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Memoirs of the Museum of Comparative Zoslogn,

AT HARVARD COLLEGE, CAMBRIDGE, MASS.

Vou ll. No, 9:

ON SOME INSECT DEFORMITIES.

By DR. HERMANN A. HAGEN.

UNIVERSITY PRESS, CAMBRIDGE,
WELCH, BIGELOW, & CO.

May, 1876.
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THE quarto publications of the Museum will hereafter be issued
under the title of “Memoirs of the Museum of Comparative Zodlogy.”
In order not to commence a second series, the numbers of the Ilus-
trated Catalogue already issued have been combined to form the
first volumes of these Memoirs. Title-pages and tables of contents
of the three volumes already completed are sent with the present

number.
ALEXANDER AGASSIZ.

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ON SOME INSECT DEFORMITIES.

In arranging the Lepidoptera of the collection, I found, among the
insects brought home from Brazil by Professor L. Agassiz, a speci-
men of Morpho Eurylochus with the head of the caterpillar. The
excellent condition of the large specimen induced me to compare all
published observations of a similar deformity. These are few, and scat-
tered in transactions not easily accessible. Therefore I concluded to
reprint the full text for the two oldest known, and to give copies of the
figures, together with those of JZ. Hurylochus. In the hope of having
information about other similar cases, I published a provisional paper in
the Stettin Ent. Zeit. (1872), p. 388, and Iam indebted to Professor Zeller
of Stettin, Professor Westwood of Oxford, Mr. M‘Lachlan of London,
and Dr. J. L. Leconte of Philadelphia, for additional information.

The rare and less known paper of Mr. C. Majoli, on a precocious
development of Bombyx Mort, and the notice of two deformities of
Coleoptera, seemed to me not out of place in this paper.

PERFECT INSECTS WITH THE LARVAL HEAD.

Phalena heteroclita.

The well-known Danish naturalist, Professor O. F. Mueller, has de-
scribed (1764) in his Fauna Fridrichsdalina, p. 47, No. 413, a new
species of Noctua, found by himself at Fridrichsdal, a locality a few
miles distant from Kopenhagen.

Phal. N. heteroclita subcristata, capite erucee, antennis nullis: alis
albis, lineis transversim undatis punctatisque marginalibus nigris. In
Epilobio. .

This description is verbatim, repeated by himself in his Prodromus
Faun Danice (1776), p. 124, No. 1428.

In the Mém. de Mathém. et de Phys. présentés a l’Acad. R. des
Sciences 4 Paris (1774), Vol. VI, pp. 508-511, pl. 1, Professor O. F.
Mueller has given a detailed account of this most remarkable speci-

6 ON SOME INSECT DEFORMITIES.

men, believing it to be a new and very curious species of Lepidopteron,
with all the characters of the order, except that the head is exactly that
of a caterpillar. In the same volume of the Mémoires, the editor (Pré-
face, p. 8) believes it more prudent not to admit Mueller’s insect as
a new species, because a fact contrary to all hitherto known must be
proved by a great number of observations before it can be adopted by
the scientific world. Professor Beckmann, the reputed polyhistor from
Goettingen, in his Physik. Oekon. Bibliothek, Vol. VI, p. 358, be-
lieves that Mueller’s insect is only a deformity. A review of Mueller’s
paper in the Comment. Lipsiens. ( Vol. X-XI, p. 466) I have not seen.

The French paper is translated by the Rev. J. A. EK. Goeze, in the
16th part of the Naturforscher (1781), pp. 208 — 212, pl. 1. The plate
is of course the same as Mueller’s plate, but somewhat inferior in exe-
cution. The translation, in some places at least, does not entirely
agree with the original, as Goeze introduces some suppositions to ex-
plain more fully Mueller’s words, which are not everywhere free from
ambiguity. But it is to be remembered that Goeze had spoken of the
whole with Mueller, at a visit paid to him by the latter in 1776. At
that time the type was still present in the collection of the author,
which was afterwards destroyed at the bombardment of Kopenhagen.
In my Bibliotheca Entom. (Vol. I, p. 556) I stated that the insect was
Bombyx dispar, which is apparently an error. Westwood (Introd.,
Vol. I, p. 356) calls it one of the Noctuidae; and Lacordaire (Introd.,
Vol. HU, p. 442), une Noctuelle. The insect is not mentioned in the
general works of Borkhausen and Ochsenheimer, but Werneburg (Beitr.
zur Schmetterl. kunde, Vol. I, p. 376), quotes it as Bombyx monacha,
and there is no doubt that this determination is a correct one.

Mueller found the insect alive, quietly sitting on a stem of /pilobiun
montanum, on July 28, 1762, pinned it, and only became aware at
home of the remarkable fact that the head of the caterpillar was still
existing on the moth. Both Mueller and Goeze give as the date June
28, apparently erroneously, as in the paper it is twice stated that the
insect lived ten days on the pin, until the 6th of August, when it died.
From June 28 to July 6 there are only nine days.

The description by Mueller is as follows :

Nearly the size of Phal. vinula ; upper wings white, with several
transverse zigzag lines, the border spotted with black; hind-wings
smaller, grayish, the border with alternate black and white spots; all
the wings blackish underneath, the border spotted with black; abdo-
men black, somewhat hairy, with five yellow rings, which are broad
above, narrower beneath, and twice interrupted; the tip of the abdo-
men pointed, yellow, with a yellow ovipositor; the prothorax densely
covered with white hairs, sprinkled with black; the thorax with four
legs, black and gray-colored ; the tibia with two spurs on the inside.

