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NOVITATES ZOOLOGICAE.
Wom, XQ0IL, I@sis—39).
NOVITATES ZOOLOGICAE.
MH Fournal of Zoology
IN CONNECTION WITH THE TRING MUSEUM.
EDITED BY
THE HON. MIRIAM ROTHSCHILD
AND DR. K. JORDAN, F.R.S.
Vor. XLI, 1938-39.
(WITH SIX PLATES.)
IssuED AT THE ZooLocicaL MusEUM, Trina.
PRINTED BY HAZELL, WATSON & VINEY, Lrp., LONDON AND AYLESBURY
1938-1939
CONTENTS OF VOLUME XLI (1988-39).
GENERAL SUBJECT.
See Siphonaptera No. 1.
MAMMALIA.
1. New or little-known Marsupials: A new species of Phascogalinae, with
notes upon Acrobates pulchellus Rothschild. G. H. H. Tate .
2. Some Arabian Mammals collected by Mr. H. St.J. B. et C.I.E. (Plate
VI). T.C. S. MORRISON-SCOTT . 5 :
3. The Races of Jaguar (Panthera onca). R. I. Boca
TRICHOPTERA.
1. The Brazilian Hydroptilidae (Trichoptera). Martin E. MosELy
LEPIDOPTERA.
1. Note on Polygonia c-album (L.) 1758, at Ashton Wold, Northamptonshire,
and the record of a notable aberration. MIRIAM ROTHSCHILD and MALCOLM
SPOONER .
2. On the remarkable variation of an Oriental Hawkmoth (Lepidopter.
Kart JoRDAN (Plate III, part)
3. On some unfigured Lepidoptera from the Oxon and Bihiopian an
(Plates III and IV). KARL JoRDAN
4. Zur Nomenklatur der beiden deutschen en mit en Fühlern.
KARL JORDAN .
On some Old-World Nee (este). KARL JORDAN.
6. On the Constancy and Variability of the Differences between the Old- World
species of Utetheisa (Lepid.: Arctiidae). KARL JORDAN
7. Notes on Oriental Theclinae Gee eed with descriptions of new
species. N. D. Rırry : é
8. On some Old-World Lepidoptera eto: KARL JORDAN
COLEOPTERA.
1. On some Old-World Anthribidae. KARL JORDAN a :
2. Descriptions of Anthribidae, mostly from Fiji. Kart JoRDAN .
3. On some Anthribidae from Africa and the Solomon Islands. KARL JORDAN
SIPHONAPTERA.
1. Where Subspecies meet. KARL JORDAN
2. On some Siphonaptera from Tropical Africa and Iraq. KARL JoRDAN
3. On some Nearctic Siphonaptera. KARL JORDAN
4
On the species of Bird-Parapsylli from the ‘tills, obtained on the
British Graham Land Expedition. KARL JORDAN 6 0 :
On some Neotropical Siphonaptera. KARL JoRDAN. : 5
On five new Siphonaptera from the Republic of Argentina. Kari JoRDAN
On some Nearctic Fleas. KARL JoRDAN 0 © 5
Siphonaptera collected by Dr. Gerd Heinrich in Be KARL JORDAN
On Rhopalopsyllus Baker 1905 (Siphonaptera). KARL JORDAN
sae also Mallophaga, No. 1.
senon
Vv
PAGES
58—60
181—211
406—422
217—239
125
126—129
130—133
212—216
248—250
251—291
355—361
433—436
140—147
423—432
437—442
103—111
112—118
119—124
134—139
164—169
292—303
.316—320
362— 375
443—448
vi CONTENTS OF VOLUME XLI (1938-39).
ORTHOPTERA.
1. A preliminary Revision of the Palaearctic species and subspecies of Thisoi- PAGES
cetrus Br. W. (Orthoptera, Acrididae). B. P. Uvarov 4 dj . 377—382
MALLOPHAGA.
1. Eetoparasites from captive Birds. THERESA CLAY AND MIRIAM RoTHScHILD 61—73
2. A note on some Mallophagan names. THERESA CLAY : 0 lee
3. Ectoparasites from captive Birds. THERESA Cray and Mae ROTHSCHILD 305—315
MOLLUSCA.
1. Some observations on the growth of Peringia ulvae (Pennant) 1777 in the
laboratory. ANNE ROTHSCHILD and Miriam ROTHSCHILD . 6 . 240—247
See also Trematoda, No. 3, Peringia ulvae.
POLYCHAETA.
1. Polychaeta of the “ Rosaura ’” Expedition. ©. C. A. Munro . . . 345—354
2. On some tropical Polychaeta in the British Museum, mostly collected by
Dr. C. Crossland at Zanzibar, Tahiti and the Marquesas. C.C. A. Monro 383—393
3. On some tropical Polychaeta in the British Museum, mostly collected by
Dr. C. Crossland at Zanzibar, Tahiti and the Marquesas. C.C. A. Monro 394—405
POLYZOA.
1. Notes on some cellularine Polyzoa (Bryozoa). Anna B. Hasrtıncs . . 321—344
TREMATODA.
1. Cercaria sinitzini n.sp., a cystophorous Cercaria from Peringia ulvae
(Pennant) 1777. Mirtam ROTHSCHILD . 0 5 5 6 : 42—57
2. Notes on the classification of Cercariae of the Superfamily Notocotyloidea
(Trematoda), with special reference to the excretory system. MIRIAM
ROTHSCHILD . : 6 6 © 0 ® 0 : 6 . 75—83
3. Further observations on the effect of Trematode parasites on Peringia ulvae
(Pennant) 1777 (Plates I and II). Miırıam RotuscuHinp . 0 5 84—102
4. The excretory system of Cercaria coronanda n.sp. together with notes on
its life-history and the classification of Cercariae of the Superfamily
Opisthorchioidea Vogel 1934 (Trematoda). Mrriam ROTHSCHILD . . 148—163
5. A note on the fin-folds of Cercariae of the Superfamily Spahr n
Vogel 1934 (Trematoda). MırIam ROoTHScHILD . 9 . 170—173
6. A note on the life cycle of Cryptocotyle aes eee 1825 (Premateda)
(Plate V). Mırıam RoTHSCHILD 4 . 178—180
7. Large and small flame cells in a Cercaria nee Mrsrhar Rorasgmn 376
NOMENCLATURE.
1. Notice & De suspension of the Rules of Nomenclature in certain cases
[A. (n.s.) 1]. Francıs HEMmMInG 6 6 6 : 5 5 0 304
OBITUARY.
1. In memory of Lord Rothschild, Ph.D., F.R.S., J.P. KARL JORDAN . 0 1—41
INDEX . 9 6 0 . 6 5 . ö ¢ . 3 . 449—463
PLATES IN VOLUME XLI.
PLATES I and II.
PLATES III and IV.
PLATE V.
PLATE VI.
Peringia ulvae.
Lepidoptera.
Cryptocotyle lingua.
Heads of Arabian Gazelles.
vii
In ein a EEl Sin on on SET SEE one tne) ins) ln elle)
ERRATA.
. 67, line 8 from above, read METAPIANA for METAPOPIANA.
. 71, line 6 from below, read (Linn.) 1766 for (Linn.) 1866.
. 73, line 8 from above, read Myrsidea for Myrsides.
. 115, line 9 from above, read Xenopsylla for Xenopsyllus.
. 125, line 3 from below, read possibly for possible.
. 154, line 1 from below, insert ) after Spuler.
. 155, line 4 of footnote 1, delete “ is.”
. 168, line 19 from above, add sp.nov. after 4. A. dilecta.
. 253, line 5 from above, read small for elongate.
. 270, line 18 from above, delete full stop after Heyn.
. 276, line 2 from below, add (type 3) after Diré Daoua.
. 279, line 10 from above, read PULCHELLOIDES for PULCHELLA.
. 313, line 28 from above, read COSSYPHA for COSSYHA,
Vili
Nr,
TR SITE m mS Ss
ne
NOVITATES ZOOLOGICAR.
MH Journal of Zoology.
BDITED BY
THE HONOURABLE MIRIAM ROTHSCHILD
AND Dr. KARL JORDAN, E.R.S.
Vor. XLI.
"No. 1.
Pages 1-73.
Issuep May 1SrH, 1938, ar tHE ZootocicaL MUSEUM, Trine.
PRINTED BY HAZELL, WATSON & VINEY, Lrp., LONDON AND AYLESBURY.
1938.
3.
. IN MEMORY OF LORD ROTHSCHILD . i . Karl Jordan
Vou. XLI.
NOVITATES ZOOLOGICAE
EDITED BY
THE HON. MIRIAM ROTHSCHILD and KARL JORDAN.
CONTENTS OF NO. I
PAGES
1-41
CERCARIA SINITZINI N.SP., A CYSTOPHOROUS
CEROARIA FROM PERINGIA ULV AH (PENNANT)
1777
Miriam Rothschild 42-57.
NEW OR LITTLE-KNOWN MARSUPIALS: A NEW
SPECIES OF PHASCOGALINAH, WITH NOTES
UPON ACROBATES PULCHELLUS ROTHSCHILD
1937
G. A. H. Tate 58-60
. ECTOPARASITES FROM CAPTIVE BIRDS. I. 1936-
Theresa Clay and Miriam Rothschild 61-73
NOVITATES ZOOLOGICAE
as) MAY 1038.
IN MEMORY OF LORD ROTHSCHILD, Pu.D., F.R.S., J.P.
BORN THE 8TH FEBRUARY, 1868, DIED THE 27TH AUGUST, 1937.
By DR. KARL JORDAN, F.R.S.
(With 13 photographs.)
IONEL WALTER, second Baron Roth-
schild of Tring, will always occupy a
place of honour in the history of Zoology.
The collections contained in the Museum he
founded and maintained are the largest ever
assembled by one man, and are in many
orders of animals unrivalled even by National
Museums, a fit monument to an enthusiastic
Zoologist. But he acquired still greater merit
by generously placing the contents of the
Museum at the service of Science. Numerous
letters of condolence from biologists testify
to the high esteem in which the deceased was
held as a man and scientist and to the deep
gratitude of the many specialists who have
profited by the Tring Museum in their
researches. Lord Rothschild was probably
the last non-professional systematist who
Kenia. amassed large collections in more than one
class of animals. Love of animals being the
original motive for the foundation of the 5 ;
Tring Museum, and the study of problems of
evolution its ultimate aim, one must expect
to find exemplified in it the various stages or
phases we observe in the development of
Zoology during the last hundred years. The
animals which lend themselves to easy pre-
servation in a dry state, such as insects,
shells, and skins of mammals and birds, were
in post-Linnean times collected and classified
largely by amateur systematists, the public
Museums of Natural History being as yet
in their early youth and the professional
zoologists of the Universities concentrating
almost exclusively on the classes of animals
which had to be preserved in liquid. In our
days the amateur systematist has to restrict
his researches carried out at home to groups
of manageable proportions, the National
Museums with their much enlarged staffs Ace 10.
1
un 16 1938
2 NovitatEes ZooLoGIcAE XLI. 1938.
embrace the whole sphere of systematic
Zoology, and the University has almost
entirely abandoned morphology and
systematics for the experimental study of
life. The next phase will bring an adjust-
ment when it is realized that the true
understanding of life depends on the
knowledge of all its aspects. In the Tring
Museum there are the series of different
species as the primary object of systematic
research, long series of specimens of the
same species for the study of variation,
distribution, and descent, and intersexes,
gynandromorphs, hybrids, and mutants as
illustrations of the modern subjects of
zoological investigations.
As with most private and public
collections, the beginning of what has
grown into an important Museum was haphazard. Like so many boys, the
Honourable Lionel Walter Rothschild, the eldest of the three children of the first
Lord Rothschild, head of the famous banking house of N. M. Rothschild and
Sons, interested himself in Coleoptera and Lepidoptera, beginning to collect at
the early age of seven, according to his own recollection. But unlike other
youths, he did not forsake his childhood’s love in favour of attractions in other
spheres of life, his devotion to natural history collections remaining with him to
the end. In the mind of the general public the name of Rothschild is so inti-
mately connected with finance that other activities of any member of the family
are quite overshadowed. Yet Walter Rothschild was no mutant. It is not
surprising, indeed, that in a family which discovered and developed national
merües wien is a very i im ortant part of the life of a nation and may be
: : claimed and acclaimed by scientists as a
branch of biology—there is a strong trend
towards natural history, manifesting itself
in various ways in different members.
Walter Rothschild’s father was keen on
botany and kept an aviary for his own
enjoyment and that of his children, all
three of whom remained equally fond of
animals. Walter’s sister breeds horses and
other domestic animals and keeps various
mammals as pets; and his brother, the
late N. Charles Rothschild, who collected
insects, acquired fame through his re-
searches in Siphonaptera, an order of blood-
sucking insects of vital importance in the
investigation of the transmission of tropical
diseases, and was the founder of the Society
for the Promotion of Nature Reserves.
AcE 20. One cousin has very extensive collections
AGE 17.
NOVITATES ZooLOGICAE XLI.
Acr 21.
1938.
3
of live plants, and another is a physician
of repute and owner of a children’s
hospital. Miriam Rothschild, eldest
daughter of Charles, makes an intensive
study of parasitic worms, and her brother,
Victor, the third Lord Rothschild, devotes
himself to Biophysics and Cytology. In
Walter Rothschild’s life, however, the
dominance of natural history was extreme,
his active interest in so many different
branches of it being inimical to wise
restriction.
He was a delicate child who could
not be exposed to the rough and tumble
of school life, and therefore was educated
entirely at home under governess and
tutor. Admiration for the intelligent
boy and early flattery were not missing,
and accustomed him to regard himself as the centre of his world and to expect
the fulfilment of his boyish wishes as a natural corollary of his important position.
Shy by nature, he became unduly self-centred as he grew up and averse to asking
advice. He had ample opportunities in London for indulging in his hobby by
buying specimens from natural history dealers, the collections gradually getting
too large for the schoolroom and being then stored in a spare bedroom at the
back of the house; these collections con-
sisted chiefly of insects, with the addition
of a few mounted mammals and birds.
The young, slender naturalist had the
great advantage of an early acquaintance
with a friend of the family, Dr. Albert
Giinther, the Keeper of Zoology in the
British Museum, with whom he remained
intimate until Dr. Günther’s death. Visits
to the Zoological Gardens and the British
Museum tended to enhance his interest,
and the admiration for all the strange
creatures he saw perhaps created in his
subconsciousness the ambition to possess
one day similar collections of his very own.
Anything large made a deep impression
onhim. His predilection for Ratite birds
and Giant Tortoises, and his pride in
having record horns and fishes in his
Museum, exemplify this trait. The last
insects he bought were some Atiacus
caesar, among which there was one speci-
-men larger than any he had of that
species in his collection. As a boy and
youth he was an assiduous field-collector
4 Novirates ZooLoGIcAE XLI. 1938.
of insects and skilful in setting even very
small specimens, a skill he lost in later
years. He never learnt to skin a mammal
or bird.
In order to widen his views and to
perfect his German—in which language
he became quite proficient, speaking it
fluently, though the grammar was some-
times too much for him—he went with a
Mr. Althaus as a companion to the
University of Bonn, and remained there
two terms, from the autumn of 1886 to
July 1887, enjoying the freedom of the
life of the ‘“ Studenten.” He bought here
a collection of German Lepidoptera, now
incorporated in the general collection
with the exception of the Micros, which
are still in the original cabinet. Little
is left of his own collecting during his stay abroad, but as a reminder of his
student days at Bonn, there were among the wall decorations of his sitting-
room two long pipes such as were much in fashion in Germany during the nine-
teenth century among students, schoolmasters, and parsons. He was proud of
them, though he was no smoker, unlike his father and brother, who enjoyed
strong cigars. Shortly after his return to England, he went up to Magdalene
College, Cambridge, for nearly two years (till June 1889), and came in closer
contact with scientists who were in sympathy with his own predilections,
particularly Professor Alfred Newton, the famous ornithologist. From this time
his life as a Zoologist for the fifty years till his death is the history of the creation
and growth of his Museum. The amateur collector who derived pleasure from
collecting turned into a naturalist whose collections had the object of increasing
our knowledge of nature. His acquaintance with Sir Walter Buller, from whom
he acquired a collection of New Zealand birds, drew his attention to the danger
of extinction threatening many mammals and birds by the spread of the European
population into the remote corners of the earth, a question which remained of
great interest to him all his life, the measures of protection of the fauna and flora
by the creation of Nature Reserves and National Trusts having his whole-hearted
support. He had at Cambridge several dozen live Kiwis, a number of which he
kept later at Tring. With the broadening of his scientific horizon at Cambridge,
his zoological aims became ambitious, embracing live mammals and birds as well
as Museum collections. Besides the aviary of his father, there were already
Kangaroos of various kinds and Emus and Rheas in Tring Park, and now enclo-
sures were built in a paddock for a small number of mammals and Cassowaries,
the beginning of a zoological gardens. This ambitious undertaking, however,
had to be given up later on; the specimens were disposed of, and the wild
population in the Park reduced to Rheas and Emus. His large Zoological
Gardens occasionally referred to in newspapers never existed. The collections
of skins and insects had already so much increased while the Hon. Walter
Rothschild was still at Cambridge that they had to be stored at Tring in rented
rooms and sheds, and it became obvious that adequate premises had to be
AGE 34.
NOVITATES ZooLOGIOAE XLI. 1938. 5
a 2
4 a. ae ee We }
wavs alee
In THE ZooLoGicaAL GARDENS, Lonpon. 1903.
provided in which they were not exposed to deterioration. His father gave him
a piece of ground on the outskirts of Tring Park. Here he built a cottage for
the insects and books and a smaller one attached to it for a caretaker; at the
same time a larger building was erected behind the cottage and connected with
it, destined for the display of mounted specimens of all classes of the animal
kingdom. The cottages were ready for occupation in 1889, when Walter Roth-
schild came of age, but the Museum took some time to arrange and the building
was not opened to the public until August 1892. Whoever advised him in the
making of the plans for this public Museum forgot that all the specimens exhibited
should be plainly visible. However, there was one advantage in making the
glass cases too high : Walter Rothschild had evidently learnt from Dr. Giinther
that it is essential for the good preservation of specimens not to expose them to
direct sunlight, and there is certainly no danger that the colours of any of the
animals and birds exhibited in these cases
will fade. Moreover, the preference he always
showed for anything big may have influenced
the construction of these high cases. Walter
Rothschild recognized in later years that they
were excellent for storing large numbers of
specimens, but that they did not come up
to his expectations as regards display. To
modify the building for the requirements of
a modern exhibition gallery will be a difficult
task.
After leaving the University, Walter
Rothschild, following tradition, entered the
firm of Messrs. N. M. Rothschild and Sons
with the object of studying finance under the
tuition of his father, and found but little
Aem time for thorough supervision of the ever-
6 NoviratEs ZootocioaE XLI. 1938.
increasing collections. The mounted vertebrates in the public gallery were
quite safe, being well looked after by the caretaker, Mr. A. Minall; but the
insects, for which one or two attendants had been engaged, were more exposed to
damage on the shelves and in the corners where the boxes were piled up. When
in 1891-92 he purchased the collection of Lepidoptera and Coleoptera of the
Burgomaster of Vienna, Dr. Cajetan Felder, containing some thousand types of
Lepidoptera, the accumulation of material became somewhat chaotic, and Dr.
A. Günther urgently advised his young friend to put a reliable zoologist as Curator
in charge, recommending Mr. Ernst Hartert, who was personally known to Dr.
Giinther and Walter Rothschild. Hartert, who was collecting bird skins at that
time on the Dutch islands off the coast of Venezuela for Count Berlepsch and
Walter Rothschild, accepted the offer of the appointment and came to Tring in
October 1892. The writer of these lines had met Hartert on several occasions
at the house of Count Berlepsch in Hannoversch Münden, and after some corre-
spondence and a visit to Tring agreed to take over the curatorship of Invertebrates
from April 1893.
The collections were already of great size and considerable scientific value
in 1893. However, there was much material of inferior quality, and some of it
of little use because the data necessary for research were not preserved. One
can hardly blame a young and enthusiastic collector with many interests for
having accepted such material ; for in those days the specimens and their names
were the main thing for the average amateur and quite acceptable, be the locality
and other data ever so vague. In the 80’s of the last century it had not yet
entered the mind of every collector that exactness of data is a primary require-
ment for a scientific collection. For instance, when a Continental insect dealer
was asked at Tring in the 90’s to give a precise locality for the Lepidoptera that
had been selected, he was very angry that “ New Guinea ” was not considered
good enough for Tring. There are some old bills for Lepidoptera supplied in
which the localities are given as S. Amerika, Afrika, Indien, the specimens
themselves bearing no labels on the pins with additional information ! Some
of this old material is being kept as an
illustration of the history of the pro-
gressive change in amateur collecting.
Nobody at school or at the University
knew, or troubled to tell a budding
naturalist, that the specimens collected
must be properly labelled. Some of the
early mounted Vertebrates at Tring also
have only general data or none.
It was the first duty of the new
Curators to see to the safety of the
collections. The cottage and the rooms
at the back of the Museum were not large
enough to take all the accumulated
material. Many Vertebrates were stored
in the Victoria Hall (at Tring) and in
sheds near the Museum; the Felder
collections of Lepidoptera and Coleoptera
were housed in the Unity Hall, and here IANGHEEO!
Novirates ZooLoGIcAE XLI. 1938. 7
were also piled up numerous boxes con-
taining mounted and unmounted insects,
some of the boxes being so badly infested
with Dermestid beetles that they had to
be burned. The collections were soon
brought into some sort of order, so that
time became available for research. It is
of interest to quote here some statements
from “Notes and Details about the Tring
Museum” which accompanied the pros-
pectus of NOVITATES ZOOLOGICAE issued
in 1893:
“The Museum contains two distinct
departments, namely :
“I. The Public Galleries, which
contain at present about 950 stuffed
Nees mammals, 3,600 stuffed birds; about
200 reptiles, stuffed and in spirit ;
about 300 fishes, stuffed and in spirit; about 1,500 insects, crustaceans
and arachnidae, of the most typical and representative forms,
mounted ; it
“II. The Students’ Department, which is entirely devoted to ornithology,
coleoptera, and lepidoptera. Here the collections are much more
considerable. Of Birds there are at present about 40,000 skins of over
7,000 species; of Beetles, about 350,000 specimens of more than 60,000
species ; of Lepidoptera the collection
contains about 300,000 specimens of
nearly 25,000 species.”
The scientific staff consisted of Walter
Rothschild, Ernst Hartert, and Karl
Jordan; and, in addition, there was
William Warren, who had already been
engaged for a while at Tring in studying
and arranging the Geometridae (Warren
was subsequently more or less regularly
employed by the Museum until his death
in 1915). A caretaker, Mr. A. Minall,
and his assistant, Mr. F. Young, saw to
the cleaning and heating of the premises,
and an odd-job man looked after the
live mammals and birds in the paddock
and park. In the summer of 1893, Mr.
Arthur Goodson was appointed assistant
to the Curators, helping them to label
and sort the specimens. The cottage
where the work in systematics was carried
2 Dr. E. Hartert, Lorp ROTHSCHILD,
out was full of cabinets, books, and boxes, Dr. K. Jorpan, AMSTERDAM, 1930.
8 Novirates Zootoaicaz XLI. 1938.
and there were only two desks
with good light available for the
three scientists ; the junior Curator
had to find a place at a corridor
window when he wished to use a
compound lens or the microscope.
The division of work had already
been planned when Karl Jordan
joined the Museum. Ernst Hartert
was to be in charge of the birds
and the general management of
the Museum ; Karl Jordan to name
and arrange the Coleoptera ; and
Walter Rothschild to attend to
the Lepidoptera and the Public
Galleries, reserving for his own
studies any group of Vertebrates
or Invertebrates that might speci-
ally interest him. In 1893 it was
further agreed to encourage outside AcE 68.
help as much as possible. The
number of new species in the collections being already very large, and interesting
material requiring recording extensive, it became at once apparent that the
manuscripts would soon be much too numerous for the existing periodicals to
accept for reasonably quick publication, and therefore Dr. Giinther, a frequent
visitor to Tring Park, suggested that the Museum should issue its own periodical,
publishing manuscripts which were entirely, or for the greater part, based on
material in the Tring Museum, the editors to depart from this rule only in very
exceptional cases. The advice was accepted, and Vol. I of NOVITATES ZOOLOGICAE
accordingly issued in 1894, appearing in five numbers containing contributions
by nine authors,
Collections had generally been bought by Walter Rothschild as and when
they were offered. But when during his Cambridge days Professor Newton had
introduced him to Ornithology as a most interesting and suitable subject of
research for a gentleman of means, and the foundation was laid of the Ornitho-
logical collection of the Tring Museum by the purchase of New Zealand birds
from Sir Walter Buller, Walter Rothschild’s interest was so thoroughly roused
that he sent Henry Palmer, a sailor who could skin birds, to the Chatham Islands
with the sole object of collecting all the species of birds occurring on the islands.
The description of a new Pigeon, in 1891, from these islands was Walter Roth-
schild’s first contribution to the literature of Zoology (and his last notes, in 1937,
were on Cassowaries). It was again Newton who drew Walter Rothschild’s
attention to the Sandwich Islands by showing him some of the remarkable birds
sent home by Scott Wilson. Seeing such birds, and no doubt listening to the
professor’s exposition of the variation of the birds from island to island, was
enough to decide Walter Rothschild on an ornithological exploration of the whole
group of islands, and he instructed Palmer to proceed to Hawaii when he had
finished his work on the Chatham Islands. Palmer stayed on the Sandwich
Islands from 1890 to 1893, and collected with great success, the account of his
Novirares ZoonoaicaArn XLI. 1938. 9
collecting being published in Avifawna of Laysan, issued in three volumes (1893-
1900), the first important work written by Walter Rothschild.
From 1893 onward the Museum entered into direct relations with many
explorers, professional collectors, and residents in tropical countries so as to
obtain material from places as yet little Known. In most cases the collectors
engaged by the Museum went abroad at their own risk and expense, often a
small sum being advanced for the payment of the initial costs, the Museum
being bound by contract to take a certain number of specimens of each species
of birds and Lepidoptera at an agreed fair price and having the right to the first
offer of specimens of other orders and classes which might be required. Such
arrangements worked very well, being advantageous to both contracting parties,
and the Museum remained always in friendly relations with Ansorge, Doherty,
Beck, Eichhorn, Everett, Hoffmann, Hose, Klages, Kühn, Meek, and a host of
other explorers whose collections came to Tring.
x Although a rapid worker endowed with a remarkably retentive memory and
a keen eye for differences, great gifts for a systematist, Walter Rothschild soon
found the time at his disposal too short for the many tasks he had set himself,
and already in 1894 the Coleopterist was asked to come to his assistance. The
first result of this collaboration was the publication in 1895 of the Revision of
the Papilios of the Eastern Hemisphere exclusive of Africa, in which for the first
time in Lepidopterology geographical races described by various authors as
distinct species were almost consistently (as much as the material at hand
warranted) reduced to the rank of subspecies and the classification thereby
much clarified. The publication had a decided influence on the methods of
systematics in Lepidoptera. It was incidentally pointed out that logically the
first described subspecies should bear a subspecific name like the other subspecies
(P. eurypylus eurypylus, P. eurypylus lycaon, etc.), but two of the editors of
NOVITATES ZOOLOGICAE were much against the duplication of the name in
subspecific designations and therefore the accidentally first-described subspecies
was termed forma typica in the Revision. Collaboration with either of his
Curators or with the specialists in the British Museum was the best method of
dealing with the rapidly increasing material of the orders in which he was most
interested ; but Walter Rothschild worked in many instances quite inde-
pendently, neither discussing the subject with the Curators, nor submitting the
manuscript. The careful observer will notice in the numerous papers entirely
composed and written by him some interesting peculiarities which might puzzle
anybody who was not intimately acquainted with the working of the mind of
the author. When describing a new species, the points of difference discovered
became so super-conspicuous in his mind that he emphasized them by the
employment of superlatives and in print often by the use of clarendon type.
Another point is the frequent lack of detail. Although he had studied at Bonn
and Cambridge at a period when the acquisition of knowledge in internal and
external morphology was one of the main objects of the Zoological curriculum,
Walter Rothschild had never taken kindly to the microscope and microtome.
Structural details easily escaped him. Like an artist, he perceived the animal
as a whole and not the details which made up the picture, and in describing a
small insect he painted in words a picture that was vividly formed in his mind
without the substructure of detail which would have been revealed by the cold
lenses of a microscope. In order to understand and interpret such descriptions,
2
10 NOVITATES ZOOLOGICAE XLI. 1938.
usually short, one must put oneself into the author’s frame of mind. It will
also strike the Lepidopterist as rather strange in an author of great experience
that he so often laid undue stress on differences of size in closely related forms
of butterflies or moths ; for it was so well known that size in Lepidoptera depends
frequently on the quality or quantity of the food of the caterpillar that it is no
criterion of distinctness unless corroborated by other differences. But here
Walter Rothschild applied experience gained in Ornithology to the systematics
of insects, the species and subspecies of birds being as a rule remarkably constant
in size and many subspecies diagnosed by a difference in wing-length only.
In 1893 and 1894 accommodation was already short, and as the collections
came in from abroad the lack of space became very embarrassing and caused
much waste of time. Asa first measure of relief a small corrugated-iron building
was put up for the overflow of Lepidoptera cabinets, and in 1897 this was followed
by a much larger building of the same type for the birds and additional insect
cabinets. And as it appeared quite impossible to build up large collections of
both Lepidoptera and Coleoptera with only one Entomologist in charge, it was
decided to dispose of the beetles ; they were gradually sold, and the Lampyridae
given as a present to Olivier, in the end only one small family (Anthribidae)
being retained, as the Curator of Entomology did not wish altogether to lose
contact with Coleopterists. Additional space and time were thus gained for
Lepidoptera.
Up to 1895 all the Lepidoptera were set by professionals in London and
elsewhere, but with the greater influx of collections it became imperative to have
an assistant on the premises who would attend to the setting of particularly
valuable material. Mr. J. W. Shipp, who had been working in the Hope Depart-
ment at Oxford, was engaged, and came to Tring in 1895 (he died in the winter
1897-8 at the home of his parents at Oxford). In the following year the staff
was further increased by the appointment of Mr. F. W. Goodson, the brother of
Arthur Goodson, and from this time on the latter worked exclusively in the
Ornithological Department and the former in the Department of Entomology.
Both had learned to set butterflies and moths, and did some of this work in
overtime ; several ladies in Tring and neighbourhood also took up setting,
working at home, so that at one time there were more than half a dozen out-
side helpers to cope with the many collections that came in.
Work on the systematics on mammals, birds, reptiles, and insects kept pace
with the increase in the collections. The more important of the numerous
publications by Walter Rothschild between 1897 and 1908, in collaboration
with the Curators, were the Monograph of the Charaxes, the Monograph of the
Cassowaries, the Revision of the Sphingidae, the Revision of the American Papilios,
and Extinct Birds. Before the century ended, his work and his Museum were
known in all quarters of the globe.
In 1908 there came a break in Walter Rothschild’s life. He retired from
the City, having neither inclination nor ability for finance, and he could now
spend his time with more freedom on scientific pursuits and travel. He had
frequently been for four or five weeks in the Alps, but had never gone outside
Europe. From 1908 to 1914 he took longer holidays, visiting various European
countries and North Africa, always with the object of increasing his collections,
Algeria being the principal country visited by him nearly every year, accom-
panied by one of the Museum’s Curators, usually Dr. Hartert. He was not one
Novirarres ZooLoGIcAE XLI. 1938. 11
of those hardy travellers who can stand any amount of discomfort, and having
suffered when a youth from an attack of pneumonia was always afraid of catching
a cold, and therefore preferred to remain within easy reach of a physician. It
was fortunate that during his first visit to Algeria he made the acquaintance of
Dr. Chr. Nissen, the Danish Consul-General, a physician who took an interest
in Lepidoptera. When staying in the interior of Algeria, Walter Rothschild’s
party was generally accompanied by Dr. Nissen. Aflou, Laghouat, Touggourt,
and, of course, Biskra, and many places in the coast region and on the central
plateau, were visited and some time spent at each for collecting purposes. A
more arduous undertaking was a journey to El Oued, in 1909, across a wide
stretch of sandy desert, Walter Rothschild on horseback and the other European
members of his caravan on camels. As an able collector of Lepidoptera
(V. Faroult) was employed for many years, staying in different districts and
collecting and breeding specimens all the year round, and as Dr. Hartert made
several independent trips to Algeria in company with an assistant who could
collect and skin, the Algerian material of birds and butterflies and moths became
very extensive ; in addition, some hundred specimens of mammals were obtained,
among them several new species and subspecies and the Wild Boar and Leopard.
It was a good documentation for a critical study of the Algerian fauna, Dr.
Hartert publishing a number of papers on the birds, Walter Rothschild several
articles on the butterflies and moths, and Charles Rothschild and Dr. Jordan on
Siphonaptera.
In 1908 a budget had been agreed upon, intended to limit the yearly expenses
of the Museum to a definite sum sufficient to meet the overhead expenses and to
leave a good margin for the purchase of further collections. However, Walter
Rothschild had always been accustomed to a large expenditure, of which the
Museum budget was only a moderate item, and it could really not be expected
that he would suddenly turn a blind eye to specimens he coveted. In the same
year the administration of the Museum was simplified for the Director by the
engagement of a Secretary-Librarian, the books scattered over several rooms
and landings were united in a library built in 1907-8, and the Lepidoptera
assembled not far from the Museum (Akeman Street) in a house altered for the
purpose, being provided with central heating and electric light (the electric light
of the Museum buildings was provided, as it still is early in 1938, by the Tring
Park Estate, the town having installed electric light only lately). In 1909 the
second corrugated-iron structure was pulled down and a large permanent building
added to the Museum ready for occupation in 1910, consisting of a basement
and two floors, the lower floor being allotted to the entire collection of bird skins
and the upper floor to an extension of the public exhibits. In 1912 another
large building, a gift from his brother, was begun and finished in 1913, with two
large halls for the accommodation of all the Lepidoptera, a basement of the
same size and several working-rooms with shelves for the Entomological Library,
ample room being provided for the extension of the collection and the Library.
The increase in the overhead expenses since 1908 and the financing of the
exploration of North Africa rendered it financially inconvenient to continue the
issue of extensive Revisions and Monographs. The researches on the Saturnian
moths, a superfamily in which Walter Rothschild was much interested, were
discontinued, only the manuscripts on some small groups being completed and
published after the war.
12 Novirates ZooLoGIcAE XLI. 1938.
From 1925 onwards Lord Rothschild, who had succeeded to the title on the
death of his father in 1915, attended several Entomological, Ornithological, and
Zoological Congresses abroad, but after his visit to Algeria in 1914 did not go
again on a collecting trip ; he felt tired too easily and preferred staying at home.
In 1930 Dr. E. Hartert retired, and in 1931 his assistant, Arthur Goodson,
died, the Ornithological Department being carried on by Lord Rothschild. We
endeavoured to find a successor to Dr. Hartert with the same enthusiasm for
Ornithology, but as there was no prospect of permanent employment and the
salary offered moderate, the negotiations came to nothing. That proved to be
fortunate, for in 1931, when arrangements were nearly completed for an expedi-
tion to New Guinea with the object of collecting birds and Lepidoptera, Lord
Rothschild was suddenly confronted with the demand to pay a debt which he
had allowed to accumulate without making provision for its ultimate payment,
a debt not incurred on account of the Museum. When he confessed his plight
and said that he would have to sell the bird-collection in order to meet the debt,
substantial assistance was at once offered, but the sum required was fairly large
for that moment of general financial depression, the outlook in the City being
very gloomy in the autumn of 1931. He tried in vain to exclude from the sale
the Parrots and Birds of Paradise in addition to the Ratites, but had to be content
with the Ratites and a few specimens of rarities not represented in the British
Museum. The collection was packed up and despatched to the American
Museum of Natural History, New York, in 1932. The loss of the collection
was a great shock to him, and though he continued to work in the Museum at
Lepidoptera, to which he had always devoted most of his time, his energy was
much diminished ; he missed the birds, and it was very hard to make him realize
that in consideration of his age and his financial position, it was useless starting
anew to build up a bird-collection. It was a disaster for him which preyed on
his mind to the end.
In May 1935, when walking from the Museum to the mansion, he slipped
in a paved tunnel in the grounds and injured his left knee very severely. The
shock brought on an attack of gout, and he had to keep in bed for five months.
He could walk again, but was very bent and obliged to use two sticks for safety.
He soon became used to his state as an invalid, however, and even attended the
1936 meeting of the British Association at Blackpool, enjoying the drives along
the coast and the visits to various gardens at Blackpool and neighbourhood.
His capacity for work, however, was practically gone, a couple of hours in the
Museum being enough to tire him. When some time after the death of his
mother early in 1935 Tring Park was shut up, the Home Farm House at Tring
was altered to suit his requirements, and here he lived the last year of his life.
Though an invalid who required constant attention, he seemed to be much
happier and more content in his new home than he had been for years. In
June 1937 his back began to trouble him, and by the middle of the month it
became apparent that his spinal cord was affected by cancer, the paralysis
extending downwards from the middle of the back. During the first few weeks
of his illness his mind remained clear and he could discuss scientific problems
and make plans with the Curator for the future of the Museum. Devoted
nursing kept him alive till the 27th August, when he died in his sleep early in
the morning.
Since the sale of his bird-collection Lord Rothschild had been a less prolific
NOVITATES ZootoGIcAR NLI. 1938. 13
author than before. The last important paper published by him, in collaboration
with Captain Guy Dollman of the British Museum, was a Monograph of the Tree
Kangaroos. This work, beautifully illustrated, was a very appropriate finish to
his activities as a mammalogist, for the Marsupials were an order of mammals
which had interested him deeply from his undergraduate days, the series of
species represented by mounted specimens and skins assembled by him in the
Tring Museum approaching completeness. His knowledge of the Marsupials
and the keenness of his eye are well illustrated by an incident that happened in
1934 and is worth recording. Driving along Brompton Road, he noticed a
chauffeur standing at the door of a car with a fur-rug over his arm ; he stopped,
looked at the rug, took the number of the car, wrote to the lady who owned the
car, and bought the rug of Tree Kangaroo pelts for £30, adding it to the collection
of skins in the Museum, where it still is, together with letters referring to the
transaction. The Marsupials had serious rivals in Lord Rothschild’s affection in
the Monotremes and the Anthropoid Apes, both groups being better represented
at Tring than in any other Museum. His favourite birds were the Ratites and
Birds of Paradise, of both of which he has described many novelties. The
Cassowaries in particular were a speciality on which he spent much time and
money, there being no less than sixty-two mounted specimens in the public
galleries at Tring, including many types, a series far in excess of anything pre-
served in other Museums. The last two notes he published, in 1937, were on
Cassowaries. -Albinism and melanism in mammals and birds had a great
fascination for him; the number of albinos in the exhibition series being
particularly large. His early connexion with Dr. Günther drew his attention to
the Gigantic Tortoises, weird creatures which aroused his lively interest because
they were so large and moreover threatened with extinction. Among Lepidop-
tera his favourites were Papilio, Charaxes, and Morpho, Sphingidae, and Saturn-
idae, and in later years Syntomidae and Arctiidae, of which he has described
hundreds of new species. His fondness for large species did not prevent him
from studying also small moths, such as Lithosiids and Pyrals, and he always
asked collectors going to the Tropics to pay special attention to small Lepidoptera,
which, he said, were much neglected by professional collectors because they had
little commercial value. Like Charles Oberthtir before him, he learnt by his
own experience in taxonomy that the old kind of collection containing of each
“ species ” a restricted number of specimens did not give any scope for the study
of variation, and he therefore advocated, like Charles Oberthür, the amassing of
long series wherever feasible. The idea was carried out by him, there being
sometimes hundreds of specimens from the same place in the Tring collection of
Lepidoptera. It was a frequent topic of conversation with visitors. He was
right in saying that systematics must rise above the mere registration of the
species and, being the basis necessary for the solution of biological problems,
must be sound. He knew, however, that the systematist is constantly faced
by the fact that the material at his disposal is inadequate and his results, there-
fore, remain often tentative and inconclusive. Long series facilitate research,
and facilitation of research was one of the driving factors in Lord Rothschild’s
activities as a collector. This idea was there from his earliest days as a syste-
matist : for he had adopted the so-called Continental style of setting insects,
which makes it easy to handle a specimen and to inspect it under a strong hand-
lens or the microscope, whereas the short pin then generally in use in Great
14 NOVITATES ZooLoGIcAE XLI. 1938.
Britain (and nowhere else) leaves insufficient room for good labelling and compels
the investigator of the legs or body to take the label off. The adoption by him
of glass-bottomed drawers for Lepidoptera was likewise a great convenience
for the comparison of the underside of long series. Though Lord Rothschild
liked nothing better than showing series of beautifully preserved specimens to
interested visitors, the possible service of the collections to science was uppermost
with him, and he wished bad specimens to be kept because they might be wanted
for research in morphology. Therefore, unlike the old type of amateur who
was most anxious that his well-arranged rows of specimens with or without those
“ugly ” bits of paper on the pins, should not be disturbed, Lord Rothschild was
always highly gratified when some new distinctive character of a species was
discovered by the dissection of specimens. He was no less liberal in lending
material to specialists at home and aboard and in according to scientific visitors
every facility for successful work in the Museum. ‘Tall and broad, he made an
impression on strangers, who at first stood in awe of the owner of all these
collections, but his friendly smile and amiable courtesy soon put them at their
ease. His liberality has influenced the policy of other Museums to the great
advantage of science. Keenness to increase his own collections did not prevent
him from being generous to other Museums, particularly the British Museum,.
where numerous specimens are labelled as being his donations. In various
instances he bought several specimens of a mammal, keeping one for his galleries
and giving the others away.
Tring Park not being his property, he had no space in which to keep live
animals ; but he was so fond of them that he frequently acquired interesting
species of mammals, birds, and reptiles and deposited them in the Zoological
Gardens in London and elsewhere. Dogs, however, were the greatest attraction
for him ; his love of them being often very embarrassing, especially in the wilds
of Algeria, where he always tried to make friends with any dog he saw ; sometimes
the dog looked as if it would bite first and then perhaps wag its tail. His interest
in natural history went beyond Zoology and embraced Botany as well as Biblio-
phily. He was one of the original holders of the Victoria Medal of Honour in
Horticulture (V.M.H.), established in 1897 with the consent of Queen Victoria
“to enable the council of the Royal Horticultural Society to confer conspicuous
honour on British Horticulturists resident in the United Kingdom”; Lord
Rothschild is survived by only one of the sixty original holders of the medal.
Before 1908 his collection of live Orchids was considerable, several species being
named after him, and the American Orchid Society electing him an Honorary
Member.
The library of the Museum contains a number of rare old books on travel
and natural history, of little use for systematic Zoology but of interest for a
book-lover, De Bry’s publications particularly taking his fancy.
His services to Zoological science were much appreciated, many Ornitho-
logical and Entomological Societies electing him an Honorary Fellow; he
became a Corresponding Member of the Academy of Natural Sciences of Phila-
delphia, and in 1898 the University of Giessen conferred on him the honorary
degree of Ph.D. In 1899 he was elected a Trustee of the British Museum, in
which institute he took as much interest as in his own Museum, and in 1911 he
received the blue ribbon of science, becoming a Fellow of the Royal Society in
recognition of his services to the natural sciences. He was President of the
NOVITATES ZOOLOGICAE XLI. 1938. 15
Hertfordshire Natural History Society in 1911 and 1912, and of the Entomo-
logical Society of London in 1921 and 1922, Chairman of the British Ornithologists’
Club from 1913 to 1918, and of the British Oological Association from 1923 to
1932 ; in 1932 he presided over the Zoology Section of the York Meeting of the
British Association, taking on this occasion as the subject of his address, “‘ The
Pioneer Work of the Systematist.”’
Few naturalists who knew Lord Rothschild’s pursuits in science were aware
that there was another active side to his life. As a young man he served as an
officer in the Royal Buckinghamshire Yeomanry, attaining the rank of Major,
and when at the outbreak of the Boer War the whole regiment volunteered for
active service, and a selection from among the officers and men was accepted,
Walter Rothschild regretted he was not numbered among them; he was at
that time already a heavy man, requiring exceptionally strong horses, and it
was obvious that neither man nor horse would for long stand strenuous campaign-
ing in a hot climate. From 1899 to 1910 he sat in Parliament as member for
the Aylesbury Division of Buckinghamshire, was made a Justice of the Peace,
and for many years represented Manchester on the Jewish Board of Deputies.
Much of his time was taken up by service as an active member or chairman of
committees of various Societies and Institutions, and he was particularly in-
terested in the Middlesex Hospital. For a number of years he was chairman of
the Tring Urban District Council, and until his death President of the Tring
Agricultural Society. He used to hunt regularly with the Rothschild Stag
Hounds before he became too heavy for this sport, and was a good shot in the
pheasant coverts and at the Tring reservoirs. He remained unmarried and
lived with his parents, his mother surviving his father by twenty years, dying
in 1935.
At the time of his death the Museum buildings had an aggregate floor-space
of nearly an acre and a half, inclusive of the basements for storage.
I. The Public Galleries contained :
1. 2,004 complete mounted mammals, 207 heads, 335 pairs of horns
and antlers, 6 large Elephant tusks, and many skeletons, and skulls,
some of the horns being record specimens. There were in this series
13 Gorillas, 25 Chimpanzees, 228 Marsupials, 24 Echidnas, etc.,
the most valuable specimen from the commercial point of view
being the Quagga.
2. 2,400 mounted birds, inclusive of 18 Apteryx, 62 Cassowaries, 62
Birds of Paradise, 520 Humming Birds, the Great Auk with skeleton
and two eggs, and a fine Korean Eagle.
3. 680 Reptiles and Amphibians, including 144 Giant Tortoises.
4. 914 Fishes.
5. Representative collections of Invertebrates.
II. The Students’ Department contained :
6. 1,400 mammal skins with skulls, including a number of types.
7. 4,470 bird skins, among them some extinct species.
8. A very large collection of birds eggs, including the best collection
of eggs of Birds of Paradise and a number of perfect eggs of
Aepyornis.
9. 300 Reptiles, dry, and a small collection in alcohol.
16 NOVITATES ZOOLOGICAE XLI. 1938.
10. Collections of unmounted skeletons of mammals, birds, and reptiles ;
numerous bones of fossil birds.
11. Some two million specimens of Lepidoptera, with numerous types
(of Geometridae alone over 6,000 types).
12. A collection of Anthribid beetles containing types or paratypes of
more than two-thirds of the known species.
III. A Library of nearly 30,000 volumes.
Lord Rothschild had placed the collections at the service of scientists during
his life, and he wished them to remain after his death equally available for
research. His frequent visits to the British Museum all his life, his intimate
connexion with it as a trustee, and his collaboration with members of its staff,
turned his mind towards that institute when, during his last illness, he thought of
the future of his own Museum. Knowing how short of space the British Museum
was, and would remain for a long time, he agreed with his Curator that the
Tring Museum made into an annexe of the British Museum would not only enrich
the national collections, but could be used immediately to relieve the congestion
from which the Entomological Department of the British Museum suffered.
He therefore sent a memorandum to the Trustees, offering the entire freehold
buildings, collections, and library to the British Museum on condition that the
Tring Museum be continued in some form or other as an Institute for Zoological
Research. It was touching to see how gratified he was when the Chairman of
the Trustees, the Archbishop of Canterbury, wrote and thanked him for his
generosity. It made him very happy to hear that the Trustees would accept
the gift, pending the consent of the Treasury and Parliaınent.
His reputation as a Zoologist will be lasting. Whatever the present genera-
tion of biologists may think of systematics and Museums, the knowledge of the
animals as created by Nature and preserved as well as can be in collections for
future generations of mankind will for ever be the subject of research for the
scientist and of admiration for the public. The founder of the British branch of
the House of Rothschild, Nathaniel Mayer, acquired such fame as a financier
that the family name still casts a spell over the imagination of man in all quarters
of the globe. His great-grandson owed much to that name, but more to his
own activity as a Zoologist. Passing along a different road, he, too, arrived
at world-wide fame.
Novirarres Zootocican NLI. 1938. 17
LIST OF SCIENTIFIC PUBLICATIONS.
The names of collaborators are placed in brackets at the end of the quotation.
I. GENERAL.
1894. First glimpses of the Zoology of the Natuna Islands.
Nov. Zool. i, pp. 467, 468. (E. Hartert.)
1899. A review of the Ornithology of the Galapagos Islands, with notes on the
Webster-Harris Expedition. L.c. vi, pp. 85-142. (E. Hartert.)
1905. On the advantage of the trinomials. Ibis, pp. 133-137.
1912. On the term “ Subspecies ” as used in Systematic Zoology.
Nov. Zool. xix, pp. 135-136.
1913. Bird-life on Oceanic Islands, and insular variation.
Trans. Herts. Nat. Hist. Soc. xv, pp. 1-14.
1914. Mimicry and protective resemblance. L.c. xv, pp. 105-120.
A zoological tour in West Algeria. Narrative.
Nov. Zool. xxi, pp. 180-186. (E. Hartert.)
1915. On Algeria: the Tell Region, the Hauts Plateaux, and the Saharan
Desert. Bull. B.O.C. xxxv, pp. 90-99.
Effects of environment on the evolution of Species, discussion.
L.c. xxxv, pp. 128-142.
1918. On the naming of local races, subspecies, aberrations, seasonal forms,
etc. Trans. Ent. Soc. Lond. pp. 115, 116.
1922. President’s Address : Algeria and its Fauna.
Proc. Ent. Soc. Lond. 1921-22, pp. 131-162.
1923. President’s Address : On some aspects of variation in Lepidoptera.
L.c. 1922-23, pp. 122-134.
1925. Nomenclatural note. Bull. B.O.C. xlv, pp. 89, 90.
1932. The pioneer work of the Systematist.
Brit. Assoc. Adv. Sci. York Meeting, 1932, pp. 89-102.
The same in Nature, cxxx, pp. 529-531.
Obituaries :
1916. Henry Ecles Dresser. British Birds, ix, pp. 194-196, portrait.
1934. Ernst Johann Otto Hartert.
Ibis, pp. 350-377, two portraits, bibliography.
II. MAMMALIA.
1892. Descriptions of two new Mammals from New Guinea.
Proc. Zool. Soc. Lond. 1892, pp. 545, 546.
1893. On Mesoplodon bidens. Ann. Mag. N.H. (6), xi, p. 439.
1894. Propithecus majori sp. nov. Nov. Zool. i, p. 666, pl. xiv.
1895. On two new species of Antelopes. L.e. ii, pp. 52, 53, pl. iv.
1897. Note on a new Antelope. Ann. Mag. N.H. (6), xx, pp. 376, 377.
Neumann’s Hartebeest. Nov. Zool. iv, p. 377, pl. xiv.
18
1898.
1899.
1900.
1901.
1902.
1903.
1905.
1906.
1907.
1908.
1909.
1910.
1911.
1912.
1913.
1914.
NOVITATES ZOOLOGICAE XLI. 1938.
Note on some Kangaroo hybrids. L.c. v, p. 4.
Notes on T'ragelaphus spekei spekei and T. speakei gratus, with description
of a new species. L.c. v, p. 206.
Dendrolagus mazimus. L.c. vi, p. 217, pl.i.
On a new race of Ibex: Capra sibirica lydekkeri subsp. nov.
L.c. vii, pp. 277, 278, pl. ii.
Notes on Bubalis. L.c. viii, pp. 177, 178.
Two new subspecies of Proteles. Lic. ix, p. 443.
Notes on Abyssinian Mammals.
In Powell-Cotton, Sporting Trip through Abyssinia, pp. 453-486.
Note on Alces bedfordiae. Proc. Zool. Soc. Lond. 1902, ii, p. 317.
Preliminary diagnosis of a new genus and species of Kangaroo.
Nov. Zool. x, p. 414.
Description of a new species of Gazelle. L.c. x, p. 480, pl. xv.
Note on Dendrodorcopsis woodwardi. Luc. x, p. 543.
Notes on Zaglossus and description of a new subspecies of Echidna
hystrix. L.c. xii, pp. 305, 306.
Note on the Eland of the White Nile. L.c. xii, p. 447, pl. xii.
Note on Macropus rufus Desm., with description of a new subspecies.
L.c. xii, p. 508.
Notes on two Kangaroos from the “‘ Northern Territory of South Aus-
tralia,’ with description of a new species. L.c. xii, pp. 509, 510.
Notes on Anthropoid Apes.
Proc. Zool. Soc. Lond. 1904, ii, pp. 413-440, pl. xxiv.
On African Forest Pigs. L.c. 1906, p. 632.
Description of a new Zebra. L.c. 1906, p. 691.
Description of a new Bush-Buck. L.c. 1906, pp. 691, 692.
Further notes on Anthropoid Apes. L.c. 1906, pp. 465-468.
Zur Nomenclatur der Menschenaffen.
Sitzungsber. Ges. Naturf. Freunde, pp. 85-87.
Further notes of Macropus magnus. Nov. Zool. xiv, p. 333, pl. ii.
Description of anew Tree Kangaroo. L.c. p. 506, pl. iv. (F. Foerster.)
On a new race of Orycteropus. L.c. p. 506.
Description of a new species and two new subspecies of Antelopes and a
new Sheep. Proc. Zool. Soc. Lond. 1907, pp. 237, 238.
Note on Gorilla gorilla diehli Matchie.
Nov. Zool. xv, pp. 391, 392, pl. xii.
Microunga angustirostris (Gill). L.c. xv, pp. 393, 394, pls. i-viii.
Nasua vittata Tsch. L.c. xvi, p. 333, pl. i.
Notes on Sea-Elephants (Micrownga). L.c. xvii, pp. 445, 446, pls. viii, ix.
On a new Marsupial. Ann. Mag. N.H. (8), vii. p. 337. (F. Foerster.)
Note on Zaglossus. The Field, December Number.
Some notes on the genera Zaglossus and Tachyglossus.
Nov. Zool. xx, pp. 188-191.
On Ovis lervia Pallas and its subspecies. L.c. xx, pp. 459, 460.
Descriptions of some new forms of Antelope, with notes.
Ann. Mag. N.H. (8), xii, pp. 574-576.
Ein neues Känguruh aus Neuguinea.
Nov. Zool. xxi, pp. 261, 262. (F. Foerster.)
Novirates Zoonoaicarm XLI. 1938. 19
1933.
1936.
1891.
1892.
1893.
Preliminary description of a new Warthog.
Ann. Mag. N.H. (9), vi, p. 416.
On two new races of Oryza. L.c. (9), viii, pp. 209, 210.
Captain Angus Buchanan’s Air Expedition —III. Ungulate Mammals.
Nov. Zool. xxviii, pp. 75-77.
On a new subspecies of Zaglossus, with remarks on the other species of
the genus. Ann. Mag. N.H. (9), x, pp. 129-131. (O. Thomas.)
On a new race of Bharal. L.c. (9), x, p. 231.
Description of a new Baboon. L.c. (9), x, P- 232.
Remarks on a Mountain Gorilla from near Lake Kiva.
Proc. Zool. Soc. Lond. pp. 176, 177.
On a new race of Bongo and of Gorilla. \
Ann. Mag. N.H. (9), xix, p. 271.
Description of a new race of Aardvark (Orycteropus afer kordofanicus).
L.e. (9), xix, p. 512.
On the skull of Gorilla gorilla halli Rothsch. L.c. (9), xix, p. 512.
New Mammals from Dutch New Guinea.
Proc. Zool. Soc. Lond. 1932, ii, pp. 1082-1084. (G. Dollman.)
On Mammals collected in Dutch New Guinea by Mr. F. Shaw Mayer in
1930. L.c. 1933, i, pp. 211-219, pls. i-iv. (G. Dollman.)
A new Tree-Kangaroo from the Wondiwoi Mts., Dutch New Guinea.—
Abstr., Proc. Zool. Soc. Lond. i, p. 40, and reprinted in Proc. Zool.
Soc. Lond. ii, pp. 540, 541. (G. Dollman.)
The genus Dendrolagus.
Trans. Zool. Soc. Lond. xxi, pp. 477-548, pls. 35-57. (G. Dollman.)
III. AVES.
Description of a new Pigeon of the genus Carpophaga.
Proc. Zool. Soc. Lond. pp. 312, 313, pl. xxviii.
Descriptions of seven new species of birds from the Sandwich Islands.
Ann. Mag. N.H. (6), x, pp.108-112.
A new Pigeon: Ptilopus salvadorii, p. 10, Jobi—Exhibition of three
new birds from the Sandwich Is.: Palmeria mirabilis, Loxops ochracea,
Hemignathus affinis, p. 16 (no descriptions)—A new Duck: Anas
laysanensis, p. 17, Laysan I. Bull. B.0.C. i.
Columba rupestris pallida subsp. nov. p. 41, Altai.
Ornith. Monatsb. i, p. 97. (E. Hartert.)
Die Formen von Fringilla spodiogenys in Nordafrika.
L.c. i, p. 97. (E. Hartert.)
Descriptions of three new birds from the Sandwich Islands.
Ibis, pp. 112-114.
The Avifauna of Laysan and the neighbouring islands ; with a complete
history to date of the Birds of the Hawaiian possessions.
Folio, London, parts i and ii; pp. i-xiv and 1-26, 56 pls. (E.
Hartert.)
Description of a new species from the Sandwich Is.: Hemignathus
lanaiensis, p. 24, Lanai.—A new genus and species of Fringilline bird
from the Sandwich Is.: Pseudonestor xanthophrys, pp. 35, 36. (E.
Hartert.)—A new species of Rail: Rallus muelleri, pp. 40, 41, Auckland
1894.
1895.
NOVITATES ZOOLOGICAE XLI. 1938.
I.— Three new birds from the Sandwich Is.: Acrulocercus bishopi,
p- 41, Molokai; Himatione newtoni, p. 42, Mauai ; Himatione wilsoni,
p. 42, Mauai.—Exhibition of a melanistic variety of the Razorbill
(Alca torda), p. 43.—A new species of Albatross: Diomedea immuta-
bilis, p. 48, Laysan.—Loxops wolstenholmei n.sp., p. 56, Oahu; Viri-
donia maculata (Cab.), p. 57, Oahu; Anous hawaviensis sp.n., p. 57 ;
Oestrelata nigripennis sp.n., p. 57, Kermadec Is.; Thalassogeron
salvini sp.n., p. 58, N. Zealand; Diomedea bulleri sp.n., p. 58, N.
Zealand. — Notes on the Genus Apteryx, pp. 59-62.
Bull. B.0.C. i.
Note on Himatione dolei Wils., p. 9.—A new species of Snipe: Gallinago
tristrami, pp. 11, 12. Antipodes I. L.c. ii.
On Turturoena sharpei Salvad. and Osmotreron everetti sp. nov.
Nov. Zool. i, pp. 40-41, pl. iii.
On Albino Swallows and Wheatears. L.c. i, p. 667.
On the Habitat of Chalcopsittacus dwivenbodei Dub. resp or:
Salvadorina waigiuensis gen. nov. et sp. nov.
L.c. i, pp. 683, 684. (E. Hartert.)
On a new Miro from the New Zealand Region. L.c. i, p. 688.
On a new Bustard from the Palaearctic region. L.c. i, p. 689. (E. Hartert).
Pseudonestor xanthophrys a Drepanine Bird. L.c. i, p. 692.
A series of Snipes from the New Zealand region, with remarks thereon,
p. 16.—Note on Palmeria dolei (Wils.) p. 25.—Exhibition of egg of Great
Auk, and eggs of Ptilorhis victoriae, p. 36.—Note on Apteryx, p. 36.—
Exhibition of three species of birds, p. 42.—Exhibition of birds from
the Talaut Is., p. 46.—On Parus owstoni, p. 46.—On Aithurus polyimus
(Linn.) and the new race Aithurus taylori, pp. 46, 47.—A Japanese
specimen of Diomedea immutabilis, p. 47. Bull. B.0.C. iii.
A new genus and species of bird from New Zealand: Traversia lyalli,
pp. 10, 11. Stephens I.—On Craspedophora mantoui Oust., p. 11.
L.e. iv.
On Salvadorina waigiuensis Roths. & Hart. Nov. Zool. ii, p. 22, pl. iii.
Note on the Zoxops of Oahu. L.e. ü, p. 54.
Further notes on the Houbara Bustard. L.c. ii, p. 54 (E. Hartert).
A new Bird of Paradise. L.c. ii, p. 59, pl. v.
Note on the Stephens Island Rock-Wren Traversia lyalli Rothsch.
L.c. ii, p. 81.
A new species and genus of Rollers. L.c. ii, p. 479.
A new species of Bower-Bird. L.e. ü, p. 480.
A new species of Rail. L.c. ii, p. 481.
A new species of Tanager. Luc. ii, p. 481.
The White Swallows of Aylesbury. L.e. ii, p. 84.
On a new Parrot. L.c. ii, p. 492.
Exhibition of Birds of Paradise of different genera and remarks thereon,
pp. 21, 22.—A new species of Bower-bird: Artluroedus jobiensis,
pp. 26, 27.—Spermophilopsis (nom. emend.) toreplace Drepanorhynchus
Dubois 1894, p. 37.—On Sterna vittata Gmel. p. 37. Bounty I.—Note on
Amblyornis inornata, Parotia carolae, and other Birds of Paradise, p. 42.
Bull. B.0.C. iv.
9
Novirares ZooLoGIcAE XLI. 1938. 21
1896.
1897.
1898.
List of three small collections from British New Guinea, mostly brought
together in the Owen Stanley Mts.
Nov. Zool. iii, pp. 8-19, pl. i. (E. Hartert.)
Notes on two more specimens of Astrapia splendidissima. Lc. iii, p. 19.
List of a collection made by Mr. Albert S. Meek on Fergusson, Trobriand,
Egum, and Woodlark Islands. L.c. iii, pp. 233-251. (E. Hartert.)
On some species in a small collection made on the Owen Stanley Mts.
in the Kaiari and Oriori distriets between Mts. Alexander and Nisbet
in January 1896. . Luc. ii, pp. 252-255. (E. Hartert.)
On some skins collected from April to June on Mount Victoria, Owen
Stanley Mts., mostly at elevations of from 5,000 to 7,000 feet.
L.c. iii, pp. 530-533. (E. Hartert.)
List of a collection made in the Aru Islands by Captain Cayley Webster
from May to July 1896. L.c. iii, pp. 534-536. (E. Hartert.)
Uratelornis chimaera Rothsch. (Description appeared in Vol. ii. p. 479.)
L.e. i, pl. ii.
Exhibition of Astrapia splendidissima and Amblyornis, p. 38.—A new
Pigeon from Sumba: Ptilopus dohertyi, p. 46.—Exhibition of Humming-
birds, p. 46. Bull. B.0.C. v.
A new Parrot from New Guinea: Psittacella picta, p. 5, Mt. Victoria,
Owen Stanley Range.—A new genus and species of Bird of Paradise :
Loboparadisea sericea, pp. 15, 16. L.e. vi.
On Differences between Güldenstadt’s Redstart and its Eastern Ally.
Nov. Zool. iv, pp. 167, 168.
Description of a new Hill-Wren from Flores. L.c. iv, p. 168.
On the Figures on Plate II. L.c. iv, p. 169.
Comatibis eremita (Linn.), a European bird.
L.c. iv, pp. 371-377, pls. vii-x. (KH. Hartert, ©. Kleinschmidt.)
Rhytidoceros everetti spec. nov. Journ. Ornith. p. 513.
Exhibition of Loboparadisea sericea, Nemophilus macgregoriae, and
Loria loriae, with a note on Loria mariae and Loria loriae, pp. 24, 25.
— Rhamphocoelus inexpectatus sp.n. p. 32, Panama.—Notes on Oestrelata
haesitata (Kuhl), Paradisea intermedia De Vis, Ruticilla erythrogastra
Güld. and Z. grandis Gould, pp. 40, 41.—On the subspecies of Paradisea
minor: typical P. minor, Dutch N. Guinea; P. minor jobiensis
subsp. n., Jobi; P. finschi A. B. Meyer, German N. Guinea, pp. 45,
46.—Notes on Ardetta neoxena Cory, Eclectus cornelia Bonap., Psit-
teuteles weberi Büttik., Ps. euteles, and Paradisea minor var. albescens
Musschenbr., pp. 53, 54.—Purchase of Brehm Collection, p. 54.
Bull. B.0.C. vi.
Three new birds from Ecuador: Crypturus berlepschi, p. 5; Odonto-
phorus parambae and Nemosia rosenbergi, p. 6.—Exhibition of various
birds and note on female of Hudynamis honorata, p. 15.—New species
from New Guinea: COyclopsittacus macilwraithi, p. 21; Pachycephala
gamblei, Pachycephala salvador n. em., and Epimachus astrapioides,
p. 22. L.c. vü.
Notes on Paradiseidae. Nov. Zool. v, pp. 84-87.
On a new Parrot of the family Loriidae. L.c. v, p. 110.
Description of a new Cassowary. L.c. v, p. 418,
bo
bo
1899.
NOVITATES ZOOLOGICAE XLI. 1938.
Notes on some Parrots. L.c. v, pp. 509-511, pl. xviii.
Casuarius loriae sp. nov. L.c. v, p. 513.
Three new species: Pitta dohertyi, p. 29, Sula Mangoli; Ptilinopus
mangoliensis, S. Mangoli, and Pt. everetti, Alor, p. 334.—Note on an
expedition to the Galapagos Is. and descriptions of six new birds:
Phalacrocorax harrisi, Narborough I.; Sula websteri, Clarion I. ;
Nesomimus hulli, Culpepper I.; Nesomimus affinis, Narborough I. ;
Certhidia becki, Wenman I.; Certhidea drownei, Culpepper I., pp. 51-53.
— Two new birds from British New Guinea: Jfrita (gen. n.) coronata
and Charmosyna atrata, pp. 53, 54. Bull. B.0.C. vii.
Pitta meeki sp.n., p. 6, Rossel I. ; Pitta novaehiberniae Ramsay, distinct
from P. mackloti, p. 7; Nesomimus carringtoni sp. n., p. 7, Barrington
I., Galapagos ; exhibition of specimens, p. 7.—Egg of Twelve-wired
Bird of Paradise, p. 13.—A new subspecies of Cassowary, Casuarius
casuarius intensus, p. 21, no loc., and a new Cormorant: Phalacrocorax
traversi, p. 21, Macquarie Is. L.c. viii.
Paradiseidae. Das Tierreich, 2. Lief., pp. vi and 52.
Birds of Paradise collected by Captain Cayley-Webster.—In Through
New Guinea and the Cannibal Countries ; Appendix II, pp. 376, 377.
Memoir on the Cassowaries. Proc. Zool. Soc. Lond. pp. 773-776.
Note on Casuarius casuarius sclateri. Nov. Zool. vi, p. 75.
A review of the Ornithology of the Galapagos Islands, with Notes on the
Webster-Harris Expedition.
L.c. vi, pp. 85-205, pls. v, vi. (E. Hartert.)
On some rare birds from New Guinea and the Sula Islands.
The genus Apieryx, with a chapter on the Anatomy of the Kiwis by F.
Beddard. L.c. vi, pp. 361-402, pls. ix-xvi. (F. Beddard.)
Nest and egg of Onemophilus macgregori De Vis., p. 26, and description of
Casuarius casuarius violicollis, subsp. n., Aru Is., p. 27.—A new Thrush:
Geocichla dumasi, Buru, p. 30.—On Casuarius casuarius slateri, the P.
cinctus group of Ptilinopus, and varieties of Lophophorus refulgens, p. 42.
—Two new subspecies of Cassowaries: Casuarius picticollis hecki,
p. 49, German N. Guinea; Casuarius uniappendiculatus aurantiacus,
p- 50, German New Guinea.—The species and subspecies of Casso-
waries, p. 55; Palaeornis salvadoru and P. derbyana, p. 56; Telespiza
cantans and T. flavissima, p. 56; Mirafra erythropygia and Cerchneis
alopex, p. 57 ; Pyrocephalus dubius and P. nanus, p. 57.
Bull. B.0.C. viii.
On LEclectus westermanni Bp., p. 2.—Exhibition of and comments on the
red-bellied species of Pitta, pp. 3, 4.—A new species of Oyster-catcher :
Haematopus reischeki, p. 4, New Zealand.—A new form of Lory: Eos
variegata obiensis ; and exhibition of Scolopax saturata, New Guinea,
and Neoscolopax rochusseni, Obi Major, p. 16.—On Hupetes geislerorum,
A. B. Meyer, p. 26. ILO, 3%,
Ein neuer interessanter Vogel aus Neuguinea.
Ornith. Monatsh. vii, p. 137.
Ein kleiner Beitrag zur ferneren Kenntnis der Ornis von Neu-Hannover.
L.c. vii, pp. 138, 139. (E. Hartert.)
NOVITATES ZOOLOGICAE XLI. 1938. 23
1900.
1901.
1902.
The Avifauna of Laysan and the neighbouring islands ; with a complete
history to date of the Birds of the Hawaiian Possessions.
London, Part iii, pp. xx and 127-320, 27 pls. (E. Hartert.)
A monograph of the genus Casuarius. With a dissertation on the
morphology and phylogeny of the Palaeognathae ( Ratitae and Crypturi)
and Neognathae (Carinatae) by W. P. Pycraft.
Trans. Zool. Soc. Lond. xv, Part v, pp. 109-290, pls. xxii-xlv.
(W. P. Pycraft.)
Lalage sharpei sp. n., p. 40, Upolu, Samoa; on Cracticus quoyi and (©.
rufescens, p. 40.—Turnix olivii sp. n., p. 44, Cooktown; Geocichla
papuensis Seebohm, p. 44.—On Aquila fulvescens, p. 51, Albania.—On
Phalacrocorax chalconoius, pp. 52, 53; on Urubitinga urubitinga, p. 53,
Caripe, Venezuela.—Albinos in the Tring Museum, pp. 67-94 (E.
Hartert).—On Burnesia gracilis, p. 100, Luxor; Parotia duivenbodei
sp.n., p. 100, Dutch N. Guinea.— Nasiterna bruijni and Chenorhamphus
grayi, p. 101, Ambernoh R. Bull. B.0.C. x.
Eggs of Bubo ascalaphus and melanistic specimen of Podicipes fluviatilis,
p. 10.—New birds: Crateroscelis pectoralis and C. rufobrunnea, p. 25,
Microeca viridiflava and Poecilodryas cyanus salvadorii, p. 26 (E.
Hartert).—On Amalocichla sclateriana and A. brevicauda, p. 26, British
N. Guinea (E. Hartert).—A new species of Crow: Gazzola unicolor, p. 29,
Banggai (E. Hartert)—On Diphyllodes gulielmitertii, p. 30.—Grallaria
parambae, pp. 36, 37, N. Ecuador. L.c. xi.
Einige Bemerkungen tiber Kasuare. Journ. Ornith. pp. 360, 361.
Notes on Papuan Birds.
Nov. Zool. viii, pp. 55-88, pls. ii-iv. (E. Hartert.)
Notes on Papuan Birds. L.c. viii, pp. 102-162. (E. Hartert.)
List of a collection of birds from Kulambangra and Florida Islands in
the Solomons Group. L.c. vill, pp. 179-189. (E. Hartert.)
List of a collection of birds from Guadalcanar Island.
L.c. viii, pp. 373-382. (E. Hartert.)
On Microeca viridiflava, Sericornis salvador, and Dacnis berlepschi,
p. 44 (E. Hartert).—A new species of Bush-shrike : Laniarius doherty,
pp. 52, 53, Nandi Escarpment.—Note on Phasianus mongolicus and
Ph. reevesi, p. 53.—A new subspecies of Kingfisher: Alcyone azurea
yamdenae, p. 65, Timor Laut Is.—A new Woodpecker: Chloronerpes
litae, p. 70, Lita, N. Ecuador.—The real male of Dacnis berlepschi
Hart., pp. 70, 71. Bull. B.0.C. xi.
On Central Asiatic Pheasants and two new species: Phasianus berezow-
skyi, Kansu, and Ph. hagenbecki, N.W. Mongolia, pp. 19-22.—New
species from Isabel, Solomon Is.: Pitta anerythra, p. 22, and Ceyz
meeki, p. 23.—Exhibition of birds from Kulambangra and Isabel,
p. 23.—New birds: Leptoptila batiyi, p. 33, Coiba I., off Panama ;
Loborhamphus (gen. n.) nobilis, p. 34, Dutch N. Guinea. L.e. xi.
On the British Dipper. Ibis, pp. 524, 525.
List of a collection of birds from south of the Issik-Kul in Russian
Turkestan. Nov. Zool. ix, pp. 161-168.
Further notes on the Fauna of the Galapagos Islands. Notes on the
birds. L.c. ix, pp. 381-418, pl. x. (E. Hartert.)
1903.
1904.
1905.
NovITATEsS ZOOLOGICAE XLI. 1938.
List of a collection of birds made on Ysabel Island.
L.c. ix, pp. 581-594, pls. vii, xi. (E. Hartert.)
List of a small collection of birds made on Treasury Island, Solomon
Islands. L.c. ix, p. 594. (E. Hartert.)
Nests and eggs from the Galapagos Is. : Certhidea olivacea fusca, Geospiza
fuliginosa, Pyrocephalus dubius, Nesomimus adamsi, pp. 46, 47.—A
variety of Bclectus roratus, p. 47.—A new subspecies: Rheindarius
ocellatus nigrescens, p. 55, Ulu Prahang, Malay Pen. —On Corvus
eremita Linn., pp. 56, 57.—Exhibition of photographs of 5 races of
Apteryx, 12 races of Casuarius, and of a hybrid between Peahen and
Guinea-fowl, p. 57.—A new species of Petrel: Oestrelata worthent,
p. 62, Pacific Ocean, not far from the Galapagos Is.—Two new birds :
Hypotaenidia kuehni, p. 75, Tukan Bessi Is.; Francolinus coqui
angolensis, p. 76, Angola. Bull. B.O.C. xii.
Birds from Cocos I., p. 6.—Exhibition of Wicranous diamesus and three
species of Gannets, p. 7.—A new species of Gannet: Sula granti, p. 7,
Galapagos Is. L.c. xiii.
Notes on Papuan Birds. Nov. Zool. x, pp. 65-116, pl.i. (E. Hartert.)
Notes on Papuan Birds. L.c. x, pp. 196-231. (E. Hartert.)
Notes on Papuan Birds. L.c.x, pp. 435-480, pls. xiii, xiv. (E. Hartert.)
Chalcurus inopinatus Rothsch. Luc. x, pl. ü.
Berichtigung. Ann. Mus. Hungar. i, pp. 447-450. (E. Hartert.)
On Janthothorax mirabilis (Reichen.), p. 31.—Two new species of birds :
Chalcurus inopinatus, p. 41, Ulu Pahang, Malay Pen.; Myzomela
kuehni, p. 42, Wetter.—Remarks on Pheasants: Phasianus torquatus
pallasi subsp. n., p. 43.—Stigmatops notabilis Finsch from Wetter,
p. 43.—A new species of Rail: Hypotaenidia wakensis, p. 78, Wake I.
Bull. B.O.C. xiii.
A new Albatross: Thalassogeron carteri, p. 6, N.W. Australia.—A
collection of birds from Hainan (with descriptions of 2 new species
and 7 new subspecies), pp. 6-9. L.c. xiv.
Microgoura meeki Rothsch. Nov. Zool. xi, pl. xxi.
On Phasianus, pp. 36-38.—On Fulica cornuta, p. 38.—On Casuarius,
with descriptions of 3 new species and 3 new subspecies, pp. 38-40.—
Hybrids of Pheasants, p. 58.—A gigantic new species of Centropus :
C. albidiventris, p. 59, Gizo I., Solomon Is.—A new Parrot from the
Philippines : Prioniturus waterstradti, pp. 71, 72, S. Mindanao.—A new
Pigeon : Microgoura (gen.n.) meeki, pp. 77, 78, Choiseul I., Solomon Is.
— A new Pitohui: P. dichrous monticola, p. 79, Upper Aroa R.—On a
series of Barn-owls, pp. 87-90.—A series of Ratite eggs, p. 90.
Bull. B.0.C. xiv.
Two new Kingfishers: Halcyon bougainwillei, p. 5, Bougainville ; Halcyon
owstoni, p. 61, Northern Mariannes.—Exhibition of Paryphephorus
dwivenbodei and Charmosyne atrata, p. 7.—A new Pitta from the Solomon
Is.: P.anerythra pallida, p. 7, Bougainville—A new Raven from Bou-
gainville: Corvus meeki, p. 21, Bougainville.—On Phoebetria fuliginosa
(Gmel.) and Ph. fuliginosa cornicoides (Hutton), p. 28. IDs AF
Further contributions to our knowledge of the Ornis of the Solomon
Islands. Nov. Zool. xii, pp. 243-268, pl.x. (E. Hartert.)
Novirares ZootocıcAu XLI. 1938. 25
A new species of Cassowary: Casuarius roseigularis, p. 32, no loc.—
Note on Eider-ducks, pp. 43, 44.—On Diomedea carteri (Rothsch.),
pp. 44, 45.—Psittacirostra psittacea deppei, nom. nov., to replace Ps.
ps. oliwacea Rothsch., p. 45.—Discussion on Dendroeca aestiva (Gmel.),
p. 47.—Albinos of Penguins, p. 57. Bull. B.0.C. xv.
Notes on extinct Parrots from the West Indies, with descriptions of 5
unnamed species: Conurus labati, p. 13, Guadeloupe ; Anadorhynchus
purpurascens, p. 13, Guadeloupe ; Anadorhynchus martinicus, p. 14,
Martinique ; Ara erythrocephala, p. 14, Jamaica; Ara gossei, p. 14,
Jamaica.—Clutch of 3 eggs of Comatibis eremita, p. 15, Mogador.
Ice. xya.
1906. Two new Birds of Paradise. Tring, 3 pp.
Two new birds: Granatellus pelzelni paraensis, Para; Hulabeornis
castanewentris sharpei, Wokan, Aru Is., p. 81.—A new species of
Polyplectron: P. katsumatae, pp. 111, 112, Hainan.
Bull. B.0.C. xvi.
On Astrapia rothschildi Foerster and Parotia wahnesi Rothsch., pp. 7, 8.
—A new Parrot: Charmosyna stellae wahnesi, pp. 27, 28, Sattelberg,
German N. Guinea.—A new Pigeon: Henicophaps foersteri, p. 28
New Britain (E. Hartert). L.c. xvii.
1907. On extinct and vanishing birds. A short essay on the birds which have
presumably become extinct within the last 500 years, and also those
birds which are on the verge of extinction, including a few which,
though not yet so far gone, are threatened with extinction in the near
future. Ornis, xiv, pp. 191-217.
Extinct birds. An attempt to unite in one volume a short account of
those birds which have become extinct in historical times—that is,
within the last six or seven hundred years. To which are added a
few which still exist, but are on the verge of extinction.
London, Fol., 244 pp., 42 pls.
Notes on Papuan Birds. Nov. Zool. xiv, pp. 433-446. (E. Hartert.)
List of a collection of birds made by Mr. A. S. Meek in the mountains
on the Upper Aroa River and on the Angabunga River, British New
Guinea. L.c. xiv, pp. 447-483. (E. Hartert.)
Some notes on Cassowaries. L.c. xiv, pp. 504-505, pls. v-vii.
The British Willow Tit (Parus atricapillus kleinschmidtı Hellmayr).
British Birds, i, pp. 44-47.
Baer’s Pochard as a British bird. L.e. i, pp. 63, 64.
A new species of Parrot: Conurus canibuccalis, pp. 48, 49, Humaytha
R., Madeira.— Pipra nattereri from the Rio Madeira, and Cinclus
schulzi from Tucuman, p. 49.—On Palaeornis intermedia, P. schisticeps,
and P. cyanocephala, pp. 49, 50.—On the threatened extinction of the
breeding home of birds on Ailsa Craig, p. 56.—A very rare Parrot:
Urochroma dilectissima Sclater, among Bogota skins, p. 77.—A new
Bird-of-Paradise : Lophorina minor latipennis, Rawlinson Mts., German
N. Guinea, p. 92. Bull. B.0.C. xix.
On Carpodacus incertus Grant, p. 9.—On a new species of Bullfinch :
Pyrrhula owstoni, pp. 9, 10, Formosa (E. Hartert)—On Calophasis
mikado Grant, and other Pheasants, pp. 22-25.—An unknown Bird-of-
1908.
1909.
1910.
‚ao,
NOVITATES ZOOLOGICAE XLI. 1938.
Paradise belonging to a new genus: Pseudasirapia (gen. n.) lobata,
p. 25, Dutch New Guinea.—On Loborhamphus nobilis and Janthothorax
mirabilis, p. 25.—Exhibition of eggs of Birds-of-Paradise, p. 36.
L.c. xxi.
The birds of Vella Lavella, Solomon Islands.
Nov. Zool. xv, pp. 351-358, pl. xiii. (E. Hartert.)
On a collection of birds from San Christoval, Solomon Islands.
L.c. xv, pp. 359-365. (E. Hartert.)
Note on Casuarius casuarius bistriatus Oort. L.c. xv, p. 392.
Exhibition of Algerian bird-skins, p. 6.—A new species of the genus
Lioptilus: L. rufocinctus, pp. 6, 7, Rugege Forest.—On Drepanornis
albertisi geisleri A. B. Meyer, p. 7.—A new species of Crombec : Sylvietta
neumanni, p. 42, West of L. Tanganyika.—On Parotia berlepschi and
P. carolae, p. 42. Bull. B.O.C. xxii.
Note sur le Gypaéte et les Aigles criards. Rev. Fr. Orn. i, pp. 105, 106.
Unusual birds in Hertfordshire. Brit. Birds, ii, p. 309.
Description of a new bird from Africa. Ibis, pp. 690, 691, pl. x.
A new species of Weaver-Finch : Lagonosticta graueri, p. 102, N.W. of
L. Tanganyika.—A new form of Wryneck: Iynx torquilla mauretanica,
p. 103, N. Algeria.—On the Little Owl, p. 104. Bull. B.0.C. xxii.
A new species of Tree-Partridge : Arboricola rolli, pp. 7, 8, Battak Mts.,
Sumatra.—Pieces of the egg-shell of a Struthious bird from the South
Algerian desert, and egg of Megalapteryx huttoni, pp. 8, 9.—A new
Sunbird: Aethopyga seheriae owstoni, p. 32, Nauchau I., S. China.
Lic. xxv.
Notes on Eagle-Owls. Nov. Zool. xvii, pp. 110-112. (E. Hartert.)
On Caprimulgus aegyptius and C. aegyptius saharae, p. 73.—Eggs col-
lected in Algeria, pp. 74, 75.—On Perdix montana Brisson, p. 81.—On
white-breasted Cormorant, p. 89. Bull. B.0.C. xxv.
A new form of Amblyornis: A. subalaris germanus, pp. 13, 14, Rawlinson
Mts., German N. Guinea.—On Cuculus jacksoni Sharpe and Corythornis
cyanostigma, p. 14.—On a new Bird-of-Paradise from the Snow Mts. in
Dutch N. Guinea: Parotia carolae meeki, p. 35; and on Paradisea
mariae Reichenow, p. 36. L.c. xxvii.
Uber einige neue Formen vom Schneegebirge in Neuguinea.
Ornith. Monatsb. xix, pp. 157, 158. (E. Hartert.)
Hine neue Form von Pachycephala. L.c. xix, p. 178. (E. Hartert.)
Preliminary descriptions of some new birds from Central New Guinea.
Nov. Zool. xviii, pp. 159-160. (E. Hartert.)
Ornithological explorations in Algeria,
L.c. xviii, pp. 456-550, pls. ix-xi, xv-xxv. (E. Hartert.)
On recently described Paradiseidae, with notes on some other new species.
Ibis, pp. 350-367, pls. v, vi.
On the former and present distribution of the so-called Ratitae, or ostrich-
like birds, with certain deductions and a description of a new form by
C. W. Andrews. Verhandl. v. Intern. Ornith. Kongr. Berlin,
1910, pp. 144-174. (C. W. Andrews.)
Preliminary descriptions of some new birds from Central New Guinea.
Nov. Zool. xviii, pp. 159, 160. (E. Hartert.)
NoVvITATES Zoonogican XLI. 1938. 27
1912.
1913.
A new Petrel and a new Spider-hunter; Puffinus obseurus atlanticus, p. 43,
near Madeira; and Melilestes novae-guineae flaviventris, p. 44, Aru Is.
(E. Hartert).—Two new Parrots: Charmosynopsis multistriata, p. 45,
Oetakwa R., and Nasiterna pygmaea viridipectus, pp. 45, 46, Oetakwa
R.—On Mr. Meek’s Oetakwa Expedition, p. 46. Bull. B.O.C. xxvii.
Two new species of Honey-eaters : Melirrhophetes foersteri and Melipotes
ater, pp. 12, 13 (E. Hartert).—A new species of Thickhead from New
Guinea: Pachycephala tenebrosa, p. 20.—Some new birds from New
Guinea, pp. 33-35 (E. Hartert).—Cyclopsitta blythi meeki, the same
as Oyclopsittacus godmani, p. 35 (E. Hartert)—On three races of
Pristorhamphus, pp. 35, 36 (E. Hartert). IL, ZOaIbx,
On some newly described Birds-of-Paradise, and some undescribed eggs
of the same group. Ibis, pp. 109-112, pl. ii.
Ornithological explorations in Algeria. List of birds collected and
observed. Nov. Zool. xviii, pp. 471-550. (E. Hartert.)
List of a collection of birds made by Mr. Albert S. Meek on the Kumusi
River, North-Eastern British New Guinea.
L.c. xix, pp. 187-206. (E. Hartert.)
List of birds collection by Mr. A. S. Meek at Haidana, Collingwood Bay,
in North-Eastern British New Guinea.
L.c. xix, pp. 207-209. (E. Hartert.)
Remarks on Cassowaries and description of a new species: Casuarius
keysseri, Rawlinson Mts., German N. Guinea, pp. 49-52.
Bull. B.0.C. xxix.
Alteration of plumage due to derangement or injuries to the genital
organs, pp. 52-58. L.c. xxix.
A new subspecies of Albatross: Diomedea culminata mathewsi, p. 70,
Campbell I., New Zealand. — Exhibition of birds’ eggs from New Guinea,
p. 117.—Exhibition of and remarks on skins of Birds-of-Paradise and
Parrots, pp. 117, 118. L.c. xxix.
A new Weaver-finch from Madagascar: Foudia omissa, pp. 26, 27.
L.c. xxxi.
On some Australian forms of T'yto.
Nov. Zool. xx, pp. 280-284. (E. Hartert.)
List of the collections of birds made by Albert S. Meek in the Lower
Ranges of the Snow Mountains, on the Eilanden River, and on Mount
Goliath, during the years 1910 and 1911.
L.c. xx, pp. 473-527. (E. Hartert.)
Obituary notices of W. B. Tegetmeier, H. J. Pearson, E. A. Wilson, and
R. Collett, p. 44.—Melanistic specimens of Pheasants and of some other
birds, albinistic birds from his collection, pp. 45-51.—Exhibition of
and remarks on hybrids exhibited from his collection, pp. 51-54.—
Remarks on eggs and birds from Algeria, pp. 106, 107.
Bull. B.0.C. xxxi.
Remarks on various Cassowaries, pp. 34, 35.—Remark on Chelidon rufula
and Ch. smithi, p. 42.—Chairman’s Address, pp. 52-56.—Remarks on
the genus Bradyornis and description of a new subspecies: Br. pallidus
sharpei, Abyssinia, pp. 65, 66.—A new species of Cassowary : Casuarius
foersteri, p. 66, S.E. New Guinea.—Remark on change in colour of
1914.
1915.
1916.
NOVITATES ZOOLOGICAE XLI. 1938.
plumage and on Müllerian mimicry, p. 67.—On immature examples of
Accipiter gularis and of three species of Birds-of-Paradise, pp. 74, 75.—
On the correct name of Montagu’s harrier, pp. 75, 76. L.c. xxx.
Notes on some Conurus of the ‘“ emops ” group.
Avicult. Mag. v, p. 153.
On a collection of birds from Goodenough Island.
Nov. Zool. xxi, pp. 1-9. (E. Hartert.)
List of a small collection of birds from the Aicora River.
L.c. xxi, pp. 10-12. (E. Hartert.)
A zoological tour in West Algeria. List of birds collected and observed.
L.c. xxi, pp. 180-204, pls. vii, vii. (E. Hartert.)
On the birds of Rook Island, in the Bismarck Archipelago.
L.c. xxi, pp. 207-218. (E. Hartert.)
On a new form of Rhea. L.c. xxi, p. 223. (C. Chubb.)
The birds of the Admiralty Islands, north of German New Guinea.
L.c. xxi, pp. 281-298, pl. x. (E. Hartert.)
Birds from the Admiralty Is., with descriptions of new species and
subspecies, pp. 104-109 (E. Hartert).—Notes on Algerian birds,
p- 140.—Note on Chersophilus duponti, p. 141.—Remarks on the nesting
habits of Passer simplex in Algeria, p. 143. Bull. B.O.C. xxxiii.
Chairman’s Address, pp. 2-5.—A new subspecies of Cassowary from Jobi
I.: Casuarius papuanus goodfellowi, pp. 5-7.—A new species of
Goshawk from New Guinea: Accipiter ( Astur) eudiabolus, pp. 7, 8;
and note on a young female Goshawk from Halmaheira, p. 8 (E.
Hartert).—A new form of Kingfisher: Ceyx solitaria mulcata, pp. 24,
25 (E. Hartert).—A new form of Dicaeum: D. geelvinkianum rosseli,
p- 32, Rossel I. (E. Hartert),—Remarks on Halcyon nigrocyanea,
p- 33 (E. Hartert). L.c. XXXv.
The birds of Dampier Island. Nov. Zool. xxii, pp. 26-37. (E. Hartert.)
The birds of Vulcan Island. L.c. xxii, pp. 38-45. (E. Hartert.)
Notes on Papuan birds. L.c. xxii, pp. 46-60. (E. Hartert.)
On the Genus Fregata. L.c. xxii, pp. 145, 146.
Note on the genus Sula, pp. 41-45, with description of a new subspecies :
S. dactylatra californica, p. 43.—Varieties of Common Partridge, p. 45.
Bull. B.0.C. xxxv.
Chairman’s Annual Address, pp. 29-34.—Remarks in a discussion on the
bearing of Oology on classification, p. 20.—Remarks on breeding in
immature plumage of various birds, pp. 40, 41. L.c. xxxvi.
On some forms of Coracina from the Solomon Islands.
Nov. Zool. xxiii, pp. 289-291. (EK. Hartert.)
A new Monarcha from Rossel Island. L.c. xxiii, p. 297. (E. Hartert.)
On Zdolvisoma incertum (Meyer) and HE. meyeri Meyer, p. 58.—Note on
Scolopax saturata Horsf. and Scolopax saturata rosenbergi Schl., p. 66.—
Descriptions of and remarks on 5 new Flycatchers from the Solomon
Is., pp. 72-74 (E. Hartert)—On Scolopax saturata Horsf., and its
subspecies rosenbergi Schleg., pp. 86, 87. Bull. B.0.C. xxxvi.
Chairman’s Annual Address, pp. 2, 3.—On Crex porzana, p. 3.—A new
Flycatcher from Australia: Microeca flavigaster laetissima, p. 4,
Queensland. L.c. xxxvii.
NOoVvITATES ZOOLOGICAE XLI. 1938. 29
1917.
1918.
1919.
1921.
Review of the forms of Lalage karu inhabiting the Papuan subregion,
pp. 15-17 (E. Hartert).—On Tyto arfaki Schleg., pp. 17-19.—A new sub-
species of Myzomela: M. eichhorni interposita, p. 38, New Georgia and
Rendova (Hartert).—On the status of Lophophorus impejanus Lath. and
L. refulgens, pp. 49-51.—Remarks on Oenianthe, p. 57. L.c. xxxvü.
Annual address of the Chairman, pp. 1-3.—A new subspecies of Game-
bird from Borneo: Melanoperdix nigra borneensis, pp. 3, 4.—Remarks
on African Pigeons, with description of a new subspecies, Turtur afer
slateri, pp. 25, 26. L.c. xxxviii.
A few additional notes on the birds of Rossel Island, Louisiade group.
Nov. Zool. xxv, pp. 311, 312. (E. Hartert.)
Further notes on the birds of Sudest Island, or Tagula, in the Louisiade
group. L.c. xxv, pp. 313-326. (EK. Hartert.)
Note on Turtur afer slateri, pp. 37, 38.—On an Eagle-Owl from North
Algeria, pp. 38, 39.—Remarks on a recently described Albatross, p. 39.—
A curious Rhipidura from Kapaur, Western New Guinea, pp. 57, 58.—
A new form of Lioptilus: L. abyssinicus ansorgei, p. 78, Benguela.
—Remarks on the status of the Wildfowl on the Tring Reservoirs,
pp. 79, 80. Bull. B.0.C. xxxviii.
Remarks on the picture of the White Dodo by Pieter Witthoos, pp. 16, 17.
—Note on Crossoptilon mantchuricum, p. 17. Luc. xxxix.
On one of the four original pictures from life of the Réunion or White
Dodo. Ibis, pp. 78, 79, pl. ii.
Remark on the Black-necked Grebe, p. 34.—On a new subspecies of
Ostrich: Struthio camelus syriacus, and a key to the males and eggs of
the five known forms of Ostriches, pp. 81-83. Bull. B.O.C. xxxix.
On New Guinea Parrots of the desmaresti group of the genus Opopsitta,
pp. 64-67.—On a new species of Blood-Pheasant: Ithaginis clarkei,
pp. 67, 68, N.W. Yunnan.—Exhibition of eggs of Paradiseidae and
Corvidae, p. 123.—On a model of Dinornis maximus Owen, p. 133.—
Note on /frita kowaldi (de Vis), pp. 134, 135. Luc. xl.
Two new birds from Asben: Myrmecocichla buchanani and Anthus
sordidus asbenaicus, p. 33. Luc. xli.
On the correct name of D’Aubenton’s “ Manucode a Bouquets.”
Ibis, p. 465.
On a collection of birds from West-Central and North-Western Yunnan.
Nov. Zool. xxviii, pp. 14-67.
On some birds from the Weyland Mountains, Dutch New Guinea.
L.c. xxviii, pp. 280-294.
A new species of Snipe from the New Zealand region: Coenocorypha
aucklandica iredalei, p. 63.—A new Thrush from Ceram: Turdus
joiceyi, pp. 74, 75 (E. Hartert)—On Oreoibis akleyorum Chapm.,
p. 111.—A new subspecies from Buenos Aires: Thinocorus apoda
venturii, p. 111.—A new species of Paradisea: P. mixta, p. 127, no loc.
—On Paradisea apoda granti, p. 127.—A new Bird-of-Paradise :
Paradisea apoda subintermedia, pp. 138, 139.—Exhibition of eggs of
Struthious birds, pp. 149, 150. Bull. B.0.C. xli.
Exhibition of eggs of Alcididae, Tetraonidae, and Phasianidae, p. 34.
L.c. xlii.
1924.
NOVITATES ZOOLOGICAE XLI. 1938.
On the names of certain Parrots of the genus Larius Bodd. = Zelectus
Wagl. Ann. Mag. N.H. (9) ix, pp. 411-412.
Note on egg of Struthiolitus, pp. 71, 72.—Exhibition of eggs of Larks
and Pipits, pp. 80, 81.—Tanisiptera danae intensa subsp. nov., p. 91,
New Guinea.—On an egg of Struthiolithus from Honan, p. 123.
Bull. B.O.C. xlii. (E. Hartert.)
Descriptions of (11) new Yunnan birds, pp. 9-12.—A new subspecies of
Flycatcher: Hmpidornis semipartitus orleansi, p. 45, Upper Nile-—On
two very rare Owls from South Abyssinia, pp. 45, 46.—On an example
of Asio capensis (Smith), pp. 45, 46.—A white variety of Scopus
umbretta bannermani Grant, p. 46. L.c. xliü.
On a second collection sent by Mr. George Forrest from N.W. Yunnan.
Nov. Zool. xxx, pp. 33-58.
An Ornithological Autumn Journey to Algeria.
L.c. xxx, pp. 79-88. (E. Hartert.)
On a third collection of birds made by Mr. George Forrest in N.W.
Yunnan. ' Lc. xxx, pp. 247-267.
The Reading Sewage Farm as a possible bird sanctuary, p. 83.—Remarks
on intermediate specimens between Paradisea apoda raggiana and P. a.
novaeguineae, p. 115.—A new bird from Buru: Madanga (gen. n.)
ruficolus, p. 117 (EK. Hartert)—On sporadic and racial Hybrids,
pp. 126-129.—Crossoptilon crossoptilon crossoptilon, p. 129, N.W.
Yunnan.—On a new Courser from Fuerteventura: Cursor cursor
bannermani, pp. 165, 166. Bull. B.0.C. xliii.
Exhibition of Chats’ eggs from Rumania and North Africa, pp. 1, 2 (E.
Hartert).—Exhibition of and remarks on eggs of Balaeniceps, Phoeni-
copterus, Megalornis, and many other birds, pp. 9-11 (E. Hartert).—
Eggs of European Titmouses and other birds, pp. 15-17 (E. Hartert).
Bull. B. Ool. Assoc. i.
Notes on Lophocerus jacksoni Grant and L. deckeni (Cab.)
Ann. Mag. N. H. London (9), xiv, p. 317.
Birds from St. Matthias I., with descriptions of seven new forms, pp. 50-
53 (E. Hartert).—On a British-killed specimen of Cwursorius cursor,
p- 93. Bull. B.0.C. xliv.
A new species of Lory and of Honey-eater from New Ireland : Domicella
albidinucha and Philemom eichhorni, pp. 7, 8 (EK. Hartert).— Centropus
violaceus from New Ireland, p. 9.—A specimen of the “ Hairy Moor-
hen,” p. 9. L.c. xlv.
Exhibition of and comments on eggs of Pittidae, Caprimulgidae, Podar-
gidae, Steatornithidae, Hurylaemidae, and Cuckoos, pp. 33-36.—Exhibi-
tion of eggs of Charadrius, Thinornis, Lymnocryptes, Capella and
Coenocorypha and remarks on some of them, pp. 47, 48.
Bull. B. Ool. Assoc. i.
On a fourth collection of birds made by Mr. George Forrest in N.W.
Yunnan. Nov. Zool. xxxii, pp. 292-313.
Muscicapa blythi renamed M. collini, p. 90. Bull. B.0.C. xlv.
A new form of Cassowary: Casuarius casuarius lateralis, p. 30, north
coast to North-East N. Guinea.—On Oenanthe seebohmi near Biskra,
p. 45. L.c. xlvi.
Noyirares ZOOoLOGICAE XLI. 1938. 31
1927.
1928.
1929.
Exhibition of eggs of larger Limicolae and remarks on some of them,
p. 58. Bull. B. Ool. Assoc. i.
On the Avifauna of Yunnan, with critical notes.
Nov. Zool. xxxiii, pp. 189-343.
Supplemental notes on the Avifauna of Yunnan. L.c. pp. 395-400.
Ein ausgestorbener Drepanidide : Sassius simplex Rothsch. & Hart.
Journ. Ornith. 74, pp. 386-388, pl. ix. (E. Hartert.)
On the variation in Centropus ateralbus (Less.), pp. 50, 51.—An appar-
ently extinct bird of the family Drepanididae: Sassius (gen. n.) simplex
sp. n., p. 51, Sandwich Is. (E. Hartert)—Two new species from New
Britain: Aceipiter luteoschistaceus and Turdus talaseae, p. 53 (E.
Hartert)—On various forms of Fulvetta and description of a new
subspecies: F. chrysotis forresti, p. 64, Yunnan.—Faked bird-skins,
p. 81. Bull. B.0.C. xlvi.
On One-wattled Cassowaries, pp. 26-28. L.c. xlvii.
Eggs of Vultures and Eagles, pp. 89, 90.—Eggs of Falcons and Buzzards,
with data, pp. 97-99.—Exhibition of Cranes’ eggs and remarks thereon,
pp. 117-119. Bull. B. Ool. Assoc. i.
Corrections and criticisms to the article on the Avifauna of Yunnan.
Nov. Zool. xxxiii, pp. 398-400.
Supplement to the Avifauna of Yunnan (Novitates Zoologicae, xxxiii,
pp. 189-343). L.c. xxxiv, pp. 39-45.
Remarks on the Carolina Parrot, Conuropsis carolinensis, p. 130.—
Exhibition of drawings and specimens of Melanistic forms of various
families of birds, and remarks thereon, pp. 138-143.—On pictures of
the White Dodo, p. 144. Bull. B.O.C. xlvii.
On two Humming-birds hitherto considered as hybrids, p. 41.
L.c. xlviii.
Exhibition of eggs of Oenanthe, various species and subspecies, pp. 36,
37 (E. Hartert)—Exhibition of eggs of Argusianus, Rheinardtius,
Numida, Carpococcyx, Cabalus, Tricholimnas, and other birds, with
remarks thereon, pp. 37, 38. Bull. B. Ool. Assoc. ii.
Wild hybrid between Mallard and Pintail.
Scottish Naturalist, pp. 89, 90.
A pure white example of the Red-necked Grebe, p. 73.—Description of a
new Cassowary : Casuarius rogersi, pp. 87, 88, no loc.—A new Ground-
Pigeon: Otidiphaps nobilis arvensis, p. 88.—Remarks on a series of
hermaphrodite and gynandromorphic birds, pp. 89, 90.—The egg of
the Flightless Rail from Inaccessible I., Tristan da Cunha, p. 121.—
Exhibition of 52 species of Rails and remarks thereon, pp. 122-124.
Bull. B.O.C. xIviii.
On a pale variety of the Kestrel and similar varieties of other birds,
pp. 4-9.—On a Great Auk and egg, p. 9.—A new race of Cassowary :
Casuarius bicarunculatus intermedius, pp. 9-10, Aru Is. ILC, llbz,
Exhibition of and short remarks on eggs of Calopezus, Tinamus, Crypturus
and many others, pp. 67-69.—Exhibition of eggs of Bustards and other
birds, with remarks thereon, pp. 80-82. Bull. B. Ool. Assoc. ii.
Ailuroedus buccoides and its subspecies.
Nov. Zool. xxxv, p. 59. (E. Hartert).
32 Novitates ZooLoGIcAE XLI. 1938.
Note on Lampribis rara and L. olivacea. L.c. xxxv, p. 81.
A British specimen of Turdus dauma aureus Holandre, p. 58, Suffolk.—
Exhibition of hybrid Ducks and remarks thereon, pp. 93-99.—Note on
Manucodia ater alter and a new subspecies: M.a. subalter, pp. 109, 110.
Bull. B.0.C. xlix.
Exhibition of drawings on Cassowaries and remarks thereon, pp. 5, 6.—
Remarks on Lämmergeier, p. 18.—On the three known species of
Xanthomelas, p. 33. L.c. 6.
On a series of eggs of various Rails from all over the world, pp. 103-106.—
Exhibition of eggs of Ragles, pp. 140, 141. Bull. B. Ool. Assoc. ii.
1930. On the breeding of the Sarus Crane (Grus antigone Linn.) at Tring.
Avicult. Mag. viii, pp. 89-91, 2 photographs.
Notes on the preceding article of Dr. Stresemann (Welche Paradiesvogel-
arten der Literatur sind Hybriden Ursprungs ?).
Nov. Zool. xxxvi, pp. 16-17.
Exhibition of Birds-of-Paradise and remarks thereon, pp. 38-40.—
Melanistic mutant of Common Pheasant, p. 40.—On his Sarus Cranes,
p. 67.—On three races of Turdus philomelos, p. 68.—The hitherto
undescribed eggs of the Tristan da Cunha Thrush and Diving Petrel
(Nezocichla eremita and Pelecanoides urinatrix dacunhae), p. 74.
Bull. B.0.C. 1.
On the hitherto unknown eggs of the Paradise-Crow (Lycocorax pyrrho-
pterus pyrrhopterus) and Phonygammus kereaudreni kereaudreni, p. 9.—
Exhibition of a series of male Pochard (Fuligula ferina) and remarks
thereon, p. 25. L.c. li.
On the eggs of Cicinnurus regius, p. 2.—Exhibition of eggs of Birds-of-
Paradise, pp. 2, 3. Bull. B. Ool. Assoe. iii.
1931. A preliminary review of the Mooruk (Papuanus bennetti) Group of
Cassowaries. Nov. Zool. xxxvi, pp. 181-183, pls. i, ii.
On a collection of birds made by Mr. F. Shaw Mayer in the Weyland
Mountains, Dutch New Guinea, in 1930.
L.c. xxxvi, pp. 250-276, pls. iii, iv.
Notes on Paradiseidae with a list of species, subspecies, and hybrids
exhibited at the Seventh International Ornithol. Congress.
Proc. 7th Intern. Ornith. Congr. Amsterdam, 1930, pp. 285-298.
Note on Nomenclature, p. 68.—A new subspecies : T'rugon terrestris mayri,
p. 69, Hollandia.—On a pair of American Wigeon, p. 99, North Uist.—
A new subspecies of Lark: Aethocorys personata intensa, p. 100,
N’Guasa Nyiro R. Bull. B.0.C. ii.
A new race of Cypseloides: C. fumigatus major, p. 36, Tucuman.— Two
hitherto undescribed eggs: Pseudorestes ferrugineus brevipennis and
Paradisea rubra, p. 37. L.e. li.
1932. Ona new Bird of Paradise. Ann. Mag. N.H. (10), x, 1932, p. 126.
Ornithologische Ergebnisse der Expedition Stein, 1931-32.
Nov. Zool. xxxvili, pp. 127-247 (partim: Paradiseidae, Psittaci,
Rallı).
A series of Pheasants bred in England and remarks thereon, pp. 80, 81.
— A new subspecies: Sylvia undata maroccana, p. 82, N.W. Marocco.—
Note on three Cassowaries, p. 82.—Perversion of plumage in Pheasants,
Novirates ZOOLOGICAE XLI. 1938. 33
1937.
1894.
1895.
1896.
1897.
1898.
1901.
1902.
1903.
1907.
1915.
1925.
1927.
1928.
1931.
1893.
p. 94.—On a specimen of Spathula clypeata, p. 102.—A correction: Sylvia
undata maroccana renamed S. u. tingitana, p. 105. Bull. B.0.C. lii.
On Casuarius uniappendiculatus rufotinctus, p. 238.—A curious variety
of the Blackbird at Tring Park, p. 238. L.c. iii.
Two new subspecies of Cassowaries: Casuarius papuanus shawmayeri,
p. 120, Kratke Mts., and C. casuarius grandis, p. 121 (2), North coast of
Dutch N. Guinea. L.e. lvii.
IV. REPTILIA.
On Giant Land Tortoises. Nov. Zool. i, pp. 676, 677, pl. xi.
Remarks and corrections relating to the living Giant Tortoise on Mauri-
tius. L.e. i, p. 690.
Further notes on Gigantic Land Tortoises. L.c. ü, p. 483.
Further notes on Gigantic Land Tortoises. L.c. ii, pp. 85-91.
Further notes on Gigantic Land Tortoises. L.c. iv, pp. 407, 408, pl. xiii.
Note on Testudo sumeiret. L.c. vi, pp. 359, 360.
A new Land Tortoise from the Galapagos Is. L.c. viii, p. 372.
Further notes on the Fauna of the Galapagos Is.
L.c. ix, pp. 373, 374, photographs. (E. Hartert.)
Note regarding Testudo elephantopus. L.c. ix, p. 448.
Further notes regarding Testudo elephantopus. L.c. ix, p. 618.
Description of a new species of Gigantic Land Tortoise from the Galapagos
Is. L.c. ix, p. 619.
Description of a new species of Gigantic Land Tortoise from Indefatigable
I. L.c. x, p. 119.
Note on Uroplates fimbriatus lichenius subsp. nov.
L.c. x, p. 490, pls. iii, iv. (A. W. Head.)
A new species of Giant Tortoise. L.c. xiii, pp. 753, 754.
The Giant Land Tortoises of the Galapagos Is. in the Tring Museum.
L.c. xxii, pp. 403-417, pls. xxi-xxxii.
On the Gigantic Land Tortoises of the Seychelles and Aldabra-Mada-
gascar group with some notes on certain forms of the Mascarene group.
L.c. xxii, pp. 418-442, pls. xxiii-Ixxvi.
Giant Tortoises of the Galapagos and Mascarene Is.
Proc. Linn. Soc. Lond. p. 40.
The Giant Monitor Lizard. The Field, Jan. 1927, p. 115, fig. 1.
Exhibition of a mounted specimen of Varanus komodensis.
Proc. Zool. Soc. Lond. p. 256.
Notes on Gigantic Land Tortoises. L.c. pp. 658-660.
V. PISCES.
The record Bream for Britain. The Field, p. 629.
The largest Bream ever recorded. L.c. p. 897.
VI. INSECTA.
A. COLEOPTERA.
On two new species of the genus Hnoplotrupes.
Ann. Mag. N.H. (6), xii, pp. 36-38. (K. Jordan.)
On some new or little known species of Coleoptera from the Hast.
L.c. pp. 452-455. (K. Jordan.)
34
1894.
1892.
1893.
1894.
1895.
NOVITATES ZOOLOGICAE XLI. 1938.
Six new species of Plusiotis and one new Anoplostethus.
Nov. Zool. i, pp. 504-507. (K. Jordan.)
B. LEPIDOPTERA.
A collection of Lepidoptera from S.W. Celebes.
Proc. Ent. Soc. Lond. 1892, pp. 14-16.
Exhibition of and comments on Neptis mimetica n.s., Cynthia aequicolor !
n.s., and other Lepidoptera. L.c. p. 21.
On a collection of Lepidoptera from Timor, Pura, Sumba, and other
islands. L.c. pp. 21-23.
On a little known species of Papilio from the Island of Lifu, Loyalty
group. Trans. Ent. Soc. Lond. p. 141, pl. iv.
On a new and also on a little known species of Pseudacraea in the collection
of the Hon. Walter Rothschild.
L.c. pp. 201, 202, pl.x. (A. G. Butler.)
Notes on a collection of Lepidoptera made by William Doherty in Southern
Celebes during August and September 1891: J. Rhopalocera.
D. Ent. Zeits. Iris, v, pp. 429-442, pls. iv-vü.
Zwei neue Charaxes-Formen. L.c. vi, pp. 348-350.
Two new species of Lepidoptera from German New Guinea.
Ann. Mag. N.H. (6), xü, pp. 455-457. (K. Jordan.)
Descriptions of new Sphingide in the collection of Dr. Otto Staudinger.
D. Ent. Zeits. Iris, vii, pp. 297-302, pls. v—vii.
Exhibition of Ornithoptera paradisea Staudg. and other Lepidoptera.
Proc. Ent. Soc. Lond. p. 15.
On a new species of the Hepialid genus Oenetus.
Ann. Mag. N.H. (6), xiii, p. 440.
Notes on Sphingidae, with descriptions of new species.
Nov. Zool. i, pp. 65-98, pls. v—vii.
