FOREWORD

"I thank Dr. Derek Sikes for his help in obtaining this translation of Pukowski's
classic 1933 paper on Nicrophorus.

The text is thorough and very detailed and a link to a succinct summary
is to be found in the Table of Contents labelled 'SUMMARY'."

A Dale,
http://www.bugsandweeds.co.uk

ENGLISH TRANSLATION OF PUKOWSKI'S 1933 NICROPHORUS PAPER

The following was translated from the German by Janet Christie and originally posted
by Derek Sikes. Refer to the original document for literature cited, tables and figures,
which are not included here. This reproduction is not to be used for any purpose other
than private study, scholarship, or research.
Ecological Investigation of Necrophorus F.

by

Erna Pukowski

Zeitschrift fur Morphologie und Oekologie

der Tiere 27(3): 518-586. 1933.

TABLE OF CONTENTS

INTRODUCTION

RESULTS

A. General Life of the Adult

1. Occurrence -- frequency
2. Environment
3. Temporal occurence
4. Nutrition

B. Reproductive Biology

1. Relationship of beetles to one another

a. Attraction of females
b. The isolation of pairs from a large number of
individuals end the resulting conclusion
c. Ecology of the fight
d. Fight with strange species or races of
gravedigger
e. Course of the fight
f. Relations between partners of the pairs

2. Relations of the beetles to their offspring

a. Brood care
b. Burying of a body
c. Rounding of the body and completion of the
crypt
d. Egglaying
e. Moisture and digestion of the body
f. Male participation in brood care
g. The brood at the time of larval development
h. Feeding of the larvae
i. Ecology of feeding
j. Defense of the larvae and their food
k. Maintenance of the crypt
l. Defense of males at the nursing
m. Development of full-grown larvae to young beetles

SUMMARY
INTRODUCTION

The striking instinct of gravediggers (Necrophorus fabricuis) to bury small animals was first
reported by Gleditsch (1752) with the help of experiments, in which development of beetles
with different species of small animal corpses at their disposal was observed. Gleditsch
succeeded in putting the burywork of the gravediggers in relation to their reproduction. On
the suggestion of these chemists Melim from Bremer (1755) who independent of the
mentioned report, followed the burial of a mole by the gravedigger and recognized in them
the remarkable broodcare, which also occupied A. I. Rosel von Rosenhof (1761) with the
remarkable life of the gravedigger. We find his result put down in the celebrated "In
sektenbelustigung" equipped with coloured pictures of adults, larvae and pupae.

From the statement of Gleditsch (1752) that a gravedigger had brought a stick to undermining
to collapse was fed on the body of a toad, Lacordaire concluded about the intelligence of this
beetle. First the French researcher J. H. Fabre destroyed the fable of the intelligence of the
gravedigger while he showed with the help of numerous carefully carried-out experiments
that the actions of this animal are fixed within the setting of instinct, but not as was first
interpreted a reasoned thought.

If the experiments of Fabre lead to greater clarity so too a new problem escaped his
observations that researchers had perceived as such, but not been able to solve. This new
problem lies in the fact that on the buried carrion there always remains a single pair of
gravediggers, highly suitably, even if more beetles worked at it. The question of how this
appearance is to be explained was discussed recently in terms of animal psychology and
sociology (Renter 1913, Schroder 1929) however until now no satisfactory solution was
found. On the suggestion of my highly revered lecturer, Herr Geheimen Regierungsratses
(private administrator adviser) Prof. Dr. zur Strarsen, I put the solution of named problem to
the test.

Fulfillment of the task required most precise knowledge of the whole digging event within
which the isolation of pairs from a larger number of individuals must take place; this could
only be achieved through extensive experiments preferably under natural conditions. For this
reason I have investigated in depth the general ecology of the genus Necrophorus.

The experiments were carried out on the following species: Necrophorus germanicus L.,
humator B1., vesplllo L., vespilloides Herbst, investigator Zett, fossor Er. (interruptus Steph.)
The species germanicus, was sent to me mostly from Danzig; I baited the other animals
around Frankfurt a.M.

RESULTS

A. General Life of the Adult

1. Occurrence - frequency.

Although around Frankfurt not all species of the genus Necrophorus occur, humator, vespillo,
vespilloides, fossor and investigator still occur with certainty. The first three species are found
in large quantity, and investigator and fossor in smaller quantity. N. germanicus is scarce and
N. vestigator even more so.

In order to catch the gravediggers, I arranged measures of bait in the following state: a pail,
whose base was perforated for the draining of rainwater filled to 10-15 cm with earth. Inside a
slightly decaying large piece of meat was laid and the entire pail was put so that it is not
exposed to direct sunbeams, yet still breezy enough for the smell of the carrion to flow about
unhindered. Beetles flying by are easily able to attain the body; but there they are caught for
the narrow limit of space left them is too little for flying away and they are not able to
clamber up the smooth and steep walls of the pail. Besides few make attempts to fly for there
they find rich booty and the soil layer offers them the possibility of shelter. Every 10-14 days
the traps were baited with inew meat, and were able to be used continuously in this way.

In order to give a rough impression of the frequency of the gravediggers, I have included 3
tables which give again the total yields of the baits. The catch was made after the meat had
layed out for 3-4 days in warm weather. On baits M.S., M.E. and M.W. named beetle species
in the following numbers were found. From this set up it is obvious that the gravediggers are
by no means as rare as one accepted from chance finding.

2. Environment

The searching of numerous baits gave me the opportunity to study the occurrence of frequent
species (humator, vespillo, vespilloides) around Frankfurt a.M. with regard to their
environment and to confirm the winning conception with a systematically employed series of
experiments. The traps were put out in different areas of the nearer and farther surroundings
of Frankfurt, and the caught animals registered with time, numbers and species. Data in Table
1 are from a large series of arbitrary catches. The baits are labelled with the numbers I-VI. All
catches given here were made at the same time of year, namely mid Mav 1931. M (bait) I and
M II were in Tannus near Eppstein. Distant mixed woodland (Pinus and Fagus) is the near
and far environment. M III and M IV were in meadows at Ginnheim and Rodelheim in
Niddatal which stretches to the foot of Tannus. M V stood in the middle of a large, very dry
confer wood area in Frankfurter Stadtwald. M VI lay in mixed woodland (Pinus, Picea,
Fagus) in Walldorf and close to the woodedge not far from extensive meadowland. So M VI
joined the environmental conditions of previously chosen places.

From Table I in mixed woodland from Eppstein at M I and II altogether 41 individuals of

humator and 36 of vespilloides were found, but not a single example of vespillo. Forty-four
examples of N. vespillo were found from meadows at Ginnheim and Rodelheim in M III and
M IV. However M III and M IV show uniform vespillo- material. The yield in pine stock
brings a new species picture. Here in M V the domination of N. vespilloides really comes to
light. Only at M VI are the three N. species commonly found.

This proves that the findings of the single catching points at the same time of year concerning
the species distribution are by no means all equal to each other. Rather it looks as though
clearly there is a factor governing the occurrence in appearance: The habitat of N. vespillo is
meadowland, while humator and vespilloides are indigenous to woodland. M VI supports as
its surroundings are mixed woodland as well as meadow, the importance of control
experiments. Correspondingly examples are found of humator and vespilloides which come
from the adjoining meadow. Those in the mid-part of the table containing beetle numbers
perhaps appear scarce. But the unquoted records of the same year (1931) as well as the
records for 1929, 1930, and 1932, extend the total to 2357 examples, and corroborate the
reported results, eliminating the probability that these patterns are due to chance alone.

Naturally, the membership of a species to a clearly defined environment from the records
must be all the clearer, all any time a habitat and untroubled from strange intruders opposed to
them. For the species vespillo it was possible to employ a trial experiment in a handy absolute
territory. It was carried out in Freistaat Danzig at a place where there is no woodland for
many kilometres in circumference. The experiment caught 64 examples of N. vespillo and
one of N. vestigator. Trapping in Freistaat Danzig, on different ground confirmed the
Frankfurt results. It was possible in Frankfurt in spite of little expansion of affiliated areas, to
construct a correlation of species and environment showing a considerable environmental
loyalty of the species. This applies particularly to N. humator and N. vespilloides, which in
general were found not more than about 100 m distance from woodedge; while vespillo
advanced a kilometer into the woods.

The results from M V in Table I indicate that N. vespilloides is found predominantly in stands
of confer wood. Hence its occurrence in mixed wood was due to the presence of scattered
pine stands while N. humator occurred particularly in deciduous woods. Although this
hypothesis is supported by numerous results, isolated results still occur which directly
contradict this. A moment on the ecology of Nicrophorus however helps to eliminate the
apparent contradiction. Because the gravedigger buries small carrion as food for the
descendants, they are dependent upon the condition of the ground which offers unequal
resistance depending upon the material covering the ground. The conifer woods, which appear
as pine woods in a large part of the Frankfurt area, stand here on predominantly very dry,
sandy ground. The ground is almost free from rooted plants, covered with an even rough
humus layer of needle litter which is interrupted here and there by small moss lawns
(Hypnum, Polystichum species). Elsewhere the ground is damp,deciduous leaf woods, which
in west Germany is mostly dark brown to black crumbling humus with a far higher moisture
content than the sandy ground of pinewoods. Its covering with leaf-litter and loosely twisted
roots of ground plants, is about average in resistance to mechanical destruction falling
between the loose leaf litter of pinewood and the dense layer of roots of meadows. The
ground of valley meadows (and only these are common here) is heavy and moist to wet. It
bears an extremely dense root layer of its predominantly Gramineae covering.

The described soils differ so strongly from one another in their condition that it was
understandable, when every N. species preferred for digging capability and behavior an
adequate environment. There is also clay-sandy deciduous wood soil and moist to marshy
pine wood soil. At such;places. the picture appears reversed exceptionally. In wet pinewood
N.humator rules but in dry deciduous wood N. vespilloldes does thus accounting for the
above contradiction. The exception offers the best proof that the ground is the primary habitat
characteristic for the-gravedigger.

Extreme differences from the rules are rare in the Frankfurt area. More frequently one finds
that the relative population between N.vespilloides and N.humator in mixed woodland shifts
in favor of one or other sides only somewhat. according to whether the experiment was put in
a dry, sandy or a more damp, humus mixed wood. For example the yield from M I and M II in
mixed woodland of the Tannus in Table I shows the proportion 41:36 where N. humator
forms the larger quantity. In mixed wood of sandy urban woods likewise both species are
found, but in totally changed proportion (Results M.S. S.520), 7 N. humator and 69 N.
vespilloides. The sample numbers of N. vespillo from adjacent meadowland were approached
in the woods.

3. Temporal occurrence

The baits were visited during the warmer times of the year for Necrophorus species. The more
or less early time appearance of beetles in the spring as with their disappearance in autumn is
naturally dependent on favorable or unfavorable weather. In 1930, the spring was warm, and
on 4 April the first gravedigger, N. humator, although the bait was on the edge of a small,
damp deciduous wood in the middle of a further stretch of meadow equally favorable to N.
vespillo. N. vespillo appeared for the first time that year 16 days later. Due to cold springs in
1931 and 1932 I cannot say what the average date of appearance is. In both cases the animals
did not appear until May and they were caught at the same time. While N. humator, N.
vespillo and N. vespilloides appear at approximately the same time and also disappear to
hibernate at the same time. N. investigator and N. fossor (interruptus) make a striking
exception. These species were not found until the end of June or beginning of July during all
three summers of the study (1930, 1931, 1932). The late appearance of these species is
connected with their individual cycles which are delayed for months in relation to the 3 other
species, N. humator, N. vespillo, N. vespilloides. On particularly warm November days one
finds here and there flying Necrophorus but as a rule, all species begin their hibernation
around the beginning to middle of Octoher.

Although gravediggers were recorded as common animals, one rarely finds them without the
aid of bait. This is based on their nocturnal life style. The gravedigger comes out of the
ground in which he has been secreted for the day, around sunset. If one observes this
happening, one sees as with a small shake of the ground two appearing antennae. Apparently
the beetle examines the environment through his smelling organs. After that the animal leaves
the sheltered province and climbs any blade of grass. In spite of his size and weight he brings
this about with dexterity. If the grass bends to the ground under his weight, another blade is
climbed. Apparently, the beetle operates by instinct, to raise himself somewhat above ground
level. In this way he manages to contact initially the light scents from possible food and
secondly he can lighten himself for take-off. There he remains sitting preening himself with
antennae moving, ready to begin his flight in search of food.

The gravediggers are pronounced dusk and night animals when living in a well maintained
terrarium of 50-60 beetles and only rarely does one appear in the daytime. If the
environmental conditions are unfavorable for the beetles, e.g. strong sunlight, high
temperature, extreme drought, narrowness of cage and lack of food, they are driven to the
surface during the daytime. They then undertake rapid running around and continual attempts
to escape in which they fly round wildly. These are however clear reactions to abnormal
conditions.

In certain cases the Necrophorus can indeed be found also during the day in natural
conditions. Every animal which has found carrion at night stays there during the day. There
they hide on the underside or slide inside in some which contain cavities within, where they
are able to devote themselves to undisturbed feeding. Often on windy but sunny spring or late
autumn days, thus shortly before or after the winter hibernation of the animals, one sees flying
Necrophorus. Probably they are driven to it by the hunger, and that at all times of the night it
is too cold to search for food.

