. Trans. Norfolk and Norwich Nat. Soc., 
vol. 12, pp. 380-381. Stomach of one contained about 729 small 
fish (lesser sandlaunces). 

Pearse, A. S. 

1915. On the food of the small shore fishes in the waters near Madison, 
Wisconsin. Bull. Wisconsin Nat. Hist. Soc., vol. 13, no. I, pp. 7-22, 
1 fig., Mar. Sixteen species, of which nine lived largely on insects 
and their larvae, two on ostracods, two on copepods, and one on 
Cladocera. 

1918. The food of the shore fishes of certain Wisconsin lakes. Bull. U. S. 
Bur. Fisheries, vol. 35 (1915-16), pp. 249-292. Report on more 
than 1,600 specimens of 32 species, with bibliography. 

1919. Habits of the black crappie in inland lakes of Wisconsin. Rep. U. S. 
Comm. Fisheries 1918, app. 3, pp. 5-16. Tabulation of contents of 
276 stomachs. 


Ig2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 





1921. Distribution and food of the fishes of Green Lake, Wis., in summer. 
Bull. U. S. Bur. Fisheries, vol. 37, 1919-1920, pp. 255-272, I map. 
Notes on 16 species; the food of all combined comprised insect 
larvae 21.7 per cent, amphipods 16.5 per cent, fish 9.6 per cent, | 
crayfishes 7.8 per cent, cladocerans 7.6 per cent, insect pupae 6.7 | 
per cent, snails 4.4 per cent, bivalves 4.1 per cent, and the following |i 
items in smaller proportions, adult insects, ostracods, oligochaetes, | 
leeches, mites, Mysts, and copepods. Sixty-seven per cent is arthro- | 
pods, composed of 31.7 insects and 35.6 crustaceans. Comparison | 
is made with the fishes of Lake Mendota. Bibliography. | 

1924. Amount of food eaten by four species of fresh-water fishes. Ecology, | 
vol. 5, no. 3, pp. 254-258, July. Order of choice, minnows, earth- | 
worms, amphipods, dragonfly nymphs, crayfishes, grasshoppers, 
snails, and caddis larvae. 

PearsE, A. S., AND ACHTENBERG, HENRIETTA. 

1917-1918. Habits of yellow perch in Wisconsin Lakes. Bull. U. S. Bur. | 
Fisheries, vol. 36, 1917-1918, pp. 297-366, pl. 83, figs. 1-35. Report | 
on 1,147 stomach examinations of which the food as a whole was | 
made up of 38.3 per cent insect larvae, 21.4 Entomostraca, 9.5 insect | 
pupae and adults, 5.5 macroscopic crustaceans, 4.5 fishes, 2.4 mol- | 
lusks, 1.4 oligochaetes, leeches and arachnids. Enemies of the perch 

include pickerel, black bass, a number of birds, and a variety of | 
parasites. Bibliography. | 
PEARSON, JOHN C. | 

1928. Natural history and conservation of the redfish and other commercial 
sciaenids of the Texas coast. Bull. U. S. Bur. Fisheries, vol. 44, 
pp. 129-214, 44 figs. Sciaenops ocellatus: Shrimps, crabs, mollusks, 
fish; Pogonias cromis: Clams, mussels, oysters, crabs, shrimps, 
fish, annelids ; Cynoscion nebulosus: Shrimps, crabs, fish; Micropo- 
gon undulatus: Shrimps, crabs, annelids, fish. 

Peck, JAMEs I. 

1894. On the food of the menhaden. Bull. U. S. Fish Comm., vol. 13, 1893, 
pp. 113-126, pls. 1-8. Food filtered from water by gill-raker 
mechanism, consists chiefly of unicellular organisms, both animal 
and vegetable. They also take ostracods, copepods, amphipods and 
other small Crustacea, and young Nerets. Composition of food the 
same as material filtered from water by mechanical contrivances: 
Diatoms, rotifers, dinoflagellates, etc. The supply of such food 
illimitable. 

1806. The sources of marine food. Bull. U. S. Fish Comm., 1895, pp. 351- 
308, pls. 64-71. Plankton, largely diatoms, the basis; notes on 
the food of the squeteague, the bluefish, sea bass, scup, and tautog. 

PETERSEN, C. G. J. 

1894. On the biology of our flat-fishes. Rep. Danish Biol. Sta., vol. 4, 
1893, pp. v-+146,2 pls., I map, 18 tables. Notes on food of young 
and adults. 

Scott, ANDREW. 

1899. Observations on the habits and food of young fishes. Proc. & Trans. 

Liverpool Biol. Soc., vol. 13, 1898-99, pp. 90-93. 





INOS 7 PROTECTIVE ADAPTATIONS—McATEE 193 


Scott, THOMAS. 

1902. Observations on the food of fishes. 20th Ann. Rep. Fishery Board 
Scotland 1901, pt. 3, pp. 486-538. Notes on 56 species. 

1903. Some further observations on the food of fishes, with a note on the 
food observed in the stomach of a common porpoise. 21st Ann. 
Rep. Fishery Board Scotland 1902, pp. 218-227, 2 figs. 

S1ptey, C. K. 

1929. The food of certain fishes of the Lake Erie Drainage Basin. Suppl. 
18th Ann. Rep. New York Conserv. Dep. 1928, pp. 180-188. 
Thirty-four species feed mainly on immature aquatic insects, es- 
pecially midge larvae, and Crustacea; eight species are pronounced 
spawn-eaters; small fish are important food of the larger species; 
food of young chiefly copepods and Cladocera. 

SMALLWoop, W. M., AND STRUTHERS, P. H. 

1927. Carp control studies in Oneida Lake. Suppl. 17th Ann. Rep., New 
York Conserv. Dep., pp. 67-83. Much on food; animal matter taken 
by adults includes fish, ostracods, phyllopods, copepods, crayfish, 
midge and caddis larvae and other insects; by young, ostracods, 
copepods, Cladocera, insect larvae, snails, worms, mites, eggs of 
snails, insects and copepods, rotifers, and bivalves. 

SmitH, HucH M. 

1896. A review of the history and results of the attempts to acclimatize 
fish and other water animals in the Pacific States. Bull. U. S. 
Fish Comm., vol. 15, 1805, pp. 379-472, pls. 73-83. Notes on food 
of a few species. Catfish, fish eggs and fry; carp, spawn; shad, 
shrimps; striped bass, carp, catfish, crabs. 

Smitu, W. Ramsay. 
1889. On the food of fishes. 7th Ann. Rep. Fishery Board Scotland 1888, 


pp. 222-258. 

1890. On the food of fishes. 8th Ann. Rep. Fishery Board Scotland 1880, 
pp. 230-256. 

1891. On the food of fishes. 9th Ann. Rep., Fishery Board Scotland 1890, 
pp. 222-242. 


1892. On the food of fishes. toth Ann. Rep. Fishery Board Scotland 1801, 
pp. 211-231. This and similar papers in three previous reports are 
based on investigations of Thomas Scott. 

STEWART, N. H. 

1926. Development, growth, and food habits of the white sucker, Catosto- 
mus commersoni Lesueur. Bull. U. S. Bur. Fisheries, vol. 42, pp. 
147-184, 55 figs. Among animal food midge larvae are most impor- 
tant at all ages; some rotifers, Entomostraca, and Protozoa are taken 
at all stages also, but dragonfly, caddisfly, mayfly larvae, and 
Mollusca are taken only by adults. Bibliography. 

SrruTHERS, P. H. 

1929. Carp control studies in the Erie Canal. Suppl. 18th Ann. Rep. New 
York Conserv. Dep. 1928, pp. 208-219. Animal food (p. 214) in- 
cludes insect larvae, snails, midge larvae, bivalves, ostracods, 
Malacostraca, copepods, Cladocera, and decapods, 


194 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


SUMNER, Francis B.; Ospurn, RAyMonp C.; AND Core, Leon J. 

1911. A biological survey of the waters of Woods Hole and vicinity. 
Bull. U. S. Bur. Fisheries, vol. 31, pt. 2. The catalogue of the 
marine fauna, fishes, pp. 734-744, contains notes on the food 
mainly quoted from Verrill, Goode, Linton, and Field. 

TAVERNER, P. A. 

1915. The double-crested cormorant (Phalacrocorax auritus) and its re- 
lation to the salmon industries on the Gulf of St. Lawrence. Can. 
Geol. Surv., Bull. 13, 24 pp., 1 pl. Food sculpins, herring, capelin, 
eel, etc., no salmon; the salmon feeds on other fishes, and crusta- 
ceans, and is cannibalistic. 

TIFFANY, Lewis H. 

1921. Algal food of the young gizzard shad. Ohio. Journ. Sci., vol. 21, 
no. 4, pp. 113-122, Feb. Mentions several game fishes that prey 
on this wholly vegetarian species. 

TURNER, CLARENCE L. 

1920. Distribution, food and fish associates of young perch in the Bass 
Island region of Lake Erie. Ohio Journ. Sci., vol. 20, no. 5, pp. 
137-152, Mar. Details of analyses of 138 stomach contents. 

1921. Food of the common Ohio darters. Ohio Journ. Sci., vol. 22, pp. 
41-62. Usually the food changes with age from Entomostraca to 
midge larvae and similar organisms, and then with maturity, to 
a varied diet in which ephemerid and other large insect larvae 
predominate. | 

1922. Notes on the food habits of young Cottus ictalops (miller’s thumb). 
Ohio Journ. Sci., vol. 22, pp. 95-96. Midge and other insect larvae. 

VerrILL, A. E. 

1873. Report upon the invertebrate animals of Vineyard Sound and the 
adjacent waters, with an account of the physical characters of the 
region. Report on Sea Fisheries of New England, pt. 1, pp. 295- 
778. Lists of species found in the stomach of fishes (pp. 514-521). 

Warren, B. H. 

1897. Fish-eating birds and mammals. Ann. Rep. Pennsylvania Dep. Agr., 
1896, pp. 297-303, I pl. Seventeen or more kinds of birds, wild 
cats, raccoons, muskrats, mink, and the otter. 

WELsH, WM., AND Breper, C. M., Jr. 

1923-1924. Contributions to life histories of Sciaenidae of the eastern United 
States coast. Bull. U. S. Bur. Fisheries, vol. 39, pp. 141-201, 60 
figs. Notes on food of eight species; it is chiefly crustaceans, next 
in order coming worms and fishes. Bibliography. Cynoscion re- 
galis: Shrimps, schizopods, isopods, amphipods, worms when small, 
fish when mature, but including shrimps and squids; Bairdiella 
chrysura: Schizopods, isopods, amphipods, worms, fish; Stellifer 
lanceolatus: Schizopods, copepods, decapods, ostracods, amphipods, 
worms; Leiostomus xanthurus: Ostracods, copepods, amphipods, 
worms, mollusks; Micropogon undulatus: Shrimps, echinoderms, 
worms, mollusks, copepods, ostracods, amphipods; Menticirrhus 
americanus: Crabs, shrimps, worms, fish; Menticirrhus saxatilis: 
Shrimps, amphipods, schizopods, worms, fish; Pogonias cromis: 
Mollusks including oysters; Eques pulcher: Small crustaceans. 











NO. 7 PROTECTIVE ADAPTATIONS—McATEE 195 


WICKLIFF, Epwarp L. 

1920. Food of young small-mouth black bass in Lake Erie. Proc. Amer. 
Fisheries Soc., pp. 364-371. Report on 313 specimens, the most 
important items being copepods found in 61 per cent of the stomachs 
and Cladocera in 39 per cent. Other commonly taken foods were 
midge larvae and pupae, adult insects, fish, and mayfly nymphs. 


AMPHIBIA 
DRAKE, Caru J. 

1914. The food of Rana pipiens Schreber. Ohio Naturalist, vol. 14, no. 5, 
pp. 257-269, Mar. Detailed account of the contents of 209 stomachs 
collected at Cedar Point, Ohio. 

Frost, S. W. 

1924. Frogs as insect collectors. Journ. New York Ent. Soc., vol. 32, 
no. 4, pp. 174-185, pl. 14, Dec. Eat worms, snails, crayfishes, 
spiders, mites, insects and frogs; insects most important. Larvae: 
Lepidoptera 9; Coleoptera 24; Diptera 13; Neuroptera 1. Adults: 
Orthoptera 1; Hemiptera 25; Neuroptera 3; Mecaptera 1; Diptera 
33; Coleoptera 242; spiders 37; pseudoscorpions I. 

GARMAN, H. 

1901. The food of the toad. Bull. 91, Kentucky Agr. Exp. Sta., pp. 60-68, 

fig. 16. Report on 20 stomach contents. 
HAmILton, W. J., JR. 

1930. Notes on the food of the American toad. Copeia, 1930, no. 2, June 30, 
p. 45. Bufo americanus. Report on food of 400 young toads: 
Diptera 22 per cent, mostly larvae; mites 15.5 per cent; ants 
12.8; beetles and their larvae 11.8, the most abundant group being 
Staphylinidae; thrips 10.1; Collembola 6.2; Lepidoptera, Hymen- 
optera, aphids, sowbugs, spiders, worms, and snails, the remainder. 

KIRKLAND, A. H. 

1904. Usefulness of the American toad. Farmers’ Bull. no. 196, U. S. Dep. 

Agr., 16 pp. Contents of 149 stomachs discussed. 
Kiucu, A. Brooker. 

1922. The economic value of the leopard frog. Copeia, no. 103, pp. 14-15, 
Feb. 15. Contents of 25 stomachs; chiefly Melanoplus femur-rubrum 
and Leptinotarsa 1o-lineata. 

Muwnz, Pur A. 

1920. A study of the food habits of the Ithacan species of Anura during 
transformation. Pomona Coll. Journ. Ent. Zool., vol. 12, no. 2, 
pp. 33-56, June. Report on 586 stomachs of eight species; sum- 
maries of results of previous investigators. 

SMALLWoop, W. M. 

1928. Notes on the food of some Onondaga Urodela. Copeia, no. 160, 
pp. 89-98, Oct. 25. Ambystoma maculatum: Centipeds, earth- 
worms, snails, sowbugs, crickets, grasshoppers, beetles; Plethodon 
cinereus: Centipeds, earthworms, snails, sowbugs, ants, beetles, 
mites, spiders, phalangids, caterpillars, grasshoppers, flies spring- 
tails; Eurycea bislineata: Earthworms, caterpillars, and beetle, 
fly, and caddisfly larvae; Triturus viridescens: Snails, water- 
boatmen, fish, earthworms, beetle larvae; bivalves, daphnia, cater- 


196 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 85 


pillars, amphibian eggs (including its own), water bugs, mosquito 
and other fly larvae, slugs, snails, leeches, spiders, springtails, 
beetles, mites. 
SurFAceE, H. A., [Ep.] 
1913. First report on the economic features of the amphibians of Pennsyl- 
vania. Zool. Bull. Div. Zool. Pennsylvania Dep. Agr., vol. 3, nos. 
3-4, pp. 67-152, figs. I-25, pls. 1-11, May-July. General discussion 
of the subject, including report on examination of stomachs of 14 
species of salamanders, two of toads, and nine of frogs. 
WricutT, A. H. 
1920. Frogs: Their natural history and utilization. App. 6, Rep. U. S. | 
Comm. Fisheries 1919, 44 pp. Notes on the food of various species, 
pp. 38-42. Enemies, pp. 42-44; invertebrates, fishes, amphibians, | 





reptiles, birds, and mammals discussed. 
WricHt, A. H., AnD Haaser, Juria M. 
1922. The carnivorous habits of the purple salamander. Copeia, no. 105, 
pp. 31-32, April 15. Feed on aquatic insects; in captivity take 
frogs and salamanders. 


Burt, Cuas. E. 

1928. Insect food of Kansas lizards with notes on feeding habits. Journ. 
Kansas Ent. Soc., vol. I, no. 3, pp. 50-68, July. Notes on stomach 
contents of seven species with compiled information on others. Of 
the total food in all lizard stomachs examined 51.92 per cent was 
Orthoptera, 11.65 Lepidoptera, 9.35 Arachnida, 8.900 Hymenoptera, 
and 6.00 Coleoptera; Diptera, Hemiptera, Trichoptera, and Mol- 
lusca in smaller amounts. 

KELLocG, REMINGTON. 

1929. The habits and economic importance of alligators. Techn. Bull. 147, 
U. S. Dep. Agr., 36 pp., 2 pls. Dec. Food (pp. 21-32), nearly 
half is crabs, crawfishes, and shrimps; spiders, insects of various 
orders, toads, smaller alligators, lizards, turtles, snakes, birds and 
mammals also eaten. 

LyYDEKKER, R.; CUNNINGHAM, J. T.; BoULENGER, G. A.; AND THomson, J. A. 

1912. Food and growth [of reptiles], reptiles, Amphibia, fishes, and lower 
Chordata, pp. 47-61, London. ‘“ The food of reptiles is very vari- 
ous,” a dictum which shows distribution of predation is as charac- 
teristic of this phylum as of others. Details in many cases. 

PACK Elead): 

1921. Food habits of Sceloporus graciosus graciosus (Baird and Girard). 
Proc. Biol. Soc. Washington, vol. 34, pp. 63-66, Mar. Report on 
the contents of 71 stomachs. 

1922. Food habits of Crotaphytus wislizenit Baird and Girard. Proc. Biol. 
Soc. Washington, vol. 35, pp. 1-3, Mar. 20. Report on the con- 
tents of 18 stomachs. 

1923. Food habits of Callisaurus ventralis ventralis (Hallowell). Proc. 
Biol. Soc. Washington, vol. 36, pp. 79-81, Mar. Twenty stomachs; 
disclosing caterpillars, coccinellids, meloids, erotylids, chrysomelids, 
weevils, grasshoppers, mantids, Hemiptera, ant-lions, Diptera, and 
spiders. 


( 
REPTILIA | 
{ 


| 





INO: 7. PROTECTIVE ADAPTATIONS—McATEE 197 


1923. Food habits of Crotaphytus collaris baileyi (Stejneger). Proc. Biol. 
Soc. Washington vol. 36, pp. 83-84, Mar. Report on 16 stomach 
examinations; Orthoptera the principal animal food, caterpillars, 
wasps, bugs, leaf-hoppers, and ant-lions also being taken. 

1923. The food habits of Cnemidophorus tessellatus tessellatus (Say). Proc. 
Biol. Soc. Washington, vol. 36, pp. 85-80, Mar. Sixty-three 
stomachs containing caterpillars, 37.7 per cent of the food, grass- 
hoppers 14.4; beetles 14.2; other insects 14.27; and arachnids 8.2. 

Surrace, H. A. [Ep.] 

1906. The serpents of Pennsylvania. Monthly Bull. Div. Zool. Pennsyl- 
vania Dep. Agr., vol. 4, nos. 4-5, pp. 115-202, pls. 14-52, figs. 5-23, 
Aug.-Sept. Includes data on contents of stomachs of snakes ot 
14 species. 

1907. The lizards of Pennsylvania. Bull. Div. Zool. Pennsylvania Dep. 
Agr., vol. 5, no. 8, pp. 235-258, pls. 30-33, figs. 26-28, Dec. 1. 
Notes on food of five species, in the case of two of them based on 
examinations of stomachs. 

1908. First report on the economic features of turtles of Pennsylvania. 
Bull. Div. Zool. Pennsylvania Dep. Agr., vol. 6, nos. 4-5, pp. 107- 
195, pls. 4-12, 16 figs., Aug.-Sept. Includes report on stomach 
contents of representatives of nine species. 

WIntTon, W. M. . 

1915. A preliminary note on the food habits and distribution of the Texas 
horned lizards. Science, n. s., vol. 41, pp. 707-8, May 28. Brief 
summary of the results of examination of 485 stomachs; agricul- 
tural ants found in 80 per cent and stink bugs in 60 per cent of 
the stomachs. 

Wricut, A. H.; FUNKHousErR, W. D.; AND BrsHop, S. C. 

1915. A biological reconnaissance of the Okefinokee Swamp in Georgia. 
Turtles, lizards, and alligators, by Wright and Funkhouser, pp. 
108-139; snakes by Wright and Bishop, pp. 139-192. Proc. Acad. 
Nat. Sci. Philadelphia, pp. 107-192, pls. I-III, figs. 1-14, Mar. 
(Apr.). Notes on food of many of the species. 


AVES 


The entries under food of birds are chiefly general papers in which bibliogra- 
phies introductory to the very extensive literature of the subject can be 
found. 


CLELAND, J. B. 

1922. The parasites of Australian birds. Trans. and Proc. Roy. Soc. South 
Australia, vol. 46, pp. 85-118. Cestodes in 86 species, adult nema- 
todes in 28, Microfilaria in 34, Acanthocephala in 25, trematodes 
in 38, fleas on 3, Hippoboscidae on 4, Mallophaga on 107, ticks 
on 4, mites on 38, Haemosporidia in 47, and haemoflagellates in 12. 

Cram, ELoIse B. 

1927. Bird parasites of the nematode suborders Strongylata, Ascaridata, 
and Spirurata. U. S. Nat. Mus. Bull. 140, 465 pp., 444 figs. About 
500 species. 





198 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


ForsusH, E. H. 

1904. Special report on the decrease of certain birds, and its causes with 
suggestions for bird protection. 52nd Ann. Rep. Massachusetts 
State Board Agr., pp. 420-543, 2 pls. Chief causes, gunners, trap- 
pers, egg collectors, destruction of environment, natural enemies, 
and the elements. 

1907. Useful birds and their protection. Massachusetts State Board Agr., 
437 pp., 56 pls., 171 figs. Capacity of birds for destroying pests, 
birds as enemies of insects, and mammals, hairy caterpillars, plant 
lice, also on natural checks upon bird life. 

1916. The natural enemies of birds. Econ. Biol. Bull. 3, Massachusetts 
State Board Agr., 58 pp., 7 pls., figs. A thorough review of the 
subject, treating enemies among domesticated animals and among 
wild mammals, birds, reptiles, amphibians, fishes, and insects. 

Gross, A. O. 

1928. The heath hen, pp. 525-526. Marsh hawk, Cooper’s hawk, sharp- 
shinned hawk, and goshawk, the snowy owl, and crows enemies 
of this species. Domestic cat the worst. 

HENDERSON, JUNIUS. 

1927. The practical value of birds, 342 pp. An exhaustive review of litera- 
ture on the economics of American birds, with a long bibliography. 
Chapters on birds as enemies of injurious insects, mammals, and 
plants; birds as scavengers, and on the destruction of birds. 

Hersey, L. J. 

1907. A naturalist’s notes on birds and snakes. Outdoor Life, pp. 481-483, 

Nov. Snakes eating birds and their eggs. 
Lewis, ELIsHA J. 

1857. [Enemies of the partridge]. The American sportsman, 3rd ed., 
Enemies of the partridge (pp. 102-4) : fox raccoon, weasel, polecat, 
serpent, hawk, crow (p. 102); sparrow hawk, pigeon hawk, 
goshawk (p. 103). 

1857. [Enemies of the wild turkey]. The American sportsman, 3rd ed. 
Wolf, fox, lynx, cougar, opossum, and wild cat. Also the larger 
hawks and owls (p. 141). 

1857. [Enemies of the ruffed grouse]. The American sportsman, 3rd ed. 
Polecats, weasels, raccoons, opossums, foxes, crows, larger hawks 
(p. 150). 

Linton, E. F 

1927. Notes on cestode parasites of birds. Proc. U. S. Nat. Mus., vol. 70, 
art. 7, 73 pp. 15 pls. Thirty-four species. 

1928. Notes on trematode parasites of birds. Proc. U. S. Nat. Mus., vol. 
73, 30 pp., 11 pls. Twenty-two species. 

LoncstaFF, T. G. 

1927. Bird’s-nesting mice and insects. British birds, vol. 20, pp. 198-1090. 
Notes certain insects (e.g. ants) attacking nestlings, and mice 
destroying eggs. 

McAter, W. L. 

1913. Index to papers relating to the food of birds by members of the 
Biological Survey in publications of the United States Department 
of Agriculture, 1885-1911. U. S. Biol. Surv. Bull. 43, 1913, 69 pp. 
Bibliography with subject index. 





INOS) 7, PROTECTIVE ADAPTATIONS—McATEE 199 


PuiatH, O. E. 

1919. Parasitism of nestling birds by fly larvae. The Condor, vol. 21, pp. 
30-38. Protocalliphora azurea in 39 out of 63 nests; parasites and 
scavengers on this dipteron. 

Ransom, B. H. 

1909. The taenioid cestodes of North American birds. U. S. Nat. Mus. 

Bull. 60, 141 pp., 42 figs. About 140 species; bibliography. 
RusseEL, J. F. 

1926. Predatory bass. Outdoor Life, vol. 57, no. 2, pp. 146-147, Feb. Black 

bass with swallow in its stomach. San Diego Co., Calif. 
Tucker, B. W. 