Some new species of Lepidoptera. L.c. i. pp. 535-540, pl. xii.
Additional notes on Sphingidae. L.c. i. pp. 541-543.
On five new Delias. L.c. i. pp. 661-662.
Additional notes on Sphingidae. L.c. i. pp. 664, 665.
On some new local races of Papilio vollenhovii Feld and Papilio hipponous
Feld. L.c. i. pp. 685-687.
On a new genus and species of Butterfly. L.c. i. p. 687.
On a new species of the Family of Sphingidae. L.c. ü, p. 28.
Notes on Saturnidae ; with a Preliminary Revision of the Family down
to the genus Automeris, and descriptions of some new species.
L.c. ii, pp. 35-51, pl. x.
Two new species of Rhopalocera from the Solomon Is.
L.c. ii, pp. 161, 162.
A new species of Theretra from the d’Entrecasteaux Is. L.e. ii, p. 162.
Revision of the Papilios of the Eastern Hemisphere, exclusive of Africa.
L.c. ii, pp. 167-463, pl. vi. (K. Jordan.)
Some new forms of the genera Bizarda and Milionia.
L.c. ii, pp. 464, 465. (K. Jordan.)
On two new Moths and an aberration. L.c. p. 482.
1 These names apparently have escaped notice.
NOVITATES Zoonoaroan XLI. 1938. 35
1896.
1897.
1898.
1899.
1900.
On Milionia and some allied genera of Geometridae.
L.e. ii, pp. 493-498, pl. vii.
Some notes on my Revision of the Eastern Papilios, exclusive of Africa.
L.e. ii, pp. 503, 504. (K. Jordan.)
On Copaxa multifenestrata (H.-S.) L.e. ii, p. 504.
Description of a new local form of Troides victoriae (Gray) from Bougain-
ville I., Solomon group. Entomologist, xxviii, pp. 78, 79.
Explanation of plates vii—x. Nov. Zool. ii (in front of pl. vii).
Two new Charazes from the Lesser Sunda Is.
Entomologist, xxix, pp. 308, 309.
Notes on Heterocera, with descriptions of new genera and species.
Nov. Zool. iii, pp. 21-62. (K. Jordan.)
Further notes on my Revision of the Papilios of the Eastern Hemisphere,
exclusive of Africa. L.c. iii, pp. 63-68.
New Lepidoptera. L.c. iii, pp. 91-99.
Notes on Heterocera, with descriptions of new genera and species.
L.c. pp. 185-208, pl. iv. (K. Jordan.)
Some undescribed Lepidoptera. L.c. iii, pp. 231, 232.
New Lepidoptera. L.c. iii, pp. 322-328.
On some new subspecies of Papilio. L.c. iii, pp. 421-425.
Descriptions of some new species of Lepidoptera, with remarks on some
previously described forms. L.c. ii, pp. 600-603, pls. xiii-xv.
On Eudaemonia brachyura Drury and E. argiphontes Kirby.
Proc. Ent. Soc. Lond. p. 37.
On a new species of Papilio from Uganda. Entomologist, xxx, p. 165.
Descriptions of some new species and subspecies of Lepidoptera.
Nov. Zool. iv, pp. 179-184.
Some new species of Heterocera. L.c. iv, pp. 307-313, pl. vii.
Notes on Heterocera, with descriptions of new genera and species.
L.c. iv, pp. 314-316, pl. iv. (K. Jordan.)
On some new Butterflies and Moths. L.c. iv, pp. 507-513.
On some new or rare Lepidoptera of the Old-World regions.
L.c. v, pp. 96-102.
Some new Lepidoptera from the East. L.c. v, pp. 216-219.
Some new Lepidoptera from Obi. L.c. v, pp. 416-418.
A Monograph of Charaxes and the allied prionopterous genera.
L.c. pp. 545-601, pls. v-xiva, text-figs. 1-38. (K. Jordan.)
Some new Lepidoptera from the East.
L.c. v, pp. 602-605, text-figs. 1-4.
Some new Eastern Lepidoptera. L.c. vi, pp. 67-71.
A Monograph of Charazxes and allied prionopterous genera (continuation).
L.c. vi, pp. 220-226, text-figs. 39-42. (K. Jordan.)
On some new Lepidoptera from the East.
L.c. vi, pp. 429-444. (K. Jordan.)
Two new species of Charaxes. Entomologist, xxxii, pp. 171, 172.
Description of the hitherto unknown female of Oenetus mirabilis.
Nov. Zool. vii, p. 24.
Some new or recently described Lepidoptera.
L.c. vii, pp. 274-276, pl. v.
36
1901.
1902.
1903.
1904.
1905.
1906.
1907.
NOVITATES ZOOLOGICAE XLI. 1938.
A Monograph of Charaxes and the allied prionopterous genera (continua-
tion). L.c. vii, pp. 281-524, pls. vi-viii, xi, xii. (K. Jordan.)
Some new Butterflies and Moths. L.c. viii, pp. 218-220.
On some Lepidoptera. L.c. viii, pp. 401-407, pls. ix, x. (K. Jordan.)
On some new or rare Lithosiinae. L.e. viii, pp. 409-424. (K. Jordan.)
Some new N.E. African Lepidoptera discovered by Oscar Neumann.
L.c. ix, pp. 595-598.
A Revision of the Lepidopterous Family Sphingidae.
L.c. ix, Supplement, pp. cxxxv and 972, 64 pls. (K. Jordan.)
Some new Cossidae from Queensland, bred by Mr. R. P. Dodd.
L.c. x, pp. 306-308.
Some new Butterflies and Moths. L.c. x, pp. 309-312.
A Monograph of Charaxes and allied prionopterous genera (conclusion).
L.c. x, pp. 326-342. (K. Jordan.)
Some new or unfigured Lepidoptera.
L.c. x, pp. 481-487, pls. xi, xii. (K. Jordan.)
Some new African Papilios. L.c. x, pp. 488, 489. (K. Jordan.)
Lepidoptera collected by Oscar Neumann in North-East Africa.
L.c. x, pp. 491-542. (K. Jordan.)
Lepidoptera from British New Guinea, collected by Mr. A. S. Meek.
A new form of Prothoé from the Solomon Islands. L.c. xi, p. 366.
New Sphingidae L.c. xi, pp. 435-440.
A new African Melanitis. L.c. xi, p. 451.
New forms of Butterflies. L.c. xi, pp. 452-455.
Two new Saturniidae. L.c. xi, p. 601.
A new subspecies of Troides victoriae. L.c. xi, p. 654.
Some undescribed Lepidoptera. L.c. xii, pp. 78, 79.
Lepidoptera collected by Oscar Neumann in North-East Africa (continua-
tion). L.c. xii, pp. 175-191. (K. Jordan.)
On some new Lepidoptera discovered by A. S. Meek in British New
Guinea. L.c. xii, pp. 448-478. (K. Jordan.)
Papilio steinbachi spec. nov. Entomologist, xxxviii, p. 125.
On a new parasitic Tineid Moth from Queensland, discovered by P. F.
Dodd. Nov. Zool. xiii, pp. 162-169. (P. F. Dodd.)
New Sphingidae. L.c. xiii, pp. 178-185. (K. Jordan.)
Two new Saturnidae. L.c. xii, p. 189.
Some new Sphingidae in the British Museum.
L.c. xiii, pp. 406, 407. (K. Jordan.)
A Revision of the American Papilios.
L.c. xiii, pp. 411-752, pls. iv-ix. (K. Jordan.)
New Sphingidae. L.c. xiv, pp. 92-96. (K. Jordan.)
Troides alexandrae sp. nov. L.c. xiv, p. 96.
New American Saturniidae and Ceratocampidae. L.c. xiv, pp. 413-342.
A new species of Sphingidae. L.c. xiv, p. 507.
Some new Himantopterinae, a subfamily of Zygaenidae.
L.c. xiv, pp. 507, 508.
Lepidopteren aus Neu-Guinea.
D. Ent. Zeits. Berlin, pp. 189-198. (K. Jordan.)
Novitares ZooLoGIcAE XLI. 1938. 37
1908.
1909.
1910.
1911.
1912.
New African Saturniidae. Ann. Mag. N.H. (7), xx, pp. 1-10.
Lepidoptera Heterocera, Fam. Sphingidae.
Wytsman, Gen. Ins. Fasc. 57, pp. 1-157, pls. i-viii. (K. Jordan.)
New forms of Oriental Papilios. Nov. Zool. xv, pp. 165-174.
Some new Sphingidae. L.c. xv, pp. 258, 259. (K. Jordan.)
New subspecies of Parnassius apollo. L.c. xv, p. 390.
A new Papilio from Africa. Ent. Mo. Mag. xlix, p. 249.
New Oriental Papilionidae. Entomologist, xli, pp. 1-4.
Description of a new form of Zygaena from Algeria. L.c. xli, p. 185.
Description of a new Lemonia. Ann. Mag. N.H. (8), iii, p. 7.
Descriptions of new South American Arctianae.
L.c. (8), iv, pp. 205-229
Mauretanisches. Ent. Zeits. Stuttgart, xxiii, p. 142
Catalogue of the collection of Parnasstiinae in the Tring Museum, with
systematic notes. Nov. Zool. xvi, pp. 1-20.
Descriptions of some new South American Arctiadae, with notes.
L.c. pp. 21-52, pls. iv-vü.
A new Aegerid. L.c. xvi, p. 132.
Descriptions of some South American Arctiadae, with notes.
L.c. xvi, pp. 268-299.
Catalogue of the Arctianae in the Tring Museum, with notes and descrip-
tions of new species. L.c. xvii, pp. 1-85, 113-171.
Descriptions of new species of Arctianae in the Tring Museum.
L.c. xvii, pp. 172-188.
Descriptions of Syntomidae. L.c. xvii, pp. 429-455.
List of the Sphingidae collected by the late W. Hoffmanns at Allianca,
Rio Madeira, Amazons. L.c. xvii, pp. 447-455. (K. Jordan.)
Some new Sphingidae. L.c. xvii, pp. 456-459. (K. Jordan.)
Notes on Arctianae and descriptions of a few new species.
L.c. xvii, pp. 504-506.
A new form of Attacus. L.c. xvii, p. 507.
Descriptions of anew Hawkmoth and some Syntomidae.
Ann. Mag. N.H. (8), v, pp. 506-511.
Some new Algerian Lepidoptera. L.c. (8), vii, pp. 231-233.
Two new Lepidoptera from New Guinea. L.c. (8), vili, pp. 233, 234.
A new Madagascar Arctiid. L.c. (8), viii, p. 255.
New Syntomidae in the Tring Museum. Nov. Zool. xviii, pp. 24-45.
New Aegerüidae. L.c. xvii, pp. 45-47.
New Syntomidae and Arctianae. L.c. xviii, pp. 154-158.
An apparently new form of Lambessa from Algeria. L.c. xix, p. 113.
A synonymic catalogue of the Syntomid genus Balacra Walk., ete., with
descriptions of new species. L.c. xix, pp. 119-122, pls. xiii, xiv.
New Bornean Aegeriidae and Syntomidae. L.c. xix, pp. 122-124.
New Mauretanian Moths. L.c. xix, pp. 125-127.
Some new Sphingidae. L.c. xix, pp. 128-135. (K. Jordan.)
New Syntomidae. L.c. xix, pp. 151-186.
New Lithosianae. L.c. xix, pp. 212-246.
Some unfigured Syntomidae, Aegeriadae, and Arctianae.
L.c. xix, pp. 375-377, pls. ii-v.
38
1913.
1914.
1915.
1916.
NOVITATES ZOOLOGICAE XLI. 1938.
Verbesserungen und Zusätze zu den Cossiden.
Seitz, Grossschmett. ii, pp. 450-452.
Expedition to the Central Western Sahara, vii, Lepidoptera.
Nov. Zool. xx, pp. 109-143.
Descriptions of two new Colias and some African Syntomidae.
L.c. xx, pp. 187, 188.
New Lithosianae. L.c. xx, pp. 192-226.
Some new Lepidoptera from German New Guinea.
L.c. xx, pp. 278, 279.
Descriptions of two new Syntomids. L.c. xx, p. 591.
Notes on Jxias undatus Butler, with the description of the unknown
female. L.c. xx, p. 279.
Expedition to the Central Western Sahara. Corrections to vii, Lepi-
doptera. L.c. xx, p. 469.
Some unfigured Syntomidae. L.c. xx, pp. 470-472, pls. xiii, xiv.
Descriptions of two new Syntomids. L.c. xx, p. 591.
A new species of T'roides. L.c. xxi, p. 262.
Description of a new Troides. L.c. xxi, p. 275.
A preliminary account of the Lepidopterous fauna of Guelt-es-Stel,
Central Algeria. Part I. L.c. xxi, pp. 299-357.
Arctiinae in Seitz. Seitz, Grossschmett. x, pp. 236-263.
Lepidoptera collected by the British Ornithologists’ Union and Wollaston
expeditions in the Snow Mountains, Southern Dutch New Guinea.
Macrolepidoptera, 168 pp., 2 pls. Issued at Zoological Museum, Tring.
On Lepidoptera from the Islands of Ceram (Seram), Buru, Bali, and
Misol. Part I. Nov. Zool. xxii, pp. 105-144.
A preliminary account of the Lepidopterous fauna of Guelt-es-Stel,
Central Algeria. Part 11. L.c. xxii, pp. 186-191.
On the Lepidoptera in the Tring Museum sent by Mr. A. S. Meek from
the Admiralty Islands, Dampier, and Vulcan Islands. Part I.
L.c. xxii, pp. 192-208. ,
On Lepidoptera from the Islands of Ceram (Seram), Buru, Bali, and
Misol. Part II. L.c. xxii, pp. 209-227.
Lepidoptera of the M’Zab country, South Algeria, collected by Dr. Ernst
Hartert and Carl Hilgert in 1914. L.c. xxii, pp. 228-243.
Notes on and descriptions of Delias. Ann.Mag.N.H. (8), xv, pp. 172-180.
On the Lepidoptera collected in 1913-1914 by Herr Geyr von Schweppen-
burg on a journey to the Hoggar Mts. Part I.
L.e. (8), xvi, pp. 247-258.
On the Lepidoptera collected in 1913-1914 by Herr Geyr von Schweppen-
burg on a journey to the Hoggar Mts. Part II.
L.c. (8), xvi, pp. 392-402.
Some new Sphingidae in the collection of the British Museum.
Nov. Zool. xxii, pp. 291-294. (K. Jordan.)
Thirteen new Sphingidae. L.c. xxii, pp. 281-290.
Some new Lepidoptera from Siam.
Journ. N.H. Soc. Siam, ii, pp. 74, 75.
Corrections of and additions to our Revision of Sphingidae.
Nov. Zool. xxiii, pp.115-123. (K. Jordan.)
Noviratres ZooLOGICAE XLI. 1938. 39
1916.
1917.
1918.
1919.
Further corrections of and additions to our Revision of the Sphingidae.
L.e. xxiii, pp. 247-263. (K. Jordan.)
Some new Arctiadae. L.c. xxiii, pp. 264-271.
Some notes of a faunal and other nature on the Lepidoptera collected by
Herr Geyr von Schweppenburg in the Hoggar Mountains.
L.c. xxiii, pp. 287-288.
Notes on Amathustidae, Brassolidae, Morphidae, ete., with descriptions
of new forms. L.c. xxiii, pp. 229-318, pls. ii-vi.
On the Lepidoptera in the Tring Museum sent by Mr. A. S. Meek from
the Admiralty Islands, Dampier, and Vulcan Islands. Part III.
L.e. xxiii, pp. 319-334.
Some new Lepidoptera from Siam and Africa.
Ann. Mag. N.H. (8), xvii, pp. 474-476.
Supplemental notes to Mr. Charles Oberthür’s “ Faune des Lepidopteres
de la Barbarie,’” with lists of the specimens contained in the Tring
Museum. Part I. Nov. Zool. xxiv, pp. 61-220.
Supplemental notes to Mr. Charles Oberthür’s “ Faune des Lepidopteres
de la Barbarie,’ with lists of the specimens contained in the Tring
Museum. Part II. L.c. xxiv, pp. 325-373.
Supplemental notes to Mr. Charles Oberthür’s “ Faune des Lepidopteres
de la Barbarie,’” with lists of the specimens contained in the Tring
Museum. Part III. L.e. xxiv, pp. 393-409.
Some apparently new Notodontidae.
L.c. xxiv, pp. 231-264, pls. ii-vii.
Notes on Metanastria digramma, with descriptions of two new subspecies.
L.c. xxiv, p. 453.
On the genera Melanothrix, Drepanojana, Melanergon, Paracydas, Cotana,
Hypercydas, Epicydas, and Nervicompressa, of the family Eupterotidae,
with descriptions of new forms. L.c. xxiv, pp. 463-474, pls. xi, xil.
Some new Moths of the families Arctiidae and Hupterotidae.
L.c. xxiv, pp. 475-492.
Catalogue of Zerynthiinae and allied genera in the Tring Museum, with
critical notes. L.c. xxv, pp. 64-75.
Catalogue of the Parnassiinae in the Tring Museum.
L.c. xxv, pp. 218-262.
On a new species of Stichophthalma. L.c. xxv, p. 310.
Some apparently new forms of African Nymphalidae.
L.c. xxv, pp. 338-345.
A new species of Ogoa. L.c. xxv, p. 365.
Explanation of Plates I and III (Lepidoptera discussed in vol. xxiv).
L.c. xxv.
Supplementary notes to the Review of Houlbert and Oberthür’s Mono-
graph of Castniinae by Talbot and Prout. L.c. xxvi, pp. 1-27.
List of types of Lepidoptera in the Tring Museum. Part I: Sphingidae.
L.c. xxvi, pp. 193-251.
Algerian Lepidoptera. (Explanation of figures on Plate I.)
L.c. xxvi, pp. 356-357.
Descriptions of a new genus and two new species of Heterocera.
Ann. Mag. N.H. (9), iii, pp. 487, 488.
1921.
1922.
1925.
1926.
NOVITATES ZOOLOGICAE XLI. 1938.
Heterocera collected in Korinchi, West Sumatra, by Messrs. H. C.
Robinson and C. Boden Kloss.
Journ. Fed. Malay States Mus. viii, pp. 107-139.
Rhopalocera collected in Korinchi, West Sumatra, by Messrs. H. C.
Robinson and C. Boden Kloss. L.c. viii, pp. 141-163.
Supplemental notes to Mr. Charles Oberthiir’s “ Faune des Lépidoptéres
de la Barbarie,”’ with lists of the specimens in the Tring Museum.
Nov. Zool. xxvii, pp. 1-127, pls. xiv—xvii.
On the genus Zlachyophthalma Feld. L.c. xxvii, pp. 321-326.
Preliminary descriptions of some new species and subspecies of Indo-
Malayan Sphingidae. Ann Mag. N.H. (9), v, pp. 479-482.
Parthenogenesis in two generations of Lasiocampa quercus.
Proc. Ent Soc. Lond. p. 54.
Exhibition of and remarks on Gynandromorphous Lepidoptera.
L.c. pp. 10-12.
Moths of Mesopotamia and N.W. Persia. PartI: Noctwidae, Lemoniidae,
and Pyralidae. Journ. Bombay N.H. Soc. xxviii, pp. 170-183.
Captain Angus Buchanan’s Air Expedition. V: Lepidoptera collected
in Northern Nigeria and the Southern Sahara in 1919-1920. Part I.
Nov. Zool. xxviii, pp. 142-170.
Captain Angus Buchanan’s Air Expedition. V: Lepidoptera collected
in Northern Nigeria and the Southern Sahara in 1919-1920. Part II.
L.c. pp. 215-229.
A preliminary list of the Arctiinae of Para, Brazil, and a few species
from other localities. Ann. Mag. N.H. (9), ix, pp. 457-494.
On a new form of Epinephele from Cyrenaica. Luc. (9), ix, p. 668.
A correction. L.c. (9), ix, p. 692.
On mimetic Lepidoptera and Hymenoptera.
Proc. Ent. Soc. Lond. p. 22.
Exhibition of Teinopalpus aureus and T. imperialis. L.c. p. 2.
Some new or noteworthy Madagascar and African Heterocera.
Ann. Mag. N.H. (9), xiv, pp. 306-317.
Descriptions of new forms of Delias, with some notes on others.
L.c. (9), xv, pp. 671-677.
On three North African Zygaenas. L.c. (9), xv, pp. 678-680.
List of Lepidoptera collected by Dr. E. Hartert and Herr Carl Hilgert in
Cyrenaica in March, April, and May 1922. L.c. (9), xvi, pp. 200-208.
A further note on Moroccan Zygaenas. L.c. (9), xvi, p. 269.
Description of a new species of Hligma (subfamily Sarrothripinae).
L.c. (9), xvi, p. 269.
Critical list of the collection of Algerian Lepidoptera of the late Captain
N. J. E. Holl. Nov. Zool. xxxii, pp. 195-229.
List of Lepidoptera collected from April to end of June 1925 by E. Hartert
and F. Young in Morocco.
Bull. Soc. Sci. Nat. Maroc. pp. 324-345.
On some African Papilios, with descriptions of new forms.
Ann. Mag. N.H. (9), xvii, pp. 112-114.
On some new species and subspecies of Milionia.
L.c. (9), xvii, pp. 114-116.
Noviratres Zoonoaicar XLI. 1938. 4]
1927.
1929.
1930.
1931.
1932.
1933.
1935.
1936.
1925.
On some varieties of Papilio machaon. Proc. Ent. Soc. Lond. p. 24.
On a new species of Delias from Mindanao.
Ann. Mag. N.H. (9), xix, p. 563.
List of Lepidoptera collected in Morocco in 1927 by Ernst Hartert and
Frederick Young. Nov. Zool. xxxv, pp. 220-24.
Lepidoptera collected in Morocco by Dr. E. Hartert in 1929.
L.c. xxxv, pp. 235-243.
List of Lepidoptera collected by Dr. Ernst Hartert and Mr. Frederick
Young in Morocco in 1930. L.c. xxxvi, pp. 191-200.
Description of the unknown 9 of Paraphrygia rectangulata Kenrick and a
new species of Phryganopteryx (Lepid. Arctiidae). L.c. xxxviii, p. 148.
Notes on Syntomidae (Amatidae) with descriptions of new species.
L.c. xxxvii, pp. 149-158.
On some new Hupterotidae. L.c. xxxvili, pp. 250-252.
On a collection of Lepidoptera from Spanish Morocco.
L.c. xxxviii, pp. 315-330.
On Opsiphanes orgetorix orgetorix Hew. and O. orgetorix magnalis (Stich.).
Ann. Mag. N.H. (10), x, p. 200
New species and subspecies of Arctiinae. L.c. (10), xi, pp. 167-194.
Some new Arctitinae. Nov. Zool. xxxix, pp. 239-250.
On a female of T'roides victoriae regis with male coloration.
Ann. Mag. N.H. (10), xvii, pp. 306, 307.
New species of Chionaema, Asura, and Miltochrista.
L.c. (10), xvii, pp. 482-483.
New species and subspecies of Zygaenodes.
L.c. (10), xvii, pp. 483-485.
New species and subspecies of Caprimima (Lithosvinae).
Nov. Zool. xl, pp. 1, 2.
VII. BOTANY.
The fertilisation of Ophris speculum, O. lutea, and O. fusca.
Orchid Rev. xxxiii, p. 99.
42 NOVITATES ZOOLOGICAE XLI. 1938.
CERCARIA SINITZINI: N. SP., A CYSTOPHOROUS CERCARIA
FROM PERINGIA ULVAE (PENNANT 1777).
By MIRIAM ROTHSCHILD.
(With 31 text-figures.)
IB 1936 I reported the occurrence of a cercaria closely allied to Cercaria sagittarius
Sinitzin 1911 in Peringia ulvae. Since the publication of that note I have
found another species pertaining to this group, which, however, is so rare that
there seems little if any prospect of solving the life-history. In consequence it is
deemed unnecessary to withhold the morphological description of this interesting
species until the publication of the writer’s fuller paper.
The cystophorous cercariae—the larvae of the Hemiuridae Lühe (Sinitzin
1905, Dollfus 1923, Krull 1935)—are among the most extraordinary and beautiful
of all known cercariae, and have never failed to excite the enthusiasm of those
helminthologists fortunate enough to discover them. Thirteen species have been
described, several of which have proved extremely rare, occurring only once or
twice in several thousand examples of the host.
Cercaria sinitzini greatly resembles C. sagittarius in general appearance and is
undoubtedly closely related to it. It can be distinguished instantly from this
species by the possession of an additional caudal appendage (see Table I, p. 42).
TECHNIQUE.
These cercariae are exceptional in that they are best studied when fixed
and stained, though certain characters, notably the excretory system, are only
visible during life. Two rediae and their contents were set aside and studied
alive. The remaining rediae were carefully opened and fixed on the slide with
Bouin Dubosq. Two slides were stained with Delafield’s haematoxylin and
another with paracarmine. The extreme anterior portion of the snail’s liver was
sectioned. The sections were cut at 8 u, and stained with Delafield’s haematoxylin
with a counterstain of eosin.
Measurements.—Measurements, unless otherwise stated, were made from
preserved material only. The size of the various appendages is very variable.
Table II is provided to afford a comparison between them and the developing
cercaria.
TABLE I.
C. SAGITTARIUS C. SINITZINI
1. Rediae not active. 1. Rediae very active.
2. No 5th caudal appendage. 2. 5th caudal appendage present.
3. Terminal portion of Appendage IV (= “Phry- 3. Terminal portion of Appendage IV (=‘‘Phry-
gian Cap’) not drawn out into long tapering gian Cap ”) drawn out into long tapering
point. point, longer than the main body of the
cap.
4. Linear arrangement of lateral cells of Append- 4. Spiral arrangement of central cells of Append-
age II (= “ Ribbon”). age II (= “ Ribbon’) and irregular arrange-
OR ment of lateral cells.
5. Encysted cercaria folded twice inside cyst, 5. Eneysted cercaria folded once inside cyst,
with extremities in upper half. with extremities in basal half.
1 T have named this species in honour of D. T. Sinitzin. The type is deposited at the British
Museum (Natural History), Cromwell Road, London.
NOVITATES ZOOLOGICAR XLI.
1938.
TABLE II
1
MEASUREMENTS OF APPENDAGES
in microns.
Body of Tail-vesicle |
Cercaria Appendage I |Appendage II Appendage IIL Appendage IV Appendage V
aspect
37 x 31 26 X 23 21x 6 12 x 10 |not developed | not developed |not developed
38 x 31 28 x 26 38 x 5 24 x 10 | not developed | not developed |not developed
44 x 31 33 x 31 45 xX 5 28 x 9 9 x 10 not developed |not developed
(at base)
44 x 33 31 x 30 38 x 4 28 x 9 not developed | not developed |not developed
45 x 30 43 x 33 79 x 3-5 37 x 9 broken off 26 x 16 [not developed
49 x 30 33 x 35 70 x 5 37 x 9 21 long 21 x 16 9 long
51 x 24 38 x 31 107 x 3 40 x 9 38 long 23 x 14 17 long
52 x 31 40 x 35 65 x 7 35 x 9 35 long 23 x 17 14 long
54 x 28 35 x 35 Broken 37 x 9-10 44 long 26 x 12 31 long
56 x 26 44 x 37 87 x7 44 x 9 40 long 24 x 16 19 long
56 x 38 40 x 35 61 long 43 x 9 17 long broken broken
57 x 40 40 x 35 52 long 26 x 9 8 long broken broken
59 x 28 40 x 37 115 x 8 58 x 8-9 37 long 23 x 12 17 long
60 x 30 |not measured) 129 x 2-5 45 x 8 40 long 32 x 12 16 long
(extended)
60 x 30 |not measured 113 x 8-9 45 x 8 40 long 8 3< Iz 16 long
(not extended)
63 x 25 52 x 48 104 x 5 44 x 9 43 long broken 23 long
12 x 23 49 x 35 122 x 3 45 x 9 49 long broken 28 long
(ventral
aspect)
73 x 23 42 x 31 56 x 7 49 x 9 31 long 38 x 16 17 long
(telescoped)
73 x 23 42 x 31 61 x 10 48 x 9 30 long 49 x 16 28 long
(telescoped)
79 x 18 43 x 35 52 26 x 9 not measured | not measured | not measured
(telescoped)
79 x 23 45 x 40 105 x 7 45 x 9 40 long 38 x 17 17 long
80 x 25 not measured 52 x 10 50 x 10 30 long 38 x 12 34 x 1-7
(telescoped)
82 x 23 50 x 40 51 50 x 9 35 long not measured 17 long
(withdrawing
into tail)
85 x 25 |not measured 53 x 11 48 x 9 35 long 36 x 16 broken
(telescoped)
87 x 21 46 x 38 33 49 long 35 long 23 long 17 long
(withdrawing
into tail)
87 x 24 52 x 44 58 x 10 Oil sz Y 30 long 35 x 13 22 long
(telescoped)
Cyst — — Appendage II Appendage III Appendage IV |Appendage V
43 x 43 — — 35 x 6 46 long not not
(lateral measurable | measurable
aspect)
39 x 39 — — 39 x 6 45 long not not
(lateral measurable | measurable
aspect) :
42 x 42 = == 40 x 6 40 long not not
(lateral measurable | measurable
aspect)
1 See Fig. 9 for lines of measurements.
Cercaria sinitzini n. sp.
The Redia.—Measurements.
Body:
Intestine :
Pharynx :
1,000 u x
125 u
33u X
The redia of this species is very distinctive, and can be
of a hand lens.
200 u. or less.
be)
AD, 35
29
be}
recognized with the aid
The body is cylindrical and a pearly white in colour.
The
44 NOVITATES ZOOLOGICAE XLI. 1938.
pharynx is well developed and leads into a short, bright yellow intestine about
3 the length of the body, which is very conspicuous in the living animal. Collar
and ambulacral processes are absent. A birth-pore is situated a little below the
pharynx. The pharynx is surrounded by digestive glands.
Owing to the scarcity of the
material the excretory system was
not worked out in detail. It is well
developed, and there are over four-
teen flame-cells on each side of the
body, which appear to be arranged
in groups of four.
A very characteristic feature of
the redia are its movements, and it
is this in conjunction with the pearly
white colour which gives these par-
thenitae such a distinctive appear-
ance. The body wall of the redia
is exceptionally muscular. When
liberated from the tissues of the
snail, the body is violently contracted,
first in one place, then in another.
Unlike the rediae of C. sagittarius
these rediae are very active. The
body cavity is packed to bursting-
point with cercariae in all stages of
development, including the encysted
forms. Over 100 were counted in
one redia. It seems probable that
the cysts are expelled through the
birth-pore by the contractions of the redia, as they themselves seem incapable
of movement. The rediae infest the gonad of the snail. Sections show the
liver to be comparatively undamaged. One infection was found in two
thousand specimens of Peringia ulvae taken from St. John’s Lake, Plymouth,
in April 1936.
The Cercaria.—Measurements of full-grown specimens :
Body (length) . ‘ ; : . 631-105 u (average 851).
Body (width) . 6 : . . 12p-25u (average 181).
Tail (length without appendages) . 43y-52u (average 48 u).
Tail (width from dorsal surface to ventral 35-42 (average 38 u).
surface) . B ;
Oral Sucker. : 2 : . 14u x l4u-15u x 4p.
Ventral Sucker . : ; , . Tex 7Tp-9e x Ow.
Pharynx . 3 : ; ; » OMS OM.
Peduncle . i : : : . 8u-9u.
The Body (fig. 4).—As in most Hemiurid cercariae the body is poorly differ-
entiated and stains in the manner of germ balls and very young individuals of
other species of cercariae. The whole animal is very transparent. The oral
sucker is markedly subterminal and leads into a pharynx, oesophagus and forked
Novirates Zoonoarcan XLI. 1938. 45
intestine. The caeca reach beyond the ventral sucker posteriorly. The ventral
sucker, situated at about the posterior third of the body, is poorly developed and
smaller than the oral sucker. The Y-shaped excretory bladder is terminal. ‘The
excretory vesicle (fig. 4) is formed by two lateral arms joining the bladder at the
anterior cornu, which pass parallel to each other
on either side of the ventral sucker and fuse
dorsally behind the pharynx. The formation of
the excretory vesicle or ““ closed circuit’ occurs
after the lateral excretory tubes have turned
posteriad (in the region of the ventral sucker)
and divided into an anterior-lateral and posterior-
lateral branch. Thus the general development of
the excretory system resembles that of ©. ephemera
Lebour 1907 (nee Nitzsch) (Rothschild 1936)
except in the very important point that the main
lateral excretory tubules only grow as far forward
as the acetabulum! before turning posteriad.
Owing to the undifferentiated condition of the
cercaria the finer branches of the excretory system
are very difficult to follow. If my observations are correct this species (and
probably all the Hemiuridae) is an exception to Dubois’ rule that cercariae with
the Mesostoma type of excretory system develop in sporocysts and not in
rediae (Dubois 1929).
The cuticle is unarmed and no cystogenous glands were observed.
A very striking feature of the body of the cercaria is the
investing primitive epithelium (figs. 1, 9, 9a). The nuclei
are arranged in pairs and are very conspicuous in stained
specimens—bulging out at intervals along the edge of the
body. This primitive epithelium is retained by the encysted
cercaria, and according to Pratt (1898) the metacercariae of
Hemiurids found in the body cavity of Copepods also retain
it for a considerable period. This fact strongly suggests that
any cystogenous material which may be required for closing
the orifice of the tail comes from the inside of the tail-vesicle
itself, and is not shed by the cercaria as suggested by Sinitzin.
A similar condition was recorded for ©. macrura Faust 1921.
The body of the cercaria is fixed in the anterior aperture
of the tail by a thin peduncle, consisting of four rows of single
cells. Early in development these arise from the base of the
ut inside of the tail-vesicle, close to the vorigin of Appendage 1
(fig. 7). The cells forming the original portion of the peduncle
degenerate early, so that only a few remain near the aperture
of the tail, forming the very short peduncle of the fully-
developed cercaria (fig. 2). The cercaria encysts by withdrawing into the tail.
The Tail—When viewed from the side the main body of the cystic tail is
shaped like a jug with a wide mouth (figs. 1, 2, 4,5). When viewed from directly
in front (fig. 3) or behind, it is egg-shaped. Almost all mounted specimens are
1 In the Azygiidae Odhner, the excretory system is of the Stenostoma type, and it seems likely
that this pseudo-Mesostoma excretory system of Cercaria sinitzini is a later development from the
former type—arising by a shortening of the lateral tubules.
3.
AL
46 NOVITATES ZOOLOGICAE XLI. 1938.
seen from the side, as the tail is compressed laterally and tends to assume this
position on the slide. The ventral half of the tail is narrower than the dorsal
half. The tail carries five appendages (fig. 1, 4).
Appendage I (the “ Arrow” of Sinitzin (1911), “ Exeretory Projection” of
Cort and Nichols (1922), Willey (1930), and the “ Delivery-Tube ” of
Krull (1933), arising from the interior floor of the tail and passing out
at the aperture, ventral to the body of the cercaria.
Appendage II (the “ Ribbon” of Sinitzin), attached externally to the
mid-ventral line of the tail, a short distance below the mouth.
Appendage III (the “ Sultan’s Plume” of Sinitzin), attached externally to
the centre of the mid-dorsal line of the tail.
Appendage IV (the “ Phrygian Cap” of Sinitzin), forming a prolongation of
the terminal portion of the tail, directed ventrad.
Appendage V, a thin outgrowth of the base of Appendage IV, directed distad
and posteriad.
The outermost layer of the tail consists of an exceedingly thin transparent
membrane which is most probably the primitive epithelium of which the nuclei
have degenerated. It invests
the vesicle proper and all the
appendages, at least in their
early stages. Below this is a
much thicker, hyaline, cuticular
layer staining a faint mauve
with Delafield’s haematoxylin.
This homogeneous layer forms
the cyst-wall. Posteriorly it
is sharply invaginated (figs. 1,
4) so that in lateral aspect the
base of the vesicular portion
of the tail is divided into
two asymmetrical halves. The
dorsal half is larger than the
ventral half both in breadth
and depth, and shows behind
the latter if the tail is viewed
from directly in front (i.e. ven-
trally). This invagination of
the cyst-wall seems more pro-
nounced than in ©. sagittarius.
Within the cyst-wall is a layer
of single cells with large oval
nuclei and granular protoplasm (figs. 2, 5). Quite early in development, before
the appearance of the cyst-wall itself, these can be traced clearly outlining its
future course, including the posterior indentation, etc. All these three layers
thin out, and are obliterated inside the aperture of the tail.
The internal structure of the tail-vesicle is extremely complicated, and I
1 Prior to encystment Appendages II and III are connected with the outer layers of the tail
only, but during some stage in development these appendages also are connected with the inner
cell-structure of the tail-vesicle.
Novirares ZOOLOGICAE XLI. 1938. 47
have been unable to gain a clear understanding of the whole organ and the
function of the various cells. There are several compact groups of cells, each
invested inathinmembrane. These give rise to Appendages II, IIT, the peduncle,
ete. Two other groups are placed near the base of Appendage I (fig. 2) and
these appear to be intimately connected with the membranous sheath which
develops round the base of this appendage after it is withdrawn into the tail-
vesicle. In addition there are numerous ungrouped cells, lying in different planes
and possessing various-sized nuclei, some of which may provide cystogenous
material. The connections between the various appendages and the groups of
cells within the tail-vesicle are lost at different times. A gradual degeneration of
all these cells takes place, but certain groups disintegrate before others.
By the time the cercaria is ready to withdraw into the cyst the inner structure
of the tail has changed considerably. The cyst-wall has thickened, particularly
at the aperture of the tail, where
it is at least double the thickness
of other regions. The nuclei of the tr
granular protoplasmic layer have
flattened out. The rest of the
cellular contents of the vesicle have
shrunk and disintegrated.
While Appendage I (the de-
livery-tube) is still growing (i.e.
before it has attained its maximum
length) the tail-vesicle is more or
less “solid.” The portion of the
delivery-tube within the vesicle is,
however, surrounded by a narrow
space. This cavity is bounded by
a stout membrane which can be
seen in sections (fig. 2). Although
at this stage it is not easy to
make out the precise relationships
of these parts, the membrane
lining the central cavity appears
to be continuous with the delivery-tube itself. By the time this appendage
has reached the “telescoping” and “withdrawing” stage (see below) the
cavity in the tail-vesicle has increased considerably in size. This appears to take
place as a result of the disintegration of the internal cell-structure of the tail
mentioned above. The innermost membrane expands with the cavity until it
forms a second lining to the cyst-wall. Thus when the cercaria withdraws, the
tail-vesicle is hollow, and it enters the cavity bounded by this inner membrane.
At this stage the tail-vesicle barely takes up any stain. ‘The cyst-wall and
the lining membrane, together with the membranes of two internal groups of
cells, are just visible, and here and there the remains of disintegrating nuclei
show up.
Appendage I (figs. 15-18a).—This appendage originates from a group of
cells on the floor of the inside of the dorsal half of the tail-vesicle, and, as already
mentioned, is homologous with the delivery-tube of Krull. Its function also
appears to be identical. It is differentiated at about the same time as the bud
48 NOVITATES ZOOLOGICAE XLI. 1938.
of Appendage II, but grows much faster. It arises as a somewhat thick double
row of cells, passing upwards and outwards by the future aperture of the tail-
vesicle, ventrally to the body of the cercaria. It consists of an inner core of
closely-packed cells, with their long axis at right angles to the long axis of the
appendage (fig. 15). The cell-walls are never very distinct. When the body of
the cercaria measures about 40 u in length, about thirteen of these cells, counted
by their nuclei, can be seen outside the body of the tail. This row of cells
terminates in a club-like swelling containing a group of numerous small deeply-
staining nuclei. At the extreme terminal portion of the swelling is one giant oval-
shaped nucleus. The appendage is invested with a delicate transparent mem-
brane which cannot be detected in later stages except in sections. Rarely, during
development, one of the nuclei pertaining to this membrane can be seen bulging
out from the surface of the appendage (fig. 15). When the appendage pushes out
from the tail-vesicle, it is surrounded by the most external cuticular or epithelial
layer of the tail. I have been unable to determine whether this layer is ruptured
or whether it grows with the appendage, adhering closely to its sides. The
appendage grows by the elongation of these cells outside the body of the tail
vesicle. These come to form a long ribbon, of single cells with their long axes
now parallel to the long axis of the appendage. The external cells still rarely
exceed 13 in number. The nuclei, particularly near the aperture of the tail,
tend to be grouped in pairs (fig. 1). At this stage the structure of the inside
of the tail-vesicle becomes less dense and the internal course of the append-
age can be seen more clearly (fig. 5). Here the long axis of the cells remains
transverse. Immediately above the posterior invagination of the tail-vesicle,
the appendage bends sharply forward and continues its course upwards in the
ventral half of the tail. At a stage when the body of the cercaria measures about
60 u-70 u. in length, the appendage is capable of considerable elongation and
contraction. It then measures 87 u-129 u in length and 2 u-9 u in width (figs.
16, 17).
When the cercaria reaches a length of approximately 73 u a marked change
occurs in the appendage. A general telescoping of the portion outside the tail-
vesicle takes place, so that either the top of one cell fits into the bottom of
the next, or the base of one fits into the top of the adjacent cell (figs. 16, 18a).
This brings the nuclei very much closer together and the appendage now
measures only 50 u-61 u. in length and 10y—11u in width. The small, deeply-
staining nuclei in the terminal bulb disintegrate. The giant terminal nucleus
also disappears. The cuticle of the whole appendage thickens somewhat, but
particularly that of the terminal bulb. The constriction between it and the rest
of the appendage is accentuated, forming a teat-shaped extremity. The entire
appendage now stains a pinkish mauve instead of a bluish purple. Vacuoles
and fibres become more noticeable in the protoplasm of the cells (fig. 18).
The onset of encystment is marked by the withdrawal of Appendage I into
the cavity of the tail. This does not appear to be the result of muscular con-
traction but due to the mechanical effect of the collapse of the basal, internal
portion of the appendage, weakened by the enlarging of the cavity inside the
tail-vesicle. The telescoped portion of the appendage is thus pulled down into
the inside of the vesicle, within the lining membrane. In C. sagittarius it appears
that the withdrawal of the appendage and the body of the cercaria coincide, but
in the case of C. sinitzini complete withdrawal of the appendage takes place first.
Novirarrs Zoonogican XLI. 1938. 49
It comes to lie in the base of the tail, irregularly coiled up, immediately above
and to the sides of the cuticular evagination. Once inside the tail-vesicle, a fairly
rapid degeneration of the nuclei and protoplasm of the cells takes place, so that the
outline of the whole ribbon can be made out only with difficulty. The cell-walls
remain intact and a hollow tube is formed. When Appendage I is thus retracted
within the lining membrane of the central cavity of the tail, the whole can be
roughly compared with a glove with a single long, inverted finger.
When Appendage I is fully everted during excystment (see below) it will be
seen that the basal third is considerably wider than the rest of the appendage,
and is invested by a voluminous ribbed sheath (figs. 27-29). This sheath is con-
tinuous with the lining membrane and is folded inside the tail-vesicle and cannot be
clearly distinguished prior to excystment. In shape and also in function it is very
reminiscent of the “ Handle ” appendage of both C. projecta and the cercaria of
Halipegus occidualis. It seems possible that is is homologous with that appendage.
Appendage IT (figs. 19-21).—The bud of this appendage is differentiated soon
after the constriction is formed between body and tail, and is seen as a com-
paratively large outgrowth on the mid-ventral surface of the tail (fig. 6). With
the enlargement of the vesicle it comes to occupy a place nearer the aperture of
the tail. The appendage when at the stage of maximum development (fig. 19)
is finger-shaped and attached to the tail by a short hyaline stalk, continuous with
the hyaline cyst-wall. It then measures 45 u-60 u. x Su-10 u. There is a large
central cavity, occupied by a column of about 32 closely-packed, single cells, of
which the walls and nuclei are very distinct. They are arranged in a loose spiral
which is sometimes lost distally, where the column is looped back on itself for lack
of space within the central cavity. The long axes of the cells are at right angles
to the long axis of the appendage.
50 Novitates ZootocicaE XLI. 1938.
Although the column lies in the centre of the cylinder, it is not closely invested
by the membranous outer wall of the appendage. This, unlike the external mem-
brane of Appendage I, appears to be a somewhat rigid structure. Its very con-
spicuous nuclei, about 13 in number, are scattered irregularly over the appendage,
either bulging out from the surface or bulging inwards into the concavities of the
spiral core (fig. 19).
In C. sagittarius it
appears that these
nuclei are arranged
regularly in 2 rows,
parallel to the cells
of the central column,
which here have no
spiral twist, but a
simple linear arrange-
ment. It is possible
that this character is
not specificand varies
from one infection to
another. The whole
appendage is in-
vested by the outer-
most epithelial layer
of the tail-vesicle.
During the earlier
stages of develop-
ment the core of the
appendage is con-
nected with the in-
side of the tail-
vesicle, but this con-
nection is severed
when the terminal
portion becomes con-
stricted and the short
hyaline stalk is
formed.
After the tele-
scoping of Append-
age I, degeneration
of Appendage II commences. The nuclei of the cells of the central core
degenerate first (fig. 20). The chromatin of the nucleus shrinks away from
the nuclear membrane, forming a dark undifferentiated mass in the centre of
the nucleus. It gradually disappears altogether, but even after encystment of
the cercaria the faint outline of the collapsed nuclear membrane of these cells
can be distinguished in stained specimens (fig. 21). The nuclei of the wall of
the appendage subsequently disintegrate completely, and the wall collapses,
remaining as a loose crumpled covering to the central column of cells (fig. 21).
The degeneration of the nuclei of all the appendages occurs in a similar
Novirates Zoonoagrcar XLT. 1938. 51
manner, but owing to the more definite structure of the cell-walls and nuclei of
Appendage II, it is most clearly seen in the case of the latter.
Appendage III (figs. 22—26).—This appendage is differentiated considerably
later than I and II, when the body of the cercaria already measures about 40 u.
in length (figs. 7, 8). About 16 cells, arising from one of the internal cell-groups
of the vesicle, situated in the centre of the mid-dorsal line of the tail, push
outwards in a flower-like formation. They are invested by the outer membrane
of the tail-vesicle. These cells gradually elongate, each nucleus remaining in the
distal portion of the cell. The elongated posterior portions become closely wound
together, eventually fusing and forming a homogeneous hyaline stalk continuous
with the thickened cyst-wall. Thus the connection between the appendage and
the inside of the tail-vesicle is lost, and the appendage now resembles a flower on
a stalk, the petal-like extremities of the cells having apparently broken free from
the investing membrane. It measures between 30 u and 50 u in length. During
and after the encystment of the cercaria the stalk elongates considerably and the
nuclei of the cells gradually degenerate. The terminal free ends become ragged,
and finally split into a great number of minute threads (figs. 26, 27). In this
condition the appendage remains attached to the cyst.
Appendage IV.—This forms a cap at the base of the cystic portion of the tail,
12 u-17 u broad at the base x 21 u-49 u in length. Its terminal portion is pro-
longed into a long tapering point, directed towards the ventral side. This taper-
ing point is as long as, or longer than, the upper portion of the cap (figs. 1, 14).
There is an internal column of 6-8 cells, in communication with the cells inside the
base of the cystic tail. The ventral portion of the cap is enlarged into a hump
just before the tapering extremity. A mass of nuclei can be seen inside this
52 Novirates ZooLocicaE XLI. 1938.
portion of the appendage, whereas in the dorsal half no nuclei are visible. The
outermost layer of the appendage is continuous with that of the cystic portion
of the tail. It is differentiated soon after Appendages I and II. Simultaneously
with the appearance of the bud of Appendage III, it can be seen asa hoop of
4-8 cells standing out from the main body of the tail, below Appendage II
(fig. 7). Simultaneously with the telescoping of Appendage I, the cells in the
cap begin to degenerate. The thickening of the cyst-wall of the tail severs the
connection with the central core of cells, which gradually disappear. The cells
in the ventral hump break down and form a spherical vacuolated mass with a
single disintegrated, deeply-staining nucleus in the centre. The tapering point
shrivels up. After the encystment
of the cercaria the appendage
remains attached to the outside
of the cyst, but consists of little
more than a structureless mass of
tissue.
It will thus be seen that
Appendage IV of C. sinitzini differs
considerably from that of C. sagit-
tarius (Table I, p. 42). The ter-
minal portion is greatly elongated,
being at least twice as long as in
©. sagittarius. There is also no
peripheral layer of cells in the
dorsal half of the appendage, but a
ventral “‘ hump ” which appears to
be absent in C. sagittarius.
In several specimens of C.
sinitzini, the terminal portion of
the cap is bifurcated (fig. 10) and
in other cases it bears a small
additional lobe at right angles to
the main prolongation.
Appendage V.—This is the last of the appendages to be developed. It is a
dorsal outgrowth of the base of Appendage IV (close to the cyst-wall of the tail-
vesicle). It is first seen as a giant nucleus pushing out at right angles to Append-
age IV (figs. 8, 9). When fully developed it consists of a thin homogeneous
hyaline stalk 16 u-34p in length, with a bulbous tip which contains the giant
nucleus measuring 4:5 x 2:3 (fig. 1). It is enveloped in the outermost
membrane of the tail-vesicle. After encystment this appendage also undergoes
considerable degeneration. The giant nucleus migrates back along the stalk
(fig. 5) and finally disintegrates at the base near Appendage IV. The stalk, ragged
and distorted in shape, remains attached to the cyst.
In some specimens an interesting abnormality was observed. A second giant
nucleus grows out at the base of Appendage V, forming a similar but much smaller
appendage at right angles to Appendage V, and parallel to Appendage IV.
Encystment of the Cercaria (figs. 10-13). The cercaria encysts by withdrawing
into its tail. As soon as encystment commences, the aperture of the tail-vesicle
draws together and it appears as if the body is pulled into the cavity by suction.
Novirares ZOOLOGICAE XLI. 1938. 53
The posterior portion of the cercaria descends into the dorsal half of the cyst,
rupturing the peduncle (fig. 10). The anterior portion of the body, while still
passing through the closing aperture of the tail, undergoes a half twist (fig. 12),
and appears to slip through into the central portion of the cyst. However, some
vital moment in the process of encystment must have been missed, for when the
process of encystment is completed it is the anterior end of the cercaria which
almost invariably lies
in the dorsal half, and
the posterior end in
the ventral half of the
eyst. During the last
stages of withdrawal
some fluid material
appears to besqueezed
out of the tail. This
may help to close the
eyst, and probably
also serves as a lubri-
cant, which assists
the cerearia in slipping
through the aperture
of the tail. The
various stages of
encystment were fol-
lowed in preserved
material only.
The Cyst (figs.
10-13, 27-29), when
fully formed, measures
39 wx 39 u-43 ux43u
in lateral aspect. The
aperture is completely
closed, but its position
is marked by a slight
thickening of the
cyst-wall. The various
appendages are still
attached to the out-
side, but in a much
reduced or altered
condition. All nuclei of Appendage II have degenerated completely. The stalk
of Appendage III has elongated and the free ends of the cells are frayed into a
great number of separate strands (fig. 27). The nuclei have degenerated. In
this condition the spread of the free ends of the appendage is about equal to the
width of the entire cyst. Appendages IV and V appear as mere strands of
tissue without a trace of nuclei. Appendage I, folded up inside the cyst, is
so transparent that it becomes almost invisible. All the cells between the lining
membrane and the cyst-wall have degenerated in the fully-formed cyst. The
position of the encysted cercaria is different from that of ©. sagittarvus. In the
54 Novirates ZooLtocicaE XLI. 1938.
latter species the body is folded twice, so that both extremities come to lie in
the top of the cyst. In C. sinitzini the body is twisted once only, and both
extremities lie in the bottom of the cyst. The anterior end of the cercaria lies
in the dorsal half of the cyst. The body is twisted above the cuticular
invagination of the cyst-wall, thus bringing the ventral sucker uppermost near
the top of the cyst. The posterior end of the cercaria lies in the ventral half of
the cyst. It should be remembered that the cyst is not spherical, but strongly
compressed laterally.
Excystment of the Cercaria (figs. 28, 29). If light pressure is brought to bear
on the cyst, the anterior end of the cercaria is pushed against the base of the dorsal
half of the cyst close to the point of origin of the delivery-tube. This releases
the delivery mechanism. Appendage I is everted—turned inside out—and thus
shot out through the bottom of the cyst at the point of origin of Appendage V.
The telescoped walls of the cells of which Appendage I is composed are thus
extended once more to their utmost limit (fig. 28). It then measures about 44
times the length of the cyst-itself. Immediately after eversion of Appendage I
the cercaria is drawn rapidly down the tube and is projected to the outside.
It was possible to fix and mount specimens in all stages of excystment.
Excysted Cercaria.mWhen released from the cyst the cercaria moves about
sluggishly. The body is still comparatively undifferentiated, but what appears to
be the appendiculate portion of the tail has begun to form. In stained specimens,
measuring 100 or over, two groups of cells in the posterior region stain more
deeply than the rest, and probably represent the rudiments of the reproductive
organs.
DIscussION.
Since the publication of Krull’s paper describing the life-history of Halipegus
occidualis Staff. 1905, the function of some of these extraordinary appendages of
Hemiurid cercariae has been demonstrated for the first time. The delivery-tube
shoots the encysted cercaria through the intestinal wall into the body cavity of a
Copepod, which serves as the second intermediate host. The other appendages
are believed to attract the Copepod, which releases the mechanism of the delivery-
tube by manipulating, touching or becoming entangled in these appendages, in its
endeavour to eat the cyst or its contents.
One of the most interesting aspects of the peculiar caudal appendages of the
cystophorous cercariae is their great variety. In this connection it is worth
recording that in the case of C. sinitzini there appears to be a general tendency for
the tail-vesicle to produce accessory appendages. As already noted, a number of
abnormal specimens were observed in which the terminal portion of Appendage
IV was bifurcated, or else carried a terminal finger placed at right angles to the
main prolongation. Other specimens showed a replica of Appendage V, set at
right angles to it—formed by the outward migration of a second giant nucleus
from the base of Appendage IV. All these appendages may possess some useful
or vital function, but it seems quite possible that one or several of them have no
function whatsoever. So long as any of the appendages came to serve a useful
purpose, the tendency to produce accessory caudal outgrowths might become
fixed in the whole group. As would be expected, the delivery-tube, the im-
portance of which is now quite clear, is the least variable of all the appendages.
It is fairly constant in structure except in the case of C. laqueator Sinitzin 1911,
Novirates ZOOLOGICAE XLI. 1938. 55
where it is drawn out into an immensely long filament and apparently never with-
drawn into the tail-vesicle.
The extraordinary degree of specialisation shown by this group of cercariae
is unique, and it is difficult to conceive how the delivery apparatus, with its
peculiar function in the life-history of the cercaria, can have arisen. It is worth
noting, however, that in the Azygiidae Odhner, a family of trematodes almost
certainly related to the Hemiuridae, the cercariae also encyst by withdrawing into
their own tails. Although this was probably the first stage in the evolution of the
present peculiar method of excystment of the cystophorous cercariae, it gives
us no hint how the extremely complicated and delicately adjusted delivery
mechanism first came into being.
REFERENCES.
Cort, W. W., and Nichols, E., 1920. A new cystophorous cercaria from California
—In J. Parasit. Urbana, 7, pp. 8-15, 2 figs.
Dollfus, R. P., 1923. Remarques sur le cycle évolutif des Hémiurides.—In
Ann. Parasitol., Paris, 1, pp. 345-351, 4 figs.
Dubois, G., 1929. Les Cercaires de la Région de Neuchatel—In Bull. Soc.
Neuchatel. Sci. Nat., 58 (N.S. 2) 1928 (1929), pp. 1-177, 17 pls., 8 figs.
Faust, E. C., 1921. Notes on South African larval Trematodes.—In J. of Parasit.,
Urbana, 8, pp. 8-21, 1 pl., 13 figs.
Krull, W. H., 1935. Studies on the life-history of Halipegus occidualis Stafford,
1905.—In Amer. Midl. Nat., Notre Dame, 16, pp. 129-143, pl. iv.
Pratt, H. S., 1898. A contribution to the life-history and anatomy of the
Appendiculate Distomes.—In Zool. Jahrb. Anat. & Ont., Jena, 11, pp. 1-40,
3 pls., 14 figs.
Rothschild, M., 1935. Note on the excretory system of Cercaria ephemera
Lebour 1907 (nec Nitzsch).—In Parasitology, Cambridge, 27, No. 2, pp.
171-174, 7 figs. :
Rothschild, M., 1936. Preliminary Note on the Trematode Parasites of Peringia
ulvae Pennant 1777.—In Nov. Zool., Tring, 39, pp. 268-269.
Sewell, R. B.S., 1922. Cercariae indicae.—In Ind. J. med. Res., 10 (suppl. no.),
pp. 1-370, i-ii, and 32 + 7 pls.
Sinitzin, D. T., 1905. Studies on the life-cycle of Trematodes. The distomes of
the fish and frogs in the environs of Warsaw (Russian).—In Mem. Soc. Nat.
Varsovie, biol., 15, pp. 1-210.
Sinitzin, D. T., 1911. Parthenogenetic Generation of Trematodes and its
Progeny in Molluscs of the Black Sea. (Translated by Alexis M. Bagusin,
under the direction of Professor Henry B. Ward.)—In St. Petersburg Mém.
Ac. Sc. (Ser. 8), Vol. 30, No. 5, pp. 1-127, 6 pls.
Willey, C. H., 1930. A cystophorous cercaria, C. projecta n. sp., from the snail
Helisoma antrosa, North America.—In Parasitology, Cambridge, 22, pp. 481-
489, 5 figs.
EXPLANATION OF FIGURES.
Fig. 1. Cercaria sinitzini. (At this stage the body stains uniformly and the
suckers are only indicated.) Tail in lateral aspect with Appendage I
expanded. The internal cellular structure of the tail-vesicle is omitted in
the figs. except figs. 2 and 5.
56 NOVITATES ZOOLOGICAE XLI. 1938.
Fig. 2. Sagittal section of the tail-vesicle of a fully-developed cercaria before
telescoping of Appendage I.
Fig. 3. Outline drawing of ©. sinitzini from ventral aspect after withdrawal of
Appendage I.
Fig. 4. Cercaria sinitzini. Body in ventral aspect and tail in lateral aspect after
telescoping of Appendage I, of which one cell is already withdrawn into the
tail-cavity. The giant nucleus of Appendage V is migrating back along the
stalk of the appendage. The other appendages also show signs of degenera-
tion. (Nuclei of primitive epithelium omitted.)
Fig. 5. Optical section of the tail-vesicle of a fully-developed cercaria
after the telescoping of Appendage I, showing flattened nuclei of sub-
cuticular layer and cell-group giving rise to Appendage III. The central
cavity is not visible in all optical sections.
Figs. 6-9. Outline drawings of cercariae in different stages of development.
Fig. 9 shows the lines along which measurements were taken for Table II.
Fig. 9a. Nuclei of primitive epithelium.
Figs. 10-12. Process of encystment. Appendages II and III have been omitted
in figs. 10-13.
Fig. 13. Encysted cercaria. Lateral aspect.
Fig. 14. Outline drawing of Appendage IV, showing maximum development of
terminal prolongation.
Figs. 15-18. Terminal portion of Appendage I, showing different stages of
development.
Fig. 18a. Diagram showing the manner of telescoping of the cells of
Appendage I.
Fig. 19. Appendage II at maximum stage of development. A portion of the
cellular structure is drawn in to show the spiral arrangement.
Fig. 20. Terminal portion of Appendage II during encystment of cercaria.
Nuclei of the spiral have degenerated. External nuclei are shrinking.
Fig. 21. Appendage II after encystment of cercaria.
Figs. 22-26. Appendage III at different stages. Fig. 26 shows the final condition
after encystment of the cercaria.
Fig. 27. Empty cyst after projection of cercaria to the outside through the
everted delivery-tube (Appendage I).
Fig. 28. Excysting cercaria about to pass down everted delivery-tube.
Fig. 29. Excysting cercaria caught in delivery-tube, which has failed to evert
normally beneath a cover-slip.
EXPLANATION OF LETTERING.
Al ‘ | : . Appendage I.
A le 1 : : . Appendage I everted.
A Il 5 : : . Appendage II.
AI . : : . Appendage III.
AV « : : . Appendage IV.
AVIV: : 3 . Appendage IV degenerating.
AV 3 : 4 . Appendage V.
AN NVGh j = . Appendage V degenerating.
a : 5 : . anterior end of cercaria.
ac : : : . anterior lateral collecting-tube.
Novirates ZooLoGIcAE XLI.
De
bc
bl.
Cc
Cum:
cw
dn
1938.
57
body of cercaria.
body of cercaria caught in delivery-tube.
bladder.
cavity.
cuticular invagination.
eyst-wall.
degenerating nucleus.
dorsal half of tail-vesicle.
encysted cercaria.
group of cells of unknown function.
giant nucleus.
intestine.
internal column of cells of Appendage IV.
lining membrane of cyst.
main collecting-tube.
nucleus.
giant nucleus of Appendage V.
nuclear membrane.
oral sucker.
pharynx.
posterior lateral collecting-tube.
peduncle.
primitive epithelium.
posterior end of cercaria.
sheath of Appendage I after eversion.
subcuticular protoplasmic layer.
tail.
tubular circuit of excretory vesicle.
terminal prolongation of Appendage IV.
ventral sucker.
ventral half of tail-vesicle.
58 NOVITATES ZOOLOGICAE XLI. 1938.
NEW OR LITTLE-KNOWN MARSUPIALS: A NEW SPECIES
OF PHASCOGALINAE, WITH NOTES UPON ACROBATES
PULCHELLUS ROTHSCHILD.
By G. H. H. TATE.
IS the Zoological Museum, Tring, in the summer of 1937, I came across two
specimens of an obviously new Dasyurid collected by A. S. Meek in 1905.
At my request Dr. Karl Jordan, Director, consulted the late Lord Rothschild as
to the desirability of permitting the new species to be described. Lord Roth-
schild kindly consented to lend the material to the American Museum of Natural
History, New York, for description, and in addition to send for study the type
skull of the unique Acrobates pulchellus Rothschild and two specimens of A. p.
frontalis (De Vis) 1887 from near the type locality. Studies were conducted also a
few months ago of the type of Acrobates pygmaeus at the British Museum (Natural
History). So I am now in a favourable position to discuss the relationships of
the three named forms of Acrobates.
My presence in Europe last summer was due to the generosity of Mr. Richard
Archbold, who has financed two expeditions to New Guinea, and who last year
set aside a sum of money to allow study of the types of Indo-Australian mammals.
1. Phascogale (Murexia) rothschildi n. sp.
Type.—Field No. 1, Zoological Museum, Tring; <g, adult; Aroa River,
Papua; May 28, 1905; collector A. S. Meek. The type is a skin and skull in
fair condition but lacking field measurements.
General Characters—A member of the subgenus Murexia Tate & Archb.
1937, but sharply differentiated from Ph. murex Thomas 1913, to the eye by
the presence of a broad dorsal stripe running from nose to tail.
Description of type.-—General colour greyish brown, becoming slightly tinged
with cinnamon on thighs ; the individual hairs with grey bases, brown subterminal
bands and fuscous tips ; occasional guard hairs shining black ; dorsal band fus-
cous, running from frons to base of tail, reaching an average width of 13 mm. ;
underparts Light Cinnamon Buff,! with grey bases ; ears and tail light fuscous ;
hands and feet buff. Pelage relatively short (4 mm.) and crisp.
Skull with the characters of Ph. murex but small throughout ; palate and
muzzle relatively short and broad; incisors and premolars with similar size
gradients. Teeth considerably smaller than those of P. murex.
Measurements.—Head and body,? 170 mm. ; tail, 180; hind foot (s.u.), 33 ;
ear from crown, 13; skull, greatest length, 43-2; basal length, 40-1; zygomatic
breadth, 24-6 ; interorbital breadth, 8-0; breadth of braincase, 16-0; mastoid
breadth, 16-0; nasals, 14-9 x 5-3; palatal length, 23-0; anterior palatal
foramina, 3:7; breadth of mesopterygoid fossa, 5-1; length of bulla, 2-9;
crowns p‘-m*, 10:8; m!?, 7:6; mi, 2-6 x 2-0.
Paratype.—No. 108106, Amer. Mus. Nat. Hist.; &, adult; collector and
locality as for the holotype.
1 Colours are those of Ridgway, Colour Standards and Nomenclature, 1912, only if capitalized.
2 From the dry skin.
Novirares ZooLoGIcAE XLI. 1938.
59
I take pleasure in naming this interesting new species in memory of the late
Lord Rothschild, who devoted so much of his life to furthering zoological know-
ledge.
2. On Acrobates pulchellus Rothschild 1892.
Forty-five years ago Rothschild described! Acrobates pulchellus from “ one
of the small islands in Northern Dutch New Guinea.”
The unique specimen had
been received from A. Bruijn, who acted as a dealer and obtained his material
(chiefly from natives) from all about the western end of New Guinea.
of origin of the animal is therefore in doubt.
Rothschild compared the external characters of A. pulchellus with those of
He did not mention A. frontalis from the-
Herbert River, Queensland, which De Vis founded upon a very young animal and
-A. pygmaeus of New South Wales.
placed originally in the genus Dromicia.
The place
The skulls of A. pulchellus and of one
adult specimen of A. frontalis have now been cleaned and their characters and
measurements checked with those of the type of A. pygmaeus.
The three named
forms are compared in the following tabular arrangement :
SKINS.
A. pygmaeus, paratype ”
A. frontalis, topotype
A. pulchellus, type
New South Wales .
Male, adult . 5 5 5 5 ö
Colour faded by alcohol ; scarcely a trace
of grey bases to the white ventral pelage ®
Tail longer than head and body
North Queensland
Male, adult
Dorsal colour Prout’s
Brown, with fuscous
bases. Ventral colour
creamy white to roots.
Tail as pygmaeus
Dutch New Guinea (?)
Sex ? adult.
Dorsal colour brown with
fuscous bases. Ventral
colour dull white, with
dark grey bases.
Tailshortened. By injury?
The white posterior part of the ear is a character common to all three types, as is the white
upper lip.
2 The female paratype corresponds externally to the male.
extracted.
3 A dried skin (A.M. 37185) from N. 8. Wales has grey bases.
Its skull has not been
SKULLS.
A. pygmacus A. frontalis A. pulchellus
6 paratype 3 topotype type
Breadth braincase
on gl 1-01) 1:01 -90
Mastoid breadth en)
Breadth mesopter. fossa 19 (+17) .20 .18
Zygomatie breadth
Palatal length
——— IE ERRE ENG 75 (79 77 79
Zygomatic breadth 2)
en. :24 (24) 27 25
palatal length
width inside m!-m! .33 (-30) 35 .34
palatal length
ken -40 (-40) 39 36
mi-3
4 Proportions in parenthesis are those of a d topotype, A.M. 37185.
Obviously we lack adequate series of the forms under consideration. Also
ignorance of the sex of A. pulchellus weakens our comparison, for the very slight
variations distinguishable in pulchellus may be entirely attributable to sexual
1 Rothschild 1892, Proc. Zool. Soc. London,"p. 546.
6
60 Novirates ZOOLOGICAE XLI. 1938.
differences. Aside from these points pulchellus agrees very closely with pygmaeus
in the colour of the ventral pelage, in size, and in most of the cranial proportions
tested. A. frontalis, on the other hand, is marked off from both pygmaeus and
pulchellus by its ventral fur being white to the roots, and by the greater size of
its teeth (see measurements above). Bearing in mind the doubtful origin of
pulchellus and the presence of a distinct geographical form between its supposed
territory and that of pygmaeus, 2,000 miles distant, the conclusion is scarcely
avoidable that by error a specimen of pygmaeus was mislabelled New Guinea. In
that case, when a sufficient series of the latter is available for study, pulchellus will
fit within the range of individual variation of that series. If such is not the case,
pulchellus can only be explained as one of the rarest of mammals living in parts of
New Guinea whose zoology, broadly speaking, has been well explored.
In conclusion, I am inclined to the view that Acrobates Desmarest is an
Australian genus comprising two species (or subspecies ?)—A. pygmaeus with grey-
based ventral pelage and smaller teeth (mt? = 2-5) in the south, and A. frontalis
with self-coloured white ventral fur and larger teeth (m! = 2-8) in the north.
The animal A. pulchellus (= pygmaeus) was perhaps carried as a pet on board
ship and obtained subsequently without proper locality by Bruijn.
Novirares ZOOLOGICAE XLI. 1938. 61
ECTOPARASITES FROM CAPTIVE BIRDS.
I. 1936-1937.
By THERESA CLAY, B.Se., AND MIRIAM ROTHSCHILD.
LARGE number of birds, obtained from all parts of the world, die in
aviaries in this country every year. It occurred to the authors that a
record of ectoparasites from some of these captive birds might prove of con-
siderable interest. Quite apart from the number of new species such a collection
is likely to provide, the conditions found in aviaries particularly favour the study
of straggling of parasites from one host to another, and the acquisition of new
parasites by foreign birds bred in this country.
In the following list the Mallophaga ! have been identified by Miss Theresa
Clay, the Siphonaptera by the Hon. Miriam Rothschild, and the Acarina by Mrs.
A.M. Hughes. Thesnitials following each record, whether positive or negative,
indicate the collector who examined the bird in question. The figure in brackets
after the name of the host indicates the number of that species examined.
Various methods were employed for collecting the parasites, but owing to
the poor state of preservation of many of the birds it was not always possible
to guarantee the negative records. Mites were not collected by the junior
author.
GENERAL REMARKS.
A total of 128 birds was examined, including 91 species pertaining to 34
different families. Of this number, 90 specimens (65 species) were confined in
captivity with various species of birds—thus providing ample opportunity for
the straggling of parasites from one to another. The type of confinement varied
from comparative freedom on a large lake to close confinement in an aviary
or cage.
It has already been stated that mites were not collected from the birds
examined at the London Zoo by the junior author, and it is therefore more
convenient to study them apart from the lice. Only one bird was found to be
harbouring fleas.
Mallophaga.—48 different species of Mallophaga were found distributed
among 42 species of birds. The number of specimens infected was 48, or a little
under half the total of the birds examined. On the whole the infections were not
heavy, a comparatively small number of parasites being found on each bird.
Notable exceptions to this were the Jay and Buzzard, which swarmed with
parasites. The Hoopoe, Rosy-billed Pochard, and Stanley Crane were also fairly
heavily infected. 10 species of birds showed infections with more than one
species of Mallophaga.
11 of the birds were bred in captivity. Of these the Hoopoe, Rosy-billed
Pochard, Emu, and Jackass Penguine were reared by their own parents. All
proved to be infected, and the parasites were obviously acquired from the
parents, except possibly in the case of the Jackass Penguin, which can have
1 Before attempting to describe any of these species a revision of several genera is necessary,
and for this purpose further material is required.
62 Novirates ZooLoGIcAE XLI. 1938.
acquired the parasite either via its parents or directly from the other species of
penguins (true hosts) with which it was confined. 6 birds were reared by foster
parents (hens). 4 of these were ducks, kept on a large lake (pinioned) fully
stocked with various water-fowl, including Geese, Oyster-catchers, Plovers,
Curlew, etc. 2 acquired parasites, and it is worth noting that in both cases they
proved to be typical duck parasites. On the other hand, one of the Madagascar
Partridges, also reared by foster parents, acquired a parasite pertaining to a
group which normally infests the Gressores.
The most highly infected group of birds proved to be the Cardinals. Out
of 8 specimens (three species) 7 were infected. These included 5 species of lice.
The least infected were the Waxbills. Out of 20 birds (four species) none were
harbouring Mallophaga. The families Phasianidae and Fringillidae both showed
a high percentage of infection ; on the other hand, the Psittacidae, birds which
are frequently infected in the wild, all proved negative.
The egg-laying habits of lice appear to vary specifically, both with regard
to the site selected and the manner of attachment of the egg capsule. This is
well illustrated in the case of Degeeriella fusca (p. 67) and Degeeriella upupae
(p. 70). With regard to the former species, it would be of interest to ascertain
whether the lice migrate directly after hatching from the egg. The Buzzard
was examined shortly after death, and no very extensive post-mortem movement
of the lice had probably occurred. That such a migration does take place was
suggested by the fact that the young stages were distributed all over the body
and not confined to the lower wing coverts upon which the eggs are laid. A
return migration to this area must presumably be undertaken by the gravid
female. It is hoped to pay more attention to the habits of the lice, and include
figures of different eggs, when making future collections and lists.
Four cases only of straggling were recorded. As already mentioned, the
Jackass Penguin had acquired a species of Austrogoniodes generally found on the
Macaroni and Rock-hopper Penguins. The Kirghiz Pheasant was infected with
two species of lice, Lipewrus caponis and Goniodes meleagridis. The former is
usually found on the domestic fowl and certain game birds, while the true
host of the latter is the Turkey. It is possible that other cases of straggling
might become apparent if more information were available. Thus, for example,
the Brüelia sp. obtained from the Dominican Cardinal could conceivably have
the Red-crested Cardinal (with which it was confined) as the true host in nature.
However, with the exception of the Madagascar Partridge infected with a single
specimen of Ardeicola sp., we have no record of a type of parasite characteristie
of one definite group of birds straggling on to a bird pertaining to another unre-
lated group. No evidence was obtained to disturb the belief that the Mallophaga
are one of the best groups of parasites for assisting the study of host relationship.
Even under such exceptionally favourable conditions the evidence of straggling
is small.
It is worth drawing attention to the specimens of Ricinus sp. obtained from
the Ruby and Topaz Humming Bird (confined with related species only). The
presence of this genus of lice on a Humming Bird, which is otherwise characteristic
of Passerine birds, is of interest in view of the controversy regarding the syste-
matic position of the Trochlidae.
Acarina.—A total of 91 specimens (58 species) pertaining to 26 different
families of birds was examined for mites (in addition to lice and fleas), of which
Novirarrs ZooLoGIcAE XLI. 1938. 63
15 specimens (13 species) were found to be infected. The number of species of
mites recorded was 14. This figure is relatively low, and it appears probable
that some infestations were overlooked, particularly in the case of birds which
were received in a poor state of preservation. As with the lice, the actual
number of parasites on each bird was generally small. In the case of the genera
Cheyletiella, Laelaps, and Liponyssus, this is explained by the fact that they are
essentially parasites inhabiting the nests of their hosts, rather than the birds or
mammals themselves. However, an exceptionally heavy infection was recorded
for the Green Magpie, literally thousands of mites being present. This bird
also provided the only record of two species of mites on the same host. 8 of
the infections were found in association with lice.
9 of the birds were bred in captivity. 2 showed infection with mites.
One of these, the Falcated Teal, was reared by foster parents and must have
acquired the mites, as it did the Mallophaga with which it was infected. Un-
fortunately these specimens, together with a small series off the Scaly Ground
Dove, were mislaid and are therefore omitted from the following list. No definite
case of straggling was recorded. The genus Laelaps is, however, found more
frequently on small mammals than on birds.
Siphonaptera.—The disappointing fact that only one record of fleas was
made is probably due to the two following reasons. Firstly, fleas leave the host
when it becomes cold, and in most cases the birds examined were collected several
hours after death. Secondly, fleas, like many genera of mites, breed in the nests
of the host, and are more likely to be collected there than on the bodies of the
birds themselves.
LIST OF PARASITES.
I. SIPHONAPTERA (Fleas).
Ceratophyllidae.
Parasite. Host.
Ceratophyllus gallinae Schranck 1803 . Dendronessa galericulata (Linn.) 1758.
II. MALLOPHAGA (Biting Lice).
Menoponidae.
Anas falcata (Georgi) 1775.
*Trinoton querquedulae 1 (Linn.) 1758 . | Anas forınasn (Cases) NIE
Menacanthus sp.? - : 6 . Turacus leucotis (Rüppell) 1835.
Menacanthus sp. - : : . Cyanops asiatica (Latham) 1790.
Myrsidea sp.? . . Paroaria dominicana (Linn.) 1758.
Chapinia africana ( (Bedford) 1929 . Bucorvus cafer (Schleg.) 1862.
Colpocephalum decimfasciatum (Boisd.
& Lacord.) 1835 . ; ‘ . Ardea c. cinerea Linn. 1758.
Colpocephalum sp.? . : : . Pavo cristatus Linn. 1758.
1 The species indicated by a star (*) show slight variation which may ultimately prove to be
subspecific.
64 NOVITATES ZOOLOGICAE XLI. 1938.
Ricinidae.
Ricinus sp. n. . : : 3 . Chrysolampis elatus (Linn.) 1758.
Philopteridae.
Goniodes meleagridis (Linn.) 1758 . Phasianuscolchicus mongolicus (Brandt)
1844.
Goniodes cervinicornis Giebel 1874 . Gennaeus leucomelanos hamiltoni (Gray)
1829.
Goniodes pavonis (Linn.) 1758 . . Pavo cristatus Linn. 1758.
Goniodes sp. n. . Diardigallus diardi (Bonaparte) 1856.
Goniocotes Blogs Giebel 1866 . Tragopan satyra (Linn.) 1758.
Physconelloides sp. ? : ; . Phaps c. chalcoptera (Latham) 1790.
Physconelloides sp. ? : ; . Geophaps smithi (Jard. & Selby) 1830.
[ Eudyptes chrysolophus (Brandt) 1837.
Austrogoniodes strutheus Harrison 1915 Eudyptes c. crestatus (Miller) 1784.
Spheniscus demersus (Linn.) 1758.
Lipeurus caponis (Linn.) 1758 . . Phasianus colchicus mongolicus (Brandt)
1844.
Lipeurus subsellatus Harrison 1916 . Gennaeus leucomelanos hamiltoni (Gray)
1829.
Oxylipeurus himalayensis (Rudow)
1869 . : 5 : : . Tragopan satyra (Linn.) 1758.
Oxylipeurus sp.n.- . . Oreortyx p. picta (Douglas) 1823.
. Otilipeurus turmalis a) 1842 . Otis t. tarda (Linn.) 1758.
Dahlemhornia asymmetrica (Giebel) Dromiceius n. novae-hollandiae (Lath-
1866 am) 1790.
Degeeriella rufa (Burm.) 1838 . . Falco tinnunculus (Linn.) 1758.
" Degeeriella fusca (Denny) 1842. . Buteo buteo (Linn.) 1758.
Degeeriella hospes (Giebel) 1866 . Squatarola squatarola (Linn.) 1758.
Degeeriella upupae (Denny) 1842 . Upupa e. epops (Linn.) 1758.
Brüelia glandarii (Denny) 1842 . Garrulus glandarius (Linn.) 1758.
[ Paroaria cristata (Boddaert) 1783.
Paroaria dominicana (Linn.) 1758.
Briielia limbata (Burm.) 1838 . . Loxia c. curvirostra Linn. 1758.
Brüelia sp. . 5 : : . Richmondena cardinalis (Linn.) 1758.
Brüelia sp. Richmondena cardinalis (Linn.) 1758.
Briielia sp. Hypochaera chalybeata (Müller) 1776.
Brüelia sp. Niltava grandis (Blyth) 1842.
Briielia sp. Cissa chinensis (Boddaert) 1783.
Paroaria dominicana (Linn.) 1758.
Loxia c. curvirostra Linn. 1758.
Brüelia trithorax (Burm.) 1838 .
DO OD ew
*Philopterus citrinelle (Schrank) 1776 .
Philopterus cursitans (Giebel) 1861 . Athene n. noctua (Scop.) 1769.
Philopterus sp. ? . : ö . Erythrura psittacea (Gmel.) 1788.
Philopterus sp. ? : : ; . Niltava grandis (Blyth) 1842.
Anatoecus sp. ? : : 3 . Metapopiana peposaca (Vieillot) 1818.
Anatoecus sp. ? : . Mareca sibilatrix (Poeppig) 1829.
Rallicola lugens (Giebel) 1874 : . Porphyrio poliocephalus (Latham) 1790.
Ornithobius fuscus Le Soeuf 1902 . Chenopis atrata (Latham) 1790.
Novirates ZooLOGICAE XLI. 1938. 65
*Oolumbicola columbae (Linn.) 1758 . Geophaps smithi (Jard. & Selby) 1830.
Columbicola sp.? . ‘ : . Scardafella sguammata (Lesson) 1831.
Columbicola sp.? . 5 : . Phaps c. chalcoptera (Latham) 1790.
Columbicola sp.? . : ‘ . Chalcophaps i. indica (Linn.) 1758.
*Hsthiopterella gruis (Linn.) 1758 . Tetrapteryx paradisea (Licht.) 1793.
Ardeicola sp. ? ; : ; . Margaroperdix madagarensis (Scop.)
1786.
III. ACARINA (Mites).
Analgesidae.
Megninia sp. ? 3 : ; . Cissa chinensis (Boddaert) 1783.
Pierolichus sp. ? Cissa chinensis (Boddaert) 1783.
Pterolichus sp. ? Recurvirostra avocetta Linn. 1758.
Plerolichus sp. ? Niltava grandis (Blyth) 1842.
Pterolichus sp. ? Melopsittacus undulatus (Shaw) 1805.
Laelaptidae.
Laelaps sp.? . ; ; ; . Acryllium vulturinum (Hardwicke)
1834.
Liponyssidae.
Liponyssus sp. ? : ¢ ¢ . Tiaris canora (Gmel.) 1788.
Liponyssus sp. ? : ‘ : . Cyanops asiatica (Latham) 1790.
Liponyssus sp. ? ; ; : . Richmondena cardinalis (Linn.) 1758.
Cheyletidae.
~~
Cheyletiella sp.
Cheyletiella sp.
Cheyletiella sp.
Estrilda melpoda (Vieillot) 1817.
Paroaria dominicana (Linn.) 1758.
Paroaria cristata (Boddaert) 1783.
m +
LIST OF HOSTS.
Dromiceiidae.
DROMICEIUS N. NOVAE-HOLLANDIAE (Latham) 1790, Emu (1). London Zoo,1
November 1937. Bred at Whipsnade. Confined with same species only.
1 &. 5 22 Dahlemhornia asymmetria. (T. C.)?
Spheniscidae.
SPHENISCUS DEMERSUS (Linn.) 1758, Jackass Penguin (1). London Zoo,
1937. Bred in captivity. Confined with various other species of penguins.
19 33, 23 99 Austrogoniodes strutheus. This infection was probably acquired
from another species of penguin as it is not the usual species of Austrogoniodes
found on this bird. (T.C.)
EUDYPTES CHRYSOLOPHUS (Brandt) 1837, Macaroni Penguin (1). London
Zoo, April 1937. Obtained from the Antarctic. Confined with various species
of penguins. 5 99 Austrogoniodes strutheus. (T. C.)
1 “ London Zoo ” refers to the Zoological Society of London, Regent’s Park.
2 T. C. = Theresa Clay, M. R. = Miriam Rothschild, E. M. = Elizabeth Meyerhof.
66 Novirates ZooLOGICAE XLI. 1938.
EUDYPTES C. CRESTATUS (Miller) 1784, Rock-hopper Penguin (1). London
Zoo, April 1937. No data. Confined with various species of penguins. 12 34,
8 99 Austrogoniodes strutheus. (T. C.)
Podicipidae.
PopICEPS c. CRISTATUS (Linn.) 1758, Great Crested Grebe (1). London
Zoo, 1937. Died on arrival. No data. No Mallophaga or Siphonaptera.
(T. C.)
Ardeidae.
ARDEA C. CINEREA Linn. 1758, Common Heron (2). London Zoo, January
1937. @ obtained in Great Britain. Confined with various species of birds ;
3 33,7 29 Colpocephalum decimfasciatum. The second specimen, obtained on the
Continent of Europe, confined with various species of birds ; no Mallophaga or
Siphonaptera. (T. C.)
BUBULCUS IBIS COROMANDUS (Boddaert) 1783, Buff-backed Heron (1).
London Zoo, 1937. Obtained from India, confined with various species of birds,
No Mallophaga or Siphonaptera. (T. C.)
Phoenicopteridae.
PHOENICOPTERUS ANTIQUORUM Temminck 1820, Flamingo (1). Lilford
Aviaries, 22.3.37. Obtained from Esypt. Free on lake for several years with
various species of water-fowl. No ectoparasites. (E. M.)
Anatidae.
CHENOPIS ATRATA (Latham) 1790, Australian Black Swan (1). London
Zoo, November 1936. Q obtained from Australia, confined with various species
of birds. 12 Ornithobius fuscus. (T. C.)
ANAS FALCATA Georgi 1775, Falcated Teal (1). Lilford Aviaries, 15.9.37.
Bred in England, reared by foster parent (hen). Free on lake with various
water-fowl. 1 3, 1 Q *Trinoton querquedulae in breast feathers. No eggs
noted. A small series of unidentified mites on the superficial head and neck
feathers. (M. R.) 5
ANAS CASTANEA (Eyton) 1838, Chestnut-breasted Teal (1). Lilford Aviaries,
16.3.37. Bred in England. Conditions as above. No ectoparasites. (M. R.)
ANAS CYANOPTERA Vieillot 1816, Cinnamon Teal (1). Lilford Aviaries,
12.3.37. Bred in England. Conditions as above. No ectoparasites. (M. R.)
ANAS FoRMOSA Georgi 1785, Clucking Teal (1). Lilford Aviaries, 9.2.37.
2 obtained from Japan or Siberia. Free on lake for about three years with
various water-fowl. 1 3 *Trinoton querquedulae in breast feathers. No eggs
noted. (M. R.)
Anas DISCORS Linn. 1766, Blue-winged Teal (1). London Zoo, 1937.
Obtained from North America, confined with various species of ducks. No
Mallophaga or Siphonaptera. (T. C.)
Arx sponsa (Linn.) 1758, Carolina Duck (1). Dudley Zoo, 17.9.37. Obtained
from North America. Confined with various species during captivity. No
ectoparasites. (M. R.)
Novirares ZooLoGIcAE XLI. 1938. 67
DENDRONESSA GALERICULATA (Linn.) 1758, Mandarin Duck (1). London
Zoo, March 1937. Confined with various species of birds. A series of dd and
9° of Ceratophyllus gallinae from breast feathers. (T. C.)
MARECA SIBILATRIX (Poeppig) 1829, Chiloe Wigeon (1). Lilford Aviaries,
27.38.37. Bred in England, reared by foster parent. Free on lake with various
water-fowl. 1 2 and 1 immature Anatoecus sp.? on head. No eggs noted.
(M. R.)
METAPOPIANA PEPOSACA (Vieillot) 1818, Rosy-billed Pochard (1). Lilford
Aviaries, 21.12.36. Bred in England, reared by own parents. Conditions as
above. A fairly heavy generalized infection of Anatoecus sp.? (M.R.)
Accipitridae.
ACCIPITER NIsus (Linn.) 1758, Sparrow Hawk (1). Dudley Zoo, 7.9.37.
Captured near Birmingham and subsequently confined alone. No ectoparasites.
(M. R.)
Burro BuTEO (Linn.) 1758, Common Buzzard (1). Lilford Aviaries,
August 1937. Captured in Devonshire, England. Confined with same species
only. Exceedingly heavy general infection with * Degeeriella fusca. Eggs con-
fined to the surface of the basal half of lower wing coverts close to the body.
As many as 160 eggs on one feather. (M. R.)
FALCO TINNUNCULUS (Linn.) 1758, Kestrel (2). One specimen for which
we have no data, infected with Degeeriella rufa. Second specimen Dudley Zoo,
22.3.37. Captured near Dudley. Confined alone. No ectoparasites. (M. R.)
Phasianidae.
OREORTYX P. PICTA (Douglas) 1823, Plumed Quail (1). London Zoo, 1937.
No data. Confined with various species of birds. Infected with Oxylipeurus
sp.n. (T.C.)
TRAGOPAN SATYRA (Linn.) 1758, Satyr Tragopan (1). London Zoo, 1937.
Obtained from India, confined with various species of birds. Infected with
Goniocotes diplogonus and Oxylipeurus himalayensis. (T. C.)
GENNAEUS LEUCOMELANOS HAMILTONI (Gray) 1829, White-crested Kaleege
(1). London Zoo, June 1937. Obtained from Western Himalayas, India.
Confined with various species of birds. 2 34, 2 22 Lipeurus subsellatus, and
8 dd, 6 29 Goniodes cervinicornis. (T. C.)
DIARDIGALLUS DIARDI (Bonaparte) 1856, Siamese Crested Fireback (1).
London Zoo, February 1937. Obtained from Siam, confined with various species
of birds. 6 dd, 10 99 Goniodes sp.n. (T.C.)
PHASIANUS COLCHICUS MONGOLICUS (Brandt) 1844, Kirghiz Pheasant (1).
London Zoo, 1937. Obtained from Mongolia, confined with various species of
birds. Infected with Lipeurus caponis and Goniodes meleagridis. The true host
of the latter species is the Turkey. (T. C.)
Pavo cristatus Linn. 1758, Peafowl (2). London Zoo, December 1936
and June 1937. 2 99 obtained from India, confined with various species of birds.
First specimen with 2 gg, 8 29, 4immature Goniodes pavonis. Second specimen
with 4 gg, 4 22 Goniodes pavonis and 1 g, 2 99 Colpocephalum sp.? (T. C.)
ACRYLLIUM VULTURINUM (Hardwicke) 1834, Vulturine Guinea-fowl (1).
Lilford Aviaries, December 1936. Obtained from Somaliland. At liberty for
68 Novirates ZooLoGIcAE XLI. 1938.
about six years in an enclosure of about three acres, together with various species
of birds. Two specimens of Laelaps sp.? (M. R.)
MARGAROPERDIX MADAGARENSIS (Scop.) 1786, Madagascar Partridge (2).
Foxwarren Park, 16.12.36 and 4.2.37. Both specimens bred in England,
reared by foster parents, and confined with various other birds. One infected
with a single specimen of Ardeicola sp. ?, a straggler from one of the Gressores.
The second bird without ectoparasites. (M. R.)
Gruidae.
TETRAPTERYX PARADISEA (Licht.) 1793, Stanley Crane (1). London Zoo,
December 1936. 4 obtained from Cape Colony, confined with another crane.
109 SS, 247 29, 246 immature *Hsthioptereila gruis. (T. C.)
Rallidae.
PoRPHYRIO POLIOCEPHALUS (Latham) 1790, Grey-headed Gallinule (1).
Lilford Aviaries, 5.2.37. Obtained from India. Confined with jungle fowl and
wild guinea fowl for about six years. A fairly generalised infection with
Rallicola lugens. ‘This species seems more firmly attached to the feathers than
is generally the case. (M. R.)
Otididae.
Ov1s T. TARDA Linn. 1758, Great Bustard (1). London Zoo, February
1937. © obtained from Hungary, confined with various species of birds. 5 34,
10 99 Otilipeurus turmalis. (T. C.)
Charadriidae.
SQUATAROLA SQUATAROLA (Linn.) 1758, Grey Plover (2). Ashton Wold,
August 1937. A pair from Great Britain. Confined together in a large garden.
One showed a light general infection of Degeeriella hospes. (M. BR.)
Recurvirostridae.
RECURVIROSTRA AVOCETTA Linn. 1758, Avocet (1). Foxwarren Park,
22.1.37. Obtained on the Continent of Europe. Heavily infected with Ptero-
lichus sp. ? The mites were situated between the barbules of the retrices, parallel
with them, close to the shaft of the feather. The white feathers showed dis-
coloration. (M. R.)
Chionidae.
CHIONIS ALBA (Gmel.) 1788, Sheath-bill (1). London Zoo, 1937. Obtained
from the South Shetlands, confined with numerous other species of birds. No
Mallophaga or Siphonaptera. (T. C.)
Laridae.
LARUS DOMINICANUS Licht. 1823, Southern Black-backed Gull (1). London
Zoo. No data. Confined with various species of birds. No Mallophaga or
Siphonaptera. (T. C.)
LARUS N. NOVAE-HOLLANDIAE Stevens 1826, Silver Gull (1). London Zoo,
1937. Bred in captivity, confined with various species of birds. No Mallophaga
or Siphonaptera. (T. C.)
Novirarrs ZOOoLOGICAE XLI. 1938 69
Columbidae.
GEOPHAPS SMITHI (Jard. & Selby) 1830, Smith’s Partridge Pigeon (1).
Foxwarren Park, 26.3.37. Obtained in Australia. Confined with various
species of birds. A fairly general infection with Physconelloides sp.? and 2
specimens of *Columbicola columbae. (M. R.)
GALLICOLUMBA JOBIENSIS (A. B. Meyer) 1875, White-breasted Ground
Dove (1). Foxwarren Park, 30.4.37. Obtained from Jobi Island. Confined
with various species of birds. No ectoparasites. (M. R.)
ZENAIDA ASIATICA (Linn.) 1758, White-winged Dove (1). Lilford Aviaries,
7.2.37. Obtained from Mexico. Confined for two years at least with various
other birds. No ectoparasites. (M. R.)
SCARDAFELLA SQUAMMATA (Lesson) 1831, Scaly Ground Dove (1). Dudley
Zoo, 2.9.37. Possibly bred in captivity. Confined with various species of
birds. One specimen of Columbicola sp.?, and a small series of mites (unidentified).
(M. R.)
CHALCOPHAPS I. INDICA (Linn.) 1758, Emerald Dove (1). London Zoo. No
data. Confined with various species of birds. Infected with Columbicola sp. ?
(T. C.)
PHAPS C. CHALCOPTERA (Latham) 1790, Common Bronze-wing (1). London
Zoo, November 1936. Obtained from Australia, confined with various species of
birds. 9 $4, 38 29 Physconelloides sp. ? and 1 g,3 99 Columbicola sp.? (T.C.)
Cuculidae.
CucuLus ©. CANORUS Linn. 1758, Common Cuckoo (2). Dudley Zoo,
8.8.37 and 23.8.37. Captured near Dudley, England. Confined separately.
No ectoparasites. (M. R.)
GUIRA GUIRA (Gmel.) 1788, Guira Cuckoo (1). London Zoo, 1937. Ob-
tained from Paraguay, confined with various species of birds. No Mallophaga
or Siphonaptera. (T. ©.)
Psittacidae.
MELOPSITTACUS UNDULATUS (Shaw) 1805, Budgerigar (3). Dudley Zoo,
13.8.37 and 6.9.37. London Zoo, 1937. The two specimens from Dudley
were bred in captivity and confined with various species of birds. One showed
a light infection of Pterolichus sp.? (EK. M.) The specimen from London Zoo
was obtained from Australia. No Mallophaga or Siphonaptera. (T. C.)
AGAPORNIS ROSEICOLLIS (Vieillot) 1817, Rosy-faced Love Bird (1). Dudley
Zoo, 6.8.37. Obtained from Africa. Confined separately since it reached this
country. No ectoparasites. (E. M.)
PLATYCERCUS ADscırus (Latham) 1790, Pale-headed Rosella (1). Fox-
warren Park, 7.1.37. Obtained from Queensland or New South Wales. Con-
fined alone. No ectoparasites. (EK. M.)
TRICHOGLOSSUS HOEMATODUS RUBRITORQUIS Vigors & Horsf. 1827, Red-
collared Lorikeet (1). London Zoo, 1937. Obtained from North-west Australia,
confined with the same species only. No Mallophaga or Siphonaptera. (T. C.)
LEPTOLOPHUS HOLLANDICUS (Kerr) 1792, Quarrion (1). London Zoo, 1937.
Obtained from Australia, confined with the same species only. No Mallophaga
or Siphonaptera. (T. C.)
70 NOVITATES ZOOLOGICAE XLI. 1938.
PSITTACUS E. ERITHACUS Linn. 1758, Grey Parrot (1). London Zoo, 1937.
Obtained from the Gold Coast, Africa, confined with another bird of the same
species. No Mallophaga or Siphonaptera. (T. C.)
Musophagidae.
TURACUS LEUCOTIS (Rüppell) 1835, White-cheeked Plantain Eater (1).
Lilford Aviaries, 1937. Obtained from Africa. Kept in large aviary with
various species of birds, for about five years. A light infection of Menacanthus
sp.? (M. R.)
Bucerotidae.
TROPICRANUS ALBOCRISTATUS (Cassin) 1848, White-crested Hornbill (2).
Foxwarren Park, 1.3.37. Obtained from Africa. Confined alone. No ecto-
parasites. (E. M.)
BUCORVUS CAFER (Schleg.) 1862, Ground Hornbill (1). London Zoo,
November 1937. Obtained from Cameroons, confined with various species of
other birds. 1 9 Chapinia africana. (T. C.)
Strigidae.
BuBo CAPENSIS (Smith) 1834, Cape Eagle Owl (1). Dudley Zoo, 2.8.37.
Wings only examined. Obtained from South Africa. Confined with other owls.
No ectoparasites. (E. M.)
ATHENE N. NOCTUA (Scop.) 1769, Little Owl (1). London Zoo, 1937.
© obtained from Germany, confined with other owls. 9 3, 26 22 Philopterus
cursitans. (T.C.)
Upupidae.
Urupa Epops Linn. 1758, Hoopoe (1). Foxwarren Park, 14.12.36. Bred
in captivity and reared by own parents. Very heavily infected with Degeeriella
upupae. Eggs laid in numbers on breast feathers, parallel with barbules, at
right angles to shaft. (M. R.)
Capitonidae.
CYANOPS ASIATICA (Latham) 1790, Blue-throated Barbet (4). Dudley Zoo,
7.8.37, 22.7.37, 11.9.37, and 18.9.37. Obtained from the Himalayas. Not
confined with other species of birds. Two specimens with a light general infection
of Menacanthus sp. ? Eggs laid in groups at the base of breast and neck feathers.
A light infection on one bird of Liponyssus sp.? (M. BR.)
Pittidae.
Pirra CYANEA Blyth 1843, Blue Pitta (1). Foxwarren Park, 16.12.36.
Obtained from Java. Confined alone. No ectoparasites. (E. M.)
Muscicapidae.
NILTAVA GRANDIS (Blyth) 1842, Greater Niltava (1). Foxwarren Park,
22.1.37. Obtained from East Himalayas. Confined alone. A single specimen
of Brüelia sp. ?, Philopterus sp. ? and Pterolichus sp.” (M. R.)
Turdidae.
KITTACINCLA MALABARICA INDICA (Baker) 1924, The Shama (1). London
Zoo, 1937. Obtained from India. Confined alone. No Mallophaga or
Siphonaptera. (T. C.)
Noviratres ZOOLOGICAE XLI. 1938. 71
Timeliidae.
HETEROPHASIA PICAOIDES Hodgson 1839, Long-tailed Sibia (2). Foxwarren
Park, 22.12.36 and 10.3.37. Birds confined together and not with other
species. No ectoparasites. (M. R. and E. M.)
Corvidae.
CIssA CHINENSIS Boddaert 1783, Green Magpie (1). Foxwarren Park,
12.12.36. Obtained from India. A few specimens of Brüelia sp.? No eggs
observed. An exceptionally heavy infection of Megninia sp. ? and Pterolichus
sp. ?> (M. RB.)
GARRULUS LIDTHI Bonaparte 1850, Lidthi’s Jay (1). Foxwarren Park,
17.1.37. Obtained in Northern Group of Rynkyn Islands. Confined with
various species of birds. No ectoparasites. (E. M.)
GARRULUS GLANDARIUS (Linn.) 1758, Common Jay (1). Dudley Zoo,
27.7.37. Captured in Britain and confined with domestic pigeons. Extremely
heavily infected with Brüelia glandarw. (M. RB.)
Tanagridae.
STEPHANOPHORUS DIADEMATUS (Temminck) 1829, White-capped Tanager
(1). Foxwarren Park, 14.1.37. Obtained from Northern Argentine. Confined
with various species of birds. No ectoparasites. (E. M.)
Ploceidae.
ESTRILDA ASTRILDA SANCTAEHELENAE Shelley 1903, St. Helena Waxbill
(12). Dudley Zoo, July-September 1937. All these birds were confined with
various species of birds. All negative for ectoparasites. (M. R.)
ESTRILDA MELPODA (Vieillot) 1817, Orange-cheeked Waxbill (4). Dudley
Zoo, 12.7.37, 23.7.37. Obtained from West Africa and confined with various
species of birds. Two specimens with a fairly generalized infection of Cheyletiella
sp.? (M. R.)
ESTRILDA SUBFLAVA (Vieillot) 1819, Zebra Waxbill (3). Dudley Zoo,
12.8.37, 24.8.37, and London Zoo, 1937. Obtained from Africa, confined
with various species of birds. No ectoparasites. (M. R. and T. C.)
ESTRILDA TROGLODYTES (Licht.) 1823, Grey Waxbill (1). London Zoo,
1937. Obtained from West Africa, confined with various other birds. No
Mallophaga or Siphonaptera. (T. C.)
OTHYPHANTES REICHENOWI (Fischer) 1884, Reichenow’s Weaver (1).
Dudley Zoo, 7.4.37. From Kenya Colony. Confined with various species of
birds. No ectoparasites. (M. RB.)
POEPHILA ACUTICAUDA (Gould) 1842, Long-tailed Finch (1). Foxwarren
Park, 14.12.36. Obtained from Australia. Confined with various birds. No
ectoparasites. (M. R.)
URAEGINTHUS BENGALUS (Linn.) 1866, Red-cheeked Cordon Bleu (2).
Dudley Zoo, 27.7.37. Obtained from Africa. Confined with various species of
birds. No ectoparasites. (M. R.)
EUPLECTES FRANCISCANA (Isert) 1789, Orange Bishop (1). Dudley Zoo,
1937. Obtained from West Africa. Confined with various species of birds.
No ectoparasites. (M. R.)
12 NOVITATES ZOOLOGICAE XLI. 1938.
HYPOCHAERA CHALYBEATA (Müller) 1776, Combassont (2). Dudley Zoo,
11.9.37. Both birds confined with various species of birds. One with a light
general infection of Brüelia sp.? The eggs laid singly, 1-3 on the same feather,
along the shaft of the breast feathers. (M. R.)
Monta MALaccA (Linn.) 1766, Black-headed Munia (1). Dudley Zoo,
30.7.37. Obtained from Ceylon, India. Confined with various species of birds.
No ectoparasites. (M. R.)
Pappo oryzivora (Linn.) 1758, Java Sparrow (1). Dudley Zoo, 12.8.37.
Confined with various species of birds. No ectoparasites. (M. R.)
Sturnidae.
SPREO SUPERBUS (Rüppell) 1845, Superb Glossy Starling (1). Foxwarren
Park, 10.8.37. Obtained from East Africa. Confined with other species of
birds. No ectoparasites. (M. R.)
LEUCOPHASER ROTHSCHILDI Stres. 1912, Rothschild’s Grackle (1). Fox-
warren Park, 7.2.37. Obtained from Bali Islands. Confined with various
species of birds. No ectoparasites. (M. R.)
Graculidae.
GRACULA RELIGIOSA Linn. 1758, Indian Grackle (1). Dudley Zoo, 30.7.37.
Obtained from Northern India. No ectoparasites. (E. M.)
Icteridae.
IcTERUS BONANA (Linn.) 1766, Martinique Oriol (1). Foxwarren Park,
4.1.37. Confined alone. No ectoparasites. (M. R.)
QUISCALUS QUISCULA (Linn.) 1758, Purple Grackle (1). Foxwarren Park,
24.2.37. Obtained from Southern America. Confined with various species of
birds. No ectoparasites. (M. R.)
Fringillidae.
Loxta €. CURVIROSTRA Linn. 1758, Crossbill (1). London Zoo, February
1937. Obtained in Europe. Confined with many various species of birds.
2 33, 4 22 Brüelia limbata and 3 33, 6 99 *Philopterus citrinellae. (T. C.)
CHLORIS CHLORIS (Linn.) 1758, Greenfinch (1). Dudley Zoo, 2.8.37.
Obtained near Oxford, England. Confined with various species of birds. No
ectoparasites. (M. R.)
ERYTHRURA PSITTACEA (Gmel.) 1788, Royal Parrot Finch (2). Mr. Part-
ridge’s Aviaries, 11.8.37. Obtained from Numea. No contact with other
species of birds. Many empty egg capsules on breast feathers. One specimen
of Philopterus sp.? (M. R.)
TIARIS CANORA (Gmel.) 1788, Cuban Grassquit (1). Dudley Zoo, 2.7.37.
Confined with other birds of the same species only. Two specimens of Liponyssus
sp.? (M. R.)
RICHMONDENA CARDINALIS (Linn.) 1758, Virginian Cardinal (4). Dudley
Zoo, 9.7.37, 23.8.37, 2.9.37, and 15.7.37. Obtained from Eastern U.S.A.
Three of the birds had light general infections of Liponyssus sp.? and three different
Briielia sp. ? were found among them. Eggs laid about the billand neck. (M. R.)
Noviratres Zoonogrcar XLI. 1938. 73
PAROARIA CRISTATA (Boddaert) 1783, Red-crested Cardinal (1), Dudley
Zoo, 6.9.37. Confined with various species of birds. Brüelia trithorax around
face and cheek. A few on the breast feathers. Eggs around face. A light
general infection with Cheyletiella sp.” (M. R.)
PAROARIA DOMINICANA (Linn.) 1758, Dominican Cardinal (3). Dudley
Zoo, 23.7.37, 16.7.37. From Eastern Brazil. Confined with P. cristata. Two
birds with Brüelia trithorax on head. Eggs around bill, throat, and top of head.
One specimen of Myrsides sp.? Also one specimen of Cheyletiella sp.? (M. R.)
On the third bird, a light infection of Brüelia sp. ? and *Philopterus citrinellae.
(E. M.)
Trochilidae.
CHRYSOLAMPIS ELATUS (Linn.) 1766, Ruby and Topas Humming Bird (1).
Ashton Wold, September 1936. Obtained from Brazil and only came into con-
tact with other species of humming birds. 1 4, 1 9, and 1 immature specimen
of Ricinus sp.n. Eggs laid singly on throat feathers. (M. R.)
EUPETOMENA MACROURA (Gmel.) 1788, Swallow-tail Humming Bird (1).
London Zoo, 1937. Confined with various species of birds. No Mallophaga or
Siphonaptera. (T. C.)
ACKNOWLEDGMENTS.
We would like to express our gratitude to all those who have so kindly co-
operated with us in preparing this list, in particular to Mrs. A. M. Hughes for
identifying the mites, and Mr. J. D. MacDonald for correcting the host names.
Our special thanks are also due to the Earl of Dudley, Mr. Alfred Ezra,
Lord Lilford, Mr. R. W. Partridge, and the Zoological Society of London for
permission to examine the birds dying in their aviaries; to Dr. H. Honigman,
Mr. A. F. Moody, and Mr. A. P. Shearing for the immense trouble they have
taken in dispatching the material and supplying data, and to Miss Elizabeth
Meyerhof and staff of the prosectorium of the Zoological Society of London for
examining birds in the authors’ absence.
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NOVITATES ZOOLOGIGAE.
H Zournal of Zoology.
BDITED BY
THE HON. MIRIAM ROTHSCHILD
AND. Dr. KARL JORDAN, F.R.S.
Vou. XULI
No. 2.
Pıgzs 75-173.
Issurp NOVEMBER 25TH, 1938, AT THE ZooLocicaL Museum, Trine.
PRINTED BY HAZELL, WATSON & VINEY, Lrp., LONDON AND AYLESBURY.
1938.
co
Ou
10.
11.
12.
13.
. FURTHER OBSERVATIONS
Vou. X LI.
NOVITATES ZOOLOGICAE
EDITED BY
THE HON. MIRIAM ROTHSCHILD and KARL JORDAN,
CONTENTS OF NO. II
. NOTES ON THE CLASSIFICATION OF CER-
CARIAE OF THE SUPERFAMILY NOTO-
COTYLOIDEA (TREMATODA), WITH
SPECIAL REFERENCE TO THE EXORE-
TORY SYSTEM ME
ON THE
EFFECT OF TREMATODE PARASITES ON
PERINGIA ULVAE (PENNANT) 1777
(PLATES I AND IM) ee
. WHERE SUBSPECIES MERT .
. ON SOME SIPHONAPTERA FROM TROPICAL
AFRICA AND IRAQ.
- ON SOME NEARCTIC SIPHONAPTERA
. NOTE ON POLYGONIA C-ALBUM (L.) 1758,
AT ASHTON WOLD, NORTHAMPTON-
SHIRE, AND THE RECORD OF A NOTABLE
ABERRATION (PLATE II, FIGS. 1 AND 2)
. ON THE REMARKABLE VARIATION OF
AN ORIENTAL HAWKMOTH (LEPIDO-
PTERA) (PLATE III, FIGS. 3-6)
. ON SOME NEW OR UNFIGURED LEPIDO-
PTERA FROM THE ORIENTAL AND
ETHIOPIAN REGIONS (PLATES III AND
IV)
. ON THE SPECIES OF BIRD -PARAPS YLLI
FROM THE FALKLANDS OBTAINED ON
THE BRITISH GRAHAM LAND EXPEDI-
TION, 1934-37 .
ON SOME OLD-WORLD ANTHRIBIDAR
THE EXCRETORY SYSTEM OF CERCARIA
CORONANDA N. SP. TOGETHER WITH
NOTES ON ITS LIFE-HISTORY AND THE
CLASSIFICATION OF CERCARIAE OF THE
SUPERFAMILY OPISTHORCHIOIDEA
VOGEL 1934 (TREMATODA)
ON SOME NEOTROPICAL SIPHONAPTERA
A NOTE ON THE FIN-FOLDS OF CERCARIAE
OF THE SUPERFAMILY OPISTHOR-
CHIOIDEA VOGEL 1934 (TREMATODA) .
Miriam Rothschild
Miriam Rothschild
Karl Jordan
Karl Jordan
Karl Jordan
Miriam Rothschild and
Malcolm Spooner
Karl Jordan
Karl Jordan
Karl Jordan
Karl Jordan
Miriam Rothschild
Karl Jordan
Miriam Rothschild
EXPLANATION OF PLATES I AND II (see pp. 101—102)
EXPLANATION OF PLATES III AND IV.
PAGES
75—83
84—102
103—111
112—118
119 —124
125
126—129
130—133
134—139
140—147
148—163
164—169
170—173
NCrmrrArHs ZOOLOGICAE
Vol. XLI. NOVEMBER 1938. No. 2.
NOTES ON THE CLASSIFICATION OF CERCARIAE OF THE SUPER-
FAMILY NOTOCOTYLOIDEA (TREMATODA), WITH SPECIAL
REFERENCE TO THE EXCRETORY SYSTEM.
By MIRIAM ROTHSCHILD.
(With text-figures 30-40.)
NUMBER of species of cercariae pertaining to the superfamily NoTo-
COTYLOIDEA (in all probability restricted to the family Notocotylidae)
are found parasitizing Peringia ulvae (Pennant) 1777 and Hydrobia ventrosa
(Montagu) 1803.
