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PARASITOLOGY
CAMBRIDGE UNIVERSITY PRESS C. F. CLAY, Manager LONDON: FETTER LANE, E.C. 4
H. K. LEWIS & CO., LTD., 1 36, GOWER STREET, LONDON, W.C. I
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CHICAGO : THE UNIVERSITY OF CHICAGO PRESS
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BOMBAY, CALCUTTA, MADRAS! MACMILLAN & CO., LTD.
TOKYO! THE MARUZEN-KABUSHIKI-KAISHA
A ll rights reserved
PARASITOLOGY
EDITED BY
GEORGE H. F. NUTTALL, F.R.S.
QUICK PROFESSOR OF BIOLOGY AND DIRECTOR OF THE MOLTENO INSTITUTE FOR RESEARCH IN PARASITOLOGY IN THE UNIVERSITY OF CAMBRIDGE
VOLUME XIV 1922
*
CAMBRIDGE
AT THE UNIVERSITY PRESS
1922^
PRINTED IN GREAT BRITAIN
/
5ll.o'5
?A
v, I ^
CONTENTS
No. 1 (April)
PAGE
Baylis, H. A. Observations on certain Cestodes of Rats, with an Account of a New Species of Hymenolepis . (With 4 Text-figures)
Baylis, H. A. Note on the Habitat and Structure of Crassicauda [Nema-
toda]. (With 3 Text-figures) .
Dudgeon, G. C. Occurrence of Hdigmosomum brasiliense Trav. in England Hopwood, A. T. An interesting case of Cysticercus fasciolar is infesting the Brown Rat. (With Plate I) •••••••
Ghosh, Ekendranath. On a New Ciliate, Balantidium blattarum, sp. nov., Intestinal Parasite in the Common Cockroach ( Blatta amcvicana).
(With 1 Text-figure) .
Cunliffe, Norman. Some observations on the Biology and Structure of Ornithodorus savignyi Andouin. (With 3 Text-figures)
Allen, J. A. and Wickware, A. B. A Preliminary Note on Parasites infesting Domesticated Silver Black Foxes in Canada
Bezzi, Professor M. On the Dipterous Genera Passoromyia and Ornitho- musca , with Notes and Bibliography on the Non-Pupiparous Myiodaria parasitic on Birds .••••••••
Pavlovsky, E. N. Description of a Box for Collecting and Transporting Living Insects, etc. (With 6 Text-figures) ......
Caldwell, Fred C. An Accidental Infection with Uncinaria . Bullamore, Geo. W. Nosema apis and Acarapis ( Tarsonemus ) woodi in relation to Isle of Wight Bee Disease .......
Hill, Laurence. Three New Species of Trichodedes from Cephalophus monticola and Procavia capensis from South Africa. (With Plate II) . Morris, Hubert M. On the Larva and Pupa of a Parasitic Phorid Fly — Hypocera incrassata Mg. (With Plate III and 4 Text-figures)
Ferris, G. F. The Mallophagan Family Trimenoponidae. (With 8 Text- figures) ...••••••••*
Boulenger, C. L. On Ascaris vitulorum Goeze. (With 3 Text-figures) Nuttall, George H. F. Notes bearing on Van Beneden, Leuckart and Sonsino whose portraits appear in Parasitology , xiv, No. 1. (Portrait- plates XII-XIV) .
1
9
13
14
15 17 27
29
47
51
53
63
70
75
87
93
532133
VI
Contents
No. 2 (June)
Nuttall, George H. F. The Molteno Institute for Research in Para¬ sitology, University of Cambridge, with an Account of how it came to be founded. (With Plates IV-VI and 6 Text-figures)
Yokogawa, Sad amu. The Development of Heligmosomum muris Yoko- gawa, a Nematode from the Intestine of the Wild Rat. (With Plates VII-XII and 8 Text-figures) ........
Haviland, Maud D. On the Larval Development of Dacnusa areolaris ,Nees (Braconidae), a Parasite of Phytomyzinae (Diptera), with a Note on certain Chalcid Parasites on Phytomyzids. (With 5 Text-figures) .
Nuttall, George H. F. Notes bearing on Dufour, von Siebold and Goodsir, whose portraits appear in Parasitology , xiv, No. 2. (Con¬ tinuing the Series begun in Vol. xm.) (Portrait-plates XV-XVII)
Ferris, G. F. and Cole, F. R. A Contribution to the Knowledge of the Hippoboscidae (Diptera Pupipara). (With 20 Text-figures) .
Boulenger, C. L. The Structure and Systematic Position of Strongylus poly gyrus. (With 4 Text-figures) .......
Leigh-Sharpe, W. Harold. A Parasitic Copepod belonging to the Genus Medesicaste (Kroyer), and its Relation to the Tumours it produces on the Fish, Trigla gurnardus. (With 7 Text-figures) . . . .
Keilin, D. Parasitism and Symbiosis. (A Review) .
Nos. 3 and 4 (December)
Kudo, R. On the Morphology and Life -history of a Myxosporidian, Leptotheca ohlmacheri , parasitic in Rana clamitans and R. pipiens. (With Plates XIII-XX, containing Figs. 1-111) . . . .
Cawston, F. G. Some Notes on the Differentiation of closely-allied Schistosomes ...........
Faust, Ernest Carroll. Notes on Larval Flukes from China. (With Plates XXI and XXII) .........
Flattely, F. W. Considerations on the Life-history of Tapeworms of the Genus Moniezia ...........
Balfour, Andrew. Observations on Wild Rats in England, with an account of their Ecto- and Endoparasites. (With Plates XXIII and XXIV, 1 Text-figure and 2 Charts) .......
Vincent, Mary. On the Life-history of a New Gregarine: Pyxinia anobii n. sp. Intestinal Parasite of Anobium paniceum L. (Coleoptera). (With 5 Text-figures) ...........
PAGE
97
127
167
174
178
206
214
220
221
245
248
268
282
299
Contents
Vll
PAGE
Richards, Major W. G. Note on Dracunculus medinensis (Guinea-worm) 307
Pavlovsky, E. N. and Anitchkov, N. N. Tetracotyle sogdiana — a New Trematode Parasite of the Fish, Schizothorax intermedius, with a description of adhesive peritonitis produced by the parasite in its host.
(With Plate XXV) .......... 309
Iwanow, E. A Contribution to the Biology of Trypanosoma equiperdum . 315
Gray, R. A. Harper. The occurrence of a Leech (Trocheta subviridis) in
an Allotment ........... 320
Warburton, Cecil. The Warble-Flies of Cattle. Hypoderma bovis and
H. lineatum. (With 3 Text-figures) ....... 322
Dobell, Clifford. The Discovery of the Coccidia ..... 342
Browne, Frank Balfour. On the Life-history of Melittobia acasta,
Walker; a Chalcid Parasite of Bees and Wasps. (With Plate XXVI) . 349
Ghosh, Ekendranath. On a New Ciliate, Balantidium ovatum , sp. nov., an Intestinal Parasite in the Common Cockroach ( Blatta americana).
(With 1 Text-figure) . . . . . . . . . .371
Kostylev, Dr N. Sur les Acanthocephales de l’Eider (Somateria mollis-
sima L.). (With 5 Text-figures) ........ 372
Nicoll, William. Recent progress in our knowledge of Parasitic Worms
and their relation to the public health . . . . . .378
Baylis, H. A. Notes on the Collection and Preservation of Parasitic
Worms ............ 402
Bulloch, Wm, Dobell, Clifford, and Nuttall, George H. F. Notes bearing on Spallanzani, Lewis and Cobbold, whose portraits appear in Parasitology , xiv, Nos. 3-4. (Continued from p. 177.) (Portrait- plates XVIII-XX) .......... 409
Index of Authors . 419
Index of Subjects . 421
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Volume XIV
APRIL, 1922
No. 1
OBSERVATIONS ON CERTAIN CESTODES OF RATS, WITH AN ACCOUNT OF A NEW SPECIES
OF HYMENOLEPIS.
By H. A. BAYLIS, M.A., D.Sc.
(Published by 'permission of the Trustees of the British Museum.)
(With 4 Text-figures.)
During some investigations in which he has been engaged on the internal parasites of rats in the British Isles, Mr G. C. Dudgeon, of the Wellcome Bureau of Scientific Research, has kindly submitted to the writer examples of all the species of adult Cestodes found.
Two species of Hymenolepis are well known to occur in the brown and black rats, viz. H. diminuta and a smaller form most commonly referred to in literature as H. murina. The latter is of exceptional interest on account of the vexed question as to its identity with the human parasite H. nana. In more or less direct connection with this question a very extensive literature has grown up, which it is impossible to review fully here. It may be said, however, that the general result of researches by numerous authors, while still inconclusive, tends to show that the forms occurring in man and the rat are morphologically identical, while on physiological grounds there is some justification for regarding them as distinct species, subspecies, or at least varieties. This view is held on account of the many unsuccessful attempts that have been made to infect rats with H. nana of human origin, and on the other hand to infect man with “ H. murina
A few of the more important contributions to the subject may be men¬ tioned. The celebrated researches of Grassi and his collaborators (1887-1892) along these lines led them to regard nana as a variety of “ murina f and they concluded that infection was capable of transmission from rats to man, though the experimental evidence was very scanty. On the other hand, on morpho¬ logical grounds, as well as from considerations of geographical distribution, other authorities (Moniez, Blanchard, von Linstow) regarded the two forms as distinct species. The last-named author (1896) gives comparative measure¬ ments and other data compiled from his own and numerous other authors’ observations, which he appears to have regarded as conclusive, but which have not proved equally convincing to later writers. Stiles (1906) favoured the separation of the forms as “host subspecies,” and renamed “//. murina " Parasitology xiv
I
2
Hymenolepis longior n. sp.
H. nana fraterna. A recent important contribution to the question is that of Joyeux (1919). This author returns to the position taken up by v. Linstow, in regarding the forms as distinct species. He proposes to call the rat-parasite H. fraterna.
The supposed morphological differences between H. nana and “ H. murina ” must be discussed more fully later. It will be well, however, first to consider Mr Dudgeon’s Hymenolepis material, which appears to have rather important bearings on this question.
Among the material there occur three forms, viz.
(1) Hymenolepis diminuta (Rud.).
(2) A form which may be called H. nana fraterna.
(3) A form very closely related to the latter, but quite distinct, which does not appear to be referable to any of the other species of Hymenolepis
Fig. 1. Hymenolepis longior. Two scolices in dorsal or ventral view: A., with evaginated rostellum; B., with invaginated rostellum.
from rats or their relatives, of which the descriptions are accessible to the writer (see list, p. 7). This form, accordingly, it is proposed to regard as a new species.
Hymenolepis longior , sp. n.
The worm is very slender, fragile and semi-transparent. Complete speci¬ mens attain a length of 45 mm., or even 60 mm. in a stretched condition. The maximum width of the strobila (usually occurring in the region of the gravid segments, but sometimes, in contracted specimens, more anteriorly), is from 0*42 to 0-53 mm. In exceptionally contracted individuals it may even reach 0*65 mm. The scolex is flattened dorso-ventrally, more or less rect¬ angular in transverse section, and assumes very different shapes (Fig. 1)
3
H. A. Baylis
according to the state of contraction. It is, in any case, considerably wider than the “neck” which follows it. Its diameter, measured from side to side, is from 0-21 to 0*26 mm., with a mean measurement of about 0-24 mm. The suckers, situated somewhat laterally, have an outside diameter of 0*075 0*093 mm. There is a well-developed rostellum, having a diameter of 0*07- 0*08 mm., and armed with a single row of hooks of the characteristic shape, 21 or 22 in number (rpost commonly 22), and 19-20 /x in length.
The “neck” is unsegmented for some little distance behind the scolex. The total number of recognisable segments in the strobila may reach 000 oi more. Rudiments of genital organs begin to be recognisable at about the 50th segment from the anterior end; “mature” segments i segments
Fi'T. 2. Hymenolepis longior. Scolex, with evaginated rostellum, viewed ev face.
containing male and female organs in a state of full functional activity — at about the 250th. The number of gravid segments at the posterior end varies between (roughly) 00 and 100. The segments are broader than long through¬ out, though the length tends to become nearly equal to the breadth in the more posterior segments, and the ratio of length to breadth is (except in greatly contracted specimens) much higher in mature segments than in corresponding
segments of //. nana or H. nana fraterna.
In a mature segment (Fig. 3) the most conspicuous organ is the ovary, which is large and compact, rather irregularly but not very deeply lobed, and occupies roughly one-third of the width of the segment, having a greatest diameter (from side to side) of about 0*15 to 0*17 mm. The ovarian ova have a diameter of 0*015 mm. The yolk-gland and shell-gland lie in the usual
1—2
4
Hymenolepis longior n. sp.
positions with regard to the ovary. The three testes are relatively much smaller than in H. nana and H. nana fraterna, and, instead of being almost equal in diameter to the antero-posterior length of the segment, they occupy only a small space close to its posterior border. They are generally more oval in shape than those of the other species, which are almost spherical. Their greatest diameter varies from 0-05 to 0-075 mm. The middle testis varies
e.
c.s.
n.
Fig. 3. Hymenolepis longior. Three mature segments; dorsal view. (From a whole preparation.)
c.s., cirrus-sac, with contained internal seminal vesicle; e., excretory canals; n., longitudinal nerve; oy., ovary; r.s., receptaculum seminis; s., shell-gland; t., testis; v., vitelline gland; v.s.e., external seminal vesicle.
01mm.
Fig. 4. Hymenolepis nana fraterna. Two mature segments; dorsal view. (From a
whole preparation.) Lettering as in Fig. 3.
in position with regard to the yolk-gland, and may lie dorsally to it or on either side of it. In the other species it seems to be always in the centre of the segment and dorsal to the yolk-gland. The genital pore is situated at about the middle of the lateral border of the segment, or very slightly in front of it. The cirrus-sac, which measures about 0-15 mm. in length, passes, as usual in Hymenolepis, dorsally to both the longitudinal excretory vessels and to the
o
H. A. Baylis
longitudinal nerve. It is club-shaped, as in the othei species, and expands towards its inner end to a maximum width of 0-025-0-03 mm. It is almost completely filled by the internal seminal vesicle. There is also a pear-shaped external seminal vesicle (Fig. 3, v.s.e.), which occupies the space between the inner end of the cirrus-sac and the ovary. V entrally to this lies a large re- ceptaculum seminis (Fig. 3, r.s.), being the expanded inner end of the vagina, which runs along the ventral side of the cirrus-sac.
The passage from mature to gravid segments is very gradual. The latter are almost completely filled by the sac-like uterus, which contains a large number of ova. These have a thin outer membrane and a thicker, chitinised inner shell, between which there is a rather deeply-staining granular material which sometimes appears to form a third membrane.” The inner shell is dis¬ tinctly lemon-shaped, as opposed to the almost spherical inner shell of H. nana and H. nana f rater na, and it possesses at or near each pole a well-developed thickening. The terminal filaments present in the other species appear to be absent. Measurements taken by Mr Dudgeon and Dr A. C. Stevenson from ova in a fresh state in the faeces of infected rats were as follows: outer membrane, 49-60 /jl x 42-48 ju.; inner shell, 28-5-35*25 /x x 22-5-28-.) p,, the knobs at the poles measure 2-4 /x; onchosphere, 24-35 /x x 21-27 /x;
length of embryonic hooks, 10-5-15 /x.
The following table of measurements and other data taken from Mi Dudgeon’s material will help to bring out some of the differences between H. nana J rater na and the new species.
Total length
Width of strobila (max.)
Width of scolex (great varia¬ tion according to degree of contraction)
Suckers, outside diameter Rostellum, diameter Number of hooks Length of hooks Number of segments
Segment at which rudiments of genital organs begin to appear (roughly)
Segment at which mature geni¬ tal organs appear (roughly)
Number of gravid segments (roughly)
Ova, diameter of outer mem¬ brane1
Ova, diameter of inner shell1 Ova, diameter of onchosphere1 Embryonic hooks, length
II. nana fraterna Up to 20 mm.
0-27 — 0-51 mm.
010 — 0-23 mm. Mean about 019 mm.
0-05 — 0 07 mm.
0 05 — 0 07 mm.
22 — 20 (usually 22 or 23)
16 - 18yU.
Up to 250 35th
75th
20—70
07-5 — 90/a x 00 — 87/x
40-5— 51m x 30—43-5 /a 30 — 45/a x 31*o — 40-5/a 15/a
H. longior
Up to 45 mm.
(00 mm. if stretched)
0-42 — 0-53 mm.
(0-05 mm. if contracted)
0-21 — 0-20 mm.
Mean about 0-24 mm.
0-075—0-093 mm. 0-07—0-08 mm.
21 — 22 (usually 22) 19—20/a
Up to, or over, 000 50th
250th
00—100
49—00/a x 42 — 48/a
28-5—35-25/a x 22-5—28-5/a 24— 35/a x 21— 27/a 10-5 — 15/a
1 These measurements were taken by Mr Dudgeon and Dr Stevenson from fresh ova in the faeces of rats.
6
Hymenolepis longior n. sp.
Many authors have given measurements of Hymenolejris nana and “ H. murina ,” and it appears to the writer that, among the individuals ascribed to the latter form, some in reality belonging to the species now described as new have frequently been included. The new species appears, in fact, to have been confused with “//. murina ” ever since the original description of the latter by Dujardin (1845), and to have been responsible for some of the alleged differences between it and the H. nana of man. The discrepancies in the measurements of total length given by different writers for “ H. murina ” are greater than could readily be accounted for by different states of contraction in complete individuals of the same species. 25 mm. is given by several authors as the maximum for H. nana, whereas “ H . murina ” is frequently said to reach 40 or 45 mm. — the latter figure agreeing with that given above for H. longior. According to Mr Dudgeon’s material, H. nana fraterna (the true “H. murina ”) rarely exceeds 20 mm. in length, unless abnormally stretched. Other measurements, such as those of the maximum width of the strobila, the width of the scolex, and the diameter of the suckers, are so variable that little reliance can be placed upon them. They are certainly, as Joyeux (1919) has pointed out, useless as criteria for distinguishing H. nana from H. nana fraterna ; and they seem to afford as little help in separating H. nana fraterna from H. longior. It may be mentioned that some of the extreme measurements given by previous writers, especially for the width of the strobila and the diameter of the suckers, find no parallel among the present material. Thus the width of H. nana has been recorded as reaching 0-7 mm., and that of “ H. murina ” 0-9 mm. Similarly the diameter of the suckers in H. nana is said to be sometimes as much as 0-105 mm. Reference to the table given above will show that nothing approaching these figures has been ob¬ served among Mr Dudgeon’s material, and it may be suspected that some of them, at least, are erroneous, or due to excessive artificial pressure on the specimens. Much, of course, depends upon technique in handling soft-bodied worms, and for this reason the value of such measurements is often questionable.
In the present case there is one character, depending only in part on measurements, which seems to be of great importance. The inner shell of the egg, being composed of a relatively hard, chitinoid substance, is not subject to alteration by pressure or the action of reagents to the same extent as the soft parts. It has been repeatedly insisted upon that the inner shell of the egg of “ H. murina ” is lemon-shaped, and provided at each pole with a well-developed knob-like thickening; whereas in the egg of H. nana the inner shell is more rounded, with the polar knobs scarcely distinguishable, but with a filamentous process at each pole. Unfortunately the measurements of the inner shell have seldom been given. Von Linstow (1896) gives the following :
nana: 0-028 mm. (exceptionally 0-029 x 0-024 mm.), usually spherical.
murina: 0-031 x 0-023 mm., lemon-shaped, with knobs at the poles.
The lemon-shaped inner shell, with polar knobs, is highly characteristic
of the ova of H. longior, and it seems almost certain that those authors who have laid stress upon this feature in their descriptions of “ H. murina ’—among them Dujardin himself— were dealing, as regards the ova, with //. longior.
There remains one point of considerable interest to be mentioned. As is well known, the researches of Grassi and his collaborators have led to a general acceptance of the theory that H. nana (and this covers also H. murina") normally passes through the whole of its life-history in the intestine of the same host, its cysticercoid living in the villi of the small intestine, and not requiring an intermediate host. Nicoll and Minchin (191 1) found in the bodv - cavity of one of the fleas that infest rats in this country ( Ceratophyllus fasciatus ) a cysticercoid, of which they observe that, unless it is that of “H. murina ,” it must be that of some undescribed form, the scolex of which is indistinguishable from that of “ H . murina ” Johnston (1913) also found a cysticercoid in Australia in Ceratophyllus fasciatus and another rat-flea, Xenopsylla cheopis, which he regarded as that of “ H. murina. Now it seems highly probable, bearing in mind the great similarity between the scolices of H. nana fraterna and H. longior , that the latter is the adult foun into which this cysticercoid develops, and that these cases cannot be taken as e\ idence that H. nana fraterna ever makes use of fleas as intermediate hosts.
The following species of Hymenolepis are recorded in Muridae1:
With armed scolex:
Hosts
nana fraterna Stiles, 1906
( — Taenia murina Duj., 1845, e.p.)
microstoma (Duj., 1845)
contracta Janieki, 1904
criccti Janieki, 1904
muris-variegati Janieki, 1904 longior, sp. n.
With unarmed scolex: diminuta (Rud., 1819)
r dicta (Zschokke, 1888)
horrida (v. Linst., 1900) proccra Janieki, 1904
Epimys rattus E. norvegicus E. alexandrinus Mas musculus M. pumilis Micromys minutus Eliomys quercinus Epimys rattus E. norvegicus M us musculus Epimys norvegicus Mus musculus Cricetus cricetus ( C . vulgaris )
Mus variegatus Epimys rattus E. norvegicus
Epimys rattus E. norvegicus E. alexandrinus M us musculus H esperomys pyrrhorhi n us Epimys norvegicus ? Mus musculus Epimys norvegicus Arvicola amphibius
Scolex undescribed:
asymrnctrica Janieki, 1904 Microtus arvalis
crassa Janieki, 1904 Epimys iwrvegicus
Mus musculus
1 The descriptions of H. arvicolina, II. diminutoxdes and II. inexspectatu Cholodkovskv, 1912, have unfortunately proved inaccessible. The first appears to occur in Microtus arvalis ( Arvicola campestris), the two last in Epimys norvegicus.
8
Hymenolepis longior n. sp.
REFERENCES.
Dujardin, F. (1845). Histoire naturelle des Helminlhes. Paris.