1926. Bird’s-nesting bank voles. British birds, vol. 20, pp. 158-160. Evi- 
dence was given that bank voles destroy birds’ eggs. This note 
was followed by a number of other communications by various 
authors, in the same journal (vol. 20, pp. 180-181, 198-199, 207, 
230, 255), which showed that various species of mice commonly 
attack birds’ eggs. 

Weep, C. M., AND DEARBORN, NED. 

1903. Birds in their relations to man, 380 pp., illus. Extensive chronological 
bibliography; chapters on birds as regulators of outbreaks of in- 
jurious animals, relations of birds to predacious and _ parasitic 
insects. 

Wairp, OF A. 
1927. Wasps destroying young birds. British birds, vol. 20, pp. 254-255. 


MAMMALIA 
Bascock, H. L. 

1914. Some observations on the food habits of the short-tailed shrew 
(Blarina brevicauda). Science, n. s., vol. 40, pp. 526-530, Oct. 0. 
Review of literature, chiefly about observations on captive animals. 

BaILey, VERNON, AND SPERRY, CHAS. C. 

1929. Life history and habits of grasshopper mice, genus Onychomys. 
Techn. Bull. 145, U. S. Dep. Agr., I9 pp., 4 pls., Nov. Animal 
food (pp. 10-19), nearly 90 per cent of the whole, largely grass- 
hoppers, crickets, caterpillars, moths, and beetles; insects of other 
orders, spiders, and mice also taken. 

Brooks, Frep E. 

1908. Notes on the habits of mice, moles, and shrews. Bull. 113, West 
Virginia Agr. Exp. Sta., pp. 89-133, 10 pls., 2 figs., Jan. Con- 
siderable on food; review of previous writings. 

Bruce, JAY. 

1925. The problem of mountain lion control in California. California Fish 
and Game, vol. 2, no. I, pp. I-17, figs. 1-5, Jan. Each mountain- 
lion costs the State $1,000 a year in deer meat, or about $15,000 
to maintain the animal during its natural existence. 

CrIDDLE, NORMAN. 

1917. Varying hares of the prairie provinces. Agr. Gaz. Canada, vol. 4, 
no. 4, p. 262, Apr. Goshawk, golden and bald eagles, and great 
horned owls serious enemies. 


200 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


Drxon, JosEPH. 

1925. Food predilections of predatory and fur-bearing mammals. Journ. 
Mamm., vol. 6, no. I, pp. 34-46, pl. 4, Feb. Wild cat: Mammals, 
birds, fish; coyote: Game, stock, rodents, insects, mammals, birds; 
mountain-lion: Deer, stock, small wild mammals; skunks: Insects, 
rodents, birds, mammals. 

DYCHE We. le: 

1903. Food habits of the common garden mole (Scalops aquaticus ma- 
chrinus Rafinesque). Trans. Acad. Sci. Kansas 1901-1902, pp. 183- 
186. Report on the stomach contents of 50 specimens. 

ForsusH, E. H. 

1916. The domestic cat. Bird killer, mouser, and destroyer of wild life. 
Means of utilizing and controlling it. Econ. Biol. Bull. 2, Massa- 
chusetts State Board Agr., 112 pp., 20 pls., figs. The most compre- 
hensive review of the subject; cats kill millions of birds annually; 
destructive also to moles, shrews, toads, field mice, wood mice, 
insects. 

GARMAN, H. 

1895. The food of the common mole. 7th Ann. Rep. Kentucky Agr. Exp. 

Sta. 1894, pp. xli-xlv. Notes on contents of 14 stomachs. 
Hamitton, W. J., Jr. 

1930. The food of the Soricidae. Journ. Mamm., vol. 11, no. I, pp. 26-39, 
Feb. Over 300 stomachs representing four species; food is insects, 
annelids, Crustacea, snails, mice, salamanders, arachnids, centipeds, 
and millipeds. Bibliography. 

JOHANSEN, FRITs. 

1910. Observations on seals (Pinnipedia) and whales (Cetaceae) made on 
the “Danmark Expedition” 1906-1908. Danmark Eksp. Gronl. 
Nordéstkyst, 1906-1908, vol. 5, no. 2, pp. 203-224, 9 figs. Includes 
some notes on food. 

Jounson, Cuas. E. 

1925. The muskrat in New York; its natural history and economics. 
Roosevelt Wild Life Bull., vol. 3, no. 2, pp. 205-320, pl. 5, figs. 48- 
87, Mar. Animal food includes bivalves, snails, crayfish, insects, 
fishes, turtles, and birds; enemies include minks, foxes, weasels, 
otters, hawks, and owls. 

Lantz, D. E. 

1905. Kansas mammals in their relations to agriculture. Bull. 129, Kansas 
Agr. Exp. Sta., Dec., 1904, pp. 331-404, I pl., 1 fig. Notes on the 
food habits of most of the groups. 

1906. Meadow mice in relation to agriculture and horticulture. U. S. Dep. 
Agr. Yearbook 1905, pp. 363-376, pls. 38-41, fig. 89. Natural 
enemies (pp. 370-373) include wolves, lynxes, foxes, badgers, 
raccoons, opossums, skunks, minks, weasels, shrews, hawks, owls, 
crows, shrikes, cranes, herons, bitterns, snakes, and domestic cats 
and dogs. 

1918. The house rat the most destructive animal in the world. U. S. Dep. 
Agr. Yearbook 10917, pp. 235-251, pls. 41-44. Natural enemies (pp. 
248-249) include domestic dog, cat, and ferret, as well as snakes, 
storks, herons, owls, hawks, skunks, weasels. 








NO. 7 PROTECTIVE ADAPTATIONS—McATEE 201 


1923. Economic value of North American skunks. Farmers’ Bull. 587, 
U. S. Dep. Agr., 24 pp., 10 figs. Food (pp. 9-14), poultry, game, 
mice, and armyworms, tobacco worms, whitegrubs, hop grubs, 
grasshoppers, potato beetles and other insects. 

L.[ucas], F. A. 

1905. The Newfoundland whale fisheries. Science, n. s., vol. 21, p. 713, 
May 5. Large whales feed almost exclusively on Euphausia; fin- 
backs upon caplin. 

Piper, S. E. 

1909. Mouse plagues, their control and prevention. U. S. Dep. Agr. Year- 
book 1908, pp. 301-310, pls. 21-25. During a plague near Humboldt 
Lake, Nevada, 2,000 predatory birds and 1,000 mammals put in 
their appearance and together consumed about 1,350,000 mice per 
month. 

1928. The mouse infestation of Buena Vista Lake Basin, Kern County, 
California, September, 1926, to February, 1927. Monthly Bull. 
California Dep. Agr., vol. 17, no. 10, pp. 538-560, figs. 91-102, Oct. 
Ring-billed gulls, short-eared owls, barn owls, various hawks, 
ravens, great blue herons, road-runners, shrikes, coyotes, skunks, 
and house cats noted as predators (pp. 550-552). 

RAtnzsow, W. J. 

1913. Food, medicines, and charms of savage man. Abstract in Rep. 
Trustees Australian Mus. 1913, p. 9. Humans feeding on spiders, 
beetle larvae, caterpillars, grasshoppers, ants, bees, wasps, termites, 
and scorpions. 

SCHEFFER, THEO. H. 

1910. The common mole. Bull. 168, Kansas Agr. Exp. Sta., 36 pp., figs. 
Natural enemies (pp. 20-21) include hawks, owls, coyotes, domestic 
dogs. On the whole has few foes. 

1927. American moles as agricultural pests and as fur producers. Farmers’ 
Bull. 1247, U. S. Dep. Agr., 20 pp., 18 figs. Animal food (pp. 7-8), 
earthworms, beetles and their larvae, spiders, centipeds, ants, 
caterpillars. 

West, JAMES A. 

1910. A study of the food of moles in Illinois. Bull. Illinois State Lab. 
Nat. Hist., vol. 9, pp. 14-22, Oct. Details of contents of 56 
stomachs; references to previous literature. 





SMITHSONIAN MISCELLANEOUS COLLECTIONS 
VOLUME 85, NUMBER 8 


MODERN SQUARE GROUNDS OF THE 
CREEK INDIANS 


(WITH Five PLATES) 





BY 
JOHN R. SWANTON 


Ethnologist, Bureau of American Ethnology 





(PUBLICATION 3126) 


| CITY OF WASHINGTON 
| PUBLISHED BY THE SMITHSONIAN INSTITUTION 
NOVEMBER 11, 1931 





The Lord Battimore Press 


BALTIMORE, MD., U. 8. As 


MODERN SQUARE GROUNDS OF THE CREEK INDIANS 
By JOHN R. SWANTON, 


ETHNOLOGIST, BUREAU OF AMERICAN ETHNOLOGY 
(WirH Five PLateEs) 


The writer has already published descriptions of many of the 
square grounds of the Creek Indians, the sacred areas where their 
busks and other annual ceremonies took place." In collecting this 
material, however, my endeavor was to learn the most ancient 
arrangement of the several grounds and the arrangement of those 
grounds no longer in use. During the summer of 1929 I visited the 
Creek country again to secure information regarding the organization 
of the extant grounds. This work duplicates and supplements the 
earlier to a considerable extent, but the main purpose was somewhat 
different. 

Besides the three Yuchi grounds, now in the vicinity of Kellyville, 
Bixby, and Depew, respectively, with which I did not concern myself, 
there are, or were in 1929, 17 square grounds, as follows: Abihka 
and Otciapofa near Henryetta ; Nuyaka north of Okemah ; Latogalga ° 
or Fish Pond and Asilanabi west of Okemah; two Tulsa grounds 
near Holdenville; Tukabahchee at Yeager; Laptako near Wetumka ; 
Alabama east of Alabama Station on the Frisco Railroad; Eufaula 
west of Eufaula; Kasihta east of Okmulgee; and Hilibi, Kealedji, 
Okchai, Pakan Tallahassee, and Wiogufki about Hanna. Abihka, 
Otciapofa, Nuyaka, Latogalga, Kasihta, Hilibi, Pakan Tallahassee, 
and Wiogufki were visited, and new information obtained regarding 
all of the others except Eufaula, of which I secured very good 
descriptions 17 years before. The Eufaula ground, that of Asilanabi, 
one of the Tulsa grounds, Tukabahchee, Alabama, and Okchai were 
visited in the winter of tg11-1912. Kasihta is the only square ground 
representing the Lower Creeks now maintained. It was not in existence 
during my earlier work in the Creek Nation, nor were Laplako or 
Kealedji... These two last and the Yuchi grounds are the only ones 
that I have not seen. I was present at part of the busks at Otciapofa, 
Nuyaka, and Pakan Tallahassee. Not much attention was devoted 


*42d Ann. Rep., Bur. Amer. Ethnol.. 1924-25, pp. 204-206, 1028. 

*In the present paper, a indicates the obscure a in such a word as ability, and 
t or £. is a surd 1 approximating thl in English. 

*But see p. 35 regarding the former. 





SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 85, No. 8 


Zz SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


to the Seminole grounds, but information was obtained regarding one 
of these, Ochesee Seminole, which had been discontinued in 1912, 
when I visited the Seminole squares, but was afterwards revived. 

In order to make the material obtained in 1929 intelligible, it will 
be necessary to give a brief outline of the Creek political, social, and 
ceremonial organization. 

The name Creek is a shortened form of Ochesee Creek Indians, 
a name which English traders from South Carolina came to apply 
to that part of the Indians of the Creek Confederation who were 
living upon Ocmulgee River in the closing decades of the seventeenth 
century and the opening years of the eighteenth. The word Ochesee 
signifies “people of a different speech” in the language of the 
Hitchiti, one of the minor constituents of the Creek Confederacy, 
being equivalent to the word Tciloki in the Creek or Muskogee 
language. It was applied to the Creeks proper or Muskogee by the 
Hitchiti along with many other tribes, but came in some way to be 
particularly associated with the Muskogee and the river upon which 
they were then living. 

Anciently there seems not to have been a single term applicable 
to all of the Muskogee, the latter name having been unknown to 
the Spaniards who first entered this section. It does not make its 
appearance until the English had settled in the Carolinas. The origin 
of the word is uncertain, but there are indications that it was derived 
from Shawnee, since a band of Shawnee lived for a time near what 
is now Augusta, Ga., and from a very early period occupied an inter- 
mediate position between South Carolina and Georgia on the one hand 
and the Creek Nation on the other. It is probable that there were 
originally several tribes speaking the same language but having sepa- 
rate names and that the necessity for a distinguishing term for all 
did not present itself until the number of non-Muskogee tribes in 
the Confederation came to be considerable. As to the names of 
these Muskogee tribes, we seem to have indications of the following: 
Abihka, Coosa, Okchai, Pakana, Tukabahchee, Hilibi, Eufaula, 
Kasihta and Coweta (or perhaps an original tribe of which the 
Kasihta and Coweta were sections). There were some other groups 
on the lower course of Tallapoosa River, such as the Atasi, Kealedji, 
and Kolomi, which cannot be definitely placed and may have been 
independent of these or early subdivisions of them. Of course some 
may represent people of wholly different connections who had become 
assimilated to the Muskogee and had lost their own language and 
customs. This is rendered probable from the fact that we have 


| 
. 





No. 8 CREEK SQUARE GROUNDS—SWANTON 


ww 


several actual cases of such assimilation in later times. However, 
so far as the tribes enumerated are concerned, this must always 
remain in doubt. 

When the tribes of the Confederation first became known to 
white people, they were distributed geographically into two main 
sections to which the names Upper Creeks and Lower Creeks have 
become attached. The former were on the Coosa and Tallapoosa 
rivers and the upper course of Alabama River in the present state 
of Alabama, the latter on the lower and middle courses of the 
Chattahoochee, which now forms part of the boundary between 
Alabama and Georgia. It was this latter division principally which 
lived upon Ocmulgee River for a time and thus gave rise accidentally 
to the popular English name for the entire people. A minor division 
also existed between those Creeks living on the middle course of 
Coosa River and those centering about the lower Tallapoosa, the 
two being sometimes designated as Upper and Middle Creeks, 
respectively. In the distribution of the original Muskogee tribes, 
the Abihka and Coosa constituted the greater part of the Upper 
Creeks, while the Kasihta and Coweta were the dominating element 
among the Lower Creeks. The Okchai, Pakana, Tukabahchee, 
Atasi, Kealedji, Liwahali, Laptako, Kolomi, and a number of towns 
descended from the Coosa, including Otciapofa, the Tulsa towns, 
and the Okfuskee towns, besides several minor groups, formed the 
bulk of the Middle Creeks. The Eufaula had the distinction of 
being connected with all three. Their oldest seat seems to have been 
in the Upper Creek country ; later they established themselves among 
the Middle Creeks and about the period of first white contact they 
formed a colony well down Chattahoochee River, among the Lower 
Creeks. To complete the story of their migrant habits, we may 
add that they seem to have furnished the first true Muskogee 
contingent to the Florida Seminole in the Red House or Tcuko Tcati 
Indians north of Tampa Bay. 

Tradition seems to be borne out by circumstantial evidence in 
pointing to the Lower Creek country as that region in which the 
tribes in question began their federation. According to the story 
this had to do on the one hand with the division of the Muskogee 
into the Kasihta and Coweta and on the other with the relations 
between them and the non-Muskogee elements, particularly the 
Apalachicola. The relations of the Kasihta and Coweta to each 
other are somewhat uncertain, for while it is at times implied that 
they resulted from the fission of a single body of people, the most 
popular traditions speak of them as having come from the west 


4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


as two distinct tribes and it is possible that the one-tribe idea may 
be the result of later rationalizing. The story goes that, having 
defeated all of their enemies, the Kasihta and Coweta instituted 
periodical ball games as a kind of ‘moral equivalent for war,” and 
afterwards, when either of them established relations of friendship 
with other Indians, whether Muskogee or not, these Indians entered 
on the same side as their friends so that this dual system soon 
became general. 

One of these sides, that of Kasihta, came to be known as the 
White or Peace side, though it did not receive that specific name; 
while the side of Coweta was the Red or War side. 

At the same early period the Muskogee entered into intimate 
relations with the Apalachicola Indians, who spoke a dialect related 
to Hitchiti. This was the outgrowth of a treaty of peace following 
upon hostilities, or to avert threatened hostilities. The Apalachicola 
were then taken into the Confederation on the same side as Kasihta. 
In some particulars, however, they are held to have been more 
representative of the White towns than Kasihta and for that reason 
their settlement came to be called Talwa Lako, “ Big Town.” Indeed, 
the migration legend related to Oglethorpe by Tchikilli implies that 
Kasihta was at least partly Red, their hearts being “ red on one side 
and white on the other.” However, in all later times Kasihta 
assumes the leadership of the White towns among the Lower Creeks, 
as does Coweta the leadership of the Red towns. Four having 
been the sacred number—the sacred formulae being gone through 
four times, four arbors or beds constituting the ceremonial buildings 
in the square ground and four sticks the number employed in the 
ceremonial fires—it is not surprising that the Creeks should select 
two towns from the Upper Creeks, taken collectively, to add to 
these two leading Lower Creek towns. The White towns of the 
Upper Creeks were represented by Abihka, the Red towns by 
Tukabahchee, the second being from that group I have called Middle 
Creeks, the other from the northernmost bounds of the Nation. 
These four towns were the “back sticks”? of the Confederation, 
and each had a special ceremonial name, viz., Kasihta Lako (“ Big 
Kasihta”’), Coweta Mahmayi (“Tall Coweta’), Tukabahchee 
Ispokogi, and Abihka Nagi. Ispokogi was the name of the culture 
heroes of the Tukabahchee and it may be a Shawnee term. It bears 
a suspicious resemblance to that of the Kispokotha band of Shawnee. 
I do not know the meaning of Nagi. The Abihka were also called 
specifically ‘“ the door-shutters ” because they protected the northern 
frontier of the Confederation. 








no. 8 CREEK SQUARE GROUNDS—SWANTON 5 


In course of time the non-Muskogee element represented by the 
Apalachicola Indians was increased, first by other groups related 
to the last mentioned—such as the Hitchiti, Okmulgee, Sawokli, and 
Tamati—who spoke closely related languages and called themselves 
Atcik-hata, a term said to have some reference to the ashes of the 
ceremonial fire in the square grounds. These Indians formed the 
greater part of the first Creek invaders of Florida who presently 
constituted the Seminole nation. The leading town in this southward 
movement was Oconee, almost certainly affiliated with the Atcik-hata, 
and the titular leadership among the Seminole remained with them 
until after the Seminole War. However, the complexion of the 
Seminole as a whole was changed from Atcik-hata or Hitchiti, to 
Muskogee by the multitudes of refugees which fled to Florida after 
the Creek War of 1813-14. The later removal to Oklahoma seems 
to have reversed the situation since more than two-thirds of the 
Indians now in Florida speak a language of the Hitchiti group. 

There is strong evidence that the Chiaha Indians originally spoke 
Hitchiti and that the Mikasuki of Florida branched off from them, 
but some early event in their history separated them from the other 
Atcik-hata and made them allies of the Coweta. This friendship 
they shared with the Osotci who seem originally to have belonged 
to the Timucua linguistic group of Florida. To the Upper Creeks 
were added the distinct but dialectically related Alabama, Koasati, 
and Tuskegee, while bands of Yamasi and Apalachee were 
temporarily connected with both Upper and Lower Creeks. The 
Alabama town of Tawasa seems to have had an origin similar to 
that of the Osotci. At a very late date the wholly alien Yuchi 
population was admitted into the Confederation, most of them 
making their home among the Lower Creeks though there was a 
small body also among the Upper Creeks. And more divergent still 
were the Shawnee, from among whom two towns made their homes 
in Creek territory for several decades during the eighteenth century. 
One of these probably continued on into the early years of the 
nineteenth century. 

It may be added that towns are known to have changed from 
one side to the other. Alabama was once a White town closely 
associated with the Okchai, but later they were affiliated with the 
Tukabahchee and came to be reckoned as Red. Wiogufki, Hilibi, 
and Wiwohka are also said to have shifted from one side to the 
other. In the case of the two last this may be partially explained by the 
fact that, if we may trust native tradition, they were built up of 
refugees from other settlements. 


6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 85 


In former times a certain aloofness was maintained by the towns 
of one moiety toward those of the other. They did not encourage 
intermarriage and did not attend each other’s annual ceremonies. 
This latter inhibition is now breaking down and it is claimed that 
men of all towns attend the busk of Otciapofa. Otciapofa, however, 
has long occupied an exceptional position. A chief belonging to the 
Bird clan of this town always delivered the principal speech when 
a new chief of the Confederation was installed. This town was also 
the residence of the Creek dictator Alexander McGillivray, and it 
was here that Crazy Snake, leader of the Creek conservatives, 
called his important councils. Evidently the functions of the White 
and Red sides in maintaining peace or bringing on hostilities were 
formerly of great importance and some White towns, certainly 
Apalachicola and Coosa, were places of refuge for murderers. The 
“regular ” ball games, as distinguished from practice games, always 
took place between towns of different sides and the supporters of 
each town marched to the encounter in much the same spirit as if 
they were going to war. 

The principal White towns were: Kasihta, Apalachicola, Hitchiti, 
Okmulgee, Sawokli, Yuchi, Abihka, Coosa, Otciapofa, Tulsa, 
Okfuskee, Okchai (including Latogalga and Asilanabi), Pakana, 
Koasati, Tuskegee, and Wiogufki. 

The principal Red towns were: Coweta (including Likatcka), 
Eufaula, Chiaha, Osochi, Tukabahchee, Liwahali, Laptako, Atasi, 
Kealedji, and Hilibi. Alabama changed from White to Red in the 
manner described. 

The people of each town were subdivided into clans which were 
usually named after animals and were invariably perpetuated in the 
female line. The only clan of importance not named for an animal 
was the Wind clan and with this the Skunk was closely associated, 
the Skunk clan having always been linked with it. I obtained the 
names of over 50 clans but some of these were known to only one 
or two informants, and a number of others were small and bound 
into phratral associations with clans of greater prominence. Some 
clans were considered as equivalents throughout the entire nation. 
The Skunk, Fish, Rabbit, Otter, and Turtle seem always to have 
been united in one phratry with the Wind; the Wolf and Salt with 
the Bear; the Pahosa with the Deer; the Wildcat with the Panther ; 
and the Turkey and Tami with the Alligator. In the same way the 
Snake, Kapitca, and Woksi were almost invariably counted in with 
the Aktayatci; the Mole, Toad, and Tcikote always went together 
and were generally allied with the Deer and Pahosa; and the 








No. 8 CREEK SQUARE GROUNDS—SWANTON A, 


Lidjami, Eagle, Hickory nut, Fox, Cane, and Muskrat * were usually 
placed with the Raccoon. 

Some phratral associations, however, were confined to one or a 
few towns and did not extend throughout the nation. Thus the 
Potato was commonly placed in one phratry with the Raccoon but 
in Tukabahchee it was separated. The Beaver was usually placed 
with the Bird, but in Alabama it was quite distinct. On the other 
hand it was sometimes classed with the Alligator. Occasionally the 
Aktayatci formed one phratry with the Raccoon, and much more 
rarely the Deer and Panther were found together. Differences of this 
kind were due in some measure to the council system. Every 
important clan in a given town, or every group of related clans, 
held meetings during the annual ceremony known as the busk and 
each listened to an address by its oldest capable male member or 
“uncle.” If an individual came to live in a town in which his own 
clan or his phratral group was not represented, he would elect to 
affiliate with one of those already in existence. It was usual for all 
of the children of each group of this kind to consider themselves 
brothers and_ sisters between whom marriage was ordinarily 
prohibited. However, the information I received shows plainly that 
sexual intimacy between individuals of linked clans was not considered 
as serious as between members of the same clan. It is specifically 
stated regarding some of these clans that “they were kin” up to 
noon, or up to midnight, and separate the rest of the time, 7. e., a 
limited taboo. was maintained against them. It is also said that a 
man would sometimes pretend that a woman whom he wished to 
marry was of a certain clan, for which he would manufacture a name, 
although she was in fact of his own, and that, if he were a man of 
influence, he often “ put over” this new creation of his. On the 
other hand, I have been told that, even though children of certain 
of the primary clans were brought up together, they would never 
be regarded as brothers and sisters. It is quite plain that all sorts 
of variations had grown up in response to unpredictable situations. 

When one eliminates the obscure and the constantly linked clans, 
about nine are left of something like major importance. These are 
the Wind, Bear, Bird, Beaver, Alligator, Raccoon, Aktayatci, Deer, 
and Panther. We should perhaps add the Potato. The Beaver, 
however, has importance mainly in one group of towns, and the 
Aktayatci appear to have been rather closely associated with the 
Hitchiti and the Seminole, but also with Hilibi, Wiogufki, and 
Eufaula. 


“ee 





*In the 42d Ann. Rep., Bur. Amer. Ethnol. (p. 116) I erroneously called this 
the Mink. 