The group formed by these cercariae is well defined, the main characters
are easily recognizable and peculiarly distinct, but until now it has not been
subdivided satisfactorily. Dubois (1929) pointed out that Sewell’s (1922)
division into an “ Ephemera” and an “ Urbanensis ”” Group, based on size in
conjunction with the number of eye-spots, is not a natural one. He has described
a species, Cercaria helvetica I Dubois 1929, which bridges the apparent gap
between the two groups. In the light of present-day knowledge it would appear
that these characters are only of specific importance. Thus, for example, the
Cercaria of Notocotylus attenuatus Joyeux 1922 (nec Rud. 1809) is devoid of any
eye-spots, a condition no doubt associated with the habit of encysting within the
first intermediate host, while the closely related Cercaria of Notocotylus attenuatus
(Rud.) 1809 (for which species it was mistaken, in the adult form) has the more
usual complement of three.
While endeavouring to separate the various species found in P. ulvae and
H. ventrosa, it was discovered that these larvae could be divided into three
groups according to the structure of the anterior portion of the excretory vesicie.
This classification can be extended to include all the cercariae of the Noro-
COTYLOIDEA if the shape of the anterior end of the body is also taken into con-
sideration. It is hoped to prove by life-history experiments that these characters
are not merely exceedingly useful for practical work with larval forms, but can
also be employed in forming a natural classification of the adult worms.
A certain number of the known species of cercariae, however, are inade-
quately described and it is therefore impossible to assign them to their correct
position at present.
THE EXCRETORY VESICLE OF NOTOCOTYLOIDEA CERCARIAE.
The development of the excretory vesicle or closed tubular circuit has already
been described for C. ephemera Lebour 1907 (nee Nitzsch) (Rothschild 1935).
The same plan is found in all the other species from P. ulvae and H. ventrosa.
75
JUN 16 1939
76 Novirates ZooLoGIcAE XLI. 1938.
After the lateral tubules have grown forward as far as the oesophageal region
or the middle of the body they turn posteriad and divide into an antero-lateral
and postero-lateral branch (figs. 30-32). The excretory vesicle is formed by an
expansion of the main lateral excretory tubes, which in the fully developed
cercaria leave the bladder at the anterior cornu. In the oesophageal
region, however, each half grows away from the tubules towards the median
line of the body, and there fuses to complete the tubular circuit. This anterior,
transverse portion of the excretory system is the last to be developed. In the
30 31
SS) AS bo Sy
fully grown cercaria the vesicle is filled with large excretory granules and the
bunches of cilia within the lumen cannot be made out.
THE CHARACTERS OF NOTOCOTYLOIDEA CERCARIAE.
The main characters of the cercariae of this superfamily are as follows :
(1) Size of body medium to large (0-25 mm.—1-25 mm.).
(2) Body spatulate, generally slightly convex dorsally and concave ventrally.
(3) Pigment present to a greater or lesser degree in the body, concentrated
anteriorly.
(4) Two pigmented eyes with spherical lenses usually present, often with
median pigment spot between them, forming the so-called third eye. Eyes
may be totally wanting.
(5) Ventral sucker absent.
(6) Cuticle generally smooth, but spiny in some forms.
(7) Excretory bladder small, terminal, either spherical, triradiate or oval
(transversely elongate). Excretory vesicle formed by two wide tubes which
open into it antero-laterally or in or near the-middle of the anterior wall. These
tubes fuse in the oesophageal region forming the characteristic tubular circuit.
Vesicle filled with excretory granules.
(8) Alimentary canal consists of medium-sized oesophagus, simple bifid
(“ trielad ””) gut, the intestinal caeca reaching beyond the middle of the body.
Pharynx wanting.!
1 This character will almost certainly have to be omitted in the future, as there appear to be
adults of this superfamily with a pharynx,
Noviratres ZooLoGIcAE XLI. 1938, zer
(9) Locomotor pockets present at the posterior u of body.
(10) Penetration glands absent.
(11) Cystogenous glands numerous, highly specialized, containing refractile
particles.
(12) Tail generally well developed, thin and tapering, but in some forms
vestigial. Fin membranes absent.
(13) Reproductive organs of Notocotylid type.
(14) No second intermediate host. Encystment occurs either free or in
first host.
(15) Development occurs in a redia with intestine, but without collar.
(16) Intermediate host : Fresh-water, brackish-water or marine Gastropods.
(17) All free-swimming forms are phototropic.
The subdivisions can be made as follows :
A. Notocotylidae.
Cercaria without aural lappets or collar.
1. The Monostomi Group (figs. 33 and 36).
In this group the anterior transverse portion of the main excretory vesicle
is situated posterior to the median eye-spot or cerebral ganglion. Although
Cercaria ephemera Nitzsch 1807 belongs to this group and was the first to be
described, Cercaria monostomi von Linst. 1896 has been chosen as the typical
example in order to avoid confusion with Sewell’s classification.
Species which can be included here are :
Cercaria monostomi von Linst. 1896.
Cercaria ephemera Nitzsch 1807.
Cercaria ephemera Lebour 1907 (nec Nitzsch).
Cercaria inkermanni Sinitzin 1911.
Cercaria zostera Sinitzin 1911.
Cercaria pellucida Faust 1917.
Cercaria fulvoculata Cawston 1919.
Cercaria plana Faust 1922.
Cercaria trabeculata Faust 1924.
Cercaria of Notocotylus seineti Fuhrmann 1919.
Cercaria vaga Szidat 1933.
Cercaria imbricata Wesenberg-Lund 1934 (nec Looss).
Cercaria sp. (undescribed) from P. ulvae.
Cercaria sp. (undescribed) from P. ulvae.
2. The Impricata Group (figs. 34, 37 and 38).
In this group, of which the first species described was Cercaria imbricata
Looss 1896, the anterior transverse portion of the excretory vesicle forms a loop
between the lateral eye-spots which passes anterior to the median eye-spot and
cerebral ganglion. In profile it can be seen that this loop extends in a ventral
direction, and is not on the same plane as the lateral portion of the vesicle.
Species which may be included here are :
Cercaria imbricata Looss 1896.
Cercaria urbanensis Cort 1917.1
1 Vide Miller’s (1937) redescription.
78 Novitates ZootocicaE XLI. 1938.
Cercaria robusta Faust 1918.
Cercaria spatula Faust 1919.
Cercaria sp. (undescribed) from Hydrobia ventrosa.
3. The YENCHINGENSIS Group (figs. 35, 39 and 40).
In this group, of which Cercaria yenchingensis Faust 1930 was the first to
be described, there is an unpaired finger-like diverticula of the anterior transverse
portion of the vesicle, extending forward generally a little to one side of the median
eye-spot or cerebral ganglion. In the full-grown cercaria it is also filled with
excretory granules. It is the very last portion of the vesicle to be formed,
developing after the fusion of the two sides of the transverse band—and con-
sequently can be overlooked if descriptions are made from cercariae obtained
from crushed snails. It is therefore probable that some of the species included
in Group 1 will eventually, on redescription, be transferred to Group 3. In
this group we can include :
Cercaria yenchingensis Faust 1930.
Cercaria triophthalmia Faust 1930.
Cercaria lebouri Stunkard 1932.
Cercaria sp. (undescribed) from P. ulvae.
Cercaria sp. (undescribed) from P. ulvae.
Cercaria sp.‘ (undescribed) from Littorina neritoides (L.) 1758.
B. Pronocephalidae.
Cercaria with aural lappets or a collar.
The Inpicae XI Group.
In this group, the transverse band of the excretory vesicle is situated posterior
to the median eye-spot. It includes:
Cercaria indicae XI Sewell 1922.
Cerearia of Macravestibulum obtusicaudum Mackin 1930.
Cerearia of Macravestibulum eversum Hsü 1937.
? Cercaria aurita Faust 1918.
Species of Uncertain Position.
As already stated a certain number of these species are both insufficiently
described and figured, so that it is impossible to assign them to their correct
eroup. Among these we must include :
Cercaria hyalocauda Haldemann 1842.
Cercaria lucania Leidy 1877.
Cercaria konadensis Faust 1917.
Cercaria of Notocotylus attenuatus Joyeux 1922 (nec Rud.).
Cercaria of Notocotylus magniovatus Yamaguti 1934.
Cercaria osafuner Faust 1924.
Cercaria nakagawai Faust 1924.
Cercaria helvetica I Dubois 1929.
Cercaria No. I Petersen 1932.
Cercaria sp. Yokogawa 1934.
1 Dr, Averil Lysaght discovered this species, and very kindly presented it to me.
Noviratres ZooLogIcAE XLI. 1938. 79
THE SPECIFIC CHARACTERS OF THE NOTOCOTYLOIDEA CERCARIAE FROM
P. ULVAB AND H. voNnTRosa.
The usual difficulty is experienced in distinguishing between the related
species of these cercariae, and it was once again found, as in other groups, that
very few descriptions of these larvae are specifically diagnostic. Thus the name
Cercaria ephemera Lebour 1907 (nec Nitzsch) probably covers seven or eight
different cercariae. Even recent descriptions! are not very helpful in this
respect and do little more than place the species in one or other of the groups
mentioned above.
1 This applies particularly to life-history studies, where very little attention is paid to the
morphology of the cercariae (Szidat 1933).
80 NOVITATES ZOOLOGICAE XLI. 1938.
It is hoped that the following list of characters which are found to differ
specifically in NOTOCOTYLOIDEA cercariae from P. ulvae and H. ventrosa may
prove a useful guide to those engaged on the difficult and tedious task of describing
these larval forms from other molluscs.
It should be noted that size is a far more important character in this
group than in, for example, the Pleurolophocerca or Ubiquita cercariae. This
applies particularly to the diameter of the cyst, which is remarkably constant and
does not increase in size after formation, or vary according to the degree of
development of the encapsuled metacercaria.
Cercaria.
(1) Size, particularly relative length of body and tail.
(2) Degree of concavity of the body and “ death attitude ” under a coverslip.
(3) Pigmentation of the body.
(a) Colour.
(6) Density.
(c) Distribution.
(4) Development of eye-spots.
(5) Excretory system.
(a) Shape of bladder.
(b) Size of excretory pore.
(c) Length of anterior diverticula of excretory vesicle in Yenchingensis
Group.
(6) Size of cystogenous glands.
(7) Degree of development of the reproductive organs.
(8) Behaviour.
(a) Length of free-swimming life.
(b) Site selected for cyst formation.
(c) Length of duration of cyst formation.
(d) Stage at which cercaria leaves the rediae to continue development
in the tissues of the snail.
Redia.
(1) Pigmentation.
(2) Relative length of intestine and body.
(3) Colour of contents of intestine.
(4) Type and number of crystal inclusions in the intestine.
(
one redia.
Cyst.
(1) Size.
(2) Situation.
Descriptions of species from other molluscs suggest that specific differences
may also be found in the structure and position of the “ locomotor pockets,” the
cystogenous glands and the internal structure of the tail.
SUMMARY.
(1) A division into two main sections is suggested for cercariae of the
superfamily NoTOCOTYLOIDEA. Cercariae of the family Notocotylidae can be
Novirares Zoonoaioar XLI. 1938. 81
further divided into three groups according to the structure of the anterior
transverse portion of the excretory vesicle.
(2) Attention is drawn to the specific differences of these cercariae from
P. ulvae and H. ventrosa.
ACKNOWLEDGMENTS.
I would like to express my gratitude to Mr. G. M. Spooner for supplying
collections of Hydrobia ventrosa, and to the Editors of Parasitology (Cambridge),
the Journal of Parasitology (Urbana) and to the Institut d’Egypte for their
great kindness in allowing me to reproduce diagrams or parts of diagrams from
their journals and publications.
EXPLANATION OF TEXT-FIGURES.
Fig. 30. Development of Excretory System of a Notocotylid cercaria of the
Impricata Group from H. ventrosa. Four flame-cell stage.
Figs. 31-32. Later stages in the development of the same species. (The
bundles of cilia in the collecting tubes, and the flame-cells, are omitted.
Pressure from the coverslip has distorted the outline of the specimen in
Fig. 31.)
Fig. 33. Diagram showing the formation of the anterior portion of the excretory
vesicle in the Monostomi Group.
Fig. 34. Diagram showing the formation of the anterior portion of the excretory
vesicle in the IMmBRICATA Group.
Fig. 35. Diagram showing the formation of the anterior portion of the excretory
vesicle in the YENCHINGENSIS Group.
Fig. 36. Cercaria ephemera Nitzsch (after Sinitzin), showing the typical forma-
tion of the excretory vesicle in the Monostomi Group.
Fig. 37. Cercaria imbricata Looss (after Looss) during development, showing
the loop of the anterior portion of the excretory vesicle. The third eye-spot
is not yet developed.
Fig. 38. Cercaria robusta (after Faust), showing the typical formation of the
excretory vesicle in the IMBRICATA Group.
Fig. 39. Cercaria yenchingensis (after Faust), showing the anterior unpaired
diverticula of the excretory vesicle.
Fig. 40. Cercaria lebouri (after Stunkard).
EXPLANATION OF LETTERING.
Antero-lateral collecting tube.
Bunch of cilia.
Cerebral ganglion.
Unpaired diverticula of anterior portion of excretory vesicle.
Excretory vesicle.
Flame-cell.
Lateral eye-spot.
Median eye-spot.
Postero-lateral collecting tube.
Main collecting tube.
Hee ee bomt
82 Novirates ZooLoGIcAE XLI. 1938.
REFERENCES.
Cort, W. W., 1914. Larval Trematodes from North American Fresh-water
Snails.—In J. Parasitol. Urbana, 1, 1914, pp. 65-84, 15 figs.
Cort, W. W., 1915. Some North American Larval Trematodes.—In Illinois
Biol. Monog., 1, No. 4, 1915, pp. 1-86, 8 pls.
Dubois, G., 1929. Les Cercaires de la Région de Neuchätel.—In Bull. Soc.
Neuchatel Sci. Nat., 58 (nouv. sér., t. 2), 1929, pp. 1-177, 17 pls.,
8 figs.
Ewarts, H. C., 1880. Cercaria hyalocauda Haldemann.—In Amer. Month.
Microsc. J., 1, 1880, pp. 230-232, 3 figs.
Faust, E. C., 1917. Life-history studies on Montana Trematodes.—In Illinois
Biol. Monog., 4, No. 1, 1917, pp. 1-121, 9 pls., 1 fig.
Faust, E. C., 1917. Notes on the Cercariae of the Bitter Root Valley, Montana.
—In J. Parasitol., 3, 1917, pp. 105-123, 1 pl.
Faust, E. C., 1918. Studies on Illinois Cercariae—In J. Parasitol., 4, No. 3,
1918, pp. 93-110, 2 pls.
Faust, E. C., 1919. The excretory system in Digenea. III. Notes on the
excretory system in a Monostome Larva, Cercaria spatula nov. spec. —In
Biol. Bull., 36, No. 5, 1919, pp. 340-344, 3 figs.
Faust, E. C., 1919. Notes on South African Cercariae—In J. Parasitol., 5,
1919, pp. 164-175, 1 pl. ;
Faust, E. C., 1922. Notes on Larval Flukes from China.—In Parasitology,
Cambridge, 14, Nos. 3 and 4, 1922, pp. 248-267, 2 pls.
Faust, E.C., 1924. Notes on larval flukes from China. II. Studies on some larval
flukes from the Central and South Coast Provinces of China.—In Amer.
J. Hyg., 4, No. 4, 1924, pp. 241-301, 2 pls., 3 figs.
Harper, W. F., 1929. On the structure and life-histories of British fresh-water
larval Trematodes.—In Parasitology, Cambridge, 21, 1929, pp. 189-219,
5 figs.
Horsfall, M. W., 1930. Studies on the structure of Cercaria infracaudata n. sp.—
In J. Parasitol., 17, pp. 43-48, 1 pl., 1930.
Horsfall, M. W., 1935. Observations on the life-history of Macravestibulum
obtusicaudum Mackin. 1930 (Trematoda : Pronocephalidae).—In Proc.
Helminth. Soc., Washington, 2, No. 2, 1935, pp. 78-79.
Hsü, D. Y., 1937. Life-history and morphology of Macravestibulum eversum
sp. nov. (Pronocephalidae, Trematoda).—In Trans. Amer. Mier. Soc.,
Menasha, 56, No. 4, 1937, pp. 478-504.
Joyeux, C., 1922. Recherches sur les Notocotyles —In Bull. Soc. Path. exot.,
Paris, 15, 1922, pp. 331-343, 9 figs.
Kobayashi, H., 1922. A review of Japanese Cercariae.—In Mitteil. med. Acad.,
Keijo, 1922, pp. 1-27, 5 pls.
Lebour, M. V., 1907. Larval Trematodes of the Northumberland Coast.—In
Trans. Nat. Hist. Soc., Northumb., 1, Pt. 3, 1907, pp. 437-454, 5 pls.
Lebour, M. V., 1912. A review of the British Marine Cercariae.—In Parasitology
Cambridge, 4, No. 4, 1912, pp. 416-456, 4 pls.
Leidy, J.,.1877. On Flukes infesting Mollusks.—In Proc. Acad. Nat. Sci.,
Phila., 29, 1877, pp. 200-202.
Looss, A., 1896. Recherches sur la faune parasitaire de Egypte. (Premiere
Partie.—In Mém. Instit. Egypt., 3, fasc. 1, 1896, pp. 1-252, 16 pls.
Noyirates Zoonoaican XLI. 1938. 83
Miller, E. L., 1936. Studies on North American Cercariae.—In Jilinois Biol.
Monog., 14, No. 2, 1936, pp. 1-125, 8 pls.
Nitzsch, D.C. L., 1807. Seltsame Lebens- und Todesart eines bisher unbekannten
Wasserthierchens.—In Kilian, Georgia, 257-262; 281-286. Cited after
Braun 1893.
Petersen, H., 1932. Cercarien der Niederelbe.—In Zool. Anz., Leipzig, 97, 1932,
pp. 13-27, 20 figs.
Rothschild, M., 1985. Note on the excretory system of Cercaria ephemera
Lebour, 1907 (nec Nitzsch).—In Parasitology, Cambridge, 27, No. 2, 1935,
pp. 171-174, 7 figs.
Sewell, R. B., 1922. Cercariae indicae.—In Ind. J. Med. Res., 10, Suppl. Number,
1922, pp. 1-870, iii, and 32 +, 7 pls.
Sinitzin, D. T., 1905. Studies on the life-cycle of Trematodes. The distomes
of the fish and frogs in the environs of Warsaw (Russian).—In Mem. Soc.
Nat. Varsovie, Biol., 15, 1905, pp. 1-210.
Sinitzin, D.T.,1911. Parthenogenetic Generation of Trematodes and its Progeny
in the Molluses of the Black Sea.—In Mem. Acad. Sci. St. Petersb., 30,
Ser. 8, 1911, pp. 1-127, 6 pls., 17 figs.
Stunkard, H. W., 1932. Some larval Trematodes from the Coast in the region
of Roscoff, Finistere.—In Parasitology, Cambridge, 24, No. 3, 1932, pp.
321-343, 13 figs.
Szidat, L., 1930. Die Parasiten des Hausgefliigels. 4. Notocotylus Diesing und
Catatropis Odhner, zwei, die Blinddärme des Geflügels bewohnende mono-
stome Trematodengattungen, ihre Entwicklung und Ubertragung.—In
Arch. Geflügelkunde, Berlin, 4, 1930, pp. 105-114, 7 figs.
Szidat, L. and U., 1933. Beiträge zur Kenntnis der Trematoden der Mono-
stomidengattung Notocotylus Dies—In Zbl. Bakt. Jena Orig., 129, 1933,
pp. 411-422, 12 figs.
Wesenberg-Lund, C., 1934. Contributions to the development of the Trematoda
Digenea. Part II. The biology of the fresh-water Cercariae in Danish
fresh waters.—In K. danske Vidensk. Selsk. Skr., 5, Ser. 9, No. 3, 1934,
pp. 1-223, 39 pls.
Yamaeuti, S., 1938. Zur Entwicklungsgeschichte von Notocotylus attenuatus
(Rud., 1809) und N. magniovatus Yamaguti, 1934.—In Z. Parasitkd., 10,
No. 2, pp. 288-292.
84 Noviratrs ZooLoGIcaE XLI. 1938.
FURTHER OBSERVATIONS ON THE EFFECT OF TREMATODE
PARASITES ON PERINGIA ULVAE (Pennant) 1777
By MIRIAM ROTHSCHILD.
(With Plates I and II and text-figures 41-47.)
INTRODUCTION.
| PESOUCEIOUT the investigations on the trematode parasites of Peringia
ulvae an attempt has been made to establish various points regarding the
relationship of parasite and host. Those noted here concern the relative changes
of growth in male and female snails, and the probable “‘ sex-reversal ”’ of the host.
Experimental evidence is required to prove both these phenomena, but as
this is not at present available the discussion is based on the results of examining
and sampling large collections of P. ulvae from different localities. Each collec-
tion has necessarily been dealt with separately and a comparison made between
the infected and uninfected portions of the sample.
One of the major difficulties is the lack of a suitable method for estimating
the age of the snails. As will be seen later, size is no criterion and so far has not
proved useful in this respect.
Pelseneer (1928) first drew attention to the fact that the males of gastropod
molluscs are more frequently infected than the females. He noted one excep-
tion, Turritella communis Lamark 1785, but Rothschild (1936) found that at
Plymouth this species conformed to the rule. Pelseneer (1906 and 1928) also
originally pointed out the reduction of the penis in infected snails, due in his
opinion to the partial or complete castration of the host by the parasites.
Wesenberg-Lund (1931) was, however, the first to put forward the view that
changes in sex might be directly due to the effect of trematode parasites. He
found that in the hermaphrodite snail Succinea putris (L.) 1758 the female organs
are totally destroyed, but the production of spermatozoa continues after this
for some time—although greatly decreased—before castration is complete.
“With some right it may be maintained that many parasitized snails pass
through a male stage, or in other words that the parasite alters a hermaphrodite
organism into a male.”
Krull (1935), apparently unaware of either Pelseneer’s or Wesenberg-Lund’s
observations, published an interesting paper on the morphology of various
snails, including Peringia ulvae (= Paludestrina ulvae), in which he suggests that
those infected specimens showing a reduced penis are in reality “ sex-reversed ”
females. He bases his belief on the discovery of specimens in which the internal
organs were only partially destroyed and which he considered were unquestionably
female, while a small functionless penis had also been developed. He does not
suggest that these snails ever function as males. “ Sex-reversal ” is therefore a
rather unsatisfactory term to employ. In default of a better one it is used
by the author of this paper.
The change in relative size of parasitized male and female snails has
not received any attention until now. In view of the gigantism produced in
P. ulvae and various other snails by trematode parasites (Rothschild 1936) it was
Novirarrs Zootogican XLI. 1938. 85
thought probable that the normal differences in size between the sexes might be
noticeably altered, irrespective of whether the males are more frequently infected
as appears to be the case with most species, or whether a certain proportion of the
infected ‘‘ males ” are in reality sex-reversed females.
THE NORMAL Sex Ratio or P. urvan.
In nature the relative number of males and females of P. ulvae seem to
vary considerably, according to both size (age) and distribution, and other
unknown factors, and therefore each sample must be examined separately.
Pelseneer (1926) states that in the Mollusca as a whole, it is usual to find the
males outnumber the females in the whole population—their numbers being
greatly in excess of the females in the young stages, approximately equal when
they become sexually mature, and somewhat fewer in the older stages.
Pelseneer himself examined 115 specimens of P. ulvae from the mouth of the
River Slack (Port d’Ableteuse) and a further 663 collected from the mouth of
the Canche (Etaples). Sixty-eight per cent. of the former and 58 per cent. of
the latter were females. This surprised Pelseneer, who thought that some of
the females might be in reality males—the penis having regressed after the breed-
ing season as happens with Littorina littorea (L.) 1758, and various other species.
He records trematode parasites, but we can infer from his remarks that the
rate of infection was low.
TABLE I.
SNAILS BETWEEN 1 AND 21 mm. IN LENGTH.
ds 92 Total
Tamar Saltings, February 1938 . 6 17 33 50
Tamar Saltings, March 1938 5 j 23 51 74
St. John’s Lake, February 1938 . F 63 137 200
St. John’s Lake, June 1937 5 13 36 49
Although the females do appear to outnumber the males in numerous
samples examined, yet my observations in no way support the theory of a seasonal
regression of the penis. Only four uninfected males have been noted with a
reduced penis, although numbers have been looked at during all seasons of the
year. ‘These four specimens can be regarded as abnormal examples, just as a
few individual snails were found with a split or reduced tentacle.
It is very doubtful, however, if the sex of specimens under 22 mm. can be
determined with certainty. Among these very small snails specimens without a
penis greatly outnumber those with one. The penis is then found in all stages
of development, and it is quite probable that many of these very small snails
are males in which the penis has not developed. In comparing the sex ratio it
has been found safer to ignore all snails under 2? mm. in size. Figures for the
small size groups are given in Table I. It will be seen that apparent females
regularly outnumber recognizable males by about 2 to 1.
The distribution of P. ulvae is patchy, but when it is present it is often
found in enormous numbers. As many as 28,400 per square metre are recorded !
1 Mr. G. M. Spooner supplied this figure in a private communication.
86 NoyirarEs ZoonocicaE XLI. 1938.
from St. John’s Lake, Plymouth. It is not at all unusual to find a considerable
difference in the sex ratio for random samples collected a few feet apart, or at
the same ground on different occasions. Thus from the same area of St. John’s
Lake, Plymouth, collections of snails gave the following results: 39 per cent.
36 (6 per cent. infection), 38 per cent. 34 (3 per cent. infection), 60 per cent.
dd (4 per cent. infection), 26 per cent. g3 (2 per cent. infection), 48 per cent.
33 (3 per cent. infection). Similar variation was discovered in samples from
Millport (West Scotland), Holbeach (Lincoln), Wittering (Sussex), Poole Harbour
(Dorset), and Blakeney (Norfolk). Extensive sampling would be necessary for
an exact estimate of the sex ratio of the whole population of any given area.
If infection is high it
is very important to
record the number of
uninfected males and
females of each sample.
Krull unfortunately
completely ignores this
point, and does not re-
cord the sex ratio of the
uninfected specimens
which he examined.
100
80
fon
=)
THE VARYING GROWTH
OF P. ULVAE AND THE
RELATIVE SIZE OF
MALES AND FEMALES.
>
o
*suemtoeds Jo zequmy
20 Examination of
samples of P. ulvae
from various habitats
shows that their size
varies greatly irrespec-
tive of the age of the
Fig.4]: sanples showing uneven spat fall. snails and the degree
Holbeach Saltings ,August. of parasitism (Pl. 1).
Bes On the whole those
populations of P. ulvae
commonly found in saltings attain a larger size than the populations on mud flats
(Pl. IL, figs. 1,2). In addition, the size of the snails varies in a most remarkable
manner from one permanent pool in the saltings to another. This must partly be
due to irregularity of spat fall, which is a feature of this type of habitat and
accounts for those pools which contain hundreds of thousands of tiny specimens—
generally fringing the edge—none of which exceed 2-3 mm. in length, and other
pools in which it is difficult to find any small snails at all, and a third type in
which there are large and small specimens, but few or no intermediate stages
(fig. 41).
Apart from this there are strong indications that the growth-rate itself
varies from one pool to another. The variation in growth referred to above is
generally associated with a difference in shell texture, a difference in shell colour
and shape and, somewhat surprisingly, in extreme cases with a definite difference
Size groups in mm.
Novirates ZootocicaAn XLI. 1938. 87
in the consistency of the soft parts of the body. It seems likely that it is
dependent on environmental conditions, since both extremes of growth and of
shell texture are found in different types of pools in the same saltings. On the
mud flats the largest shells are not found at all; it is unusual to discover a
specimen exceeding 5} mm. in samples from this habitat.
The difference in growth greatly complicates the study of the effect of para-
sitism on the various populations, as it makes it almost impossible to gauge the
age of the snails. It is difficult to say with certainty, unless observations are
made over a long period, whether a pool is populated with all comparatively young
specimens or whether growth conditions in the pool are particularly poor.
Irrespective, however, of whether the environmental conditions tend to
produce large or small snails, those individuals infected with trematode parasites
are relatively larger. Thus, in certain areas of the saltings of the River Tamar,
Plymouth (higher up the estuary than St. John’s Lake), very large-sized uninfected
samples are found with snails ranging up to 64 mm. in length. In this area
heavily infected pools are stocked with P. ulvae displaying super-gigantism, where
shells of 10-102 mm. are found quite frequently (Pl. I, rows G and K).
With many uncertain factors such as age, growth rate, uneven spat fall,
ete., to take into account, it is difficult to decide whether there is a true difference
in the size and growth-rate of the sexes, such as is demonstrated for Littorina
httorea (H. B. Moore, 1937) or whether the difference in size between the largest
males and females of P. ulvae is due to the longer life of the latter. Pelseneer
(1926) believes that this accounts for the greater size of the females in many
species of molluses—the males dying off before they attain the dimensions of the
largest females.
It appears to be a fairly general rule with P. ulvae that, whatever the relative
number of the sexes of the whole sample may be, the largest uninfected specimens
are females, and that the uninfected females outnumber the uninfected males
in the largest size groups.
The three following examples are typical. A random sub-sample of 530
from a collection of 10,000 snails from St. John’s Lake, Plymouth (June 1937),
showed 332 (62 per cent.) females and 198 (38 per cent.) males. Forty-six of the
largest specimens were selected. Twenty-eight were uninfected. Of these
24 (85 per cent.) were females and 4 (15 per cent.) males. This 6: 1 ratio con-
siderably exceeds the 1-7: 1 ratio for the whole sample and the difference proves
statistically significant. A random collection of several hundred snails from
Wittering showed that in the largest size groups, 64-5 mm., there were only
35 per cent. males, but among the smaller snails (43-33 mm.) males outnumbered
females by 6 to 1. Several hundred specimens from Millport displayed 68 per
cent. males for the whole sample: of the 30 largest uninfected snails opened, 22
(73 per cent.) were females.
In a collection of several thousand snails from Poole Harbour Zostera-beds
(August 1937) the uninfected females outnumbered the uninfected males by 3: 1
in the largest size groups. Several hundred snails from a pool in Burnham
Saltings (August 1937) showed a 4: 1 excess of females among the largest unin-
fected snails. A similar figure was shown for a Holbeach sample (August 1937)
from grass sweepings. It is perhaps worth noting two exceptions. A collection
of snails from a pool at Blakeney (August 1937) showed 16 males and 15 females
among the largest size group. The ratio of the rest of the sample was unfor-
88 Novirates ZOOLOGICAE XLI. 1938.
tunately not recorded. In the sample from the Tamar Saltings, with 100 males
and 80 females, 12 uninfected males and 11 females were found in the largest
size groups.
Thus even in pools in saltings, a habitat where the irregularity of spat fall
might well produce some inconsistency in the results, the largest uninfected
females were usually larger than the uninfected males. As will be seen in these
instances the mean size of uninfected specimens of the whole sample, or of the
largest size groups, was sometimes approximately equal.
ANALYSES OF SAMPLES OF P. ULVAR.
The samples described below were selected for the following reasons. They
were collected from different parts of Britain, and somewhat different habitats.
In addition they showed the greatest contrast in the degree of parasitism by the
various groups of cercariae.
The snails were collected by scooping them up with the mud from the
pools in the saltings or off the mud flats. Collections from the plant roots on the
saltings were made by sweeping the grass, and from Zostera by washing the snails
from the plant in fresh water. Snails below 23 mm. in length were subsequently
discarded for the reasons given above. Fig. 41 includes the measurements of
entire sub-samples from single pools in the saltings. These were obtained by
sieving the mud from permanent pools with a fine-mesh sieve. It illustrates the
irregularity of spat fall in this habitat—the intermediate, large, or small sizes
being sometimes absent.
Hunterdon Sands (Sample C) (Pl. I, Row D).
This sample was collected in May 1937 from the edge of the sands, along a
rather extended front, and not from one small pool. The various sizes were
found mingled together. After discarding the smaller snails, 239 living specimens
remained for examination. The mean size of the snails was not great (4:58 mm.),
but a number of them attained dimensions and variation in shape only associated
with heavy infections and good growth conditions. The shells were of the
hard grey type. Eighty-four specimens (34 per cent.) were infected, and of these
no less than 57 proved to be harbouring Echinostome cercariae (see Appendix).
Almost all these infections were very heavy and many cercariae were emitted.
There was a striking difference in the size of infected and uninfected snails, the
former ranging between 71-33 mm., with a mean length of 5-62 mm., the latter
between 51-2} mm. with a mean length of 4:02 mm. This suggests that the
infections were acquired at an early age.
Sixteen of the infected snails were required for life-history experiments and
were put aside after their infections had been identified. The rest of the sample
was dissected and it was found that 153 were males (i.e. snails with a penis) and
70 females. Most of the females were gravid and some laid eggs before they were
killed. The size of the uninfected males and females showed the usual difference
referred to above. The largest specimens were females, no uninfected males
reaching 54 mm. in length. The mean length of the females was also larger,
4:26 mm. compared with 3-82 mm.
Of the 68 infected snails which were dissected 64 proved to be males and
only 4 females. When the mean lengths of the males and females of the whole
sample are compared it will be seen that the males are larger (4:5 mm. for 3g,
Novirates ZooLoGIcAE XLI. 1938. 89
4-2 mm. for 99). This sample therefore displays a complete reversal of the
difference in size generally exhibited by the sexes. Yet if the uninfected snails
alone are considered it is perfectly normal in this respect.
Another suggestive feature was the comparative lightness of the female
infections ; eggs were present, indicating that these were of comparatively recent
origin—the destruction of the gonads not being far advanced. In three out of
four of these infections the rediae and sporocysts had not reached the cercariae-
producing stage. This, taken into consideration with the fact that the rest of
the infections were heavy, inevitably leads one to suspect that some of the
snails with a penis were in the first place females which had “ reversed” as the
infection developed.
In all these infected males, the penis was more or less reduced—a feature
which is discussed in a later portion of this paper.
SUMMARY OF HUNTERDON SANDS (SAMPLE C).
(Mean size with standard error of the mean given in brackets.)
(god 89
J (3:82 + 0:063 mm.)
Uninfected 155 22 66
(4:02 + 0-051 mm.) U (4:26 + 0-074 mam.) | 153 g3 with 64 infec-
Larger size of | tions = 42 per cent.
Total 239 Lah }
(4-58 mm.) B ee 70 99 with 4 infections
| ag 64 = 6 per cent.
(5:65 + 0:125 mm.)
224
11 mm.) 1 ( ca. 4:44 mm.)
mie
Not examined 16
L (5:80 mm.)
Marked differencein Numbers of 99 too Very marked difference in
size. small for adequate infection-rate of the two
estimate of mean sexes.
SIze.
Holbeach Saltings (Sample A) (Pl. I, Row J).
This sample contained a very high percentage of the Pleurolophocerca group
of cercariae. It was collected from a permanent pool on the extreme upper
edge of the saltings at Holbeach (Lincoln) in August 1937. Two scoops of mud
were made from the centre of the pool. The sample contained 326 living
snails over 24 mm. in length; 33 of these died before the dissections were
completed.
The shells were very dark reddish brown with the spires well preserved.
There was a certain amount of individual variation in shell texture. The growth
conditions were obviously good on the whole, the mean length of the snails
being large (5:40 mm.) and the maximum length attained 8! mm. The contrast,
however, between the uninfected and infected portions of the sample was not so
great as in the collection from Hunterdon Sands. The percentage of infection
was about 50. Of the 148 infections 90 pertained to the Pleurolophocerca
group of cercariae (see Appendix). Fifty-five of the largest specimens were
spared for experiments. The remaining snails of the whole sample revealed on
dissection 104 males and 134 females. The uninfected females outnumbered the
males by 91 : 54, and, as is usually the case, the largest snails of this portion of
the sample were females. No male attained a greater length than 6} mm.,
whereas females reached 61 mm., and the five largest size groups contained 31
females compared with 14 males.
90 Noyirates ZooLocicaE XLI. 1938.
The disproportion in infected females is slight compared with the sample
from Hunterdon Sands. Also no reversal of the normal size of the sexes is
shown in this case and both the individual and mean sizes of infected female
snails are greater than those of the males. However, not much importance
should be attached to this, owing to the fact that the sex of 55 of the largest
infected snails was not determined.
SUMMARY OF HOLBEACH SALTINGS (SAMPLE A).
(Mean size with standard error of the mean given in brackets.)
40
Ope!
We)
m
[ Uninfected 145
(5:20 + 0:060 mm.) (5-20 mm.) 104 3g with 50 infected
= 48 per cent.
Di ifference in size not
SOLO Cui | 134 99 with 43 infected
Total 293 [ 3g 50 = 32 per cent.
(5-40 mm.) 4 (5:32 mm.)
(5-36 mm.)
Difference in size not
Infected 148 significant,
| (5-60 + 0-063 mm.)
Not examined 55
L (6:05 mm.)
Larger mean size of Difference in percentage
infected significant. infection significant
(even though largest
specimens not = ex-
amined).
Holbeach Saltings (Sample C) (Pl. I, Row L).
A collection of about 800 snails was made from a permanent pool in the middle
zone of the saltings, alongside another pool in which very large specimens had
been found. This sample contained only relatively small shells, and their friable
nature and shape, together with the condition of the soft parts of the bodies of
the snails, suggested poor growth conditions rather than a collection of young
snails only. Twenty-five of the largest specimens were selected and examined.
These ranged in size from 54-3} mm. Ten were not infected and of these 8
were females and 2 males (see Appendix). The largest uninfected specimen
was as usual a female.
Of the fifteen infected snails ranging between 54-4 mm. in length, 8 were
females and 6 males. The sex of one specimen was not determined. Thus con-
siderably more males were infected than would be expected from a comparison
of the sex ratio. The mean size was 4-73 mm. as against 4-38 mm. for infected
examples.
Holbeach Saltings (Sample E) (Pl. I, Row F).
This collection of about 1,000 snails was made by sweeping the grass round
a permanent pool in the middle zone of the saltings, where the percentage of
infection had been found to be high, and growth conditions favourable. The
snails obtained from the grass sweepings were of the same type, but not so large.
This habitat does not favour a high infection-rate and the snails rarely, if ever,
reach the dimensions of those found in many of the pools.
Twenty-five of the largest snails, ranging between 63-5} mm. in length,
were selected. Four only were infected. Of the 21 uninfected specimens 16
were found to be females and 4 males, ‘The sex of one snail was not determined.
Novirates ZooLoGIcAE XLI. 1938. 9]
Three of the infected specimens were males and one female. It will thus be seen
that although the females in the largest size groups outnumber the males by 4: 1,
yet the infected males outnumber the females by 3: 1.
Uninfected females reached a size of 64 mm., but the only males to reach
these dimensions were infected.
Burnham Saltings (Sample A) (Pl. I, Row E).
A collection of about 1,000 snails was made in August 1937 from a permanent
pool in the middle zone of the saltings near Burnham (Norfolk). Populations of
P. ulvae from this area showed an exceptionally low incidence of infection.
The growth conditions of this pool appeared favourable and the shells were
of the hard grey type, with very well preserved spires. Twenty-five of the largest
were selected, of which two were empty. The range of the remaining 23 was
71-51 mm. Two specimens, one male and one female, were infected. Seventeen
of the uninfected snails were females, with a mean length of 6-76 mm., and only
4 males, with a mean length of 5:75 mm. Here again the females outnumber the
males by over 4: 1, yet there was one male among the two infected specimens.
Tamar Saltings (Samples Al and A) (Pl. I, Rows G and K; Pl. II, fig. 1).
It was discovered in October 1937 that growth conditions were very favour-
able for P. ulvae in certain pools in the upper levels of the saltings on a stretch of
ground locally known as “ Egypt ” (see Map, Hartley & Spooner, 1938). Here
uninfected snails reached 64 mm. in length. The shells were of the very hard
type, red in colour with blunted spires. A search was made for a highly infected
sample, and this was ultimately found in a permanent pool with a very soft
muddy bottom. Here large specimens, ranging up to 102 mm. in length, were
discovered lying on the surface of the mud. About fifty of these were collected
by hand and all proved to be infected. Scoops were made round the edges of
the pool, but there appeared to be a complete absence of the smaller sizes of
snails. In March 1938 a larger and strictly random collection was made in the
same pool, and on this occasion a number of smaller specimens were found, but
there was a marked absence of intermediate sizes (fig. 41).
The total collection (March 1938) numbered 2,200 specimens. The shells
were red, exceptionally hard, with blunted spires and corroded surface in the
larger sizes. Some of the smaller shells were grey with well-preserved spires.
A random sub-sample of 254 specimens was dissected (see below). After the
sub-sample had been taken, 33 of the largest snails were selected from the rest of
the collection and opened (Sample Al). They ranged in size from 101-7 mm. in
length, with a mean of 8-24 mm. All were infected (see Appendix). Twenty-
five were males and 8 were females. Two of the females contained developing
eggs.
Seventy-four of the sub-sample were under 2? mm. and were discarded.
Details of these smaller specimens are given in Table I. The mean size of the
remaining 180 specimens, ranging between 9-2} mm., was 5-48 mm. One hundred
were males (mean length 5-86 mm.) and 80 females (mean length 5-00 mm.).
The uninfected females, however, outnumbered the uninfected males in the
ratio 69 : 55.
The sample displayed a 31 per cent. infection. Forty-five of the infected
specimens were males (mean length 6-96 mm.) and only 11 were females (mean
8
92 NOVITATES ZOOLOGICAE XLI. 1938.
length 7:36 mm.). The difference in the proportion of the sexes infected is
clearly significant. The gigantism displayed by infected snails (mean 7-04 mm.
compared with 4-77 mm.) is very marked, but on the other hand there is no
evidence regarding the reversal of the normal differences in size displayed by
the sexes.
Attention should here be drawn to a unique and peculiar feature of the
samples from this area. With one exception (see p. 93) no hermaphrodite
examples of P. ulvae have been found by me. However, among the largest
size groups from this pool and a neighbouring uninfected pool, a few gravid
females were discovered with a swelling or papilla in the region of the penis,
sometimes to the left of the body. No trace of such a swelling is generally visible.
It was therefore thought safer to regard 50 per cent. of infected specimens with a
papilla as females. Ten specimens fell into this category.
SUMMARY OF TAMAR SALTING (SAMPLE A).
(Mean size with standard error of the mean given in brackets.)
Uninfected snails 124 [ 38 55
(4:77 + 0:12 mm.) (4:96 + 0-17 mm.)
69
(4:63 + 0:16 mm.) ) 100 33 with 45 infec-
Total 180 tions (45 per cent.)
80 9° with 11 infections
bh 45 (13 per cent.)
Infected snails 56 (6:96 + 0:14 mm.)
(7:04 + 0-125 mm.) Oll
(7-34 + 0-20 mm.)
Difference in size Difference in each Difference in proportion
clearly significant. case not significant. of sexes infected signi-
ficant.
St. John’s Lake (Sample A) (Pl. I, Row B, PI. II, fig. 2).
This collection contained approximately 10,000 snails and was obtained from
the mud flats of St. John’s Lake (Trevol Range), Plymouth, in June 1937. Of
these 530 were examined to determine the sex ratio. There were 332 females
and 198 males. The percentage of infection was low (3 per cent.), and in order to
avoid the dissection of a large number of specimens 46 of the largest snails of the
whole sample were selected. Eighteen, about 40 per cent., were found infected
(see Appendix). As usual the females outnumbered the males among the largest
uninfected specimens of the sample: on this occasion in the proportion 6: 1, as
compared with under 2:1 for the whole sample (see p. 87).
Twelve of the infected snails were females and 4 males. The sex of one
specimen was not recorded. In this sample therefore the proportion of infected
males is only slightly more than would be expected if infection was governed
solely by chance, and indeed is not statistically significant.
The majority of the infections (11) pertained to the Oocysta group of
cercariae.
THE EFFECT or PARASITISM WITH TREMATODES ON THE PENIS OF P. ULVAE.
Very little attention has hitherto been paid to the effect of trematode
parasites on the external sex organ, although the destruction of the gonads has
been described constantly. I have found that the penis of infected P. ulvae
is invariably abnormal, It is almost always possible to recognize an infected
Noyirates Zoonogicakn XLI. 1938. 93
specimen by the casual examination of this organ. However, in light infections,
or young infections, this is much less marked.!
The abnormality consists of reduction in size, which may be slight, or so
pronounced that the penis is replaced by a small, almost shapeless papilla.
(Figs. 42-47 show both normal and abnormal penis). In advanced stages of
reduction the vas deferens cannot be distinguished, although in a penis less than
half the normal size (fig. 44) it is still functional. Change of shape is also quite
common. On one occasion an abnormally large penis was developed, with a
highly sculptured border. The heavier the infection the greater the reduction
in size. In those cases in which some of the seminal vesicles are still producing
sperm the organ is generally of fair dimensions. One gravid female containing
many eggs was found to be lightly
infected with C. ubiquita Lebour
1907. A small penis had been
developed. This was the only
definitely hermaphrodite specimen
found.
It is highly improbable that
all the snails displaying small or
abnormal penis are “‘sex-reversed ”
females as Krull suggests. ‘Some
are almost certain to be males in
which the normal growth has been
inhibited, or in which the tissue
has atrophied. In fact it is quite
possible and indeed probable that
certain specimens devoid of a penis,
in which the gonads are also de-
stroyed, are not females but totally
castrated males. It is only when
the gonads are recognizable that
one can be certain of the sex of the
parasitized specimens.
Sh
mm
GENERAL REMARKS.
It will be seen from the
samples of P. ulvae described
above that the males were found to be more frequently infected than the females.
This is invariably so if the sex ratio of uninfected snails is taken into consideration.
Several causes may be suggested to account for this. The miracidia may exercise
sex selection in their free-swimming stage, or, having either penetrated from
without or been ingested within their egg capsule, they may only develop further
in a male host. Infected females, on the other hand, may have a much higher
rate of mortality than the males. On the whole, however, what little evidence
1 Krull considers that these infected specimens with a “‘ normal ” (= large) penis showing a
vas deferens are probably true males. He also notes these snails are not so large and the infections
less heavy. The specimens with a reduced and often functionless penis without a vas deferens
he suggests were originally female. It should be noted that in the author’s experience the unusually
small penis is not only associated with the heaviest infections, but also with the most marked
gigantism and shell variation.
94 NOVITATES ZOOLOGICAE XLI. 1938.
we have tends to support the view that a certain number of snails with a penis
are in reality sex-reversed females. Both Krull and the author have found infected
females which have developed a penis. Moreover the altered condition of the
penis of all infected snails emphasizes the profound effect these parasites exert
on the sexual organs.
It will be noticed that the ratio of infected males and females differs con-
siderably in the different samples. Several explanations may be offered to account
for this, but here again no proof is available.
P. ulvae, particularly in the saltings, is exposed to infection very sporadically,
for most of its trematode parasites are found in the sexually mature stage in birds.
This is well illustrated by the populations of adjacent pools in the saltings, which
will often harbour a completely different trematode fauna. This fact, together
with the irregularity of the spat fall so characteristic of this habitat, suggests
that the age at which the snails of any given pool become infected is also likely to
vary. It is quite possible that changes in the sex of the host are only brought
about by infections which enter the snail at an early age and are of long standing.
Another factor which may be of importance in this respect is the effect of
the snail’s environment upon the parasites. If conditions are somewhat un-
favourable the parasites are few in number, small in size, and the output of
cercariae is greatly reduced. Often the infections are moribund. The parasites
react surprisingly quickly to loss of vigour or sluggishness on the part of the host,
and, for example, in the laboratory (Pl. II, figs. 3 and 4) fare much worse than
the snails themselves. In habitats where growth conditions are poor it is possible
that a number of infections do not develop sufficiently to produce sex-reversal.
It is also possible that the tendency to grow a penis is more pronounced in female
snails living under maximum growth conditions.
Whatever is ultimately proved to be the cause of the excess of parasitized
males over females, it seems certain that this phenomenon in conjunction with
the gigantism produced by infections is capable of disturbing or reversing the
normal differences in size displayed by the sexes. It is therefore necessary to
employ great caution when studying this aspect of sexual dimorphism in P. ulvae,
and probably the same applies to any heavily infected species of snail.
TABLE II.
PERCENTAGE OF THE DIFFERENT GROUPS! OF CERCARIAE FOUND IN INFECTED P. ULVAE
FROM THREE TyPES OF HABITAT.
Infected | : B [a i | Specimens from
2 fe | Specimens from | Specimens from z
, ; . specimens arante les Hunterdon and
Groups of Cercariae examined mee | & | Fairlie Sands
Number | per cent.) Number per cent.| Number | per cent.) Number | per cent.
Pleurolophocerca 0 246 29-6 20 8-2 | 167 56-4 59 20:2
Echinostome . . 197 23-8 11 4:5 15 5:0 171 58:5
Ubiquita . 0 0 154 18:6 | 58 24:0 16 22-6 30 10-2
Oocysta . 5 ® 127 15:3 | 115 47-7 9 3:0 3 1:0
Notocotylid . é 26 31 | 6 2-9 6 2-0 14 4:7
Metentera 3 : 10 12 | 4 1:6 os = | 6 2-0
Sagittarius ; 3 04 | 2 0:35) | — | 1 0:3
Undescribed or un- | | |
identified . a | 66 | 79 | 25 10-3 33 11-1 8 2-7
Total . | 829 — | 241 = 296 = 292 =
1 The term “ group ” indicates a collection of related cercariae, but whereas one group may
include an entire superfamily (Pleurolophocerca), another may contain only a few related genera
(Oocysta or Ubiquita),
NovITATES ZooLoGIcAE XLI. 1938. 95
The numerous groups, and even species of cercariae, probably exert both
a different and varying degree of influence on the host. In the laboratory, for
example, the Echinostome infections are by far the most lethal. The impression
is also gained that the penis is altered to a greater extent in these infections than
in any other, and that sex-reversal occurs most frequently in snails parasitized
by Echinostomes. Further investigations are being carried out on this aspect
of the problem.
The relative degree of parasitism by the various groups of cercariae is given
in the Appendix and Table II. It is clearly shown that the Oocysta group is
commonest on mud flats, and is in fact comparatively rarely found in P. ulvae
from saltings, despite the higher percentage of infection met with in this
habitat. The adult worm of Cercaria oocysta is nevertheless a bird parasite
and develops in the Black-headed Gull into an undescribed species of Maritrema
Nicoll 1907 (Rothschild 1936). The Ubiquita group is found commonly in both
habitats, but relatively more frequently on the mud flats. The heaviest infec-
tions of Echinostome larvae are met with in the samples from Hunterdon and
Fairlie Sands, where they have always proved the most numerous group, while
the Pleurolophocerca cercariae which develop into Heterophyid flukes are found
in highest numbers in the saltings. The rarer species, such as the Notocotylid
and Metentera groups of cercariae, are sparsely but more evenly distributed.
It is perhaps worth noting that in P. ulvae, as in many other species of
molluses, the larger the individual snail, the greater the number of cercariae
produced by an infection. Apart from the environmental changes already
referred to, it would appear that the amount of cercariae produced is directly
controlled by the amount of food available. The gigantism of the host, which
involves an increase in the soft parts of the body as well as the shell, is thus
of great advantage to the parasite. The faculty of producing this increase in
size is a character which is presumably most susceptible to selection and its
widespread occurrence among the Trematoda is therefore not surprising.
SUMMARY.
(1) The normal sex ratio of P. ulvae varies from sample to sample irrespective
of age and size.
(2) The growth of P. ulvae is also very varied and the size of the shell is
no criterion of the age of the snail. The variation in growth, shell texture, shell
shape and shell colour, and the consistency of the soft parts of the body, are
probably closely linked with environmental factors.
(3) Female specimens attain a larger size than do males, and usually out-
number them in the largest size groups of any given sample.
(4) P. ulvae conforms to Pelseneer’s rule that more male than female snails
are infected with larval Trematodes. In some samples the ratio of infected males
to females is as high as 16: 1.
(5) The higher rate of infection among males, together with the gigantism
produced by the parasites, can combine to obscure or even reverse the normal
difference in size generally displayed by the sexes.
(6) The penis of all infected P. ulvae is abnormal, and usually more or less
reduced in size. It is possible that some specimens without a penis are totally
castrated males. An infected gravid female, with a small penis, was discovered.
96 NOVITATES ZOOLOGICAE XLI. 1938.
(7) There is some evidence to support Krull’s suggestion that infected
females of P. ulvae grow a penis. This would explain satisfactorily the excess
of parasitized males over females. It is, however, highly improbable that all
infected snails with reduced penis are sex-reversed females.
(8) There are indications that the metabolism of the snail is more seriously
disturbed by some species of cercariae than by others.
(9) The parasites are extremely sensitive to the condition of the host.
Under favourable growth conditions the parthenitae increase in size, number,
and fertility. The larger the snail the greater the number of cercariae emitted.
It is therefore of considerable advantage to the parasites to produce gigantism
in the host.
ACKNOWLEDGMENTS.
I would like to express my gratitude to Miss Anne Rothschild for her
assistance with the measurement and collection of samples of snails and to
Mr. G. M. Spooner for his invaluable help and advice.
APPENDIX.
Hunterdon Sands (Sample A), September 1935.
Hard grey type shell. Spires well preserved.
Range. Mean.
Total, 101 specimens . : 2 : . 9 -4 mm. 5-97 mm.
89 infected specimens . : : : . 9-44, 6-16 ,,
12 uninfected specimens i 2 Ä . 534 ,, 4:50 ,,
Group. Range. Mean.
Echinostome . 3 . 48 9 -43 mm. 6-37 mm.
Pleurolophocerca 5 . 20 TiO. 6-05 ,,
Ubiquita . : : . iz Gi 4.83 ©
Notocotylid : ; HB ER 5-57 „
Undescribed . ö a GB Br.
Hunterdon Sands (Sample B), March 1936.
Hard grey type shell. Spires well preserved.
Range. Mean.
Total, 131 specimens 83-41 mm. 6-27 mm.
119 infected specimens BA 6-40 ,,
12 uninfected specimens 6-44 ,, 4-92 ,,
Group. Range. Mean.
Echinostome . 5 . 66 83-4} mm. 6:77 mm.
Pleurolophocerca : . 35 N 6-17 ,,
Ubiquita . 9 53-43 5-22 ,,
Notocotylid 4
Metentera 2 TR 6:56 „
Oocysta 4
Unidentified 1
Novirates ZooLoGIcAE XLI. 1938. 97
Hunterdon Sands (Sample C), May 1937 (see p. 88).
Range. Mean.
Total, 241 specimens 71-2} mm. 4-57 mm.
84 infected specimens . 71-3} ,, 5.63 „
157 uninfected specimens . : ; . 5423 ,, 4:00 ,,
Group. Range. Mean.
Echinostome . : . 57 (5434,19, 4
unopened) 71-43 mm. 5:98 mm.
Ubiquita . 9 (593g, 4 un-
opened) 64-4 ,, 4:89 ,,
Pleurolophocerca 4 (not opened) 64-41 ,, 6-00 ,,
Metentera 4 (14,3 unopened) 53-4 _,, 4-75 ,,
Notocotylid 3 (333) Dee An
Oocysta 1 |
Sagittarius 1 7 (33d, 329, 1
Undescribed : mate amopenadl) 51-33 4.57
Unidentified sporocysts and Be ch
rediae . ! : Po ee
Holbeach Saltings (Sample A), August 1937 (see p. 89).
Range. Mean.
Total, 288 specimens . : : ; . 84-22 mm. 5-42 mm.
148 infected specimens : ; : . 814 ,, 5.63 ,,
145 uninfected specimens. | 5 . 62-23 „ 5-20 „,
Group. Mean.
Pleurolophocerca : . 90 (2288, 1999, 49 not opened for
sex determination) 5-66 mm.
Ubiquita . : : . 18 (639g, 822, 4 not opened) 5:79 ,,
Oocysta i : . 6 (896; 329) 6:00 ,,
Notocotylid . : 2 — =
Echinostome . i RS — —
Not identified ! : . 29 (1494, 1399, 2 unopened) 5-28 ,,
Holbeach Saltings (Sample B), August 1937. Hard grey type shell.
A hand-picked selection of 12 of the largest snails from a very small sample
from a permanent pool in the middle zone of the saltings. One snail
dead. One female (52 mm.) not infected. Range of infected speci-
mens 63-5 mm. Mean: 5-82 mm.
Group. Range. Mean.
Pleurolophocerca : 6 63-54 mm. 5-92 mm.
Ubiquita . : ; 8 645 , 5:58 „,
Echinostome . : a 6 as
1 A number of these snails were kept under cold conditions in the laboratory for several months
before dissection. Consequently numbers of these infections showed only rediae containing germ
balls, but no cercariae. Many were of the Pleurolophocerca type.
98 Noyirates ZooLoarcar XLI. 1938.
Holbeach Saltings (Sample C), August 1937 (see p. 90).
Total, 800 specimens. 25 of the largest selected.
Range. Mean.
15 infected specimens . 5 : ‘ . 54-44 mm. 4-73 mm.
10 uninfected specimens . ! 5 » 8 BS, 4-38 ,,
Group. Range. Mean.
Pleurolophocerca é é 8 (334, 529) 5-4, 4-50 mm.
Echinostome . : 2 & (2866; 2e9, I
unopened) 5i-44 ,, 5:00 _,,
Ubiquita . 5 : lle(D) 51 oe a
Oocysta 5 3 le) om 5 gen
Holbeach Saltings (Sample D), August 1937.
Red medium-hard shells. Spires blunted. A selection of 15 of the largest ı
snails hand picked from a permanent pool in the middle zone of the
saltings. Allinfected. Range: 7-54 mm. Mean: 6-38 mm.
Group. Range. Mean.
Ubiquita 7 63-54 mm. 6-25 mm.
Pleurolophocerca 6 7 Bh 5, 6-50 ,,
Echinostome 2 63-61 „, ER
Holbeach Saltings (Sample F), August 1937.
Very hard grey type. Spires perfectly preserved. A selection of 12 of the
largest snails hand picked from a dried-up pool in the lower zone of
the saltings, around which sweeps were made and Sample E (see p. 90)
collected. All infected. Range: 8-53 mm. Mean: 7-19 mm.
Group. Range. Mean.
Ubiquita . : : eS 8 -63 mm. 7-41 mm.
Echinostome . : 5 8 74-64 ,, TE SER
Pleurolophocerca : See 53 5 =
Tamar Saltings (Sample Al), March 1938 (see p. 91).
Total, 1,946 specimens. 33 of the largest snails selected ; all infected.
Range: 104-7 mm. Mean: 8-57 mm.
Group. Range. Mean.
Pleurolophocerca 21 (1788, 499) 104-7 mm. 8-62 mm.
Ubiquita . 6 (394, 329) 9-71 ,, 8-50 ,,
Echinostome 2 (236) BE Pr:
Notocotylid 2 (18,19) 9 e en
Undescribed 2 (288) yp =
Tamar Saltings (Sample A), March 1938 (see p. 91).
Range. Mean.
Total, 180 specimens . 5 : 6 . 9 -22 mm. 5:47 mm.
56 infected specimens . \ 4 5 9-43 7-04 ,,
124 uninfected specimens. ; : N 4-77 ,,
1 The large specimens were lying on the mud in the middle of the pool. Around the edges
which were not under water were hundreds of thousands of very small specimens. A scoop from the
middle and the edge gave completely different size groups.
Novirarres ZOOLOGICAE XLI. 1938.
Group. Range.
Pleurolophocerca ; . 35 (2784, 329) 83-51 mm.
Ubiquita . ; : . 17 (1434, 399) 9 -54 ,,
Ooeysta 1
Notocotylid 1+(493) Waals
Undescribed 2
Tamar Saltings (Sample B), October 1937.
Hard red type shell. Spires blunted.
Total, 1,000 specimens, 16 of the largest selected.
| Range.
7 infected specimens 8 -64 mm.
9 uninfected specimens 64-6 ,,
Group. Range.
Ubiquita . : ! = 16 8 -64 mm.
Oocysta aD el! 74 >
St. John’s Lake (Sample A), June 1937 (see p. 92).
Total, 10,000 specimens. 46 of the largest selected.
Range.
18 infected specimens . 51-41 mm.
28 uninfected specimens 51-41 ,,
Group. Range.
Ooeysta 11 (799, 346;
unopened) 51-41 mm.
Ubiquita . 2 (299) 43 ss
Pleurolophocerca 2 (292) 515,
Echinostome 1 (13) 5 A
Undescribed 2 (299) 5-4 „,
St. John’s Lake (Sample B), December 1933.
Red hard shell type.
99
Mean.
7:23 mm.
6-88 „,
6:00 _,,
Mean.
7:71 mm.
6-33 ‚,
Mean.
7:83 mm.
Mean.
4-89 mm.
4-88 ,,
Mean.
4-82 mm.
Several thousand shells selected, of which 182 of the
largest were taken and roughly divided into two groups, one containing
the largest specimens.
lst Group. 2nd Group.
100 largest shells. 42 infections. 82 of the smaller shells.
Group. Group.
Oocysta . ; ; 6 25 Ubiquita
Ubiquita . 7 Oocysta
Pleurolophocerca 5 Pleurolophocerca
Notocotylid 2 Notocotylid
Metentera : 2 Metentera .
Too young to identify 1 Echinostome
St. John’s Lake (Sample C), December 1934.
Hard type grey shells. A random sub-sample Range.
of 206 shells over 2$ mm. 54-3 mm.
18 infected specimens . BE
23 infections.
I
Se Oe DS
Mean.
4-27 mm.
4-42 ,,
100
Group. Range.
Oocysta 8 5 -31 mm.
Ubiquita . 5 DE Ar en
Pleurolophocerca 3 Aa
Echinostome 1 44 5
Unknown. 1 44
„>
St. John’s Lake (Sample D), January 1935.
Hard type grey shells.
infected specimens.
Group.
Oocysta 10
Un. 4 No measurements taken.
Pleurolophocerca 1
Notocotylid 1
St. Andrews, Esturine Mud (East Scotland), 1935.
NOVITATES ZOOLOGICAE XLI.
35 of the largest shells of 51-5 mm. selected.
1938.
16
Very soft grey type shell. Several thousand shells, of which 83 of the
largest selected.
Range. Mean.
24 infected specimens . 41-31 mm. 4-07 mm.
61 uninfected specimens Ei pp 3:70 ,,
Group. Range Mean.
Oocysta 12 41-32 mm 4-17 mm.
Ubiquita . 5 4-31 ,, 3:90 ,,
Echinostome 3 41-32 ,, 4:00 „,
Notocotylid 5 1 4 ie =
Too young for identification 3 = =
St. John’s Lake (Sample E), July 1935.
Hard grey type shell. About 10,000.
A certain number of each size group were dissected.
Total 1,385. Range: 54-1 mm. Mean: 3-09 mm.
100 infected specimens. Range: 54-1 mm. Mean: 3-85 mm.
| ; | Too
0 . Pleuro- :
: - Number Total au Ubi- | Echino- Sagitta-| Noto- | Unde- | young
Size Group in | „yam- | infec- © Se quita | stome ee rius |cotylid |scribed | for
um: ined | tions zeup Group | Group Crom Group | Group | Groups| identi-
| P fication
|
5mm. . 47 19 10 6 = = = 1 1 1
4mm. . 574 59 27 17 3 2 2 = 2 6
3mm. . 335 12 Gy 2 — 1 — = — 4
2 mm. . 321 8 1 3 1 3
lmm. . 108 2 D
Total 1,385 100 43 28 3 3 2 1 4 16
Range in mm. | 53-1 54-1 54-2 54-2 4 4-3 4 5 5% 5-1
Mean length in
mm. . . | 3:09 | 3:85 | 4:07 | 3-93 4 3:67 4 5 3:75 | 3-06
Novirarrs Zoonoarcar XLI. 1938. 101
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Krull, H., 1935. Anatomische Untersuchungen an einheimischen Proso-
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biol. Ass., Plymouth, 21, No. 2, 1937, pp. 721-742, pl. III, 10 text-figs.
Pelseneer, P., 1906. Trématodes Parasites de Mollusques Marins.—In Bull.
Scientifique, France and Belgique, 40, 1906, pp. 161-186, pls. VIII-XL.
Pelseneer, P., 1926. La proportion relative des Sexes chez les Animaux et
particuliérement chez les Mollusques —In Acad. Roy. Belgique, Mémoires,
8, No. 11, 1926, pp. 1-258.
Pelseneer, P., 1928. Les Parasites des Mollusques et les Mollusques Parasites.—
Bull. Soc. Zool., France, 58, 1928, pp. 158-189.
Rothschild, M., 1935. The Trematode Parasites of Turritella communis Lmk.
from Plymouth and Naples.—In Parasitology, Cambridge, 27, No. 2, 1935,
pp. 152-170, 22 text-figs.
Rothschild, M., 1936. Gigantism and Variation in Peringia ulvae Pennant 1777,
caused by Infection with Larval Trematodes.—In J. Mar. biol. Ass.,
Plymouth, 20, No. 3, 1936, pp. 537-546, 2 text-figs.
Sewell, R. B. Seymour, 1922. Cercariae indicae.—In Ind. J. med. Res., 10,
Suppl. number, 1922, pp. 1-370, pls. I-X XXII, 6 text-figs.
Wesenberg-Lund, C., 1931. Contributions to the development of the Trematoda
digenea. Part I. The Biology of Leucochloridium paradoxum.—In Mem.
Acad. R. Sci. Lett. Danemark, 4, No. 3, 1931, pp. 89-142, 6 pls., 7 text-figs.
EXPLANATION OF PLATES AND TEXT-FIGURES.
Plate I. Peringia ulvae from various habitats (x 2).
Row A. Some of the largest specimens, black, soft-shelled type about
four years old, from a collection bred in the laboratory. The eggs were
obtained from a typical red hard-shelled sample of snails from the
mud flats, St. John’s Lake.
Rows B and C. The largest snails from two collections of several thousand
snails from the mud flats, St. John’s Lake.
Row D. Some of the largest snails from Hunterdon Sands (Sample C).
Row E. Ditto, Burnham Saltings (Sample A).
Row F. Ditto, Holbeach Saltings (Sample E) (grass sweepings).
Row G. Ditto, Tamar Saltings (Sample A1).
Row H. Ditto, Poole Harbour.
Row I. Ditto, Estuarine Mud, St. Andrews (see Appendix).
Row J. Ditto, Holbeach Saltings (Sample A).
Row K. Ditto, Tamar Saltings (Sample A).
Row L. Ditto, Holbeach Saltings (Sample C).
102 NovirarEs ZooLocicaE XLI. 1938.
Plate IT.
Fig. 1. A portion of sample A from the Tamar Saltings (x 2).
Fig. 2. A portion of sample A from St. John’s Lake (x 2). Note the
contrast in size between the specimens of the two samples.
Fig. 3. Longitudinal section of very heavily infected P. ulvae (x 18) killed
after it had been kept some time in the laboratory, showing the reduction
in number, size, and fertility of the Echinostome rediae.
Fig. 4. Section of infected P. ulvae (x 18) killed soon after it had been
collected, showing the vigorous condition of the Echinostome rediae.
Text-fig. 41. Irregularity of spat fall in samples of P. ulvae.
Text-fig. 42. Normal penis of a specimen of P. ulvae measuring 6 mm. in length.
Text-fig. 43. Somewhat reduced penis of a specimen measuring 63 mm. in
length, infected with Cercaria oocysta.
Text-fig. 44. Reduced penis of a specimen measuring 84 mm. in length, infected
with the cercaria of Cryptocotyle jejuna Nicoll 1907.
Text-fig. 45. Greatly reduced penis of a specimen measuring 8 mm. in length,
infected with an Echinostome cercaria.
Text-fig. 46. Papilla-like penis of a specimen measuring 81 mm. in length,
infected with the cercaria of C. jejuna.
Text-fig. 47. Papilla at the terminal portion of a normal penis of P. ulvae.
This is lost in most reduced examples.
Novrranes ZooLoGIcAE NLI. 1938. 103
WHERE SUBSPECIES MEET.
By KARL JORDAN.
(With text-figures 48-53.)
IAS a paper read at the 6th International Congress of Entomology which met at
Madrid, September 6th to 12th, 1935, I dealt at some length with this subject.
In March 1937 the General Secretary of the Congress informed me that 400 pages
of the Transactions had been printed before war broke out and that the other
manuscripts were set up in type. No progress has been made since. The General
Secretary being involved in the clash of factions, it is most unlikely that the
Proceedings and Transactions of the 6th Congress will be published before the
end of the struggle in Spain. As my investigations of 1936 into the subject
were a continuation of those recorded in the Congress paper, I repeat here some
of the earlier observations before referring to my later results.
Innumerable species live side by side. However close they may agree in
their morphology, there is a barrier between them which prevents interspecific
promiscuity, keeping the specific populations pure. In a large percentage of
species the area of distribution is subdivided into minor areas, each subarea
being inhabited by a subspecific population. In those groups of animals where
the systematist has advanced to the study of subspecies of the dead material at
his disposal in museums, the diagnoses of subspecies are based on morphological
distinctions, which are of the same kind as in the case of species. The two
systematic categories being. in this respect essentially alike, differing only in
degree, the question arises as to whether the aversion to intercrossing which keeps
the species apart also exists among the subspecies of one and the same species
wherever they come into contact with one another. Where this aversion does
not exist, the subspecies, so closely related to each other morphologically, must
be expected to hybridize and to form hybrid populations in districts of contact.
And conversely, the absence of hybrid populations may be taken as evidence
that the subspecies in question are kept apart by a physiological barrier as are
the species. Such conclusions may, in a given instance, be correct or not, being
based on the evidence of circumstances which are not known in their entirety.
Field researches and experimental breeding will have to supplement the investi-
gations of the museum’s systematist in order to make the results conclusive.
A most suitable subject for this kind of research in Europe is the widely
distributed mouse-flea, Ctenophthalmus agyrtes Heller 1896, which is common
on Apodemus sylvaticus and Evotomys glareolus and has developed into a number
of subspecies. The species offers the student the great advantage that the differ-
ences between the subspecies are structural and definite, at least in the male.
Moreover, a survey with the object of ascertaining the exact distribution of each
subspecies of this flea would also be a survey of the fauna of mice, extensive
collecting thus obtaining results in two classes of animals. I have collected
most of the subspecies of Ctenophthalmus agyrtes ; but it was only in 1931 that
I for the first time devoted my holiday to obtaining an answer to the question :
where do subspecies of Ct. agyrtes come into contact with one another? I
concentrated on trapping mice; collecting other hosts and other fleas was
104 NOVITATES ZOOLOGICAE XLI. 1938.
incidental. The fleas were studied at the hotel, so as to be certain of the progress
of the investigation. The distribution of the three subspecies concerned was
known at that time to be as follows :
(1) The central European Ct. agyrtes agyrtes Heller 1896. Norway,
Denmark, Germany, southward to the Lake of Geneva and westward to Seine
Inférieure.
(2) The north-western Ct. agyrtes nobilis Roths. 1898 (= Ct. a. celticus
J. & R. 1922). Great Britain and Ireland, Brittany and southward to the
Puy-de-Döme district and Charente.
(3) The French alpine Ct. agyrtes provincialis Roths. 1910.
Hautes Alpes, Zermatt and Bex-les-Bains in the Swiss Rhone valley.
My first investigation aimed at the discovery of the districts where Ct. a.
_ agyrtes and Ct. a. provincialis meet. As I had previously collected agyrtes above
Montreux and provincialis at Bex-les-Bains, I expected to find both subspecies
somewhere in between the two places. I went in the spring, which proved to
be a mistake ; mice were very rare from Villeneuve, at the upper end of the
Lake of Geneva, to Aigle, while slugs and snails abounded and sprung the traps.
At Villeneuve I found only Ct. a. agyrtes ; but at Aigle, a little east of the town,
on the left side of the river coming down from Les Ormondes, I obtained Ct. a.
provincialis; and much higher up the right bank of the Rhone valley towards
Leysin, the only specimen caught was a ¢ intermediate between the two and
suggesting that the plateau north of the main chain of the Bernese Oberland is
inhabited by a special intermediate subspecies or by a hybrid race. We have
no fleas from these higher districts of Vaud and Valais; on the north side of
the mountains Ct. a. agyrtes is found. Poor as the results of several weeks’
trapping were, they proved that the two subspecies meet round about Aigle.
A renewed comparison of the fairly extensive material from Bex with the speci-
mens from Zermatt and the French Alps has shown us that some of the Bex
males are less typical, inclining a little towards Ct. a. agyrtes, which agrees with
the suggestion that the two subspecies hybridize.
On the south side of the lake I collected at La Roche-sur-Foron, where I
found only Ct. a. agyrtes, while in the Chamounix district only Ct. a provincialis
was obtained, in evidently typical specimens. The two fleas probably come
into contact on the escarpment from the lower to the higher level, and good
centres for this investigation would be Servoz in the Arne valley and St. Germain
in the Montjoie valley. On the south side of the Col de Bonhomme Ct. a.
provincialis may be expected to occur at higher altitudes (as it does on the
north side), and Ct. a. verbanus J. & R. 1920 farther down. Courmayeur would
appear to be a convenient place for trapping up and down the Dora Baltea in
search of the two subspecies.
As stated above (p. 104), two subspecies were known from Northern France,
and it was evident from the occurrence of the one in Seine Inférieure and the
other in Brittany that their areas of distribution must meet somewhere in
Normandy, and Bagnoles-de-l’Orne, nearly in the centre of Normandy, seemed
indicated as a good starting-point for the investigation. ‘Trapping behind the
bathing establishment on a wooded knoll covered with boulders produced Ct. a.
agyrtes ; but outside the wood, in the hedges and meadows, I found Ct. a. nobilis.
The results remained constant during the fortnight’s stay in the place; only
agyrtes in the wood and only nobilis in the open country : a few hundred yards
Valescure,
Novirares ZooLoGIcAE XLI. 1938. 105
from the wood to the nearest trap in a hedge. The intermediate ground was
occupied by houses and gardens where I could not put up any traps. The
restriction of the two subspecies each to a definite biotope at Bagnoles is interest-
ing and instructive. If the specimens had not been studied on the spot, but
been labelled in the usual way with locality, date and host, and then been ex-
amined at home, the fact that the two insects are separate in space at Bagnoles
would have been concealed. The areas of distribution do not overlap, but
dovetail. Whether that is so in other districts of Normandy remains to be
ascertained. There is no indication in the morphology of the Bagnoles specimens
that intercrossing occurs.
Ct. a. nobilis was found by Mr. J. F. Cox at Ruffec, Charente, and in the
Puy-de-Döme district in 1931, and the Hon. Miriam Rothschild obtained it at
Royat-les-Bains. In these districts it occurs together with another flea, Cten-
ophthalmus arvernus Jord. 1931, which is nearly related to Ct. agyrtes, but evidently
specifically distinct. Miss Rothschild found the two insects actually on the
same host at the same spot. There is no trace of hybridization in the fairly long
series obtained.
As we had no collections from farther south and west in France, we did not
know what happens to Ct. agyrtes south of the Auvergne. Does Ct. a. nobilis
occur there or is it replaced by another subspecies ? To provide an answer to
this question I visited South-west France in 1936, selecting four widely separate _
places at which to spend some time: Figeac (Lot), Brassac (Tarn), St. Jean-
Pied-de-Port (Basses Pyrénées) and Gavarnie (Hautes Pyrénées).
As I had found in North Spain, in 1935, a subspecies of Ctenophthalmus
agyrtes which is in some details nearer to Ct. a. agyrtes than is Ct. a. nobilis, I
expected to discover in one or the other of these French localities a race like the
Spanish one or intermediate between it and Ct. a. nobilis, and was much sur-
prised that I obtained no subspecies of Ct. agyrtes at all, but only Ct. arvernus
in all four places. I have certainly not covered the whole country, and there
exist possibly pockets between the Auvergne and the Pyrenees where Ct. a.
nobilis may occur ; but so much is certain that the ordinary flea of Apodemus
and Zvotomys in South-west France is Ct. arvernus, which takes here the same
place in the fauna as Ct. a. nobilis in Brittany and Great Britain and Ct. a.
agyrtes in Central Europe. A connection between the area of Ct. a. nobilis and
that of the Spanish subspecies may be discovered to exist along the coast of the
Bay of Biscay ; or the pine woods of Les Landes may harbour a race of their
own. We do not know. Our knowledge of the distribution in Europe, not only
of fleas but of large numbers of other animals, is so meagre and it would be in
many instances so easy to fill in the lacunae that I here appeal to local naturalists
to take an interest in this kind of research. I suppose it is too much to hope
for a well-organized survey of the fauna of Western and Central Europe.
I append here the list of Siphonaptera collected in South-western France,
to which is added the description of the Spanish race of Ctenophthalmus agyrtes
above referred to, and that of another flea from Sardinia.
Collecting was concentrated on Apodemus sylvaticus, Evotomys glareolus
and Talpa europaea. As the Spanish race occurs on the Mole, there was a
possibility that the same or a similar race of Ct. agyrtes was parasitic on Talpa
in the Pyrenees ; the result was negative. Shrews were taken incidentally, but
there were hardly any fleas on them, which was contrary to the usual experience.
106 NOVITATES ZooLoGIcAE XLI. 1938.
No attempt was made to obtain a large and varied collection; nevertheless,
several subspecies and one species are new to science. Some of the mice collected
are still rare in museums. The specimens from Figeac, Brassac and St. Jean-
Pied-de-Port were collected in June and those from Gavarnie and neighbourhood
in July.
1. Citellophilus occidentis sp. nov. (text-fig. 48).
Gave d’Ossoue, Gavarnie, 1,460 m., 13.vii.1936, on Microtus nivalis aquita-
nius, one 9.——The mouse was caught in a place where Hliomys quercinus
occurred ; this is possibly the true host of the new flea. The only example of
Eliomys caught was dead in the trap, and there were no fleas on it.
9. Somewhat aberrant in this genus. C. danubianus Roths. 1909 is geo-
graphically the nearest species we have in the collection. The new species
differs from it particularly in having 3 antepygidial bristles, no row of bristles
on the outer surface of mid- and hindfemora, a small stigma on tergite VIII,
and a long body and short tail to the spermatheca. In Wagner’s Catalogue of
the Palaearctic fleas C. danubianus is placed as a synonym of C. simplex Wagn.
1902 ; but this can hardly be right, if Wagner’s figures are absolutely correct as
he affirms. The digitoid of C. danubianus bears a spiniform bristle not shown
Novirates Zoonocroan XLI. 1938. 107
in Wagner’s figures and the other bristles of the digitoid are much longer in his
figures than in our specimens of ©. danubianus from Rumania and Bulgaria.
Rostrum extending to middle of trochanter, shorter than in ©. danubianus.
In front of eye a row of 3 long bristles, no other bristles on frons ; on occiput a
long bristle above middle of antennal groove, subapical row widely interrupted
above the long bristle placed at the lower angle, above the wide interspace only
2 or 3 bristles in the row each side. Eye with hardly any pigment except at
periphery (accidental ?). Bristles of segment IT of antenna much longer than club.
Pronotal comb with 19 spines inclusive of a small ventral one. On mesono-
tum a posterior row of 10 long bristles (the two sides together) and a row of 11
small ones, no further lateral bristles except at anterior margin, and a row of
3 or 4 dorsal ones a short distance behind the basal ones, on inner surface 11
bristle-like spines ; on mesopleura 7 long and longish bristles and anteriorly 10
or 12 small ones. Bristles on metanotum 15, 10; on metepimerum 6.
Apical spines on abdominal terga (the two sides together) : I 3, II 4, III 3;
bristles in two rows, no additional dorsal bristles : I 13, 10, II 10, 15, III 9, 14,
IV 7, 14, V 7, 14, VI 8, 18, VII 4, 12; onsterna: II 0, 2, III 0, 7, IV, 0, 8, VI 0,
8, VIL 12, 10.
Apart from the subapical subventral bristle, on outer surface of forefemur
8 lateral bristles (and 2 near apex), on inner surface 2 lateral ones ; on midfemur
on outside no lateral bristles, on inside a row of 5; on outside of hindfemur no
lateral bristles, on inside a row of 7 on one femur and 9 on the other. On hind-
tibia 7 dorsal notches (inclusive of apical notch), third notch with a single heavy
bristle, between fifth and sixth a single subdorsal heavy bristle ; on outer surface
of hindtibia 9 dorsolateral bristles, including an apical one; the longest apical
bristle of segment II of hindtarsus reaching to apex of III; in all tarsi of
C. danubianus segment V with 2 apical ventral bristles and proximally of them
1 or 2 similar but smaller bristles, these latter absent in the new species.
Three antepygidial bristles, upper one 3 of middle bristle, lower one 3,
approximately.
Modified Segments.—Sternum VII narrowing apicad, the upper margin
slanting-incurved (text-fig. 48, VII st.), apex broad, rounded-truncate, upper
apical angle a little over 90°. Above stigma of VIII t. 3 bristles each side ;
stigma much smaller than in C. danubianus, below stigma 2 short and 3 long
bristles, on lower portion of outer surface 17 bristles, inclusive of those at apical
margin, on inner surface 2. Sensilium measured dorsally shorter than its distance
from stylet. Some of the bristles of anal sternum (X. st.) stout and curved.
Spermatheca (R.s.) : body twice as long as broad, almost of even width, ventrally
concave, dorsally convex, orifice terminal-subdorsal, tail measured in a straight
line a very little shorter than body ; bursa copulatrix with conspicuous glandular
tissue, as in C. danubianus, blind duct long.
Length 3-0 mm. ; hindfemur 0-48 mm.
2. Nosopsyllus fasciatus Bose 1801.
Brassac, on Apodemus sylvaticus, 1 &, and on Mus musculus, 1 6.
3. Ctenophthalmus arvernus Jord. 1931 (text-fig. 49).
Figeac, a series on A. sylvaticus and Evotomys glareolus. The best
collecting ground at Figeac was a steep damp lane above the railway, with rank
9
108 NOVITATES ZOOLOGICAE XLI. 1938.
vegetation and a hedge each side. Here the two mice and Crocidura russula
were rather abundant. On the dry higher ground of each side of the valley
hardly any mice were found; but I obtained on the right side well above the
town a specimen of Pitymys pyrenaicus, the second record of the species from
Central France.
Brassac, a series on A. sylvaticus and E. glareolus, and 1 3 on Mus musculus.
St. Jean-Pied-de-Port, on A. sylvaticus, a small series. Mice were very
scarce ; the hills too dry and the valley too populated. No Evotomys were
found.
Gavarnie, on A. sylvaticus, 2 3,
1 °.—Only one specimen of this mouse
was trapped and no Hvotomys.
Gave d’Ossoue, above Gavarnie, on
Pitymys pyrenaicus, 4 II, 2 29. One
specimen of this mouse taken in a
mole run.
The marginal projection of VII. st. of
2 is always short (text-fig. 42, the segment
spread out), in Ct. a. nobilis twice as long
or over.
4. Ctenophthaimus nivalis. nivalis
Roths. 1909.
Gave d’Ossoue, above Gavarnie, on
Microtus mivalis aquitanius, 1 9. As
sternum VII of the specimen is injured,
I cannot say with certainty whether the
specimen agrees in this segment with Ct.
nivalis nivalis from the Hautes Alpes and
Haute Savoie. I obtained only two Snow
Mice; one, without fleas, was trapped
among the large boulders a short distance
beyond the first bridge over the brook,
the other at the other side of the water in
x
the wood near a meadow beyond the second bridge.
5. Ctenophthalmus agyrtes hispanicus subsp. nov. (text-fig. 50).
A single 3 from the nest of Talpa, near Lake Enol, above Covadonga,
Asturia, 3 Sept. 1935. Mole-hills were plentiful in a meadow, and here
and there the feet of cattle had sunk into the ground, a sure sign that there was
a nest underneath. I took several nests, but contrary to my expectation (and
usual experience) there were no flea-larvae in them and only one imago ; the
ground had been perhaps too wet.
3. Nearest to Ct. a. nobilis Roths. 1898, but differing in the genitalia,
particularly in the ventral lateral lobe of the phallosome. Manubrium (M) of
clasper strongly curved upwards, nearly as in Ct. baeticus Roths. 1910 from
Portugal. The ventral rounded corner of clasper more projecting downwards
than in Ct. a. nobilis and its lowest point farther distant from digitoid F. Process
P* of clasper obliquely truncate-sinuate as in some specimens of Ct. a. nobilis,
Novrrares Zoorocıcan XLI. 1938. 109
the posterior angle but little projecting ; in the majority of French specimens
of Ct. a. nobilis the apex of P? is more deeply incurved, the lower angle therefore
projecting considerably ; this is also the case to the same extent in a minority
of British specimens, an indication of the commencement of subspecific separa-
tion. Ventral, horizontal, arm of sternum IX as narrow as in Ct. a. nobilis,
but its ventral margin feebly angulate at some distance from tip at the point
where there is a cluster of bristles. Phallosome with large, rounded, ventral
flap each side (Lo), densely covered anteriorly and ventrally with semiovate,
somewhat scale-like, swellings directed frontad ; in Ct. a. nobilis, on the other
hand, the flap is turned upwards (occasionally turned down in mounted
specimens) and its upper margin very distinctly denticulate (text-fig. 51) ;
nobilis differs therein from all other subspecies of Ct. agyrtes, and from the
purely morphological point of view therefore has some claim to specific
distinctness.
110 NoviratEes ZooLoGICAE XLI. 1938.
6. Rhadinopsylla mesa Jord. 1920.
Gave d’Ossoue, Gavarnie, 1,460 m., 13.vii.36, 1 g on Talpa europaea.——
The specimen differs slightly from the only Swiss 3 (the type) we have; the
difference is either individual or geographical ; at least one more ¢ is necessary
for a tentative decision. The genal comb of the Pyrenean ¢ consists of 5 spines
on each side ; in the Swiss ¢ there are 5 spines on one side and 6 on the other ;
in our two Swiss 29 6 on each side. The uppermost spine of this comb is, in the
Pyrenean 4, very little broader at the base than the lowest spine and a trifle
longer, not shorter as in the Swiss 3.
In the Key to the species of Rhadinopsylla published in Nov. Zoou., xxxv,
pp. 183, 184 (1929) it is stated under a‘that the bristles of VIII. st. are apical
in Rh. isacanthus ; it should read subapical, the row of bristles being placed
some distance from the apical margin.
7. Typhloceras favosus rolandi subsp. nov. (text-fig. 52).
Basses Pyrénées: St. Jean-Pied-de-Port, 150 m., on Apodemus sylvaticus,
28.vi.36, one Q.
©. Agrees with our two Algerian 9? in the eye being smaller than in
T. poppei Wagn. 1903, thorax and abdomen more distinctly reticulated, bristles of
metanotum less numerous, many of the bristles of the body less pointed, the
interspace between eye and upper genal spine broader than third genal spine,
NOVITATES ZOOLOGICAE XLI. 1938. 111
hindtibia with about 25 lateral bristles (in 7. poppei over 40). The Pyrenean
specimen differs from the Algerian ones in the pronotal comb containing 22
spines instead of 20, and in the sinus of sternum VII not being deeper than in
T. poppet : triangular lobe above this sinus narrower than in 7’. poppet (narrower
on right side than on left figured) and not longer than lower lobe, which is broad
and rounded (text-fig. 52).
8. Typhloceras favosus asunicus subsp. nov. (text-fig. 53).
The above-described subspecies is connected with T. favosus favosus J. & R.
1914 by an intermediate one from Sardinia, of which we have also but one 9,
collected by Dr. A. H. Krausse at Asuni off a mouse (and mentioned in Nov.
ZooL., xxi, p. 236 (1914)). Pronotal comb with 22 spines as before. Sinus of
sternum VII narrow (text-fig. 53), the lobe above it decidedly longer than the
broad lower lobe and twice as long as in the preceding subspecies, but one-third
shorter than in 7’. f. favosus, in which the lower lobe, moreover, is nearly as long
as the upper. As VII. st. is injured on the left side, we figure the right side ;
the long bristles are the same on both sides.
112 ; Novirares ZoonocicaE XLI. 1938.
ON SOME SIPHONAPTERA FROM TROPICAL AFRICA AND IRAQ.
By KARL JORDAN.
(With text-figures 54-64.)
1. Xenopsylla bantorum sp. nov. (text-figs. 54, 56, 57).
A SERIES from Uganda: Tororo, on Aethomys, March 1938.
A most interesting discovery, for which we are indebted to G. H. E.
Hopkins, Agricultural Laboratories, at Kampala. The flea is X. cheopis Roths.
1903 with a difference. If the two were geographically separated units, I should
treat them as subspecies. They were subspecies before X. cheopis had overrun
the tropical world. Mr. Hopkins’s attention was drawn to the specimens not
by any apparent difference from X. cheopis in their make-up, but by the fact
that they occurred on field rodents and not on rats in the huts where one would
expect X. cheopis to be the rat-flea. These hut rats were infested with X.
brasiliensis Baker 1904 instead of X. cheopis, which occurs also in tropical
Africa. A close study of the morphology of the field-rat flea proved the males
to differ from X. cheopis $3. We find, therefore, in the same country two fleas
so closely related as if they were subspecies, and the hasty conclusion might be
drawn that the African cheopis population had been modified on the indigenous
field-rats into the new population, which I name X. bantorum : a species being
split into two species without isolation. ‘The present case is a very instructive
illustration refuting that assumption. The existence of two very closely related
species side by side cannot be explained satisfactorily without a knowledge of
the distribution of each and without reference to the geological history of the
fauna, a fauna being composed of elements which have arrived at different
Novıraras Zoonogican XLI. 1938. 113
periods. In the case of X. cheopis, the history of distribution does not go back
very far in time.
X. cheopis is essentially a flea of the domestic rats, Rattus rattus and R.
norwegicus, and its distribution depends and has depended on that of these hosts.
In recent times the rats have been incidentally brought to all parts of the world,
and where the conditions were favourable for the flea, it has established itself
and has spread inland with the ships’ rats and found temporary or even perma-
nent new hosts. But whence did X. cheopis, now cosmopolitan, originally come ?
Its true homeland was, of course, the country from which the rats went forth to
conquer the world: Iran and Iraq and adjacent countries. The rats are not
mentioned with certainty in Greek history—
the Greek language has no name for them ;
but may have invaded the Nile valley at
the time when armies crossed Syria from
Mesopotamia. The area of distribution of
the rats and therefore of X. cheopis com-
prised at that historical period South India,
Persia, Mesopotamia, Syria and the Nile
valley ; the indigenous rats of tropical East
Africa had their own flea, X. bantorum. If
we could switch our life back to that period,
the systematist would name the two fleas
X. cheopis cheopis and X. cheopis bantorum.
And the distribution of these fleas being now
what it is, the employment of trinomials in
this case, though they express the true
status of the two fleas, might be confusing,
and as nomenclature has been invented
majnly to simplify and clarify the survey of
the confusing mass of different creatures, it
appears to me advisable to treat the new flea as
specifically distinct. And thus arise species.
d. There are apparently no other
differences than those presented by the
genitalia, the sclerites in question being
represented by text-figs. 54 (X. bantorum)
and 55 (X. cheopis). Process P! of clasper is short and truncate in all specimens
of X. bantorum, the four sketches a, b, ¢ and d illustrating the variability of P!
in our series; in X. cheopis the process also varies, but is always longer and
much more triangular, the upper margin being strongly excurved ; the portion
above the bristles is not readily seen in X. cheopis, because the margin is very
transparent, the dorsal area being but feebly chitinized. The grouping of the
bristles on P! is also different, the group being arranged longitudinally in X.
cheopis and more obliquely and transversely in X. bantorum. A further difference
is found in the ventral arm of sternum IX ; in X. cheopis this segment broadens
towards apex, more or less so than in our fig. 55, whereas in X. bantorum it
widens but slightly and is more pointed and curved up. Fig. 55, IX. st., is taken
from the type of X. cheopis, fig. 55, P!, from a dissected paratype ; fig. 54a and
IX. st. from type of X. bantorum. Some of our few Kenya specimens from
114 Novirates ZoonocicaE XLI. 1938.
indigenous Muridae are X. cheopis, others are X. bantorum, and others again
are intermediate, indicating hybridization. If hybridization does take place,
the native X. bantorum will probably be absorbed by further invasions of
X. cheopis. Our material from Kenya is insufficient for the investigation of
this question.
2. No difference has been observed between the 99 of X. cheopis and X.
bantorum, as is frequently the case with subspecies, but also occurs with some
species of which the 3G are very different. The spermatheca varies in the
body being more rounded in some specimens than in others and in the base of
the tail sometimes not being strongly ventricose (text-figs. 56, 57).
2. Ctenophthalmus modicus elgonensis subsp. nov. (text-fig. 58).
Uganda: Bulambuli, Mt. Elgon, Lophuromys aquilus August 1934 (J. Ford
and G. L. R. Hancock), and Sipi, Mt. Elgon, on same host and one specimen on
Rattus rattus, September 1934 (G. H. E. Hopkins) ; 6 3d, 3 99.
3. Ventral process P? of clasper longer than in Ct. m. modicus Jord. 1933
from Kenya, its inner incrassation (touching the digitoid F) projecting beyond
the outer apical margin of P?, which is not the case in Ct. m. modicus (text-fig.
59) ; in some specimens P! and P? are closer together than in the type figured,
and the apical margin between the two processes is sometimes hardly at all
incurved ; the row of bristles dorsally more oblique than in Ct. m. modicus ;
the long bristle on the side of P? has a more ventral position than in most speci-
mens of Ct. m. modicus, as shown in the figures. The ridge (ri) of the digitoid F
against which the processes P! and P? play does not extend so near the ventral
margin of F as in Ct. m. modicus, and is dorsally farther away from the apical
margin of F than ventrally, whereas in Ct. m. modicus the ridge inclines more
or less ventrad.
9. The three specimens agree with what we took with some doubt to be
this sex of Ct. m. modicus ; cf. Nov. Zoou., xxxvili, p. 350, text-fig. 70 (1933).
In one example the spermatheca has the shape as figured in 1933, whereas in
the two others the body is much more swollen, which is also the case in one of
our examples of Ct. m. modicus from Nakuru ; this specimen was purposely not
mentioned in the description of 1933, as I was uncertain as to whether it really
was a Ct. modicus. The Mt. Elgon 99 varying in the same way may be taken
as conclusive evidence that all these 9? belong to Ct. modicus.
Novrrates Zoonoaican NLI. 1938. 115
3. Stivalius afer timanus subsp. nov. (text-figs. 60, 61).
Sierra Leone : Freetown, on rats (D. H. S. Davies), 1 8, 2 29.
In making a survey of the rats at Freetown and their fleas on behalf of
the Liverpool School of
Tropical Medicine Mr.
Davies obtained these
3 specimens among a
multitude of Xeno-
psyllus brasiliensis and
X. cheopis.
Chaetotaxy as in
S. afer afer Roths. 1908,
apart from the modified
terminal segments of
the abdomen. Z
&. Abdominalster- Uf
num VIII with few |! il
lateral bristles, the cen-
tral area bare, the
bristles numerous ven-
trally and subventrally
(text-fig. 60, VIII. st.),
there being on each side
of VIII. st. about 30
bristles. Dorsal side of
apical portion of digitoid
F more rounded, the
highest point evenly
rounded, not subangu-
late. Dorsal side of paramere
(Par) less rounded than in
St. afer afer, and its apical,
ventral, strongly chitinized pro-
jection (pr) short, not conical-
pointed. Widened distal
portion of ventral arm of IX.
st. broader, more abruptly
widened, its apex not pointed,
and its ventral bristles more
numerous.
®. Abdominal sternum
VII (text-fig. 61) deeply sinu-
ate, as in St. afer afer, but the
upper lobe longer and much
more pointed. Glands (g) near
orifice of bursa copulatrix
vertical.
Length: & 3-5; 9 4-1-4-3 mm.; hindfemur ¢ 0:56, 9 0-61 mm.
The specimens are somewhat extended.
116 .Noviratres Zoonocicar XLI. 1938.
4. Stivalius afer nigeriensis subsp. nov. (text-fig. 62).
Nigeria : Lagos, on bush rat (Mrs. 8. L. M. Counal), 1 9.
This single specimen agrees neither with St. torvus Roths. 1908, nor with
St. afer Roths. 1908. It may represent a separate species ; but as the gland at
the base of the bursa copulatrix is narrow and vertical as in St. afer, I describe
the specimen provisionally as subspecies of that species.
Smaller than St. afer. Pronotal comb with only 20 spines. On mete-
pimerum 10 long and 5 short bristles on one side and 11 and 6 on the other. On
basal abdominal sternum 15 lateral bristles on left side and 18 on right. Upper
antepygidial bristle a little less than one-half the long lower one, in St. afer afer
and St., afer timanus a little longer than one-half. Sternum VII (text-fig. 62)
more nearly resembling that segment of St. torvus ; but the upper lobe shorter,
with but a slight indication of a projecting angle ; three long bristles above sinus,
the lowest of them near margin of upper lobe. Above stigma of VIII. t. four
bristles each side ; on lower widened area of VIII. t. 21 bristles each side, 1 on
inner side above long marginal one ; the marginal lobe above the long marginal
bristle almost effaced. Body of spermetheca (R.s.) without dorsal hump; gland
near orifice of bursa copulatrix narrow, vertical.
Length: 9 3:1 mm. (somewhat extended) ; hindfemur 0-47 mm.
The Iraq Expedition of Field Museum of Natural History, Chicago, obtained
several species of fleas around Bagdad, among which there is an undescribed
Nosopsyllus. The specimens were all collected by Yosuf Lazar and represent
five species :
(a) Xenopsylla cheopis Roths. 1903, 1 g, on Rattus norwegicus, i. 36.
(b) Pulex irritans L. 1758, a series, from an Arabian woman, ii. 36.
(c) Ctenocephalides canis Curtis 1916, a few specimens on a dog, i. 36.
(d) Ctenocephalides felis felis Bouché 1825, a series on a cat, xii.35; also
on jackal, viii. 35, and on dog, i. 36, and:
Novirares Zoonogican XLI. 1938. 117
5. Nosopsyllus medus sp. nov. (text-figs. 63, 64).
Iraq: Bagdad, on Rattus norwegicus, 1 3, 2 99, and on a “ mouse,”
1 & (type figured) ; other specimens, also from Bagdad and “ Mesopotamia,”
in coll. N.C.R. Type at Tring, paratypes SQ in the Field Museum.
$9. Near N. londiniensis Roths. 1903, the 3 distinguished by the long
digitoid, and the 2 by the shape of abdominal sternum VII. Chaetotaxy almost
the same in the two species.
On frons an anterior row of 6 to 8 bristles in 3, of 4 (rarely 5) in 9, and a
posterior row of 3 (rarely 4) in both sexes. None of the bristles of segment II of
antenna reach apex of club. Pronotal comb with 17 to 20 spines, the short
ventral ones inclusive. Apical bristles of mid- and hindtibiae and of segments
I and II of hindtarsus as long as in N. londiniensis.
Modified Segments.—g. At dorsal margin of VIII. t. (text-fig. 63) four long
strong bristles, proximally of the row a smaller bristle, the margin curved upwards
beyond the last long bristle, an apical lobe being formed as in N. londiniensis,
the dorsal and distal margins if prolonged would form an acute angle. Process
P of clasper narrower than in N. londiniensis ; distances of the lowest point
of the movable digitoid F to the tip of P and tip of F very different from the
same measurements in N. londiniensis, the figures being on an average in N.
medus 32: 44 and in N. londiniensis 22 : 29 ; measured on the frontal side process
118 Noyirares Zoonocicar XLI. 1938.
P is about as long as the distance of its tip from the tip of F. Digitoid very
slender, about five times as long as broad at the widest point, which is much
above middle ; posterior margin incurved below this point, where there is a
longish stout bristle, from this bristle to the tip the margin moderately convex ;
a second bristle, somewhat thinner and shorter, at about one-third from first
long bristle to tip. Widened apex of vertical arm of IX. st. in our other Jg more
evenly rounded on posterior side than in type figured. Q. VIL. st. (text-fig. 64)
ventrally less deeply sinuate than in N. londiniensis, the distance of the most
ventral long bristle from ventral apical angle of segment somewhat longer and
in N. londiniensis somewhat shorter than segment V of hindtarsus ; the apical
lobe broader than in N. londiniensis, the lower portion of the margin less strongly
receding. Spermatheca not different, its body somewhat variable in width.
Noyirares Zoonoaicarn XLI. 1938. 119
ON SOME NEARCTIC SIPHONAPTERA.
By KARL JORDAN.
(With text-figures 65-75.)
HE fleas dealt with under Nos. 1, 2 and 3 were sent to me by Dr. R. Parker,
Director of the Rocky Mountain Laboratory, Hamilton, Montana, and the
Assistant Parasitologist, Mr. Wm. L. Jellison, to both of whom I express here
my gratitude. The types of the new fleas have been returned to Dr. Parker,
while one or more
paratypes, where
available, have been
retained at Tring.
1. Malaraeus peni-
cilliger dissimilis
subsp. nov. (text-
figs. 65, 66).
Ceratophyllus penicilliger
(Grube 1852), Jordan,
Nov. Zool., xxxv. p. 36,
no. 24 (1929) (Flat,
Alaska, on Microtus) .
The species is
widely distributed,
being known to us
from Lapland to the
Alps and to the
western mountains
of Transylvania, and
from England to
Japan. It has de-
veloped into several
geographical races, of
which we describe
here the Alaskan one. A small series is before us as follows :
Alaska: Rapids, on Hvotomys, and Fairbanks, on Microtus, July 1937,
7 34,3 22. The two specimens recorded by me, l.c., belong to this subspecies,
the sketch I made of the genitalia of the specimen in the collection of the
U.S: National Museum agreeing with the fresh material sent by Dr. Parker. I
have adopted the name proposed for the new subspecies by Mr. Wm. L. Jellison.
Chaetotaxy as in European specimens, the number of bristles being variable :
on metepimerum from 7 to 12, on tergite VIII of ¢ from stigma downwards 20 to
33, of which 3 to 6 are subventral in one or two vertical rows.
Modified Segments.— 3. Process P of clasper (text-fig. 65) longer than in the
Old World races, and the acetabular bristles, of which there are sometimes 3
120 NOVITATES ZooLoGIcAE XLI. 1938.
(as happens occasionally also in European 3), thinner and shorter. Digitoid
F as broad as in M. p. penicilliger at the widest point, but much longer and
therefore appearing slenderer, the dorsal rounded bay between the short pedicel
and the main body of F wider than in Old World examples. Proximal, elongate-
oblong partition Lot of IX. st. longer and its subspiniform bristle shorter and
thinner. Sternum VIII narrower, especially the widened apical portion, and
this portion and the widened basal one farther apart, i.e. the narrow median
portion longer. Process of paramere (Par) longer and narrower than in M. p.
penicilliger. 2. Sternum VII (text-fig. 66 a and b) deeply sinuate, the upper
lobe variable, broad, dorsally rounded.
For comparison I give a figure of sternum VII of a 9 from Lapland (text-
fig. 67), which is M. p. penicilliger. One of our Siberian 99 has practically the
same sternum, whereas
67 in others the lobes are
shorter. In the specimens
from farther south in
Scandinavia, the Alps
and Great Britain the
sinus of sternum VII is
shallow, sometimes almost
effaced, very rarely deep.
The geographical varia-
tion in Europe of M.
penicilliger is now being
studied. Apart from the
Alps, we have no speci-
men from France and
Germany, which is worthy
of note; does the species
occur in the central and northern provinces of these countries? It is common
in England.
2. Monopsyllus exilis Jordan 1937 (text-figs. 68, 69).
& Megabothris exilis Jordan, Nov. Zool., xl, p. 264, no. 3, text-fig. 46 (1937) (Montana).
In the two ¢¢ on which the species was based the antepygidial stigma
could not clearly be seen, and as at the time I studied the species my mind was
much perturbed by Lord Rothschild’s illness and the uncertain future of the
Tring Museum, I made the mistake, otherwise inexcusable, of placing the species
into Megabothris instead of Monopsyllus, where it belongs, the species being near
M. ciliatus Baker 1904.
In the single additional 3 kindly submitted by Messrs. Parker and Jellison
the two apical spiniforms of the digitoid F (text-fig. 68) are thinner than in my
figure of 1937; each is drawn out into a sharp point, the lower one especially
resembling a thin bristle. The subapical thin bristle is longer in this specimen,
and sternum VIII bears a few more bristles, there being a row of 4 in proximal
two-thirds, and a subapical bristle (each side) as in paratype.
The @ is distinguished from M. ciliatus by the shape of sternum VII. This
segment is sinuate, as it is in M. ciliatus, but the lobe above the sinus is short,
very broad and rounded, and sometimes projects very little; text-fig. 69, a,
Novirates ZOOLOGICAE XLI. 1938. 121
b, c, illustrates this variability. Chaetotaxy almost as in M. ciliatus. In the
Montana 9° there is no incrassation below or in front of the sinus of VII. st.
The marginal bristles on inner side of widened area of VIII. t. situated below the
long marginal bristle of the outside are thinner and longer than in M. ciliatus.
Stylet longer. Spermatheca (R.s.) larger, the tail shorter in proportion to its
width. The striae of the basal abdominal sternum of 9 as deeply curved back-
wards in middle of side as in M. ciliatus, and very close together in apical area.
The Montana specimens were found in a nest of Spreo tyto in Beaverland
county in June 1937.
Dr. Parker sent also some specimens from Arizona and Colorada, which are
slightly different from the Montana examples and, I think, represent two further
local races.
a. M. exilis opadus subsp. nov. (text-fig. 70, d, e).
Arizona: Yavapai, April 1937, on Dipodomys olii subsp., 1 g, 3 29, and
on Onychomys leucogaster capitulatus, 1 2.
6. In the ¢ sternum VIII is as in our figure of 1937; but distance from
lower spiniform to
long marginal bristle
below it the same as
to marginal bristle
above the spiniforms.
The lower apical spini-
form of digitoid F is
narrower and much
shorterthantheupper,
but not thin and
bristle-like as in the
Montana specimen
above described. The
vertical arm of ster-
num IX subapically
more strongly widened than in the 1937 figure. Sternum VII of 9 (text-fig.
70, a, b) with short rounded upper lobe; lower lobe very broad, rounded-
truncate, the apical margin vertical, not incurved asin M. exilis ewilis. Type 3.
122 NoviratEes ZooLoGIcAE XLI. 1938.
b. M. exilis triptus
subsp. nov. (text-
a Se s Nee, WIL hy Pec
; Colorado:
Rugger, June 1937,
on Dipodomys or
Onychomys, 2 29.
®. In these two
©9, which are some-
what smaller than
those from Arizona
and Montana, the
sinus of VII. st. is
deeper and nar-
rower, the upper
lobe projecting
farther distad; the
lower lobe in one
specimen nearly as
in the Arizona 99,
except that it is
longer, in the other
specimen this lobe
is narrow and its
short apical margin
oblique. Text-fig.
71 g from type.
M. ciliatus is not known to us from Colorado and Arizona ; M. exilis perhaps
takes its place.
3. Megabothris asio orectus subsp.
nov. (text-figs. 72, 73).
Montana: Ravalli county, October
1935, in nest of a mouse, 2 $4, 299;
type 3.
39. A western subspecies. In the ¢
the apex of the digitoid F more produced
backwards than in M. a. asio Baker 1904,
and the terminal margin between this
nose (which resembles that of Monops.
cihatus) and the anterior apical angle
less rounded ; the prominent ventral lobe
of sternum IX narrower, and sternum
VIII has fewer ventral bristles. In the 9
the upper lobe of sternum VII is slightly
longer than in the specimens from the
states of New York and Massachusetts,
and the distance of the third or fourth bristle of the posterior row of this segment
from the one above it greater.
Novitates Zoovocıcar NLI. 1938. 123
4. Foxella ignotus ignotus Baker 1895 (text-figs. 74, 75).
Ceratophyllus ignotus ignotus (Baker 1895), Jordan & Rothschild, Betoparasites, i, p. 55, text-fig. 57
(1915) (3g, type of Typhl. americana) ; Jord., Nov. Zool., xxxv, p. 32, no. 13 (1929 (32).
Until recently the only specimens of this subspecies which I had seen were
Baker’s original 2 $¢ and 3 99, from Ames, Iowa. I have now received a fine
series of both, very generously sent to me by Mr. C. C. Sanborn and Mr. K. P.
Schmidt, of the Field Museum of Natural History, who collected the specimens
at Pembroke, Kankakes county, Illinois, on Geomys bursarius illinoisensis, and
to whom I am very grateful for this kind help.
Sternum VIII of the Illinois $¢ agrees with our fig. 57 in Ectoparasites,
l.c., differing from that of the various other subspecies in the vertical arm meas-
2
x
ured from the ventral margin to the tip being as long as the ventral arm measured
from the anterior margin to the tip of the apical projection ; the variability in
the proportions is slight. The long bristle of this segment is always very stout
proximally, thicker than in any other subspecies. Proximally of this bristle
there are one or two small bristles each side and usually indications of minute
hairs. The apex of the basal vertical arm of VIII. st. as slender as in text-fig. 58
of Hctoparasites, but curves forward. Process P of clasper varies a little in
width and length. As in F. 7. franciscanus Roths. 1910, it is always ! shorter
than the distance from the angle above the acetabulum to the ventral curve of
the acetabulum, whereas in all the other subspecies the process is longer than the
vertical diameter of the acetabulum. The digitoid F resembles that of F. 7.
recula J. & R. 1915, from British Columbia ; its apex is more rounded in some
specimens than in others ; the posterior margin is feebly incurved above middle ;
the width is somewhat variable, and if the digitoid is moved away from P, it
appears narrower than if lying vertically along P or partly inside it, as shown in
text-fig. 74; the position of the upper long bristle is not quite constant. The
1 “ Always” in a diagnosis means “all the specimens before the author.”
10
124 Novirates ZooLoGICAE XLI. 1938.
paramere is long-nosed, as in F. 7. albertensis J. & R. 1915. Spicules on dorsal
area of inner surface of VIII. t. more conspicuous than in other subspecies.
I mentioned in 1929, l.c., that the sternite VII is deeply incurved in the Iowa
92. Inall the Illinois 99 before me the apex of this segment is likewise incurved,
but not so deeply in any of them as in our Iowa ® (cf. text-fig. 75, a from Iowa,
b and c from Illinois) ; there is a good deal of variability in the Illinois series of
99, the ventral apical lobe projecting much more in some than in others. The
bursa copulatrix is as short as in F. i. recula. Body of spermatheca either
almost globular or ovate.
Epitedia gen. nov.
Like Catallagia Roths. 1915, but with genal spines as in Neopsylla Waen.
1903. Differs from the latter genus in the vinculum of basal abdominal sternum
being absent, in segment V of fore- and midtarsus bearing 4 lateral pairs of
plantar bristles and a proximal ventral pair, whereas this segment of hindtarsus
has only the 4 lateral pairs. Body of spermatheca longer than its tail, convex
below, concave above, line of demarcation between body and tail distinct above
and below, tail deeply projecting into lumen of body. Upper margin of pro-
pleurum with sinus where the posterior end of the (internal) vinculum between
head and prosternite touches it; this sinus absent in Neopsylla. Abdominal
terga II to III with two rows of bristles ; sternum VIII of & without bristles at
and near apical margin. Marginal incrassation of metasternum (squamoid, in
the membrane closing sternal lumen in front) longer than broad, in Neopsylla
broader than long or vestigial. ‘The pointed endoskeleton (a fork of which one
sees one prong only in lateral aspect ; the fork supports the chain of ganglia)
of the metasternite is directed straight upward, being placed in between the
meral rods, whereas in Neopsylla it has a slightly more forward position, being,
in mounted specimens, generally more or less directed obliquely upward-
forward. Genotype: Ctenophthalmus wenmanni Roths. 1904. Here also
belong Neopsylla faceta Roths., 1915 and Neopsylla testor Roths. 1915.