Grassi, B. (1887) (a). Die Taenia nana und iiire medicinische Bedeutung. Centralbl. f. Bakt. i. 4. 97.
Grassi, B. and Calandruccio, S. (1887) ( b ). Einige weitere Nachrichten iiber die Taenia nana. lb. n. 10. 282.
Grassi, B. (1887) (c). Entwicklungscyclus der Taenia nana. lb. n. 11. 305.
Grassi, B. and Rovelli, G. (1889). Embryologische Forsehungen an Cestoden. lb. v. 11, 12. 370, 401.
- (1892). Ricerche embriologiche sui Cestodi. Alii Ace. Gioenia Sci. Nat. Catania (4),
iv. 15.
Johnston, T. H. (1913). Notes on some Entozoa. Proc. Roy. Soc. Queensland, xxiv. 63. Joyeux, Ch. (1919). H ytnenolepis nana (v. Siebold, 1852) et Hym. nana var. fralerna Stiles, 1906. Bull. Soc. Path. exot. Paris, xn. 5. 228.
Linstow, O. von (1896). Ueber Taenia ( Hymenolepis ) nana v. Siebold und rnurina Duj. Jena. Zeitschr. xxx. 571.
Nicoll, W. and Minchin, E. A. (1911). Two species of Cysticercoids from the rat-flea ( Ceratophyllus fasciatus). Proc. Zool. Soc. 9.
Stiles, C. W. (1906). Illustrated Key to the Cestode Parasites of Man. Bull. 25, Hyg. Lab. U.S. Pub. Health and Mar.-Hosp. Serv. Washington.
Addendum.
Since the preparation of the paper on Hymenolepis from rats, the author has seen descriptions of H. diminutoides, H. inexspectata and H. arvicolina given by Cholodkovsky in Ann. Mus. Zool. Ac. Sci., Petrograd, xviii. (1913), ppL 227-229. Although the names are here marked “spec, nova," they appear to have been first published in a catalogue of parasitic worms of the Army Medical Academy of Petrograd, in the previous year. It is this publication which was referred to as inaccessible, the redescriptions of 1913 having escaped notice.
H. diminutoides and H. arvicolina belong to the unarmed group of species. The description of H. inexspectata is very brief, and scarcely suffices to deter¬ mine whether this species is distinct from that described by the writer as H. longior. The number and size of the hooks, the size of the suckers and the arrangement of the testes, as described, indicate differences which may be of specific importance, but the other differences are such as to be accounted for by a greater degree of muscular contraction, and it is impossible to lay any stress upon them.
H. diminutoides is recorded from Microtus arvalis (Arvicola campcstris) as well as from the brown rat. H. straminea (Goeze, 1782), from Cricetus cricetus, should be added to the list of forms with armed scolex.
NOTE ON THE HABITAT AND STRUCTURE OE GBASSICAUDA [NEMATODA].
By H. A. BAYL1S, M.A., D.Sc.
I
(Published by permission of the Trustees of the British Museum.)
(With 3 Text-figures.)
Complete; specimens of this nematode are difficult to obtain, owing to the manner in which the worms bury themselves in the tissues of the urinogenital system of the Cetacea in which they are found. Mr A. G. Bennett, to whom the writer has previously been indebted on more than one occasion for Crassicauda material, has recently sent home further interesting specimens from South Georgia, accompanied by notes and a sketch which throw some light upon the burrowing habits of the worm. A special effort was made to
obtain complete specimens by dissection.
In a previous note, in which an attempt was made to collect the lecoids of the occurrence of this remarkable genus, the writer (1916) quoted an inter¬ esting passage from a report by Mr J. B. Hamilton, on the foimation of “connective tissue masses” by the worms in the kidneys of Balaenoptera physalus. From Mr Bennett’s observations it appears that “masses” or nodules of a similar nature may also be formed in other tissues. I he ac¬ companying diagram, which is adapted from Mr Bennett’s drawing, shows the course taken by one of the worms in the tissues of the penis of a Balaeno- ptera (probably B. physalus). The caudal end of the worm hangs freely in the lumen of the urethra, through a perforation in the wall of which its body passes into the dense surrounding tissue. Through the kindness of Mr R. H. Burne, the writer has been enabled to compare Mr Bennett s sketch with a preparation of the penis of a Balaenoptera in the Museum of the Roy al C ollegc of Surgeons. This makes it clear that the actual tissue invaded by the parasite is the dense fibrous sheath of the corpus cavernosum.
According to Mr Bennett’s notes, the free portion of the worms vaiied from two to four inches in length. After traversing the fibrous tissue for some distance, the body passes into a dense nodule (A), where it becomes flattened and much coiled. The substance surrounding the worm in this nodule is in some cases putty-like, in others hard and apparently calcareous. The worm may be traced on again beyond this nodule for a longer or shorter distance, but eventually it passes into a second nodule (B) ol pus and fibrous
1 0
Crassicauda
tissue, in which the head is found. It is easy, therefore, to understand the difficulty hitherto experienced in obtaining an unbroken specimen of Crassi- cauda. The nodules, formed in the dense tissue of the penis or in other solid tissues, effectually prevent it from being pulled out; while its tortuous course renders the task of dissecting out the worm, without cutting it, a very difficult one, especially if the surrounding tissue has been hardened by reagents.
Mr Bennett succeeded, by cutting out one of the hard nodules (A) with the worm, in extracting the latter entire, and on examination it proved to be a male C. crassicauda (Crepl.). Anterior portions of females, almost certainly of the same species, were also obtained. This material enables some further details to be added to the description of C. crassicauda. The writer (1916,
Fig. 1. Diagram, adapted from a drawing by Mr Bennett, of an excised portion of the penis of Balaenoptcra, with a specimen of Crassicauda crassicauda in situ. The upper part of the draw¬ ing is in transverse section. In the lower part, the skin has been cut longitudinally over the urethra and reflected on either side. The urethra is represented as slit open longitudinally, c., cavernus core of corpus cavernosum; /., fibrous sheath of corpus cavernosum; s., skin of penis; u., urethra; X, caudal end of worm hanging freely in urethra; A, B, successive nodules formed by the worm.
1920) has published some notes on the anatomy, but the characters of the anterior end were still very imperfectly known. The cephalic papillae of a specimen thought to be C. crassicauda were described in the former paper, but in the second it was shown that at least two species exist, and this rendered the conclusions of the first less certain. In the light of the material now received, which is proved by the characters of the male tail to be C. crassi¬ cauda , it is possible to say that the characters of the head previously described agree with those of C. crassicauda. It is not, of course, possible to say whether they differ from those of C. boo pis, which is only certainly known from headless specimens. The accompanying figures (Figs. 2 and 3) show that the same papillae are present, and that they are arranged in the same way, as in the head of which an “ en face ” view was previously (1916) figured.
H. A. Baylis 1 1
The structure of the oesophagus and other organs of the anterior region has not hitherto been described. The mouth, which is laterally compressed, leads into a similarly compressed buccal cavity, with very thick cuticulai walls. This is followed by an oesophagus consisting of a relatively short
i.l.p.
Fig. 3. Crassicauda cvcissicuuda. Anterior end of male; lateral view. Lettering as in tig. 2.
anterior portion, which is scarcely, if at all, muscular, and a very long posterior portion, which is partly muscular and partly glandular. The latter portion may double upon itself several times in its course. An excretory pore has not
been detected.
12 Crassicauda
Measurements made from the complete male and an incomplete female
were as follows (in millimetres) :
<? $
Length of buccal cavity . . . 0*18 0-23
Length of anterior portion of oesophagus 1-85 1-95
Thickness of anterior portion of oesophagus 0-11 0-16
Length of posterior portion of oesophagus about 25-0 about 31-0 Thickness of posterior portion of oesophagus 0-3 — 0-4 0-47 — 0-8
Distance from anterior end to nerve-ring 0-5 0-4
(neck of worm contracted)
The genus Crassicauda has hitherto been assigned tentatively to the family Filariidae, but the posterior position of the vulva is a character difficult to reconcile with this arrangement. It may be remarked that in this feature, as well as in the general structure of the oesophagus and buccal cavity, now described for the first time, Crassicauda somewhat resembles Tetrameres (— Tropidocerca, Tropisurus), a genus usually included in the Spiruridae, but placed by itself by some authors in a family Tetrameridae. The globular form characteristic of the mature females of Tetrameres is not, however, seen in Crassicauda.
REFERENCES.
Baylis, H. A. (1910). On Crassicauda crassicauda (Crepl.) [Nematoda] and its Hosts.
Ann. and May. Nat. Hist. (8), xvn. 144-148.
- (1920). Observations on the Genus Crassicauda. Ibid. (9), v. 410-419.
OCCURRENCE OF HELIGMOSOMUM BRA SILIENS E
TRAV. TN ENGLAND.
By G. C. DUDGEON, C.B.E., F.E.S., etc.
Wellcome Bureau of Scientific Research.
In making the examination of a series of 400 rats, Helignwsomum krasiliense was found in 6 per cent, of those examined. The rats were all of the one species, Epimys norvegicus, and were chiefly from London and suburbs; the exceptions being one from Berkshire, one from Hampshire and two from
Hertfordshire. ,
The worms are found, generally, in the upper third oi the small intestine,
and the measurements, where they differ from those previously described for
this species, are given. ,1-00
Male. In all cases, measured, the average length of the male is -.-8 mm.
and the width 0-087 mm., giving a mean ratio of 32 : 1. The spicule averages
0-561 mm. The lobes of the bursa are asymmetrical in form; the latero-
ventral and externo-lateral rays of the left lobe run parallel, and generally
contiguous, throughout their length. The right lobe is much thickened and
normally enrolled into a bar. The dorsal lobe is short and bears three rays,
the central one terminating in two branches, which are again divided, the
inner digitation bearing a small node on its outer edge, the outer digitations
being slightly curved outwardly.
Female. The female averages 3-7 mm., with a thickness of 0-106 mm., giving a mean ratio of 34 : 1.
The worm has 14 longitudinal ridges visible in the median section ; those of the lateral area are deeper and more prominent than either the dorsal 01 ventral ones. These ridges diminish in number towards the anterior end, and appear to be all present posteriorly, but of more uniform and much reduced
dimensions. .
A comparison of the measurements of British with those of Australian
specimens may be of interest: Heligmosomum brasiliense Trav. was described
and figured by T. Harvey Johnston, M.A., etc., in the Proceedings of the
Royal Society of Queensland, vol. xxx. p. 56, for 1918, but with some rather
remarkable differences in measurement. The general form of the worms from
both continents is the same.
14
AN INTERESTING CASE OF CYSTICERCUS F ARC 10- LARIS INFESTING THE BROWN RAT.
By A. T. HOPWOOD,
Assistant in Zoology , West Virginia University , Morgantown , W. Va.
(With Plate I.)
Cystic erous fasciolar is, the larval stage of the common tapeworm Taenia taeniaeformis ( T . crassicolis), was recently found in great numbers in the liver of a brown rat. The rat measured 18 cm. in length and weighed about 275 g. ; it was vigorous and apparently healthy when caught, but when killed and examined, its liver was found to be heavily infested with the cysticerci.
This cestode in itself is common enough, but the great number of cysticerci attracted attention. On the ventral surfaces of the ventral lobes of the liver which are shown in the photograph (PL I), 108 of the cysts were visible and this region was the least infested. The dorsal surfaces of these lobes as well as the entire surfaces of the others harboured cysts almost to their full capacity. It is difficult to conceive how many more of the larvae could possibly have found room to encyst themselves. There was visible a total of 256 larvae, about 50 of which were visible from both surfaces. Many more may have been embedded in the tissue and invisible from either surface.
Possibly not more than nine-tenths of the liver was incapacitated and the bile ducts were not greatly obstructed, which probably explains why the animal was still apparently healthy. The cysticerci were identified in the laboratory and their determination was confirmed by reference to the Zoo¬ logical Division of the Bureau of Animal Industry of the U. S. Department of Agriculture.
RASITOLOGY, VOL XIV. NO. 1
PLATE I
Brown Rat infested by Cyxticercu a Jaaciolaris.
THE LIBRARY OF THE
UNIVERSITY Or ILLINOIS
*
15
ON A NEW CILIATE, BALANTIDIUM BLATTARUM, SP. NOV., INTESTINAL PARASITE IN THE COMMON COCKROACH ( BLATTA AMERICANA).
By EKENDRANATH GHOSH, M.Sc., M.D.,
Professor of Biology , Medical College , Calcutta.
(With 1 Text-figure.)
The parasite herein described was found in the intestinal contents of Blatta americana at Calcutta. The species seems to be comparatively rare and was observed twice only. It may be named B. blattarum sp. nov.
Diagnosis: Body irregularly pyriform, slightly less than twice as long as its greatest transverse diameter, and circular in transverse section. Anterior
Length 0*09 mm. (See Text-fig.)
Balantidium Clap, and Lach., was emended by Biitschli (in Bronns, Tierreich, Protozoa, 3 Abt., pp. 1723-1725), by the removal of two species (B. duodeni and B. rotundum) which were placed together in a separate genus. Balantidiopsis. He characterised Balantidium by its round transverse section and its habitat in the large intestine, and Balantidiopsis by its flattened body and its habitat in the small intestine.
16
Balantidium blattarum n. sp.
Scheier (Arb. d. St Petersburger naturf. Gesellsch. xxvm. No. 4) defines the two genera in the following manner: Balantidium: body egg-shaped or cylindrical; contractile vacuoles numerous; macronucleus oval or horseshoe- shaped. Balantidiopsis : body broadly oval; contractile vacuole single, posterior; macronucleus spherical.
This year I described two new species of Balantidium and Balantidiojpsis (Bull. Carmichael Med. College, No. 2) to which I appended synopses of the known species of both the genera. But the recognition of a large number of species of both the genera has made all the distinguishing characters fail one after another, viz. the number of contractile vacuoles, the form of the macro¬ nucleus and the nature of the habitat. Lastly, the discovery of species with oval transverse section of the body breaks the last distinction between the two genera. Under these circumstances it seems necessary to reunite the two genera into Balantidium , as already suggested by Bezzenberger (Arch. f. Protistenk. m. 154-156).
As regards habitat, the already known species of Balantidium have been found in the gastric cavity of Hydra (B. hydrae, Entz, Arch.f. Protistenk. 1912, xxvii.) and medusa, and in the intestinal canal of the mollusc and of many vertebrates, specially the amphibians. The present species seems to be the first described from the intestinal tract of an arthropod.
17
SOME OBSERVATIONS ON THE BIOLOGY AND STRUCTURE OF ORNITHODORUS SA VIGNY I,
ANDOUIN.
By NORMAN CUNLIFFE, M.A. (Cantab.).
Christopher Welch Lecturer in Economic Zoology , University of Oxford, and formerly Student in Medical Entomology , Quick Laboratory , Cambridge.
(With 3 Text-figures.)
CONTENTS.
Introduction .
Section I. Biology of O. savignyi ........
Experimental records relating to females kept at different temperatures Oviposition ...........
No evidence of parthenogenesis .......
Longevity of female tick ........
Duration and number of nymphal stages at 30° and 37 C.
(а) Experimental data ........
(б) Discussion of data ........
Duration and extent of engorgement ......
The influence of moisture on vitality and ecdysis ....
Section II. Structure of 0. savignyi •.....•
Dimensions of egg ....••••••
Dimensions of unfed stages . . . ...
Changes in external anatomy undergone during development (a) hypo stome, ( b ) leg and spiracle ....•••••
PAGE
17
18 18
20
20
20
21
21
21
22
22
23
23
23
24
INTRODUCTION.
Some observations on the biology and structure of Ornithodorus moubata have been recorded in a recent communication to this journal (Cunlifie, 1921) and in the following pages all references to this species refer to these observations, unless otherwise stated. Prof. Nuttall received living material of 0. savignyi from Dr J. H. Ashworth in 1911 (from Aden, Quick Lab. Cat. No. 1575) and again in 1913 (no data, Cat. No. 2011). These stocks being available for breeding experiments, a few observations on the biology and structure of this species were made in the years 1913-14.
Even at the present time, details of the life-cycle of O. savignyi appear to be unknown. Howlett (1916) and Fletcher (1919) have both stated that this
Parasitology xiv -
18
Ornithodorus savignyi
tick was being studied at Pusa, but their results are apparently as yet un¬ published. Nuttall and Warburton (1908) only note that “there are at least two nymphal stages, if not more.” These facts and also that 0. savignyi is a potential disease carrier justify the publication of these incomplete notes, made under laboratory conditions. This investigation did not yield the same amount of information as was obtained in the case of 0. moubata, as the two were run together, and it was not possible to rear successfully large numbers of individuals of both species. As would be expected, the results of this second investigation are, for the most part, in agreement with those of the first, and where confirmation is complete, discussion of the records is omitted for economy of space, the reader being referred to the paper on 0. moubata.
SECTION I. BIOLOGY OF 0. SAVIGNYI.
It is unnecessary to recapitulate the experimental procedure, which was the same throughout as that adopted for 0. moubata. Similar also was the aim of the investigation, namely, the determination of the number of nymphal stages passed through by this tick before reaching maturity, of the effect of temperature and moisture on the duration of the stages and of the changes taking place in the external anatomy of the ticks at each stage of develop¬ ment. As before, some notes were made on oviposition, copulation, longevity and engorgement. Females, with their progeny, from each stock were reared separately, but their records, being similar, are not shown separately in the following synopses of results.
Experimental records relating to females kept at different temperatures.
Three series of female ticks were maintained at 22°, 30° and 37° C. re¬ spectively, but only the second series bred at all successfully, as shown in Synopsis I.
Females and progeny maintained at 22° C.
A series of six females, which emerged from the last nymphal stage between May and July, 1913, although they fed well and were fecundated on one or more occasions, failed to oviposit until about the 400th day. The larval yield was too small to promise success in rearing and the experiment was discon¬ tinued. Females which emerged about the same time from the same stock, reproduced very successfully at 30° C. However, it is considered that a repetition of this experiment would show that 22° C. is not too low a tempera¬ ture for reproduction by this species.
N. ClJNLIFFE
19
Synopsis I. Females and progeny maintained at 30° C.
Time reckoned in days from date of emergence of female.
Commence¬ ment of Oviposition
Copu- oviposition t - A -
|
No. of |
Date of |
Times of |
c? |
lation |
after |
Period |
No. of |
No. of |
Death |
|
$ |
emergence |
feeding |
added |
observed |
added |
of life |
eggs |
larvae* |
of ? |
|
40 |
20. vii. 13 |
14 |
37 |
37 |
47 |
84-85 |
29 |
nil |
393 |
|
_ |
38 |
_ . |
— |
— |
105 |
4 |
nil |
— |
|
|
_ |
79 |
_ |
— |
— |
135 |
6 |
nil |
— |
|
|
_ |
115 |
_ |
— |
— |
148-150 |
32 |
nil |
— |
|
|
172 |
- |
— |
— |
191-195 |
77 |
10 |
— |
||
|
_ |
277 |
_ |
— |
— |
295-296 |
26 |
17 |
— |
|
|
41 |
26. vi. 13 |
5 |
69 |
70 |
15 |
84-85 |
43 |
27 |
412 |
|
_ |
69 |
_ |
— |
— |
88 |
18 |
11 |
— |
|
|
_ |
103 |
_ |
— |
— |
92-94 |
3 |
nil |
— |
|
|
_ |
_ |
138 |
_ |
— |
— |
123-124 |
44 |
30 |
— |
|
_ |
. |
194 |
_ |
— |
— |
129-130 |
23 |
17 |
— |
|
_ |
_ |
301 |
_ |
— |
— |
145 |
6 |
5 |
— |
|
_ |
_ |
— |
— |
156 |
3 |
3 |
— |
||
|
_ |
_ |
— |
— |
212-219 |
59 |
49 |
— |
||
|
42 |
26. vi. 13 |
5 |
69 |
— |
16 |
85-94 |
128 |
90 |
— |
|
_ |
69 |
_ |
— |
— |
120 |
20 |
15 |
— |
|
|
_ |
_ |
103 |
_ |
— |
— |
125-126 |
25 |
11 |
— |
|
_ |
_ |
139 |
_____ |
— |
— |
155-158 |
80 |
49 |
— |
|
_ |
_ |
196 |
— |
— |
— |
229 |
4 |
nil |
— |
|
_ |
_ |
298 |
— |
— |
— |
— |
— |
— |
— |
|
43 |
23. vi. 13 |
2 |
74 |
74 |
20 |
94 |
8 |
6 |
420 |
|
_ |
72 |
_ |
— |
— |
97 |
10 |
8 |
— |
|
|
_ |
_ |
106 |
_ |
— |
— |
100 |
9 |
9 |
— |
|
___ |
_ |
144 |
_ |
— |
— |
128 |
29 |
25 |
— |
|
___ |
_ |
197 |
_ |
— |
— |
134-136 |
58 |
37 |
— |
|
_ |
_ |
303 |
_ |
— |
— |
165-166 |
45 |
21 |
— |
|
_ |
_____ |
-- |
— |
182 |
10 |
nil |
— |
||
|
_ |
_ , |
_ |
— |
229 |
10 |
nil |
— |
||
|
44 |
20. v. 13 |
17 |
103 |
104 |
17 |
120-124 |
68 |
47 |
249 |
|
_ |
_ |
103 |
_ |
— |
— |
155-157 |
21 |
7 |
— |
|
_ |
_ |
137 |
_ |
— |
— |
165 |
2 |
nil |
— |
|
_ |
_ |
175 |
— |
— |
— |
189 |
9 |
nil |
— |
|
— |
_ |
228 |
— |
— |
— |
— |
— |
— |
— |
|
16 |
25. iii. 13 |
24 |
19 |
20 |
20 |
49-50 |
89 |
77 |
242 |
|
— |
— |
78 |
— |
— |
— |
90 |
25 |
17 |
- — |
|
— |
— |
155 |
— |
— |
— |
— |
— |
— |
— |
|
— |
— |
196 |
— |
— |
— |
— |
— |
— |
■ |
|
— |
— |
234 |
— |
— |
— |
— |
— |
— |
— |
|
17 |
20. iii. 13 |
29 |
37 |
37 |
17 |
54-55 |
82 |
65 |
396 |
|
_ |
55 |
_ |
92 |
— |
67 |
48 |
38 |
— |
|
|
____ |
160 |
_ |
_ |
— |
69-72 |
64 |
46 |
— |
|
|
_ |
. |
201 |
____ |
_ |
— |
75 |
3 |
nil |
— |
|
_____ |
295 |
_ |
_ |
— |
95-96 |
46 |
43 |
— |
|
|
_ |
___ |
_ |
— |
101 |
13 |
13 |
— |
||
|
_ |
_ |
_ |
_ |
— |
179-180 |
91 |
76 |
— |
|
|
____ |
_ |
_ |
_ |
_ |
214 |
16 |
11 |
— |
|
|
_ . |
_ |
_____ |
- |
_ |
— |
251-252 |
39 |
32 |
— |
|
_ |
_____ |
_ |
_ |
_ |
— |
264 |
15 |
7 |
— |
|
18 |
25. iii. 13 |
24 |
24 |
25 |
16 |
40-41 |
55 |
41 |
394 |
|
— |
_ |
50 |
— |
108 |
— |
49-50 |
62 |
57 |
— |
|
_ |
78 |
____ |
_ |
— |
60-61 |
76 |
42 |
— |
|
|
_ |
- |
155 |
_ |
— |
— |
65 |
34 |
12 |
— |
|
_ |
196 |
- |
— |
67 |
20 |
12 |
— |
||
|
_ |
_ |
203 |
__ |
— |
96 |
26 |
12 |
— |
|
|
_ |
_ |
290 |
_ |
_ |
— |
118 |
8 |
nil |
— |
|
_ _ |
_ |
____ |
_ |
_ |
— |
173 |
19 |
14 |
— |
|
— |
— |
— |
— |
— |
— |
207 |
11 |
nil |
— |
|
* The subsequent history of these larvae is summarised in |
Synopsis 11. |
20 Ornithodorus savignyi
Females and progeny maintained at 37° C.