8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


The question naturally arises whether some of these clans may 
not have been brought in with formerly independent tribes. All we 
can say is that certain clans are more prominent in some of these 
tribes than among the true Muskogee but whether they were brought 
in by them we do not know. Thus, as just mentioned, the Aktayatci 
was particularly prominent among the Hitchiti, as were the Snake, 
Kapitca and Woksi, and in a more pronounced manner the Toad, 
Mole, and Tcikote. The Daddy-long-legs and Salt were similarly 
associated with the Alabama, the former, indeed, being hardly known 
outside of that tribe. 

Besides this division into phratral groups all of the clans were 
ranged in two moieties called respectively, Hathagalgi, ‘‘ White 
People,” and Tcilokogalgi, “People of a different speech.” The 
Wind and Bear with their phratral associates were almost invariably 
White, and the Raccoon and Aktayatci and their allies almost 
invariably Tciloki. The Bird is usually White but among the 
Alabama and Koasati it is Tciloki. The Beaver is also White usually, 
but when it is associated with the Alligator and when the Alligator 
is not a White clan, the Beaver often becomes Tciloki. The Alligator 
is most often Tciloki but in a number of towns it is White. The 
Deer is usually Tciloki but it is White in a few towns. Today the 
Panther is almost always Tciloki but some of the oldest myths and 
some of my best informants assert that it was anciently White. 

When I first went among the Creeks, I was told that in one or 
two towns the clan moieties were exogamous, but the greater number 
of my informants held the contrary opinion. I was much surprised, 
therefore, during my last visit to have most of my informants 
maintain that they were exogamous. This much is certain, that there 
were striking exceptions to this law in comparatively early times, 
for instance, in the case of the famous Creek speaker Hobohit Yahola. 
Probably it will never be possible to say whether this phratral 
exogamy was breaking down or growing in times known to us. In 
recent years the principal function performed by these moieties has 
been to determine the line-up of the players in practice games within 
the town. The important bearing the mere character of a name 
may have in social evolution is shown by the fact that, on account 
of the name, persons of European blood were usually reckoned as 
“friends ” of the Hathagas, and in consequence the latter acquired 
a reputation as “ progressives,’ while the Tcilokis were considered, 


oe : 93 
conservatives. 


and acted like, 
Besides the clans, phratries, and moieties there were certain groups 


in each town which had official functions. Some of these were 

















no. 8 CREEK SQUARE GROUNDS—SWANTON 9 


determined by the individual ability—usually military—of those who 
belonged to them. Thus a man might start his. public career as a 
common warrior or tasikaia, a word now often translated “ citizen,” 
be promoted to the position of an imata labotski, or “ Little Imata,”’ 
then to that of an imata tako, or “* Big Imata,” and finally become a 
tastanagi, or “‘ war leader.” If sufficiently prominent he might be 
made a tastanagi tako or hohbonaia, “‘ war speaker,” though of these 
there was never more than one in a town at any one time. There 
were ‘“ beds ”’ or seats in the square grounds for each of these classes, 
but not all were promoted into them. Men who belonged to the clan 
of the chief (miko) would be given seats in his section and form 
the mikalgi, ‘‘ chiefs,” who acted as a kind of special executive 
council. If they belonged to a certain clan known as henthalgi, they 
would be given seats in another place. The functions of the henthalgi 
are somewhat uncertain but they were concerned largely with the 
maintenance of peace and charged themselves with the internal 
prosperity of the tribe. The henthalgi were almost always formed 
of the Wind clan, and if, for any reason, the Wind clan could not 
be used, the Bird or Beaver, or at least some clan considered White, 
would take their place. There was also a class of men called ist- 
atcagagi, the old, experienced men from all tribes, retired from 
active service but keepers of the tribal lore. It seems fairly evident 
that a correlation existed anciently between White towns, White 
clans, and the henthalgi, and that the miko of a White town was 
generally chosen from a White clan. It is even possible that the 
miko of a Red town was formerly chosen from a White clan. 
Certainly there is a marked tendency to choose chiefs from the Bear 
clan, even in Red towns, though many of them are also from the 
Raccoon and the Aktayatei. 

A word must now be said regarding the ceremonial grounds. 
Originally every Creek town had such a ground which at a still earlier 
period was probably the ceremonial ground of a small tribe. As a 
tribe increased in numbers, however, the ground often became too 
small to accommodate all of its members comfortably and so it split 
into two or more. Undoubtedly, before the organization of the 
Creek Confederation, there was great diversity among these grounds 
and a‘ certain element of diversity has persisted until the present day, 
but on the other hand considerable standardization has undoubtedly 
taken place. Creek legend asserts that the first ceremonial ground 
was given to the Coweta, the Kasihta, or the Tukabahchee by the 
Breath Controller or by other supernatural beings, and copied by the 
remaining towns from them. While this represents a modern 


‘ 


LO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


rationalization, there can be no doubt that the earlier ceremonial 
grounds of the constituent members were altered in many particulars 
in conformity with the prevailing pattern. 

All the grounds known to us originally consisted of three elements, 
a tcokofa or community hot house used in bad weather or for secret 
ceremonies, a “square ground,” and a “chunkey yard,’ or ball 
ground. The name “ chunkey yard ” is derived from an old pastime 
which consisted in rolling a stone disk along a level plot of land 
and throwing certain long poles after it, the game turning on the 
relative nearness of the poles to the roller after all had come to 
rest. There was a single pole in the middle of this yard surmounted 
by a cow or horse skull or by a wooden figure, and about this men 
and women played against each other in a kind of ball game. This 
game was mainly confined to people of the town and was social in 
character while the great ball game, similar to our game of lacrosse, 
was played by men only and was highly ceremonial. The fact that 
the “chunkey yard” was a part of the ceremonial ground may 
indicate that the single pole game formerly had more religious 
significance than was the case in later times. 

The tcokofa has long been out of use, though at Tukabahchee fire 
was until recently lighted in the middle of a circular offset of the 
ceremonial ground where this structure would stand if it were still in 
existence, and one such building was put up at Pakan Tallahassee 
after the Civil War. 

The most important part of the ceremonial area today is the 
“square ground,” so called because in the largest towns there were 
on it four long cabins or arbors, in native parlance “ beds,” forming 
four sides of a square. Partly from tribal idiosyncrasy and still 
more on account of failing numbers, several of these grounds now 
lack one cabin, and the Alabama ground lacks two. Today the 
cabins consist merely of two or three rows of split logs to serve 
as seats and an arbor of boughs to shield their occupants from the 
direct rays of the sun, but anciently the seats consisted of mats 
woven out of cane raised upon short posts and the cabins were 
provided with a back and roof of wattle or split shingles plastered 
with clay. The arbors in the largest modern towns are supported 
on eight posts, four in front and four behind, but some have only 
six, and most of the Seminole towns only four. On the other hand, 
a sketch of one of these cabins made by a Frenchman early in the 
eighteenth century shows ten posts, five in front and five at the 
back. Today, however, the eight post arrangement seems to be 
considered orthodox, and the three sections marked off by these 





No. 8 CREEK SQUARE GROUNDS—SWANTON It 


posts are used for the seating of as many clans, groups of clans 
or related officials. 

Considerable variation in the ancient and intended plan has been 
brought about by the attrition which the tribe and its several 
divisions have undergone, loss of the keepers of the sacred lore, 
and other factors, but it is plain that normally one of the four cabins 
was mainly devoted to the miko and his clan. Hence it was called 
mikalgi (or mikagi) intupa, the “ Chiefs’ bed.” Another was devoted 
mainly to the henihalgi and was named from them, another to the 
higher class of warriors, the tastanagalgi, and so received their name, 
and still another to the novitiate warriors or youths from whom it 
was called tasikaialgi intupa or tcibanagalgi intupa. The positions 
of these in the square ground varies considerably. It should be 
stated in the first place that the cabins are placed normally toward 
the four points of the compass, but that for some unexplained reason 
in the square of Tukabahchee the entrances are toward the cardinal 
points. In the old Kasihta square, which seems to have set the 
fashion for many other towns, the mikos’ cabin lay west, the henihas’ 
south, the tastanagis’ north, and the tasikaias’ east. In the Okfuskee 
towns, of which Nuyaka is an example, the mikos’ cabin lay north, 
the henihas’ east, the tastanagis’ west, and the tasikaias’ south. In 
Pakan Tallahassee the mikos’ cabin is north, the henihas’ south, the 
tastanagis’ west, and there is no east cabin. In Eufaula the mikos’ 
cabin is north, the henihas’ east and the tastanagis’ south, the west 
cabin being missing. In Alabama, where there are but two cabins, 
the mikos’ cabin is east. 

The other variations in the arrangement of the squares will come 
out in the subjoined material, but before presenting it something 
must be said regarding town officials. 

The miko, as already stated, was chief of the town, its head 
presiding officer and responsible executive. Like many of the other 
officers he had a special companion or heniha taken from the Wind 
clan or whatever clan occupied the position of henithalgi. This heniha 
is sometimes called miko apokta, “ second chief,” but the latter name 
is also given to still another functionary who shares the burdens 
of state with his superior. The chief had one or more yatikas or 
interpreters who also bore the name asimbonaia, ‘‘ speakers.” Unless 
wanted to make an announcement or to send upon an errand, they 
sat with their clan or other group to which they normally belonged. 
The tastanagi tako and holibonaia have been mentioned. The position 
of hotibonaia, “ war speaker ” was the most exalted military position, 
and it is possible that there was only one such official at a time in 


WZ SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


the entire Creek Nation. It was the position occupied by Hobohit 
Yahola, famous leader ot the Creeks during their removal west and 
all of their subsequent troubled history until the Civil War. He 
was not a miko even of one town, but his influence was actually 
greater than that of any miko, or any number of them. There were 
two ta‘pala, whose chief function was to act as messengers before 
and during the women’s dance, and there were two singers for the 
women (inyahaikalgi, “ singers for them”), who sat at the end of 
one of the cabins just in front of the spot where the women began 
dancing. They were usually selected for their knowledge of songs 
rather than on account of their clan affiliations. The hilis haya was 
the head priestly functionary. He supervised the preparation of the 
medicines and gave them their final potency and he ordered 
everything in any way touching upon the supernatural. He was 
assisted by one or more men called hilis tcalaba or “ medicine 
mixers,’ whose functions are defined by the name, and by two or 
more young men called hilis hoboia, ‘ medicine gatherers,’ who 
collected the red root, pasa, and other plants that went into the 
sacred medicine. Some towns seem to have had a separate official 
called tutka didja, “ fire builder,” to start the fire, but in others the 
hilis haya did that, and there was instead a tutka oktididja, “‘ gatherer 
of wood” for the fire. A number of boys known as oidjawalgi 
brought water for the medicines. In one town we learn of ahaga 
haiyalgi, “‘law makers,” who are said to carry out the instructions 
of the tastanagis and may be identical with the imatas elsewhere 
mentioned. There were also officers called simiabaia, or “ leaders.” 
The hoktagi immiko, “ 
have been identical with the ta‘palas. Anciently the tastanagis and 
their assistants acted as town police, but nowadays three or four 
light horsemen are selected at random to police the square ground 
during ceremonies, and there are boys called “‘ dog whippers ”’ with 
long whips stationed at each opening into the square during the 
women’s dance to drive away dogs. Many of these officers were chosen 
for four years only. If one died before the expiration of his term, 
a substitute was selected from the same clan, which seems to indicate 
that the position was something of a clan prerogative. 

Of the ceremonies which took place on these grounds, only two 
have survived. One, called by the popular local name of “ stomp 
dance,”’ was confined to the people of the town and was simple in 
character, the miko hoyanidja (red willow) being ordinarily the only 


chief of the women,” and his henitha, may 


medicine used during it. There were no dances other than the common 
and relatively secular ones, no ceremonial lighting of the fires, no 








no. 8 CREEK SQUARE GROUNDS—SWANTON 13 
ceremonial complications of any sort. It seems to have been usual 
to hold three of these dances in the spring and early summer, a 
month apart, the series paving the way for, and leading up to, the 
second ceremonial, the “ busk,” which was the great annual ritual. 
This last is usually considered as lasting four days, though in that 
four are included the days of assembling and departure. The 
principal event on the second day is the women’s dance. On the 
third day the men fast, take medicine four times, and near its close 
march down to the creek and bathe. After they return, they are 
dismissed to their camps and break their fast. Later they are 
summoned to the square again where they dance four times, and 
then the dance becomes “‘ common,” visitors from friendly towns 
being admitted to it. A fire is kept up all night in the center of the 
square and dancing continues about it until it is nearly day. In 
former times many, perhaps all, of the towns extended their busks 
over eight days, but from what can be learned of these longer 
ceremonials they seem to have been in the main a simple doubling 
of the shorter ceremony, except for a few features like the kindling 
of the new fire which took place but once annually. 

Let us now turn to the new material regarding the square grounds 
and the ceremonies conducted there. Under each heading I give the 
notes obtained from native informants belonging to the square or 
town in question. 


ABIHKA 


Figure I gives the general arrangement of the square of Abihka, 
‘or Talladega, and Plate 1, Figure 1, gives a view of the ground itself 
near Henryetta, Okla., as it appeared in 1912. The exposure was 
made from the southwest. 

The medicines were taken first by those in the Bears’ bed, then 
in succession by those in the beds of the Raccoon on the west side, 
the Raccoon on the east side, the Deer, and those in the south bed. 
The asimbonaia in the northern section of the east cabin acted more 
particularly as the chief’s messenger; the one in the western 
compartment of the south cabin called first the women and then 
the children to come to take their medicine from the vessel at the 
north end of the east cabin. The ta‘palas were changed every four 
years. The women gathered preparatory to their dance at a tree on 
the edge of the tadjo (the ridge of sweepings that makes the edge 
of the ceremonial ground) and entered’ the section about the fire at 
its northwestern corner. Five pots of medicine were prepared at 
the north end of the west cabin (fig. 1, 11). Afterward one pot 


14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


was placed a little farther north and west for the boys (12) and 
another was carried around to the north end of the east cabin for 
the women and children (13). The ingredients of this medicine 


N 


* 
5 





Fic. 1.—General arrangement of the square of Abihka. 


A. Chieis’ Bed (mikalgi or mikagi intupa); I, miko (Bear); 2, heniha 
(Raccoon) ; 3, hilis haya (Raccoon) ; 4, hilis tcalaba; 5, yahaikas (any clan). 

B. Henihas’ Bed (henihalgi intupa): 6, asimbonaia (Alligator); 7, ta‘pala 
(Panther) ; 8, ta‘pala (Wind). 

C. Warriors’ Bed (tastanagalgi intupa) : 9, asimbonaia (Raccoon) ; 10, tutka 
didja (Deer). 


II, medicine pots (Ist position) ; 12, medicine pot for boys (2d position) ; 
13, medicine pot for women (2d position); 14, place where medicines were 
piled immediately after they were brought in; 15, ball post (pokabi). 
It is to be noted that, in all of these diagrams, the ball post was actually much 
farther from the center of the square than is indicated. 


were miko hoyanidja (“red root”), pasa (“ buttonsnake-root ’’), 
wilana (“ wormseed”), and hobaga (“maypop’’); tcato hatki 
(‘‘ white stones”) were added. After the ceremony was about over and 
the fasters were ready to go down to bathe in the creek, what was left 














no. 8 CREEK SQUARE GROUNDS—SWANTON T5 


of the medicine was poured on the fire. It is said that one gallon of 
spring water was brought for all five pots. This must mean a gallon 
for each. Following are the busk names of the present officers : 


ATUL Oey cote erenck retort ce neeh iors ate a one aie ors Tctktcat Heniha 

eMMiliah By: Ae-pert sensi eee cise keccte eine ae Itchas Hadjo 

nilis*tcalabay ac access oe. saiseles Konip Yahola 

tals ail dea ragetenete baeyeters witiehe se Sherer tiece sic Katca Tastanagi 
el NEST Ser as steiodte octet weaxcyeus.e eens etene he Kona Yahola 

EUTECRGIC |i ae sepa re atniaie Sere *.... Tastanakutci 

ASI DOMlal awe ete ie eieioe es cleyen is aietere Wotko Fiksiko 


“ 


Bia RPM SiGe TELS Kapitca Hadjo 


The Hathagalgi of this town are Wind and Bear; the Tcilokogalgi 
are Panther, Raccoon, Deer, Beaver, Alligator, and Bird. 

My information regarding this ground was obtained mainly from 
Jim Star who described the Talladega ground to me in 1912, the 
plan of which is in the Forty-second Annual Report of the Bureau 
of American Ethnology, page 205. The different aspect of the east 
cabin is mainly due to the fact that the earlier account gives a more 
ancient arrangement, when the warriors were graded into tastanagis, 
and big and little imafas. The position assigned to the hilis haya 
in the earlier plan is probably erroneous. The other differences are 
due mainly to the more extensive information obtained on my last 
trip. My new information disagrees with the older, however, in 
assigning the Bird, Beaver, and Alligator clans to the Tciloki side 
as was said to be the case at Abihka-in-the-West instead of to the 
White side as was given me for Talladega and the old Abihka town 
near Eufaula. It is probable that the new information is correct 
since the last mentioned square was given up when the man from 
whom I obtained data regarding it was a boy. However, it must be 
remembered that these allocations are not invariable and probably 
changed at times even within the same town. 


OTCIAPOFA, OR HICKORY GROUND 


Figure 2 shows the square ground of Otciapofa, popularly known 
as Hickory Ground. 

The hilis tcalaba was changed every four years and was not taken 
invariably from the same clan. 

At the southernmost front post of the east cabin were fastened 
two poles with black feathers tied to the ends and at every other 
front post were three similar poles but with white feathers. These 
were carried by the men in the “ feather dance.” 


2? 


16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 85 


In the women’s dance there is but one leader who carries a notched 
stick called atasa, the old name of the war club, from the middle 
of which depends an eagle feather. 

Besides the invariable miko hoyanidja or “red root,’ the busk 
medicine contained tutka hiliswa (“fire medicine’), wilana 
(““ wormseed ’’), tutka-tcok-hissi (a place on the ground where wood 
has been burned and moss has sprung up), and hobaga (‘‘ maypop”’). 


‘ 


< AKTAYATC/ 
iK AND 


BEAVER 
AND 
ALLIGATOR 

° 


\ 
‘ 
‘ 
' 
‘ 
‘ 
‘ 
t 
t 
t 
t 
‘ 
' 
t 
t 
' 
v 
' 
’ 
‘ 
‘ 





Fic. 2—The square ground of Otciapofa, or Hickory Ground. 


A. Chiefs’ Bed: 1, miko (Beaver); 2, miko apokta (Beaver); 3, hilis haya 
(any clan; at present Wind); 4, hilis tealaba (changed every 4 years; now 
Beaver). 

B. Henihas’ Bed. 

C. Warriors’ Bed. 

5, medicine pots (Ist position); 6, medicine pot for boys (2d position) ; 7, 
medicine pot for woman (2d position); 8, log on which medicines were laid 
and macerated with a small wooden pounder; 9, place where medicines were kept; 
10, point where women gathered preparatory to their dance; 11, point where 
women made a final stop before entering to dance; 12, where the ashes from 
the central fire were deposited every year; 13, sweepings from the square ground 
(tadjo) ; 14, ball post. 


As usual, the doctor perfected this medicine by blowing into it 
through a hollow reed. Two men took medicine at the same time, 
using gourd dippers. After all were through a dipperful was poured 
On the fire: 

For a drum they use a stout jar and there are two coconut-shell 
rattles. 

There is a line of tadjo.(sweepings) around the four cabins but 
it does not include the ball post. In playing the single pole game a 











no. 8 CREEK SQUARE GROUNDS—SWANTON 7 


hit on the skull at the top counts five and a hit on the pole above 
a certain mark counts two when it is struck twice in succession. 

The Hathagas consisted of the Beaver, Alligator, Bird, Bear, 
Skunk, Wind, and Rabbit; the Tcilokis of the Raccoon, Deer, 
Panther, and Aktayatci. The Rabbit, Wind, and Skunk formed one 
phratry. 

The information regarding this ground which I obtained in 1912 
was particularly incomplete. It is therefore gratifying to find 
that there are no serious discrepancies between the plan based on 
that (42d Ann. Rep., Bur. Amer. Ethnol., p. 211) and the present 
information. Some clans are given in one and omitted from the 
other, but where the same ones appear in both they have practically 
the same positions except for the Bear, which, according to the 
earlier description, sat in the west cabin and according to the later 
at the east end of the north cabin; and the Tami, which the former 
places at the west end of the north cabin and the latter at the back 
of the northern section of the west cabin. The location of the 
miko, miko apokta or heniha, and hilis haya is probably more exact 
in the later plan which also adds many more details. The informants 
differed somewhat regarding the Hathagas and Tcilokis, the earlier 
authority placing the Beaver among the Tciloki and the Panther 
among the Whites, allocations exactly reversed by my later informant. 


Dies. PU Ss 


Next we come to one of the two divisions into which the Tulsa 
have recently split, this being known as Little Tulsa (fig. 3). Plate 1, 
Figure 2, shows the old Tulsa ground in 1911 from the southeast, 
before the fission had occurred. 

It is said that all of the offices are filled from particular clans. 
The following is a list of the present officers; exclusive of the miko 
and his heniha: 


ASTANA CAN AAO 2 --cer eh cisiseclerse seaman Ispani Tastanagi 
RUG frUllal Clete enero mites ree mene et set arora Kapitca Tastanagi 

A NES RMR tate o<as Toate gues ’s, ah areas. as aie Lata Miko (controls the two above) 
ahaga haiyalgi (“law makers”)...... Kapitca Tastanagi (Aktayatci), and 

(messengers for the tastanagis ) Tami Tastanagi (Tami) 
SUIT UT cl Del AR ea er eyeiae rele iehe exert hae iercnare wise 30/6 Kantcati Miko (busk name) or 
Nokos Hadjo (common name) 

IITs iach Vicia Meet ates res os cin iat chee once Miko Tcapko (Beaver) 
Inilisiatcalalawesvere s skteeie nes «arenes one Tami Yahola (Tami) 


faticao atic ticle vemta sites: soa satus ties se. Tamatakutci (Tami) 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 





YOUTHS 





\ARTAYATC / 


Oo 





B 


Fic. 3.—Little Tulsa square ground. 


A. Chiefs’ Bed: 1, miko; 2, heniha; 3, simiabaia. 
B. Henihas’ Bed. 

C. Warriors’ Bed. 

D. Youths’ Bed. 


4, old location of medicine pots; 5, modern location of medicine pots. 





No. 8 CREEK SQUARE GROUNDS—SWANTON TQ 


AED all AMR versyareye.sls nxcpnlehanciahes creas stereiae Nokos Fiksiko (Bear) 
SMU EGER, a 0ss te of cys aig Ae. olde di ovayah ote leheue stots Tami Hatkutci (Tami) 
AAU AST ave cars. eesctie ox Bis Sreisieie! s oie auetaa ave Kapitca Fiksiko (Aktayatci), and 
Pin Hadjo (Tami) 
Captain of the Light Horsemen........ Yahola Tcapko (Tami) 
Oktcan Hadjo (Bear) 
To hit MAOUSEMIe! ee sree nee le eileen .. Fas Yaholute (Beaver) 


Kapitcutci (Aktayatci) 


The ta‘palas functioned at the women’s dance, the yahaikas sang 
at the women’s dance and at the feather dance. 

My original Tulsa data, published in the Forty-second Annual 
Report of the Bureau of American Ethnology (p. 213), was obtained 
from an old man and was intended to reflect the most ancient 
arrangement he could remember. Since that time the Tulsa Indians 
who used to meet at the Little River ground have divided and 
maintain two distinct squares. The general agreement between the 
older and the later plan is therefore surprisingly close. The principal 
difference seems to be in the position given the Aktayatci who appear 
in the north bed in the earlier plan and in the south bed in the later 
one. The earlier plan may also be in error in the position given the 
medicine pots but this was subject to change from town to town and 
during the ceremony itself. The Eagle clan, which appears on the 
older plan, died out so long ago that it can barely be remembered 
by any living Creeks. 

NUYAKA 

The plan of this ground is given in Figure 4 and a view of it as 
it appeared some years ago in Plate 2, Figure 1. 

The positions of hilis haya, hilis tealaba, and tutka didja were held 
for four years when the man and clan were changed so as to teach 
others the duties of these offices. The tastanagis and imatas had 
become confined to one or two clans. The hoktagi immiko and his 
heniha controlled the women’s dance and were called istatcagas. The 
term hola‘ta was applied to a certain class at the square ground 
in some towns, sometimes to the Tcilokis, but its application here is 
not explained. The first ta‘pala acted under the women’s chief. He 
was taken from the Bear clan, or, failing that, from the following 
in order of preference: Wind, Raccoon, Tami. 

In taking the medicines they drank of the miko hoyanidja first 
and pasa second. : 

A rock was placed under the miko’s seat “‘ to make the seat heavy.” 
Anciently there was a tcokofa northeast of the square. Nowadays 
the ground is hoed off only once a year. 


20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


B/RD 


AND 


BEAVER 


TASTAIVVAG/S 


HOLA TAS | ALLIGATOR 


o1i3 
Fic. 4.—Nuyaka square ground. 