Neopsylla— Epitedia— Catallagia are an interesting chain of genera.
Tamiophila gen. nov.
The giant among Neopsylla disturbs the diagnosis of the genus and is better
segregated from the rather uniform series of species. It agrees in most essentials
with Neopsylla, but differs in various details, being easily separated by the
hairiness of the underside of the basal abdominal sternum. There are but few
submarginal bristles on the inner surface of the hindcoxa and no short spini-
forms. The patch of very dense striation on the side of the basal abdominal
sternum usually very conspicuous in Neopsylla (as it isin Epitedia and Catallagia)
indicated by the striation being denser than dorsally, ventrally and anteriorly
on the segment, but does not contrast very much. On mid- and hindtibiae 8
dorsal notches, each having a pair of bristles, between sixth and seventh pairs an
additional strong dorsal bristle. Genotype: Typhlopsylla grandis Roths.
1900.
Novrrares ZooLoGicAau XLI. 1988. 12
ou
NOTE ON POLYGONIA C-ALBUM (L.) 1758, AT ASHTON WOLD,
NORTHAMPTONSHIRE, AND THE RECORD OF A NOTABLE
ABERRATION.
By MIRIAM ROTHSCHILD AND MALCOLM SPOONER.
(With Plate III, figs. 1, 2.)
| A History of British Butterflies (F. ©. Morris, 1857) the Comma is recorded
from Ashton Wold, but during the period 1900-1921 it was not seen in that
locality. In the early spring of 1922 one of us (M. R.) was collecting in the
Home Wood with the late N. C. Rothschild and saw two specimens on the wing.
A single female was captured the next summer in August. During the years
that followed the species became increasingly common, as elsewhere in England.
It is, however, curious that during the summer of 1937 this butterfly should
have been more abundant than in any other year, for it proved the worst season
in living memory for species such as the Small Tortoiseshell (Aglais urticae
(L.) 1758). On different days during August, from five to ten pairs of the
Comma were counted in the garden, and one very remarkable aberration was
captured on Heliotrope. This specimen,! as will be seen from the photograph
(Pl. III, figs. 1, 2), essentially resembles the variety known as var. Symond’s
Yat (Frohawk, Natural History of British Butterflies, Plate 21, figs. 18-19).
It would appear that these are the only two records of this variety in Great
Britain. On the Continent a very similar aberration is known as ab. f-albwm
Esper (= j-album Spuler (Macrolepidoptera of the World, Seitz, Vol. 1, Pl. 63)
from the fact that the “c” takes the form of a small “f.” In the specimen
recorded here the mark is intermediate in form.
1 Placed in the British Collection, Tring Museum.
126 NOVITATES ZooLoGIcAE XLI. 1938.
ON THE REMARKABLE VARIATION OF AN ORIENTAL HAWK-
MOTH (LEPIDOPTERA).
By KARL JORDAN.
(With text-figures 76-79 and Plate III, figs. 3-6.)
A NEW subspecies of Polyptychus trilineatus Moore 1888 received from Celebes
agrees so closely with the various other subspecies in colour and pattern
and differs so markedly in the male genital armature that the re-examination of
these organs in the entire species (as far as represented in the Tring collection)
promised to be of interest. In order to be able to refer to the Celebes subspecies
by name I insert here the description before giving an account of the variation
of the species.
Polyptychus trilineatus kalisi subsp. nov. (text-fig. 79; Pl. III, fig. 4).
Central-West Celebes: Gunong Tompoe, Paloe, 2,700 ft., i., 11.1937 (J. P. A.
Kalis), 2¢¢; 4 other gg obtained by Mr. Kalis in the Maros district, South-
West Celebes, ii. 1938.
6. As dark as fresh specimens of P. t. undatus (North India), the grey areas
of forewing rather glossy and contrasting with the black areas ; there is nothing
definite in the pattern in which the subspecies differs from the others; the white
transverse bar of hindmargin of hindwing close to anal angle (part of a discal
line) more oblique than in Indian specimens, but that is also the case in the
races from the Philippines and Andamans. The white colouring of underside
variable in extent. Upperside of mid- and hindtibiae nearly entirely blackish, or
more or less buff. Size as in ¢¢ from the Philippines and Andamans.
Genitalia : Anal tergite (X. t.) broader than in other subspecies, the apical
hook broad, apically rounded ; anal sternite (X. st.) a narrow ridge. Ventral
division of clasper (Cl) ending in a single, strongly curved, hook. The pair
of horns present in other subspecies below X. st. absent. Ventral process (ve)
of penis-funnel straight in terminal half, with the apex subtruncate or irregularly
rounded ; right dorso-lateral process (ri) of penis-funnel very short, left process
absent. Length of forewing 43-45 mm., width 17-19-5 mm.
We know now 9 subspecies :
P. t. luteatus R. & J., Revis. Sphingidae, p. 273 (1903), Ceylon.
P. t. sonantis Jord., Nov. Zool., xxxvi, p. 2 (1930), South India.
P.t. trilineatus Moore, Proc. Zool. Lond., p. 290 (1888), North-West India.
P. t. undatus R. & J., l.c. p. 238 (1903), North-East India.
P. t. costalis Mell, Stidchin. Sphing., ü, p. 130 (1922), South China.
P.t. mincopicus Jord., l.c. p. 3 (1930), Andaman Is.
P.t. kelanus Jord., lic. p. 3 (1930), Sumatra.
P.t. philippinensis R. & J., l.c. p. 239 (1903), Philippine Is.
9) P. t. kalisi subsp. nov., Celebes.
The species is not yet represented in the Tring collection from Formosa,
West China, Indo-China, Burma, Siam, Malay Peninsula, Borneo, Palawan, Java,
where it may certainly be expected to occur.
Novırares ZooLogIcar XLI. 1938. 127
The hawkmoth described as P. t. chinensis R. & J. 1903 from China we now
consider to be a distinct species ; the mutilated 2 on which the name was based
is still the only example at Tring.
The genital armature of the 3 is somewhat complicated, and in 1903 we
were uncertain about the homology of some of the sclerites ; the comparison of
several subspecies discovered since has cleared up the doubtful points. In order
to demonstrate ad oculos the interesting differences we give small diagrammatical
figures of the organs of P. t. luteatus and of P. t. wndatus, but omit terga IX and
X, and for comparison with P. t. kalisi from
Celebes we give a figure of equal size taken
from a specimen of P. t. philippinensis.
The 9 subspecies fall into 4 groups according
to the development of the genital armature :
I. P. t. luteatus, Ceylon (text-fig. 76),
and P. ¢. sonantis, South India. Tergum
X shorter than in the other subspecies.
Ventral division of clasper rounded at apex,
with a slight sinus and on the concave inner
surface a sharp ridge, more oblique and
shorter in luteatus than in sonantis, swollen
ventral margin with an upward projection,
the pair of horns (h) below X. st. present ;
these horns are projections from the con-
tinuation mesad of the upper margin of the
upper division of the clasper, and occur also
in some other species of Polyptychus ; in
function they probably replace X. st., which
is a low ridge and evidently of no great service
in the lock and key arrangement of the sexes.
Ventral process of penis-funnel (ve) widened
at apex and here curved towards the right
side (in Revis. Sphing., pl. xxxiv, fig. 10, the
process is seen from the upperside and points =
to the left, which is the right side of the yy
body) ; the right dorso-lateral process (ri) is ‘
short and the left (le) still shorter in the t
specimen from which the figure is taken, but both may be entirely absent or
vestigial.
II. P. t. trilineatus, North-West India, P. ¢. undatus, North-East India,
P. t. costalis, South China, P.t. mincopicus, Andamans, P. t. kelanus, Sumatra.
The area of distribution of this group embraces no doubt all the countries from
North India and China south and east as far as Borneo (and probably Palawan)
and Java. Tergum X longer than in Group II; X. st. a narrow ridge (text-
fig. 77). Ventral division of clasper (Cl) apically divided into two short, more or
less pointed and curved, claw-like processes, which are different in the various
subspecies of this group ; ventral margin without projection about middle. The
pair of horns (h) present. Ventral process (ve) of penis-funnel longer than in
Group I, along middle membranous as in I, apically divided into two projections
and resembling a fish-tail, the right projection longer or shorter than the left
128 Novitates ZOOLOGICAE XLI. 1938.
according to subspecies ; right dorso-lateral process (ri) long, and the left much
shorter.
III. P. t. philippinensis, Philippines. We have studied only the armature
of Luzon males ; the Mindanao 3 mentioned in the Revision, p. 240, may differ,
and if that is the case, Mindoro specimens also will show differences ; we may,
therefore, expect that at least three subspecies belong to this group. X. t.
with long and narrow apical hook. X. st. a narrow undulated ridge. Ventral
division of clasper (Cl) with two lobes, one apical, rounded at apex, concave on
inner surface, appearing in a view from inner side (text-fig. 78, A) narrower than
in ventral aspect (text-fig. 78) ; the upper margin of this lobe gradually dilated
upward into a triangular projection, the apex of which is smaller than 90°. Pair
of horns (h) present, rather
strongly arched. Ventral
process of penis-funnel (ve)
long, at apex gradually
narrowing to apoint, lateral
margins minutely denticu-
late, apex slightly turned
towards the left, not to the
right as in group I; right
process (ri) very long,
reaching to apical fifth of
broad process ; left process
(le) indicated by a suture
and a small notch, not
separated from ventral
process.
IV. P. t. kalisi, Cele-
bes. The distinctions
have been given above ;
the most remarkable differ-
ences being the broad apical
hook of X. t., the absence of
the pair of horns and the
abrupt narrowing of the apex of the ventral division of the clasper into a
strongly chitinized single long hook (text-fig. 79). The ventral process (ve) of
the penis-funnel has proximally a deep median channel. X. st. is a narrow
ridge or fold which has the appearance of being pulled anad by the longitudinal
median ridge which connects it with the anal cone (An). Similar subspecies
probably occur on the Sula Is. and Buru. In one of. the 4 specimens from
the Maros district there is a small marginal tubercle some distance from the
hook of the clasper.
The first point of general interest is the very striking contrast between the
geographical modification of the colour and pattern on the one side and the male
genital armature on the other. The female organs of copulation also are different
in Groups I, II and III, but as we have no 9 of Group IV, I abstain from referring
to the structure of this sex. I find nothing in colour, pattern, size and wing-
contour which quite definitely distinguishes one subspecies from all the others,
or one group from the other. The 4 photographs, figs. 3-6, on Plate III illustrate
Novirates ZooLoGIcAE XLI. 1938. 129
this statement ; individuals, of course, are always different from each other in
some way or other. The male genital armature, on the contrary, is so strongly
modified in the 4 groups of subspecies that there would be great justification for
treating each group as being a distinct species. It is obvious that the causes of
evolution underlying the development of P. trilineatus into geographical modifica-
tions have affected the genital armature very strongly and the colour and pattern
of the insect hardly at all; the one set of somatics has gone ahead and the other
has remained stagnant, there being no co-ordination between them and no
interdependence.
The groups of subspecies
we are considering represent
each other geographically just
as do the subspecies within
each group. The differences
between the groups are large
and those between the sub-
species of groups I and II
small—if in future Groups III
and IV are found each to
comprise several subspecies,
what we have said of I and II
will doubtless apply to all four.
But the differences are of the
same kind; it is a matter of
degree. Each group inhabits
an area known to be a major
zoogeographical unit of the
Oriental Region. The separa-
tion of one such unit from the
other took place at an earlier
period than the separation of the minor divisions within these zoogeographical
units, and we may look upon the 4 groups of P. trilineatus as representing
the early development of the species into 4 subspecies. These were the ancestral
subspecific divisions of P. trilineatus, which became further modified and, at a
later period, themselves split up into subspecies. The process of evolution is
here evident. The older the original area, the longer the isolation, the greater
the differences of the subspecies.
Although the 4 groups may already have attained a degree of independence
of each other which equals the independence existing between species that live
side by side, we keep them as divisions of one species with the object to emphasize
the process of their origin and at the same time to simplify classification.
Plate III, fig. 3, P. t. sonantis, South India, type.
Plate III, fig. 4, P. t. undatus, type.
Plate III, fig. 5, P. t. philippinensis, Luzon.
Plate III, fig. 6, P. t. kalisi, Celebes, paratype.
130 Noyirates ZOOLOGICAE XLI. 1938.
ON SOME NEW OR UNFIGURED LEPIDOPTERA FROM THE
ORIENTAL AND ETHIOPIAN REGIONS.
By KARL JORDAN.
(With Plates III and IV.)
1. Delias itamputi luctuosa Jord. 1912 (Pl. III, fies. 7, 8).
3g. D.i. 1. Jordan, Nov. Zool., xviii, p. 584, no. 4 (1912) (Mt. Groome, 11,000 ft.).
The specimen, a 4, is still the only one recorded of this subspecies, which is
distinguished by the great extent of the black borders.
2. Charaxes tectonis Jord. 1937 (Pl. III, figs. 9, 10).
&. Ch. t. Jordan, l.c., xl, p. 323, no. 1 (1937) (Cameroon: Bamenda).
No further specimens have come to hand.
3. Daphnusa ocellaris ailanti Boisd. 1875 (Pl. IV, figs. 11 g, 12 2).
3. Smerinthus ailanti Boisduval, Spec. Gen. Lep. Hét., i, p. 28, no. 16, pl. 3, fig. 2 gd (1875).
6. Daphnusa ocellaris ailanti, Rothschild & Jordan, Revision of Sphingidae, p. 285, no. 2415 (1903).
Although Boisduval stated that his specimen had been obtained from a
chrysalis found by Lorquin at the foot of Ailanthus glandulosa at Hongkong,
the true locality is Celebes, probably the Minahassa district where Lorquin had
been collecting. We have lately received several JS and a 9 from Mr. J. P. A.
Kalis, who obtained them inland from Paloe Bay, West Celebes, at various
places from November 1937 to April 1938, at altitudes from 900 ft. to 3,700 ft.
As stated in the Revision, the distal margin of the forewing is more convex
below middle than in D. o. ocellaris Walker 1856 and the apex of the hindwing
more broadly rounded. Moreover, the tornus of the forewing projects less far
backward, particularly in the &, and the hindwing is somewhat differently
marked at the anal angle, bearing here close to the distal margin a short broadish
brown continuous band.
4. Hippotion moorei moorei Jord. 1926 (Pl. IV, fig. 13 3).
3. H. m. Jordan, Nov. Zool., xxxiii, p. 383, no. 11a (1926) (Tanganyika Terr.: Mwanza).
5. Hippotion exclamationis robur subsp. nov. (Pl. IV, figs. 14, 15).
Q. Larger and more robust than H. e. exclamationis Fawe. 1915 from Kenya
and H. e. austrinum Jord. 1930 from Zululand. Occiput, upperside of thorax
and abdomen and the wings above and below much more shaded and irrorated
with blackish olive, the base of the abdomen being particularly darker than in
the other subspecies.
On upperside of forewing the discal dots form the proximal boundary of a
diffuse olive band, which is more or less conspicuous ; in one paratype (fig. 15)
there is rufous scaling within this band ; dots at apices of veins well-marked.
Hindwing very dark, blackish olive, the pinkish submarginal diffuse band of the
other subspecies absent or vestigial.
Novirarrs Zoonocicamn XLI. 1938. 131
On underside the black irroration strongly developed on both wings; the
transverse row of dots and the submarginal line variable on both wings; in
type the apex of hindwing black, this colouring extending to apex of cell.
Length of forewing 29-31 mm., breadth 11-12 mm.
Tanganyika Territory: Banagi Hill, Musoma, 3 99 at light on April 14,
15, 18, at 7.30 and 8.30 p.m. after heavy rain. We have to thank Mr. and Mrs.
M. S. Moore for this interesting discovery and for having carefully noted the
time of appearance at the lamp.
6. Hippotion rebeli R. & J. 1903 (Pl. IV, fig. 16).
62. H.r. Rothschild & Jordan, Revision of Sphingidae, p. 761, no. 715, pl. v, fig. 12 9 (1903) (Sudan ;
Obock).
Besides the two original specimens there are now several others in collec-
tions. The British Museum has received the species from S.W. Arabia, and at
Tring there is a pair from Diré Daoua, Abyssinia, collected in March and May
1936 by H. Uhlenhuth, the ¢ of which we figure. The species evidently repre-
sents H. exclamationis in the Sudan, Abyssinia, Somaliland, and South Arabia.
It is distinguished in colour from the more southern species, the body and the
upperside of forewing and underside of hindwing being clayish buff, and the
hindwing above and the greater part of the forewing below more or less tawny
ochraceous with a slight pink tint.
7. Hippotion butleri Saalm. 1884 (Pl. VI, figs. 17, 18).
9. Panacra butleri Saalmüller, Lep. Madag., p. 118, no. 275, pl. 5, fig. 51 (1884) (Nossibe).
We have received two dd of this rare Malagassic species, of which only
one specimen, a 9, has been recorded as far as we know. The upper & (fig. 17)
was obtained by Monsieur R. Catala in the Forét d’Isaka, near Fort Dauphin,
Nov.-Dec. 1934 ; it is in good condition and shows the pale diffuse submarginal
band of the forewing but faintly on the upperside. The second ¢ was collected
by Monsieur and Madame d’Olsoufieff in March 1935 at Périnet, east of Tanana-
rivo ; it is somewhat worn, having lost some of the scaling of the upperside, in
consequence of which the conspicuous pale submarginal band of the underside
is also very distinct above. The triangular spot before the anal angle of the
hindwing is less deep buff than in Saalmiiller’s figure and less sharply defined.
8. Temnora grandidieri Butl. 1879 (Pl. IV, fig. 19).
6. Diodosida grandidieri Butler, Ann. Mag. N.H. (5), iv, p. 234 (1879) (Madagascar).
6. Temnora grandidiert (Butler), Rothschild & Jordan, Revision of Sphingidae, p. 570, no. 495,
pl. vii, fig. 8, 3.
When we wrote the Revision we had seen only the 3 $& in the British
Museum and the 2 3 in coll. Oberthür. The Tring collection now contains a
series of 26 SS from the neighbourhood of Périnet, 140 km. east of Tananarivo,
collected by M. and Mme. d’Olsoufieff in x., xi., xii., i. and 11.1930 to 1935, and
2 Sg obtained by M. R. Catala at Ankarampotsy, near Fianarantsoa, col de
Tantamaly, xii.33. Among the series sent by M. d’Olsoufieff there was a single
specimen of the following species. We figure a specimen of T. grandidieri for
comparison.
132 Novirates ZooLOGICAE XLI. 1938,
9. Temnora engis Jord. 1933 (Pl. IV, fig. 20).
d. T.e. Jordan, Nov. Zool., xxxviii, p. 342, text-figs. 58-61 (1933).
The difference from 7’. grandidieri in the shape and markings of the forewing
is well shown in the photograph. The hindwing is tawny, with a rather ill-
defined darker border.
10. Ludia arida sp. nov. (Pl. IV, fig. 21).
3. In colouring half-way between L. corticea Jord. 1922 and L. hansali Feld.
1874, but differs much in the genitalia from these and all the other species of Ludia.
Body broccoli brown, upperside of thorax blackish, the pale lanceolate or
spatulate hairs less numerous, less widened at apex and less contrasting in
colour with the normal hairs than usually. Pronotum and base of tegulae buffish
cream, nearly ecru drab, contrasting very strongly with the dark colouring of
the mesonotum. Antenna with 32 segments, 16 of them quadripectinate, but
the branches of the 16th quite short.
Wings, upperside. Forewing: submedian transverse line evenly excurved
from costal margin to below cell as in L. hansali, then straight to hindmargin,
but this posterior half slightly incurved on submedian vein; proximal area up to
this line the same pale colour as collar, the pale colouring extending costally beyond
the line, behind cell shaded with broccoli brown ; submarginal transverse line
scalloped, without the conspicuous pale band of ZL. hansali on its distal side ;
marginal area finely irrorated with broccoli brown ; termen undulate, fringe pale
between the veins; upper vitreous mark a little dilated at both ends, slightly
curved, lower mark projecting distad beyond upper mark, its lower end very
thin and vestigial. Hindwing : the black area surrounding ocellus connected
with costal margin by a very faintly blackish and narrower continuation of the
dark area, a small subapical red patch being thereby separated from the proximal
red area; marginal area as on forewing irrorated with broccoli brown ; termen
undulate, rounded tooth at M! more projecting than the others; fringe pale
between veins.
Underside almost uniformly pale drab, minutely irrorated with darker scaling,
costal margin of both wings with light and dark spots, which are more apparent
on hindwing ; submarginal line inconspicuous on both wings ; red area of fore-
wing diffuse distally and not distinctly reaching the submarginal line.
Neuration : the three subcostal veins of forewing on a long stalk, first and
third branching off from second at the same point.
Genitalia : Median lobe of sternite VIII more broadly rounded than in any
other species. Anal tergite divided. Clasper very broad; its ventral lobe
short and pointed, apical lobe broad, rounded at apex as in JL. corticea, with a
few small teeth, measured from the sinus which separates the lower lobe the
apical lobe nearly as broad as long.
Length of forewing 30 mm., width 14-5 mm.
Angola : Murro de Pundo, 60 km. north-east of Lobito, 300 m., 21 May 1934
(K. Jordan), 1 &.
The specimen came to the lamp after midnight.
11. Ludia orinoptena monroei subsp. nov. (Pl. IV, fig. 22).
9. Antenna with 27 segments, 16 of them bipectinate, but teeth of 16th
very short. Termen less rounded in both wings than in the other subspecies of
Novirates ZooGoGIcar NLI. 1938. 133
L. orinoptena, forewing narrower, submarginal line of hindwing more parallel
with termen and posteriorly a little farther away from anal angle than in 9°
from tropical East and West Africa, and rounded median lobe of vaginal plate
less projecting. Subcostal branches of forewing on a shortish stalk, first and
third branching off from second at the same point. Upper vitreous spot of
forewing thin at lower end, on right wing covered with scaling.
Length of forewing 33 mm., width 16 mm.
North Transvaal: Woodbush, Pietersberg district, 25 February 1936
(E. Munroe), 1 9.
Named in honour of Mr. E. Monroe, who very kindly presented the specimen
to me.
12. Argyrolepidia stevensi sp. nov. ; Agaristidae.
9. Head hairy, abdomen banded with yellow and hindwing with two yellow
spots, the outer one large, nearly central.
Cone of frons high, with feeble indication of rim ; sides of frons white mixed
with black, and bearing long hairs overhanging eye, similar hairs behind eye in
border composed of white erect scaling. Antenna setiform, with a regular row
of white spots on upperside. Segments I and II of palp white beneath, black
at side, with long black hairs, I beneath and at upper apical angle washed with
yellow, III longer than usual, as long as II, rather strongly incrassate at apex.
Pronotum narrowly margined with pale yellow in front and white laterally as
well as posteriorly and in middle line. Meso-metanota isabella-colour, on meta-
notum a pale yellow lateral tuft. Abdomen black, rather rough-hairy, regularly
banded with yellow at apices of segments, at base a large black dorsal tuft, with
some yellow scale-hairs at the sides, anal tuft black externally, hairs beneath this
covering yellow. Breast and coxae orange-yellow, paler at side, abdomen
beneath banded with orange-yellow. Legs black, femora with yellow hair on
upperside, on tibiae some white scaling, especially at tip; dorsal side of spurs
of mid- and hindtibiae basally more or less white ; on tarsi a few white scales
at apex of I and II or only on I.
Wings, wpperside, black, with a very feeble blue tint. Forewing : with
three metallic blue, very thin, lines, one near base, second about half-way between
base and lower median vein, forming a shallow arc below cell, third discal, curving
basad-posticad from lower median, in cell beyond middle a grey spot shaded with
brown, on disc a similarly coloured narrow band, about 2 mm. broad, well separate
from cell, nearly straight from near costal margin, opposite lower angle of cell
curved backwards to lower median, where it ends ; fringe white at apex of wing.
Hindwing deeper black than forewing; fringe of anterior angle white ;
below cell close to base of lower median a pale orange spot about 14 mm. wide,
outside the cross-veins a larger spot, 5 x 7 mm., somewhat oblique, the same
colour, irregularly rounded.
Underside of both wings with blue scaling at base, markings as above, but
no metallic transverse lines on forewing and the cell-spot and abbreviated discal
band pure white.
Length of forewing 23 mm., width 12 mm.
Papua: Morobe district, 7,000 ft., xi. 1932 and iii.1934 (H. Stevens), 2 99.
134 Novirates ZoonocicaE XLI. 1938.
ON THE SPECIES OF BIRD-PARAPSY LILI FROM THE FALKLANDS
OBTAINED ON THE BRITISH GRAHAM LAND EXPEDITION,
1934-37.
By KARL JORDAN.
(With text-figures 80-84.)
HE specimens were collected on Kidney Island by Mr. Brian Roberts, who
informs me that fleas were very common in the burrows of Spheniscus
magellanicus, but rarely found on the birds themselves ; they occurred occasion-
ally also on the bodies of Hudyptes cristatus, and were found in the burrows of
Procellaria aequinoctialis, but not on the birds, although about 15 birds were
examined. ‘‘ These fleas,” says Mr. Roberts, i.l., “ are well known locally as
Jackass fleas, Jackass being the local name of Spheniscus magellanicus. They
were so common that, while studying these penguins, it was necessary for me to
strip three or four times daily in order to get rid of them. Their bites are
particularly irritating to the men who regularly go out to the island to cut tussock
grass for fodder.’ We are very grateful to Mr. Roberts for having preserved
some specimens of what is evidently a local pest. As we had already a good
series of specimens from the same island, sent to us by Mr. A. G. Bennett, all of
which were Parapsyllus longicornis Enderl. 1901, I expected Mr. Roberts’s
material also to belong to that species. But when they were mounted and
could be compared, I found three species instead of one. It was a surprise.
All three are almost black and look to the naked eye identical, for which reason
probably Mr. Roberts considered a dozen and a half a sufficiently large number
of specimens of one species. I should have been still more grateful to him for
a couple of hundred, or as many as he could have crammed into the tubes. We
systematists require quantities, but have generally to be content with what we
get. The two species discovered by Mr. Roberts have some characters in common
with the mammal-Parapsylli, which makes it almost certain that these bird-
fleas are derived from mammal-fleas. The diagnosis of group A of Parapsyllus
which we gave in Ectoparasites, i, p. 352 (1923) does no longer apply ; both new
species have a larger number of bristles than P. longicornis and P. australiacus
on which the description of group A was based, and in both new species the
mesonotum bears a row of false bristles, which are absent from the other two
fleas ; moreover, the bristles of the second segment of the antenna are short,
not long as in the other two species. All four bird-Parapsylli contrast with the
numerous species occurring on mammals of the Andesian countries in being
larger and more strongly chitinized, the internal incrassation from the base of
the antenna dorsad being always strongly marked.
P. longicornis and P. australiacus Roths. 1909 evidently represent each
other, the former being known from St. Paul Island and the Falklands (Kidney
Island), and the latter from various islands along the south and west coasts of
Australia and from Chile; that is to say, the one is Atlantic and the other
Pacific. One would much like to know what flea or fleas occur on the penguins
of the islands of South Africa.
Novirates ZooLoGIcaR XLI. 1938. 135
1. Parapsyllus longicornis Enderl. 1901.
Kidney Island : a series off Spheniscus magellanicus and Hudyptes cristatus,
Nov.-Dec. 1936 (Brian Roberts). Also received from Sparrow Cove, Falk-
lands, off Pygoscelis papualis, Oct. 1933, and Kidney I., off Hudyptes cristatus,
Noy. 1934 (A. G. Bennett).
In most specimens there are a few small bristles in front of the postmedian
row of the meso- and metanota. On the whole larger than P. australiacus, the
digitoid of & longer, the parameres of a different shape, the sinus of sternum VII
of 2 wider and the tail of the spermatheca longer.
2. Parapsyllus magellanicus sp.nov. (text-figs. 80, 81, 82).
Kidney Island: Spheniscus magellanicus, 28 Nov. 1936 (Brian Roberts),
Wy i @
82. Eye smaller than in P. longicornis, genal process behind eye longer,
As
bes z
N
|
|
less rounded ; in front of eye 3 bristles ; farther forward a row of 6, the most
ventral one near the frontal tubercle. On occiput an anterior, a median and a
posterior long bristle and on side an oblique row of 3 to 5, near posterior margin
4 to 6 (each side) besides the long ventral one. Bristles of antennal segment II
short.
136 Novirates ZoOoLOGICAE XLI. 1938.
Bristles on pronotum (both sides together) in g 17, 9 18; mesonotum in ¢
16, 14, in 9 26, 16; metanotum in ¢ 13, 15, in 2 21, 17. False subapical bristles
of mesonotum in g 12, in 2 14. Bristles on metepimerum in 3 2, 2 and 3, 3,
in © 3, 5 and 4, 6. On abdominal terga in g on II 19, 18; III 21,17; IV 18,
IGS WAVE NO, Ile sy © on UL 2, Bile IN a9, Ms IN, 87,183 WIL 25, iv | Om
sterna in don III 4,7; IV 3,7; V 3,7; VI 6,6; VII 4,8; in 9 on III 13,
15; IV 2,13; V 8, 11; VI 11, 11; VII 21, 23. In both sexes some minute
hairs in addition on sterna. Basal abdominal sternite with 21 to 23 small bristles
on the lateral surface.
On outer surface of hindfemur, not counting the subbasal ventral pair, in
g 15 on one femur and 20 on the other, on inside a row of 9; in 9 the bristles
82 2
more numerous. On outer surface of hindtibia 9 slender subdorsal bristles, a
larger number near and at ventral margin. The longest apical bristle of hind-
tarsal segment II reaches in both sexes to apex of IV or beyond ; longest of III
extending in 3 to second notch of V, in 9 not quite so far.
Modified Segments — 3. On sternum VIII an irregular posterior row of 12
bristles, farther forward about 22 bristles (on the two sides together). Posterior
margin of clasper (Cl) convex in ventral half, the notch (n) in both clasper and
digitoid subapical ; at the ventral margin near base of manubrium a larger
number of bristles than in P. longicornis. Apex of vertical arm of IX. st.
broader than in P. longicornis, and the ventral arm wider and more strongly
narrowed from middle to apex. Parameres very different : dorsal apical sclerite
not rounded at apex, but dorsally and ventrally angulate, the ventral angle more
or less sharp; second sclerite, which is irregularly elongate-elliptical in P.
longicornis with the apex round, is replaced by an acute hook which is proximally
very broad. ®. VII. st. truncate, the apical margin somewhat undulate,
Noviratres Zoonogican XLI. 1938. 137
without the narrow sinus present in P. longicornis and P. australiacus. Apical
margin of VIII. t. angulate below middle, concave above and below this rounded
lobe; in P. longicornis and P. australiacus the margin is deeply incurved sub-
ventrally, the ventral angle projecting as a rounded lobe. Head of spermatheca
(text-fig. 81) very slightly longer than broad, much more square than in P.
longicornis and P. australiacus.
Length: & 2:5 mm., 2 3-3 mm. ; hindfemur : 3 0-52 mm., ? 0:64 mm.
A second 9, smaller and paler (hindfemur 0-56 mm.) agrees with the above 9,
but the false setae of the mesonotum are absent except a dorsal one and the
eye-row of frons contains | or 2 more bristles. Fig. 82 is taken from this specimen.
Tergum VIII bears rather fewer marginal and submarginal bristles, and the
head of the spermatheca is shorter. The specimen suggests that the number of
false setae is very variable in this species; individuals with few pseudosetae
may even preponderate. A fair series of specimens will be required to settle
this point. The specimen cannot be a hybrid between P. longicormis and P.
magellanicus; apart from
the reduction of the row
of pseudosetae there is
nothing in the morphology
which points to inter-
crossing. The left stylet
of the specimen is not quite
normal; it is somewhat
reduced in size and its
base partly fused with
tergite IX.
3. Parapsyllus robertsianus
sp. nov. (text-figs. 83,
84).
Kidney Island: in a
burrow of Spheniscus
magellanicus and Procel-
laria aequinoctialis, 3 Dec.
1936 (Brian Roberts),
1 9.—tThe two birds, Mr.
Roberts informs me, often
have separate burrows
branching from the same
entrance. The true host
of this new species, which I have much pleasure in associating with the
discoverer’s name, is probably the Procellaria.
2. Frons with an anterior row of bristles, but this row less far away from
the eye than in P. magellanicus, the most ventral bristles of the row standing
some distance below the frontal tubercle, not at the side of it. The bristle above
the tubercle shorter than in the two previous species. Genal process behind eye
as short as in P. longicornis, but less rounded, the genal area ventrally enlarged
by a feebly chitinized expansion which contrasts with the strongly chitinized
stripe that bears the bristles. On occiput the bristles as in P. longicornis, the
138 NOVITATES ZoOLOGICAE XLI. 1938.
additional bristles present on the lateral surface in P. magellanicus being absent
in P. robertsianus. DBristles on antennal segment II short. Bristles on
pronotum 18 ; mesonotum 59, 15 ; metanotum 31, 16, the lowest of the posterior
row close to metepisternum, on which there are 10 on one side, 8 on the other ;
metepimerum 4, 5. Pseudosetae on mesonotum 10. The number of anterior
bristles of mesonotum includes some bristles placed at the anterior ventral angle,
which are absent from the other species here dealt with.
Bristles on abdominal terga II 40, 19; III 35, 19; IV 23,15; VII 19, 9;
on sterna III 32, 15; IV ? 10,15; IV 9,13; VI 17,13; VIL 15, 15. On side
of basal sternum 14 one side, 17 the other.
On outer surface of hindfemur 10 bristles, on inner surface a row of 13 or
14. Mid- and hindtibiae with 7 instead of 6 dorsal notches bearing pairs of
K VIL st.
strong bristles (apical pair inclusive) ; on outer surface of hindtibia 7 or 8 sub-
dorsal bristles (inclusive of apical one), and a small number at and near the
ventral margin. Lateral bristles of segments I and II of hindtarsus somewhat
more numerous than in P. longicornis, the longest apical one of III reaching to
middle of V; the ventral surface of V studded with small bristles from third
lateral pair to apex in hindtarsus, in fore- and midtarsi these bristles more
numerous and the hairy area extending farther towards base. In left foretarsus
V has on one side 5 instead of the usual 4 lateral bristles. Midtarsal I a little
longer than II. Apical margin of sternite VII (text-fig. 84) strongly slanting,
subventrally rounded, no lateral sinus. On VIII. t. 10 bristles above stigma on
one side, 12 on the other ; from stigma down 12 (10 on other side), separated by
Novirates ZooLoGIcAE XLI. 1938. 139
a space bare of bristles from the subventral row, which extends along apical
margin and contains 12 (10) bristles, there being an additional long bristle on
the side some distance from apical angle ; upper and lower portions of apical
margin nearly forming a right angle; on inside of this triangular area 11 (9)
bristles. Head of spermatheca much larger than in the previous species, longer
than broad, ovate, but with the posterior surface slightly curved in ; tail shorter
than in the other species.
Length: 3-5 mm.; hindfemur 0-69 mm.
11
140 NOVITATES ZooLoGIcAE XLI. 1938.
ON SOME OLD-WORLD ANTHRIBIDAE.
By KARL JORDAN.
(With text-figures 85-88.)
1. Acorynus laenatus sp. nov.
©. Near A. coomani Jord. 1928 from Tonkin, but frons broader, dorsal carina of
pronotum angulate in centre, lateral black area of elytra much larger and grey
median ring of tibiae narrower.
Brownish black. Head and rostrum creamy buff, on occiput a pair of
elongate brown spots extending on to frons, on cheek a diffuse brown spot ;
rostrum flat in middle, with two carinae each side in basal half, edge of antennal
groove cariniform, longer than in A. coomanz, directed towards side of prosternum
(in A. coomani towards underside). Frons a little less than half the width of
apex of rostrum (5:12). Segment IIT of antenna longer than any other, one-
half longer than IV.
Creamy median area of pronotum densely pubescent, irregular at sides, but
rather sharply defined, occupying not quite one-third of the surface, narrowed
to a spot in front of scutellum, within it a pair of blackish dots at apex, close
together, and another pair a short distance from carina farther apart, at base of
a short median sideward projection another dot; within the blackish lateral
area a grey median dot and some diffuse grey pubescence more or less condensed
at lateral carina as well as in front of grey dot and as an irregular stripe be-
tween dot and median area; sides punctate; carina somewhat convex, but
medianly distinctly angulate, lateral angle a little over 90°, less rounded than
in A. coomani ; basal longitudinal carinula distinct, horizontal.
Elytra finely punctate-striate, for the greater part densely creamy, shoulder
angle and a large lateral area black, this area extending from behind shoulder to
base of abdominal sternum ITI, slightly narrowing from margin to stripe V, then
abruptly to stripe III, this upper portion somewhat rounded; on subbasal
callosity an irregular transverse line of 5 dots, behind black area from near
suture to near margin 5 dots and on apical declivous area an indication of an
irregular brown band.
Pygidium and underside creamy grey, metepisternum with diffuse brown
stripe. Grey median ring of tibiae occupying less than one-third of upperside.
Burma: Yanaungmyin, Pyinmana, |.ix.1937 (R. Hla Ogh), 1 9 ex Albizzia
lebbek.
In colouring rather closely resembling the subspecies of Hucus bicolor here-
after described.
The species is not a true Acorynus, and might with as much justification be
placed in Tropideres. I have not yet arrived at a satisfactory classification of
the numerous, described and still undescribed, species allied to Acorynus,
Litocerus and Tropideres of Schönherr.
2. Hucus bicolor numatus subsp. nov.
39. Like H. b. bicolor Jord. 1937, from Assam ; but dorsal carina of pro-
notum medianly more convex. In the dark-brown lateral area of pronotum a
Novirates Zoonocican XLI. 1938. 141
distinct narrow grey line from apex to base, a little nearer to median grey area
than to lateral carina, or in other words the blackish lateral area divided by a
thin line into two vittae. Elytra: two subbasal dots on each, before middle a
round blotch on suture, at the side of which a dot contiguous with the blotch and
either free laterally or connected with the upward extension of the blackish
lateral area, on apical fourth three small dots, two of them before apical declivity
and one half-way to apical margin.
A little larger than the only known specimen of H. b. bicolor.
Burma: Kahilu Residency, Salween, 13.iv.1932, ex Croton oblongifolius,
one pair, type 9, the antennae of 3 being missing except segment I (Mg. San
Thwin).
3. Litocerus timius spec. nov.
oe. Nearest to L. planirostris Jord. 1894 (French Congo, Rio Muni, Came-
roon), but carina of pronotum with acute lateral angle as in Hucus Pasc. 1859.
Rostrum without dorsal carinae. Blackish brown, upperside with creamy grey
vittae.
Rostrum coarsely rugate-punctate, narrowest in middle, where the antennae
are inserted, a little more than the apical half nearly a transverse oblong, apical
margin with a very small median sinus, upperside thinly pubescent except the
lateral area from antenna to eye, which is densely greyish white, the rufescent,
brown median area narrowing backwards. Frons somewhat broader than
segment II of antenna is long. Eye margined with creamy grey, oblique, large
its outline straightened anteriorly-laterally. Occiput with thin creamy grey
median stripe. Antennae rufescent, paler at base, reaching to (2) or beyond
(8) base of elytra; segment IX in 3 somewhat longer than IV and as long
as XI, X shorter, not quite twice as long as broad, in 9 IX as long as IV,
very little longer than X and shorter than XI. No lateral groove in front
of eye.
Pronotum minutely coriaceous, widest at base, here one-half broader than
at apex, sides straight, a little over one-half as long as base is broad, with 5
creamy grey vittae, lateral one narrowest, separated from grey underside from
apex of carina forward by a brown stripe, sublateral vitta broadest, connected
posteriorly with lateral vitta ; dorsal carina convex, slightly undulate, distant
from base in middle, lateral angle sharply marked, acute, lateral carina
straight.
Elytra widest close to base, first gradually and then more strongly rounded-
narrowed, base depressed, but behind subbasal swelling a mere indication of a
depression, punctate stripes distinct, but not conspicuous, each elytrum with
three creamy grey vittae, first and second united about middle and continued in
interspace III as a single vitta to sutural angle, third vitta from base across
shoulder (which is bare) to apex, where it joins the dorsal vitta, in middle
a short branch forward to lateral margin above epimerum of metasternum,
this small branch sometimes a thin short isolated marginal line. Pygidium
creamy grey.
Underside greyish white, a lateral stripe from prothorax to end of abdomen
very densely pubescent. Legs rufescent.
Length : 5-0-6-5 mm.
Congo Belge: Kala, iii. and iv.1936 (J. Ghesquiere), 1 3 (type), 2 99.——
Type in Musée du Congo Belge.
142 NOVITATES ZOOLOGICAE XLI. 1938.
4. Nessiara vatia sp. nov.
32. Similar to N. mosonica Jord. 1904 and N. cethis Jord. 1911 ; pronotum
very densely and minutely coriaceous, not punctate, subbasal swellings of elytra
higher, hindtibia of 3 curved.
Head and rostrum punctate-reticulate, most of the meshes longitudinal,
small on occiput. Relative length of rostrum in & 30, width 55, width of frons
22; in 2 length of rostrum 30, width 50, width of frons 25 ; in centre of rostrum
a rudimentary carina, flattened, reaching neither apex nor base, more distinct
in 2 than in d.
Minute ridges of pronotal surface transverse, close together; markings
buffish grey, five spots at apical margin, middle one largest, reaching half-way to
carina, about twice as long as broad, across middle seven spots, namely a patch
at lateral carina and each side of disc three confluent spots, the middle one of
the three projecting backwards, in front of carina a sublateral spot and medianly
three spots connected with the basal median spot, this sublateral spot joined to
the middle one of the subcentral cluster of three and the lateral one of the ante-
carinal cluster of three joined to the dorsal spot of the subcentral cluster ; all the
spots well defined, no blackish spots.
Elytra tessellated, the russet spots small on suture, in interstice III a longish
russet median dash before and behind which a buffish-grey dash of about the
same length as the russet one, spot larger in V than at suture, between shoulder
and middle a black marginal streak bearing a rounded buffish-grey spot in middle ;
subbasal swelling higher in ¢ than in 9, prominent in both sexes.
Outside of femora black from base to middle (foreleg), or to two-thirds
(mid- and hindlegs), midfemur blackish also on inner surface. Hindtibia of &
distinctly curved, the apex gradually turned downwards as well as body-
wards.
Length: ¢ 5:7, 2 5-0; width: & 2-6, 9 2:2 mm.
Fast Java: Samberbajem, Res. Kediri, one pair (C. J. Louwerens).
5. Sintor molops sp. nov. (text-fig. 85).
9. Near 8. zamboangae Heller 1918; rostrum shorter, elytra more convex
and each with a black-brown streak from interspace III to near apex of VII and
more or less far continued forward in III. In the colouring of the elytra the
species is similar to S. suturalis Jord. 1895, but in that species the grey median line
is broader in middle and is accompanied by some blackish spots, and the elytrum
has an additional blackish mark before the apex, moreover the club of the antenna
is narrower in S. suturalis than in the new species.
Upperside grey-buff with black-brown stripes, underside whitish grey.
Rostrum one-ninth longer than apically broad ; from close to apical margin to
near base a median carina, base with median groove which extends on to head,
between carina and sides the surface depressed, less so than in S. zamboangae.
Head with broad brown stripe divided by a thin grey-buff median line, the brown
stripe continued on to rostrum, where it is narrow, non-divided, and ill-defined.
Antenna pale testaceous, club brown or brownish, segment IX triangular, as long
as broad, a little narrower than X and XI, X transverse, ? broader than long,
XI short-ovate, „'; longer than broad, obtuse, widest near base, proportional
length of the three segments of club: 10, 7, 13 (text-fig. 85). Blackish brown
median stripe of pronotum broader than buff-grey sublateral stripe and divided
Novirates Zoonoarcan NLI. 1938. 143
by a thin buff-grey median line, black or blackish lateral stripe extended on to
underside.
Pubescence of elytra in lines of punctures and an elongate basal sutural
spot whitish grey, interstices I and III blackish brown, V and VII with dots
of the same colour, those of V more or less confluent in type specimen, the
black-brown post-
median oblique streak
somewhat variable in
width, margin black
from base to middle,
the streak followed by
a row of dots, which
are confluent in one
ofthe paratypes.
Pygidium unicolorous,
buff-grey. Legs rufes-
cent grey, apical row of
spines oftibiaeblackish.
Length: 5-3-5-7; 85
width: 2-5-2-7 mm.
East Java : Tjomanoek, Res. Kediri, 1 9, type; Besoeki, Res. Kediri, 1 9;
Bredjo, Zuider Geb., 1 2 (C. J. Louwerens).
87 88
6. Gulamentus signatus ealanus subsp. nov.
3. Differs in the extent of the grey pubescence of the upperside. Rostrum
densely grey as in G. s. signatus Jord. 1923, from Elisabethville, but the posterior
margin of this area more straight, the area somewhat narrower. On pronotum
the central brown area much reduced by the appearance of grey pubescence, the
lateral area on the other hand less densely grey. Scutellum grey, sometimes
brownish at side. The brown median area of elytra reduced to a smallish patch
on each elytrum, grey pubescence of lateral half or two-thirds arranged in stripes
which are composed of spots and short lines, sometimes one or two stripes nearly
uninterrupted.
Congo Belge: Hala, iii. and iv. 1936 (J. Ghesquiére),3 gg, 2 22, type (3)
in Musée du Congo Belge.
7. Gulamentus depictus sp. nov.
6. Elytra distinctly flattened and punctate-striate ; pygidium triangular,
its apex rounded and strongly convex ; anal sternum without groove.
A little over twice as long as broad (pygidium inclusive). Derm black,
pubescence white, grey and black, with a shade of cinnamon on elytra. Head
white, frons and an equally broad area on occiput brownish black, separated from
eyes by some diffuse white pubescence, in centre of frons a short white line.
Space between antennal groove and eye more than twice as wide as segment IT
of antenna. Eye ovate. Frons a little less than half as wide as base of rostrum.
Antenna (3) black, reaching to basal fourth of elytra, apex of end-segment pale
testaceous ; segment II a little shorter and much thinner than I, III a little
longer than II, III to VII about equal in length, VIII slightly shorter, but of
the same shape, club long, flat, nearly as long as III to VI together, IX triangular,
144 Novirates ZOOLOGICAE XLI. 1938.
about one-third longer than broad, X a little broader than long, XI about one-
half longer than broad.
Pronotum rather more than one-third broader than long, whitish grey, a
central area as broad as dark area of occiput blackish, not sharply defined, shaded
with grey and bearing two short lines indicating a grey median line; dorsal
carina convex half-way between middle and lateral angle. Scutellum black,
subvertical in front, its apical (dorsal) area nearly semicircular.
Elytra flattened in middle, with distinct stripes of punctures except in
convex basal-sutural area; grey, suture and alternate interstices dotted with
black, a black lateral stripe from line VIII of punctures to margin commencing
abruptly at basal fourth and running to apical fourth, where it turns obliquely
dorsad and broadens, terminating in interspace III ; pubescence adjacent to this
black mark dorsally and posteriorly at suture more or less cinnamomeous, a lateral
patch behind the black mark white. Pygidium one-fifth broader than long,
triangular, apex rounded, the reflexed margin faintly straightened in centre,
carinae flanking basal median groove not extending half-way to apex, apical half
of pygidium convex, highest in front of apical margin, between this margin and
the hump an indication of a groove.
Underside greyish white, centrally less densely pubescent than laterally ;
abdominal segment I with black streak at lateral margin, on II to IV a minute
lateral basal dot, centre of V black (denuded), apex of foretibia, entire foretarsus
and segments II to IV of mid- and hindtarsi black. Antecoxal area of pro-
sternum longitudinally about half as wide as forecoxa. Apex of abdominal
segment V slightly incurved ; the segment medianly somewhat flattened.
Length : 9-2 mm.
Belgian Congo: Buta, 1 ¢ (at Tring).
Key to the species of Gulamentus Jord. 1895, genotype; G. cylindricus
Jord. 1895.
When both sexes are known of all the species, the classification of the genus
will probably be based to a great extent on the shape of the pygidium, which
often differs much in the species and in the sexes. Another interesting point
in the morphology of Gulamentus is the total disappearance in the majority of
species of the stripes of punctures usually present on the elytra of Anthribidae.
In some species the stripes are well developed and in some others vestigial, and
sometimes the stripes are traceable in a few specimens of a species, but not in
the others. Knowing nothing of the habits of the species, we can make no guess
at the reason of the loss of the stripes. The enlarged hindfemora and short
hindtibiae of several species also may have been acquired in connection with
specialized habits.
1. Lateral carina of pronotum not reaching to apical margin. Space between
eye and antennal scrobe about as wide as segment II of antenna is long
Pygidium carinate in apical half, the carina prominent in ¢ and ending in
a projection, low in 2 ; G. albinasus Jord. 1922; Madagascar.
Lateral carina complete : : 2.
2. Lateral carina ending with a onen aerate paper Prosternum in
front of coxa much longer than the coxa is wide. Segment II of antenna
short, stout, less than one-half longer than broad.
G. major Jord. 1922 ; Congo.
Novirates ZooLoGIcAaE XLI. 1938. 145
10.
11.
. Eye broadly margined with creamy buff.
Lateral carina not forming a prominent tubercle at anterior margin of pro-
notum. Prosternum in front of coxa shorter than the coxa is wide. 3.
. Serobes of antennae a little nearer together than are the eyes posteriorly 4.
Scrobes of antennae farther apart than are the eyes posteriorly . : 5.
. White, with black spots. Pygidium of 9 rounded (4 not known).
G. laetus Jord. 1922 ; Natal.
Cinnamomeous, with blackish and greyish dots. Apex of pygidium subtrun-
cate in g, rounded in 9.
G. pardalis Jord. 1901 ; Cameroon, French Congo.
. Pronotum with well-defined vittae. Apex of ae gidium subtruncate in 3,
roundedin? . , 5 , 5 6.
Pronotum without ikea ronan apex ko Ben : : i ; 5 To
G. taeniatus, Jord. 1924 ; Cameroon.
Eye not margined with creamy buff on frontal side, creamy grey rostrum
sharply contrasting with frons . G. virgatus Jord. 1930 ; Congo.
. Scutellum more than twice as broad as long (i.e. the free part not covered by
pronotum). Apex of pygidium rounded in 4, truncate-emarginate in 9.
Segment II of antenna shorter than I, XI longer and broader than
IX+xX. : ; ; F : ; 8.
Scutellum less than irre as Brandl. as Tore : : B 6 „ 10,
. Rostrum the same dark colour as frons and occiput.
G. melius Jord. 1922; Ogove R., Spanish Guinea, Congo.
Rostrum white, sharply contrasting with frons and occiput ; tarsi black.
G. lujai Jord. 1925 ; Congo.
Rostrum white ; tarsi pubescent grey . : : 9.
. Median area of roman blackish, with thin, dl Ankemayatradl, grey median
line ; on elytra a large grey basal patch more or less completely separate
from a smaller postmedian patch.
G. signatus signatus Jord. 1923 ; Katanga district, Upper Congo.
The grey pubescence of pronotum and elytra much extended ; median stripe
of pronotum complete, the grey patches of elytra joined together.
G. signatus ealanus Jord., p. 143 ; Lower Congo.
Eye rounded, sinuate, the sinus parallel with margin of scrobe. Segment II
of antenna longer than I. Pygidium of 3 acuminate, carinate, of 9
truncate-emarginate . : h alle
Eye oblique, ovate, the sinus if essen mer mare ith ede of scrobe 12.
Base of pronotum russet ; before middle of elytra a transverse wood-brown
band slightly shaded with russet, conspicuous.
G. fasciatus Jord. 1901 ; Spanish Guinea.
A cluster of subbasal lateral spots on pronotum, an antemedian spot near
suture and another at apex of elytra yellow.
G. tricolor Jord. 1914 ; Congo.
A yellow spot on shoulder extending on to pronotum and another at apex of
elytra 5 F : . G. alveatus Jord. 1922 ; Spanish Guinea.
Elytra olivaceous, “ dloiiied with black, across suture an ainkarmadkem black
patch Ä Ä G. guttatus Jord. 1930 ; Cameroon.
Elytra sepia colon, dotted end mottled with greyish white.
@. cylindricus Jord. 1895 ; Cameroon, Gaboon, Congo.
146 Novrrates ZOOLOGICAE XLI. 1938.
12. Space between eye and scrobe about as wide as segment II of antenna is
long . ; B Sls:
This interspace Shan: as W Hie, as Be IL is breed ; ‘ . 16.
13. Elytra flattened, with a sharply defined broad lateral stripe turning dorsad
before apical declivity. Rostrum white, contrasting with frons and
occiput. : : G. depictus Jord., p. 143 ; Congo.
Elytra convex, w ithout conspicuous black lateral stripe rostrum not
contrasting with frons and occiput : : h „14,
14. Segment XI of antenna longer than IX, VIII of 3 very little broader than
VII; eye slightly incurved. Apex of pygidium rounded, in 39 a vestige
of a median carina at apex.
@. brevis Jord. 1922 ; Mandated East Africa, Ukerewe.
Segment XI of antenna smaller than IX, VIII of 3 broad, forming part of
club ; scutellum with transverse ridge before apex. : F 5 lw,
15. Upperside of elytra more or less cinnamomeous shaded with grey. Pygidium
(8, the 2 not known) one-third broader than long, apical margin straight-
ened. : : e G. kohli Jord. 1914 ; Congo.
Upperside olivaceous, dorteds and mrouied with grey; club of antenna
somewhat narrower. eve one-fifth broader than long, apical margin
rounded . 5 : . G. tephrus Jord. 1922 ; Fernando Po.
16. Segment XI of antenna twice as broad or nearly.
G. nanus Jord. 1920 ; Fernando Po, Cameroon, Spanish, Guinea,
Gaboon, Congo.:
Segment XI of antenna little longer than broad 5 . : Blris
17. Elytra one-half longer than broad . G. picatus Jord. 1930 ; Cameroon.
Elytra at most one-third longer than broad.
G. sellatus Jord. 1901 ; Cameroon, Ogové R.
8. Mauia candida sp. nov.
2. In shape and size like M. subnotatus Bohem. 1859, but conspicuously
decorated with white.
Head and rostrum dark brown or clayish, with numerous white scales,
centre more or less brown. Antenna very pale buff, club a little darker, shaft
thinner than in M. subnotatus. Pronotum with basal and lateral borders of
white scales, basal border medianly widened into a patch, behind apical margin
also some white scales, which are most numerous in centre, derm varying from
buffish clay to blackish brown, surface somewhat smoother than in M. subnotatus,
angle of carina more rounded, and lateral carina longer, extending to # (about).
Scutellum black.
Elytra likewise black or brownish black, basal margin and a transverse
band behind subbasal swelling white, the band extending to impressed stripe VII,
and connected with basal border (or nearly) at suture, in interspace III and
above shoulder, behind shoulder a submarginal elongate spot and some additional
white marginal scales, behind middle another transverse band, somewhat variable,
not quite reaching to margin and usually interrupted at suture (scales fallen off ?),
before apex irregular white scaling. Pygidium somewhat longer and less rounded
than in M. subnotatus.
Noviratres ZOOLOGICAE XLI. 1938. 147
Underside and the very pale buff legs pubescent white, the hairs rather
broad, but thinner and longer than the scales of the upperside.
Length : 2-7; width: 1:2 mm.
East Java: Samberbajem, Res. Kediri ; 3 99 (C. J. Louwerens).
9. Misthosima subtuta sp. nov. (text-fig. 88).
SP. In colour like M. virilis Jord. 1928. Distinguished in the 3 by the
pygidium, foretibia and -tarsus and the anal sternum. The 92 of M. virilis is not
known and is possibly very close to that sex of the new species. The type of
M. virilis, from Tonkin, is a somewhat abraded specimen ; we have two good dd
from Sumatra (no 99) which agree in structure with the Tonkin specimens.
In these gg the brown median area of the pronotum is divided by a broadish
grey median line into two stripes each of which bears some grey pubescence
before middle, the stripes nearly being broken up into two patches each ; between
them and lateral carina an indication of another brown stripe. On elytra a large
blackish sutural patch, irregular ; this patch is represented in the type-specimen
by a diffuse blackish sutural area, the grey pubescence surrounding the patch
and making it very conspicuous lost in type of virilis.
In M. subtuta the pronotum bears a subapical and a subbasal transverse
row of four diffuse brown spots each, the spots of the same row partly united.
Black sutural patch of elytra as in M. virilis.
In & of M. subtuta the pygidium truncate, with the angles rounded, in M.
virils distinctly emarginate ; abdomen flattened from base to apex, the posterior
segments impressed, apex of V emarginate, foretibia very little humped at apex ;
very different from that of M. virilis. In the latter species the apex of the fore-
tibia distinctly enlarged on the underside and somewhat curved ; the underside
partly flattened longitudinally and bearing along the outside of this flattened
area a comb of short spear-head spines from base to near middle, the comb
continued at apex in a curve towards the inner side (text-fig. 86, sketched from
inner side, text-fig. 87 from underside; diagrammatical). In M. subtuta the
comb is absent except for a few short obtuse spines at apex (text-fig. 88, under-
side). Segment I of foretarsus of g in M. subtuta shorter and in M. virilis
longer than II to IV.
In @ the pygidium triangular, about one-fifth broader than long, sides
slightly rounded, apex pointed, but the tip of the projection rounded off ; segment
I of foretarsus nearly as long as II to IV.
East Java: Tjomanoek and Pelitoeran, Res. Kediri, 1 g, 3 22 (C. J.
Louwerens).
M. angulata Wolfr. 1934, from Mindanao, which I know only from the
description of the 9, is evidently very close to M. subtuta.
Text-fig. 85.—Sintor molops, p. 142.
86, 87.—Misthosima virilis, p. 147.
88.—Misthosima subtuta, p. 147.
148 NoviratEs ZOOLOGICAE XLI. 1938.
THE EXCRETORY SYSTEM OF CERCARIA CORONANDA!N. SP.
TOGETHER WITH NOTES ON ITS LIFE-HISTORY AND
THE CLASSIFICATION OF CERCARIAE OF THE SUPER-
FAMILY OPISTHORCHIOIDEA VOGEL 1934 (TREMATODA).
By MIRIAM ROTHSCHILD.
(With text-figures 89-107.)
A NUMBER of larval Trematodes pertaining to the Pleurolophocerca and
Parapleurolophocerca Groups (Sewell 1922) are known to parasitize Peringia
ulvae (Pennant) 1777 (Rothschild 1938). These cercariae develop into adult
worms of the superfamily OPISTHORCHIOIDEA Vogel 1934.
One of the species from P. ulvae, discovered unfortunately on only two
occasions, attracted special attention, as all the important morphological details,
in particular the excretory system, could be distinguished without the slightest
difficulty.
This cercaria is a typical Heterophyid larva, and despite the aberrant
arrangement of the collecting tubes and flame-cell pattern it is impossible to
doubt its relationship with other Opisthorchioidea cercariae. It will be seen
from the description given below that in addition to all the recognized characters
of the group it also agrees in a number of apparently minor morphological
details with Pleurolophocerca cercariae.
C. coronanda develops into a metacercaria pertaining to the subfamily
Neochasminae Van Cleave and Mueller 1932, flukes which inhabit the
digestive tract of fish.
DESCRIPTION OF THE EXCRETORY SYSTEM (figs. 89, 90, 93-95).
The excretory bladder of C. coronanda is typical and resembles that of the
Cercaria of Oryptocotyle lingua (Creplin) 1825, the Cercaria of Opisthorchis felineus
(Riv.) 1884, the Cercaria of Clonorchis sinensis (Cobbold) 1875, ete. Owing to
the deeper caudal pocket of C. coronanda, the form of the bladder is almost
butterfly-shaped when the body is contracted. The walls are very thick and the
giant nuclei of the cells particularly noticeable in this species, even in living
specimens.
There is a short caudal excretory tube, extending about one-eighth the length
of the tail. It terminates in a fork, each branch opening to the exterior on the
lateral edges of the tail.
The main collecting tubes leave the bladder at the anterior cornu, and after
a short convoluted course divide into an antero- and postero-lateral branch
below the level of the rudimentary ventral sucker. Each antero-lateral tube
proceeds forward, parallel to the sides of the body, and receives the canaliculi of
two flame-cells situated on a level with the penetration glands. They terminate
in a second pair of flame-cells in the region of the eye-spots and oral sucker.
The postero-lateral collecting tubes pass backwards over the corners of the
! I would like to thank the Master of the Rolls (the Rt. Hon. Sir Wilfrid Greene) for suggesting
this apt and attractive name.
149
Novirates ZOOLOGICAE XLI. 1938.
bladder (fig. 90) (the only point where they are rather diffieult to follow) and each
receives the canaliculi of two flame-cells situated in this region. They terminate
-Fia. 89.—Cercaria coronanda, showing flame-cell pattern (x 257).
Fic. 90.—Portion of the excretory system, showing where the postero-
lateral collecting tube passes over the corner of the bladder.
Fie. 91.—Tail of C. coronanda (lateral view), showing dorso-ventral fin
membrane and anterior cuticular expansions (x 233).
Fie. 92.—C. coronanda ventral view (tail contracted x 154).
(For explanation of lettering see p. 161.)
in a second pair of flame cells in the posterior end of the body. The flame-cell
pattern is thus : AD 4 Dy ©. | = 16,
The development of the excretory system is shown in figs. 93 and 94. In
150 Novrrares ZooLoGIcAE XLI. 1938.
the earliest stage observed, the 4-flame-cell stage, the excretory tubes are quite
separate. It is in this stage that the division into postero-lateral and antero-
lateral collecting tubes (fig. 95) can be most clearly seen. Later on the junction
is sometimes obscured by the convolutions of the short main collecting tube.
md
Fıcs. 93, 94.—Development of excretory system of C. coronanda.
Fic. 95.—Main division of the lateral collecting tube at 8-flame cell stage.
Fie. 964.—Ventral sucker of cercaria (before it leaves the redia), showing giant
nuclei.
Fie. 968.—Papilla protruding from the centre of the developing ventral sucker in
lateral aspect.
Fic. 97.—Diagram showing the apertures of the penetration gland ducts, and the
piercing spines in the mouth of the anterior sucker.
Just before a constriction between body and tail is formed the collecting tubes
meet and fuse in the region of the future bladder. This is probably at the 8-flame-
cell stage. From then onwards no great change occurs and the excretory system
gradually assumes the form described above.
In the metacercarial stage a striking development takes place in the bladder.
This grows forward until it almost reaches the level of the ventral sucker where
it divides into two branches each extending to the oesophageal region, thus
Novirarrs Zoonoarcan XLI. 1938. 15]
forming a large Y. Owing to the scarcity of material the flame-cell pattern was
not worked out in detail at this stage, but from a cursory examination no change
appeared to have taken place.
98
Fic. 98.—‘‘ Death attitude’ of Pleurolophocerca and Parapleurolophocerca cercari».
(a) Cercaria coronanda (6) ©. of Oryptocotyle jeyuna
(c) Cercaria sp. (d) C. of Cryptocotyle lingua
(all drawn to the same scale x 175)
Discussion ON THE EXCRETORY SYSTEM.
The type of excretory system found in (©. coronanda is very unexpected.
The Y-shaped vesicle of the metacercaria, extending into the anterior end
of the body, is described for adults of the Neochasminae, and certain other
Heterophyid flukes from fish, together with the Metacercaria of Cercaria flori-
152 NOVITATES ZOOLOGICAE XLI. 1938.
densis McCoy 1929, but it is the only known example of any member of the
OPISTHORCHIOIDEA in which the main division of the collecting tubes occurs
below the level of the ventral sucker. In general features, such as the arrange-
ment of the tubules and flame-cell pattern, it resembles the excretory system of
Heterophyes heterophyes (Siebold) 1852 (fig. 99) more closely than any of the other
known forms, such as Cryptocotyle lingua (fig. 102), Rossicotrema donicum Skrjabin
1919 (fig. 101), Opisthorchis felineus (fig. 100), ete.
In both Heterophyes heterophyes and C. coronanda the antero- and postero-
lateral collecting tubes receive an equal number of flame cells, although in the
former species there is already a manifestation of asymmetry, and an elongation
of the postero-lateral portion.
Stunkard (1929) pointed out that the flame-cell pattern of certain genera
of the Heterophyidae Odhner 1914 were strikingly different, and concluded that
too much importance must not be attached to this character for purposes of
classification. Although several of the examples he used as illustrations can no
longer be included in this family or even the superfamily OPISTHORCHIOIDEA, the
genera Heterophyes and Cryptocotyle alone are quite sufficient to prove the point.
La Rue (1938) commenting upon this says: “‘ Stunkard’s words of caution .. .
should be heeded in all future work.” The excretory system of C. coronanda
more than justifies this point of view.
A much harder task will be to decide which of these systems is the more
primitive, and how they are derived. When further material of this family
is available for study and comparison it may be possible to revise the whole
question of the evolution of Mesostoma and Stenostoma types. For the moment
there is little to be gained from a purely speculative discussion.
CHARACTERS OF THE PLEUROLOPHOCERCA AND PARAPLEUROLOPHOCERCA GROUPS
OF CERCARIAE.!
Additional knowledge concerning the morphology and life-histories of the
superfamily OPISTHORCHIOIDEA necessitates a revision of Sewell’s (1922) and
Vogel’s (1934) definitions of these groups.
A modified list of characters is given below.
(1) A pair of pigmented eye-spots is generally present, but may be lacking
altogether.
(2) The anterior end of the body, or the whole body, is armed with backwardly
directed spines.
(3) The anterior sucker is modified to form a protrusible organ of penetration.
(4) The acetabulum may be absent or only poorly developed.
(5) A pharynx is always present, but the alimentary canal is absent or only
poorly developed.
(6) Penetration glands, with ducts opening anteriorly, are present.
(7) The bladder is reniform or roughly globular, with thick walls. The
lateral excretory tubes enter the bladder antero-laterally.
(8) The tail is longer than the body, and powerful, and is provided with
cuticular fin-folds or a cuticular sheath.
1 These two groups grade into one another, and at present it is thought there are insufficient
grounds for separating them. A more complete knowledge of the excretory systems and life-
histories of the Parapleurolophocerca cercariae and their adult forms (flukes pertaining to the
subfamily Haplorchinae), may eventually make a satisfactory subdivision possible.
1938,
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154 Novirates ZooLoGIcAE XLI. 1938.
(9) The cercariae are phototropic. They swim by short dashes, followed in
each case by a pause, during which the body hangs below the tail in the water.
(10) Development occurs in rediae with an intestine and birth pore but no
locomotor appendages. The cercariae leave the rediae at an early stage and
continue development within the tissues of the first intermediate host.
(11) First intermediate host : fresh water, brackish water or marine gastro-
pod. Second intermediate host: generally a fish, but exceptionally an amphi-
bian. Encystment may occur in the first intermediate host.
THE SPECIFIC CHARACTERS OF PLEUROLOPHOCERCA AND PARAPLEUROLOPHOCERCA
CERCARIAE.
Before describing ©. coronanda and its metacercaria it is advisable to con-
sider the specific characters of these larvae. I have frequently drawn attention
to the difficulty of distinguishing between closely related species. This fact is
forcibly brought home to anyone who happens to discover a host infected with
several cercariae pertaining to the same group, but it is apt to be otherwise
overlooked. It thus happens that scarcely any descriptions of Opisthorchioidea
cercariae exist which can be termed specific.
Measurements, although important for supplementing descriptions, cannot
be relied upon for a variety of reasons. The complete excretory system of a
Pleurolophocerca or Parapleurolophocerca cercaria has only once before been
worked out, and is, generally speaking, too difficult a matter to serve as a practical
diagnostic character. The number of penetration glands varies and may prove
extremely useful, as in the case of C. coronanda, but on the other hand many of
these larvae (see Table I) have the same complement of glands arranged in the
same manner. The spination, development of the ventral sucker and intestine,
transparency of certain organs, etc., are all “ specific” characters, but the
differences they display are questions of degree rather than kind, and they do
not lend themselves well to a precise written description.
The characters which have so far proved most useful in distinguishing closely
allied species are :
(1) The shape and size of the body under a coverslip at death.
(2) The precise extent, position and shape of the caudal fin-folds.
(3) The pigmentation of the body.
(4) The behaviour and length of life of the cercaria.
The “ death attitude ” (fig. 98) is obtained by making a series of camera-
lucida outline drawings of the body immediately bubbles begin to break through
the cuticle. It is important not to make the drawing when the cercaria merely
becomes quiescent. Owing to the varying degree of development of the ventral
sucker some species stop moving about under a coverslip long before they have
become fully and characteristically extended in a “ death attitude.”’
The variation in the fin-folds and cuticular expansions of the tail, and the
method of studying them are described elsewhere. The precise outline, length
and position of each fin is of great importance, together with the degree of
folding and fluting it displays.
The size of the pigment granules and their distribution should be noted,
particularly in the eye-spots and on the dorsal surface of the body.
The behaviour and length of life of the cercariae will possible prove the most
NOVITATES ZOOLOGICAE XLI. 1938.
Species of Cercarlae.
A. Examined by the author.
Cercaria of Cr yptocotyle lingua (Creplin)
C. of Cryptocotyle jeyjuna Nicoll 0
C. of Oryptocotyle sp.
C. of Haplorchid fluke
Cerceria coronanda ;
Cercaria sp. (from P. ulvae)
Cercaria sp. (from Littorina littoralis)
B. From various authors.
. of Metagonimus yokogawai (Katsurada)
. of Metagonimus sp. Takahashi .
. of Apophallus venustus (Ransom)
. of Haplorchis taihokui F. and N.
. of Haplorchis taichui F. and N.
. of Stamnosoma armatum ? (Tanabe) .
. of Pygidiopsis summus Nishio .
Cercaria lophocerca Fil.
Cercaria floridensis McCoy
Cercaria vogeli Cable!
Cercaria indicae XXXI Sewell
Cercaria indicae L Sewell
Cercaria indicae III Sewell .
Cercaria indicae VIII Sewell
Cercaria indicae VII Sewell
Cercaria translucens Faust .
C. of Opisthorchis felineus (Riv.)
C. of Metorchis intermedius Heinemann
C. of Clonorchis sinensis ? (Cobbold)
Cercaria nicobarica I Sewell . .
Cercaria pleurophocerca Sons.
Cercaria parvomelaniae Tubangui .
QAQ2QAQR2QaQ
Total Number of
Penetration Glands.
Number of Ducts
per Bundle.
4, 5, 6, 4
3, 4, 4, 3
3, 4, 4,3
3, 4, 4, 3
4, 6, 6, 4
linear arrangement
4, 5, 5, 4
er
3, 4, 4,3
3, 4, 4, 3
4,4, 4,4
linear arrangement
linear arrangement
useful characters of all. The manner of swimming is common to the whole group.
Short periods of activity are followed by short passive periods when the cercaria
hangs in the water with the body below the tail.
These larvae are also all photo-
tropic. The differences observed fall under the following headings :
The site of encystment.
) Length of free-swimming life (Table II).
) Relative length of passive and active periods of swimming.
3) The nature and degree? of the phototropic response.
) The intermediate host (or hosts) selected.
)
)
The behaviour of the tail after it has been cast.
7) The stage of development of the cercaria when it leaves the redia.
1 At the time of sending this paper to press the author had not seen Cable’s redescription of
Cercaria vogeli (‘‘ Studies on larval trematodes from Kentucky with a summary of known related
species.” —Amer. Midl. Nat., 19, No. 2, pp. 440-464, 1938), in which this larva is stated to possess
14 penetration glands, not 12, the ducts of which is are arranged in bundles of 3, 4, 4, 3.
2 An attempt was made to compare the degree of phototropism in a simple manner, the
detailed results of which will be published in the writer’s fuller paper on the Trematode parasites of
P.ulvae. A dish was filled with 4 inch of water and 50 cercariae introduced at one end. A 60-watt
lamp was placed at the opposite end of the dish (if the cercariae were positively phototropic) and
the time recorded for (a) the first cercaria, (b) the majority (40) of the cercariae, (c) the entire
number of cercariae to reach the opposite end of the dish. The larvae were inserted at the lighted
end if they were negatively phototropic.
Sufficient members of C. coronanda were not available
for this experiment, but a single specimen approximated}ito the time taken by the Cercaria of
C. lingua—i.e. about 4 a minute to traverse a dish 5 inches wide.
12
156 Novirates Zoonocicar XLI. 1938.
HALB II.
Approximate | | Average Length | Average Length
Length of Reaction to of Active of Passive
Free-swimming Light. Period of Sinking
Life. Swimming. Period.
k |_ abe Bee is
Cercaria of C. lingua o || 42 hours positively | 24 seconds | 15} seconds
| phototropic
Cercaria of C. jejuna | 8 hours positively | — | —
| | phototropic |
Cercaria of Cryptocotyle sp. 3 | 8hours | negatively | 7 seconds 8 seconds
phototropic | |
Cercaria of Haplorchid fluke. 44 hours negatively 13 seconds 14 seconds
phototropic
Cercaria coronanda . . : 52 hours | positively 2; seconds | 20 seconds
| | phototropic
| I |
EN 60- watt eleetrie ehe was ited en omg this, table!
It is of interest to find that in the Opisthorchioidea cercariae from P. ulvae,
Littorina littorea (L.) 1758, and L. littoralis (L.) 1758, the length of life is longest
in those species with the longest passive phases relative to the active swimming
phases. It is remarkable that those species with a short active phase are both
more strongly phototropic and less specific with regard to the choice of the
second intermediate host.
Cercaria coronanda n. sp. (figs. 89-97 and 98a).
Measurements in Microns.
Body (length) ! ; . 355-225. Average: 286
Body wide 000857 By 143
Tail (length) : . 565-404 +, 458
Tail (width, without ag) . 24-40 a 32
Oral sucker . : : . 52 x 57-52 x 45. Average: 52 x 51
Pharynx . ; . 19 x 26-17 x 19
Bladder Kernen : > Bal
Specific Diagnosis.—Opisthorchioidea cercaria with body more rounded (when
alive) than is usually the case. Almost heart-shaped in death attitude (fig. 98).
Broadest about level of ventral sucker, width decreasing on either side of this
point. Length approximately twice maximum breadth. Tail (fig. 91) nearly
twice the length of body, inserted in very deep caudal pocket which is generally
deeper than width of tail (without fins) at its widest point. Dorso-ventral fin-
fold and anterior cuticular expansions present. Dorsal fin extending to the level
of the lateral anterior cuticular expansions, widest ;°- from the anterior end of
tail, where it is usually 4 wider than tail at its greatest width (uncontracted).
Ventral fin extending a little over 4 the length of the tail, widest about 4 from
the tip of the tail. At its widest point nearly 4 the width of dorsal fin at its
maximum. Dorsal fin with undulations (folds under a coverslip) along its whole
length. Ventral fin with distinct folds at the posterior tip only. Anterior
cuticular expansions visible laterally for about +, and ventrally for 4 the length
of the tail. Colour of body uniformly yellowish white. No individual pigment
granules discernible even under -!; objective. Eye-spots roughly rounded, not
markedly rhomboid or square,
Novirates Zoonoarcar XLT. 1938. 157
Cuticle with long fine sensory spines, at some distance from each other but
small spines very poorly developed. Anterior organ of penetration with 4 acicular
boring spines in the dorsal lip and an accessory row of 6 finer spines behind these.
Pharynx situated posterior to level of eye-spots, no matter what state of contrac-
tion the body may be in.
Twenty lobed, granular penetration glands, with ducts arranged in 4 bundles.
Lateral bundles with 4, median bundles with 6 ducts each. Ventral sucker
rudimentary ; lip composed of six cells with giant spherical nuclei A distinct
papilla projects from centre of sucker during development. Sucker very much
more distinct during development than at any other time (fig. 96).
Walls of bladder composed of giant cells with large spherical nuclei. Main
collecting tubes divide below ventral sucker into antero- and postero-lateral
branches (mesostoma type). Flame-cell pattern: 2 [(2 + 2) + (2 + 2)] = 16.-
Short caudal excretory canal extending 4 length of tail, opening laterally by
two pores.
Behaviour.—Cercaria positively phototropic. Stimulated to activity by a
sudden decrease in hght but upward movement in response to a shadow not very
pronounced. Average length of passive period 9 times length of active period
(Table II). Duration of free-swimming life approximately 52 hours. Tail not
continuously active after detachment from body. Tip curled upwards (fig. 92)
when it comes to rest. Encystment occurs in muscles of second intermediate
host. Cercaria leaves redia to continue development in tissues of host at very
early stage.
Redia (fig. 107) colourless with brown intestine, about ~, the length of
the body. Maximum-sized specimens 688 x 182, with pharynx 28u x 22u.
Fertility low. Only one developing cercaria with eye-spots in redia at same
time. Site of infection, liver and gonads of snail.
Host: Peringia ulvae (Pennant) 1777. Two infections found in several
thousand snails collected on the saltings of the River Tamar, Plymouth, England,
in June 1937.
COMPARISON WITH OTHER RELATED SPECIES.
At present there is no difficulty in separating C. coronanda from all other
known Pleurolophocerca and Parapleurolophocerca cercariae, as it is the only
species, apart from the Cercaria of Opisthorchis felineus, with 20 penetration
glands with the ducts arranged in four bundles. It is quite likely, however, that
other species of the same or related genera will be found to possess a similar
number and arrangement, and a closer comparison will then be required. The
only other cercaria with a fin-fold pertaining to these groups which is known
to develop a crown of spines in the metacercaria is C. floridensis. Table III
compares these three species.
LIFE-HISTORY EXPERIMENTS.
The majority of the Pleurolophocerca and Parapleurolophocerca cercariae
from P. ulvae will encyst in the Spotted Goby, Gobius ruthensparri (Euphras.)
1786. This species also serves as the second intermediate host of C. coronanda.
It is more than likely that a variety of fish are attacked in nature, but owing to
the scarcity of material it was impossible to experiment with any others.
1 A precisely similar condition was discovered in a species from Littorina littorea, and it is quite
possible that these six cells with giant nuclei are present in all Opisthorchioidea cercariae, but are
not easily detected in most species, The same applies to the caudal excretory tube.
. NOVITATES ZOOLOGICAE XLI.
TABLE III.
C. of Opisthorchis felineus.
Individual pigment granules on
dorsal surface.
Comparatively shallow caudal
pocket.
Pharynx conspicuous.
Twenty penetration glands.
Ducts arranged in bundles of
4, 6, 6, 4.
Flame-cell pattern :
2[(5) + (5 +5 +5 + 5)] = 50
Free-swimming life 40-53 hours.
?
Site of encystment principally
fins. Also skin, scales, muscles, |
etc.
Pigment cells formed round cysts.
Bladder does not extend beyond
ventral sucker in metacercaria.
No crown of spines develops
round oral sucker in metacer-
caria.
Eye-spots no longer visible in
fully developed metacercaria.
©. floridensis.
1938.
C. coronanda.
Individual pigment granules on
| dorsal surface.
2
No pharynx observed.
| Fourteen penetration glands.
Ducts arranged in bundles of
| 8.4524 8h
°
| Free-swimming life 36 hours.
| Active phase from about 0-6-1
sec. Passive phase from
about 0-9 sec.
Site of encystment principally
the fins and beneath the skin.
No pigment cells formed round
cysts.
Bladder extends into anterior
region of body in metacer-
caria.
Crown of 27 spines develops
round oral sucker in metacer-
caria.
Eye-spots not retained in full- |
grown metacercaria.
No individual pigment gran-
ules on dorsal surface.
| Deep caudal pocket.
Pharynx conspicuous.
Twenty penetration glands.
Ducts arranged in bundles of
4, 6, 6, 4.
Flame-cell pattern :
2[(2 + 2) + (2+ 2)] = 16
| Free-swimming life 50-52 hours.
| Active phase (average) 2-2 sec.
Passive phase (average) 20
sec.
No cysts formed in fins or
under skin, but in muscles.
No pigment cells formed round
cysts.
Bladder extends into anterior
region of body in metacer-
caria.
Crown of 15 spines develops
round oral sucker in meta-
cercaria.
Eye-spots retained in metacer-
caria.
A Spotted Goby was isolated together with the two specimens of P. ulvae
infected with C. coronanda. ‘The penetration of the cercariae was observed
under the dissecting microscope.
The tail is cast the instant the cercaria comes
into contact with the host, and not, as is often the case with the Cercaria of
Cryptocotyle jejuna, some while after penetration has commenced.
At the end of three weeks one of the snails died and it was thought advisable
to examine the rediae and young stages from the other snail immediately. The
infection experiment was therefore terminated and the Goby killed.
A graduated series of cysts of C. coronanda in various degrees of develop-
ment was found embedded in the lateral muscles, not beneath the skin or in the
fins. There was no pigment formation around the site of the cysts. It was
possible to see the coronet of spines through the cyst-wall, and all the fully
developed cysts except one were hurriedly fed to a laboratory-reared gull (Roths-
child 1936) under the mistaken impression that the ’metacercariae pertained to
Pygidiopsis Looss 1907 or an allied genus. It was no surprise, in the light of
subsequent knowledge, to find no trace of adult worms when the gull was
eventually autopsied.
The fully developed cyst (304 x 174) is oval in shape and of a rather
pearly appearance. It is easily separated from the tissues of the host. Neither
the fine outer layer nor the thicker inner layer of the cyst wall is very tough.
The metacercaria does not fit closely inside the cyst, and there is a space in
which the dissecting needle can be planted without damaging the worm.
The following description of the metacercaria is based on the single specimen 1
removed from one of the largest cysts. As already stated, those fed to the gull
were only examined through the cyst-wall.
1 Deposited at the British Museum (Natural History), Cromwell Road, London.
Novyirarrs ZoonoaioaAn XLI. 1938. 159
The oral spines, excretory bladder and gonotyl were examined while the
worm was alive. It was subsequently fixed in Bouin Dubosq and stained with
Delafield’s Haematoxylin.
The specimen is mounted partially on its side and has elongated somewhat
during fixation (compare figs. 103 and 104). The alimentary canal is pushed to
one side and the body-wall slightly damaged.
METACERCARIA OF C. coronanda (figs. 103-106).
Measurements in Microns.
Body (length) ‘ : N ! : . 860
Body (width, maximum) i : 5 . 183
Oral sucker . ; : i ‘ . 75x 75 (at widest)
Oral aperture iramsrenseih) 4 : é 33
Prepharynx en : : : : > Ol
Pharynx: . 5 : ; : . 23 x 19 (at widest)
Ventral sucker : : : . 53 x 57 (at widest)
Gonotyl (transversely, in ee asien) . 46x 14
Oral spine (length). : : : : . 26
Eye-spot 5 ; é : : : . 4x7
Body elongated, about equal width along its whole length, tapering some-
what anteriorly, a little more than 4 times as long as broad. Hye-spots present
with long axis longitudinal. Cuticle armed with very small spines. Anterior
sucker (reminiscent of the anterior sucker found in Pygidiopsis, and allied
genera) terminal and funnel shaped, encircled by single row of 15 blunt spines.
These appear to be larger in comparison with the oral sucker than the spines
of species of Allocanthochasmus and Neochasmus figured by Van Cleave and
Mueller (1934). Mouth terminal, leading into long prepharynx, followed by
well-developed pharynx. There is a bulb-like enlargement of the prepharynx
anterior to the pharynx, most noticeable in the living animal. Oesophagus
divides immediately above ventro-genital sac, each intestinal caeca extending to
posterior end of body.
Ventral sucker situated a little more than + from anterior end of body,
moderately well developed with 9 raised ridges on its surface. These are not
visible in the mounted preparation, but were quite distinct during life. Excre-
tory vesicle Y-shaped, forking immediately below ventral sucker, and branches
extend forward to the oesophageal region. Excretory pore terminal, opening
by narrow sphincter muscle to exterior. Anlage of testes, situated tandem in
posterior end of body. Vasa efferentia, represented by a chain of cells, leave
testes on left side and pass upwards to meet and form vas deferens about 4 from
posterior end of body. Anlage of ovary forms a pretesticular band, uahbelhihr
lobed in mature worm. Germinal duct leaves it on left side. Uterus, represented
by a chain of cells, can only be traced anterior! to ovary. It passes forwards and
joins vas deferens some distance posterior and to the right of ventral sucker. This
hermaphrodite duct passes close to right border of ventral sucker and opens to
exterior by genital pore above and slightly to right of the median line of ventral
sucker. Genital pore surrounded by small transversely elongated oval sucker or
1 In the mature worm there are probably uterine loops in the posterior end of the body.
160 NOVITATES ZOOLOGICAE XLI. 1938.
gonotyl (fig. 104). Vitellaria undeveloped, but what is probably anlage of
vitelline ducts can be traced on either side of ovary, passing obliquely towards
FıG. 103.—Metacercaria of C. coronanda drawn from the living
specimen, showing the excretory bladder.
Fic. 104.—The same specimen mounted and fixed, showing
Anlage of reproductive system. (The alimentary canal is
much distorted by pressure.)
Fie. 105.—Oral spines of metacercaria of C. coronanda (ventral
view). The ninth (corner) spine is not figured.
Fic. 106.—Oral spines of Metacercaria of C. coronanda (dorsal
view).
Fia. 107.—Redia of C. coronanda.
lateral edge of body.
It will be seen from
the foregoing description
that, unfortunately, the
reproductive organs are in
a somewhat undeveloped
stage, although the char-
acters of the metacercaria
are such that it can with
some confidence be placed
in the subfamily Neo-
chasminae.
SUMMARY.
(1) The excretory
system of a new species
of Opisthorchioidea cer-
caria, Cercaria coronanda,
is described.
(2) The characters of
cercariae of the super-
family Opisthorchioidea
Vogel 1934 are discussed
and a specific diagnosis of
Cercariacoronandaisgiven.
(3) The first inter-
mediate host is Peringia
ulvae (Pennant) 1777, and
the second intermediate
host Gobius ruthensparri
(Euphras.) 1786.
(4) The metacercaria
pertains to the subfamily
Neochasminae Van
Cleave and Mueller 1932.
ACKNOWLEDGMENTS.
My thanks are due
to Miss A. Rothschild for
her invaluable assistance
in finding the second
specimen of P. ulvae in-
fected with this interesting
cercaria. I would also like
to express my deep gratitude to the Editors of the Journal of Parasitology, the
Zeitschrift fiir Parasitenkunde, and Zoologica, Professor H. W. Stunkard and
Professor H. Vogel for their kind permission to reproduce diagrams and drawings.
NOVITATES ZOOLOGICAE XLI. 1938. 161
EXPLANATION OF LETTERING.
anlage of ovary 5 : 5 : 5 : ON
anlage of testes : 5 é 5 1 Ww
anterior cuticular sxqenetona of tail , R 6 , ae
antero-lateral collecting tube . 3 5 ; . at
birth pore 2 5 : \ ; ö oe bp
bladder . : : ; : 5 : 3 5
boring spines. : : : E : 5 lox
caudal excretory Kalbe ; : : i : 5 OF
dorsal fin . ; : : ; ; : ; td
excretory pore . 2 . : © 6 © „ep
eye-spots . © : ¢ : : © : o ©
genital pre . : . : ; 4 { . gp
germinal duct . : F ; : h > esd
gonotyl . : : : \ : ; 6 . go
hermaphrodite duct . : : : : : Ah
intestine . 5 : : : a)
lateral ducts of qaneination elenmdk ; : : . Id
main collecting tube . 3 : ; : om)
median ducts of penetration clog ö : ; . md
nuclei of cells of developing sucker . ; : om
oesophagus 5 : \ 3 ; Ä 2 . oe
oral spines : ‘ i 5 ; ö : . os
oral sucker 5 ‘ ‘ | : : : 5 ©
penetration glands . B B : : : > &
pharynx . : J : : : 5 9
postero-lateral Bollecting aba ; 5 : ; =) pt
prepharynx : : : : i : © . pp
uterus. . : 6 . . . © .au
vasa efferentia . : : 4 ; : ; > W®
vas deferens . 5 : : 5 : - vd
ventral sucker . : : ; ; : ; SNe
ventral fin : : 4 ; : : : Aafv;
vitelline duct ? . i ; : b : ‘ . vie
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POSTSCRIPT
At the time of sending this paper to press the author had not seen Yama-
guti’s redescription of the Cercaria of Centrocestus (Stamnosoma) armatus. (Zur
Entwicklungsgeschichte von Centrocestus armatus (Tanabe) mit besonderer
Berücksichtigung der Cercarie—In Z. Parasitkd., Vol. 10, No 2, pp. 293-296,
1938). The excretory system of this larva is shown to be similar to that of C.
coronanda, although, of course, the bladder does not extend beyond the level of
the ventral sucker in either the metacercaria or mature worm.
164 NOVITATES ZOOLOGICAE XLI. 1938,
ON SOME NEOTROPICAL SIPHONAPTERA.
By KARL JORDAN.
(With text-figures 108-114.)
ASS some specimens of fleas obtained in Venezuela by Dr. G. H. H. Tate,
American Museum of Natural History, New York, and very kindly pre-
sented to our collection, is a specimen of Adoratopsylla Ewing 1925 which does
not quite agree with any we have. In order to ascertain whether it belongs to A.
bisetosa Ewing 1925 or not, this species had to be compared, and Dr. Edward A.
Chapin, Curator of Insects in the U.S. National Museum, Washington, D.C., has
very kindly lent mea Jj and a 9 of the original
three pairs on which A. bisetosa was based. I
very sincerely thank both Dr. Tate for his
welcome present and Dr. Chapin for his help.
Adoratopsylla is very closely related to
Tritopsylla Cunha 1929, but easily distinguished
from it.
Tritopsylla Cunha 1929 (text-fig. 108).
Syn.: Stenopsylla Cunha 1914 nec Kuwana 1909-10.
32. Head shorter than in Adoratopsylla,
more strongly rounded ; lower spine of genal
comb not longer than the next one. In Q
(text-fig. 108) the lower half or two-thirds of
frons with incrassations extending inward asin
Craneopsylla Roths. 1911 and allied genera,:
but these structures shorter; in & there isa
very faint trace of them close to frontal margin.
Both sexes with 3 antepygidial bristles. Stigma
cavity of abdominal tergum VIII very large.
g. Clasper apically divided by a sinus
into two processes ; tendon of ductus ejaculatorius exceptionally long, with 4
convolutions.
2. Duct of spermatheca not enlarged near bursa copulatrix, of even width
throughout.
Only one species (consisting of several subspecies) is known as yet: 1’. inter-
media Wagner 1901.
Adoratopsylla Ewing 1925.
32. Lower genal spine narrower and longer than the next one (text-fig. 113).
Internal incrassations (or channels) from frontal margin of head inward rudi-
mentary. ‘Two long antepygidial bristles, upper one the longer. Stigma cavity
of tergum VIII smaller than in Tritopsylla.
Sg. Clasper not divided by a deep apical sinus into two processes ; tendon
of ejaculatory duct short, little longer than lamina of phallosome, but slightly
curved upwards at proximal end, not rolled up.
Novrrares Zoonoaican XLI. 1938. 165
2. Duct of spermatheca widened from apex of bursa copulatrix for some
distance, the widened portion (text-fig. 112) somewhat resembling a pea-pod.
Three species have been described, one of them as yet doubtful; I add a
fourth. The species resemble each other closely, evidently being distinguished
only by the genitalia.
1. A. antiquorum Roths. 1904.
Described as Ctenophthalmus, it was placed by N. ©. Rothschild in 1915
together with 7’. intermedia into Doratopsylla ; Ewing put it into Adoratopsylla
in 1925, where it belongs. We know two subspecies :
a. A. a. discreta Jordan 1926.
6, Doratopsylla antiquorum discreta Jordan, Nov. Zool., xxxiii, p. 392, no. 12, text-fig. 18 (1926)
(Colombia).
The proofs of the paper quoted were evidently read hastily by me and not
properly corrected. After the manuscript had gone to the printers I decided to
employ the generic name Stenopsylla for both intermedia and antiquorum, but
forgot to replace Doratopsylla on one of the pages of the proofs by Stenopsylla.
Only the g is known. It differs from the Brazilian && particularly in the
nose of the posterior margin’ of the clasper being moved farther upwards.
6. A. a. antiquorum Roths. 1904 (text-figs. 109, 110).
SQ. Ctenophthalmus antiquorum Rothschild, Nov. Zool., xi, p. 643, no. 32, pl. xiv, fig. 72, pl. xv, figs.
80, 82 (1904) (Brazil).
Doratopsylla antiquorum Rothschild, Ectoparasites, i, p. 25, no. 2 (1915); Pinto, Arthrépodes Para-
sitos, 1, p. 374, no. 7 (1930).
The illustrations (figs. 109, 110) are here published partly as a supplement
to the figures of 1904 and partly to facilitate comparison of the next species
with A. a. antiquorum. The main feature in the clasper of the ¢ is the peculiar
166 Noyirates ZOOLOGICAE XLI. 1938.
nose (n), an incrassation on the inner surface, resembling the head of a bird ;
the nose is concave in the centre as figured and strongly chitinized at the margins ;
opposite it the digitoid F bears a rounded impression for the reception of the
tip of the nose.
Sternum VII of the 2 has not yet been figured. Unfortunately we have
only a single specimen of this sex and in this, still more unfortunately, the sternum
of the left side is ventrally torn, the specimen evidently having suffered in being
mounted. Both the left (A) and the right (B) side are here figured. The two
sides agree well, apart from minor differences usually observed in the two sides
of an individual. The small triangular excision in both halves, which at first
sight gives one the impression of being an accidental tear in the margin, must be
looked upon as the homologue of the larger sinus of the other species of Adorato-
psylla. Body of spermatheca not quite twice as long as broad.
In the collection from South-east Brazil, 4 3g, 1 9, off Didelphys marsupials
aurita. -
2. A. cunhai Pinto 1925.
Stenopsylla cunhai Pinto, Bol. Inst. Bras. Sci., i, no. 3 (1925) (Est. de Rio).
Tritopsylla cunhai id., Arthrépodes Parasitos, i, pp. 351, 377, text-figs. 139, 183, 184 (1930).
On Didelphys opossum. The figures published by Dr. Pinto apply equally
well to all the known
Brazilian species of
Adoratopsylla ; sternum
VII is not figured. As
Dr. Pinto did not know
that A. antiquorum be-
longed to this genus, and
therefore did not com-
pare his cunhai with it,
=18 his Didelphys flea may
turn out to be the same
y as A. a. antiquorum or
N as A. bisetosa. If it
equals bisetosa, the name
Vill.st. cunhai may have priority.
\e 3. A. bisetosa Ewing
va 1925 (text-figs. 111, 112).
A. b. Ewing, Journ. Parasito-
logy, xii, p. 44 (1925) (Rio
Ce 4 Branco); Pinto, Arthropodes
N Parasitos, i, p. 373, no. 1
N (1930).
According to in-
formation received from
Dr. E. A. Chapin, the six
specimens on which the
original description was
based were [all on one slide. It is therefore probable that all six are co-types ;
if that is the case, a type should be selected and labelled as such. Specific
differences between 99 being difficult to see, while those between 33 are easily
Novirates Zoonoarcan XLI. 1938. 167
observed, a male should be selected as type. Dr. Ewing, like some other
naturalists, evidently prefers to describe the 9 of a new species first and then
the g. A description has the object, I take it, to enable other workers to
identify the new species; therefore in any group where the 9 shows the
specific differences best, this sex should stand first, and where the g-characters
serve more conveniently for the identification of the species, I should give the
& priority of place, and select a g as type in order to simplify identification.
In the present case a description of the 2 which does not include sternum VII
is of little value for the identification of the species. Dr. Ewing says that,
in the 9, the number of spines in the pronotal comb is 16 to 18; I definitely
exclude a specimen with 16 pronotal spines from being selected as the type ;
the normal number being 18 in this genus, an example with 16 pronotal spines
may possibly represent another species. Dr. Ewing’s description of the
seventh abdominal tergite is that of the eighth, a mere pen-slip. Some of the
main differences in the g-organs between A. bisetosa and A. antiquorum are
correctly stated by Dr. Ewing, and his description of them left no doubt in my
mind that A. bisetosa was a species distinct from A. antiguorum. The pair loaned
to me by Dr. Chapin amply confirms the specific distinctness of A. bisetosa, as a
comparison of figs. 109 to 112 will prove to the reader.
3. Process P of the clasper (text-fig. 111) broadly conical, with the anterior
and posterior margins somewhat rounded, the apical portion of the anterior margin
from the pair of long subapical bristles upward more oblique, the apex itself
subacuminate rotundate ; the nose of A. antiquorum absent, the notch n apical.
Digitoid F broader than in A. antiquorum, much more ventricose, and the pair of
bristles much more ventral. Apex of ventral arm of sternum IX _ broader,
168 NoviraTEs ZoonocicaE XLI. 1938.
its anterior apical margin less incurved; ventral arm as slender as in.
A. antiquorum, but with more bristles, the most proximal of them farther from
apex on the arm of the right side than on the left figured. Dorsal apieal project-
ing angle of phallosome (Par) broader than in A. antiquorum.
9. Sternum VII (text-fig. 112) deeply and broadly incurved, quite unlike in
shape from what we see in A. antiquorum (text-fig. 110) and the species here
following ; each side with 10 bristles, of which 4 are subapical and long and strong.
Apical margin of tergum VIII sinuate much below middle, strongly rounded above
sinus, farther upwards slightly incurved and then again rounded for a short
distance ; above stigma 3 bristles each side, in sinus 2 small ones ; on lower area a
submarginal row of 5 on one side, 6 on the other, 2 of these almost on a level with
the sinus and in front of them 3 small bristles as stated by Ewing. Spermatheca
(R.s.) very similar to that of A. antiquorum, its tail more bulbous, its body nearly
twice as long as broad. The two specimens examined are a little larger than
A. antiquorum: hindfemur in A. antiquoram 3 0-39, 2 0-43, in A. bisetosa & 0-40,
2 0-45 mm.
The specimens were obtained at Santa Maria, Rio Branco (Minas Geraés) off
Monodelphis brevicaudata.
4. A. dilecta (text-figs. 113, 114).
A single 2 obtained by Dr. G. H. H. Tate on the Phelps Venezuela Expedi-
tion on the Anyantepui plateau, 1850 m., off Marmosa murina.
The geographically nearest Adoratopsylla is A. antiquorum discreta Jord.
1926 from Cundinamarca, Colombia, off Peramys adustus. The 9 of A. a. discreta
not being known, I was at first inclined to place the Venezuelan 2 with that sub-
Novirates ZooLoGIcaE XLI. 1938. 169
species ; but a close comparison with the 9 of A. a. antiquorum convinced me
that the specimen represents a different species.
The head (text-fig. 113) agrees with that of the other species ; the inward
extensions of the frontal margin so distinct in the 9 of Tritopsylia intermedia
(text-fig. 108) are indicated in the lower half of the frons. The lower genal spine
of the left side is broken at the tip, the dotted line indicating the length of the
spine of the right side. The specific distinctions are found in the abdominal
sternite VII and the spermatheca. This sternite (text-fig. 114) has a narrow, but
rather deep sinus
the bottom of which
is rounded ; above
and below the sinus
the margin of the
segmentis rounded,
the upper lobe more
strongly so, pro-
jecting farther than
the lower one; there
is a subapical row
of 4 bristles on one
side and 5 on the
other ; in front of
the row 6 smaller
bristles on each
side. Apicalmargin
of tergum VIII far
below middle with
a shallow sinus at which there are two small bristles as in the other species,
but the upper bristle much longer than the lower one ; on left side a subapical
row of 4 bristles in front of which 2 small ones, on right side 5 subapical
bristles and 4 small ones; above stigma 3 bristles. Anal sternite apically
concave as usual in this genus and Tritopsylla (not in Doratopsylla), but the
cavity small, less deep, therefore a larger number of the small bristles are placed
on the outside of the segment in front of the cavity. Spermatheca much
broader posteriorly than near its tail, which may partly be due to the organ
perhaps not presenting a view from the side, orifice in middle, not subventral ;
tail with a small appendage and much less bulbiform than in the other species ;
duct of spermatheca destroyed in the process of clearing.
Length 3-1 mm. ; hindfemur 0-43 mm.
170 NOVITATES ZOOLOGICAE XLI. 1938,
A NOTE ON THE FIN-FOLDS OF CERCARIAE OF THE SUPER-
FAMILY OPISTHORCHIOIDEA VOGEL 1934 (TREMATODA).
By MIRIAM ROTHSCHILD.
(With text-figures 115-124.)
(ae importance has been placed on the structure of the tails of cercariae,
which, as Stunkard (1930) points out, is accepted by many authors as a
morphological feature of much phylogenetic significance.
Vogel (1934) states that it is unlikely a different arrangement of tail fin-
membranes could be found in one family or even superfamily of Trematodes,
and suggests that the apparent differences in the fin-membranes of the cercariae
of the superfamily OPISTHORCHIOIDEA are due to faulty descriptions. Cable
(1938) apparently accepts all these descriptions, and states that: “The Opis-
thorchiidae and Heterophyidae as well as the Allocreadiidae present a perplexing
problem in the taxonomy and relationships of Trematodes. The diversity
of cercarial types in these families makes it exceedingly difficult to apply the
recapitulation hypothesis in determining relationships.”
So far only two types of cercariae have been definitely proved to belong
to the superfamily OPISTHORCHIOIDEA, namely those pertaining to the Pleuro-
lophocerca and Parapleurolophocerca groups (Sewell 1922). The only important
difference between these two larval forms is found in the position of the tail
fin-folds.1_ In the former the membrane (if present) consists of a median dorso-
ventral fin-fold (text-figs. 115-119) and in the latter a continuous lateral
fin-fold (text-fig. 120).
However, among the typical Heterophyid cercariae from Peringia ulvae
(Pennant) 1777, several species have been discovered which combine the characters
of both these groups. A median dorso-ventral fin-membrane is present together
with a continuous lateral fin-membrane (text-figs. 121-124).
There seems little doubt that, in early descriptions, several of the forms with
a dorso-ventral fin-fold have been erroneously described as possessing a lateral
fin-membrane. This mistake can arise very easily owing to the displacement of
the fins under pressure of the coverslip (text-figs. 115, 118, 124). The Cercaria ?
of Clonorchis sinensis (Cobbold) 1875 is almost certainly a case in point, as
suggested by Vogel. Nevertheless, if we entirely disregard all doubtful
descriptions we can still trace a graduated series in the tails of the cercariae
of the Pleuro- and Parapleurolophocerca groups, ranging from the four
continuous fin-folds of Cercaria sp. from Peringia ulvae (text-fig. 121) to Cercaria
indicae III Sewell 1922, and the Cercaria of Centrocestus armatus (Tanabe) 1922,
1 Other minor morphological differences may be noted. The Parapleurolophocerca cercariae
have the acetabulum, intestine and cuticular spines relatively more developed, and the ducts of the
penetration glands are generally arranged in a line and not in bundles. The adults pertain to the
subfamily Haplorchinae Looss 1899. These differences are not constant, however, and in some
Parapleurolophocerca cercariae penetration-gland ducts are arranged in bundles, etc.
2 Yamaguti (1935) redescribed the Cercaria of Clonorchis sinensis and found, contrary to Faust
and Khaw (1927), that the fin-fold was dorso-ventral, resembling that of the Cercariae of Opisthorchis
felineus. Unfortunately he does not offer the necessary experimental proof that the cercaria he
examined was in fact this species.
Novitrates ZooLoGIcAE XLI. 1938. 171
where the tail is provided with a cuticular sheath, but is totally devoid of fins.
It would, therefore, seem more correct to say that within the superfamily OPrıs-
THORCHIOIDEA there occur differences in the structure of the caudal fins of the
cercariae rather than
“two distinct larval
types.”
A group of Tre-
matodes such as this
superfamily will often
present a character-
istic and distinctive
type of cercaria, the
importance of which
cannot be over-
estimated as a guide
to their natural re-
lationship. This fact
has frequently misled
authors into forming
a far too rigid classifi-
cation, based on a
combination of larval
characters, such as
fins, eye-spots, pene-
tration glands, stylets,
etc., displayed by the
more typical species.
A slight departure
from the usual life-
cycle by any members
of a family or even
genus is frequently
associated with mor-
phological changes in >
these very characters.
Thus in the two cases
recorded in the super-
family OPISTHOR-
CHIOIDEA in which the
second intermediate
host is not a fish,
the cercariae display
marked modifica-
tions. The Cercaria of
Euryhelmis monorchis
Ameel 1938 which enemas fi in tadpoles and frogs is devoid of eye-spots, and the
dorsal fin is lacking in the Cercaria of Metagonimoides sp. (Ingles 1935) which
encysts within the first intermediate host. The variation in the fin membranes
of this group may also be associated with some aspect of the life-cycle, which is at
13
172 NOVITATES ZOOLOGICAE XLI. 1938.
present unknown. Parallels to this can be found elsewhere among the Trematoda.
MacMullen (1937), in his admirable paper on the PLAGIORCHIOIDEA, has already
drawn attention to the fact that too much stress should not be placed on single
larval adaptations.
It seems possible that the cercariae with continuous lateral fin-folds are
restricted to the Haplorchinae Looss 1899 which may eventually be raised to
family rank. It would not, however, be surprising to find a parallel tendency
towards reduction or multiplication of fin-folds in the cercariae of various sub-
families cf the Heterophyidae.
There seems insufficient evidence to justify Cable’s (1938) statement that
“ The application of the principal that related digenetic trematodes have similar
cercariae unites dissimilar miracidial and adult forms in the same superfamily
in the case of the Heterophyidae and Opisthorchiidae. ... .” Not more than
fourteen life-cycles of the Pleuro- and Parapleurolophocerca groups have been
completely and satisfactorily worked out, but (despite variation in fin-folds)
from the available data the impression is gained that there is no discrepancy
between the cercariae and adult forms. On the contrary, a very characteristic
type of larva is found, confirming the relationship of these Trematodes, which,
on morphological considerations, have been grouped together in the superfamily
OPISTHORCHIOIDEA.
EXPLANATION OF TEXT-FIGURES
115. Ventral view of tail of Cercaria of Cryptocotyle jejuna Nicoll under a
coverslip, showing displacement of the fin-folds.
116. Dorsal view of tail of Cercaria of ©. jejuna.
117. Lateral view of tail of Cercaria of C. jejuna.
118. Dorsal view of tail of Cercaria of C. lingua (Creplin) under a coverslip,
showing displacement of fins.
119. Lateral view of tail of Cercaria of C. lingua.
Novirares Zootoaicarn XLI. 1938. 7c}
120. Tail of a Parapleurolophocerca cercaria showing lateral fin-folds only.
121. Ventral view of tail of Cercaria sp. from Peringia ulvae with four con-
tinuous fin-folds, lateral and dorso-ventral.
122. Lateral view of tail of Cercaria sp. from P. ulvae with continuous lateral
fin-fold and reduced dorso-ventral fin-fold.
123. Ventral view of tail of the same species.
124. Ventral view of tail of the same species greatly compressed under a cover-
slip, showing displacement of fins and anterior cuticular expansions.
EXPLANATION OF LETTERING
Anterior cuticular expansion.
Dorsal Fin.
Lateral Fin.
Tail.
Ventral Fin.
<q fa] IS lel @
REFERENCES
Ameel, D. J., 1938. The Morphology and Life-History of Huryhelmis monorchis
n. sp. (Trematoda) from the Mink.—In J. of Parasit., Urbana, 24, No. 3,
pp. 219-223, 5 figs.
Cable, R. M., 1938. Studies on Larval Trematodes from Kentucky with a
summary of known related species.—In Amer. Midl. Nat., Notre Dame,
19, No. 2, pp. 440-464, 14 figs.
Faust, E. C., Khaw, O. K., Ke-Fang, Y., and Yung-an, C., 1927. Studies on
Clonorchis sinensis (Cobbold), with a consideration of the Molluscan hosts
of Clonorchis sinensis (Cobbold) in Japan, China and South-eastern Asia,
and other species of Mollusca closely related to them, by Bryant Walker.—
In Amer. J. Hyg., Baltimore Monogr. Series No. 8, pp. xi and 284, 14 pls.,
33 figs.
Ingles, L. G., 1935. Notes on the development of a heterophyid trematode.—
In Trans. Amer. Mic. Soc., 54, pp. 19-21, 1 pl.
McMullen, D. B., 1937. A discussion of the taxonomy of the family Plagiorchiidae
Lühe, 1901, and related Trematodes.—In J. Parasit., Urbana, 28, No. 3,
pp. 244-258, 1 pl.
Sewell, R. B. S., 1922. Cercariae indicae—In Ind. J. med. Res., 10 (suppl. no.),
pp. 1-370, i-iii, and 32 + 7 pls.
Stunkard, H. W., 1930. The life-history of Cryptocotyle lingua (Creplin), with
notes on the physiology of the Metacercariae.—In J. Morph., Philadelphia,
50, pp. 143-190, pls. i-iv.
Vogel, H., 1934. Der Entwicklungszyklus von Opisthorchis felineus (Riv.)
nebst Bemerkungen über die Systematik und Epidemiologie. —In Zoologica,
Stuttgart, 33, Hf. 86, pp. 1-103, 28 figs., 8 pls.
Yamaguti, S., 1935. Uber die Cercarie von Clonorchis sinensis (Cobbold).—
In Z. Parasitenk., Berlin, 8, Hf. 2, pp. 183-187, 4 figs.
Yamaguti, S., 1938. Zur Entwicklungsgeschichte von Centrocestus armatus
(Tanabe) mit besonderer Berücksichtigung der Cercarie.—In Z. Parasitkd.,
10, No. 2, pp. 293-296.
101 and 102),
EXPLANATION OF PLATES III AND IV
Polygonia c-album, upperside
i en underside
Polyptychus trilineatus sonantis 3, type
undatus 3, type.
55 iy philippinensis 3
6 n kalisi 3, paratype
Delias itamputi luctuosa 3, type
Charaxes tectonis 3, type
Daphnusa ocellaris ailanti 3
2 2» » 2
Hippotion moorei moorei 3, paratype
a exclamationis robur 2, type
5 55 „ 9, paratype
re rebeli &
butleri 5
> mC
Temnora grandidieri 3
engis 3, type
Ludia arida 3, type
» orinoptena monroe 2, type .
re)
no}
id te) te TSI i to ts) to te as) fe) fe) ts) is
. 125
125
127
127
128
130
CE, Vou. XLI.1938.
ZooLocı
NovITAtEs
nr, mr |
ee Et ea
Novirates Zoo.ocic#, Vou. XLI.1938. Pl. IT.
