Bull1Iss6ArtII


THE FOOD RELATIONS OF THE CARABIDlE AND 

COCCI ELLIDlE. 

BY . A. FORBE . 

A group or a ociation of animals or plants is like a single 
organi m in the fact that it bring to b ar upon the outer world 
onl the surplus of force remaining after all conflicts interior to 
it elf have been adjusted. Whatever expenditure of energy is 
ne e ary to maintain the exi tino- internal balance amounts to so 
much power locked up, and rendered unavailable for external use. 
In many groups thi latent energy is so considerable and is liable 
to uch fluctuation , that a knowledge f its amount and kinds, 
and of the laws governing its distribution, is extremely important 
to on interested in mea uring or for eeing the sum and charac­
ter of the outward-tending activitie of the class. 

This eem e pecially true of the insect world. If the checks 
upon the multiplication of insects and upon their average length 
of life which are due to in ects themselves were to be suddenly 
removed, there is mu b reason to suppose that the total external 
effect of the class would be very greatly intensified, at least for a 

time. 
v hether our purpose be mnely to understand the internal 

economy of insect life as a part of the general system of nature, 
or to apply such knowledge to a regulation of the depredation 
of insects upon pll!,nt and animals, it is equally necessary that we 
hould know the character and extent f t.he conflicts which pre­

vail within the class, and should under tand how the various sub­
ordinate groups limit each other's numbers and activity, either 
indirectly by competition, or directly by destruction. 

The following notes are a contribution to a more exact knowl­
edge of this subject than bas hitherto prevailed. The view of 
the functions of the two principal predaceous families of Coleop­
tera (Carabidre and Coccinellidre) which is common among 



34 The Food Relations of the Carabidce and Coccinellidce. 

entomologists, is largely due to a hasty generalization, based upon 
insufficient data. Observations of the food of these beetles have 
hitherto been left almost wholly to chance, and have nowhere 
been systematically pursued-from which it has resulted that we 
know their habits only in the most conspicuous situations, and 
have not a fair idea of the general average of their food . 
Neither have observations of any kind been numerous enough to 
enable us to detect clearly differences of food habit in different 
species or genera of these families; but,. with slight occasional 
exceptions, all Carabidre and Coccinellidre have been classed 
together as essentially carnivorous. 

Besides insufficient observation, a tendency to reason too confi­
dently from structure to function is responsible for many mistaken 
notions-a tendency particularly liable , to mislead when applied 
to the habits of animals. It is frequently assumed that the most 
prominent and peculiar adaptive structures are necessarily indica­
tive of the most important and customary habits, and that struct­
ures especially fitted for one function are thereby incapacitated for 
every other. 

The first of these assumptions ignores the fact that many 
adaptive structures are acquired for the sake of the advantage 
derived, not in ordinary, but in extraordinary circumstances. The 
struggle for existence is one of greatly varying intensity, and the 
really decisive moments of the conflict are often only brief and 
occasional. The time spent in actual combat by very belligerent 
and very powerful animals, is doubtless but a small fraction of 
their whole lives; and yet by far the most prominent and import­
ant of the structural peculiarities which serve to distinguish them 
from their more peaceful allies, may be those which enable them 
to triumph in these occasional but critical instants. Likewise the 
pinch of starvation must commonly be felt only at rare intervals, but 
no structures will be more thoroughly elaborated or carefully pre­
served than those serving to give the animal the advantage 
during these brief periods, since the continued existence of the 
species depends on these no less than on those of constant use. 
From the prominent adaptive structures we may safely infer, as a 
general rule, what the animal will do in the stress of a life and 
death struggle, but not necessarily what are its ordinary practices. 

The second of the above assumptions is also negatived, occa-



The Food Re/,ation oj th arabidce and Coccinellidu,. 35 

ionally at 1 a t b the principl of natural lecti n e p cially , 
a· applied to the macbin r r f food prehen ion. vYhat v r 
d "partur from the primitiv v Y tarian habit of animal any 

group ha acquir d wa of cour e initiat cl to nabl it to tlra,' 
u other food re ource than th e previou ly open to it. But a

animal foo 1 i u uall le abundant and 1 s generally di ·tribute(l 
than veg table it would not at fir t, be to the ach·antag ( f any 
that th boul 1 be· m xclu ·iv ] d pend nt upon th' former; 
th ir inter st woulcl b 1 e t ·erv c1 by ·uch modifi.cati n of 
·tru ·tur and habit as woul l nabl th 111 to draw upon ne or th •
oth r tore, accorclinCY t circum tance ·. AcquirinCY sorn p ,v r 
t captur and ma ticate aJJimal food, they woulcl not wholly I P 

that f appropriating- ·rney table fooL1 al.·o · and l1ow v�r well fitt ,t1 
their preh n il an l li · tive rgan. might becom for the form 'r 
function w hould expe ·t that th · would not alto ·eth r l · 
th ir fitn for th latter. It , ould he only a: comp titiun in 

thi hio·her plane incr a· d t th prebsure point, that a few 111 111-

b r of the differ ntiatino- -roup w ulc.1 be forced to the high st 
plane of complete l pend nc on animal f od alone. 

The fir t r �ult of an attempt at a more exact and ex.hausti ,·e 
inve ·tiaation of thi ubj ·t wer giv •n by the writer in a hri f 
paper publi h cl in Bull tin 3 of thi · s ries in X ovemb 'r 1 'O::� 

In anoth r paper b l\Ir. F. ::\I. i\ b�ter in the ame Bulletin t a 
ummary of previou ly r conled ob· rYati w wa · giYen, too·ether 

with man , additional and rio-inal fi kl n te.·. -� few oth •r item· 
ha ·in ·e been published b others, hut confined, a far a· known 
t m to hanc ob ·erYation on in

i:')
le in::; <.:t�. 

The metho l here follow cl a in the pap r abov mentiorn'<.1, 
ha b n that of di · tion. Th alim ntary canals of b t1, · 
taken in a o-reat variety of ituations, at various season· and at 
tliff rent tim s of day, have h n r moved, placed in gly erine on 

mi ro ope slid ancl open cl with mall kniv s and mount d 
11 lle a to di play th ·ontent ·ompletely. The have 
th n been tudi 1 with whate r pow r of the mi ro cop wa · 
n ce · ar and mount cl a mi r cop lide for permanent pr ·­
ervation and repeateJ ->xami11ation. Th amount of information 

1''-r\otei, on Insectii1orou· Colwptera. By '. A. Fo:rbe. Illinoi: tate 
Laboratory of �atural Hi tory, Bulletin .Jo. :3, pp. 1!'i3-160. 

tPp. H9-152. 

5 



BG The Food R elations of the Carabid e and Coccinellidrl'. 

whi h ould thu b acquired by patient tudy, was ften quite 
·nrprising. Whil it was of course rar ly possible to distinguish 
·pecies, or even genera, all th fragments ould usually be classi­
fi ed wit h some fair d gre of definiteness ; and there was om­
monly no difficulty in making satisfactory estimates of the ratios 
of the cliff r nt food "lements present. 

In ome of the most important case , the facts elicit cl were f 
the highest degre of exactness. everal collections of pre la­
ceous beetl s were made in situations wh re some particular 

. sp cies of noxious insect was especially abundant, with a view to 
determining to what extent the latter was preyed upon by its 
supposed enemies. In such cases it was not difficult to tell with 
certainty, even fr m very minute fragm nts, whether th g iven 
insect had been eaten or not. Even where no solid structur s 
were present, and the cont ents of the alimentary canal w re 
entirely fluid, it wa still usually possible to say whether these 
fluids had an animal or a vegetable origin. After many observa­
tions and som~ exp riments, it was found that partially digested 
animal food in the stomach of a beetle was commonly bathed in a 
black juice, which, when examined under a high power of the micro­
scope, was seen to contain nothing but a minutely divided flocculent 
matter, probably composed of irregular aggregations of 
fat droplets and other organic particles. This fluid was never 
found in connection with purely vegetable contents, but some­
times filled the stomach alone, and contained nothing to indicate 
its origin . In all th latte r class of cases I have regarded it as 
proof that the food had been derived from animal sources, proba­
bly usually consistin<J' of the juices of ins cts recently captured. 

For the determinations of the fungi mentioned herein, I am 
indebted chiefly to Prof. T. J. Burrill, of the Industrial University 
at Champaign. 

The insects dissected for this paper were partly obtained in the 
course of miscellaneous collecting, and partly secured for me 
especially for -the purpose, by one of my entomological assistants, 
Mr. F. M. W ebster, who kept careful notes of the situations in 
which the specimens were taken, the hour of the day when they 
were captured, and the objects upon which it seemed probable 
that they had lately fed. Examples of the latter w re also fre­
quently bottled with the specimens, for comparison. The special 



Th,. Ponrl Relation.~ of th Carabi<l// ancl 1or·f'i11ellicl(I'. 7 

)ll tion · from th' or hard infrst<)d by C"ankcr-worms, and the 
rn-fi. Id. at .la.ck ·onvilk and ~ ormal o,, 'ITllll hv <"hinl'h-lrno·.·, ·' :-, 

w r mad , b - my lf. 

In th followino· cliscn iou, •ach g nus i · taken up ' 'paratcly, 
and the <l tails of its f o l ar' giv n both unc.1 'l' g neral c·irrum­

. tan e a . h nvn by p' ·im n · from mis e l Ian Pons situations, and 

al nncl r th vanou p' 'uliar ·onditions illu trate l Ii · the 
p uial ollections, madP f r the purpo · f xhibiting th' f ocl of 

th s ins, ,t. a · relat cl to particularly i11juriou ·p i ·, at cl the. e 
ar follow ,c.1 l y a ·umma.r_y ancl tlis ·u · ·i 11 of the food of ach 
family, tak n a a unit. Th fa.ble exhibit, fir ·t, the fo cl of the 
family und r or linary ir nm. tauces; e ouc.1, und r pe ·uliar · n­
dition · and, thir 1, und r all th,_. cir um tan · s, taken tog th r. 

F MILY R BlDLE. 

JI y not upon th f ocl of this family are derive cl from the 
di section and study of ne hun lred and seventy-fiv spe irnens, 
repre enting thirty-eio-ht sp ies and twenty genera. Eighty-two 
specimen were colle ted in mis ellan ou. situations, tw lv w r 
tak n in a fi ld infested by cabbaa -worm ·, ten in a rn--fielcl 
overrun b chincb-bug , and venty-one in an orchard whi ·h 
was being de troye 1 by cank r-worms. The first collection of 
eighty-two pecimen from various ituation · repres nted thirty­
two pecies belonging to eighteen g nera. They were obtained 
in different part of the tate, from D Kalb County in th north 
to Tnion in the outh, and at all ea on· 0f the year, from pril to 
October· a.ml doubtless repr ent fairly well the fo d of the 
family in Illinoi · during· the entire y ar. The collections illus­
trating the f od of the arabid::e a · re lat d to the cab bag -worm 
were made in a fi.el 1 of young plants at Normal, 111., in April, 
1 2 wh re th larv, of Af!J'Otis w111e.1·a were abundant ancl 
de tructi e. The coll tion howing th fool of this family in 
the' pres nee of the chinch-bu o·, consi ted of ten spe im n of a 
single spe ·i s found in July, 18 2, very abundant about th roots 
of corn in a field wher the bases of the stalks w r larg ly c v­
ered by y ung chinch-bug . The thircl pecial colle tion con-
i ted of venty-on in ects, repre enting nineteen sp cies, b-

tainerl in May of two ·ur . siv " y "ar · (1 1 alH1 1 82) in au 



3 The Food Relation of the Carabidre and Cocciuellidce. 

orchard whi h had been infested for several years with the canker­
worm to such an extent as to cause the total destruction of a 
large part of the trees. 