A. Chiefs’ Bed: 1, miko (Bear); 2, heniha (usually Wind; Deer in 1929) ; 
3, hilis haya (changed every 4 years; Turkey in 1929); 4, hilis tealaba (Alli- 
gator); 5, tutka didja (Raccoon). 

B. Henihas’ Bed. 

C. Warriors’ Bed; 6, two tastanagis (or law makers). 

D. Tcilokis’ Bed (Tcilokogalgi intupa), or Tasikaias’ Bed (Tasikaialgi 
intupa): 7, hoktagi immiko (Aktayatci) ; 8, heniha for hoktagi immiko (Alli- 
gator); 9, Ist ta‘pala (Bear preferentially) ; 10, 2d ta‘pala (Alligator). 


11, medicine pots; 12, point at which women enter; 13, ball post. 








No. 8 CREEK SQUARE GROUNDS—-SWANTON 2] 


In this town the Turkey and Alligator clans belong in one phratry 
and so do the Wind and Skunk. 
The present officers are as follows: 


5100) COMES rey oan yh con tree ee Nokos Miko 
Weta Meat ects ciciereveletersia sretersee? ofetenttciers Miko Tcapko 
HntlsS Una yaar eese he araee ates al rotons coats. ohs. crepe alas Tastanakutci 

EUG ae © cts C11Cl ici parapet eas ore elles sewtedate ete vee 2) Hotalgi Hadjudji 
Hig tSpec Gall lly daeareteuste exe rstons coleasietivein ie teins Lodja Yahola 
Moktaei 1mMMiIkOWs. a. .<cessis es seen. Wotko Yahola 


When the present Nuyaka data are compared with that which | 
obtained in 1912 for Nuyaka and the related towns Okfuskee, 
Abihkutci, Talmutcasi, and Tcatoksofa, the agreement is found to 
be close except in the cases of the two last where the square grounds 
had long been given up and were described by individuals from 
memories of their early years. The main correction is in locating 
the miko, heniha, and hilis haya and the difference here is not great. 


PAKAN TALLAHASSEE 

Figure 5 gives the plan and Plate 2, Figure 2, and Plate 3, Figure 1, 
views of the ground, one showing the three cabins, or arbors, and the 
other the chunk yard and ball post. 

The tutka oktididja, hilis tealaba, hilis haya, and oidjawas, were 
appointed every four years from any clan. The ta‘palas and hilis 
hoboia were appointed every four years from the same clan. The 
miko and asimbonaias held their positions for life. 

Here we seem to meet some strange innovations. The Bears’ 
section of the south cabin receives one name connected with war, 
tasikaialgi intupa, and the section of the Birds, Beavers, and 
Alligators another, tastanagalgi intupa, while, at the same time, they 
are White or Hathaga clans and their cabin is called hathagalgi 
intupa. Yet one section of the west cabin is called tastanagalgi intupa 
also, and the whole cabin receives the unusual name of laksafaskalgi 
intupa, “bed of the Blacks,’ the Blacks being evidently the clans 
elsewhere called tcilokogalgi. 

The tcokofa was to the northwest and this was the last town to 
put up such a structure. 

The Birds’ section was called istatcagagi intupa. The Deer and 
Pahosalgi were formerly called the imatagalgi. 

Three poles with white feathers attached were fastened to each 
of the front posts for use in the feather dance. 

Back of the tastanagis’ section of the west cabin was a little 
structure in which to inclose the medicine pots when they were not 
in use. 


No 
bo 


SMITHSONIAN MISCELLANEOUS COLLECTIONS 





voL. 85 
N 
oT TO O 
196 
© 
© 
48 
© 
© 
1 23 
Mt | 
e | 
15 
14% a | 
12 





% 
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een tt 


Fic. 5.—Square ground of Pakan Tallahassee. 


A. Chiefs’ Bed (mikalgi intupa): 1, miko (Bear); 2, hilis haya (Panther 
in 1929) ; 3, yahaikas (singers for women) ; 4, dog whipper. 

B. Whites’ Bed (hathagalgi intupa): 5, asimbonaia (Bird); 6, ta'‘pala 
(Bear) ; 7, hilis hoboia (Bird) ; 8, oidjawa (Bird) ; 9, oidjawa (Bear) ; 10, dog 
whipper. 

C. Blacks’ Bed (laksafaskalgi intupa): 11, asimbonaia (Deer); 12, ta‘pala 
(Bird; father Raccoon) ; 13, hilis tealaba (Deer in 1929); 14, tutka oktididja 


(Raccoon in 1929); 15, hilis hoboia (Raccoon in 1929) ; 16, oidjawa (Panther 
in 1929); 17, dog whipper. 


18, medicine pots (Ist position) ; 19, medicine pot for boys (2d position) ; 
20, medicine pot for women (2d position); 21, drum; 22, box for tobacco; 23, 
place where medicine was laid before being used; 24, woodpile; 25, ball post. 
The dotted line marks the course pursued by the women when they entered to 
dance. 





no. 8 CREEK SQUARE GROUNDS—SWANTON 


to 
© 


In this town the Hathagas were the Bird, Alligator, Beaver, Bear, 
Tami, and Wind. The Tcilokis were the Deer, Raccoon, Panther, 
and perhaps Aktayatci. The two moieties were anciently exogamous. 

The particular opponents of this town in regular ball games were 
the people of Atasi but they also played against [ufaula Hopai, 
Alabama, Hilibi, and Upper Eufaula. The Koasati Indians are said 
to have divided up in the ball games, some playing on each side. 

The names of the present officers are: 


TTT <Ouseiewls em eis es Nokos Miko (‘“ Bear Miko”) (Bird) 
ASiMVONAIA® “Vaan Tastanak Imata (Deer) 
or. Tf apeta. dhe Tastanak Hadjo (Bird) 
Ihilis haya >....:. Katcutci (‘‘ Little Panther”) (Panther) 
hilis tcalaba ..... Nokos Hadjutci (‘“ Little Bear Hadjo”) (Deer) 
oidjawalgi ....... 1. Fus Yaholotci (“Little Bird Yahola”) (Bird) 


No 


. Hotalgutci (‘Little Wind”) (Bear, father Wind) 

3. Halak Hopai (“ Potato Hopai”’) (Panther, father Rac- 
coon, Raccoon and Potato belonging to the same 
phratry ) 

hilis hoboia ...... 1. Itco Ilutci (‘‘ Little Deer Foot’’) (Raccoon, father Deer) 
2. Talsi Yahola (Bird) 
Easel Mieteteveseteee eee 1. Halak Yahofla (“Potato Yahola”) (Bird, father Rac- 
coon) 
* weeeeeeeee 2 Hotalgi Hadjutci (“Little Wind Hadjo”) (Bear, father 
Wind) 
tutka oktididja ...Itco-ili Imata (“Deer Foot Imata’’) (Raccoon, father 
Deer) 
Valiaikal Sipe meresre rie 1. Pahos Fiksiko (Raccoon, father Deer with which the 
Pahosa is affiliated) 
Be) Reece. 2. Miko Yahola (Bear) 


If one town wished to play a match game with another, said my 
Pakan Tallahassee informant, they sent a man to that town with a 
ball stick, and when thé people of town number two had reached an 
agreement they sent the ball stick back. My informant said that he 
then had a ball stick hanging up in his house which had been sent 
by the Alabama. 

In the women’s dance, the atasa held by the leading woman is 
red and has an eagle feather attached to it; that of the second is 
white and has a feather of the fus hatki (“‘ white bird’), a bird 
found down by the creeks, attached to it. They used from 12 to 
14 terrapin rattles. During the women’s dance the two ta’palas stand 
about where the two pots nearest the west bed are in the plan. Each 
carries a wand called si’dik-kika having a white feather fastened 
to the end. 

The dog whippers used in this dance are taken from any clan. 


24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


About 54 men were present at the last preceding busk and 25 
women and girls participated in the dance. 

At the top of the pokabi in this town was a horse skull. In scoring 
for this game they draw a line from the ball post to the nearest 
corner of the south cabin. When the skull is struck, 4 are scored; 
when the pole under the skull is struck, it counts 2 on the way out 
and 1 on the way back. They may count it as a game, however, by 
agreement when they reach the corner post. The women’s tallies are 
marked on one side of the line, the men’s on the other. 

There is little difference except in detail between the above plan 
of Pakan Tallahassee and the several I recorded in 1912. 


WIOGUFKI 

The plan of Wiogufki is given in Figure 6 and a view of the ground 
from the southwest in Plate 3, Figure 2. 

There is a little log house on the grounds in which the pots are 
stored when not in use to keep them from being broken. There 
never was a north cabin so far as my informant knew. On the upper 
end of the ball post is a cow skull. 

The women walk four times around the fire; then their leaders 
stop opposite the singers and they begin to dance. 

The hilis tcalaba holds office for four years. In this town the place 
of the miko’s heniha is taken by a tastanagi. Indeed all of the 
tastanagis are considered the same as the henihas. They are called 
“the people who are named” and are of the nature of lawmakers 
and assistants to the miko. The miko’s tastanagi is also the same 
thing as the yatika. There are no istatcagagis (retired leaders who 
acted as councillors), and no Creek town now has a holibonaia. There 
are five water boys picked at random. At the front posts of each cabin 
are four poles with feathers tied to the ends for use in the “ feather 
dance.” 

The two leading women in the women’s dance carry atasa. The 
principal function of the ta‘palas is to call the women up for their 
dance which they do four times. Each has a wand with a little white 
feather at the end. Their official positions do not end with the 
women’s dance but continue to the end of the busk. 

No medicine is now put on the firesticks but it was formerly done. 
The only medicine they use at the busk is the miko hoyanidja, to 
which nothing is added. 

The Hathagas and Tcilokis were the same as in Hilibi. They 
were exogamous, and if the exogamic law was violated the ears of 
the culprits were cut off. 








“= 


no. 8 CREEK SQUARE GROUNDS—SWANTON 25 


The towns of Wakoka1, Tukpafka, and Talahasutci were all one 
with this. In regular ball games they always played against Alabama. 
Comparison between the plan given above and the two obtained 
in 1912 shows considerable differences, but since one of the former 




















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TASTANAGIS a A aN 
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Roo hes 10 
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BiRPD own-: BEAR AND 
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SAT WEITHER, HE RE 


B 


Fic. 6—Square ground of Wiogutki. 


A. Chiefs’ Bed: 1, miko (Alligator); 2, hilis haya; 3, hilis tealaba (Bear 
and Deer); 4, hilis hoboia (Panther); 5, a tastanagi (instead of heniha) 
(Raccoon). 

B. Whites’ Bed: 6, yahaikas (Deer). 

C. Warriors’ Bed: 7, ta‘pala (Raccoon); 8, hilis hoboia (Deer); 0, tutka 
oktididja (Deer). 


10, medicine pots (near bed A for men; near bed C for women and boys) ; 
II, point where women enter to dance; 12, point where women begin dancing; 
13, woodpile; 14, ashes of old fires; 15, ball post. 


was obtained from the same man, I think the explanation lies in 
the smallness of the town and the weakness of many of the clans 
which has resulted in many changes within a comparatively short 
time. Nevertheless there is a general correspondence and _ the 
allocation of clans to the moieties also agrees except in the case of 


26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


the Panther clan. This is explained by the fact that the Panther 
was anciently considered a White clan and later came to be regarded 
as Tciloki. 

OKCHAI 


Figure 7 shows the arrangement of this square and Plate 4, Figure 
1, shows a view of it taken in winter. 





SIDYNULS UL 


SILL 4 ILV7 M/ 
‘SUSOH YJ 





O | SIDYNYLSUL 


B14 
© 
V/SITORS 
0D £XCEPT O 
6 
19 





Fic. 7—Arrangement of square ground of Okchai. 


A. Chiefs’ Bed: 1, miko (Raccoon formerly; Bear in 1929); 2, heniha 
(Deer); 3, yatika (Deer formerly; Wind in 1929); 4, hilis haya; 5, ta‘pala 
(any clan) ; 6, hilis tcalaba (Bird) ; 7, tutka oktididja (Wind). 

B. Henihas’ Bed: 8, ta‘pala (any clan); 9, hilis hoboia; 10, yahaikas (any 
clan). 

C. Warriors’ Bed. 


D. Youths’ Bed: 11, oidjawa (any clan). 


12, medicine pots; 13, medicine pot for boys; 14, medicine pot for women; 
15, point where women assemble preparatory to their dance; 16, point where 
women start dancing; 17, ashes of previous fires; 18, woodpile; 10, ball posts; 
20, a spring lies in this direction. 


The hilis tcdlaba, ta‘pala, and tutka oktididja changed every four 
years. The hilis haya was reappointed every four years. 

Wiley Buckner, an old informant of mine, was the former yatika. 
The tutka oktididja in 1929 was Hotalgi Miko. 








no. 8 CREEK SQUARE GROUNDS—-SWANTON 27 


The medicine was pounded up just in front of the pots at the 
north end of the chiefs’ bed. 
_ The fire was brought from Alabama in the great migration and 
new fire was lighted from it. They dug up the earth as deep as the 
arm would reach, put the fire in there, and made the new fire on 
top. This is the very spot at which they first placed their square 
in Oklahoma and it has not been moved. The squares of the kindred 
towns, Latogalga and Asilanabi, are not so old. 

There were two poles with feathers on the ends at each of the 
front posts of the cabins all the way round. 

Atasa were borne by the two women who led in the women’s dance. 

The tastanagis were law makers. They had charge of the rules 
governing the taking of medicine and if anyone broke one of these 
regulations they made him stay in the square ground all night without 
eating instead of breaking his fast that evening as was usual. 

In the various beds of this square ground the “‘ sons of the clan’ 
can sit with their fathers. 

The Hathagas and Tcilokis were the same as at Hilibi. 


Their principal opponents in the ball games years ago were the 


Hilibi. \ 9 

This arrangement agrees substantially with that obtained by me 
in 1912 (42d Ann. Rep., Bur. Amer. Ethnol., p. 234), the principal 
difference being in the location of the Bear and Bird clans. My 
earlier informants placed the Bird and Aktayatci in the middle of 
the south cabin and the Bear at the east end of the same, while the 
later ones said nothing of the Aktayatci but placed the Bird clan in 
the southern section of the west cabin and the Bear clan in the center 
of the south cabin, the sons of the Bear being located next to them 
on the east. The arrangement of this square must be taken in 
connection with the plans of Latogalga and Asilanabi. 


, 


LALOGALGA, OR FISH POND 

For a plan of this ground, see Figure 8. 

The tutka oktididja is appointed every four years but always from 
the Aktayatci clan. The hilis hoboia are chosen every four years and 
ordinarily from the Aktayatci and Bear clans but this is not necessarily 
the case. However, if one dies before his four years have expired 
he is replaced by someone of his clan. The hilis hoboia also act as 
the hilis tealaba. 

Ta‘palas are appointed only when needed. There are no special 
water carriers, the hilis hoboia calling upon any boys for this purpose 
whenever water is required. 


28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


The two leaders among the women carry atasa, that held by the 
first being colored red, that by the second white. 

One of the two medicine pots contains pasa; the other the miko 
hoyanidja along with wilana, tutka hiliswa (“ fire medicine”), which 


Be chars 


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us : 
4 

Re nt 


> 
U 


es 
6 
ALLIGATOR S8EAR 


Oe 
B 


Fic. 8—The square ground of Latogalga, or Fish Pond. 


A. Chiefs’ Bed: 1, miko (Raccoon); 2, heniha (or 2d chief) (Wind); 3, 
asimbonaia or yatika (head speaker, drawn from any clan); 4, hilis haya (any 
clan) ; 5, hilis hoboia. 

B. Citizens’ Bed (Tasikaialgi intupa) ; 6, yahaikas (no fixed clan). 

C. Warriors’ Bed: 7, tutka oktididja (Aktayatci). 


8, medicine pots; 9, boys’ medicine; 10, point where women assembled pre- 
paratory to the dance. 


grows in wet places in swamps and has red flowers, tcato hatkutci 
(four “little white rocks”’), and tutka-tcok-hissi (green moss from 
an old fireplace). 

If one has eaten new roasting ears he is given a little pasa root 
to chew and then a little pasa in cold water of which he must take 
four drinks. Then he can take all of the medicines like the others. 





no. 8 CREEK SQUARE GROUNDS—SWANTON 29 


Some of the medicine is swallowed, the rest spit out. The doctor sees 
that the medicines are taken and fines those who neglect to do so. 

There is a town policeman called istikona’ha who carries out the 
orders of the miko against those who have refused to obey him, and 
collects fines from them. The incumbent in 1929 was Maxcy Alakotci. 
When I interviewed them they were using the tastanagi as the 
town miko. 

The Hathagas are the Bear, Wind, Bird, and Alligator; the 
Tcilokis are the Raccoon, Aktayatci, Deer, and Potato. 

The Aktayatci are said to have formed one phratry with the 
Raccoon. 

In match games, they played against Tukabahchee, Atasi, Laplako, 
Eufaula, Hilibi, and Kealedji. Alabama was formerly of the same 
fire but later drew away. 

This year (1929) they did not use the pasa. The Asilanabi square 
ground is arranged just like this one, but the Okchai differ from these 
two a little in the use of their medicines. It is thought that Asilanabi 
is older than Latogalga and that the latter branched off in order to 
get the extra money that was paid to its six representatives in the 
national assembly. 

Except that there is more detail, the arrangement given here differs 
only slightly from that which I recorded in 1912 (42d Ann. Rep., 
Bur. Amer. Ethnol., p. 236). The only noteworthy divergence is 
in the position assigned to the Deer clan by my earlier informants, 
but this may be attributed to the fact that the miko apokta was then 
a Deer and his clan was probably brought over to the west cabin 
for that reason. As we should expect, the agreement is also close 
with the arrangement of the Asilanabi ground though there are 
minor divergencies in the allocation of clans to the south cabin. All 
of my authorities agreed well in assigning clans to the two moieties, 
but the oldest of all of them thought that the Beaver and Alligator 
were probably Tciloki. This may have been the ancient arrangement. 


TURABAHCHEER 


The plan of Tukabahchee square appears in ligure 9 and a view of 
it as it looked in 1912 is given in Plate 5, Figure 1. 

The number of tastanagis is indeterminate. The toba mawidine 
were officers not otherwise named who always remained in the cabins. 
There is one in each of the 12 beds except a part of the southeast 
cabin as indicated. They were selected from any clan, given names 
taken from the father’s clan, and seated with the latter. Thus, if 


30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 85 


a man were a Deer and his father a Raccoon, they would give him 
a name from the Raccoon clan’s names and seat him with them. 
After being so seated these officers were not obliged to do any further 
work connected with the busk. 


9 


“wy 
t 
~ 
Q 
< 
S 


-OL DTSITOM/LV 


SNY7I SIOIYA GA 
ASO SHAGLILNW 


Oo 


IKK KE XK 

a ae 

RACCOON, BEAR 
vax Se 

BIFfPD 5 


' 
' 
| 
' 





Fic. 9—Tukabahchee square ground. 


A. Chiefs’ Bed: 1, miko (Raccoon); 2, mikos (substitutes) (Raccoon) ; 
3, henthas (Wind); 4, henihas and yatikas; 5, hilis haya; 6, ta‘pala. 

B. Potato Bed; sometimes also called Youths’ Bed: 7, 7, 7, toba mawidine. 

C. Warriors’ Bed: 8, ta‘pala; 9, tastanagi takalgi. 

D. Youths’ Bed (Tcibanagalgi intupa): 10, yahaikas; 11, point where women 
assemble preparatory to the dance; 12 point where the women start dancing. 


The istatcagagis, most of whom sat in the southeast bed, were like 
a jury, or like a committee, to settle matters of routine. They com- 
prised both men and women. The admittance of women to this select 
group is something which I heard from no other informant. 





no. 8 CREEK SQUARE GROUNDS—-SWANTON 31 


A man was picked out of the bed of the henihas to speak for 
others, 7. ¢., the yatikas were selected from the henihas. None was 
used in the stomp dance. As payment for his services the yatika 
was given a deer hide and a ribbon. When he was wanted by the miko 
he was summoned four times, but he did not start until he had 
received the fourth call. He set a basket on the ground into which 
all of the tasikaias threw bits of tobacco to be taken to the singers 
for the women as payment for their services. 

The hotibonaia was a special speaker used in the ball game and 
in war. The term asimbonaia was about equivalent to that of yatika. 
He was a head man chosen from among the tastanagis for almost 
any purpose. He listened to any message which the miko wished to 
give out and then repeated it aloud. 

There were two hilis tcalaba who sat with their clans and were 
summoned whenever needed. One was a Raccoon and one a Bear. 

There were two oktididja. who sat with their clans or wherever 
they belonged until summoned to attend to the fire. 

Four hilis hoboia, selected from any clans, were sent to gather 
the medicine. 

All of the tasikaias shared in getting water. 

The famous Tukabahchee plates were taken out and cleaned four 
years in succession and then left covered for four years. 

The feather dance was discontinued at Tukabahchee when my 
informant was a small boy. 

The three leading women in the women’s dance carry atasa. The 
ieading woman has an eagle feather on her atasa, the second a white 
crane feather, and the other the feather of a third bird, perhaps 
a goose. They start one at a time and when the third moves all the 
rest follow. After they have walked round the fire four times they 
begin to dance and circle the fire again six times. 

The Hathagas were Bird, Wind, Bear, and Beaver; the Tcilokis 
were Turkey, Alligator, Raccoon, Deer, Panther, and any others. 

In olden times the moieties were exogamous. The Raccoon and 
Potato were then brothers, or rather half brothers, but intermarriage 
between them is now common. Even today, however, the Beaver 
and Bird will seldom intermarry. The Bird was the “uncle” of 
the Beaver and the Beaver the former’s “ nephew.” 

A man’s children called his father’s clan “ fathers.” One can say 
anything he wishes to, however disrespectful, about his own clan, 
but he must not speak against his father’s clan or permit anyone 
else to. 


32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


The general arrangement is the same as that obtained from the 
father of my informant in 1911-12 (42d Ann. Rep., Bur. Amer. 
Ethnol., p. 244), but the stations here assigned to some of the indi- 
vidual officers are probably more nearly correct, particularly the 
locations of the mikos. 


KEALEDJI 


Figure 10 shows the Kealedji square. 

The hoktagi immiko, “‘ women’s chief,’ supervised the women’s 
dance. There are now no regular water carriers. The imatas 
formerly sat in the north cabin, but now there are no officers so 
called. The beds are called by the names of the clans which occupy 
them. If the asimbonaia is wanted to deliver a speech, he is called 
over to the miko’s seat for that purpose. For tastanagis the best 
men are chosen. The ta‘palas are changed every four years. In 
former times they had a regular four days’ busk, but now it lasts 
for but one day and there is no feather dance. 

In the match games they played against Okchai, Wiogufki, and 
Tulsa. 

The Hathagas are Wind, Bear, Panther, and Turkey; the Tcilokis 
are Raccoon, Deer, and Aktayatci. In ancient times these moieties 
were exogamous. 

The names of the present officers are as follows: 


TIKOwese Ae na arene orate Kasihta Yahola (Raccoon) 
henthaaprorinc Sema eccr ts Heniha Imatutci (Wind) 
miilkcoOuapoktamermc cece Kayomulgi (Raccoon) 
hoktagi*immikor 2.22. cces =» sok Wiwohka Yahola (Raccoon) 
henihaicesaec een eine ere Oikas Hadjo (Wind) 
asin bonalay eee nee ee occ Kosa Fiksiko (Bear) 
taipalae ac PR ascent: one Talmutcas Hadjo (Bear) 

Se he | wc Ce PrN Ror RT Alak Hadjutci (Turkey) 
lnilbis: Wemleloey skoknodsecodoucuane Kan Tcati (Wind) 

See RY. antares el eae: Itco Imata Fiksiko (Deer) 
tutka oktididj¥ ........0000.0.004 Ahali Imata 
lapbicy 1oVO\OR), sceacassobcosaonnc Tatk6na Hadjo (Wind) 

- Tee Rave ere oe et Hotalgi Fiksiko (Wind) 
tAStANACIN ces cnka cree erties Tastanak Tcapko 

SN eA eet COM ORIN OO Lotete Tastanakutci 


“ 


(ces ete ieee eR Inheniha (= heniha) 
Sse EERE neha eeoeeioererte Tatkona Hadjo 


The plan of the revived square ground of the Kealedji agrees 
in all essentials with the one given me in 1912 from memory by a 
very old man, except that in recent times the Deer clan seems to 
have lost its importance or died out. It is interesting to find that old 


no. 8 CREEK SQUARE GROUNDS—SWANTON a2 
and recent informants agree in assigning the Panther clan to the 
White moiety. They differ, however, regarding the position of the 





RACCOON 





xX 
10 





AMTAYATC!' TURKEY 617 


Fic. 10.—The Kealedji square ground. 
A. Chiefs’ Bed: 


I, miko (Raccoon); 2, heniha (Wind); 3, miko apokta 
(Raccoon) ; 4, hoktagi immiko (Raccoon); 5, heniha (Wind); 6, hilis haya 
(usually Raccoon; Wind in 1929); 7, hilis tealaba (Deer and Wind) ; 8, hilis 
hoboia (Wind). 