4
f = > wo [4 j \ > x
Me» X PN F DR Koh ‚6
ae mn ' non Se yf
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2
re
Novirares Zoouocic#, Vou. XLI.1938. Pl. III.
Hilda Jordan photo John Bale Sons & Curnow, [td London
Novirates Zootocic#, Vou. XLI, 1938. Pl. IV.
Hilda Jordan photo John Bale Sons & Curnow, Ltd London.
PER u a
i " Bun ;
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if
Vi
NOVITATES ZOOLOGICAR,
\ CN ¢ At
AN ANLEIOAN
ed A N C=
Vor =
by Man ICh Wee
ares a
AH Fournal of Zoolog :
EDITED BY
THE HON. MIRIAM ROTHSCHILD
AND Dr. KARL JORDAN, F.RS.
No. 3.
Pacss 175-304.
PLATES V AND VI
Issurp Aucust 31, 1939, ar THE ZootocicAL MUSEUM, Trine.
PRINTED BY HAZELL, WATSON & VINEY, Ltp., LONDON AND AYLESBURY.
1939.
Or
10.
. ON
Vor. XLI.
NOVITATES ZOOLOGICAE
BDITED BY
THE HON. MIRIAM ROTHSCHILD and KARL JORDAN.
CONTENTS OF NO. III
. A NOTE ON SOME MALLOPHAGAN NAMES .
. A NOTE ON THE LIFE CYCLE OF CRYPTO-
COTYLE LINGUA (CREPLIN) 1825 (TRE-
MATODA) (PLATE V) . BNE Oy de
. SOME ARABIAN MAMMALS COLLECTED BY
MR. H. ST.J. B. PHILBY, C.LE. (PLATE V1) .
. ZUR NOMENKLATUR DER BEIDEN DEUT-
SCHEN PROCRIS-ARTEN MIT SPITZEN
FÜHLERN .
. THE BRAZILIAN HYDROPTILIDAE en
PTERA)
. SOME OBSERVATIONS ON THE GROWTH OF
PERINGIA ULVAE ae 1777 IN THE
LABORATORY :
SOME OLD-WORLD AGARISTIDAE
(LEPID.)
. ON THE CONSTANCY AND VARIABILITY
OF THE DIFFERENCES BETWEEN THE
OLD WORLD SPECIES OF UTETHEISA
(LEPID.; ARCTIIDAE)
. ON FIVE SIPHONAPTERA FROM THE RE-
PUBLIC OF ARGENTINA .
NOTICE OF POSSIBLE SUSPENSION OF THE
RULES OF NOMENCLATURE IN CERTAIN
CASES [A.(n.s.)1]
PAGES
Theresa Clay 175—177
Miriam Rothschild 178—180
T. C. S. Morrison-Scott 181—211
Karl Jordan 212—216
Martin E. Mosely 217—239
Anne Rothschild and
Miriam Rothschild . 240—247
Karl Jordan 248—250
Karl Jordan 251—291
Karl Jordan 292—303
Francis Hemming
304.
NOwWwiEnATES ZOOLOGICAE
Vol. XLI.
AUGUST 1939. 7 No. 3.
A NOTE ON SOME MALLOPHAGAN NAMES.
By THERESA CLAY.
I. Nirzscu’s Nomina NUDA AND MANUSCRIPT NAMES.
IR 1818 Nitzsch published a number of specific names of Mallophaga giving
the hosts but no descriptions,! and when he died in 1837 he left a manu-
seript containing the descriptions and names of a large number of species ; both
these nomina nuda and manuscript names have led to endless confusion in the
nomenclature of the Mallophaga. Burmeister, working on the Nitzsch collection,
published (1838) his own descriptions of a number of species using these Nitzschian
names, and Denny (1842) also applied many of the names to this species attributing
them to Nitzsch. Giebel, who had access both to Nitzsch’s collection and manu-
script, proceeded to publish from 1861 onwards Nitzsch’s specific descriptions,
giving the Latin diagnosis taken from the Nitzsch manuscript and also, in certain
cases, adding a German description of his own. Giebel thus published the
descriptions of all the species which had in the meantime been described by
Burmeister, Denny, and other authors, attributing them to Nitzsch. This has
caused complications both as to the date and author of the name and to the
interpretation of the species. Also in some cases Burmeister, in describing a
species in the Nitzsch collection, had used a different name from that found in
the Nitzsch manuscript, e.g. Nirmus sellatus Burmeister 1838 and Nirmus selliger
Nitzsch 1866, both with the same type specimens. Again in some cases authors
had applied a Nitzschian name to what was obviously a different species from
that meant by Nitzsch, e.g. Nirmus cameratus de Haan 1829. In order to obviate
.the difficulties and confusion which may arise it is necessary to adhere strictly
to the law of priority. Thus any author who described a species for which he
used a nomen nudum contained in Nitzsch’s 1818 list or a name taken from the
Nitzsch manuscript, and published prior to Giebel’s publication of the Nitzsch
manuscript, must be considered as the sole author ? and his interpretation and
spelling must stand. Names and descriptions taken from the Nitzsch manuscript
and published for the first time by Giebel should stand under the authorship of
Nitzsch.
Additional complications arise in the case of Lyonet’s descriptions, since in
his unpublished manuscript he described several species, but referred to them by
1 This does not apply to names with which there is a reference to an earlier description and
figure as in the case of Nirmus minutus (1818, p. 281); such names must date from 1818.
2 It is unfortunate that Dr. S. Kéler, who is working on the Nitzsch collection, does not conform
to this rule and thus reduces the value of much of his excellent work.
14 175
JAN 1.9 1940
176 NovirarEes ZooLoGIcAE XLI. 1939.
French names only. In 1829 de Haan published these descriptions, and in addi-
tion applied Nitzschian nomina nuda to Lyonet’s species. In this case, therefore,
de Haan must be considered as the author and the synonymy of such species
should run as follows :
COLPOCEPHALUM FLAVESCENS de Haan 1829.
Colpocephalum flavescens Nitzsch, 1818, p. 298 (nom. nud.).
Pou d’aigle, Lyonet, 1829, p. 262, pl. iv, fie. 1.
Liotheum (Colpocephalum) flavescens de Haan, 1829, p. 309, pl. iv, fig. 1.
II. EsTHIOPTERUM HARRISON 1916 OR ESTHIOPTERELLA HARRISON 1937.
The genus Lipeurus Nitzsch 1818, as originally described, contained a large
and heterogeneous collection ofspecies. In 1916 Harrison restricted the genus to
a group of species found mainly on gallinaceous hosts. The residue was placed
in a new genus, Hsthiopterum, with E. ebraeum (Burmeister) 1838 as a genotype.
Harrison, however, included in his list of species of Esthiopterum the genotype of
the genus Pseudonirmus Mjöberg 1910 which he did not at that time consider a
valid genus. Harrison (1937, posthumous publication) argues that as the geno-
type of Pseudonirmus was included in his genus Esthiopterum, this latter genus
isa synonym of Pseudonirmus and therefore proposes the new name Hsthiopterella
for Esthiopterum. As the two genotypes of Pseudonirmus and Esthiopterum are
quite distinct and obviously belong to different genera, it is unnecessary to discard
the name Esthiopterum. For this opinion I am indebted to Dr. Karl Jordan.
III. GENERA IN HARRISON’S POSTHUMOUS PUBLICATION.
After Harrison’s death in 1928 selections from his manuscript were published
by G. B. Thompson in a series of papers in the Annals and Magazine of Natural
History."
Recently (1937) a posthumous paper by Harrison has appeared containing
the majority of genera and subgenera which had already been published under
Thompson’s name, the descriptions being identical in the two cases. Since
Thompson’s descriptions were taken from Harrison’s manuscript, the names must
stand under the authorship of Harrison with the date as published by Thompson.
Below are given a list of these genera and subgenera :
Genera. Episbates Harrison 1935.
Perineus Harrison 1936.
Halipeurus Harrison 1936.
Subgenera. Epipelecanus Harrison 1935.
Philichthyophaga Harrison 1935.
Epifregata Harrison 1935.
I would like to express my gratitude to Dr. Karl Jordan for his invaluable
help and advice.
REFERENCES.
Nitzsch (1818). Die Familien und Gattungen der Thierinsekten. Germar’s
Magaz. d. Ent., iii, pp. 261-316.
1 Thompson, Ann. Mag. Nat. Hist., ser. 10, vol. xv, p. 555, 1935; vol. xvi, pp. 148 and 485,
1935 ; vol. xviii, p. 40, 1936 ; vol. xx, p. 538, 1937.
Novirates ZooLoaicar XLI. 1939, 177
Burmeister (1838). Handbuch der Entomologie, Berlin, Bd. ii, pp. 418-433.
Denny (1842). Monographia Anoplurorum Britanniae, London.
De Haan (1829). Lyonet’s Anatomie de differentes especes d’insectes. Mem.
du Mus., xviii, pp. 233-312.
Harrison (1916). The Genera and Species of Mallophaga. Parasitology, vol. ix,
pp. 1-156.
Harrison (1937). Mallophaga and Siphunculata. Australasian Antarctic Expedi-
tion, 1911-1914, Scientific Reports, ser. c, vol. ii, pt. i, pp. 1-47.
178 Novirates ZooLoGIcAE XLI. 1939.
A NOTE ON THE LIFE CYCLE OF ORYPTOCOTYLE LINGUA
(CREPLIN) 1825 (TREMATODA.)
By MIRIAM ROTHSCHILD.
(With Plate V.)
iE 1930 Professor H. W. Stunkard demonstrated experimentally that a Pleuro-
lophocercous cercaria parasitising Littorina littorea (L.) (probably identical
with Cercaria lophocerca Lebour 1907 nec Fil.) encysted in the fins and skin of
certain fish, and when fed to white rats, guinea pigs and a kitten developed into
the Heterophyid fluke Cryptocotyle lingua (Creplin) 1825.
In 1935 attempts were made to repeat the life-history of ©. lingua experi-
mentally with material obtained in Great Britain.
A prolonged study of the cercariae (pl. V, fig. 6) infecting Littorina littorea
convinced the author that only one species of Pleurolophocerca cercaria parasitises
this mollusc in the Plymouth district, although a very closely related and almost
indistinguishable species was found in Littorina littoralis (L.) in precisely the
same locality. Slight morphological differences were noted between the- species
from L. littorea and Professor Stunkard’s description of the Cercaria of Crypto-
cotyle lingua, but it seems ‘probable that these differences are more apparent
than real.
Special attention was paid to the behaviour of the cercariae, as this feature
is of importance for purposes of specific diagnosis. The method of swimming is
characteristic of the Pleurolophocerca Group. Active periods (averaging 2}
seconds) in which the cercariae swim towards the light, alternate with passive
periods (averaging 15} seconds) during which they hang suspended, tail upper-
most, and then sink vertically through the water. They are strongly photoposi-
tive and activated by a shadow or by touch. On account of the latter characteris-
tic they accumulate on the lighted side of a vessel far more rapidly in shallow
than in deep water, as the passive sinking phase is abruptly terminated by contact
with the bottom of the dish.
The fertility of a single infection is very high. Littorina, in the laboratory,
have emitted cercariae for periods extending over three years, averaging about
300 cercariae per day! Emissions seem unaffected by light or darkness, but
inhibited by a relatively small fall in temperature.
If the host is kept entirely submerged, the majority of cercariae emerge at
a certain definite period of the day, but the peak varies by several hours according
to individual snails. The time of emergence is therefore of little or no value as a
specific character in the cercaria of ©. lingua. If the snail is removed from the
water for several hours and then replaced, emissions can generally be induced
at any time. This type of behaviour is to be expected when the host is an inter-
tidal form, such as Littorina. Another type of variation is also recorded. In
some individual snails a certain number of cercariae are emitted fairly regularly
over long periods, whereas in other cases short and intense periods of production
alternate with periods of complete inactivity during which the shedding of
cercariae ceases altogether.
Novtrates Zoonoarcarm XLI. 1939. 179
The fish employed experimentally as second intermediate host by Professor
Stunkard was Tautogolabrus adspersus, the Cunner, which does not occur in
British waters. A search was therefore made for another suitable intermediate
host.
Cryptocotyle lingua, unlike C. jejuna, is reported to encyst in a wide range of
fish. Many species collected at Plymouth appeared to be naturally infected with
encysted Cryptocotyle lingua, but owing to the remarkable similarity of the meta-
cercariae of several species of this genus, it is unwise to record them definitely as
intermediate hosts. It was found that in the laboratory the cercariae will attack
almost any fish with which they are brought in contact, although they penetrate
certain species far more readily than others. Among these were several in-shore
forms, such as Wrasse (Labrus bergylia Ascan., Ctenolabrus rupestris (L.)), Rockling
(Onos mustela (L.), O. tricirratus Bloch), Blennies (Blennius pholis L., B. gattoru-
gine Bloch) and the Butterfish (Pholis gunnellus (L.)), which are almost certainly
natural hosts.
It was interesting to find that in C. lingua encystment of the cercaria does
not necessarily follow penetration. Thus, for example, the larvae readily invade
the freshly laid eggs of certain fish. The tails are shed and the cercariae actively
work their way into the egg. After moving about for a varying period they
gradually force their way out again and drop to the bottom of the dish. The
cercariae will also penetrate the soft parts of molluscs with similar results.
On account of the ease with which they can be kept alive in the laboratory,
the Spotted Goby, Gobius ruthensparri (Euphras.) and Ballan Wrasse, Labrus
bergylta Ascan., were selected for infection experiments. Large invasions of the
cercariae generally killed the Spotted Gobies immediately ; therefore the numbers
to which they were exposed had to be carefully controlled. It was noticed that
in the Wrasse (pl. V, fig. 5), the cysts generally became surrounded by the usual
mass of pigment cells, whereas no pigment whatsoever was formed around the
cysts in the Goby, which were marked only by a whitish, opaque area in the fins
or tail (pl. V, figs. 1 and 3). This appeared the more remarkable when it was
noted that the tissue surrounding the cysts of Cryptocotyle jejuna Nicoll 1907
(Rothschild 1938), became heavily pigmented in the same host (pl. V, figs. 2
and 4). It is possible that the pigment response is indicative of the degree of
mutual adaption of host and parasite. The Spotted Goby is probably rarely
attacked in nature by the Cercaria of Cryptocotyle lingua. It is, however
almost certainly a natural host of the Cercaria of C. jejuna. The location of
the cysts, namely in the fins and beneath the skin and scales of the body, is
identical in both cases.
After infection the fish were kept a fortnight before feeding experiments were
commenced.
In 1935 and 1936 only laboratory-reared albino rats were employed as the
final host. The rats were sacrificed at varying intervals, but all these experiments
proved negative. Simultaneously, two rats were fed cysts believed to be those
of C. lingua from naturally infected Rockling. These also failed to develop. In
1937 laboratory-reared albino rats, laboratory-reared ducks and laboratory-
reared Black-headed Gulls (Rothschild 1936) were used. Again no flukes were
recovered from any of these animals.
In 1938 the experiment was repeated, using the same intermediate hosts and
with two of the gulls, aged six months. Apart from the fact that the birds were
180 NOVITATES ZOOLOGICAE XLI. 1939.
deprived of the artificial vitamins generally added to their food, the experiments
were precisely similar to those carried out in previous years, and which hitherto
had met with no success.
The first bird was fed cysts continuously from November 10th to December
10th and was then sacrificed. About 200 specimens of C. lingua (pl. V, fig. 7) in
varying degrees of development were recovered from the intestine. The second
bird was fed cysts at different intervals from August 1st until November 10th,
when cysts were given continuously until December 10th. An autopsy was
performed on January 2nd and about 60 specimens of C. lingua recovered
from the intestine. None of these flukes contained more than thirty eggs in
the uterus.
ACKNOWLEDGMENTS
I would like to express my gratitude to Dr. Stanley Kemp, F.R.S., for the
facilities he afforded me for carrying out this work at the Marine Biological
Laboratory, Plymouth.
REFERENCES
Rothschild, M., 1936. Rearing Animals in Captivity for the Study of Trema-
tode Life-histories—In J. Mar. Biol. Assoc. Plymouth, 21, No. 1, pp.
143-145.
Rothschild, M., 1938. Preliminary Note on the Life-history of Cryptocotyle
jejuna Nicoll, 907 (Trematoda)—In Ann. Mag. Nat. Hist. London, 1,
Ser. 11, pp. 238-239.
Stunkard, H. W., 1930. The Life-history of Cryptocotyle lingua (Creplin), with
Notes on the Physiology of the Metacercariae.—In J. Morph. Philadelphia,
50, pp. 143-190, 4 pls.
Noviratres Zoonoatcan XLI. 1939. 181
SOME ARABIAN MAMMALS COLLECTED BY
WOR. Lely Sind), PEIITEBSSELONERT
By T. C. 8. MORRISON-SCOTT.
Department of Zoology, British Museum (Natural History).
(With Plate VI and 8 text-figures.)
INTRODUCTION
RABIAN mammals were first discussed systematically by Cretzschmar
(1826), Lichtenstein (1827-34) and Hemprich & Ehrenberg (1828-33).
Since then little was written till Thomas (18948) wrote on the mammals of Oman.
This was quickly followed by discussions of the Aden mammals by Yerbury &
Thomas (1895) and Thomas (1900). Since that time there have been many papers
(see Appendix I) on individual forms, notably by Thomas, but only three papers
dealing with any considerable collections : Cheesman & Hinton (1924) on mam-
mals collected by Cheesman between Riyadh and the east coast, Dollman (in B.
Thomas 1932) on collections by Bertram Thomas in the south-east and Pocock
(19354) on Thomas’s specimens once more and on a collection from various
parts of Arabia presented to this museum by Philby.
Now Philby has sent home another fine collection, the results of his recent
journey to the Rub al Khali (Philby 1938), and it is with these specimens that
this paper is principally concerned though the earlier material has been re-
examined.
Apart from the fact that Arabia is still relatively unknown zoologically,
natural history collections from that country are of the greatest interest from the
zoogeographical point of view. Arabia les between the Ethiopian, Palaearctic
and Indian regions and the extent to which each region has contributed to the
fauna has been a matter of speculation ever since 1866 when Murray wrote on
the geographical distribution of mammals.
The Red Sea is almost certainly a rift valley and the land connexion between
Aden and Somaliland does not appear to have been broken until late in the Ter-
tiary period. Pilgrim (1919, Pl. I; 19254, p. 73; 19258, p. 202) shows that the
land bridge across the Straits of Hormuz between Oman and Persia lasted at all
events until the end of the Pliocene and probably till the late Pleistocene.
In view of the above it might be expected that south-eastern Arabia should
show faunistic affinities with Africa and that Oman should have Persian and Indian
relationships, and this is in fact the case. The Arabian Tahr (Hemitragus jayakari
Thos.), a goat-like animal living in the mountains of Oman, is an interesting case.
It is a close relative of the Tahr which is found on the southern slopes of the
Himalaya and the one in the hill ranges of southern India. It is essentially a
mountain animal and there do not appear to have been mountains along the land
route which the Tahr would have had to have followed if there had been no bridge
across the Straits of Hormuz. Oman has not been explored from the mammal
point of view since Dr. Jayakar’s time, and it would be interesting to find out if
182 Novirates ZOOLOGICAE XLI. 1939.
there are any Oriental forms there; that is, truly Oriental as opposed to
Indian.
Some elements of the fauna are common to India and Africa. Philby’s latest
collection includes some gerbils of the genus Tatera (a new record for Arabia) which
ranges as far as Ceylon on the one hand and S. Africa on the other, but the con-
nexion between these two extremes may be independent of land bridges across
the south ends ofthe Red Sea and the Persian Gulf, as the genus has been recorded
from the top end of the Persian Gulf.
_ The largest of the Arabian foxes (Vulpes vulpes L.) is common to N. Africa,
Arabia and N.W. India, but it is not to be regarded as a true Ethiopian or Indian
form, but as a Palaearctic intruder.
Lastly there are some Arabian mammals such as the hyrax (Procavia) which
are found elsewhere only in Africa, Palestine and Syria.
The Arabian fauna, therefore, is of mixed origin and offers many interesting
problems. More specimens are needed to solve these, apart from the entirely
philosophical object of finding out what mammals live in Arabia, and any speci-
mens sent to the British Museum will be gratefully acknowledged. ;
Throughout this paper the use of parentheses for the names of authors of
species which have been moved to another genus has been discontinued—for
reasons given by Dr. Osgood (1939), Science, 89: 9. Conventions used with
regard to certain measurements have been explained in Appendix II and a list
of the co-ordinates of those Arabian localities which are hard to find in maps
has been given in Appendix III.
I should like to take this opportunity of thanking my colleagues in the
Mammal Room for their various suggestions and help,’ especially Mr. R. I.
Pocock, F.R.S.
1. (?) Procavia syriaca jayakari Thos.—Hyrax.
Procavia syriaca jayakari O. Thomas 1892, Proc. Zool. Soc. Lond., p. 63, type loc.: Dofar; id.,
1894, l.c., p. 455, Dofar ; id., 1900, 1.c., p. 104, Abyan Mts. ; R. I. Pocock, 1935, Ann. Mag. Nat.
Hist. 15: 455, Ilha’ az & Deriyot.
Procavia habessinica jayakari Thos., H. Hahn, 1934, Z. Säugetierk. Berlin, 9: 302.
|
148 Q | 384 57 W.Hanaka | VIII? | 24.xi.1936 — 60 7-1
147 3 = = W. Hanaka Vv 24.x1.1936 68 52 6-2
107 3 357 56 Najran IV-V | 12.vii.1936 67 50 6-0
Type Q — Dofar VIII 1891 77 59 6:3
There is much variation in the colour of these specimens. Number 107
agrees fairly well with the type, but numbers 147 and 148 show more buff and
black amongst the hairs and agree better with two of Bertram Thomas’s specimens
from Deriyot (B.M. 34. 8. 4. 22-23) and one of Bury’s from Wasil in Yemen
(B.M. 13. 6.19.11). None of these specimens is so uniformly coloured dorsally
as the type and their bellies are creamy buff and not ochraceous as in the latter.
In fact, some of these agree better with specimens of ruficeps and burtoni than with
jayakari.
1 The identifications of the bats are due to Mr. R. W. Hayman.
2 For growth stages see Thomas, 1892, Proc. Zool. Soc. Lond., p. 53.
NovItates Zoonoaican XLI. 1939. 183
2. Procavia sp. —Hyrax.
Number 149, a young & (growth stage II) from Jadaliya, 26 November,
1936, and number 57, & (growth stage V-VI) from Dha’f Hills, Wadi Fatima,
25 January, 1935.
These specimens are alike in colour and differ widely from jayakari. The
following description is of number 57.
Dorsally—finely speckled black and buff, the hairs being very dark at the
base, buff in the middle and black at the tip, giving a brindled effect. The buff
dorsal spot is very distinct. The hairs are much shorter than in jayakari, 20 mm.
as opposed to 30mm. I have examined many skins of Procavia from Egypt and
the Sudan, as well as from Sinai and farther north, and the nearest specimen
to this one, so far as colour is concerned, is one from Rejaf, S. Sudan
(B.M. 28. 3. 11. 71), which was left unidentified.
Ventrally—silvery grey with a slight buff wash.
Hands and feet—as back.
Skull—proportionately broader in the lachrymal region than any other
Arabian specimen I have seen (text-fig. 125 B). Hahn (1934, p. 294), who makes
jayakari a subspecies
of habessinica, says of
the latter species: “ P,
verkümmert. Stets
einzelwurzelig.”’: The
specimens which Ihave
called jayakarı agree
with this but number
57 has P, well deve-
loped and two-rooted ;
number 149 has Pd,
strongly developed and
two-rooted. y Trxt-Fic. 125.—Skulls of Arabian Procavia : (A) the type of P.
The dimensions of syriaca jayakari Thomas (stage VIII) from Dofar ; (B) Mr. Philby’s
number 57 are: head No. 57 (stage V-VI) from Dha’f Hills; (C) Mr. Philby’s No. 147
Baile dy 376. hindfoot (stage V) from Wadi Hanaka. Drawn to the same scale, measure-
d
ments in text.
61, ear 30 millimetres.
Skull: condylobasal length 71, zygomatic breadth 43, postorbital width 26,
maximum width across frontals 35, mandibular length 55, breadth of M* 6-25
millimetres.
Hommel, in Die Namen der Sdugethiere bei den südsemitischen Volkern,
Leipzig, 1879, says (p. 322) : “ Die Araber beschrieben es als ein Tierchen, kleiner
als die Katze, braun, mit schönen Augen, ganz kurzem Schwanz, welches in
Häusern gezogen und gegessen wird, weil es das Gemüse abfrisst ; es gehöre zum
Geschlecht der Wiesel und die Leute nennen es ‘ ganamu bani Isra’il ’ (Kleinvieh
der Kinder Israel).”’
1 Itis as wellto add that many of the forms which Hahn includes in habessinica may show a two-
rooted P, and so belie his “‘ stets einzelwurzelig.” I have seen two specimens of burtoni from Eirerib,
Sudan ; one of shoana from Shoa ; the type of butleri from Jebel Fazogli, and one of jacksoni from
Nakusa, B.E.A., all of which have P, two-rooted and strong. These specimens are all in stage VIII,
so there is no question of P, being mistaken for P,.
184 NoviratEs ZooLoGIcAE XLI. 1939.
Hyraxes are no longer kept in captivity,! but their great variability may be
due in part to feral forms of ancient domestic stocks.
I therefore propose to wait till more specimens come to hand before adding
more names to this already crowded genus.
3. Capra hircus sinaitica Hempr. & Ehrenb.—Ibex.
Capra sinaitica Hemprich & Ehrenberg 1830 (1833), Symbolae Physicae Mammalium, 2: folio kk
recto, type loc.: Upper Egypt and Sinai; Yerbury & Thomas, 1895, Proc. Zool. Soc. Lond., p.
555, Aden ; Thomas, 1900, Proc. Zool. Soc. Lond., p. 104, Abyan Mts.
Capra Mengesi Noack 1896, Zool. Anz., 19: 353, type loc.: Hadhramaut; Noack, 1897, l.c., 20:
365 ; Sclater, 1897, Proc. Zool. Soc. Lond., p. 900 ; Noack, 1899, Zool, Anz., 22: 13.
Capra hircus nubiana F. Cuv., Schwarz, 1935, Ann. Mag. Nat. Hist., 16: 435.
Four specimens, horns only. Collector’s number 100b, 3, Najran, July
1936, outside curve of horns 34 and 35 inches (86 and 89 centimetres). Two
$3, no numbers or locality, outside curve of horns 27 inches (68-5 centimetres)
in one head and 221 inches (56-5 centimetres) in the other. One 9, no
number, Jabal Tuwaig (near Riyadh), September 1936, outside curve of horns
8 and 83 inches (20-3 and 21-6 centimetres).
Gazella Blainville 1816—Gazelles.
Mr. Philby has brought back a fine series of gazelles which give a clearer
understanding of the differences between the “ Idmi”’ gazelle (arabica) and the
“ Afri ” gazelle (saudiya), differences which are at first sight not apparent in
individual specimens unless they are at the extreme ends of their variation range.
There are also specimens of the horns of the “ Rim ” gazelle (marica) in this
collection, but unfortunately no skins.
The distribution of these three gazelles is interesting. This is what Mr.
Philby writes: “ Idmi passim in all suitable country, i.e. valleys of foothills,
open rock or sand plains and in true sand areas (nufud), but not so plentiful in
sand tracts as elsewhere. Afri confined (?) to sandy plains and overlapping into
sand areas. Rim only in sand areas and fringes of sandy plains near them.” So
from “idmi ” to “rim” represents a progressively more barren habitat, and
this is correlated with an increase in horn length from the former to the latter
(see Pl. VI). It seems to be only partly correlated with a lightening of coat
colour because though “ afri ” is lighter than “ idmi ” yet “rim ”’ has a darker
body 2 colour than “ afri.”
T have avoided following Pocock and Schwarz in calling the Arabian gazelles
subspecies of Gazella gazella because though arabica and saudiya may often be
seen on the same ground they appear to be distinct animals and the herds (of from
4 to 20 individuals) seem never to mix. Further, such a procedure would not
express the inter-relationships of the Arabian gazelles as I see them. I have
left the question of subspecies open for the moment and have called each form a
“species in an arbitrary manner.
The following is my interpretation of the available material: saudiya is
related to arabica, the former being a desert form and the latter a not-so-desert
form ; marica bears the same relationship to muscatensis. These two latter
gazelles show the lyrate shape of horn in the gg which is lacking in the two
1 Philby in litt., 21. vi. 1937.
2 “Rim ” has, however, a paler face than “‘ afri.’
ce
Noviratrs ZooLogIcAE XLI. 1939. 185
former, and marica has longer and more slender horns than muscatensis in the
same way that saudiya has longer and more slender horns than arabica. There
is, however, a ¢ gazelle (B.M. 34. 8. 4. 29), collected by Bertram Thomas at
Jurbaib, which is intermediate between arabica and muscatensis so far as the
horns are concerned.
The only other Arabian gazelles I have seen are from Aden. The horns of
these Jg are much shorter, stouter and straighter than those of arabica illus-
trated in Lichtenstein’s and in Hemprich & Ehrenberg’s plates and than
Philby’s specimens, and I do not quite agree with Pocock (1935a) who, considering
the skins only, said that he did not think the Aden specimens were sufficiently
different from the typical ones to merit subspecific rank. I am not at all sure
that Neumann’s name erlangeri should not stand for the Aden specimens.
It is to be hoped that more specimens of Arabian gazelles will be collected,
more especially of marica, and that no time will be lost. Motor-cars and modern
rifles are pressing hard on these animals and they may soon be wiped out.
4. Gazella arabica Hempr. et Ehrenb.
Antilope arabica Hempr. et Ehrenb., M. H. C. Lichtenstein 1827-1834 (1827), Darstellung Säugethiere,
pl. vi and text, type loc.: Hast coast of Red Sea.
Antilope arabica Hemprich et Ehrenberg 1828 (pl. 1828, text 1833), Symbolae Physicae Mammalium,
1: folio r verso.
17 gg and 2 22 from western and southern Arabia.
Native name—“idmi.” Hemprich & Ehrenberg have a note: “ Striatam
Antilopam, quam Arabum scriptores et Lexicographi Adm vel Idma vocant et
ex cuius nomine Shaw Lidmeae nomen male fabricasse videtur (nam el Idma Lidma
pronuntiari solet), ad nostram Antilopam arabicam spectare, nullus dubito.” On
the other hand, native names of gazelles should be regarded with suspicion. For
instance, the Atlas Gazelle (G. cwvieri Ogilby) is also known as ‘“‘edmi” and
Lichtenstein says that the natives call arabica “ariel.” Moreover, in Arabia
there seems to be some confusion between the names “ afri ” and “rim ” (see
under saudiya and marica), and Loder’s Gazelle (G. leptoceros loderi Thos.), a
Tunisian form, is also known as “ rim.”
There has been some difference of opinion as to the type locality of arabica.
Lichtenstein, who first published the name from Hemprich & Ehrenberg’s manu-
script says: “ . östliche Küste des rothen Meers .. . Sie bewohnt dort die
Vorhöhen in der Nähe der Küste und findet sich sogar auch auf den derselben nahe
liegenden grösseren Inseln, z.B. auf der . . . Insel Farsan.” Since no restricted
locality is mentioned by name save Farsan, some writers have taken this as the
type locality. But Lichtenstein does not emphasize this island more than the
mainland, and when we turn to Hemprich & Ehrenberg we find: “... ex
vallibus sinaiticis, Arabia deserta littorali, et ex insula Maris Rubri Farsan,
Arabiae felicis urbi Gisan proxima.” The plate which goes with this text has
“Sinai ” on it, so some writers have taken this to be the type locality. In the
absence, however, of any evidence as to which skin the original description was
based on there seems little point in trying to pin down the type locality any closer
than “‘ east coast of the Red Sea.”
Mr. Philby’s fine series shows considerable variation in coat colour and in
shape of horns, and with this new material before me I am unable to agree with
186 NoviratEes ZooLogIcAE XLI. 1939.
Pocock (1935a, p. 460), who believed that the differences which he observed were
seasonal.
The dorsal colour is a sandy brown with a slaty tinge, the hairs being
grey at the base. The flank band is variable, being a dark brownish slate in
some and in others only slightly darker than the dorsal colour; there are
intermediate conditions. Not only are these variations apparently unconnected
NUMBER
OF 5
SPECIMENS
i}
35335 BAÄRERE TR RL GEAR ES BU SE EE GD
= CONDYLOBASAL LENGTH CMM)
NUMBER
OF Wa
3 SAUDIYA
SPECIMENS Z
4
ARABICA m)
3
2
I
Swe wm ae 2
% Gb ie @
a Rey AG NEE NE
1-5
bo-s
[7
®
58-5
~
55.5
Ss
>
+
[0
53-5
BREADTH ACROSS HORN-CORE BASES CMM.)
Histograms of two skull characters of adult 5d of Gazella saudiya and @. arabica.
with seasonal change, but I am unable to correlate them with sex, age or
locality.
The horns of the gg are very variable, some being stoutly built, short and
strongly sigmoid, and others being relatively long and straight (see Pl. VI).
Some of the horns are like those of @. saudiya (“‘ afri ”’), but contrary to what might
have been expected this similarity is not always found in the specimens with the
lightest coloured coats. In the plate which accompanies the description of
saudiya the authors have selected the shortest, stoutest and most hooked
horns to illustrate arabica and the longest, most slender and straightest to
illustrate saudiya, thereby giving the impression that the two forms can
be easily distinguished by their horns which, as stated above, is not always
the case.
The following characters, however, serve to distinguish these two forms :
3 skulls.
G. arabica G. saudiya
1. Premaxilla does not make a long 1. Premaxilla makes a long junction
junction with nasal (text-fig. 126 A). with nasal.
2. Width of combined nasals usually 2. Width of combined nasals usually
greatest at proximal end. greatest at distal end.
3. Breadth across the bases of the horn- 3. Breadth across the bases of the horn-
cores (teat-fig. 126 B and the histo- cores 53:3-60-7 mm.
gram) 62-2-69-0 mm.
4. Larger, heavier skulls, condylobasal 4. Smaller, lighter skulls, condylobasal
lengths 171-190-5 mm. (see the his- lengths 166-173 mm.
togram).
Novirates ZooLoGIcAE XLI. 1939. 187
© skulls.
5. Horns have sigmoid curve of the 5. Horns either straight! or with
males. Length in a straight line a scimitar-like backward curve.
about 100 mm. Length in a straight line 200-250
mm. -
126 a
zZ
. TexT-FıG. 126.—(A) The rostral regions of the skulls of (i) Gazella arabica and (ii) G. saudiya.
(B) The character referred to in the text as ‘“ breadth across the bases of the horn-cores ” is indicated
by A-A.
Skins (both sexes).
It is difficult to distinguish between saudiya skins and those skins of arabica
which have the flank band almost absent, as in these cases the tips of the hairs
are the same colour. But the bases of the hairs are still different :
G. arabica G. saudiya
Bases of hairs smoky to slate. Bases of hairs lighter than in arabica,
greyish pink or pinkish sandy to
pinkish slate.
Mr. Philby tells me that these two forms are easily distinguished by coat
colour in the field. Presumably in life the hairs stand out in such a way as to
expose the bases somewhat.?
1 The horns of 2 saudiya are used by Arab women as hair curlers.
2 Since this was written I have received a letter from Mrs. Dickson at Kuwait, who had been
told the following by a Badawin. ‘‘ Rhim ” is the biggest and most handsome of the gazelles and
their meat is the best to eat. The face has white cheeks with a band of sandy red going down the
centre of the face and merging into a black nose. The horns are not very long. Two other gazelles
are: “ Idmi,” which is reddish in colour with less white on the belly and “‘ Afri,” which is the smallest
of the lot. These three are distinguished by their colour and size, as the length of the horns and
ears in each are similar.
188 NOVITATES ZOOLOGICAE XLI. 1939.
Measurements of skulls of adult 3 Gazella arabica :
5 Breadth acros:
Number. Locality. en Zygomstic (etraentin e). nano a
A Rakba Plain (?) 187-5 80-4 238 65-3
B Rakba Plain (?) — 81-4 241 65-6
12 Rakba Plain 174-5 81-8 210 64:9
13 Rakba Plain 171 77:5 237 62-2
14 Rakba Plain 188-5 84-0 200 69-0
15 Rakba Plain — 80-1 230 65-2
35 Dhalm 179-5 | 82-5 256 65-9
70 Hudhaiyib al Aranib 183 | 84-0 204 65-2
123 Taraf al Ain 132-5 81-8 252 62-5
128 Ugqla Plain | 190-5 86-7 221 68-8
130 Haraj Plain 177 80.3 | 224 64-1
138 Sudaihat 182 82-1 226 64:3
B.M. 69. 10. 24. 100 | Mocha 183 84-5 205 64-6
B.M. 96. 12.4. 1! Aden (? juv.) — | 83-8 215 60-2
B.M. 96. 12. 4. 21 Aden | — | — 235 66-1
5. Gazella saudiya Carruthers & Schwarz.
Gazella gazella saudiya Carruthers & Schwarz 1935, Proc. Zool. Soc, Lond., p. 155, type loc.: Dhalm.
Native name—“afri.”’ There is a certain amount of confusion as the names
“rim ” and also ‘“ khadhri ” seem to be applied to saudiya occasionally. More-
over the name “‘rim”’ is also used for Loder’s Gazelle, a pale-coloured, long-
horned form from N. Africa. For the true “ rim ” of Arabia see under marica
below.
Mr. Philby has sent six 33 and nine 22 from Dhalm, Jau al Khudaif, Taraf
al ‘Ain, Sirr al Yamani, W. Markha, Ruwaik Tract, ‘Alam Abyadh and ‘Arq Abu
Da’ir.
The 98 are hard to distinguish from those 33 of arabica which have the
flank band obsolescent and have long horns. For the characters which may be
used to separate saudiya from arabica see above under G. arabica. The 2%
are easily distinguished by the fact that they have horns at least twice as long
as those of 9 arabica.
6. Gazella marica Thos.
Gazella marica O. Thomas 1897, Ann. Mag. Nat. Hist., 19: 162, type loc.: Nejd, Central Arabia ;
Lydekker & Blaine, 1914, Cat. Ungulate Mamm., 8: 60; Cheesman, R. E., 1920, J. Bomb. Nat.
Hist. Soc., 27 : 343 ; Pocock, R. I., 1935, Ann. Mag. Nat. Hist., 15: 463.
Native name—‘rim.” To this species I assign a 3 frontlet complete with
horns (Pl. VI, fig. 14) which Mr. Philby found half buried in sand at Afalil on 30
September, 1936, collector’s number 136, and also several odd horns without
locality.
The identification of this species is not easy because the type is an immature
¢g and the only mature 3 which Thomas saw had its horns sawn off leaving
stumps 100 mm. long ; the ends were missing. However, Dr. Jayakar, who sent
the type to Thomas, described it as a “‘ reem ” gazelle, which agrees with the name
“yim ” supplied by Mr. Philby, but what is more important the proximal 100 mm.
of the horns of Philby’s specimen agree well with those of the sawn-off horns.
Philby’s horns are lyrate in shape and the mean length (measured along the
outside of the curve) of the two horns of the Afalil specimen is 336 mm. Thomas
1 Frontlets only.
Novrrares ZooLoGIcAE XLI. 1939. 189
had estimated that the horns of the sawn-off specimen would have been about
250 mm. long on the outside of the curve, but too much attention need not be
paid to this seeing that he had never seen an adult head complete. ‘The distance
between the tips of the horns of the Afalil specimen is 93 mm. and the maximum
outside breadth of the “lyre ” is 174 mm.
Lepus L.—Hares.
Twenty-three hares (in addition to those sent on previous occasions) from
south-west Arabia.
The Arabian hares have affinities with those of Egypt and are quite different
from those of Syria, Palestine and Persia, which are larger animals with darker
coloration and longer fur and have the characteristic “‘ long hairs,’ which pro-
trude far beyond the general coat, much more pronounced ; these hairs are usually
absent from Egyptian specimens and always from Arabian ones.
The zoogeographical affinities of the hares are thus different from those of
the foxes (Vulpes vulpes), which seem to be more closely related to those of
southern Persia, Baluchistan and India than to Egyptian foxes.
Arabia produces three hares, a sandy-coloured one and two dark ones, the
first being found in very barren country and the last two in not-so-barren country.
The particular interest of this latest collection is that it contains seventeen speci-
mens of one of the dark forms which I have little hesitation in referring to Lepus
arabicus arabicus.
7. Lepus arabicus arabicus Hempr. et Ehrenb.
Lepus arabicus Hemprich & Ehrenberg 1830 (1833), Symbolae Physicae Mammalium, 2: folio r recto,
type loc.: “ Ex Arabia Deserta prope Gumfudam! et ex Arabia Felici prope, Loheiam” ;
Yerbury & Thomas, 1895, Proc. Zool. Soc. Lond., p.555 ; Thomas, 1900, Proc. Zool. Soc. Lond.,
p. 104.
Seventeen ? specimens from that part of the west coast hill-country which lies
between latitudes 16°-22° N.
There is a curious feature in some of these hares, particularly No. 42, a 2
from Shariya, 17 September, 1934. The hairs of the back are worn in places
and some of these patches are rust-coloured. The unworn hairs are not rust-
coloured at any point along their length. Perhaps Hemprich and Ehrenberg are
alluding to this when they say: “ vellere cinereo aut fulvescente nigro rufoque
variegato.”
The following are the measurements in millimetres of six adult gg and
seven adult 29 of arabicus :
[
Extremes. Average. Standard deviation.
|
3 2 3 | 2 3 2
Head and body (collector’s) . 6 338 -410 323 -402 381 | 368 24:6 25:6
Hindfoot (in skin) 6 6 3 89-5— 99 86-5- 97-5 94 91 3:3 3:6
Ear (in skin) 5 6 ö 6 91 -ı18 87 —-112 105 101 | OG 8-1
Skull (over all) . 6 6 0 76-3— 83:7 78:9— 82-6 | 80-4 80-7 | 2-4 1-2
1 The modern spelling of Gumfuda is Qunfidha.
2 Mr. Philby has sent an extra specimen, 2, number 151, from Wadi Luwai (25° 0’ N. 40° 39’ E.)
dated 22.v.1938. Head and body 388, hindfoot 90, ear 110 millimetres. Skull broken.
190 Noviratrs Zootocrcan XLI. 1939.
I am not sure whether L. omanensis Thos.1 should not be regarded as a race
of arabicus. Thomas says of omanensis : ‘“ This interesting little hare is at once
distinguished from all other allied members of the genus by its slender proportions
and its extremely small size. Its hindfoot is nearly an inch less than in any
species described by Hemprich & Ehrenberg. . . .” With regard to Thomas’s
statement about the hindfoot Hemprich & Ehrenberg give 4 German inches as
the “planta ’’ measurement for arabicus, and this is only 16 mm. longer than
Thomas’s measurement for omanensis. Further, it is probable that Hemprich &
Ehrenberg included the claws which Thomas did not. The type of omanensis, a
©, B.M. 94. 3. 9. 36, has head and body 360, hindfoot 89, and ear 99, all of
which measurements fall within the limits of the above specimens collected by
Mr. Philby.
The skull of the type of omanensis does seem on the small side with an over-all
length of 74-5 mm. This may be due to a sub-adult condition and I should like
to see more specimens ; only three have been recorded so far.
8. Lepus arabicus cheesmani Thos.
Lepus omanensis cheesmani Thomas 1921, J. Bombay Nat. Hist. Soc., 28: 28, type loc.: Dohat al
Salwa ; Pocock, 1935, Ann. Mag. Nat. Hist.,15: 445.
Though it is an open question whether omanensis should be regarded as a
race of arabicus or not, it is now clear that cheesmani is a desert form of
the latter. The transition from the dark igneous formation of the west coast
to the light sandstone of the interior is quite abrupt and along the dividing
line are found the lighter coloured of the arabicus and the darker of the
cheesmant.
Six specimens. The following are the measurements in mm. (the head and
body length was taken in the flesh, and the hindfoot and ear in the skin) :
Head and body. Hindfoot. Tar. Skull (over all).
| us|
3 | 109 Wadi Khabb 4,200 ft. 386 934 1104 82:5
Q 110 Wadi Khabb 4,200 ft. 328 85 844 68:3
3} 90 Husainiya 4,000 ft. 362 85 | 111 81-0
ie) 91 . Husainiya 4,000 ft. | 423 90 99 81-4
3 116 Mushainiga 3,700 ft. 360 96 1134 =
3 Type Dohat al Salwa 380 84 94 70-5 2
Mr. Philby’s number 90 (killed in June) agrees with the type (killed in
March) save that it is a little pinker * and has larger ears. Number 109 (killed
in July) is similar. Number 110 (killed on the same day as 109) is pure sandy-
coloured with none of the buff of the type and numbers 91 (killed in June) and
116 (killed in August) are pink-sandy.
So these hares seem variable not only in ear and hindfoot length, but also
in colour. These specimens fit in well, however, with those which Mr. Philby has
sent from other parts of Arabia and which have been discussed by Pocock (1935,
p. 445).
1 O. Thomas 1894, Proc. Zool. Soc. Lond., p. 450, type loc.: Ziki in Oman.
_? This is Thomas’s measurement—I make it 71-1.
3 Buxton (1923, p. 146) mentions seeing a pinkish buff hare at Qazvin in N.W. Persia.
Novirates Zoonoaican XLI. 1939. 191
9. Jaculus jaculus L.—Jerboa.
Mus jaculus Linnaeus 1758, Syst. Nat., ed. x, p. 63, type loc.: “ Aegyptus, in montibus Aegyptum
ab Arabia dividentibus ”—see Hasselquist, Acta Scientiarwm Upsaliensis, 1744-1750 (1751),
p. 17.
Mr. Philby has sent seventeen specimens, numbers 78-84 from Raushan and
73-76 from Dailami (both in Wadi Bishe), 58-60 from Rass (N.W. of Riyadh),
113-114 from Matau (S.W. Rub al Khali) and number 13 from Jidda.
The more I look at these jerboas the more difficult it becomes to decide what
to call them. The types of J. florentiae Cheesman & Hinton 1924, from Jabrin,
J. f. oralis of the same authors, from Koweit, and J. loftusi Blanford 1876, from
near Basra, can all be lost among the Raushan specimens alone.
These animals seem to be very variable and I can find no constant features
in the skins or the skulls. Their general dorsal colour varies from ashy-buff to
sandy with a pinkish tinge. The only remarks I can make are of a general
nature ; the Matau specimens are on the whole perhaps a little yellower than the
Raushan and Dailami ones and the Rass specimens are on the whole rather darker
(though the darkest Raushan specimens are as dark as the Rass ones). The Jidda
specimen is an exact match of the type of J. loftws:.
Jaculus jaculus is very plastic so far as colour is concerned and turns up pale
in very desert areas and dark in not so-desert-areas. For instance, the type of
J. j. airensis Thomas & Hinton 1921, from the Air Mts. in French West Africa,
is strikingly like the type of J. florentiae and specimens collected by W. Ruttledge
in Kordofan in 1929. These localities are widely separated and there are darker
forms of jerboa in the intervening country.
The pale and dark forms of Jaculus jaculus are not distributed according to
conventional ideas of geographical races, but have a mosaic distribution. The
conventional system of according separate names to each separated group
overlooks this fact and has led to far too many descriptions.
10. Acomys dimidiatus Cretz.—Bicoloured Spiny Mouse.
Mus dimidiatus Cretzschmar in Rueppell 1826 (1827), Atlas Reise Nördl. Afrika, Pt. 1, p. 37, type
loc.: Sinai.
Acomys dimidiatus Cretz., Sundevall, 1842 (1843), K. Svenska Vetensk. Akad. Handl., p. 222.
One specimen, collector’s number 168, g, Harra Rahat, 4,000 ft., 19
November, 1938. Head and body 103, tail missing, hindfoot 18, ear 19, total
length of skull 30-4, condylobasal length 28-2, zygomatic breadth 14-2, maxillary
alveoli 4:7, mandibular length 14-6.
11. Acomys russatus Wagner—Golden Spiny Mouse.
Mus russatus Wagner 1840, Abh. Bayer. Akad. Wiss., 3: p. 195, pl. 3, fig. 2, type loc.: Sinai.
Three specimens from the Najran district. Number 142 is the same colour
as the specimen depicted in Wagner’s plate,! but the other two are darker, i.e.
they have more grey-brown about them. Also a specimen (2?) from ‘Aqda,
near Hail, number 154, June 1938.
These specimens extend the range of russatus in a southerly direction, no
specimens having previously been recorded from peninsular Arabia. Thomas
described Acomys dimidiatus homericus from El Khaur, Aden (Ann. Mag. Nat.
Hist. 12. 1923. 173), but russatus is quite distinct. The latter has a relatively
1 The British Museum’s copy.
15
192 NoviratEes ZooLoGICAE XLI. 1939.
small skull, the tail is shorter than the head and body and the belly is a dirty
grey, whereas homericus has a larger skull, the tail is 10-20 mm. longer than the
head and body and the belly is white.
Mr. Philby’s notes say, “ Skulking among granite rocks ” ; the measure-
ments in mm. are :
Oollec- | | Head Hind Total | Condylo- | Zygo- Mandi-
tor’s | Sex. | Locality. and | Tail. foot Ear. length basal matic bular Date.
number. | | ody. | of skull. | length. | breadth. | length.
142 3 | Hadhba 100 GQ |. 17 154 26-8 _ 13-2 14-7 10 Oct., 1936
144 3 | Shaib Arjan} 106 57 16 18 26-4 — — 14-4 5 Nov., 1936
208 3 | Najran 111 64 17 164 28-2 26-0 14:0 15:0 22 Oct., 1936
12. Rattus rattus L.—Rat.
Two specimens. Collector’s number 1, 9, Jidda, 27 January, 1934, and
number 86, 3g, from Suda Village, near Abha, 9 June, 1936. The skull of
number 86 was too damaged to preserve.
These rats are not R. r. flaviventris or R. r. frugivorus, but may be R. r.
alexandrinus. The dorsal surface is grey-brown in both and the belly of number
86 is grey with a cinnamon wash, whereas the belly of number 1 is grey-white.
13. Psammomys obesus dianae subsp.n.—Miss Philby’s Fat Sand-rat.
Type locality —Dailami, 20° 20’ N. 42° 40’ E., 3,900 ft.
Type.—Adult 3, collector’s number 77, 26 May, 1936, in the British Museum.
Paratypes.—Juv. 9, collector’s number 10, Samuda, 21 June, 1934. Juv.
3, collector’s number 36, Jidda, 28 August, 1934. Adult 9, collector’s number
63, Dawadami, 9 March, 1935. Adult g, collector’s number 71, Dailami,
26 May, 1936.
Description.—Pale ashy sandy colour; rusty yellowish tinge dorsally,
strongest on head and after-end of back ; yellowish tinge on the flanks. Yellow
patch on the shoulders. Belly greyish white with a tinge of yellow on the middle
line. Feet, hands and tail pale ashy sandy ; tip of tail pencilled on the dorsal
side only—pencil black.
Paler than P. obesus Cretz.! and P. terrae-sanctae Thos., 1902 (type loc. : Dead
Sea) ; rather paler even than P. vexillarıs Thos., 1925 (type loc. : Bondjem, Tripoli).
Measurements in mm.—Considerably smaller than P. terrae-sanctae. Type :
head and body 133, tail 118, hindfoot 34, ear 14, condylobasal length 34-50,
zygomatic breadth 20-65, maxillary tooth row (alveolar length) 6-24, diastema
(upper) 10-21, interorbital width 6-14, nasals (maximum length) 14-10, depth of
rostrum at level of anterior end of tooth row 10:06, mandibular length 20-46,
greatest diameter of bulla 14-00 (this last measurement was taken from number
71, since the bullae of the type are missing).
Meriones Illiger 1811—Jirds.
These are rat-like animals about 250-300 mm. (10-12 inches) long, about
half of which are tail. They are mostly ‘ desert-coloured ” and the tip of
1 Cretzschmar in Rüppell, Atlas zu der Reise im nördlichen Afrika—Säugethiere, 1826 (1828).
Type loc. ; Alexandria,
Novirares Zoonoaican XLI. 1939.
193
the tail has a dark brush.
in jerboas.
The following forms have been described from Arabia :
1. M. longifrons Lataste 1884 (type loc. : Jidda) is a smallish form, medium
pale.
2. M. buryi Thos. 1902 and M. rex Yerbury & Thomas 1895, both from
Aden, are very dark forms.
3. M. pelerinus Thos. 1919 (type loc.: Tebuk, on the Hedjaz railway) has
very much enlarged bullae. Aharoni (1932, p. 197) makes this species a synonym
of M. crassus Sundevall 1842, described from Sinai.
4. M. arimalius Cheesman & Hinton 1924 (type loc.: Jabrin) is the palest
of all the Arabian Jirds.
5. M. ismahelis Cheesman & Hinton 1924 (type loc.: Hufuf) is a pale form
with very large bullae like pelerinus.
6. M. erythroura edithae Cheesman & Hinton 1924 (type loc.: Khudud
Spring, Hufuf) is a darkish form with a reddish brown tail. For M. erythroura
evelynae of the same authors see below under M. erythroura edithae.
The character of the large bullae is shown in text-fig. 127. (A) shows a
form with small (comparatively speaking) bullae and (B) shows a form with
larger bullae which project well beyond the occipital region of the skull.
The hind legs are not modified for jumping as
14. Meriones erythroura ‘ edithae Cheesman & Hinton—Edith Cheesman’s Jird.
Meriones syrius? edithae Cheesman & Hinton 1924, Ann. Mag. Nat. Hist., 14: 555, type loc. :
Khudud Spring, Hufuf ; Cheesman, 1926, In Unknown Arabia, p. 360 (a reprint of the original
description with additional matter).
Thirteen specimens. Those from Riyadh and Khufaifiya, and some of
those from Shari Wells, closely resemble edithae, though their tails are not
quite so long? (the description says that
the tails of edithae are “ ochraceous [27 i NEN 5
salmon,” but I prefer to describe them -
as reddish brown; I have examined
the original series and I can see no ;
salmon about them). The rest of the
Shari Wells specimens exactly match
the paler-coloured form M. syrius
evelynae * Cheesman & Hinton, but it
is significant that these specimens are a
not fully grown. The skulls of the
type and paratypes of evelynae have
the basi-occipital/basi-sphenoid suture
still visible as have the pale coloured
Text-Fie. 127.—(A) Meriones erythroura
edithae, collector’s number 163, Shari Wells.
(B) M. ismahelis, collector's number 158,
Shari Wells.
Shari Wells specimens, but the type of edithae and the darker Shari Wells
specimens have this suture obliterated.
1 Gerbillus erythroura Gray 1842, Ann. Mag. Nat. Hist., 10: 266, type loc. :
With this in mind and the fact that
Afghanistan.
2 Meriones syrius Thomas 1919, Ann. Mag. Nat. Hist., 8 : 268, type loc.: Karyatein.
3 The type of edithae has head and body 143 and tail 181, and the largest of Mr. Philby’s speci-
mens has head and body 162 and tail 173, so it may be that the longer tail of the former is due
to differences in styles of measurement and making up.
4 Meriones syrius evelynae Cheesman & Hinton 1924, Ann, Mag. Nat, Hist., 14: 555, type loc. :
Khorosan Spring, Hufuf,
194 NOVITATES ZOOLOGICAE XLI. 1939.
the Shari Wells specimens were all caught on the same day (see the table
of measurements) it looks as though evelynae turns into edithae with age.
The following are the measurements in millimetres :
Meriones erythroura edithae
| Collec- | | Head rn Total | Condy-| Zygo- | Max. | Mail Aandi-
Sex. | tor’s Locality. Date. and | Tail. foot, Ear. | length | lobasal| matic | diam. a bular
number! body. a skull. | length. breadth| bullae.| $y), | length.
3 43 | Riyadh 27.1x.1934 | 136 161 34 17 — — )214| — 6-4 | 20-5
3 49 | Khufaifiya | 13.xi.1934 | 160 162 39 21
3} 50 | Khufaifiya | 14.xi.1934 | 162 173 37 21 42-2 | 39:0 | — | 16-5] 6-8 | 22-1
2 51 | Khufaifiya | 14.xi.1934 | 148 140 34 18 40-2 | 37-2 | 21-8 | 15-3 | 6-6 | 20-1
3 | 155 | Shari Wells 4.vi.1938 | 148 156 38 17 — — — 5-7 | 16-5
3 | 156 | Shari Wells} 4.vi.1938 | 1504 | 149 36 19 — — | 23:0} — 6-9 | 21-8
& | 157 | Shari Wells | 4.vi.1938 | 132 1454 | 37 18 = — | 20-5 | 16:0] 64 —
& | 159 | Shari Wells) 4.vi.1938 | 1284 | 142 | 33 17 — | — [19-1] | — | 5:8 | 18-8
3 | 160 | Shari Wells | 4.vi.1938 | 111 122 324 | 14 — — — — 5:6 | 16-4
& | 161 | Shari Wells| 4.vi.1938 | 111 1174 | 324 | 174 | — | — /212|161|56 | —
2 | 163 | Shari Wells} 4.vi.1938 | 149 147 35 19 37-6 | 34-3 | — | 15-2] 6-0 | 18-6
& | 164 | Shari Wells; 4.vi.1938 | 135 1384 | 35 15 39-4 | 36-5 | 22-3 | 15-6 | 6-4 | 20-3
& | 165 | Shari Wells | 4.vi.1938 98 111 32 14 30-8 | 27-7 | 16-5 | 12-3 | 5-4 | 15-4
15. Meriones ismahelis Cheesman & Hinton—Nocturnal Jird.
Meriones ismahelis Cheesman & Hinton 1924, Ann. Mag. Nat. Hist., 14: 553, type loc.: Hufuf
Oasis ; Cheesman, 1926, In Unknown Arabia, p. 357 (a reprint of the original description with
additional matter).
Two specimens from Shari Wells, 4 June, 1938. Collector’s number 158,
&, head and body 111, tail 120, hindfoot 29, ear 16, total length skull 36-9,
maximum diameter of bullae 16-4, maxillary alveolae 5-4, mandibular length 17-5
millimetres. Collector’s number 162, 3, head and body 111, tail 122, hind-
foot 31, ear 164, maxillary alveolae 5-5, mandibular length 17-6 millimetres.
Dipodillus Lataste 1881—Gerbils.
The systematics of the Arabian gerbils of this genus are in a rather unsatis-
factory condition, so a few notes on some of them may be useful.
D. dasyurus Wagner 1842, Arch. Naturgesch. Berlin, Jahrg. 8, Bd. 1, p. 20,
type loc.: West coast of Arabia.
Wagner gives “ Schwanz 3" 011 ” but Thomas, who inspected the type, has
“Tl. (c) 100 ” (in MS.). Since Miss Aharoni (1932, p. 206) says that the tail of
the type is broken, I take it that Thomas’s MS. note means that the tail must
originally have been about 100 mm. long: that is, about 4inches. So it looks as
though Wagner had overlooked the fact that the tail was broken when he described
it as being as long as the head and body, and this has led to confusion. As
a matter of fact dasyurus has the tail considerably longer than the head
and body and with a good pencil. The name dasyurus suggests a rough-
haired tail but poecilops has a rougher tail. The bullae in dasyurus are
relatively small.
D. famulus Yerbury & Thomas 1895, Proc, Zool. Soc. Lond., p. 551, type
loc. : Lahej.
This species has very large ears and a very long tail, with a much bushier
pencil than the other Arabian Dipodillus. The bullae are large and when viewed
Noviratres ZOOLOGICAE XLI. 1939. 195
dorsally project backwards beyond the foramen magnum, though perhaps not
quite so markedly as in D. arabium.
D. lixa Yerbury & Thomas 1895, Proc. Zool. Soc. Lond., p. 550, type loc. :
Shaik Othman.
Not a satisfactory species : based on immature specimens. The authors say
that the type is “a slightly immature 9.” I have examined the skull and
would not have qualified the word “immature.” The teeth of gerbils present
an adult appearance very early, even at irruption, due to the fact that the
crowns of the molars have little or no enamel on them at the apices of the
tubercles.
D. mimulus Thomas 1902, Ann. Mag. Nat. Hist., 9: 362, type loc.:
Lahej.
This species bears a strong resemblance to poecilops, but is considerably
smaller.
16. Dipodillus arabium Thos.—Arabian Gerbil.
Dipodillus arabium Thomas 1918, Ann. Mag. Nat. Hist., 2: 61, type loc.: Tebuk, Hedjaz Railway.
One specimen, collector’s number 12, 9, from Ashaira, 4,000 ft., 21 March,
1934. The tail is browner than that of the type and the tail-pencil is a little
fuller, but not so fullasin famulus. Otherwise there is goodagreement. Measure-
ments in mm.: head and body 95, tail 125, hindfoot 25, ear 14, over-all length
of skull 29-4, condylobasal length 26-1, maximum diameter of bullae 12-0,
diastema 7-4. The bullae are large and in dorsal view project backwards beyond
the foramen magnum.
17. Dipodillus poecilops Yerbury & Thos.—Spiny-tailed Gerbil.
Dipodillus poecilops Yerbury & Thomas 1895, Proc. Zool. Soc. Lond., p. 549, type loc.: Lahej.
Three specimens from Hadda. 4, collector’s number 8, 2 March, 1934,
and &, No. 66 and @ juv., No. 67, 15 Feb., 1936. These have pencil-less tails
with spiny hairs. The bullae, viewed dorsally, do not project backwards beyond
the foramen magnum.
18. Gerbillus arduus Cheesman & Hinton—Jafura Gerbil.
Gerbillus arduus Cheesman & Hinton 1924, Ann. Mag. Nat. Hist., 14: 551, type loc.: Jafura.
The following skins and skulls were collected by Mr. Philby :
120 & Ramlat Sab’atain . . 7.viil.1936
124 & Ras al Khabb F . 19.ix.1936
128 © Sahel 5 020711936
126 ¢ Sahlal . 6 5 . 20.1x.1936
135 — ‘Arain . é ; . 30.ix.1936
These specimens agree well with the typical series and extend the known
range of the species in a south-southwesterly direction. Pocock (1935, p. 444)
mentions a specimen from Hawiya some 180 miles south of the type locality, but
the above specimens come from localities some 650 miles from Jafura and on
the opposite side of the Rub al Khali.
196 NoviratEs ZooLoGICAE XLI. 1939.
Tatera Lataste 1882.
Wroughton (J. Bombay Nat. Hist. Soc., 25: 1917: 40) made a new genus,
Taterona, for the African gerbils formerly included in Tatera. He distinguished
the two genera as follows :
Tatera.—Tail dark above and below with a light-coloured stripe along each
side of it and the vertical part of the parieto-squamosal suture about the same
length as the anterior horizontal part thereof.
Taterona.—Tail dark above and pale below (save in nigricauda which has an
entirely black tail) and the vertical part of the parieto-squamosal suture about a
third the length of the anterior horizontal part thereof.
Wroughton gave other characters besides, but admitted that they were not
constant. Mr. Philby’s specimens are of great interest in that according to the
shape of the parieto-squamosal suture they would be put into T'atera, while
according to the tail character they belong to T'aterona. Hollister (Bull. U.S.
Nat. Mus., 99 : 1919: 28) has indicated that he does not consider T'aterona a good
genus and the above specimens, coming from a country where Indian and Ethio-
pian elements meet, provide a good link between Tatera and Taterona ; I intend
to drop the latter name.
19. Tatera philbyi sp.n.—Philby’s Gerbil.
Type.—Adult 9, collector’s number 95, Najran (17° 30’ N. 44° 20’ E.) 4,400
ft., 29 June, 1936, skin and skull.
Paratypes.—, collector’s number 38, Taif, 6 September, 1934; 92, number
67, Ashaira, 5 May, 1936; $, number 72, Dailami, 26 May, 1936; 3, number
85, Shaib Hanjur, 2 June, 1936; 9, number 93, Najran, 26 June, 1936.
Description.—General colour olivaceous buffy brown, head and spine rather
warmer tone than rest of back. Hairs of the back about 10 mm. long, basal two-
thirds slate-coloured, terminal third buffy brown, some tipped with black.
Under-parts dirty white with a yellow tinge in places. Hands, feet, ears and
dorsal surface of tail same colour as spine, ventral surface of tail lighter. The
skull is much like that of Persian and Indian specimens of T'atera.
Measurements.—Type: head and body 153, tail 157, hindfoot 36, ear 19,
condylobasal length 37-1, zygomatic breadth 21-0, greatest length and breadth
of combined nasals 16-7 x 4-7, anterior palatal foramina 7-1, upper diastema
10-4, crowns M!”3 5-4. The type is not the largest of the series. Paratype :
number 85, head and body 162, tail 160, hindfoot 38, ear 21, condylobasal
length 40-9. This specimen, though larger than the type, was rejected because:
the hairs of the back are badly worn.
Discussion.—Tatera philbyi is easily distinguished from neighbouring species
by the fact that the latter have dark tails with a light stripe down each side,
whereas philbyi has the tail dark above and light below. The two nearest neigh-
bours are T. taeniura Wagner 1843, described from Syria, and T. bailwardi
Wroughton 1906, type locality Karun River. T. philbyi is decidedly smaller than
T. taeniura and slightly smaller than 7’. bailwardi. The nearest African form is
T. robusta Cretz. 1830/31, described from Kordofan. This form is about the
same size as philbyi but lacks the olivaceous tinge of the latter and has the belly
almost pure white. Moreover there are differences between the skulls of robusta
and philbyi.
Novıtates ZooLoGIcAE XLI. 1939. 197
20. Caracal caracal Müll.—Caracal Lynx.
[73
Felis caracal Müller 11776, Natursyst. Suppl., p. 30, type loc.: . . en Barbarie, en Arabie & dans
tous les pays qu’habitent le lion, la panthére & l’once.”
Felis caracal Güldenstädt in Schreber, ©. Thomas, 1895, Proc. Zool. Soc. Lond., p. 548, Aden; O.
Thomas, 1900, l.c., p. 100, Lahej.
Native name— Tifa.”
A sub-adult 9, collector’s number 39, Qaim near Taif, 5,200 ft., 7 September
1934.
The skin is a sandy cinnamon dorsally with a silvery grizzling, the total effect
being a purplish-pink tinge on a sandy ground colour. The head is somewhat
darker. The outside of the ear and the ear-tuft are black with a white admixtute,
the rim of the ear and the hairs on the inside of the ear are white. The fore- and
hindlegs are the same colour as the back. The face has some white on each side
of the bridge of the nose and on the lower eyelids. The throat, armpits and groins
are white. The belly is the same hue as the back but several tones lighter ; there
are irregular cinnamon to brown spots on the chest. There are three irregular
brown transverse bars on the upper inner surface of each foreleg.
Measurements in millimetres: head and body 630, tail 227, hindfoot 157,
ear 74, over-all length of skull 110-7, condylobasal length 100-9, zygomatic breadth
76-2, inter-orbital width 29-3, maximum length of upper carnassial 14-35.
21. Felis constantina * syriaca Tristram—Wild Cat.
Felis syriaca Tristram 1867, The Natural History of the Bible, p. 67, type loc.: Syria.
Felis maniculata Cretz. in Riippell, Yerbury & Thomas, 1895, Proc. Zool. Soc. Lond., p. 547, Aden ;
Thomas, 1900, ibid., p. 100, Lahej.
Felis ocreata Gmelin, Pocock, 1935, Ann. Mag. Nat. Hist.,15: 455, Umm al Qurun.
Native name— “ Hyr-Birry ” (fide A. G. Griffin). The classical Arabic name
seems to be “ Hirr.”’
Three specimens :
(a) Adult 3 (but basi-sphenoid pre-sphenoid suture still open), collector’s
number 145, ‘Iraq Well, 4,300 ft., 6 November, 1936. The spots are very in-
distinct. The measurements in mm. are: head and body 498, tail 390, hindfoot
131, ear 56, over-all length of skull 98-6, condylobasal length 89-9, zygomatic
breadth 68-9, maximum length of upper carnassial 11-1, mandibular length 64-8.
(db) Adult © shot by A. G. Griffin 15 miles north of Jidda at sea-level,
7 January, 1938. There are no flesh measurements but Mr. Griffin writes: “ It is
the biggest I have seen.” The skin has the spots well developed. The skull has
the occipital region missing, but the mandibular length gives a comparison with
(a) and (b). The measurements in mm. are: zygomatic breadth 60-7, maximum
length of upper carnassial 10-2, mandibular length 58-8.
(c) Young 9, collector’s number 52, 15 miles N.E. of Jidda at sea-level,
1 October 1934. The skin of this kitten does not show the spots at all distinctly.
Skull measurements in mm. are: over-all length 82-9, condylobasal length 74-2,
zygomatic breadth 57-6, maximum length of upper carnassial 11-9, mandibular
length 56-2.
It will be seen that (c) has a slightly smaller skull than (b) but that its upper
1 For the authorship of caracal see Matschie, 1912, S.B. Ges. Naturf. Fr. Berl., pp. 55-67.
2 The name constantina (Forster 1780, Buffon’s Naturgeschichte, 6: 313) has line priority over
lybica, which appears on the same page.
198 Novirarrs ZooLoGicAaE XLI. 1939.
carnassial is much bigger. Further, (c) has P? well developed, whereas (6) has
no P?. Now (b) is by no means aged—the teeth being strong and unworn—and
I can find no explanation for its comparatively small teeth. It is possible that
it may be a feral form of domestic cat.
MONGOOSES
There are three mongooses known at present from Arabia.
22. Ichneumia albicauda G. Cuv.
Herpestes albicaudus G. Cuvier 1829, Régne Animal, 2nd edit., 1: 158, type loc.: Senegal.
Ichneumia albicauda G. Cuv., I. Geoffroy, 1837, Ann. Sci. Nat. Paris (Zool.), 8: 251.
This is a large mongoose with coarse hair and a white tail. Head and body
about 184 inches (470 mm.), tail about 14 inches (358 mm.). It has been recorded
from the Muscat district (Thomas, 1894, Proc. Zool. Soc. Lond., p. 450) and may
exist round Aden (Yerbury & Thomas, 1895, Proc. Zool. Soc. Lond., p. 548, say :
“A Mongoose was seen at Haithalhim. The white-tailed species has been re-
corded by Thomas from Muscat, and no doubt occurs at Aden ”’).
The Muscat specimens seem to have a rather more strongly bowed zygoma
than the Sudan specimens I have seen, but this may be an age character. The
skins are slightly darker as well, but Abyssinian skins are darker still. I have
no good reason for separating the Arabian specimens from the Sudan ones in
spite of their geographical separation.
23. Herpestes javanicus pallipes Blyth.
Mangusta pallipes Blyth 1845, J. Asiatic Soc. Bengal, 14: 346, type loc.: Kandahar.
Herpestes persicus Gray 1864, Proc. Zool. Soc. Lond., p. 554, type loc.: Rhugistan & Mohammerah,
Herpestes javanicus pallipes Blyth, Pocock, 1937, J. Bombay Nat. Hist. Soc., 39: 242.
A small mongoose, head and body about 12 inches (305 mm.), tail about
10 inches (255 mm.).
Cheesman (1920, J. Bombay Nat. Hist. Soc., 27 : 332) says that this form is
common from Fao to Baghdad. It may occur round Kuwait.
24. Herpestes edwardsii ferrugineus Blanf.
Herpestes ferrugineus Blanford 1874, Proc. Zool. Soc. Lond., p. 661, pl. 81, type loc.: Larkhana,
Sind.
Herpestes edwardsii ferrugineus Blanf., Pocock, 1937, J. Bombay Nat. Hist. Soc., 89: 217-222.
Mr. Philby has sent one specimen from ‘Uquair, a & collected on 22
August, 1938, collector’s number A.
This form is intermediate in size between albicauda and pallipes; Mr.
Philby’s specimen measured (in mm.)— head and body 368, tail 371, hindfoot 69,
ear 18. The skull (occipital portion damaged) measured— zygomatic width 36-8,
inter-orbital width 15-0, maxillary width 14-0, mandibular length 47-3, pm‘
(maximum external length) 7-7, m! (maximum diameter) 7:6.
Mr. Philby tells me that these mongooses are very local, in fact he has only
seen them round the customs shed down by the quay, where they eat fish, chickens,
eggs, etc. This suggests that they have been introduced ; they appear to be
plentiful.
Novirates Zoonoarcar XLI. 1939. 199
FOXES
There are three species of fox in Arabia. The largest is Vulpes vulpes arabica
Thomas. It is related to the British fox, but is smaller and paler. It has
been recorded from the southern half of Arabia as far north as Riyadh, but is
doubtless found all over Arabia and probably into Palestine.
Vulpes ruppelli sabaea Pocock is a medium-sized fox, paler than the preceding
species and distinctly smaller. Not many specimens have been recorded, but it
is known from S.E. Arabia, the S.W. corner of the Rub al Khali and Riyadh.
The smallest of the three is a fennec fox, Fennecus sp., which is about the size
of a cat! and has huge ears. It is the palest of the three. The only other speci-
men on record besides Bertram Thomas’s is one which was presented to the
London Zoo by Mrs. Dickson of Kuwait, near where the animal was captured.
This specimen died in July 1935 and is now in the British Museum—B.M. 25. 8. 4. 1.
25. Vulpes vulpes arabica Thos.—Arabian Fox.
Vulpes leucopus Blyth, O. Thomas, 1894, Proc. Zool. Soc. Lond., p. 450, Muscat. See also Anderson
& de Winton, 1902, p. 222.
Vulpes nilotica Geofir., O. Thomas, 1895, Proc. Zool. Soc. Lond., p. 548, Aden.
Vulpes vulpes arabica O. Thomas 1902, Ann. Mag. Nat. Hist.,10: 489, type loc.: Muscat; J. G.
Dollman in B. Thomas, 1932, p. 339. R. I. Pocock, 1935, Ann. Mag. Nat. Hist.,15: 450.
Taif © Collector’s number 4 23.v.1934
Jidda 3 " Kl 5 19.11.1934
x 3 ” = 7 26.1.1934
a 3 a i" 10 9.ii.1934
Kharj fe) As a 44 13.x.1934
es Q _ i 45 13.x.1934
Abha 3 * = 88 10.vi.1936
Najran 3 = 4 94 29.vi.1936
i 3g i N 96 29. vi. 1936
Dhahran 9 ms 146 20.x1.1936
Riyadh & ” 5 166 6.v1l.1938
I am in some doubt as to whether these Arabian foxes should not be included
in Vulpes vulpes pusilla Blyth. Thomas originally referred some foxes from
Muscat to Vulpes leucopus Blyth which Pocock (1936, p. 46) shows to be asynonym
of pusilla. A year later Thomas tentatively referred two specimens from Aden
to Vulpes nilotica Geoffr. on account of their size and drew attention to the fact
that some of the Muscat specimens were like the Aden ones. Later still he de-
cided to call them (i.e. the South Arabian foxes) Vulpes vulpes arabica, and in his
description of this form he says that the size is intermediate between V. vulpes and
V. leucopus. There were only six skulls available at the time, from Muscat ? and
Aden, three 2 2 with condylobasal lengths 126, 126-5, 123-5 and three 3 4,
135-3, 140, 120-4 millimetres. From these figures it would appear that the
Arabian Vulpes vulpes are distinctly larger than lewcopus, but more material
which has since come to hand shows that Thomas’s specimens were exceptionally
large ones; the average condylobasal length seems to be 127-2 for males and
1 Bertram Thomas (1932, p. 236) says that it was referred to as “ hirr,”’ which is classical Arabic
for “ cat.” His specimen came from the Rub al Khali in longitude 51° E.
2 Blanford, 1888, Fauna of British India—Mammalia, p. 152, mentions leucopus from Muscat.
200 Novirates ZooLoGIcAE XLI. 1939.
118-7 for females. Specimens of pusilla from Baluchistan, S. and W. Persia and
‘Iraq fit in with these dimensions though pusilla from the type locality (Salt
Range, Punjab) are certainly smaller. The Salt Range, however, is the extreme
eastern end of their range.
The description of arabica says: “ ear-backs rarely deep black, more often
greyish, and sometimes scarcely darker than the head.” The words “ rarely,”
“often ” and “ sometimes ”’ are rather misleading seeing that there were but six
specimens. There are now twenty specimens from S. Arabia and ten from Central
Arabia in the British Museum, and the only ones which lack the deep black ears
are three of the paratypes. Taking the above into consideration it may well turn
out that the Arabian foxes of the species vulpes should be included in the race
pusilla, but for the moment I prefer to regard this as a speculation.
The Arabian Vulpes vulpes do, however, appear to be distinguishable by size
from the Egyptian ones. Unfortunately many of the earlier specimens of foxes
bear no information about the sex of the animal so that they are not very helpful,
but I have seen a fair number of adult sexed specimens and the following are their
measurements :
| Condylobasal
Numb f Condylobasal lengths— Standard
Se | ee age | Genen
V. v. aegyptiaca Sonn. :
Males o 0 9 5 123-5-140-3 133-9 7:0
Females 3 3 : 10 115:7-129-3 126-4 4-4
V. v. arabica Thos. :
Males . 5 5 E 11 116-6-140 127-2 6-3
Females 2 R S| 10 113-6-126 118-7 4-5
In Palestine V. v. arabica seems to meet a more northern form. Some of the
skins from Palestine have smokier flanks than arabica and there is one specimen
from Ramleh—B.M. 19. 4. 11. 8. ?— which is much greyer than any of the other
Palestine skins and which Thomas made the type of V. v. palaestina.‘ Then
there are other skins from Palestine which exactly match the Arabian foxes.
Bodenheimer (1935, p. 111) says: ‘‘ Several Foxes occur in Palestine, each of
which represents one zoogeographical region. In the southern and eastern
deserts and steppes the slender, dull reddish-grey Egyptian Fox (Vulpes nilotica)
is frequently found. Its burrows have been observed to the south of the Arabah.
This form still prevails in the southern Philistine plain and the mountains of S.
Judaea. The prevailing form of the Mediterranean sections from Ramleh and
Jerusalem up to the Lebanon is the reddish-grey Palestine Fox (V. palaestina).
The Tawny Fox (V. flavescens) 2 is an Irano-Turanian intruder in Galilee. It is
the largest of the three forms. It is of a bright, light-yellowish colour, has black
ears and a splendid brush.”’
There are no skins in the British Museum from Palestine south of Jerusalem
and it is interesting to hear that the Egyptian Fox comes up here ; moreover, the
only specimens of flavescens which I have seen are from Persia. It would seem
that the Palestine specimens which I assign to arabica are included by Boden-
1 Ann. Mag. Nat. Hist., 5: 1920: 122.
2 I.e. V. flavescens Gray 1843 not V. flavescens Blyth 1853, which is a synonym of V. v. pusilla
Blyth.
Novirates ZooLoGIcAE XLI. 1939. 201
heimer in palaestina. If this is so, it is interesting in view of the fact that Pocock
(1935, p. 450) gives palaestina as a probable synonym of arabica.
26. Vulpes ruppelli sabaea Pocock—Arabian Sand-fox.
Vulpes ruppelli sabaea R. I. Pocock 1934, Ann. Mag. Nat. Hist., 14: 636, type loc.: Rub al Khali;
id., 1935, l.c., 15: 452.
Three specimens. A <4, collector’s number 46, Riyadh, 5 November, 1934,
skin and skull; a 4, collector’s number 61, Khafs, 23 February, 1935, skin and
skull; collector's number 140, a skull picked up in a valley at Alam Abyadh.
These specimens extend the range of the subspecies in a north-westerly
direction.
Pocock had four specimens at the time when he wrote his description, and
as one of the above specimens is slightly larger than any of his it is worth recording
its measurements : Number 46, 4, head and body 451, tail 315, hindfoot 107, over-
all length of skull 108-3, condylobasal length 104-0 mm.
It is quite likely that this is the fox mentioned by Yerbury & Thomas (1895,
p. 543): “‘ At the same time Yerbury believes that a second Fox and a second
Hare are to be found in the district ”—i.e. Aden. These authors record “ Vulpes
nilotica Geoffr. ? ” (V.v. arabica). They also record Lepus arabicus, so the second
hare is very likely L. a. cheesmani Thomas (q.v.).
27. Canis lupus arabs Pocock—Arabian Wolf.
Canis lupus arabs R. I. Pocock 1934, Ann. Mag. Nat. Hist., 14: 636, type loc.: Ain, S.E. Arabia ;
id., 1935, Ann. Mag. Nat. Hist., 15: 449 (amplified description).
Canis hadramauticus Th. Noack 1896, Zool. Anz., 19: 356, type loc.: Hadhramaut ; id., 1897, Zool.
Anz., 20: 368 (amplified description ; Noack’s species is a composite one made from a jackal
and a wolf. The jackal was chosen as lectotype by Matschie).
Canis lupaster Hemprich & Ehrenberg 1830 (1833), Matschie, 1897, S.B. Ges. Nat. Fr., p. 73,
Aden.
Canis lupus subsp. (? aff. pallipes) Schwarz, 1926, Senckenbergiana, 8: 43-47, Hadhramaut (this is
Noack’s wolf specimen).
Native name is Dyb. It is interesting to note that the Wolf-like Jackal of
N.E. Africa, Canis lupaster Hemprich & Ehrenberg, is known in Egypt as Deeb
(Flower 1932, p. 395) or as Deib or Dib (Anderson 1902, p. 214).
An adult 2 shot by Mr. A. G. Griffin, 15 December, 1937, 18 miles N. of
Jeddah, altitude 40 ft.
This specimen extends the range of arabs in a north-westerly direction.
The skin is sandy-coloured and there is an admixture of black on the back,
flanks, face and tail. There is also a slight black streak on the foreleg above the
wrist. The longer, coarser hairs down the spine form a sharply defined band
some 100 mm. wide (wider between the shoulders) ; this band, together with the
tail, has the black admixture, which is weak elsewhere, strongly developed. The
belly is whitish.
The skull measurements are of interest and are comparable with those given
by Pocock (1935) which include the wolf from the Hadramaut which, together
with a jackal, was called hadramauticus by Noack. The following are the skull
measurements of Mr. Griffin’s specimen in mm.: over-all length 192, condylo-
basal length 179-5, zygomatic width 107, inter-orbital width 34, post-orbital
width 38, P* 21-4, Mt 14-0 x 18-2, M1 + M? 20-9.
202 NOoVITATES ZooLtocicaE XLI. 1939.
28. Suncus sp.—Shrew.
(?) Crocidura murina L., Thomas, 1894, Proc. Zool. Soc. Lond., p. 449, Muscat.
Two specimens in alcohol from Jidda. Collector’s number 4, 3, 5 February,
1934; no number, 2, 6 February, 1934.
Paraechinus—Hedgehogs.
Paraechinus Trouessart 1879, Rev. Mag. Zool., p. 242.
Colour is an unreliable guide as these hedgehogs show a distinct tendency to
melanism and a certain tendency to albinism. In one form, dorsalis, the belly is
blotched dark brown and white, and the variation ranges from entirely brown to
entirely white. I have considered the skulls more than the colour.
The Arabian members of the genus and those which are geographical neigh-
bours seem to be divisible into the following species :
(1) dorsalis Anderson & de Winton 1901, Ann. Mag. Nat. Hist., 7: 42;
type loc. : Hadhramaut.
This species has the skull comparatively short in the rostrum and broad in
the pterygoid region and across the zygomatic arches. Bullae are large. P® is
a mere peg or is missing. The spines of the dorsal area are dark and those of the
flanks pale so that there is a marked dark band along the back. To this species
belong ludlowi,1 albatus* and albior.3
There are two specimens of dorsalis (B.M. 97. 1. 14. 1-2) which were collected
at Muscat by Dr. Jayakar in 1896 and which had been put away in spirit labelled
“ Erinaceus.” These must have been overlooked by Anderson and de Winton,
who make no mention of them in their description of dorsalis (1901). There is
another Muscat specimen of dorsalis (B.M. 1938. 9. 8. 1) which was collected in
December (year ?) by Major Trevor. This skin resembles albior.
(2) aethiopicus Hemprich & Ehrenberg 1830 (1833), Symb. Phys. Mamm.,
2: sheet k recto; type loc.: “in desertis dongalanis.”’
This species has a short-snouted skull with large bullae and is broad across
the zygomata, but is very small, much smaller than the preceding species. P® is
a peg or is two-rooted. To this species appears to belongs oniscus,‘ a melanistic
form only known from the type.
(3) deserti Loche 1858, Cat. Mamm. et Oiseaux d’ Algérie, p. 20, type loc. :
Algeria.
The skull of this species closely resembles that of dorsalis, but the skin lacks
the dorsal band of the latter. P® has two roots.
Anderson & de Winton (1901, p. 44) say that the skull of dorsalis resembles
in general that of aethiopicus. I cannot agree ; the skulls of the typical series of
dorsalis are far larger than those of aethiopicus and resemble much more closely
deserti. It may be that dorsalis will turn out to be a subspecies of deserti.
On the other hand, I have seen two specimens of dorsalis (diagnosed by the
dark dorsal band) whose skulls, though apparently fully adult, are small and
more nearly resemble those of aethiopicus. These are Major Trevor’s specimen
from Muscat (1938. 9.8.1) and one of Dr. Jayakar’s Muscat specimens (4,
97. 1. 14. 2).
1 Thomas 1919, J. Bombay Nat. Hist. Soc., 26 : 748, type loc. : Hit, on the Euphrates.
* Thomas 1922, Ann. Mag. Nat. Hist., 9: 144, type loc.: Tanb Is., Persian Gulf.
3 Pocock 1934, Ann. Mag. Nat. Hist., 14: 636, type loc. : W. Dhimir, Geradun, S. Arabia.
4 Thomas 1922, Ann. Mag. Nat. Hist., 10 : 307, type loc. : Fayush 7 miles N. of Sheikh Othman.
Novirates Zoonocrear XLI. 1939. 203
(4) hypomelas Brandt 1836, Bull. Sci. Acad. Imp. St. Petersbourg, 1: 32 ;
type loc.: “pays des Turcomans”’! (macracanthus Blanford 1875 from Car-
mania and amir Thomas 1918 from Afghanistan are synonyms).
This species has a long narrow skull, narrow in the pterygoid region and with
small bullae. P® is three-rooted. There are wholly black and wholly white
phases. To this species belong niger? and sabaeus.?
(5) blanfordi Anderson 1878, J. Asiat. Soc. Beng., 47: 208; type loc. :
Rohri in Sind.
This species has small bullae. P® is three-rooted but the shape of the skull
is intermediate between that of hypomelas and dorsalis. 'To this species belongs
seniculus,* the black Tanb Is. form which Thomas described as a subspecies of
niger.
29. Paraechinus dorsalis dorsalis And. & de Wint.
The following specimens may be identified thus. A couple, gj and ®, from
Taif, 31.v.1934, and three from S.W. Arabia, 1936 A, B and c. Only one of
these is fully adult (1936 A, 2) and its skull measurements in mm. are: over-all
length 50-3, zygomatic width 32-4, post-orbital constriction 12-5.
30. Paraechinus sp.
There is a specimen from Hadda, 9, 2.ii1. 1934, whose skull (sub-adult) agrees
fairly well with those of deserti. The skull measurements in mm. are: over-all
length 48-9, zygomatic width 28-3, post-orbital constriction 11-0.
This skin has the distal half of the spines dark brown with a short light tip
save on the back of the neck and between the shoulders where the distal halves
of the spines are entirely dark brown with no light tip. The face is entirely deep
brown ; the lower part of the inside of the ear is white ; there is a white mark at
the angle of the jaw but the chin is black ; the belly is very dark brown, and so
are the legs and feet. I am not sure where this specimen should be placed and
prefer to wait till more specimens come to hand.
Mr. Philby has also sent two very young hedgehogs from W. Tasrir,
6.vili.1931. These have a cinnamon-coloured skin and uniformly coloured,
ginger spines.
BATS
31. Pipistrellus sp.—Pipistrelle.
One 9, collector’s number 47, Riyadh 1,800 ft., 8 November, 1934. This
is not P. kuhli ikhwanius Cheesman & Hinton.
32. Asellia tridens E.. Geoff.—Trident Bat.
Rhinolophus tridens E. Geoffroy 1813, Ann. Mus. Hist. Nat. Paris, 20: 260, type loc.: Egypt.
Ten dd and 3 99 (no numbers) from a coral rock quarry near Jidda,
22 April, 1934.
1 Ognev 1927, Zool. Anz. Leipzig, 69 : 210-211, says that the type loc. is N. Persia.
2 Blanford 1878, J. Asiat. Soc. Beng., 47: 212, type loc.: ‘‘ juxta Muscat.”
3 Thomas 1922, Ann. Mag. Nat. Hist., 9: 143, type loc.: El Kubar, 60 miles N. of Aden.
4 Thomas 1922, Ann. Mag. Nat. Hist., 9: 142, type loc.: Tanb Is., Persian Gulf.
204 Novirates ZootocicaE XLI. 1939.
33. Rhinolophus clivosus Cretz.—Horseshoe Bat.
Rhinolophus clivosus Cretzschmar in Rüppell 1826 (1828), Atlas zu der Reise im nördlichen Afrika, 1,
Sdug., p. 47, type loc.: Mohila.
Two gd and 7 29, collector’s numbers 19-21, 23-26 and two without numbers,
Taif, July 1934.
One J, no number, Mecca, December 1934.
One 3, B.M. 1937. 9. 20.1, collected by W. H. Ingrams at Zeil, Seiyun,
Hadhramaut, December 1936.
34. Rhinolophus hipposideros minimus Heug.—Horseshoe Bat.
Rhinolophus minimus Heuglin 1861, Nova Acta Leop. Carol., 29: 6, type loc.: Cheren.
One 4, collector’s number 40, Taif, 5,500 ft., 12th September, 1934.
35. Nycteris thebaica adana K. And.—Long-eared Bat.
Petalia thebaica adana K. Andersen 1912, Ann. Mag. Nat. Hist.,10: 548; type loc.: Myba, near
Aden.
One g and 1 9; no numbers, near Jidda (?).
Three 33, collector’s numbers 92, 105, 106, Najran 4,400 ft., June and July
1936. Local, name Mussa.
36. Taphozous perforatus haedinus Thos.—Tomb Bat.
Taphozous perforatus haedinus O. Thomas 1915, J. Bombay Nat. Hist. Soc., 24: 62, type loc.: Chan-
ler Falls, Juaso Nyiro, Kenya,
Two specimens (sex ?), corral quarry near Jidda.
Two specimens (sex ?), collector’s numbers 64 and 65, Shumaisi, 450 ft.,
28 January, 1936.
37. Liponycteris nudiventris Cretz.—Naked-bellied Tomb Bat.
Taphozous nudiventris Cretzschmar in Riippell 1826 (1830/31), Atlas zu der Reise im nördlichen
Afrika,1, Säug., p. 70, type loc.: Giza.
One 9, collector’s number 97, Najran 4,400 ft., 30 June, 1936.
38. Rhinopoma sp.—Mouse-tailed Bat.
Four young specimens (unnumbered), 3 34, collector’s numbers 92, 93 and
97, and one 9, number 95, all from Najran, 4,400 ft., 30 June, 1936.
39. Otonycteris hemprichi Peters—Long-eared Bat.
Otonycteris hemprichi Peters 1859, Monatsbericht Akad. Berlin, p. 223, type loc.: “ex Hemprich &
Ehrenberg’s collection.” ı
One 9, collector’s number 165, ‘Anaiza 2,300 ft., 10 June, 1938. Two
immature specimens, 153 from Hail 3,500 ft., 31 May, 1938, and 164 from
‘Anaiza, 10 June, 1938.
Novirares ZOooLoGIcAE XLI. 1939. 205
APPENDIX I.
A SHORT BIBLIOGRAPHY FOR THE STUDY OF ARABIAN MAMMALS.
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de Winton, W. E., 1903. Hair whorl in a gazelle from Aden. Proc. Zool. Soc.
Lond., 2 : 317-318.
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Thomas, p. 339. London.
Dollman, J. G., 1933. A list of mammals in “The Empty Quarter” by H.
St.J. B. Philby, p. 394. London.
Doughty, C. M., 1889. Travels in Arabia Deserta. 2 vols. Cambridge.
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2: 99-144. (Two proof impressions 1813.)
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le genre des Rhinolophes (pp. 254-266). Ann. Mus. Hist. Nat. Paris, 20.
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Hist. Nat., 2 : 733-743.
Gray, J. E., 1864. A revision of the genera and species of Viverrine animals
(Viverridae), founded on the collection in the British Museum. Proc. Zool.
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Harris, W. B., 1893. A Journey through Yemen. London.
Hart, H. C., 1891. Some account of the fauna and flora of Sinai, Petra and
Wädy ‘Arabah. London. (Mammalia, pp. 233-238.)
Hemprich, F. W., & Ehrenberg, C. G., 1828-1830 (1828-1833). Symbolae
physicae seu icones et descriptiones mammalium, 1 & 2. Berlin.
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Hommel, F., 1879. Die Namen der Säugethiere bei den südsemetischen Völkern.
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289-312.
Lataste, F., 1882. Mammiferes nouveaux d’algérie. Naturaliste, 2 : 27-28.
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London, 5 vols.
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melte und beobachtete Säugethiere. S. B. Ges. Naturf. Fr. Berlin,
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Matschie, P., 18998. See Peters, W., & Matschie, P.
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5: 517.
Morrison-Scott, T. C. S., 1939. The identity of Acomys megalotis (Lichtenstein),
described from Arabia. Ann. Mag. Nat. Hist., 3 : 238-240.
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(Appendix VII, pp. 411-412, a letter from W. G. Palgrave on the Arabian
fauna.)
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16
208 NOVITATES ZOOLOGICAE XLI. 1939.
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Novirares ZooLoGIcAE XLI. 1939. 209
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APPENDIX II
MEASUREMENTS.
Total length or over-all length of skull as used in this paper means the total
length of the skull between perpendiculars, including the incisors. Pocock does
not include the incisors.
Mandibular length is the distance in a straight line from the posterior surface
of the mandibular condyle to the ventral rim of the alveolus of I,.
Standard deviation. —It is not often that a collection of mammals contains a
sufficient number of any one species to be sure that the distribution curve for any
factor is a normal one, but the standard deviation may still give some idea of the
variation from the mean and saves having to give the measurements at length.
210 NoviratEs ZooLoGIcAE XLI. 1939
In some cases, therefore, the extremes, mean and standard deviation have been
considered sufficient. The standard deviation has been worked as follows :
re \/ (deviations)?
n
where “‘ deviation ’’ means the difference between the measurement of each speci-
men and the mean measurement, and ““n ” is the number of specimens. (For
further details see—The British Association for the Advancement of Science,
Biological Measurements, 1935, p. 12.)
Hindfoot.—This measurement does not include the claws.
APPENDIX III.
LOCALITIES REFERRED TO IN THIS PAPER.!
N. E. N. E.
Abha . 5 . 18°20’ 42° 20’ Haraj Plain . . 14°50’ 46° 40’
Abyan Mts. . . near Aden Harra Rahat . 21° 537 402 207
Afalil . : . 46° 30° 15° 45’ Hasa Oasis . . just E. of Hufuf
Ahl al Dhib . . 18°15° 42°55’ Hawiya : 22° 10° 49° 30’
Alam Abyadh . 16°00’ 45° 42’ Hudhaiyib al emt 20° 30° 42° 40’
Anaiza. 6 . 26°05’ 43° 59° Hufuf . 6 = 257220 24955304
Aqda . : . few miles N. of Husainiya . . 18°00’ 44° 30’
Hail Ilha‘az. 5 . 17°10’ 54° 10°
Arain . . 15°35’ 46° 45’ ‘Iraq Well . . 17°40’ 44° 00’
Arq Adu Da’i ir . 16°40’ 45°15’ Jabrin Oasis . 28°15’ 48° 55°
Ashaira : . 22°00’ 41°00’ Jadaliya : 17° 25’ 44° 00°
Dailami Ä . 20°20’ 42° 40’ Jafura Desert (centre)24° 15’ 50° 00’
Dawadami . . 24° 40° 44° 30° Jaual Khudaif . 15°15’ 46° 35’
Deriyot A . 17°10’ 54°10’ Jauf, W. : . 15245 452107
Dha’f Hills . . 21° 30° 39° 30° Jau Mulais . 7 NG RE EGS? B55’
Dhahran : . 17°40’ 48°30’ Jidda . : . 21° 30’ 39° 00’
Dhalm . © . 22° 50° 41°40’ Jurbaib : 1720027542107
Dhirawiya . . 17°27° 43° 37’ Karyatein . . 840 115) 30157
Dhofar. : . 17° 00° 54°00’ Khabb, W. . . 16°50’ 45° 10’
Dofar . : . see Dhofar Khafs . 6 . 25° 20’ 46° 30’
Dohat al Salwa . 25°10’ 50° 40’ Kharj . : . 23° 55° 47° 30/
Farsan Is. . . 16°50’ 42°00’ Khaur, El . 5 IB IG A
Ghumair, W. . 16°37’ 45°07’ Khufaifiya . . 24° 35’ 45° 00’
Hadda. 3 . 21°30’ 39° 30° Khurma, W.. . 22° 00° 42° 00°
Hadhba 3 . 17°00’ 45° 00° Lahej . : . 138° 00° 44° 50’
Haill . i . 27° 31’ 41° 44’ Loheia. 5 . 15°40’ 42° 30’
Hamdha : . 19°05’ 438° 45’ Luwai, W. . . 25°00’ 40° 39’
Hammam, W. . 14° 50° 46° 36’ Madina (station) . 24° 28’ 39° 36’
Hanaka, W. . 5 lief BIS yay Wendt, Wao o A576
Haqu . 5 . 17° 40' 42° 40’ Maswara : 2 17° 46’ 43° 24"
1 Some of these co-ordinates are only rough.
Novirares ZOOLOGIOAE XLI.
Matau .
Mathnat
Mocha .
Mushainiqua.
Najran.
Naq’a, W.
Nejd
Ogair .
Qaim
Qariya .
Qunfidha
Rakba Plain.
Ramlat Sabatain .
Ras al Khabb
Rass
Raushan
Rikani .
Riyadh
Ruwaik
Sahlal .
1939.
N. E.
16° 30° 45° 38’
part of Taif
13° 15° 43° 15’
16° 30° 45° 55’
17° 30° 44° 20’
15° 30° 47° 15’
a large district of
Central Arabia
25° 38° 50° 13’
21° 20° 40° 45’
17° 12’ 43° 10’
19° 00° 41° 00’
22° 30’ = 41° 40’
15° 40’ 47° 00’
15° 48’ 47° 50’
25° 50’ 43° 40’
20° 00° 42° 40’
21° 30’ 39° 30’
24° 35° 46° 40’
15° 55° 46° 10°
15° 35° 47° 30°
Samuda
Shabwa
Sha’ib Arjan.
Sha’ib Hanjur
Shaka .
Shari ;
Shariya (= Sharaya)
Shukra
Shumaisi
Sirr al Yamani
Suda
Sudaihat Tract
Sumaika, W.
Maiti): :
Taraf al Ain .
Tasrir, W.
Thar, W.
Ugla
Wasil .
APPENDIX IV.
FORMS DESCRIBED AS NEW: Psammomys obesus dianae
Tatera philbyv
211
N. E.
on Rakba Plain
15° 25’ 47° 00’
17° 40’ 44° 00°
18° 15’ 42° 45’
13° 00’ 44° 45’
DI elt 43° 27
21° 35’ 40° 05’
13° 25’ 45° 40’
212 25! 39° 40’
16° 20’ 46° 50’
18° 20’ 42° 20’
16° 00’ 45° 50’
17° 56’ 43° 48’
21° 10’ 40° 45’
15° 50° 47° 40’
between Riyadh
and Dawadami
see ‘Iraq Well
15° 25°
15° 00’
46° 50’
43° 39’
bo
pay
bo
Novitates ZOOLOGICAE XLI. 1939
ZUR NOMENKLATUR DER BEIDEN DEUTSCHEN PROCRIS-
ARTEN MIT SPITZEN FUHLERN.
Von KARL JORDAN
EFT 1 der Stett. Ent. Zeit. 1939, S. 149-153, bringt einen Aufsatz von
Dr. B. Alberti, in dem der Verfasser seiner Ansicht nach beweist, dass
ich in Seitz, Grossschmett. II (S. 8, 1907) einige Verwirrung in der Nomenklatur
der Procris-Arten mit spitzen Fühlern angerichtet habe. Nun, alle Zweige der
Wissenschaft sind voller Fehler ; das ist nicht sonderlich aufregend, denn durch
Ausmerzen der Irrtümer kommt man doch allmählich zur Wahrheit. Mein
Versuch einer Bearbeitung der Gattung Procris für Seitz, bei dem mir nur recht
geringes Material zur Verfügung stand, enthält in der Tat einige Fehler und man
kann mir mit Recht den Vorwurf machen, dass ich sie trotz des grossen Materials,
das sich seitdem in Tring angesammelt hat, nicht verbessert habe. Der Wille
war schon da, aber die Zeit wurde anderweitig in Anspruch genommen und die
Notizen über meine Untersuchungen sind nie zusammengefasst und druckfertig
gemacht worden.
Albertis Artikel ist so klar geschrieben, dass man in seiner Darstellung ohne
Weiteres die Lücken und Fehler sieht, die ihn zu falschen Schlussfolgerungen
verleitet haben.
Die zu beantwortenden Fragen sind diese : Welche von den im jetzigen Gross-
deutschland vorkommenden Procris-Arten mit spitzen Fühlern ist Hübners
globulariae und welcher Name ist der richtige für die zweite deutsche ähnliche
Art?
Irrtümlicherweise beginnen alle Autoren die Geschichte des Namens globu-
lariae mit Band II von Hübners Sammlung Europäischer Schmetterlinge. Der
Name erschien aber schon drei Jahre früher in Hübners Sammlung Auserlesener
Vögel und Schmetterlinge (1793). Das Büchlein enthält 100 farbige Tafeln, auf
denen je ein bunter Vogel und ein europäischer Schmetterling abgebildet sind ;
die Falter sind zum Teil Mikros. Die 16 Seiten Text geben nur die deutschen
und lateinischen Namen, ohne Beschreibung und ohne Fundort. Das Werkchen
ist sehr selten ; das einzige vollständige Exemplar in England kaufte ich Anfang
des Jahrhunderts von Dr. G. Kraatz für 100 Mark. Auf Taf. 63 ist Sph. Immac.
globulariae abgebildet. Das Bild ist verfehlt ; die Vorderflügel sind zu schmal
und die Fühler dünner als beim Männchen irgend einer Art, an der Wurzel dünn,
bis zu etwa 2/3 dicker und dann sehr dünn. Das Bild ist aber sicher für ein
Procris Männchen mit spitzen Fühlern gemeint.
Der Name erscheint dann zunächst wieder in 1796 auf Taf. 1 des 2. Bandes
von Hübners Samml. Europ. Schmett. Die Fig. 4 dieser Tafel ist globulariae
. genannt ; das Original stammte aus den “ nahen Gebirgen Wiens.” Zwischen
1803 und 1805 gab Hübner nochmals eine Taf. 1 heraus, die von der ursprüng-
lichen Taf. 1 beträchtlich abweicht. Fig. 2 und 3 sind auf dieser Tafel als
Männchen und Weibchen globulariae bezeichnet und der Fundort ist diesmal
““ Deutschland, auf Heiden.” Die Fühler der Fig. 2 sind viel dicker als in Fig. 4
der Tafel von 1796. Nun wissen wir seit etwa 30 Jahren, dass es in Deutschland
NoviratEs ZooLogIcAE XLI. 1939. 213
zwei spitzfiihlerige Arten gibt, die bei der Deutung der Hiibnerschen Figuren in
Frage kommen und in Oesterreich drei (von denen budensis erst 1858 von Speyer
als Art erkannt wurde). Nach Hübners Figuren zu urteilen. stellen die Bilder
von 1793 und 1796 die dünnfühlerige Art und das Bild von 1903-5 die dicker-
fühlerige dar. Da es aber sehr schwer ist, korrekte Abbildungen dieser Falter
zu liefern, so kann man immerhin noch im Zweifel sein, ob Hübner mit jenen
Figuren zwei oder drei Arten abgebildet hat oder nur eine. Die Autoren des
vorigen Jahrhunderts konnten über Hübners Bilder nicht ins Klare kommen,
weil sie nicht wussten, dass ihre globulariae ein Gemisch von zwei oder selbst
mehr Arten war.
Zeller, von dem Dr. Alberti merkwürdigerweise sagt, dass er die globulariae-
Frage schon in 1847 entschieden habe, war betreffs der Anwendung des Namens
ganz und gar im Ungewissen. Wir lesen in /sis 1847 S. 394 die folgende Darstel-
lung dieser Frage: ,, Von At. globulariae besitze ich nur 3 Männchen, 2 Weibchen
aus der Wiener Gegend. Ich muss hier auf ihre genauere Beziehung eingehen,
damit ich mit meinen Siciliern verstanden werden kann. Meine At. globulariae
ist gleich den kleinsten hiesigen Exemplaren von statices .... Fühler kaum 3 ”
lang, die männlichen gekämmt mit zwey gedrangten Reihen Kammzähne und
dadurch dick aussehend, am Ende spitz... . Die Vorderflügel sind schmal,
erweitern sich wenig und laufen ziemlich spitz zu... . “
,, Ein österreichisches Männchen ist sicher specifisch verschieden. Es ist
viel grösser .... hat zater beschuppte Vorderfliigel . . . . Fühler fast 4” lang—
mit langen, aber dünnen Kammzähnen, daher im Ganzen von schmächtigerm
Aussehen... . “
,, Auf dieses Männchen so wie die kleinen von Mann und Fischer v. Röslerst.
erhaltenen Exemplare passt Ochsenheimers Beschreibung sehr wenig und
Hübners Tafeln habe ich nicht zum Vergleich. Ich lasse es also unentschieden,
ob das einzelne Männchen At. globulariae und die kleinen Exemplare Hübners
At. chloros sind.“
Zellers Sammlung ist im British Museum (siehe Horn, Ueber den Verbleib
der Entomologischen Sammlungen der Welt, 1926). Einige kleine, von Mann
erhaltene Exemplare sind in Zellers zierlicher Handschrift globulariae etiquettiert ;
sie sind chloros Hübner 1808-13. Das grosse Stück habe ich nicht in der Samm-
lung gefunden.
Laut Alberti soll Zeller durch Beschreibung der neuen Art notata aus Sizilien,
die mit der dünnfühlerigen deutschen Spezies artgleich ist, diese aus dem globu-
lariae-Komplex eliminiert haben. Die Ansicht steht im Widerspruch mit den
Regeln der zoologischen Nomenklatur. Eine Eliminierung hätte nur stattgehabt,
wenn Zeller gesagt hätte, dass seine notata artgleich mit der dünnfühlerigen
deutschen Art wäre. Er wusste ja aber gar nicht, dass zwei deutsche Arten
unter globulariae steckten und konnte daher nichts eliminieren. Die Zellersche
Arbeit hat Dr. Alberti vielleicht nicht vorgelegen; denn Zeller beschreibt darin nicht
eine Art, sondern zwei: (1) tenwicornis nov. sp. Die Genitalarmatur des Typus
ist von Mr. W. H. T. Tams präpariert ; die Haftklappe hat einen langen Basal-
dorn ; die Art ist mit jener dickerfühlerigen deutschen Procris meines Erachtens
artgleich, die ich im Seitz als cognata behandelt habe. (2) notata nov. sp. habe
ich im Seitz als mit der dünnfühlerigen Procris Deutschlands artgleich aufgeführt.
Nach Albertis Prinzip hätte also Zeller beide deutsche Arten eliminiert und
globulariae müsste auf eine dritte Art bezogen werden.
214 Novirates Zoonocicar XLI. 1939.
Die Kenntnis der Procris-Arten wurde um einen grossen Schritt durch
Staudingers Bearbeitung der Gattung in Stett. Ent. Zeit. XXIII, 1862, weiter-
gebracht. Leider waren ihm die spezifischen Charaktere der Genitalarmaturen
noch nicht bekannt und er musste sich daher durch äussere Unterschiede leiten
lassen, die bei solch nahe verwandten Arten oft unzuverlässig und soweit sie die
Fühler betreffen in den Abbildungen nie genügend zum Ausdruck gekommen
sind. Staudinger kannte das Werkchen von 1793 nicht ; aber er hatte wenigstens
beide Ausgaben der Taf. 1 von Hübners Band II der Samml. Europ. Schmett.
vor sich. Nach kritischer Besprechung der Hübnerschen Abbildungen und was
man sonst noch bis dahin über globulariae publiziert hatte, gesteht er: ,, Ich bin
leider trotz aller Bemühungen nicht im Stande, diese Schwierigkeiten zu lösen.
Zunächst ist es um so unmöglicher, mit Gewissheit anzugeben, welche Art die
Hübner’sche globulariae sei, als dieser darunter zwei sehr verschieden scheinende
Thiere abbildet ”—(nämlich Taf. 1, Fig. 4, 1796, und Taf. 1, Fig. 2, 1803-5). —
,, Ich verstehe unter globulariae die Art, bei welcher die Kammzähne....an den
männlichen Fühlern nicht sehr lang sind .... Sichere Exemplare dieser In.
globulariae habe ich nur von... .. Glogau, Jena, Augsburg, Wien, Mehadiah und
Paris.“ Was hier mit “ Kammzähne nicht sehr lang” gemeint ist, geht aus
Staudingerts Beschreibung von cognata hervor, deren Männchen “ sehr lange
Fühlerkämme ” hat.
Als ich die Gattung Procris für Seitz, Grossschmett., bearbeitete, musste ich
mich in der globulariae-Frage nach dem vorliegenden Material und den Litera-
turangaben richten :
(1) Ich hatte aus Deutschland und der Schweiz nur eine Art mit nicht sehr
. langen Kammzähnen und einfacher Haftklappe (3) vor mir. Exemplare mit
diesen Charakteren hatte ich auch aus Oesterreich, Frankreich und Italien.
(2) Aus England, Frankreich und Italien waren in der Sammlung Exemplare
einer zweiten Art, mit viel längern Fühlerkämmen und einer mit einem Basaldorn
bewaffneten Haftklappe ().
(3) Hübners Abbildung 1793 hat sehr dünne Fühler, ein offenbar fehlerhafter
Stich ; die Bilder des Männchens von 1796 (mit mässig langen Kammzähnen,
aus der Wiener Gegend) und des Männchens von 1803-5 (mit sehr stark gekämm-
ten Fühlern, aus Deutschland) wurden von Staudinger angezweifelt.
(4) Staudinger (1862) bezieht den Namen auf Exemplare mit “ nicht sehr
langen Kammzähnen ” (3).
(5) Alle Autoren geben nur eine einzige Art mit spitzen Fühlern aus Deutsch-
land an. Keiner erwähnt eine Art mit sehr langen Kammzähnen aus Oesterreich
oder Deutschland.