PERFECT INSECTS WITH THE LARVAL HEAD. a

“La tete”” (I give purposely Mueller’s words), “cette étrange partie,
est grisatre et arrondie, plate au devant; elle est composée, comme le
sont ordinairement les tétes de chenilles, de deux lobes latéraux, gri-
satres et pointillés en noir, lesquels se joignant par-dessus, laissent au
milieu une figure triangulaire et brune. C’est une membrane mince,
qui a Taide dune loupe, laissait entrevoir une liqueur transparente,
agitée dun mouvement continuel. Il y a au bas du triangle deux
petits corps ovales, qui avancent sur deux organes noirs, lesquels se ré-
pondent exactement et se choquent, au milieu de ’embouchure, comme
deux marteaux. On voit a c6té deux organes émoussés de couleur
jaune, qui dans les chenilles sont communement garnis d’un_ poil fin, ce
qui manque ici; plus bas, il s’avance des cotés deux crochets coniques
et jaunatres qui se touchent au milieu de la bouche ; a l’'entour on voit
quelques taches incarnates et grandes; plus a coté quelques points
brillants et par-ci par-la quelques petits brians de poils.”

The moth lived ten days, and deposited a number of green eggs,
most of them on the first and second days, some on the latter days up
to the 6th of August, when it died. The eggs were not developed.

Mueller repeats a second time (p. 511): “On voit clairement le
mouvement peristaltique de la liqueur sous la membrane triangulaire,
aussi bien que le mouvement des organes de la bouche; il ne s’en
trouve pas la moindre trace des antennes et trompe.”

It would not be justifiable to consider this statement pure and simple
an error, inasmuch as Mueller was undoubtedly one of the most promi-
nent naturalists, and must have known very well the importance of the
described facts. If the statement of Mueller is accepted as correct, the
specimen is an exception, and differs considerably from all others as
yet recorded. It must have been an imago, with the head of the cater-
pillar preserved; not only with the skin covering the head of the
imago preserved, but with a real head of the caterpillar, in which the
circulation of the blood is still taking place, and the maxillary organs
are still movable. Such a condition of the parts is contrary to all our
present knowledge of the anatomy and the development of insects. It
is remarkable that the forelegs have not been developed, as the su-
perior part of the prothorax is similar to that of the imago. But
Mueller records and figures only the four posterior legs.

The opinion of Mueller, that his moth represents a new and some-
what intermediate genus and species, is of course an erroneous one.
The supposition by Kirby and Spence, Introd., Vol. II, p. 121 (transl.
Oken), that the head was damaged perhaps in the caterpillar state by
some parasite, and the caterpillar therefore was unable to cast off its
skin, needs no refutation. I cannot give any probable explanation of
the fact; perhaps it was a monstrosity, never observed but in this iso-
lated specimen.

§ ON SOME INSECT DEFORMITIES.

Mueller’s observation was accepted by the prominent naturalists of
the time, — for instance, by Bonnet. Later J. F. Meckel (Handb. der
pathol. Anatom., Vol. I, p. 55) explained the fact as an arrested devel-
opment of the insect. Dr. Stannius (Mueller’s Archiv., 1835, p. 296)
accepts this explanation. J. van der Hoeven (Tijdskr. voor natuurl.
Gesch., Vol. VI, p. 274) believes the case to be just like that of Wes-
mael and rejects the opinion of Mueller concerning the head; but I
think he has not given strict attention to Mueller’s statements.

Nymphalis Populi.

Professor Wesmael gives in Bull. Acad. Bruxelles, 1838, Tom. IV,
p- 359, reproduced partly in Ann. Scienc. Nat., Ser. 2, Vol. VII, p. 191,
a description and colored figure of this insect. If I am not mistaken, I
saw, in 1870, the type in the Museum in Brussels. He caught the
specimen in July, near this city. The insect had the thorax, abdomen,
legs, and wings perfectly well developed and colored, but with the
head of the caterpillar. The insect turned the curious head to the
right and left, and tried, by a quick motion of the forelegs, to push it
off. Mr. Wesmael, in dissecting the left side of the head, discovered
underneath the external skin a second one much thinner than the
first, and beneath the second one the well-developed eye of the butter-
fly. The parts around the eye were covered,as commonly, with scales.
Therefore Wesmael considers the second skin as that of the chrysalis,
and believes the deformity originated by the imability of the caterpillar
to cast off the head.

Underneath the head of the caterpillar, and just above the skin of
the chrysalis, was the left antenna, coiled up, but without an apical
knob. The antenna was covered by a very fine membrane, which was
to a great extent diaphanous, and transversely striated with brown.
The left palpus was free, normally developed, and turned horizontally
backwards. ‘The right palpus seemed broken off; the place of its in-
sertion was clearly visible. Mr. Wesmael says nothing about the pro-
thorax ; as the forelegs were free and movable, it must have been with-
out any covering.

This buttertly differs essentially from the moth described by Mueller.
The head shows only the skin of the caterpillar, which really has gone
through the transformation into the head of the chrysalis, and later
into the head of the imago, retaining throughout the skin of the former
stages, one above the other. The recorded movement of the head was
apparently done by the movement of the head of the imago. Wesmael
makes the following conclusions : —

1. The insects which are obliged to undergo transformation may
have only a partial one, which does not prevent the total transforma-

PERFECT INSECTS WITH THE LARVAL HEAD. g
tion of other parts; even if the untransformed parts are important to
the life of the animal. He believes this to be a natural consequence of
the segmentation of the body of the arthropods.

2. The accidentally ‘covered parts nevertheless go through all trans-
formations which are necessary for the insect to arrive at the imago
state. The second conclusion is of course not to be accepted, if the
facts recorded by Mueller are adopted.

Morpho Eurylochus.