Their life in the dark of night or the soil makes the gravedigger correspondingly sensitive to
bright light, yet the degree of sensitivity varies with the individual. Most times they retire into
the soil with sudden lighting with white light individual answers to a strong light attraction
done with Thanabos. Others again are not disturbed in their occupation. In strongly softened
light one can observe almost all individuals undisturbed. For this reason this sort of
illumination was employed chiefly for observation.

4. Nutrition

In quick flights, thereby frequent and uninterrupted in varying direction, the gravediggers
wander through woods and meadow till the wind carries to them the scent of carrion. At once
the flight stops, flying round the source of the smell in narrow circles and finally land on it or
in their near surroundings. In the last case one sees the beetle hurrying to his find with
swinging antennae.

Although the Necrophorus lets himself be enticed to the scent of carrion, they are not
exclusively carrion eaters, but predominantly predators. Already more than lOO years ago an
English researcher, Bell, observed how gravediggers seized and consumed the fly larvae from
a carcass. The description of this event, which was first published in 1873, has remained
unnoticed and still long afterwards Necrophorus is reported as a typical carrion eater. Later
Ch. U. Clark (1895) and W. T. Davis 1915 often have called attention to the predatory life
style. Most recently F. Steele (1927) showed this in a perfect series of experiments on the
North American gravediggers describing the facts of eating charmingly and vividly.
My experiments showing the preference for diptera larvae by European species is of value. It
was especially pointed out that also N. vespilloides is no exception to the pattern of nutritional
habits. The fact that this species was sometimes found in rotten woodland fungi, gave
occasion to the statement (Chenu and Desmarest 1851), that N. vespilloides did not feed from
carrion but only from fungi. But that is not true. Although these gravediggers were enticed
through the smell to decomposing fungi, they were still only hunting for the living diptera
larvae in the fungus. I saw a hungry male of this species consume 17 maggots in 35 minutes
after which the meal was interrupted.

The predatory nature of the genus Necrophorus is clearest expressed in the species N.
germanicus. In the night, this large, extremely agile animal hunts for Geotrupes unless fly
maggots are within reach. Klungelhoffer (1843) and Schmidt (1883) always observed a fierce
struggle between N. germanicus and Geotrupes (mutator) stercorarius and both concluded
from that the predatory life style of N. germanicus. This conjecture I was able to confirm. I
went along the following train of thought: If those observed cases from the above metioned
authors are not chance records, particulary N. germanicus, rather instinctive hunting for
Geotrupes it must be expected that it reacts to the smell of horse dung the typical habitat of
their prey--as N. vespilloides reacts to the smell of fung. In order to prove this, the following
experiment was set up.: In a round terrarium (lOOx94x58 cm) in which several N. germanicus
were held, I brought some horse dung. The Necrophorus, which until this moment were
wandering round apparently aimlessly in the cage, stopped with antennae beating keenly or
others immediately began their direction of marching and hurried to the dung heap. This is
possibly explained by the water or bacterial content in the fresh dung attracting the
Necrophorus to approach. That however contradicts the behavior of the beetle when they have
approached the source of the smell. None of them bite into the heap with their mandibles,
rather all run as if searching it over and turn around now and again a large piece of dung with
head and thorax. One cannot keep the false impression as these animals search the dung for its
occupants. Single individuals withdraw, the majority however remain at the site and bury
themselves superficially in the dung. One particularly lively female hid for example under a
lump of dung. Only the head with perceiving antennae, was still to be seen. At this point I
would like to take the opportunity to mention an observation of V. Lengearcken, who found a
N. germanicus under a ball of horse dung lying in the wild. Animal and dung ball were
already sunk somewhat into the ground. Undoubtedly it had to do with the same event which I
was able to observe in the terrarium. There also I saw here and there how the dung was sunk
somewhat into the earth through the agitation beneath the beetle.

So the beetles remain to a certain extent in ambush, only alternating now and then hastily
leaving his place. This behaviour of the beetle then follows without there being any Geotrupes
in the cage or in the vicinity! After a fresh dung pat is searched through for the Geotrupes by
N. germanicus, they wait for the moment which enticed hy the smell of dung, happens with
all probablity in a short time.

Now I placed about 60 Geotrupes into the terrarium. All belonged to the species G. sylvaticus,
which I was able to collect most easily in the surroundings of Frankfurt. Nearly all the dung
beetles groped about for a long time at the nearest way of the dung heap. Scarcely had the
first reached their goal when those still inanimate predators became lively, appearing in
masses. Everywhere the large predatory gravediggers claimed their prey which to begin with
chirped loudly, but desperately, to defend themselves. Also the previously mentioned female
left its place under the dungball and hurried smartly towards the near prey by the shortest
route. N. germanicus attacked its prey not from ambush as one would presume from its initial
behavior but hurried openly to his prey and seized it with legs and mandibles. Every caution
appeared totally useless as the observer did not receive the impression that Geotrupes
sylvaticus was able to recognize his enemy as such. One sees on the contrary the dung beetles
marching on the Necrophorus and even in direct proximity no impression was noticeable.
That what took place now between the attacker and his prey can by no means be described as
a struggle. For both animals are too unequal in strength and quickness. N. germanicus seized
the weaker, G. sylvaticus with both legs and thereby threw itself on the back or the side. This
pose was taken willingly as the eating of large fly maggots, so that more than half of the
gravediggers can be found after a feed with these larvae, the prey held in front of them
between first and second pairs of legs. Small fly larvae are nevertheless held free in the
mandibles.

Through the extremely tight hold of predators, Geotrupes sylvaticus becomes pressed against
the ventral side of its opponent and can be eaten conveniently. One hears loudly between the
weakening chirping of the dung beetles cracking of the chitin armor under the strong
mandibles of the gravediggers. At some point the outer skeleton becomes broken. In the
majority of observed cases the prey was slit open on the ventral side of the area between pro-
and meso-thorax. In other cases the predator succeeded in destroying the strongly chitinized
(thorax) neck plate. Once the body is open, the organs are pulled out and eaten. In the course
of half an hour I saw one N. germanicus male seize and consume five Geotrupes in the
described way. The others who were no less lucky in their hunting so that the remainder of the
Geotrupes lay around everywhere in the terrarium. The N. germanicus seizes the dung beetles
so quickly when hungry, but they are indifferent to them in the satiated state, hurrying by
without any attempted attacks on the Geotrupes.

The experiment was performed several times with the same results: N. germanicus habitually
hunts for Geotrupes! To be sure, fly maggots and Geotrupes were offered to them at the same
time and as a rule the fly maggots were preferred.

In spite of their preference for living prey Necrophorus show only a slight inclination for
cannabalism. Healthy individuals did not cannabalize others when very hungry and under
confined living conditions. An exception to this may arise in fresh, recently colored young
beetles which at times consume uncolored individuals--usually brothers and sisters. This is
seen only in rearing under artificial conditions,indicating environmental conditions as
causative factors. While in nature the beetle immediately goes out in search of food after
leaving the pupal stage, in the terrarium he is held back by enormously limited room. From
deficiency in other food the animal seizes conspecifics which are convenient prey for him.
This was easy to prevent; shortly before the hatching of the beetles some meat was put in the
cage. Still in nature, cannabalism can occasionally be observed, when sick or hadly injured
individuals are attacked and consumed.

Besides the predatory nutritional ways all here-mentioned, Necrophorus species are also
carrion eaters. As Steele (1927) already mentioned for the North American species, he
showed the little decayed food of the strongly decomposed. Largely independent of the type
of meat, the Necrophorus accept every kind of carrion that is offered to them. Meat or internal
organs of vertebrates such as sheep, pigs, beef, horse or even tiger and turtle soup whose
bodies were placed at my disposal by chance served the gravediggers for food just as the
bodies of small vertebrates or even many invertebrates like earthworms, slugs, and larger
insects. When one puts a hungry animal on a piece of meat, it tests the food here and there
with antennae and mouth appendages. In a somewhat damp and soaking place one sees it stick
to and drive in its mandibles. The choice of place to eat corresponds to completely the
preference of the gravedigger, on small vertebrate bodies firstly the body opening, eyes or
possibly wounds as this also Steele (1927) was able to establish for those species which he
had examined. Yet without having a visible ingredient cut off, the jaw becomes open again, in
order to immediately grip to new dense area behind the place again, a match that is repeated
so long with between the mandibles rises a weak border of ordered meat. One sees the mouth
appendages working this border again and again. After a long time, often after an hour, the
beetle turns to another place, without that, that border would have completely disappeared.
If one now disturbs the animal in some way, it vomits immediately a part of the food taken
up, in which from microscopic examination are traced no formed ingredients. Therefore the
meat is not devoured in small pieces, particularly rather milled through and pressed out by
continual work of the mouth appendages to then take up the food in liquid form. The tough
and hard tissue stays behind, as it also happens in the-consumption of a diptera larva whose
cuticle stays behind--when also in young larvae till crushed to an unrecognisable state.
Besides the mechanical work to gain liquid food, it would be possible that the material
becomes liquified through the effect of a deposited extraintestinal gut secretions. Although
the vomiting of a digestive secretion was not able to be explained up till now, certain
observations and considerations make the surmised events probable: A fresh piece of insect
from which a Necrophorus had eaten showed for a long time at the eating place distinct
alteration. Brown color and jelly-like blurred contrasted the place clearly with almost
unaltered surroundings. Further speaks the admitted factor, that disturbed gravediggers vomit
intestine contents, respectively intestinal secretions for the operation of a preoral digestion
during meals; to us this appearance is faced only in beetles with extraintestinal digestion.
Further evidence for this species the food in-take by the nearest relations of the gravediggers,
the Silphinae, is wide, as Heymons and V. Lengercken pointed out (1926-1932).

B. Reproductive Biology

1. Relationship of beetles to one another.

a) Attraction of females

In May the species N. vespillo, N. vespilloides, N. humator, and N. germanicus, in August the
species N. investigator, N. fossor are already in reproduction. At this time the beetles begin
with that activity to which their name gravedigger points: they bury small carrion in the soil in
order to receive it as food for their descendants. Their gonads have developed immensely to a
mature state, and both sexes happen now on the search for carrion that corresponds to the
demand of their broodcare instincts, and describes at the same time the meeting-place for
males and females. When a body is found and a pair are together then the preconditions for
reproduction are fulfilled.

If only a male arrives at the carrion, one can make the following highly remarkable
observation: the males which initially set about the digging work, break it off after a short
time in an effort to reach a high point in the immediate vicinity of digging. The male climbs,a
blade of grass, a stone or even a small heap of soil, for want of a high lying point. In all
observed cases this is commonly a very surprising pose which

the male assumes. The animal orients itself so that the head is as low as possible but the
abdomen stands as high as possible in space. The head is thereby bowed so low that the
mandibles often rest on base, grass stalk, stone or soil. The whole body, which normally
shows in every Necrophorus in front of as well as behind a distinct slope from the ventral
aspect, becomes extended in a slope from behind causing a steep angle. The body weight
thereby rests on fore and middle tarsi, while the hindlegs are only lightly placed or even held
freely suspended. The hind end of the body is so strongly extended that the abdomen appears
longer and more slender and the last segment which normally lies hidden in the body appears.
In this position the male persists, refraining from short breaks without clearly visible
alteration for more than an hour. Only the tip of the abdomen betrays slight swinging or
rotating movement; the last segment stretches away from the others occasionally revealing the
intersegmental membrane. I would like to describe this strange operation as the "Ventilation"
of gravediggers.
What ecological meaning comes from the "ventilation" of male gravediggers? I was able to
observe it 13 times, the results of which are summarized below.

1) The "ventilation" was regularly performed in the immediate vicinity of a carcass.

2) Ventilating animals were without exception males with ripe, plump full testes.

3) Never was a female observed in the vicinity.

4) Only on warm summer evenings (June, July, August) after sunset was ventilation able to be
observed.

These facts suggest "ventilation" of male gravediggers involves the females. Presumably
flowing from either the tip of the abdomen or the intersegmental membranes are sex or
species specific scent to signal passing females and to guide their approach. This assumption
is supported in that at times one sees both sexes coming to the carrion's scent. When the
carrion is buried superficially by the male, thereby checking the enticing smells, the sex scent
of the males was able to lure a female; a peculiar profit results for the prey is protected from
any other carrion eaters, and remains reserved in all probability for the species.

The declared explanation also found experimental confirmation in degrees: after a series of
failed experiments I succeeded finally in two cases in bringing to puberty males isolated on a
carcass to "ventilate." That this experiment frequently failed, I believe goes back to the
artificial environmental conditions. Out of the obvious assumption that the above mentioned
points 1-4 are taken from, the size of the cage (my successful exp. were carried out in a cage
of the dimensions of lOOx94x58 cm) and above all good ventilation are important in success.

So the term "ventilation" in the gravedigger ought to be authorized. Still it ought to be pointed
out that a Necrophorus is not designed for a such extreme placement of the abdomen in
ventilation. Although the movement of the hind body is not equal to that of Hymenoptera--a
still striking apparent convergence finds the ventilation of gravediggers remaining like the
termites. From Tollin (1862) a male termite that has found no partner through swarming who
scarcely had begun digging work, stretched the abdomen in the air and stuck to it for many
hours each day.

b. The isolation of pairs from a large number of individuals and the resulting conclusion.

When a male and female of the same species meet on a suitable body for broodcare (or the
body has been found by an already paired female (vgl. S 539) the carrion is buried and serves
the larvae as food, till they are full grown and move further into the soil for the pupation. At
this time the store is consumed until from insignificant amount remaining, has sufficed even
for the brood's parents. But what happens if the same quantity of food has to suffice for the
brood of several females? Without doubt the development of all larvae would be greatly
endangered through shortage of food. This danger is likely -for it happens often enough, that
through the smell of the carrion several females are attracted with mature~ ovaries. How is
the threatened danger averted?