Daher war es doch wohl selbstverständlich, dass ich mit diesen Tatsachen
vor mir den Namen globulariae nur auf die mir vorliegende Art mit mässig langen
Fühlerkämmen und einfacher Haftklappe bezog. Um es unmöglich zu machen,
dass Hübners Name globulariae von 1793 auf eine dritte Art bezogen wird, erkläre
ich hiermit, dass die Figur eine misratene Zeichnung der P. globulariae mit
unbewehrter Haftklappe ist und wähle als Heimatland der Originale dieser Figur
und der Fig. 2 und 3 der Tafel 1 von 1803-5 die weitere Umgebung von Augsburg.
Für die zweite Art nahm ich den Namen an, unter dem ich sie bei Staudinger
in 1862 und in seinen Katalogen aufgeführt fand : cognata. Bei der Angabe der
Verbreitung fügte ich hinzu: “Ob auch in Deutschland und Oesterreich ? ”
Das hatte seinen Grund ; denn vor Ablieferung des Manuskriptes erhielt ich
Novirates ZooLogIcae XLI. 1939. 215
direkt aus Süddeutschland einige Exemplare dieser langkiimmigen Art. Leider,
so muss ich heute sagen, hatte mich ein Entomologe aus Deutschland, der hier
zu Besuch war, gewarnt, dass man sich auf die Fundortszettel des betreffenden
Sammlers nicht immer verlassen könnte. Immerhin deuteten diese Stücke die
Möglichkeit an, dass die Art mit bewaffneter Haftklappe auch in Deutschland
vorkäme, und daher wünschte ich die Aufmerksamkeit der deutschen Lepidopter-
ologen durch jene Frage auf diese zweite Art zu lenken, womit ich ja auch Erfolg
gehabt habe. Bis dahin hatte niemand geahnt, dass es zwei solch ähnliche Arten
in Deutschland gibt.
Nach Alberti soll cognata H.-S. durch cognata Lucas präokkupiert sein.
Bibliographen würden sich sehr für einen Beweis dieser bis jetzt leeren Behauptung
interessieren. Irrtümliche Kritik aber hat einigen Wert, wenn sie, wie in diesem
Falle, zur Aufklärung einer Sache Anlass gibt. Ich schrieb Herrich-Schäffer
Priorität zu, weil Erichson’s Bericht für 1847-48 (Archiv Naturg.) die Angabe
enthielt, dass in 1847-48 vom 2. Bande der System. Bearbeit. Schmett. Europ.
die Hefte 21-36 ausgegeben wären und Heft 21 den Abschluss der Spinner bildete.
Damit war gesagt, dass die Fig. 94 u. 95 von Procris cognata auf Taf. 13 des 21.
Hefts in 1847 erschienen waren. Das ist späterhin bestätigt worden. Das Jahr
der Ausgabe und der Inhalt jedes Heftes waren auf den Umschlägen angegeben ;
glücklicherweise ist in England ein Exemplar des Werkes, in dem die Umschläge
erhalten sind. Diesem Exemplar nach sind die Bilder der Zygaenen in den
Jahren 1843-47 wie folgt ausgegeben : 1843 Taf. 1 u. 2; 1844 Taf. 4-7; 1846
Taf. 8-11; 1847 Taf. 12 u. 13. Vergl. Supplem. Cat. Libr. Brit. Mus. (Nat. Hist.)
1922, S. 457-8, und Hemming, Hübner 1937, 8.580. Der Text zu den Zygaeniden-
Bildern erschien in 1847 ; globulariae und cognata sind nicht darin beschrieben
und erstere nur ganz beiläufig unter chloros erwähnt und cognata gar nicht ; die
Beschreibungen (ohne Bedeutung) dieser beiden Procris finden sich im Nachtrage
1852, S. 42 (daher wohl der Irrtum in Kirby, Cat. Het. 1892; in Staudingers
Katalogen fehlen vielfach die Hinweise auf die Erscheinungszeit). Der Fundort
der cognata ist von Herrich-Schäffer nicht angegeben. P. cognata Lucas datiert
von 1849, ist also um zwei Jahre jünger als cognata H.-S. Damit ist aber die
Nomenklatur der Art noch nicht geklärt ; denn in 1847 erschien auch die Beschrei-
bung von P. tenwicornis Zeller, welche die bewaffnete Haftklappe der langkämmi-
gen deutschen Art hat (vergl. S. 213 Zeile 6 v. u.), also wie gesagt meiner unmass-
geblichen Ansicht nach artlich mit der cognata H.-S. identisch ist. Welchen der
beiden Namen von 1847 sollen wir für die Art nehmen ? Für tenwicornis sprechen
folgende Punkte : sie wurde mit genauer Angabe der Herkunft im April 1847
beschrieben und der Typus ist erhalten und untersucht, sodass gar kein Zweifel
besteht, um welche Art es sich handelt. Andererseits finde ich nur den einen
Grund, Herrich-Schäffers cognata die Priorität zu geben, dass ich,den Namen in
diesem Sinne im Seitz gebraucht habe. Gegen cognata sprechen aber das Fehlen
der Fundortsangabe, die ungenaue Figur und recht oberflächliche Beschreibung,
die unpräzise Angabe 1847 als Datum des Namens und das Fehlen des Typus,
wodurch uns die Möglichkeit genommen ist zu beweisen, was Herrich-Schäffer
wirklich abgebildet hat. Ich gebe also tenuicornis Priorität. Der Name erschien
zuerst als nomen indescriptum in Boisduval, Index Method. 1840, S. 54, wo wir als
No. 449 angegeben finden: ,, cognata Ramb. Faun. And.—Hispan. mer.“
Lucas, Explor. Alg. III, S. 373 (1849), zitiert Boisduval. Da Herrich-Schäffer
denselben Namen im selben Dezennium gebraucht, ist es nicht unwahrscheinlich,
216 NOVITATES ZooLocicaE XLI. 1939.
dass seine Exemplare von cognata aus Spanien kamen, woher auch Staudinger
(1862) Material hatte. Ich bezeichne daher hiermit Andalusien als Land der
Herkunft von Herrich-Schäffers cognata. Ob die Exemplare aus Zentral-Europa,
England, Frankreich und Spanien wirklich identisch mit tenwicornis aus Sizilien
sind, kann ich nicht sagen ; vielleicht besteht die Art aus mehreren Subspezies
oder es mag mehr als eine Art mit ähnlichen Genitalien wie tenwicornis geben.
Früher hatte ich kein Material, um diese Feinheiten zu studieren und jetzt ist
anderes zu tun.
Was Alberti über die Häufigkeit und Verbreitung der. beiden Arten sagt,
ist für die Nomenklaturfrage ganz ohne Bedeutung. Eine seltene Art kann am
richtigen Flugplatze recht häufig sein und was vor 150 Jahren häufig war, mag
heute in derselben Gegend selten sein. Das von Staudinger in 1862 als globulariae
erwähnte Material mag sehr wohl ein Gemisch beider Arten gewesen sein ; wer
kann das wissen? In der Zellerschen Sammlung stecken deutsche und öster-
reichische Exemplare beider Arten zusammen als globulariae ; seine Stücke aus
Jena sind tenuicornis. Herrich-Schäffer (Nachtrag, S. 42, 1852) stellt fest, dass
bei Regensburg nur eine Art vorkommt, die er für globulariae hält ; im British
Museum befindet sich ein Männchen von dort, das die mässig langen Kamm-
zähne und unbewehrte Haftklappe der ächten globulariae hat. Aber daraus
folgere ich noch nicht, dass bei Regensburg wirklich nur die eine Art zu finden
ist. Im Hügellande der Göttinger Gegend fliegt die eine spitzfühlerige Art links
der Leine und die andere hat man rechts gefunden, was vielleicht auf Zufall
beruht. Das Auffinden hängt eben davon ab, ob man zu rechter Zeit am rechten
Orte sammelt. Wie aus Seitz II hervorgeht, habe ich damals tenuicornis Zeller
nicht erkannt. Was ich über die Art sagte, war von Staudinger abgeschrieben
(——in der Beschreibung sind versehentlich vor “ Oberseite ” die Worte
“ Wurzel der ” ausgelassen. ). Der Typus war zwar im British Museum,
aber ich konnte ihn nicht näher untersuchen und verliess mich auf Staudinger,
dem die Zellerschen Typen vorgelegen hatten. In Wirklichkeit hatte ich in
turatii Bartel die tenuicornis schon vor mir, war mir dessen jedoch nicht bewusst.
Durch den spätern Ankauf der Sammlungen von Ragusa und Stauder erhielten
wir beträchtliches Material aus Sizilien, das mich über tenuicornis aufklarte. Da
Procris turatii Bartel 1905 schon unter cognata im Seitz erwähnt war, kam es mir
nicht in den Sinn, dass jemand der Art einen überflüssigen Namen geben könnte.
Ich fühle mich wirklich nicht für die Handlung von Sr. Agenjo verantwortlich ;
er hätte sich über die im Seitz noch in der Luft hängende tenwicornis durch Anfrage
beim British Museum orientieren sollen. Ich habe in den letzten ein oder zwei
Jahren mit Agenjo über Procris korrespondiert und ihm auch einige Exemplare
geschickt ; das mag aber zu spät gewesen sein. Die Jugend hat grosse Eile,
wir Alten nehmen uns Zeit.
Noviratrs ZooLoGIcAE XLI. 1939. 217
THE BRAZILIAN HYDROPTILIDAE (TRICHOPTERA).
By MARTIN E. MOSELY
(With 93 text-figures.)
| Nae from a few larval cases which were described by Fritz Miller in 1880
only one Brazilian Hydroptilid species has so far been known in the adult
state. This is Mortoniella albolineata Ulmer 1906.
As a consequence, the collection made by Herr Fritz Plaumann, in Nova
Teutonia, is of considerable importance.
It includes eighteen new species contained in eleven genera, of which five
are new to science, one distributed throughout North America and Mexico, four
erected originally to take Mexican species and, finally, one genus, Hydroptila,
which is ubiquitous. Records of species in this genus have been made from
Europe, Asia, the Far Kast, India, South Africa, North Africa (with one species
from an Egyptian oasis), Madeira, the Canaries, North America, Mexico, Australia,
New Zealand and now, Brazil. This Brazilian species, H. producta, is closely
related to our own simulans, and it is interesting to find that the scent-organ
apparatus at the back of the head, so far restricted to Hydroptila, is fully developed
in the South American species.
Of the new genera, particular reference must be made to Abtrichia, which
contains two species. Of these, sguamosa is noteworthy as the first Hydroptilid
to be discovered in which the vestiture of the wings consists of scales as well as
of hairs, a condition which prevails in other Trichopterous families. These scales,
mainly confined to an overlapping fold of the costa of the anterior wing, are of
the nature of androconia and form, probably, part of some scent-distributing or
generating apparatus in connection with this fold.
In both the species in the genus there is a considerable modification of the
form of the head and also of the basal joint of the antenna, particulars of which
are given with the generic description. A similar modification of the antenna
occurs in the Mexican genus Zumatrichia Mosely 1937, and, as in this genus, the
ocelli number only two.
A word must be added as to the nature of the terrain in which the captures
were made. Nova Teutonia is situated in western Santa Catharina, in a moun-
tainous region near the River Uruguay. Small mountain torrents, waterfalls,
bogs and other forms of water abound and, as the station from which collecting
is made, is situated at an altitude of between five and seven hundred metres, with
a not unduly high temperature, conditions are eminently favourable for the
existence of Trichoptera.
It may be added that Herr Plaumann has sent to the British Museum con-
siderable numbers of Caddis flies from families other than the Hydroptilidae,
though, as might be expected, the Phryganeidae and the Limnophilidae are not
represented.
All the new species here described were collected by Herr F. Plaumann
in Nova Teutonia, and the types, in most cases also paratypes, are in the
collection of the British Museum.
218 Novirates ZoontocicaE XLI. 1939.
1. Diaulus ladislavii Fr. Müller.
Diaulus ladislavii Fr. Müll., Arch. Mus. Nac. Rio Jan., 3, p. 118, 1880; id., Zeits. Wiss. Zool., 35,
p. 68, 1881.
This species is described from a larval case, the adult remains unknown.
2. Rhyacopsyche hageni Fr. Müller.
Rhyacopsyche hageni Fr. Müll., Arch. Mus. Nac. Rio Jan., 3, p. 121, 1880; id., Zeits. Wiss. Zool.
35, p. 72, 1880; Thienemann, Zeits. Wiss. Insektenbiol., 1, pp. 287-289, 1905; Ulmer, Deuts.
Ent. Zeits., p. 385, 1913.
The species was described from a larval case. In 1905, some larvae and
pupae were obtained by Thienemann, who was able to give a description and
figures of the male genitalia from a mature pupa. A figure is given by Ulmer of
the wings of a Paraguan insect which is doubtedly referred to this species.
3. Peltopsyche sieboldi Fr. Müller.
Peltopsyche sieboldi Fr. Müll., Arch. Mus. Nac. Rio Jan., 3, p. 124, 1880 ; id., Zeits. Wiss., Zool., p. 74,
1881.
The species was described from a larval case, the adult remains unknown.
4. Peltopsyche maclachlani Fr. Müller.
Peltopsyche maclachlani Fr. Müll., Arch. Mus. Nac. Rio Jan., 3, p. 133, 1880; id., Zeits. Wiss.
Zool., 35, p. 74, 1881.
The species was described from a larval case, the adult remains unknown.
5. Mortoniella albolineata Ulmer.
Mortoniella albolineata Ulm., Notes Leyd. Mus., 28, p. 98, fig. 104, 1906 ; id., l.c., 29, p. 44, 1907.
This species is not a true Mortomella and in all probability should find a place
in the new genus Antoptila described below.
Sta. Catharina, Rio Janeiro, Organ Mts., Tijuca, 500-900 m., E. R. Wagner,
1902 (in the Paris Museum).
Canoptila gen. n.
Ocelli present ; antenna with the basal joint long, second short, third and
other joints each nearly twice the length of the second ; maxillary palpi with the
first joint short, the second slightly shorter and globular, third longer than the
first, fourth slightly shorter or as long as the third, fifth as long as the third ;
anterior wing with an oblique apical margin, costal and posterior margins more
or less parallel, neuration fairly regular ; in the posterior wing, the costal margin
is excised towards the centre ; spurs 1, 4, 4, that of the anterior leg very fine and
transparent. d
Genotype: Canoptila bifida sp. n.
6. Canoptila bifida sp. n. (text-figs. 128-133).
General characters as detailed in the generic description.
Genitalia g.—The ninth tergite is strongly produced in a large plate whose
apical margin, widely excised from above, slopes downward, from the side, in a
Novrrarrs Zoonoarcan XLI. 1939. 219
concave sweep, lower margin straight ; from near the hinder margin of the seg-
ment arises a pair of spine-like processes, incurved from above and beneath ;
penis membranous ; upper penis-cover helmet-shaped ; there are two pairs of
very strong, black processes whose function in the genitalia is uncertain ; lower
Canoptila bifida, g.—Fig. 128. Wings.—Fig. 129. Basal joints of the
antenna.—Fig. 130. Maxillary palpus.—Fig. 131. Genitalia, dorsal—Fig. 132.
The same, lateral.—Fig. 133. The same, ventral. i
penis-cover, seen in a wide excision of the ninth sternite, is comparatively narrow
at its base, apical margin very wide, truncate, with the apical angles produced
and bifurcate, partly membranous ; a long, pointed process to the sixth sternite.
Length of the anterior wing ¢ 2:25 mm. 1.x.1937; one dg.
Antoptila gen. n.
Spurs, 0, 4, 4. Ocelli present. Antennae twenty-three- or twenty-four-
jointed in the &, basal joint long, slightly curved, next joint short, remaining
joints all long, but not so long as that at the base. Maxillary palpi with the two
basal joints short, the second globular, shorter than the first, each of the remaining
joints about as long as the first and second together. Anterior wing long, costa
and sub-costa more or less parallel, apical costal margin oblique, neuration regular
but varying slightly in individuals in respect to the length of the forks ; posterior
wing long and narrow, costal margin slightly concave, fork no. 3 wanting.
Genotype: Antoptila brasiliana sp. n.
The genus is closely allied to Mortoniella Ulmer 1906. It differs in the pos-
terior wing, where fork no. 3 is present in that genus.
Novitates ZOOLOGICAE XLI. 1939.
7. Antoptila brasiliana sp. n. (text-figs. 134-138).
Anterior wings chocolate-brown with no white markings as occur in Mor-
toniella. Other characters as given in the generic description.
Genitalia 9.—Ninth tergite considerably produced, apex triangular, from
134
Antoptila brasiliana, 3.—Fig. 134. Wings.—Fig. 135. Maxillary
palpus.—Fig. 136. Genitalia, dorsal.—Fig. 137. The same, lateral.—
Fig. 138. The same, ventral.
above ; from the side, the upper margin is sinuous, terminating in a tailwardly
and upwardly directed, acute point ; the apical margin is obliquely truncate, with
the lower angle produced in a pair of processes, set close together, the upper longer
and more slender than the lower and curving over it; penis obscure ; lower
Noviratrs Zoonoagicarn XLI. 1939. 92]
penis-cover long and sinuous from the side, with a projecting angle on its lower
margin towards the base ; the apical margin of the eighth sternite is exeised at
the centre with the margin on each side of the excision slightly elevated and
bearing four extremely long, black, wire-like bristles which extend to the apex of
the genitalia ; a very strong process to the sixth sternite.
Length of the anterior wing 32:75 mm. ix-x.1936, ii. 1937, iv.1937, v.1937,
vii.1937, x.1937 ; numerous examples.
8. Protoptila dubitans sp. n. (text-figs. 139-142).
The shape of the wings and neuration as is usual in the genus.
Genitalia &.—The apical margin of the eighth tergite evenly rounded,
sternite produced in a long, tapering process with a narrow, excised apex ;
the ninth segment is somewhat retracted within the eighth; the dorsal
\
Protoptila dubitans, §.—Fig. 139. .Wings\—Fig. 140. Genitalia, dorsal.—Fig. 141. The
same, lateral—Fig. 142. The same, ventral.
margin is produced in a large plate with angular projections at the sides and
a deeply excised apex, the excision commencing at the extreme apical angles ;
from the side, the plate is deep with a blunt hook towards the centre of the lower
margin and a projecting, rounded process, fringed with hairs towards the basal
margin about midway ; arising from the lower basal angle, seen from the side, is
a strong, black appendage (? intermediate) with a dilated, truncate apex and
sinuous, lower margin ; penis membranous with a large hook on its under-surface,
directed basally ; inferior appendage (?) arising from a broad, membranous base
222 Novirates ZooLoGIcAE XLI. 1939.
and terminating in a strong hook whose lower margin is heavily fringed ; this
hook appears to be inserted in its base in a socket ; a strong process to the sixth
sternite.
Length of the anterior wing ¢ 3mm. 1.x.1937; one ¢.
9. Mexitrichia ormina sp. n. (text-figs. 143-148).
Wings slightly narrower than those of M. unota and M. teutona, with the
anterior wing less dilated towards the apex.
Genitalia 3.—Dorsal plate deeply excised at its apical margin to leave large,
elongated lobes on each side of the excision ; from the side, the plate is deep, with
an angular projection along the upper margin towards its base ; the plate tapers
to a fairly broad, rounded apex ; beneath it, seen in a balsam preparation, is an
Mexitrichia ormina, 3.—Fig. 143. Wings.—Fig. 144. Genitalia, dorsal.—Fig.
145.—Upper penis-cover, etc., dorsal—Fig. 146. The same, lateral.—Fig. 147.
Genitalia, lateral.—Fig. 148. The same, ventral.
upper penis-cover, excised at its apex ; the centre of this excision, as seen from
the side, is produced in an upwardly directed, sinuous spine ; beneath this, lying
above a membranous penis, is a fine, straight spine ; penis sheaths, from above,
caliper-shaped, sinuous from the side ; lower penis-cover formed of two wide
plates ; from beneath, each inner apical angle is produced in a pair of fine spines ;
there is a deep excision between the two plates with a rounded base ; from the
side, the inner angle appears as a stout, upcurving hook and there is an angular
Novirarrs ZoOLoGICAE XLI. 1939. 293
projection of the lower margin of the cover near the base ; a strong process to
the sixth ventral segment.
Length of the anterior wing, & 2 mm. 22.x.1937; one d.
10. Mexitrichia teutona sp. n. (text-figs. 149-152).
Insect brownish.
Genitalia &.—Extending beyond the margin of the ninth tergite is a dorsal
plate with the apical margin sharply excised ; the plate is pent-shaped and from
above, the sides project outwards in small, irregular shelves ; from the side, the
distal margin is excised with a deep, rounded excision ; beneath, is a spatulate
Mesxitrichia teutona, $.—Fig. 149. Wings.—Fig. 150. Genitalia, dorsal.—Fig. 151. The
same, lateral. Fig. 152. The same, ventral.
upper penis-cover with a dilated apex, curving upward from the side ; penis mem-
branous with a pair of caliper-shaped sheaths, inclining slightly upward, from the
side ; there is a single spine arising on the lower surface ; lower penis-cover widely
excised at its broad apical margin, with a large, asymmetrically directed spine
arising from the base of the excision ; from the side, the penis-cover is very deep,
the spine arising from the lower angle ; a stout process to the sixth sternite.
Length of the anterior wing, &3 mm. ix-x. 1936; 8. ii. 1937 ; 22.11. 1937 ;
ll. v. 1937; 1. xii. 1937 ; numerous examples.
11. Mexitrichia unota sp. n. (text-figs. 153-156).
Insect brownish.
Genitalia ¢.—The apical margin of the ninth tergite is rounded ; beyond it
is a large dorsal plate, narrowing to its apex which is deeply excised to leave a
17
224 Novirates ZoonocicaE XLI. 1939.
pair of finger-like projections separated by a U-shaped excision ; the plate is
pent-shaped, and, seen from the side, bears a large, rounded projection on its
lower margin, separated from the finger-like process by a wide excision ; beneath
the dorsal plate may be seen a narrow, straight upper penis-cover which is blade-
shaped and curves upward from the side ; penis membranous with a strong spur
on its lower surface ; penis sheaths divergent from beneath, curving downward
Mexitrichia wunota, &.—Fig. 153. Wings.—Fig. 154. Genitalia, dorsal.—
Fig. 155. The same, lateral—Fig. 156. The same, ventral.
Abtrichia, 9.—Fig. 157.—Anterior wing.
from the side ; lower penis-cover (?) with a pair of wings, directed outwardly
from beneath and upwardly from the side ; from this aspect, the wing is triangular,
its apex produced in a hooked spur directed distally, the lower distal angle pro-
duced and notched ; the whole structure arises from a stem which is slender and
constricted from the side, wide from beneath ; a strong process to the sixth
sternite.
Length of the anterior wing 4 mm. ix-x. 1936, 22. ii. 1937, 11. v. 1937,
8. vill. 1937, 1. xii. 1937 ; a series.
Abtrichia gen. n. (text-fig. 157, 9).
Insects large, with long, acuminate wings. Ocelli in both sexes two in
number. Antennae very short, twelve jointed in the ¢, seventeen in the 9. In
the 3, a large process arises from the side of the basal joint bearing an elaborate
structure having the appearance of regular or irregular rows of minute round pits ;
Novirates ZooLogIcAE XLI. 1939. 225
this process is much larger than the basal joint and has one or more membranous
leaves attached to it; the basal joint is itself long, next joint very short, sub-
sequent joints each about double its length. On the vertex of the head is a large
Abtrichia squamosa, 3g.—Fig. 158. Head, from in front.—Fig. 159. Head,
palpi, etc., lateral, antenna removed (a = socket of antenna)—Fig. 160. Shield on
the vertex of the head, viewed obliquely.—Fig. 161. Basal joints of the antenna
much enlarged.
plate-like shield, from the side, with a down-curved, acute beak, and it bears on
the dorsal surface a deep furrow down the centre ; above the labrum is a plate
which projects upwards towards the shield on the vertex and appears, from the
side, as a strong spur with an acute apex, triangular from in front. Maxillary
226 Novirates ZOOLOGICAE XLI. 1939.
palpi with the two basal joints short, the third nearly twice the length of the first
and second together, fourth a little shorter than the third, fifth about as long as
the third. In the anterior wing g, the costal margin is bent over the wing
towards its base for
MZ nearly half its length
SSS to make a fold or
pocket which, in
162 squamosa, is filled
= with small scales, in
antennata, with coarse
N hairs ; in the 9, and
in the posterior wing
in both sexes, the
wing is normal; neu-
ration as shown in
the figures. Spurs
1, &
Genotype: Ab-
trichia antennatasp.n.
The genus is
evidently allied to the
Mexican Zumatrichia
Mosely 1937, which
is furnished with a
plate to the basal
joint of the male
antenna and also has
but two ocelli.
12. Abtrichia squa-
mosa sp. n. (text-
figs. 158-166).
Anterior wing
clothed with brown
or blackish hairs,
intermixed with
irregular patches of
coarse white hairs,
Abtrichia squamosa, 3.—Fig. 162. Wings.—Fig. 163. Genitalia, .
p = an
dorsal.—Fig. 164. The same, lateral.—Fig. 165. The same, ventral.— d DU mern
Fig. 166. Penis, ventral. scales, partieularly
beneath the over-
lapping fold of the costa ; apex of the wing acute but not particularly produced ;
posterior wing without scales. Other characters given in the generic description.
Genitalia g.—The apical margin of the ninth tergite deeply and roundly
excised, the apical angles of the tergite produced in black, flattened, vertical
ridges outside which the tergite is again produced in large, wide side-pieces ; in
the excision is a long, dorsal plate with rounded apical margin ; penis stout and
straight ; in a balsam preparation are seen four bunches of feathery spines either
165
Noviratrs Zoonoaican XLI. 1939. 227
on its under-surface or perhaps contained within its walls ; these bunches are
arranged in two pairs, one pair between, and shorter than the other ; lower penis-
cover long, tapering to a blackened, hooked apex ; inferior appendages branched,
the branches long and slender, upper branches with the apices bent nearly at right
angles towards each other ; lower branch long and sinuous; margins of both
eighth and ninth sternites widely excised, ninth possibly open at the base of the
excision ; a short, stout process to the seventh sternite, acute from the side.
Length of the anterior wing, & 4mm. 1.xii.1937; one 3.
|
|
Abtrichia antennata, 3.—Fig. 167. Wings.—Fig. 168. Basal joints of the antenna,
much enlarged.—Fig. 169. Genitalia, dorsal.—Fig. 170. The same, lateral.—Fig. 171.
The same, ventral.—Fig. 172. Penis, ventral.
13. Abtrichia antennata sp. n. (text-figs. 167-172).
Anterior wing clothed with brown or blackish hairs intermingles with patches
of coarse, white hairs but without scales ; the fold of the costa is slightly shorter
228 Novrrarrs ZooLocIcAaE XLI. 1939.
than in squamosa; apices acute and produced in both anterior and posterior
wings; antennae and palpi as in sguamosa, but with a slightly different and
more irregular pattern of pitting on the lobe of the basal joint of the antenna.
Genitalia §.—Highth sternite considerably produced and strongly chitinized ;
ninth sternite broadly excised, the apical angles each bifurcate to make a pair of
rounded forks of which the inner is blackened ; outside these, the tergite is pro-
duced in large, rounded side-pieces which slope downward, from the side, with ~
sinuous upper margins ; in the excision is a membranous dorsal plate with trun-
cate apical margin ; penis straight, furnished either internally or on its under-
surface with two pairs of strong, black spines, the basal the longer ; lower penis-
cover, from the side, globular at its base, apex produced in a blackened hook, the
sides curving up over the dorsal surface of the penis ; inferior appendages un-
branched, rather short, straight and transparent ; a strong process to the seventh
sternite, blunt from the side but acute from beneath.
Length of the anterior wing, $ 4mm. 1.xii.1937; one £&.
There are in the collection two or three female examples belonging to either
one or other of the species in this genus, but I can find no means of associating
them with their respective males.
Acostatrichia gen. n.
Spurs 1,3, 4. Ocelli present. Antennae about nineteen-jointed in the 3,
basal joint long, second short, remaining joints each longer than the second, nearly
as long as the first. Maxillary palpi with the first and second joints very small,
third slightly longer than the fourth, fifth nearly twice the length of the fourth.
Anterior wing with a pouch along the costa near the base, filled with broadened
hairs, neuration as figured. In the 4, the seventh sternite bears a pair of small,
acute processes, one situated immediately behind the other.
Genotype.— Acostatrichia plawmanni sp. n.
Acostatrichia is closely related to the Mexican genus Costatrichia Mosely 1937,
differing in neuration, more particularly in that of the anterior wing, and also in
the normal antennae which, in the latter genus, are considerably dilated at their
centre. This may, however, be a specific rather than a generic character.
14. Acostatrichia plaumanni sp. n. (text-figs. 173-178).
Anterior wing bearing large patches of coarse, white hairs. Other characters
as detailed in the generic description.
Genitalia g.—The apical margin of the ninth tergite excised with a nearly
circular excision, the edges of which are partly obscured by a membranous dorsal
plate or, perhaps, an upper penis-cover ; beyond this is a more strongly chitinized
structure with a broad, excised apical margin, probably the penis and lower penis-
cover together ; from the side, the eighth segment is cut back so that its apical
margin is very oblique ; the ninth segment, from the side, slopes slightly down-
ward, its apical margin slightly excised to form a downwardly directed finger at
its lower apical angle ; on its lower margin, from the same aspect, towards the
centre, is a pair of finger-like processes, the lower the longer and bearing a thick,
black spine which is the most noticeable feature of the genitalia ; inferior appen-
dages small and triangular ; apical margin of the eighth sternite not excised at
its centre ; a paired process to the seventh sternite.
Novirarrs ZooLOGICAE XLI. 1939. 229
Length of the anterior wing, & 2:5 mm. ix-x.1936, ii.1937, iv. 1937, viii.
1937, xii. 1937; a series.
173
Acostatrichia plaumanni, §.—Fig. 173. Wings.—Fig. 174. Basal
joints of the antenna.—Fig. 175. Maxillary palpus.—Fig. 176. Genitalia,
dorsal.—Fig. 177. The same, lateral—Fig. 178. The same, ventral.
I have pleasure in dedicating this species to Dr. Fritz Plaumann, who has
collected extensively for the British Museum in Brazil.
15. Acostatrichia simulans sp. n. (text-figs. 179-182).
Anterior wing resembling that of A. plawmanni in being variegated with
patches of white ; the costal pouch is more conspicuous than in that species and
bears a larger mass of yellow hairs.
Genitalia ¢.—The apical margin of the ninth tergite excised as in plaumanni ;
the bases of the large, black spines of that species are, in simulans, represented by
long, transparent, slender processes, each with a short, stout, black and slightly
curved spine situated towards the apex; these processes curve towards each
other, caliper fashion, so that their apices nearly meet ; a second finger-like process
230 Noviratrs ZooLoctcar XLI. 1939.
is situated above as in plawmanni and is furnished with one or a pair of fairly long
bristles ; inferior appendages short and triangular ; apical margin of the eighth
sternite excised at its centre ; processes of the seventh sternite paired, the distal
the longer.
\
Acostatrichia simulans, &.—Fig. 179. Wings.—Fig. 180. Genitalia,
dorsal.—Fig. 181. The same, lateral.—Fig. 182. The same, ventral.
Length of the anterior wing, & 2 mm. ix-x.1936, 8.11.1937, 25.iv.1937,
1.xi1.1937; several examples.
The black spines of the processes of the ninth segment are easily shed so that
it has been difficult to find an example in the collection having both present.
When the spines are wanting, the species is less easy to recognise.
Betrichia gen. n.
Ocelli present. Antennae ¢ short, only nineteen-jointed, basal joint long,
next three joints shorter than each of the remaining joints. Maxillary palpi
wanting in the single example of the species available. Wings elongate with
acute and somewhat produced apices, neuration as figured. Spurs 1, 3, 4.
Genotype, Betrichia zilbra sp. n.
Novirarres ZooLoGIcAE XLI. 1939. 231
16. Betrichia zilbra sp. n. (text-figs. 183-186).
General characters as given in the generic description.
Genitalia 4.— The apical margin of the eighth tergite deeply excised ; in the
excision, from above, is seen the ninth tergite whose apical margin is shallowly
excised, the apical angles produced in a pair of rounded and irregular lobes ;
beyond these is a pair of processes whose attachments are obscure ; from the side,
Betrichia zilbra, &.—Fig. 183. Wings.—Fig. 184. Genitalia, dorsal.—
Fig. 185. The same, lateral.—Fig. 186. The same, ventral.
the eighth sternite is strongly produced, the upper margin sloping sinuously
downward to a truncate apex whose margin is serrate and fringed with a few long
bristles ; from the side is seen a membranous penis furnished with a pair of stout
sheaths, strongly fringed at their apices; lower penis-cover long and sinuous ;
from beneath, the eighth sternite is strongly produced, apical margin truncate
and somewhat concave, with long bristles at the apical angles; process of the
seventh sternite long and slender, very acute from the side.
Length of the anterior wing, ¢ 3 mm. ix-x.1936; one d.
232 NOVITATES ZOOLOGICAE XLI. 1939.
17. Exitrichia noteuna sp. n. (text-figs. 187-192).
Genitalia ¢.—The apical margin of the ninth tergite produced at its centre
in a rectangular dorsal plate ; in the angles formed between the base of the plate
and the margin of the tergite appear the long, finger-like side-pieces of the tergite ;
these, from the side, are broad and rounded, arising from constricted bases ;
beneath the dorsal plate is the apex of the penis, furnished with a pair of black
188
Exitrichia noteuna, &.—Fig. 187. Wings.—Fig. 188. Genitalia, dorsal.—Fig. 189. The
same, lateral.—Fig. 190. The same, ventral.—Fig. 191. Lower penis-cover, ventral.—Fig. 192.
Penis, lateral.
spines ; lower penis-cover large, with a truncate apex and with a large, down-
wardly directed process arising from its under-surface towards the centre; the
sides of the penis-cover, from beneath, are turned under to overlap partly the
under-surface ; the overlapping part is triangular with the apex deeply excised ;
inferior appendages long, broad at the bases, tapering to subacute apices ; the
margin of the ninth sternite produced at its centre in a small process with a
rounded apical margin.
Length of the anterior wing, d 2 mm. ix-x. 1936, 15. iv. 1937, 11. v. 1937,
8. vill. 1937, 1. x. 1937; numerous examples.
18. Exitrichia novara sp. n. (text-figs. 193-197).
Insect very small and brownish.
Genitalia g.—The apical margin of the ninth tergite slightly produced and
rounded ; side pieces of the segment with the apices only slightly dilated ; penis
slender, with a black spine, apex dilated from above ; lower penis-cover with a
truncate apical margin with a bristle at each apical angle and an angular, down-
wardly directed process about the middle of the under-surface ; inferior appen-
dages black, branched, broad and triangular from beneath ; from the side, the
Novrrares Zoonoatcan XLI. 1939. 933
upper branch is very stout, apex greatly dilated, with a truncate apical margin,
lower branch comparatively slender ; apical margin of the ninth sternite produced
at its centre in a slender triangle.
Length of the anterior wing, d 2 mm. ix.-x. 1936, i. x. 1937; two Id.
196
197
193 195
Exitrichia novara, &.—Fig. 193. Genitalia, dorsal.—Fig. 194. The same,
lateral—Fig. 195. The same, ventral.—Fig. 196. An inferior appendage, lateral.
—Fig. 197. Penis, lateral.
19. Exitrichia ovona sp. n. (text-figs. 198-202).
There appears to be an additional fork in the anterior wing of ovona as com-
pared with the neuration in the genotype, the Mexican species H. anahua, but the
neuration is so indistinct that, as the new species is represented by only a single
example, I have considered it better to retain it in Haitrichia until more material
should be available for study. The genitalia, however, are scarcely reconcilable
with the typical Exitrichia pattern.
Genitalia $.—From above, the apical margin of the ninth tergite is produced
in a rectangular dorsal plate whose apical margin is cut away in a wide, semi-
circular excision to leave the angles of the plate produced in slender processes
with down-hooked apices ; beneath this are slightly longer processes which may
234 Novirates Zootocicar XLI. 1939.
be the produced side-pieces of the ninth tergite ; penis long, with a pair of black-
ened hooks directed laterally, rather widely separated and situated towards the
apex ; lower penis-cover triangular with its apex turned down in a broad hook
with a single bristle on each side; inferior appendages short and triangular ;
apical margin of the ninth sternite bearing a small, triangular projection at its
Exitrichia ovona, 3.—Fig. 198. Wings.—Fig. 199. Genitalia, dorsal.
—Fig. 200. The same, lateral.—Fig. 201. The same, ventral.—Fig. 202.
Penis and lower penis-cover, lateral.
centre and with its lateral angles produced in slender processes ; from the side,
a pair of processes may be seen, elbowed towards the base and directed upward ;
these may be connected with the inferior appendages, but there is not sufficient
material available for a complete investigation of these structures.
Length of the anterior wing, ¢ 2mm. 1.x. 1937; one &.
The figures of this species are not entirely satisfactory owing to lack of
material.
Novirates ZooLoGIcAE XLI. 1939. 935
20. Exitrichia tertia sp. n. (text-figs. 203-206).
Genitalia ¢.—The apical margin of the ninth tergite very widely excised ;
in the centre of the excision is a nearly square, membranous dorsal plate ; the
angles of the excision are produced in straight processes with the apices concave,
the upper and lower apical angles acutely pointed ; side-pieces of the ninth seg-
ment long, gradually dilating from a slender base to a broad, rounded apex ;
penis retracted, furnished with a pair of stout, black apical claws emerging from
Exitrichia tertia, &.—Fig. 203. Genitalia, dorsal.—Fig. 204. The same, lateral.—Fig. 205.
The same, ventral.—Fig. 206. Penis and lower penis-cover, lateral.
an obliquely cut sleeve ; lower penis-cover trifurcate, the central fork, from the
side, appearing as a large, downwardly directed hook ; a single bristle on each of
the outer forks ; inferior appendages very large, slightly smuous, apices approach-
ing each other and rounded, with a small, projecting angle on each under-surface
near the apex ; apical margin of the ninth sternite deeply excised with the centre
produced in a triangular process.
Length of the anterior wing, {2 mm. ix-x. 1936; two 3d.
21. Dolotrichia (?) dubitans sp. n. (text-figs. 207-209).
The species dubitans departs somewhat from the typical pattern in the shape
of the dorsal plate as the apical angles are not produced in strongly chitinized
hooks.
Genitalia g.—The apical margin of the ninth tergite is produced in a large
dorsal plate whose apical margin is excised to leave two wide, rounded lobes ;
from the side, the apex is produced and slender ; beneath the plate is a long penis
236 NOVITATES ZOOLOGICAE XLI. 1939.
with an obliquely truncate apex or sleeve from which protrudes a single, black-
ened spur; there is an internal duct whose apex slightly projects beyond the
margin at the apex; beneath the penis is a pair of stout, blackened processes
whose apices are out-turned ; these processes are very deep from the side, with
hooked apices ; inferior appendages from beneath, bifurcate, outer forks nearly
Dolotrichia dubitans, $.—Fig. 207. Genitalia, dorsal.—Fig. 208.
The same, lateral.—Fig. 209. The same, ventral.
twice the length of the inner, with serrate inner margins towards the apices ; the
inner forks are each armed at the apex with a single bristle ; side-pieces of the
ninth segment bifurcate ; from the side, the upper fork is widely separated from
the lower by a rectangular excision and is twice its length ; the apices of each
armed with long bristles ; apical margin of the ninth sternite widely excised and
with the centre produced in a long triangular process.
Length of the anterior wing, jf 2mm. 25. iv. 1937; one 3.
Although this species scarcely conforms with the generic diagnosis, yet, as it
is represented by a single example only, it seems undesirable to erect a new genus,
until more material shall be available.
22. Hydroptila producta sp. n. (text-figs. 210-215).
The species belongs to the sparsa group. Antennae, thirty-seven-jointed in
the 3, basal joint large, irregularly dilated at one side of its apex, second joint
Novirarrs ZOOLOGICAE XLI. 1939. 237
about twice as long as the third, which is slightly shorter than each of the remain-
ing joints; scent-organ consisting of a pair of eversible filaments fringed with
unusually broad scent-hairs ; the scent-organ cap of the usual acorn-pattern, the
cup being much larger than the acorn.
Genitalia &.—Dorsal plate very large and broad, the apical margin excised
from the extreme apical angles ; an angular projection towards the centre of the
210
Hydroptila producta, §.—Fig. 210. Wings.—Fig. 211. Basal joints of the
antenna.—Fig. 212. Scent-organ cap and scent-hairs.—Fig. 213. Genitalia,
dorsal.—Fig. 214. The same, lateral.—Fig. 215. The same, ventral.
dorsal basal margin ; side-pieces of the ninth segment small; penis long and
straight, with a single long spine lying more or less parallel with it and slightly
shorter, the usual, twisted, apical hook wanting ; inferior appendages with the
apices, from the side, considerably dilated, with a blackened apical hook on the
upper apical angle, directed outward as seen from beneath, margins fringed with
widely spaced bristles ; lower penis-cover rounded, with a pair of minute bristles
slightly below the centre of the apical margin ; an acute process to the seventh
sternite.
Length of the anterior wing, ¢ 2 mm. 8. ii. 1937, 11. v. 1937, 1, xii. 1937 ;
several specimens.
bo
=
wm
NOVITATES ZOOLOGICAE XLI. 1939.
23. Loxotrichia zilaba sp. n. (text-figs. 216-220).
Neuration conforming with the Mexican pattern.
Genitalia. $.—The eighth tergite is very hairy with the apical margin deeply
excised from the outer angles ; in the centre of the excision, is a straight penis,
below which, on each side, is an elongate, oval process with a slender, out-
Loxotrichia zilaba, 3.—Fig. 216. Wings.—Fig. 217. Genitalia, dorsal.—Fig. 218. The same,
lateral.—Fig. 219. The same, ventral.—Fig. 220. Penis, etc., with the ninth sternite.
stretched spine arising from each apex ; ninth sternite produced and narrow, its
apical margin excised as seen from above and beneath, forming a short claw from
the side ; eighth sternite with a deep V-shaped excision ; a strong, pointed pro-
cess to the seventh sternite.
Length of the anterior wing, ¢ 2:5 mm. ix—x.1936; one {.
INDEX.
Abtrichia antennata sp. n.
5 squamosa sp. n.
Acostatrichia plaumanni sp. n.
% simulans sp. Nn.
Antoptila brasiliana sp. n.
Betrichia zilbra sp. n.
Canoptila bifida sp. n. .
Diaulus ladislavii Fr. Müller
Dolotrichia dubitans sp. n.
218
. 235
SoU spe p es
bo
bo
S
NOVITATES ZooLoGIcAE XLI. 1939.
18
Exitrichia noteuna sp. n.
ss novara sp. n.
ovona sp. n.
ns tertia sp. n..
Hydroptila producta sp. n.
Loxotrichia zilaba sp. n.
Mexitrichia ormina sp. n.
55 teutona sp. n.
59 unota sp. n.
Mortoniella albolineata Ulmer
Peltopsyche sieboldi Fr. Müller
iS maclachlani Fr. Miller
Protoptila dubitans sp.n. .
Rhyacopsyche hageni Fr. Müller
BB > "U SH CB DUP SS
232
232
233
235
236
238
222
223
223
218
218
218
221
. 218
240 Novirates ZooLoGICAE XLI. 1939.
SOME OBSERVATIONS ON THE GROWTH OF PERINGIA
ULVAE (PENNANT) 1777 IN THE LABORATORY.
By ANNE ROTHSCHILD Ann MIRIAM ROTHSCHILD.
(With 3 text-figures.)
INTRODUCTION.
(DUBING the study of the effect of Trematode parasites on the growth of
Peringia ulvae, an attempt had been made to estimate the age of individual
snails by some external feature. Great variation was recorded in the size, shell
colour, shell shape and shell texture of this molluse (Rothschild 1938). It seemed
probable that this variation was linked with environmental factors, thus rendering
the size of the snails an unsuitable criterion for estimating their age. The
following experiments were undertaken with the hope of obtaining some definite
data on this subject, and to prove whether, in the laboratory at any rate, some
slight differences in environment produced any marked contrast in the growth
rate, shell colour, etc., of the snails. It was also thought that, simultaneously,
the growth of specimens with Trematode infections might be compared with
those free from parasites.
A number of specimens of P. ulvae ranging from 0-6 mm. to 5:3 mm. in
length were collected at random from two different habitats, the mud-flats of
St. John’s Lake and a permanent pool in the saltings at “ Egypt,” Plymouth
(see Map, Hartly & Spooner, 1938). A certain number of each were placed in
finger bowls and test tubes respectively, and their growth compared by measure-
ments taken periodically.
TECHNIQUE.
The snails were collected by placing scoops of mud in dishes and allowing
the specimens to crawl out. Any infected individuals were included purely
accidentally, as they were not isolated before the commencement of the experi-
ment in order to determine whether they were parasitized.
The glass test tubes and finger bowls in which they were placed were filled
to within two inches of the top with a mixture of half sea and half fresh water.
The snails were provided with a very plentiful supply of ulva—a seaweed upon
which they frequently feed in nature. The water in both types of vessel was
changed once a month. All were kept at room temperature upon a wooden
shelf in the laboratory. The positions of individual bowls and tubes on the shelf,
as well as all bowls and tubes en bloc, were changed from time to time.
From two to six specimens were placed in each bowl, and from two to four
in the tubes. Under these conditions the snails appeared healthy and quite
active. The survival rate was in fact remarkably high,! only four snails dying out
of a total of 152.
Measurements were taken once a month from December 1937 to December
1938 with the aid of a micrometer. The larger size groups were measured,
1 In this connexion it is worth noting that in brevet jars containing several hundred P. ulvae
—even if the water is changed daily and the food supply is unlimited—the mortality is high, perhaps
more than 50 per cent, of the snails die off within six months,
Noviratrs Zoonogican XLI. 1939. 241
less accurately, with a ruler marked in quarter millimetres. In the case of the
subsidiary tube experiment, which is considered separately, measurements were
made at long irregular intervals over a period of three years.
THE GROWTH OF P. ULVAE IN Bowls AND TUBES
It is, of course, well known that several species of fresh-water Limnaca are
dwarfed if kept in small vessels. The exact cause of this dwarfing is evidently
not properly understood. The result of the experiment with P. ulvae shows
clearly that specimens from both habitats grew considerably less when kept in
tubes than in bowls. Measurements are given below in Tables I and II.
TABLE II.
GROWTH OF INFECTED AND UNINFECTED SPECIMENS COMPARED.
(Numbers in brackets indicate number of specimens.)
Locality, Mean Initial Final Size Final Size Sex of Infected
Bowl and Tube No. | Size (Dec. 1937) Uninfected Infected Snails
St. John’s Lake :
Bowl 3. . 5 0-67 (2) 4-45 (2) Zi =
Bowl 5 . $ 0-79 (4) 4-76 (3) 5-70 (1) ©)
Bowl 4 0-87 (4) 5-11 (3) 6-11 (1) 3
Bowl 2 0-91 (2) 4-46 (2) an =
Bowl 13 -. 5 1-44 (4) 5-19 (4) =e pis
Bowl 10 . : 1-71 (4 5:29 (3) 6-22 (1) d
Bowl 14 . 3 1-93 (4) 5-35 (4) = a
Bowl 12 . 5 1-95 (3) 4-86 (2) 4-64 (1) 3
Bowl 11 . j 2-13 (4) 5-35 (4) en ais
Bowl 18 . 3 3-75 (4) 4-99 (2) 5-44 (2) 32
Bowl 18a 4-19 (4) 4-97 (3) 5-21 (1) 3
Bowl 19 4-63 (4) 4-87 (3) 6-85 (1) 3
Tube 7 . : 1-32 (3) 2-33 (3) — —
Tube 15 . ; 1-63 (4) 2-69 (4) — >
Tube 16 . 2 2-13 (4) 2-73 (4) ne u
Tube 17 . 5 2-94 (4) 3-94 (3) 4-95 (1) Q
Egypt Saltings :
Bowl 20. , 1-09 (3) 4-43 (3) — —
Bowl21 . : 1-76 (3) 5:35 (3) _ —
Bowles) 0. 1-87 (5) 4-97 (5) — —
Boul 2 5) 5 2-07 (3) 4-60 (3) = =
Bowl 23. ; 2-11 (4) 5-35 (4) = en
Suspected of
Infection
Bowl 27. ! 2-53 (5) 5-37 (4) 5-93 (1) 8
Bowl 28. : 3-92 (6) 5:23 (5) 5-60 (1) 3
Bowl 29. : 4-71 (6) 5-39 (6) = =
Tube 24 . a 2-02 (3) 2-61 (3) — —
Tube 25 2-24 (4) 3-28 (4) © — —
242 Novirares ZooLoGIcAE XLI. 1939.
It will be seen that as a rule specimens kept in bowls, measuring under
2-6 mm. at the commencement of the experiment, had increased at the end of a
twelve-month period to a mean length of about 5 mm., whereas tubed snails
attained a mean of only approximately 3 mm. Even though snails with an
initial measurement of over 3 mm. do not grow as rapidly as the smaller speci-
sy
23 Sa esa ESO T8
z
z
z
es;
Dec | Jan | Feb | Mar | Apr | May | June | July] Aug | Sep | Oct Nov | Dec |
Text-Fic. 221.—GROWTH-RATE oF P. ULVAE FRoMm Ecyrt Savrınas. In Bowis (———) AND
TUBES (.)
Measurements at monthly intervals from December 1937 to December 1938.
B5: 2 bowls, 7 specimens T7, 3 specimens
B6: 1bowl, 3 35 T8, 3 oo
B7: 3 bowls, 11 = T9, 4 on
B8: 2 bowls, 12 5 T10, 4 9
B9: 1bowl, 6 = T11, 4
B10: 1 bowl, 6
Bll: 1 bowl, 6 a
iraphs B9, B10, B11, T10, T11 are smoothed freehand curves (see text).
mens, the contrast between the growth rate of specimens in bowls and tubes
remained constant. A comparison, for example, of bowls Nos. 10, 11, 18 and
19 with tubes Nos. 15, 16, 17a and 31 show that the average increment of snails
from the bowls is three to four times as great as that of the tubed snails.
Four snails from tube 6 were changed to a bowl in April 1938. During this
four-month period in the tube the mean size of the snails had increased by only
Novirates ZooLoGIcAE XLI. 1939. 243
0:05 mm. Snails of a similar initial size in a bowl increased by an average of
1:02 mm. during the same period. After the change to a bowl three specimens
(the fourth died) rapidly increased in size, eventually reaching a mean of 4:38 mm.
in December 1938.
In text-figs. 221 and 222 the curves marked B—representing the average of
aoe PSS ER ASSS
7
Feb | Mar | Apr | May | June | July | Aug | Sep | Oct | Nov | Dec |
Text-Fic. 222.—GROWTH-RATE OF P. ULVAE FROM ST. JOHN’S LAKE. In BowLs (———) AND
TUBES (mem).
Measurements at monthly intervals from December 1937 to December 1938.
Bl: 4 bowls, 13 specimens T1, 3 specimens
B2: lbowl, 4 » Te, Zh 5
B3: 4 bowls, 15 54 T3, 4 ns
B4: 3 bowls, 12 5 T4, 4 Ss
TB, tube and bowl, 3 55 T5, 4 Ne
a number of bowls containing specimens of approximately the same initial size
—are compared with specimens from tubes (T). In order to simplify the
diagram, smoothed freehand curves have been given for specimens of 4 mm. and
over (which were measured with the aid of a ruler only). Many of these large
individuals grew very little during the twelve months and the fluctuations in the
measurements, which are noted in the Tables, can probably be accounted for by
chipping of the spires.
It is interesting that, apart from the different rapidity of increase in size,
the specimens in bowls and tubes display certain differences in their growth
curve. Thus the growth rate of the specimens in bowls falls abruptly in August
244 NovITATEs ZooLoGIcAE XLI. 1939.
irrespective of age and size. Even those specimens of under 2 mm. which were
started in April instead of December display this feature. On the other hand,
the tubed snails tend to stop growing from April to July—the period of maximum
growth of the snails in the bowls—but to recover with a slow steady growth
subsequently.
RESEMBLANCE BETWEEN THE GROWTH OF SPECIMENS FROM MUD-FLATS (ST.
JOHN’S LAKE) AND SALTINGS (EGYPrT).
In nature, random collections of P. ulvae from the mud-flats of St. John’s
Lake and permanent pools from the saltings at Egypt, display certain marked
Dec | Jan | Feb | Mar | Apr May | June | July Aug | Sep | Oct | Nov Dec
Texr-FıG. 223.—GROWTH-RATE OF P. ULVAE FROM Eaypr SALTING (———) AND ST. JoHN’S
LAKE (wenn).
Obviously abnormal specimens with retarded growth have been omitted (one specimen from
Bl, and one from B7), and the aggregates modified accordingly.
differences (Rothschild 1938, Plates I & II). The former are of medium size
(maximum infected specimens reaching about 5-8 mm. in length) with a medium
hard, grey or reddish shell, showing fairly well-preserved spires. The latter are
very large (maximum infected specimens reaching 10-5 mm. in length) with an
exceptionally hard, thick, very red shell, showing blunted spires in all large
individuals. In the laboratory, snails collected from these two habitats exhibited
similar growth curves. These are compared in text-fig. 223, in which single
specimens of obviously abnormal growth have been omitted. The two sets of
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Noviratrs ZooLoGIcAE XLI. 1939. 245
curves are, in fact, remarkably alike, and it would seem safe to suppose that the
differences of growth shown by P. ulvae in nature are not due to inherited
characters, but environmental factors.
In both groups of snails the specimens measuring 1 mm. in December (which
were kept in bowls) reached 4:50 mm. by the following August, and those of
1-50 mm.—2 mm. reached 5 mm. in the same period. Thus they overtake speci-
mens which were 4 mm. in length at the start of the experiment. Any size
groups which could be distinguished in December are merged into a single group
by the following summer.
The shells of all snails were grey in colour at the end of the twelve-month
period, and rather more friable than in normal specimens from St. John’s Lake,
but, on the other hand, the spires were better preserved. In those snails which
had been started at 4 mm. or over, the spires were inclined to chip off or blunt.
It is interesting to find that the specimens from Egypt, which were put in
bowls and tubes at 4 mm. or over, failed to grow appreciably, although in the
wild they would have presumably added 2-24 mm. to their length. They
remained at about the size of maximum uninfected specimens from St. John’s —
Lake, and also lost their red colour and thick, hard shells.
This may have been due to a sudden change of environment at a relatively
late stage, but it also seems probable that the growth conditions in the laboratory
are inferior to those of many of the permanent pools of the saltings, but superior
in bowls, at any rate, to those pertaining on the mud-flats.
COMPARISON BETWEEN UNINFECTED AND INFECTED SPECIMENS OF P. ULVAE.
When the growth experiment was terminated in December 1938 all the snails
were dissected and examined for the presence of Trematode parasites. Twelve
infected specimens were found in the material from St. John’s Lake and two
from the permanent pool in the saltings. A third specimen from Egypt was:
suspected of infection, but died a day or two before dissection.
The size of these infected snails was quite consistently larger than that of
uninfected specimens as shown by the data presented below in Table II. (The
final figures represent the mean of the November and December measurements.
A few specimens from the bowls in which growth was abnormally low have been
omitted. These were all uninfected snails.) The evidence obtained in the
laboratory therefore supports the theory (Rothschild 1938) that infection with
Trematode parasites causes increased growth.
Out of a total of fourteen infected snails eight proved to be males and six
females. The total number of uninfected males was fifty-two, and of females
sixty-six. However, some of the smallest specimens may have been males in
which the penis had not yet developed. These figures are therefore inadequate
for the purpose of studying the question of possible sex reversal of infected
females. It is, however, worth noting that an infected male attained a length of
6-85 mm., whereas the maximum uninfected male reached only 6 mm.
As in nature, the penis of all infected snails showed abnormalities in size
and shape.
COMPARISON IN GROWTH RATE OF MALES AND FEMALES.
Although the figures at our disposal are not very large there is sufficient
data to show the average growth rate of uninfected. females during the period
246 NovitatEs ZooLoGIcAE XLI. 1939.
of this experiment was higher than that of uninfected males. A comparison of
both sexes is made in Table III. (Tubes in which there was only very little or
no growth are omitted.) This agrees with the evidence obtained by examining
samples of P. ulvae from various habitats and tends to confirm the suggestion
that in this species the larger size of the females is due to increased growth rate
rather than a longer life (Rothschild 1938).
TABLE III.
GROWTH OF MALES AND FEMALES (UNINFECTED).
(Figures in brackets indicate number of specimens.)
Mean Final Mean Final
Locality and Bowl and Tube No. Initial Length Length of Length of
Males Females
Egypt : Tubes 20c, 24,25 . . | 1:90-2:60 3-25 (5) 2-91 (3)
St. John’s Lake : Tubes 17, 17a, 17b . | 2-50-5-00 3-91 (3) 4-32 (7)
Egypt : Bowls 20, 20a, 20b . 0 under 1-25 4-24 (4) 4-58 (6)
St; John's Lake: Bowls) 3,4,5,6 0-65-0-95 4-56 (6) 4-85 (5)
Egypt: Bowls 21, 22, 23, 26, 27 . 1-55-2-77 4-52 (10) 5-61 (9)
St. John’s Lake: Bowls 10, 11, 12, 13, 14 1:37-2:27 4-97 (6) 5-39 (11)
Egypt : Bowls 28, 29,30. 5 3-50-5-50 5-21 (6) 5-67 (10)
THE SUBSIDIARY TUBE EXPERIMENT.
From April 1936 to March 1939 nine specimens of P. ulvae collected from
St. John’s Lake were kept in tubes and measured at several months interval.
One snail died in December 1937. The individual measurements are given in
Table IV. It will be seen that the snails grew most rapidly during the first twelve
months and from thereafter only increased very slowly, but steadily, in size for
eighteen months. During the final six months of the experiment only two snails
increased at all, and then only by 0-10mm. Three specimens remained stationary
and three became reduced in length, probably by chipping off the spires. It
would therefore appear likely that under these conditions the snails reach their
limit of growth by the end of the third year.
TABLE IV.
SUBSIDIARY TUBE EXPERIMENT.
|
5 April Oct. May Dec. | March Oct. March
Tube No. 1936 1936 | 1937 1937 1938 1938 1939
1 v8 | 0 2-2 3-2 3-2 41 41
2 0:6 2-] 2-8 3:3 3:5 42 4-2
3 0-6 2-7 3-2 3-9 41 44 4-2
4 1 | 8d 3-4 4-0 4-2 45 46
5 12 | 3.2 3-7 4-2 43 4-9 5-0
6 1-9 | 3-5 3:8 4-0 4-2 4-7 46
7 1-9 3:8 4-0 4-0 4-3 4-8 4:8
8 20 | BME Beil aM 4-5 4-9 4-8
Novirates ZooLoGIcAE XLI. 1939. 247
SUMMARY.
1. A comparison is made between the growth rate of P. ulvae, collected from
two different habitats and kept in the laboratory in (a) glass bowls, (b) glass tubes.
2. The snails kept in bowls grow considerably faster than snails kept in
tubes ; their size is consequently no criterion of their age.
3. The growth curve is different in specimens kept in bowls and in tubes.
4. When reared in the laboratory, specimens collected from the mud-flats
and saltings, which in nature differ from each other in size, shell shape, shell
colour and shell texture, show no such variation. These differences are therefore
thought to be due to environmental rather than genetic factors.
5. Snails parasitized by Trematodes grow faster than uninfected specimens.
6. The uninfected females grow faster than the uninfected males.
ACKNOWLEDGEMENTS.
Our best thanks are due to Mr. G. M. Spooner for his invaluable help and
advice.
REFERENCES.
Hartley, P. H. T., and Spooner, G. M., 1938. The Ecology of the Tamar Estuary.
. I. Introduction. —In J. Mar. Biol. Ass., Plymouth, 22, No. 2, pp. 501-508,
pl. xviil.
Rothschild, M., 1938. Further Observations on the Effect of Trematode Para-
sites on Peringia ulvae (Pennant) 1777.—In Nov. Zool., xli, pp. 84-102,
7 figs. 2 pls.
248 Novirarrs ZooLoGIcAE XLI. 1939.
ON SOME OLD-WORLD AGARISTIDAE (LEPID.).
By KARL JORDAN.
(With 2 text-figures.)
1. Mimeusemia econia hedya subsp. nov.
39. Abdomen black above and at sides, not banded with yellow, underside with
broad yellow stripe from base to yellow apical tuft. Segment I of foretarsus
yellow, the other segments black, with white apical spot; mid- and hindtars
black with indication of white scaling at tips of segments.
Wings as in M. e. econia Hamps. 1900 (from Christmas Island, in the Tring
collection also from West Bali) ; the spots of the forewing purer white and as
a rule a little larger, especially the median spot, which is not transverse, but
irregularly rounded.
Anal sternite of ¢ dilated at some distance from apex ; harpe a moderately
curved pointed hook, which is somewhat broader in middle than in M. e. econia.
Hab. West Celebes : Goenong Rangkoenan, Paloe, 900 ft., xi.36 (J. P. H.
Kalis), 3 &&, 2 99.
2. Mimeusemia limbata sp. nov.
3. Similar to M. econia Hamps. 1900, but larger and the forewing with
two instead of three large creamy white spots. Frons and occiput with black
median stripe instead of being entirely white. Abdomen black, apex orange ;
in ¢ basal fourth of underside orange-yellow, this colouring extended laterally
farther back than medianly, in 2 a whitish lateral spot at base. Breast and legs
orange, hindtibia and all tarsi black, first foretarsal segment orange, tips of
tarsal segments white, hindtibia with a submedian spot and the tip yellow, fore-
and midtibiae with distinct or vestigial subapical black dot.
Upperside of wings black. Forewing with the usual metallic markings; a
short subbasal streak below cell and a larger one along hindwing, a narrow
diffuse streak above M? and a broader one below M? and a marginal border about
24 mm. broad from apex to tornus chestnut-red ; a small subcostal dot near
base, an antemedian oblique spot touching base of M?, 5-6 mm. long and 2} mm.
broad, a small spot in upper angle of cell, and a discal spot somewhat larger
than antemedian one, 6 mm. long and 31-4 mm. broad, neither reaching costal
margin nor M*, creamy white. Fringe white at apex of wing and above tornus.
On hindwing a rounded costal spot outside apex of cell, 44 mm. long and
3-33 mm. broad, and a larger patch in front of abdominal margin white, this
patch entering cell, distally rounded, longest posteriorly (about 8 mm.), the
extreme base and the fringe of abdominal margin remaining black ; fringe of
distal margin white at apex and before tornus, mixed white and black in between.
On underside the white markings larger than above ; the chestnut colouring
absent ; hindwing with a white basal subcostal streak.
Anal sternum of 3 much narrower than in M. econia ; harpe longer, more
abruptly curved, on the whole somewhat narrower, but less pointed.
Length of forewing: 911, 9 11-5 mm.
Novrrarrs ZoonoarcaAn XLI. 1939. 249
Hab. West Celebes: hinterland of Paloe Bay, Lindoe, 3700 ft., iv.37
(J. P. A. Kalis), 1 g, 1 9.
3. Mimeusemia eumelas Jord. 1912 (text-fig. 224).
A small collection of Heterocera acquired by the late Lord Rothschild from
M. E. Le Moult and labelled Toeelo, Halmaheira, contains a series of 15 Mimeu-
semia which, at first sight, look as if they all belonged to one somewhat variable
species. On closer inspection two of the specimens prove to be different in the
palpus, pattern, and genitalia. These two are a subspecies of Mimeusemia
simplex Lucas 1891, and the other 13 are-Mimeusemia eumelas Jord., described
from a ¢ and a Q
from Halmaheira in
Seitz, Macrolep. xi,
p. 18, pl. 3, f. (1912),
as M.simplexeumelas.
The present material
proves eumelas to be
a separate species.
In pattern M.
eumelas is distin-
guished from M. sim-
plex by the white
colouring of the frons
occupying less space
than the black central
scaling, the white
stripes of the thorax
being thinner, the
third segment of the
palpus entirely black,
in the & elongate-
elliptical, not ovate as in the Moluccan subspecies of M. simplex, segment II
entirely black on outside, without white scaling externally at apex, the
creamy band on upperside of forewing narrow, not quite reaching to M?, but
frequently followed by an isolated spot below M2, fringe of hindwing entirely
white, the proximal scales not being black as in M. simplex. Anal tergite of 4
broader in lateral aspect, but more sharply pointed ; the long sternum divided
as in M. simplex, broader, the sides not incurved, and the teeth of the apical
area more numerous and shorter. Clasper (text-fig. 224) much broader than in
M. simplex ; harpe a simple, slender, sharply pointed hook devoid of the teeth,
projecting upward from dorsal margin in M. simplex (text-fig. 225, M. s. buruensis
Jord. 1912); the basal median fold (m) without the rounded hump (h) of M.
simplex. 'The two projections of penis-sheath without brush of bristles.
Text-Fic. 224.—Mimeusemia eumelas.
Tuxt-Fic. 225.—Mimeusemia simplex buruensis.
4. Argyrolepidia aequalis repens subsp. nov.
Q. Nearest to A. a. aequalis Walk. 1864, but hindwing with one small white
central spot. Dorso-lateral stripe of segment II of palp deeper orange ; white
borders of abdominal segments narrow, on upperside little more than indications
250 NOVITATES ZooLoGIcAE XLI. 1939.
of bands, on underside distinct, but much narrower than the black portions of
the segments.
Upperside of forewing : whitish blue spots reduced ; in cell a minute white
dot ; on disc an oblique white band as in A. a. aequalis, at widest point a little
over 2 mm. broad, posteriorly 6 mm. from tip of fringe; the latter white at apex
of wing and bearing a few white scales at tornus. Hindwing : fringe white-
spotted as in A. a. aequalis ; outside discocellulars a small white spot, diameter
about 14 mm.
On underside the blue markings small as on upperside, except the discal
spots of hindwing, which are about as large as in A. a. aequalis ; on forewing
the white dot in cell and the discal band somewhat broader than above ; the white
central spot of hindwing truncate on proximal side, size about 24 x 3 mm.
Length of forewing: 24 mm., width 14 mm.
Hab. Halmaheira : Toeelo, 1 9 received from M. E. Le Moult.
Novirares ZoonoaicaAn XLI. 1939. 251
ON THE CONSTANCY AND VARIABILITY OF THE
DIFFERENCES BETWEEN THE OLD WORLD SPECIES OF
UTETHEISA (LEPID.; ARCTIIDAE).
By KARL JORDAN.
(With 30 text-figures.)
ke pretty Arctiid speckled with black and scarlet on the ereamy-white
ground of the forewing which favours this island by an occasional visit,
and makes the heart jump in the breast of collectors old and young who know
what is the white flake they see flutter in the air, belongs to Utetheisa, a genus
of fourteen species, three of them being American and the others occurring in the
Old World, mainly in the tropics. They are all closely related to one another,
perhaps with the exception of U. galapagensis Wallengr. 1860, which stands
apart, because its female (the ¢ is not known to me) has the antenna dentate,
whereas in that sex of all the other species the antenna is simple. With this
exception, the structure and colouring prove the species to form a compact
natural genus, the relationship being unmistakable. On closer investigation
the genus proves nevertheless to be a bundle of contradictions which obscure the
real relationship of the species with one another and have led the systematists
astray in their attempt to group the species in a natural order. But putting
the counters into their right places is only the beginning of a game. The species
have arisen from a common basis, each species representing the ancestor in a
specifically modified form. The differences acquired and the similarities retained
are facts before us that ask for an enquiry into their meaning. Speaking of the
genus in the aggregate, the specific distinctions (or “ specifics ” for short) refer
to the colour and pattern of the wings (hereafter called coloration), the genital
armature of the male (segments IX and X of the abdomen) and female (sternum
VII), the tarsal claws, and, in the male only, to the antenna, the hindleg and a
scent-organ on the hindwing.
The pattern of the wings of Lepidoptera being under the influence of the
neuration, it probably consisted originally of ill-defined stripes along the veins
and in the folds between the veins; these stripes broke up into spots forming
rows between the veins and across them. Such a groundwork of the pattern is
preserved or indicated in Utetheisa and many other Arctiids (inclusive of Hypsids).
The original pigment derived from the haemalymph was probably black and red,
amalgamated into brown or separated as red and black patches or spots, unpig-
mented scaling filled with air being white. The primitive colours red and black
or the modifications of red (orange, yellow and white pigments) are characteristics
of many Arctiids. In this respect the most primitively coloured Utetheisa
evidently is the American U. bella L. 1758. This common insect presents the
three principal stages in the evolution of wing-colour, the forewing being either
scarlet or yellow and the hindwing scarlet, yellow or white. The two American
U. bella and ornatrix L. 1758 would be excellent subjects for the experimental
study of colour-development and variation ; they are easily bred and produce
several generations a year. Somebody might take up these insects instead of
Drosophila.
252 Novrrares ZooLoGIcAE XLI. 1939.
The upper- and undersides were alike in the ancestral Utetheisa, as they still
are in the females of Argina Hübn. 1819. The black spots of the upperside
became reduced and acquired pale rings in U. bella and in the males of Argina.
The uppersides remain at some such early stage in bella, pulchella, pulchelloides
and lotrix, while in the other species the modification of the pattern proceeded
farther. As in most moths, the evolution of the exposed upperside of Utetheisa
has not been the same as on the concealed underside. The highest development
of the Utetheisa coloration is found in elata (inclusive of fatua from the African
Continent), in which, in contradistinction to all the other species, the two sides
are almost identical; evolution having started with similarity of the upper-
and undersides terminates in this species with both sides strongly modified, but
again nearly alike.
The Old World species nearest the ancestral coloration are U. pulchella L.
1758, U. lotrix Cram. 1777 and U. pulchelloides Hamps. 1907. For the sake of
simplification these names cover in this general survey all the subspecies as well
as those geographical representatives which we consider to have attained specific
rank. In all three species the forewing is creamy white speckled with black and
red ; there are exceptions with predominantly red forewing. The hindwing is
white with black distal border, which is missing in most specimens of the Sumatran
U. sumatrana with red forewing. The spots of the forewing are in the same
position in pulchella, pulchelloides and lotrix, and although they are variable in
size and number within each species, no differences in coloration have developed
by which one could with certainty separate one of these three specific populations
from the other. Mr. W. H. T. Tams pointed out to me about a year ago that
U. pulchella bears behind the base of the cell a black dash not found as such in
U. lotrix, and that the last two marginal spots are elongate in lotrix and short
and round in pulchella. This combination of differences would be a good guide
if it applied to all the specimens. Unfortunately quite a large proportion of
pulchella is without the basal dash, and the specimens of lotrix from New Caledonia
and Lifu and many examples from other localities possess a black spot instead of
the dash of pulchella (very rarely a dash). This spot, however, is usually placed
on the median vein, not behind it; in pulchella, on the other hand, the dash
extends rarely on to the vein, and if it is abbreviated, its remnant is usually at
some distance from the base. As regards the posterior marginal dots, they are
elongate in nearly every Oriental specimen of lotrix, but in the African lotrix
rarely larger than the dots in front of them and often smaller, while in pulchella
the posterior spots are frequently distinctly enlarged. In spite of the instability
here pointed out, there is something distinctive : the presence of well-elongate
posterior marginal spots and the absence of the basal dash are a combination
found frequently, in certain districts normally, in lotrix, but not in pulchella+
* In Curtis’s figure, Brit. Entom., pl. 169 (1827), the two spots are elongate, less so in the original
drawing (in the Tring library) than in the published copy ; the basal dash is present in both. The
scarlet spots of the original drawing appear to the naked eye dark olive-tawny in the published
plate, being under the lens pale scarlet shaded with purple. The great convenience of the Tamsian
distinguishing characters for a rapid examination of material was demonstrated to me when I
looked at the series of 16 specimens placed as pulchella in the British collection at Tring; one of
them caught my eye on account of the elongate posterior marginal spots: it was U. lotria, not
pulchella. The specimen, a 9, is labelled ‘‘ Dartmouth, Sept. 16th, 1892. F. Jackson.’ The label
has several pinholes, and I doubt that the specimen is the original one. It came to Tring with
the Gibbs collection. The nearest locality of U, lotrix is Senegambia (a subspecies recognizable
in the male only),
Novirates ZooLoGIcAE XLI. 1939. 253
The specimens of U. pulchelloides can be separated, more or less, into four
groups according to the features discussed : (1) basal dash present, posterior
marginal spots small ; (2) basal dash present, posterior marginal spots elongate ;
(3) basal dash absent, posterior marginal spots elongate ; (4) basal dash absent,
posterior marginal spots elongate. The second combination occurs only among
the specimens of U. pulchelloides, of the three species here discussed.
These unstable distinctions do not affect the appearance of the moths at
all; in the specimen at rest or on the pin they are only discovered after a close
scrutiny and are invisible in a flying specimen, which looks white. The persist-
ence of minute somatics which as such have obviously no function are a feature
common to many species. For instance, the posterior end of an oblique discal
band on the forewing, or of a row of spots, often remains present in Arctiids ; spots
which can be seen neither in resting nor in flying specimens are a character of
the same kind: a remnant carried along because it is innocuous in the life of
the individual. The two marginal dots in front of the tornus of the forewing
remaining usually separate on the upperside in these three Utetheisa are an ancestral
relic probably unique in Arctiidae, or at any rate very rare, as I have not seen
another example outside Utetheisa in cursorily searching for one in the Tring
collection. Elsewhere in the family the two have often coalesced into one spot,
which is frequently larger than the one in front of it.
A similarly inconstant distinction between U. pulchella and its allies is met
with on the underside of the forewing. The species similar to pulchella in pattern
(we exclude U. antennata from this discussion, its pattern being different) have a
black discal band which projects towards the apex of the cell. The black
discocellular bar is either connected or amalgamated with this projection or
remains completely separate. The connection consists often of only one or two
lines on the veins, and the bar is not infrequently free on one wing and con-
nected or amalgamated on the other. In the following Table I we unite under
U. pulchella and U. pulchelloides all the subspecies, because these show no essential
differences in the percentages (the actual figures will be given in a later table
when dealing with the subspecies) ; U. lotrix, however, is here divided into four
sections on account of an interesting difference in the percentages. The table
demonstrates several facts in variation of which the significance will be appreci-
ated if we consider that amalgamated elements of pattern represent a later stage
of evolution than separate original elements.
‘TABLE I.
PERCENTAGES OF FOREWINGS IN WHICH THE DISCOCELLULAR BAR OF THE UNDERSIDE IS CONNECTED
OR AMALGAMATED WITH THE DISCAL BAND.
Z a U. pul-
an U. salo- | chelloides, | U.lotrix | U.lotrix | U.lotrix | U. lotrix
al U. amhara, monis, Oriental lepida, lotriz, lotriz, stigmata,
Palaearctic Africa. Solomons, Region Ethiopian | Australia New Solomons,
Regions, etc. to Mada- Region. to India. Guinea. | etc.
India, gascar.
Number of forewings . 2,790 378 56 1,132 146 540 54 46
Percentages 4%, 14% 87% 80%, 0% 2% 44%, 41%
In this particular detail of pattern U. pulchella, U. lotrix lepida and U. 1.
lotrix are primitive.
The low percentage of U. amhara, which represents
pulchelloides in Africa, but has no tuft on the male hindwing, is interesting,
254 Novirarrs ZooLoGIcaE XLI. 1939.
contrasting very strongly with the percentages of U. salomonis and U. pulchel-
loides. Although U. amhara has the same type of genitalia and male antenna
as the two others, the relationship is evidently not very close ; it appears as a
primitive in the table, whereas the two others are advanced. U. salomonis, on
the other hand, which agrees with U. amhara in the absence of the scent-organ on
the male hindwing, does not contrast at all with U. pulchelloides, the various
subspecies of which show a variation from 75 to 100 per cent. ; it is geographic-
ally surrounded by subspecies of U. pulchelloides and fits very well into the
series, apart from the missing tuft of the male hindwing. The relationship with
U. amhara is much more remote.
U. lotrix is divided into four sections in the table ; the African subspecies
again shows the lowest percentage, there being no connection between bar and
band (as is also the case in the Angolan U. pulchella kallima) ; in Indo-Australian
U. 1. lotrix the percentage is only half that of U. pulchella. It is therefore very
surprising that in New Guinea (not in Australia !) the percentage rises to 44 per
cent., though there are no other differences from U. lotrix lotrix. Approximately
the same percentage is found in U. lotrix stigmata from the Solomons and neigh-
bouring groups of islands, but here the rise is accompanied by other colour
differences, the population having attained the status of a subspecies, while the
New Guinea population has not.
An attempt to find in the neuration of Utetheisa some distinctions corroborat-
ing and supplementing the specific and subspecifie differences has incidentally
had the unexpected result that the specimens proved the current diagnosis of
Utetheisa to be only partially correct. The neuration is very variable in all the
species, especially the subcostal veins of the forewing. It is generally stated
that the forewing of Utetheisa has an areole or even two areoles ; only Lord
Rothschild, in Seitz, vol. x, p. 260 (1914), varied the diagnosis in stating that
an areole is frequently present. In order better to understand the variability
and its meaning for systematics, we insert here seven diagrams (text-fig. 226) ;
B to G are based on actual specimens of U. pulchella ; A represents a phase in
the pupal wing in which vein 10 is still free (in this phase 6 to 9 would not yet
have acquired the position they have in the figure ; but we may overlook that
inaccuracy, as we only wish to illustrate the subsequent relation of 10 with 8.9.
We also employ here Herrich-Schäffer’s system of numbering the veins, because
most diagnoses of Utetheisa are written that way). In a later pupal phase 10
bends down, approaching 9 beyond the end of the cell, amalgamating with it for
a short distance or becoming connected with it by a bar, an areole (ar) being
formed (figs. B and E). We can group the specimens of U. pulchella in two
series; in the left series B represents an individual in which the coalescence of
9 and 10 is short ; in C it has gone so far that the areole is much reduced ; in
D the areole has entirely disappeared. We find examples of all intermediate
stages between B, C and D. In the right-hand series fig. E is a specimen with
the bar developed ; fig. F another in which the bar has remained incomplete ;
and in fig. G a third with no trace of the bar, this being a more ancestral phase
than any of the others, and D the most advanced. If one follows Hampson’s
Key to the Genera of Arctiidae in his great work, Lep. Phal., iii, p. 2 (1901), the
characters of B and E would lead to p. 8 of the Key, where Utetheisa is found ;
perhaps C and F might also lead to p. 8; but D would land us at b? on p. 4 and
G at con p. 5, far away from Utetheisa. These are not the only modifications
Noyirates Zoonoarcan XLI. 1939. 255
of the neuration considered in Hampson’s Key and his descriptions of genera as
being of generic value. Vein 11 sometimes branches off from 10, which modifica-
tion leads us to Pachydota and Baritius without areole or to T'yria with areole.
The very long and narrow areole present in a large percentage of U. bella and
ornatrix leads to Rhodogastria, specimens of these American species with a shorter
areole to Utetheisa and those without an areole to Protomolis. The position of
vein 7 also is not so constant as in our figures B to G ; in specimens represented
by B, C, D and E its point of origin is sometimes on a level with that of 10 or more
proximal, and occasionally 7 arises from the posterior side of the areole.
-11
226
Oo No ©
11 10 9
9
7 8
T
11
Spuler, Schmett. Hurop., ii, p. 149 (1906), and Rebel, Berge’s Schmetterlingsb..,
9. Aufl. p. 435 (1910), attribute to U. pulchella a double areole. I have found
an anomaly of this kind only once in over 2,800 forewings of Utetheisa examined,
and in this case the areole was divided by a transverse bar. In none of the
species of Utetheisa have neurational characters been consolidated into constant
specifics. Nevertheless, there is here something of the nature of Tams’s semi-
specific recognition marks of U. lotrix described on p. 225, as is very apparent
in Table II on p. 256.
The vertical divisions of this table are the same as in Table I. The hori-
zontal divisions are grouped into two sections : with areole and without areole.
This grouping is artifieial, but is here adopted because the presence and absence
of the areole are generally considered of very great weight in the classification of
Arctiids (and other moths). The stalk of vein 10 with 8.9 is counted as long if it
is at least half the length of the transverse diameter of the cell (at its widest
point). Vein 10 frequently arises from or from close to the apex of the areole
together with the stalk of 8.9; if the distance of 10 from the tip of the areole is
less than twice the width of the short vein common to 10 and 8.9, vein 10 is
19
256 NOVITATES ZOOLOGICAE XLI. 1939.
counted as arising from the areole ; if in doubt, 10 is counted as short-stalked
with 8.9.
TABLE II.
THE VARIATION OF THE POSITION OF VEIN 10 OF THE FOREWING EXPRESSED IN PERCENTAGES OF
THE WINGS EXAMINED.
U. pul- io, rile
Gea U. sal heiloid V.lotrix | U.lotrix | U.lotrie | U. lotri
Ethiopian . sat 0- Ci ELLOU eS, . OLFIE . ol nay . ree 5 OW
d U. amhara, montis, Oriental lepida, lotrix, lotrix, stigmata,
P tose t Africa. Solomons, Region | Ethiopian | Australia New Solomons,
an NS etc. to Mada- Region. to India. Guinea. etc.
ee 2 gascar. |
Number of forewings |
examined A 5 880 400 54 _ 46
With areole : |
(a) Vein 10 from
areole , 119%, 159% 82% 78%
(b) Vein 10 on short |
stalk with 8.9 . 41% | 19% 50% 20%
(c) Vein 10 on long
stalk with 8.9 . 31% 1% 2% 0%
A | es)
Without areole :
(d) Vein 10 stalked
with 7.8.9 x 11595 10% 0% 0%
(e) Vein 10 direct
from cell . ol 296 4% 11% 2%
The most primitive of the five modifications (a)-(e) embodied in the com-
prehensive survey presented by Table II is (e), then follow (a), (b), (c) and (d),
the most advanced modification being (d) (cf. text-fig. 226, D, on p. 255). In
many wings the areole is reduced to a small groove, (c) being but a little less
advanced stage than (d). It is evident from the percentages that U. pulchella
has a singularly isolated position as regards neuration. The very low percentage
of (a) contrasts very strongly with the high percentages of the other species.
Still more striking is the difference between U. pulchella and the others in the
most advanced modifications (c) and (d), the two together amounting to 46 per
cent. in pulchella as compared with at most 4 per cent. elsewhere. There are
two other points in the table worthy of note : in U. amhara and the New Guinean
population of U. lotrix lotrix the number of wings with modification (e) is com-
paratively high, and in the New Guinean lotrix, moreover, modification (6) is
much rarer than in U. lotrix stigmata from the Solomons (whereas in Table I the
two agree closely).
An interesting point not brought out in the table is the frequency of a
difference in the two forewings ; the left wing may belong to one category and
the right wing to another. That is the reason why Tables I and II are based
on wings and not on specimens. In U. amhara, for instance, 28 per cent. of the
right and left wings are different. Conspicuous asymmetry is known of the
pattern of some Lepidoptera, for instance Uraniidae, but as a rule the differences
between right and left wings are small, and may be attributed to the incidents
of growth, variability prevailing everywhere. But vein 10 arising in one forewing
from the cell and being stalked with 7.8.9 in the other is a difference which the
strict neurationist would have to consider of generic value and cannot be
waved aside as coming under the general rule of slight individual variability.
The reason for vein 10 stopping at stage (a) in one wing and going on to stage (c)
Novirates Zoonoaroamn XLI. 1939. 257
in the other will probably be found in the equilibrium of the development in the
pupa being upset by external factors acting upon the caterpillar when changing
into the pupa or upon the chrysalis at the critical moment in its development.
The pupa lies in a flimsy cocoon on the ground among debris. We cannot
attribute the asymmetry to a faulty action or an unequal distribution of a direct-
ing hormone. That assumption would mean assigning the variability of the
neuration to the unexplained variability of a hormone, which would leave us
in ignorance.
However that may be, the example here demonstrated of the instability of
the neuration within a genus of closely related species teaches the taxonomist
the primary lesson that classification should not be mainly based on an organ
the variability of which has not been previously studied. As the range of varia-
bility can only be ascertained from a sufficiently large material, it follows that
collections consisting of a limited number of specimens are inadequate for obtain-
ing reliable results, for which reason we mention in the tables the number of
wings examined.
When some 20 years ago I looked at Utetheisa with a view to ascertain the
real status of some types in the Tring collection, I was greatly astonished to find,
after the elimination of the males with pectinated and dentated antennae, that
the remaining mass of seemingly unclassifiable material similar to pulchella was
very readily separable into three groups according to the external structure of
the genitalia of both sexes. An impossible task if coloration only were examined
became quite easy with the help of the binocular microscope. Text-figures 227—
234 require no description at this point, the differences staring one in the eye. I
was further struck by observing: (1) that subspecies different in coloration or
even in some structure did not yield any difference or only a slight one in the
genitalia ; and (2) that individual variability, so frequent and conspicuous in
the coloration and neuration, was hardly noticeable in the genitalia. The
genitalia and the coloration vary independently, as I had found out long ago
from the study of the individual and geographical modifications in the genitalia
of Papilio (1896).
With the result of the examination of the three externally so similar
species pulchella, pulchelloides and lotrix before me, I expected the externally
dissimilar species to exhibit even greater differences in the genitalia. The
expectation was fulfilled by U. semara Moore 1859 from Java, Celebes and
Lombock (text-figs. 230, 234); but the remaining Old World species, all from the
Malagassic subregion (elata, cruentata, lactea), tell a very different and surprising
tale: their genitalia are alike. In these species evolution has gone ahead in
the coloration, whereas the genitalia have remained stationary ; just the opposite
from what we find in pulchella, pulchelloides and lotrix. The two Americans,
bella and ornatrix, are likewise different in colour and alike in the genitalia.
The contrast between the Malagassic species on the one hand and the three
species with the pulchella-pattern on the other does not entirely conform with
the relationship of the species. Superficially it looks as if we had here two
natural groups in each of which the species were closely related ; but that is
not so, for pulchella does not belong to the group to which the forewing coloration
would seem to assign it. The genitalia of pulchella female are indistinguishable
from those of the three Malagassic species and the male genitalia show only slight
differences. This is the more unexpected because the males of the Malagassic
258 Novitatrs ZOOLOGICAE XLI. 1939.
Utetheisa differ further in bearing on the hindwing a scent-organ which is absent
from pulchella, and in having slightly different antennae. That pulchella is more
nearly related with the Malagassic elata, cruentata and lactea than with the other
Utetheisa is not really strange, considering that it is an essentially African insect.
What we have said about the coloration in the previous lines applies equally to
both sexes.
The abdominal segments modified for facilitating a firm union of the sexes
NOVITATES ZooLoGIcAE XLI. 1939. 259
form in Utetheisa five different lock-and-key arrangements (galapagensis, no
doubt, presenting a sixth, not yet known), and, though each such arrangement
may be regarded as a single unit from the viewpoint of function, the somatie
contribution of the sexes is different and does not necessarily show the same
amount of variability and variation. In Utetheisa this genital armature is
remarkably constant in the individuals from widely different districts. But
while the female armature remains the same throughout each species, the male
claspers are modified to a slight extent in some subspecies of U. pulchella and
U. lotrix and in some geographical representatives of U. pulchelloides. This
difference of the sexes in the evolution of the lock-and-key arrangement is probably
more apparent than real ; the male organs being much the more complicated of
the two, a difference is more easily noticed. A “law,” however, that the male
is here in advance of the female should in any case not be based on the examina-
tion of dry specimens only. The clasper of the American type is much narrower
than in pulchella, broader than in semara, to which it is similar, and simpler in
outline than in pulchelloides and lotrix. The basi-dorsal process (P!) is much
longer than in any Old World species, as is also the anal tergum ; the dorsal
margin is formed, as in semara, by a strongly chitinized tube, distally somewhat
bent down and projecting beyond the membranous apical margin ; the ventral
half of the innerside is densely covered with hairs and scales. In pulchella we
find the dorsal margin enlarged upwards, and both lotrix and pulchelloides have
a subapical (P?) and an apical process (P®), which differ very conspicuously in
the two species. In the female it is sternum VII which is modified, the general
type remaining the same in all species inclusive of galapagensis. That of the
American 29 is the most primitive type : the median sinus is small and shallow ;
the deeper sinus of the Old World Utetheisa is probably a later acquisition.
U. galapagensis has a somewhat deeper sinus than bella (and ornatrix). It is not
necessary to comment further on sternum VII ; the specific differences obtaining
in that segment will be described in the systematic section of this paper. There
is only one point to which I should like to draw attention : in lotrix there is at
the lateral apical corner of this segment a small smooth groove concealed by the
apical margin of sternum VI and appearing in the inside of the body as a smooth
dome. The groove is not present in any other species and its function is obscure
to me ; it is probably sexual.
Apart from the marginal projections, the male clasper, after having been
denuded, appears from outside as a more or less elongate simple flap. In reality
it is a flat box. There is a large invagination from the outer surface, the mouth
of which is easily overlooked. The central area of the clasper, generally more
strongly chitinized, as indicated in the figures by stippling, is somewhat convex
and forms the outer wall of the pocket. Below this sclerite is a longitudinal
slit, closed in the dry specimens and diagrammatically emphasized in the drawings
by a black line. The slit is an outlet rather than an entrance ; for the cavity
contains a scent-organ and is filled with modified scaling, a large tuft projecting
from the base distad consisting of two different types of scales: a transparent
scale on a long stalk appears white, and a small more or less elongate-elliptical
one is filled with a waxy yellow substance. The scale of the long type (text-
figs. 235-237) differ in some of the species. In the American Utetheisa, for
instance, it is very narrow from the base to three-fifths and then gradually
widens, the scale not being unlike an oar with the tip more or less asymmetrical
260 Noviratrs Zootocicam XLI. 1939.
(text-fig. 235). The stalk and the blade are densely studded with elliptical
tubercles, which appear at the lateral margins as small domes. ‘The scale of
semara is similar, but is much shorter and narrower, and the tubercles are replaced
by thin short ridges which are confined to the blade. In pulchella (text-fig. 236)
the stalk is thinner than in ornatrix and devoid of tubercles or ridges ; the apex
is more abruptly widened, at least one-half broader than in the American
species, and bears short ridges; its outline is variable as indicated by text-
fig.236. The scales are alike (or perhaps only very little different) in pulchelloides
and lotrix, resembling those of pulchella, with
the apex somewhat broader as a rule, more
symmetrical and bearing longer striae, and with
the apical margin more distinctly dentate (text-
fig. 237, U. lotrix). This scent-organ presumably
comes into play when the male is hunting for a
female. Glands in the clasper occur elsewhere
in Arctiidae, for instance in Rhodogastria Hübn.
1819. I have not studied the organ any further ;
that should be done with material adequately
preserved in a fixative liquid. The resemblance
in the clasper and its scent-scales of the Malayan
U. semara to the American species instead of
some Oriental one is surprising, the explanation
possibly being that both claspers are of a more
ancestral type than the claspers of the other
species. The shapes of sternum VII in the
females of semara and bella point the same way.
I know of only one more specific distinction
found in both sexes, at least in one species:
asymmetrical tarsal claws. The pair of sickles,
each with a tooth about the centre, are equal in
size and shape in both sexes of Utetheisa with
the exception of U. pulchella, pulchelloides and
representatives and U. lotrix. The asymmetry
is very slight in pulchella and pulchelloides and
confined to the male, being more noticeable in
the hindtarsus than in the other legs ; in lotrix
it is rather well pronounced in the male and
slight in the female, but also in this sex sufficiently marked for distinguishing
under the microscope the females of pulchella and lotrix by their claws. We
find this asymmetry sporadically in other Lepidoptera, the best-known example
being the Acraeine butterflies, where it is a male character occurring in most
species. In Utetheisa and Acraea the posterior (or inner) sickle is the longer
of the two. The distinction was evidently originally linked with the male sex
only and at a later stage appeared in the female lotrix as a rudiment, comparable
to the glossy coloration of male butterflies being occasionally more or less
indicated in the female. The function is no doubt sexual, but why it is the
posterior half of the claw that is the longer, whereas in Rutelid beetles, for
instance, the anterior sickle is the longer, is a mystery to me. The slight
asymmetry in the female claw of lotrix has probably no function at all; it is a
Noyrratrs ZooLoGIcAE XLI. 1939. 261
phylogenetic budding-on, not occasioning any disturbance and therefore not
eliminated.
The remaining specific distinctions of the Old World Utetheisa are confined
to the males and are found in the antenna, the hindwing and the hindleg.
As in Heterocera with pectinated antenna, the female antenna is frequently
also pectinate or dentate or serrate, the processes being occasionally (in some
Saturnioid Ceratocampidae) longer than in the male, I expected to come across
some dentate female antenna among the material from localities where the
males have fairly long pectinations. All the females, however, I have seen have
simple antennae, with the exception of U. galapagensis, which shows a short,
but noticeable, dentition ; the male of this species should, therefore, be expected
to have longer pectinations than any other Utetheisa, if evolution were bound
to follow the demands of human logic. However, it is better to be an agnostic
when dealing with nature; prophecies are only quite safe if the wording is
sufficiently ambiguous.
The antennae of the males of the two other American species and of pulchella,
lotrix and semara are simple, with the usual long cilia and a long bristle (or two).
In elata the antenna is almost exactly the same as in pulchella, but in cruentata
and lactea the segments are shorter and constricted at the base, the sides projecting
a little, these rudimentary teeth being best developed in lactea. Uniting all the
other males with dentated or pectinated antennae under U. pulchelloides, with
the exception of the African representative amhara, the Nicobar species antennata
and salomonis, we find the lateral projections varying from a short tooth to a
rather long pectination (text-figs. 249-257) and the distribution of these varia-
tions so erratic that it is difficult to decide in the absence of any other structural
distinctions whether long, medium and short projections indicate species or
subspecies. The question is much complicated by the occurrence, in some
localities, of specimens with long and short projections. The only solution of
the puzzle I can suggest is based on the habit of migration observed at least in
. pulchella. This African species ranges from the Mediterranean countries to the
Mascarene Islands and across Asia to China, Ceylon and Burma. It occurs
frequently in Central Europe in favourable years, and is occasionally seen farther
north. Its distribution is nearly the same as that of two Ethiopian Hawkmoths,
Acherontia atropos L. 1758 and Deilephila nerii L. 1758. All three are wanderers.
Swarms of U. pulchella have been seen, and the species has recently managed by
some means or other to reach South America, Miss Lucy I. Clarkson as well as
the Rev. Arthur Miles Moss having caught and bred it at Pernambuco.
It is hardly far-fetched to assume that also some of the other species are
migrants, at least in years favourable for the release of the urge to seek other
quarters. If that is the case with one or the other form of U. pulchelloides, the
occurrence of two of them in the same locality would be explained. That,
however, raises another question. Subspecies which differ in no other structure
but the male antennae might be expected to interbreed and form a hybrid
population. We have from the small island of Guam a short series of males
(4 dd, 2 22) with pectinated antennae, and a larger series (7 gg, 9 29) with
a short dentition instead, and one intermediate 3 from that place, presumably
a hybrid. The dentate form of the male is the only one we have from the
Philippines, Hainan, Indo-China and India ; our series being very large means
that the moth is common in that area and therefore quite likely to travel far
262 Novirates ZooLOGICAE XLI. 1939.
afield and occasionally to reach Guam. Hampson’s pectinata with long pectina-
tions is based on a single specimen from Port Essington, North Australia. All
our Australian males have short teeth instead: but our series from Moa,
Tenimber and Key consists of males with long pectinations, suggesting that the
Port Essington specimen was an immigrant, probably from Tenimber. A few
specimens (Key, Koor) have the antenna subdentate and may be immigrants,
possibly from the Moluccas. Another equally long-pectinated form inhabits the
Nicobars (U. antennata). Males with shorter and broader pectinations are
known to me from the Louisiades (south-east of New Guinea), and the Seychelles
and Chagos. The males from Tuamotu, Tonga, Ellice and the Marshall Islands,
inclusive of Wake Island (and presumably all the more outlying Pacific islands),
have dentate antennae, the teeth being broad and prominent. With the excep-
tion of the localities mentioned, the area from Ceylon to China and eastwards
to Australia and New Guinea has a male with very short and broad teeth which
strongly contrast with the pectinations (compare text-figs. 252-254). This widely
distributed antenna, nearest to the antenna of lactea, was probably the original
one from which the male antennae with longer processes developed, the growth
of the teeth into pectinations having taken place sporadically and independently.
The inconvenient fact that two pulchelloides forms occur together on Guam
and Tenimber has above been explained away by referring it to migration. In
the absence of the observation of migratory habits in pulchelloides—which pre-
supposes that the observer has learnt to distinguish between pulchelloides, lotrix
and pulchella—the explanation is at most plausible, because it is based on the
assumption that pulchelloides possesses the migratory habit known of pulchella.
The plausibility of the explanation may have misled me to accept it as an easy
way out of a difficulty, and I should be wrong if pulchelloides did not migrate.
I emphasize the point, because I wish to draw attention to the importance of a
knowledge of the habits of the species for research in systematics.
I touched above, also in connection with migration, on hybridism between
two subspecies found on Guam, and I think there can be no doubt that hybridiza-
tion occurs also in the subspecies of pulchella. This moth is common in North,
Kast and South Africa, but is more rarely found in the West African forest region,
being an insect of the open country. It is represented in Angola and Katanga
by a conspicuously coloured form described as a distinct species, U. kallima, by
Swinhoe in 1907. Rothschild 1910 and Hampson 1920 concurred in this opinion.
Weymer, on the other hand, described it in 1908 as U.pulchella completa, and I
agree with him that kallima and pulchella are subspecies of one and the same
insect. The amount of scarlet on the forewing and the black border of the
hindwing are not so constant as one might infer from the descriptions of kallima,
and we have some Palaearctic specimens nearly as scarlet as some of the Angolan
ones. Most specimens from the Lake Kivu district, whence we have a good
series, and from the region of Lake Tanganyika, approach U. p. pulchella and
have been described as U. callima dilutior Roths. 1910, some examples being
similar to the least red Angolan individuals, others coloured like certain specimens
of pulchella pulchella from Tanganyika Territory and Kenya. The intergradation
is complete from extreme kallima via dilutior to pulchella. There being no line
of distinction and the area of dilutior being adjacent to the areas of pulchella and
kallima, we have every reason to look upon dilutior as an indefinite hybrid
population.
Novirarrs Zoorocıcan XLI. 1939. 263
“a
The area inhabited by the Angolan U. p. kallima being quite small as com-
pared with the area in which U. p. pulchella is found, one might wonder why
U. pulchella is here subspecifically modified and nowhere else in its enormous range.
An enquiry as to which status kallima occupies in the evolution of U. pulchella
partly clears up the anomaly. As stated on p. 251, the ancestral Utetheisa were
scarlet and black, the creamy-white pigmentation being a later acquirement. This
opinion is supported by the colouring of some crippled pulchella from South
France in the Tring collection : these specimens, bred and not fully developed,
show an amount of red much exceeding the average, approaching kallima closely.
This points to arrested development of the colouring. Therefore I look upon
pulchella kallima as an older form than the whiter pulchella pulchella; as an
Angolan relic in danger of being slowly absorbed at the eastern outskirts of its
range by the migratory pulchella pulchella, which is a Mediterraneo-Ethiopian
modification. A slight difference in the male genitalia of kallima is also found
in some dilutior ; other specimens of dilutior intergrade with pulchella or do
not differ.
In the account of the antennae of the forms of U. pulchelloides (p. 261) we
purposely did not include the males from Africa (U. amhara) and the Solomon
Islands (U. salomonis), because they occupy a special position. The African
antenna is like the Indian one, and that of the Solomon male similar to, but in
dorsal aspect a little wider than, the antenna of pulchelloides from the Seychelles
and Chagos. Both moths, however, differ from the other modifications of
pulchelloides in quite another somatic. 5
All the males of the various subspecies of pulchelloides bear on the hindwing
a tuft of long hair projecting from a submedian fold. This scent-organ is a truly
secondary sexual male character, which has no equivalent in the female. Besides
pulchelloides and antennata the organ is present only in the Malagassic elata,
cruentata and lactea. It is undoubtedly an acquired organ superimposed on the
ancestral Utetheisa and not correlated with the specific or subspecific distinctions
in the genitalia, antennae and coloration, and not of fundamental taxonomic
value. A grouping of the species according to the presence or absence of this
organ is unnatural, as proved by the forms representing pulchelloides in Africa
and on the Solomons, New Hebrides and Lifu. If all the species of Utetheisa
possessed the fold and tuft, I should not hesitate to regard the absence of the
organ in the districts mentioned as due to loss. But was the scent-organ really
acquired by all species and then lost in the majority of them, the male hindwing
having returned to an ancestral state ? Or did only the Malagassic species and
pulchelloides acquire the organ with the exception of the African and Solomon
Islands representatives of pulchelloides, the hindwing of these exceptional males
having retained its original state ? Without further evidence one way or the
other, one can answer such questions with yes or no according to one’s preferences
or prejudices. For, though all roads lead to Rome, all lead also away from it.
In which direction did the evolution of the scent-organ travel? The starting-
point is indicated. The scent-organ is placed in the fold between veins la and 1b,
and the fold is enlarged so as to cover the organ. In all the species without the
organ the membrane between la and 15 is likewise folded, but the fold is simple
and can be entirely flattened out. U. pulchella, U. bella and ornatrix and U.
semara bear in this fold a row of hairs as thin as those of the hairy area extending
from 15 to the cell and base. The “ tuft ”’ of the scent-organ has evidently been
264 Novrrarres Zoonocicarn XLI, 1939.
formed by a concentration and enlargement of these thin hairs of the pulchella-
fold. Therefore pulchella and the other species mentioned as having this rudiment
of the scent-organ must be regarded as more ancestral in respect to this organ
than the species with the tuft fully developed. If that is so, which position take
the representatives of U. pulchelloides which have no scent-organ, but otherwise
agree closely with the tuft-bearing forms of it ? I have examined 70 males from
Africa and 17 from the Solomons, New Hebrides and Lifu, i.e. 174 hindwings
altogether, and have found them all devoid of the primitive row of hairs. This
complete absence suggests that the hindwing of these two forms is not primitive,
but has lost the scent-organ: the two insects are derivations from the tufted
U. pulchelloides.
Moreover, the African representative of pulchelloides is confined to Abyssinia
and Northern Somaliland (as far as we know at present), which indicates an
extension of the Oriental fauna, i.e. an extension of the range of the tufted
pulchelloides, which is equivalent to saying that the tuft has been lost. The
home of U. salomonis, the far-eastern form without tuft, is surrounded by
pulchelloides-forms in which the tuft is in evidence ; salomonis differs somewhat
in the coloration and in the male antenna from its neighbours, but there is no
apparent reason why the male tuft should be absent. One would like to examine
the wings in the pupa for vestiges of the tuft. It is of fairly frequent occurrence
in Lepidoptera that geographically widely separated species or subspecies have
a characteristic in common independently acquired either by accretion or reduc-
tion ; U. amhara (Africa) and U. salomonis (Solomons, etc.) are illustrations of
this phenomenon.
The last species without a scent-organ on the hindwing of the male is
U. lotrix. In the specimens from the Oriental region and the Seychelles there
is likewise no trace of the primitive row of hairs in the fold, which I interpret as
being due to reduction as in the case of the two previous forms. But it is here
not the loss of the tuft, but the loss of the primitive row of hairs ; the ancestor
of U. lotrix had no scent-organ on the hindwing. I arrive at this conclusion
from the fact that in some African specimens the primitive row of hairs in the
fold is present or indicated and that the presence or absence in the Africans
seems to be correlated with the degree of development of another type of organ,
probably also a scent-organ, situated in the hindleg. In all eastern males this
modification of the hindleg is very strongly developed, and the primitive hairs
of the fold are absent. In Africa the row of hairs is lost in the examples which
have the hindleg more modified than the specimens which have the row pre-
served ; this statement, however, requires confirmation, the number of African
males (16) before me not being large enough for an enquiry of this kind. The
number of Oriental males in the Tring collection is very large.
The modification of the hindleg of U. lotrix above referred to is instructive in
more than one way. The leg combines three distinguishing characters confined
to the male. The most conspicuous one of them, which catches the eye at once
under the microscope, is the peculiar scaling of the inner- and underside of the
femur and tibia (text-fig. 255). The scales covering these sides are short, ovate,
arranged like flat tiles on a roof, but in less regular rows, contrasting sharply
with the longer and rougher scaling of the rest of the leg. The tibia is a little
more swollen than in other species ; but the swelling is generally less noticeable
in the African specimens than in the Oriental ones ; both the modified scaling
Noyirarrs ZoonogicaAr XLI. 1939. 265
and the slightly modified tibia depend probably on the degree of development
of a gland. However, I have not found the gland. I do not wish to infer that
the presence of a gland in the tibia of a moth is inevitably accompanied by this
peculiar scaling ; far from it. The tile-scales are a special modification of lotriz,
the meaning of which we do not yet know. We are here on the borderland of
correlation.
It is, I think, generally agreed that there are two kinds of specific distinc-
tions: (1) independent distinctions, which are not necessarily affected when
other characteristics become modified ; and (2) correlated distinctions, which
are so bound up each with an independent one that the correlate exists and
varies only along with its master character.
In a discussion on the Species Problem at the Linnean Society of London I
said that correlation in the different organs was rare.1 According to Professor
W. Robyns’s contribution to the discussion ?: “ There is a certain correlation
between the different characters of a species or group, so that a few main charac-
ters may be recognized, which by itself will bring up some minor characters of
secondary value. That seems to be the result of the use of different criteria.”
I am afraid the word “ correlation” is often used in a somewhat loose sense. A
specific distinction is either correlated with another distinction or it is not; it
is not a matter of opinion, but should be a matter of fact ascertained by the
study of variation. For purely diagnostic work, of course, all constant distinc-
tions are on the same level and for the purpose of facilitating the determination
of a species an author may seize upon any distinctions he considers most suitable ;
but in questions of relationship, i.e. evolution, a minute specific may be of much
greater importance, because it is an ancestral relic, than a conspicuous distinction
acquired at a much later stage in the evolution of the species. It is so easy to
sweep away quantitatively small differences as “‘ minor ” specifics correlated
with “ major ” ones and therefore “ negligible.” The status of a specific must
be known before it can be used in any argument. More than 40 years ago I
came to the conclusion from the study of the genitalia of Papilio that “ the
direction of the variation of the genital armature within a species of Papilio is
entirely independent of the variation of the wings.’ I have found no exception
in other Lepidoptera, and what I said above about the pattern of Utetheisa
pulchella, pulchelloides and lotrix is an affirmation of the statement of 1896.
Professor S. Maulik in a paper lately published 4 made a mistake in regarding the
inconstant distinctions in the pattern of U. pulchella and lotrix (cf. p. 225) as
correlated with the genitalia. Characters correlated occur together; but
characters occurring together are not necessarily correlated. An old barn I saw
here in the Chiltern Hills had a hole in the roof and broken windows, and on the
floor was a small heap of smashed tiles and splintered wood; three features
occurring together. The hole in the roof was caused by beetles having so
weakened some laths that these could not support the weight of the tiles and
snow, and the broken panes were due to boys who, passing that way and seeing
the dilapidated building, had shied stones at the windows, as any normal boy
would. Two features originated at different times and by different forces ;
1 Proc. Linn. Soc. Lond. 1938, p. 241. On p. 244, line 5 from top, ‘“ co-ordination ” is printed
instead of “ correlation.”
2 Ibid., p. 250.
3 Nov. Zool., iii, p. 499 (1896).
4 Ann. Mag. N.H. (11), iii, p. 232 (1939).
266 Noviratrs ZooLoGIcAE XLI. 1939.
the heap of rubble on the floor, however, was correlated with the hole in the
roof, and the solitary unbroken pane was an ancestral character, a relic. All
the features were acquired in a process of decay, an evolution downwards. The
features of a species also are acquired in a process of evolution. Sometimes
correlation is easy to recognize. If in a genus of beetles a species is distinguished
by abnormally convex elytra, the interspaces between the rows of punctures are
wider than on the normal elytra of other species. And in a species with a
tubercle on the elytrum, the interspace bearing the tubercle is widened in that
place according to the size of the tubercle. In both cases there are two distinc-
tions, but the presence of one depends entirely on the presence of the other.
U. lotrix presents us with an interesting illustration of correlation of a less
obvious kind. The swollen tibia and the modified scaling are not the only
peculiarities of the male hindleg. The two pairs of spurs likewise are more or
less different from those of the other species of Utetheisa, being often distinctly
shortened in the Oriental males of lotrix, which all have modified scaling ; while in
Africa, where the extent of the scale-modification varies, the spurs vary from
being normal to being slightly shorter. Swollen hindtibiae carrying a scent-organ
are not infrequent among Geometridae, the swelling being here accompanied by a
reduction in the size of the spurs, which sometimes disappear altogether. This
coincidence cannot be fortuitous: the cause of the swelling is the development
of a gland, and the swelling causes a reduction of the spurs. It appears to be
a matter of growth, the material used for the enlargement of the tibia not being
available for the growth of the spurs to the normal size.
TABLE II.
DISTRIBUTION OF CHARACTERS ALIKE AND UNLIKE IN TEN OLD WoRLD SPECIES.
Coloration. SP, Genitalia. JQ | Antenna. 5 |Tarsal Claw. 32 Hindwing. 3 | Hindleg. &
| |
U. pulchella a a a | a a a
U. elata b (a) (a) 1 a
U. cruentata . | c (a) (a) (a) l a
U. lactea 6 ; d (a) (j) (a) 1 a
U. antennata . | e g j a 1 a
U. pulchelloides j a 1 a
U.amhara . ’ a j a a a
U.salomonis . .\ (a) (g) J a a a
U. lotrix RER a | h a k a m
Gh sumatrana eer: | n h i a | k a m
U. semara 6 - | f i a (a) a a
The distinctions constantly present at least in one sex of the species of
Utetheisa are found in six organs : coloration of wings 39, genitalia SQ, antenna
g or 89, tarsal claw Z or $9, hindwing scent-organ 4, hindleg g. The two
American species bella and ornatria differ from each other in one specific only,
coloration ; the two together are distinguished from all the Old World species
by two specifics, coloration and genitalia. In the Old World series the lowest
species as regards the number of constant specifics peculiar to it alone is U. pul-
chella, which only differs from every other species in a slight modification of
Noviratrs Zootoaican XLI. 1939. 967
the male genitalia, every other distinction being shared with one or the other
species, and U. lotrix is the highest, being different from every other species in
the genitalia (SP), the tarsal claw (SQ) and the hindleg (3). Designating each
of the six distinctions of U. pulchella as “‘a,” those of elata which differ but
slightly as “ (a) ”” and those which differ more strongly as “ b,” and continuing
in the same way with the other species, the preceding table gives an approximately
correct survey as to which distinctions are peculiar to a species and which each
species shares with one or more of the other species.
Hxample.— U. semara differs from U. pulchella strongly in the coloration
and genitalia, and slightly in the tarsal claw, and agrees with it in the antenna,
hindwing (no scent-organ) and hindleg.