Two separate series of females were maintained at this temperature, the first set being started in November 1912, and the second in July 1913. Out of a total of 11 females only four oviposited, the records being as follows: 9 1, 92 eggs after 19-28 days (from date of emergence) followed by 14 eggs after 69 days; 9 2, 100 eggs after 16-17 days; 9 3, 8 eggs after 46 days followed by 56 eggs after 78-86 days; 9 4, 71 eggs after 69-74 days. These females were fed for another six months without further oviposition taking place and they were then discarded. Larvae emerged from 65 per cent, of the eggs, but reference to Synopsis II, in which their subsequent history is summarised, will show that none of them succeeded in passing the third nymphal stage at this temperature.
Oviposition.
When first deposited the egg is yellow in colour; it rapidly turns brown, more slowly becomes pitchy-brown and finally, in the course of three or four days, becomes jet-black. The agglutinative coating soon loses its efficacy in this species. The data, relating to oviposition, are summarised in the following table, minimum, maximum and mean results being shown:
Ticks maintained at 30° C.
, - A - >
Minimum Maximum Mean
No. of days before oviposition occurred after $ was allowed 15 20 17
access to 9 (aberrant case 47)
No. of eggs deposited after each feed (when oviposition occurred) 4 1 74 62
No. of days between dates of feeding and oviposition commenc- 5 35 18
ing or recommencing
No. of days over which oviposition extended after each feed 1 50 9
No. of eggs deposited bya9 100 417 219
Percentage of fertile eggs 62 %
The mean results are very closely comparable with those obtained in the case of O. moubata (at 30° C.), except that the number of eggs oviposited by the female is less by 40 per cent.
No evidence of parthenogenesis.
Parthenogenetic reproduction did not occur during the course of these experiments. Virgin females, which emerged in October 1913, were fed re¬ peatedly, and maintained at 30° C. between meals, but they failed to oviposit during the following eight months. On the other hand, it has been shown that gorged females oviposit regularly about 17 days after the males are allowed access to them (at 30° C.).
Longevity of the female tick.
When performing their normal functions and maintained at 30° C., seven females lived for minimum, maximum and mean periods of 292, 420 and 358 days respectively. Under similar conditions at 22° C., the female had an average life of 775 days (three individuals).
N. CUNLIFFE
21
Duration and number of the nymphal stages at .>0 and 37 C.
(a) Experimental Data.
Some of the progeny of each female were reared in separate batches to the adult stage, to establish the duration and number of the nymphal stages. The records are summarised in Synopsis II, minimum, maximum and mean periods, together with the numbers of individuals observed, being given. As in 0. moubata, the larva1 passes into the first nymphal stage without previous engorgement; from the first nymphal stage onwards, the periods required for ecdysis are reckoned from the date of the previous meal and not that of the previous moult.
Synopsis II. Duration of stages in days.
Stage
Esg to larva Egg to 1st ® 1st to 2nd ® 2nd to 3rd ®
3rd to 4th \ ® or adult J
4th to 5th \ ® or adult f
5th to 6th \ ® or adult f
6th to 7th \ ® or adult J
Ticks kept at 30° C. Ticks kept at 37° C.
A _ _ A -
|
, „ |
No. of ' |
r |
No. of |
||||
|
individuals |
individuals |
||||||
|
Min. |
Max. |
Mean |
observed |
Min. |
Max. |
Mean |
observed |
|
7 |
28 |
12 |
709 |
7 |
11 |
8 |
231 |
|
13 |
32 |
19 |
630 |
11 |
20 |
15 |
198 |
|
8 |
30 |
11 |
348 |
5 |
11 |
8 |
121 |
|
9 |
18 |
12 |
184 |
7 |
11 |
8 |
8* |
|
® 7 |
49 |
14 |
116 |
||||
|
3 21 |
33 |
29 |
3t |
||||
|
® 11 |
25 |
15 |
63 |
||||
|
3 13 |
22 |
17 |
29 |
||||
|
$ 16 |
20 |
17 |
6 |
||||
|
® 15 |
26 |
17 |
6 |
||||
|
3 14 |
23 |
18 |
14 |
||||
|
$ 15 |
19 |
17 |
12 |
||||
|
® |
No record |
||||||
|
$ 15 |
— |
15 |
2 |
* These eight individuals were the only survivors out of 121 second stage nymphs and as third stage nymphs they refused to feed, t Probably abnormal.
(6) Discussion of Data.
As with 0. moubata , males appeared after four to six and females after five to seven ecdyses, 63 per cent, of the males appearing after the fifth ecdysis and 60 per cent, of the females after the sixth ecdysis. As, however, only 66 individuals were reared to maturity, these percentages may require correc¬ tion. The mean minimum periods required for the metamorphosis of an individual from egg to adult, at 30 C., under laboratory conditions, are
(1) for males, 60 ( corrected ), 73 or 89 days according to whether the indi¬ vidual undergoes four, five or six moults;
(2) for females, 73, 88 or 103 days, the number of moults varying from
five to seven.
These periods are longer by an average of 27 days than in the case of
1 The larva normally frees itself from the egg shell, but is otherwise inactive. In some cases, however, the species approaches O. moubata , inasmuch as the larva may undergo ecdysis without throwing off the shell. The feeble development of the hypostome, legs and claws and also the absence of eyes is doubtless correlated with the passiveness of this stage.
22
Ornithodorus savignyi
0. moubata, reared under similar conditions. An increase in temperature of 7° C. (from 30° C.) only reduced the period required for the production of third stage nymphs by 26 per cent.
Duration and extent of engorgement .
An experiment, similar to that carried out with 0. moubata, was conducted with this species, ten nymphs of different stages being kept under observation until they reached maturity. Their weights before and after engorgement and the times of attachment to the host were accurately recorded. Again there was no regularity in the time of attachment, observation on 29 individuals of all stages giving minimum, maximum and mean times of 10, 74 and 26 minutes respectively, at room temperature, about 16° C. The figures are in agreement with those of Drake-Brockman (1915). The mean time recorded for 0. moubata was 48 minutes, thus 0. savignyi is apparently a more rapid feeder. Of the ten nymphs under observation, only four reached maturity, one being male and three female. In the female series, the mean increase in weight in grams was (a) before the final moult, 0-0647, and (b) after the moult 0-2202; the corresponding figures for the solitary male were (a) 0-0528, and (b) 0-0246. None of these figures were reached by 0. moubata.
The influence of moisture on vitality and ecdysis.
A similar experiment to that carried out with 0. moubata proved that, for this species also, excess of moisture is injurious to the individual. A batch of 33 first stage nymphs from the same female was divided into three lots, which between meals were maintained at 30° C. in saturated, moist and dry atmospheres respectively. A summary of the experimental records1 is given in the following table, in which are shown the minimum and maximum number of days required for ecdysis, with the mean in brackets and the number of individuals observed in square brackets :
|
% of ticks |
||||||
|
which |
||||||
|
Atmosphere |
2nd ® |
3rd ® |
4th ® |
5th ® |
6th ® |
matured |
|
Saturated |
9—13 (11) |
11—14(13) |
23—48 (40) |
21 |
— |
9% |
|
[11] |
[11] |
[7]1<J |
[1] |
|||
|
Moist |
7—8 (8) |
9—32 (12) |
10—18 (12) |
11—17(14) |
— |
27% |
|
[11] |
[11] |
[11] 2 ^ |
[4] 1 3 |
|||
|
Dry |
7—9 (7) |
9—13 (10) |
11—16(12) |
10—14(12) |
13—17 (15) |
45% |
|
[11] |
[8] |
[8] 1 c? |
Of r-H *6 <N O |
[2] 19 |
Excess of moisture was therefore a decidedly unfavourable factor, as under this condition only 9 per cent, of the ticks matured, whereas in the dry atmosphere 45 per cent, matured. The ecdysis period was not markedly affected by the presence of moisture ( vide Synopsis II), until after the third nymphal stage was attained, when the lack of vitality was indicated by a lengthening of this period.
1 The 1st nymphs were fed on 17. i. 13 and the 5th nymphs were fed or offered a meal on 13. vi. 13.
N. ClJNLIFFE
23
SECTION II. STRUCTURE OF 0. SAVIGNYI.
Observations on the dimensions of the egg and unfed specimens of the different stages and the changes in form undergone during development by the hypostome, the fourth tarsus and the spiracle are included in this section.
Dimensions of the egg.
A total of 179 eggs were measured immediately after deposition by four different females, the minimum, maximim and mean measurements of length x breadth being 1-0 x 0-9, 1*8 x 1-4 and 1*4 x M mm. respectively. The minimum and mean measurements of the eggs of this species equal the mean and maximum measurements of those of 0. moubata.
Dimensions of unfed stages.
As no records of the dimensions of the different stages appear in the litera¬ ture on this species, measurements of material bred under control from one stock are summarised in Synopsis III. The number of individuals preserved was, in several cases, too small to give very accurate results, although as far as possible extreme examples were chosen for preservation.
Synopsis III. Dimensions of stages.
Measurements in mm. to nearest tenth, from unfed specimens preserved shortly
|
after emergence in 70 % spirit. |
No. of |
|||
|
Min. |
Max. |
Mean |
individuals |
|
|
Stage |
Length x Breadth Length x Breadth Length x Breadth |
observed |
||
|
Larva |
1-2 x 10 |
1-5 x 1-2 |
1-4 x 1-1 |
52 |
|
1st ® |
1-3x10 |
1-8 x 1-4 |
1-5 x 1-2 |
107 |
|
2nd ® |
1-5x11 |
2-6 x 1-9 |
2-1 x 1-5 |
119 |
|
3rd ® |
2-2 x 1-5 |
3-6 x 2-6 |
30x2-1 |
16 |
|
4th ® |
3-1 x 2-3 |
5-6 x 3-9 |
4-4 x 3-1 |
14 |
|
o from 3rd ® |
5-2 x 3-8* |
6-0 x 4-3 |
5-7 x 4-0 |
3 |
|
5th ® |
3-9 x 2-8 |
6-5 x 4-6 |
5-6 x 3-9 |
7 |
|
from 4th ® |
5-2 x 3-5 |
6-7 x 4-6 |
6-0 x 3-9 |
25 |
|
$ from 4th ® |
7-0 x 4-9* |
7-8 x 5-6 |
7-4 x 5-2 |
4 |
|
6th ® |
5-0 x 3-4 |
8-5 x 5-7 |
6-7 x 4-8 |
6 |
|
<$ from 5th ® |
5-8 x 3-8 |
7-4 x 5-6 |
6-5 x 4-7 |
15 |
|
$ from 5th ® |
6-7 x 4-7 |
9-2 x 6-7 |
8-1 x 5-7 |
14 |
|
7th ® |
— |
— |
— |
|
|
$ from 6th ® |
6-4 x 4-4 |
9-3 x 6-7 |
7-8 x 5-5 |
2 |
|
* Probably not true minima. |
||||
|
As in 0. moubata , there is |
considerable variation in the size |
of individuals |
of any one stage, maximum measurements in one stage exceed minimum measurements in the succeeding stage, and males of any one stage are on 1 \ slightly larger, whereas females are distinctly larger, than nymphs of the equivalent stage. In 0. moubata, the greatest growth took place in the second nymphal stage but in this species it would appear to be spread equally over the second and third nymphal stages, the percentage increases in length being 7-3 (larva to 1st O), 40-0, 42*8, 46-7, 27-3, 19-7 (5th to 6th O), while the
24
Ornithodorus savignyi
corresponding increases in breadth are 9-1, 25-0, 40-0, 47-7, 25-8 and 23-1 per cent, respectively. The percentage increases in length and breadth from nymphal to adult stages correspond closely in both species, e.g. the percentage increases in length x breadth in 0. moubata, from fourth nymph to male, are 32 x 28— -while in 0. savignyi, the increases are 36 per cent, x 26 per cent.
Fig. 1. Ornithodorus savignyi, hypostomes in ventral aspect: ( b ) to (g) of 1st to 6th stage nymphs;
(*) and U) of females from 4th and 6th stage nymphs; (k) and (l) of males from 3rd and 5th stage nymphs. (N. C. del.)
Changes in external anatomy undergone during development.
(a) The Hypostome. Fig. 1.
The changes in the dentition of the hypostome during the development from first nymph to adult are indicated in Fig. 1, (6) to ( g ) representing the nymphal organs arranged in order of development, (i) and ( j) the hypostomes of females emerging from fourth and sixth nymphs, and (k) and (l) of males from third and fifth nymphs respectively.
On comparison with the hypostomes of 0. moubata , it will be observed that in this species also several proximal rows and the distal row of teeth are equally poorly developed and their arrangement difficult to determine, that in the first and second nymphal stages the number and arrangement of the teeth are very similar, but that in the later stages the number of teeth is considerably less, owing to the presence of smaller numbers of rows and files. The changes taking place during development consist of an increasing com-
25
N. OUNLIFFE
plexity of dentition, correlated with an increasing size of hypostome and they are of much the same order in both species.
In many cases the teeth are not arranged symmetrically on the two sides of the hypostome, the rows on one side appearing to alternate with the rows on the other side.
(b) The Legs and Spiracle. Figs. 2 and 3.
Outlines of the terminal portions of the fourth legs and of the spiracular plates are shown in Figs. 2 and 3 respectively, for the following stages, namely, first to sixth nymph (a) to ( / ) ; males from third and fifth nymphs (h) and (i) ; and females from fourth and sixth nymphs ( j) and (k).
1 m m. a to c
Fig. 2. Ornithodorus savignyi, tarsi of fourth legs, in lateral aspect: (a) to ( / ) of 1st to 6th stage nymphs; ( h ) and ( i ) of males from 3rd and 5th stage nymphs; (j) and (k) of females from 4th and 6th stage nymphs. (N. C. del.)
The degree of development of these structures at each stage is similar in the two species, 0. savignyi and 0. moubata and needs no further discussion.
SUMMARY OF RESULTS.
1. The biology, as studied under laboratory conditions, of 0. savignyi is very similar to that of 0. moubata.
2. Females may oviposit over 400 eggs, of which at least 60 per cent, may be fertile; parthenogenesis does not occur.
3. An increase in temperature of 8r C. (from 22° C.) decreases the longevity
26 Ornithodorus savignyi
of the female from 775 to 358 days, i.e. by 45 per cent.; an increase of 7° C. (from 30° C.) reduces the period required for the production of third stage nymphs by 2b per cent. At 30° C., the mean minimum periods required for metamorphosis are 60 days for males and 73 days for females (cf. 0. moubata ,
06 days for males and 45 days for females). Reproduction was inhibited at 37° C.
4. Moisture is an unfavourable factor, decreasing the vitality of the individual at each stage of growth.
o. Changes in external anatomy undergone during development are similar to those already described for 0. moubata.
Fig. 3. Ornithodorus savignyi, outlines of cribriform plates of spiracles: (a) to (/) of 1st to 6th stage nymphs; ( h ) and ( i ) of males from 3rd and 5th stage nymphs; (j) and ( k ) of females from 4th and 6th stage nymphs. (N. C. del.)
REFERENCES.
Cunliffe, N. (1921). Some observations on the biology and structure of Ornithodorus moubata, Murray. Parasitology, xm. 327-347.
Drake-Brockman, R. E. (1915). Some notes on the Bionomics of Ornithodorus savignyi in British Somaliland. Bull. Entom. Research, London, vi. 195-196.
Fletcher, T. B. (1919). Report of Agric. Research Inst. Pusa, 1915-16, 78-84.
Howlett, F. M. (1916). Ibid. 1914-15, 74-77.
Nuttall, G. H. F. and Warburton, C. (1908). Ticks, a monograph of the Ixodoidea, Pt. i. Cambridge.
27
A PRELIMINARY NOTE ON PARASITES INFESTING DOMESTICATED SILVER BLACK FOXES IN CANADA1.
By J. A. ALLEN, B.V.Sc.,
Animal Pathologist , in charge of Fox Research Station, Charlottetown, P.E.I.
AND A. B. WICKWARE, V.S.,
Animal Pathologist , Biological Laboratory, Ottawa, Ont.
( From the Health of Animals Branch, Dept, of Agriculture, Canada.)
The growing importance of the silver black fox industry in Canada, and latterly in other countries, has been so marked, that scientific workers have turned their attention to a study of the maladies affecting this species of fox.
A survey has been made by one of the writers (J. A. A.) of the benign and contagious forms of disease to which domesticated foxe.. ate subject, and papers of a popular and scientific nature have already been published in
various journals.
In the course of recent investigations dealing with a distemper-like foim of disease causing a high mortality amongst foxes, a determination of the species of internal and external parasites was undertaken.
As pointed out by Riley in his recent paper2, "An Annotated List of the Animal Parasites of Foxes,” very little literature is available on this subject, and it is for the purpose of adding a small contribution that the parasites
found by us to date are herewith recorded.
We take especial pleasure in acknowledging the courtesy shown b} Dis B. H. Ransom, Chief Zoologist, Bureau of Animal Industry, Washington, D.C. ; Maurice C. Hall, Senior Zoologist, Washington, D.C. ; and F. C. Bishopp, Entomologist, Bureau of Animal Industry, Dallas, Texas, in identifying the listed parasites, and to the various fox ranchers who have provided us with animals for autopsy.
PROTOZOA.
Coccidium bigeminum — Isospora bigemina (Stiles, 1891).
Small and large intestine.
ACARINA.
Sarcoptes scabiei vulpis ( Fiirs ten berg, 1861) Railliet.
Body of host.
Otodecies cynotis (Hering, 1838). Ears of host — external meatus.
1 Published with the approval of Dr F. Torrance, Veterinary Director General.
2 Riley, W. A. (in. 1921), Parasitology , xiii. 86-96.
28 Parasites infesting Domesticated Black Foxes in Canada
SIPHONAPTERA.
Ctenocepholus canis (Curtis, 1826). Body of host.
NEMATODA.
Eucoleus aerophilum (Creplin, 1839). Trachea and large bronchi. Eggs found constantly
in trachea, oesophagus and faeces.
Belascaris cati (Schrank, 1788)
Bela.s'caris marginata (Rudolphi, 1802) Small intestine.
Toxascaris limbata (Railliet and Henry, 1911)
|
Uncinaria polaris (Looss, 1911). |
Sometimes B. marginata penetrates the bile duct and develops in the liver. Immature ascarids found in fox pups less than one week old. Quite common to find intestinal canal from stomach to large colon completely packed with ascarids in young foxes only three to four weeks old. One seldom finds more than one or two ascarids in adult foxes. Small intestine. |
|
Ascocotyle longa (Ransom, 1920). Echinochasmus sp. |
TREMATODA. Small intestine. Small intestine. |
29
' ON THE DIPTEROUS GENERA PASSE MOM Y I A AND ORNITHOMUSCA, WITH NOTES AND BIBLIOGRAPHY ON THE NON-PUPIPAROUS MYIODARIA PARASITIC
ON BIRDS.
By Professor M. BEZZI, Turin, Italy.
CONTENTS.
I. Passeromyia and Ornithomusca
II. Protocalliphora
III. Philornis ....
IV. Camus ....
V. Chortophila and N eottiophilum
VI. Conclusions
VII. Bibliography
PAGE
29
32
3(3
39
40
41 41
I. PASSEROMYIA AND ORNITHOMUSCA.
In 1851 Macquart described two new species of the Dipterous genus Cyrtoneura from Tasmania, under the names of Cyrt. longicornis and Cyrt. analis, both species being very different from all others of the same genus in having hairy eyes and the first species also owing to its very long third antennal joint.
They were never recorded by subsequent writers until 1916 when Tylei Townsend describing his new Australian genus Ornithomusca expressed doubts as to the relationship of the above mentioned Macquart’s species to his own genus (p. 145). In 1919 the late Prof. Stein in his Catalogue of the Anthomyidae 'of the World (p. Ill) recorded these two species, showing however that they do not belong to the genus Muscina (which now comprises several species of the old genus Cyrtoneura) and that they cannot be placed in any known genus of Anthomyidae.
I have recently received from Mr E. W. Ferguson two specimens of a fly, one bred at Sydney, N.S.W., from a larva found in a nestling of the New Holland Honeyeater, the other caught in the open. Both these specimens I have recognised as belonging to the species Cyrtoneura longicornis Macquart. On the other hand, the type specimen of Ornithomusca victoria Tyler Town¬ send has been also found in nests of Pardalotus sp. and according to the description differ from longicornis only in the colour of the dust of the head, palpi, calypters, the base of the wings and the hairs of the body so that it is
are
30 Parasitic Diptera
very probable that the names Cyrt. longicornis and Orn. victoria synonymous1.