GENUS CALOSOMA. 

This genus is represented by three specimens of C. sc1·utato1·, col­
lected in the orchard with the canker-worms, and by nine of 
C. caliclwn, which were variously distributed. The C. scrutato1· was 
found to have eaten only animal food, about two-thirds of which 
was recognizable as of insect origin. The remaining third was 
due to the occu rrence of liquid animal food, or the fluid to which 
I have given this interpretation. In the stomach of one of the 
beetles the insect food consisted only of minute particles of a 
reddish brown crust which it was impossible to classify further. 
A single C. caliclum, taken in May in Central Illinois, contained 
only liquid animal food. Seven specimens, taken in the orchard 
above-mentioned, had likewise fed upon animal food alone, forty 
per cent. recognizable as insects, and the remainder not otherwise 
determinable. As far as can be judged from the contents of the 
alimentary canal in these thirteen specimens, the species of this 
genus are strictly carnivorous, and have the habit either of suck­
ing the juices of their prey, or of selecting only those parts most 
easily masticated, reducing these to indistinguishable fragments. 
Certainly there was not the slightest trace of vegetable food in 
any of these beetles.-l<-

GENUS ScARITES. 

Two specimens of S. subten·aneits, taken m 1882, one at 
Normal and the other at Anna, in Southern Illinois, had eaten 
only animal food, one-half of which was unrecognizable, and the 
remainder insects. Four specimens of the same species, taken in 
the cabbage-field, have a precisely similar record. 

These nineteen specimens, belonging to three species, wen ~ the 
only examples of Oarabidr,e prope1· whose food was studied, aBd 
all agreed in a strictly carnivorous character. 

*Mr. F. M. Webster has seen a G. calidum eating a small grasshop­
per. 



The Food Relations of the arabidc, and Coccinellidrr'. 39 

GEXU H BRACHYN R. 

A single sp cimen of Bmchynus fwncms, caught 111 Central 
Illinois, in May, had taken only liquid animal food. 

GE GALERITA. 

Seventeen specimens of Galen:ta }({nus, four collected in vari­
ous situations, and thirteen in the o~charcl in Tazewell County, 
had made a much more varied record. All of the group first 
mentioned had eaten insects, which amounted to eighty-eight per 
cent. of their food, nearly all caterpillars of undetermined species. 
The remaining twelve per cent. con isted of v getable food eaten 
by two of the specimens, and. was apparently derived chiefly 
from th eeds of grass. A larger ratio of animal food is 
noticed in the thirteen taken where cank r-worms abounded. 
Here vegetation amounted t only six per cent., all of exogenous 
origin, as shown by the branching bundles of spiral cells in the 
vegetable fragments noticed, while the animal food amounted to 
ninety-four per cent. Ins cts stand at eig hty-five per cent., seven 
per cent. beinD' Diptera, one per cent. unrecognizable insect larvre, 
and th whole of th e remainder caterpillars. The last were 
nearly all asily letermined as canker-worms, which amounted to 
a little over half the food. Seven individuals of the thirteen had 
eaten these worms. Five per cent. of the food (taken by three of 
the specimens) consisted of spiders, and four per cent. (taken by 
a single specimen) was animal food, not otherwise determinable. 
The remains of a caterpillar in the stomach of a single beetle 
were clearly distinguished as those of a noctuid larva (cutworm). 

If from the ratios of animal food taken by the examples from 
the orchard we subtract the ratio of canker-worms (fifty-two per 
cent.) the remamder is just seven times the ratio of vegetation eaten. 
Recalling the percentages of animal and vegetable food taken by 
the four specimens first mentioned, we :find that here also the 
former is almost exactly seven times the latter. This shows beyond 
question that the canker-worms eaten were in addition to the ordi­
nary ratio of animal food taken by this species under the usual 
conditions. 



JO 1'he F oocl R elations of th Cambid(r and C0Nin eltidr1'. 

G EN . . LOXOP E ZA. 

Three specinP 11 s of t his g enus were studi ed, a.11 b -longing to 
the specjes L . 11,tl'in(' ntris, collect d in .July ancl September jn 
Northern and C ntral Illinois. One of these ha<1 at n immense 
numbers of minute, oval, binucleate cells, clet ' rmin cl by Prof. 
Burrill as spores of Sphn.·ronemei, probably Phuma, a fungus 
which forms small bl ack specks upon dead wood, ·te rns of weeds, 
etc. A second specimen had eaten some undet r-min ed insect, 
and about equal qu antiti es of three elements, namely : the above 
spores of Phorna and pollen and anthers of g rass,- doubtless blue 
grass, upon which the insect was taken. A fpw spores of Hel­
minthosporium were likewise noticed. The cro1 of a third speci­
men, taken at N onnal, was distended with an oily liquid, but 
contained nothing else except a few spores of H elminthosporium. 
This specimen had · probably been sucking the juices of some in­
sect. The ratios of animal and veg etable food, as nearly as I 
could estimate them, were as forty-four to fifty- six. A specimen 
of thi.s species, captured in the orchard, had not recently taken food. 

GEN U ' C .A.LATHUS. 

Six examples of ( 1• r11·er;m·iits, three from DeKalb County and 
three from the orchard, are th e only representatives of this genus. 
One-third of the foo d of those first mentioned consisted of cater­
pillars, a second third of other insect larvre, and the remainder of 
the pollen of oTass. The food of the second group was extremely 
similar, a third consisting, as before, of vegetation, anoth er thfrd 
of canker-worms, and the remainder of insect frag ments not 
further determinable. 

GENUS P L ATY N UR. 

The stomach of a single P. clecm·its, taken in the orchard, con­
tained only liquid animal food. Two examples of P. linibcttus, 
both from Southern Illinois, in April, had derived about four­
fifths of their food from the veg etable kingdom, partly seeds of 
grass and partly the parenchyma of exogenous plants. The 
remainder consisted entirely 0£ Aphides (plant-lice). These 
specimens were doubtless too few to give a correct idea of the 
average food of the g enus as a whole. 



The Food Relation8 of I he <'arubidu• ,wd occi11clliclll'. 41 

E ... :-; E ARTJIH,U:-;. 

Five pe ·imen f E. r·o/ ,-;sir.-.;, tak 11 a anou da,t s an<l 
plac had d rived about on -tenth f th ir f o<l from endo · n , 
and th r 'mainder wholl Tw >nt p "r nt., eat 11 

b on f th b tl niz0cl a.· ·at rpillar . arab, ,_ 
id} ar credit <l with an ther tw nty p •r nt., and un<l tennine<l 
larvre of oleopt ra with ab utan -<1ua.l rati . :\Iinute quantiti 
of funai wer n ti cl in th stoma ·h · of two of the· b tl , 
and tra s of un<l termin cl Al ·re in 11 

Two exampl s of E. snrl<llis, taken in the 'l'az well County 
orchard, had con urned nly in c t ·, all ·anker-w rms, xc pt 
trace of an ant and a ingl anat. 

Th insect ratio of th o·enu a, r pr ·ellt 0 d by these sev n 
·p cim n , tand at ninety-thr e per nt. 

GE ... us PTERO, 'TI ·uu:-;. 

Thirteen ·p 1mens w r dis· ctecl, repre · nting P. p ,1·mw1-

du , P. ay'i,, and P. lucttblwul11~. 

The number of ea h ·pe i s i · not ·ufficieut to aiv di tin ·tive 
food characters, an l the genu may th refor best be treat c1 as a 
whole. Seven of th " pecimeu , taken in miscellaneous situa­
tions in Central Illin i in April, May, a,ud eptember, had foun<l 
about one-fourth of th ir f d in th v 'o· table kingdom, about 
one-third of which consisted of uncl tennined fungi, ancl th 
remainder chiefly of xogen us plants. A f 'W por s of Helmin­
thosporium, probably accidental, wer notir- 'U in th st ma h of a 
ingl b etle. Forty-thre per ent. of th" foocl consi ted of 

insect among which Hymenoptera only were r "cogniz <l. A 
singl mite oc urred in ne of the h etl . . 1 lute sp cim "n 
taken in the orchard inf st <l by canker-worm ·, had eaten ndo­
gen u Yegetation, t th amount of about on -fifth f th ir foocl. 
Caterpillar mad 1 ven p r c nt., ancl und--termin cl iu ects tw 
per ·ent., th r maining ratio h 1 ing ac ·ountetl for by th pr ·en ',e 
of liquid animal f od. 'I'wo-_thj 1'(1s of the contents f three 
spe im n tal 11 among th abl>ag s con.·i ·ted or animal matter, 
half of whi ·Ii wa clearly re(.; gnize<l a , tl1, 1arv~ of ., Lyrotis 
<fl11te,1'll inf ·tinu· th<' fi 1<1. Th remaining tl1ird · mp ·ing th 



42 The Food Relationi, of t/ie Uarabidce and Coccinellidce. 

entire food of one of the beetle , consisted wholly of fragments 
of grass.* 

GENUS A IARA. 

Six specimens of this genus were dissected, three of .A. ca1·i­
nata, one of A . anr;ustata, and two of A. impuncticollis. Three 
specimens of A . ccwinctta, taken in Southern Illinois in April, 
1882, had eaten only vegetation, partly derived from gramina­
ceous plants, and partly consisting of seeds and exogenous tissues. 
About one-fourth of the food was recognizable as fungi, chiefly of the 
genus P eronospora. Ninety per cent. of that of a single A. ctn­
gustata, taken in June, consisted of m~tes, the remainder being 
fragments of grass. An A . impu1ncticollis, taken in the orchard 
with the canker-worms, had eaten only vegetable food, chiefly 
undetermined, but with traces of fungi. Another of the same 
species from the cabbage field, had derived its food about equally 
from plant and animal sources, that from the former consisting 
chiefly of grass. 

GENUS DIC.LEL us. 

Three examples of Diccelus elongatits had taken only animal 
food, as indicated by the fluid contents of the stomachs. One of 
these was found in the orchard, and the other in Central Illinois. 

GENUS OHL.LENIUS. 

This genus is represented by twenty-three individuals, the next 
to the largest number studied of any genus of Carabidre. Six 
examples from Southern Illinois, collect d from April to Septem­
ber, belong to the species C. elf/finis, C. ne11w1·aUs, and C. tomen­
tositS. The animal food of these was about three times the veg­
etable. Two-thirds consisted of insects, of which caterpillars 
alone were determinable; and earth-worms eaten by one of the 
beetles made about eight per cent. More than half the vegeta­
ble food consisted of fungi, which included fou rteen per cent. of 
some fleshy fungus, apparently Coprinus, together with spores of 
Dematiei. Fragments of exogenous plants were recognized in 
one of the beetles. A sing le C'. cli(ffinis, taken among the cab-

-:;; A specimen of P. liicublandus was seen by Mr. F. M. Webster 
making a meal from a dead P. sayi. 



