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Psyche 7:95-99, 1894.

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Psyche, Vol. VII., 1894.
Plate UI.
AFRICAN NOCTUE (HOLLAND).
tREDUCED ONE FIFTH.)




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Psyche, Vol. VII,, 1894.
Plate IV.
AFRICAN NOCTUK (HOLLAND).
(REDUCED ONE FIFTH.)




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AFRICAN NOCTUE (HOLLAND).
(REDUCED ONE FIFTH.)




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PSYCHE.
ON THE PIGEON MITE, HYPODECTES FILIPPI.
,
BY HENRY B. WARD, PHSDa, LINCOLN, NEBR.
The November, 1892, number of
Insect Life contained a note by Prof.
D. S. Kellicott on a mite found in the
thymus of a pigeon ; on the authority of Murray ('77) it is said to be Hypoderas
coZumbae. Last spring while working
in the zoological laboratory of Harvard
University, I obtained specimens of this same form through the kindness of Dr.
W. McM. Woodworth, who had found
them some time previously in the loose
peritracheal tissue of a pigeon. Study
of the parasites and of the literature
bearing upon the group revealed so
many points of interest, especially con- cerning this stage in the life history,
that it seems proper to present here an
abstract of the work of previous investi- gators since the number of names under
which this form has been described
renders its study difficult. The appar-
ent lack of information among Amer-
ican students as to its interesting and
complicated life history is clue also, no doubt, to the inaccessibility of most of the papers bearing upon the subject.
Together with this review is given as
complete a bibliography as it has been
possible to prepare. For assistance in
this I am indebted to Mr. A. D.
Michael of London, Eng. and to Profes-
sor J. A. Lintner of Albany, N. Y.
Reference has been made in the bibli-
ography to reviews, abstracts and trans- lations of the original articles so far as known to the writer, but the list is
probably not complete.
The first published account of this
parasite seems to have been that of
Montagu ('08) whose description and
figures leave no doubt as to the close
relationship of the form he observed to
that found in the pigeon, while at the
same time the specific identity of
the two forms must always remain un-
certain on account of the incoimplete-
ness of liis description. He named the
form which he found in the gannet,
CeZZuZaria Basso&.
Two Italian naturalists, G611e ('48)
and Filippi ('61), were the next to
record observations on similar mites.
Filippi made a new genus, Hyfodectes,
to include them and described five spe-
cies parasitic in the areolar tissue of
various birds, each species receiving a
name from its host.*
I11 the same year Giebel ('61), with:
out knowledge of this last paper, pub-
lished a posthumous article by Nitzsch
*His synopsis of these species together with their hosts is to be found in Mkgnin ('79, p. 131).