Among about a dozen specimens of this butterfly brought home by
Professor L. Agassiz from Brazil, one male has retained the head of the
caterpillar. The specimens are from Canta Gallo, communicated by
Dr. Teuscher. Their perfect condition leads me to suppose they were
reared from the chrysalis.

The quoted male is in perfect condition, and, as all others, entirely
well developed in size and in colors. The head of the caterpillar is
retained and perfectly preserved in shape and in color; the minute
yellow hairs which cover the head are in good condition, and the
spines are scarcely crumbled. Beneath the head the mentum is
broken off near the prothorax. Its lateral sutures are separated, and
the mentum hangs down as a kind of trap-door, being united with the
head only by a small anterior lobe. This kind of adjustment leads me
to suppose that the mentum was broken by the pushing out of the
spiral tongue of the imago. The opening is large enough to show that
the head of the caterpillar is empty inside. The skin between the
head and the prothorax is still preserved in the shape of a contracted
ring, which is open only for a small space beneath, where the mentum
is separated. The large dorsal plate of the prothorax is present, and
covers loosely the thorax of the imago; on the left side the external
third is wanting. The palpi are rejected to the thorax, but the right
palpus has the two basal thirds covered by the skin of the caterpillar,
which is connected with the dorsal part of the prothorax. Behind the
palpus and rather near to it can be seen the free foreleg of the right
side. Its limbs are well developed, neither as stout nor as hairy as in
the other specimens. The left palpus, though not covered, seems to be
shorter and less hairy than the right one. The left foreleg is covered
by the femur of the middle ieg. JI am not able to state whether any
part of the skin of the chrysalis, either beneath the dorsal plate of the
prothorax on the middle and on the right, or on the entirely free left
side of the thorax, is present. Perhaps the skin of the chrysalis 1s
broken off just at the ring formed between the head and the prothorax.
I am unable to see the skin of the chrysalis inside of the head of the

10 ON SOME INSECT DEFORMITIES.

caterpillar through the small opening of the mentum. As the skin of
the chrysalis must have existed, I did not deem it necessary to dissect
the specimen, especially as Wesmael’s dissection of Nymphals Populi
has sufficiently explained the fact.

The head of the caterpillar resembles very much the figure in Merian
Surinam. Lepid., pl. 25. The color is leather-yellow, with two brown
bands on each side. There are two yellow finger-shaped horns on the
top, and three similar ones on each side; they become successively
smaller. The last one is very short.

The specimen has doubtless lived long enough to get the colors per-
fectly developed, and to break down the mentum with the spiral tongue.
It differs from Wesmael’s butterfly in having retained the dorsal part
of the prothorax, though somewhat distant to allow a view of the tho-
rax of the imago. In Wesmael’s butterfly the palpi were not covered.
I have quoted erroneously, in the Proceed. Bost. Soc. N. H., 1868, Vol.
XII, p. 163, the Brazilian specimen as Morpho Iloneus, and Mueller’s
specimen as Dicranura vinula.

Vanessa Antiopa.

Professor Zeller has described in the Isis, 1859, p. 259, a specimen
with the head of the caterpillar, raised by himself together with about
150 others. The specimen differs from them only by the presence of
the head of the caterpillar, which is in a vertical position, just as in thie
caterpillar. The mouth is closed. Having cut a part of the left side,
the Professor could observe a hollow space between the head ‘of the
caterpillar and the remaining parts of the insect. Behind the head
and not connected with it the two anterior plates of the chrysalis are
retained. The butterfly made its transformation in the absence of the
Professor, and was pinned at the same time with all the others. It was
impossible to find its chrysalis skin.

Vanessa Atalanta.

Mr. Bond exhibited in the Entomological Society in London, Febru-
ary 6, 1871, a specimen bred by a metropolitan collector, which still
bore the larval head. The specimen, as I am informed by Mr. M‘Lach-
lan, was very perfect.

Pieris Rape.

Among a number of chrysalids which had not transformed, I found in
the fall of 1871, in Cambridge, one of an extraordinary appearance.
In casting off the skin of the caterpillar only the thoracic part of the

PERFECT INSECTS WITH THE LARVAL HEAD. aia

chrysalis was developed; the head of the caterpillar was still present,
but its sutures were separated. The dorsal split of the skin reaches the
first segment of the abdomen, and the skin of the abdomen is retracted,
but still present.

A similar specimen was observed last fall by Mr. S. H. Scudder.

Zygena exulans var. Vanadis.

Dr. Staudinger, in a paper on the Lepidoptera of Lapland in Stett.
Entom. Zeit., 1861, Vol. XXII, p. 359, records a larval-headed male of
this species, found, July 11, near Bossekop. The specimen is fully
developed, with the head of the caterpillar. The mouth parts were
immovable in the living insect; the head was fastened to the protho-
rax, and moved only by the motion of the prothorax. The latter is
fully developed beneath, and with its legs; above there is a horny
black vaulted ring, somewhat hairy on the left side. Mr. Staudinger
believes it impossible that the head of the imago is enclosed in this
larval head.

Sphinx spec.

Mr. Trouvelot assured me that he had caught in Cambridge several
years ago, a Sphinx with the head of the caterpillar. The specimen is
no longer in existence.

Professor Van der Hoeven, in his quoted paper, p. 274, records that
he has seen a caterpillar of Sph. Tilie which had not been able to cast
off in the last moult the skin covering the spine of the tail. The cater-
pular died before the transformation.

Bombyx Mori.

Mr. J. J. Bruinsma, in Tijdschrift voor Natuurl. Geschied, 1840, Vol.
VU, pp. 257 — 270, pl. 1, has published detailed observations concerning
the same deformity, accompanied by figures.