'l.'he Food Relatfo11s of the arabidce and GoccinellidtP. 43 

bage-worm , had aten only in ct , hie fly a at rpillar, and a 
larva of a beetle. mere trac of nd g nou vegetation wa 
also detected. Of ixteen p cimens coll te 1 amono- the anker­
worm , three w re ryt/0·01 ll , and thirteen 1

• d(ffini ·. ut­
worm made about ne-third of the fool of th fir t, an d eartl 1-
worms the remaining two-thirds. Th latter wer easily listin­
gui hable by the p culiar pines mix d with dirt in the stomachs 
of the beetles. bout ninety p r cent. of the food of the ther 
pe ies wa of animal origin, and ab ut half the vegetable food 

was fun ·i. In ect made seventy-two p r 'ent., nearly half cater-
pillar f which th greater part (thirty-one per cent.) was anker-
worm Fragm nts of a fly were observ d in one of the heetl , 
and another had eat non of the T, leplwl'iclrP . Mite an l m ria­
pod (Geophilus) had al o been devoured by one. 

GEN AGONODERUS. 

Fifteen specimen of Agonoderus were stu lied, ten f whi h 
wer tho ·e already referred to as representing the food relation of 
these beetles to the chinch-bug. Fragments of that insect 
amounted to about one-fifth th e foocl of all, and w re found in 
four of the beetle ; and plant-lice, taken by half that number, 
amounted to about eight per cent. A single ant, Lasius fl tVus, 
eaten by one wa rated at five per cent.; and ther inse ts 
brought the general average of the class up to tbirty-five p r 
cent. Vegetation made just half the food, all fragm ents of the 
higher plants except one per cent. each of H elmintho porium 
and Perono pora. A single Agonod rus, taken among the ab­
bages, had eaten only undeterminable animal food . F our speci­
mens from various situations had made a similar r cord, differing 
only by the presence of a few mites in the stomach of one of the 
beetles. Eleven per c nt. of fungi, taken by the group last 
mentioned, was deriv d from Ramularia and Coleosporiurn. The 
circumstances of capture, togeth r with the ontents of the 
stomach of one of these beetles, in dicat cl that it 11ad made its 
meal chiefly from the seeds of Jun grass, but the rernaind r of 
the vegetable food could not be mor d finitely classified. 

(i 



44 The Food R elations oj the Carabidre and Coccinellidce. 

GENUS ANISODACTYLUS. 

This large and abundant genus is represented by thirty-one 
specimens, belonging to six species. Five specimens of A. ritsti­
cus were examined, captured in McLean and D eKalb Counties in 
May, June, and July. Two of these had taken only liquid animal 
food, but the remaining three had eaten no animal matter at all. 
Among the fungi found, Cladosporium and Peronospora were 
recognized, and fragments of H epaticre were noted in two of the 
beetles. T wo specimens of A. ha1"1·isi, taken in Union County 
in April, 1882, had eaten only vegetation, all seeds of grass and 
of other plants. A single A . discoideus from McLean County in 
June, contained nothing but liquid food. Seven examples of A. 
baltimorensis, widely distributed in time and place, had derived 
only about fourteen per cent. of their food from the animal king­
dom, all taken by one of the beetles, whose stomach contained 
only chyme. About half of the eighty-six per cent. of vegeta­
tion, composing the en tire food of the remaining six specimens, 
was demonstrably obtained from the seeds of June grass, upon 
which several of the insects were taken. Two examples of A. 
sericeus from Northern Illinois had made about three-fourths of 
their food of grass, and the remainder of unrecognizable insects: 
In the stomachs of two specimens of A. opaculits, fragments of 
seeds and other vegetation were the only objes:ts found. 

Taking together the nineteen specimens of this genus above 
mentioned, collected in various places, we find that animal food 
made about one-fourth of the total, and that the vegetation as far 
as recognized was chiefly derived from June grass and other 
graminaceous plants. 

The record of ten specimens taken from the canker-worm 
orchard, is not especially different from that of the foregoing 
group. Only ?ne of these had eaten animal matter at all, ninety 
per cent. of the food of this consisting of undetermined Diptera. 
Here, again, the recognizable vegetation was chiefly graminaceous, 
only ten per cent. being clearly derived from exogenous plants. 
Two specimens from the cabbage field afford no occasion for 
special remark. The stomach of one was distended with liquid 
animal food; that of the other contained vegetation only. 



The Food R elations of the Carabidw and Coccinellidce. 45 

GENUS AMPI-IA, IA. 

Four examples of A. interstitiali· indicated that this speci 1s 
almost strictly vegetarian, only three per cent. of th food con­
sisting of insects. Of the remaining ninety-seven per cent., little 
can be said except that it was certainly of vegetable origin . 

GENUS BRADYCELLU ·. 

A single specimen of B . clich1·ou · had eaten only in ects, whi h 
could not be forth r classified.* 

GEN s HARPALUS. 

Nineteen specimens of Harpalus wer studied, belonging t 
the three species ca.,ligiriosus, pennsylNtnicus, and lie1·bivagus. 
Two individuals belonging to the first of these species, from 
.r ormal and Towanda in August and September, had taken about 
one-tenth of their food from insects (caterpillars and Diptera). 
Twenty per cent. of unrecognizable animal food an<l five per cent. 
of mites bring the general average up to thirty-five per cent. 
The sixty-five per cent. of vegetation eat n consisted chiefly of 
tissues of grass. A little pollen of Compositre, an<l other exog n­
ous structures were likewise recognized. Three per ceut. was 
fungi, all spores of Helminthosporium. even specimens of H . 
pennsylv inicus, caught in orthern, entral, and Southern Illi­
nois, in April, Augu t, and September, had taken about one p r 
cent. of their food from the animal kingdom. This included an 
ant eaten by one of the beetles, and a few mites tak n by another. 
About half the vegetable foo<l was not further r cogniz,ahle. 
Twenty-nine per cent. was the pollen of rag-weed, taken by two 
beetles captured upon that p lant, and fourteen per cent. wa 
derived from June grass. Fungi made eight per cent. of th • 
food of these beetles, a little of it Helminthosporium, but chiefly 
P eronospora. Three examples of II. Ii 1·biVltgus, taken in North­
ern Illinois, had eaten only vegetation, about one-third of it _ 
graminaceous, and another third fungi. Only seven per cent. of 
the food of the above twelve specimens of this genus, taken from 

1"Mr. ·w ebster reports a ::;pecirn n of B. n 1pPMris taken in 1881 in 
the act of levourin g an earth-worn1. 



46 The Food R elations of the Carabidce and Cocciuellidce. 

ordinary situations, consisted of animal food, of which a little less 
than half was insects. Fungi made thirteen per cent., and the 
remaining vegetable food was about equally divided between 
grasses and exogenous plants. Three specimens of H. caliyino­
sus and H. p nnsylvcmicus, taken among the canker-worms, had 
derived one-third of their food from those caterpillars, while the 
other two-thirds consisted of vegetation, sixteen per cent. being 
Peronospora, and the remainder chiefly seeds and exogenous 
tissues. Four specimens of H. lie1·bivagus, collected in the cab­
bage fi eld, in April, had eaten none of the cabbage-worms, and 
only ten per cent. of insects (Diptera). -The remainder of the 
food consisted apparently of fragments of seeds, as indicated by 
the contents of the cells of the fragm ents and by other micro­
scopic eharacters. A piece of the epidermis of grass was noticed 
in one of the beetles. Taking the g enus Harpalus as a whole, as 
far as these nineteen specimens can be supposed to indicate its 
food, we find that only about one-eighth of it consisted of animal 
substances. Insects stand at nine per cent., two-thirds of them 
caterpillars,-ants and Diptera making up the balance. Among 
the items on the vegetable side of the account, we find fungi and 
pollen of Oompositre each eleven per cent. and seeds and other 
tissues of g rasses, fourteen per cent. 

GENUS p ATROBUS. 

Two specimens of P. longico?'nis, one from Central and the 
other from Southern Illinois, had eaten nearly twice as much 
vegetation as animal food. The latter consisted chiefly of cater­
pill~rs, and included in fact nothing else but traces of plant-lice, 
eaten by one of the two. A little of the vegetation was derived 
from grass, but the source of the remainder could not be satisfac­
torily traced. 

THE FAMILY AS A UNIT. 

W e have now t o treat the various collections of Oarabidre upon 
which this paper is based, as distinct and unbroken groups, with­
out reference to the g enera of which they are composed. The 
eighty-three specimens of all the species obtained in miscellane­
ous situations, are found to have derived forty-two per cent. of 
their food from the animal kingdom, while the seventy specimens 



The Foocl Relations of the Cai-abiclce and Coccinellidw. -:1:7 

captured in th orchard o ft n mention d took sev nty-sev n p r 
cent. of their food from the am our s. Th individuals from 
the cabbage field, however, sh w no such excess of animal food 
as tho ju ·t mentioned, the rati . ta11<ling for them at forty-one 
per cent. If we seek to account for this striking surplus h wn 
by the eco11d group, we shall find, in the fir t place, a <lifferen ·e 
of more than sixteen per cent. betw en the ratios of insects eaten 
by the first and second group respectively- a fact clearly due to 
the presence of canker-worms where the second group wa ·ol­
lect cl. This species was eaten by sixteen of the seventy beetles, 
and composed about one-fifth of the contents of all the aliment­
ary canals. This accounts, however, for only about half th dif­
ferenc noted, the remainder appearing in the larger ratios of the 
other insects, of mollusks, of earth-worms, and of undetermined 
animal food. 

This indicates either that other forms of animal life than the 
canker-worms were superabundant in th orchard, or else that th 
miscellaneous collections do not correctly represent the ordinary 
food of the Carabidre. Th truth I robably lies between the two. 
The extraordinary wetness of the ea on, together with the amount 
of rubbish on the ground in the or ·har 1, gave these be t l s an 
unusual opportunity to capture slugs and earth-worm , and 
afforded excellent harborage for all sorts of insects. On the 
other hand, many of the beetles from other situations wer pr -
erved especially for dissection because the circumstances of th ir 

capture made it seem probable that they were feeding upon veg­
etation. 

These tables indicate one interesting and important fact with 
regard to the preferences of this family, namely, that where an 
extraordinary abundance of any kind of animal food appear d, 
with a consequent increase in the percentage of that kind appro­
priated by the beetles, this increase was compensated, not by a 
decrease in the other animal elements, but in the ratios of vege­
tation nly-a fact which clearly shows that the preferenc s of 
the Cara bidre are for animal food. It should be noticed, howev r, 
that thj argument does not apply to all the genera, as is se n, 
for example, by recalling the re ord of Anisodactylus. The ten 
specimens of this genus taken in the orchard had eaten much 
more veaetation than the nineteen fr m various other place . 



48 The Food R elations of the Carabidce and Cocc:inellidce. 

The combination of these various tables into the final one given 
will tend to correct the defi ciencies of the separate exhibits, and the 
averages of that table will consequ ently be found to represent 
more closely t he g eneral food of the family than either of the 
others. 

Continuing the comparison of th e three separate tables, we find 
that the beetles r epresented by the first had taken insects to the 
amount of t wenty-six per cent.; that those from the orchard had 
about doubled this ratio; while those from the cabbage field fell 
a little short of it. This last fact is probably related to the time 
of the year when these beetles were taken- the middle of April 
in a very late spring , when insect life in g eneral was but just 
beginning to stir abroad. The ratios of Diptera, Ooleoptera, and 
H emiptera, were but trivial in all these groups, and not worth 
separate mention. The extraordinary difficulty of determining 
the elements of the veg etable food from the minute fragments 
found in the stomachs of these beetles, makes it impossible to 
enter into much detail with respect to this. The miscellaneous 
collections and those from the cabbage field had found a little 
over half their food in the structures of plants, while those from 
the orchard had obtained from this source somewhat less than a 
quarter. Pollen of exogenous plants, which will be found to form 
so large a ratio of the food of the family next to be considered, 
appeared here only in three of the specimens, and amounted to 
but three per cent. of the entire food of the first group. These 
beetles fed much more largely on graminaceous plants, the recog­
nizable tissues of which amounted to about seventeen per cent. 
in the first group, and eight in each of the special col­
lections. Fung i were reckoned at about one-tenth of the food of 
the beetles included in the first collection, and only two per cent. 
of those from the orchard. The spores of the omnipresent Hel­
minthosporium make the most important contribution to this 
element of the food, but a number of other genera were recog­
nized. 