J. H. Fabre (1899), as I have already mentioned in the introduction, made the striking
statement that he never found several females on buried carcasses. Always he mentions only
one pair there, even if beforehand more than two individuals had been working at the digging
at the same time. My first task was to examine this declaration of Fabre for its accuracy.
Fabre, known as a sophisticated observer, remains to be considered for this observational
experiment on Necrophorus which was carried out in an aviary, therefore under artificial
conditions, with only a limited number of beetles, namely 14 individuals. The question which
I had to answer ran accordingly: is only one single pair really found on buried carrion in all
cases, including under natural conditions?

All my experiments were made correspondingly in natural conditions--thus they were not
successful. Through laying out suitable pieces of meat to attract the grave diggers, they were
allowed to bury the prey undisturbed and after a known time the number of beetles remaining
on the carrion was checked by digging it up. Digging up is too crude a method. Through the
warning of shaking the ground, many beetles hide further into the soil and cannot be found
and cautious digging is very time- consuming.

In order to avoid this disadvantage, 200 flower-pots (12 cm high, 14 cm diem. at top) were
distributed in woods and meadows in the surroundings of Frankfurt. They were sunk in the
soil to the upper edge and filled with dug material till full, after the base of the vessel was
covered with a fine mesh wire net, to prevent the escape of beetles from beneath (Abb. 3). A
small piece of meat about the size of a mouse was laid on the soil in the flower pot. On a
small adjacent piece of wood a number was marked under which the results were entered in
the records. The experimental site was visited several days in succession and observations
made on them continuously registered. After the meat was sunk into the ground, through the
work of grave diggers. I waited 2-3 days, then dug the flower pots from the ground and over
turned them on a firm base. Its contents lay exposed in this experimental procedure and could
be searched through without trouble, which is also of importance for later mentioned
observations. Escape of beetles was excluded through this arrangement.

Although not all 200 experimental sites were in operation simultaneously (for the meat was
eaten frequently by other animals or carried off by the gravediggers), I was still able to
examine in the course of Summer 1930, 254 cases in the described way. An experiment was
only counted then, when the carrion was found already in a subterranean hole, which Fabre
called the crypt, there in the presence of this crypt for sole proof for the termination of
digging work is recognized. The examination gave the following results.

In 197 of the mentioned cases a pair was found on the carrion or in the near vicinity within
the flower-pot; in 53 cases, I was able only to find a female. In 4 cases I found in the near
vicinity of the crypt 2 females (which by further investigation proved to be sexually mature).
My experiment then proved perfectly that really only 1 female remains behind on the buried
carrion and it has in the great majority of cases the male with it. The number of exceptions is
so small that they may be neglected. Also it ought to be emphasized that the results were not
being influenced substantially perhaps by the lack of competitors: in most cases more distant
individuals were found in a small distance outside the flower pot in the soil.

If the initially asked question is answered in the affirmative, we are faced secondly with the
problem of how the separation of a single pair is brought about from a large number of
available candidates. This looks as if it involves animal psychology. But the apparently
puzzling occurrence is clarified through broader consideration. The grave diggers dispute
their right to possession of the carrion by fighting one another, and it is predominantly
individuals of the same sex which fight.

The description of a typical start to an experiment comments on the cause of events in detail.
In a spacious terrarium, 4 pairs of the species N. vespillo were introduced. Every female was
marked with a color mark on the pronotum. All males received at the same time in the same
color (white, light blue, light green and yellow) a mark on the elytra so one was able to
recognize again at a glance the sex of the animal and at the same time the individual. The
pairs were first confined to this experiment after they, separated from the others, had proved
their sexual maturity through desire to dig.
On the day when the animals were committed resting onto the soil, I layed a dead mole on the
soil surface in the middle of the cage. At sunset the first female (bright blue) appeared and
hurried to the mole, climbed and clambered around on it, in order to disappear. Here & there
the shaking of the dead body showed that the female was digging. Meanwhile the light green
and yellow females also turned up. While the first ran here and there the corner of the cage,
the yellow male marched smelling to the carrion. Arriving there it soon slipped under the
body. While for several minutes nothing happened, till both animals, the bright blue female
and the yellow male by chance at the same time came forth from under the mole, each at a
different place. Now the female scent appears to come from the new comer (male). The
female pauses visibly and runs directly to the male with quick steps, hesitates, reaches with
the antennae the abdominal tip, that aggravates him, as the male has quickly hurried over by
the approach of the female. Scarcely has the female succeeded in testing the hind end of male
with tasting tips of the antennae, than it turns away. Then the male begins to follow the
female. With a few, hasty skips it stops, a little shake with the antenna and with the loud
chirping of both animals, mating takes place. After that the pair turn toward their work again.

In the corner of the terrarium, the second female appears level with the soil. It bears a white
color on the neck plate and is already well known to me because of its strength and
savageness. This beetle also goes to the carrion immediately and meets there with the female.
Both go immediately one after another and both appear to recognize the smell of the hind end
of the body of the sex of opponents. The movement of the grave digger at their work
particularly on warm summer evenings was described already, and quickly mentioned, yet the
aggressive savageness surprises so, with those in the situation the two females rushed at one
another. The fight begins.

But, scarcely begun, it is already finished. The bright blue female, which at first had taken
possession of the mole, left the place of the fight in a rush to escape. It searched the walls of
cage and clambered at them, and as this did not succeed, it buried itself in the furthest corner.
It was to be seen no more for the rest of the evening. The strong female with the white mark
now crept under the mole where meanwhile the male was pursuing his work. The victorious
female thus thrust aside the loser, and took possession of the prey itself. Soon it emerged
again a few cm distance from the body. Close examination showed that it had created a
passage, which began under the mole and ran to the surface here. One is reminded of passages
which lead from the inside of old fortresses to freedom. In this security the female remains
sitting till hidden to head and forelegs in the soil, and throws itself from here out onto every
grave digger which wants to approach the carrion. First it is the light blue female that comes
that way. Although after itself, the female has information about the sex of the approaching
animal, it creeps under the carrion, and turns up there shortly again in the mouth of its
subterranean passage. The newly arrived male occupies himself meanwhile with the digging
work and has soon disappeared in the soil. An approaching female (yellow) is treated less
peacefully. Scarcely had it shown its identity through its scent as such, than it was attacked
savagely, and was driven away after a strong fight. The bright green female did not come off
any differently. From this the white maintained its place. During the watch the guarding
animal stayed directly in the soil. I sat the yellow, already once expelled female, with forceps
on the mole. Then it had plenty of time to work, and was mated with the bright blue male. By
accident the white female came into its vicinity, the fight was repeated, then again the yellow
animal left. Now it was no longer possible, to bring the defeated female onto the mole and for
it to remain there. I tried it so often, but these animals were good at escaping from these
places. The strong female even set back the yellow, and then also attacked it, when I held it
tightly with the forceps. Both layed into one another so quickly, that I was able to lift both
fighters together without them separating. During these maneuvers a newly arrived male
(white) succeeded in getting under the body unnoticed by the white female.

We see how well a female fared in the field of other females! How do the males behave
among one another? The two worked during the whole duration of the experiment on the
same carcass without mutually disturbing each other; the mole is sunk further into the soil
through their common vehicles. Finally even a third male arrived. Suddenly, a loud chirping
becomes audible and in a wild chase 2 males appear under the carrion, the third (white) close
behind them. This comes (as in the females) at last usually to a fight, from which the white
male comes off victorious and turns back to this work, while the fugitives appear to keep
away from the mole now. After a further hour in which nothing more in particular happens
and the mole is rapidly fully covered with soil, the observations were discontinued. On the
following morning by digging it up, I found on the carrion the white pair alone, whose
isolation had taken place in the described way.

If the described events are valid as typical, then they also show certain peculiarities. For
example, the sort and manner as the victorious female creates an ambush in the form of a
subterranean passage, do not describe the rule, although this was also observed in other
females now and again. Frequently the female briefly undertook trips radiating out from the
carrion but always returning to the body. I would like to interpret this behavior of the female
as a species watch (guard) service, during this the close vicinity of possessions at risk is
searched for strange females and prevents an unnoticed intruder going by. Further the
experiment demonstrates that the second female was superior to all remaining in strength and
drive to attack. If this were not the case, then the carrion could be conquered still by the third
or even fourth female and so several times change owners.

The experimental description has shown in which way the fight between grave diggers of the
same sex brings about the peculiarity of a single pair from a large number of individuals. It is
true that the described mechanism is not absolutely clear, but still works with greater certainty
except with a very small number of exceptions (4 out of 254 cases) in which two females
were found on one and themselves buried the carrion.

But how is it, that 53 times only females were found. So far it is not a matter of cases in
which a female mated elsewhere had layed out the burrow; this is the result of one female
changing instinct after completing the passage digging. The female that puts up with a
digging male, however, attacks after completion of the crypt of the partner and drives him
away. Most of the females enter the change in behavior shortly after egg laying, yet can
before; after this time the male can be expelled from the brood chamber. Under these
circumstances, different sexes of the same species of grave diggers alone fight. I was never
able to observe, under the same conditions, fighting between different sexes of individuals in
N. vespilloides.

The fighting instinct of males and females has a certain distinction. The female behaves
during the time of passage digging in a vigilant, and limiting manner and neglects the digging
work more or less. The male however turns exclusively to the work. First if they by chance
meet at their work, the fighting instinct is aroused. Then their fight does not differ from that
of the female.

The circumstance that temporarily several males are able to be found digging at once on the
same carcass, is naturally very conspicuous in the biology and has given occasion to far-
reaching, theoretical remarks. One remark by Fabre says that the researcher saw, on average,
more males to use the help of pairs, often also the effect of social instinct. "Un couple etait-il
dans l'embarras, avertus par le fumet, des aides surviennent, servants des dames, qui se
glissent sons la piece, la travaillent de l'echine et de la patte, l'enterrent, puis s'en vont en
laisent a leurs joies les maitres de caens". Quite similarly Renkev (1913) interpreted this
appearance as "the first germ of an altruistic instinct", while Schroder (1929) in his even
wanted to recognize a " particular case of the use of strange foreign expediency".

Alverdes (1925) expressed the presumption the opposite way, that the assembly of several
grave diggers at carrion suitable for broodcare showed only "association", in a sociological
respect, whose origin lead back to environmental factors, not the existence of social instincts.
This explanation neglects the crucial point of the problem, namely the question of what
causes the surplus individuals to leave the carrion. So far, however, only the carrion work of
the males from my results confirms Alverdes' prediction.

For inspite of the common work which in truth is only based on the chance adding up of
single results, in fact no relationship exists between the digging males. Every male works for
himself alone and for himself personally. His advantage lies in the chance to reproduce
successfully and this possibility is never abandoned without a fight in favor of a competitor.
Every male looses much more than every female, first then his place and if it is forced to do
that by a mate of the same sex, which is superior to him in strength and skill. Through this
therefore, the prediction of Renkers' (1913), as also the explanation of Schroder's 1929, is
without foundation.

c. Ecology of the fight.

The main value of the fight between gravediggers of the same sex is frequently that a single
pair remains behind at the carrion and their descendants find sufficient food. The existence of
the instinct to fight is already sufficiently motivated through this.

But it appears to me that this fighting still raises a further use: An effective choice within the
species. This opposes us more clearly in the fights of females. Any female which is in
possession of a body has to prove, as it were, before a new approaching female by way of a
fight its strength, skill and the normal functioning of certain instincts. Then the carrion passes
into possession of the superior animal and so that the possibility of egg laying on the piece of
carrion in question remains only for the better equipped female, and obviously encourages the
propagation of the stronger and healthier female. While one is allowed to accept that this
stronger selection puts the reproduction of weaker or even genotypically retarded females in
question.

The translation of these results in the fighting of males with one another however meets with
difficulties. While with the driving away of female animals also these germ-cells become
distant, when any female first proceeds to egg-laying after the burying of the carrion and these
long winded preparations, the numerous males are chased away often first because of the
small desire to fight off male animals after they have already mated with healthy females of
the same carrion. In such cases their violent removal becomes meaningless as a means of
eliminating inferior quality. A cause of selection, through fighting, therefore can be found in
the males only in one case, in which the chased male did not succeed in mating. The possible
drawback in this for the species made up for through the gain would result from the adding up
the work performance of present male at the same time.

This remark "natural breeding choice", which is recognized in the fighting between
gravediggers of the same sex appears suitable from consideration to permit a classification in
the narrower conception of "sexual breeding choice". Although those fights, at least by the
females, determine the permission of participation of individuals in reproduction, nevertheless
the lack of connection of fighting animals to the representatives of the different sexes
excludes the sexual breeding choice. The fighting of gravediggers is for possession of the
carrion, not for possession of females, and males respectively. Any Necrophorus fights only
in the direct vicinity of a suitable body for the broodcare. At meetings of the sexes under other
conditions, for example of large carcasses, mating pairs are frequently found without
observations of hostile attitudes or fights between individuals of the same sex.
d. Fight with strange species or races of gravedigger

Gravediggers are sometimes also ready to defend the stores of food committed to the brood
against representatives of strange Necrophorus species. Naturally it gives in easily for in
districts with mixed environmental characters several species are attracted to a carcass. Yet
fights do not always result from this meeting; the weaker species usually hastily retreats
before interacting with a larger species. This is established particularly frequently in N.
vespillo and N. humator. A N. vespillo female leaves the prey it has fiercely defended against
females of its own species, when approached by N. humator of either sex. Even strong N.
vespillo females, which in size are scarcely smaller than weak example of N. humator, retreat,
leaving the carrion to a N. humator. Occasionally particularly aggressive individuals of N.
vespillo fight against an enemy of a strange species with variable results, as I observed in
eight cases. The fight developed then as between the same species, however it went ahead
independent of the sex of the two participants. More males and females attack representatives
of the weaker species, for example N. vespillo or N. vespilloides, without any hesitation
provided that these do not avoid the attack through hurried escape.