Having read Wesmael’s paper, Mr. Bruinsma concluded to try his
own observations in raising silk-worms. In the course of the summer
he found some specimens, which did not agree exactly with Wesmael’s
butterfly, but seemed interesting for publication. But shortly after
Mr. 8. Van Leuwen, also interested in the same kind of observations,
communicated to him, August 9, a chrysalis with the upper part of the
larval head still remaining. The chrysalis was fourteen days old, and
had been taken out of a cocoon, in which the skin of the caterpillar
was found. The skin showed nothing unusual, except that the head,
which commonly remains united with the skin, was broken off.

12 ON SOME INSECT DEFORMITIES.

The chrysalis was of medium size, rather lively, and perfectly devel-
oped. To the upper part was fastened with a collar the caterpillar’s
head, split in two lateral parts, which are united together in the com-
mon caterpillar head. The mouth parts of the caterpillar were still
remaining, but between them another prolongation could be seen. The
nympha is figured in different views, Figs. 1-4.

The chrysalis transformed, August 26, into a moth with the caterpil-
lar head. The chrysalis had the skin split as usual on the dorsal part.
So the moth left the chrysalis in the usual way, and was perfectly
developed (Fig. 5), except that the right foreleg was smaller than the
left one, but otherwise well formed. ‘Therefore the moth stood some-
what obliquely. It was a male, rather lively in his movements, and
used both forelegs to push off the caterpillar head, by which it was
seemingly annoyed. The head of the caterpillar covered exactly the
place where the head of the moth should be, so that nothing was to be
seen of it, nor of its antenne or eyes. On the prothoracic border the
same elongated part was to be seen as in the chrysalis, without any
hairs, consisting only of a brownish membrane ‘The side parts of the
head were fastened to it just as in the chrysalis. The right part was
taken off, and beneath it the right antenna was discovered, well formed,
but coiled up. In taking off more of the skin, a well-formed eye of the
perfect insect appeared.

Mr. Bruinsma explains the fact by the difference of the last moult
from the four preceding ones. He states that in the last moult the
skin splits near the tail, and the chrysalis comes out backwards, by
which process the head is sometimes not able to follow. I confess that
this statement is entirely new to me, and disagrees with Malphigi’s
description.

Mr. Bruinsma concludes his paper with some observations made on
caterpillars which, after having spun the cocoon, were taken out and
obliged to spin a second cocoon. This was only imperfect, so that a
full observation of the caterpillar was possible. Mr. Bruinsma observed
that one of them formed a chrysalis with the head of the caterpillar.
But the chrysalis died very soon. Two of the caterpillars are figured.

Professor J. Van der Hoeven has published a paper in the same jour-
nal, following that by Mr. Bruinsma, pp. 271-275, about perfect Lepi-
doptera with the larval head remaining. He draws attention to an old
observation by J. Jonston, published in his Hist. Natur. de Insectis,
Amstelod, 1657, p. 123 (Edit. Heilbr., 1768, p. 176). The account by
J. Jonston is very detailed, and concerns a male and a female of B.
Mori. The male was in the chrysalis state, the anterior part covered
with the parts of the caterpillar. The imago was not able to cast off
the skin, died, and was dissected. The head of the imago was found to
be fully developed.

PERFECT INSECTS WITH THE LARVAL HEAD. 13

The female showed perfectly well the head of the caterpillar, and
beneath it the skin of the chrysalis, containing the head of the imago.

“Ttaque ibi senectee caput [head of the caterpillar], nymph ver-
tex et necydali [imaginis] conjuncta conspiciebantur ; qua conjunctio
retinebat senectam [skin of the caterpillar] in ventre, ne potuerit
potius avelli et destringi. Ideo et cohawrebat, cum alvi acumine, non
aliter ac si quis sacco fuit inclusus; et circa caput astrictus ; facto vero
in tergo foramine dorsum extraxisset quidem, sed adhue heereret capite
et podice, ita jacens incurvus et exanimis. Sic habebat senecta (the
caterpillar]. Ex hac prominebat et aurelia [chrysalis], quod attinet
partem superiorem. Ex aurelia vicissim necydalus [imago] fere totus
eluctatus erat, fracto putamine in dorso, solitaque regione ; sed capita
coherebant indivulsa, sicut et alvi extrema. In ventro exorto magna
copia ovorum conspiciebatur colore flavo.”

The statement of the skin of the caterpillar split on the dorsum dis-
agrees with the statement by Mr. Bruinsma.

Professor Van der Hoeven (ibid., p. 274) communicates, in a letter to
Mr. A. Brants, November 26, 1839, that Mr. Kinodhven, in his silk-worm
nursery at Brummen, Holland, had observed several times imagines
with the head of the caterpillar. Mr. Brants was able to take out of
one of them the perfectly developed head of the imago. The antennx
were coiled up, covering the eyes of the insect.

Mr. Bond exhibited in the Entomological Society in London, Febru-
ary 20, 1871, a specimen of B. Mori retaining the larval head. The
specimen was somewhat crippled and very small, as Iam informed by

Mr. M‘Lachlan.

Gastropacha quercifolia.

A specimen with the larval head is recorded by Professor Westwood
in Entomol. Month. Magaz., No. 82, p. 239.

Zerene adusta.

I am indebted to Professor Zeller for the details of this. Among a
number of caterpillars of this species, one transformed in a chrysalis,
with the head of the caterpillar. The chrysalis died, perhaps because
it was kept too dry. Otherwise probably a moth would have been
reared, as the chrysalis was perfectly developed. The head of the cat-
erpillar was in perfect condition, but placed so far beneath that the
chrysalis had a hunchbacked appearance. The face and the ventral
side met in an acute angle; a collum was wanting, but the head was
round, separated more deeply below. As all parts of the head of the
chrysalis are covered by the head of the caterpillar, there are no an-

14 ON SOME INSECT DEFORMITIES.

tennal covers visible. The furrow in which the antenne should have
been placed begins very shallow on the prothorax, near the head, and
runs between the front margin of the anterior wings and the hind
legs, somewhat longer on the right side. This furrow is largest along
the tibia. ‘The covers of the palpi and the tongue are wanting.