B. Youths’ Bed: 9, ta‘pala (Turkey) ; 10, yahaikas. 

C. Warriors’ Bed. 

D. White Bed: 11, asimbonaia (Bear) ; 12, ta‘pala (Bear). 

13, medicine pots (Ist position); 14, medicine pots for those fasting (2d 
position) ; 15, medicine pot for boys (2d position) ; 16, medicine pot for girls 


(2d position) ; 17, medicine pot for women (2d position) ; 18, point where women 
assemble preparatory to the dance; I9, point where women start dancing; 
20, ball post. 


Turkey, which the earlier informant considered Tciloki and the later 
White. But this clan, together with the Alligator and Tami, is known 
to have been on one side in some towns and on the opposite in others. 





34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


EAPLAKO 


Figure 11 shows the plan of this ground. 
The two pots of medicine were prepared where they are shown 
(nos. 4 and 5) and remained there all of the time. The Bird clan sat 


N 
© 
4 


© 
J 








Fic. 11.—Square ground of Laptako. 


A. Chiefs’ Bed: 1, miko (Bear) ; 2, heniha or miko apokta (Bird) ; 3, yatika 
(sometimes seated here). 

ie Warriors’ Bed. This was usually occupied by middle aged men of various 
clans. 

C. Citizens’ Bed (Tasikaialgi intupa). Occupied by various classes, includ- 
ing persons not properly citizens of the town but taking medicine there, men 
married in the town or adopted into it, and men whose fathers belonged to it. 


4, a large pot of medicine for the fasters; 5, a small pot of medicine for the 
women and children; 6, rack for ball sticks. 
anywhere. The yatika belonged to no regular clan, and at the present 
time there is no such officer. There are two hilis tcalaba who occupy 
no special seats, and one hilis haya who is taken from any clan of,a 
friendly town. Towns belonging to the other town moiety do not 





no. 8 CREEK SQUARE GROUNDS—SWANTON 35 


take medicine with them and in fact are not invited except in the 
case of individuals married into the town. 

The Hathagas are the Bear (the most important of them), Bird, 
Beaver, and Wind; the Tcilokis are the Raccoon, Deer, Potato, 
Aktayatci, Panther, Alligator, etc. Here the Alligator, Turkey, and 
3eaver can intermarry; in Eufaula they could not. But probably this 
means that the Beaver could intermarry with the other two, as the 
Alligator and Turkey were seldom allowed to marry under any cir- 
cumstances. The Bird and Beaver were classed together in Laplako, 
but this group does not include the Bear or Wind. The Hathaga clans 
had the reputation of being progressive while the Tcilokis were full- 
bloods and reactionaries. This characterization probably followed the 
coming of the white people. 

The towns of the opposite group were called Talipota, which means 
“ foreign but not unfriendly.” 

The word Laptako indicates a place where there are many marshes 
filled with canes. This town square, which had been discontinued, 
was revived in the year 1903 in this way. They had to prepare a 
ground in order to take medicine before a game with the Nuyaka 
Indians next year. Later this was improved with regular cabins, but 
it must have been inconspicuous or have been considered unimpor- 
tant, as I heard nothing about it in 1912. 

Lapltako and Atasi are now nearly fused on account of the number 
of marriages between individuals belonging to them. Before the 
Civil War the Laplako had an Atasi Indian named Hotalgi Hadjo 
married among them as their hilis haya. One of the great men of 
Laptako in former times was Jim Boy (Tastanagi Imata) whom my 
informant remembers to have seen. He thinks he died just before 
the Civil War broke out. McKenney and Hall ( History of the Indian 
Tribes of North America, vol. 2, pp. 71-74) give a portrait of this 
chief and a considerable account of his life. He was born in what is 
now Alabama in 1793 and accompanied the warriors of his town 
during the Creek War of 1813-14, but was too young for active 
participation. In the war with the Seminole he was one of the leaders 
of the Creek contingent which aided the Americans. The exact date 
of his death seems to be unknown. 

The arrangement of the modern ground differs more from those 
described to me in 1912 (42d Ann. Rep., Bur. Amer. Ethnol., pp. 
254 and 255) than any of the others. The first of the latter was 
obtained from a very old man who should have known the ancient 
arrangement well, but of course my interpreter and I may have 
misunderstood him. The cabins are at different points of the compass 


30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


and more confusion is shown in the later organization, but they 
agree in stating that the miko belonged to the Bear clan and in 
placing that clan in the south cabin. My older informant allocated 
this clan with the Tciloki. This was probably an error on his part 
but he seemed to insist upon it. 

They have had no busk since the Civil War and no women’s dance, 
and the pasa is no longer used, only the miko hoyanidja. The dances 
in the square are three stomp dances. Seven days before one of 
these the “broken days” are sent out and on the day when the 
sixth stick is thrown away they are all to be at the ground, while 
on the seventh they are supposed to be taking medicine. Seven days 
before the dance they also meet and pick out four clean young 
men, called hilis hoboia, men whose wives are not pregnant, who 
are not given to intoxicants, and who have not attended to the 
digging of a grave during the preceding month. These men gather 
four bundles of medicine (miko hoyanidja) which they lay down 
with their tip ends toward the west. First they spread out a bed 
of leaves called lodja issi, “ turtle leaves,’ which should be taken 
from hickory trees. The medicine is laid on top of these and more 
leaves are spread over it. On the morning of the fast day all of 
those who are to take medicine are supposed to present themselves 
at the square ground. The fire is built up so that it will not go out 
all day. Early in the morning the two hilis tcalaba prepare the 
medicine, first the medicine for the women and children and then 
that for the adult men. This is taken four times during the day, 
the fourth time between one and two o’clock. Before they take the 
medicine the yatika announces, on behalf of the chief, that each of 
those intending to take it is to get a stick and throw it into a blanket. 
These sticks are counted and the yatika announces the number. Four 
times (or sometimes twice) during the day the men who are to 
take medicine with the exception of the officials (the mikalgi, yatika, 
hilis haya, and the two ta‘pala) go out to get firewood so as to be 
ready for the dance that night. Four men are selected to keep watch 
of the fasters during the night, to see that no one sleeps or breaks 
his fast, or drinks or goes with a woman. That is why the numbers 
are taken. These four men are called istikona’ha, “men taking 
away,” because they take away the hat of anyone found sleeping 
(and treat similarly anyone who breaks the taboos in other ways). 
The hat they carry to the miko and, when the owner comes to get it, 
he is fined. 

After they have taken medicine for the fourth time, they go to 
the creek for a bath and then return to the square. The yatika 





no. & CREEK SQUARE GROUNDS—SWANTON . 37 
talks to them telling them to take care of themselves all of that 
night, and then they scatter to the camps to eat. Before sundown, 
however, they are supposed to be back on the ground. ‘The yatika 
calls to them four times and by the fourth time they are to be in 
their seats. When all are in their places, the two ta‘pala are selected. 
The yatika makes a speech on behalf of the miko, calling upon his 
hearers not to act in an unfriendly way toward the outside friends 
who are about to be admitted to the dance, not to take liquor, to 
behave themselves all through the night, not to fight, and so on. 
This speech is addressed to the town people and outsiders alike. 
After it the fasters dance four times and then the visitors are 
admitted. That “kills the fast.” All through the day the square is 
to be kept clean with the idea that the fasters will in consequence 
be clean. They go up to take the medicine two by two, and those 
who are ceremonially unclean take the medicine last. They dance 
until about daybreak. Then all belonging to the town go down to 
the branch and bathe, after which they return and sit in their 
respective beds. Then a collection may be taken to defray their 
expenses and they settle other matters. The night before, the chief 
of a friendly town may have announced a dance, and, if so, the 
announcement is now made and advice given as to how they are 
to help their friends. This advice is uttered by the yatika, speaking 
for the miko, and he then gives a general talk, advising his people 
not to use liquor, not to break the laws, and to be good citizens in 
every sense of the word. Then they disband for the year. 


HILIBI 


A plan of Hilibi square is shown in Figure 12 and a general view 
of the ground as it appeared in the winter of Ig11-12 in Plate 5, 
Figure 2. 

Two poles with white feathers attached were at each front post. 
These were used in the “ feather dance ” and were called ‘ the path,” 
because the path was to be white. 

When there are visitors the owners of the east cabin move 
elsewhere in order to make room for them. When the south cabin 
is overcrowded, some of its occupants move into the east cabin. In 
this town the clans were always considerably mixed up in the beds. 
The ta‘palas can sit anywhere. The tutka oktididja does not have 
a particular seat on account of his official position. He is appointed 
every four years. The tutka didja, who starts the fire, is identical 
with the hilis haya. 


38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL. 85 


If a man whose father is of the Aktayatci, the miko’s clan, is given 
a busk name, he is brought to the mikos’ bed and given a seat there. 

The Hathagas are the Bear, Bird, Alligator, Beaver, Turkey, and 
Wind; the Tcilokis are the Panther, Potato, Deer, Aktayatci, and 
evidently the Raccoon. The Alligator and Bird are considered as 
practically one clan. 

The two leaders in the women’s dance carry atasa. 


drs 





Fic. 12.—Square ground of Hilibi. 


A. Chiefs’ Bed: 1, miko (Aktayatci); 2, heniha (Alligator); 3, hilis haya 
(Raccoon in 1929); 4, ta‘pala (Bear); 5, hilis tcalaba (Bear in 1929). 

B. Raccoons’ Bed: 6, hilis hoboia; 7, tutka oktididja (Raccoon in 1929) ; 
8, yahaikas (any clan). 

C. Cabin used by visitors: 9, ta‘pala (Alligator) ; 10, hilis hoboia (Alligator). 


II, medicine pots; 12, medicine pot for boys; 13, medicine pot for women; 
14, woodpile; 15, ball post. 


The history of this town is as follows. It was founded by a man 
of the Aktayatci clan who went off to live by himself and then put 
up a ball post. Many women belonged to his family and he had 
numerous visitors, some of whom married these women so that it 
soon grew into a large band of which he was probably the first miko. 
Anyhow that is the way the Aktayatci came to have the town. After 
stomp dances had been held there for some time, more visitors came 








no. 8 CREEK SQUARE GROUNDS—SWANTON 39 


to join them and it grew still larger. Because it was built up very 
rapidly, its founder called it Hilibi, which means “ hurry ” (hila’pkis, 
‘“T make haste”). Since it was an “ illegal” band, a talwa fatcasigo 
(“town deviating from correctness’”’), all the clans do not have 
regular places, having been drawn from so many other bands. 

The following notes on some personal names contained in the town 
roster give an interesting insight into the manner in which totemic 
names were bestowed : 

Fos Hatki Imata (“ White Bird Imata’’), so named because he 
belonged to a White clan, the Wind. 

Halak Hopaie. He belonged to the Bear clan, but his father 
belonged to the Raccoon, hence the name Halak or Ahalak (“ Po- 
tato’’), since both these clans are Tciloki. 

Pahos Fiksiko. He belonged to the Wind clan, but his father was 
a Deer and the Pahosa is of the same phratry as the Deer. 

As shown by the native story above given this town was not 
supposed to be ancient or to have a firmly fixed town organization, 
and, while it was older than the Hilibi people themselves believed, 
it seems to have preserved the irregularities which might naturally 
be associated with a new town. Not improbably the tradition of 
irregularity preserved the fact. At the same time there is a general 
agreement between the plan here given and that which I obtained in 
Toi2 (42d Ann: Reép., Bur. Amer. Ethnol., p» 258): The mikos’ 
cabin is to the west, and the mikalki and henihalgi were of the same 
clans, Aktayatci and Alligator respectively. The Alligator and Tur- 
key were classed as Whites from association with the Bird by 
my later informants, but the earlier ones gave them as Tciloki. 


ALABAMA 

This was a very simple square of exceptional arrangement as shown 
in Figure 13. 

The Alabama were one of the incorporated tribes with a language 
distinct from Creek. The clan names in Alabama are: Mahaleha 
(Wind), Sawaha (Raccoon), Aktayatciha (Aktayatci), Hatcuntco- 
baha (Alligator), Konoha (Skunk), Nitaha (Bear), Kotha 
(Panther), Fociha (Bird), Fitoha (Turkey), Ofataha (Beaver). 

All of the officers were brought over to the chiefs’ bed. Men of 
the Alligator, Bear, and Aktayatci have been mikos and a man of 
the Skunk clan was once the heniha. There is no regular rule for 
either position. 

The Hathagas were the Wind, Bear, Panther, Skunk, and Raccoon ; 
the Tcilokis were the Bird, Beaver, Turkey, Alligator, and Aktayatci. 
The Wildcat was the same as the Panther. 


4O SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


One of the greatest Alabama Indians now remembered was 
Kantcati Yahola (Alabama name Tcisoki), who was the hilis haya. 
He was born in Alabama and came west with his tribe. He lived 
until about 1866. 

The above plan and the three others I obtained in 1912 and the 
years immediately following (42d Ann. Rep., Bur. Amer. Ethnol., 
pp. 263-264) show considerable minor variations but all agree in 
locating the chiefs’ bed in the east and the warriors’ bed in the west. 
Most of them also place the Bear and Panther clans in the latter 
and the Wind, Aktayatci, and Deer in the former, where they are 
noted at all. In allocating the clans the above informant agrees with 


Oo 
PANTHER 


BIRD, 


TORREY 
AND 


BLAVER 


[THE WORTH ii 
END OFRIGI- 

NALLY BELON 
GEO TO THE 
BEAR CLAN) 





oO 





B 


Fic. 13.—Alabama square ground. 


A. Chiefs’ Bed: 1, miko (Wind); 2, heniha (Raccoon). 
B. Warriors’ Bed. 


the older ones except regarding the Panther which the men first 
consulted asserted was Tciloki while it is here given as a White clan, 
but this is a clan which has been placed on both sides. 


KASIHTA 


For the plan of Kashita square ground, see Figure 14. 

There should be two hilis tcalaba, drawn from the Alligator clan, 
but they are not employed now. Four hilis hoboia for the pasa, four 
for the miko hoyanidja, and one tutka oktididja are chosen by the 
miko without reference to clan. There is no definite body of water 
carriers. Two ta‘palas are selected from any clan to serve just for 
the night. They carry sticks called reels, and their function is to 
invite the dance leaders to lead dances and see that all take part. 
There is only one singer for the women’s dance. He sits behind 
the miko. 


no. 8 CREEK SQUARE GROUNDS—-SWANTON Al 


In rgor the old square ground was given up and, the new one was 
established in June, 1920. Because the new generation was weak 
one cabin was cut out. The Coweta square is said to have been the 


same as that of Kasihta. 
Every time anything is brought in or anything repaired they dance 


ll night because thus the two things are joined together, just as 


0 — SS? oO 
TASTANAGIS RACCOON, 
Ano AMTAYATC! 
MIALAS AND PANTHER 
{= 1 
Cc 5 


SEAR 
a TREES 


o ‘ 
‘ 


Bien f POTATO, s 


‘ 
‘ 
t 
t 
' ‘ 
' 
' 
' 
; i 
266 
3 
: iW 
E LLIGATOR )) 
\ St * ‘ 
: F | 
; t 
' 
‘ 
' 
0 
' 
‘ 
' tf WIND ANS B/FAD BEAVER 4° DEER 5 





12 
® 


Fic. 14.—Kasihta square ground. 
A. Chiefs’ Bed: 1, miko (Bear); 2, heniha (Alligator); 3, hilis haya (seat 
Ww Fa on duty) ; 4, asimbonaia. 
Whites’ Bed (Henihalgi intupa), or “ Cabin of the Greater Kings.” 

c Warriors’ Bed: 5, asimbonaia (Aktayatci). 

6, medicine pot; 7, point where women assemble before dancing; 8, trash 
heap; 9, where ashes of old fires were placed; 10, split log where are seated 
those young men who have broken the rules and are in consequence placed 
here as a punishment before being allowed to mix with the rest of the people; 
11, line of sweepings (tadjo) marking limit of square ground; 12, ball post. 
two days are united by the night on which they dance. Formerly 
dances were held every Saturday night, but hard times have put an 


end to them. The dance which was being held when I paid my visit 


was because they were then putting up a ball post. 
The Hathagas are the Wind, Beaver, Bird, and Deer; the Tcilokis 
are the remaining clans, yet it is said that the Bear, Alligator, and 





Mae 


42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 
Wind were of the same “ class.” 
marry outside of third cousins. 

The Kasihta defeated the Coweta three times in the ball game, 
the last time in 1878, and after that they took them under their 
jurisdiction and they have played on the same side. 

Comparison of the plan of the new Kasihta with the plans of the 
earlier grounds secured by Gatschet and myself (42d Ann. Rep., Bur. 
Amer. Ethnol., pp. 266-268) shows that the old order has been fairly 
well maintained allowing for the disappearance of some clans, 
particularly the Fish clan. Incidentally I wish to correct an erroneous 
statement in my report in which I misquoted Gatschet to the effect 
that the Kasihta miko belonged to the Alligator clan. While the 
Alligator clan occupied half of the chiefs’ cabin, the miko himself 
has always been taken from the Bear and was stated to have been 
so by Gatschet. The allocation of clans to the moieties by my recent 
informants contains a number of difficulties, for they seem to have 
placed the Bear among the Tciloki, which is unlikely, especially as 
it was said to belong to the same “class ” as the Alligator and Wind. 
My own earlier authorities also classed the Beaver and Deer as 
Tciloki. 

The following information will be interesting to those who wish 
to study acculturation processes in intangibles. It was told me by 
the Indian considered best able to speak for the town. 


Under the present law they can all 


The original four cabins represented the New Jerusalem with its 12 gates. 
The busk goes back to the time when Jacob set up the altar at Bethel and is 
traced from him and his 12 sons. All of the Indians in America entered in two 
migrations, one at the time of Jacob (1500-2000 B. C.) and the second 600 years 
later, at the time when Jerusalem was destroyed by Nebuchadnezzar. Then 
they talked face to face with the Great Spirit because they were more obedient 
to God than any other tribe, but about 700 years after the Messiah they got 
away from the original law on account of desire for riches. Then they lost the 
old law and asked for a new government, and by holding a ceremony in mid- 
summer, in the month of July, it was given to them. That new law taught 
them to tell the truth and be honest with their fellowmen and to raise their chil- 
dren in such obedience, not to touch anything that did not belong to them, not 
to make a false statement. That is the law which we are trying to follow. 


He added: 


We have a hard time because the white men have failed to fulfill their part 
of the agreement. They have strong laws that we can’t begin to understand and 
our customs are about choked out through grafters who claim to have bought the 
claims of the allottees on which our squares are located. In order to hold their 
grounds several towns have to pay rent year after year. If the law makers 
would cooperate with us and give us full privileges we would raise more sub- 
stantial, law-abiding young men and young women. That was the custom and 
the wish of our forefathers. 





no. 8 CREEK SQUARE GROUNDS—SWANTON 43 


OCHESEE SEMINOLE 


This is the only Seminole ground from which I obtained informa- 
tion during my recent trip. The plan of it is shown in Figure 15. 

There are two pots of medicine for the miko hoyanidja and pasa 
respectively, but the latter is introduced only at the busk. The women, 


N 


© 
7 


DEER, PANTHER, 
POTATO ALLIGATOR 


(/NCLUDING TURKEY) 





ee oe Ne Ss 
” . Gre 
ve ‘ q = 
‘ ‘ S Q\ 
° aa | Cee 
\ ‘ i aw 
i : Ah SN 
eee a7 x a & 
- k ~ 
~ aK 
\ ‘ 
ae \ 














AKTAYATC/, 
WIND, SNAKE 

AND HAPITCA 
(2) 


B 


Fic. 15.—Ochesee Seminole square ground. 
A. Chiefs’ Bed: 1, miko (Bear) ; 2, heniha (Bird) ; 3, yatika (Deer in 1929) ; 
4, hilis haya (Wind in 1929). 
B. Warriors’ Bed: 5, yahaikas. 
C. Warriors’ Bed. 
D. Youths’ Bed. 


6, medicine pots; 7, medicine pot for boys (containing red root) ; 8, point where 
women gather before dancing; 9, point where women start to dance; 10, ashes 
of old fires; 11, ball post. 





however, use wormseed (“ wilana”’). After they are through with the 
medicines, whatever is left is poured out at the place where they have 
been taken. 

The miko is chosen if possible from the Bear clan, but if they do 
not find a suitable man of that clan, they select someone from the 
Beaver clan. 


44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL.. 85 


The yatika and hilis haya may be selected from any clan, the former 
being chosen for his oratorical gifts. 

There is now no heniha. The last they had went to live with the 
Creeks and did not come back. 

Two ta‘palas are used in the women’s dance and hold their position 
for four years. Other ta‘palas are chosen temporarily for the other 
dances. In the stomp dances they change these ta‘palas several times 
during the night. 

There are four hilis hoboia taken from the west, north, and south 
cabins but from any clan. They keep their positions as long as they 
choose to serve. 

There is one tutka didja who can be of any clan. 

Five or six boys bring water to the ground. 

The two leaders among the women carry atasa which are painted 
with white clay annually just before the dance. Women do not take 
the wilana internally; they merely wet their faces and other parts 
of their bodies with it. The boys use only the miko hoyanidja. 

The ashes of the old fire are removed from the square and the 
new fire lighted on the morning of the fast day. 

The Hathagas are the Bear, Bird, Beaver, Wind, Otter and Skunk; 
the rest are Tcilokis. The Bird and Beaver belonged in one phratry, 
and so did the Wind, Otter and Skunk; the Alligator and Turkey ; 
and the Aktayatci, Kapitca, and Snake. 

The name of the present miko is Nokos Miko, and his father 
belonged to the Deer clan. The last heniha was named Henitha Miko, 
and his father was of the Bear clan. The busk name of the yatika 
(my informant) is Pahosa Tastanagi; his father belonged to the 
Aktayatci, and his father’s father to the Bear. 

They have a ball post surmounted by a wooden fish. A hit on this 
fish counts 4, and on the post above a certain mark 2. There have 
been no regular match games between towns in the lifetime of my 
informant, but about two years ago the old men and the young men 
played against each other. 

The above plan of the Ochesee ground agrees closely, naturally 
enough, with that which I obtained from the man whose advice was 
particularly resorted to in reestablishing it some years ago (42d Ann. 
Rep., Bur. Amer. Ethnol., p. 283), for in 1912 it had been given up. 
The main differences are in the seating of the Potato and Alligator 
clans. In the matter of the moieties the only change is in the case 
of the Raccoon clan, which I previously set down as White. This is so 
exceptional, however, that I have always believed that I must have 
misunderstood my informant. 





No. 8 CREEK SQUARE GROUNDS—SWANTON 4 


wm 


GENEALOGIES 


The two brief genealogies which follow will illustrate in some 
measure the influence of the clan system on marriage. The first is 
the genealogy of Jeff Canard of Laplako; the second that of Earnest 
Gouge of Hanna, the latter of especial interest because it includes 
the famous orator Hobohit Yahola. 











paternal paternal maternal maternal 
grandfather grandmother grandfather grandmother 
(Bear) (Alligator ) (Wind) (Beaver ) 
father mother 
(Alligator ) (Beaver ) 
a 
self 
(Beaver) 
paternal 
grandfather maternal 
(this was paternal grandfather maternal 
Hobohit Yahola) grandmother (Potato of grandmother 
(Potato) (Raccoon) Abihka) (Bird) 
a 
father’s = father mother 
second (Raccoon) (Bird of Otciapofa ) 
wife __— = = x - 
(Raccoon?) self wife’s wife’s 
(Bird of father mother 
Otciapofa) (Bear) (Alligator ) 
ee 
wife 
(Alligator) 
CONCLUSION 


The busk undoubtedly represents a long period of development, 
and as the stages through which it passed and the elements entering 
into it have been lost, we can never understand it fully. Its main 
purpose, however, was evident. It was to restore the connections 
of the tribe with the universe which a year of civil or profane living 
had tended to rupture. Hence the new fire, extracted from its 
abiding place in the wood and not as yet sullied by contact with 
humanity. Hence the rigorous fast accompanied by administration 
of medicines divinely revealed to the ancestors, the general pardon 
of offenders, the sabbath calm prescribed for that period of 
regeneration, the use of white paint, and the employment of the 
term white—‘ the white day,” “the white smoke,” “the white 
drink ”—in various parts of the ritual. The one discordant note 
seems to have been provided by the women’s dance, since the leaders 


46 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


of that dance carried representations of war clubs, which there is 
every reason to believe were anciently adorned with scalps, and some 
of these were painted red. But I suspect that this dance was an 
attempt to represent war as a protective institution and to thank the 
being or beings who preside over human destiny for having so well 
defended them against assaults of the—as usual—perfidious foe. 
Possibly some element of propitiation also entered into this dance. 