A remarkable observation is inserted here. In the surroundings of Frankfurt the species, N.

vespilloides shows considerable variation concerning elytral markings. The black crossband
which normally goes through the elytra unbroken, slightly behind the middle and often
extends so far that the distally lying red yellow crossband, is reduced to two almost round
spots, is displaced and in several examples is so strongly reduced that only a row of small
black dots or a long line remains between which both red yellow marks come together. There
the existing various different kinds of elytral marking show all transitions, missing any
possibility which separates stronglv differing forms with the help of named characteristics as
hereditary race of the species N. vespilloides

The investigation offered me the opportunity to establish whether really only one pair from a
large number remained behind on a buried carcass. I observed that females with strong elytral
markings either always paired alone or with a male differing from the norm by the same
degree. This observation is based on twelve recorded cases which were all positive. I never
observed a negative case, and I can rely on several more positive but not recorded
observations. If one considers the relative scarcity of this extreme observation, it appears that
it could not be due to chance; one explanation is that the abnormally marked females fight or
drive away the normal males and only leave males similar to themselves unmolested. This
supposition assumes a physiological difference of those females from the normal which can
only be based on hereditary mutation. Then would mention the possibility for it that the
weaker differences of all degree is considered genotypical without being isolated
physiologically from the stem type. The extremely different form had the value of a beginning
species.

e. Course of the fight

After establishing which individuals fight each other we now move towards the kind of fight.
The typical fight of a gravedigger happens with such speed that it is impossible at the first
observation to grasp in this confusion fast moving legs of individual actions. Repeated
observations of the incident supported by instantaneous photographs finally allowed
recognition of the real stages of the fight. How quickly the movements were carried out
comes from special difficulty of the photographic technique. At first, when the exposure time
was reduced to under 1/100 sec the picture received sharp contours. Most were therefore
taken with an explosive of 1/250 sec. I operated with "Agfa instant flashes." It offers the
advantage that at the moment of the brightest illumination which lasts for the duration of
1/250 sec., the adjusted shutter of the camera was released electronically. The duration of the
fight amounted to the quickness of movement corresponding to a few seconds. Only rarely for
example with very fatigued animals did it take a longer time.
The beetles usually simultaneously came upon the lip of the abdomen of others scent, began
the fight in opposite positions. Next the opponent throws himself at the side and seizes the
opposite side of the head and thorax. In this attitude one sees how each of the fighters works
the abdomen of the adversary with its mandibles. The closed jaws slide repeatedly from distal
to proximal part and bite, where they push off the hindrance as at times on the segmental
boundary. To this point the fight still remains undecided. Soon the firm hold of the beetles
appears to loosen. Each individual tries to move his adversary with his legs, for this the
longitudinal axis is put across its own. If one of the fighters succeeds in this, the fight is soon
decided. The beetle which at first squeezes his opponent with his legs, now has an easy
match. He presses his victim together many times with such strength that one hears the chitin
cracking. The animal which is stuck under the firm grip of the legs in a living vice is unable to
offer resistance while the limbs are shackled to him. Fig. 5 shows this phase of the fight with
which its end is reached. Again free, the defeated gravedigger takes to flight immediately.

The fight takes another start, if one of the escaping animals, which ought to be withdrawn or a
young beetle appeases his hunger on the carrion intended for the brood. In both cases the
persecutor tries to seize with his forelegs from behind the last or second last pair of his
opponents and these pulled close with a jerk. Here the animals soon lie against one another on
their sides. In contrast to the above situation they have an equal tendency to be righted as Fig.
6b illustrates. The rest of its course follows as described above.

Finally still the result of the fight remains to be considered from the individual concerned.
The weapons of the gravediggers which are employed in the fight are above all the legs, in
particular the last pair, the mandibles being of secondary importance. Although the fight
advances on strength and wildness one hears at times loud cracking of the chitin. I have never
seen a beetle emerging hurt from it. Conspicuously indeed the comparatively high percentage
of mutilated individuals remains, which one comes across at the time of reproduction. Mostly
it concerns missing tarsi, sometimes also missing tibiae or antennae. It is also possible that
these animals have lost these parts of the body through their difficult digging work; I
nevertheless incline towards the assumption that also during the fight, injuries can originate.
On the one hand, the observations speak for this interpretation for I repeatedly found injured
individuals full ovaries with mature eggs. This suggests that the animals in question probably
still had not even dug. Also, the mutilated animals are particularly aggressive; they are more
frequently willing to fight the others, especially strange species and perhaps stronger
individuals.

A recurring effect of fighting is the speedy escape of the weaker animal which as result of
receiving attraction a clear change of behavior can be observed. The digging instinct which
rules the beetle up to his defeat, becomes displaced by the flight instinct. In by far the
majority of cases the escaping beetles collected in the nearest vicinity in the earth without
venturing a new attack during the following hours. On the contrary one sees them either
clearly avoiding the body or in their search for a hiding place not giving it more consideration
than an indifferent hindrance. Still, the duration of this happening appears to be dependent on
the intensity of the sustained defeat.

Undoubtedly the escape instinct supports, so far as it concerns fights between individuals of
the same species, the fighting instinct with regard to its biological significance in most
favorable ways. Then firstly from the cooperation both results in every advantage for the
species which were already reviewed in the last chapter.
f. Relations between partners of the pairs.

That the relationships between male and female during digging are extremely loose has
already been said. The fight between beetles of the same sex round the carrion can in effect
bring about a change in the pair without releasing a visible reaction to the fight by the
indifferent remaining partner . Only the presence of the body holds both sexes together at this
time. If the carrion is removed the animals then leave the digging place and furthermore take
no notice of one another.

Nevertheless, the following observation allows one to assume an association between male
and female: the distinctly audible chirping that the male as well as the female produces with
the help of shell organs--most irregularly--arouses an impression now and then, that as if the
partners mutually answer one another. In most cases the chirping proceeds in the way that the
verse of the animal which is composed of several shrill noises of the same kind start the verse
of the partner; schematically it can be represented in the following way at which the number
of the quoted shrill noises added is of no importance:

Male ------------------------

Female -------------------------

While this incident in the male is already over after several seconds and repeated first after a
considerable length of time, I was able in three cases which were observed completely in the
wild, to overhear the loud utterances of the gravediggers which really differ from the first type
described. It is a matter of a continuous common chirping of males and females which lasts
minutes long and sounds very strange. The unusual sound picture arises that each alternates a
chirp of one animal with that of the other, schematically expressed:

Male - - - - - - - - - - - -

Female - - - - - - - - - - - - -

Whether a firm task falls to the chirping of the gravedigger at the digging and which it is, I
cannot say with certainty. In all probability the partners influence (perziperen) their mutual
chirping in some way. Copulation took place regularly in the course of digging, suggesting
that the mutual chirping is an expression of sexual excitement. Moreover the possibility exists
that the chirping operates as a warning noise and serves to frighten away related species or
genera. The common minutes-long chirping of males and females especially give this
impression. It was strengthened owing to a pair chirping in this way being constantly in
regular activity: while the male digs one frequently sees the female radiating from the prey,
going out investigating.

As soon as the digging work is thriving over the first start, the mating happens above ground.
The male climbs the female in a manner that in normal important relations his foretarsi get a
grip of the corners of the shoulders of the female and the middle tarsi grip round roughly the
epipleurons. During the actual union of both beetles the male performs a brushing movement
with his foretarsi which appears broadening in relation to the female. At the end of 3-4
seconds copulation is terminated. That same pair can however still proceed to renewed mating
before the conclusion of the digging work.
2. Relations of the beetles to their offspring.

a) Brood care

Examination of the food selected for the brood. As the gravediggers are not choosy
concerning their own food, they are also satisfied with any carrion for the provision of the
brood. Indeed the size sets the upper and lower limit. Beetles are not able to bury large pieces
and small ones do not supply a sufficient quantity of food for the numerous hungry larvae.
However, between those there remains considerable latitude. If we disregard the strongest
member of the genus, N. germanicus (this species is rare in the surroundings of Frankfurt and
so therefore too small for experimentation) carrion from the size of three day old kittens to
pieces of meat 1 cc were buried. They were usually distributed such that N. humator naturally
found the largest, N. vespilloides the smallest pieces.

Unable to detach pieces from large carrion, the beetles are directed in their care for the
offspring predominantly to the bodies of small vertebrates. When a sexually mature
gravedigger has located a carcass, he first reacts in a wholly determined way. The dead body
is climbed, the mandibles beat here and there loosely in the carrion with the maxillary palps
clearly shaking, the antennae moving in a swinging motion. At least twice the beetle steps in
nearly vertically one upon another, standing directly on the body. Thereupon the beetle slips
under the carrion and its activity can only be guessed. He raises the prey a little from the
ground.

The single phases 1. Examination of the find with mouth parts and antennae. 2. Inspection of
length and breadth. 3. Lifting of the carrion, take place in succession, being constantly
observed before the beetle begins the real digging work. One gets the impression that the
animal deals with the act of examination of his find in this capacity. It is conceivable that
initially after the find through each constantly recurring action the chemical condition, size
and shiftability are examined, thus first from the determined combination of doses of
excitement, the digging instinct is released. A confirmation of this presumption is seen in the
fact that a Necrophorus not once attempted bury carrion whose size exceeded his capacity.

b. Burying of a body

Naturally the direct observation of the digging activity was impaired to a large measure
because the largest part was carried out in the soil and therefore remained invisible to the
observer. This circumstance explains why the analysis of the digging technique of the burying
beetle is still outstanding. In order to be able to follow the occurrence without a break the
investigation must be completed in the wild as such under known artificial conditions. To this
end, sexually mature beetles were brought pairwise into narrow terraria to dig so that a direct
observation of the activity was possible through the glass walls.

The terraria (24 cm long, 18 cm high, 2.5 cm broad) have wood borders on the narrow side-
walls which are cut crossway square (2.5 cm x 2.5 cm). On this wood border is screwed, from
below the soil on the long side, a right angled aluminum strip. The long walls consist of glass.
Each glass plate leans on the right as well as the left woodborder so that every time

they are attached through one of the outer sides of the wood border and held by the right
angled aluminum strip underneath the glass plate on the ground is a prop. Both long walls are
held so tightly only by the metal strip that they try to slide slightly here and there as in a
rabbet. This is implicitly required in order to be able to change the glass plates during the
observation without strong shock. That often proved to be necessary against the work of the
beetle held together dirt through small soil particles or carrion. The vessel top was closed with
wire gauze.
However, under these conditions which differ strongly from the natural, some of the beetles
did not set about digging, but rather exhausted themselves through continuous attempts to
escape. In order to be able to use these animals for experimental purposes, I attempted a
further trick. A box of zinc plate was made of the size that therein holds the glass vessels put
against one another, tightly. All single cages were filled full with soil and above that still a 2
cm thick layer of earth spread out. It originated in this way a roomy surface under which the
glass terraria lay hidden. At the start the beetles were in no way confined to their work. In due
course they came across the glass wall in the soil, but this hindrance meant no more then a
stone or a hard root in the wild which the animals are able to avoid.

Actually all gravediggers went nearly immediately to the work. They fall on the way in the
depth with their prey necessarily in one of the glass boxes which then with help from the earth
surface over-topping is raised out and the observations made. The glass cage being used was
replaced immediately by another, and consequently the described experimental order is ready
for use so long as it does not lack sexually mature beetles.

If a beetle found a carrion which proved after complete examination to be suitable then the
digging work began. Next the beetle scraped more soil out from under the body. The
gravedigger thereby proceeds so that he carries soil particles from behind with forelegs which
is further given to the second pair and then reaches the last. From there the strong hind legs
shove out the mass from under the body. Though frequent repetition of the activity arises the
already mentioned small earth wall of Fabre (1899). However the dead body is not ring-like,
but only grasped round partly. Correspondingly, only a part of the carrion is also only dug
under.

More frequently than on bare, soft ground the gravedigger finds the carrion on hard leaf. This
is especially true for the the meadow-inhabiting N. vespillo. Under these conditions are
experienced the application of the already well known (Fabre 1899) ability of the beetles to
cut through grass and roots with their mandibles. If one perhaps removes the small animal
body an hour after work begins, one sees an approximately round place on which the grass
stalks are pressed on the sides and are bitten through the base so that from the small area all
sort of roots still travel through, laying free in the soil. The size of the prepared place always
is dependent on but, importantly, smaller than the object to be buried. Therefore, also in this
case the whole body is not dug under.

From this stage onwards the digging work is continued independent of the existence of a
ground covering always in the same way. Even after the

number of hindrances which it rescues from the ground in the shape of roots or small stones,
it merely fluctuates the working hours. The beetles push out more and more soil from under
the carrion and thereby deepen the already started hole without broadening it at the same time.
The part of the body which is dug under begins to sink under its own weight (Fig. 8a).