A few words will suffice for the final table, summarizing the 
data relating to all the collections, from whatever source derived. 
This table presents the ratios from one hundred and seventy-five 
specimens, and as already remarked, a little over half the food of 
all consisted of animal matter, about one-third being insects, 



The Food R elation of the arabidce and Coccinellidce. 49 
I 

while mollu k , earth-worm · myriapod and rachnida make up 
the remainder. 

All order of in e t are repr ented on the li t, with the excep­
tion of Orthoptera an d N europt ra. The ratios of none of the e 
are of any pecial importance, except that of the Lepidoptera, 
which tand at fifteen per cent. Hymenoptera and Diptera are 
each one per cent., and Coleoptera and H emiptera each two. 
Among the Coleoptera, only carab, idre and Telephoridre were 
recognized· among the Hymenopt ra only a single ant; and 
among the H emiptera, plant-lice and chinch-bugs only. About 
half the vegetabl e food could be di tinauished as exogenous or 
endogenou the remainder being of too indefinite a character to 
be a i ned to either cla . As far as known, the endogenous 
food wa more than twice as abundant as the exogenous, and con-
isted almo t wholl of gras or gra -like plants. The fungi, 

which make somewhat more than a fourth of the food, require no 
further , pecial mention. 

If di carding the ratios given above, we look only to the nu·m­
ber of specimens in which the various food elements are detected, 
we reach irnilar re ult . One hundred and seventeen individuals 
of the one bun lred and seventy-five represented by this final 
table had eaten animal food, and ninety- even had taken vegeta­
tion. In ects were recognized in eighty-two, Lepidoptera in thirty­
one (about one-half of which had eaten canker-worms), Diptera 
and oleoptera in nine and four respectively, and Hemiptera in 
seven. Earth-worms were found in five myriapods (Geophilus) 
in but one, and Arachnida (mites and spjders) in nine. Grass­
like plant were taken by thirty-six, and fungi by twenty-nine. 

Scanning the totals for each genus on this final table, a few 
results are noted which are worthy of special remark. First, we 
observe that at least two very abundant genera, represented by 
specimens enough to give us a fair probability that the average 
food is correctly exhibited, can hardly be classed as carnivorous 
insects at all, namely, Harpalus, with its nineteen specimens and 
twelve per cent. of animal food, and Anisodactylus, with its 
thirty-one specimens and twenty-one per cent. of the same. 
Amara and Amphasia should probably be placed in the same cat­
egory, six specimens of the first and five of the second having 
taken but twenty-three per cent. and seven per cent., respect-



50 The Food R elations of the Carabidce and Coccinellidce. 

ively, of food of animal 01·igin. The excessively abundant Agon­
oderus ranks but little higher as a carnivorous insect, fifteen 
examples having derived only about one-third of their food from 
animal sources. On the other hand, twenty-three specimens of 
Chlrenius, and seventeen of Galerita had taken a bout nine-tenths 
of their food from insects, mites, myriapods and earth-worms. 
Thirteen specimens of Pterostichus had obtained three-fourths of 
theirs from similar sources, while Evarthrus and Calathus, repre­
sented by seven and six specimens respectively, had averaged 
ninety-three per cent. and sixty-seven per cent. 

The fact has already been alluded to that the Carabic1::e proper 
had ea.ten only animal food, and that nearly all this was of a fluid 
character. 

Second, we find the Carabidre dividing into at least three tol­
erably distinct groups as respects their food: first, those which 
seem usually to seize their prey and suck its juices, and take veg­
etation rarely, if at all; second, those which take a much larger 
ratio of animal food than of vegetable, but masticate and swallow 
it, as a rule, including indigestible fragments; and thirc1, those whose 
habit is essentially vegetarian, but which still take solid animal 
food in diminished ratios. A fourth group, consisting of Lebia 
and its allies, is perhaps obscurely indicated by the facts relating 
to the three specimens of Loxop eza atr·iventr·.is studied. This 
will probablY. be found to feed largely upon pollen and fungus 
spores, after the manner of the Coccinellidre ; and the fossorial 
Carabidre will, perhaps, constitute a fifth. 

If we look now 1+) the structures of these beetles for some 
explanation of their differences of habit, we shall find correspond­
ing variations in the form and structure of the mandibles. Where 
the mandibles are long and curved, and are destitute of basal 
molar processes, but are provided at or near the middle of the cut­
ting edge with processes relatively long and sharp, the beetle seems 
to feed substantially upon soft or liquid animal food. If they are 
of medium length, somewhat slender, broad at base and tapering 
distally, with the tip acute, and provided with basal processes which 
are not especially prominent or sharp, the food is chiefly animal, but 
solid structures are masticated and swallowed, and some vegeta­
tion appears in the alimentary canal; while, finally, if they are short 
and quaclrate, blunt at the tips, and provided either with strong 



The Food R elations of the arabulce and Coccinellida,. 51 

ba al proce se or broall uµpo · d urface v getabl food 
i f mH.l to pre<lominat . ~alu oma i ~ an xample of th fir ·t f 
the e cla Chlreniu of the econ<l, anc1 ... ni oda ·tylu, f th 

third. The eeminQ· x ption to thi · o·en ralization Lown b 
o n 

the table at the lose of th pap r, ar found among those g 'll 'ra 
of whieh too f , sp ·imen · hav been studied to warrant o· neraJ 

h 

·onclu ion re pecting th ir foocl. 

FAMILY O "CI.i:TELLlD.Ji. 

Thi · farnil share with the pre ·edi1w th c: retlit of lirnitiug 
the multipli ·ation of ther in , ·ts, but "·a hown in the Bulleti11 
of th Laborator · pr viou 1 mention ed, apparently to (lep >11cl 
larg·el)· while in th au.ult tao· upon fungi antl other v ,a table, 

U ~ h 

food . The rn te in the paper men tion cl r ferre 1 h nveYer, tu so 
mall a num ber of p ·im n · as to make thi · eon lusion ( f lou bt­

ful Yalue. :N" umerou di e ·tion of Coe inellid,t> mad, inee that 
tim ha, e afford d the material for a mu h more cum pr hen ivc arnl 
thorouo-h treatment of th u bj et and the result · of a car ful stud 
of thirt ' -nin e lides ar h r with ,?'iven. The A phi -eati1w habit of 
th oc inelli l.e i a fa t f such a y obsen'ation, and is ·o 
thorough! · well kn wn that I ha,· not thouo-ht it worth wliiJt, to 
in Ye tio·ate espe ially the f o 1 of be ~tl of tbi · family taken 
among plant-lice. 

Th colle ·tion from whi 'h the pre ent 11ot · are derived ,,re 
from a vari ty of miscellaneou ituatiun ' ancl al o from a ·orn­
fiel l mentione l in the note on the food of the prece(ling fami ly 
in which hinch-bugs were sup rabundant, the purpose of tl1 
latter collection being to determine th food relations of the Co ·­
cinellidre to those insects . It so happened t hat the same fi ,1c1 
was infested by the corn Aphis i11 g reat numbers, and the speci­
mens obtaine l therein consequently illustrate to some extent th 
foocl of the lady-bugs in the pr ·en e of plant-lice. It was in 
this last situation only that larv~ were collert"'d, and the fa t h re 
given consequently relate almost wholly to th adult b tlP . 

GEN HIPPOD.,UIIA.. 

Eleven specimens of I-I. m, tculata taken in orthen , entral, 
and outhern I llinois at variou ason of the year from April to 

7 



52 The Foocl R elations of the Ca-mbidce and Coccinellidce. 

September, give an average of forty -six per cent. of animal food, 
all insect excepting a few mites eaten by three of the beetles, 
ancl amounting to only one per cent. of the food. The insect 
ratio, as far as recognized, with the exception of a single Podura, 
·onsisted wholly of p lant-lice, which amounted to thirty-five per 
cent., while the fifty-four per cent. of vegetable food contained 
only pollen of plants and spores of lichens and fung·i, the pollen 
an<l spore occurring in about equal quantities. The former was 
chiefly from flowers of grass and composite plants, about seven 
per cent. of the first and fifteen per cent. of the second. One 
per cent. of the pollen of Polygonum, and a trace of the pollen 
of pine, both eaten by a single beetle, are the only other items 
under this head. Lichen spores, including Pbyscia, were reckoned 
at two per cent., and those of fungi at twenty-five per cent. At 
least two-thirds of the latter, eaten by nearly half the beetles, 
consisted of spores of H elminthosporium. 

Three specimens of this species, taken in the corn-field at 
Jacksonville, had eaten much smaller ratios of animal food, which 
amounted to only thirteen per cent., all insects. Traces of plant­
lice were recogni:ted, but no structures of chincb-bugs occurrec1. 
All but five per cent. of the vegetable food was derived from 
spores of fungi, very largely Claclosporium. Helminthosporium 
amounted to nine per cent. Macrosporium and Septoria were 
also found. Three per cent. of the spores of Physcia and other 
li chens, and twu per cent. of th pollen of rag-wee(l and other 
Composit..e, complete th r 'Cord. 

Four examples of H . r·onverven!:i, all taken at onnal in 
August and September, liad eaten about the same amount of 
animal food as the preceding species (forty per cent.), but differed 
in the distribution of it by the fact that on of the specjmens had 
eaten a myriapod (Geophilus), and that a caterpillar had been 
taken by another. Insects proper amounted. to but twenty-five 
per cent., over half plant-lice. The vegetable food of this species 
stands at fifty-six per cent., as compared with fifty-four of the 
preceding, and the ratios under this head ar very similar to those 
just given for the other species. Pollen of Compositre ( dandeliou) 
makes thirteen per cent. that of grass makes five per cent., spores 
of lichens two, and those of fungi thirty-three per cent. As in 
H. maculata, Helminthosporium was by far the . most important 



fnngu · < 1 m nt. Th0 ther µ; 11 'nl rt' ·ognizPtl ,,ere' ~t"ptoria, 
T~tiJaa ~Ia ro ·p wium 'ol o:p rium P<>r n ·p ra ~l ni. p ra 

an<l por of ph~r n m i an l ~Iyxoga. tr . 
Fi,P adult tak n at Jack n,ille, "- . re fou11cl to hav • 

maclP about m ' -thircl of th ir f od of in ect , ( c1ua 11:v 
c1iYic1 11 between plant-Ii an(l hinch-bng , a 11 aten b on of 
th l tle -. Tlw Y g tation on i. t 1 a u ual of poll n of 
Compo it:"\:' (el ven p r nt.), pore · of Ji he,n (two per ent.) 
and f fnngi ( ,- nt. ·-one per c nt. . Th Ii t f th la t inrlud . 
, ept ria - tila:-- , H lrninth p rimn, ~Ia rosponum lad . po­

rium. antl Perono pora. 
Tw larv,i> f thi · peci tak1'n at th0 anw plae ancl time 

diff ,n cl hut littlp in food, t my :urpri '< frnm th e ac1ults ju t 

menti n <L ( 'hinch-hug , plant-Ji e arnl catPrpillars, in about 
equal ratio. , with tra of unr 'C'< gnizahl0 insPct , arnonnt t 
tw nt ·-thrC'e P' r ·ent. Poll n of Compo. it;e stands a n,·0 
per · nt . lidH'n . por : at . e,· n, an(1 . pores of fnng-i at ·ixt_r-five, 

in Juclino- th amr a 1wra a. tho •p 1·11 . t nwntion cl, exc:C'1)t e ron-, n . 

o. pnra ancl .~eptoria. 
H. !fl,, ,. ;,,t,.• wa r pr ·entPd l :r four ·ppcinwns akPn in the 

c·orn -tir·ld. The> cliff renc · I, w en th '1r food and tl1at of J-1. 
r·1J111 •,,,.!f' n,.; werP pmely triYial. In . ·t~ amount to thirty per 
e nt. all ·liirn-h-l ugs and pJa1 t-JieP tm ln' p <> r 'nt. f th form r 
and ei.., bte n r tl e latter. 'l'he . ,. nty per ·ent. of v g table 
food i · divid cl abont a befor , b tw( en poll n of ,ompo it,-e, 

· yen p e r cent. an1l . por of fungi fi.fty-rn1' per c nt. Licht n 
por were tak nm r fre ely howen•r and were e timat d at 
w ]ye p r cent., aten by all th etl ··. The fun ·i w re m tl -

CJado ·porium (f rt -thre p r ent.) hut eptoria, r <lo, Hel-
mintho. porium ancl P rono:pora likewi. e o cm. 