The universe with which the Creeks sought reconciliation was not, 
however, a material one. What they had in mind was rather the 
mind or minds believed to be operant there. While we know of some 
supernatural beings connected primarily with the busk and numerous 
spirits associated with natural objects were anciently believed in, 
it seems fairly certain that the peculiar patron of the ceremony was 
a solar, or rather celestial, being generally called Hisagita-immisi, 
“the breath controller,’ and also Ibofanga, “the one above,” and 
that the busk fire was in some way an earthly representation of the 
great solar fire overhead. While it is probable that Hisagita-immisi was 
not in ancient times the monotheistic deity he has now become, there 
is every reason to think he was already, before White contact, the 
supreme being of the Creeks. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL... 85, NO. 8; PEs 1 





1. Abihka or Talladega square ground. 





2. Tulsa square ground from the southeast. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL.|85, NO=8, Pie 





1. Nuyaka square ground. 





2. The three beds of Pakan Tallahassee, from the southwest. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85, (NO.-8; PL. 3 





1. Pakan Tallahassee square ground looking between west and south beds, show- 
ing chunkey yard and ball post. 





2. Wiogutki square ground from the southwest. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85, NO. 8, PL. 4 





1. Okchai square ground in winter from the southwest. 





2. Fish Pond square ground. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLE. 85; NO. 8. -PE. 5 





1. Tukabahchee square ground. 





— 


2. Hilibi square ground in winter from the southwest. 

















SMITHSONIAN MISCELLANEOUS COLLECTIONS 


VOLUME 85, NUMBER 9 


Roebling JFrund 


THE DETERMINATION OF OZONE 
BY SPECTROBOLOMETRIC 
MEASUREMENTS 


(WiTH THREE PLATES) 


BY 
OLIVER R. WULF 
Smithsonian Institution and U. S. Bureau of Chemistry and Soils 





(PUBLICATION 3127) 


CITY OF WASHINGTON 
PUBLISHED BY THE SMITHSONIAN INSTITUTION 
NOVEMBER 30, 1931 





Tbe Lord Waltimore Dress 


BALTIMORE, MD., U. S. A. 





Roebling Fund 


THE DETERMINATION OF OZONE BY SPECTROBOLO- 
METRIC MEASUREMENTS 


By OLIVER R. WULF 
SMITHSONIAN INSTITUTION AND U. S. BUREAU OF CHEMISTRY AND SOILS 


(With THREE PLateEs) 


In the fall of 1930, at the suggestion of Dr. C. G. Abbot, measure- 
ments of the transmission of visible light by ozone were made on the 
solar spectrometer of the Smithsonian Institution at Table Mountain, 
Calif. This determination of the absorption of ozone in the region 
of its very weak absorption, practical in the laboratory only by the 
method of photographic photometry, by which it has been done by 
Colange,’ can be accomplished by direct energy measurements on the 
spectrobolometer because of its great sensitiveness and the extreme 
intensity of the source. At the same time fluctuations in weather 
conditions are likewise registered with great sensitiveness and consti- 
tute a serious source of error in the measurements. But on the other 
hand it is favorable that observations may be made of the absorption 
of controlled amounts of ozone placed in the sun’s beam with all other 
conditions of operation identical with those of the regular solar 
observations. One of the useful results of this work has been the 
selecting of a large number of points on the solar bologram which may 
be satisfactorily used for ozone determination. The present paper has 
to do for the most part with the results of these measurements as 
forming a basis for the determination of atmospheric ozone from 
solar bolograms and not with the results of their application. 

The essential apparatus used in this work auxiliary to the spec- 
trometer is shown in Plate 1. A cylindrical glass cell, 20.0 cm. long 
and approximately 13 cm. in diameter, was mounted in the front 
room of the spectrometer tunnel in such a way that it could be easily 
moved in or out of the solar beam as it came from the coelostat mir- 
rors on its way to the first slit of the instrument. The circular windows 
of this cell were made of high-grade plane plate glass. These windows 


*Colange, G., Journ. Phys. et le Rad., ser. 6, vol. 8, p. 254, 1927. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 85, No. 9 


2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


of the cell were covered with diaphragms having vertical rectangular 
openings sufficient to pass a beam that considerably more than covered 
the first slit of the spectrometer. The absorption cell was sealed with 
all glass seals in connection on its inlet side with the ozonizers and 
on its outlet side with an analytical apparatus for ozone. No altera- 
tions of any sort were made in the regular observing conditions except 
for the interposition of the cell. Tank oxygen was supplied to the 
ozonizers under a small constant pressure through the capillary of a 
glass flowmeter having nujol as the manometric liquid, this flow- 
meter being calibrated in a series of independent measurements. The 
exit ozone was analyzed by the method described by Wulf and Tol- 
man. The samples, whose volumes were known from the time and 
rate of gas flow, were collected over potassium iodide solution, set 
aside, and subsequently analyzed. It is evident that, in the filling of 
such a cell by sweeping at the low rates of gas flow necessary in such 
a system for producing ozone of relatively high concentration, a con- 
siderable amount of time will elapse before the exit gas attains practi- 
cally the concentration of the entering gas. To study this circumstance 
a tube was also brought to the analytical apparatus from before the 
cell as the gas came from the ozonizers allowing a sample of the inlet 
gas to be taken, and by means of this it was possible to determine the 
time sufficient for the exit gas to rise nearly to the concentration of 
the inlet gas. The concentration of the exit gas was taken, in view 
of the processes of diffusion and mixing going on in the cell, as 
representative of the ozone concentration in the cell. The ozonizers 
were of the familiar silent discharge type. For part of the work one 
alone was used, while for the rest of the work two were used in series 
and both water-cooled, these giving the largest concentrations em- 
ployed. Ordinarily over the period of the taking of four ozone bolo- 
grams six analytical samples of the ozone were collected. 

Knowledge gained from earlier work on ozone permits a descrip- 
tion of the character and position of the absorption.” A small fraction 


*Wulf, Oliver R., and Tolman, Richard C., Journ. Amer. Chem. Soc., vol. 40, 
p. 1650, 1927. 

“a. Colange, G., Journ. Phys. et le Rad., ser. 6, vol. 8, p. 254, 1927. 

b. Wulf, Oliver R., Proc. Nat. Acad. Sci., vol. 16, p. 507, 1930. 

c. Ladenburg, Erich, and Lehmann, Erich, Ann. Phys., ser. 4, vol. 21, p. 305, 
1906; Verh. Deutsch. Phys. Ges., vol. 8, p. 125, 1906. 

d. Schoene, E., Journ. Russ. Phys.-Chem. Soc., vol. 16, pt. 9, p. 250, 1884; 
Journ. Chem. Soc., vol 48, pt. 2, abstracts, p. 713, 1885; Chem. News, vol. 69, 
p. 289, 18094. 

e. Chappuis, J., Ann. l’Ecole Norm. Sup., ser. 2, vol. 11, p. 137, 1882; Compt. 
Rend., vol. 91, p. 085, 1880; Compt. Rend., vol. 94, p. 858, 1882. 





NO. Q DETERMINATION OF OZONE—WULF 3 


of the solar energy will be cut out by the ozone as is illustrated in 
Figure 1, which represents a normal spectral energy curve roughly 
similar to that of the sun showing the approximate area removed by 
the atmospheric ozone in the visible. 

However, the actual observing of this reduction of intensity must 
be made on the complicated solar curve as shown in Plate 2, which is 
composed of a series of typical bolograms notched by many Fraun- 
hofer absorption lines. The discontinuities in the curves are due to 
the insertion of rotating sectors in the path of the solar beam to cut 
down the intensity in the regions of great intensity, to values such 
that the galvanometer deflections will still fall upon the photographic 


A 


Fic. 1.—Spectral energy curve of black radiator, approximating that of 
the sun, showing the atmospheric ozone absorption in the visible. 


plate on which these deflections are being continuously recorded in 
the form of these curves. In addition, at two points a shutter is 
inserted for the purpose of determining the base line, that is, the line 
of zero deflection. Owing to the scattering of the rays by the earth’s 
atmosphere the apparent maximum of the sun’s intensity is shifted 
to longer wave lengths. The strong atmospheric absorption in the 
deep red is conspicuous, while over the region of ozone absorption 
chiefly the Fraunhofer lines are in evidence. 

In spite of these complexities, however, comparison of two bolo- 
grams, one without ozone in the cell and one with ozone, should show 
the reduction in intensity caused by the ozone, providing weather 
conditions remained sufficiently constant between the two. If, for 
example, we were to take the ratio of ordinates at corresponding 
points on two such bolograms under ideal conditions this ratio should 


4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


be unity in regions not influenced by ozone, and somewhat greater 
than this where the intensity was reduced by the ozone absorption. 
In view of the exponential decrease in intensity of light passing 
through an absorbing medium, it is the logarithm of the ratio of 
ordinates which is proportional to the amount of ozone in the path. 
This will be zero evidently for those cases where the ratio is unity, 
and positive or negative as the ratio is greater or less than unity. This 
proportionality is based on the assumption that Beer’s law may be 
used here. There appear to be no causes concerned with the apparatus 
and structure of the spectrum which could bring apparent false devia- 
tions from this law, since the spectrum is composed of broad diffuse 
bands,’ or perhaps more accurately fluctuations in the absorption co- 
efficient without any apparent discontinuities. The question as to 
whether the ozone absorption actually obeys Beer’s law is an impor- 
tant one and, so far as the author is aware, has not as yet been 
satisfactorily answered, although attention has been called to it by 
Ladenburg.” However, this assumption is contained in all previous 
determinations of ozone such as we are employing here and must be 
similarly contained in the present work. Actually, instead of a single 
bologram in these measurements, four consecutive bolograms were 
taken with the cell containing no ozone and four more with the cell 
containing ozone. The average values of the ordinates at correspond- 
ing points were compared. 

A difficulty enters, however, because of the time which elapses 
between the taking of the ozone bolograms and the uninfluenced ones 
which we will call the oxygen bolograms, since a very appreciable 
change in air mass, or amount of atmosphere traversed, occurs be- 
tween the two as well as possible weather changes. This was unavoid- 
able with the apparatus at our disposal at the top of Table Mountain 
because of the nature of the ozone technique, and is not easy to avoid 
under any circumstances. Under these conditions, the ratio of 
ordinates outside of ozone absorption would be far from unity and 
the logarithm far from zero. Standard correction for air mass was, 
therefore, applied to the average ozone ordinates to bring them to the 
air mass of the oxygen observations. It is hardly possible, however, 
to make correction for all differences between the two sets with 
accuracy sufficient to bring the logarithm of the ratio of ordinates to 
zero within a quantity small compared to the small ozone effect, and 
it is not necessary because it is a difference in the logarithm for 
different wave lengths that is of interest. 


* See footnote 2) on page 2. 
*Ladenburg, R., Gerlands Beitr. Geophys., vol. 24, p. 40, 1920. 


NO. 9 DETERMINATION OF OZONE—WULF 5 


The points shown on the bolograms were chosen as far as possible 
with respect to the known position of the ozone bands. These bands 
are shown in Plate 3 as they were obtained in the work described in 
footnote 2b, on page 2. The attempt was made to find points on 





1000 5000 6000 7000 8000 9000 


Fic. 2—Diagrammatic example of ozone area. 


the solar bologram suitable for measurement which lay in every ozone 
absorption maximum and minimum. 

From the above considerations, if a plot of the logarithm of the 
ratio of ordinates is made against wave length, the type of result to 
be expected is something of the form shown in Figure 2. The failure 
to make accurate correction for changes in atmospheric transparency 
or sensitiveness of apparatus will result in the points outside of ozone 


6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


absorption being something greater or less than zero, here illustrated 
as less than zero, but this will not obscure the increased value of the 
ratio in the region of ozone absorption showing as an increase above 














4000 5000 6000 7000 8000 3000 


Fic. 3.—Typical ozone area. 


the smooth curve passing through the values of the points on both 
sides of the absorption. It is the area under this curve above the 
smooth curve which is closely proportional to the amount of ozone 
in the path. 

Figure 3 shows the results of one typical day’s observations. For 
clearness of illustration Figure 2 was made similar to this. In this case 


NO: 9 DETERMINATION OF OZONE—WULF Wi 


an amount of ozone approximately equal to the average atmospheric 
quantity was present in the cell. Failure to correct exactly for all 
differences between the two sets of bolograms led to the smooth curve 
of the points in the absence of ozone absorption being below zero, but 
the area due to ozone appears clearly and can be measured with 
sufficient accuracy to be of much use. Several days’ determinations 
were made with larger quantities of ozone in the path, giving areas 
which can be measured with greater percentage accuracy. On other 
days smaller amounts were used in order to observe how well such 
areas could be determined. Included in figure 3 are also three points 
lying far off the curve which are illustrative of unsuitable points. 
They lie in weak atmospheric water absorption in the red, which ab- 
sorption may vary considerably over short intervals of time, rendering 
the points evidently unsuitable for aiding in ozone determination. 

The area under the observed curve should evidently be limited 
between ordinates whose values are still large compared with the un- 
certainty in placing the base line. In the blue this has been taken as 
the value at 4750 A, while in the red it was necessary to terminate 
the area at 6135 A because of the uncertainty in the point at 6335 A. 
This limitation was caused by instrumental circumstances which can 
be altered in the future to include a greater area. 

These areas were determined for 10 independent sets of observa- 
tions, the amount of ozone in the cell being known in each case from 
the analytical work carried on at the time of the measurements. From 
these results, shown graphically in Figure 4, a value sufficiently accu- 
rate to be useful can be had for the amount of area per unit path 
length of pure ozone at o°C and one atmosphere pressure, the 
common meteorological form of expressing atmospheric ozone. The 
least-squares solution, assuming the ozone concentration values essen- 
tially free from error compared to values for the areas, which method 
automatically weights the individual values in proportion to the area, 
leads to the result 21.5 sq. cm. of area per mm. of ozone at standard 
conditions of temperature and pressure when a plot is made to the 
scale ordinates 0.001 per cm., abscissa 200 A per cm. This area can 
thus be stated as 4.30 A independent of the scale to which it is plotted. 
Plotting the data and planimetering the area is a procedure which has 
the decided advantage of giving a visual record of the amount of 
ozone which can be judged approximately at a glance. From the 
results of Colange’s data on the absorption coefficient of ozone one 
can compute this same area, and it may be estimated directly from his 
published curve of the absorption coefficient. One finds thereby that 
the value obtained in the present work is about 4 per cent higher than 


8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


that given by Colange’s results, a difference which lies within the 
limits of error of the present work. With a larger number of sets of 
observations than the 10 of the present work, and particularly at high 
ozone values, the accuracy of the determination could, of course, be 
greatly increased. 


AREA 





/ 2 3 4 5 6 7 8 
PATH mm 0, 


Fic. 4.—Character of the results of the absorption of ozone. 


Anarea such as that illustrated in Figure 3, and pertaining to atmos- 
pheric ozone only, can be obtained from atmospheric transmission 
coefficients, if combined with the knowledge that there is no important 
atmospheric absorption except ozone across this spectral region. That 
atmospheric transmission coefficients show unmistakably the ozone 
absorption has been pointed out and used by Fowle and others.’ 


*Fowle, F. E., Smithsonian Misc. Coll., vol. 81, no. 11, pp. 1-27, 1929. Caban- 
nes, J., and Dufay, J., Journ. Phys. et le Rad., ser. 6, vol. 7, p. 257, 1026. 


NO. 9 DETERMINATION OF O0ZONE—WULF 9 
Figure 5 shows an area analogous to the one shown earlier, deter- 


mined from the transmission coefficients for March 24, 1929, as a 
typical day used simply as an illustration. The logarithms of the 


0.025 





0.0/0 


0.005 





° 





0 ; 7 
° 


co 4 
| 








4000 5000 6000 7000 8000 5000 
Fic. 5.—Ozone area for March 24, 1920. 


transmission coefficients for this day, read for all the wave lengths 
selected in this work, were plotted against wave length and a smooth 
curve passed through the values lying to both sides of the ozone 
absorption, and the differences of the points lying in the ozone region 
from the smooth curve were read. These differences were then 
plotted against wave length to the scale used above, yielding the area 


IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


shown in Figure 5. Thus the ordinates are the logarithms of the ratio 
of the intensities before and after passing through the atmospheric 
ozone. Referring this area so obtained to the results of these present 
measurements pictured in Figure 4 affords a method of determining 
the atmospheric ozone based on direct intensity measurements, and 
yields for this day over Table Mountain, Calif., an ozone value of 
3.8 mm., while the value given by Dobson for Table Mountain on this 
same day was 3.42 mm. A somewhat more satisfactory extension of 
this method of determining the ozone transmission may be employed, 
whereby the curved base is reduced to a straight line. Instead of 
plotting the logarithm of the atmospheric transmission coefficients 
against wave length and determining the difference of the observed 
points from the smooth curve in the ozone region, the logarithm of 
the logarithm of the transmission coefficients may be plotted against 
the logarithm of the wave length, yielding very closely a straight line, 
except for points in the ozone region. This fortunate circumstance 
is due to the approximate inverse A" dependence of the logarithm of 
the transmission coefficients on the wave length. The ozone trans- 
mission can be determined from this plot in a similar way from the 
difference of observed points from the straight line. The trans- 
mission coefficients for March 24 were treated independently in this 
second way. The area resulting was very closely the same as that 
shown in Figure 5, yielding 3.9 mm. ozone path. It is believed that the 
application of this method to data of days which give good trans- 
mission coefficients affords a satisfactory method for determining 
atmospheric ozone from direct intensity measurements. 

In order to make a somewhat more extensive comparison of ozone 
values determined by this method with those previously known from 
observations made with the Dobson apparatus, a series of eight days in 
1928 and 1929, for which values by the Dobson method have been 
obtained at Table Mountain, was treated according to a somewhat 
abbreviated form of the above method. For these days there were 
available the atmospheric transmission coefficients as regularly read 
and already computed from the “ long method ” observations at Table 
Mountain. These values are not given at all points used in the deter- 
mination of the area defined in this present work, but at the regular 
spectrum points ordinarily determined in the solar-constant work. 
These relatively few points are scattered over the spectrum in such 
a way as to outline, somewhat less accurately to be sure, essentially the 
same area as that defined in the present work. In particular there were 
but four ordinates lying in the ozone region, but if these values were 
sufficiently accurate the area of ozone absorption outlined by them, 
and lying between the wave lengths specified above, would be suf- 


NO. 9 DETERMINATION OF OZONE—WULF ll 


ficiently close to the area defined in the above work to yield a satis- 
factory determination of the ozone. 

For these eight days, then, the logarithms of the logarithms of the 
transmission coefficients were plotted against the logarithm of the 
wave length and a straight line passed through the values lying to 
both sides of the ozone absorption. The differences of the anti- 
logarithms of those points lying in the ozone region from the anti- 
logarithms of the corresponding points on the straight line were read. 
These differences were then plotted against wave length to the same 
scale as that described above and the areas planimetered and divided 
by the area corresponding to 1 mm. of ozone path, determined as the 
result of the work described in this paper. Thus an approximate 
determination of the ozone on these days was afforded, utilizing an 
area practically the same as that described above, defined by the 


TasLe I.—Atmospheric Ozone for Eight Days 


Path mm. o» 





Date Present work Dobson 
PICA LOZ Oe sarees seen see Blin atewarete tists acute eee 27 
OCE sha LO2S rere eee ereeie fh tere DB Oiaasste Sut teat s7k 22520 
INOvaedaelO2Sretertem ine erro ccs PAE rowed onar eaheyeites oh orers 1.97 
DECI Ga LOZO Memes oo eiecs TPO Vass siete erect eae 1.08 
Dece 18; 1028 seataecere gece e TeGyaiicemem eines 5s 
TD CGE TO LOD Siprcacrat hacrateyela, Hees 2A cate So 60 vivartye eo. 2.43 
Mary 245 O20 nies aoe ee ees oe Oe eee ee 3.42 
NDT 23 A TO2O mrrensveieacske oe ia/e vec DEAN e re ee erie 3.08 

MICA TI raceeecrer eerie se eee ge 2.35 2.49 


positions in which these points lay. The results are shown in Table 1. 
For these eight days the values of the ozone over Table Mountain as 
determined by the Dobson method were also available and they are 
given in this table for comparison. From these results it appears that 
the average amount of ozone given by the two methods is essentially 
the same. The independent values, however, are not in good accord, 
which may be entirely due to the uncertainties in the present determi- 
nations. It must be emphasized that this is due to the insufficiency of 
the data used to make such a determination, which had been collected 
from solar-constant work, and not to the method employed. It is to 
be noted especially that differences in the relative values for the eight 
days obtained by the two methods of Dobson and of Wulf cannot be 
due to the method of the ozone absorption measurements described in 
this paper, but must be contained in the roughness of the data. For 
quite independent of the evaluation of the areas in terms of ozone, 
these areas should be closely proportional to the ozone values on these 
days. That they are not is actually due largely to the uncertainties in 


I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


the transmission coefficients, which in this case define the ozone area. 
This is shown by a comparison of the March 24 value in this set of 
eight with the value for this same day as given above, which was 
determined from the transmission coefficients at all the points chosen 
in the above work. The former value is 2.8 mm., while the latter 
3-9 mm., and the latter is, as stated above, believed to be a satisfactory 
determination. The cause of this discrepancy appears to lie chiefly in 
the fluctuations in the few observed transmission coefficients. To 
summarize: The use of the few single values of the transmission 
coefficients as regularly determined in the solar-constant work only 
suffices to define an area which gives the approximate amount of ozone 
and is not ordinarily competent to show the fluctuations from day to 
day within an error small compared to the fluctuations. 


SUMMARY 


The transmission of ozone for visible light has been determined 
from spectrobolometric data using the solar spectrometer of the 
Smithsonian Institution at Table Mountain, Calif., with the sun as 
the source and introducing chemically determined quantities of ozone 
in the path of its rays. The results are in close accord with the labora- 
tory results of Colange. Using the results of this study the amount 
of ozone over Table Mountain for one typical day as an illustration 
has been determined. By an abbreviated method, using only the trans- 
mission coefficients normally measured in the regular solar work, the 
value for the ozone over Table Mountain has been determined for a 
series of eight days. The mean of these eight days presumably gives 
a good value for the mean amount of ozone, but this abbreviated 
method is not ordinarily sufficient to show the fluctuations in the ozone, 
since the possible error in a single determination is of the order of 
the fluctuations. The mean value for the eight days differs but about 
6 per cent from the mean value for the same days determined by the 
method of Dobson. It is very interesting that the bolographic method 
depends on ozone absorption in the yellow, while Dobson’s photo- 
graphic method employs the ultraviolet ozone absorption. 

The author wishes to express his sincere thanks to Dr. C. G. Abbot 
for suggesting the problem and for his continued interest and help 
throughout the work, and to Mr. J. A. Roebling for a financial grant 
which made the work possible. The efforts of a number of people 
have contributed directly to the completion of this work, especially 
Mr. Alfred F. Moore, Mrs. Beatrice J. Wulf, Mr. Fred Greeley, and 
Mr. George Cox. The author is grateful to the members of the 
Smithsonian Astrophysical Observatory for their frequent kind 
assistance. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, 85, NO. 9, PL. 1 





Two views of absorption cell and auxiliary apparatus. 





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SMITHSONIAN MISCELLANEOUS COLLECTIONS 


VOLUME 85, NUMBER 10 


HUMAN HAIR AND PRIMATE 
PATTERNING 


(WITH FIvE PLATEs) 


BY 
GERRIT S. MILLER, JR. 


Curator, Division of Mammals, U.S. National Museum 





(PUBLICATION 3130) 


CITY OF WASHINGTON 
PUBLISHED BY THE SMITHSONIAN INSTITUTION 


DECEMBER 19, 1931 


The Lord Gattimore Press 
BALTIMORE, MD., U. S. A. 





HUMAN HAIR AND PRIMATE PATTERNING 


By GERRIT S. MILLER, JR., 
CURATOR, DIVISION OF MAMMALS, U. S. NATIONAL MUSEUM 


(WitH Five PLates) 


Few problems have caused more perplexity to anthropologists, 
physicians, and zoologists than those presented by human hair. Why 
is it that only some relatively small areas of the human skin are 
normally capable of bearing a hair growth dense enough to be in any 
way comparable with the fur of other mammals? Why do men have 
beards and women not? Why are beards better developed in some 
races than in others? What is the cause of baldness, and why is there 
no certain cure for it? Why does baldness commonly occur on the 
crown and rarely on the sides of the head? Why do we turn gray? 
Why does grayness usually show itself first on the temples or in 
the beard? Why does the moustache often remain dark after the 
beard has turned gray? Why do we have hairy eyebrows, and why, 
when there is a difference in color between the hair of the eyebrows 
and that of the head, are the eyebrows usually the darker of the two? 
Why is the hair of the scalp often different in quality from that of 
other parts of the body? Why are there several types of hair—kinky, 
curly and straight? 