The habit of the above-named Australian fly Cyrt. longicornis living upon the bird nestlings recalls that of the African species belonging to the recentlv established genus Passeromyia, and the comparative study of the specimens of the Australian and African forms shows that they are very closely allied. The wide distribution of the African species indicates moreover the im¬ portance of a careful study of specimens of this species from the Oriental region. In fact, to this group belongs the species Muscina longicornis from Java, previously described by Stein (1909, p. 221). He noticed in this species the elongated third antennal joint, but has overlooked the hairiness of the eyes. In 1915 Dr Villeneuve described under the name Muscina heterochaeta a new African fly showing aberrant characters. The larvae of this fly have been previously described by Rodhain (1914, p. 214). The same year (1915) Rodhain and Villeneuve established for this African species the new genus Passeromyia and in 1916 appeared a very important paper by Rodhain and Bequaert on the structure and life-history of this fly ( Passeromyia heterochaeta ).
Professor Stein in a recent letter (1920) informed me that the specimens of Passeromyia heterochaeta sent to him by Dr Villeneuve, are identical with his paratypes of Muscina longicornis. I was able myself to compare the paratype of longicornis of my collection with Australian specimens and found that they were strictly congeneric; it is even a matter of some difficulty to distinguish the Australian species from the African. It can be concluded now that the genus Ornithomusca Tyler Townsend 1916 is synonymous with the genus Passeromyia Rodhain and Villeneuve 1915, and that the type species heterochaeta Villeneuve 1915 is synonymous with longicornis Stein 1909. As the last name is already preoccupied by Macquart’s species, A llleneuve’s name heterochaeta must be retained.
The two species of the genus Passeromyia can be distinguished as follows :
1 (2). The dust of the entire body, chiefly that of the abdomen of the male, is of a distinct bluish tint; third antennal joint about nine to ten times longer than the second joint, and extending almost to the mouth border; bristles of all femora less numerous, shorter and thinner; hind tibiae shortly ciliated ; fourth longitudinal vein with the twro portions on either side of the bend, of nearly the same length; average size 7 mm. longicornis Macquart.
2(1). Dust of the body grey or slightly bluish on sides of the abdomen of the male; third antennal joint seven or eight times longer than the second
1 Another question arises in connection with the species Cyrtoneura analis Macquart. This species was described from a type specimen with the second and third joint of antennae missing, and it is therefore not certain that this species belongs to the genus Ornithomusca. The description of the abdomen recalls the fly Synthesiomyia nudiseta which was originally described as Cyrtoneura and which is present even in Australia. The hairy eyes and the black first antennal joint of analis do not agree with nudiseta. Bigot in 1877 (p. 250) placed the species analis in the genus Graphomyia but removed it in 1887 (p. 584) to the genus Dasyphora.
M. Bezzi
31
and not reaching the border of the mouth; bristles of the underside of the femora longer, stronger and more numerous than in previous species; hind tibiae with longer ciliation; the portion of the fourth vein in front of the
bend is longer than the hinder portion; average size 8-9 mm.
helerochaeta Villeneuve.
Passeromyia.
Rodhain and Villeneuve, 1915, p. 592; Rodhain and Bequaert, 1916, p. 219. Ornithomusca, Tyler Townsend, 1916, p. 45.
1. Passeromyia longicornis (Macquart).
Cyrtoneura longicornis , Macquart, 1851, p. 228 (255), pi. 23, fig. 8; Stein,
1919, p. 111.
Ornithomusca victoria , Tyler Townsend, 1916, p. 45.
Geographical distribution. Australia; Victoria (in nest of Pardalotus sp.), New South Wales, Sydney; Tasmania.
2. Passeromyia heterochaeta (Villeneuve).
Muscina longicornis , Stein, 1909, p. 221 (non Macquart, 1851); 1919, a, p. Ill, and b, p. 68.
Larva (without a name), Rodhain, 1914, p. 214, fig. 1.
Muscina heterochaeta , Villeneuve, 1915, p. 225, fig. 1.
Passeromyia heterochaeta , Rodhain and Villeneuve, 1915, p. 593; Rodhain and Bequaert, 1916, p. 250, figs. 1-6, pi. xix, fig. 2; Roubaud and Van Saceghem, 1916, p. 765; Roubaud, 1918, p. 428; Stein, 1919, p. 86; Patton,
1920, p. 30, pi.; Engel, 1920, p. 258.
Geographical distribution. Africa: Nyasaland, Port Herald; N.W . Rhodesia, Chilanga; Katanga, Elisabethville ; British East Africa, Mombasa; Belgian Congo, Ouelle, Leopoldsville, Boma, etc. I have in my collection specimens from Usambara, Nguelo.
Asia: India, Madras; Java, Batavia; China, Kamsi. I have received numerous specimens from S. China, Canton, by Prof. C. W. Howard.
Ethology. The species has been found in the nests of various species of birds of the genera Passer , Hirundo, Cinnyris, Spermestes , Sitagra, Ploceus , etc.
According to Rodhain and Bequaert the larvae of Passeromyia have the same habits of intermittent haematophagy as observed in the floor maggot and other African Calliphorines belonging to the genera Auchmeromyia and Chaeromyia which live upon the blood of bare-skinned mammals.
It is interesting to note that Passeromyia, as a bird-parasite, is widely spread over the whole tropical region of the Old World only. There are however true Calliphorine flies the larvae of which live upon birds and have the same habits of intermittent haematophagy as is the case in Passeromyia. These flies belong to the well-known genus Protocalliphora which we shall examine presently.
32
Parasitic Diptera
IT. P ROT OCA LLI PHORA .
There is a marked tendency both in Europe and in N. America to place the three species of haematophagous Calliphorine flies ( caerulea R. D., azurea Falk, and metallica Tyler Townsend) in the genus Phormia.
The genus Pliormia was established by Robineau-Desvoidy in 1849 and accepted by Coquillet in 1910 (p. 589) for the species Musca regina Meigen 1826 which differs markedly from the above mentioned calliphorine species.
The calliphorine bird parasites cannot therefore be placed with the species regina Meigen in the same genus Phormia. As to the new generic name Euphormia proposed by Tyler Townsend in 1919 (p. 542) for the genotype regina it becomes quite superfluous, and was introduced only through mis¬ interpretation of Villeneuve’s statements (see 1911, p. 84).
The right generic name for the bird parasites is that of Protocalliphora Hough 1899, as it has already been used by me in the third volume of the Katalog der palaearldischen Dipteren, pp. 544-545. The true genus Phormia thus remains a monotype, with the species regina, while in the genus Protophormia Tyler Townsend 1908 must be placed the species terraenovae Robineau-Desvoidy 1830 (= groenlandica Zetterstedt 1838) 1 and Boganidae Erichson 1851.
There has been some discussion about the validity of the described species of Protocalliphora. Hendel in 1901 (p. 29) distinguished in this genus five different species, while Hough, Villeneuve and Bequaert accepted only two species which were subsequently united by Roubaud (1918) into a single species. However, Kramer (1911, p. 43) has already shown that the species can be distinguished by the structure of the male genitalia; and recently Engler (1920) clearly established the existence of the two distinct European species by the study of the structure of the larval, pupal and adult stages. In addition to the two European species Tyler Townsend (1919) has added a third North American species: metallica. Of the two European species caerulea is the commoner. It shows however in addition to sexual dichroism, some variation in the breadth of the frons in the male and in the colour of the calypters. Being often the unique species of Protocalliphora present in col¬ lections, it was erroneously subdivided into two or more species under the names of sordida, azurea, hraueri, etc. ; the other species azurea is more uniform but very often wanting in collections.
The three known species of Protocalliphora may be distinguished as follows :
1 (4). Parafacialia without golden spot above.
2 (3). Parafacialia (viewed obliquely from above) smooth and uniformly covered with whitish pollen or dust. The two sexes differ in colour: the male has the thorax and abdomen of metallic dark blue coloration with scarce
1 In the Catalogue of Palaearctic Diptera I have erroneously reported this species under the name caerulea R. D., which belongs to Protocalliphora.
M. Bezzi
33
pollen; the female showing golden-greenish thorax and metallic green abdomen with bluish reflects and a thin whitish pollen covering all the segments except the last one. Frons of the male more or less narrow, not broader than the ocellar triangle; male genitalia in lateral view show acute paralobes and straight mesolobe. Females with the middle frontal stripe three times bioadei than one of the parafrontalia; average size 9-11 mm. . caerulea R.-D.
’ 2 (1). Parafacialia with whitish pollen distributed in irregular rows with
black interspaces. In both sexes the thorax is of a dark blue metallic colour with scarce pollen and the abdomen of a glittering bluish-green colour de\ oid of pollen and with a distinct dark longitudinal median stripe upon the second and third segments. Frons in the male distinctly broader, being about twice as broad as the ocellar triangle ; the male genitalia with the paralobes broadly obtuse and with curved mesolobe. Parafrontalia of the female broader, the middle frontal stripe being twice as broad as one of them; average size 11-14 mm. .....••••• azurea hall.
4 (1). Parafacialia each with a conspicuous golden spot at the upper end. Thorax metallic, greenish-black to bright green colour with grey pollen and three nearly equal longitudinal stripes of the ground colour. Abdomen metallic dark bluish-green to bright cupreous or golden-green colour with scarce silvery pollen ; anal segment varies always from cupreous to golden- green colour. Calypters white to buff yellow. Size 7-8*5 mm. Only female known metallica T. T.
The synonymy of the three above described species of Protocalliphora is as follows :
Protocalliphora.
Hough, f899, a, p. 66 and b, p. 289.
Avihospita, Hendel, 1901, p. 29 and 68; b, p. 2101; Aldrich, 1901, p. 68.
1. Protocalliphora caerulea (Robineau-Desvoidy).
Phormia caerulea , Robineau-Desvoidy, 1830, p. 466; 1863, p. 846.
Lucilia caerulea , Macquart, 1835, p. 256; Meigen, 1838, p. 295; Schiner, 1862, p. 591; de Meijere, 1902, p. 682, fig. 54.
Protocalliphora caerulea , Falcoz, 1921, p. 139.
Musca azurea, Meigen (nec Fallen!), p. 63; ? Rossi, 1848, p. 59, note. Calliphora azurea , Schiner, 1862, p. 585; Nowicki, 1867, p. 44; Brauer, 1883, p. 74; Yerrall, 1886, p. 231; Strobl, 1893, p. 104; Brauer and Bergen- stamm, 1894, pp. 546 and 568; Pandelle, 1896, p. 214; Grimshaw, 1901, p. 27.
Protocalliphora azurea , Hough, 1899, p. 289, fig. 11; Aldrich, 1905, p. 524; Bezzi, 1907, p. 544, p.p. ; Villeneuve, 1910, p. 313; Nielsen, 1911, p. 205. Avihospita azurea, Hendel, 1901, p. 29.
1 The Catalogue of Palaeai die Diptera, p. 444, contains also the genus Philo inis Meineit as a synonym of Protocalliphora. The late Dr Nielsen, 1911, p. 207, pointed out this error, which is not mine (only pp. 1-597 of vol. hi being my work) but that of Brauer, 1894, p. 568.
Parasitology xiv '*
84
Parasitic Diptera
Phormia azurea, Villeneuve, 1911, p. 84 and 1913, p. 132; Surcouf and Gonzalez-Rincones, 1912, p. 135; Rodhain and Bequaert, 1916, p. 245; Rodhain and Villeneuve, 1915, p. 593; Roubaud, 1918, p. 420, pi. v; Plath, 1919, a, p. 30, b, p. 191, c, p. 373; Patton, 1920, p. 30.
Musca sordida, Zetterstedt, 1838, col. 657, 1845, p. 1332, and 1859, p. 6185; Bonsdorff, 1866, p. 150.
Pollenia sordida, Rondani, 1862, p. 198; E. Corti, 1897, p. 140.
Avihospita sordida, Hendel, 1901, p. 29.
Protocalliphora sordida, Villeneuve, 1910, p. 313; Kramer, 1911, p. 43, pi. iii.
Phormia sordida, Villeneuve, 1911, p. 84 and 1913, p. 132; Roubaud, 1914, p. 27, and 1915, a, p. 77, b, p. 94, fig.; Rodhain and Villeneuve, 1915, p. 593; Rodhain and Bequaert, 1916, p. 244; Roubaud, 1917, p. 434; Ville¬ neuve, 1918, p. 158; Engel, 1920, p. 256, fig. 8.
Lucilia dispar, Leon Dufour, 1845, p. 205, pi. ii; Zetterstedt, 1849, p. 3269, ohs.
Phormia dispar, Robineau-Desvoidy, 1849, p. iv.
Calliphora nidicola, v. Heyden in Nowicki, 1867, p. 44, note.
Phormia nigripalpis, Robineau-Desvoidy, 1863, p. 846.
Phormia corusca, Robineau-Desvoidy, 1863, p. 849.
Avihospita braueri, Hendel, 1901, p. 29.
Geographical distribution. The species has been found throughout Europe, from Lapland to Sicily. Its distribution in North America is not yet thoroughly known, but from Plath’s observations it seems that the species is prevalent in the west of the United States. The species was also recorded from Hawaii; if this is not an importation by man, the fly must have a very wide distribution.
Ethology. The habits of the larvae were first described by Leon Dufour and more recently Dr Roubaud has established their habit of intermittent haematophagy and that they cannot live as subcutaneous parasites1.
The species has been observed in nests of Corvus, Passer, Hirundo, Cotyle, Pams, Ruticilla, etc. in Europe, and of Pipilo, Hylocichla, Merula, Ampelis, Dendroicha, Petrochelidon, Passer, Carpodacus, Zonotrichia, Melospiza, Astra- galinus, etc., in North America. All these birds belong to the Passeres.
2. Protocalliphora azurea (Fallen).
Musca azurea, Fallen, 1816, p. 245 and 1821, p. 46; Zetterstedt, 1838, col. 657 and 1845, p. 1334; Bonsdorff, 1866, p. 150.
Pollenia azurea, Rondani, 1862, p. 197.
Protocalliphora azurea, Bezzi, 1907, p. 544, p.p.; Villeneuve, 1910, p. 313, p.p.; Kramer, 1911, p. 43, fig.
Phormia azurea, Engel, 1920, p. 257, fig. 9.
1 According to Roubaud the puparia of this fly are sometimes parasitised by a Hymenopteron Nasonia brevicornis Ashm.
M. Bezzi
35
Musca chrysorrhoea , ? Meigen, 1826, p. 60.
Calliphora chrysorrhoea , Macquart, 1835, p. 263; Schiner, 1862, p. 585; Brauer, 1883, p. 74; Strobl, 1894, p. 70; Brauer and Bergenstamm, 1894, ). 546.
Protocalliphora chrysorrhoea , Hough, 1899, p. 289; Aldrich, 1905, p. 524.
Avihospita chrysorrhoea , Hendel, 1901, p. 29.
, Avihospita n. sp., Hendel, 1901, p. 30.
Phormia chrysorrhoea , Tyler Townsend, in Plath, 1919, pp. 374 and 380.
? Calliphora splendida , Macquart, 1845, p. 324.
Phormia caerulea , Kramer, 1917, p. 27.
Geographical distribution. This fly has apparently the same distribution in Europe as the other species, but is much more rare and there are only a few orecise records, those of all the earlier writers being in my opinion very ioubtful. It has been assumed here that the descriptions in which the sexual dichroism is not clearly indicated, apply to the true azurea , because Fallen’s original description refers to a species in which the males and the remales are equally coloured. But even in this last case a confusion with Vrotophormia terraenovae , R.-D., is always possible. The species is also recorded :rom North America, its distribution there being not yet determined.
Ethology. The larval stages were described by Engel in 1920, and compared with those of caerulea. The habits of these larvae seem to be the same. Engel has obtained the species from subterraneous nests of Riparia riparia, and this is the unique precise record, which indicates a very peculiar habitat.
3. Protocalliphora metallica (Tyler Townsend).
Phormia metallica, Tyler Townsend, 1919, p. 379; Plath, 1919, p. 376.
? Protocalliphora azurea, Coutant, 1915, p. 145.
Geographical distribution. This species as far as it is known is exclusively a North American one; its range extends throughout the United States chiefly in the East, being very rare in the West. It is possible that the above mentioned splendida, Macquart, from Texas, may be synonymous with the present species.
Ethology. Plath found the larvae in nests of Merula migratoria. The larva from nests of Corvus americanus fully described by Coutant, may possibly belong to the present species; at any rate this larva cannot be that of azurea or of caerulea owing to the very important differences pointed out by Roubaud, 1918, pp. 423-424.
* * * *
The larvae of the above named species of Passeromyia and Protocalliphora live in bird nests, and attracted by positive thermotropism to the naked bodies of the young nestlings, they can suck their blood. But this type of intermittent haematophagy makes it impossible for them, as shown by Dr Rou¬ baud, to become permanent parasites. There are however several records of fly-larvae found in subcutaneous tumours on young birds and attributed
3—2
36
Parasitic Diptera
erroneously to Prolocalliphora. These records are enumerated here separately, as they do not belong to the above named flies.
In Europe they are as follows:
Schneider, 1866, p. 89, and Kirsch, 1867, p. 245, pi. iii, figs. 3-4, on Passer domesticus, attributed to Prot. caerulea.
Portschinsky, 1887, p. 17, pi. i, figs. 9-10, on Anthus pratensis, attributed to Prot. azurea ( Pollenia ).
Meinert, 1889, p. 315, on Emberiza miliaria , attributed to Prot. caerulea.
Brauer and Bergenstamm, 1894, p. 546, on Passer domesticus , attributed to Prot. caerulea , and on Riparia riparia, attributed to Prot. azurea.
Pavay-Vajna, 1909, p. 288, on Motacilla alba , attributed to Melinda cognata ( Onesia ).
Heinroth, 1916, p. 158, on Motacilla , attributed to Prot. caerulea.
And in North America:
Henshaw, 1908, p. 87, on Sialis sialis, attributed to Prot. azurea.
Miller, 1909, p. 1, the same.
Plath, 1919, p. 377, on Astragalinus tristis.
It is probable that there are here more than one kind of parasite, belonging to the Calliphorine or other flies.
The genera Onesia and Melinda must be excluded from this list, as Dr Keilin’s researches show that the former is parasitic in earthworms and the latter lives in snails (1915 and 1919). The gen. Lucilia seems to be questionable, as Hesse’s observations of 1921, p. 154, are not convincing.
If we have to deal with some Calliphorine larva, the gen. Nitellia must be taken into consideration. This genus was established in 1830 by Robineau- Desvoidy, but was united with Pollenia by all subsequent writers except Hendel and Tyler Townsend. It differs from Pollenia in having the common basal stem of the Radius ciliated above, as in Protocalliphora, with which it shows moreover a notable resemblance in the shape of thorax, in the flattened body and in the whole facies. The type species (vespillo Fabricius, 1786) is common in Europe and is present even in North America; its bionomics are still unknown.
III. PHI LORN IS.
Muscid larvae, typically living as subcutaneous parasites on birds, have been found only in the Neotropical Region. They belong to the Anthomyidae and not to the Calliphorinae.
It seems that in Central and South America there are several species which at present are not well distinguished. The first case was recorded by Macquart in 1853 from San Domingo under the name of Aricia pici parasitic on Picus striatus1. The species was subsequently reported under the same name by Blanchard in 1896 who on Brauer’s authority established its synonymy with Mesembrina anomala Jaennicke. The birds infested were Oriolus cayennensis
1 Macquart’s assertion that Salle had seen similar tumours in Mexico on other birds ( Icterus ) and even on squirrels is doubtful; the latter case may have been due to some Cuterebrine larva
M. Bezzi
37
and Or. mexicanus from French Guiana. Blanchard had already recorded it in 1895 under the name of Spilogaster anomalus as a subcutaneous parasite of man.
In 1901 A. de Miranda Ribeiro accepted the above synonymy on the authority of Prof. Mik of Vienna, to whom he had sent specimens for identi¬ fication. These specimens were caught on the bird Peristera ruf axilla.
Meinert in 1889 had already described, from an unidentified Brazilian bird nestling, a muscid larva as Philornis molesta which evidently belongs to the same group. Other allied forms were described in 1895 by Tyler Townsend from the West Indies under the name of Mydaea sperm ophilae, found on birds of the genera Spermophila and Mimus.
In two papers of 1911 and 1913 the late Dr J. C. Nielsen has studied very thoroughly two species of bird parasites under the names of Mydaea anomala and Mydaea torquans ; the material examined was obtained from Argentina, and the larvae were found as subcutaneous parasites in tumours on various birds, both old and young ones; torquans occurring on birds of the genera Spermophila , Mimus, Homorus and Pitangus , and anomala on Xiphocolaptes albicollis.
The Rev. J. Aiken (1913) has described Mydaea pici from British Guiana and has recognised four different species belonging to the present group. On the contrary Lutz and Neiva in 1912, mentioning that subcutaneous larvae are frequent on young birds in Brazil, think that they all belong to a single species, Mydaea pici Macquart. Engel in 1920 expressed a similar opinion.
In 1916 Neiva and Penna recorded Mydaea pici from numerous birds in Central Brazil, saying that the larvae are so frequent as to be commonly named i(berro: ; they were found on birds of the genera Cassicus, F urnarius, Molothrus, Paroaria, Amazona, Pionus, etc., mostly belonging to the Passeres.
The late Prof. Stein in 1918 revised Mik’s types of 1901 under the name of Mydaea anomala ; and in the World Catalogue of 1919 stated that anomala and torquans form a single species.
In my opinion there are in the Neotropical Region several species of Anthomyidae, the larvae of which live in subcutaneous tumours of birds. This opinion is supported by the fact that in a collection of Brazilian flies belonging to this group sent to me some years ago by Dr Lutz, Prof. Stein has recognised the following species of Mydaea: brevipectinata Stein, lati- palpis Stein, sparsiplumata Stein and tinctinervis Stein, all described in 1918. It is probable that some of these species were bred from subcutaneous larvae.
As a conclusion, we can provisionally place the South American An¬ thomyidae with the larvae forming the subcutaneous tumours in birds together under the generic name of Philornis.
38
Parasitic Diptera
Philornis.
Meinert, 1889, p. 315.
1. Philornis pici (Macquart).
Aricia pici , Macquart, 1853, p. 657, pi. xx, 2.
Hylemyia pici, Osten Sacken, 1878, p. 167; Aldrich, 1905, p. 553.
Mydaea pici, Miranda Ribeiro, 1901, p. 156, pi. i; Busck, 1906, p. 2; Lutz and Neiva, 1912, p. 133; Aiken, 1913, p. 3 (sep.), figs. 1-11; Rodham and Bequaert, 1916, p. 247; Neiva and Penna, 1916, p. Ill; Keilin, 1917, pp. 399 and 436; Stein, 1919, p. 123; Engel, 1920, p. 250.