Botys fuscalis.

Mr. Stainton exhibited in the Entomological Society in London, a
specimen with the head covered by a part of the puparium, caught on
the Isle of Man. It was flying briskly when captured, and was other-
wise perfectly developed. The antennz and the haustellum were free,
and the case of the latter projected downwards like the rostrum of a
Panorpa. I am indebted to Mr. M‘Lachlan for the communication of
this case.

Of insects not belonging to Lepidoptera, only four with a similar de-
formity are known.

Cybister limbatus.

Mr. Smith (Proceed. Entom. Soc., London, Ser. 2, Vol. IV, p. 34) ex-
hibited a specimen with the larval head, caught swimming near Hong-
kong in China.

Dytiscus marginalis.

Professor Westwood (the Entom. Month. Mag., No. 82, p. 239) stated
that he had seen a specimen with a larval head.

Hydaticus bimarginatus.

I am indebted to Dr. John L. Leconte for the information that a
specimen of this beetle, with the larval head, is in the collection of Dr.
Helmuth in Chicago.

Syrphus spec.

Professor Westwood (ibid.) states that he had observed one specimen
with a larval head.

I am indebted to the same professor for the communication that he
is about to publish a dozen cases of perfect insects with the larval head,
all of which he has figured.

The specimens mentioned above are : —

Lepidoptera. — 1. Vanessa Atalanta.
2. Vanessa Antiopa.
3. Nymphalis Popul.

PERFECT INSECTS WITH THE LARVAL HEAD. 15

4. Pieris Rape.
5. Morpho Eurylochus.
6. Zygeena exulans.
7. Sphinx spec.
8. Bombyx Mori (several times).
9. Liparis Monacha.
10. Gastropacha quercifolia.
11. Zerene adusta.
12. Botys fuscalis.
Coleoptera. — 13. Dytiscus marginalis.
14. Hydaticus bimarginatus.
15. Cybister limbatus.
Diptera. — 16. Syrphus spec.

Only the fact of the presence of the larval head is known for the
Nos. 7, 10, 18—16. More or less sufficient details are known for Nos.
1, 3, 8, 12, but the publication by Professor Westwood will give doubt-
less a full information about them. The Nos. 4, 11, were only in the
chrysalis state, and do not strictly belong here. But just those cases
are interesting, as all the others must have passed the chrysalis state in
the same manner.

The Nos. 1, 2,8, and probably 5, were bred by home raising ; Nos. 3,
6, 7, 9, 12, 15, were caught living.

The interesting fact that the larval head is sometimes retained in the
perfect insect is proved by the quoted observations. Probably all cases
except Noctua heterochta belong to the same kind of deformity. The
head of the imago is contained in the head of the larva, which the in-
sect was not able to cast off in the transformation. It must be admit-
ted that circumstantial details are known only for V. Populi, V. Antiopa,
M. Eurylochus, Z. exulans. The prothorax of the two last-mentioned
species, and even one foreleg in M. Lurylochus, is still covered by the
larval skin. The antenne are free in B..fuscals ; the palpi are rejected
in V. Antiopa, V. Populi, M. Eurylochus ; in Z. exulans they are free. All
specimens were fully developed im size, shape, and colors, except B.
Morr.

Perhaps such deformities are not so rare as the small number of
known cases would lead us to believe. Such deformed specimens are
more easily caught and destroyed by their natural enemies, or they die
sooner for lack of food. Nevertheless, the very large number of Lepi-
doptera bred and raised during the past hundred years allows us to con-
clude that at least in home raising such deformities occur very rarely.
Mr. Trouvelot observed many times 7’. Polyphemus caterpillars casting off
the larval head with more or less difficulty, and sometimes not at all.
The last case proved to be fatal. As insects of course are developed
more easily in liberty than in captivity, the rare occurrence of perfect

16 ON SOME INSECT DEFORMITIES.

insects retaining the larval head may depend on the larger mortality of
their caterpillars.

There is very little known concerning the physiological and mechani-
cal processes shortly before or during the act of transformation of
arthropods. Nearly all entomological works state that the larvae moult
or change their skin several times, that the larvae become restless some
days before the change, stop eating, and desert their food; later, the
skin splits, and the insect perfects its transformation.

I believe there exist few men who have not seen and observed once
in their life this wonderful spectacle. The proceedings are so common,
and always so easily performed, that observers are not induced to think
about the manner in which transformation is effected, nor about the
mechanical acts providing the possibility of such a change. Concerning
the mechanical acts, so far as I know, nothing is published. . An animal,
or even a man, covered with an artificial skin well fitted to the whole
body, obliged to go out of the skin through an aperture made of similar
size and relation as that in insects, would scarcely be able to do it with-
out a violent use of the limbs. Insects use their limbs very little or
not at all in the beginning of transformation, but nature has provided
some help in the necessary coincidence of certain physiological pro-
ceedings just at the time of transformation.