Up to this time one is able to follow the work of the Necrophorus from the shaking of the
body, then suddenly this ceases and remains completely motionless for a long time. Through
that the observer receives the impression that the animal has ceased its activity, but in reality
the work goes ahead briskly and is continued in another way. The working animal has begun
from the hitherto cultivated, shallow depression to dig out a passage sloping from beneath in
the soil. There the diameter of the hole behind the original cavity falls behind with its
increasing length more and more so an automatic after sinking of the body does not come into
question (Fig. 8b). By this it appears that it is able to arrive during the work of the beetle with
no shaking of the body. As tools for the digging of the pit, the animal is served principally by
legs, which forward the soil in the already described species and manner. Now and again one
also sees the gravedigger push large lumps of soil with their neck plates.
The newly excavated cavity reached finally a length of 3-4 cm, with very small carrion from
about 2 cm, so begin again a new phase of digging activity. The carrion now becomes forced
into the prepared hole. This represents an important achievement for the beetles. For the body
must be brought to a small volume, otherwise it does not manage to pass along the narrow
passage. The start of this working phase the beetle orients itself under the carrion so that its
back is turned towards the underground, the abdomen tip towards the newly excavated,
funnel-shaped hole. The animal now grips the body with the tarsi and hauls it in with accurate
movements of the legs from behind and like-minded movements from the head and thorax
over itself off deeper into the hole. In most cases, the gravedigger grasps the carrion in the
middle, and pulls it into the passage while the LAST SENTENCE CUT OFF WHEN
XEROXING! this way, out of it the advantage results that the carrion is pressed together in a
very much concentrated mass. This is the first step in the rounding of the body which is
experienced later in the crypt. The more the carrion squeezes itself in the slope in the cavity
leading to the funnel, the more effective the effort of the working beetles which in order are
firmly it refutes gains on the funnel wall.

How successful this working method is, is shown by a chance observation. A N. humator
female buried a piece of horse meat some 6-8 cm in a sunk flower pot filled to the higher edge
with earth. By aiming at the base of the vessel, the female pressed its find through holes in the
base of the flowerpot which were big enough to allow through the N. humator female a small
example. By digging up the pot, the meat was held tightly only still with a small point, while
the remaining part had already passed the narrow opening. Only with difficulty did I succeed
in pulling back the meat the same way with forceps which is able to give an idea of the
working capacity of the beetle.

Still before the carrion has arrived at the end of the prepared passage it is extended in the
beginning successful way. Fig. 8d. Scarcely does this happen round a tiny piece, so the animal
shoves itself along between carrion and soil and walks round the prey and in doing so gives it
the advantage of a smooth body shape. In this way all parts of the body become dragged here
which still stick out of the main lump as for example tail and legs. Thereby the whole carcass
is stuck together more and more. Fig. 8e. Both activities, gradual lengthening of the passage
and hauling back the body with rounding off at the same time, alternates at length with one
another till the carrion has reached a certain depth. The thrown out material is, as already
Fabre (1899) observed, in the meantime thrown together. Only a small soil accumulation free
of plant growth betrays the activity of the gravediggers.

From the given description one presumes understandably that the carrion is not vertically
under the small hillock. Moreover, one regularly finds several centimeters laterally from the
small heap, loose earth distant in the soil (Fig Bb), an establishment, which again allows a
conclusion about the working method of the gravedigger that through digging under the prey
alone would hardly succeed in this place.

In the same way as N. vespillo goes about its work. so too does N. humator, N. germanicus,
N. investigator and N. fossor; N. vespilloides is an apparent exception. This inhabitor of drier
woods buries his finds only under mosslawns, leaf or needle litter which puts plenty of living
room at his disposal. His performance confines him thereupon to carrying the body through
the often several cm high ground covering till the carrion is put on free earth. If one finds
such a buried carcass, one can be certain to have the work of a N. vespilloides, Fig. 9. The
instinct of this species is so well adapted to the normal conditions of his environment that N.
vespilloides in a cage without ground covering is very rarely brought to reproduction as I was
able to experience with my experimental animals. However as soon as one puts plenty of
moss or needle litter in the rearing cages, they are no longer opposed to the natural course.
Thereby showing that the different phases of the digging event, which were observed in the
work of the other species in the essentials are also the same in N. vespilloides, only on
account of the loose condition of his materials differing in appearance.

A proof that the above described manner of digging is not confined alone to the named native
species, is given in an observation of Osten-Sacken (1862). According to him, N. americanus
ought to produce a long sloping in the soil leading to a pit in which he slips in the carcass.
There this beetle ought to be instructed in the main point of snake carrion, Osten-Sacken
considered this as he believed from that of other species, differing in digging method as
adaption to the shape of prey animals. However this assumption was invalidated for other
Necrophorus species proceed independent of the shape of the body likewise. Then the
extension of the prepared hollow with increasing size of the buried object takes root, becomes
understandable if the long passages which are manufactured as a rule in N. americanus to the
taking up of snakes, is seen, while they were able to ignore decidedly shorter cavities in the
native gravediggers average for small carcasses.

The depth up to which the gravediggers bury their finds, varies with the species. In some 60
cases the distance of the soil surface vertically to the roof of the crypt was measured. Thereby
proving that N. germanicus digs the deepest. This species was only observed in captivity
however, in which the animal moved the carrion without exception to the hottom of the cage
which was 20 cm deep; it is possible that this animal reaches greater depths in the wild. A
buried carcass of N. humator was found normally at 7.4 cm depth; the highest measured value
for this species amounted to some 13 cm. In the smaller interval from the soil surface one
finds the brood holes of the remaining species.

The time which the burying of the carcass requires is dependent on the size of the object, on
the condition of the ground of the physiologica competence and finally the number of
working beetles. The following observations which refer to a female N. vespillo give an idea.
The animal had carried a mouse in loose garden soil, with peat mixed through to a typical
depth for the species N. vespillo in 3 hours. In natural conditions in which a turf is penetrated
through, roots and stones are dealt with; this work takes up 8-10 hours in the above described
narrow glass terraria the beetles required likewise a whole night during which several long
rest pauses were observed. If several gravediggers take part in the work, so the working time
shortens in proportion to their number.

c .Rounding of the body and completion of the crypt.

One finds clumsily formed balls of carrion after the finish of the digging activity in the
ground. Fig lOa-f. show carrion taken from the crypt. From all forms shows the instinct of the
beetle to adjust the carrion as far as possible to a ball shape. The skeleton of vertebrate
animals frequently resists the rounding; from figures lOb and c this problem is obvious with a
bisected frog. Boneless pieces of meat which can be shaped anyway by the beetles keep a
perfect ball shape well. The expediency of this round form is obvious: with the smallest
surface area the ball protects the largest volume and therefore is particularly favorable to
preserve the food from desiccation.

Rounding of the carrion and building the crypt are closely connected in their origin, and arise
as a result of one and the same activity. For hours on end the beetle wanders round the buried
prey step by step in all possible ways. With its feet on the body and its back against the
surrounding earth, he lifts the mass continuously through stretching of its strong bent legs
away from itself. Through the counter pressure of the soil which opposes the back of the
beetle as well as the opposite lying side of the body, the carrion is strongly pressed together
and obtains the ball shape from the constant repeated kicking on all sides unless the strong
skeleton resists it. Necessarily through the constant pressure of the surrounding soil, a cavity
arises with hard walls in which the fully rounded animal body rests. It is the crypt. The
diameter is dependent on the size of buried object as well as that of the working beetles.
As was recognized already by Fabre (1899), one finds the small mammal's bodies hairless, the
bodies of birds featherless also without tail or steering feathers in the crypt. How this
appearance ought to be explained, as though the results of wind decay or as the result of
active work of the beetle, Fabre left open. However, none of these possibilities alone give the
truth; first through the interlocking of both factors arise several admitted results. Through the
ungentle treatment which the body receives during the digging activities in the passages of the
narrow cavity on the one hand the sedimentary hairs or feathers which are only still loose in
the skin of the body, fall out without direct action of the beetles; on the other hand the animals
subscribe directly to their removal: after rounding the body and at the same time completing
the crypt, the beetle continues to walk round over the carrion, however with renewed activity.
Like a spatula its head now stretches with slightly open mandibles over the surface of the ball
and free the food provisions of particles for the larvae and also still attached hair or feathers.
For hours on end, only broken by short rest pauses, one sees the beetle performing these
movements which are brought about through bending in of the initial thrusting out of the
head. In this way every square millimeter is cleared stepwise--apart from direct observations--
as is recognized from an enormous quantity of tiny points which were left behind by the
mandibles in the soft substrate and which by now cover the whole carrion ball. The falling
away--be it hairs or feathers--gather on the base of the hole and there become pressed against
the walls of the brood rooms by the gravedigger so that he covers them with a carpet similar
to the base of the crypt. This was observed at its most perfect on carrion which were buried in
soft peat, free from all hindrance. Under the conditions which in the wild are most frequent,
one finds traces of hair or of feather which have been lost for the large part already on the
way through the firm soil.

d. Egglaying

When the carrion is rounded, cleaned and resting with a smooth surface in a firm-walled crypt
(which to reach this portion of necessary activities requires in total between 12-48 hours) the
female proceeds to egglaying while the male repeats from time to time the above quoted
work.

As first Hain (1927) and V. Lengerken (1928) have stated independently the eggs of the
gravedigger are not layed on the carrion or in the carrion, but individually in the soil, as I
likewise confirmed. Unexceptionally I found in more than 100 cases, the eggs of included
species, singly in small earthholes which show a striking arrangement. They lie, as I was able
to observe in the species N. vespillo (76 cases) in some 6-10 mm intervals to both sides of a
round shaped, smooth-walled against shock, indeed little resistant passage which the mother
animal has dug from the crypt out into the soil. It is analogous to the parent gallery of the
barkbeetles, a space is quite big enough to let the female move through it unhindered. The
passage runs horizontal with weak bends to the crypt in the soil (fig. 11 and 12). Its length
depends on the number of its flanking eggs, whic in N. vespillo can fluctuate between 1-24,
but averages about 14-15 eggs(Table 3.)

The small chambers in which the eggs lie, are built in all probability through the activity of
the extended laying apparatus of the female, which releases frothy drops of secretion at the
same time the small hole receives an egg. The barrel shaped eggs are white and shining. The
size directly after egglaying, an average of 2.958 x 1.836 mm, a result which was calculated
from the following values (table 4).

e. Moisture and digestion of the body

After egglaying the female hurries back again to the crypt and moves the pile of earth from
building the mother passage to the side while it wanders round the carrion in the explained
way and presses the loose earth to the wall. This as all previous earth working of the
gravedigger requires for success a certain moisture of the ground which the animals find in
the wild at their living places and which is carefully supplied in the terrarium.

After the hole is tidied up, the female begins with a remarkable activity: at the highest point of
the carrion ball the animal digs a circular hole which on average measures 1/2 cm in diameter.
First the surface of the body is bitten to pieces with the mandibles then the fore tarsi grip
extended into the gaping split. With help from head and forelegs which represent, with the
strong tibial spurs, excellent instruments, the cutting circle is pulled apart as with hooks so the
opening enlarges to a small crater. In this hollow the beetle lowers its head and begins to eat,
as is concluded from the movement of the mouthparts. It is the first meal taken from the
buried body. This was certainly not remarkable for an eating place and was found repeatedly
at the corresponding point of every carrion ball and at the same time. The place was at that
where later the young sliding larvae gather. At the end of about 1/2 hour the beetle appeared
satiated. Still she never moved away without shutting the hole and finally smoothing the
surface for a long time with the underside of the head. In fact this precinct in no way differs
from her later surroundings. The excrement of the beetle, after completion of this work, was
distributed now on the upper half of the ball. The animal which put the anal opening of the
carrion surface to this purpose, proceeded slowly over it then while the abdomen carried out a
sideways pendulum movement in this way the liquid excrement succeeded in making a snake-
like track on the carrion which is recognizably shining with moisture and first disappears after
some time through blending.

In the change with hours of peace which the animal spends pressing tight the meatball, this
work fills up the period in which the brood passes through embryological development.

The question of whether this instinctive action is of use for the descendants ought perhaps to
be answered in the affirmative. The smearing of the liquid excrement on the ball of carrion
doubtless serves the use on the one hand that the food provisions, already protected against
the danger of drying out to a certain degree by their shape, remain moist with greater
probability as this then also found, always independent of the weather, moisture glittering in
the crypt. On the other hand, for this reason, it is possible for the carrion ball to become dried
through with the scent of animals which, is demonstrated in return for our ability to smell the
gravedigger in the excrement. Be it that the Necrophorus scent acts to scare away other likers
of carrion or also the attraction of the small larvae which emerge from the egg scattered in the
neighboring soil, which would also conceivably be an advantage in this respect.

But also the taking up of food of females represents an action of broodcare. Presumably, if the
already previously mentioned supposition of a preoral digestion of the gravedigger is right,
the food begins to decay with the feeding activity of parental animals sharing digestive
enzymes with the environment. Thereby, the crater becomes deeper and deeper with the
passing of time on the inside of the ball of carrion and the female renews the deposition of
fresh secretion which accelerates the otherwise slowly advancing process.