GEx s oc IXELJ,A. 

1x pe ·unens f thi g nu· wer stucli cl, three of ( '. :J-notata, 
andthr eof C'.3-11otota. llw refrom "ntr:1lillinoi exc pton, 
which wa from .Jack om-ill . Exdudin · th la t, the ratio of 
animal food eaten by these spe ·imen wa not far from two-third 
of the total, all plant-lice. Only a trace of pollen f Composit,, 
wa noti d in one of the in cts. Funo-u pores amounted t 
thirty-two per c nt. (ah ut half H lrninth ):porium an(l .,. tilag1 ), 



. 4 The Food Relations of the Carabidce and Cocciuellida,. 

and lichen spores to four per cent. The .J acksonvme specimen 
had eaten on ly fungi. 

GENUS CYCLONEDA. 

In the corn-field with the chinch-bugs, three specimens of C. 
san,quinea were collected, which had eaten plant-lice, pollen of 
Compositre, lichen spores and spores of fungi. The first made 
about one-third of their food, the pollen grains were estimated at 
nearly half, and lichen spores at three per cent. The eighteen 
per cent. of fungi were of the usual character. 

THE FAMILY AS .A UNIT. 

A summary and comparison of the food of these two groups, 
taken singly without reference to their genera, develops some in­
teresting and unexpected facts. Although the corn-field in which 
the second collection was made was teeming with insects of the 
kinds especially tempting to the Coccinellidre, an'd although these 
beetles themselves were there in truly surprising numbers, it is 
not easy to believe, considering the tables upon which this dis­
cussion is based, that the Coccinellidre were attracted to the field 
by the abundance of insects available for their food. The beetles 
of the first group are seen to have eaten nearly twice as many 
insects as those from the field of corn, while the fungi eaten were 
as thirty-six to fifty-six respectively. Only eighteen specimens 
were dissected, out of the large number collected in the corn-field, 
but the contents of their stomachs were of so uniform a character 
that there was every reason to suppose that they illustrated cor­
rectly the food of the family at that time and place. It would 
therefore seem possible that these beetles were attracted rather 
by the stores of fungi in the field, than by the chinch-bugs and 
Aphides. The condition of the leaves and stalks of the corn, 
drained and deaden ed by insect depredations, was such as to 
afford an excellent nidus for the development of those fungi 
which spring up every where spontaneously upon dead and decay­
ing Yegetation, and these were in fact extremely abundant. An 
alternative explanation is perhaps more probable. The condition 
of the field gave abundant evidence that the plant-lice had been 
very much more numerous some time before; and it is possible 
that, as a consequence of this decrease of food, and the increase of 



T!ie Fooll RelationR of thr> arabidce and Coccinellidrr. , 5 

the Co cinellidc them lYe , th latter had r a he l an x iv 
number, for whi h the ·upply of plant-lice wa rea lly in ·uffi ci nt, 
and that for thi r a on th y hacl re ·ort cl to fungi. 

Th bin ·h-bug taken hy th p cimens f th sec011c1 group 
amountec1 to nly ight 1 r c nt. f th ir ntire foo<l, and plant­
li e to fourte n p r nt. - le than half tho taken by the other 
pecimen , which stancl at thirty- ix p r c nt. Th pollen eat n 

by ea h group wa thirte n p r cent.,- the sam in both. If w 
combine the two coll ction , and treat th thirty-nine specim n 
of both as a who] , w find that in ect foo<l. is ab ut a third of th 
entire amount, and that the oth r animal lements ar only trivial. 
The fun tion of the beetles of this family of limiting the multi­
pli ation of plant-Ii e is expre sed by th fact that these insects 
·ompo a fourth of the food of thi entir collecti n. Th p ll n 
of gra es an l Compo itre make fourteen p r ent., the pores f 
lichens four per ent., and those of fungi nearly half th wbole 
(forty-five p r cent.) . The list of genera, a far as recognized, 
and the relative importance of these, may b found by reference 
to the table at the end of thi paper. 

FFICIENCY OF DATA. 

The f od of the Coccinellich se ms to be, on the whole, remark­
ably simple and uniform, consisting almo t wholly of spores of 
the low r cryptogam ·, pollen grain ·, an<l. plant-li , aml varying 
but littl fr m one germ to anoth "r. This s imilarity is likewi · . 
r fleeted in the m uth part , which agre as closely in f rm and 
structure as do the ratios of the food. I have consequently littl 
doubt that th data derived from the thirty-nine sp irnens h r 
dis ussed, wrn b found sufficient for a · rrect general idea of tli 
food of the family und r ordinary circum tanc . 

With respe t to the Carabidre, we have oth r pro f. 'In th 
preliminary paper in Bulletin 3 alr ady r f rred to, based on. an 
examination f only twenty- ight specimen belonging to seven­
teen p cie , the conclu ion wa announ , l that about one-half of 
the food of this family consisted f v getati n, an<l one-third of 
insects; and th veg tation was th ught to be about qually divided 
between cryptogam ·, grasses and exogens. If these figures or 
those of the pre ent paper were far wrong, the probabilities 
would be very slight ind ed that the two estimate would agree, 



Th Food R elrzt ion.~ of th r ambidr1' and octindlid(I' . 

:p iall · as n c mpari on what"v r wa rnac1 ' of th two 
f data until th "' table· wer ' mplet ~ l in their pr " ent form. 

n, th refor , we find that th on hundred a.11d sev nty-fiv 
p imens f the prese11t paper, b I ngino· to thirty- eight p ci s, 

were estimated to l ave taken fifty-seven per cent. of animal food 
and thirty- ix of ins cts, and that the ratios of cryptoga m , gram­
ina eous plants and xogens are r sp ctively fiv e, I v n, anrl fiy e, 
we must conclude that th se fio·ures ar a fair aYerage f the 

,., < 

ordinary food of the family. 

The foregoing pages have set forth th relations of th ( arab­
iclre and th Cocci11 ellid:v t the species upon whi ch they feed, 
and a few general statem nts will now be proper con cerni 11 g tlie ani­
mals which prey upon them in turn. Predaceou ground-beetles 
are peculiarly exposed to birds whi h commonly seek their foocl upon 
the ground, and we need not be surprised to fin cl that tht'y enter 
largely into the food of su ·h species as the thrush( s ancl th e blue­
bird . Carabidre were found to furnish about fiv or six per cent. 
of the food of four bunrlrctl and twenty-three specimen s of th ese 
birds, as stated in a pa.per 0 11 that ubj ect in the t l1inl Bulletin of 
this series, but Coccin ellirl m did JJ Ot occur at all. I nde0t1, in the 
foo<l of more than four hundred other birds, of various famili es, 
Coccinellidm were found only in R egulns, ·yyJiere a single specie: 
was reckoned at one per cent. of the food. 

The great differences in th e food of the Carabida:, dis losecl hy 
this paper, give c nsirl rabl e importan ce to the question of the 
kinds of these beetles m st freely eaten by birds, allcl th follow­
ing list of specit · and genera recognized in th e food of the col­
lection of thrusl1e anc1 bluebirds above mention e(l is given as an 
answer. 

It will be see n tl1at there is a very wide difference between the 
number of Carabidm proper taken by these birds, and the number 
of Harpalidre, representatives of the form er group occurring in 
only six specimens, and of the latter in one hunched and sixteen. 
On the other hand, fifty-nine of the birds had taken Harpalids 
which may be fairly classed with the second group established in 
this paper, and fifty-seven had taken those belonging to the third 
group, or phytophagous Carabida:-. The genera most preyed 



Th Foocl R elalio11. of !lie ambid1 c and o,·<'inellidrr. 57 

up n a.r Harpalu tak n b tw ·- iaht f th bird ni ·oda -

tylu b · ·hte f urte n, Crnta an hu by thir-
t, '11 t r ti an 1 Evarthru h · •lev n · numb r 

which r pr ' n fairl w 11 th r lati,· abun lan · ~ f individual· 
taking th vY 11 t , ho, v r, a r marka­
l I d fi •ie>n y of th, highly c l r cl g 11 ra, u h a. G-al rita, 

Brach nu , L bia, Platyuu ·, ~lih·niu de. whi h ar ~ith 'r 
ab · nt r f und but rar 1 in the · birds foocl. Evid ntl th · 
more ·h wy bee ti ar, prot t d by ·om m r ff cti v , 
m an than b urity of color. Tn th-' foll win · list, th " figm 
pr din · th , 11am of "ach ·p ei ... of bird Ll n te th numb r 

f p im n · in whi h tlw in · c:t 111 ntion cl was found: 

IX' T OF GEXER A:ND :-:\Pb ' IE. ' . F 'I lXDELID.E AXIJ AR.\.B­

lD}E EATEX BY 423 OI•' THE TlIRl'.'HE,' AMD THE BLUEBIRD. 

1. Ci indela le ont i: 1 .Jlimns ·arolin n~is. 
•) Carabus palustri : l .., iali<1, siali ·. 

;3, carit , sp .: l Harporh n ·u: rnfus. 
-!. Di hyrius globule u ·: ] '1\m1u palla ::;i. 
,). ~ pidoglo ·sa ·ubang-ulata: 2 Iinrns car 1iuen is. 
H. "ilivina bipu tulata: 1 Hal'porhyncu.- rnfus. 
7. Platynu, sp.: 11Iirnu: ean lim•n ·is, 1 Harporhyn ·us rufu , l 

ialia sial i ·. 
Evarthru ·, sp.: l Tunlu: mustt>linus, 1 'I'. migratoriu.- :Z. Mimus 

ar linensi , ,., 'ialia sia li:. 

tl. E. olo · ·u ·: 1 Turdus palla ·i, I Harporhyn ' U · rnfu . 

10. Pl 'rosti •hus ·p.: l Turclu · 111ust -, Jill\1 ·, L T. palla. i 1 T. rni-
grat riu 1 Harporhyn c u · rufos, ;> .._ ialia ·iaJi ·. 