? Hylemyia angustifrons, Loew, 1861, p. 41.
Mesembrina anomala, Jaennicke, 1867, p. 377, pi. ii, fig. 4.
Spilogaster anomala, Brauer and Bergenstamm, 1893, p. 209, note 114, 1; Blanchard, 1895, p. 118 and 1895, p. 652, pi. 17, figs. 5-9.
Mydaea anomala, Nielsen, 1913, p. 252, figs. 1-4; Aiken, 1913, p. 3 (sep.); Keilin, 1914, p. 114 and 1917, p. 399; Stein, 1918, p. 212.
Philornis molesta, Meinert, 1889, p. 315, pi. vi.
Geographical distribution. If all the above records belong really to one species, it must have a wide range of distribution over Neotropical Region, from Cuba, San Domingo and Central America to Argentina.
Ethology. Well illustrated by Meinert and others. The larva pupates in a cocoon of earth cemented by a white substance. This fact seems to be un¬ common in Anthomyidae, but is observed also in the tropical Synthesiomyia nudiseta, v. d. Wulp, besides the cases recorded by Dr Keilin, 1917, p. 437.
2. Philornis torquans (Nielsen).
Mydaea torquans , Nielsen, 1913, p. 252, fig. 3; Aiken, 1913, p. 3 (sep.); Keilin, 1914, p. 114 and 1917, p. 399.
Mydaea anomala, Nielsen, 1911, p. 195, figs. 1-14.
Geographical distribution. At present only known from Argentina. Ethology. The larva pupates freely, without producing a cocoon.
3. Philornis spermophilae (Tyler Townsend).
Mydaea spermophilae, Tyler Townsend, 1893, p. 381; 1894, p. 173 and 1895, p. 79; Aldrich, 1905, p. 543; Aiken, 1913, p. 3 (sep.).
Mydaea spermophila, Keilin, 1917, p. 399.
Geographical distribution. Only known from Jamaica; but Townsend, l.c., records a different species, which infests birds in Trinidad.
Ethology. The larva does not make a cocoon for pupation.
These three flies can be distinguished as follows:
1 (4). Third antennal joint about twice as long as the second1; abdomen
1 The figure given by Blanchard, 1896, pi. 17, figs. 5—8, if really belonging to pici {anomala), must be somewhat inaccurate, showing, in opposition with the figures of Macquart, Jaennicke, Miranda Ribeiro and Aiken, a much longer third antennal joint and a bare arista.
39
M. Bezzi
more or less yellowish at the base; legs as a rule entirely yellowish, at least in the male.
2 (3). Frons more narrow, the parafrontalia of the male almost touching
above, and the middle stripe of the female being not broader than one of the parafrontalia; sides of the face yellowish; legs entirely yellow even in the male . . Picz Macquart.
3 (2). Frons broader, the parafrontalia of the male well separated to the vertex and the middle stripe of the female broader than one of the para¬ frontalia; sides of the face silvery; legs of the female with blackish femora.
torquans Nielsen.
4(1). Third antennal joint about three times as long as the second; abdomen entirely black and the silvery pollen forms a somewhat marmorate
pattern; legs browmish, with basal portion of femora darker.
spermophilae T. T.
IV. CARNUS.
The unique non-pupiparous Dipteron, which in the adult stage is parasitic on birds, is the strange Camus hemapterus ; the fly is haematophagous and is to be found chiefly on young birds, while the larva lives in the nests, but is
saprophagous.
The fly was described and figured by Nitzsch about a century ago, was redescribed and figured by Egger m 1854 under the same name, and owing to a misinterpretation of the original description w^as in 1862 renamed b\ Schiner as Cenchridobia eggeri. Thus for a long time it has been believed that there were two species, one belonging to the Pupipara, and one to the Acalvpterata of the family Borboridae (Schiner), or Sepsidae (Brauer).
Collin in 1911, in a short but important paper, has cleared up the matter recognising but a single species and locating it in the family Milichiidae.
In the subsequent year there appeared the paper of Prof, de Meijere, with the confirmation of the singular fact that the fly emerges from the puparium fully winged, and loses subsequently the wings, short stumps of which remain only as in the Pupiparous gen. Lipopteua. Thus in my papei of 1916 on the reduction of wings in the Diptera, I have included Camus, with Lipopteua, Echestypus and Ascodiptevon, in no. i of my graduation,
p. 108.
A supposed second species of the gen. ( avuus wras described in 1913 bv Stobbe, but its distinction seems to be very doubtful. Engel in his recent paper has added some new records, and therefore this inteiesting flv is at present known to occur on birds of the genera Aquila , halco, Iicus, lynx, Sylvia, Stumus and Coloeus. I have received from Prof. B. Grassi a couple of specimens caught at Maccarese, near Rom, by Luigioni on a nestling of Falco ; these specimens are dark coloured and bristly as in setosus.
40
Parasitic Diptera
Carnus.
Nitzsch, 1818, p. 305 and p. 283.
Cenchridobia, Schiner, 1862, p. 435, and 1864, p. 334.
1. Carnus hemapterus, Nitzsch.
Carnus hemapterus, Nitzsch, 1818, p. 305; Germar, 1822 (?), pi. 24 and 25; Egger, 1854, p. 3, pi. ii, figs. 7-11; Bezzi, 1900, p. 522; Collin, 1911, p. 138; Bezzi, 1911, p. 115; de Meijere, 1912, pp. 1-18, figs. 1-12; Melander, 1913, p. 237, Stobbe, 1913, p. 193; Bezzi, 1916, pp. 108 and 177; Wasielewski and Wuelker, 1918, p. 117; Engel, 1920, pp. 249 and 258; Frey, 1921, p. 151, pi. x, fig. 125.
Cenchridobia eggeri, Schiner, 1862, p. 436 and 1864, p. 335; Brauer, 1880, p. 117, 3 and 1883, pp. 40 and 87 ; Bezzi, 1900, p. 521 ; Becker, 1905, p. 36. Carnus setosus, Stobbe, 1913, p. 193.
Geographical distribution. Only known from Europe : Finland, Germany. Austria, Hungary, Roumania and Middle Italy.
V. CHORTOPHILA AND NEOTTIOPHI L UM.
It may be finally recorded that in bird nests are to be found many other species of Myiodaria, which are not parasitic, but only inquiline, living prob¬ ably on excrement or on organic refuse. Falcoz in 1914, p. 60, has pointed out the great number of dipterous larvae found in bird nests. It is to be noted that these flies, as far as is known, belong to the lower forms of Anthomyidae and even to the Acalypterata.
Of the former there are only the two following species, one from Europe and one from North America:
1. Chortophila eannabina, Stein, 1916, p. 169; Kramer, 1917, p. 49; Engel,
1920, p. 258.
Germany; bred by Kramer from larvae in nest of Acanthis eannabina.
2. Chortophila nidicola (Aldrich).
Hylemyia nidicola, Aldrich in Plath, 1919, p. 380.
State of Washington, U.S.A. ; bred by Plath from larvae found in various bird nests.
Among Acalypterate flies there is the following very interesting genus :
N EOTTIOPHIL UM.
Frauenfeld, 1868, p. 894.
1. Neottiophilum praeustum (Meigen).
Dryomyra praeusta, Meigen, 1826, p. 257.
Neottiophilum praeustum,, Mik, 1882, pp. 98 and 194, 1887, p. 34 and 1894, p. 166; Brauer, 1883, pp. 40 and 86; v. Roeder, 1892, p. 204 and 1895, p. 270; Strobl, 1894, p. 86; Becker, 1902, p. 219 and 1905, p. 38; Bezzi, 1911, p. 115; Kramer, 1917, p. 58; Engel, 1920, p. 250.
M. Bezzi
41
N eottiophilum fringillarum, Frauenfeld, 1868, p. 895.
Blepliariptera cartereaui , Bigot, 1881, p. 370.
Geographical distribution. At present known only from Central Europe: France, Germany and Austria. The adult fly seems to be rare, and was found mostly in houses, on windows ; it lives probably on trees.
Ethology. The fly was bred from puparia found in nests of Fringilla coelebs and of Passer domesticus, but almost nothing is known about the larva.
VI. CONCLUSIONS.
1 . The Myiodaria living with birds show a parallelism between the grades of their parasitic adaptation and their systematic position.
2. The lower forms — Acalypterata have saprophagous larvae, living in the nests of several orders of birds: Scansores, Passeres and Raptores. In the larval stage they feed upon decaying organic matter, while in the adult stage they are, in some cases, blood-sucking (Camus).
3. The intermediate forms — Anthomyidae show two grades of adaptation : (a) lower forms, the larvae of which are mainly saprophagous or phytophagous (Chortophila) and which, like the Acalypterata , live in the nests upon de¬ caying substances; (6) higher forms the larvae of which are mainly carnivorous and have adapted themselves to two modes of life: (A) as subcutaneous para¬ sites ( Philornis ) of Scansores, Columbae, and Passeres; (B) as intermittent haematophaga, on Passeres (Passeromyia).
4. The higher Myiodaria — the Calliphorinae show in their larval stage the two last types of parasitic adaptation, i.e. (a) intermittent haematophagy (Protocalliphora), and (b) possibly a subcutaneous mode of life on Passeres only.
5. The adult flies of all the intermediate and higher Myiodaria are non¬ bloodsucking. It seems to be a rule among the Diptera that the forms with haematophagous adults have non-haematophagous larvae and vice versa.
6. All these facts have to be taken into consideration in the study of other parasitic Myiodaria and especially the heterogeneous groups like Pupipara and Oestridae which, undoubtedly, are of polyphyletic origin, and are derived from lower, intermediate and higher Myiodaria.
VII. BIBLIOGRAPHY.
Aiken, J. (1913). A subcutaneous larva of Mydaea. B. O. Medical Annual, pp. 1^, 3 pis. Aldrich, J. M. (1901). Synonymische Notiz. Wien. Ent. Zeit. xx, p. 68.
- (1905). A Catalogue of North American Diptera ( or Two-winged Flies), pp. 680.
Washington.
Becker, Th. (1902). Die Meigen’schen Typcn der sogen. Muscidae acalypterae (Muscaria holo- metopa) in Paris und Wien. Zeitschr. fur Hym. und Dipt, ii, pp. 209-256, 289-320, 337-349.
- (1905). Katalog der paldarktischen Dipteren, iv, pp. 328. Budapest.
Bezzi, M. (1900). Sulla presenza del genere Chionea Dalman in Italia, e !a riduzione delle ali nei ditteri. Rendic. Istit. Lomb. Milano, xxxiii, pp. 511-526.
42 Parasitic Dip ter a
Bezzi, M. (1907). Katalog der paldarktischen Dipteren. hi, pp. 1-597. Budapest.
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Arch, de Parasit. Paris, xv, pp. 110-143, 15 figs.
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pp. 85-182, 11 pis.
Bigot, J. M. F. (1877). Dipteres nouveaux ou peu connus. xi. Notes et melanges dipt^ro- logiques. Ann. Soc. Ent. Fr. (5) vn, pp. 260-262.
— — (1881). Dipteres nouveaux ou peu connus. xxvi. Ibid. (6) i, pp. 363-371.
- (1887). Dipteres nouveaux ou peu connus. Muscidi (J. B.). Bull. Soc. Zool. Fr.
xn, pp. 581-617.
Blanchard, R. (1895). Note in Bull. Soc. Zool. Fr. xx, p. 118, seance du' 14 Mai.
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YAricia pici Macq. Bull. Soc. Ent. Fr. lxv, pp. 652-654, pi. xvii, figs. 5-9.
Bonsdorff, J. E. (1866). Finlands Twavingade Insekter ( Diptera ), etc-. Andra delen, pp. 306. Helsingfors.
Brauer, F. (1880). Die Zweifliigler des Kaiserlichen Museums zu Wien. i. Denkschr. d. math.-naturwiss. Cl. der K. Akad. d. Wiss. Wien, xm, pp. 105-216, pi. 6.
- (1883). Die Zweifliigler des Kaiserlichen Museums zu Wien. in. Ibid. XLvn, pp. 1-
100, pi. 5.
Brauer, F. and Bergenstamm, J. v. (1893). Die Zweifliigler des Kaiserlichen Museums zu Wien. vi. Vorarbeiten zu einer Monographic der Muscaria schizometopa (exclusive Anthomyidae). Pars hi. Ibid, lx, pp. 89-240.
- (1894). Die Zweifliigler des Kaiserlichen Museums zu Wien. vn. Vorarbeiten zu
einer Monographic der Muscaria schizometopa (exclusive Anthomyidae). Pars iv. Ibid, lxi, pp. 537-624.
Busck, A. (1906). Mydaea pici Macq. Proc. Ent. Soc. Washington, vm, pp. 2-3.
Collin, J. E. (1911). On Camus hemapterus Nitzsch ( Cenchridobia eggeri Schiner) and its systematic position among the Diptera. Novit. zoolog. xvm, pp. 138-139.
Coquillett, D. W. (1910). The type-species of the North American genera of Diptera. Proc. U.S. Nat. Museum , xxxvn, pp. 499-647.
Corti, E. (1897). Aggiunte alia Fauna ditterologica della provincia di Pavia. Terza Cen- turia. Bull. Soc. entom. ital. xxix, pp. 136-143.
Coutant, A. F. (1915). The habits, fife-history and structure of a blood-sucking Muscid larva ( Protocalliphora azurea). Journ. of Parasii., Urbana, III. i, pp. 134-150.
Dufour, L. (1845). Histoire des metamorphoses de la Lucilia dispar (Diptere). Ann. Soc. Ent. Fr. (2) in, pp. 205-214, fig.
Egger, J. (1854). Beitrage zur bessern Kenntniss des Camus hemapterus Nitzsch. VerhandL zool.-bot. Ver. Wien, iv, pp. 3-7, pi. i, figs. 7-11.
Engel, E. O. (1920). Dipteren, die nicht Pupiparen sind, als Vogelparasiten. Zeitschr. f. wiss. Insektenbiol. xv, pp. 249-258, 9 figs.
Falcoz, L. (1914). Contribution a V etude de la Faune des Microcavernes. Faune des terriers et des nids, pp. 185, 1 pi., 38 figs. Lyon.
- (1921). Materiaux pour 1’ etude de la faune plioleophile Premiere. Note: Dipteres.
Bull. Soc. Ent. Fr. pp. 137-142.
Fallen, C. F. (1816). Beskrifning ofver de i Sverige funne flugarter, som kunna foras till slagtet Musca. Vetensk. Acad. Handl. pp. 226-254.
- (1821). Diptera Sueciae. Vol. n, Monographia Muscidum, Pars iv, pp. 41-48 (resp.
Sundewall). Lundae, 1818-1825.
Frauenfeld, G. (1868). Zoologische Miscellen. xv. Verb, der zool.-bot. Ges. Wien, xvm, pp. 885-902.
Frey, R. (1921). Studien liber den Bau des Mundes der niederen Diptera Schizophora, etc. Acta Soc. Faun, et Flora Fenn., 48, pp. 3-245, pi. x.
M. Bezzi 43
Germar, E. F. (1822-?). Fauna Insector um Europae. Ease, ix, tab. 24-25. Halae, 1817 -1840.
Gilbert, P. A. (1919). A Dipterous Parasite of Nestling Birds. Emu , xix, pp. 48-49.
Grimshaw, P. H. (1901). Fauna Hawaiiensis or the Zoology of the Sandwich (Hawaiian) Isles, etc. Vol. m, Part i, Diptera, pp. 1-77, 3 pis. Cambridge.
Heinroth, — (1916). Journ.f. Ornithologie, I, pp. 158-159.
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_ ( 1901 b). Ueber einige neue oder weniger bekannte europaische Muscaria schizometopa.
Verh. zool.-bot. Ges. Wien, li, pp. 198—211.
Henshaw, W. H. (1908). A parasitic fly injurious to our native birds. The Auk, Cambridge, Mass. (N.S.), xxv, pp. 87-88.
Hesse, E. (1921). Bemerkung zu: E. Engel, Dipteren, die nicht Pupiparen sind, als Vogel- parasiten. Zeitschr. f. wiss. Insektenbiol. xvi, p. 154.
Hough, G. de N. (1899 a). Some North American genera of the Dipterous group Calli- phorinae Girschn. Entom. News, x, pp. 62-66.
_ (1899 6). Synopsis of the Calliphorinae of the United States. Zoolog. Bull, n, pp.
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Jaennicke, F. (1867). Neue Exotische Dipteren aus den Museen zu Frankfurt a. M. und Darmstadt. Abhandl. der Senckenb. Ges. vi, pp. 311-407, pi. 2.
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Kirsch, Th. (1867). Ueber zwei Fliegenlarven aus dem Nacken eines jungen Sperlings. Berl. ent. Zeitschr. xi, pp. 245-246, pi. m, figs. 3^.
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- (1917). Pie Musciden der Oberlausitz. Ibid, xxxm, pp. 1-96, figs.
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Metgen, J. W. (1826). Systematische Beschreibung der bekannten Europaischen zweifliigeligen Jnsekten, Fiinfter Theil, pp. 412, pi. 13. Hamm.
Meijere, J. C. H. de (1902). Ueber die Prothoracalstigmen der Dipterenpuppen. Zoolog. Jahrbiich. xv, pp. 623-692, pis. 32-35.
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zu Konigsberg, Lin, pp. 1-18, figs. 1-12.
Meinert, Fr. (1889). Philornis molesta, en paa Fugle snyltende Tachinarie. Vidensk. Meddel. nat. Foren. pp. 304-317, pi. vi.
44 Parasitic Diptera
Melander A. L. (1913). A Synopsis of the Dipterous Groups Agromyzinae, Milichiinae, Ochthiphilinae and Geomyzinae. Journ. N. Y. Ent. Soc. xxi, pp. 219-300, pi. vrn.
Mik, J. (1882). Ueber die Dipteren-Gattung N eottiophilum Frnfld. Wien. Ent. Zeit. I, pp. 194-197. The synonymy was already given in the same vol. p. 98, note 2.
(1887). Dipterologische Miscellen, iv, p. 21. Ein seltenes Dipteron. Ibid, vi, pp. 34—35.
- (1894). Dipterologische Miscellen (2 Serie), v. Ibid, xm, pp. 164-168.
Miller, — (1909). Worthington, Soc. Bird Life, no. 2, pp. 1-8.
Miranda Ribeiro, A. de (1901). Sobre a Mydaea pici Macq. Archiv. do Mus. nac. de Bio de Janeiro, xi, pp. 153-156, pi. i.
Xeiva, A. and Penna, B. (1916). V iajem cientifica pelo Norte da Bahia, sudoeste de Pernambuco, sul do Piauhi e de norte a sul de Goiaz. Mem. do Instit. Osw. Cruz, Rio de Janeiro, vin, pp. 74-224, 28 pis.
Nielsen, J. C. (1911). Mydaea anomala Jaenn., a parasite of South American birds. Vidensh. Meddel. fra nat. Foren. lxiii, pp. 195-208, figs. 1-14.
- (1913). On some South American species of the genus Mydaea, parasitic on birds.
Ibid, lxv, pp. 251-256, 4 figs.
Nitzsch, C. L. (1818). Die Familien und Gattungen der Thierinsecten ( insecta epizoica) als ein Piodromus der Xaturgeschichte derselben. Gerrnars Mag. d. Entomol. hi, pp. 261-316.
Nowicki, M. (1867). Faunistische Miscellen. Verh. der nat. Ver. Briinn, p. 44.
Osten Sacken, C. R. (1878). Catalogue of the described Diptera of North America (Second Edition), pp. 276. Washington.
Pandelle, L. (1896). Etudes sur les Muscides de France. Rev. d? Entom. Caen, xv pp 214-215.
Patton, W. S. (1920). Note on the occurrence of Passeromyia heterochaeta Villeneuve, in India. Ind. Journ. Med. Res. vm, pp. 30-31, 1 pi.
Pavay-Vajna, Fr. (1909). Onesia cognata als Vogelparasite. Aquila, xvi, pp. 288-290.
Plath, O. E. (1919 a). Parasitism of nestling birds by fly larvae. The Condor, xxi, pp. 30—39.
(1919 b). A Muscid larva of the San Francisco Bay Region which sucks the blood of nestling birds. Univ. Calif. Publ. Zool. xix, pp. 191-200.
- (1919 c). The prevalence of Phormia azurea Fallen (larva parasitic on nestling birds)
in the Puget Sound Region and data on two undescribed flies of similar habit. Ann. Ent. Soc. Amer. xn, pp. 373-381. With Appendices by Townsend and Aldrich.
Portschinska , J. (1887). Diptera europaea et asiatica nova aut minus cognita (cum notis biologicis). Hor. Soc. Ent. Ross, xxi, pp. 3-20, pi. i.
Robineau-Desvoidy, J. B. (1830). Essai sur les Myodaires. Mem. Savants etrang. Acad. Paris, ii, pp. 1-813.
(1849). Note sur la synonymie de la Phormia regina (Diptere). Ann. Soc. Ent. Fr. (2) vn, Bull. p. 4.
(1863). Histoire Naturelle des Dipteres cles environs de Paris, n, pp. 920. Paris.
Roder, V. \. (1892). Ein neuer Fundort der Dipteren N eottiophilum praeustum Mg. und Acyglossa diversa Rond. Entom. Nachr. xvm, pp. 204-206.
(1895). Xeue Fundorte der Diptere N eottiophilum praeustum Meig. Wien. Entom. Zeit. xiv, p. 270.
Rodhain, J. (1914). Sur une larve de Muscinae vivant dans le nid de Passer griseus au Congo. Rev. zool. Afric. hi, pp. 213-217.
Rodhain, J. and Villeneuve, J. (1914). Passeromyia, genre nouveau des Anthomyidae (Dipt.), a larve hematophage parasite des jeunes oiseaux. Bull. Soc. Path. exot. vm pp. 591-593.
Rodhain, J. and Bequaert, J. (1916). Histoire de Passeromyia heterochaeta Villen, et de Stasisia {( ordylobia) Rodhaini Ged. Bull. Sci. de la Fr. et de la Belg. (7) xlix pp 236-289, 1 pi., 14 figs.
M. Bezzi 45
Rodhain, J. (1919). Nouvelles observations sur la biologic de Passeromyia heterochaeta Villeneuve. Bull. Biol, de la Ft. et de la Belg. lii, pp. 499— ol.O, 2 figs.