That it is a matter of the disclosed consideration, not pure speculation is proved by the
behaviors of the female several hours before the brood emerges. At this time the up to now
calm movements of the animal become hasty--nearly abrupt as at the end clockwork. The ball
of carrion hurriedly climbed on and the eaten area opened with impetuous movements of the
head, which allow the onlooker to suspect rapid destruction of work. Without turn, as hitherto
again to close the beetle now walks through, in many cases softly chirping, the mothering
passage from which it turns back to the crypt after several minutes. After the beetle has
walked through the whole subterranean cavity one soon finds her sitting and eating again on
the ball of carrion at the crater thereupon arranging the opening in a regular circle. This
activity of the female which coincides roughly with the end of embryonic development in the
brood, is closely connected with the appearance of the descendants. The fact that the crater is
opened at this time and is no longer shut, proves that external digested food is intended for the
larvae. The walking through the mother passage which is repeated several times in the course
of 20-30 minutes is suitable for facilitating the hatching larvae in the soil in finding the
carrion. On the one hand the scent of the female makes it possible for the larvae to orient
better, on the other hand the beetle roams out in the space of passages, throwing pieces of
earth. Thereby the young larvae opened a way in which they are successful in obtaining a
quick and certain food source.

f. Male participation in brood care

Brood care is independent of the sex in sexually mature gravedigger. Every male just as every
female, is able to bury carrion as experiments in which the sexes were isolated have shown.
Observations in the wild are however in contradiction to this result. Males which set about the
digging work without the presence of a female stop their work after a short time in order to
attract a female through "ventilation". If there is no result, the carrion remains uncovered or
buried only shallowly. This indicates the dependence of the male on the presence of the
female. We verify this supposition with an observation occurring when a female was removed
after termination of the digging activity and the male was left alone. In this case the body
remained unaltered, a fact whose meaning is unmistakable. A mistake by the female affecting
the brood and long preparation of the body would cause a significant loss of time. The pair
remains together so that the male participates in all the females work. However, in the
majority of observed cases the male left the crypt in the interval between egg-laying and the
hatching of the descendants--either of its own free will or in the face of a hostile attitude from
the female. Only exceptionally is the male still found together with the young brood, perhaps
because the female is not always successful in driving the male away. However, N.
vespilloides males are the exception to the rule because males are found with broods in nearly
half the cases.

Fabre (1899) supposed that after preparation of the body for the brood the pairs left the crypt.
The majority of obtained observation in the wild oppose this (Table 5.) In the woods and
meadows in the surroundings of Frankfurt suitable carrion was laid out for burying. Roughly
48 hours later I visited the same place and noted which carcasses were buried by the
gravediggers which was recognized by the slight but typical alteration in the ground. After a
further 7-9 days during which the building of the crypt, the preparation of the carrion is
completed the larval development must be more or less advanced, the broodrooms were
cautiously opened and the finds resulted as follows--table 5.

After the combination the larvae from I to III are to be found and the last stage nearly always
in the company of a pair or single female in the crypt. The moderate number of exceptions (4
out of 59) lost most meaning when one thinks how easily the results could be negatively
influenced by digging.

Also Fabre once found by opening a crypt of a Necrophorus pair together with the larvae.
This observation which contradicts his acceptance that the parents shall leave the crypt before
the hatching of the brood, he explains in the following way: "La periode de la parse est
maintenant fise, et la victuaille est copieuse. N'ayant pas aube chose a faire, les romanicus se
sort attables a cote des noumsaous" Fabre (1899). The view arrived at in this quotation that at
the end of the brood period many parents stay with the larvae on the carrion merely to satisfy
their own feeding requirements has till today prevailed. Through my establishment that the
parents, above all the female, are found regularly and during the whole breeding period in the
company of the larvae, this is refuted. But this presentation becomes entirely valid, if itself
restricted a close relationship between the female and her descendants is disclosed over the
care of the brood. Was the species already as the mother attracted the scarcely hatched larvae,
already unusual in the beetle order, thus excelling in the activity, which happens in crypt
between the female and her brood all that one knows up to now in this respect about
indigenous beetles.
The following observations were described using the cage in Fig. 7, a glass cage which made
insight into the crypt possible. After completion of digging activity the cage was brought from
its vertical position to a side position, so that the beetle was given the necessary room in the
horizontal to build the crypt. The larval nest was drawn from the natural brood room and
brought together with the pursuing female into a small flower pot, which was shut from above
by a damp cloth. During the observation in softened light this was removed or replaced by a
glass disc so as not to disturb the female. Before I was able to turn towards the mentioned
activity the brood must be accurately taken into consideration.

g. The brood at the time of larval development.

In the extraordinarily small time interval of 7 days for the development of beetle larvae, the
necessary weight and size increase of Necrophorus larvae to form the adult occurs. The speed
with which the development proceeds becomes more astonishing if one takes into
consideration the achieved performance of growth in this time. While the increase in length
which normally is in no way exceeded (the N. vespillo larvae grow an average from 0.5 to 2.8
cm), the increase in weight is in proportion to developmental time. The enclosed curve Fig.
14, whose values were obtained through daily weighing of the larvae with the help of a
Sarbornis weighting analyzer, gives an insight into the weight increase of three N. vespillo
larvae during the 7 day developmental period. On the X-axis is the age of larvae - calculated
from the moment of hatching-stated in hours. The Y-axis bears the weight in mgs. The first
weighing was carried out immediately after the hatching and before the first feeding and was
for larva I 0.0035 g. After 7 hours the starting weight had doubled. The last weighing took
place shortly after the wandering of the larvae from the remains of the food carrion, and
showed for larva I 0.3037 g, where afterward the total increase in weight was roughly 100
times the initial weight.

Disregarding the statement of absolute values, the graphical representation gives us

information about the rate of increase of weight of age stages. Two rapid molts H1 and H2,
separate the three larval stages I, II & III from one another (Fig. 15 for N. germanicus.) Larval
stage III not only takes up the longest time in the larval life of Necrophorus, explaining why
one encounters mostly larvae of this age in the wild but also has the greatest rate of increase
in weight, which probably during the first 48 hours was somewhat held back through the
energy consumption in the ,short interval following moulting. The drop in the curve towards
its end (from L1 from 0.3573 - O.3037 g) has a double cause. Firstly, the larvae cease feeding
shortly before leaving the crypt and more importantly they empty their previously full guts.

The three larval stages, which do not significantly differ with regard to morphology, can be
recognized through certain clear differences in their behaviors. At the end of embryonic
development, which takes 5 days, the larvae of the grave digger hatch from the egg. In order
to free the sticking egg cover from part of their body, the animals try with intaking
movements of their longitudinal axis, in which they succeed in about 1 min. Thereafter the
egg chamber is left immediately and the larvae walk to the stored up food. The newly hatched
larvae are white, very agile creatures, which with the help of six legs and their hind segments
which function to push behind, move away smartly and restlessly with antennae held
upwards. The majority break through the earth-layer which separates the egg chamber from
the mother passage following the leveled path of the female and reach the crypt in this way in
a comparatively short time, where they are attracted to the scent of the carrion and that of the
female. Single individuals also try to succeed directly through the soil by the shortest route to
the provisions of food where they likewise succeed. Indeed then the small larvae require 2-3
hours to cover a stretch of 4-5 cm which it does in constant motion, only stopping for a few
short rests. Every small stone poses an obstacle to advance and must be gone round, till
finally the animal falls into the subterranean crypt by shoving through the crypt wall. At once
it climbs the ball of carrion to the highest point and there enters into the crater together with
its brothers and sisters, which follow the same instinct to the same place.

That the freshly hatched larvae, in fact react chemotactically to the smell of the carrion which
forces its way through the mother passage and the soil to them, is shown by the following
experiment. A wall of carrion taken from one of the crypts was laid on a piece of filter paper
and the center formed a circle of 25 cm radius from the periphery in regular formation were
distributed 10 N. vespillo larva of the first stadium. Each larva was covered with a small glass
dish under which they were free to move and could be put in any course. The experiment was
carried out in a room in which no noticeable air current was recognized. The illumination of
the otherwise dark room was done with a lamp with strongly diverted light. After lifting off
the glass dish all 10 larvae wandered in roughly circular courses to the ball of carrion.
Thereby with the exception of 6 larvae, all experimental animals appeared at the carrion after
short searching movements in the direction, which on the whole were detained. Only 6 larvae
initially moved themselves in the opposite direction in order to later hurry to the carrion in
nearly straight movement. In the immediate vicinity of the carrion they met several larvae.
These came to the residence shortly, whilst these move round one another, making their
separate ways without exception to the source of the smell. The required times for the ten are
given in the following table.

The experiment was unsuccessful when repeated with larvae III. Then their movement
resembled a random walk in various directions which took the larvae away from the carrion,
as well as toward it. Often the smell of the carrion was not received by these larvae over large
distances, although when in the immediate vicinity these animals appear to be able to perceive
the carrion and walk into it.

The aimless wandering around, which restricted the experiment involving older larvae, caused
the animals to meet frequently. Contrary to the behavior of larvae I, larvae III remained
closed to one another when they came into contact; frequently they were attracted by their
own smell of carrion. This behavior gives rise to a cluster of larvae, which is in constant
motion through the circling around one another of the larvae and as a whole rolls slowly from
the place. Fig. 16 shows one of these groups of larvae which originated in this way. Although
in this experiment the food is only 9 cm distance from the larvae heap, the animals remain
crowded together, till after an hour the carrion was put back 2 cm distance. First in this
moment the closely pressed group loosened and the animals clambered singly onto the
carrion. This observation shows for the first time that the old larvae are attracted to the smell
of carrion only from a short distance, in striking contrast to larval I stage.

With increasing age of larvae, weakness of the ability to smell can occur in connection with
the altering condition of the larval environment. Thus the needs of the larvae correspond
throughout but the problem of finding carrion from greater distances falls to the young larvae;
larvae II and III suffice with the kind of attraction which holds them together in close
combination with the food.

The portion of the ball of carrion from which the attraction radiates is the higher pole,
prepared by the parents, with intestinal secretion to form the crater. Here the young larvae
appear and the old larvae remain. From here the consumption of the carrion starts. How this
takes place is again remarkable. While the brood are in stage I and II they contribute, through
their feeding, to the broadening of the crater in which the young larvae lie horizontal in the
casing with their heads pointing outwards. Soon the larvae advance to their second moult
inside the body. They rummage about and feed themselves deeper and deeper into the carrion.
As long as there is no lack of food during the 7 day development, instinct allows the larvae to
leave the outside of the ball of carrion undamaged -indifferent to conditions - so that they
finally rest in a nest with both shelter and food. In the wild in which the grave digger directs
in his brood care on small bodies, the center layer is formed from the skin of the corpse giving
mechanical protection against advancing robbers. The rounding of the ball therefore retains
also the shape first bestowed on him. presentation is given in Fig. 17 of how the larvae are
able to disappear into th depth of the body. If the provision of food for the brood is less
plentiful than in the quoted example, the larvae are not able to completely conceal themselves.
They then put only the front of the body in the carrion, while the hind end remains visible
(Fig. 18).

The larvae II stage is also found if the ball of carrion was big enough for them to allow a
scattering of the larvae inside its mass, nestled uniformly crowded with one another. It is true
the appearance was able simply to be gone back on, that the larvae from a close undescribed
precinct from the crater in which they lie close together, begun the advance into the ball of
carrion. The following observations however give us the right to perceive in every appearance
the utterance of a particular instinct.

It appears namely that larvae III are qualified for the pursuing of oral drips : if one lays a
larvae in a cupped hand, they leave every where small brown drops where they touch their
closest surroundings, tasting with the head. That allows conclusion of extra intestinal
digestion. It appears that the quantity of preorally digested food I suppose is used for the
young larvae not for the older brood. If however the larvae III really digest preorally then it is
able to be of help to the animals, if they press together with their heads in a small circle to
feed. Why?

We start from the condition of an isolated single larva. The deposited digestive secretion from
it is pressed constantly in all ways loses very soon with distance from outlet point the required
concentration for effectiveness, through a certain part of the digestive secretion going for the
conversion of food. Otherwise by some sense pressing together of the brood. Now the
resulting fall in concentration from the oral drops of each larva overlaps with the original fall
in concentration around the quantity of secretion of the neighboring larvae. At the place of
overlap however much too small quantity of enzymes reach effective concentration to be able
to contribute in this way to the quick formation of the food.

This presentation finds yet further support in the fact that one finds also other extraintestinal
digesting insect larvae in the same group. Larvae of Miastor metraloas were found to lie
radially, the head reaching from the middle of the circle, by Springer (1917) who showed the
above mentioned advantage in their position.

If, however, the habit of remaining bundled together of older burying-beetle larvae is an
advantageous instinct, it is doubtful through which kinds of attraction they would intervene. It
maybe a matter of mutual thigmotaxis of the larvae. It is possible that the ball on the

outside of the carrion ball happens in older larvae is not explained merely as a reaction to the
outgoing attracting smell, but from mutual attractive contact playing a part. The thigmotactic
attraction of the larvae prevent in a suitable way any dispersion of the animals. Actually it is
extremely rare to observe a larva leaving the nest prematurely. Only from time to time one
can surprise the larvae as they hang their hind ends far out from the crater edge, Fig. 19. This
is the attitude which the older larvae maintain for defecation. The emerging pellet of
excrement falls off on the outside of the ball of carrion containing a mixture of food and
excrement and with that prevents a reduction in the nutritive value of the carrion and at the
same time strong soiling of the larvae is avoided.

Further, a most remarkable instinct of the larvae should now be mentioned in the next chapter
in connection with certain instincts of the mother and the parents respectively.
h. Feeding of the larvae.