Further and more detailed research will show whether the various species
are really like U. pulchella in the organs represented by “a” in the table. I
may have missed minute but constant distinctions.
SYSTEMATICS.
The genitalia offering the most reliable distinctions in all the species of
the genus, we group the species as follows (compare figures on p. 258) :
A. Pulchella Group. Four species. g. Clasper (text-figs. 227, 231)
dorsally convex, no gap dorsally between the two claspers, or a very narrow one,
except basally ; dorsal margin more or less corrugate; basi-dorsal process Pı
concealed in a view from outside except its base ; no separate process P?; apex
(P®) curved downwards.
9. Apical sinus of sternum VII extending forward less than halfway to base
of segment ; apex of lobes rounded ; sides of sternum rather sharply marked,
incurved (text-fig. 231).
(a) 3. Basi-dorsal process P1 of clasper reaching a little beyond the median
joint (n) of inside. Hindwing with fold between veins la and 1b proximally of
middle on upperside, the fold containing a large tuft of yellowish buff stout
and long hairs packed closely together in a row.
1. U. elata, composed of three subspecies p- 268
2. U. cruentata. . : 5 : : : ; . p. 270
3. U. lactea, composed of two subspecies. Ä B ; > Ds Mal
(6) 3. Basi-dorsal process of clasper reaching to median joint (n) of inside
(text-fig. 238). Hindwing without tuft in fold. Tarsal claw slightly asym-
metrical.
4. U. pulchella, consisting of two subspecies ; ; 1
B. Pulchelloides Group. Four species. 6. Clasper (text-figs. 228, 239-
242) with two dorsal processes: basal one (P1) directed backward-upward, its
apex not concealed ; postmedian process (P?) a broad lobe ; apical process
(P®) curved down ; a wide gap between the two claspers below the postmedian
lobe (text-fig. 240). On inside the ventral area of clasper with short blackish
ridges and at and near ventral margin with blackish bristles. Antenna dentate
or pectinate. Tarsal claw slightly asymmetrical,
268 Novirates ZooLoGIcAE XLI. 1939.
9. Groove separating sternum VII (text-fig. 232) from the tergum less
defined than in the pulchella group; median sinus as in that group or
narrower. At least the first ten segments of the antenna about as broad
as long or broader.
(c) 3. Hindwing with scent-organ.
5. U. antennata i : : ; : : : ; . p. 278
6. U. pulchelloides, composed of 8 subspecies : 2 : eps 2i79
(d) 3. Hindwing without scent-organ.
7. U.amhara sp. nov. : ; F ; : ; : . p. 276
8. U. salomonis i : : : : : 5 : 5D. Bad
C. Lotrix Group. One species. Sg. Claspers dorsally not contiguous
(text-figs. 230, 234); with three processes: P! broad, entirely visible from
outside ; P? likewise broad, directed distad, below it a sinus separating it from
P®, which is directed upwards, not downwards as in the previous groups, and is
studded with tawny bristles. The ventral area of inside of clasper densely hairy.
Anal tergum (text-fig. 229, X.t.) strongly widened downward before apical hook.
Hindleg with the tibia more or less swollen and the scaling partially modified
into small rounded scales covering each other like tiles (text-fig. 255). Tarsal
claw asymmetrical. Hindwing without scent-organ.
@. Sinus of sternum VII extended to before middle (text-fig. 233) ; apex of
lobes broad, oblique from sinus outward ; at base of segment close to tergum a
small sharply defined, well-chitinized, round groove concealed under the apical
margin of sternum VI. Tarsal claw less asymmetrical than in male.
9. U. lotrix, composed of four subspecies . : : : . p. 286
10. U. sumatrana . : ; ; : : . ; . p. 289
D. Semara Group. One species. d. Clasper narrow (text-fig. 230), the
two claspers widely apart dorsally ; basal process P! long and broad, tongue-
shaped ; P? apical, rounded; P®? represented by a small projection downward
of P? ; dorsal margin nearly straight in lateral aspect, strongly chitinized, tubular ;
lateral sclerite (wall of cavity) white, feebly chitinized, glossy ; scales in cavity
with long narrow blade. Antenna simple. Tarsal claw symmetrical. Hindwing
without scent-organ. Anal tergum rounded-widened from before middle to
short apical hook.
®. Sternum VII basally very broad, with large lateral groove (text-fig. 234),
the segment swollen at the lateral side of this groove ; medium sinus short.
11. U. semara . ; ; : ; : ; F j . p. 290
Here would follow the Bella Group, containing the two American species
bella and ornatrix, and then galapagensis with the antenna of the female dentate.
1. Utetheisa elata Fabricius 1798.
Bombyx elata Fabricius, Ent. Syst. Suppl., p. 440, no. 222-3 (1798) (‘in India orientale ” falso).
The largest species of the genus. Forewing much more extended black
than in other species, with large costal and submarginal red spots ; the amount
of red and white in basal two-thirds varying individually.and subspecifically ;
Novirares Zootocroarn XLI. 1939. 269
beyond middle a white band, usually large, consisting of a patch in cell, another
below it and as a rule a small dot ; fringe black, rarely with indications of white
dots. Hindwing white, with black distal border, or the white reduced to a
small diffuse patch, with intergradations.
Antenna simple, in male the segments a little more strongly rounded than
in U. pulchella. Tarsal claw symmetrical.
6. Median sclerite of clasper much longer than in U. pulchella; apical
blade of scales in cavity wider than in that species, more like the scales of
U. lotrix and U. pulchelloides.
Neuration of U. elata. No. of wings examined 174. (a) With areole ;
vein 10 from areole in 60 per cent. (6) With areole ; vein 10 on short stalk with
8.9 in 31 per cent. (c) With areole ; vein 10 on a long stalk with 8.9 in one
wing (0-6 per cent.). Without areole ; vein 10 from 7.8.9 in no wing.
Without areole ; vein 10 direct from cell in 9 per cent. Vein 7 more often from
areole and more proximal than 10 than in other species, except lactea.
Hab. Malagassic Subregion and Usambara.
(a) U. elata elata Fabricius 1798.
B.e. Fabr., l.c. -
Uthetheisa (!) venusta Hübner, Zutr. Exot. Schm., iii, p. 29, figs. 521, 522 (1827-31) (“ Martinique ”
falso, probably slip for Mauritius).
Euchelia formosa Boisduval, Faune Madag., p. 85, no. 2 (1833) (Maurice, Bourbon, Madagascar ;
new name to replace venusta Hübn.).
Utetheisa venusta, Saalmüller, Lep. Madag., p. 159, no. 336 (1891) (“ Ind. S. Afr.” falso).
Deiopeia venusta, Hampson, Fauna Brit. Ind., Moths, ii, p. 55, no. 1280 (1894) (partim ; ““Sikhim,
Sumatra” falso).
Utetheisa elata, Hampson, Lep. Phal.,iii, p. 480, no. 2084, text-fig. 215 (1901) (Madagascar, Mauritius,
Johanna); Strand, Ann. Soc. Ent. Belg., liii, p. 357, no. 2 (1909) (Madagascar: Betsileo,
Majunga ; Gr. Comoro; Isle de France); id., in Wagner, Lep. Cat., pars 22, p. 360 (1919) ;
Gaede, in Seitz, Macrolep., xiv, p. 93, pl. 12a (1926) (Madagascar).
Deiopeia diva Mabille, Ann. Soc. Ent. Fr., 1879, p. 305 (Madagascar).
Hind-
Basal third of forewing above red marked with black and white.
wing with broad black distal border.
In the abnormally coloured specimen described by Mabille as a distinct
species, diva (in the Brit. Mus. ex coll. Oberthür), the red marginal and sub-
marginal spots are small and the others are replaced by white slightly shaded
with red, the forewing appears at first sight black patched and marmorated with
white ; hindwing with the black border extending to apex of cell, abdominal
area shaded with black. Mabille lays much stress on the presence of an additional
dot at the apex of the forewing ; this dot, but small, is marked in one of our
specimens (?) from Reunion in which the white borders to the red spots are
rather more conspicuous than is usual.
Hab. Madagascar ; Bourbon ; Mauritius.
In the Rothschild collection from Madagascar: Morondawa ; Mahanoro,
Ambinanindrano ; Lac Aloatra, Imerinandrosa ; Antsihanaka ; Perinet, Forét
d’Anamalazotra ; Baie d’Antongil; “Madagascar”; 15 dd, 30 99.
Réunion, 10 3g, 4 29. Mauritius, 7 gg, 5 99.—Comoro Is.: Anjouan,
6 3d, 5 99.
Specimens from Mauritius and Bourbon have on the underside of the
forewing frequently more scarlet in the centre.
270 Novirates ZOOLOGICAE XLI. 1939.
(6) U. elata fatela subsp. nov.
Utetheisa elata, Fletcher, Trans. Linn. Soc. Lond., xiii, p. 270 (1910) (liter. excepted ; Mahé).
Utetheisa elata ab. diva, l.c. (err. determ.).
©. Forewing, upperside: the red markings near base and on disk much
reduced in size and number ; costal and submarginal red spots about the same
size as in U. e. elata; beyond middle an oblique white band composed of two
patches and a dot, partially and slightly bordered with red. Hindwing
suffused with black (type), nearly in middle a suffused white spot or patch ;
sometimes the hindwing white, but the black border broader than in U. e. elata,
about as broad as in ab. diva.
On underside the hindwing white to apex of cell.
Upperside of abdomen suffused with black.
Hab. Seychelles.
In the Rothschild collection from Mahé, Cascade, 1 2; in the Brit. Mus. a
small series of 99 from the same place ; specimens with the white on hindwing
extended from Barbarons, west side of the island.
A very interesting form, connecting elata with fatua.
(c) U. elata fatua Heyn. 1906.
Deilemera fatua Heyn, Ent. Zeits. Guben, xix, p. 213, text-fig. (1906) (Usambara, type in Berlin Mus.).
Utetheisa fatua, Strand, Ann. Soc. Ent. Belg., liii, p. 356 (1909) (Usambara: Sigi, Derema, Amani,
ix-xi, i, iii), and zbid. ab. devittata and ab. mediomaculata ; id., in Wagner, Lep. Cat., pars 22,
p. 360 (1919).
Utethesia (!) fatua, Hampson, Lep. Phal. Suppl., ii, p. 20845, pl. 68, fig. 15 9 (1920) (Magila ; “ Zanzi-
bar ” falso ?).
Utetheisa fatua, Gaede, in Seitz, Macrolep., xiv, p. 93, pl. 13d (1926).
Forewing above with red costal and submarginal spots as in fatela, no red
on disk, only occasionally some red scaling at the margins of the white band,
usually the margins of this band buff or tawny ; the band variable, being some-
times much reduced.
Hab. Usambara.
In the Rothschild collection from Usambara, 1 3, 4 99.
2. Utetheisa cruentata Butler 1881.
Deiopeia cruentata Butler, Illustr. Het. Brit. Mus., v, p. 38, pl. 86, fig. 10 (1881) (Mauritius ; “ North
India ” falso).
Deiopeia venusta, Hampson (error determ.), Fauna Brit. Ind., Moths, ii, p. 55, no. 1280 (1894).
Utetheisa cruentata, Hampson, Lep. Phal., iii, p. 481, no. 2085 (1901) (Mauritius ; “locality Sikhim
for this species erroneous”); Strand, in Wagner, Lep. Cat., pars 22, p. 359 (1919); Gaede, in
Seitz, Macrolep., xiv, p. 93, pl. 12b (1926).
Forewing above: the black of a dull olivaceous tint, the white markings
of U. elata replaced by dull buff; the black distal marginal area broken up into
spots. Hindwing white, with black distal border which does not quite reach
to near anal angle as in U. elata ; sometimes a small discocellular spot present,
which evidently does not occur in U. elata.
Segments of male antenna more convex at sides than in U. elata, rudimentary
teeth being indicated.
Sternum VII of female with black spot in middle as in U. elata.
Hab. Mauritius.
In the Rothschild collection 1 g, 5 29.
Noviratrs ZOOLOGICAE XLI. 1939. 2a
3. Utetheisa lactea Butler 1884.
Deiopeia lactea Butler, Rept. Zool. Colls. Alert, p. 577 (1884) (Providence).
Utetheisa lactea, Hampson, Lep. Phal., iii, p. 482, no, 2086, p. 50, fig. 6 (1901) (Providence) ; Fletcher,
Trans. Linn. Soc. Lond., xiii, 2, p. 268 (1910) (variability discussed) ; Strand, in Wagner,
Lep. Cat., pars 22, p. 360 (1919) (Fletcher’s article missed and in consequence two new names
introduced) ; Gaede, in Seitz, Macrolep., xiv, p. 93, pl. 125 (1926).
In size like smallish U. pulchella. Forewing above varying from being
similar to that of U. pulchella to being almost devoid of black and red spots ;
black distal marginal line broken up into dots ; fringe entirely white. Black
distal margin of hindwing broad only at apex, very narrow or absent from about
middle ; no discocellular bar.
Segments of median third of male antenna distinctly subdentate, broader
than long in ventral aspect.
Hab. Islands north and south of the 10° S. Lat. between Madagascar and
the Comoros in the south-west and the Amirantes in the north-east.
A most interesting species with a very restricted area. It is a dwarfed
branch of the same stock from which arose U. cruentata and U. elata. Its great
variability is partly individual and partly geographical according to the very
valuable detailed account given of the species by T. Bainbrigge Fletcher, 1.c.,
and it appears to me probable that another expedition to these islands will not
only confirm that the coloration of the species is in flux, but also prove that the
distribution of the colour-varieties is unstable. However, in systematics we can
only go by what we know, and the facts presented by Bainbrigge Fletcher seem
to me to indicate that there are two subspecies, one fairly constant and restricted
to Aldabra, and the other very variable and distributed over a number of islands,
sometimes a particular colour-form being dominant on an island.
(a) U. lactea aldabrensis Fletcher 1910.
U. lactea aldabrensis Fletcher, Trans. Linn. Soc. Lond., xiii, p. 268 (1910) (Aldabra) ; Hampson,
Lep. Phal. Suppl., ii, p. 510, sub no. 2086 (1910) (Aldabra; “ Assumption” not this subspecies
according to Bainbrigge Fletcher).
Utetheisa lactea var. aldabrae Strand, in Wagner, Lep. Cat., pars 22, p. 361 (1919).
Similar in aspect to U. pulchella pulchella ; fringe of forewing white.
Hab. Aldabra.
In the Rothschild collection 1 g, 1 9; several specimens in the Brit. Mus.
(6) U. lactea lactea Butler 1884.
U. 1. Butler, l.c. (Providence).
U. lactea lactea, Fletcher, !.c.; Hampson, Lep. Phal. Suppl., ii, p. 510 (1920).
Bainbrigge Fletcher distinguishes three varieties according to the colour of
the forewing :
(61) Form lactea Butler 1884.
reduced, often nearly entirely absent.
(6?) Form nigrosignata Fletcher, l.c.
pulchella pattern preserved; no scarlet spots.
Scarlet spots absent, black spots much
On all the islands except Aldabra.
The black spots of the usual
Predominant on South
Providence.
(63) Form rubrosignata Fletcher, l.c.; var. assumptionis Strand, l.c.
Black markings reduced, scarlet spots enlarged—-Rather sparingly with form
20
272 NoVITATES ZooLoGicaE XLI. 1939.
lactea on Farquhar and S. Providence; appears quite to replace form lactea on
Gloriosa and Assumption.
Hab. Assumption, Cerf, Farquehar, Gloriosa, S. Province.
In the Rothschild collection f. Jactea, 1 g kindly presented by T. Bainbrigge -
Fletcher ; f. nigrosignata, 1 & from Cerf Is.; f. rubrosignata, 1 g, 1 2 from
Gloriosa and Farquhar.
F. rubrosignata is what Staudinger used to call “ var. et ab.,” the distinctive
character occurring in a small percentage of specimens in one locality and in the
whole population in another district. Such cases require close scrutiny based on
large material.
4. Utetheisa pulchella L. 1758 (text-figs. 226, 227, 231, 238).
Phalaena Tinea pulchella Linnaeus, Syst. Nat., ed. x, p. 534, no. 228 (1758) (in Europa australi,
Mauritania).
The Mauretanian insects received by Linnaeus from Brander came from the
neighbourhood of Alger ; therefore I regard Alger as “ terra typica ” for pulchella,
Linnaeus’s additional locality ‘“ Europa australis ” being too vague.
Upperside of forewing speckled with black and scarlet on a creamy-white
ground ; very variable, sometimes the black, sometimes the red reduced or
increased ; in most specimens a black dash at base behind cell; the last two
black marginal dots rarely more elongate than the dots in front of them.
Hindwing with a black distal border of variable width ; most specimens with
black discocellular spot.
On underside the area of forewing between cell and hindmargin with pale
scarlet markings ; the black discocellular bar of forewing well separated from
the black discal band, only in 4 per cent. connected with it.
g. Antenna simple, at least the distal segments longer than broad, the sides
of the segments more rounded than in female, but not enlarged into a tooth.
Lateral sclerite of clasper not much longer than broad, anteriorly broadly con-
nected with the dorsal chitinized area. Widened apex of long scales in pocket of
clasper usually asymmetrical.
Q. While the ¢ is easy to distinguish from similarly coloured specimens of
U. pulchelloides and its African representative U. amhara, the separation of
the females requires sometimes closer inspection.
Neuration (cf. p. 254, text-fig. 226, p. 255) as variable as in other species of
Utetheisa, but the proportions of the different modifications of the subcostal
system of the forewing deviate remarkably from those found elsewhere in the
genus. Whereas in U. lotrix and U. pulchelloides vein 10 arises from the areole in
over 50 per cent. of the forewings, the figure is 11 per cent. in U. pulchella; and
whereas in the former two species 10 is on a long stalk with 8.9 in 2-4 per cent.
of the wings, the figure in U. pulchella is 46 per cent. See Table II on p. 256.
In a very small number of specimens (2 per cent.) 10 arises direct from the
cell and is usually more proximal than 7.
Although the specimens of U. pulchella from Tropical Africa are not sub-
specifically separable from Palaearctic pulchella, with the exception of U. p.
kallima, the populations are by no means identical, as shown in Table IV, in
which 400 Mediterranean wings taken from the collection at random are com-
pared with 400 wings of Tropical African pulchella and 80 of U. p. kallima (all
we have at Tring from Angola).
Novirarrs Zoonocican XLI. 1939. 273
TABLE IV.
NEURATION OF U. PULCHELLA,
en.) Tropical Aria. U. p. kallima.
Number of wings . o 5 2 : 400 400 80
With areole : j 5 5 NE
(a) Vein 10 from areole c . © 8% 15% 10%
(6) Vein 10 on.a short stalk with 8.9 . ‘ 34%, : 49% 33%
(c) Vein 10 on a long stalk with 8.9 . 32% 26% 51%
Without areole : ER
(d) Vein 10 from 7.8.9 B . 2 23% 9% 5%
(c) Vein 10 direct from cell. . . 3% 1% 1%
The long stalk of category (c) and (d) being mostly due to the reduction of
the areole, the two modifications are separated but by a very short step in the
individual development and may be counted as one. In that case Mediterranean
pulchella with the percentages 8, 34, 55 and 3 agrees very closely with kallima,
whereas Tropical African pulchella differs in the high percentage of (b) and the
low percentage of (c) + (d).
Hab. Southern districts of Central Europe south- and eastward to the
Malagassic Subregion, Burma and China, farther north as a migrant; recent
arrival at Pernambuco, Brazil.
The records of U. pulchella from outside the Palaearctic Region must be
discarded: those from India and Ceylon refer to pulchella or pulchelloides or
lotrix ; those from farther east than Burma refer to pulchelloides, lotrix or
salomonis ; records from Tropical Africa refer to pulchella, amhara or lotrix lepida.
Moore, Lep. Ceyl., ii, p. 104, fig. 5, represents a form of U. pulchelloides (“ antenna
very finely pectinated ”), and Hampson’s text-fig. 28 in Fauna Brit. Ind.,
Moths, ii, p. 55, is evidently the same insect.
Three subspecies :
(a) U. pulchella pulchella L. 1758 (text-figs. 226, 227, 231, 238).
Phalaena Tinea pulchella Linnaeus, l.c.
Noctua pulchra Denis & Schiffermüller, Syst. Verz. Wien, p. 68, no. 9 (1775) (pulchella renamed).
Bombyx pulchella, Fabricius, Syst. Ent., p. 586, no. 102 (1775) (partim).
Lithosia amabilis Trost, Klein. Beytr. Entom., p. 63 (1801).
Lithosia pulchella, Haworth, Lep. Brit., p. 150, no. 11 (1809) (York; “ Georgia ” falso).
Eyprepia pulchra, Ochsenheimer, Schmett. Europ., iii, p. 304 (1810).
Utetheisa pulchra, Hübner, Verz. bek Schm., p. 168, no. 1733 (1819).
Deiopeia pulchra, Curtis, Brit. Ent., iv, pl. 169 (1827).
Huchelia pulchra, Boisduval, Index Meth., p. 39 (1829) (partim).
Deiopeia pulchella var. candida Butler, Trans. Ent. Soc. Lond., p. 361 (1877) (Congo, South Africa
(type), Natal, Cape).
Deiopeia thyter Butler, l.c., p. 361 (1877) (Turkey, Punjab (type)).
Deiopia (!) pulchella, Hampson, Fauna Brit. Ind., ii, p. 55 (1894) (partim ; text-fig. 28 alia spec.).
Utetheisa pulchella, Hampson, Lep. Phal., iii, p. 482, no. 2088 (1901) (partim ; text-fig. 217 alia
spec.) ; Strand, in Wagner, Lep. Cat., pars 22, p. 362 (1919) (partim).
Utetheisa pulchella v. (ab.) pallida Spuler, Schmett. Hurop., ii, p. 143 (1906) (prevalent in S.W.
Germany).
Utetheisa pulchella ab. fasciata Spuler, l.c. (locality not specified).
Utetheisa pulchella ab. semisignata Spuler, l.c. (locality not specified).
Utetheisa pulchella ab, melampyga Spuler, l.c. (Dalmatia,— _Discoloured ?),
bo
74 Novirares ZoonocicaE XLI. 1939
Dejopeja (!) pulchella-bicolor Oberthür, Lép. Comp., v, i, p. 182 (1911) (Biskra).
Deiopeia pulchella-bicolor Oberthür, l.c., p. 335, pl. 82, figs. 764, 765 (1911) (Biskra).
Deiopeia pulchella-arcuata Oberthür, l.c., p. 335, pl. 82, fig. 766 (1911) (Ain Draham),
Deiopeia pulchella ab. flava Bang-Haas, Horae Macrolep., i, p. 58 (1927) (Darjiling ——This species ?
More probably U. lotrix).
The coloration of this Palaearctic-Ethiopian subspecies varies considerably
in the proportional development of black, white and scarlet. Extreme modifica-
tions are more often observed in Europe and North Africa than in Tropical
Africa, which is probably due to
collectors in the former areas rearing
larger numbers of specimens than
collectors in the Tropics. The neu-
ration is likewise very variable, but
its variability does not affect that of
the coloration. For instance, in 13
Mediterranean specimens with little or
no black on the upperside of the
forewing I found nearly all the same
modifications in the neuration as in
13 Mediterranean examples with the
black dots strongly developed. Vein
10 from areole or on a short or a long
stalk with 8.9, more proximal or more
distal than 7; if the areole is absent,
10 arises direct from cell or from
7.8.9, etc. As such distinctions are
generally used for the diagnoses of
genera, while differences in coloration
are as a rule considered at most as
of specific value, we may well ask why
only individual deviations from the
average colouring of U. pulchella have
been emphasized by “names.” It is
tradition, Ithink. Lepidoptera having
been contemplated with the naked
eye, or at most been studied with a
hand-lens till recent times, the results of the old method loom very large in the
Catalogues and keep the method dominant. In research on behaviour and
external adaptation to the environment the picture presented to the naked eye
is certainly of paramount importance ; but evolution may have made the picture
so deceptive as to mislead both the insectivore and the systematist. There is no
a priori reason why colour should be given preference in systematics. Reliability
should be the criterium. Hampson’s action (Lep. Phal. Suppl., ii, pp. 195, 220
(1920)) in placing names of species or subspecies “ described on differences in
genitalia only ” as synonyms is based on a queer misconception on the part of
an author who had such great faith in the constancy of another morphological
character, neuration, that he relied on it implicitly. If the lepidopterist requires
“names ” for colour-forms, he should also require them for neuration-forms
(and the modifications of any other variable organ), the system of naming leading
Novirates ZOoOLOGICAE XLI. 1939. 275
to a diagnosis of this sort: U. p. pulchella ab. areolata quadrifida septima-proxima
bicolor. Each word after “‘ ab.” is a descriptive term and should be treated as a
term, not as a name.
It is interesting to note that the most persistent black markings of the forewing
are the marginal dots and, on the underside, the discal band. The black marginal
band of the hindwing is sometimes broader than usual and occasionally includes
a white spot homologous with the median sinus of the normal border ; black
discocellular spot often absent, but not in examples with the black spots of the
forewing strongly developed. Some specimens have a small diffuse red costal
patch on the underside of the hindwing.
Hab. Palaearctic and Ethiopia Regions, extending eastward to Burma.
In the Rothschild collection some 1,100 specimens from : Palaearctic Region
(with the exception of the northern districts, East Siberia and Japan) ; Ethiopian
Region: Sierra Leone, Gold Coast, Nigeria (various places from Kano to the
Niger Delta), Cameroon, Kassai, Uganda, Kenya, Tanganyika Territory,
Nyasaland, Transvaal, Union of S. Africa, Southern Angola, Comoro Is., Mada-
gascar, Mauritius; Oriental Region (only 15 specimens): Ceylon, Cochin,
Nilgiris, Bangalore, Madras, Kulu, Sikhim, Silhet, Toungoo (Burma), Shan States.
(6) U. pulchella dilutior Rothschild 1910.
Utetheisa callima dilutior Rothschild, Nov. Zool., xvii, p. 182, no. 50 (1910) (Lake Kivu, type ;
“Germ. East Africa”: Urundi, Mpwapwa, Kilimanjaro; Nyasaland: Ft. Johnston) ; id. l.c.,
xviii, pl. 6, fig. 29 (1911) (paratype, ®).
Utetheisa kallima v. dilutior, Strand, in Wagner, Lep. Cat., pars 22, p. 360 (1919).
Utetheisa kallima subsp. dilutior, Hampson, Lep. Phal. Suppl., ii, p. 512 (1920) (Katanga, Lake Kivu).
Probably a hybrid population, intergrading with U. p. pulchella and U. p.
kallima, there being neither geographical nor morphological definite boundaries.
Cf. p. 262. In most specimens there is more scarlet on the forewing above and
below than in the majority of U. p. pulchella. In some the scarlet transverse
bands are complete, there being no white in the cell except the rings of the black
spots. Such examples agree closely with U. p. kallima or are identical with it.
Similar specimens are found among U. pulchella, but very rarely. In the type
specimen the black border of the hindwing broad. Very unsatisfactory as a
subspecies and perhaps better united with U. p. pulchella.
Hab. Districts bordering on Eastern Angola.
In the Rothschild collection a fairly large series from Kissenyi, Lake Kivu ;
Urundi and other inland places of Tanganyika Territory ; Nyasaland.
(c) U. pulchella kallima Swinhoe 1907.
Utethesia (!) kallima Swinhoe, Ann. Mag. N.H. (7), xix, p. 202 (1907) (Angola).
Utetheisa pulchella completa Weymer, Deuts. Ent. Zeits., p. 731 (1908) (Benguela).
Utetheisa callima, Rothschild, Nov. Zool., xviii, pl. 6, fig. 30 (1911); Hamp., Lep. Phal. Suppl., ii,
p. 512 (1920) (Angola, type).
Utetheisa pulchella subsp. completa, Hampson, l.c., p. 512 (1920) (Angola).
Cf. p. 262. The scarlet bands of forewing above and below complete, only
the fourth very narrowly interrupted above by a short creamy streak below
middle ; the narrow interspace between the sixth and seventh bands more or
276 Novirates ZooLocIcAE XLI. 1939.
less completely creamy.—— Black border of hindwing broad, of nearly uniform
width ; fringe completely (or nearly) sooty black ; in one of the specimens from
the Cuanza valley the border as in U. p. pulchella sinuate in middle and more
deeply in submedian fold, the long scales of the fringe white as in dilutior and
pulchella ; in a second example from the same district the lower sinus is repre-
sented by a white spot in the black border ; 13 other specimens from that locality
are normal kallima. Our two examples from Katanga have the border bisinuate
and the fringe pale.
Hab. Angola and Katanga.
In the Rothschild collection 40 specimens from the interior of Angola ;
several places in the Quanza Valley, and Bihé, Bailundu, Cubal R.; 2 from
Katanga, Congo Belge.
5. Utetheisa amhara spec. nov. (text-figs. 239-241).
Probably in collections as U. pulchella.
The African representative of U. pulchelloides. Antenna of & dentate,
genitalia similar to those of U. pulchelloides ; sinus of sternum VII of 2 broader ;
hindwing of & without tuft of hairs.
Most specimens larger than Indo-Malayan U. pulchelloides, but some quite
small. No constant distinction in coloration from U. pulchella and U. pulchelloides.
Black and scarlet spots of upperside generally small ; black basal dash before
hindmargin thin and incomplete and mostly absent ; posterior marginal spots
small in almost every specimen ; the scarlet spots not confluent to form
transverse bands in any of our specimens; on underside no scarlet between
cell and hind-margin (but the spots of the upperside shining through), and the
black discocellular bar nearly always separated from the black discal band,
whereas in Oriental pulchelloides they are rarely completely separated ; cf.
Table I on p. 253. Black margin of hindwing narrower in most specimens
than in U. pulchella ; discocellular spots mostly absent or small, rarely the
two spots connected.
d. Segments of antenna somewhat broader than long, except a few distal
ones, rather strongly constricted at base, triangular in ventral aspect, apical
lateral projection short, but distinct, very little more prominent than in Indo-
Malayan U. pulchelloides with dentate antenna. Clasper (text-fig. 239) as in
U. pulchelloides, process P? somewhat larger ; anal tergum (X.t.) broader before
apical hook. Tarsal claw slightly asymmetrical.
9. Sinus of sternum VII somewhat shorter and broader than in U. pulchella,
much broader than in U. pulchelloides, about twice as long as broad in
middle.
Occurs together with U. p. pulchella and U. lotrix lepida; the male is
easily distinguished from both by the antenna and clasper, and the female
differs much from lepida in sternum VII; but the separation of the females
of pulchella and amhara is not always easy, if the abdomen is much com-
pressed.
Hab. Abyssinia and northern districts of Somaliland ; Kenya (migrant ?).
In the Rothschild collection 170 specimens from Somaliland: Dubar ;
Abyssinia : Harar, Diré Daoua, Kumbi, Hauash R. west of Addis Abeba ; Cheren
in Eritraea ; Sokoke near coast of Kenya (1 @).
Novitarrs ZooLoGIcAE XLI. 1939. DH,
6. Utetheisa salomonis Rothschild 1910 (text-fig. 242).
Utetheisa pulchella ab. 4, Hampson, Lep. Phal., iii, p. 484 (1901) (New Hebrides, Solomons),
Utetheisa pulchelloides salomonis Rothschild, Nov. Zool., xvii, p. 181 (1910) (Solomons: Guadalcanar
and Guizo).
Utetheisa pectinata ruberrima Rothschild, l.c., p. 182, no. 49 (1910) (Loyalty Is.); id., in Seitz,
Macrolep., x, p. 260, pl. 13% (1914) (Friendship Is. falso).
Utetheisa lotrix salomonis Rothschild, in Seitz, Macrolep., x, p. 260, pl. 24g (1914).
Utetheisa lotrix var. salomonis, Strand, in Wagner, Lep. Cat., pars 22, p. 361 (1919) (Solomons,
New Hebrides).
Utetheisa pectinata var. ruherrima, Strand, l.c., p. 362 (1919) (Freundschaftsinseln falso).
Utetheisa pulchelloides from salamonis (!), Hampson, l.c., p. 511 (1920).
Uletheisa pectinata subsp. ruberrima, Hampson, Lep. Phal. Suppl., ii, p. 510 (1920).
Asin U. amhara the & without tuft on hindwing. - Coloration very variable,
but the black of fore- and hindwing much more extended than in any specimen
of U. amhara. On forewing, above, the black dots of one or the other row
united to form a band, in type (from Guadalcanar ; the specimen figured in Seitz
is a paratype) the fifth band exceptionally broad, also the third band ; the other
extreme is represented by some specimens in which the black dots are nearly all
isolated ; the posterior two marginal spots elongate, mostly confluent, and a
basal black dash present in front of hindmargin in all our specimens. Black
border of hindwing broad, sinuate in middle or the posterior portion of the
border connected on the discal side with the anterior portion, the bottom of the
sinus being isolated as a white spot within the black border ; this is the case in
about 25 per cent. of our Solomon Island specimens and in the 7 from the Loyalty
Islands and New Hebrides.
On the underside of the forewing the discocellular black bar is nearly always
united or connected with the discal band.
More variable in pattern of forewing than an equal number of specimens of
any other Old World subspecies collected at random, perhaps with the exception
of U. lactea; the black bands of some specimens very conspicuous. In most
278 Novirares ZooLoGIcAE XLI. 1939.
males there is an interesting anomaly in the antenna : in pectinated or dentated
antennae the projections of the outer side, i.e. those towards the wing, are longer
than those on the inner side. In nearly all males from the Solomons the proximal
segments up to the 7th or Sth are an exception. This anomaly is also not rare
in the subspecies of U. pulchelloides, the direction of development being unstable
(outside influence on the pupa ?).
Our two specimens (92) from New Britain are small and not in perfect
condition as regards colouring ; in one the black spots ofrow 4 are merged together,
forming a regular line which is angulate at the lower angle of cell spots of row 6
likewise confluent ; in the second specimen the spots are nearly all separate.
In both examples the black border of the hindwing with deep median sinus. The
specimens fit fairly well into the Solomon Islands series.
3. Antenna strongly dentate, the projections shorter and broader than in
U. pulchelloides pectinata, much longer than in U. pulchelloides papuana and but
little larger than in U. p. marshallorum. Claw of tarsus slightly asymmetrical.
Process P? of clasper broad and the sinus at its distal side almost effaced (text-fig.
242), therefore P? as such posteriorly not defined. Hindwing without fold and tuft.
9. Apparently no structural difference from U. pulchelloides 99.
Hab. Loyalty Is., New Hebrides, Solomon Is. and New Britain.
In the Rothschild collection from Loyalty Is.: Lifu, 1 g, Maré, 1 4;
Solomon Is.: Guadalcanar, 12 $3, 4 99, Guizo, 3 gg, 1 9; New Britain:
Telesea, 2 99.
In the B.M. 3 3g, 2 22 from the New Hebrides.
7. Utetheisa antennata Swinhoe 1893.
Deiopeia antennata Swinhoe, Ann. Mag. N.H. (6), xii, p. 215 (1893) (Camorta, Nicobars, 3 33,1 9).
Utetheisa antennata, Hampson, Lep. Phal., iii, p. 482, no. 2087, text-fig. 216 3 (1901); id., Ann.
Mag. N.H. (7), xix, p. 239, sub no. 2086a (1907) ; Roths., in Seitz, Macrolep., x, p. 260, pl. 13k
(1914) ; Strand, in Wagner, Lep. Cat., pars 22, p. 358 (1919).
Placed by Hampson, l.c., into his section II without tuft and fold on the
hindwing of the male ; he probably examined at that time only the type specimen,
in which the tuft is concealed in the fold, as it often also is in U. pulchelloides ;
he corrected the mistake in 1907, l.c.
Structurally not different from the subspecies of U. pulchelloides which have
pectinated antennae, but much modified in coloration. Counting the black dots
near base as first row, the sixth row of forewing absent, the submarginal area
occupied by a broadish pale scarlet band, and the fourth row strongly angulate at
vein 6.——Black marginal border of hindwing reduced to a line from before
middle, not reaching anal angle and devoid of a distinct enlargement behind
lower median vein.
On underside the forewing devoid of the black postdiscal band, which is
always present in the U. pulchelloides.
Hab. Nicobars : Camorta.
Not represented in the Rothschild collection ; but in the B.M. 2 33, 1 9
(inclusive of type g and paratype 9).
Table V gives a survey of the neuration of the various subspecies of
U. pulchelloides. For comparison we have added the percentages of the two allied
species U. amhara and U. salomonis, which were already given in Table II.
Novirates Zoonoaican XLI. 1939. 279
U. antennata is left out, because only 3 specimens are known, of which but one
is before me. The number of wings of several subspecies is much too small
to give fairly reliable results. The subspecies are arranged according to the
length of the dentition of the male antennae, as far as possible. It will be noticed
that the two subspecies wmata and pectinata with the longest dentition do not
agree in the percentages of the various modifications of the subcostal venation,
whereas pectinata agrees very closely with its geographical neighbour, the
Australian population of vaga with very short teeth.
TABLE V.
NEURATION OF U. PULCHELLA AND ALLIED SPECIES WITH DENTATE MALE ANTENNAE.
U. p.
U. U.p. Ichel U
lo- Gate || WM: Be U. p. U. p. U.p. U. p.
U. ar U.p. P loides, |aphanis, | marshal- oe) = =
amhara, a umala, 1s oe. Chagos, | Louisi- | lorwm, darum, re hee y Tae!
Africa. 00 | Guam. | ,°:,° Sey- ades, | Pacific | “ns Kn Daun a
mons, Arafura chelles, An Ts. Is. Guinea. lia. Malaya.
ete. Sea. Gio,
Number of wings 378 56 12 80 38 20 62 24 80 34 400
With areole :
Vein 10 from
areole . . | 56% | 75% | 25% | 76% | 26% | 50% | 56% | 79% | 52% | 77% | 51%
Vein 10 on a
short stalk
with 8.9 . | 29% | 21% | 67% | 17% | 50% | 50% | 26% | 17% | 40% | 23% | 40%
Vein 10 on a |
longstalk with
8.9 : 5 1% 2% 8% 0% 5% 0% 3% 0% 3% 0% 4%
Without areole :
Vein 10 stalked
with 7.8.9 . 2% 0% 0% 2% 8% 0% 0% 0% 10% 0% 208
Vein 10 direct
from cell o |) 1294 2% 0% BO || 119% 0% | 15% 4% 4% 0% 3%
8. Utetheisa pulchelloides Hampson 1907 (text-figs. 228, 232, 244-254).
Utethersa pulchella, Hampson, Lep. Phal., iii, p. 483, no. 2088, text-fig. 217 (1901) (partim).
Utetheisa pulchelloides Hampson, Ann. Mag. N.H. (7), xix, p. 239 (1907) (partim ; typefrom Chagos
Is.: Peros Banhos); id., Lep. Phal. Suppl., ii. 510 (1920) (partim).
Utetheisa lotrix, Rothschild (err. determ.), in Seitz, Macrolep., x, p. 260 (1914) (partim) ; Strand, in
Wagner, Lep. Cat., pars 22, p. 361 (1919) (partim).
We unite under this name all the forms with dentate or pectinate male
antennae, with the exception of U. antennata, which is distinguished by the
reduction of the black markings, and U. salomonis and U. amhara differing
chiefly in the loss of the scent-organ of the male hindwing. These two exceptional
populations are likewise modifications of the pulchelloides-type, but are here
treated as distinct species for the sake of simplifying the rather involved syste-
matics of this group of Utetheisa and because there is an easily recognized definite
gap between them and pulchelloides.
Base of forewing with a dash behind la, with very rare exceptions, and
posterior two marginal spots elongate in a very large percentage of specimens.
On the underside the black discocellular bar of the forewing nearly always
connected or more or less completely united with the black discal band ; see
p. 253.
Most specimens small, but many fully as large as U. pulchella: length of
forewing 12-21 mm.
280 NOVITATES ZOOLOGICAE XLI. 1939.
For genitalia, see p. 267. External median sclerite of clasper elongate ; the
apex of the scales inside the pocket broad, more symmetrical than in U. pulchella.
The sinus of sternum VII of 2 short (text-fig. 232, on p. 258), varying in width.
Hab. The species occupies the whole of the Oriental Region (perhaps with
the exception of the southern districts of Austalia ?) and extends westwards to
Madagascar. Some of its subspecies are well defined and may already have
attained the status of species ; but I keep them together in order to emphasize
their geographical and morphological connection. As explained on p. 262, their
distribution is puzzling.
The chief distinctions of the subspecies are found in the male antennae,
which are pectinate, subpectinate, dentate or subdentate ; the subdentate
antenna is the most widely distributed one, the others being more localized.
The figures represent segments 25-29. The teeth of segments 18-25 are
somewhat longer as a rule.
(a) U. pulchelloides pectinata Hampson 1907 (text-figs. 244-247).
Uletheisa pectinata Hampson, Ann. Mag. N.H. (7), xix, p. 240 (1907) (Port Essington); Roths.
in Seitz, Macrolep., x, p. 260, pl. 13% (1914) (partim ; ruberrima not this subsp.) ; Strand, in
Wagner, Lep. Cat., pars 22, p. 362 (1919) (partim); Hamps., Lep. Phal. Suppl., ii, p. 509,
no. 2084a, pl. 68, fig. 17 (1920).
Scarlet dots along anterior side of subcostal vein of forewing above short,
the third rarely longer than broad. On underside the postmedian projection
of the black border separated from margin in cellule 2 by the white scaling
penetrating backwards to vein 2; the black discocellular bar of forewing nearly
always completely united with the discal band ; in many specimens there are
one or two scarlet spots at the costal margin of the hindwing.
Noviratrs ZooLoGIcAE XLI. 1939. 281
&. Antenna pectinate (text-figs. 244-247), the ciliate ventral area of the
median segments a narrow ridge, which strongly slants upwards proximally ;
width from tip to tip of projections more than twice the length of segment.
Q. Sinus of sternum VII rather wide, at most one-third deeper than apically
broad ; apex of lobes more or less truncate, wider than the sinus is deep.
Hab. Islands of the Arafura Sea.
In the Rothschild collection from Moa, Dec. 1902, 5 gg, 3 92; Dammer,
Nov. 1898, 1 9; Tiandoe and Tam, July 1898, 3 gg, 2 92; Tenimber: Selaru,
March-April 1897, 14 gg, 5 99; Little Kei, May, Jan.-March, 2 33, 5 99.
(6) U. pulchelloides umata subsp. nov.
Like U. p. pectinata ; red spots of forewing above somewhat larger, especially
those in front of vein 1b, the third from the base being much longer than broad
and extending across Ib. The postmedian section of the border of hindwing
broader, the sinus in front of it smaller, in most wings separated from the white
disk by a bar connecting the two sections of the border, or almost entirely sup-
pressed. On underside the white sinus of the border likewise small and there
is only a trace of a backward extension or none at all (5 examples), or a thin
whitish line shaded with black connects the sinus with vein 2; no minute scarlet
spots at costal margin of hindwing.
&. Pectinations of antenna a trifle longer than in U. p. pectinata. The
dorso-apical margin of clasper from apex of process P? to apex of P® about one-
sixth shorter than in pectinata.
9. Sinus of sternum VII narrow, narrowest at apex, lobes of sternum also
narrower than in pectinata, rounded from near sinus outwards.
Hab. Mariana Is. : Guam.
In the Rothschild collection from Guam, Oct. 1894, April 1895, 4 3g, 2 99
(type 3).
(c) U. pulchelloides aphanis subsp. nov.
In coloration like U. p. pectinata, but rarely with a red costal dot on under-
side of hindwing. Genitalia also the same. Differs in the male antenna being
dentate : middle segments twice or nearly twice as broad as long.
Of some taxonomic importance on account of its geographical position, its
nearest neighbours having subdentate male antennae.
Hab. Islands off south-east New Guinea.
In the Rothschild collection from the Louisiade Archipelago : Sudest Is.,
Jan.-Feb. 1916, 2 33, 5 29 (type 3); Trobriand Is.: Kiriwini, March-May
1895, 1 & (another $ in B.M. ex coll. Oberthür) ; Yanarba: Egum Is., Feb.
1895, 1 d.
(d) U. pulchelloides pulchelloides Hampson 1907 (text-figs. 248, 249).
Utetheisa pulchella Hampson, Lep. Phal., iii, p. 483, no. 2088 (1901) (partim).
Utetheisa pulchelloides Hampson, Ann. Mag. N.H. (7), xix, p. 239, no. 2086a (1907) (partim ; type
from Chagos Is.); Fletcher, Trans. Linn. Soc. Lond. (2), xiii, 2, p. 268, no. 2 (1910) (partim :
Chagos, Cargados Carajos, Coetivy, Poivre, d’Arros, Desroches, Eagle, Seychelles) ; Hamps.,
Lep. Phal. Suppl., ii, p. 510, no. 2086a, pl. 68, fig. 18 (1920) (partim).
Utetheisa lotrix, Rothschild (error determ.), in Seitz, Macrolep., x, p. 260 (1914) (partim) ; Strand,
in Wagner, Lep. Cat., pars 22, p. 361 (1919) (partim).
Like U. pulch. aphanis, but the teeth of the male antenna shorter, in the
median segments the distance from tip to tip of the projections only one-half
282 NOVITATES ZOOLOGICAE XLI. 1939.
longer than a segment (text-fig. 248) ; apical margin of clasper from lobe P? a
little shorter than in aphanis and in about half the specimens corrugated. Sinus
of sternum VII of female somewhat asymmetrical, a little less than twice as long
as broad, lobes broad, subtruncate, outer angle more strongly rounded than inner
one.
The percentage of wings in which vein 11 arises from the areole near its
base is much higher (26 per cent.) than in other subspecies of U. pulchelloides.
On the Amirantes and Cargados Carajos the majority of specimens belong
to the xanthochrome form, which is very rare elsewhere in P. pulchelloides.
Hab. Chagos Is. ; Seychelles ; Amirantes and Cargados Carajos Is.
In the Rothschild collection from Seychelles : Mahé, 3 dd.
Other specimens examined from various islands in coll. T. Bainbrigge
Fletcher ! and the British Museum. The teeth of the male antenna are a trifle
longer in the Seychelles than in the other localities, segments 18-25 being almost
subpectinate on the outer side.
(e). U. pulchelloides marshallorum Rothschild 1910 (text-figs. 250, 251).
Deiopia pulchella, Hampson, in Blanford, Fauna Brit. Ind., Moths, ii, p. 55, no. 1279 (1894) (partim).
Utetheisa pulchella, Hampson, Lep. Phal., iii, p. 483, no. 2088 (1901) (partim).
Utetheisa pulchelloides marshallorum Rothschild, Nov. Zool., xvii, p. 182, no. 52 (1910) (Marshall Is.).
Utetheisa lotrix marshallarum (!) Rothschild, in Seitz, Macrolep., x, p. 260, pl. 24g (1914).
Utetheisa lotrix var. marshallorum, Strand, in Wagner, Lep. Cat., pars 22, p. 361 (1919).
Utetheisa marshallorum, Hampson, Lep. Phal. Suppl., ii, p. 510, sub no. 2086a (1920) (quotation
as species erroneous).
Utetheisa pulchelloides form marshallorum, Hampson, l.c., p. 511 (1920).
Scarlet markings well developed, taken altogether larger than in U. p.
pulchelloides ; in front of vein 1b a row of 5 spots, of which spots 4 and 5 are
united in most specimens, spot 4 extends forward across submedian fold, there
being 2, sometimes 3, scarlet spots in front of this fold, the first of them placed
behind the cell; the submarginal spots often forming an uninterrupted band ;
the black dots of the postdiscal row small as a rule, rarely several of them united ;
posterior two marginal spots little larger than the dots preceding them, sometimes
even smaller. Hindwing as in U. p. pulchelloides ; sometimes a white trans-
verse linear spot in the apical portion of the marginal band (in type and two
other examples from the Marshall Is.) ; in one of the Henderson I. specimens the
distal margin a little more distinctly incurved below apex than in most specimens
of the various preceding subspecies.
On underside 8 of our 9 specimens from the Marshalls have a scarlet spot
at the costal margin of hindwing, in 4 of them there is in addition an ill-defined
scarlet streak in front of vein 5.
3g. Antenna dentate, the shaft broader than in U. p. pulchelloides, the teeth
shorter, the ciliate non-compressed portion of the underside somewhat larger.
Apical margin of clasper from process P? more or less corrugated.
Q. Sinus of sternum VII narrow, directed distinctly towards the right side
as in the two subspecies following.
Vein 11 from costal side of areole or from basal point of it in 4 wings. The
frequency of 10 arising free from the cell is noteworthy (Table V).
Hab. Marshall Is. We place here also the specimens from Gilbert, Ellice,
Tonga groups and the islands farther east. We have not seen specimens from
1 T am much indebted to Mr. T. Bainbrigge Fletcher for the loan of specimens.
Noviratres Zoonocrcan XLI. 1939. 283
Fiji and Samoa. The examples available, with the exception of those from the
Marshalls, are mostly rather worn.
In the Rothschild collection from: Marshall Is. 6 gg, 3 92; Wake Is.,
Feb. 1903, 2 gg, 19; Tonga Is.: Hapai, Jan. 1911, 12 gg, 1 2; Tuamatu :
Makemo, Takapolo, Hikuero, Anaa, Aug. 1925, 6 34, 2 99; Society Is.:
Moorea, 1 3.
Additional localities of B.M. specimens : Union Is., Henderson Is., Friendly
Is. Occurs probably on all the islands east of Fiji, most likely as a migrant.
(f) U. pulchelloides darwini subsp. nov.
Diopaea sp. Darwin, Journ. Voyage Beagle, 2nd edit., p. 485 footnote (1890) (Keeling Is.).
Utetheisa pulchella ab. 5, Hampson, Lep. Phal., iii, p. 484 (1901) (partim ; Keeling Is.). ,
Utethersa pulchelloides Hampson, Ann. Mag. N.H. (7), xix, p. 239 (1907) (partim) ; Fletcher, Trans.
Linn. Soc. Lond., xiii, 2, p. 268, no. 2 (1910) (partim) ; Hamps., l.c. Suppl., ii, p. 510, no. 2086
(1920) (partim).
Utetheisa lotrix ab. papuana Strand, in Wagner, Lep. Cat., pars 22, p. 365 (1919) (partim).
Most specimens large. - Forewing narrow, scarlet spots reduced and, as a rule,
many of them replaced partly or entirely by olive, sometimes only the four
proximal costal spots remaining crimson ; the subspecies contrasting with U. p.
marshallorum, which it resembles in size and shape ; black spots of fringe sharply
marked, generally nearly as black as the marginal spots below as well as above.
Sinus of marginal band of hindwing nearly always narrow.
&. Antenna dentate as in U. p. marshallorum, the convex ventral ciliate
area a little more rounded.
©. Sinus of sternum VII narrow, as in marshallorum.
Length of forewing 17-20 mm.
Hab. Keeling Is. (= Cocos Is.).
In the Rothschild collection 5 &&, 5 99, type d. In the BM. 2 3d
collected by Charles Darwin, and other specimens.
(g) U. pulchelloides papuana Strand 1919.
Utetheisa pulchella ab. 5, Hampson, Lep. Phal., iii, p. 484 (1901) (partim).
Utetheisa pulchelloides Hampson, Ann. Mag. N.H. (7), xix, p. 239 (1907) (partim).
Utetheisa pulchella ab. papuana Strand, in Wagner, Lep. Cat., pars 22, p. 365 (1919) (name for
‘ Hampson’s ab. 5, which is a mixture).
Utetheisa dorsifusa Prout, Ann. Mag. N.H.(9), v, p. 286, no. 1 (1920) (New Guinea: Arfak).
I accept the designation papuana for this subspecies and select Arfak as
terra typica and as the type of the name one of the two 3 collected by Pratt
and recorded by Hampson. [If it should finally be decided that names given to
“aberrations ” have no claim to priority, dorsifusa would take the place of
papuana.
The specimens from Arfak and the mountains back of Huon Gulf are dark ;
spots on head and thorax large, abdomen with an admixture of olive scaling.
Black dots of forewing more or less enlarged and showing a tendency to become
elongate, between them generally some black smudges ; the olivaceous borders
of the scarlet spots on the whole broader than usually. Sinus in terminal
border of hindwing smaller on upperside, large below ; on fold of male scent-
organ a more or less distinct diffuse olivaceous streak.
At lower altitudes in mandated territory and at low and high altitudes in
284 Novirarrs ZooLocicar XLI. 1939.
former British New Guinea the body and forewing are much less smudged with
olive, the black dots are smaller on the whole and the scent-fold of the male
has no olive streak. The extremes are connected by intergradations ; there is
also no sharp distinction from the next subspecies ; but the series of the smaller-
spotted Papuans contrasts nevertheless with the series of Australians in the
latter having on the whole less conspicuous black dots.
g. Antenna subdentate (as in text-fig. 252), the segments smaller and more
rounded than in any of the preceding subspecies, the teeth very short and obtuse
and barely visible in a dorsal view of the antenna. Apical margin of clasper
corrugate.
Q. Sinus of sternum VII as in darwini narrow and oblique.
Neuration : in one wing 7 stalked with 8, and 9 with 10, no areole; in
another wing 11 vestigial, being thin and short; in 5 wings 11 from base of
areole.
Hab. New Guinea, main island.
In the Rothschild collection 20 $3, 29 99 from Dutch N.G.: Arfak Mts. ;
Mandated Territory: Astrolabe Bay, Krätke and Cromwell Mts. (Eastern
Finisterre Range), Waria R.; “ British N.G.” : Hydrographer Mts., Biagi on
the Mambaré R., Aroa R.
(h) U. pulchelloides vaga subsp. nov. (text-figs. 228, 232, 252-254).
Deiopeia pulchella Auct. vetust. partim.
Deiopia pulchella, Hampson, in Blanford, Fauna Brit. Ind., Moths, ii, p. 55, no. 1279, text-fig. 28
(1894) (partim).
Utetheisa pulchella, Hampson, Lep. Phal., iii, p. 483, no. 2088, text-fig. 217 (1901) (partim).
Utetheisa pulchelloides Hampson, Ann. Mag. N.H. (7), xix, p. 239, no. 2086a (1907) (partim).
Utetheisa lotrix, Rothschild, in Seitz, Macrolep., x, p. 260 (1914) (partim) ; Strand, in Wagner, Lep.
Cat., pars 22, p. 361 (1919) (partim).
Agrees in structure with U. p. papuana, and there is also no definite line of
separation in coloration ; but the darkening of body and forewing characteristic
of typical papuana does not occur. It is possible that the dark specimens of
papuana are the true New Guinean subspecies and that the non-darkened
specimens from the eastern half of New Guinea are the result of an influx of
small-spotted Australians. Is this moth a migrant in Australia ?
Though individually variable in coloration, and to some extent also in
structure, there is no indication of geographical variation, all the specimens with
subdentate male antennae, found from Australia to Madagascar and northward
to Japan being vaga. In several localities vaga occurs together with subspecies
which have pectinated male antennae. I have explained this fact on p. 261
by assuming that vaga is migratory.
As in papuana the six distal segments of the male antenna longer than
broad ; in the female the segments of the distal two-thirds longer than broad,
but occasionally the middle segments shorter than usual and only the distal ones
longer than broad. One of the 8 males from Guam has the antenna conspicuously
dentate and the forewing nearly as brightly coloured as U. p. wmata from the
same small island. I consider the specimen to be a hybrid.
Most specimens are small, average of our 23 examples from Java 15-5 mm.
forewing length ; the longest wing in this series 17 mm., in our Manila series
19 mm. ; the shortest wing, 11 mm., in our Hainan series,
Novirares ZooLoaicar XLI. 1939. 285
Neuration essentially as in papuana ; cf. table V on p. 279. Vein 10 direct
from the cell in 12 of 400 wings examined.
Hab. Madagascar (migrant), Ceylon northward to Japan, eastward to the
Moluccas, Kei and Australia; New Zealand (migrant probably). Not yet
obtained on Keeling Is., but found on Tenimber and some other islands of the
Arafura Sea where another subspecies occurs (U. pulch. pectinata).
I
:
Tn
In the Rothschild collection from New Zealand, 1 9 (ex coll. Felder) ;
Australia, 10 gg, 8 22: S. Australia, N.S. Wales, Queensland, West Australia ;
Dutch Timor: Oinanisa, 2 99; Portuguese Timor: Dili and Suai, 2 gd, 5 99;
Adonara, 1 29; Sumba, 1 g, 7 99; Koor, 1 9; Kei Toeal, 1 3 (together with
U. p. pectinata) ; Ceram Laut, 1 &; Batjan, 3 99; Sula Is.: Mangoli, 1 9;
Toekan Bessi Is.: Binongka, 2 99; Buton, 1 g; Central Celebes: Paloe and
Kalewara, 5 3g, 3 99; Saleyer, 3 99; Kalao, 1 ¢g, 1 9; Djampea, 2 99;
Philippine Is.: Panay, 4 gg, 6 29, Luzon, 21 gg, 47 99; Guam, 7 3d, 9 29,
1 g hybrid ; Palawan, 1 ¢; Sulu Archipelago: Bongao, 1 9; British N. Borneo:
Kina Balu, Mt. Marapok, Tenom, 5 dd, 4 99; Sumbawa: Tambora, 1 4;
Java, 9 33, 14 99, type of vaga a g from Semplak, W. Java; (none from
Sumatra); Malay Peninsula: Mt. Tahan, 29 34, 20 99, Penang, 1 g, 4 99;
Saigon, 1 g; Ceylon, 2 gg; Bombay, 1 &; Hainan, various places, 37 3,
39 22; Hongkong: Kowloon, 1 g; Formosa, various places, 18 gg, 41 99;
“ Japan,” 2 dd, 2 99.
Some additional localities in the B.M. collection: Nicobar Is.: Camorta,
1 8; Madagascar (ex coll. Ch. Oberthür), 1 ¢.
286 Novirares Zoonocicar XLI. 1939.
9. Utetheisa lotrix Cramer 1777 (text-figs. 229, 234, 237, 243, 255).
Phalaena Tinea lotrix Cramer, Pap. Exot., ii, pp. 20 and 149, pl. 109, figs. E, F (1777) (partim;
Coromandel, Tranquebar, nec Cyprus).
Utetheisa pulchella auctorum.
Though both sexes are easily distinguished from U. pulchella and U. pulchel-
loides, the species has only lately been recognized as distinct. When Cramer
figured this species in 1777, he was not aware of Linnaeus’s description of
Ph. T. pulchella 1758 ; but at the end of vol. II, on p. 151, we find under “ Addi-
tions ” the reference: “Pl. CIX, E. F. La Phal. Tinea Pulchella. Linn. Syst.
Nat. pag. 884, n. 349. On la trouve pareillement en Hongrie et dans la Russie.”
Cramer identified his lotrix with pulchella. This identification, perhaps unfortu-
nately, was erroneous ; the figure represents an Oriental specimen of the species
which has been correctly identified as lotrix in Mr. 8. Maulik’s paper in Ann.
Mag. N.H. (11), iii, p. 232, text-fig. x, pls. 8, 9 (1939) ; the original drawings in
the British Museum (Nat. Hist.) leave no doubt. The correct identification is
due to W. H. T. Tams.
Antenna simple in $9. Tarsal claw asymmetrical, more so in ¢ than in 9.
Coloration as in U. pulchella, there being no difference applying to all the speci-
mens ; but the majority distinguished by the last two terminal spots of the
forewing being elongate and, in addition, by the black dash at base behind sub-
median vein being absent. It is perhaps worthy of note that one finds in most
specimens a black dot on the extreme base of the median vein and another a
little farther from base on the submedian, the second dot being often shifted to
the posterior side of the vein and sometimes prolonged into a short dash. The
posterior terminal dots are short in most African specimens.
For differences in the genitalia cf. p. 268. Long scales in pocket of clasper
(text-fig. 237) broader at apex, more symmetrical and more distinctly dentate
than in U. pulchella. In order to see the tawny bristles of the apical turned-up
lobe of the clasper, it is sufficient to turn the long hair-scales aside with a brush.
The anal tergum dorsally carinate, being dorsally more strongly compressed than
ventrally ; the downward widening distally rounded, separated from the apical,
dorsal, hook by a sinus, and this widened portion ventrally broader than in the
other species of the genus. Sternum VII of 2 more or less extended concave
around the bottom of the sinus. Hindwing of 4 without scent-organ ; hindleg
of $ with the modified scaling (text-fig. 255) less extended in African specimens
than in Oriental ones.
Hab. West Africa eastward across Tropical and Subtropical Asia to New
Caledonia and the Loyalty Is.
The variability of the neuration is given in Table II on p. 256.
(a) U. lotrix lepida Rambur 1866 (text-fig. 243).
Deiopeia lepida Rambur, Cat. Lep. Andal., p. 225 footnote (1866) (Bourbon “ je crois ””).
The only point of importance in Rambur’s description of lepida is the refer-
ence to the genitalia: “ Les pinces génitales sont bilobées a l’extremite avec le
lobe inférieure plus saillant, le stylet est comprimé de haut en bas, bifide, obtus.”
The description fits the clasper of U. lotrix. What Rambur says about the
anal tergum, his “stylet,” is only understandable if we assume that he had a
specimen in which the tip of the tergum was broken off or in which the tergum
was lying on its side.
Novıtates Zoonocican XLI. 1939. 287
Spots of upperside rarely scarlet (in one of our specimens and two in the Hope
Department) usually more or less deep salmon, paler on the upperside than below.
On underside the red markings between cell and hindmargin as distinct as in
U. pulchella, forming in many examples bands, which are particularly conspicuous
in Madagascar specimens, more rarely in Continental ones.
Modified scaling on hindleg of 3 very variable in extent, sometimes practi-
cally absent, in others the scales nearly as numerous as in U. 1. lotrix, but the
area much less sharply defined.
Hab. Ethiopian Region, inclusive of Arabia ; not known from the southern
half of the continent.
In the Rothschild collection from Gambia: Bathurst, 5 jg, 11 99; Liberia,
12; Oasis Bilma, Nov. 1922, 2 49,5 99; Upper Egypt: Assuan, 1 9; Gara,
Nov. 1816; Upper Congo: Lukolele, 1 9; Uganda: Entebbe, Oct. 1900,
2 99; Kenya: Escarpment, Nov. 1901, 1 9; Abyssinia: Sekwala, south of
Addis Abeba, Nov., 1 3, Maki R., north of L. Zuai, Nov., 2 33g, Gorgoru Ganale
R., April, 1 9, Dabele, Ganale R., April, 1 3, 1 9, Malka-Ré, Dana R., May, 2 29,
Habrona, Merehan, May, 1 9, Solole and Hanole, Ganale R., June, 1 J, 1 9,
Diré Daoua, May 1935, 1 & ; Madagascar: Antanosy, 1 3,1 9, Lambomakandro,
Sakaraha, Tulear, March 1935, 1 9, “ Madagascar,” 5 99,
In Hope Department, Oxford, from Dahomey: Cotonou, June 1914, 1 &;
Nigeria: Lagos, 1 3,19; Egypt: Harkeko and Akeek (from the Meldola coll.),
2 34,12; Sudan: Khartum, Feb. 1909, 1 g, 1 9, El Dueim, Feb. 1909, 2 32,
Nuba Mts., Nov. 1917, 2 gg; S.W. Abyssinia: Tirma, April 1925, 1 9.
Some additional localities in the B.M. collection—Arabia: Jidda, Feb.
1936 and 1939 ; Hejaz, Dec. 1927.
I have not seen any specimens from Bourbon and Mauritius, but expect
them to be like Madagascar ones. Our 8 examples from Madagascar do not
differ from every one of our Continental series, but are remarkably uniform in
size, coloration and neuration. The scarlet is strongly developed above and
below, and the black dots are larger than in most Continental specimens ; length
of forewing 18-20 mm., on the Continent 15-21 mm.
(b) U. lotrix socotrensis subsp. nov.
Deiopeia pulchella, Butler (error determ.), Proc. Zool. Soc. Lond., p. 179 (1881) (Socotra) ; Dixey,
ibid., p. 372 (1898) (Socotra).
Utetheisa pulchella, Hampson, Lep. Phal., iii, p. 483, no. 2088 (1901) (partim ; Socotra); id., in
Forbes, Nat. Hist. Socotra, p. 322, no. 3 (1903) (Socotra, very common),
Deiopeia pulchella, Rebel, Denkschr. Math.-Nat. Kl. K. Akad. Wiss. Wien, lxxi, 2, p. 53, no. 66 (1907)
(one of the commonest Lepidoptera in the southern half of Socotra; one 9 extended vermilion).
Utetheisa pulchella, Strand, in Wagner, Lep. Cat., pars 22, p. 362 (1919) (partim ; Socotra).
Nearest to U. 1. lotrix; modified scaling of hindleg of ¢ as extended, and
the area occupied by it on the tibia as sharply defined, as in U. 1. lotrix, differing
therein markedly from U. I. lepida. Scarlet markings of upperside strongly
developed, with a tendency to form transverse bands, the wing appearing more
scarlet than creamy white and contrasting much with the forewing of U. 1. lepida
and the series of U. 1. lotrix from the Seychelles, Ceylon and British India ; behind
vein 1b a series of 3 or 4 scarlet or olive spots, which are frequently present also in
U. 1. lotrix, but are hardly ever indicated in Continental specimens of U. 1. lepida.
Hab. Socotra: Adho Dimellus, Hombil, Jena-agahan, Ardahan. In B.M.
1 3 (type), 3 92; in the Hope Department 3 33, 1 9.
21
288 € Novirates ZooLoGIcAE XLI. 1939.
(c) U. lotrix lotrix Cramer 1777 (text-figs. 229, 233, 237, 255).
Phalaena Tinea lotrix Cramer, Pap. Exot., ii, pp. 20 and 149, pl. 109, figs. E, F (1777) (partim ;
Coromandel, Tranquebar).
Utetheisa pulchella auctorum (partim).
Utetheisa pulchella form tenuella Seitz, Macrolep., ii, p. 73, pl. 13k (1910) (“‘ East Asia,” no locality
specified).
Utetheisa pulchella v. tenuella, Strand, in Wagner, Lep. Cat., pars 22, p. 365 (1919) (East Asia),
Utetheisa lotrix, Maulik, Ann. Mag. N.H. (11), iii, p. 232, text-fig and pls. 8, 9 (1939).
Utetheisa pulchella ab. flava Bang-Haas, Horae Macrolep., i, p. 58 (1927) (Darjiling).
The figure in Seitz, l.c., named tenuella, represents this subspecies; East
Asia being very vaguely given as the home of this “‘ form,” I select as terra typica
Kowloon, near Hongkong.
The specimen with yellow instead of scarlet spots named ab. flava by Bang-
Haas most probably belongs to this subspecies ; we have three such specimens,
one each from Sumatra, Java and Celebes. A & from Ajmere corresponds in
colouring to U. pulchella ab. bicolor.
In nearly every specimen the two posterior terminal spots of forewing
elongate, or at least longer than the spots preceding them. The scarlet spots
sometimes deep salmon as in U. 1. lepida, in none of our specimens forming more
or less complete bands ; the creamy stripe in cell broader than in U. 1. socotrensis.
On underside the scarlet colouring almost completely confined to the costal spots
and submarginal band, the scarlet discal spots absent or so
covered with white scaling that they look very pale pink. On
hindwing below always a black spot or smudge at or close
to anal angle, which is absent from many specimens of
U. 1. lepida.
The subspecies is not so constantly small as one might
infer from Seitz’s remark about his tenwella, l.c., many speci-
mens being larger than the average U. pulchella; length of
forewing 14-20 mm. I have not found any differences in the
material from widely separate districts except New Guinea. The area of
modified scaling on the hindleg of the & is sharply defined in all the localities
(text-fig. 255).
Hab. Seychelles, Ceylon, China and Japan eastwards to New Guinea and
Australia ; also on New Zealand (immigrant).
In the Rothschild collection from Seychelles : Mahé and St. Anne, all months
July to Dec. 1906, 5 $3, 11 99; Ceylon, 3 3g, 3 92; India: Travancore,
Nilgiris, Madras, Malabar, Ganjam, Muttra, Karachi, Ajmere, Sikhim, Khasia
Hills, 12 34, 17 99; Burma, 2 39,19; S.E. China: Kowloon, Taipoo and
Hongkong, 6 33, 7 99; Japan, 1 g; Formosa: Anping and Tainan, 3 33,
4 99; Annam: Phuc-Son and Xon-Gom, 1 9,19; Singapore, 1 3,19; West
Sumatra : Benkoelen and Padang Bovenlanden, 16 $4, 19 99; Java: Buiten-
zorg, Garoet, Mt. Gedeh, Tjilatjap, Pengalengan, Trettes, Nonkodjadjar, 17 $3,
31 92; Bali: Gilmanoek, Prapetagoeng, Mondoktoempang, “ Bali,” 5 33,
6 92; Lombok: Sapit and Sambalun, 7 3g, 4 92; Christmas Is., 1 3; Sam-
bawa, 1 g; Alor, 2 92; Pura, 2 99; Dutch Timor: Oinanisa, 1 9; Kisser,
6 33,299; Tenimber, 2 44; Koer, 2 gg; Banda, 1 3; South and Central
Celebes, 5 $3, 5 99; North Borneo: Labuan, 1 3; Palawan, 1 3; Philippine
Is.: Luzon, 1 3, 1 9, Panay, 2 3g; New Guinea: Astrolabe Bay, 3 3g, 4 99,
Aroa R., 1 3; Dampier I., 3 33, 4 22; Vulcan Is., 4 gg, 4 99; French Is.
Novirarrs ZooLoGıcAE XLI. 1939. 289
(Witu), 1 2; Goodenough, 1 9; Sudest Is., 2 $&; North Australia: Port
Darwin, 2 99, Eureka, 6 gg, 3 92; West Australia: Sherlock R., 1 9, Roe-
bourne, 1 9, Perth, 1 2; Queensland: Cairns, 2 99, Toowoomba, 2 33, 3 29,
Bowen, 1 3, Mackay, 2 99, Brisbane district, 2 gg, 1 2, “ Queensland,” 5 gd,
5 22; New Zealand (coll. Felder), 1 9.
Some additional localities in the B.M. collection—Nicobar Is. ; Camorta ;
West China: Moupin and Pu-tsu-fong ; Amboina (1 , ex coll. Orberthür).
The 27 specimens in the Rothschild collection from New Guinea and the
islands nearby on the eastern side are remarkable as a series in as much as in
44 per cent. of the forewings the discocellular black bar on the underside is
connected with the discal band, whereas this is the case only in 4 per cent. of the
specimens from the Malay Archipelago. The specimens do not seem to differ
in any other way except in neuration (see Table II, p. 256).
It is rather surprising that we have no specimen of U. lotrix from Hainan
and only a few from the Philippines whence there are in the collection large
numbers of U. pulchelloides, and that the Moluccas proper are represented but
by a single example in the long series of U. lotrix examined. The species seems
to be commoner in the southern than in the northern districts of the Oriental
Region, and its range is more restricted in the Pacific than that of U. pulchelloides.
(d) U. lotrix stigmata Rothschild 1910.
Utetheisa pulchelloides stigmata Rothschild, Nov. Zool., xvii, p. 182, no. 48 (1910) (N. Caledonia,
Loyalty Is.) ; id., l.c., xviii, pl. 6, fig. 27 (1911).
Utetheisa lotrix stigmata Rothschild, in Seitz, Macrolep., x, p. 260, pl. 13k (1914) (““ Freundschafts-
inseln”’ in error for Loyalty Is.).
Utetheisa lotrix var. stigmata, Strand, in Wagner, Lep. Cat., pars 22, p. 361 (1919).
Utetheisa pulchelloides form stigmata, Hampson, Lep. Phal. Suppl., ii, p. 511 (1920).
The black dots of forewing above larger than usual, at least some of them
merged together ; discocellular spot of hindwing large, black border broad.
Black markings of underside likewise enlarged, particularly the black discal band
broader than usual and frequently connected with the discocellular black bar ;
the proximal costal spot of the hindwing not rarely continued by a line across
the cell.
The red specimen mentioned by Lord Rothschild in 1910 and termed by him
ab. rubra in Seitz belongs to the next species. The type of stigmata, a 9, is from
Lifu.
Hab. Loyalty Is.; New Caledonia ; New Hebrides; Fiji.
In the Rothschild collection from New Caledonia : Nouméa, 6 22; Loyalty
Is.: Lifu, 1 g, 4 99.
In the B.M. from the New Hebrides (nearly all collected by Miss Cheesman) :
Aneityum, Tanna, Hog Harbour, 5 ¢ 4, 6 92; Fiji, 1 & (colouring of forewing
very dull).
10. Utetheisa sumatrana Rothschild 1910.
Utetheitsa sumatrana Rothschild, Nov. Zool., xvii, p. 182, no. 51 (1910) (Delhi, N.E. Sumatra) ; :d.,
l.c., zviii, pl. 6, fig. 31 (1911); id., in Seitz, Macrolep., x, p. 260, pl. 24g (1914) ; Strand, in
Wagener, Lep. Cat., pars 22, p. 366 (1919); Hampson, Lep. Phal. Suppl., ii, p. 512, no. 2089a
(1920).
Utetheisa lotrix stigmata ab. rubra Rothschild, l.c.
Forewing above Chinese orange, a little deeper red towards base, markings
more or less obliterated, usually a white streak in cell. Hindwing white,
290 Novirates ZooLoGIcAE XLI. 1939.
the black border varying from being absent to being nearly like that of
U. pulchella.
Underside of forewing white from middle of cell to hindmargin ; black discal
band indicated or distinct. Apex of hindwing with an ill-defined Chinese-
orange patch, distal margin more or less washed with the same colour.
In the specimen called ab. rubra by Rothschild the forewing is an orange-
scarlet, with the dark markings more extended and more numerous ; fringe with
olive-brown spots corresponding to the marginal spots of the same dull colour ;
hindwing with discocellular bar and narrow black margin which has a broad
projection behind lower median, this projection longer than broad. On
underside the hindwing with orange-scarlet apical patch like normal specimens.
This specimen was received from Messrs. Watkins and Doncaster together
with other set material labelled Lifu, Loyalty Is. I consider the specimen to
have got mixed up with the Lifu material on the setting boards.
U. sumatrana is a very puzzling species. It is so different in coloration
from U. lotrix that the close agreement in structure with that species is surprising.
Hab. N.E. Sumatra.
In the Rothschild collection 2 34, 2 22 from Deli, N.E. Sumatra.
In the Dohrn collection at Stettin 8 specimens ; one of the females with the
hindwing nearly as in U. pulchella, according to the information very kindly
sent me by Dr. Urbahn.
11. Utetheisa semara Moore 1859 (text-figs. 230, 234).
Utethesia (!) semara Moore, in Horsf. & Moore, Cat. Lep. East-Ind. House, ii, p. 307, no. 712, pl. Vila,
fig. 12 (1859) (Java).
Utetheisa semara, Hampson, Lep. Phal., iii, p. 485, no. 2089 (1901) (Java) ; Rothschild, in Seitz,
Macrolep., x, p. 260, pl. 24g (1914) ; Strand, in Wagner, Lep. Cat., pars 22, p. 366 (1919) (Java).
For distinctions in the genitalia, see p. 258. Forewing above olive-brown,
with dark-red streak along costal margin, elongate spots of the same colour
between the veins and more or less elongate black spots, a dull creamy-white
streak in cell from base on to disk. Hindwing white, without discocellular
spot ; olive-black distal border nearly even in width.
TABLE VI.
NEURATION OF U. SEMARA,
Celebes Wings. Lombok Wings. Java Wings. eee ln all
Number of wings : 6 3 . 30 16 6 52
With areole: Vz | Gi
(a) Vein 10 from areole : “| 2 | — | 3 10%
(6) Veinl0onashort stalkwith8.9| 6 | mo | =, 30%
‘(c) Vein 10 on a long stalk with 8.9 i a RT 8%
Without areole : | ]
(d) Vein 10 from 7.8.9 : : — 1 — 2%
(e) Vein 10 direct from cell. . 2 i 4 PET 50%
(f) Vein 11 from areole or from 10. | 17 8 We aed 56%
The numbers of specimens we have are not large enough to make the com-
parison of the neuration of the Lombok and Celebes examples reliable ; never-
theless, the result shown in the table is interesting, if compared with Table II on
Noviratres Zoonoarcar XLI. 1939. 291
p. 256: vein 10 rarely arises from the apex of the areole and very frequently
free from the cell (50 per cent.), and the latter is much more often the case in
Celebes than in Java and Lombok ; vein 11 is in 17 of the 30 Celebes wings on
a short stalk with 10 and in 8 of the 16 Lombok wings, whereas in U. pulchella
and U. lotrix this happens rarely.
Hab. Java, Lombok and Celebes.
In the Rothschild collection from West Java, 1 ¢; Lombok: Sapit and
Sambalun, 3 gg, 5 99; South Celebes : Bua-kraeng, Lampa-Battan, Bonthain,
4 33,7 22 (the other 4 specimens examined are in the B.M. coll.).
EXPLANATION OF TEXT-FIGURES.
Fig. 226. Neuration of Utetheisa pulchella : > F 09255
227. Clasper of Utetheisa pulchella, outer as 258
228. Rn ” pulchelloides, outer surface 258
229. an ik lotrix, outer surface 258
) 230. ae semara, outer surface 258
231. ical segments of abdomen of U. pulchella 2
232. Sternum VII of U. pulchelloides 2 258
233. i“ „ U. lotrix 2 258
234. „ U. semara 2 258
235. Seales from pocket of clasper of U. bella
236. ry, < A a U. pulchella 260
ZBI. en Ir in u U. lotrix . 260
238. Clasper of U. pulchella, inner surface 375
239. 5% U. amhara, outer surface 375
240. Claspers of U. amhara, from anal side
241. Clasper of U. amhara, inner surface
EB U. salomonis, outer surface
243. na U. lotrix lepida, inner surface 5
244. Segments 25 and 26 of male antenna of U. ee
>25 SU, CH CH Pee ee Pe
bo
fer)
(=)
pectinata, dorsal aspect ; Selaru, Tenimber p. 280
245. The same, lateral aspect p- 280
246. The same, ventral aspect . p. 280
247. Apical surface of segment 27 of U. p. an p- 280
248. Segments 25 and 26 of male antenna of U. p. Perel aes
ventral aspect ; Chagos d . 6 p. 285
249. Apical surface of segment 29 of the same p- 285
250. Segments 25-27 of male antenna of U. p. nee um,
lateral aspect ; Marshall Is. . ; 5 : . p. 285
251. Segments 26 and 27 of the same, ventral perce ; ; p- 285
252. Segments 27-29 of male antenna of U. p. vaga, lateral a ;
Java : : . p. 285
253. Segments 27 and 28 af the same, van aspect : p- 285
254. Apical surface of segment 29 of the same p- 285
255. Diagram of scaling of outer surface of hindtibia of U. ige
lotrix . 6 : c 5 6 : & : . p. 288
292 NOVITATES ZOOLOGICAE XLI. 1939.
ON FIVE NEW SIPHONAPTERA FROM THE REPUBLIC
OF ARGENTINA.
By KARL JORDAN.
(With 10 text-figures.)
u species and subspecies here described form part of a collection sent to
the British Museum by Dr. J. M. de la Barrera, of the Instituto Bacterio-
logico at Buenos Aires. The specimens being obtained in western districts of the
Republic whence we have received but little material, it is not surprising that
several of the fleas are new. The three Parapsylli here described are particularly
interesting, widening our knowledge of the morphology of the genus in more than
one direction. The species I have the pleasure in naming after Dr. Barrera is
distinguished from all the other Parapsylli by the long proboscis being 8-seg-
mented ; another species differs in the tarsal claws being strongly asymmetrical
in both sexes ; and the third species is exceptional in having more than 80 pits in
the sensilium of the ninth abdominal tergite. The collection further contains
an example of geographical
variation of a species of flea
within the Republic, the speci-
mens of Rhopalopsyllus platensis
from the foothills of the Andes
being different from those we
have from east and west of the
Rio de La Plata. A collection
like the present makes one hope
that the survey of the Argen-
tinian Rodent fauna will be
continued and further material
of Siphonaptera be forthcoming.
1. Hectopsylla gemina sp. nov.
(text-fig. 256).
@. Agrees with H. eskeyi
Jord. 1933 in the subbasal
position of the pits of the sen-
silium (text-fig. 256, C) and with
H. stomis Jord. 1925 in the
shape and small size of the
spermatheca (text-fig. 256, R.s.) ;
differs from all the species in
the metepimerum and tarsi.
Bristles of metepimerum (4) and
abdominal tergites I and II stronger than in the allied species ; upper posterior
angle of metepimerum (text-fig. 256, A) produced into a hook, which varies in
size, but is much more curved down than in H. stomis and H. eskeyi in all
NOVITATES ZOooLoGIcAE NLI. 1939. 293
7 specimens. Stigmata of abdominal segments I to VII slightly smaller than in
H. stomis and much smaller than in 4. eskeyi, the diameters measuring 11, 13
and 17 in the three species. Bristles at apical margin of tergite VIII variable
in size, 2 to 5 of them being larger than the others. Segment II of midtarsus
longer than I and in hindtarsus not much shorter than I, the measurements
being : midtarsus 7, 9, 7, 5, 12; hindtarsus 16, 12, 8, 6, 13. Segment V bears
in all tarsi only 3 pairs of plantar bristles (text-fig. 256, B, midtarsus) ; in
H. stomis there are very occasionally also 3 pairs or 3 bristles on one side and
4 on the other in one or the other tarsus, in H. gemina the number is constantly
3 in the 7 specimens before me.
Hab. Argentina: Fortin Uno, Rio Negro, August 1937, on Microcavia
australis, 2 99 (type 1); Santa Rosa, Mendoza, February 1938, on the same host,
5 99.
2. Rhopalopsyllus platensis cisandinus subsp. nov.
39. A western modification of Rh. platensis Jord. & Roths. 1908, distin-
guished by the reduction in the number of bristles on the thorax, abdomen and
legs. Genitalia the same as in Rh. pl. platensis, of which we have a series from
eastern districts of Argentina and from Uruguay.
Bristles on pronotum in two rows, occasionally a single additional dorsal
bristle in front of the first row: in ¢ 12 to 15, 15 or 16, in 9 11 to 15, 15 to 18.
On mesonotum in ¢ 16, 13 to 15, in 2 13 to 18, 13 to 15. On metanotum three
rows, in ¢ 7 to 11, 15 to 18, 14, in 9 8 to 10, 16 to 19, 15 or 16. On mesopleura
in ¢ 5, in 9 5 or 6; on metepimerum two rows, in g 4 or 5, 3 to 5, in 9 3 to 5
(usually 4), 4.
On abdominal tergites I in ¢ 1, 11 to 14, 10 to 13, in @ 1 to 3, 11 to 14,
11 or 12; Ilin g 13 to 16, 16 to 18, in 2 16 to 19, 17 to 21,; II. in & 10 to 13,
16 to 18, in 2 16 to 18, 17 to 20; IV in & 8 to 12, 15 to 19, in 9 12 to 17, 17 to
19; Vin ¢ 5 to 10, 15 or i6, in 2 15 or 16, 17; VI in ¢ 5 to 8, 14 or 15, in
99to 14,16; VILin 9 5to7, 9 or 10, in 29 or 10, 12 or 13. Lateral bristles on
each side of basal sternum of abdomen in & 6 to 10, in 9 17 to 24; on the other
sternites, the two sides together : on Ill in ¢ 4 to 6, 6 to 9, in 9 9 to 15, 10 to 13;
IVin gO orl, 6to 8,inQ@0orl, 8toll; Vin 90, 8,in 20, 8to 10; VI in
6 0, 7 to 9, in 20, 10 to 12; VILin G0, 8, in 9 (the two rows together) 15 to 19.
On outer surface of hindfemur a subventral row of 7 to 9 bristles in both
sexes and above the row 2 to 5; on inner surface a subventral row of 5 or 6 in
& and 6 or 7 in 9. On outer surface of hindtibia in g 11 to 14 subdorsal and
lateral bristles and 6 or 7 subventral ones, in 9 12 to 15 subdorsal and lateral
and 6 to 8 subventral.
Above stigma of tergum VIII of ¢ and 9 9 to 11 bristles on the two sides
together ; from stigma downwards in 9 on outer surface of each side 34 to 41,
on inner surface at and near margin 13 to 17; on sternum VIII of ¢ 10 to 15
on right and left sides together.
In Rh. pl. platensis the basal abdominal sternum of ¢ bears each side more
than 15 bristles ; the numbers are on the two sides together on metanotum in
& 10 to 12, 21 to 26, 16 to 18, in 2 10 to 14, 19 to 28, 17 to 21; on abdominal
tergum III in & 15 to 22, 21 to 23, in @ 20 to 28, 21 to 26; VII in $7 to 12,
15 or 16, in 2 11 to 17, 14 or 15; abdominal sternum VII of 2 19 to 25 (usually
1 The word “ type ” in brackets means here (and in other cases) that one of the specimens has been
selected as the type, the other specimens mentioned being paratypes.
294 Novirares ZooLoGIcAE XLI. 1939.
more than 20). On outer surface of hindfemur in & a subventral row of 9 to 12
and above it 6 to 8, in 9 a row of 9 to 11 and above it 7 to 10 (rarely fewer than 7).
Hab. Argentina: Mendoza, January 1938, on Microcavia australis, a series
(type) ; Telen, La Pampa, April 1936, on Graomys griseoflavus, 1 3, on Microcavia
australis, 1 3.
3. Parapsyllus galeanus sp. nov. (text-figs. 257-260).
39. Nearly related to P. budini Jord. & Roths. 1923, but differs from all
known species in the claw being strongly asymmetrical in all tarsi.
In front of eye 4 bristles, of which the first and third counted from below
are long, in & 2 or 3 small additional bristles farther upwards, below eye in & 3
and in 9 2 bristles, behind eye 2 or 3 small ones, of which the lowest is the longest.
On occiput a longitudinal row of 3 in ¢ and only 1 in 9, which is placed near
middle ; a subapical row of 11 or 12 (the two sides together) in g and 9, with an
additional bristle below the long ventral one in 2; above antennal groove the
usual dense row of short bristles in g, only a few such bristles posteriorly in 9.
Segment I of antenna much longer and broader in ¢ than in 9, with an apical
row of bristles which do not reach the middle of the club in 4, whereas in 9 they
are much longer, some of them extending beyond the club ; segment II for the
greater part covered by I, visible only at the sides, where it bears some small hairs.
Proboscis reaching to near apex of forecoxa in 9, a little shorter in ¢; labial
palpus consisting of 5 segments, V being as long as or a little longer than III + IV.
Proportional length of segments of maxillary palpus in ¢ 11, 14, 9, 16, in 2 the
segments a little shorter, slightly variable in both sexes.
On thoracic tergites two rows of bristles, with a few small subbasal dorsal
hairs on mesonotum ; the numbers are in J on pronotum 6 to 10, 11 to 13, on
mesonotum 9 to 11, 10 or 11, on metanotum 9 to 13, 9 to 12, the anterior row
containing in @ 2 or 3 fewer bristles ; on mesopleura 5 and on metepimerum 2 or
3, 3 in both sexes. Apical area of metanotum more (9) or less ($) membranous.
The abdominal tergites bear two rows of bristles, except the posterior
segments of 3, the numbers being on I in ¢ 10 to 13, 8 to 10, in 2 8 to 12, 8;
Il in $7 to 13, 13 to 16, in 9 9 to 12, 12 or 13; Ill in ¢ 2 to 11, 13 to 15, in
2 10 to 12, 12 to 14; IVin J 0 to 4, 13 or 14, in 2 10 to 12,13; Vin 60 to 2,
13 or 14, in 2 9 to 11, 12 or 13; VI in 30, 12 to 14, in 2 9 to 13, 12 to 14;
VIL in 3 0, 11 to 13, in 2 9 to 12, 8 to 12. On basal sternite (each side) in & 0
to 2 small lateral bristles, in 9 12 to 17, besides the few bristles of the subapical
row ; the numbers on the other sternites are (the two sides together as in the case
of the tergites) : IIIT in g 0, 6 to 8, in 9 13 or 14, 9 to 12; IV in 30, 6 to 8, in
©2to 5,10 0r1ll; Vin 9 0,7 or 8, in 91 to 4, 13 to 16; VIlin ¢ 0, 6 to 8,
in 9 7 to 9, 16 to 18.
The forecoxa bears 28 to 38 bristles in g (not counting the short basal ones)
and 22 to 26in 9. The mid- and hindfemora have a subventral row on outer and
inner surfaces and no lateral bristles above these rows, the numbers being on out-
side of hindfemur in & 10 to 15 and in 9 9 to 13, on inside in g 9 to 14 and in 2 9
to 12; as a rule these bristles are more numerous on outside in ¢ and on inside
in 9, with exceptions. Foretibia with 6 dorsal notches and an additional stout
apical bristle ; mid- and hindtibiae with 7 or 8 dorsal notches ; on outer surface
of hindtibia in 3 20 or 21 dorsolateral bristles, in 9 12 to 16; the longest dorsal
and ventral apical bristles of hindtibia reach beyond apex of tarsal segment I.
Novirarres ZooLoGIcAE XLI. 1939. 295
Tarsi very distinctive : V in all tarsi only twice as long as broad (claw excluded)
(text-fig. 257); I and II of midtarsus almost equal in length ; I of hindtarsus
about half as long as hindtibia, three times as long as broad, II short, triangular,
only half as long again as broad, IV in all tarsi broader than long. Measurements:
midtarsus, in & 12, 11 or 12, 8, 5 or 6, 17, in 9 11, 12, 8 or 9, 5, 16; hindtarsus
in d 32 or 33, 13 or 14, 9 or 10, 7, 17, in 2 28, 13 or 14, 9, 6, 17. Bristles of
hindtarsus very long, longer in ¢ than in 9 (text-fig, 257, 9); in g I with 5
notches on posterior side bearing each a
long bristle and a short stiff one, the
proximal long bristles reaching to apex of
V, the others beyond it; II with 3 long
bristles, two on the posterior side and one
anteriorly, all 3 apical, on the anterior side
also a stiffer one which extends to near
middle of V; in 2 I with 2 or 3 long
bristles, the long apical one reaching to near
apex of V, the longest of II extending to
apex of V, not beyond. Inner claw nearly
as long as V (foreshortened in text-fig.
257), the tooth much enlarged in both
sexes, broad and about 2 the length of the
claw ; outer claw small (text-fig. 258).
Modified Segments.—d. Of the same
type as in P. budini, but different in detail.
Sternum VIII with two processes each
side, both narrow and pointed (text-fig.
259), the upper one (L!) the broader and
shorter, more or less curved upwards, the
lower one (L?) curved downwards: at end,
both somewhat variable ; at some distance
forward 2 or 3 bristles each side and farther
up a vertical row of 3 long ones, of which
the lowest is the shortest and often absent.
Apex of clasper (Cl) less rounded than in
P. budini, the angle being about 90° or less,
distal margin rounded, obtusely angulate in
middle, often more rounded-elbowed than
in the specimen figured ; at dorsal margin
many small bristles, especially numerous
on inner side, on lateral surface about 8, nearly all long, from upper angle to
median elbow a marginal row continued downward by a submarginal row of
longer ones, the entire row containing some 20 bristles, of which the upper
ones are small; ventral spur (Sp) of clasper proximally more curved than in
P. budini, the horizontal portion dorsally almost straight or feebly concave,
ventrally convex basally and apically and nearly straight in middle; the spur
has no bristles on outer surface, but about 20 on inner side, of which 5 or 6 are
long and placed at the ventral apical margin ; inner side of clasper with many
small bristles in apical half. Digitoid (F) not projecting beyond the margin of the
large clasper, elongate-conical, with the apex drawn out into a point, anterior
296 NOVITATES ZooLoGIcAE XLI. 1939.
margin more or less convex, posterior margin partly concave, bearing in upper
half a row of 6 to 8 thin bristles and a lateral row of small ones ; posterior portion
of base rounded, bearing a row of small bristles. Manubrium (M) very broad,
broader than the lamina (Lam) of the phallosome. Vertical arm of sternite IX
elbowed, apex wider than in P. budini, both the forward and backward projec-
tions longer; ventral arm nearly straight dorsally, slightly curving upwards,
apical angle rounded, not pointed as in P. budini, apical fourth of segment
abruptly enlarged ventrad, this area bearing on outer side some 20 longish bristles
which are more or less compressed, resembling slender blades of grass. Para-
meres of phallosome ending in a pair of short pincers curved towards each other,
the lower one less pointed than the upper and ventrally widened into a rounded
lobe ; dorsal hood rounded at apex.
9. Sternum VII broadly rounded, the margin ending at the ventral bristles
of the posterior row ; at this angle 3 or 4 long bristles rather close together, on
each side of the body. Apical margin of tergum VIII (text-fig. 260) rounded,
without sinus below apex, above stigma 4 to 7 bristles each side, from stigma to
Novirarrs ZooLoGIcAE XLI. 1939. 297
ventral margin 21 to 24 bristles, most of them long, at apex 3 or 4 long marginal
bristles on outer side and 17 to 20 marginal and submarginal ones on inside, most
of which small. Pleural sclerite of segment IX (IX.p.) very distinct above
dorso-apical margin of VIII.t. On side of anal tergum, between the small papilla-
like bristle, ventral margin and stylet 4 or 5 bristles. Body of spermatheca
<---VIILst.
(R.s.) short-ovate, sometimes almost globular, as long as the posterior margin
of tail (margin on anal side).
Length: 3 2-4 to 2-3 mm., 9 2-9 to 3:2 mm. ; hindfemur : & 0:55, 9 0-50
to 56 mm.
Hab. Argentina: San Juan, May 1938 on Galea leucoblephara, a series of
both sexes.
4. Parapsyllus barrerai sp. nov. (text-figs. 261, 262).
62. The most conspicuous distinguishing character of this species is the long
proboscis, which extends below the base of the forefemur, the labial palpus being
anteriorly divided into 8 segments. This distinction, though very interesting, is
not of generic value, at least not in a classification based on relationship. The
new species is undoubtedly closely akin to P. budini and allies with 5-segmented
labial palpi. Moreover, the 9 of P. onychius Jord. & Roths. 1923 has 6 segments
and the ¢ only 5, and in the present new species two of the segments are separated
only on the anterior side, all of which shows that the segmentation is in flux and
not of primary importance in this case.
In front of eye 3 bristles of which the middle one is short and the others long,
farther forward a longish bristle in ¢ only ; below and behind eye a row of 7 or
8, of which some are small. On occiput in ¢ the usual row of short bristles
(about 24 in this species) above antennal groove and a longitudinal row of 3 long
298 NOVITATES ZOOLOGICAE XLI. 1939.
ones, in 9 a median bristle and a ridge which bears posteriorly a few small ones ;
in both sexes a subapical row of 12 (on the two sides together), with an additional
short bristle below the long ventral one in 9. Bristles of antenna quite short in
gand ®. Length of segments of maxillary palpus in g 11, 15, 11, 19, in 2 15, 21,
14, 22. In & segments 4 and 5, in 9 6 and 7 not separated on posterior side
(in this pair).
Bristles on thoracic and abdominal nota in two rows: on pronotum in & 7,
16, in © 5, 16; on mesonotum in ¢ 7, 12, in 2 5, 13; on metanotum in & 9,
12, in 9 5, 14; on mesopleura in & 5, in 9 7; on metepimerum in & 2 or 3,
3, In 2 2 or 8, 4.
Bristles on abdomen : on tergites [in ¢ 6, 11, im 2 12, 14; Ilin 310, 15,
in 9 20,18; IIIin 1, 14, in 2 20, 16; IV in 3g 0, 14, in 9 20,16; Vin &0,
14, in 919, 15; Vl in & 0,13, in 9 23, 15; VII in 0, 12, in 924,13. On
basal sternum in & 5 lateral ones on one side, 6 on the other, in 9 34 and 44; on
the other sternites, the two sides together: on III in ¢ 2, 8, in 9 16, 12; IV in
31, 8,in 92,10; Vin ¢0,8,in 91,10; VIin $0,8,in 96, 12. Striation
of basal sternite not dense.
Forecoxa with about 25 bristles in ¢ and 38 in 9. On outer side of hind-
femur a subventral row of 6 bristles in ¢ and 1 lateral bristle, in 9 the row con-
taining 7 to 9 and there being 3 lateral ones ; on inner side the numbers are 9 or
10 in gf and 12 to 15 in Q, no lateral bristles. On foretibia 5 dorsal pairs of strong
bristles, the subbasal pair absent ; mid- and hindtibiae with 6 dorsal notches,
two of them in basal area, two about middle and two at and near apex, in between
second and third notches 1 or 2 small bristles, between fourth and fifth 1 in g and
2 or 3 in 9; on outer surface of hindtibia in $ 7 to 9 dorsolateral bristles and
5 or 6 subventral ones, in 9 10 dorsolateral, 1 or 2 lateral and 7 or 8 subventral ;
the long anterior as well as posterior apical bristle of midtibia extends well be-
yond apex of tarsal segment I, in hindtibia to apex or close to it. Hindtarsal II
with 3 long apical bristles, of which the longest of the posterior side reaches
beyond (3) or to (2) apex of V, the second beyond middle of V and the third to its
base ; V in hindtarsus more than three times as long as broad ; claw long and
slender, in fore- and midtarsi as long as V, in hindtarsus shorter than V, basal
tooth not broader than base of claw. Length of segments: midtarsus in & 21,
17, 9, 6, 16, in 2 34, 29, 14, 9, 21; hindtarsus in ¢ 40, 28, 13, 9, 19, in 3 60, 40,
17, 11, 23.
Modified Segments.— 3. Above stigma of tergum VIII 5 bristles each side.
Sternum VIII with one subventral lobe (L) each side, the lobe triangular, pointed,
dorsally convex, ventrally almost straight, measured dorsally about as long as it is
broad in middle ; beyond middle of segment a vertical row of about 20 bristles
on the two sides together, the upper ones long, lower ones small (text-fig. 261).
Dorsal apical angle of clasper (P) projecting, triangular, posterior margin from
this lobe to the ventral spur rounded ; from base of lobe to ventral spur a row of
8 or 9 long bristles, the upper 3 marginal, the others submarginal ; on dorso-
apical lobe one longish bristle and a number of smaller ones, on outer surface
8 long and 3 or 4 near dorsal margin shorter, at dorsal margin a number of small
bristles. Ventral spur of clasper (Sp) in shape like a scythe, gradually narrowing
to a sharp point, upper margin slightly concave, ventral margin convex, at
ventral margin 12 bristles, 4 in apical third and 2 at base of spur about as long as
the spur is long dorsally, the others much smaller, between spur and vertical arm
Novirates ZooLocıcan XLI. 1939. 299
of IX.st. about 6 small bristles. Manubrium (M) narrower than vertical arm of
IX.st., apex sharply pointed and turned up. Digitoid (F) narrow, more than four
times as long as broad, narrowed from middle to apex, which is sharp, in apical
fourth of posterior margin some thin bristles and on lateral surface a few very
short ones ; F extends above the margin of clasper, but does not reach the dorso-
apical angle; the digitoid of the other side (not indicated in figure) is moved
farther away from the dorso-apical lobe and appears narrower than the digitoid
figured. Vertical arm of sternum IX almost straight, not elbowed, its apex
posteriorly evenly rounded, the anterior lower projection also round ; ventral
261
VIllst.
“VIILst.
arm gradually widened, its dorsal margin slightly concave and the ventral margin
feebly convex, apex dorsally strongly and evenly rounded, ventrally produced
into a hook; proximally of this hook 4 moderately long marginal bristles and
farther forward 3 small ones, on lateral surface a small number of minute hairs,
apex inclusive of hook more than one-fourth as broad as the ventral arm is long.
Parameres (Par) of phallosome with two apical teeth, of which one is lateral ‘in
our specimen and the other ventral.
9. Apical margin of sternum VII (text-fig. 262) rounded subventrally, above
this lobe incurved, first more strongly and then very feebly, no sinus being formed.
On tergum VIII above stigma 8 bristles on one side and 9 on the other ; apical
margin of widened area sinuate subventrally ; the rounded lobe above this sinus
bears about 30 bristles, of which about 12 are submarginal on the inner surface ;
on the lobe below the sinus a subventral row of 3 or 4, and on the lateral surface
300 NOVITATES ZOOLOGICAE, XLI. 1939
between this row and the stigma 13 or 14 bristles arranged in two irregular rows,
the anterior ones much the smaller. Spermatheca (R.s.) not in a good position,
the tail pointing obliquely towards the observer and therefore somewhat fore-
shortened ; body subglobular, a little broader than long in this position.
Length: g2mm., 93:5 mm. ; hindtibia: ¢ 0:4 mm., 9 0-7 mm.
Hab. Argentina: San Juan, May 1938, on Galea leucoblephara, one pair, type 3.
Named in honour of Dr. J. M. de la Barrera, to whom I am very grateful for
his additions to the National collection of Siphonaptera.
5. Parapsyllus andricus sp. nov. (text-figs. 263-265).
g. Although the single specimen is much damaged, it is worth describing,
as it represents a new species which is exceptional in the long thoracic bristles,
the very large number of pits in the sensilium, the length of segment I of the mid-
and hindtarsi, etc.
In front of eye an irregular row of 6 bristles, of which the upper 2 are small,
before the row a single moderately long bristle, below the eye 5 longish ones and
at the apex of the genal process 3 small ones. On occiput a long bristle behind
base of antenna, 2 in middle, one obliquely above the other, and a fourth farther
back ; a subapical row of 14 on the two sides together ; above antennal groove
the usual row of small bristles (here over 40). Bristles of antenna short. Length
of segments of maxillary palpus 12, 14, 10, 18. Proboscis reaching to apex of
trochanter ; length of segments of labial palpus 14, 12, 9, 13, 23.
Novirates ZOOLOGICAE XLI. 1939. 301
Bristles on thorax and abdomen longer than usual. The arrangement of the
20 bristles in the pronotal row different from that in other species ; on each side
a subdorsal row of 6 (text-fig. 263) which are placed close together, the bristles
as long as the occiput measured from base of antenna to subapical row of bristles,
in the dorsal interspace between the two sets of bristles two shorter and some very
small, the row continued downwards by 3 bristles, interspace between lowest and
next double the width of that between second and third, and this interspace
5 Vllt.
ee
double that between the third and fourth ; in front of the row a small dorsal
bristle and a larger ventral one. Bristles on mesonotum 9, 12, those in posterior
row not quite so long as on pronotum, but the subdorsal ones rather closer
together than is usual in the genus. On metanotum 8, 12, the posterior ones
shorter than on mesonotum and almost normal in position. On mesopleura 5,
on metepimerum 2, 2 bristles.
Bristles on abdominal tergites: I 6, 12; II 8,17; III 2,17; IV 2, 17;
V 0,18; VI0,17; VIL0, 16; none of the bristles of the anterior row are dorsal,
one of them placed below stigma, as are 2 of the posterior row. No lateral
bristles on basal sternum; subapical ones (the two sides together) on III 2,
IV to VI 4, VII 6.
302 NOVITATES Zoonocicak XLI. 1939.
The forecoxa bears about 24 bristles, not counting the small basal ones. On
inner side of midcoxa a submarginal row of 11 or 12 slender bristles ; the row
containing 1 or 2 more on hindcoxa. On outer side of midfemur a row of
(apparently) 8 and on inner side a row of 10 or 11 bristles; on hindfemur the
outside row contains 13 and the inside one 12 or 13. Dorsal margin of foretibia
(text-fig. 264) with 5 notches and in between third and fourth 2 stout bristles ;
on midtibia 2 notches near base, 3 close together in middle, and 2 at apex, with
3 stout bristles between median and apical notches ; on hindtibia the notches
nearly as on midtibia, but in the interspace between middle and apex 4 or 5 stout
bristles. Posteriorly at apex of midtibia 3 long bristles, one reaching far beyond
apex of tarsal segment I, the others to its apex ; on hindtibia the longest median
bristle as long as tarsal segment I, the longest apical one not quite reaching to
apex of I; 10 dorsolateral and lateral bristles on hindtibia, no subventral ones.
Segment V preserved only in foretarsus : twice as long as broad, the third pair of
plantar bristles much closer together than the first pair ; segment of hindtarsus
with 3 (or 4 ?) long apical bristles, the longest probably extending to or beyond
V; the distal segments of mid- and hindtarsi missing. Measurements: fore-
tarsus 12, 12, 9, 7, 10; midtarsus 30, 25; hindtarsus 60, 40. Claw slightly
asymmetrical ; basal tooth of inner claw slightly the larger.
Modified Segments.— g. Anterior margin of sternum VIII strongly chitinized,
appearing as a curved vertical ribbon ; ventral margin also more chitinized than
usual, with a rounded, strongly chitinized, hump below the base of the ventral
arm of IX.st.; at this point, on each side, a row of 5 bristles, of which 4 are
apically curved and recurved (text-fig. 264) ; the segment divided into two lobes
each side, one (L!) short, elongate, apically rounded, the other (L?) very large,
irregularly triangular, its upper angle pointed, the posterior margin first incurved
and then rounded-truncate, ventral margin nearly straight. Clasper (Cl) broadest
ventrally, gradually narrowing upwards, but remaining broad (P), the upper angle
being broadly rounded ; outer margin nearly straight from below apex to 4/5,
then obtusely angulate (pa) and slanting downward-forward from this point to
the ventral angle (va), which is about 90°, with the tip rounded off; ventral
margin incurved. The portion of the clasper above the ventral angle corresponds
to the spur of the preceding new species. Anterior (or dorsal) margin of clasper
with numerous bristles on the out- and inside ; at posterior margin a row of long
bristles beginning at some distance from apex with a submarginal one (missing,
like most bristles of the clasper); on outer surface about 20 long bristles.
Manubrium (M) much narrower than vertical arm of IX.st., widest at apex.
Digitoid (F) broadest at apex, much longer than broad, anteriorly concave,
posteriorly almost straight, and apically slightly convex ; posterior angle rounded
off, anterior one acute ; the sclerite projects a little above the highest point of the
clasper and bears at the posterior margin some slender bristles and at the anterior
one a row of very short hairs. Vertical arm of sternum IX slightly convex on
posterior side, concave anteriorly, the upper end rounded (outline of anterior
portion of dilated apex obscure in the specimen); ventral anterior angle of
vertical arm about 90°, not rounded ; angle formed by vertical and ventral arms
acute ; apical third of ventral arm slightly curved up, its apex rounded ; ventral
margin somewhat convex in middle and studded on and near the convexity with
about 10 bristles ; farther distally a hump which appears dentate and probably
bears bristles in a perfect specimen; a short distance from apex a small but
Noviratss ZOOLOGICAE XLI. 1939. 303
distinct nose ; on side of apical area several thin longish bristles. Sensilium of
pygidium larger than usual in Parapsyllus, with more than 40 pits on each side,
which is quite exceptional in this genus.
Length: 2-9 mm. ; hindfemur 0-67 mm.
Hab. Argentina: San Juan, May 1938, on Galea leucoblephara, 1 &.
SYNONYMICAL Norr.—According to the descriptions and figures Hectopsylla
mazzai Costa Lima, Rev. med.-cirurg. xliii, p. 251, figs. (1935), is the same as
H. broscus Jord. & Roths., Thomps., Yates & Johnst. Labor Rept. vii. 1, p. 60,
pl. 4, fig. 29 (1906), and Maailhopsylla Lillo: Schreitt & Shann., Bol. Mus. Hist.
Tucuman i, p. 6, fig. (1927) (on inner surface of ear of Pronops perotis Wied.,
Tucuman) is Rhynchopsyllus pulex Haller, Arch. Naturg. xlvi, p. 72, pl. 6
(1880).
EXPLANATION OF TEXT-FIGURES.
The lettering of the figures is explained in the text of the descriptions.
Fig. 256. Hectopsylla gemina, 2 j p. 292
257. Parapsyllus galeanus, kitodhensen, 3 p. 295
258. r Er claw of hindtarsus, 3. p- 295
259. 55 Ms genitalia of 3 p- 296
260. a > nf no p- 297
261. Parapsyllus barrerai, genitalia of & p- 299
262. e £ AO p. 300
263. Parapsyllus andricus, pronotum of & p. 301
264. = a foretibia . p. 301
265. 50 55 genitalia . p. 301
22
304 NOVITATES ZOOLOGICAE XLI. 1939,
NOTICE OF POSSIBLE SUSPENSION OF THE RULES OF
NOMENCLATURE IN CERTAIN CASES [A.(n.s.)l].
[X accordance with a Resolution adopted by the International Zoological
Congress at their Ninth Meeting held at Monaco in 1913, prescribing that
not less than one year’s notice is to be given by the International Commission
on Zoological Nomenclature of all applications received for the “‘ Suspension of
the Rules,” the attention of the zoological profession is hereby invited to the
fact that requests for the “‘ Suspension of the Rules” have been received by
the Commission in the undermentioned cases :
(a) ECHINODERMATA.— Diadema Humphreys, 1797 (type Echinometra setosa
Leske, 1778), to be added to the Official List of Generic Names (see Mortensen,
1937, Ann. Mag. Nat. Hist. (10), 19 : 463-469) (reference Z.N. (S.) 52).
(b) Insecta, Neuroptera.—To be added to the Official List of Generic
Names with types as shown in brackets : Hemerobius Linnaeus, 1758 (Hemerobius
humulinus Linnaeus, 1758) ; Chrysopa Leach, 1815 (Hemerobius perla Linnaeus,
1758) (see Cowley and others, 1937, Generic Names of British Insects, Pt. 4,
(reference Z.N. (S.) 42).
(c) IssectA, Lepidoptera.—To be added to the Official List of Generic
Names with the type as shown in brackets: Actinote Hübner, 1819 (Papilio
thalia Linnaeus, 1758) (see Hemming, 1936, Proc. R. Ent. Soc. Lond. (B), 5: 56-57)
(reference Z.N. (S.) 63).
(d) Repritia.—Bitis Gray, 1842 (type Vipera (Echidna) arietans B. Merrem,
1820), to be added to the Official List of Generic Names, and Cobra Laurent,
1768, to be suppressed (Stejneger, 1936, Copeza, 3 : 140) (reference Z.N. (S.) 121).
2. In adopting the Resolution referred to above, the International Zoological
Congress expressly stated that their object was thereby to render it possible for
zoologists, particularly specialists in the group in question, to present to the
Commission arguments for or against the suspension of the rules proposed.
Any such representations should be furnished to the Secretariat to the Commission
(British Museum (Natural History), Cromwell Road, London, S.W.7) as soon as
possible and in any case within one year of this day’s date. Every such com-
munication should be clearly marked with the Commission’s reference number
as given above.
By Order of the Commission,
(Signed) Francis HEMMING,
Secretary to the Commission.
SECRETARIAT OF THE COMMISSION,
British MUSEUM (NATURAL History),
CROMWELL RoAD, Lonpon, S.W.7.
27th June, 1939.
bo
EXPLANATION OF PLATE V.
Gobius ruthensparri heavily infected with C. lingua (note complete
absence of pigment around cysts).
Gobius ruthensparri heavily infected with C. jejund, showing much pig-
ment formation round cysts.