Rondani, C. (1862). Dipterologiae italicae Prodromus, v, pp. 239. Parmae.
Rossi, F. (1848). Syslematisches V erzeichniss der Zweifliigelichen Insecten ( Diptera ) des Erzherzogthumes Osterreich mit Angabe des Standortes, der Flugzeit und einigen andern physiologischen Bemerkungen. pp. 86. Wien.
Roubaud, E. (1914). Etudes sur la Faune Parasilaire de V Afrique occidental Frangaise.
Premier Fascicule, pp. 250, 4 pis., 70 figs. Paris.
_ _ (1915 a). Hematophagie larvaire et affinites parasitaires d’une mouche Calliphorine,
Phormia sordida, parasite des jeunes oiseaux. Bull. Soc. Path. exot. vin, pp. 77-79. _ (1915 6). Les Muscides a larves piqueuses et suceuses de sang. C.R. Soc. Biol. Paris,
lxxviii, pp. 92-97, fig. 2.
_ (1917). Observations biologiques sur Nasonia brevicornis Ashm. chalcidide parasite
des pupes des muscides. Bull. Sci. de la Fr. et de la Belg. (7) l, pp. 425— 439,
1 fig.
_ (1918). Precisions sur Phormia azurea Fall. Muscide a larves hemophages paiasites
des Oiseaux d’Europe. Bull. Biol, de la Fr. et de la Belg. LI, pp. 420—430, pi. v. Roubaud, E. and Van Saceghem, R. (1916). Observations sur quelques lnsectes et Acariens parasites du Betail au Congo Beige. Bull. Soc. Path. exot. ix, pp. 763—767. Schiner, J. R. (1862). Fauna austriaca. Die Fliegen {Diptera). I Thcil, pp. 674, 2 pis.
Wien.
_ (1862). Vorlaufiger Commentar zum dipterologischen Theile der “Fauna austriaca.”
v. Wien. Entom. Monatsschr. vi, pp. 428—436.
- (1864). Fauna austriaca. Die Fliegen {Diptera). II Theil, pp. 658. Wien.
Schneider, 0. (1866). Notiz in Isis, Sitzungsber . Dresden, p. 89.
Stein, P. (1909). Neue Javanische Anthomyiden. Tijdschr. voor Entom. lii, pp. 205-271.
- (1916). Die Anthomyiden Europas, etc. Archiv f. Naturgesch. lxxxi, pp. 1-224.
_ (1918). Zur weitern Kenntnis aussereuropaeischer Anthomyiden. Ann. Mas. nat.
Hung, xvi, pp. 147-244.
_ (1919 a). Die Anthomyiden Gattungen der Welt, analytisch bearbeitet, nebst einem
Kritisch-systematischen Verzeichnis aller aussereuropaischen Arten. Archiv f. Naturg. Lxxxm, pp. 85-178.
_ (1919 6). Anthomyiden aus Java, Sumatra, Waigeoe und Ceram. Tijdschr. voor
Entom. lxii, pp. 47-86.
Stobbe, R. (1913). Zur Kenntnis der Gattung Camus Nitzsch (= Cenchridobia Schiner) mit 1 nov. sp. (Dipt.). Deutsch. Ent. Zeitschr. pp. 192-194.
Strobl, G. (1893). Beitrage zur Dipterenfauna des osterreichischen Littorale. Wien.
Ent. Zeit. xn, pp. 89-108.
- (1894). Die Dipteren von Steiermark. II Theil. Mittheil. d. Nat. Ver. fur Steierm.
xxx, pp. 1-152.
Surcouf, J. M. and Gonzalez-Rincones, R. (1912). Essai sur les Dipteres I uliieiants du V enezuela. Deuxieme Partie, pp. 240, 100 figs. Paris.
Townsend, C. H. Tyler (1893). Notes from the Museum, Institute of Jamaica, Nov. 22,
no. 70.
- (1893 a). A bot-like Anthomyid, parasite of a bird. Journ. Instit. Jamaica, i, pp.
381-382.
- (1894). The grass-quit bot, an Anthomyid parasite of nestling birds. Ibid, ii, pp.
173-174.
- (1895). Contributions to the Dipterology of North America, n. Trans. Am. Ent.
Soc. xxn, pp. 55-80.
- (1916). On Australian Muscoidea, with description of new forms. Insec. Insc.
Menstr. iv, pp. 44-45.
46 Parasitic Diptera
Townsend, C. H. Tyler (1919). New genera and species of Muscoid flies. Proc. U.S. Nat. Mus. lvi, pp. 541-592.
Verrall, G. H. (1886). A hundred new British species of Diptera. Ent. Monthly Mag. xxn, pp. 179-182, 199-202, 230-234.
Villeneuve, J. (1910). Notes synonymiques sur quelques Dipteres — Types. Deutsch. Ent. Zeitschr. pp. 311-313.
- (1911). Notes dipterologiques. i. Sur le genre Phorrnia R. D. Wien. Entom. Zeit.
xxx, pp. 84-85.
(1913). De quelques Tachinaires a grande extension geographique. Feuille des jeunes Natur. xliii, pp. 131-132.
— — (1915). Dipteres nouveaux d’Afrique. Bull. Soc. Ent. Fr. lxxxiv, pp. 225-227, fig.
- (1918). Sur Phormia sordida Zett. Ibid. no. 10, pp. 158-159.
Wasielewski, T. von and Wuelker, G. (1918). Die Haemofjroteus-lrdcktion des Turm- falken. Beih. zum Arch. f. Schiff.s-. u. Tropen-Hyg. xxn, pp. 117-212, 4 pis., 11 figs. Zetterstedt, J. W. (1838). Insecta Lapponica descripta. col. 1140. Lipsiae, 1838-1840.
- (1845). Diptera Scandinaviae disposita et descripta. Tomus quartus, pp. 1281-1738.
- (1849). Ibid. Tomus octavus, pp. 2935-3366.
- (1859). Ibid. Tomus tredecimus, pp. 4943-6190.
47
DESCRIPTION OF A BOX FOR COLLECTING AND TRANSPORTING LIVING INSECTS, ETC.
By E. N. PAVLOVSKY, M.D., Sc.D.,
Professor at the Military Academy of Medicine , Petrograd.
K (With 6 Text-figures.)
When collecting small living animals, particularly insects or other small Arthro¬ pods, for dissection, or search for parasites, etc., one encounters difficulties in choosing a suitable and convenient arrangement in which the animals may be kept without injuries until they are brought home. The usual entomological corked glass tubes and wide-mouthed jars with a gauze cover are not prac¬ ticable when a large number must be carried ; their bulk hinders free movement, and their fragility renders it necessary to pack them carefully in a box or other suitable containers.
In the summer of 1911, I was shown, at the Natural History Museum, Simpheropol, a very convenient apparatus for the collection and dispatching of Arthropods, which had been sent from Uriev by Prof. K. K. Saint-Hilaire. Unfortunately, I have not been able to discover the name of the inventor of the original model, a description of which I have never met with in the literature. Having convinced myself, during four years’ experience of its use, of the valuable qualities of this apparatus, I have made some alteration in its construction, which I believe will render its application wider, and have decided to publish a description. If ever I should succeed in discovering the name of the author of the original model, I shall publish a supplementary note to that effect.
The model received from Prof. Saint-Hilaire consisted of a flat wooden box, 48*5 cm. in length, 34-5 cm. in breadth, and 5*5 cm. in depth. The interior of the box was divided, by eight longitudinal and five transversal partitions, into 40 rectangular compartments (Fig. I). The bottom of the box was formed of a sheet of wire gauze of 0*5-1 mm. mesh, while the lid was constructed and firmly fixed to the frame of the box. In the wood at points corresponding to the centre of each of the 40 compartments a circular aperture, 2*8 cm. in diameter, had been bored. Each of these openings was closed with a cork. The apparatus, as described, is very convenient for collecting and transporting Arthropods of cannibal habits, e.g. spiders, scorpions, myriapods,
48
Box for Collecting hieing Insects
etc., while the conditions of ventilation of the inner chambers are such as to render it a safe means for the transport of sensitive insects, as bumble-bees and other Hymenoptera, which will survive imprisonment in such a box much longer than in glass jars with gauze lids. In the summer 1914 I brought scorpions from Algeria and Tunis to Petrograd. The scorpions remained in the boxes from the 7th of July until the 28th of August, and the percentage of individuals which succumbed was very small, notwithstanding the un¬ favourable conditions of the voyage.
In order to render the apparatus of wider application I have made some modification in its structure which I now proceed to describe.
The external dimensions of the improved box are: length 45 cm.; breadth 31 cm.; and depth, including thickness of lid, 5-5 cm. The internal depth of the box is 4-6 cm. as in the original model.
The wooden lid is in three sections which are hinged and can be opened independently of each other (Fig. III). Each section is fastened by a pair of hooks screwed to the front wall of the box, suitable pins or eyelets being fixed into the edge of the lid for the hooks to engage (Fig. II). For greater security an additional hook is fixed at each end of the box.
The interior of the box is divided by eight continuous fixed transverse partitions (Fig. Ill, F) with equidistant vertical slots for the reception of quadrangular pieces of wood which divide' those spaces between the con¬ tinuous partitions into six compartments; space slots at the ends of the continuous partitions (Fig. Ill, D) serve for the reception of the quadrangular movable partitions when it is desired to increase the capacity of a compart¬ ment by the removal of one of these. If desired, pieces of tin plates may be used in place of wood for the movable partition, but in this case, the slots in the continuous partitions must be cut to fit accordingly. With all the movable partitions in their usual positions, the box is divided into 54 com¬ partments each measuring 4 cm. x 4 cm. x 4-6 cm. In Fig. Ill the endmost series of compartments shows a replacement of the movable partition which results in the formation of two double compartments ( C ) and two single compartments. Such double compartments are suitable for the reception of solpugids or large scorpions. By further replacement it is possible to obtain compartments of triple capacity or more, and by removing all the movable partitions of one transverse compartment, space available for the reception of lizards, etc. may be obtained. The lid of the box is furnished with apertures (M) closed with corks, like that of the original model. The desired arrangement of the movable partitions should be made before the specimens are placed in the box and the latter should be introduced through one of the openings by removing the cork. It is obvious that no section of the lid should be opened when the corresponding part of the box already contains specimens.
The floor of the box is formed of wire gauze, as in the original model, and may be secured in position by a fillet of wood running round the lower edges of the box and screwed to the lateral end walls (see Fig. IV).
E. X. Pavlovskv
49
Fig. I. Bottom of original model, showing wire-gauze floor and partitions.
Fig. II. Lateral view of improved model showing hooks for fastening lid sections.
Fig. III. Plan of improved model showing lid sections, one of which is open: A and B, com¬ partments subdivided into two or four parts respectively by the insertion of diagonal plates; C, double compartment formed by removal of one movable partition; D, space mortice; K, movable partitions; M, aperture of a compartment.
Fig. IV. Detail of the interior in perspective. The lid is not represented. The wire-gauze floor and wooden fillet are clearly shown.
Fig. V. Tin plates for diagonal partitions of compartments: A, incision for reception of part of cork protruding beneath the lid of the box.
Fig. VL Pair of tin plates interlocked for fourfold subdivision of compartment.
Parasitology xiv 4
ilO
Box for Collecting Living Insects
The rectangular compartment of the box may be subdivided by the use of tin plates, cut to the suitable shape shown in Fig. V. These plates are in¬ serted diagonally, either singly (Fig. Ill, A) or cross- wise in pairs (Fig. Ill, B). Such small compartments serve for the reception of small spiders, myriapods, ticks, etc. The broad incision in the upper edge of each of the diagonal division plates should be cut to fit closely to the part of the cork protruding through the aperture of the lid.
Several such boxes, placed one upon the other in such a manner that the gauze bottom of one rests upon the corks of the other, are bound with travelling straps for transport. For ordinary excursions a single box is usually quite sufficient and may be carried conveniently slung by a shoulder strap.
AN ACCIDENTAL INFECTION WITH UNCINARIA
By FRED C. CALDWELL, S.B., MJ).,
State Director , International Health Board , Panama, Republic of Panama.
In the course of a study of a method for the recovery of Uncinaria larvae from the soil, one of the employees of the laboratory became infected under circumstances closely simulating experimental conditions. The history of the case is as follows :
On November 22, 1920, a quantity of faeces containing Uncinaria ova in large numbers was placed on the ground where it would be exposed to the sun and the rain. On December 8, many living larvae were recovered from the soil. These were placed in a tightly corked bottle out of the way on a high shelf, and all employees were warned that the specimen was not to be handled carelessly.
In spite of instructions, however, on December 11, one of the men took the specimen to demonstrate the larvae to some friends. While he was examining the preparation under the microscope, the fluid ran over the edge of the slide and came in contact with his hand. He is accustomed to hold the slide between the thumb and middle finger, allowing his hand at the base of the middle finger to rest upon the stage of the microscope.
Immediately after examining the specimen he became conscious of an intense itching at the tip of the thumb and tip and base of the middle finger.
On the following day, December 12, the three areas were inflamed and swollen and contained numbers of minute bright red points. There was no evidence of pus.
On December 13 the man noticed a marked tenderness under the arm in the region of the axillary glands. There was no evidence of lymphangitis, and the lesions on the hands remained the same as on the preceding day.
On December 14 a marked bronchitis developed. The stool was found to be negative for Uncinaria at this date and remained so for more than a month. Because he had been engaged as microscopist in our hookworm campaign in Panama, he had examined his stool many times during the previous year for ova of Uncinaria, with negative results.
The lesions on the hand had practically disappeared by December 24.
The stool was negative on January 10, 1921, but became positive for Uncinaria on January 1 7, thirty-eight days after the infection. On Januarv 26 the ova were increased in number. The man left the city for the interior on January 26.
4—2
52
Accidental Infection with Uncinaria
In answer to a request for information in regard to his condition in June, the following reply was received:
I have made other examinations and found several ova of Uncinaria. I am experiencing well-marked symptoms of the disease, such as general weakness, dizziness, and palpitation of the heart. My bronchitis is almost the same as when I left Panama. I have not as yet taken the treatment because of lack of time.
After repeated advice in regard to treatment, on October 31 I received the following letter:
When I received your telegram about treatment, I had already taken one treatment with chenopodium and collected the worms which were passed. I am enclosing a sample.
The sample mentioned contained about twenty hookworms. On November 15 the following statement arrived:
After a second treatment I was unable to find any worms. Repeated examinations for ova have been negative. I am now feeling quite well, except for palpitation of the heart.
From the above history it seems certain that in this case the larvae after passing through the skin, travelled up the lymphatics, through the axillary glands, into the blood stream and into the lungs in three days. The parasites had reached the intestines and had matured within thirty-eight days. One treatment with chenopodium had evidently sufficed to cure this particular case.
53
NOSE M A APIS AND ACARAPIS (T ARSON EMUS) WOODI IN RELATION TO ISLE OE WIGHT BEE DISEASE.
By GEO. W. BULLAMORE,
Molteno Institute for Research in Parasitology , Cambridge.
CONTENTS.
PAG E
I. Some outbreaks of bee disease previous to 1900 .... 53
II. The outbreak of 1906 commonly called Isle of Wight Disease . 55
III. The symptoms of disease in adult bees ..... 55
IV. Attempts to ascertain the cause of Isle of Wight Disease . . 56
A. Bacillus pestiformis apis ....... 56
B. Nosema apis ......... 56
C. Acarapis ( Tarsoncmus ) uoodi ...... 58
V. Conclusions ........... 61
I. Some Outbreaks of Bee Disease previous to 1906.
In former times when this country was dependent upon bees for its supply of sugar, a heavy mortality among these insects was of sufficient importance to be placed on record. In his Animal Plagues Fleming (1871) gives a number of references to such losses. In Ireland there was a ‘‘mortality of bees'’ in 950 a.d. and again in 992 a.d. there was a “great mortality upon men, cattle and bees.” In 1035 a.d. the destruction of bees afflicted the whole of Bavaria.
An eclipse of the sun in 1124 a.d. was followed by a great pestilence amongst oxen, sheep, pigs, and bees. During the time of the “Black Death,” also, * there appears to have been heavy losses of bees and at the Manor Court of Heacham in Norfolk, a statement was made on oath by the steward in the forty-fifth year of the reign of Edward III (1372 a.d.) to the effect that ten out of eleven stocks of bees had perished from the murrain. There is little doubt that this entry in the manorial court rolls refers to epidemic disease, but, as Fleming points out, “there is evidently no relationship between the morind of the bees and that of the sheep and cows.”
The year 1443 a.d. was rainy and tempestuous after May which “much hurted both bees and sheep in Ireland”; while in Italy in 1690 a.d. “bees extracting no sweetness from the calyces of the flowers, but a bitter poison, either died or left the country.” A great mortality among bees and carp is also recorded in 1717 a.d. in Silesia.
A well-known writer on bees, Dr Bevan (1837), states that
in the winter of 1782-3, a general mortality took place among the bees in this country, which was attributed to various causes; want of honey was not one of them; for in some
54
Bee Disease
hives considerable store was found after the bees were gone. Some were of opinion that it .oose fiom the preceding being a bad breeding year and thought that the bees died of old age. Otheis attributed it to the moistness of the spring of 1783 which rendered the pro¬ viding of pollen difficult, for without pollen no brood can be raised.... The fatal influence ascribed to the wetness ol the spring of 1782 seems to be improbable; though it might have affected the quantity of bees bred, it was not likely to put a stop to their breeding altogether, and the young bees ought at any rate to have escaped the desolating evil, if it were old age alone; yet wherever the mortality once made its appearance, every bee became its victim.
In the early sixties of the nineteenth century the columns of The Field newspaper supply evidence of the existence of trouble which beekeepers of the day described in language almost identical with that used to describe the Isle of Wight disease at a later period. Occasional references to a similar trouble are to be found in the bee literature of this country until the beginning of the present century.
In other countries, also, epidemic bee disease may still be heard of from time to time. Berlepsch records that in the spring of 1859, after the bees had made good use of the sallows, wholesale losses took place. The bees were to be seen in heaps, their bodies swollen with watery excrement. Many stocks were one day healthy, the next day half were dead, and the day following that all the bees were dead. The disease appeared in many places about Easter, and in others not till W hitsuntide. It was similar to human cholera, ravaged
the whole of south Hanover and the lands adjoining and even extended into Denmark.
According to Dadant (1907) during the years 1901-05 entire apiaries were depopulated in the province of Ancona, Italy, by mal de Maggio, a disease of the adult bee. The losses occurred just at the opening of the honey harvest.
In Australia, whole apiaries are at times extinguished from some disease the cause of which is unknown (Garrett, 1910).
In Brazil hundreds of stocks perish in March and April, the affected bees dying outside the hives. The beekeepers attribute the trouble to the poisonous properties of the nectar of certain plants which are in bloom at the time1 (Hannemann, 1909).
In Canada and the United States epidemics occur and the following quota¬ tion from the American Gleanings in Bee Culture is an interesting description of virulent disease by Critchlow (1904):
is this a new and strange bee disease, or is it a very malignant type of paralysis? Possibly some purely local cause is responsible for the great mortality among the bees. Who can give us some light? There seems to be great danger in Utah of a total loss of the entire bee industry. I shall give you as nearly as I can the conditions, both in the past and at the present time.
In this valley and in the one fifty miles north of here, called Cache Valley, there have been foi many years a great many bees, both in the hands of skilled operators and in the
1 Very many plants have come under the suspicion of beekeepers at one time and another in various parts of the world when there was no local spraying of fruit-blossom, etc., to which the losses could be attributed.
G. W. Bullamore
55
hands of many farmers and small owners. A year ago this spring there were upwards oi 2000 colonies of bees lost. It was thought a year ago that it was owing to the very cold winter weather and changeable weather in the spring; but the strange part of it is that in some localities not any warmer, but if anything, colder, the bees survived all right, with scarcely any loss at all.
I lost last year 300 colonies of bees, and in some localities in Cache Valley there were upwards of 500 colonies that went under; and in this valley (Salt Lake) last year there were fully 800 colonies that perished. It was thought by all those who were interested that it was due to the hard winter and cold spring.
This writer, who had lost nearly all his stocks, bought 225 colonies in a district that disease had not visited, and moved them into his apiary. He continues:
I was there yesterday, and a large part of the entire 225 colonies are affected with what appears to be paralysis. While there is no trembling, they drop down in the grass in front of the hives, and are unable to fly; they seem to mount the grass and twigs with great difficulty, and in taking them up in my hands they were unable to fly away, and, if thrown into the air, would drop to the ground. They seemed to have no desire even to sting. It appears to be contagious, for it seems to affect a certain part of a row, while another section of the row seems to be strong and swarming. A number of hives have all gone under. The entire yard, of course, is exposed for the reason that a few, perhaps twenty, colonies from what were left of the lot last year were put with them without any thought of anything being wrong, except that they were weak.
I am fully satisfied now that the loss last year, which would number at least 2000 colonics in Cache Valley, was due entirely to this condition.
II. The Outbreak of 1906 commonly called Isle of Wight Disease.
In 1906 we first began to hear of the bees in the Isle of Wight suffering from “paralysis,” a disease of world- wide occurrence and of which the cause is unknown. On account of the deadly nature of the Isle of Wight visitation, however, it was afterwards decided that the disease was a new one and the name of Isle of Wight disease came into use. It was found to have been pre¬ sent on the Isle of Wight in 1904, so it was consequently assumed to have then originated and to have spread thence to the mainland. It usually mani¬ fested itself by the presence in the apiary of numbers of crawling bees with their abdomens distended with undischarged faeces. In a few months all the colonies in the apiary were dead.
III. The Symptoms of disease in Adult Bees.
The whole subject of bee diseases teems with difficulties, and it is improb¬ able that any method of differentiating them other than the demonstration of the causal organism is likely to prove of value. Inability to fly, which is the chief symptom in any disease of the adult bee, may originate from a variety of causes, e.g. the ingestion of fungi, mineral poisons, fermenting honey, syrup or fruit juice, or from weakness due to starvation.
Inability to fly leads to a retention of the faeces, which are voided normally by the bee in the open air while flying. We find that a crawling bee may be
•><> Bee Disease
slow and sluggish, may run rapidly, may tremble, present a hairless appear¬ ance, drag its hind pair of legs helplessly, may be subject to dysenteric dis¬ charges, or may impart a yellow colour to the interior of the hive by passing constantly a tiny threadlet of faeces. Possibly the varying manifestation may be due to the multiplication of bacteria of different species or of other parasites in the bowel of the affected bee. Whether the condition is sometimes due to infection it is impossible to decide.