11. P. In ublam1us : 1 Turdu mu:t ,Jinus, l ::-;ialia iali ·. 
12. P. :ayi: 1 Mirnu · carnlin -' n ·is. 
] ;-3_ Amara sp.: ] Tm lu · pa.lla:i l 'I'. ·wains ni , T. mig ra.t -

rius 2 "'ialia iali ·. 
1-t-. Bnt ·bylobus li th philu ·: ;3 'l'unlu · 1uigTa.t< riu ·. 
13. 'ht ~nius, p .: 1 Tunlu migratoriu · 1 Sialia iali ·. 
16. '. tom ' nt ·u : 1 ialia . iali ·. 

17. Laclino -r pi parall lu : I Tunlus migratlll·iu ·. 

1 . G pinu in ·ra atus: 1 'l'urdus rnigratoriu~. 

] U. -, ratacanthu · dubiu : 1 Tur<lu. must linu , 3 'l. mi rat riu., 
2 Mimus -~1rolin n i , Harporhyn ·1 1.- rufo.-, "' , ialia siali:. 



58 The F ood R elatio111:1 of the Ual'abidw and Ooccinellidce. 

20. A on c1eru , sp.: 2 Tur lu · mi ratorius, l Mi mus carolinensis, 
1 Harp rbyncus rufu · 1 iaJia sialis. 

21. A. pallipes: 2 Tur<lu · rnig ra.toriu , 1 Mimus carolinensis, 1 
Harporhyn ·u rufu · :3 ia.Jia si lis. 

22 . partiarius : 1 Turdu · miara.torius. . n 

23. nisodactylu , :f·: 1 Turdu · must linu , 1 1'. swainsoni, 2 T. 
migratoriu ·, ,., H arporhy11 us rufos, (3 ialia sialis. 

24. A. rusticus: 1 ialia ialis. 
25. . <liscoi<leus : 1 Tunlus pallasi. 
26. A. baltirnorensi : 2 Tunlus rnigratorius, 1 Mimus carolin n-

sis, 1 Sialia sialis. · 
27. Xestonotus lugubris : 1 Turdu s mu telinus, 1 Sialia siali s. 
28. Harpalus, sp.: 1 Turdus mu ·t linus, 7 T. migratorius, 4 Mirnus 

carolinensis, 8 H arporhyncus rufus, 6 ialia sialis. 
29. H. herbivagus: 1 Turdus mig ratorius. 
30. H. pennsylvanicus: ] Mirnus c:arolin n ·1s. 
31. tenolophus, sp.: 2 Turdu · palla i. 

In the followiu o- tables, the •lements of the food, arranged in 
systematic order, are placed at the left of each page, while a ver­
tical column of the table is assig ned to each genus of beetle. 
The upper figure of each couple indi cates the number of speci­
mens in which the given element was found, while the lower fig­
ures ( decimal) show the ratio of the element to the entire food of 
of all the xamples of the g enus. The dagger has been used 
to indicate a trace too small to figure in the percentages, 
usually less than one-half per cent., ,and an asterisk 
denotes that tbe element against which it is placed was present in 
the food, but that the ratio was not estimated. 



The Food Relations of the arabidce and CocC'inellidCP. 59 

ARABID.1K- MI CELLANEOOS LLECTION . 

en rn 
::I 

::I rn >, ..c: ::I ad ..., ;..; ..., 
d Q) c) ·;;; 
o;l 'O o;l 
c) 0 'O <IS 

<IS 0 0 ..c: _., 
0 "' p. 

<IS b.O ·~ 8 ;..; 
:.) <l < <l 

""o . of pecimens ..... . 1 4 19 4 

NUMBER OF SPECIMENS AND RAT IOS IN WBHJH EAC H ELEMENT OF FOOD 
W AS FOUND. KINDS OF FOOD. 

----------
1 21422 1 5 1 61 2 1 5 1 3 11 1 141 43 

ANIMALFOOD .... . 1.001.001.00 SE.44 65.17 .90.73.221.00.75 ... 01 .24 .031.00 .07 .33.42 
1 4 1 2 1 5 j 4 5 2 1 1 1 2 1 30 

I. INSECTA ... . . . ... .... . .. . . 50 ..... 8 11 .~5 .17 · i l 43 ...... ·f ... .. .. 03 031.00 .03 .33 -~6 

Larvai...... . ... . ...... . .. . .. .. .. .. .. . .32 .... 10 ........... OE .. . .. .... . . . ....... . .. . 02 

1. Hymenopte1·a....... . . . . .. . .. .. .. . .is .. . .. . .. ..... .. . ....... .. . . ~1 1 .... ~1 

Formicidre . . . . . . . . . . . . . . . . ... .... . .. . : . . . . . . . . . . . . . . . . . . . . . . . . . ..... .. ..... 01 .. . + 
3 1 1 1 1 1 1 8 

2. Lepidoptera (l arvrej . ...... . ...... 63 .... 33 .. . 20 ... .17 .... .. ....... . 02 .30 .0 
1 1 

3 . .Diptera.... .. .. . . . .. . . . .. .. .. · .. · .. · .. · .. · · · 2 · .. .. .. · .. · + .. · t 
4. Ooleoptera ...... .... ............. .. .. ....... . 39 .............. . .. . .. .. ..... . . . . ... 02 

1 1 

:::.:~,~~ : : : : -• • • • • • • : • • •:: , kl •: • --· 1 : • • - , : • • • •:: : •: , i: 
~i.H1~fJ;-:rb~phidef ..... ... . .. .... ..... .. 17 .. ;·i· .i ... .. . .. . i'J ... .. .... . 2 .. 03 -~1 

(Mites) .. . \ ..... .. . . .. ... 1.01 .22. i - . .. . 01 .......... 01 . ... 01 

III. vrr:!tricus) .. ! .. .. .. .. . . . . . .. . .. .. . .. . .. . . . .. . 6 ... ... I.. . .. .. .... . 61 
2 3 2 2 1 4 4 2 1 4 15 4 12 2 58 

VEGETABLE FOOD . .. . . .. .. .... 12 .56 351.83 .10 27 .78 .. 251.00 .99 .76 .97 .... 93 .67 5 

seeds ........... . .. . .. .. .... ........ . ... 
1 

. .. 1
2 1

1 ..... ' . .. J ; ' .i6.it· ·6s .~o .~o 

1. Eroo(Jens ..... ........ .. . .. . .. ... 1.50
1
. .i4 25 .. :. is ... I ... ' .. . ~ .... .. i7 .. . . ~7 

~::;~·~~~~·~;~;;~~;: :: :::: :::: ::: ; :: : .::1::: :::1 ·:: + .. . ... !:::1::: :: :1.:: .f1 ::: i 
Ambrosia. ... .. ...... . ... ... .. ·1 ·i ·' ·2 · .i . ·i f . . i .. ).i . . 7 I !7 .i . -g3 
2. Endo(Jens .. . ........ . ......... 10 .11 .35 30 .10 ... 28 ' .25

1 

.34 .. ·1· ... . 23 .10 .17 
. . I 1 1 ~- ! 1 1 7 4 1 1 

G1ammere .......... . . ... .. ......... 10 .11 .3:> .30 .. . . ... 28 .......... . 25 .34 ........ 23 .10 .17 
1 ; l j 1 4 

Seeds .. . ........ ...... ................ . ... . .. 1. 30 . .. . . . ......... 
1

.. .05 . .. . .. + ... . 02 

:::~:m ;,~;;,,) : • • -•: • _____ : :, ;, ::: •• • -•• J --- ----;, :i __ _ _ __ I: 
2 2 

. .. . 01 t 
1 2 

3. H epaUcre .. . . . . . . . . . . . . . . . . . . . . . . . . . . .. ... . .. .. 
1 I 

t . .. ... .. .. + 
1 1 

4 . .A l(JaJ ............. . .. .... ... .. .. . .. . .. . + 
t .. .. . . ... . .. ... . + 
1 I 1 2 3 5 1 23 
141.00 .11 .04 ..... .. 13 .07 .09 
1 1 

Protococcus ...... .......... ... ..... i . . 
3 
. .. . . i . . i . 

3 
. i .... . 

5. Fwngi ..... .. .. ... ......... ... .. t .45 ... 03 + .07 .21 .. . 

.14 .... .... .. . ............. 01 
1 

Coprinus . .. ....... . .... ...... . . . ..... . 
2 

Phoma ... ... .......... .. . .. ... ....... .45.. . . . . . . . . . . . . . . . . . . . . . . . . .. . ..... . .... .. . ... 02 
1 1 

... . .. . .. .. . ... . .... .. ..... . .. t . .. . .. t 
1 , 1 

. ...... t .. ....... . . .. .... . ..... t 
2 1 1 2 6 

Helminthosporium . ... ........... . ... t ..... . . .. t .. . ....... 1.00 . .. . .. . ... .. . t ... . 01 
1 1 2 

C-Oleosporium . ........ ..... .. . 

Dematiei. ... .... ... .. .. .. .... . 

Cladosporium. .... . .. . . . . . . . . . . .. . . . . .. . .. . . . . . . . .. . . . . .. ..... 01 .01 .. . . . . . . . .. t 
1 1 1 3 

::::~:::r~::: ::::::: :::: :::. ·::: ::: ::: ::: ::: ::: ::: :1~ :::· ::: ::: : j~ -~.1: :: :::: :~~ ::: :A: 



60 The Food R elations of the Carabidce and Coccinellidce. 

CARABID.iE AND CANKER -WORM', 

KINDS OF FOOD. NUMBER OF SPECIMENS AND RATIOS IN WHICH EACH ELEMENT 
OF FOOD WAS FOUND. 

10 2 I 4 13 3 1 2 3 1 Hi 1 1 2 59 
ANIMAL FOOD .. ... 1.00 1.00 1.00 .94 .681.00 1.00 .80 .. 1.00 .92 .09 .25 .35 .77 

I. MOLLUSCA ........ 
5 2 13 3 2 1 

II. INSECT A ...... .. ... .50 .. .. .50 .85 .68 .... 1.00 .13 .. .. 

Larvre . . .... ... ... .... . 

1. Hymenopte1·a (ants) 

2. Lepidoptera (larvre) .. . 

Anisopteryx v ernata . 

Noctuidre . .. .. .... ... .. 

3 . .Diptera . .. . ... .... .. 

Culicidre . .. .. . .. ... .. . 

4. Ooleoptera . ..... .. . . 

Telephoridre .... . .. ... . 

III. A RACHNIDA ... .. . 

Spiders ...... . . ........ .. . 

Mites . .. . . .. . .. . .... . . 
IV. MYRIAPODA 1 

(Geophilus) .. I 
v. VERMES (Lum-1 

bric us) .. . . . . I 

2 
.11 

8 1 
.56 .33 .. .. 
7 1 

. 52 .33 . . .. 
1 

.01 . .. ... . 
3 1 

... . . 07 .02 

3 
.... 05 ... 

1 
.05 ... 
2 
t 

2 1 

1 
.05 .... 
2 1 

.\:10 .11 .. .. 
2 

.90 .... 

1 
.05 .... 
1 

.05 .... 

1 1 

1 1 
1.00 .... . .. .02 

12 1 1 2 42 
.64 .09 .25 .35 .50 
1 3 
.06.... .04 

8 
.41 .... 
5 

.26 .. .. 
2 

.12 .... 
1 1 

.01 .09 .... 

1 
.06 .. .. 
1 

.06 .... 
1 

.02 .. .. 

1 
.02 .. .. 
1 

.01 .... 
4 

10 1 

1 
+ 1 21 

.83 .26 
1 16 

.33 .21 

2 

3 
.03 
7 

.03 
1 
+ 
1 

.01 
1 

.01 
4 

.01 
1 

.01 
3 
+ 
1 
+ 
4 

.06 
23 

VEGETABLE FOOD .... .. .. , .06 .32 ... .20 1.00 .. .. 