Soon after the first larvae hatch and have appeared in the crypt the female employs the
attraction of the brood from the mother passage and makes for the ball of carrion in which the
larvae lie thickly pressed in the crater-like cavity. In permanent motion they crawl over and
through one another. Their mouthparts touch the carrion - but only tasting. They never eat
from there; the transparent gut remains empty. One gets the impression that the lengthily
prepared substrate will be in no way be the searched for food of the brood.

Immediately however the observer presents himself with an unexpected scene. Scarcely has
the female approached the crater when all larvae lift up the front of their bodies steeply, so
that their legs grasp in space. The beetle remains standing directly over the brood and waves
her forelegs with shaking movement on the carrion or sometimes on the larvae, which flock
round her head. Now the female closes her mandibles and rapidly lays the head of a larva
between the jaws itself pressing closely to the mouth opening of the female. If allowed the
chance, then one sees a tiny drop of brown liquid on the mouth of the mother in which the
larva infringes. But already after a few seconds the female turns and immediately tries to
reach another larva with his mouth held high. Without doubt, the brood is fed by the female.
Correspondingly the initially white, almost see-through larvae after contact with the mouth of
the mother become dirty brown, a change which gradually progresses from the front right to
the hind end of the larvae.

However, with what does the female feed the brood? Is the food source the gut or a
specialized gland. Specialized glands, which are rare in beetles, was not found, the feeding
juice is derived from the gut. The administered substance is also-gut contents presumably a
mixture of food and gut secretions. In fact, the female takes several feeds of carrion at the
crater shortly before the appearance of the brood. The quantity of food taken up must be
exhausted with time both from feeding the larvae and from feeding the beetle itself. To
continue the activity of feeding for several days, the female is compelled to fill the foregut
anew. With increasing age of the larvae, the female fed with increasing frequency
immediately before passing food at the crater, an activity which claimed up to 1/4 hour.

The feeding can be observed during the first 48 hours of larval development, but is seldom
repeated later when the brood gradually are made independent. The young larvae beg
impetuously for the food juice of the female. Three to four larvae often reach her at the same
time crowded before her mouth and dispute their right against each other to this place. Fig. 2,
which allows a glance into the inside of the crypt, shows directly the feeding of highly erect
larvae. A second is attacking to drive away the first. Larvae which by chance find the hind
end of the beetle, climb about carelessly as if searching for the mouth. Most however,
immediately set out for the head of the female. Either the larvae leave the crater to this
purpose, in order to lay back outside themselves hurriedly the prey or they make their way
along on the ventral side of the female. Here by the larvae showed themselves, lying on their
backs, forward under the beetles and more likely did not rest till their mouths had found the
provision of food. Fig. 21 shows this frequent performance. Rarely and only in the Larvae I
stage one catches sight of an initial attempt to reach the mouth of the mother, namely from
those forelegs held out. However, what I saw never led to the result: The small animals
clambered with great skill onto the tibia, and lifted themselves with their sucker - like after
segment onto the femur. Stretched forward, one then sees the larvae searching for the mouth
of the mother.

Fowlers declaration (1912), which characterized the larvae of the grave digger as inactive
maggots, is then in no way correct for the young larvae. However, larvae were not constantly
attentitive to females. Moreover, a pattern of use and disuse develops in relation to the
molting pattern.
Newly hatched larvae are apparently completely dependent on the female for food. They beg
extremely impatiently and follow the female for short distances over the carrion with great
dexterity and ability. When the young brood scatter themselves even more often over the
upper surface part of the ball of carrion, they gather again in an even shorter time in the crater
as soon as the female turns back there. At the age of 5-6 hours the larvae then begin to feed
independently. With heads turned outwards They lie close to one another in the crater. Their
mandibles, whose points have meanwhile colored light brown, beat continuously in the
carrion. These larvae are still ready in the vicinity of the female to receive food, and will act
as in the first hours of life. Yet they rarely attempt to follow the mother in her walks around
the carrion; rather they use her absence in order to eat independently. Also they stay
permanently on the inside of the boarder of the crater until the first moult.

Rarely is this executed so the larvae arrange themselves again searching upwards, without
also only undertaking an attempt to take up food independently so behaving just as in the first
hours of life. After the first moult there is a second period of greater dependence on the
female followed, 2-3 hours later, as in larva I, by greater independence. After the second and
last moult the larvae is directed in feeding by the parent, in order to quickly complete female
care, and finally to grow completely independent.

Conditions during the feeding time, which stretches roughly over the first 60-72 hours of
larval development, are summarized in Fig. 28. The ordinate shows the changing dependence
of the larvae on the female for food. On the abscissa is marked the moment of hatching, the
first and second moults, and the end of the feeding period. Three peaks show that the time
span is distributed over three periods in which the brood taxes the female for food: after
hatching, and after their first and second moults. Between periods of greatest dependence on
the female the larvae feed independently. The female accomplishes certain typical repeated
instinctive actions during the entire feeding time in the interval of 10-30 mins. she appears at
the crater. The female usually starts feeding, by moving her first pair of legs, comparable to a
running on the spot. It appears similar to (The "Mittern") an alarm signal as a sign for the
brood to begin feeding. If this hypothesis is correct, the Mittern ought particularly to be for
the older brood, hidden deep in the carrion, can be attracted through quick shock waves. In
fact the larvae III which are of no further interest to the female also appear at the food after
the Mittern, if they have not already outgrown motherly care in this respect.

After the brood is alarmed the female proceeds immediately to feeding. The most remarkable
of the activities has already been revealed, only a little still remains to be supplemented. The
female's pose during feeding, varies, depending upon the most favorable height for the
begging brood. While she bends deeply to young larvae, her head and neck plate show a
strong inclination downward, the long axis of the body climbs forward moving around to feed
the higher reaching older larvae. The weight of the female rests in this position on the last pair
of legs. The forelegs are either free and used to ward off obstructive larvae or grip the tails of
larvae being fed (Fig. 20). That the female holds tight to the larvae on this occasion shows in
the following chance observation. An N. germanicus larvae III was successful in begging for
the food juice of the female at the edge of the nest. During the feeding they fell down. The
female, continued to feed the larva which was lying on its back held tightly between the
female's tarsi. Indeed, this pose of N. germanicus is nothing exceptional; they often take up
other poses in the consumption of fly maggots.

The length of each feed to a larva, which was measured by the period in which the head of the
larva rested between the mandibles of the female, is small. In N. germanicus the feeding of
older larvae can claim up to 18 seconds; but for all Necrophorus species including N.
germanicus, feeding times are generally 2-4 seconds and not longer than 7 seconds. Feeding is
terminated by a scarcely visible sideways movement of the female's head. The mouth of the
female consequently slips from the larva and becomes free for another larva. Five to six
individuals are fed in immediate succession. After a break the female again begins feeding
exactly the same way. The larvae of a nest are provided with roughly the same amount of
food juice from the female. This is the suitable succession of a typical feeding method.
Everytime the beetle turns away after a short feed the equal distribution of food to a normal
sized number of individuals is favored. Yet the instinct concerning this is rigid and she allows
herself to be converted easily into useless activity by the experimentor, who reduced the
number of larvae. Nevertheless, the female acts in completely the same way. Mittern feeding
and again turning away following one another at typical rates at which then the beetle often
enough holds her open mandibles in space, and just turns away in the moment in which one of
the larvae approaches.

i. Ecology of Feeding.

Now the question is raised of what importance for the brood is feeding by the females. For the
larvae, is it a matter of an activity necessary for living or of more secondary importance? This
question can be decided very easily experimentally. If the female is removed before the first
larval feeding, and if the larvae are completely dependent on the female, death of the entire
brood must result. It appears that such a radical result does not happen. Most of the larvae
grow up as if fed by the female.

The other hand points to two facts that the absence of the female in any case operates
disadvantageously. Whilst in 8 rearings, which together included 89 larvae, under the care of
females all individuals completed the 7 day development. From 147 orphaned larvae which
were distributed over 14 nests, 16 animals in stages I & II died, so that in each rearing one
dead larva was found on average. It is almost certain that the early death of particular larvae is
connected with the absence of females, primarily pointing to the loss of food. Secondly it
appears that without the female grown up larvae are able to pupate but very rarely do young
beetles hatch from the pupae; from 33 such controlled pupae only one hatched. Under the
unfavorable conditions of artificial rearing, the brood perishes in the pupal stage in roughly
half of all cases even though fed by the female. The very much higher death rate of the brood
raised without the female is certainly attributed to absence of parental care. Therefore it
appears that the larvae are able to develop normally without the help of a parent animal, but
that through feeding they are favored or protected.

Is what exists now an impairment from deficiency of food? Next comes the idea of something
qualitative. The feeding female was able to supply the larvae some substance - if only a small
quantity - which is important for development. This assumption was not refuted but it was
unpredictable. It is assumed, that the deficiency of a specific substance in the course of
development represents here completely expressed and believed to itself more serious
aftermath if this is in fact the case in orphaned larvae; then still the larvae are able to complete
their development to adult without the food of the female. Moreover it is not very probable, as
p.570 showed, that the quality of food differs from that which the female has prepared in the
area of the crater: presumably both show the same kind of mixture of carrion and gut
secretion.

More likely the impairment is quantitative. Possibly the quantity of food which the female has
already mixed through with gut secretion before the appearance of the larvae, does not suffice
for the brood and must therefore be supplemented through feeding by the female. However,
the feeding of the brood begins immediately after the appearance of the larvae when a
deficiency cannot yet exist in the predigested food.

A second more distant possibility exists, why a sufficient quantity of food was missing for the
larvae in the female's absence. The gravedigger used only small carrion as provisions of food
for the brood. This specialization brings two-fold advantage: firstly, small carrion is more
numerous than large: secondly. the beetles are only able to bury and work bodies of a small
size.With these advantages however must be taken a distinct disadvantage: the rapid
transiency of small carrion. The decaying process which proceeds from the surface towards
the inside as oxidation events are tied to the equivalent of oxygen, in small carrion it leads in
time to complete destruction, which would not be the case in large bodies. That only small
carrion are used has been proved; this assumes that the development of larvae is highly
accelerated. In fact the development time of Necrophorus larvae is strikingly short; a 7 day
development with more than one year of each beetle larva subject to quality food material.
Even the carrion and meat eating larvae of closely related Oecoploma thoracica took an
average 34 days for their development (Heymons). This larva, unlike the Necrophorus larvae
is not solely a carrion feeder, but can always breed on new food when freely moving around.
Thus, the 7 day development period of Necrophorus larvae depends not perhaps on quality or
quantity of food, but is fixed genotypically so that with or without the female, development
takes 7 days.

The necessary and obligatory shortness of the available time to develop however requires
complete exploitation. Just how are hatched or freshly moulted Necrophorus larvae unable to
independently take up food because of the soft cuticle of their mouth parts. Through this
condition repeated failure in the supply of food was effected, for the quantity of food which
the larvae can take up in the time allowed in their condition, sinks below the optimum. It is as
Fig. 22 shows, just the named development phase in which the brood is fed exclusively by the
female. Therefore it is apparent that the female prevents through the feeding a break in the
supply of food. The benefit lying therein is by no means insignificant. During the 60-72 hours
feeding period the larvae are fed copiously by the female half the time. We see also that all
appearances after the feeding instinct of the female serve the purpose, to secure for the larvae
the necessary quantity of food for their development despite the--offered of other ground--
shortness of their development time.

The short development time results in yet a further use for the species. The carrion which the
grave digger must visit to feed to maturity and reproduce are distributed sparsely in distant
parts. The problem of finding carrion becomes all the greater the longer time placed at the
beetles disposal to visit the body; this time however becomes extended through an
acceleration of development. Above all the short duration of development is of use for any
species, such as N. vespillo or N. vespilloides which rears two generations in a year. The
beetles of the second brood, which in N. vespillo hatch in the middle to the end of September,
still find at this time favorable conditions to feed to maturity, which would become absent for
them with later hatching, so that they would have to overwinter without feeding to maturity.

j. Defense of the larvae and their food.

Also in the period between feedings or after the conclusion of the entire feeding period one
sees the nursing female sitting mostly over the crater (Fig. 23). Very conspicuous are the
sudden movements with which the female turns here and there as the single entrance to the
larvae might be protected by the towering female in all ways against possible enemies.
However what are the enemies of Necrophorus larvae? Above all the male is mentioned. The
male is already drawn attention to by Soffel (1904) as an occasional carrion eater. Although I
was never able to catch him red-handed, no doubt the male ate up carrion and larvae in very
many cases of my experimental nests. From an area of about 1 hectare he had expelled
outright, after robbing in 3 days not less than 36 buried carcasses which had fallen victim, as
was gathered from the existence and development of typical male tunnels.

Naturally what happens when this superior enemy comes face to face with a defense of the
brood does not need to be questioned. This is still of value for the hostile insects which either
as predators hunt after the Necrophorus larvae themselves or as carrion-eaters were able to
impair the closely measured food for the brood.
One of the predators is the large Staphylinus olens Mull. It was found in 6 cases in
experimental nests of N. vespilloides, among them 2 times in 2 examples. In all cases the nest,
in which I had established a normal number of young larvae (I & II) only a few hours before,
was found empty and with no trace of the larvae in the near surroundings. On the other hand
in two of these cases elytra and legs lying around gave away the fact that females had also
fallen victim to the superior enemy. Therefore the successful defense of the brood against this
enemy in the event does not arise.