To all of these questions so many and such unsatisfying answers 
have been suggested that it would be a huge and useless task to try 
to list them. Variable and inconclusive though they are, most of the 
answers possess one quality in common, namely, they have in their 
background the tacit assumption that all these peculiarities of human 
hair are things that arise from man’s special constitution and its 
reaction to the natural environment or to the artificial conditions that 
man has imposed upon himself. It has, for instance, been urged that 
the general bareness of the human skin comes from the widely preva- 
lent habit of wearing clothes; that baldness comes from barbers and 
tight-fitting hats; that women have less baldness than men because 
women have for centuries taken better care of their scalps than men 
have; that graying hair is the result of a lessening bodily energy 
supposed to go with increasing civilization or “ domestication ;”’ that 
the axillary and pubic tufts of hair were once useful for babies to 


SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 85, No. 10 


. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


to 


cling to; that eyebrows exist for the purpose of keeping sweat from 
running down into the eyes; that men are bearded to protect their 
throats from cold weather; that women are beardless because they 
look better that way. All of which gives evidence of ingenuity if of 
nothing else. 

But not one of the explanations that I have been able to find in 
print has taken into consideration the zoological possibility that many 
features of the human hair system may be generalized primate 
traits instead of specifically human developments. By this I mean 
the possibility that they may be characteristics that are forced on 
man because they are common property of the Primates, the animal 
group to which man pertains. Their explanation, in that event, would 
have to be made less in terms of human activities and requirements 
than in terms of the great heritage of characteristics that man shares 
with all his primate relatives. Each one of these creatures has modi- 
fied his portion of this heritage in such a way as to make it his own; 
or, in more technical language, each one of the 800-odd kinds of 
living apes, monkeys, and lemurs has developed “ specific ” characters 
by which it can be distinguished from all the others while remaining 
none the less a primate among primates. That.man should have done 
the same thing would be far from strange. 

This paper is a brief summary of a study on which I have been 
engaged for several years with the result that I have become con- 
vinced that the chief peculiarities of human hair are best and most 
simply explained as special examples of primate ‘ patterning.” 


WHAT IS MEANT. BY “PATTERNING: 


Patterning is familiar to every systematic zoologist because it is 
seen in every group of animals. It consists in the arrangement of 
(a) contrasted colored areas on the surface of the body, or (b) 
contrasted long and short outgrowths from the surface of the body, 
or (c) combinations of colors and outgrowths, in such a manner that 
the resulting patterns of color or form are sufficient to distinguish 
one related species from another. 

Familiar examples of patterning are furnished by the color mark- 
ings of butterflies, or of the American wood warblers, by the minute 
surface sculpture on the shells of some mollusks, and by the spiny 
outgrowths on the back and head of the different species of iguana 
and horned-toad. Among mammals, striking instances are provided by 
cats and squirrels with their diverse stripes, spots, mottlings, and plain 
colors, and by African antelopes with their stripes and spots as well 
as their maned necks, fringed throats and briskets, and tufted tails. 


NO. IO HUMAN HAIR AND PRIMATE PATTERNING—MILLER 3 


The making of patterns appears to be a process quite distinct from 
that by which a general harmonizing of animals with their natural 
surroundings has been effected. Nearly allied, pallid, desert species, 
for instance, may be distinguished from each other by details of indi 
vidual pattern as obvious as those that serve to mark richly colored 
species living in humid forests. General types of color and surface 
may have their relations to the surroundings in which animals pass 
their existence; but the special patterns of the species that conform 
to any one type cannot be shown to have such relations. It may be 
plausibly argued that the blotched and spotted color schemes of 
arboreal warblers and the streaked color schemes of grass-living 
finches have something to do with the unlike surroundings in which 
warblers and finches pass their lives. But this argument would not 
apply to the differences between the patterns of Blackburnian and 
black-throated green warblers nesting together among the same ever- 
greens, nor to those between savannah and grasshopper sparrows 
living in one meadow. Still less would it be possible to explain, on 
grounds of special needs, why species of horned-toad differ from 
each other in the number and form of the spiny outgrowths on the 
head, or why one species of gnu has a fringe on the brisket and 
another has not. Patterning, therefore, seems to be something physio- 
logically inherent in animals rather than something that the environ- 
ment has imposed upon them. 


PATTERNING IN PRIMATES 


Though patterning occurs in all groups of mammals—even in 
rhinoceros, hippopotamus, elephant, and cetacean—it is among the 
primates that the tendency attains its greatest development. In no 
other group does it make such full use of its chief materials, namely, 
the color of the skin, the color of the hair, and the contrasts that 
can be obtained from differences in quality and length of hair. No 
better example of this process could be given than the one furnished 
by the head markings of monkeys grouped on Plate 1. The animals 
there represented are nearly related species that live under essentially 
uniform surroundings in the great African forest belt. No two of them 
have the same arrangement of dark and light areas on the head; three 
have conspicuous white stripes over the eye; one has a black stripe in 
the same place ; in five the cheeks are white, while in three they are not 
white ; one (fig. 2) is bearded, while seven are not; one (fig. 4) has a 
moustachelike mark of white in the skin of the upper lip; another 
has a boldly contrasted spot of fine white hairs on the nose. Other 
patterns in primates come from lengthening, shortening, and varying 


4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


the direction of growth of the hair on different.parts of the crown, 
also from varying the length and quality of the hair on the chest, 
shoulders, tail, and legs, and from making contrasts, often more 
striking than the one seen in the white-lipped guenon shown on 
Plate 1, in the color of different portions of the skin itself. All of 
these elements of color and hair growth are combined and recombined 
in a variety that seems to be without end. 

On no part of the primate form is patterning so conspicuously 
developed as on the head, where strikingly marked color designs of 
both hair and skin are profusely exhibited, and where tufts, beards, 
moustaches, whiskers, and crests are brought into varied contrast with 
areas of short hair and bare skin. 


PATTERNING ON THE HUMAN HEAD 


In conformity with this universal primate trait human patterning 
shows itself more conspicuously on the head than on the body or limbs. 

The human head pattern is not exactly duplicated by any other 
primate, but all the elements that enter into it can be easily found in 
nonhuman members of the order. The usual head pattern of the young 
adult Caucasian is shown in Plate 2, Figures 1 and 2. Characteristics 
that both sexes have in common are the completely haired cranium, 
the bald forehead, nose, and upper median part of the cheeks, and 
the presence of a narrow transverse hairy strip on the forehead over 
each eye. The female’s pattern differs from the male’s in an extension 
of the bare area downward over the entire lower part of the face and 
sideways to the ears. 

In most primates the forehead and face, except the region im- 
mediately bordering the eyes, nose and mouth, are thickly haired. 
The first step in the process of baring the forehead is shown by one 
of the Celebean macaques, Magus hecki (pl. 2, fig. 3). Other steps 
have been taken by some of the South American monkeys; while 
an essentially human forehead can be found in the orang (pl. 2, figs. 
4, 6). The bare or nearly bare lower part of the face seen in the 
females of all human races, and in the males of those races in which 
the beard is slightly developed, is presaged by the very common 
occurrence among other primates of a short-haired, nearly bare area 
around the mouth (shown by all of the monkeys represented on pl. 1). 
Extensions of this bare area on the cheeks may be seen in the great 
apes. It is carried farther in some of the South American monkeys, 
culminating, apparently, in the “ cotton head,” Oedipomidas oedipus 
(pl. 2, fig. 5), which has reached a stage slightly more advanced than 
that of the human female. 


NOs 1O HUMAN HAIR AND PRIMATE PATTERNING—MILLER 5 


Though a bare or nearly bare condition of the mouth area is the 
usual condition among nonhuman primates, it is not universal. Beards 
like those of the male Caucasian or Australian occur in the orang 
(pl. 2, fig. 6), in the bearded African guenon shown at the top of 
Plate 1, and in a South American monkey (Pithecia chiropotes) of 
which I have not been able to obtain a photograph. Moustaches are 
not common among primates. Even that of the full-bearded orang 
is poorly developed. But the South American Mystax imperator 
(pl. 2, fig. 7) goes far to make up for this deficiency. While neither 
moustache nor beard is peculiar to man the strong development of 
both together appears to be a human specialty. 

Nothing exactly like the human eyebrows is known in other pri- 
mates, but the brow region is one where patterns are made in great 
profusion. Sometimes these brow patterns take the form of light 
or dark stripes (as shown in pl. 1 and in pl. 4, figs. 10 and 13) ; some- 
times they are made by lines of hair differing in quality and direction 
from that of the head (pl. 2, fig. 8), thus showing a near approach 
to the condition found in man. Human eyebrow hair, as is well known, 
often differs in color from the hair of the crown. In such cases 
(pl. 4, figs. 11, 12) it is usually darker than the crown hair, after 
the manner of the gray-cheeked mangabey (pl. 2, fig. 8) or the 
Himalayan langur (pl. 4, fig. 10) ; rarely if ever is it conspicuously 
lighter, after that of the white-browed gibbon (pl. 4, fig. 13). 

This normal human pattern does not always remain constant 
throughout life. Changes of two kinds usually take place; and the 
courses of both kinds tend to follow lines that can be traced through 
the group of primates at large. 


TURNING BALD 


With arrival at full maturity a considerable percentage of human 
males undergo a modification of their hair pattern that serves to 
differentiate them still further from the females. The forehead line 
begins to rise, either uniformly along its entire extent (pl. 3, fig. 1), 
or, more commonly, by pushing back a blunt reentrant wedge on each 
side (pl. 3, fig. 4). Frequently a bare spot begins to form at the 
same time on the top of the crown (pl. 3, fig. 6), and the hair of the 
entire median part of the crown becomes sparse. These changes may 
continue until the bald forehead area has been carried back over the 
dome of the head, leaving a well-haired border extending around the 
sides and across the nape (pl. 3, fig. 8). 

This series of maturity changes in the hair covering of the human 
male head has been the subject of endless speculation. By a few 


6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


writers it has been recognized to be, like the beard, a secondary sexual 
character,’ but, so far as I am aware, no one has hitherto shown that 
it follows the lines laid down by the patternings of other primates. 
The uniform raising of the forehead line can be found as a specific 
character in the bald chimpanzee ; it is exactly paralleled by the pattern 
of short and long hairs on the head of one South American monkey, 
Pithecia monachus (pl. 3, fig. 2), and by the color pattern of another, 
Cebus hypoleucus (pl. 3, fig. 3). The development of the two blunt 
wedges results in a pattern much like the one present in the Celebean 
black ape, Cynopithecus niger (pl. 3, fig. 5). The bald spot on top 
of the crown is an occasional character of the toque macaque, 
Macaca pileata (pl. 3, fig. 7). The completely developed human 
bald area (pl. 3, fig. 8) is perfectly outlined in the South American 
monkey known as Cacajao rubicundus (pl. 3, figs. 9 and 10). The 
long dark hair at the side and back of the head of this animal occupies 
the area that remains haired in normal human baldness, while the 
light hair on the median area corresponding with the human bald spot 
is so short and sparse that it does not conceal the skin of the scalp 
in the living animal (two have recently been on exhibition in the 
National Zoological Park). Finally it is to be noticed that the human 
bald area follows the outline of the dark cap of the West African 
gorilla (pl. 5, fig. 5) as well as that of a color pattern not infre- 
quently seen in blond men. This pattern, (which appears to occur 
in women also, but is obscured by long hair) is produced by an 
obviously paler tint of the hair that grows on the bald-spot area. 
It is visible as a faint but accurate picture of the color pattern made 
by a bare scalp contrasted with dark side hair. At the Harvard com- 
mencement exercises of 1930 I saw it on the heads of seven of the 
young men awaiting the conferring of their degrees. 


TURNING GRAY 


Another change in the human hair that begins at or slightly after 
the attainment of full maturity is seen in the familiar process of turn- 
ing gray ; this may lead in the end to a stage when all the pigmented 
hairs of the entire body have been replaced by colorless ones. 

This loss of color, like baldness, has given rise to conjecture with- 
out end. And, as in the case of baldness, its near relation to primate 
patterning seems to have passed unnoticed. Nevertheless, it can be 


*The examination of many hundreds of photographs makes it appear prob- 
able that the males of races with strong beards tend to show the highest per- 
centage of baldness, thus differentiating themselves most fully from the females. 





NO. 10 HUMAN HAIR AND PRIMATE PATTERNING—MILLER 7 
shown to have the same tendency to follow the main lines of primate 
pattern making. 

When gray hairs begin to replace the pigmented ones they do not 
appear uniformly all over the body. “ A vigorous man just beginning 
to show a touch of gray on the temples ”’ is an often-heard phrase that 
unconsciously recognizes this fact. When beards were common among 
us everyone knew how usual it was for them to turn gray before 
the scalp. 

As they increase in numbers the gray hairs tend to form patterns. 
These are sometimes nothing more than faint sketches or suggestions. 
Often, however, they develop into striking color contrasts. The 
faint and fugitive human patterns are not always easy to correlate 
with the patterns of other primates, but the definite ones rarely pre- 
sent any such difficulty. 

Eight of these well defined human color patterns with their primate 
homologues are shown on Plates 4 and 5. 

The first and second (pl. 4, figs. 1 and 3), consisting of a white 
beard contrasted with a dark crown, are frequently seen. In the first 
the mouth area is white. In the second it is dark. Both occur in 
many species of monkey, two of which, the African Erythrocebus 
pyrrhonotus and Cercopithecus Uhoesti, are shown in Figures 2 and 4. 
The identity is so obvious that it requires no comment. 

The third pattern (pl. 4, fig. 5), consisting of a white chin beard 
sharply contrasted with dark whiskers and head, is less common. 
Sometimes the white involves the moustache. It is then exactly the 
same as the white area in the African monkey Cercopithecus brazzae 
(pl. 4, fig. 6). I have seen several examples of this human pattern 
with white moustache, but have not yet secured a photograph. 

The fourth pattern (pl. 4, figs. 7, 8, 9) is merely a dark mark on 
the cheek margin of a gray beard accompanying a gray or bald head. 
Insignificant though this marking may seem, it is surprisingly com- 
mon. On April 21, 1930, I visited the Jewish pushcart market district 
in New York City, one of the few convenient places where many 
full beards can now be seen, to look for this mark. I found it in no 
less than 47 out of 55 men with gray or white beards. The same 
dark line at the edge of the longer hair on the cheeks is found in 
many of the monkeys that have a partly bare median facial area. An 
example is shown in Plate 4, Figure 10, the Himalayan langur 
(Pygathrix schistacea). It may be easily observed in immature 
Japanese macaques, animals that are often exhibited in zoological 
gardens. 


8 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 85 


The gray human temple spot (pl. 5, fig. 1) is a common feature 
of primate color patterning. It is particularly well developed in the 
gelada baboon (pl. 5, fig. 2). 

As they increase in size the temple spots often extend backward 
along the sides of the head until they cover the entire area that re- 
mains haired in normal baldness. The pattern thus formed—dark cap 
contrasted with grizzled sides and back of head (pl. 5, figs. 3, 4)— 
is a common one among non-human primates. It is particularly well 
developed in the West African gorilla (pl. 5, fig. 5. The specimen 
represented by this photograph is an unmounted skin with the head 
not filled out to natural form). Occasionally this pattern may be 
seen reversed. The grizzling is then confined to the area of the normal 
bald spot, while the hairs at the sides and back of the head remain 
dark. When this happens the color scheme of the cacajao monkey 
(pl. 3, fig. 10) is reproduced. 

White locks situated on or near the forehead line (pl. 5, figs. 6, 7) 
are not uncommon, but on other parts of the head they are rare. They 
may be present without other signs of the graying process (as in fig. 6) 
or they may appear as a step in that process (as in fig. 7). In either 
event they are usually confined to some part of an area where pattern- 
ing occurs in nonhuman primates (pl. 4, fig. 6, pl. 5, fig. 87). 


PATTERNING ON OTHER PARTS OF THE HUMAN BODY 


The process of turning gray usually begins on the head and extends 
gradually downward over the body. As it advances it often passes 
through a stage, particularly well represented in Figure 9 of Plate 5, 
in which the gray area ends abruptly at the middle of the chest, leav- 
ing the hair of the arms and lower part of the body dark. The general 
lines of a pattern found in an African colobus monkey, Colobus poly- 
comos, and in an Asiatic macaque, Macaca albibarbata (pl. 5, fig. 10), 
are then closely followed. 

Turning to other parts of the human body we find that the same 
correspondence with widely distributed primate tendencies holds good. 

The pubic region is an area of pattern formation in widely sepa- 
rated nonhuman primates. Young chimpanzees have a white pubic 
patch contrasted with the black surrounding hair. It disappears by 
becoming black before the animals reach full maturity. Some species 
of gibbon have no pubic mark whatever. Others display a black spot 


* The spider monkey represented in fig. 8 of pl. 5 has a band of white extend- 
ing along the entire frontal border of the true head-hair. The forehead-hair 
is also white, but it differs from the head-hair in quality and in direction of 
growth. Before photographing this skin I darkened the forehead-hair with ink. 


NO. I0 HUMAN HAIR AND PRIMATE PATTERNING—MILLER 9 


that stands out against its pale surroundings. Still another gibbon 
has the hair of this region so greatly lengthened that, in adult males, 
it may form a tassel reaching almost to the knees. A South American 
monkey (Oreonax hendeet) has a long yellow hair tuft in the male 
and two shorter tufts in the female. In both sexes the tufts are 
rendered very conspicuous by contrast with the dark belly and thighs. 
By specializing the hair of this region man has, therefore, merely 
followed one tendency of his tribe. 

The last conspicuous hair-pattern feature of man is the tuft in the 
arm pit. This, perhaps, comes the nearest of all the patterns to being 
an exclusively human trait; I have not yet seen an exactly similar 
development in any other primate. But, on gently blowing the hair 
of the axillary region of a freshly dead African monkey, Cercopithe- 
cus aethiops, I once found that the hairs growing in the deepest part 
of the pit tended to separate themselves from the surrounding fur 
by a slight difference in quality and in the direction of growth. More 
recently I have been able to see, in several adult chimpanzees,’ that 
these animals have a definitely specialized axillary tuft confined to 
the region of greatest glandular activity. To produce the human 
condition it would merely be necessary to suppress the long surround- 
ing hair. 


SOME OTHER FEATURES OF HUMAN PATTERNING 


A few other points about human patterning require brief mention. 

(a) The general bareness of the human body. 

Why the human body lacks a protecting general coat of fur is a 
question that has been often asked and variously answered. A final 
explanation seems to be as remote now as ever; but it is possible 
to recognize the fact that human bareness is only an exaggeration 
of a tendency that is found in other primates,’ and that it is no more 
essentially mysterious than the bare face of one tropical American 
monkey (pl. 2, fig. 5) when compared with the fully haired face of 
another (pl. 2, fig. 7). In neither instance can it be shown that a 
special need of the species is served by the bare skin; but in both it 
is evident that the tendency found throughout the primate group to 
form patterns by contrasting long-haired areas with short-haired areas 
has been carried to an extreme. 


2 At the Yale University Anthropoid Experiment Station, Orange Park, Fla., 
an opportunity for which I have to thank Professor Yerkes and Dr. Tinklepaugh. 

?On this subject see Schultz, Human Biology, vol. 3, pp. 303-321, September, 
1931, and Sci. Monthly, vol. 33, pp. 392-393, November, 1031. 


IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


The general distribution of longer and shorter hair on the body of 
gorillas rather closely coincides with the human scheme. By continu- 
ing the process along the lines marked out in this great ape a stage 
would eventually be reached in which the body would become bare 
while the arms and legs retained traces of their original coat. 

(b) The different face pattern of men and women. 

The sexual hair pattern on the human face is another subject of 
age-long speculation. No one has ever been able to show that its 
presence has aided man’s career as a species. Equally impossible would 
it be to show that the analogous sexual patterns in other primates have 
given these species any advantage over their relatives that lack them. 
But it seems clear that in this respect man has developed in the same 
general way as the white-cheeked gibbon of Siam, the orangs of 
Borneo and Sumatra, the black howler monkey of South America, 
and the macaco lemur of Madagascar, all of which have sexes that 
differ from each other in appearance. That is to say, man and these 
other primates have followed a tendency that may crop out anywhere 
in the group of animals to which they all belong. 

(c) Racial differences in hair pattern and in general color of the 

hair. 

It is well known that not all races of man are exactly alike in hair 
pattern. Some have better developed eyebrows, beards, pubic patches, 
or axillary tufts than others; some appear to be not as subject as 
others to grayness and baldness. Racial tendencies toward darker or 
lighter colored hair are also well known. These racial characteristics 
have never been satisfactorily explained on the basis of the special 
needs of different peoples. On the other hand, as examples of the 
slight differences that are everywhere found among races of primates 
nearly related to each other they are readily understood. 

The differences between the two races of orang, for instance, are 
of this nature. The United States National Museum contains 6 males 
and 6 females of the Sumatran orang, 6 males and 10 females of the 
Bornean race, all adult or nearly adult. These two series show the 
same kind of differences that are shown by races of men. In the first 
place, the beards of the males are much better developed than those 
of the females. Then, when the beards of the Sumatrans are compared 
with those of the Borneans they are at once seen to be larger, so much 
so that an adult male from either island can usually be recognized 
at once by this feature alone. Finally there is a general difference 
in the color of the hair on body and head, this being more tawny in 
the Sumatran race, more mahogany brown in the Bornean. 


NO. 10 HUMAN HAIR AND PRIMATE PATTERNING—MILLER Ld 


These two races of orangs inhabit separate parts of one climatic 
zone, exactly as Caucasians and Mongolians inhabit opposite ends of 
another. Therefore the differences in hair growth can be no more 
attributed to the influence of unlike natural surroundings in the second 
instance than in the first. But it seems clear that the two races of 
orang and the two races of men are both in early stages of species 
differentiation, and that the manner of their differentiating is one that 
is common to the whole primate group. 

(d) Total graying. 

Often, though not invariably, the process of turning gray culmi- 
nates in a stage of complete whiteness. But even when a human being 
has turned gray over the entire body or even has lost all hair color 
he has done nothing that is essentially new or peculiar for a primate. 
Light gray or nearly white species of primates have arisen in both 
Asia and South America. These animals are not albinistic nor in any 
way individually abnormal. Their near relatives, living in the same 
regions, are richly colored; and there is nothing to indicate that either 
light or dark has any advantage over the other. General graying and 
whitening in man seems likely to be nothing more than another ex- 
ample of human submission to a rule that some other primates have 
followed. Therefore the strong tendency present in the “ white race ” 
of man for the hair to lose its color at an early age may be part of 
a racial process of depigmentation that has already almost whitened 
the skin and that may be destined, in the future, to bring about 
permanent whitening of the hair as well. 


CONCLUSION 


For the present I wish to avoid detailed discussion of the published 
attempts to explain those peculiarities of human hair that have just 
been passed in review. Most of the authors who have considered the 
subject have done so from the view-point that these peculiarities must 
have originated from conditions (pathological or cultural) or needs 
(physiological or esthetic) that pertain exclusively to man. That this 
view-point is wrong seems to be sufficiently indicated by the evidence 
here selected from the large mass that I have assembled. This evidence 
points to the probability that man has these characteristics because, 
as a primate, he cannot avoid them. They are common property of 
the great group of mammals to which he pertains, and neither he nor 
any other member of this group can wholly escape from the tenden- 
cies imposed on all of them by their primate heritage. With regard 
to no nonhuman primate can it be shown that the possession of any 


I2 , SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


special assortment of these characteristics makes a species better, 
more efficient, or more at ease in the world than one that has another 
assortment. So also with man. 

The similarities that I have shown to exist between some hair 
characteristics of man and those of particular monkeys and apes must 
not be supposed to indicate any special relationship between man and 
these other primates. When superficial features of this kind are 
common to a whole group they will often appear in almost identical 
form in two animals whose relationship is shown by their anatomical 
structure to be remote. 


EXPLANATION OF PLATES 
All figures greatly reduced, not to scale 
PLATE I 


Color patterns on the heads of eight species of African guenon (Cercopithe- 
cus). From Elliot, after Pocock. 


PLATE 2 


The human head-hair pattern and its characteristics as they occur in other 
primates. 


Fics. 1, 2. Young adult Caucasian. 


Fic. 3. Partly bare forehead of a Celebean macaque (Magus hecki). 
Fic. 4. Bare forehead of an orang. 

Fic. 5. Bare face of a “cotton head” (Oedipomidas oedipus). 

Fic. 6. Beard of an orang. 

Fic. 7. Moustache of a marmoset (Mystax imperator). 


Fics. 8, 8a. Eyebrows of a mangaby (Cercocebus albigena). 


PLATE 3 


Types of human baldness and the corresponding conditions in other primates. 


Fic. 1. Raised human forehead line. 

Fic. 2. A South American monkey (Pithecia monachus) with hair pattern 
corresponding with the raised human forehead line. 