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96 PSYCHE. [June-Sept. 1894.
in which a number of species were fig-
ured and described under the
name of
Hypoderas, n. g.
In a note some time
later Giebel ('63) acknowledged the
identity of Nitzsch's genus Hypoderas
with Hypodectes Fil., which had been
brought to his attention and which
might justly claim priority. In this
note Giebel criticizes the plan of Filippi in naming the species after the host
since, as Nitzsch emphasizes, the same
species may be found in more than one
host and, on the other hand, the same
host may harbor more than one species
of the parasite.
Robertson ('66) published a short
account of the discovery of a mite para- sitic in the English pigeon, but he
appears to have been entirely ignorant
of the work of his predecessors, except
Montagu, whom he quotes. He gives
a short account of the anatomy of the
mite, recognizes it as an imperfect form, remarks upon its evident relationship to the Sarcoptidae and announces his in-
tention of pursuing the subject further, a purpose which seems never to have
been fulfilled.
Gersticker ('67) makes a brief men-
tion of Robertson's work and identifies
this mite as a species of Hy-podectes
Fils*
A full account of the anatomy and
histology of Hyjodectes coIz~mbae, -n.
sp. is to be found in Slosarsky ('77) .f In this paper, read before a congress of Russian naturalists at Warsaw in 1875,
*The species which Robertson examined is said by Mkgnin ('79, p. 131) to be the same as Filippi's Hypo- dectes nycticomcis.
+ On the authority of Mkgnin ('79).
the author adds a new species to the
genus Hypodectes Fil. He found no
internal structure at all except the
narrow muscle bands just under the
cuticula by which the movements of the
body and appendages are effected.
Further than this the interior of the
body consisted of a granular vesicular
mass in which cell structure could not
be demonstrated*
To the researches of Mdgnin is due
the greater part of our knowledge of the life history of this form.
In a series of
papers ('73-'79) he established the
larval nature of a number of mites
without mouth-parts which had previ-
ously been regarded as distinct genera,
and showed them to be merely abnor-
mal ( ?) stages in the life history of
other known species. With Ch. Robin
(Robin et Mkgnin '77) he investigated
among others the form found in the
pigeon and showed it to be an abnormal
(adventitious or "hypopia1"t) nymph
of Rerolichus falciger M6gnin.
* In sections cut last spring I was equally unable to find either nuclei or cell walls in this granular mass. Claparkde ('63) has shown that the metamorphoses of the Acarina are not simply ecdyses, but that the body undergoes an extended hystolysis.
+ Hypopial, adj. derived from Hybopus. Megnin ('73, p. 402) says "The conclusions to be drawn from my observations is that the genera Hypopus, Homopus and Trichodactyizts, and the flumerous species which have been established as subdivisions of those genera, must be stricken from our zoijlogical nomenclature. The word Hypopus may he retained, but only as a common name serving to designate the curious cz~z'rassecf, heteromorphoiis and adventitious nymph of the Tyro- glyphi whose office is the preservation and dissemina- tion of the species to which it belongs." The name was afterwards used by the same author for similar nymphs in other families, e. g. Pterolichus (Robin et Mkgnin, '77' P- 403)-




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June-Sept. 1894.1 PSYCHE. 97
Later Megnin alone ('79) published
an extensive memoir on the mites of
the cellular tissue of birds. Therein he' reviews the work of many of his prede-
cessors and adds to his own previous
contributions a more extended account
of this stage and of its relation to the life history. Megnin had found in
Lo/hyrus coronatus Vieill. not a
single form but two, one of which he
showed to be the male nymph of Ptero-
fichus falciger; the other he was
inclined to regard as the female nymph
of the same species on account of its
strong resemblance to the normal
female. M6gnin explains the occur-
rence of these forms in the cellular
tissue in the following way :
So long as the conditions of existence
remain the same, the succession of
stages in the life history of the mites is invariably egg, larva, normal nymph,
male or female, and then the egg again.
If, however, the conditions change, if
the food or shelter necessary to life
begin to disappear, the colony seems
doomed to destruction. Certain species
escape that fate, thanks to a curious
biological phenomenon which we dis-
covered and named adventitious or
hypopial metamorphosis. This has
been found to occur in four species of
TyrogIy$kus and in one of Pteroli-
chm, P. falciger, a plumicolous sar-
coptide of the pigeon. . . . When
a pigeon invaded by a colony of these
mites begins to pull out its feathers, the mites are deprived of shelter and of the secretions which serve them as food,
the normal life cycle is arrested and the normal nymph instead of giving rise to
a male or female increases in size and
brings out of its skin a new form, fitted to a new mode of life. This form is
worm-like and has been described and
figured under the name of an adventious
or hypopial nymph ; it introduces itself into the follicles of the feathers, or even by the respiratory organs and reaches
the cellular tissue, especially the peri- tracheal, which is very loose in birds.
Here it lives and grows by absorption,
for it has neither orifices nor internal organs, and returns to the surface only
when the normal conditions are estab-
lished.
Megnin found a normal nymph of
Pterolichus falcig'er enormously de-
veloped and in process of metamorpho-
sis; it contained the larger hypopial
nymph. As to the nature of the
smaller form which is supposed to be
the hypopial nymph of the female, he
was not able to furnish any direct
proof.
Ti-ouessart et Mkgnin ('85) in their
revision of the group, have placed the
adult, of which Hypodectes is the
hypopial nymph, in a new genus, Pal-
cifer.
Murray ('77) refers to the form found
in the pigeon under the generic name of
Hypoderas; but his description and
figures are entirely inadequate for iden- tification.
In the United States this form has
been reported several times.
H. Gar-
man ('84) was the first to record its
occurrence. He described it under its
proper name, and recognized its nym-