Again, the normal life of the individual bee may vary from eight weeks to as many months according to the activities of the colony, and it may be that organisms which do little harm in two months cause serious trouble where the necessities of the colony require the extension of the lives of its individual members for a longer period.
The difficulty of tracing a connection between a disease and any parasitic organism that may be found in the bee becomes very real when the investiga¬ tion of disease is attempted. If, while the external conditions are favourable, the organism is fed to a colony as an experiment, the bees may fail to die or to develop symptoms; while if unfavourable conditions prevail at the time of the experiment, death may be due to unrecognised causes, such as infection with N . apis (see below). Apparently healthy bees, if prevented from flying, will sometimes develop symptoms (crawling with bowel distention) indistin¬ guishable from Isle of Wight disease.
IV. Attempts to ascertain the Cause of Isle of Wight Disease.
A. Bacillus pestiformis apis.
The first to associate a definite causal organism with Isle of Wight disease was the late Dr Walter Malden (1909). He found no macroscopic appearances in the diseased bees that were not to be found in bees from healthy colonies. Microscopically, “no changes were discovered in the salivary glands, brain, fat-body, heart, tracheae, air-sacs, Malpighian bodies, or honey stomach.” Changes were found in the chyle stomach, however, and attention was con¬ centrated on that organ. In film preparations made from small portions of the chyle stomachs of diseased bees, teased out on glass slides and stained with methylene blue, a bacillus was found with darkly staining ends and a lightly staining central band, resembling Bacillus pestis in general appearance. This was suggested as having a causal relationship to the disease and the name Bacillus pestiformis apis was proposed. Cultures of this bacillus fed in sugar to bees did not appear to have any harmful effect and the view that it is the organism that caused the disease has been abandoned.
B. Nosema apis.
In 1907 Dr Enoch Zander (1911) discovered a protozoon in bees which was recognised as being closely related to Nosema bomb yds, a parasite which did enormous damage to the silkworm industry in France about the middle
Gr. AY. Bullamore
0/
of the nineteenth century. The organism found in bees was named Nosema apis and Zander looked upon it as the cause of heavy losses in Bavaria. The essen¬ tial feature of the trouble was a sudden and extensive mortality among the bees inside and outside the hive. Dysentery was an occasional accompaniment. BerlepsclTs description of the epidemic of 1859 (see p. 54) is considered by Zander to be a typical description of the ravages of unchecked Nosema disease.
The discovery attracted considerable attention. Maassen (1911) found Nosema to be widely prevalent in German apiaries, but although he looks upon the parasite as pathogenic he considers that unfavourable conditions are necessary for the manifestation of disease.
Nussbaumer (1912) found Nosema in association with heavy losses in Switzerland.
Beuhne (1916) states that Nosema was first discovered in Australia in 1909, but that observation showed that it was
doubtful whether the presence of the parasite is in itself necessarily fatal, or that it greatly interferes with the productiveness of the hives excepting under certain conditions due to climatic influence.... In fact under ordinary conditions the disease is endemic, and becomes epidemic only when the vitality of the bee is impaired by the malnutrition during the bees’ larval development which is caused by a dearth, or the inferior quality of the nitrogenous food which bees obtain solely from the pollen of the flowers of plants.
As the result of investigation at Cambridge (Graham-Smith, Fantham, Porter, Malden, and Bullamore 1912) a report was issued showing that Nosema was present in a large percentage of stocks affected with Isle of Wight disease and the disease was attributed to the presence of this parasite.
Later, Anderson and Rennie (1916) working on the Isle of Lewis took up the question of Nosema and Isle of Wight disease. They were unable to cor¬ roborate the findings of the Cambridge investigation and state that Nosema was found to be present in stocks without disease symptoms appearing. They did not find Nosema distributed in the bees of all the stocks but only in a few stocks at any one time, and when it occurred “Isle of Wight disease was not present.”
The result is somewhat surprising as, assuming that Nosema is a com¬ paratively harmless parasite, its non-occurrence in stocks suffering from Isle of Wight disease seems to require some explanation.
While the work on the Isle of Lewis was being carried on, continuous importations of bees were being made from all parts of the country. Bees, assumed to be healthy, were introduced into the apiary from Scotland, Eng¬ land, Wales, Ireland, America, Switzerland, Austria, and Holland. These bees developed crawling symptoms at periods varying from a week to a month or more after arrival. Consequently it is difficult to determine the disease with which Anderson and Rennie were actually working or to feel sure that im¬ portations of other diseases were not being made. The following are the particulars of what is described as a spontaneous recovery.
58
Bee Disease
I hree .stocks of bees, one Italian, one Carniolan and one black (British or Dutch?) were imported into the apiary from Devonshire on June 4th, 1915.
During June these three stocks gave off four swarms and the seven colonies were showing crawling symptoms on June 30th. Three of the colonies died o\\ mg to faulty cpieens and another was destroyed by robber bees. A fifth was reinforced by the addition of three pounds of bees (source not stated) and an Italian queen was also given to this colony. Some driven bees from Wales were placed in the apiary. In December, 1915, there were four stocks alive showing no signs of disease.
Accounts of further work by Dr Rennie and Miss Harvey (1919 a and b) arc given in the J ournal oj the Scottish Board of Agriculture. With regard to A osema the conclusion arrived at is that it "is always a weakening factor, and in the presence of other adverse conditions favourable to the development of dysentery it may become seriously pathogenic to bee stocks.” in ordinary circumstances it vras not found to destroy colonies in the rapid and virulent manner generally found in Isle of Wight disease. The authors consider that in the latter disease there is no recovery of sick bees nor of affected stocks.
Mention is also made of a temporary sickness wdiich sometimes affected all the bees of a district. ‘'In such cases it would appear to be due to some disturbing factor in the nectar or pollen.”
C. Acarapis (Tarsonemus) woodi.
In November, 1920, a paper was read before the Royal Society of Edin¬ burgh in which Dr Rennie, in association with Mr Bruce White and Miss Harvey, described a new species of mite from the tracheae of hive bees. This mite, which was named Tarsonemus ivoodi, was put forward as being the cause of Isle of Wight disease, which was henceforth to be known as “acarine disease.” Hirst (1921) considers that the new mite differs sufficiently from Tarsonemus to form a new genus and proposes that the species shall be renamed Acarajns woodi.
The mites gain entrance to the tracheae by means of the first pair of thoracic spiracles. Crawling of bees, followed by the death of the affected colony then takes place. The presence of the mite is not necessarily fatal however, for we read in the Report to the Royal Society of Edinburgh that
Of 140 stocks believed by their owners to be healthy, 50 or nearly 30 per cent, har¬ boured the parasite. Concurrent with such discoveries we ascertained by direct examination ourselves of flying bees (1) which were members of colonies in which the disease was definitely established and (2) which were taken from colonies believed to be healthy and showing no indications otherwise, that amongst these were to be found considerable numbers harbouring the parasite. This was further complicated by the fact that in those infected flying bees certain of those pathological conditions — e.g. the blackening and hardening of the tracheal tubes — were very marked. As an example it may be quoted that this condition was found in bees entering the hive carrying pollen or nectar, both belonging to stocks in which crawling and other symptoms were well established and also in those reputed healthy stocks.
G. W. Bitllamore
59
In the record of another stock we read that “the hying workers were frequently more heavily parasitised than were the bees of the same stock which were unable to fly.”
In an interesting case which was under the writer’s own observation in April, 1921, the crawling symptom was manifested but no mites were to be found. The crawling disappeared during the summer but reappeared in the autumn when the symptom was accompanied by the presence of mites in the tracheae.
As an example of the recovery of a stock we have the following from a letter received by Dr Rennie (1921):
On January 9th T received your report that the bees from one of my stocks had Tarso- nemus woodi.
From this stock reported diseased on that date 1 had a swarm on the 23rd of May and this swarm swarmed on the 28th of June. 1 have taken off 150 sections and have three very strong stocks.
Dr Rennie’s explanation is that early diagnosis enables us to recognise such recoveries and that formerly this was not possible. We only recognised the presence of the disease after it was irretrievably established. This “gave us an erroneous idea as to the gravity of the disease.”
Such an explanation is not altogether satisfactory. When Isle of Wight disease reached an apiary the loss of colonies was usually 100 per cent, and the margin of error in forming an estimate of its gravity must have been very slight.
In the Report of the Hants and Isle of Wight Beekeepers’ Association for 1906 we read:
Twenty-five years’ acquaintance with bees, bee men and bee life has not revealed any¬ thing so deadly or mysterious as this so-called bee paralysis of the Island.
Silver (1907), who toured the Island in 1907, gives his impressions in these words :
The sight of whole apiaries of 10 to 20 hives standing desolate and deserted in the middle of May is a most distressing one, and standing as I did, under a horse-chestnut tree in full blossom, in the grounds of the Rev. John Vicars, of Colbourne, situated in the centre of the Island, not a bee was visible on a beautiful spring day.
Complete apiaries died out in May and June just after swarming and when the hives must have been tenanted with young bees.
In support of the thesis that T arsonemus woodi has been entirely responsible for the losses known as Isle of Wight disease, Dr Rennie suggests that T. woodi is at present a parasite of bees in this country only. He assumes that Isle of Wight disease has never been clearly shown to exist in any other country and that no such persistent losses have ever occurred before in this or any other country, and discusses the possibility that a new disease has arisen through the migration of the mite from some other insect to the hive bee as host.
The evidence that the mite is not to be found in bees of other countries is confined to the negative results obtained by the examination of a few
60
Bee Disease
hundred bees that have accompanied queen bees sent to this country by post. But if the mite is the cause of a serious disease its occurrence in such bees would be rare. It is in the dwindling and unprofitable apiaries of other countries that search must be made, and until we get definite statements from foreign workers that the mite cannot be found in any of the troubles classed together as paralysis, the chances are in favour of its occurrence.
As to the long duration of Isle of Wight disease as compared with any other epidemic, we have no knowledge as to the number of years that apiaries have been re-stocked where paralysis is causing annual losses. In this country steady losses were formerly confined to districts. That such losses now occur all over the country is due to the altered method of beekeeping. And is it safe to assume that such losses have always been due to the same cause for the last seventeen years in this country and to some other cause in the various countries from which we have imported bees? Since Isle of Wight disease was first reported nearly all the bees of this country have disappeared. Their places have been taken by foreign bees, many of which have also died and the present bees of this country are mainly the descendants of recent importations from various parts of Europe, Asia, Africa, and America.
The German epidemic of 1859 followed the importation of foreign bees.
Our own losses in the early sixties of the last century came shortly after we commenced to import foreign bees and had started a crusade against the sulphuring of weak and redundant stocks. Prior to the starting of the epidemic in the Isle of Wight that island was a beekeepers’ paradise, where the bees increased and gave surplus with little or no assistance from their owners. Nearly every village contained a number of skep beekeepers whose surplus colonies were sulphured in the autumn or exported as driven bees. With modern methods a steady importation of foreign bees took place. From 1898 to 1904 stocks and queens of foreign races were imported into the district where the disease is supposed to have originated. All the bees in that district were destroyed and the epidemic advanced across the island. Imms (1907) states that he was informed that the disease was so virulent that healthy swarms imported from the mainland were badly diseased within a week1.
Importations of foreign bees have been so heavy and continuous that it is difficult to believe that no diseases have been introduced with these bees during the present century. In reviewing the work that has been done on Isle of AVight disease in Scotland, the possibility suggests itself that at one time
1 The danger from trading and consequent intermixing appears to have been thoroughly grasped by our forefathers who recognised that it was unlucky to trade in bees.
The monks were great beekeepers and their migrations with their bees appear to have brought much bad luck. The traditional ceremonies that have been handed down among peasant , beekeepers are often of pagan origin. The connection between Christianity and lack of honey is well shown in an old German adage given by Langstroth (1S68):
“Bells’ ding-dong and choral song Deter the bee from industry;
But hoot of owl and wolf’s long howl Incite to moil and steady toil.”
G. W. Bullamore 01
what may have been a bacterial disease was being investigated as the Isle of Wight disease1.
Tinsley (1918), in a bulletin issued by the West of Scotland College of Agriculture, states that he succeeded in infecting healthy bees with Isle of Wight disease by feeding them with sugar syrup in which the liquid contents of the intestines of sick bees had been incorporated. In one of Dr Rennie’s earlier experiments (1919 a) Nosema spores in candy were fed to healthy bees in May. Crawling without Nosema was recorded as being present in June and the bees were found dead the following January. In 1915 also, pulped diseased bees were fed in honey to a stock on June 28th. The stock swarmed and both lots showed crawling in October and died out. The nearest bees were two miles away and remained healthy (Anderson and Rennie, 1916).
We assume that the Nosema spores were obtained from sick bees and that bacteria and other organisms were therefore unavoidably present in the candy. The results obtained by Dr Rennie may thus have been due to the organism that was present in the cases recorded by Tinsley. That organism is unknown, but the results suggest that it was situated in the alimentary canal.
V. Conclusions.
Acarine disease appears to be less virulent than the disease which swept across the Isle of Wight in the early years of this century. That the mite was causing damage at the same time is very probable but the investigations were centred on the acute and virulent disease.
It may be that most of the stocks affected with mites, but showing no symptoms of disease, die out sooner or later. But this does not demonstrate the existence of a new disease. It merely emphasises the soundness of the older system of beekeeping which considered it undesirable to retain any stock after the third season, the less desirable colonies being sulphured at an earlier period.
Although it may not be the cause of the Isle of Wight disease the dis¬ covery of the mite is of economic importance, revealing, as it does, one of the causes of the failure of modern beekeeping. Ever since the introduction of the “humane” system which saved the redundant bees and distributed them as “driven bees” throughout the length and breadth of the land, there has been a steady increase in disease which has helped to render the industry of honey production an unprofitable one. Before the rise of the Isle of Wight epidemic the losses were attributed usually to foul brood, although there was
1 The difficulty in classifying bee disease by symptoms is well shown by the following instance.
While the work on Nosema in its relationship to Tsle of Wight disease was being carried out at Cambridge there ensued a heavy mortality of humble-bees which was found to be associated with the presence in the Malpighian tubes of a protozoon closely resembling Nosema. In the year immediately past a similar mortality has been noticed in humble-bees, but the protozoon could not be found. The organism accompanying the mortality in 1921 was a nematode worm Sphaerularia bomhi which undergoes development in the body cavity of the bee and eventuall}' gives rise to huge numbers of larvae. Tn both years, the symptom of the trouble was inability to fly.
02
Bee Disease
much difference of opinion as to the amount of harm caused by this latter malady. It is possible that Tarsonemus was present in the more severe cases and, consequently, a fresh series of observations are necessary.
It is the opinion of the writer that the mite will prove a comparatively harmless parasite in countries where two or more honey harvests and con¬ stant breeding activity are the rule. In some such districts we may expect to find the endemic centres of the disease.
If this opinion is confirmed, the mite will also be found in Australia, America and other countries as it is unlikely that we have been the only country to import it. In America there is a large amount of unexplained paralysis, and of winter and spring losses, while in Australia the subject of bee mortality is much discussed without anything very definite being known. Some of these troubles may be accompanied by mites.
In this country bees parasitised by mites are to be found all over the kingdom from Land’s End to John o’ Groats. The recognition of the dangers arising from modern procedure may eventually give rise to wiser and better methods of beekeeping and the discovery of the Aberdeen investigators may thus have far-reaching and unforeseen consequences.
REFERENCES.
Anderson, J. (1916). The connection of Nosema apis and ;‘Isle of Wight” Disease in Hive Bees. Proc. P. Physical Soc. of Edin. xx, 16.
Anderson, J. and Rennie, J. (1916). Observations and Experiments bearing on Isle of Wight Disease in Hive Bees. Ibid, xx, 23.
Berlepsch, Baron von (1869). Die Biene und Hire Zuchf. Mannheim.
Beuhne, F. R. (1916). Nosema apis in Victoria. Journ. of the Dept, of Agric. of Victoria , xiv,
Bevan, E. (1827). The Honey Bee. London.
Critchlow, B. P. (16. vii. 1904). Gleanings in Bee Culture Medina, Ohio, xxxu, 692. Dadant, 0. (1907). Langstroth on the hive and Honey Bee. Revised by Dadant. Illinois, U.S.A.
Fleming, G. (1871). Animal Plagues. London.
Graham-Smith, G. S., Fantham, H. B., Porter, A., Malden, W. and Bullamore, G. W. (1912). Report on the Isle of Wight Bee Disease (Microsporidiosis). Supplement No. 8, Journ. of the Bd. of Agric. xix.
Hannemann (1909). Cited by Zander (1911).
Hirst, S. (1921). On the mite Acarapis woodi (Rennie) associated with the Isle of Wight Bee Disease. Annals and Mag. of Nat. Hist. Ninth Series, No. 42, 509.
Langstroth, L. L. (1868). On the Hive and Honey Bee. Third Edition. Philadelphia. See also Dadant (1907).
Maassen, A. (1911). Zur Aetiologie und Epidemologie der Ruhr bei den Bienenvolkern.
Mitteilungen a. d. K. Biolog. Anstalt f. Land- und Forstwirtschaft, 50.
Malden, W. (ii. 1909). Further Report on a Disease of Bees in the Isle of Wight. Journ. of the Bd. of Agric. xv, 809.
Nussbaumer, T. (1912). Einige Erfahrungen Uber die Noscmakrankheit. Schiveizerische Bienenzeitung ( Reprint ) .
Rennie, J. and Harvey, E. (1919 a). Isle of Wight Disease in Hive Bees. Journ. of the Scottish Bd. of Agric. n, 176.
- - (1919 b). Nosema apis in Hive Bees. Ibid, n, 511.
Rennie, J., White, P. B. and Harvey, E. (1920). Isle of Wight Disease in Hive Bees. Trans. Boy. Soc. Edin. m, 737.
Rennie, J. (1921). Notes on Acarine Disease. Bee World, m, 1 17.
Silver, J. (1907). Bee Disease in the Isle of Wight. Irish Bee Journal, vii, 10.
Tinsley, J. (1918). Preliminary Report on the Isle of Wight Bee Disease. Bulletin 85 IT.
of Scotland Agric. Coll. Glasgow.
Zander, E. (1911). Handbuch der Bienenkunde.
THREE NEW SPECIES OF TRICHODECTES FROM CEPHALOPHUS MONTICOLA AND PROG AVI A CA PEN SIS FROM SOUTH AFRICA.
By LAURENCE HILL.
(Pietermaritzburg, N atal . )
(With Plate IT.)
• «
Trichodectes bedfordi, n.sp. Plate II, figs. 1-3.
Numerous males and females collected from a Blue Duiker (Ce'phaloyhus (Guebei) monlicola) at Ngome Forest, Mt Ngwibi, Natal, by Mr J. Tustin.
Female. Total length 1-73 mm., length of head 0-37 mm., length of pro¬ thorax 0-09 mm., length of metathorax 0-11 mm., length of abdomen M6 mm. Width of head 0-30 mm., width of prothorax 0-25 mm., width of metathorax 0*30 mm., width of abdomen 0*42 mm.
Head. Longer than broad. Forehead elongated with a moderately deep median notch, which has a deep marginal band of chestnut-brown chitin divided into two parts by a narrow median longitudinal split, and produced into prominent trabecula-like processes in front of each antenna. General colour of head a pale yellowish-brown, with deep chestnut mandibles and bands. Antennal sinuses .rather small and shallow. Ocular projections ex¬ tending only very slightly beyond the temples, which are rounded. Temples with two short widely spaced hairs. Occipital margin slightly convex, rounded at the points where it is met by the faint occipital bands. Antennal bands of medium width, not reaching the base of the antennae, and tapering towards the anterior margin of the head, then broadening out into a wide clear space, interrupted by a deep narrow longitudinal line. Occipital bands joining the antennal bands just in front of the trabecula-like processes. Antenna long and narrow, extending well beyond the posterior margin of the head; third joint the longest; first and second joints of equal length. Eye pronounced. On the dorsal surface of the forehead there are three short pustulated hairs situated in a straight transverse line on each side near the antero- lateral margin of the head.
Thorax. Same colour as the head. Pronotum bare, with sides diverging, and a broad chestnut band at the lateral margins. Metathorax as wide as
04
New Species of Trichodectes
the head, sides diverging, and rounded at the postero-lateral margins, with a broad band at the lateral margins and three small pustulated hairs on each side in a transverse line near the postero-lateral margins.
Abdomen. Elongated and narrow, white in colour, with crenulated lateral margins. Segments widely diverging towards postero-lateral borders, and with a yellowish-brown transverse band on the dorsal surface of each segment. On the dorsal surface, at the posterior margin of the band on first segment, one short pustulated hair on each side of the meson; last segment bilobed at the apex, with three to four median hairs on each lobe, and with a pair of short pustulated hairs situated on each side just within the lateral margin, and another pair mid-way between these towards the posterior indentation'; remaining segments with a transverse row of minute pustulated hairs ; segments two to seven with a small spiracle at each lateral angle.
Male. Total length 1*56 mm., length of head 0-40 mm., length of pro- thorax 0-07 mm., length 'of metathorax 0*14 mm., length of abdomen 0-95 mm. Width of head 0-28 mm., width of prothorax 0-22 mm., width of metathorax 0*26 mm., width of abdomen 0-43 mm.
Head. With the median notch deeper than that of the female. Antennal sinuses deep. Ocular projections small, not well defined. Temples narrow, smoothly rounded. Occipital margin markedly convex, emarginate wThere it is met by the occipital bands. Eye pronounced. Antennae large, and back¬ ward pointing, reaching the metathorax; the first joint large and wide (sacu- lated) longer than the second and third joints combined; second and third joints of approximately equal length; on the upper surface of the first joint of the antenna, a longitudinal row of eight small pustulated hairs one beside the other, and extending from the proximal to the distal extremity; third joint slightly curved, and at the distal end two or three short stout dark- brown denticles. Mandibles situated well forward on the head.
Abdomen. Closely resembles the female in general outline, with the last segment oval, and having a row of medium sized hairs at the tip. On the dorsal surface, the transverse brown bands of segments four to seven are deeper (somewhat telescopic in appearance), and with well marked projecting somewhat rounded edges almost reaching the spiracles at the lateral angles. (Fig. 3 gives an excellent representation of these bands.)