.25 
5 

.08 .91 .75 
3 

.65 .23 

Seeds ... .. .. . ... ... . . .. . 

1. E mogens .......... . .. 

2. Endogens . . . . . .. .. . . 

Graminere . .. . .. . ... ... . . 

Seeds . . .... . . .. .. ... . . 

3. Fungi .... .. .. .... .. 

Peronospora . . . . . . . . . . . 

Ascomycetes . . . . ... . 

2 
.06 . 

1 
.20 .... 

1 
t 

.20 .. .. 
1 

.10 .... 
1 5 
.01 .50 .... 

4 

2 

.40 .. . 
4 

.30 ... 

·04 ... 

1 
.02 

1 5 
.32 .04 
1 4 

.17 .03 
1 1·0 
t .08 

4 
.06 
4 
.04 

1 4 
.16 .02 
1 1 

.16 .01 
1 
t 



The Food R elations of the Carabidce and oc inellidce. 1 

ARABTD,K A ' D 'ABBAGE WORM ND 'Hl 'H -H ;s, 

en 

II en ,;, = = h ~ .c = ;::; en ;.... .., 
en i,:.. <.) a., <.) a., 

'..3 = 'O Cl:! = ;.... 
'O ILJ ~ 0 'O cd ~ 
0 0 ~ 

s:: A 0 0. s A ;.... ~ -~ a., e 0 ;.... 8 0 .., :;:; bl) s:: cd = bl) 
~ <1 C,) <1 <1 p:1 (/) <1 ---- -

Number of pecimen Examined .. 3 1 1 1 2 4 12 10 

KI ' D OF FOOD . NUi\[BER OF SPECIMENS AND RATIOS l N wnicm EACU 

ANIMA.L FOOD .. .. ............ .. 

I. IN ECTA .... . . .. ...... .... . .. .. .. . 

L n.rvre . .... . . .... ........... . .. . ... . 

1. Hymenoptera . . . . . . .. . .......... . 

Lasius flavus .. .. ......... .. ... .. 
1 

2. Lepidoptera . ... . ................. . .33 

Caterpillar . .. . . ..... . .. .......... . 
1 

Agrotis annex a . ... . ... ..... ... . .33 

3 . .Diptera .......... ....... . .... . ... .. 

4. Coleoptu·a (larvre) . ... .. ...... . . . 

5. Uemiptera . .... . ..... . ... .. . ..... . 

Aphide ..... . .. ... . ..... . ..... . .. · 

Chinch-bugs ... .... .. . ... .......... . . 
1 

VEGETABLE FOOD ...... . . .... .. .33 

ecc1 . ..... ...... . ................. . . 

1. E wogens . . ..... . . . . .. . ....... . .... . 
1 

2. E ndogens . ........ ... . ...... .... . . .33 
1 

Grass .. . ... .. .. .... . . . . .. . . .. . ... ... . .33 

3. Fungi ... .... .. . . . .......... .. .... . 

Ilelminthosporium .......... . .. . . . . 

Peronospora . . . .. .. ...... . .... . 

ELEMENT OF l•' OOD WAS 1'' 00ND. 
1 1 1 1 i--7 
.50 1.00 I .. .. • .50 .10 .41 

1 1 4 
1 00 .10 .20 

1 2 
.60 .13 
1 1 
.60 .05 

1 
.. .. ·-- . .... .0 

1 1 
.10 .03 

1 1 
.30 .03 

1 1 1 1 4 9 
.50 t 1.00 .50 .90 .59 
1 1 4 6 
* * .90 .30 

1 1 1 4 
* t t .0 
1 1 3 
* t .0 

.51 
3 

.36 
1 

.02 
1 

.05 
1 
.05 

5 
.20 
2 

.0 
4 

.21 
7 

.40 

2 
.13 
2 

.11 

2 
.02 
1 

.01 
1 
.01 



62 The Food R elations of the Carabidce and Coccinellidce. 

No. of Specimens . .... 

KINDS OF FOOD. 

CARABIDlE. GENERAL TABLE. 

u;; u;; 
u;; ,,; ::, u;; ::, en ::, ,,; en ::, .d ::, ~ -~ E ~ .d ;::l ~ .d """ r... """ ,,; t,'.. ,,; ,,; ::, -~ en ~ Q) C) 

~ ,,; 
E C) ,,; ::, ~ N ::, ::, ;.., ,,; ::, o:l 'O cs! rn ::, ::, ;.. 

c;! Q) ~ """ Q) 
~ .d """ ci ::, ·a 0 'O o:l C) .; .0 

o;I 
0 .§ ;:: .d ·;::: p. .d """ 

rn ;.. C) 
0 .d ~ s rn C) 0 """ ~ r... 0 c;! i o:l ~ p. 'O p. 0 

0 ;.. Q) o:l """ o:l r... 8:l """ 0 rn ;.., r... s cd "' o:l cs! cd ~ .; o:l Q) s C) :a o:l bl) ·a s c,j 
c,j """ 0:: C) ;.., 0 p::; I> """ A ;.. ;.., cs! ::, u p.. UJ P=l C!, ~ (.) ~ p.. ~ Q Q < < ~ ~ ~ p.. UJ 

11 2 6 1 17 3 ti 3 7 13 6 3 23 1 15 31 5 1 19 2 175 

NUMBER OF SPECIMENS AND RATIOS IN WHICH EACH ELEMENT OF FOOD 
WAS FOUND. 

11 2 6 1 17 2 5 2 7 12 2 3 22 9 5 2 1 7 1 117 
ANIMAL FOOD ..... 1.001.001.001 .00 .93 .44 .67 .45 .93 .73 .231.00 .88 .... . 34 .21 .07 1 .00 .12 .33 .67 

1 1 
I. MOLLUSCA (slugs). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .33 . .. . . . .. ..... . .... . .... . . .. 01 

5 3 17 1 5 1 7 7 18 6 3 2 1 5 1 82 
II. INSECTA .. .. . ....... .45 ..... 50 . .... 86 .11 .67 .12 .93 .34 . . .... . . 67 ..... 24 .05 .071.00 .09 .33 .36 

2 1 1 2 1 7 
Larvre ... . .... .. ..... .. . . ... .. .. ... ... . . 09 .... 16 .. . . 07 . .. . .. . .. . . 06 . .. .. 02 . ... .. . .. . . . . .. . . 03 

1 1 1 1 4 
1. Hymenoptera . ..... . . . . . .. .. .. . . ... . .. . . . . . .. .. .. 02 .07 ... .. . . ... ... . 03 . ..... . .... 01 .. . . 01 

1 1 3 
Formicidre . ....... .. . . ... . . .. . .. . . . ... .. 03 . .... . . .... 01 . .. t 

1 1 
L asius flavus ... . . . . . . . . . . . . . ...... .. . ... . . . . . 03 . . . .... .... .... . t 

11 2 3 2 10 2 1 31 
2. Lepidoptera ... . .. .. . .... •. • . . . . ... . 58 .... 33 . .. .40 .10 .. . . . ... 36 . . . . .... ... ... . ... 06 .30 .16 

5 1 1 1 3 1 1 13 
Larvre . . . ......... .. .. . .. ... .... . .. . ... . 18 . ... 17 .... 14 .03 .... .... 05 ....... . . . . ...... 01.30 .04 

7 1 2 4 1 16 
Anisopteryx vernata .. .. .. ... ...... . . . 39 . ... 16 ... . 26 . . . . .. .. . . . 18 ..... ... . .. . .. .. .. 05 . ... 08 

1 2 4 
Noctuidre (larvre). .... . . . . . . . . . . . . . . . . 01 .... .. . .. ... .... . . . . . . . 09 . ... ... ... .. . . ..... ..... 02 

1 1 
Agro tis annexa . . . .. . . · · · · . . . . . . . . . . . . .. . 07 .. .. . .. .. . ...... . . .. . . ....... .... . 01 

3 1 1 1 1 1 9 
3. J)iptera .... . ..... .. . · · · · . . .. . .... 05 .... 01 . ... 01 . .. . ...... t . .... .. . 03 . . ...... 02 .. .. 01 

1 1 
Culicidre . .. .. . . .. .. ... . .. . . . . ...... .. 01 ... .. . .. . .... .. .. .... . . . . . . .. . . ..... t 

2 2 4 
4. Ooleoptera . . ... .. . . . .. .. . . . ......... . .. .. .. . . . ... 28 ... . .. ... .. 06 . ..... .. .. .. .. ... . . .... . 02 

1 1 2 
Larvre . ..... . . . . . ... . . . . . .. ..... . .. . .. . . .. .. . . . .. .. . 14 . .. . . . ... . . 01 . . ... ..... . . ...... .. . ... 01 

1 1 
Scarabreidre . . . . . . . . . . . . . . . . . . . . . . . . . . . . .... . . . . ... 14 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . t 

1 1 
Telephoridre .. ... .. ....... ...... ... ... . . . .... . . . .. . .. . ...... . ... 05 . ..... . ....... . . . . ... ... 01 

1 5 1 7 
5. Herniptera . . . ..... .. .. .. . . ... . ............... . 12 . . . .. .. . ......... . . . . 19 ...... .... . .. . 03 .02 

1 2 1 4 
Aphides . . ........ . . . ....... ... . . ... . . .. .. . . . ... . 12 . ..... ..... .. .. .. ... . 05 . .. ...... . . . . . 03 .01 

4 4 
Chinch-bugs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .14 ... . .. ........ ... 01 

3 11 1 1 2 9 
III. ARACHNIDA .... . .. .. . .. ... ... .. . ... 04 . ...... ... ... 01 .15 ..... 01 . . . . t . .... ...... 01 . .. . 01 

1 1 
Spiders . . . . .. . .. .. . ..... . ........... .. .. 04 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . t 
Mites .. . ........ ..... . . 
IV. MYRIAPODA l 

(Geophilus) . . j 
V. VERMES (Lum-i 

bricus) . . 

2 11 1 1 2 8 
. . . . . . . . . . . . . . . t ... .... .... .. 01 .15 ... . . 01 . . . . t ..... .. .. .. 01 .... 01 

1 1 
... ... . .. .... .... . . . .. ... . .... .. .. , .... . .. 01 . . . . ... . .. . . . . .. . . .... . t 

5 5 
.. ...... .. .. ........ . ... . .. . . .. . .. .. . . ... . 19 .. .. . . . ... . . .. . ...... ... 02 

4 3 3 2 1 6 5 8 1 12 26 5 18 2 97 
VEGETABLE FOOD . ... .... . ... ..... 07 .56 .33 .55 .07 .27 .77 ..... 121.00 .56 .79 .93 . .. . . 88 .67 .43 

2 2 7 1 6 1 19 
Seeds.............. . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * ... .. .... ... 13 .16 .18 .. . .. 29 .50 .08 

2 1 2 1 1 2 1 4 14 
1. Ea,ooens . . . ... .. .... . .... . . . . ... .. .... 04 ... .. .. 33 ... . 08 .16 ..... 02 ..... 09 .03 .... . . .. 14 .... 05 

1 1 
* * Seeds .. . .... . ... .. ... . . . . . .. . .. . .... .... ... . .. . .. . . . . .. ::'.:==================== 



The Pood E el,ation of the Cambiclce and occinellidce. 63 

No. of pe imen .... 

KINDS OF FOOD . 

.ARABID.l~. 'E B RAL TABLE- onclud d. 