I have observed many times and in different insects, before transfor-
mation, a very accelerated and excited action of the dorsal vessel. The
same fact is recorded by other observers, for instance by Mr. Weismann
and Mr. W. Blasius. After observations of Mr. W. Blasius (Zeitschr.
f. wiss. Zool., Vol. XVI, pp. 135 -— 177), during the transformation of the
caterpillar into the chrysalis, the action of the dorsal vessel increases
successively in the first three hours, and reaches its maximum in the
last half of the fourth; after that time the action begins to decrease,
and becomes in the eighth hour equal to the action in the third hour.
The consequence of an accelerated action of the dorsal vessel is an in-
creased circulation of the blood, going from the tail to the head. This
sudden rush of an unusual quantity of blood to the head and the tho-
rax, without any corresponding arrangement for convenient emanation,
swells those parts, pushing them forward at the same time. Finally
the skin bursts, and one of the most important acts of the moult is per-
formed.

I suppose that the frontal bladder, which is observed in transforming
Diptera and Odonata, is the consequence of the rush of blood; never-
theless, an observation recorded by Weismann seems to disagree with
such a supposition. It should not be overlooked that some other purely
mechanical proceedings seem to accompany and help the propulsion of
the insect by the rush of the blood in a very easy manner. I have
described long ago a related fact in regard to the moult of Ephemera,

PERFECT INSECTS WITH THE LARVAL HEAD. Vi

Stettin. Ent. Zeit. (1849), p.365. The segments of the abdomen possess
on each side an apical spine. By a continuous movement of the abdo-
men to the right and left, those spines press successively against the
loosened skin, forcing forward the transforming insect. Probably simi-
lar arrangements will be found in other insects.

The causes why such a rush of blood originates just at the time of the
moult, I find nowhere recorded. I think it not sufficient to consider it
as a simple consequence of an action of the nervous system, especially
as I believe myself able to give a more plausible explanation.

The crust of insects consists of the external chitinized epidermis, and
the internal soft hypodermis. Above the latter, which becomes some-
what separated from the epidermis, the new skin is to be formed, at
first without impediment to the functions of the insect. As long as the
more or less isolated parts of the new skin allow a free circulation of
the blood around and between them to feed the old epidermis, the
action of the dorsal vessel follows its regular way. By and by the iso-
lated parts of the new skin become larger, and partly united, until
finally the whole new skin is already formed and chitinized. The circu-
lation is at first only disturbed; later, it is impossible for it to flow in
the old way and to the old skin, and the blood, obliged to turn in
another direction, rushes naturally in the easiest one, to the dorsal
vessel. This is the moment of the beginning of the rush of the blood to
the head ; of course the nervous system, Teeted by the mush, will help
to Ua ane more the action of the dorsal vessel.

It is obvious that the new skin, at least in some parts of the body,
must exercise a more or less strong pressure against the old skin. I
am of the.opinion of Dr. Gerstaecker, that the moult is not alone a con-
sequence of such pressure ; but in some parts, for instance in the head,
the pressure is obviously prevalent, and must originate a partial resorp-
tion of the old epidermis, as that of the thicker sutures. At least, thus
the splitting of the sutures in many insects could be explained; I say
purposely in many insects, because a large number transform in a dif-
ferent way. In some Lepidoptera the skin of the head does not split.
Mr. Trouvelot (Americ. Natur., I, p. 87) records for Zelea Polyphemus
that the skin splits transversely under the neck just at the end of the
head, and perhaps in some way laterally, and probably behind or
through the whole prothorax. “When about one half of the body
appears the shell of the old head remains like a cap enclosing the
jaws; then the worm, as if reminded of this loose skull-cap, removes it
by rubbing it on a leaf.” ;

I was able to verify Mr. Trouvelot’s observation on a cast-off skin of
T. Polyphemus. However, in the nearly related species Att. Cecropia, Pro-
methea, Yama-mai, 1 found that the sutures of the head always split in —
the regular manner. According to Kirby and Spence, the manner of

18 ON SOME INSECT DEFORMITIES.

splitting is not the same for all Lepidoptera. Most of them split at
first the dorsum of the second and third segment of the caterpillar.
Pieris crategi is stated by Bonnet to split its skin only in the head; and
Reaumur records for Zygana filipendule that the caterpillar bites off
pieces of the old skin and puts them aside. Besides those old observa-
tions there exists a large number of recent ones. Generally it is true
that Coleoptera, Orthoptera, Pseudoneuroptera, Diptera, Hemiptera, at
least the Homoptera, and the larger part of Lepidoptera, split first the
frontal sutures of the head.

It is of interest that the well-known Limulus Polyphemus splits in the
moult the frontal sutures similar to the insects, and goes out forward
of the old skin. I find nothing published about the splitting of the
skin of the Crustacea. But the Decapods split along the border of
the cephalophorax, and are obliged in moulting to take out the parts
backwards. As the systematic position of Limulus is still a matter of
dispute, I think this fact is of some value.

The restlessness of the insects before the moult, and the fact that
they do not need food, are easily understood. It is known that the
tracheze participate largely in the moult. At certain times the inner
skin must begin to separate, and it is obvious that through it the respi-
ration will be impeded. Perhaps this difficulty has something to do
with the acceleration of the blood. Besides the. trachez a large part
of the digestive canal changes its skin; the anterior part to the ven-
triculus, and the posterior to the colon. Here also, at a certain time,
the inner skin begins to separate, and the natural consequence will be
the impossibility of taking any food, or even of ejecting the superfluum
contained in the intestines.

All insects, as I stated before, go forwards out of the old skin in
transformation. But I find one case of the contrary quoted by Profes-
sor Westwood (Brit. Cyclop. Article Insect, p. 844). He says that Coc-
cus comes out backward, the wings rejected above the head.

A very remarkable fact is that the females of the Ephemerous genus
Palingenia do not change the skin of the subimago. Swammerdam
says the females for the most part do not change the skin; but a very
large number of them examined by myself possessed the subimago
skin, and I never saw one without it. They undergo copulation and
lay eggs without having arrived at the state of a perfect insect.