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9 8 PSYCHE. [June-Sept. 1894..
phal character. Leidy ('go) noted very nymphs of Tyroglyphus and some briefly its occurrence in the blue allied genera. heron.* Kellicott's note ('92) has 3. That it is not all individuals that already been referred to ; and to this list "become Hypopi^ but only a few. must be added its occurrence in the
pigeon at Cambridge, Mass., as found
by Woodworth. It will thus be seen to
be widely distributed and will no doubt
be found in most localities. It may be
hoped that more extended observations
will furnish further evidence on its life history.
This review should not be terminated
without calling attention to a paper by
Michael ('84) which, though it does
not deal with the mite found in the
pigeon, is of great interest from its bear- ing on the hypopial stage in other mites. The author first gives a very complete
review of the liteniture on Hypofis
and tabulates at the close of that part of his paper the eight different views as to what a Hypo/us really is. His own
experiments are then given and after
showing their entire incompatibility
with six of the diverse theories, he says (p. 379) : "My own opinion decidedly
confirms M6gnin's view that the true
Hyfopus is a heteromorphous nymphal
form of Tyrogly/hzts, and possibly of
some allied, or other, genera." At the
close of his paper (p. 389) Michael
summarizes his conclusions thus : "It
appears to me :
I. That the true Hytopi are not
adult animals, but are a stage in a life history.
3. That they are heteromorphous
*There is no evidence, however, that this is the same species.
4. That the hypopial period takes the
place of that between two ecdyses in the ordinary life history.
5. That, in those species which I
have examined, the hypopial stage
commences with the second nymphal
ecdysis.
6. That the change to Hypo$us is
not caused by unfavorable circum-
stances, and is not any extraordinary or exceptional circumstance, but is a pro-
vision of nature for the distribution of the species occurring irrespective of
adverse conditions.
7. That, in the present stage of our
knowledge, we can no more say why
one nymph becomes a Hypopus and
another does not, than we can say why
one ovum produces a male and another
a female."
It will be seen from this .that while
observations and experiments of Michael
serve to establish and fix with greater
precision M6gnin's view as to the occur- rence of such a stage, the conclusions of the two authors as to the cause of its
occurrence are widely at variance. To
be sure Michael's work does not touch
the hypopial form found in the pigeon
and it is difficult to see how the explana- tion he offers with evident probability
for the other forms, i. e. "the distribu- tion of the species irrespective of adverse conditions," could possibly apply to the case of a hypopial form which like that
from the pigeon lives in an inactive