NOlIBER OF SPE AND RATIOS IN WlllCl:I EACH ELEMENT OF FOOD 
WAS F OUND . 

-.,----,--~--,---,---,--- - - I I 3 I 3 
ompo it;_e '. pollen)... . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . .. . . . . . . . . . . . .. .. . . i\ · .. g1 

Ambrosia .. . ...... ········ ... . ···· ·i · ·i · ·2· ·i · ·i' ·2· ·i · · · 4-1-· ··, ·s· i2 ·· · ··· ·i\ i · · ~~ 
2. E ndouen ..... . ... .. .. ....... . ..... 02 .11 .17 .20 .07 .12 * .01 . .. .. 14 .37 ..... 14 .10 .13 

1 1 2 1 1 i 1 I 2 36 5 I 1 26 
Gra.minero . ... .. ... ........... . . ... . . . . 02 .11 .17 .20 .... 08 * . ... 10 .34 . . .... 14 .10 .11 

1 1 5 1 S 
eect .... ... ... ... . .... . . .. .... . . . . ........... 20 . . . . . . * ........ ...... . 13 ... . .. . + ... . 03 

1 2 4 
Pollen ... . ....... . ...... .. ......... . . 11 .17 . ...... .. . .... 1 ...... . .... . .. . ..... . .. . ... 01 

Phleum (seeds). .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . / . ..... .. 67 .i3 .......... .... 83 
2 2 

3. Depaticce . .... . ... . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + ............ . 1" 
1 1 2 

4. A l(lm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . t . . . . . . . . . t 1 · . . . . . . . . . . . . . . . . . . . . . . 1" 

Protococcn . ..... . ................. . .. i .:.s'.. 1 1 .a' ·q· ... t .. i . . 4. ·3· .. . ..... 6 .. i t 
5. Fun(li ... .... ... . .. .. . . . . . . . . . . . . . . . . + 

1

.45 ... . 02 + .04 .1 4 .... ~61.00 .04 .03 . . . ..... 11 .07 .~5 

Coprinus .. ..... . ........ . .. . .. .... . . .................. . . . ...... . 04 ......... . . . ... ........ t 
2 1 

Pboma . .. . .. .... . ......... ... ... . . .. . . .. .45 ..... ... .... . .... . .. . ... . .... . .... . ........... 01 
I I i i 

Coleosporium .. . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . .. .. . . . . . . t 

Dematiei. ... . ... .. ...... .. ...... . ... .. .... 2. , . • • • ., .. • • • • • • · ···.I t .. i · ·i · · · · · · · · · · · ·
2
· · · · l 

Helminthosporium ........ . . . ...... . . . .. + . .. . . . . .. . . . . . . . . . . 1. 00 .~1 .i .. .. .... + 
1 

. .. . g1 

Cladosporium... . ..... . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . ......... .. ... . + .01 .. .. . .. . . .... t 
1 1 1 2 5 

Peronospora. . . ............ .. . . ... .. . . . .. . ...... . . . ...... 12 . . . . . . . . . . . + + .. . .. .. . 05 . ... 01 
1 1 2 

:a::::~~e~::: : ::::: :: : ::: : : :· : ::: :: : : ::: ::: ::: ::: ::: .~. ::: :::: 1:0\::: j~ ::: ::: ::: : ::: :: : f. 

• 



• 
64 The Food Relations of the Cambidce and CocG'inellidce. 

co INELLIDJE. 

Misce llaneous. Chioch-lln ~·s. General. 

~ d 
d 

.d .d ..., 
ad .~ ..., 

A ·s A 
8 d d 

~ ~ ad >, 8 ~ ad c! p:, ...... c,) c,) 
0:: a) d ;... d ...... 'O ;... d a) 'O <s! ;... 
~ h d "O C1) C1) 

i:3 "O C1) h d 
0 A .d 8 0 A A s 0 A A .d a 0.. ·c3 c,) 0.. ·c3 0 0.. ·c3 .9 0.. d ·r ...... 

8 0.. 
c,) 

8 c,) c,) c,) c,) c,) d s 0 ;... p ·~ 0 h ~ ~ 0 h ;... A 0 P=l : : u 0 Cl) ::.> 0 P=l Cl) 

Number of Specimen · · ····· 15 5 1 1 3 18 29 (i 3 1 39 

KINDS OF FOOD. 
NUMBElt OF SPEUIMENS ANO RATIOS IN WHICH EACH 

ELEMENT OF FOOD WAS FOUND . 

12 5 17 12 2 14 24 5 2 31 
ANIMAL FOOD . ........... .44 .64 . . . . .47 .25 . . . .32 .25 .35 .53 .32 . ... .37 

11 5 16 12 2 14 23 5 2 30 
I. INSECT A ............ . ...... .39 .64 . ... .43 .25 . .. . .32 .25 .32 .53 .32 . . . . .35 

1 1 1 1 2 2 
1. Lepidoptera ... ... .. . ...... t .... .... t .01 .. .. . .. . .01 .01 . ... .... . . . . .01 

1 1 1 1 
Larvre ........ ....... ......... .. . . . .. . . ... . .. . .Ot . . . . .. . . .01 .01 . .. . . . . . .01 

8 5 13 7 2 9 15 5 2 22 
2 Hemiptera .. .. . ······ .... .29 .64 .... .36 .21 . ... .32 .22 .25 .53 .32 . ... .29 

8 5 13 4 2 6 12 5 2 19 
Aphides ................ . ..... .29 .64 . . . . .36 .11 . ... .32 .14 .20 .53 .32 . ... .26 

2 1 3 2 1 3 
Siphonophora granarire ..... . .. . .... ... . .. . .05 . .. . .24 .08 .02 . .. . .24 . ... .03 

4 4 4 4 
Chinch-llugs ..... .. ........ .. .... . ... . . . . . ... .10 . .. . ... .08 .05 . . .. . . .. .... .03 

1 1 1 1 
3 Neuroptera (Podura) . . .. . . .01 ... . .... .01 . . . . . .. . . . . . ' .. . t . . .. . . . . . ... t 

4 4 2 2 6 6 
I I. ARAC HNIDA (Mites) . .. .. .01 ... . .. . . .01 t . .. . .... t .01 . . .. ... . .. .. t 

1 1 1 1 
I II. M Y RIAPODA (Geopbilus) .04 . . .. .03 . .. . . . . ... . .02 . . . . . ... . .. . .02 

13 2 1 16 14 1 3 18 27 3 3 1 34 
VEGETABLE FOOD .. . .. .. .56 .86 1.00 .53 .75 1.00 .68 .75 .65 .47 .68 1.00 .63 

1 1 2 1 1 2 
Pollen . . .. .......... .... .. ... . .03 t .... .02 .. . . ... . ... .01 -~ . ... . ... .01 

7 7 13 3 16 20 3 23 
1 Emo(Jens (Pollen ) . .. . .... .13 .... . .. . .09 .07 '. , . .47 .13 .10 . .. .47 . ... .11 

6 6 13 3 16 19 3 22 
Compositro ....... .. .. . . . .. .. .13 . . . . ... . .09 .07 .... .47 .13 .10 . ... .47 . .. . .11 

1 1 1 1 
Taraxacum .. .. .. .. .. . . .07 . .. . .... .05 . . .. . ... . .. . . ... .04 . .. . .... .. .. .03 

2 2 2 2 
Ambros ia .. . .. .... ... .... .... .... .... ... . * . .. . . .. . t * . . . . * . . .. ... . 

1 1 1 1 2 2 
Polygonum ...... ... .. ...... t .. .. .. . t * . .. . .. .. t t . .. . * .... . ... 

1 1 1 1 
Coniferre . ... . . .. ......... . ... t .. .. ... . t . .. . . .. .. . . . .. . t . . . . .. .. . ... t 

1 1 1 1 
2. Endo(Jens ...... . . .. .... .. . . .05 . . .. . ... .04 . .. . . ... . ... .... .03 . ... . ... . . . . .02 

1 1 1 1 
Graminere .......... .. .. . . . .. . .05 .... . . .. .04 . . . . . . . . ... . . . .. .03 . ... ... . . ... .02 

1 1 1 1 
Pollen . . .. .. . . . ... . . . .. ... .05 . . . . . . . . .04 . .. . . ... . ... . ... .03 . ... ... . . ... .02 

4 1 5 11 2 13 15 1 2 18 
3. Lichenes .. . .. .. . ..... .. .... .02 .04 . ... .02 .06 . ... .03 .05 .04 .03 .03 . ... .04 

1 1 6 1 7 7 1 8 
P hyscia . ................. . . .. .01 . . .. .. .. t * . ... t t .01 . ... t . .. .01 

11 2 1 14 14 1 2 17 25 3 2 1 31 
4. Fun(Ji .. .... .. .. ..... .. . . ... .33 .32 1.00 .36 .61 1.00 .18 .56 .46 .44 .18 1.00 .45 

1 1 1 1 
Myxogastres . ......... . .. . ... * . . .. . . . . t . .. . .... . . . . . ... t . . . . .. . . ... . * 1 1 1 1 
Sphroronemei .. . . . . . ......... * . .. . . ... .01 .. . . .. . . .. * . . . . . .. . .... * 1 1 7 1 8 8 1 9 
Septoria . ..................... "' .... .. .. .01 * . ... * * * . . . . * . . . . * 

1 2 3 2 2 3 2 5 
Ustilago ......... ..... .. .... .. * .08 . . . . .02 * . ... ... . * * .07 .... . ... * 1 1 1 1 
Uredo ... .. .. . .... . ..... .. ... . . ... . . .. * . . . . .. . . * t ... . . .. . . .. '" . ... . . .. 

6 1 1 8 11 1 12 17 1 1 1 20 
Helminthosporium . ..... ... . * .10 1.00 .17 * . ... * * . .08 * 1.00 * 

3 3 7 1 8 10 11 
Macrosporium .. . ..... . ...... * .... .04 * . . .. * ,/I * . ... .. .. .... * .... 

3 3 13 2 15 16 18 
Cladosporium ... ..... . ... .. .. * ... . .02 "' . .. . * * * . . .. .... . ... * .... 

3 3 7 7 10 10 
Peronospora .. .. .............. * .... .... .01 * . ... .. . . * * .. .. . ... .. .. * 

1 1 1 1 
Menispora ........... .. . . ... . . * ... . . ... t .... .... . .. . ... . * .... . . .. . ... * 



ERRATA. 

Page 6, line 12 from bottom; page 8, line 15; page 11, line 2; 
for Cyclniclce, read Pentatomidce._ 

Page 17, line 9, before Vireo; omit ancl. 

Page 23, above ARACHNIDA, for Cyclnidce, read P entatomiclce. 

Pages 25 and 27, above Orthoptera, for Cyclniclce, read P enta-
tomidce. 

Page 28, lines 2 and 8, for Graphorhinus vados1ts, read Epi-
cce1·us imbrricatus. · 

Page 64, under Hemiptera, for Siplwnophora granarice, read 
Aphis maidis. 

Page 69, line 5 from bottom, for jresh-wate1·, read local. 

Page 78, line 1, after all, insert the. 

Page 82, line 7, for cha1·acte1·, read charctcte1·s. 

Page 91, line 5, for consisted, read consists. 

Page 92, line 2 from bottom, for more, read most. 

Page 97, line 11, for f01·y-six , read f01·ty-six . 

Page 99, line 2, for witn, read with. 

Page 101, lines 12 and 13 from bottom, for stritcture, read 
structitres. 

Page 105, line 23, for aeration, read ae1·cttion .