The instances where insects in the last moult are not able to throw
away the skin and carry it with them are in Ephemera not very rare,
as the long sete are sometimes fastened in the old skin. But I caught
a Libellula flying with its nympha skin fastened to the end of the abdo-
men. The specimen, Diplax secotica, is still in my collection. I am
indebted to Mr. A. Agassiz for a knowledge of the fact that larve of
Radiates are sometimes found with parts of the foregoing state still
attached to the skin.

PRECOCIOUS DEVELOPMENT OF THE CATERPILLAR. 19

a

ON THE PRECOCIOUS DEVELOPMENT OF THE CATERPIL-
LAR DIRECTLY INTO THE IMAGO STATE.

Bombyx Mori.

Mr. Cesare Majoli, in Giornale di Fisica, Chemica, Storia Naturle del
Reeno Italico di L. Brugnatelli, Pavia, 1813, Bim. V, p. 599, describes
a curious instance of a precocious development of the well-known silk-
moth. The book seems to be wanting in all libraries of the United
States, and is not even common in Europe. Iam obliged to Professor
Pelzel in Vienna, Austria, for a written copy of the notice, which I find
recorded only in J. F. Meckel’s Archiv f. Physiol. 1816, Vol. II, p. 542,
and in Lacordaire’s Introd., Vol. II, p. 443, and by Stannius in Muel-
ler’s Archiv, 1855, p. 297. As both differ from each other in some im-
portant statements and from the text, I prefer to reprint the original.

Straordinario fenomeno di anticipata trasformazione in farfalla del verme da
seta.

I] Sign. A. Farini di Forli ha communicato al Sign. Barzoni un’ osser-
vazione interessante descritta dal Sign. Lettore Cesare Majoli in un
opusculo M.S. sulla vita, costumi ed educazione del filugello. Sovente
aveva sentito raccontare da chi educava i bachi da seta, che pure
qualche volta accadeva svolgersi essi in farfalla prima che incomincias-
sero a filare il bozzolo, cioé dopo la quarta muta. Le reputava favole
femminili, giacché nissuno aveva parlato di un tale fenomeno. Ma si
é convinto del fatto nel 1792, allorché chiamato a rendere ragione di
esso trovd che due cannicci e stuoje di bruchi erano isfarfallati nella
notte antecedente senza formare il bozzolo deludendo cosi la speranza
del coltivatore. In qualcuno degli anni antecedenti accadde pure un
somigliante fenomeno, e, nel 1811 il Sign. Dott. Siboni gli mando due
di questi aborti volanti generati in una casa di proprieta della Signora
Rosatti che lo stesso Sign. Farini ha osservato. Questa farfalla diffe-
risce dalla falena bombice per li seguenti caratteri. Ha il capo piccolo,
due occhj neri reticolati, il torace quale se fosse il terzo anello dopo il
capo del bruco; ha il corpo del bruco istesso all’ epoca della quarta
muta, pari numero di anelli a quello del bruco; le ali superiori alquanto
lunghe e ristrette, le inferiori pit’ corte e strette; ha le antenne al-
quanto cenerognole in confronto di quelle della falena vera bombice.
Il Sign. Majoli espone una conghiettura sopra la cagione del fenomeno
mentovato e inclina ad attribuirlo al calore eccessivo del luogo in cui
esistevano qué bachi da seta, per cui nel momento in cui filugello sta

20 ON SOME INSECT DEFORMITIES.

per compiere la sua metamorfosi dello stato di bruco, ne altera siffatta-
mente il sistema primitivo, ne promuove una traspirazione straordinaria
de’ fluidi esistenti nel bruco e soprattutto di quello che € necessario a
formare il bozzolo, e ne acceleri cosé la sua metamorfosi d’ isfarfallare.
Sarebbe stato a desiderare che per confermare in qualche modo I’ accen-
nata opinione si fosse tentato artatamente di attenere lo stesso effetto
col sottoporre diversi bruchi ad una temperatura calda, allorché erano
vicini alla quarta muta. Interessante sarebbe il sapere se le farfalle
che abortirono avevano gli organi della generazione ben formati e
capaci come la falena bombice di accoppiamento e di mettere le uova
‘atte a sviluppare a suo tempo il bacolino.

[ TRANSLATION. |

An extraordinary case of a precocious transformation of the caterpillar of Bom-
byx Mori into the moth.

Mr. A. Farini, in Forli, has communicated to Mr. Barzoni an interest-
ing observation, published by Mr. Cesare Majoli, about the life, the
manners, and the education of the silk-moth. He was told by men
occupied with the breeding of silk-worms that it sometimes happens
that caterpillars before spinning a cocoon, therefore after the fourth
moult, were transformed into moths. He believed it to be only talk, the
more so as nobody had published anything about such a case. But he
convinced himself of the fact in 1792. Invited purposely to convince
himself by his own observation, he saw two boards filled with caterpil-
lars transformed in the preceding night into moths without having spun
cocoons, to the regret and deception of the owner. Recently the same
fact was observed, and in 1811 he received by Dr. Siboni two such
winged deformities, bred by Mrs. Rosatti, and examined by Mr. Farini.
These differ from the regularly developed silk-moth in the following
characters: The head is small, with two black compound eyes; the
thorax similar to the third segment behind the head of the caterpillar,
and the abdomen similar to that of a caterpillar at the time of the
fourth moult, with just as many segments as the abdomen of the cater-
pillar; the fore-wings somewhat elongated and narrowed; the hind-
wings shorter and narrower; the antennz grayish, compared with those
of the regular silk-moth. Mr. Majoli gives an hypothesis for the cause
of such a transformation, and believes it to be the excessive warm tem-
perature of the breeding-room. ‘The caterpillar ready for transforma-
tion is prevented by the heat from producing the exudation of the
fluids which are necessary for the formation of the chrysalis, and is
obliged to transform directly into the moth. To ascertain his hypothesis
he should have made experiments to produce such a transformation in
an artificial way, by exposing caterpillars shortly before the fourth

PRECOCIOUS DEVELOPMENT OF THE CATERPILLAR. o1

moult to a high temperature. It will be of interest to know if the
moths possessed well-developed genital parts, fitted for copulation and
for the deposition of fertile eggs.