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June-Sept. 1894.1
PSYCHE.
state in the tissue of the body of a
bird.
Note. Since the above was written
a year ago, there have been further
references to similar parasites. I wish
to call attention only to the parasite of the Goura, found by Dr. L. Karpalles,*
which does not seem to have been
identified by the finder. It is probably this genus.
University of Nebraska,
Lincoln, Neb.
Mch. 1894.
BIBLIOGRAPHY.
BARALDI, GIOV.
'75. Stato particolare di una ninfa d'Aca- ride, Hypodectes carpo'phagae, n, sp.
Atti Soc. Tosc. sci. nat., tom. i, p. 87-103, I taf.
CLAPAR~DE, ED.
'68. Studien an Acariden. Zeitschr. fur
wiss. zool., bd. xviii, 4, p. 445-546, taf. xxx-XI.
FILIPPI, F~LIPPO DE.
'61. Note zoologiche. I, Hypodectes, novo genere di Acaridi proprio degli ucelli.
Archiv. zool. anat. e fissiol. (Genova)
fasc. i, p. 54-60, tav. v.
GARMAN, H.
'84. Pterolichus falciger, Megnin, observed in the United States. Amer. nat., vol.
xviii, p. 430-431, I wood cut.
GENE, G.
'48. Brevi cenni su un acaridio del genere dei Sarcopti che vive sulla Strixflammea, Scritto postumo, Torino 1848.
GERSTACKER, A.
e'67. Bericht fiber die wissenschaftlichen leistungen im gebiet der entomologie
wahrend des jahres 1865-66, ii. halfte,
* Ver. 2001.-bot. gesellsch. Wien, bd. xlii, p. 46-7. Archiv fur naturgeschichte, bd. xxxiii.
theil 2, p. 305-533.
GIEBEL, C. G.
'61. Die Milbenarten der gattung mode-
ms Nitzsch. Zeitschr. f. d. gesammt.
naturw., bd. xviii, p. 438-444.
'63. Hypoderas Nitzsch. = Hypodectes
Filippi. Zeitschr. f. d. gesammt. naturw., bd. xxi, p. 79-80.
KELLICOTT, D. S.
'92. Hypoderas columbae. A note. Insect
life, vol. v, no. 2, p. 77-78, I fig.
LEIDY, Jos.
'go. Hypoderas in the Little Blue Heron. (Florida caerulea.)
Proc. acad. nat. sci.
Philad., 1890, p. 63.
MEGNIN, P.
'73. Sur la position zoologique et le r61e des Acariens parasites connus sous les noms
d'Hy$optis, Homo'ptis et Trichodactylus., Comptes rendus., tom. Ixxvii, p. 129-1323 p. 492-3. Translated in Ann. mag. nat.
hist., ser. 4, vol. xii, p. 429-30.
'74. Memoire sur les Hytopus (Dugds)
Acariens parasites encore nommds Homo-
å´pu Koch et Trichodactylus L. Dufour.
Jonrn. anat. et physiol., tom. x, p. 225- 254-
'76. Note sur la faculty qu'ont certains Acariens, avec OLI sans bouche, de vivre sans nourriture pendant des phases entiere de leur existence, et meme pendant toute leur vie. Comptes rendus, tom. Ixxxiii,
p. 993-995. Translated in Ann. mag.
nat. hist., ser. 4, vol. xix, p. 270-271. '79. Les Acariens parasites du tissu cellu- laii-e et des reservoirs afiriens chez les oiseaux. Journ. anat. et physiol., torn. xv, p. 123-1153, pi. vii-viii.
MICHAEL, A. D.
'84. The Hy-pofis question, or the life
history of certain Acarina. Journ. Linn. soc. Lond., Zool., vol. xvii, p. 371-394, pi. xv.
MONTAGU, G.
'08. Observations on some peculiarities
observable in the structure of the gannet ;



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[June-Sept. 1894.
and an account of an insect discovered
to inhabit the cellular membrane of that bird. Mem. Wernerian nat. hist. soc.,
vol. i, p. 176-193, pi. vii, figs. 1-3.
MURRAY, A.
77. Economic entomology, Aptera. Scrib-
ner, Welford, and Armstrong, N. Y.
ROBERTSON, C.
'66. Note on an undescribed species of
Acarus found in the pigeon, CoZumba
Zivia. Quart. journ. micr. sc., n. s. vol. vi, p. 201-203, 4 figs. Abstracted in
Amer. nat., vol. iii, p. 389.
ROBIN, CH., AND MEGNIN, P.
'77. Mdmoire surles Sarcoptides plumicoles. Journ. anat. et physiol., tom. xiii, p.
209-248, pi. xi-xii, p. 391-429, pi. xxii- xxv, p. 498-521, pi. xxvi-xxix, p. 629-656, pi. xxxvi-xxxviii.
SLOSARSKY, A.
'77. On the anatomy and system at?^ posi- tion of Hypodecfes columbae, n. sp.
Warsaw, 1877, 14 pp., I pi. (In the
Russian language.)
TROUESSART E.-L. ET MEGNIN, P.
'85. Les Sai-coptides plumicoles ou Anal- &sines. I. Les Pt4roliches. Journ. de
tnicrogr., tom. viii, p. 92-101, 150-157, 2 I 1-219, 257-266, 331-338, 380-385, 428- 436, 527-5339 572-5797 tom. ix, P. 63-70,