Fic. 3. A South American monkey with color pattern corresponding with the 
raised human forehead line. 

Fic. 4. The two reentrant forehead wedges in man. 

Fic. 5. The two reentrant forehead wedges in the Celebean crested macaque 
(Cynopithecus niger). 

Fic. 6. Bald spot at middle of crown—human. 

Fic. 7. Bald spot at middle of crown—toque macaque (Macaca pileata). 

Fic. 8. Complete, normal, human bald crown area. 

Fics. 9, 10. Nearly bald crown area in a South American monkey (Cacajao 
rubicundus ). 


NO. I0 HUMAN HAIR AND PRIMATE PATTERNING—MILLER 13 


PLATE 4 


(a) Human color patterns formed during the process of turning gray and 
the corresponding patterns in other primates. 


Fic. 1. White face contrasted with dark head in man. 

Fic. 2. White face contrasted with dark head in an African monkey (Eryth- 
rocebus pyrrhonotus). 

Fic. 3. White face contrasted with dark mouth area and dark crown in man. 

Fic. 4. White face contrasted with dark mouth area and dark crown in an 
African monkey (Cercopithecus l’hoesti). 

Fic. 5. White chin and lower lip contrasted with dark face and head in man, 

Fic. 6. White chin and mouth area contrasted with dark face and head in an 
African monkey (Cercopithecus brazzae). 

Fics. 7, 8, 9. Dark area at edge of light cheek hair in man. 

Fic. 10. Dark area at edge of light cheek hair in an Asiatic monkey (Pyga- 
thrix schistacea). 

(b) Eyebrow patterns, human and simian. 

Fics. 10, 12. Dark eyebrows contrasted with light head hair. 

Fic. 11. Dark eyebrows contrasted with hair that has turned white. 

Fic. 13. White eyebrows contrasted with black head (white-browed gibbon, 
Hylobates hoolok). 


PLATE 5 


Human color patterns formed during the process of turning gray and the cor- 

responding patterns in other primates (continued). : 

Fic. 1. The human gray temple area. 

Fic. 2. The gray temple area in the gelada baboon (Theropithecus gelada). 

Fics. 3, 4. The human gray temple area extended around the head. 

Fic. 5. Color pattern on the head of the East African gorilla. 

Fics. 6, 7. White locks on the human forehead line. 

Fic. 8. White stripe along the forehead line in a South American spider monkeys 
(Ateles hybridus). 

Fic. 9. Gray area extending downward from head to middle of chest in man, 

Fic. 10. Gray area extending downward from head to middle of chest in an 
Asiatic monkey (Macaca albibarbata). 


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SMITHSONIAN MISCELLANEOUS COLLECTIONS VOE85;, NO. 107,"REt 





Patterning on the heads of eight species of African monkey. 


(For explanation see p. 12.) 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. (85; NOs 10); PR? 





Human face patterns compared with the analogous patterns of five 
kinds of nonhuman primates. 


(For explanation see p. 12.) 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85, NO. 10, PL. 3 





Human baldness patterns compared with the analogous patterns of five kinds of 
nonhuman primates. 


(For explanation see p. 12.) 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85; NO 10; (Piet 





Fics. 1-10.—Human erayness patterns compared with the analogous patterns 
of four kinds of nonhuman primates. 
Fics. 10-13.—Eyebrow patterns, human and simian. 


(For explanation see p. 13.) 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOET 65. (NO. 10, (PE. 


) 





Human grayness patterns compared with analogous patterns of four kinds of 
nonhuman primates. 


(For explanation see p 3.) 














SMITHSONIAN MISCELLANEOUS COLLECTIONS 


VOLUME 85, NUMBER 11 


(End of Volume) 


Hodgkins JFund 


SUPPLEMENTARY NOTES ON 
BODY RADIATION 


BY 
L. B. ALDRICH 





(PUBLICATION 3131) 


CITY OF WASHINGTON 
PUBLISHED BY THE SMITHSONIAN INSTITUTION 
FEBRUARY 2, 1932 


The Lord Baltimore Dress 


BALTIMORE, MD., U. 8 A. 





Hodgkins Fund 


SUPPLEMENTARY NOTES ON BODY RADIATION ® 
By L. B. ALDRICH 
WALL TEMPERATURES AND BODY RADIATION 


In present-day ventilation, three basic factors are considered: 
(1) air temperature, (2) relative humidity, and (3) air movement. 
The results of my previous report indicate that, in addition, considera- 
tion should be given to a fourth factor, the temperature of the walls 
and surrounding objects. 

For normal indoor conditions, with the surrounding objects all at 
the temperature of the air in the room and with the subject clothed 
and at rest, the radiation loss of a human subject is nearly one half 
of his total heat loss. This radiation emitted from skin and clothing 
has been shown to be nearly that of a “ black body.” We may assume 
that, by virtue of repeated reflections from the other walls and sur- 
rounding objects ina closed room, the radiation from the walls to the 
subject is also nearly “ black.’ Then the radiation loss of the sub- 
ject is proportional to the difference of the fourth powers of the 
absolute temperatures of the subject and the surroundings, in accor- 
dance with the Stefan-Boltzmann law. Suppose the mean surface 
temperature of a clothed subject to be 32° C. and the mean tempera- 
ture of surrounding walls and objects to be the same as the air 
temperature, 23°C. The difference of the fourth powers of the 
absolute temperatures is 977 < 10°. Now imagine the air temperature, 
humidity, and air movement to stay constant and the wall tempera- 
ture to be lowered 10°. The difference of the fourth powers becomes 
1963 X 10°, an increase of 100 per cent in the radiation loss. On a 
winter day the temperature of exposed walls might easily be IO” 
below air temperature, and the inner surface of window panes prob- 
ably would be considerably more than 10° below air temperature. 
Thus a subject, particularly if on the exposed side of the room, would 
radiate at least twice as much on one side as on the other, and his 
total loss of heat would be increased some 25 per cent or more. 

Of even greater importance is the consideration of surrounding 
objects which are at higher than air temperature. As before, suppose 


See A study of body radiation, Smithsonian Misc. Coll., vol. 81, no. 6, 
1928. 
SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 85, No. 11 


2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


the air temperature, relative humidity, and air movement to remain 
normal but let the surrounding walls be raised to a temperature of 
32° C. Radiation loss from a human subject would now be negligible, 
the normal balance between heat produced and heat lost would be 
destroyed, and until readjustment is made, a condition of discomfort 
results. In actual schoolroom conditions, a student near an unshielded 
steam radiator or other artificial heat source is exposed to a tempera- 
ture much higher than his surface temperature. In classes, the student 
is surrounded by other students and the summation of the solid angles 
subtended at a point on one student by the other students may be 
very appreciable. 

As a rough example, assume a class of students placed in rows, 
with spaces of 2 feet between students in a row, and the same dis- 
tance between rows. To simplify matters, imagine each student to 
be cylindrical, 1 foot in diameter and 4 feet high. The four students 
nearest to a given student would occupy roughly 1o per cent of the 
total space to which the central student is radiating. The four next 
nearest students exposed to the given student would occupy an addi- 
tional 5 per cent, and the eight next nearest another 4 per cent. Sum- 
ming up, the amount of space occupied by surrounding students would 
be about 20 per cent of the total space to which the central students 
radiate. If we reduce the space between students to only 1 foot 
instead of 2 and proceed to sum up in a similar manner, the area 
occupied by the other students increases to about 35 per cent of the 
whole. For a spacing of 3 feet between students it reduces to only 
10 per cent. In other words, when students are spaced 1 foot apart, 
the total radiation loss of each student is some 35 per cent less than 
if he were alone in the room. When the spacing is 2 feet between 
students the radiation loss is 20 per cent less than if he were alone, 
and when the spacing is 3 feet the radiation loss is 10 per cent less. 
These rough figures serve in a general way to show the relationship 
between the spacing of students and the radiation loss of individual 
students. 

For a given wall temperature, what air conditions produce maxi- 
mum comfort? Evidently if the walls are cold an increased air tem- 
perature is indicated, and vice versa. A further study of the effect on 
a subject of various wall temperatures under controlled air conditions 
is needed. Such a study should tell us to what extent one’s radiation 
loss may be altered without producing discomfort and should furnish 
evidence as to the minimum spacing advisable in classrooms without 
injurious reaction resulting from decreased heat loss. 


NO. II BODY RADIATION—ALDRICH 3 


ACCURACY OF SKIN TEMPERATURE MEASUREMENTS 


In the study of body radiation above referred to, skin and clothing 
temperatures were measured by a special thermoelement device sug- 
gested by Dr. C. G. Abbot. For convenience I quote the following 
illustrated description of the instrument from my previous publi- 
cation: 


For the direct measurement of skin and clothing temperatures, a special device 
was prepared with the help of Mr. Kramer, the Observatory mechanician, and 
embodying Dr. Abbot’s suggestions. The device is shown in Figure 12 It con- 
sists of a specially mounted copper-nickel thermoelement of fine drawn wire. 
A frame of German silver is bent as shown in the figure and fastened in a 
wooden handle, 1”. Two silk threads are stretched to form a cross between the 





Fic. 1—Thermoelement device for measuring surface temperatures. 


F—Fibre rings. P—Spring steel projection. 
S—German silver frame. U—Silk thread. 
W—W ooden handle. T—Thermoelement. 


four spring-wire posts, p. The thermoelement wires are fastened symmetrically 
to these silk threads with the junction straddling the lengthwise thread. The 
wires lead out through fibre rings, Ff, and through the wooden handle. The 
copper wire (see fig. 2) leads through a switch to a sensitive type Leeds and 
Northrup D’Arsonval galvanometer and thence to the constant temperature 
junction in a stirred kerosene bath as shown in Figure 3. The Cu-Ni wires are 
sufficiently long so that all desired positions can be reached without moving 
the constant temperature bath. Holding the device by the wooden handle, one 
presses lightly the four prongs of spring wire f/ upon the surface whose tem- 
perature is desired. This places the junction in excellent contact with the sur- 
face. There is no backing to the junction save a single silk thread, and thus no 
possibility of heat piling up and causing too high temperatures. For about 4 cm. 
on each side of the junction, the wire also touches the surface and assumes 
the surface temperature, thus eliminating error due to cooling of the junction 
by conduction along the wires. 


*The figure numbers of the original publication have here been changed to 
accord with the arrangement in this paper. 


SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 





T C 


Fic. 2——Diagram of electrical connections of copper-nickel thermoelement. 


G—Galvanometer. ; 
T—Thermoelement junction. 4 
C—Constant temperature junction. 













y q 
Ni cd N V 
N Mea aN 
Nit Che N 
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Fic. 3—Bath for constant temperature junction. 


Th—Thermometer. 
D—Stirring device. 
K—Kerosene bath. 
V—Vacuum flask. 


NOw LL BODY RADIATION—ALDRICH 5 


The instrument has recently been recalibrated. In mounting for 
calibration and comparison, the constant temperature junction was 
fastened against the bulb of a mercury thermometer. The ther- 
mometer was then inserted in a metal tube and lowered into a stirred 
kerosene bath, surrounded by a vacuum flask. In calibrating, the 
thermoelement device was placed in a well-stirred kerosene bath whose 
temperature was measured with a second mercury thermometer. A 
sensitive-type Leeds and Northrup D’Arsonval galvanometer was used 
with the instrument. The calibration curve, plotting galvanometer 
deflections against temperature differences, is nearly a straight line. 

As certain systematic discrepancies had been noted between skin 
temperatures observed with this thermoelement and corresponding 
ones computed from observations of body radiation with the me- 
likeron, it was desired to ascertain whether the thermoelement was in 
any considerable degree influenced by air temperatures in making such 
measurements. The instrument was accordingly tested in the follow- 
ing manner by measurements on a skinlike membrane of known 
temperature. 

In the vertical copper calorimeter previously used (see Smithsonian 
Misc. Coll. vol. 81, no. 6, p. 15) three holes were made in the side at 
equal altitude, each 6 cm. in diameter. These holes were closed with 
rubber diaphragms, cemented in with waterproof cement. The thick- 
ness of the diaphragms was as follows (determined with micrometer 
gauge) : 

0.18 mm. (thinnest dental dam) 


0.36 mm. (sheet rubber) 
1.20 mm. (composite sheet rubber used for gaskets): 


Rubber was chosen because it is pliable, simulating the surface pre- 
sented by the skin or clothing. The calorimeter as before was filled 
with water kept thoroughly stirred and a record of its temperature 
determined by a mercury thermometer. 

It is evident that the surface of the thickest diaphragm will be 
appreciably lower in temperature than the water in the calorimeter, 
and that the thinner the diaphragm the more closely the surface tem- 
perature approaches the temperature of the water. By obtaining a 
series of surface temperatures of the various diaphragms, a curve may 
be plotted and extrapolated to zero thickness. The more nearly cor- 
rect the thermoelement temperatures, the more closely the zero dia- 
phragm value will approach the calorimeter water temperature. 

A series of comparisons is summarized in Table 1. Each value in 
the table is the mean of three separate determinations. Air motion was 
produced by a fan in the same manner as in the body-radiation experi- 


6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VoL, 85 


ments above referred to, and velocities (given in feet per minute) 
determined as before with the Hill katathermometer. Before drawing 
conclusions from the data in Table I it appeared advisable to obtain 
more comparisons with other thicknesses of diaphragm. Pieces of 
rubber of the 0.18 and 0.36 mm. thickness were stretched for several 
days and. then cemented into the calorimeter holes previously filled by 
0.18 and 0.36 diaphragms. The new thicknesses measured 0.12 and 
0.27 mm. A series of comparisons with these new diaphragms is sum- 
marized in Table 2. 


TABLE I. 
Calorimeter Water temp. minus thermoelement Air 
Date Room Water temp. at diaphragm thickness of motion 
1929 temp. Temp. oo (in ft. per 
Aug. C. Gc .18 mm. -36 mm. 1.20 mm. min.) 
3 20°50 31°82 °92 1°32 2°46 0 
21.40 32.47 1.03 1.50 1.80 
4 2510 31.50 73 TevLO 1.43 
26.83 28 . 32 .40 .40 357 
5 20.70 35.63 1271 2.43 3.87 
6 26.20 35.73 .97 1.70 2.20 
24.77 33-30 .80 1.33 Bos, 
10 231.27 34.00 27; 2.73 3.07 80 
24.00 30.80 73 1.67 2.57 
27.80 29.83 257 .93 1.40 
7 22.57 32.73 e277 Bn27, aly, 180 
22.43 31.07 1.20 2.90 4.63 
8 23.73 30.60 1.00 2.07 3.00 
23.63 29.27 1.00 Tey 2567 
24.17 26.97 | 270 1.03 1.53 
9 24.97 25507 2.00 3.80 5.63 280 
22.30 28.30 1.20 2.20 35.07) 
2297, 260.50 -47 Tgp 1.43 
23.93 20.37 1.40 2.73 3:27 


From Tables 1 and 2, preliminary plots were made of the differences 
calorimeter temperature minus room temperature and calorimeter 
temperature minus thermoelement temperature, for each thickness of 
diaphragm, and for the four conditions of air velocity, viz, 0, 80, 180, 
and 280 feet per minute. As would be expected, the difference 
between the calorimeter temperature and the surface temperature 
determined by the thermoelement appeared to be a linear function 
of the difference between the calorimeter temperature and the sur- 
rounding room temperature. For each of the plots the best straight 
line was drawn through the points and the origin. From each of the 
plots values of the calorimeter temperature minus thermoelement 
temperature were read off at two places, 5° and 10° calorimeter tem- 
perature minus room temperature. These values were then replotted 


NOo it BODY RADIATION—ALDRICH 7 


as shown in Figures 4 and 5, using thickness of diaphragm as abscissae 
and calorimeter temperature minus thermoelement as ordinates. 
Partly from experimental error and partly because of differences 
in conductivity of the various diaphragms, the individual points in 
Figures 4 and 5 do not all lie on the curves. Smooth curves are drawn 
however with fair certainty. In each case the extrapolation to zero 
thickness yields a zero value of the difference calorimeter temperature 
minus thermoelement temperature. This result is gratifying since it 
indicates that the thermoelement device measures correctly the surface 


TABLE 2. 
Water temp. minus thermoele- 
Calorimeter ment temp. at diaphragm Air 
Date Room water thickness of motion 
1929 temp. temp. SE (in ft. per 
Sept. Cc c .I2 mm, .27 mm, min.) 
22 17°00 24°00 247 293 oO 
26 29.60 37 .97 -43 “73 
28 22.87 28.47 33 .57 
24.63 27.70 20, -40 
25.97 36.67 57 -97 
30 20.50 32.70 .93 2.00 80 
Oct. 2 23.30 27.43 .40 1.30 
5 22.77 33.50 .87 2507, 
22.30 29.63 .60 1.77 
Sept. 30 21637 28.10 “73 1.83 180 
Oct, 2 23.43 ZA, eh 1-2 
5 22.90 33.13 1.10 2.90 
22330 20.37 Ted 2.33 
2 23.50 26.87 -43 1.33 380 * 
5 23.07 B25c3 1.10 2.07 
22.30 29.03 1.33 2.80 


1 This velocity was intended to be 280 ft. but through an error was, found to be 100 ft. 
per minute too great. In Figures 4 and 5, the calorimeter minus thermoelement temperature 
was adjusted to an air velocity of 280 ft. per minute. 
temperature. It confirms satisfactorily the substantial accuracy of the 
skin temperature measurements reported in my previous paper cited 
above. 

The following conclusions also are drawn from the surface tem- 
perature measurements summarized in Tables 1 and 2: 

(1) With the Smithsonian thermoelectric device, the flexibility of 
the surface measured is an important factor as the air motion in- 
creases. On a soft, flexible surface the instrument appears to give 
nearly correct temperatures for all air motions. On a stiff surface it 
probably reads nearly correctly for zero air motion but increasingly 
too low as air motion increases. 

(2) In appreciable air motions, the device shows large, irregular 
drift, making readings difficult. 


8 SMITHSONIAN MISCELLANEOUS COLLECTIONS voL. 85 


IN 


W, 





°o 


NM 


°o 


DEGREES C. BELOW CALORIMETER TEMPERATURE 


DIAPHRAGM THICKNESS —MM. 


Fic. 4—Calorimeter minus Room Temperature = 10° C. 


NOSE 


DEGREES Cc. BELOW CALORIMETER TEMPERATURE 


BODY RADIATION—ALDRICH 





DIAPHRAGM THICKNESS — MM. 


Fic. 5.—Calorimeter minus Room Temperature = 5° C. 


IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


TRANSMISSION OF RADIATION THROUGH THE SKIN 


In measurements previously made comparing temperatures by the 
thermoelement device with temperatures computed from melikeron 
radiation measurements, it was noted (see Smithsonian Misc. Coll., 
vol. 81, no. 6, p. 19) that in measurements on the uncovered skin the 
computed temperatures were about 1° C. higher than those measured 
by the thermoelement. In measurements on clothing and calorimeter 
this difference appeared to be much smaller. It was thought that pos- 
sibly the skin was sufficiently transparent to long-wave radiation so 
that the melikeron in reality received radiation from a warmer layer 
below the outer surface. To test the transparency of the skin the 
following arrangement was prepared: 

Pyranometer S. I. 8 (for description and use of pyranometer see 
Smithsonian Misc. Coll., vol. 66, nos. 7 and 11) was mounted without 
glass hemisphere and with the absorbing strip vertical. A grid, cut 
from platinum foil and blackened, served as a source of low tempera- 
ture radiation. The resistance of the grid at room temperature 
(22.5° C.) was 2.68 ohms. A voltmeter measured the potential fall 
across the grid, and an ammeter measured the current flowing. The 
temperature of the grid was roughly determined from its increase in 
resistance as computed from the voltmeter and ammeter readings. A 
doublewalled screen close to the grid exposed 8 sq. cm. of grid sur- 
face. The distance from grid to pyranometer was 10 cm., which per- 
mitted the interposition of two filters and a double-walled shutter. 

The accepted procedure with the pyranometer is to use the first 
swing of the galvanometer as proportional to the incident radiation. 
When the shutter is opened, exposing radiation to the pyranometer 
strip, the galvanometer spot immediately starts to move and, if the 
radiation remains constant, swings to its maximum deflection in a 
definite time. In the galvanometer used (Leeds and Northrup Type 
R) this first swing required 3.53 seconds (mean of many trials). It 
was noticed that when certain more or less opaque filters were inter- 
posed the galvanometer spot did not start to move immediately and 
took appreciably longer than 3.5 seconds to reach maximum deflection. 
This delayed deflection was due to a combination of the direct radia- 
tion transmitted by the filter and of the radiation from the filter 
itself due to its increased temperature when exposed to the grid. To 
minimize this indirect heating effect, a stop watch was used and only 
those readings retained in which the maximum deflection was reached 
within 4 second of 3.5 seconds. Temperatures of the grid source 
were varied in the range 75° to 170° C. 


NO. LT BODY RADIATION—ALDRICH II 


deflection with a filter interposed 


- : z is a measure of the 
deflection without the filter 


The ratio: 


direct transmission of the filter, plus a small quantity diffusely trans- 
mitted. Tests of the transparency of various screens were made. 
These are summarized in Table 3. 


TABLE 3. 


Transmission of various substances. Temperature of source between 
. 75° and 170° C. 


Material Thickness % Transmitted 

Rock-salt ......:...% (\Quaatiiiti eae seers terete fore tye 8s. 

BIUOFIte. 2465.2 0 de ins CCE WW ch whee is oon enue: 44. 

WhiGal sara cees 1 hee 03 ee pe ocr hee 50. 

Tissue paper ........ [OSs aa Je det oea sees DOut, 45: 

Blotting paper ...... TAY Mos Pedi oss Sawai ee Negligible. 

Hard rabber 20.24 BEA ge. Weare tis ata Seatee Very small. 

Rubber: dam J. ses. 17, A We sorte eh ciner ys Gus Less than 10. 

Lampblack ..sssss07. One coat, painted on 6 (partly due to pin holes) 
rock-salt. of rays transmitted by 

R. S. 

Weampolack NG. oteas es Two coats, painted at Less than 3% of R. S. rays. 
right angles, on R. S. 

Camphor smoke ....Smoked on R. S. plate, 20% of rays transmitted by 


so thick a lamp filament R. S. plate. 
is invisible through it. 


Camphor smoke .... Very thick coat, flaking 6% of R. S. rays. 
off. 
Skin, freshly removed. About 2 mm. ............- Negligible. 


Through the interest of a surgeon in a local hospital, a piece of 
human skin was obtained immediately after removal from the body. 
Its transmissibility was measured before it had materially lost its 
moisture. The piece obtained was about 2 mm. in thickness, with some 
fatty tissue adhering to it. When inserted as a screen in the arrange- 
ment described above, its transmissibility was found to be wholly 
negligible. Bazett and McGlone in a paper entitled ‘“* Temperature 
gradients in the tissues in man” (Amer. Journ. Physiol., vol. 82, 
no. 2, p. 415, 1927) have shown that in general an increase of 1° above 
surface temperature is found at a depth of something over 3 mm. 
below the skin. Forsythe and Christison (General Electric Rev., 
vol. 34, no. 7, p. 440, 1931) and others have pointed out that flesh, 
since it consists largely of water, would be practically opaque to the 
longer wave lengths, just as water is. It seems evident then that the 
higher melikeron skin-temperature values are not due to the instru- 
ment receiving radiation from deeper and warmer layers beneath the 


surface. 


I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 85 


The melikeron is an instrument which responds sluggishly and is 
rather difficult to manipulate. Furthermore, temperatures computed 
from its readings depend upon the Stefan radiation constant and upon 
the assumption that the radiation measured is similar to that of a 
black body. For these reasons the melikeron-computed temperatures 
should not be given equal weight with those measured by the thermo- 
element, and the 1° difference noted may not be entirely real. There 
are, however, three factors each of which tends to make the melikeron 
skin temperature higher than the thermoelement values on the skin, 
namely : 

(1) Due to the ridges and roughness of the skin surface, the 
thermoelement touches the outer and cooler parts of the ridges, 
whereas the melikeron views both ridges and hollows. 

(2) As shown by Bazett and McGlone (loc. cit., p. 433) the tem- 
perature I mm. below the surface of the skin may be as much as 
.6° C. higher than the surface temperature. Since the outer layer of 
skin is scaly and comparatively dry, it may well transmit a small but 
appreciable amount of radiation coming from the moist and warmer 
layer below. — 

(3) Each measurement with the melikeron requires several min- 
utes. The involuntary, psychological reaction resulting from so long 
an exposure of skin near the instrument aperture may tend to raise 
the temperature of the exposed skin. 

Our conclusion then is that the 1° higher temperatures on the skin 
resulting in the mean from the melikeron observations would probably 
be reduced to about 4° if all experimental error were removed. Due 
to the combination of the three tendencies just mentioned, tempera- 
tures at least several tenths of a degree higher than those measured 
by the thermoelement appear to result from the melikeron readings 
on the skin. 





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