Genitalia. Very conspicuous. The basal plate longer than the parameres, and consisting of two thickly-chitinised bars slightly thickened and rounded at their posterior end. Parameres long and narrow, of even thickness through¬ out their length (concavo-convex in shape), and markedly diverging pos¬ teriorly. Penis long and stout. Beneath the penis a wedge-shaped plate with ‘ the posterior end bifid. (Plate II, fig. 3.)
This species would seem to be closely allied to Tr. lineatus n.sp., de¬ scribed by Bedford in Part II, Anop. from South African Hosts (1920). The shape, however, of the abdominal segments of both male and female, and
L. Hill Ho
terminal segment and genitalia of the male enables this species to be readily listinguished from that described by him.
My thanks are due to Mr G. A. H. Bedford of the Veterinary Research Laboratories, Onderstepoort, Pretoria, for the kindly help and assistance vvhich he has always been most ready to give me, and I have great pleasure n naming this species after him.
Trichodectes lindfieldi n.sp. Plate II, figs. 4-6.
Numerous males and females taken from a Cape Hyrax (Procavia capensis) shot at Mtabamhlope, Estcourt District, Natal, by Mr P. Barnes.
Female. Total length 1-47 mm., length of head 0*43 mm., length of pro- bhorax 0-12 mm., length of metathorax 0-15 mm., length of abdomen 0-77 mm. Width of head across temples 0-42 mm., width of prothorax 0-30 mm., width oi metathorax 0*40 mm., width of abdomen, at third segment 0-71 mm.
Head. Light yellowish-brown. Almost as broad as long. Forehead dome¬ shaped, emarginated in front, with the sides slightly convex, and produced with a prominent trabecula-like process in front of each antenna. Antennal bands narrow and dark, terminating approximately midway between the interior margin of the head and the trabecula-like processes. Frontal sinus deep, semicircular in section, with a marginal band of dark brown chitin divided into two parts by a median longitudinal narrow space; one hair situated on each side just within the emargination. Antennal sinuses small and shallow with rounded inner margin. On each side of the forehead, there are three hairs, one in front, one in the middle, and one above the antennal sinus. Ocular projections prominent, extending well beyond the temples which ire rounded. Occipital margin slightly convex, very slightly emarginate at the points where it is met by the occipital bands. Temples rounded with a narrow dark marginal band and three short marginal hairs. Occipital bands 3onspicuous, connected at their bases by a broad band, and meeting the antennal bands just above the trabecula-like processes. Antennae long and slender, reaching beyond the posterior margin of the head ; the third segment the longest with four minute hairs; first and second segments of equal length each with two short hairs. On the dorsal surface of the forehead there is a semicircular row of six small pustulated hairs ; and below this semicircular row one small hair on either side just within the antero-lateral border; also two small and two medium hairs situated in a line with the trabecula-like processes. On the hind-head a semicircular row of four short hairs in the middle, and two on each temple.
Prothorax. Much broader than long, with a broad marginal band of deep chestnut-brown; lateral margins almost parallel, the posterior margin markedly convex, with medium postero-lateral emarginations, and one short pustulated hair on each side of the meson near the anterior margin.
Mesothorax. With three pustulated hairs on the dorsal surface.
Parasitology xrv
5
New Species of Tricliodectes
Metathorax. Wider than prothorax, with prominent projecting lateral angles reaching nearly as far as the postero-lateral angles of the first abdominal segment. On the dorsal surface two longish hairs at each lateral angle, and a row of fourteen small hairs along the posterior margin. Posterior margin markedly concave.
Abdomen. Crenate, somewhat longer than wide, widest at the third seg¬ ment, with a median dark transverse band on all the segments. On the first three segments there is a large dark brown pleurite on each side, these are succeeded on the remaining segments by a smaller pleurite of the same colour. On the dorsal surface, the first segment has two pairs of small pustulated hairs on each side of the meson near the anterior margin, and twelve hairs in a row along the posterior margin; second segment with a transverse row of twenty small pustulated hairs along the posterior margin; and a lateral group of five hairs. Above, and on the inner side of this lateral group, there is a further distinct separate group of three hairs; third segment with a transverse row of twenty hairs along the posterior margin, a lateral group of five hairs, and on the inner side above each lateral group, a further group of five hairs; fourth segment with a row of twenty-four pustulated hairs, a lateral group of three hairs, and on the inner side of this group a pair of hairs ; fifth segment with a transverse row of twenty-four pustulated hairs, and a pair of hairs just within the lateral margin; sixth segment with a continuous row of thirty- six pustulated hairs; seventh segment with a row of twenty-four hairs; last segment bilobed at the apex, with a transverse row of twelve hairs, and two small and two longish hairs on each lobe.
On the ventral surface segments one and two each with a row of twelve hairs ; segment three with a row of fourteen hairs ; segment four with a trans¬ verse row of thirty- two hairs divided into three groups, the median group consisting of twenty hairs, and each lateral group of six hairs; segment five with a row of thirty hairs, the median group consisting of twenty hairs, and each lateral group of five hairs; segment six with a row of twenty-eight hairs, the median group of twenty hairs, and each lateral group of four hairs; seventh segment with a pair of small pustulated hairs at each lateral margin; last segment with nine to ten minute pustulated hairs scattered on each side of the meson.
Male. Total length 1*52 mm., length of head 0*39 mm., length of pro¬ thorax 0*12 mm., length of meta thorax 0-15 mm., length of abdomen 0-86 mm. Width of head across temples 0*39 mm., width of prothorax 0*28 mm., width of metathorax 0-35 mm., width of abdomen at third segment 0-63 mm.
Head. The general outline of the head mostly resembles that of the ' female. The antennal sinuses are deeper and wider. Ocular projections scarcely visible. Temples bluntly rounded, meeting the occipital margin at an angle. At the posterior margin of the temples there is a small prominent chitinised protuberance (absent in the female). Antennal bands broader than in the
L. Hill
67
female, becoming fused into the occipital bands in front of the trabecula-like processes. Antennae large, and directed backwards, reaching well beyond the occipital margin of the head; the first segment broad, and longer than the second and third combined; second segment longer than the third. On the hind-head a semicircular row of four small pustulated hairs, and three on each temple.
Prothorax. With lateral angles bluntly rounded.
Abdomen. Oval, widest across the third segment. Resembles the female in regard to colour, shape of pleurites, etc. On the dorsal surface, last segment with a semicircular row of twelve pustulated hairs, and two small hairs at the posterior margin.
Genitalia. Conspicuous, the basal plate consisting of two chitinous bars considerably longer than the parameres. Parameres consist of short bands, broad at the base where they are rounded, and narrowing posteriorly, becoming fused at the tip. Penis short and stout. Beneath the penis, an elongated plate with proximal end rounded. (Plate II, fig. 6.)
Trichodectes serrations n.sp. Plate II, figs. 7-9.
Several males and females taken in company with the previous species from a Cape Hyrax (Procavia cajpensis ), shot at Mtabamhlope, Estcourt District, Natal by Mr P. Barnes.
Female. Total length T25 mm., length of head 0*33 mm., length of pro- thorax 0-09 mm., length of metathorax 0-12 mm., length of abdomen 0-71 mm. Width of head 0-33 mm., width of prothorax 0-22 mm., width of metathorax 0*23 mm., width of abdomen 0-49 mm.
Head. Light yellowish-brown, as broad as long. Forehead rounded, with four hairs on each side and a long shallow median notch produced into a medium trabecula-like process in front of each antenna. Antennal bands dark, narrow at their base, broadening out into a deep serrated band of dark brown towards the centre of the forehead. Ocular projections pronounced extending well beyond the temples which are rounded. Occipital margin slightly convex. Temples rounded, with a narrow dark marginal band, and with three short pustulated hairs on each side. Occipital bands narrow, con¬ nected at their bases by a broad, dark brown band, and meeting the antennal bands in front of the trabecula-like processes. Antennae long and narrow, reaching just beyond the posterior margin of the head; third joint the longest, with three small hairs; second joint nearly as long as the first, and each bearing two small pustulated hairs. Eye pronounced. On the dorsal surface ’ of the forehead, there are four small hairs on each side situated just within the antero-lateral margin. On the hind-head a small pustulated hair on each side close to the ocular projection, and a semicircular row of six small hairs near the posterior margin of the head.
Prothorax. More than twice as broad as long, with lateral bands of dark
i) - 2
(58
New Species of Trichodectes
brown; marginal band faint and interrupted in the middle. Posterior margin very slightly convex, becoming deeply curved towards the lateral margins, which are slightly rounded.
Metathorax. Slightly wider, and longer than the prothorax, the lateral margins curved and bordered with a narrow dark band, the posterior margin slightly convex.
Abdomen. Oval in shape, broadest at third segment, whitish in colour, with a small brown transverse band on the dorsal surface of segments 1-6; seventh segment with the transverse band involving the whole of the segment. On the dorsal surface, first segment bare, with a deep marginal band of dark brown; second segment with a median group of four hairs widely spaced, and one pair at each lateral margin ; third segment with twenty pustulated hairs ; fourth segment with a row of sixteen pustulated hairs; fifth segment with a row of fourteen pustulated hairs; sixth segment with a row of fourteen hairs and two short hairs at the angle of the postero-lateral margin; seventh segment with a semicircular row of twelve pustulated hairs; last segment bilobed at the apex, with a row of eight pustulated hairs, and two short hairs on each lobe.
Male. Total length T16 mm., length of head 0*30 mm., length of prothorax 0*11 mm., length of metathorax 0-12 mm., length of abdomen 0-63 mm. Width of head 0-30 mm., width of prothorax 0-21 mm., width of metathorax 0-22 mm., width of abdomen 0*40 mm.
Head. The head generally resembles the female in outline, except that the antennal sinuses are much wider. Ocular projections extremely small. Antennae large and backward pointing, extending well beyond the occipital margin of the head, the first segment large and broad; second segment practically the same length as the first; the third segment the longest. On the dorsal surface of the head, a row of six small pustulated hairs close to the anterior margin, two small hairs situated in the centre of the head, mid-way between the bases of the antennae and the ocular projections, and a group of three to four hairs on the temples.
Prothorax. Produced sharply posteriorly, with a deep emargination at the posterior margin; lateral margins slightly rounded.
Abdomen. Oval, widest across the fourth segment. First segment entirely lacking the deep marginal broad band of the female ; a brown transverse band on the dorsal surface of segments one to five; segments six to eight with an irregularly shaped and fused transverse band; last segment oval, with five to six short hairs along the posterior margin ; first segment with a row of eight hairs; second segment with a row of fourteen hairs; third segment with a * row of sixteen hairs ; fourth segment with an unevenly spaced row of fourteen hairs; fifth and sixth segments with a row of fifteen to seventeen hairs; last segment with a closely set row of ten hairs.
Genitalia. Conspicuous, as shown in Plate II, fig. 9.
THE LIBRARY OF THE
UNIVERSITY Or ILLINOIS
PARASITOLOGY, VOL. XIV. NO. 1
PLATE II
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DESCRIPTION OF PLATE II.
Trichodectes bedfordi n.sp.
Fig. 1. Female.
Fig. 2. Head of male.
Fig. 3. Last abdominal segments and genitalia of male.
Trichodectes lindfieldi n.sp.
Fig. 4. Female.
Fig. 5. Head of male.
Fig. 6. Last abdominal segments and genitalia of male.
Trichodectes serraticus n.sp.
Fig. 7. Female.
Fig. 8. Male.
Fig. 9. Last abdominal segments and genitalia of male.
70
ON THE LARVA AND PUPA OP A PARASITIC PHORID ELY —HYPOCERA IN CRASS AT A MG.
By HUBERT M. MORRIS, M.Sc., F.E.S.
(Entomological Dept., Institute of Plant Pathology, Rothamsted Experimental Station, Harpenden.)
(Plate III and 4 Text-figures.) CONTENTS.
PAGE
1. The Larva ..... 70
2. The Puparium .... 73
3. The Pupa . ... 73
4. Emergence of Adult from Puparium 73
PAGE
5. Comparison with Larvae of other
Phoridae ..... 74
6. References to Literature ... 74
7. Description of Plate ... 74
On Jan. 27th, 1920, among a number of nearly fully grown larvae of Bibio marci which had been kept in the laboratory about two weeks, a few individuals were observed to be unhealthy or dead. These latter were found to contain parasitic Dipterous larvae, which eventually proved to be those of Hypocera incrassata Mg. Certain of these larvae were observed to be leaving the bodies of other larvae of Bibio marci at the same time.
The larvae of Hypocera incrassata pupated in the soil immediately after leaving their hosts, and only a single parasite was observed in each of the latter.
The pupae were kept in soil in the laboratory, and the adults emerged between June 24th and July 1st.
The larva and pupa of Hypocera incrassata appear to be undescribed. It is noteworthy that H. vitripennis is recorded as having been reared from humble- bees’ and wasps’ nests (4), and H. vectabilis from dried specimens of Coleoptera (l).
The present species is of additional interest as being the first definite record of an insect parasitic on Bibionid larvae.
I am indebted to Dr D. Keilin for the identification of the adult fliess and for the information regarding previous records of the rearing of species of Hypocera. I am also much indebted to Dr A. D. Imms for advice during the preparation of this paper.
1. THE LARVA.
The larvae were not observed until they were leaving, or about to leave, the body of the host, and were then fully grown. At this stage they are about 8-0 mm. to 8-5 mm. in length and about 2-0 mm. in breath, and are of a yel¬ lowish-white colour.
H. M. Morris
71
External Form. The larva (Plate III, fig. 1) is composed of a small cephalic region, and eleven body segments, the segmentation being fairly distinct, but somewhat obscured by the presence of supplementary folds.
On the ventral surface of the cephalic region is a small opening, the mouth,
cl
•\
/ \
Text-fig. 1. Cephalic and prothoracic segments of larva of Hypocera incrassata Mg. Latero- ventral view, a., antennal organ; as., anterior spiracle; cl., cephalic lobes; ms., mandibular sclerite. x 32.
Text-fig. 2. Sensory structures on cephalic lobe, a., antennal organ; b., club-shaped projecting structure; c., group of papillae, x 670.
Text-fig. 3. Anterior spiracle of larva of Hypocera incrassata Mg. x 670.
Text-fig. 4. Posterior spiracle of larva of Hypocera incrassata Mg. x 670.
from which projects the single mandibular sclerite (Text-fig. 1). Dorsal to the mouth are a pair of hemispherical lobes, bearing sensory structures. The most conspicuous of these structures is the antennal organ, which consists of two segments, the basal segment of which is cylindrical, and the distal segment dome- or bell-shaped (Text-fig. 2 a).
72
A Parasitic Phorid Fly
Close to tlie antennal organ, but rather ventral to it, is a minute slightly curved and somewhat club-shaped projecting organ; near the latter there is also a small group of minute papillae (Text-fig. 2 b and c).
The first segment of the body is small and rather conical, and near its posterior margin is a pair of projecting lateral spiracles which are brown in colour. Dorsally this segment bears a slight median transverse depression.
The remaining segments, except the last, are cylindrical, and increase in size to the sixth, after which they are approximately equal in size. The second and third segments are each divided dorsally into two parts by a median trans¬ verse sulcus. On each of the following segments, from the fourth to the ninth, there are three similar sulci. The second and third of the regions so formed in each segment are less clearly defined than the first and fourth, and it may be added that the third and fourth regions increase slightly in prominence on the posterior segments.
The tenth segment is divided, dorsally, into three regions, of which the third is the most conspicuous.
The terminal segment is in the form of a blunt cone, and is divided dorsally into three regions, of which the first is the largest and bears the pair of small posterior spiracles laterally. The terminal portion of this segment is rather flattened.
\ entrally, the first segment is undivided ; the second segment is very faintly divided, and the third segment more distinctly divided into two regions. The fourth to tenth segments are divided into three regions, of which the first is very slightly the more prominent.
The terminal segment is similarly divided into three parts, and bears the conspicuous aperture of the anus on the foremost division. On each side of the anus is a rather prominent rounded fleshy process.
Buccopharyngeal Armature (Plate III, figs. 4 and 5). The buccopharyngeal armature is divided into three parts: (1) the pharyngeal or basal sclerite, (2) the intermediate sclerites, and (3) the mandibular sclerite.
(1) The pharyngeal sclerite is prolonged posteriorly (dorsally and ventrally) as a very delicate chitinous structure, the dorsal prolongation being much the longer. Its anterior end is strongly chitinised and bears on each side a hollow for the articulation of the intermediate sclerite.
(2) The intermediate sclerites are stout and strongly chitinised and are not fused together. They articulate with the hollows at the anterior end of the pharyngeal sclerite by means of a hook-like prolongation at their posterior end.
(3) The mandibular sclerite is a single very stout structure. It articulates with the intermediate sclerites by means of a pair of backwardly directed • divergent arms at its posterior end. Ventrally, near the base, it bears several slight transverse ridges suggestive of vestigeal teeth.
Articulating with a slight projection at the anterior external angle of either intermediate sclerite is a delicate process, more strongly chitinised at its base,
73
H. M. Morris
and having the form shown in Plate III, fig. 6. At its distal extremity each process bears several projecting spines. These processes bear some resemblance to those described by Keilin(3) as Baguettes orales ’ (oral rods) in carniv orous Anthomyid larvae.
Tracheal System. The tracheal system is amphipneustic, a pair of spiracles being situated on both the first and last segments of the body.
Both pairs of spiracles are very alike in size and structure and are brown, chitinised outgrowths from the body-wall: each spiracle communicates with the exterior by means of four oval apertures situated at its apex (Text-figs. 3 and 4).
2. THE PUPARIUM.
The puparium is brown, with the segments fairly clearly marked. It is about 5-0 mm. in length.
Dorsally there are several conspicuous constrictions. The prothoracic spiracular horns of the pupa emerge from the puparium through a pair of holes situated in a somewhat lateral position on its dorsal side, at the posterior end of the fourth body segment of the larva (Plate III, fig. 2).
These horns are about 0*23 mm. in length, and their distal half bears two rows of minute openings, about sixteen openings in each row. Both pairs of larval spiracles remain as conspicuous dark projections from the puparium, and the mandibular sclerite also projects at the anterior extremity.
3. THE PUPA.
The pupa, seen on removal from the puparium is white, with the ap¬ pendages of the adult fly closely adpressed to the body. Its general characters can be seen on referring to Plate III, fig. 3, and scarcely need detailed description.
The head is of moderate size, and the antennae are large and are produced in a long slender process. The palpi are conspicuous and conical in shape. The labrum is short and flattened. The legs lie side by side, the tarsi of the posterior pair projecting slightly beyond the apex of the abdomen.
4. EMERGENCE OF ADULT FROM PUPARIUM.
When the adult fly emerges from the puparium, the latter is split in the following way. A circular cap, consisting of the skin of the cephalic region and thoracic segments of the larva, is split off by means of a fissure passing round the body between the third thoracic and first abdominal segments. A quadrangular plate is split off in addition, and it consists of the whole skin of the dorsal surface of the first three abdominal segments.
This plate may become completely detached, or may remain slightly ad¬ hering at its posterior end to the anterior margin of the fourth abdominal segment.
74
A Parasitic Phorid Fly
5. COMPARISON WITH LARVAE OF OTHER PHORIDAE.
The larva of Hyjpocera incrassata differs considerably from the larvae of Phora Bergenstammi Mik., P. rufijpes Mg., and P. ruftcornis Mg., as described by Keilin(2).
The buccopharyngeal armature is very different as may be seen by com¬ parison with Keilin’s figures. The numerous sensory structures on the cuticle of the latter larvae are not present in the case of Hyjpocera incrassata.
The absence of sensory structures may be an adaptation to a more com¬ pletely parasitic existence, as may also be the simpler buccopharyngeal armature, with the fusion of the usual two mandibular sclerites into an unpaired organ.
The study of this larva lends support to the opinions of Brues, de Meijere and Keilin that the position of the Phoridae in the classification of the Diptera should be among the Cyclorrhapha.
REFERENCES.
(1) Brues, C. T. (1913). A New Species of Phoridae reared from dried Coleoptera. Ann. Mus. Nation. Hung. xi.
(2) Keilin, D. (1911). Recherches sur la morphologic larvaire des Dipteres du genre Pliora. Bull. Sci. France el Belg. 7me serie, T. xliv, i.
(3) Keilin, D. (1917). Recherches sur les Anthomyides a larves carnivores. Parasitology , ix, 3.
(4) Wood, J. H. (1906). On the British Species of Phora (Part I). Ent. Mon. Mag. xvn.
DESCRIPTION OF PLATE III.
Larval and Pupal Stages of Hypocera incrassata Mg.
Fig. 1. Fully grown larva. Dorsal view, as., anterior spiracle; ps., posterior spiracle, x 20.
Fig. 2. Puparium. Lateral view. Is., jirotruding spiracular horn of pupa, x 20.
Fig. 3. Pupa removed from puparium. Lateral view, s., prothoracic spiracular horn, x 20. Fig. 4. Buccopharyngeal armature of larva. Lateral view. i. , intermediate sclerite ; m. , mandibular sclerite; o., “baguette orale”; p., pharyngeal sclerite. x 126.
Fig. 5. Buccopharyngeal armature of larva. Ventral view. Lettering as in previous figure, x 126. Fig. 6. “Baguette orale” of larva x 730.
PARASITOLOGY, VOL XIV. NO. 1
PLATE
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Lvv.-S.
//. M. Morris del.
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75
THE MALLOPHAGAN FAMILY TUI M EN OP ON I DAE.
By G. F. FERRIS, M.A.,
Stanford University , California.
(With 8 Text-figures.)
Three species of two-clawed Mallophaga have heretofore been recorded from South American mammals. Concerning one of these species, Menopon ex- traneum Piaget, nothing is known aside from the information included in the original description and, judging from this and the accompanying figure, it appears not to differ from the usual type of bird-infesting members of this group. The other two species have each been made the type of a genus and these two genera have been considered by Harrison as representing a well- marked family, the Trimenoponidae.
Through the courtesy of the authorities of the United States National Museum and the Field Columbian Museum of Chicago, the present writer has been accorded the privilege of examining for parasites the mammal skins in the collections of these two institutions. Among the wealth of material thus discovered there have appeared three new species from South American mammals that are also referable to the family Trimenoponidae. In connection with the description of these three species, which necessitates the description of two new genera also, it has seemed desirable to review the family as a whole.
The tvpes of the three new species are deposited in the collections of Stanford University. Paratypes of two of them, as indicated below, are deposited in the collections of the United States National Museum.
In the accompanying figures the left half