To the predators is further counted the Staphylinid Philonthus whose determination Herr.
Prof. Dr. van Ender undertook in a loving way. Necrophorus eggs in the immediate vicinity of
an N. vespillo nest in the soil were brought up together with the Necrophorus eggs but
without the presence of Necrophorus females. They hatched out extremely lively Philonthus
larvae clambered onto the ball of carrion, seized and consumed any Necrophorus larvae which
had already arrived there. After 2-3 hours pause, in which the predator concealed itself in a
hiding place, further attacks on N. larvae were observed. In 24 hours the whole N. brood was
destroyed. A further look about by the visibly growing Philonthus larvae at this time failed as
any suitable food was absent. Carrion and pieces of meat were scorned as were as older fly
maggots. Suitable food, especially Necrophorus larvae, were unfortunately not at that moment
at my disposal. Finally the restless, hunting Philonthus larvae died with wide open mandibles.
However by this time the establishment of the species was frustrated so a possibility to
determine the species of the larvae of the genus Philonthus did not exist at the time.

Philonthus eggs were found later several more times in the vicinity of freshly laid
Necrophorus vespillo nests, in total 17 times. The danger to the Necrophorus brood through
predatory Philonthus larvae cannot be totally averted through the vigilant female, but still
substantially reduced. For in the wild I saw repeatedly, as the beetle succeeded now and
again. she seized and killed the visible predator.

The female had further success in defending her brood against the hostile carrion-eaters than
against predators. Additionally, one is justly allowed to mention all those insects which feed
themselves or their descendants on carrion. From such were used the following species for
experiment:

Necrophorus vespillo in 32 experiments

Necrophorus vespilloides in 32 experiments

Geotrupes sylvaticus 18

Oeceophoma thoracica 2

Silpha obscure 1

Lucilia caesar 1

In a weakly lit room a ball of carrion taken from a crypt, in whose insides were N. vespillo
larvae of stage III together with the attendant female were put in the middle on the floor of an
uncovered glass terrarium (52 x 40 x 27 cm) whose floor was covered up to a height of 1.5 cm
with earth. The experimental arrangement allowed the hostile insect to be brought into the
vicinity of the nest without the nursing female being disturbed through other than from the
new arrival attraction.

All experimental results agree that a reaction of the female first becomes perceptible if the
enemy was or even had climbed in the immediate vicinity of the ball of carrion. With their
particular generic name the defending female made without exception the usual way with
Necroporhus in the fight. From this same species were the dung beetles (G. sylvaticus) driven
away, while Oeceophoma thoracica and Silpha obscura were driven away only several times
in all cases and for this reason finally expelled. Only Lucilia caesar did not let itself be finally
expelled. Although the Necrophorus female repeatedly rushed on the fly with quick
movements, and frightened it away for moments, they always moved back again, till it finally
succeeded in laying some eggs on the carrion ball. These were sometimes eaten later by the
female as also fly maggots which developed in the ball of carrion are consumed by the
female. How effective in fact must be the warding off of enemies of this species for in the
Frankfurt woods are found almost on any free-lying carrion Geotrupes, Silphidae and often
insects among them not rarely also feeding Necrophorus. As soon as digging Necrophorus are
present, the remaining visitors disappear. So have I then, never during the whole time of my
observations been able to discover any intruder within an intact crypt. This observation which
covers also the very shallowly buried carrion of N. vespilloides, is all the more remarkable as
Geotrupes was also found on the carrion, that I had covered several cm deep with earth and
leaves.

k. Maintenance of the crypt.

The necessary opening of the crypt in the wild for observation led to the establishment of a
remarkable regulatory instinct of the grave digger. The animal repairs the disturbed brood
room. The following experimental descriptions serve to show the below surface observed
activities.

A larval nest which belong to N. investigator becomes laid free through cautious removal of
the crypt roof. The female which in the first moment was escaping, soon appears again,
hurries round about the carrion, clambers up on it and tastes with the antennae the edge of the
crater. Then it suddenly leaves the ball of carrion, climbs the crypt wall sidewaysand
advances into the adjoining soil. Already the grave digger is half hidden in it, then she pauses
and with stronger and from backwards directed movements of the hind legs hurls back loose
earth. At the same time also the upper half of the beetle falls back into the lying soil. Then the
female pauses, turns back to the crater and tastes alternately with the antennae the freshly
raised material and the still partially uncovered crater. Then still once again begins the
described spectacle at another place. Now is also plain to been seen, that the working animal
through raising and lowering her thorax shakes the earth mass lying over her and through that,
brings it sliding off. Soon the ball of carrion is completely covered with earth. The female
ceases working and disappears under the spilled material. Once more opening the brood hole
at the end of 4 hours shows that the crypt is again completely made: the ball of carrion lies in
a cavity with smooth walls as before the disturbance.

The typically expedient behaviour of the grave diggers remains throughout in the frame of
demands which in the wild are put on the animal. That the Necrophorus sank carrion only a
few cm in the earth, may have occurred frequently enough that from the natural reasons,
perhaps through the step of a grazing animal or a heavy downpour, the crypt fell together and
the brood often lay there abandoned from damaging weather and hostile persecution.

It is unknown which kind of attraction caused the repair instinct. Three different kinds of
attraction which singly or in combination were able to work, lie in the realm of possibility.

1. With the disturbance of the crypt often a sudden lighting of an entirely dark brood room
occurs, consequently the possible attraction to an attractive light exists.

2. The stagnant air in the crypt over-ladened with waste products, for example CO2 and H2O,
would essentially entirely change through the inflow of fresh outside air. The causal attraction
would in this instance be primarily chemical.
3. A crypt can be sunk without connection with the outside world, thus offering another kind
of attraction. As a result of the original kind of crypt the nursing female fits in the space
between the ball of carrion and surrounding earth, touching the crypt wall only lightly with
the back. By collapse of the hole, the light on the back of the beetle either suddenly intensifies
or is completely lost. Any intensive change of this kind is able to be the cause of attraction.

Occasional support that, for the mentioned attractions of contact for the activation of the
repairing instinct are of decisive meaning. A final difference however falls requires thorough
experiments which ought to be made in the coming summer.

1. Defense of males at the nursing.

Although the males in the majority of cases were no longer found together with the larvae in
the nest (Table 5), N. vespilloides and N. germanicus were observed feeding, and presumably
males of the remaining species do so as well. The fact that male grave diggers are also
equipped with a broodcare instinct, is not surprising as the participation of the male in the
broodcare in beetles is a known occurrence. One is reminded of the group of coprophagous
lamellicomier amongst whose representative are found numerous genera with highly
differentiated broodcare in which the male of many species participates. I]hy, inspite of the
qualification of male grave diggers for broodcare, was contribution of the male only observed
in exceptional cases. Two explanations are possible:

1. The male was able after fulfillment of his task, which on the one hand in the capacity of
working to bury the body, on the other making the female to fly away in search of another
carcass and through renewed ingression of task be useful at another place.

2. Certain signs suggest that broodcare instinct of the male is less pronounced than that of the
females. It is conspicuous that feeding male offers his food juices to the brood at greater
intervals than the nursing female. While the young beetles groomed themselves often and
leaning of the nursing grave diggers was confined to the cleaning of eggs and antennae. When
the larvae leave the nest, one sees the female respectively the pair completely dirty (Fig. 23)
and often with mites sitting dotted in the rest of the body. Without feeding or cleaning the
parent animals spend for 2-3 hours of the carrion, till they finally remove themselves from the
crypt and return no longer.

m. Development of full grown larvae to young beetles.

Shortly before the conclusion of the 7 day period of development the brood ceases to take up
food. Up to this point the near white looking larvae are colored yellowish and at the same
time become broader and flatter. The animal, which at this stage can also be described as
"prepupal", shows a different behavior from the larvae III stage. The continuous massing
together of the larvae under one another ceases. The animals bore through the outer layer of
the ball of carrion and advance singly into the soil to pupate. Fig. 25 illustrates an abandonded
carrion which remains behind hollowed out and with a hole through it. Over the depth as far
as to the dug in Necrophorus-larvae, Xambeu (1892) Chenn and Desinarest (1851) have
already made statements. According to Xambeu (1892) the N. fossor pupae are found in 15
cm depth, from Chenn and Desinarest N. humator in 60 cm. My results differ from the named
works: the depth, up to which the larvae were able to advance to pupate is dependent naturally
on the respective depths of the crypt. From the brood cavity the pupae move predominantly in
the horizontal plane up to 30 cm from the nest and in doing so only bore themselves a little
deeper. So one finds for example the pupae of N. vespilloides 1-5 cm under the earth surface,
however in a particular case directly under the most respectively raw humus layer. But also
for the deeper digging species N. humator the estimated values of Chenn and Desinarest are,
at least for the individuals living in the surroundings of Frankfurt, regarded as clearly too
high. At the end of their wanderings the prepupal stage pauses to build the pupal cradle which
originates through long hours of rotation of the larva on its long axis, as already shown by
Fabre.

Four of the mentioned Necrophorus species, namely N. germanicus, N. humator, N. vespillo

and N. vespilloides pupate after a pause for rest of 12-17 days and complete their
development still in the same years. N. fossor and N. investigator are different. There the
newly emerged adults _ of these species appear in the surroundings of Frankfurt all at the end
of June at the earliest, so the young generation also run through the larva and development
mostly first in September. After that the animal enters a standstill in development: the
prepupal stage overwinters in the pupal cradle. Xambeu (1892) had already stated this for N.
fossor and Main (1927) also noticed that at least one species overwinters as a larva. That the
break in development is not a direct result of environmental factors for example low
temperature, but view as fixed genotypical development rhythm, is shown in the following
experiment. The wandering larvae of both these species were put at the beginning of October
1930 in their rearing cage in a greenhouse in which the temperature of 19_C , prevaded.
Although under the same experimental conditions other ~ Necrophorus species had already
developed up to the pupa, the larvae of N. fossor and N. investigator remained standing at the
pre-pupal stage and died off gradually during the winter. Control animals taken from the same
nest put in the smallest_flower pots, filled with earth, shut above with wire gauze, buried a ~
w cm deep in the Botanical Gardens of the University of Frankfurt and the~_oil-there covered
with a layer of leaves. In April of the following year (1931) these animals were found in
strong health in the prepupal stage, pupated at the end of May and emerged at the conclusion
of the pupal rest to adult.

The pupal rest of the grave digger lasts 14-15 days. The pupa rests only on the strongest
bristles which are found on the eyes and the pseudocerci and the ventral side turned
downwards, in the small subterranean cavity. At the irritation of touching the abdomen the
bristles beat or make circular movements. Two days before the hatching of the young beetles,
the initially white pupa becomes brownish. The young beetle which frees itself from the pupal
covering, already has gold/brown extremities and a likewise colored thorax and scutellum.
The elytra, are white/yellowish, are only indistinctly marked. Under normal conditions, the
animal remains in the protection of his subterranean room. until 4 days after the casting of the
pupal skin when the complete coloring and hardening of the chitinous skeleton occurs. First
then the young beetle breaks through the wall of his pupal cradle and works his way through
to the surface of the earth. The light color and the silk smoothness of this armor betrays the
young beetle, which goes off without delay in search of food. After a maturing feed, which is
completed in 10-14 days in N. vespillo and N. vespilloides, the young beetle is ready to
reproduce and qualified for the combined performance.

SUMMARY

The investigation extended to the species: Necrophorus germanicus, L., N. humator, A., N.
vespillo L., N. vespilloides Herbst, N. fossor Er., N.investigator Zett.

1. N. vespillo is native in meadows. N. humator and N. vespilloides are confined to wood

areas. N. humator is found predominantly in humid deciduous woods, N. vespilloides in dry
coniferous woods.

2. All species feed themselves on carrion and diptera larvae; N germanicus also eats
Geotrupes.
3. Extra intestinal digestion of the adult is probable.

4. Before the burial every carcass is examined for chemical condition, size and ease with
which it can be shifted.

5. Sexually mature males, which find themselves alone at the burial of a suitable body' attract
the female of their species through "Ventilation".

6. The isolation of a single pair from a large number of commonly burying individuals is
based not on social instinct (Fabre 1988, Reuter 1913, Schroder 1929) but on fighting.

7.The burying does not advance alone through digging under the body. This is rather
conveyed in the course of the digging activity in a slanting hole leading into the depths at
whose base lies the ball shaped brood chamber (crypt - Fabre 1~899).

8. Simultaneously with the construction of the crypt, the rounding and cleaning of the buried
body advances.

9. The eggs are laid singly on both sides of a horizontal outgoing passage from the crypt into
the soil, the mothering passage.

10. In the majority of cases the males which shared broodcare with the female are driven
away out of the crypt after completion of egg-laying.

11. After egg-laying the ball of carrion is damped with excrement from the female and on its
upper pole mixed through with gut secretions.

12. At this point, which is deepened by the female into a circular crater shortly before the
appearance of the descendants, gather the hatched larvae at the end of 5 days embryonic
development, which distinguish themselves in the first stage through good ability to smell.

13. The larvae go through two moults and are fully grown in 7 days.

14. A female is regularly found by the brood in the crypt, rarely a pair. The grave digger
practices highly developed broodcare:

a) The brood is fed by the female with gut contents. The time of feeding extends over the
first 60-72 hrs. of larval development.

b) The brood is defended against enemies.

c) The crypt is repaired after destruction.

Males which are not expelled by the female, participate in broodcare.

15. At the end of a 7 day period of development the larvae leave the carrion and
predominantly withdraw from the brood room in a horizontal plane.

16. The species N. germanicus, N. humator, N. vespillo and N . vespilloides overwinter as

adults, N. fossor and N. investigator in the prepual stage.

17. The pupal stage lasts 14-15 days.