I have given purposely, in full, the original, as the two records ex-
isting disagree on some important facts. Lacordaire seems to have
seen the original, as his record contains some statements not given by
Meckel. Nevertheless, he has surely seen, and partly translated,
Meckel’s record, as is proved by the words: “les deux yeux noirs
rapprochés,” a verbal translation of the “ Zusammengesetzten Augen,”
by Meckel. The presence of fore-wings only is recorded by Meckel, of
hind-wings only by Lacordaire.

The statements given by Majoli are probable, except that the thorax,
which is said to be similar to the third segment of the caterpillar, has
four wings. The identity of the abdomen with the abdomen of the
caterpillar consists perhaps only in elongation, as the presence of other
parts is not mentioned.

Lacordaire considers the fact as proving the development of certain
parts of an insect by precocity, though the other parts follow the com-
mon rule of development.

The fact would be a rather interesting one if it was beyond doubt.
As silk-worms are raised every year by millions, I should have sup-
posed that the observation would have been oftener made and pub-
lished. Nevertheless, it is astonishing that such a fact, fillng im some
way the gap between insects with an incomplete metamorphosis, and
those with a complete one, is not used by evolutionists.

A paper by the well- oan Lepidopterologist, Mr. EK. J. C. Esper, in
Hoppe. Entom. Taschenbuch (for 1796), pp. 183 — 188, which I have not
at hand, may possibly treat the question of a precocious development.

Majoli’s observation is briefly reproduced (after Meckel) by Professor
Van der Hoeven, in his quoted paper, p. 272. He remarks that no re-
lated observations are known, and that his observation rests only upon
its own merits.

DEFORMITY OF THE ELYTRA.

Strategus (Geotrupes) Julianus.

The late Professor J. Wyman observed a specimen of this large
beetle flying around in Florida in 1874. The movements were in some
way strange and unusual, and induced Professor J. Wyman to catch
the beetle. To his astonishment he found it perfectly developed, but
the elytra wanting. The specimen, preserved in alcohol, he presented
shortly before his death to the collection.

29, ON SOME INSECT DEFORMITIES.

The specimen is a female, of thirty-six mm. length, and perfectly de-
veloped. The wings are in good and perfect condition, but there is no
trace of the elytra; which are entirely wanting on both sides. Pro-
fessor J. Wyman thought at first that the elytra had been removed by
somebody, and the insect put again at liberty But a careful examina-
tion of the living insect as well as of the alcoholic specimen by Dr. J.
L. Leconte, and by myself, showed no lesion whatsoever in the place
where the elytra should have been inserted. The anterior border of the
mesothorax is horny and smooth, and near the scutellum exists a small
membranous place without any wound. The legs, the prothorax, the
upper part of the mesothorax, the metathorax, the scutellum, and the
whole abdomen, are perfectly developed.

This case of deformity belongs to the “monstres ectroméliens” of
Lacordaire, but as far as I am able to ascertain, no similar case is
recorded. The fact that the beetle was able to fly without elytra is of
additional interest.

Prionus coriarius.

This remarkable case is twice recorded, but later entirely overlooked.
As I believe this kind of deformity of prominent value, I give a trans-
lation of the origin)! communication by Dr. Saage in Preussische Pro-
vinzial Blaetter (1839), Vol. XXII, p. 191.

“ One of my school-boys brought me to-day a male Prionus coriarius, the
thorax of which presents a curious deformity. The horny dorsal cover
of the mesothorax is wanting, and instead of elytra there are inserted,
just in their place of articulation, two perfect legs, directed above and
behind. The metathorax has the wings as usual, and the abdomen is
of the same horny character as commonly, when covered with elytra.
In attempts at flight the insect moved, together with the wings, the ab-
normal dorsal legs. ‘The scutellum is wanting, and the prothorax has
only two spines; all other parts are perfectly developed.”

“ Braunsberg, Prussia, July 10, 1859.”

This communication is reprinted in Stett. Ent. Zeit., Vol. I, p. 48. The
specimen was seen and examined afterwards by Professor von Siebold.

I have always considered this case to be a striking proof of the
homology of the wings with the legs. No similar case has been re-
corded.

EXPLANATION OF THE PLATE.

Figs. 1—5.— Morpho Eurylochus.

Fig. 1, side view. Fig. 2, the head of the same, magnified, showing beneath the separated men-
tum. Fig. 3, front view of the head. Fig. 4, the same from below, showing the forelegs. Fig.
5, head of the perfect imago.

Figs. 6 —-9.— Phalena heteroclita.

The figures are copied from the original in the Mémoir. Prés. Acad. Paris, Vol. VI. The plate
given by Goeze is not exactly reproduced.

Fig. 6, the moth from above. Fig. 7, the same from below, Fig. 8, front view of the head.
Fig. 9, the eggs.

Figs 10,11.— Nymphalis Populi.

The figures are copied from the Bull. Acad. Bruxelles, Vol. IV.
Fig. 10, the butterfly from above. Fig. 11, front view of the head.

The plate was intended to contain all figures given for this kind of deformity. When it was
made, Mr. Bruinsma’s paper was not known to me.

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