The tail of the specimen shown in Fig. 1.
The tail of the specimen shown in Fig. 2.
Tail of Wrasse infected with C. lingua, showing pigment formation round
cysts.
Cercaria of C. lingua (x 80) (permanent preparation).
Adult C. lingua (x 64) recovered from the intestine of laboratory-reared
Black-headed Gull.
Novirates Zootocice, Vou.XLI. 1939. P1.V,
JohnBale Sons & Curnow, Ltd London
Life-cycle of CRYPTOCOTYLE LINGUA (Creplin) 1825.
EXPLANATION OF PLATE VI.
HEADS OF ARABIAN GAZELLES, pp. 184-189.
The type of G. saudiya
. saudiya, Philby’s no. 132
. saudiya, Philby’s no. 33
. arabica, Philby’s no. A
. arabica, Philby’s no. 35
. arabica, Philby’s no. 123
AANNRNRN
. arabica, Philby’s no. 14
. arabica, Philby’s no. 12
. arabica, Philby’s no. 70
RRWR
. saudiya, Philby’s no. 29 ; female
R
. arabica, Philby’s no. 111; female
. muscatensis, BM. 94.3.9.8; female
. arabica, BM. 34.8.4.29
. marica, Philby’s no. 136
RRQ NR
The type of G. marica
G. muscatensis, BM. 94.3.9.7
The type of G. muscatensis
Gazelle from Aden
Gazelle from Aden
Novirates Zootocice, Vou. XLI.1939. Pl. Vi.
John Bale Sons & Curnow, Lt@ London.
ü h ae yon
v,
. ,
i
{
A Journal of Zoology.
EDITED BY
THE HON. MIRIAM ROTHSCHILD
and Dr. KARL JORDAN, F.RS.
Vor. XLI.
No. 4
Pages 305-463.
IssueD OCTOBER 31, 1939, ar THe ZooLosıcaL MUSEUM, TRING.
PRINTED BY HAZELL, WATSON & VINEY, Lrp., LONDON AND AYLESBURY.
1939.
NOVITATES ZOOLOGICAE.
10.
11.
12.
13.
14.
15.
Vor. XLI.
NOVITATES ZOOLOGICAE
EDITED BY
THE HON. MIRIAM ROTHSCHILD and KARL JORDAN.
CONTENTS OF NO. IV
. ECTOPARASITES FROM CAPTIVE BIRDS
. ON SOME NEARCTIC FLEAS . :
. NOTES ON SOME CELLULARINE POLYZOA
(BRYOZOA)
. POLYCHAETA OF THE “ROSAURA” EX-
PEDITION .
. NOTES ON ORIENTAL ZHECL INAE (LEP.
LYCAENIDAE) WITH DESCRIPTIONS OF
NEW. SPECIES :
. SIPHONAPTERA COLLECTED BY DR. GERD
HEINRICH IN BURMA .
. LARGE AND SMALL FLAME CELLS IN A
CERCARIA (TREMATODA)
„A PRELIMINARY REVISION OF THE PALAE-
ARCTIC SPECIES AND SUBSPECIES OF THI-
SOICETRUS BR. W. (ORTHOPTERA, ACRI-
DIDAB 6 ee Cee) eae ra
. ON SOME TROPICAL POLYCHAETA IN THE
BRITISH MUSEUM, MOSTLY COLLECTED BY
DR. C. CROSSLAND AT ZANZIBAR, TAHITI
AND THE MARQUESAS .
ON SOME TROPICAL POLYCHAETA IN THH
BRITISH MUSEUM, MOSTLY COLLECTED BY
DR. C. CROSSLAND AT ZANZIBAR, TAHITI
AND THE MARQUESAS .
THE RACES OF JAGUAR (PANTHERA onc):
DESCRIPTIONS OF ANTHRIBIDAE, MOSTLY
FROM FIJI
ON SOME OLD-WORLD LEPIDOPTERA HETER-
OCERA
ON SOME ANTHRIBIDAE FROM AFRICA AND
THE SOLOMON ISLANDS 5
ON RHOPALOPSYLLUS BAKER 1905 5 (SIPHON:
APTERA)
INDEX
PAGES
Theresa Clay and
Miriam Rothschild 305—315
Karl Jordan 316—320
Anna 'B. Hastings 321—344
C. C. A. Monro 345—3b4
N.D. Riley _ 355— 861
Karl Jordan 362—375
Miriam Rothschild 376
B. P. Uvarov 377—882
C. C. A. Monro 383-393
©. C. A. Monro 394—405
R. I, Pocock 406—422 -
Karl Jordan .. 423—432
Karl Jordan 433—436
Karl Jordan 437442
Karl Jordan 443—448
449—463
NOWVAEATEHS ZOOLOGICAE
Vol. XLI.
OCTOBER 1939. No. 4.
ECTOPARASITES FROM CAPTIVE BIRDS.
II. 1937-1938.
By THERESA CLAY, B.Sc., AND MIRIAM ROTHSCHILD.
COE SR ASTIES have again been collected from various birds dying in
captivity. The same procedure has been adopted as in 1936-37 (Clay
and Rothschild, Nov. Zool., XLI, p. 61, 1938). The Mallophaga! have again
been identified by Miss Theresa Clay and the Acarina by Mrs. A. M. Hughes.
As in the previous list the initials following each record, whether positive or
negative, indicate the collector who examined the bird in question, and the
figures in brackets after the name of the host refer to the number of specimens
received.
GENERAL REMARKS.
A total of 129 birds was examined, pertaining to 85 species and 27 different
families. Of this number 98 were confined in captivity with various species of
birds, but despite the opportunity thus afforded only one doubtful case of
“ straggling’ is recorded. The Crested Cariama (Cariama cristata) is probably
not the natural host of Goniocotes sp. and it may have acquired the infection from
some game bird with which it came into contact before arrival in this country
(see p. 312). Owing to the absence of negative data from the London Zoo,
and because from the hosts in question no mites were collected, it is more
convenient to consider these birds separately. Twenty-four species (26 specimens),
of which 11 were confined separately, yielded 30 species of Mallophaga. These
included two double and one treble infection. No very outstandingly heavy
infestations were noticed.
A total of 62 species of birds (104 specimens) were examined from aviaries
other than the London Zoo. Of these 23 species (28 specimens) were found to
be infected with ectoparasites. Twenty-two species (25 specimens) harboured
Mallophaga and 5 species Acarina. The former included 28 species and the
latter 7. Four double infections and 2 treble infections of Mallophaga and 2
double infections of Acarina were recorded, while 2 birds harboured both lice
and mites. It seems probable that a number of infections of mites were over-
looked on account of the rather poor state of preservation of some of the hosts.
The Buzzards again showed very heavy individual infections, but the most
striking example of great numbers of parasites was seen on the Stanley Crane.
Only the wings of this bird were examined, but these were crawling with thousands
1 Before attempting to describe any of these species a revision of several genera is necessary,
and for this purpose further materia] is required.
23 305
MAR 5 1940
306 Noyirates Zootocicar XLI. 1939.
of specimens of Esthioplerum gruis. Among them was a single specimen of
Heleonomus sp.? It was regrettable that the distribution of these parasites on
the rest of the body could not be determined.
A rather disappointing number of eggs of the Mallophaga were collected, but
those available for study tend to confirm the suggestion that, apart from their
morphological characters, the manner of attachment, position, and distribution
will prove of value for purposes of classification.
In our previous list the Cardinals had been shown as the most highly parasit-
ized group of birds, 7 out of 8 specimens being infected. Only 4 of these birds
were examined during 1937-38, of which 3 were parasitized. The Psittacidae
were again negative for Mallophaga, but two infections occurred among the
Waxbills which had previously been invariably negative.
Only 18 species (18 specimens) of these birds were confined separately or
with related species only. Of these Oxyphaps lophotes had been bred in captivity
and obviously acquired its parasite from the parents. Among the remaining
birds no case of straggling of Mallophaga is recorded. The evidence so far
obtained from the study of captive birds suggests that “ straggling ’ in Nature
must be a very rare occurrence.
LIST OF PARASITES.
I. SIPHONAPTERA (Fleas).
Parasite.
Host.
No records.
II. MALLOPHAGA (Biting Lice).
Menoponidae.
*Trinoton querquedulae ' (Linn.) 1758
Menopon gallinae (Linn.) 1758
Menopon tumidum Piaget 1885
Menopon ridibundum Denny 1842
Menopon sp.? .
Menopon sp. ?
Amyrsidea sp. ?
Myrsidea sp.? .
Menacanthus sp. ?
Menacanthus sp. ?
Menacanthus sp. ?
Menacanthus sp.? . : 5
Kurodaia alternatus (Osborn) 1902
Erdmaniella brevipalpe (Piaget) 1880. .
Colpocephalum kelloggi Osborn 1902
Colpocephalum sp. ?
Colpocephalum sp. ? .
Colpocephalum sp. !
Philacte canagica (Sewast.) 1802.
Gallus lafayetii Lesson 1831.
Sarkidiornis melanota (Pennant) 1769.
Larus r. ridibundus (Linn.) 1758.
Bubo bubo (Linn.) 1758.
Turtur chalcospilos (Wagler) 1827.
Pavo muticus Linn. 1766.
Passerina leclancheri (Lafr.) 1840.
Estrilda melpoda (Vieillot) 1817.
Estrilda subflava (Vieillot) 1819.
Temenuchus pagodorum (Gmel.) 1789.
Steganura paradisea (Linn.) 1758.
Cathartes aura falklandica (Sharpe)
1873.
Phalacocorax a. aristotelis (Linn.) 1761.
Cathartes aura falklandica (Sharpe)
1873.
Circaétus g. gallicus (Gmel.) 1788.
Haliastur 1. indus (Boddaert) 1783.
Haliaétus I. leucocephalus (Linn.) 1766.
! The species indicated by a star (*) show slight variation which may ultimately prove to be
subspecific.
Novirates ZOOLOGICAE XLI. 1939. 307
Heleonomus sp. ? : : : . Anthropoides paradisea (Licht.) 1793.
Heleonomus sp. ? 5 A : . Balearica p. pavonia (Linn.) 1758.
Ricinidae.
Ricinus fringillae Degeer 1778 . . Fringilla c. coelebs Linn. 1758.
Philopteridae.
Goniodes pavonis (Linn.) 1758 . . Pawo cristatus Linn. 1758.
Goniodes parviceps Piaget 1880 . . Pavo muticus Linn. 1766.
Goniodes eurygaster Piaget 1885 . . Lophophorus wmpejanus (Latham)
1790.
Coloceras sp. ? . k : j . Tympanistria tympanistria (Tem. and
Knip) 1808.
Coloceras sp.? . : . Turtur chalcospilos (Wagler) 1827
Goniocotes haplogonus Nitzsch 1866 . Lophophorus impejanus (Latham)
1790.
Goniocotes sp. ? Diardigallus diardi (Bonaparte) 1856.
Goniocotes sp. ? : : : . Lophura rufa (Raffles) 1822.
Goniocotes sp. ? 5 : k . Oxyphaps lophotes (Temminck) 1822
Goniocotes sp. ? Cariama cristata (Linn.) 1766.
Lagopoecus ne (Mégnin) 1880 . Lophophorus impejanus (Latham)
1790.
Otilipeurus turmalis (Denny) 1842 . Otis t. tarda Linn. 1758.
* Degeeriella fusca (Denny) 1842 . . Buteo buteo (Linn.) 1758.
Degeeriella numenii (Denny) 1842 . Numenius a. arquata (Linn.) 1758.
Degeeriella punctata (Burmeister) 1838 . Larus r. ridibundus (Linn.) 1758.
Degeeriella sp. ? ; : ; . Falco columbarius aesalon Turnstall
1771.
Degeeriella sp. ? 5 : : . Butastur teesa (Franklin) 1831.
en Be. m | } Pluvianus a. aegyptius (Linn.) 1758.
Bruéliasp.? . : ; 2 . Junco h. hyemalis (Linn.) 1758.
Bruelia sp.? . ; . Paroaria cristata (Boddaert) 1783.
Rallicola lugens (Giebel) 1874 F . Porphyrio poliocephalus (Latham)
1790.
Crasspedorrhynchus platystomus (Bur-
meister) 1838 R 5 : . Buteo buteo (Linn.) 1758.
Crasspedorrhynchus sp. n. . : . Uroaétus audax (Latham) 1807.
Cummingsiella testudinarius (Denny)
1842 . ‘ 6 : : . Numenius a. arquata (Linn.) 1758.
Saemundssonia sp. ? . Larus r. ridibundus (Linn.) 1758.
Eustrigiphilus a Da) 1842 Nyctea nyctea (Linn.) 1758.
Anatoecus sp. ? : : . Bucephala islandica (Gmel.) 1789.
Anatoecus sp. ? Nyroca nyroca (Guldenstadt) 1769.
Anatoecus sp. ? a : E . Cereopsis n. hollandiae Latham 1801.
Anatoecus sp. ? 5 : . Branta b. bernicla (Linn.) 1758.
* Hsthiopterum gruis (iain. ) 1758 . . Anthropoides paradisea (Licht.) 1793.
Anaticola subsignatum (Giebel) 1866 . Phoenicopterus antiquorum Temminck
1820.
Columbicola sp. ? 5 : 5 .. Turtur chalcospilos (Wagler) 1827.
308 NOVITATES ZoontocicaE XLI. 1939.
Falcolipeurus marginalis (Osborn) 1902 Cathartes aura falklandica (Sharpe)
1873.
Struthiolipeurus rheae Harrison 1916 . Rhea americana albescens Lynch 1878.
III. ACARINA (Mites).
Laelapidae.
Hypoaspis ferulis Mégnin 1876 . . Pluvianus a. egyptius (Linn.) 1758.
Liponyssidae.
Liponyssus sylvarum Canestrina and
Fanzago 1877 : 5 0 . Bubo lacteus (Temminck) 1824.
Liponyssus sp. ? 6 5 2 . Richmondena cardinalis (Linn.) 175°8
Tyroglyphidae.
Tyroglyphus farinae Koch 1841 . . Melopsittacus undulatus (Shaw) 1905.
Glycyphagidae.
tlycyphagus domesticus Degeer 1778 . Pluvianus a. egyptius (Linn.) 1758.
Falculigeridae.
Protolichus lunula Robin and Mégnin
Sr: 5 ; 5 ; . Melopsittacus undulatus (Shaw) 1905.
Proctophyllodidae.
Joubertia microphylla Robin and Mégnin
NSW. : : : : . Fringilla c. coelebs Linn. 1758.
LIST OF HOSTS.
Rheidae.
RHEA AMERICANA ALBESCENS Lynch 1878, South American Rhea (1).
London Zoo,! August 1938. Obtained from South America. Confined in a
paddock with various species of birds. Heavy infection of Struthiolipeurus
rheae. (Z.) ®
Phalacrocoracidae.
PHALACROCORAX A. ARISTOTELIS (Linn.) 1761, Shag (2). Dudley Zoo,
3.8.38 and 5.8.38. One caught in Great Britain (Birmingham). Confined
with other Shag purchased in Holland. High infection of Hidmaniella brevipalpe.
(R.1.)
Ardeidae.
ARDEA CINEREA Linn. 1758, Common Heron (2). Dudley Zoo, 5.11.37
and 21.9.38. Locality of capture unknown. Confined with Gulls and Pelicans.
Wings only examined. No ectoparasites. (M. R.)
EGRETTA ALBA (Linn.) 1758, Great White Egret (1). Lilford Aviaries,
13.5.38. Probably obtained from Central Asia. Not confined with other birds
during several years at Lilford. No ectoparasites. (E. M.)
1“ London Zoo ” refers to the Zoological Society of London, Regent’s Park.
2M. R. = Miriam Rothschild, E. M. = Elizabeth Meyerhof, R. I. = Ruth Italiener, Z. = Staff
of the Prosectorium, The Zoological Society of London, Regent’s Park.
Novirates ZooLoGIcAaE XLI. 1939. 309
EGRETTA GARZETTA (Linn.) 1766, Little Egret (1). Dudley Zoo, 30.12.37.
Locality of capture unknown. Confined with various other birds. No ecto-
parasites. (E. M.)
Phoenicopteridae.
PHOENICOPTERUS ANTIQUORUM Temminck 1820, Flamingo (1). London
Zoo, December 1938. Obtained from Egypt. Confined with other Flamingoes
and Ducks. Light infection of Anaticola subsignatum. (Z.)
Anatidae.
BUCEPHALA ISLANDICA (Gmel.) 1789, Barrow’s Golden-eye (1). London
Zoo, December 1938. Obtained from Iceland. Confined with other Ducks and
Waders. Light infection of Anatoecus sp.? (Z.)
TADORNA TADORNA (Linn.) 1758, Common Sheldrake (2). Dudley Zoo,
13.10.37 and 26.11.37. Locality of capture unknown. Confined with various
birds. No ectoparasites. (M. R.)
MARECA PENELOPE (Linn.) 1758, Wigeon (1 chick). Dudley Zoo, 3.7.38.
Bred in captivity. Confined with various birds. No ectoparasites. (M. R.)
SARKIDIORNIS MELANOTA (Pennant) 1769, India Comb Duck ¢ (1). Lilford
Aviaries, 24.1.38. Obtained from India. Confined with Great White Egrets.
A fairly heavy generalized infection of Menopon tumidum. No eggs noted.
(E. M.)
Nyroca nyroca (Guldenstadt) 1769, Ferruginous Duck (1). London Zoo,
December 1938. Obtained from India. Confined with other Ducks. Medium
heavy infection of Anatoecus sp.? (Z.)
METOPIANA PEPOSACA (Vieillot) 1818, Rosy-billed Pochard g (1). Dudley
Zoo, 26.1.38. Locality of capture unknown. Confined with various species
of Duck. No ectoparasites. (E. M.)
Atx sponsa (Linn.) 1758, Carolina Duck ¢ (1). Dudley Zoo, 26.1.38.
Locality of capture unknown. Confined with various species of Duck. No
ectoparasites. (EH. M.)
BRANTA B. BERNICLA (Linn.) 1758, Brent Goose (1). Lilford Aviaries,
30.1.38. Captured in Lincolnshire, England. Confined separately. Three
specimens of Anatoecus sp.? No eggs noted. (E. M.)
CEREOPSIS N. HOLLANDIAE Latham 1801, Cape Barren Goose (1). Dudley
Zoo, 19.2.38. Locality of capture unknown. Confined with various species
of birds. Light general infection of Anatoecus sp.? No eggs noted. (E. M.)
NEOCHEN JUBATA (Spix) 1825, Oronoco Goose ¢ (1). Lilford Aviaries,
4.3.38. Locality of capture unknown. Four years in captivity without contact
with other birds. No ectoparasites. (EH. M.)
PHILACTE CANAGICA (Sewast.) 1802, Emperor Goose (1). London Zoo,
August 1938. Obtained from Siberia. Confined with various species of birds.
Very light infection of *Trinoton querquedulae (Linn.) 1758. (Z.)
Cathartidae.
CATHARTES AURA FALKLANDICA (Sharpe) 1873, Falkland Turkey Buzzard (1).
London Zoo, February 1938. Obtained from Falkland Isles. Confined with
same species only. A light infection of Falcolipeurus marginalis and Colpocepha-
lum kelloggi, and a heavy infection of Kurodaia alternatus. (Z.)
310 Novirares Zoonoeicat XLI. 1939-
Accipitridae.
Burro BUTEO (Linn.) 1758, Common Buzzard (2). Dudley Zoo, 12.10.37
and 9.2.38. Locality for the first bird unknown, the second bird captured at
Oxford, England. Confined with Vultures, Eagles and Storks. Both heavily
infected with *Degeeriella fusca. Eggs confined to the surface of the basal half
of lower wing coverts. The second bird with two specimens of Crasspedorrhynchus
platystomus on the head. (M. R. and E. M.)
CIRCAETUS G. GALLICUS (Gmel.) 1788, Short-toed Eagle (1). London Zoo,
February 1938. Obtained from South Africa. Confined separately. Heavy
infection of Colpocephalum sp.? (Z.)
HALIASTUR 1. Inpus (Boddaert) 1783, Brahminy Kite (1). London Zoo,
December 1938. Obtained from India. Confined separately. Heavy infection
of Colpocephalum sp.? (Z.)
HALIAETUS L. LEUCOCEPHALUS (Linn.) 1766, Bald Eagle (1). London Zoo,
December 1938. Obtained from U.S.A. Confined separately. Heavy infection
of Colpocephalum sp. (Z.)
Uroaktus AUDAX (Latham) 1801, Wedge-tailed Eagle (1). London Zoo,
August 1937. Obtained from Australia. Confined with other Eagles. Heavy
infection of Crasspedorrhynchus sp.n. (Z.)
Falconidae.
FALCO COLUMBARIUS AESALON Turnstall 1771, Merlin (1). London Zoo,
December 1938. Obtained from Europe. Confinedseparately. Heavy infection
of Degeeriella sp.? (Z.)
Cracidae.
Mirv mirvu (Linn.) 1766, Razor-billed Curassow (1). Dudley Zoo, 21.10.37.
Locality of capture unknown. Wings only examined. Confined with various
species of birds. No ectoparasites. (M. R.)
Phasianidae.
DIARDIGALLUS DIARDI (Bonaparte) 1856, Siamese Crested Fireback (2).
London Zoo, December 1937 and July 1938. Obtained from Siam. Confined
with other Firebacks. Both heavily infected with Goniocotes sp. (Z.)
LoPHURA RUFA (Raffles) 1822, Vieillot’s Fireback (1). London Zoo, Decem-
ber 1938. Obtained from Siam. Confined separately. Light infection of
Goniocotes sp. % (Z.)
LopHOPHORUS IMPEJANUS (Latham) 1790, Impeyan Pheasant (1). London
Zoo, December 1938. Obtained from the Himalayas. Confined separately.
Heavy infection of Goniodes eurygaster, a medium heavy infection of @oniocotes
haplogonus and a light infection of Lagopoecus heterotypus. (Z.)
Pavo cristatus Linn. 1758, Peafowl (1). London Zoo, January 1938.
Obtained from India. Confined separately. Light infection of Goniodes pavonis.
(Z.)
Pavo muticus Linn. 1758, Burmese Peafowl (2). London Zoo, January
and February 1938. Obtained from Burma. Confined separately. Both
heavily infected with Amyrsidea sp.? (Z.)
Novrratrs Zoonoaicam XLI. 1939. 311
GALLUS LAFAYETIL Lesson 1831, Ceylon Jungle Fowl (1). London Zoo,
December 1938. Obtained from Ceylon. Confined with other Jungle Fowls.
Medium heavy infection of Menopon gallinae. (Z.)
ACRYLLIUM VULTURINUM (Hardwicke) 1834, Vulturine Guinea Fowl (1).
Lilford Aviaries, 9.3.38. Captured in Somaliland. Six years in captivity in a
large-sized aviary with two other species of Guineafowl. No ectoparasites.
(M. R.)
Gruidae.
ANTHROPOIDES PARADISEA (Licht.) 1793, Stanley Crane (1). Dudley Zoo,
December 1937. No data. Wings only examined. Very heavy infection of
* Hsthiopterum gruis and one specimen of Heleonomus sp. ? Noeggs found. (M.R.)
BaLEARICA P. PAVONTA (Linn.) 1758, Crowned Crane (1). London Zoo,
December 1938. Obtained from West Africa. Confined with other Cranes.
Light infection of Heleonomus sp. (Z.)
Rallidae.
AMAURORNIS PHOENICURUS (Pennant) 1769, White-breasted Water-Hen (1).
Lilford Aviaries, 13.12.37. Imported from India. Confined in a large aviary
with various birds for four years. No ectoparasites. (KH. M.)
PoRPHYRIO P. POLIOCEPHALUS (Latham) 1801, Grey-headed Gallinule (1).
London Zoo, December 1938. Obtained from India. Confined with various
species of birds. Heavy infection of Rallicola lugens. (Z.)
Cariamidae.
CARIAMA CRISTATA (Linn.) 1766, Crested Cariama (1). London Zoo, Decem-
ber 1938. Obtained from South America. Confined separately. Medium heavy
infection with Goniocotes sp.? This species is most probably a straggler, but at
present nothing definite is known concerning the Ischnoceran Mallophaga of the
Cariamidae. (Z.)
Otidae.
OTIS T. TARDA Linn. 1758, Great Bustard (1). London Zoo, January 1938.
Obtained from Hungary. No data. Medium heavy infection of Otilipeurus
turmalis. (Z.)
Glareolidae.
PLUVIANUS A. AEGYPTIUS (Linn.) 1758, Egyptian Plover (3). Foxwarren
Park, 8.12.37 and 4.5.38, and London Zoo, August 1938. Captured in Egypt
and the Gold Coast. Confined with various species of birds. The first specimen
with a heavy infection of Degeeriella sp. n. and Degeeriella sp. and the second with
only slight infections of Degeeriella sp. n., Glycyphagus domesticus and Hypoaspis
fenilis. The eggs of Degeeriella were found principally on the breast feathers.
Several eggs on one feather, confined to the posterior half, and fixed near
the central shaft parallel to the barbules. The specimen from the Zoo with a
medium heavy infection of Degeeriella sp.n. (Z., E.M. and M. R.)
Scolopacidae.
NUMENIUS A. ARQUATA (Linn.) 1758, Common Curlew (1). Lilford Aviaries,
2.3.38. Captured in Lincolnshire, England. Confined with various species of
312 Novirarrs Zoonocicar XLI. 1939.
birds in large enclosure. A fairly heavy infection particularly on the wings of
Degeeriella numenii and Cummingsiella testudinarius. No eggs noted. (E. M.)
CALIDRIS C. CANUTUS (Linn.) 1758, Knot (1). Lilford Aviaries, 1938. Cap-
tured in Lincolnshire, England. Kept in an aviary with various other waders
for about three years. No ectoparasites. (E. M.)
Laridae.
LARUS R. RIDIBUNDUS (Linn.) 1758, Black-headed Gull (1). Ashton Wold,
5.9.38. Captured in Warwickshire, England. Confined alone. General but
light infection with Menopon ridibundum, Degeeriella punctata and Saemundssonia
sp.? No eggs noted. (M. R.)
LARUS A. ARGENTATUS Pontopp 1763, Herring Gull (1). Dudley Zoo,
21.9.38. Reared in captivity. Confined with Herons and Pelicans. Wings
only examined. No ectoparasites. (M. R.)
Columbidae.
GEOPELIA CUNEATA (Latham) 1801, Diamond Ground Dove (2). Dudley
Zoo, 25.11.37. Catpured in Australia. Confined with various species of birds.
No ectoparasites. (KE. M.)
TYMPANISTRIA TYMPANISTRIA (Temm. and Knip) 1808-11, Tambourine Dove
(1). Lilford Aviaries, 26.11.37. Obtained from Africa. Confined separately
for a month, and for a time previously with other doves. A light infection of
Coloceras sp.? No eggs noted. (E. M.)
ZENAIDA A. AVIATICA (Linn.) 1758, White-winged Dove (1). Lilford Aviaries,
18.2.38. Bred in captivity. Confined with various species of doves. No
ectoparasites. (EK. M.)
OXYPHAPS LOPHOTES (Temminck) 1822, Australian Crested Dove (1). Lil-
ford Aviaries, 3.6.38. Bred in captivity. Not confined with other species of
birds. A heavy infection with Goniocotes sp. ? principally confined to the breast
feathers. No eggs noted. (M. R.)
TURTUR CHALCOSPILOS (Wagler) 1827, Emerald-spotted Dove (1). Lilford
Aviaries, 6.9.38. Locality of capture unknown. Confined with six other species
of doves. A general light infection with three species of Mallophaga, Coloceras
sp. ?, Columbicola sp.? and Menopon sp.? Of the two latter only immature
specimens were found. No eggs noted. (E. M.)
Psittacidae.
MELOPSITTACUS UNDULATUS (Shaw) 1805, Budgerigar (12). Dudley Zoo,
1937-1938. Green, Yellow and Blue varieties. Probably all bred in this country.
With one exception all confined with various birds. One specimen with light
infection of T'yroglyphus farinae and Protolichus lunula. (K.M., R. I. and M. R.)
AGAPORNIS FISHERI (Reichenow) 1887, Fisher’s Lovebird (1). Dudley Zoo,
4.12.37. Obtained from Africa. Confined with various birds. No ecto-
parasites. (M. R.)
AMAZONA AESTIVA (Linn.) 1758, Blue-fronted Parrot (1). Dudley Zoo,
12.11.37. Locality of capture unknown. Confined with various species of
birds. No ectoparasites. (E. M.)
Novirates Zoonoaicar XLI. 1939. 813
AGAPORNIS PULLARIA (Linn.) 1758, Red-headed Lovebird (1). Dudley Zoo,
16.4.38. Locality of capture unknown. Confined with same species only. No
ectoparasites. (E. M.)
K.AKATOE ROSEICAPILLA (Vieillot) 1817, Roseate Cockatoo (1). Dudley Zoo,
25.10.37. Locality of capture unknown. Confined separately. No ecto-
parasites. (E. M.)
ARA MARACANA (Vieillot) 1816, Llliger’s Macaw (1). Lilford Aviaries,
27.1.38. Locality of capture unknown. Confined in a large aviary with other
species of parrots. No ectoparasites. (E. M.)
Bucerotidae.
TROPICRANUS ALBOCRISTATUS (Cassin) 1848, White-crested Hornbill (1).
Dudley Zoo, 23.1.38. Locality of capture unknown. Confined with
Lophoceros semifasciatus. No ectoparasites. (E. M.)
LOPHOCEROS FLAVIROSTRIS LEUCOMELAS (Lichtenstein) 1842, Yellow-billed
Hornbill (1). Dudley Zoo, 8.7.37. Locality of capture unknown. Confined
separately. No ectoparasites. (E. M.)
Strigidae.
Buso BUBo (Linn.) 1758, Great Eagle Owl (1). Dudley Zoo, 24.10.37.
Locality of capture unknown. Confined separately. Wings only examined.
One specimen of Menopon sp.? (E.M.)
Buso LAcTEUS(Temminck) 1824, Milky Eagle Owl (1). Dudley Zoo, 24.10.37.
Locality of capture unknown. Confined separately. Wings only examined.
Heavy infection of Liponyssus sylvarum. (E. M.)
NYeTEA nycTEA (Linn.) 1758, Snowy Owl (1). London Zoo, January 1938.
Obtained from Greenland. Confined separately. Medium heavy infection of
Eustrigiphilus ceblebrachys. (Z.)
Turdidae.
CossYHA NIVEICAPILLA (Lafres) 1838, Snow-headed Robin-chat (1). Dudley
Zoo, 22.11.37. Obtained from Africa. Confined separately. No ectoparasites.
(E. M.)
SIALIA MEXICANA Swanson 1832, Mexican Blue-bird (1). Foxwarren Park,
20.3.38. From “ Pacific States.” Confined separately. No ectoparasites.
(E. M.)
KITTACINCLA MALABARICA INDICA (Baker) 1924, Indian Shama (1). Fox-
warren Park, 6.6.38. Obtained from India. Confined with various species of
birds. No ectoparasites. (M. R.)
Timelidae.
LIOTHRIX LUTEA (Scopoli) 1786, Pekin Robin (3). Dudley Zoo, 27.9.37
and 6.12.37. Locality of capture unknown. The first pair confined with various
birds, the third alone. No ectoparasites. (E. M. and M. R.)
LIoPTILA CAPISTRATA (Vigors) 1831, Blue-headed Sibia (1). Foxwarren
Park, 8.12.37. Bred in captivity. Confined with various species of birds. No
ectoparasites. (M. R.)
314 Novrrares ZooLocIcae XLI. 1939.
ARRULAX ALBOGULARIS (Gould) 1835, White-throated Jay-thrush (1).
Dudley Zoo, 12.7.38. Locality of capture unknown. Confined with Garrulax
leucolophus. No ectoparasites. (M. R.)
Ploceidae.
EUPLECTES FRANCISCANA (Isert) 1789, Orange Bishop (3). Dudley Zoo,
18.10.37 and 21.2.38. Locality of capture unknown. Confined with various
species of birds. No ectoparasites. (E. M. and M. R.)
ENODICE MALABARICA (Linn.) 1758. Indian Silver-bill (1). Dudley Zoo,
27.9.37. Exact locality of capture unknown. Confined separately. No ecto-
parasites. (M. R.)
Monta striata (Linn.) 1766, Japanese Munia (1). Dudley Zoo, 27.4.38.
Bred in captivity. Confined with various species of birds. No ectoparasites.
(E. M.)
ORYZIVORA ORYZIVORA (Linn.) 1758, Java Sparrow (7). Dudley Zoo, 1937—
1938. Locality of capture unknown. Confined with various species of birds.
No ectoparasites. (E.M. and M. R.)
AMADINA FASCIATA (Gmel.) 1789, Cut-throat Finch (4). Dudley Zoo,
13.2.38, 12.3.38 and 23.6.38. Exact locality of capture unknown. Confined
with various species of birds. No ectoparasites. (E. M.)
TAENIOPYGIA CASTANOTIS (Gould) 1837, Zebra Finch (1). Dudley Zoo,
25.10.37. Exact locality of capture unknown. Confined separately. No
ectoparasites. (M. R.)
ESTRILDA ASTRILD SANCTAEHELENAE Shelley 1903, St. Helena Waxbill (3).
Dudley Zoo, 18.10.37 and 6.11.37. Locality of capture unknown. Confined
with various species of birds. No ectoparasites. (M. R.)
ESTRILDA SUBFLAVA (Vieillot) 1819, Zebra Waxbill (1). Dudley Zoo,
4.12.37. Locality of capture unknown. Confined with various species of birds.
A light general infection with Menacanthus sp.? (M. R.)
ESTRILDA MELPODA (Vieillot) 1817, Orange-cheeked Waxbill (3). Dudley .
Zoo, 20.10.37, 6.11.37 and 3.12.37. Obtained from West Africa. Confined
with various species of birds. Two birds with a light general infection of
Menacanthus sp.? No eggs noted. (E. M.)
URAEGINTHUS BENGALUS (Linn.) 1766, Red-cheeked Cordon Bleu (3).
Dudley Zoo, 25.11.37 and 23.6.38. Locality of capture unknown. Confined
with various species of birds. No ectoparasites. (EK. M. and M. R.)
STEGANURA PARADISEA (Linn.) 1758, Paradise Wydah (2). Dudley Zoo
4.12.37. Exact locality of capture unknown. Confined with various species
of birds. Very light infection of Menacanthus sp.? No eggs noted. (E. M.)
Sturnidae.
TEMENUCHUS PAGODORUM (Gmel.) 1789, Pagoda Starling (2). Dudley Zoo,
3.12.37 and 13.2.38. Exact locality of capture unknown. Confined with
same species only. One of the birds with heavy general infection of Mena-
canthus sp. Eggs laid at the base of the breast feathers, several eggs on each
feather. (E.M. and M. R.)
STURNUS VULGARIS (Linn.) 1758, Common Starling (1). Dudley Zoo.
23.6.38. Obtained in England. Confined with various species of birds. No
ectoparasites. (M. R.)
or
Noviratrrs Zoonocicar NLI. 1939. 31
Fringillidae.
PASSERINA LECLANCHERI (Lafres) 1840, Rainbow Bunting (1). Dudley
Zoo, 5.6.38. Locality of capture unknown. Confined separately. Light
infection of Myrsidea sp.? No eggs noted. (E. M.)
JUNCO H. HYEMALIS (Linn.) 1758, Slate-coloured Junco (1). London Zoo,
March 1938. Obtained from Newfoundland. Confined with various species of
birds. A light infection of Bruélia sp. (Z.)
FRINGILLA C. COELEBS (Linn.) 1758, Chaffinch (3). Dudley Zoo, 8.3.38.
Captured near Dudley, England. Confined with various species of birds. One
specimen with light infection of Ricinus fringillae, and Joubertia microphylla on
the primaries. The eggs of Ricinus fringillae were laid 1-3 at the base of the
breast feathers. (M. R.)
RICHMONDENA CARDINALIS (Linn.) 1758, Virginian Cardinal (2). Dudley
Zoo, 13.10.37 and 2.12.37. Exact locality of capture unknown. One speci-
men confined with various species of birds, the other alone. The first with a
light infection of Liponyssus sp.? (M. R. and E. M.)
PAROARIA CRISTATA (Boddaert) 1783, Red-crested Cardinal (1). Dudley
Zoo, 25.11.37. Exact locality of capture unknown. Confined with various
species of birds. A very light infection of Brueliasp.? Noeggsnoted. (M. R.)
PAROARIA DOMINICANA (Linn.) 1758, Dominican Cardinal (1). Dudley Zoo,
24.10.37. Exact locality of capture unknown. Confined separately. No ecto-
parasites. (M. R.)
Trochilidae.
THALURANIA FURCATOIDES (Gould) 1849, Amazonian Wood-nymph (1).
Foxwarren Park, 5.8.38. Exact locality of capture unknown. Confined
separately. No ectoparasites. (M. BR.)
ACKNOWLEDGMENTS.
We would again like to express our gratitude to all those who have so kindly
co-operated with us in preparing this list, in particular to Mrs. A. M. Hughes for
identifying the mites, and Mr. J. D. MacDonald for correcting the host names.
Our special thanks are also due to the Earl of Dudley, Mr. Alfred Ezra, Lord
Lilford, Mr. R. W. Partridge, and the Zoological Society of London for permission
to examine the birds dying in their aviaries; to Dr. H. Honigman, Mr. A. FE.
Moody and Mr. A. P. Shearing for the immense trouble they have taken in
dispatching the material and supplying data, and to Miss Elizabeth Meyerhof,
Miss R. Italiener and the staff of the prosectorium of the Zoological Society of
London for examining birds in the authors’ absence.
316 Novirares ZooLoGIcAE XLI. 1939.
ON SOME NEARCTIC FLEAS.
By KARL JORDAN.
(With 6 text-figures.)
1. Opisodasys robustus Jordan 1925 (text-fig. 266).
HE species was originally described from 99 only. In a small series of the
species lately received there is a male, of which we here figure the genitalia.
The tergite VIII is large as in the allied species and bears at the upper portion
of the posterior margin a row of 9 long bristles and at the ventral margin 10 or 11,
266
LE fo “_VIILst.
it st.
of which several are exceptionally long ; behind the stigma the segment forms a
sharp angle directed upwards ; the posterior marginal area of this projection is
densely studded with teeth on the inner side. Sternum VIII (text-fig. 266) quite
small, its apical half lanceolate, with a shortish bristle at the top and a smaller
one on the side. Process P of the clasper nearly straight on the frontal side and
gently rounded on the posterior one, not quite attaining the tip of the digitoid F.
NOVITATES ZooLoGIcAE XLI. 1939. 317
This sclerite F apically obliquely truncate, the anterior angle being lower than the
posterior one ; apart from its pedicel the sclerite is broad, its anterior margin
being straight and the posterior margin slightly incurved and then farther upwards
somewhat rounded, the sclerite becoming gradually narrower at apex ; oblique
apical margin about one-fourth as long as the ventral margin ; posterior ventral
angle rounded, prominent; near this angle a large spiniform, below middle of
posterior margin a smaller spiniform, both of these bristles sharply pointed, at
about two-thirds a third spiniform of a very different shape, its basal third being
broad and the rest obliquely curved upwards and gradually narrowed to a sharp
point. Ventral arm of sternite IX characteristic : its proximal half with hardly
any bristles, whereas the distal half bears numerous short pale spiniform ones
along the ventral margin (the proximal half of the series are on the inner side
of the long lobe which projects forward and are indicated in the figure by
dotted lines).
We are much indebted for this series to Professor R. A. Flock, who obtained
the specimens in the Catalina Mts., Arizona, on Sciurus arizonensis catalina.
2. Orchopeas sexdentatus cascadensis subsp. nov. (text-fig. 267).
Based on the 9. The range of variability in the females from the western
districts of Oregon is most instructive. Whereas in O. sexdentatus agilis Roths.
1905, from Alberta and British Columbia, sternum VII of the abdomen varies
from being strongly rounded, with a more or less distinct indication of a sinus, to
being deeply sinuate, with a long obtuse process above the sinus, in O. s. cascadensis
267
the process is always sharply pointed and varies from being long to being absent ;
text-fig. 267 illustrates this variability, the six figures representing selected
specimens obtained at the same locality on the same host. If sufficient specimens
are examined the disappearance of the process is found to be gradual. In
O. s. agilis the rounded segment is ancestral ; in O. s. cascadensis, on the contrary,
the segment without process is the most advanced modification.
Oregon: Odell Lake, on Neotoma fuscipes fuscipes, 31. vil. 36, a series
(type) ; from other western places also on Neotoma cinerea occidentalis. Collected
by Professor C. A. Hubbard.
Phaneris gen. nov.
9. Near Rectofrontia Wagner 1930, of which it is a very interesting and, for
the understanding of relationship, important modification. The genal spines of
318 Novirates ZoOLOGICAE XLI. 1939.
Rectofrontia and short spiniforms on the inner surface of the hindcoxa are lost.
The bristles of the abdomen are very numerous and long.
Frontal tubercle below middle as in R. fraterna Baker 1895; genal margin
expanded ventrad, incurved between maxilla and forecoxa, a rounded lobe being
formed, which is rather obscured by the maxilla lying under it ; no vestige of eye.
Bristles of thorax as compared with those of abdomen short and few ; apical
area of meso- and metanota short and strongly chitinized, especially that of
metanotum ; episternum of metathorax longer than in Rectofrontia, very oblique
(text-fig. 268, est) ; upper anterior angle of metasternum acute ; metepimerum
with patch of distinct striae (striarium) as in Rectofrontia and allied genera.
Apical spines of abdomen longer than in Rectofrontia. Genotype: Ph. hubbardi
sp. nov.——Atriarium diagrammatical in fig. 268.
3. Phaneris hubbardi sp. nov. (text-figs. 268, 269).
9. Onfrons2long bristles, one of them on incrassate margin of antennal groove,
the other some distance from ventral margin, farther forward a row of 5 small
ones, of which the one near antennal groove is the longest and strongest. On
occiput three rows, containing each
5 bristles on both sides. Maxillary
palpus reaching to end of forecoxa,
268 da Win a little longer than proboscis, the
ZANE latter with curved bristle at apex.
- Bristles on thorax : on pronotum 15
in one row on the two sides together ;
on mesonotum 11 or 12 and in front
of this row about 15 more or less
irregularly scattered ; on metanotum
8, 12. On mesopleura 5, on mete-
pimerum 3 ; on mesonotum two sub-
dorsal spines each side on under
surface, these spines rather large, not
like bristles in shape.
Many of the bristles of cn
not gradually thinning to a long fine
point, but remaining rather stout to
near end, which is suddenly narrowed
to a point as in a straight sword.
Apical spines, the two sides together :
on 7 4: 11° 42:7 ih 2 Vie Vee
Bristles more irregularly placed than
is usual, the posterior row of the
tergites very oblique, numbers in two
specimens: 1 9 or 18, 12 or 19; II 19
or 22, 19 or 20; IIL 30 or 24, 20;
IV 25 or 26, 19 or 18; V 23 or 25, 17 or 18; VI 19,18; VII 27 or 26, 19 or 20
(text-fig. 269, A). Some of the bristles in front of the posterior row as large as
the posterior bristles, and those in between the long ones of the row longer than
usual ; on terga II to VI 3 or 4 (2 on one side of one segment) and on VII
4 or 5 below the stigma; lowest bristle of VI as long as hindtibia. Bristles
mt
Noviratres Zoonoaican XLI. 1939. 319
on sterna: II 7, 12; IV 7,15; V 10,15; VI 9, 14; VII 20, 17 in paratype,
more in type figured (text-fig. 269, B).
Legs inclusive of coxae somewhat slenderer than in R. fraterna ; chaetotaxy
nearly the same, the longest bristles somewhat longer, the longest of hindtarsal
segment I reaching well beyond II, and that of II much beyond IV ; proportional
length of tarsal segments
as in R. fraterna.
Side of sternum VII
(text-fig. 269, B) with
rounded ventral sinus, the
lobe above the sinus broad
andround. Apical dorsal
angle of widened portion
of tergum VIII projecting
as a lobe, which is different
in size and shape in the
two specimens. Hump of
anal tergite indistinct ;
bristles in proximal area
of anal sternum more
numerous than m AR.
fraterna. Spermatheca
(R.s.) smaller than in that
species, almost gradually
and but slightly narrowed
from posterior end for-
ward.
Length 3-1mm., hind-
femur 0-4 mm.
Oregon: Springwater,
20. vu. 31, on Aplodontia
rufa, 1 9 type, returned
to Professor Hubbard ;
5.iv.36, on the same host,
1 Q retained for the
National collection.
One of the many dis-
coveries of Professor C. A. Hubbard, from whom we may expect a publication on
Oregon fleas in the near future. I have much pleasure in associating his name
with this remarkable species.
4. Leptopsylla hamifer longiloba subsp. nov. (text-fig. 270).
9. Two 29 from Alaska sent to me for description by Mr. Wm. L. Jellison,
Assistant Parasitologist at the Rocky Mountain Laboratory, Hamilton, Montana,
agree closely with L. hamifer Roths. 1906, as stated by Mr. Jellison in litt., but
differ considerably in sternum VII of the abdomen. As this segment is the same
in L. h. hamifer from Alberta and L. h. vigens Jord. 1937 from Montana, whereas the
dd of these two subspecies are distinguishable by the genitalia, we may expect
that the & of the new flea will, like the 9, be more conspicuously different from
5
320 NOVITATES ZooLÖGICAE XLI. 1939,
N j
L.h. hamifer thanisthe Sof L.h.vigens. I describe the new flea‘as a subspecies of
L. hamifer in order to emphasize the relationship. The $ may prove this flea to
represent a distinct species.
In L. h. hamifer and L. h. vigens the sinus of VII. st. is not very deep and
the lobe above it triangular, less than half as long as its distance from the ventral
margin of the segment, sometimes more pointed than in text-fig. 271 (from an
271
270
Alberta specimen) ; in the new subspecies the sinus is much deeper, the margin
below it more oblique and the lobe more than twice as long as in the other sub-
species ; the shape of the lobe varies a little in the two specimens as well as on
the two sides of the body. Chaetotaxy as in the other subspecies, but in both
specimens only one long bristle below the stigma of VIII. t. In type 7 ante-
pygidial bristles on both sides, in the paratype 5. Some of the bristles of VII. st.
very stout.
Alaska: 1 mile camp, Valdez Creek trail, 3,500 ft., 23 September 1937,
1 2 off Microtus sp., type, in collection of the Rocky Mountain Laboratory ;
Fairbanks, 11 September 1937, on “ Rabbit,” 1 2 paratype in the National
collection at Tring.
EXPLANATION OF TEXT-FIGURES.
The lettering of the figures is explained in the text of the descriptions.
Fig. 266. Opisodasys robustus, genitalia of . : : . p. 316
267. Orchopeas sexdentatus cascadensis, sternum VII of Orn ing . p. 317
268. Phaneris hubbardi, metathorax and first abdominal tergum . p. 318
269. Ke Eu A, dorsal area of abdominal terga VII
and VIII. B,sternum VII. . . p. 319
270. Leptopsylla hamifer longiloba, sternum VII. : 5 . p. 320
271. Leptopsylla hamifer hamifer, the same : : : . p. 320
hi ace oe |
CORRIGENDUM.
ER
Novitates Zoologiicae Vol. M (4): 321-344. Hastings: Cellularine ‘Polyzoa.
The scales gi i figures are divi i i
ae S given with the figures are divided into tenths and twentieths of a
The decimal poin+ has been omitted in Figs. 272, 275, 276 a-n, 277, 278, 279 a
279c. ‘The seales for Fiyrs. 276
: R us. 276 F and 279 B each represent a whole milli i
are correct as printed, i i a a
Novirarres Zoonoargan XLI. 1939. 321
NOTES ON SOME CELLULARINE POLYZOA (BRYOZOA).
By ANNA B. HASTINGS,
Department of Zoology, British Museum (Nat. Hist.).
(With 8 text-figures.)
FE Polyzoa discussed in this paper have been examined in the course of
my work on the collections of the Discovery and other antarctic expedi-
tions, but do not come within the scope of my unpublished report.
_ The genus Emma and a section of the genus Bugula are revised, with de-
scription of new species, and examination of various recorded specimens ; the
Challenger material of Caulibugula has been re-examined, and a new species of
‘the genus described from New Zealand material collected by the Discovery ;
Dimorphozoum Levinsen is shown to be a synonym of Beania. Bugula expansa
sp. n., a New Zealand species from the Terra Nova collection, has a curious
structure at the base of the colony, which I have referred to as the foot. Its
morphology could probably only be made out by examining developmental
stages.
The classification of the Polyzoa has not yet reached a point where sub-
species, forms, races, etc., can be adequately discriminated, and I therefore use
the term variety in a wide sense to cover all such categories.
In the statements of distribution of the species a published record is indi-
cated by the author’s name, the reference being given in the statement of
synonymy. Specimens in the British Museum are indicated by their registered
number, sometimes with the name of the collector or donor added ; specimens
from the Waters Collection in the Manchester Museum are indicated by “‘ Man-
chester Mus.”, and in the Liverpool University Museum by “ Liverpool Mus.”’.
“ Terra Nova ” means British Antarctic Expedition (1910), and “ Discovery ”
the Discovery Investigations (1925 onwards).
I am very grateful to the Riksmuseum, Stockholm, the University Museums
of Zoology at Berlin, Cambridge and Liverpool, and the Manchester Museum
for lending specimens, and to Dr. C. Crossland for his material from Ghardaga,
Red Sea.
I should also like to thank Sir Sidney Harmer, K.B.E., F.R.S., for his help
and encouragement.
Emma Gray, 1843.
My examination of type and other authentic specimens of Emma in the
British Museum shows that three species have been confused under EZ. crystallina,
and that Waters’ variety of E. cervicornis is distinguished by quite definite
characters.
All the species of Emma have branches that spring from the frontal surface
of the zooecia and originate in a uniserial joint, in addition to those formed by
bifurcation, which, except in H. cyathus, are biserial from the start. In my
descriptions I have called them frontal branches.
Waters (1887, p. 88) mentioned the Straits of Magellan in the distribution
of E. crystallina, on the evidence, no doubt, of a specimen in the Busk collection
24
i)
l
322 NOVITATES ZOOLOGICAE XLI. 1939.
(99.7.1.672), which proves, however, to belong to E. rotunda. Busk received
the specimen from Miss Gatty, from whom he obtained material from many
parts of the world. The genus being otherwise recorded solely from Australia
and New Zealand, and being absent from the S. American collections of the
Discovery Expedition, I hesitate to accept the evidence of this one slide, although
there is nothing in the appearance of the slide, nor in its history as far as known,
to throw any doubt on the correctness of its locality.
KEY TO THE SPECIES OF EMMA.
1 if All joints uniserial . : a E. cyathus (not discussed here 1)
| Joints at bifurcation biserial, tere at origin of frontal branches uniserial 2.
2 [ All internodes of 3 zooecia ; : ; 5 Bo
ı Non-fertile internodes of 2 zooecia ea a Bienen En be ud
Scuta simple on non-fertile, De or forked on fertile zooecia, 1 or 2 spines
33 in axil 5 B : : N . 6. E. tricellata
Scuta clavate, larger on Bereit zooecia, 2 to 4 spines in axil . 7. E. buskia
[ Opesia roundly triangular, the largest spine opposite its apex, an internal
spine for attachment of parietal muscles, fertile internodes of 2 zooecia
with 1 or 2 ovicells not immersed in zooecia (? in kenozooecia), scuta only
| found on fertile zooecia, placed near ovicell . ‘ . 2. E. triangula
Opesia semicircular or oval, large spines not placed in definite relation to its
symmetry, no internal spine, fertile internodes ? of 3 zooecia, with 1 ovicell
which is immersed in a zooecium, scuta, when present, placed near proximal
end of opesia ; : 3 5 : = : : ; ED:
Lateral avicularia paired (i.e. one on each zooecium of internode), their palatal
surface facing outward, no scuta, no frontal avicularia 1. E. erystallina
Lateral avicularia paired, or single (i.e. only one to an internode), or absent,
palatal surface facing more or less towards apex of branch, frontal avicularia
and scuta present (sometimes with very limited distribution in colony) 6.
paired when present, aperture oval, 5 or 6 distal spines . Br: 7l;
Scuta only on fertile zooecia, unbranched, lateral avicularia single except on
fertile internodes, aperture round, 3 or 4 distal spines . 3. E. rotunda
Lateral avicularia present on some internodes at least, internodes tapering
quickly, frontal avicularia absent except on fertile internodes, larger spines
often pod-like 6 ; : 3 4. E. cervicornis
Lateral avicularia absent, Antsaasıks sigh tubular proximal portion, no
frontal avicularia on fertile internodes but sometimes present on median
L
| Scuta present on most zooecia, cervicorn or forked, lateral avicularia usually
| zooecium at bifurcation, spines not pod-like 5. E. cervicornis var. watersi.
1. Emma crystallina Gray (text-fig. 272, C).
Emma crystallina Gray 1843, p. 293; Harmer 1923, p. 357 (part).
Distribution.—New Zealand (Gray ; 35.2.28.2, the type specimen).
The zooecia of the type specimen of E. crystallina are turned away from
the axis of the branch, so that a line bisecting the opesia forms a greater angle
1 See Harmer, 1923, p. 357. 2 Unknown in E. crystallina.
Novirates Zoonogican XLI. 1939. 323
with the main axis of the branch than it forms in 1. rotunda, where the turning
is less marked (see angle a in text-figs. 272, Cand B). The opesia is almost semi-
circular. In all but the lowest internodes of the branch there is a small lateral
avicularium on each zooecium, with the palatal surface nearly parallel to the
longitudinal axis of the branch, thus facing outward. . The two outer spines of
the distal series of 4 or 5 on each zooecium are conspicuously larger than the
Text-ric. 272.—A. Emma triangula sp.n., an internode from the type specimen, 87.12.10.44b.
s, spine with parietal muscles. Internodes without rootlets are more nearly symmetrical.
B. Emma rotunda sp.n., an internode from Hooker’s specimen from Campbell Is., 99.7.1.670.
C. Emma crystallina Gray, an internode from the type specimen, 35.2.28.2. D. Emma cervi-
cornis MacG., an internode from 99.5.1.334. One of the lateral avicularia is broken. E. E.
cervicornis var. watersi var. n., an internode from the type specimen, 88.1.2.3.
rest. The position of the enlarged spines bears no constant relation to the
symmetry of the opesia. The cryptocyst of the median zooecium at the
bifurcations is very oblique. Scuta are absent and the specimen has no ovicells.
I have seen no other specimen that can be attributed to this species.
Busk (1852b) confused two species, both of which appear to be distinct from
E. crystallina Gray, and are discussed below (2. E. triangula, 3. E. rotunda).
MacGillivray also used the name E. crystallina for E. rotunda, and Levinsen
used it for #. triangula.
2. Emma triangula sp.n. (text-fig. 272, A).
Emma crystallina Busk, 1852a, p. 373; 18525 (part), p. 28, pl. XL, figs. 1-3 (Rattlesnake) ;
Harmer, 1923, p. 357 (part).
Menipea crystallina Busk, 1884, p. 23; Levinsen, 1909, pp. 132, 133, pl. II, figs. la, 1b.
Not Emma crystallina Gray.
Not Menipea crystallina MacGillivray (= B. rotunda sp.n.).
Distribution.—Port Phillip (Busk ; 87.12.10.37 and 440; 88.11.14.271
and 372; 97.5.1.282 and 283); St. 161, Challenger (Busk; 87.12.9.102);
324 NOVITATES ZOOLOGICAE XLI. 1939.
Bass Strait (Busk ; 99.7.1.5738, Rattlesnake ; 34.2.16.10; 1938.12.14.1);
Palliser Bay, Wairarapa, New Zealand (1938.1.20.2); Cape Maria Van Diemen,
New Zealand, 35-40 fms., 13.1x.1911 (Terra Nova St. 144, 1939. 2.2.4);
34° 11’ S., 172° 8’ E., New Zealand, 84 m., 17.vii.1932 (Discovery St. 935;
1939.2.2.6); 34° 11’ S., 172° 10’ E., New Zealand, 92-98 m. (Discovery St.
934, 1939.2.2.7); New Zealand (Terra Nova, 1939.2.2.3).
Type.—87.12.10.44b.
This species was well figured by Levinsen, who showed the ovicells, and by
Busk. It resembles typical E. crystallina in its avicularia, but the opesia is
roundly triangular rather than semicircular. One or two spines in the distal
series of 3 (or occasionally 4) are enlarged, and the largest is placed symmetrically
opposite the apex of the triangle. Where there are 3 spines this is the middle
one. The other spines are small and pointed, but not placed on the basal surface
as in Busk’s figure 3. The cryptocyst of the median zooecium at the bifurcation
is oblique as in E. crystallina. The outline of the aperture varies, sometimes
being nearly symmetrical as in my figure, more often drawn out at the proximal
corner as in Busk’s figure 2.
The parietal muscles form two groups, one in each proximal corner of the
opesia (text-fig. 272, A). One bundle is attached to a little thickening of the
outer wall, the other to a blunt-ended spine (s), which springs from the transverse
wall separating the distal zooecium, and frequently projects beyond the edge
of the cryptocyst so that its head is quite conspicuous in frontal view.
As far as my observations go, the fertile internodes are always on the frontal
branches. They consist of two zooecia, both of which usually bear ovicells,
though sometimes only one is fertile. Levinsen described the ovicells as im-
mersed in kenozooecia. The fertile zooecia commonly bear a spine-like scutum
placed near the ovicell.
Busk’s material from Bass Straits, both in the Rattlesnake and Challenger
collections, belongs to this species. His specimens from New Zealand (B.M. Cat.)
belong to E. rotunda. It is impossible to tell to which species the New Zealand
specimens of Livingstone (1929, p. 55) belonged.
3. Emma rotunda n.sp. (text-fig. 272, B).
Menipea crystallina MacGillivray, 1881, p. 31, pl. LVIII, figs. 2-25.
Emma crystallina Busk, 1852b, p.28 (part, New Zealand) ; Harmer, 1923, p. 357 (part) ; Hastings,
in Cranwell and Moore, 1938, p. 395.
Not Emma crystallina Gray.
Distribution. —Queenschff (MacGillivray) ; Poor Knights Is., New Zealand
(Hastings; Miss L. B. Moore, 1937.4.6.2); Campbell Is. (1939.4.22.1:;
Hooker, 99.7.1.670); New Zealand (1938.5.2.2; 99.7.1.671); Great Swan
Point, Tasmania (46.8.5.18) ; Tasmania (Manchester Mus.) ; New South Wales
(1938.9.2.1); Bondi Bay, New South Wales (Manchester Mus.) ; ? Straits of
Magellan (99.7.1.672, see p. 321 above).
Type.—1937.4.6.2.
This species, described and figured by MacGillivray as E. crystallina, differs
from E. crystallina Gray in the shape of its zooecia. They are less turned from
the median axis of the branch (text-fig. 272, B), the aperture is circular rather
than oval and the granular cryptocyst, whose proximal part descends steeply,
1 1939.4.22.1. This is the British Museum Catalogue specimen (51.1.10.12) re-registered.
Noviratrrs Zoonoarcar XLI. 1939, 325
appears of fairly even width all round. The opesia is semicircular. In dried
specimens the orifice is sometimes seen to be stretched open, forming the circular
hole in the frontal membrane figured by MacGillivray. The cryptocyst of the
median zooecium at the bifurcation is much less oblique than in H. crystallina.
Simple spine-like scuta are sometimes present on the fertile zooecia, and there
may be a frontal avicularium on the median zooecium at the bifurcations. Many
internodes have no lateral avicularia, but there is often one on the more distal
of the two zooecia of an internode. The lateral avicularium projects more from
the lateral wall of the zooecium than in Z. crystallina and, its position appearing
to be related to the angle of the zooecia, its palatal surface faces obliquely distally
instead of straight outwards. Non-fertile internodes do not have more than
one avicularium, but in fertile internodes the fertile zooecium, which is the first
zooecium of the internode, may also have one. It is a little larger than the
avicularium on the second zooecium, and is placed more frontally. The ovicell
is immersed in the third zooecium of the internode.
The difference between this species and E. triangula is clearly shown in the
figures of MacGillivray and Levinsen. The fertile internodes, in particular, are
markedly different (see key).
E. rotunda resembles E. cervicornis in the structure of its fertile internodes,
except that there is no frontal avicularium. It also resembles H. cervicornis in
the shape and direction of the lateral avicularia, but differs in their being single
on non-fertile internodes, H. cervicornis usually having two to an internode.
E. cervicornis further differs in its cervicorn scuta, and in the absence of frontal
avicularia at the bifurcations. E. cervicornis usually has 5 or 6 spines, of which
2 or 3 may be pod-like. The number in Z. rotunda is 3 or 4, of which 1 to 3 may
be large, but not pod-like. The little outer spine directed frontally, which is
common in E. cervicornis (text-fig. 272, D, E), is usually absent in Z. rotunda,
but is present in the Tasmanian material.
One of the two specimens from Campbell Is. (99.7.1.670) is typical (text-
fig. 272, B). The other approaches E. cervicornis var. watersi in having the
internodes rather long and tubular proximally, and no lateral avicularia. It
differs from the variety in the shape of the aperture and cryptocyst, in the
number and arrangement of the spines, and in the larger zooecia, in all of which
it agrees with H. rotunda. It is labelled “ Emma crystallina Gray MS.” in Gray’s
writing, and Busk has also labelled it #. crystallina, but it is clearly distinct from
both the type specimen of that species and from the species figured in the B.M.
catalogue as E. crystallina (= E. triangula).
4. Emma cervicornis MacG. (text-fig. 272, D).
Emma cervicornis MacGillivray, 1869, p. 127; Harmer, 1923, p. 357.
Menipea cervicornis MacGillivray, 1881, pp. 34, 32, pl. LVIII, figs. 4-45; Levinsen, 1909, pp. 59,
132, 133, pl. II, figs. 4a, 4b.
Distribution —Queenscliff (MacGillivray); Australia (99.5.1.334; Man-
chester Mus. from E. C. J. [Jelly]) ; Port Phillip (87.12.10.36).
There is also a specimen from MacGillivray, without locality, 97.5.1.256.
Typical E. cervicornis, as represented in MacGillivray’s figures and by one
of his specimens (97.5.1.256), has an oval or semicircular opesia. There are
two lateral avicularia on most internodes except the lowest in the branch, where
there may only be one, but they are larger and more prominent than those of
326 Novirarrs ZooLoGIcAE XLI. 1939.
E. crystallina and their palatal surface faces more distally so that their beaks
project more from the branch. Frontal avicularia are only found in relation to
ovicells. There are 5 or 6 distal spines, the 6th when present being the outer one,
which is very small, is not in line with the others and is directed frontally
(text-fig. 272, D, and MacGillivray, fig. 4b). The other 5 are graded in size, the
outer two being the largest and often being pod-like. There is sometimes a small
spine on the inner border of the aperture. In the fertile internodes the first of
the three zooecia is fertile and its ovicell is immersed in the third, whose
avicularium surmounts the ovicell. The scutum of the fertile zooecium is apt
to be particularly richly branched.
The variety recognized by Waters appears to be clearly distinguished (see
below).
5. Emma cervicornis var. watersi var.n. (text-fig. 272, E).
Menipea cervicornis var. Waters, 1887, p. 88, pl. IV, fig. 1.
Distribution.—Shark Is., Port Jackson (Waters; Manchester Mus. and
88.1.2.3, parts of the type material) ; Port Phillip Heads (88.11.14.415); off
Port Phillip, 33 fms. (99.7.1.669) ; Kermadec Isles (55.12.7.175).
Waters’ variety resembles H. cervicornis in possessing branched scuta.
Many are, however, simply forked. The general distribution of its spines is
similar, but there are not more than 5, and they are usually slender. Var.
watersi resembles H. rotunda in the presence of a frontal avicularium on the
median zooecium at the bifurcation (not seen in 88.11.14.415) and in the
absence of frontal avicularia with the ovicells. It differs from both #. cervicornis
and EH. rotunda in the complete absence of lateral avicularia, and in its smaller
zooecia with longer tubular proximal portion.
6. Emma tricellata Busk.
Emma tricellata Busk, 1852a, p. 373; 18525, p. 28, pl. XLI, figs. 1 and 2; Harmer, 1923, p. 357.
Menipea tricellata MacGillivray, 1881, p. 34, pl. LVIII, figs. 5-5d.
Distribution. —Bass Strait, 45 fms. (Busk; 99.7.1.6520 Rattlesnake, the
type specimen) ; George Town, Tasmania, Hooker (54.11.15.71; 99.7.1.729,
730 and 5576); Queenscliff (MacGillivray) ; Port Phillip (88.11.14.258, 346 ;
97.5.1.253, 254, 275-277, 279 and 281); Sydney, 10 fms. (99.7.1.729A) ;
New Zealand, Hooker (Busk ; 99.7.1.731 and 732).
There is also a specimen from MacGillivray without locality (97.5.1.278).
The chief differences between H. tricellata and E. buskii (p. 327) can be
seen in MacGillivray’s figures, namely the longer internodes and simple scuta of
E. tricellata. In both, the scutum is more developed on the fertile zooecium.
In E. buskii this development takes the form of an enlargement of the clavate
head (cf. Levinsen, fig. 3a). In E. tricellata the scutum is notched or forked.
in contrast to the simple spine-like scuta of other zooecia, or, in colonies with
few scuta, it is present as a simple spine on the fertile zooecium and absent
from its neighbours. EB. tricellata has, on the average, a smaller number of
distal spines on the median zooecium at the bifurcation. Stout rootlets are
present in both, and I have been unable to satisfy myself that there is any
constant difference in the position of the avicularium. The fertile internodes
are similarly constructed in the two species.
E. tricellata, as separated by these characters, is rather variable. At one
extreme the specimens have internodes very little longer than those of H. buskit,
Noviratrs Zoonoaican XLI. 1939. 327
the median zooecium at a bifurcation has a scutum with a blunt and slightly
expanded end, suggesting the clavate scutum of H. buskii, the non-fertile zooecia
have usually 4 outer and 1 inner distal spine as in E. buskii. The median
zooecium usually has 4 spines, of which two are apical and show in the axil, but
it may resemble H. buskii, which generally has 5, with 3 apical. MacGillivray’s
figure represents a specimen approximating to this type. In one such specimen
many of the scuta have a slightly wider tip (88.11.4.346).
At the other extreme are specimens with longer, more tapering, internodes.
There is usually 1 apical spine in the axil (sometimes 2) and the axillary zooecium
has a simple, spine-like scutum. Some of the non-fertile zooecia are without
scuta, and they have only 2 or 3 outer spines of which not more than one is large.
Busk’s figure, drawn from his Tasmanian material, which is variable, approxi-
mates to this type, and so does the type specimen. The material from Port
Phillip forms a series connecting these two types.
At one extreme A. tricellata approaches H. buskii, whose geographical range
falls within the larger area occupied by £. tricellata, and it may well be that
E. buskii is no more than the extreme term in the variation of E. tricellata. The
two forms are, however, distinguishable in the material before me.
7. Emma buskii (Wyv.-Thom.).
Menipea buskii Wyville-Thomson, 1858, p. 144, pl. XII, fig. 1; MacGillivray, 1881, p. 35, pl. LVIIL
figs. 6-65.
Menipea buski Levinsen, 1909, pp. 131-133, pl. II, figs. 3a-c.
Emma buskii Harmer, 1923, p. 357.
Distribution —Bass Str. (Thomson ; Thomson, 99.7.1.660); Queenscliff
(MacGillivray); Port Phillip Heads (87.12.9.121; 97.5.1.251 and 252);
‘Hobson Bay, Australia (99.7.1.661 and 658).
This species is discussed under ZL. tricellata.
Dimorphozoum Levinsen, a synonym of Beania (text-fig. 273, A).
The genotype of Dimorphozoum Levinsen (1909, pp. 96, 107) is Flustra
nobilis Hincks (1891, p. 288), described from specimens obtained by Jelly from
Port Elizabeth, S. Africa. Levinsen introduced his genus after examination of
material from the same source. He described it as consisting of two layers back
to back, a cheilostome-layer and a ctenostome-layer. Specimens from Port
Elizabeth given to the Cambridge Museum ! by Miss Jelly appear to be part of
the same material ; the two layers are present and, as with Levinsen’s specimens,
there are colonies of Chaperia capensis. The ctenostome-layer contains numerous
brown bodies. Levinsen regarded it as an integral part of the colony, chiefly
because it was present throughout and was connected with the zooecia by
rosette-plates. This layer is, however, not present on all the colonies in the
Cambridge material, nor does it cover the whole basal surface of those on which
it is found, moreover, small patches of it are present at two points on the frontal
surface, where they obliterate the underlying zooecia. The supposed rosette-
plates prove to be minute basal spines resembling those of Flustra echinata
Kluge (1914, p. 658) but very much smaller, and usually unbranched. They are
developed irrespective of whether the ctenostome-layer is present or not, and
in Hasenbank’s material (1932, p. 335) they were branched.
1 Part of this material is now in the British Museum, registered 34.4.6. 1.
Noyrratrs ZOOLOGICAE XLI.
Texr-ria. 273.—A. Beania nobilis (Hincks), Locality unknown, 91.10.16.1. nat.
size. B. Beania bilaminata (Hincks), New Zealand, 99.7.1.6614. nat. size.
C. Bugula neritinoides Busk MS. sp.n. 99.7.1.4650. x 2.
1939.
Novirarrs ZoonoaicaAn XLT. 1939. 329
There is thus no reason for supposing that the ctenostome-layer is anything
but an incrustation of another species. As all the material is dry it is probably
impossible to identify the species, but the two-lipped orifice appears to relate it
to Flustrella and Elzerina (see Harmer, 1915, p. 39), rather than to Aleyonidium,
The cheilostome-layer, which is the whole zoarium of Hincks’ species, agrees
very closely with the description and figures given by Hasenbank, who found no
trace of a ctenostome-layer. It resembles Beania in the tubular connections of
the zooecia, and their shape. The small tooth-like spines on the distal edge, the
branched marginal spines, the basal spines, the pedunculate avicularia, the
differentiated operculum, the very shallow ovicells and the flustrine colony are
all to be found in species of Beania. Text-fig. 273, B, shows, for example, the
flustrine colony of Beania bilaminata (Hincks, 1881, p. 157), a species closely
related to B. magellanica Busk. The attachment of the avicularia to the proximal
part of the zooecium is unusual in Beania, but is found in Beania regularis
Thornely. Dimorphozoum thus becomes a synonym of Beania.
In material in the British Museum (Text-fig. 273, A) marginal spines are
exceedingly rare, but a few are present. No basal spines are present and the
basal surface is free from the encrusting Ctenostome. The ovicells can only
partially accommodate the large embryo which projects into the cavity of the
zooecium.
Bugula Oken, 1815.
I have examined a number of species of Bugula (see key below), including
three which I believe to be undescribed, which agree in having pedunculate,
obliquely placed, globular ovicells, no true spines, though the distal corners of
the zooecia may be pointed, the avicularium when present (it is absent in
B. neritina and B. neritinoides) placed on the proximal part of the zooecium, and
zigzag lines on the basal surface as described in B. robusta by Harmer (1926,
p- 436). These lines are not always visible throughout the colony, but can be
seen on the older parts, at least, in all the species. Most of these species have
brownish pigment which persists in spirit, and often also in dried specimens.
Traces of it can be seen in B. vectifera and B. subglobosa, and the only form in
which I have found no sign of it is B. robusta var. capensis, which I have not seen
in spirit. Some of the species have a short stalk consisting of more or less
elongated zooecia, but this is not a constant feature of the group. If the type
of bifurcation, as defined by Harmer (1926, p. 433), is considered, these species
fall into two groups as follows :
Bifureation 5. B. robusta MacG., B. robusta var. capensis Waters,
B. neritina (Linn.).
Bifurcation 4. B. neritinoides (Busk MS.) sp.n., B. subglobosa Harmer,
B. vectifera Harmer, B. scaphoides Kirk., B. minima Waters, B. crosslandi sp.n.,
B. expansa sp.n.
In B. neritina the connection of zooecia G and H at the bifurcation is some-
times rather obscure, owing in part to a projection from the side of F into the
proximal part of G, which I take to be a swollen rosette-plate. The other
features of the bifurcation are of characteristic type 5, the split reaching the
distal end of E and the fork of H passing down the side of F into the axil, from
which it separates F.
B. uniserialis Hincks (see Hastings, 1930, p. 705) possesses several of the
characteristics of this group. It differs chiefly in its bifurcation, which is of
330 Novirarrs ZooLoGIcAE XLI. 1939.
type 3, and in having a long tubular proximal portion to its zoocia. Preserved
specimens are colourless. The method of attachment of its colonies is unknown.
KEY TO THE SPECIES OF BUGULA CONSIDERED IN THIS PAPER.
1 If Bifurcation of type 5 5 R © B 6 3 ; 5 0%
| Bifurcation of type 4 : : : : e ‘ 6 : . 4.
5 [ Avicularia absent . ; : : : : : ; B. neritina
Avicularia present . : i : : : 5 é : =, 82
3 [ Avicularia long-headed? . i 5 0 : 3 B. robusta
Avicularia short-headed . : ; B I, B. robusta var. capensis
[ Colony robust, ovicells larger than zooecia, avicularia absent
4 2. B. neritinoides
Colony delicate, ovicells smaller than zooecia, avicularia present . 5 &
5 { Ovicells reticulate. : : : 5 : . 3. B. scaphoides
Ovicells smooth ; i : : ; ; ; : : 6:
6 J Avicularia short-headed . 6 6 : 6 ; : 5 Sewanee
| Avicularia long-headed . : 3 é : : : : 8.
[ Avicularia attached to proximal expansion of zooecium, colony with foot
6. B. expansa
dst 5 3 6 : :
Avicularia on proximal part of zooecium, but distal to proximal expansion,
| attachment of colony unknown . 4 f : . B. subglobosa
8 { Avicularia almost level with proximal end of opesia . : 5 N
Avicularia proximal to opesia . 5 : : : 5. B. crosslandi
[ Colony delicate straggling, avicularia small, beak forming less than half their
9 length, ovicell with narrow border . 6 : : B. vectifera
Colony relatively stout and compact, onflanllodta large, beak forming more
| than half their length, ovicell with broad border . . 4. B. minima
1. Bugula robusta var. capensis Waters (text-figs. 274, D, E, 275 A, B).
Bugula capense Waters, 1887, pl. IV, fig. 17 (mandible).
Bugula robusta (B. capensis Busk MS.) Waters, 1909, p. 137, footnote.
? Bugula neritina var. tenuata Thornely, 1912, p. 142, pl. VIII, fig. 3.
Distribution.—Cape of Good Hope (Busk Coll., 99.7.1.118, 119, 120, 305,
306) ; Port Elizabeth (89.1.1.8; 99.5.1.1347 ; Manchester Mus. ; Jelly Coll.,
Cambridge Mus.) ; Grahamstown (Manchester Mus.) ; ? Cargados (Thornely).
This variety chiefly differs from typical B. robusta (MacGillivray, 1869,
p. 129; Harmer, 1926, p. 435) in its avicularia, which have a more rounded
head and a much shorter beak without cusps (cf. text-fig. 275, B, C). The
colony when dry is a pale yellowish colour, or almost white, showing no signs of
pigmentation, and the branches, which in typical B. robusta are straight, are
curved and rather straggling in appearance. The zigzag line and joints (text-fig.
274, D) are as in B. robusta, but basal calcareous processes have not been seen.
1 Comparison of text-fig. 275, B and C, or text-fig. 277, A and B, will make clear the sense
in which I use the terms long-headed and short-headed.
Novirarrs Zoonoaican XLI. 1939. 331
The zooecia are similar except that the outer distal corners of the inner zooecia
above the bifurcations are rounded. As the pointed distal corners have a
tendency to curve forward, the zooecia without them look flatter frontally, and
where the internodes are short this makes the whole branch look flat compared
with typical B. robusta. When an avicularium is present on one of the axillary
zooecia it is smaller and rather different in shape (text-fig. 275, A), having no
TeExT-FIG. 274.—Tracings of hitherto unpublished pencil drawings by Busk :
Bugula neritinoides sp.n. (Busk MS.). A, Frontal view ; the upper bifurcation is the more typical.
B, Frontal view with ovicell. C, Basal view.——D, E. Bugula robusta var. capensis Waters.
D, Basal view. EH, Frontal view. The figures were drawn to the same scale, indicated by Busk
as “‘2 inch.” According to the average of several measurements this represents a magnification of
23.5, now reduced to ec. 17.5.
A, B, C.
concavity in its profile at the beginning of the beak. The ovicells are like those
of B. robusta.
Waters’ first use of the name Bugula capense [sic], for a figure of a mandible.
is almost a nomen nudum. He later published Busk’s manuscript name, Bugula
capensis, in a footnote, without formal description, but with indications that it
possessed the specific characters of B. robusta, and that it came from S. Africa.
All the S. African specimens of B. robusta that I have examined show the varietal
differences enumerated above, and there is no evidence that typical B. robusta is
ever found there. It may perhaps be assumed from this that the 8. African
specimens on which Waters based his remarks belonged to the variety, and if so
the name capensis Waters would be available for the variety. Examination
of the specimens in the Waters Collection confirms this assumption.
332 NoVITATES ZOOLOGICAE XLI. 1939.
Although not known from S. Africa, typical B. robusta is found on the
E. African coast, for examination of Waters’ specimens confirms Harmer’s opinion
(1926, p. 435) that B. neritina var. minima Waters (1913, p. 471, but not 1909, see
below) is a synonym of B. robusta. On the other hand, the specimens recorded
as B. robusta by Waters on the same page are rather unlike the typical form. I
have examined a specimen of Waters’ B. robusta (text-fig. 275, D) lent me by
the Manchester Museum. As the ancestrula and basal parts of the colony are
not there it is not the slide figured by Waters (1913, pl. XIX, fig. 15) and may
not be part ofthe same colony. I have been unable to trace the whereabouts
oO Smm
TEXT-FIG. 275.—A. Bugula robusta var. capensis Waters 99.7.1.305, showing small avicularium
at bifurcation. DB.a typical avicularium from the same specimen. C. B. robusta MacG., avicu-
larium from 79.5.27.1, Port Jackson. ——D. Bugula sp. (B. robusta Waters 1913) Waters Collection
Wasin, British E. Africa, 501.
a, lower head-angle. f, mandibular flange.
of the figured specimen. The zooecia are about the same size as those in the
figure, but have more pointed distal corners. The avicularia appear to be smaller.
Harmer (1926, p. 443) suggested that the figure might represent B. scaphoides.
The specimen examined by me (text-fig. 275, D) does not agree with that species.
It unfortunately has no ovicells. In its bifurcation of type 5 it resembles
B. robusta, and its avicularia have the general shape of those of B. robusta var.
capensis, but the lower head-angle is more acute (compare text-fig. 275, B and D)
and the mandible has a shoulder-like flange. The avicularia agree very closely
with those of B. subglobosa Harmer, but the zooecia are not so narrow proximally
and their inner distal corner is often pointed. The zooecia are nearer those of
B. subglobosa than B. robusta in size, but the bifurcation of B. subglobosa is of type 4.
The specimen from the Arabian Sea discussed on p. 337 suggests that the zooecia
of B. robusta may sometimes be smaller than usual, and it may well be that the
specimen from Wasin is a form of B. robusta.
Bugula neritina var. tenuata Thornely, which I have not seen, probably
Novitates Zoonoagicar XLI. 1939. 333
belonged to B. robusta var. capensis. The avicularian stalks are exceptionally
long and slender in the figure, which otherwise agrees closely with var. capensis.
The description also agrees except for a statement that the zooecia are “ almost
uniserial,”’ which is disproved by the figure. Thornely’s comparison of the
zooecia with those of var. rubra further confirms the suggested synonymy,
for var. rubra, which is represented in the British Museum by type material
(1936.12.30.165), is a synonym of B. robusta (cf. Harmer, 1926, p. 435). The
resemblance of var. tenuata to B. robusta was recognized by Harmer, who gave
it as a synonym of that species.
2. Bugula neritinoides sp.n. (Busk MS.) (text-figs. 273, C, 274, A-C, 276, F.).
Type. —-Tasmania, Mrs. Gatty (Busk Coll. 99.7.1.4648 and 4650).
Description. —Colony robust, rich brown in colour, biserial, with branching
oftype 4. Zooecia very flat frontally, without spines, outer corners very acutely
pointed and with no tendency to turn forward. Opesia occupying at least
three-quarters of frontal surface. Avicularia absent. Ovicells larger than
zooecia, attached to inner distal corner of zooecium, globular, pedunculate, a
calcareous border to the aperture, rest of ectooecium membranous.
Remarks.—According to Busk’s draft description,‘ the colony of this species
“spreads dichotomously into a circular expansion about 4 inches every way,
strongly curled inwards at the edge [tips] of the branches.” This remarkable
colony is not in the Busk Collection, but the type slides were evidently made
from it.
The long, pointed corners and very regular zooecia give the branches a
characteristic feathered appearance (text-fig. 273, C ; the apparently pale colour
is due to the use of a filter for the photograph). The ovicells resemble those of
Bugula neritina and related species in their general shape, but are gigantic.
In the dried state, which is the only one in which I have seen the species, the
ectooecium is collapsed and wrinkled (text-fig. 276, F), but even so the ovicells
are distinctly larger than the zooecia that bear them. The figure is drawn to
such a scale that the ovicell appears about the same size as those of other species
in the same figure. Comparison of the size of the zooecia emphasizes the rela-
tively gigantic size of the ovicell of B. neritinoides. The ovicell is closed by a
dark brown membrane continuous with the frontal membrane of the zooecium,
arising from a point proximal and lateral to the operculum.
In the general size of the zooecia and the robust scale of the colony
B. neritinoides resembles B. neritina and B. robusta, but it has bifurcations of
type 4.
3. Bugula scaphoides Kirkpatrick.
Bugula scaphoides Kirkpatrick, 1890, p. 18, pl. IV, fig. 1; Harmer, 1926, p. 443, pl. XX XI, figs. 7,
8, text-fig. 235.
? Bugula neritina var. ramosa Thornely, 1912, p. 142, pl. VIII, fig. 3.
Distribution.—China Sea (Kirkpatrick; 89.8.21.13, 68, 69, type speci-
mens; 1937.1.6.1); off New Guinea (Harmer; 28.3.6.289 and 290);
? Amirante (Thornely) ; ?Ghardaga, Red Sea (Dr. C. Crossland, 1937.9.28.36) ;
? Mauritius (34.10.12.8).
B. scaphoides differs from the other species considered here in its sculptured
ovicells.
1 The description is written on the back of the drawing traced in figs. 274, A-C.
334 NOVITATES ZOOLOGICAE XLI. 1939.
Ortmann (1892, p. 669) recorded a specimen with punctate ovicells, from
Dar-es-Salaam, as B. dentata var. africana. His comparison with B. dentata
implies the presence of spines and excludes the specimen from B. scaphoides,
in which the ovicell is, in any case, better described as reticulate than as
punctate.
On the other hand, it seems probable that B. neritina var. ramosa Thornely,
described as having ovicells with a pitted surface, is a synonym of B. scaphoides.
The type material of B. scaphoides is uniserial in part, and, both the type and
the Siboga material, show lateral buds projecting from the sides of the zooecia
at right angles, as in B. neritina var. ramosa. In the type of B. scaphoides this
bud has given rise to a branch. Stout rootlets, as shown in Thornely’s figure,
are present in both sets of material. The zooecia agree in shape with those of
B. scaphoides, and the dots on the ovicell in the figure might be a poor represen-
tation of the reticulation. I have not, however, seen a specimen of var. ramosa.
A fragment from Ghardaga, Red Sea, resembles B. scaphoides in having
reticulate ovicells (text-fig. 276, E), but they are smaller. The zooecia are also
much smaller, and have a shorter proximal tubular part. The avicularia on the
other hand, are about the same size, and, as in B. scaphoides, are attached to an
unusually long peduncle, or outgrowth of the zooecial wall, which remains
projecting quite conspicuously if the avicularium falls off (see spine-like projection
from behind two of the ovicells in Harmer’s fig. 7).
The specimen from Mauritius closely resembles the one from the Red Sea
in its zooecia and avicularia, but has no ovicells. It differs from B. crosslandi
with which it might, in the absence of ovicells, be confused, in the pointed outer
(and sometimes inner) distal corners of the zooecia, in the long stalks of the
avicularia, and in the presence of lateral branches and stout rootlets. The colony
spread over the roots of a hydroid and was attached by the stout, thick-walled
rootlets which spring from the basal surface, and are branched at their tips.
4. Bugula minima Waters (text-figs. 276, A-C, and 278, C).
Bugula neritina var. minima part Waters, 1909, p. 136, pl. XI, figs. 4, 6,7; part (at least) Thornely
1912, p. 141; part Marcus, 1921, p. 1, pl. I, fig. 1.
Distribution.Mersa Makdah, Red Sea, 5 fms. (Waters ; Liverpool
Museum); Ghardaga, Red Sea, seaward edge of Outer Reef, low water spring-
tide (Dr. C. Crossland, 1937.9.28.37) ; Dar-es-Salaam (Daressalam) (Marcus ;
Stuhlmann, Berlin Mus. 1944 ; 1939.4.18.2) ; Providence, 50-78 fms. (Thornely ;
Thornely Coll., 1936.12.30.166) ; Ceylon (99.7.1.4608).
Type.—Mersa Makdah (Liverpool University Museum).
Description.—Colony, biserial with branching of type 4. Zooecia with outer
distal corner pointed, inner corner rounded or pointed. Opesia occupying
nearly the whole frontal surface. Brown pigment in tissues persisting in spirit.
Avicularium springing from side of zooecium, distal to proximal expansion, and
level with proximal end of opesia, large and long, strongly curved dorsally,
upper head-angle variable, hooked beak forming at least half the total length.
A few very large avicularia of similar shape to those with the more obtuse upper-
head-angle, or with even flatter head (text-fig. 276, B, C). Ovicells attached to
inner distal corner of zooecium, oval, pedunculate, with thickened band round
aperture.
1 Name of locality corrected, Waters, 1910, p. 254.
Noviratres Zoonoarcam XLI. 1939. 335
Synonymy.—Waters used the name B. neritina var. minima for specimens,
which I have examined, of three distinct species.
1. The specimen from Mersa Makdah shown in Waters’ figures? 4, 6 and
7, to which I restrict Waters’ name, giving it specific rank (text-fig. 276, C).
2. The specimen from Khor Dongola shown in his figure 5 which belongs to
B. crosslandi.
Figs. A-E
Citititstitstatstitry
0 Fig. F
TEXT-FIG. 276.—A-C. Bugula minima Waters. B and C show large and small avicularia with
parts of surrounding zooecia. A and B from specimen from Ghardaga, 1937.9.29.37. C from type
specimen from Mersa Makdah, Red Sea. D. Bugula crosslandi sp.n. part of Waters’ specimen from
Khor Dongola (Liverpool University Museum). E. Bugula ? scaphoides 1937.9.28.36, from:
Ghardaqa. One ovicell incomplete. Note long avicularian stalks. a. avicularium partially hidden
by ovicell. F. Bugula neritinoides sp.n. part of type specimen 99.7.1.4648.
a. (in B and C) upper head-angle.
3. The specimens from Prison Is. and Ras Osowamembe, Zanzibar (1913),
which belong to B. robusta, as stated by Harmer, 1926.
As noted by Waters, the specimens from Khor Dongola have
smaller zooecia and much smaller avicularia ” than those from Mersa Makdah.
In addition, B. crosslandi differs from B. minima in its shorter opesia and the
“ somewhat
1 Waters gives Khor Dongola as the locality for his figures 4 and 5, but the actual zooecia figured
can be recognized in his slides, and figure 4 is drawn from the same specimen from Mersa Makdah
as figure 6.
336 NOVITATES ZooLoGIcAE XLI. 1939.
position of the avicularium relative to it, in the less-pointed distal corners of
the zooecia, and the uniform size of the avicularia. The differences between the
two species are quite obvious when actual specimens are examined, and can
mostly be distinguished in Waters’ figures. Harmer attributed all these forms
to B. robusta, but B. minima and B. crosslandi are more delicate forms with
smaller zooecia of different shape, smaller avicularia, and different bifurcation.
Marcus also confused more than one species under B. neritina var. minima.
I have examined his specimens and find that he had :
1. Specimens, from Dar-es-Salaam, of typical B. minima as here understood.
2. Material of B. neritina which was mixed with the B. minima from
Dar-es-Salaam.
3. Specimens of B. robusta from Bagamoyo (Berlin Museum, 1949) and
from Gaspar Straits (Riksmuseum, Stockholm, No. 693).
His specimens of B. robusta agree very closely with a specimen from Siboga
St. 164 (28.3.6.272), in which the avicularia are borne on long stalks at about
the middle of the side of the zooecium and cusps are absent or evanescent.
Marcus’ figure represents a form in which the zooecia are about half the size
of those of his specimens of B. robusta, and the avicularia, which have a much
flatter head (upper head-angle obtuse in contrast to the acute angles of the
specimens of B. robusta), are attached by short stalks near the proximal end of
the zooecium. It was evidently drawn from the material from Dar-es-Salaam
and was, indeed, recognizable as B. minima without the examination of specimens
which has since confirmed this conclusion. The structures in the figure which
might be taken for cusps like those of B. robusta appear to be the median sense-
organs.
The specimen from Providence, recorded as B. neritina var. minima by
Thornely (1912), agrees very closely in the shape of avicularia and ovicells, but
does not possess any of the exceptionally large avicularia. The zooecia are
rather slender and do not widen so much distally, and the outer distal corner is
correspondingly more obtuse. Of the specimens recorded by Thornely (1905,
1907, 1912) as B. neritina with avicularia, I have only seen the specimen from
Amirante (Cambridge Museum) which belongs to B. robusta. Harmer puts them
all in B. robusta. I have no evidence about the specimen similarly recorded by
Philipps (1899).
Specimens have also been recorded under this varietal name by Osburn
(1914, p. 187), and by Okada and Mawatari (1938, p. 451), but these identifications
need confirmation now that the name is more strictly defined.
Remarks.—This species differs from related forms in the size, shape and
position of its avicularia (see key) and the shape of the zooecia. There are no
ovicells in the type, but those of other specimens have a broad thickened border
to the ectooecium (text-fig. 276, A).
In the type specimen the small avicularia are smaller and have a rounder
head (more acute upper head-angle) than those of the other specimens (cf. text-
fig. 276, B and C). The large avicularium in text-fig. 276, B, is smaller than
others on the same specimen, which shows as great a contrast between the two
sizes as in the type. In Marcus’ material some of the large avicularia are even
bigger than those figured, and have a more obtuse upper head-angle. The small
avicularia are like those from Ghardaqa. The fragments from Ceylon have avicu-
laria of the same shape as the small ones of the type, and have no large avicularia.
Novirares Zoonoarcan XLI. 1939. 337
The colony from Ghardaga springs by a short stalk from rootlets ramifying
in a sponge. The stalk (text-fig. 278, C) is an elongated zooecium with lateral
thickenings like those of certain related species. It has a blunt thickened end,
whose appearance suggests that it may have healed after breakage. It is thus
not clear whether the stalk is an ancestrula or has been budded from the tangle
of stout rootlets in the sponge. One stout rootlet arises from the stalk and
passes into the sponge.
5. Bugula crosslandi sp.n. (text-figs. 276, D, and 277, A).
Bugula neritina var. minima part Waters 1909, p. 136, pl. XI, fig. 5; part Hastings 1930, p. 704,
1. I, fie. 6.
Not ale neritina var. minima Thornely 1912, p. 141 (part at least = typical B. minima) ; Waters
1913, p. 471 (= B. robusta) ; Marcus 1921, p. 1 (= typical B. minima and B. robusta).
Distribution. —Abhu Shaar, Red Sea, 4-1 fm., May 20, 1933 (Dr. C. Crossland,
1937.9.28.35); Khor Dongola, Red Sea (Waters ; Liverpool Mus.) ; Zanzibar
(Hincks Coll., 99.5.1.407) ; Gorgona (Hastings ; 29.4.26.43, 245).
Type. —1937.9.28.35.
Description. —Colony delicate, biserial, with branching of type 4. Zooecia
without spines or strongly pointed corners. Opesia occupying at least three-
quarters of frontal surface. Avicularium springing from proximal gymnocyst,
at a point distal to proximal expansion, but proximal to opesia, with rounded
head and hooked beak of moderate length. Ovicell attached to inner distal
corner of zooecium, globular, pedunculate, with thickened band round aperture.
Synonymy.—The agreement of Waters’ specimen from Khor Dongola
(text-fig. 276, D) with the type is very close. The British Museum specimen
from Zanzibar (see Hastings 1930) also agrees closely with the type, but the
rather more robust one from the Arabian Sea (99.5.1.406) appears to belong to
B. robusta. Although it is not so robust as most specimens of that species, it
much resembles, both in size and shape, Harmer’s figure (1926, pl. XX XII, fig. 3)
of a specimen with evanescent cusps. It agrees with B. robusta in its bifurcation
and the shape of the zooecia. The avicularia are placed, in the position com-
monest in B. robusta, on the proximal gymnocyst just distally to the constriction.
The presence of this marked constriction is in itself a point of agreement with
B. robusta. In B. minima and B. crosslandi the zooecium tapers until it rather
suddenly widens to the proximal expansion. In B. robusta the lateral wall
curves outward a little before turning sharply inward to a constriction that
marks off the proximal expansion. Both the position of the avicularium and
the outline of the zooecium as here described can be seen in my figure of
B. robusta var. capensis (text-fig. 275, A).
It is rather surprising to find B. crosslandi in the Pacific, but the specimen
from Gorgona (text-fig. 277, A) agrees quite closely with the type. The avicularia
are a little smaller and more slender. The specimens from Galapagos 9
(29.4.26.246) are young colonies of some other species, in which the bifurcation
is of type 3, the avicularia are attached at the side of the opesia at about the
middle of the length of the zooecium, and spines are present.
Remarks.—The points in Waters’ drawings which might be taken as a poor
representation of the avicularian cusps of B. robusta are evidently intended for
the condyles, to which the mandible is articulated, which are rather conspicuous
in his specimen,
25
338 Novirates ZooLoGIcAE XLI. 1939.
B. crosslandi is distinguished from B. expansa by the absence of the foot-like
basal attachment, and by its smaller avicularia, with flatter head and longer
beak, attached to the proximal gymnocyst at a little distance from the proximal
expansion (cf. text-figs. 277, A, B).
With the exception of Hincks’ specimen from Zanzibar, all the material of
this species known to me has been collected by Dr. C. Crossland, after whom
it is named.
6. Bugula expansa sp.n. (text-figs. 277, B, and 278, B).
Type.—British Antarctic Expedition (“ Terra Nova ’’), St. 134, Spirits Bay,
near North Cape, New Zealand, 20-37 m., August 31, 1911, 1939.2.2.2.
ovicells are shown.
Description.—Colony biserial, with branching of type 4, attached by large,
flat, more or less pear-shaped foot (text-fig. 278, B). Foot with very thin
calcareous wall, and thick crust-like calcareous border which easily breaks
away. Interior of foot filled with yellow, granular material. A thick-walled,
yellow tube rising vertically from narrow end of foot, connected with first
zooecium by a joint. Rootlets sometimes attached to substratum by similar,
but smaller and more irregularly shaped, feet. First zooecium elongate, with long
opesia and no avicularium, giving rise to two normal zooecia. Zooecia (text-
fig. 277, B) with outer distal corner slightly pointed, inner distal corner usually
rounded, sometimes pointed. Opesia extending nearly to proximal end of
zooecium. Avicularium springing from outer side of proximal expansion of
zooecium, Head of avicularium round, beak hooked, shorter than that of
Noviratres Zoonoctcar XLI.
1939. 339
B. crosslandi. Ovicells attached to inner distal corner of zooecium, globular,
pedunculate, with a thickened band round aperture.
Remarks.—A number
of small colonies were
growing on the concave
surface of shells.
The zooecia of this
species are similar to
those of B. crosslandt,
and the two species agree
in their ovicells and pig-
mentation. The avicu-
laria of B. expansa are
larger, have a rounder
head and shorter beak,
and are attached to the
proximal expansion of
the zooecium. The two
species also differ in the
characters of the base of
the colony (cf. text-fig.
278, B, and Hastings,
1930, pl. II, fig. 6).
Caulibugula Verrill 1900.
1. Caulibugula
zanzibariensis (Waters).
Stirparia zanzibariensis Waters,
1913, p. 469, pl. LXVIII,
figs. 1, 2, pl. LXIX, fig. 14.
‚Stirpariella zanzibariensis Mar-
cus, 1925, p. 53.
Caulibugula zanzibariensis
Harmer, 1926, p. 460,
pl. XX XIII, figs. 5-10.
Bicellaria glabra Busk, 1884,
p. 35, pl. VI, figs. 1, 1a (not
Stirparia glabra Hincks).
Distribution. —Chu-
aka, Zanzibar (Waters) ;
Kurrachee (83.9.13.33) ;
Java (Harmer; 28.3.6.
306, 307, 308); off Bahia
(Busk; 87.12.9. 168,
169).
The Challenger speci-
Z1
TEXT-FIG. 278.—A. Caulibugula annulata. 97.5.1.345, Port-
land Victoria. Septum between two main stalk-kenozooecia and
bases of two lateral stalk-kenozooecia. so, septum in optical section.
sf, fringed insertion of septum on lateral wall. B. Bugula
expansa spn. Base of type colony 1939.2.2.2. Calcareous crust
shown black, broken away at b. C. Bugula minima Waters,
base of colony from Ghardaga, 1937.9.28.37. The first
zooecium is bent and the stalk or ancestrula is broken at b. r,
rootlet.
men from off Bahia, recorded by Busk as B. glabra (Hincks), differs from the
type specimen of Hincks’ species in the shape of its opesia, in the number and
position of its spines, in the shape and position of its avicularia, in the way in
which the first zooecia place themselves back to back, forming a cone with the
340 Novirares ZooLoGIcAE XLI. 1939.
zooecia facing outward, and in its more delicate appearance. Several of these
points are distinctly shown in Busk’s figure, and in all of them the specimen
resembles (©. zanzibariensis, agreeing very exactly with Harmer’s description,
except perhaps in the absence of stem vesicles. Short rootlet-like structures
with slightly inflated tips are, however, present.
In addition to the unmounted specimen of C. zanzibariensis (87.12.9.169)
and two slides of its kenozooecia (87.12.9.169 part and 168), the Challenger
material labelled C. glabra and purporting to come from Bahia includes a slide
of C. annulata (87.12.9.167), as noticed by Waters (1913, p. 468). Waters, who
evidently did not see the unmounted specimen, was puzzled by not finding
material corresponding to Busk’s figure, and by the presence of C. annulata.
Noticing that the slide of C. annulata had been relabelled, apparently in haste,
by Busk, I examined the underneath labels. This revealed that the slide was
originally labelled Port Jackson, but this had been crossed out and Bahia sub-
stituted. In view of this evidence of uncertainty about the locality, I think
there is no good reason to accept C. annulata as having been obtained off Bahia.
It is also worth noticing that Busk expressly states that he only had one specimen
of his supposed C. glabra.
C. annulata was obtained by the Challenger from St. 161 (Port Phillip,
1938.11.24.1 and 2), but this material was left unnamed by Busk.
The young colony from the Barrier Reef compared by Hastings (1932,
p. 408) with C. zanzibariensis is further distinguished from that species by the
first zooecium of the fan which is Bugula-like and not markedly different from the
succeeding zooecia, cf. the W. Australian specimen discussed by Harmer (1926,
p. 461).
2. Caulibugula tuberosa sp. n. (text-fig. 279, A-C).
Type.—Discovery St. 934, 34° 11’ S., 172° 10’ E. (New Zealand), 17. vii. 32.
98 m. (1939.2.2.1).
Description. —Colony attached by stout, vesicular, rootlet-like kenozooecia.
Stalks short, stalk kenozooecia stout, annulated proximally without calcareous
thickenings. Bifurcation of Harmer’s type 5. Zooecia bicellarielliform, opesia
almost circular, occupying considerably less than half the length of the zooecium,
sometimes with thin raised border. Spines long and curved, 0-1 proximal,
0-4 outer, 0-1 distal, the latter directed basally, zooecia bordering axil usually
spineless. Ancestrula turbinate, tubular portion fairly long, annulated proxi-
mally, with spines on proximal and lateral borders of opesia. First zooecium of
fan turbinate, but short, annulated or merely constricted proximally, with
variable spines." Avicularia absent. Ovicells unknown.
Remarks.—The material consists of two colonies growing on a stone. They
were apparently more or less surrounded by a sponge, now mostly cleared away.
Text-fig. 279, A and B, shows the bases of the colonies, and in B the remains of
the sponge in the interstices are indicated. The smaller colony (text-fig. 279, A)
has a turbinate ancestrula, attached by proximal rootlets and giving rise distally
to a fan and a stalk-kenozooecium. It is curious that the stalk-kenozooecium
appears to spring from the opesial surface of the ancestrula, being encircled by
the spine-bearing border of the aperture. The first zooecium of the fan has a
proximal constriction, and the ancestrula and stalk-kenozooecium are both
1 N U r - 1 1 i i
There are, for example, 4 spines on one side of the aperture and one on the other in one instance,
and in another (text-fig. 279, B) 9 spines are ranged in a single series.
Novrrares Zoonoarcan XLT. 1939. 341
A. Base of younger
TEXT-FIG. 279.—Caulibugula tuberosa sp.n. Typespecimens 1939.2.2.1.
colony. a,ancestrula. b, buttress (torn). fs, fringed septum. n, rootlets. sk, stalk-kenozooecium
zi, first zooecium of fan. B. Base of older colony. e, broken base of second erect tube. v, detached
vesicle of buttress (b) from thinner part of stalk. C. Zooecia from the younger colony.
342 Novirarres Zoonocicar XLI. 1939.
annulate proximally. From the side of the ancestrula arises a stout rootlet-like
structure which drops, like a buttress, to the stone and there expands into a
vesicle (torn in mounting as shown in figure).
The older colony (text-fig. 279, B) has a stout, annulate stalk, with thick,
dark brown walls. From its truncated end springs, as if by regeneration, a more
slender thin-walled kenozooecium which gives rise in its turn to a fan and a stalk,
the latter consisting of two kenozooecia and ending in the first zooecia of a fan.
Buttresses from the thick part of the stalk have thick walls and form large
thick-walled darkly pigmented vesicles on the stone. From these arise thin-
walled vesicles, which are also encrusting, and one erect tube (e), which has
formed a small buttress, but is broken short. A thin-walled buttress (b) and
vesicle (v, now broken) arise from the thinner part of the main stalk.
It is generally agreed that the stalk segments of Caulibugula are kenozooecia,
and the Californian species ©. ciliata (Robertson) affords some support to this
view, as its kenozooecia have a small vestigial opesia, often with marginal spines.
In the relation of the kenozooecia to the fans and in the general structure of the
colony C. ciliata agrees with other species of Caulibugula. I have examined a
specimen of C. ciliata sent to the British Museum by Dr. Amy Blagg (1938.11.
30.8).
The stem kenozooecia of C. tuberosa resemble those of C. annulata (text-fig.
278, A), but are much smaller. In both species the transverse septum between
one kenozooecium and the next has so irregular a line of attachment to the
lateral wall that it can only be described as fringed (cf. C. caraibica Levinsen
1909, pl. IIT, figs. 2-5). It is at once noticeable that the buttresses and vesicles
of C. tuberosa spring from similar fringed discs, and I therefore conclude that
they, too, are probably to be regarded as kenozooecia.
Harmer (1926, p. 463) described the much more root-like basal structures of
C. exilis and concluded that they were composed of kenozooecia. Traces of
fringing can be detected both in the type material from Port Nepean (97.5.1.347,
348) and in Harmer’s material (28.3.6.309), and it seems probable that
C. tuberosa represents a condition of the kenozooecial system intermediate between
those of ©. exilis and ©. annulata, and affords valuable confirmation of Harmer’s
conclusion. The specimens of C. exilis were immersed in sponges up to the
base of the fans. One is tempted to relate the condition in C. tuberosa to its less
complete immersion.
In zooecial characters C’. tuberosa is near C. annulata, which it resembles in
the rounded opesia and the number and distribution of the spines (text-fig.
279, C). The zooecia are very much smaller, with a relatively longer tubular
portion, and they do not have the forked thickening in the basal wall. The
whole growth is much more delicate. C. exilis has a longer opesia with a different
arrangement of the spines.
The four species mentioned here (C. annulata, C. tuberosa, C. exilis, C. ciliata)
agree in the absence of longitudinal calcareous thickenings in the walls of the
kenozooecia. In this they apparently resemble the form ascribed by Osburn
(1914, p. 188) to C. armata Verrill. Thickenings are, however, shown by Marcus
(1938, p. 29) in what he believes to be the same species. In any case, Osburn’s
species is distinguished from C. tuberosa by the shape of the opesia, and the
number and arrangement of the spines. It has avicularia. Fringing is not
described in the kenozooecia of C. armata and is absent in C. ciliata.
Novirates ZootoGican XLI. 1939. 343
REFERENCES.
Busk, G. 1852a. An Account of the Polyzoa and Sertularian Zoophytes. In
J. MacGillivray, Narrative of the Voyage of H.M.S. Rattlesnake, I. Appendix,
59 pp., 1 pl.
— 1852b. Catalogue of Marine Polyzoa in the Collection of the British Museum,
I. London.
1884. The Cheilostomata. Rep. Zool. Chall. Hup., XXX.
Cranwell, L. M., and Moore, L. B. 1938. Intertidal Communities of Poor
Knights Islands, New Zealand. Trans. Roy. Soc. N.Z., LX VII, pp. 375-407,
2 pls. 3 figs., 1 table.
Gray,J.E. 1843. Additional Radiated Animals and Annelides, in Dieffenbach,
E. Travels in New Zealand, II, pp. 292-295.
Harmer, S. F. 1915. Entoprocta, Ctenostomata, Cyclostomata. Rep. Siboga,
Exp., XXVIIIa.
—— 1923. On Cellularine and other Polyzoa. J. Linn. Soc. London, XXXV.
pp. 293-361, 4 pls.
—— 1926. Cheilostomata Anasca. Rep. Siboga Exp., XXVIII.
Hasenbank, W. 1932. Bryozoa der Deutschen Tiefsee Expedition, Teil I.
Wiss. Ergebn. Deutsch. Tiefsee-Exped., XXI, 2, pp. 319-380, 1 pl. 35 text-figs.
Hastings, A. B. 1930. Cheilostomatous Polyzoa from the Vicinity of the
Panama Canal. ..... Proc. Zool. Soc. London, 1929, part 4, pp. 697-740, 17 pls.
—— 1932. The Polyzoa.... Rep. Barrier Reef Exp. 4, No.12, pp. 399-458, 1 pl.,
20 text-figs.
Hincks, T. 1881. Contributions towards a General History of the Marine
Polyzoa, V. Ann. Mag. Nat. Hist. (5), VII, pp. 156-161, 3 pls.
— 1891. South African and other Polyzoa. Ann. Mag. Nat. Hist. (6), VII,
pp. 285-298, 2 pls.
Kirkpatrick, R. 1890. Report upon the Hydrozoa and Polyzoa collected .. .
in the China Sea. Ann. Mag. Nat. Hist. (6), V, pp. 11-24, 3 pls.
Kluge, H. 1914. Die Bryozoen. . . . Deutsche Südpolar-Exped., XV, 7, pp. 601-
678, 8 pls., 47 text-figs.
Levinsen, G. M. R. 1909. Morphological and Systematic. Studies on the
Cheilostomatous Bryozoa. Copenhagen.
Livingstone, A. A. 1929. Bryozoa Cheilostomata from New Zealand. Vidensk.
Medd. naturh. Foren. Kjob., LXX XVII, pp. 45-104, 2 pls., 7 text-figs.
MacGillivray, P. H. 1869. Descriptions of some new Genera and Species of
Australian Polyzoa. ... Trans. Proc. Roy. Soc. Vict., IX, pp. 126-148.
1881. In McCoy, Prodromus Zool. Victoria, Dec., VI, pp. 27-46, 4 pls.
Marcus, E. 1925. Uber Stirpariella mortenseni und das Genus Stirpariella.
Vidensk. Medd. Dansk. Naturh. Foren., LX X XI, pp. 37-55, 12 text-figs.
—— 1938. Bryozoarios Marinhos Brasileiros, II. Bol. Fac. Phil. Sc. Letr.
Univ. S. Paulo, IV, Zool. 2, pp. 1-137, 29 pls.
Okada, Y., and Mawatari, S. 1938. On the Collection of Bryozoa along the
coast of Wakayama-ken, the middle part of Honsyu, Japan. Annot. Zool.
Jap., XVII, pp. 445-462, 1 pl., 7 text-figs.
Ortmann, A. 1892. Die Koralriffe von Dar-es-Salaam und Umgegend. Zool.
Jahrb. Jena, Syst. 6, pp. 631-670, 2 text-figs.
Osburn, R. C. 1914. The Bryozoa of the Tortugas Islands, Florida. Pap.
Tortugas Lab. Carn. Inst., V, pp. 183-222, 23 text-figs. (Carnegie Publ. 181).
344 Noyrratrs ZooLoGIcAE XLI. 1939.
Philipps, E. G. 1899. Report on the Polyzoa from the Loyalty Islands, New
Guinea and New Britain. Willey’s Zool. Res., IV, pp. 440-450, 2 pls.
Thomson, Wyville T.C. 1858. On New Genera and Species of Polyzoa. ... Nat.
Hist. Rev. London, V, Proc. Societies, pp. 134-147, 5 pls.
Thornely, L. R. 1905. Report on the Polyzoa. . . . in Herdman, Rep. Pearl
Oyster Fisheries Gulf of Manaar, IV. Suppl. Rep., X XVI, pp. 107-130, 1 pl.
—— 1907. Report on the Marine Polyzoa in the collection of the Indian Museum.
Rec. Ind. Mus., I, pp. 179-196, 8 text-figs.
—— 1912. The Marine Polyzoa of the Indian Ocean from H.M.S. Sealark.
Trans. Linn. Soc. London, XV, pp. 137-157, 1 pl.
Waters, A.W. 1887. Bryozoa from New South Wales, N. Australia, ete. Pt.1.
Ann. Mag. Nat. Hist. (5), XX, pp. 81-95, 1 pl.
—— 1909. Reports on the Marine Biology of the Sudanese Red Sea... . I.
J. Linn. Soc. Zool., XX XI, pp. 123-181, 9 pls.
—— 1910. Op. cit., II. J. Linn. Soc. Zool., XX XI, pp. 231-256, 2 pls.
—— 1913. The Marine Fauna of Brit. E. Africa and Zanzibar. . . . Bryozoa.
Proc. Zool. Soc. Lond., pp. 458-537, 10 pls.
Noviratrres Zoonoarcarn NLI. 1939. 345
POLYCHAETA OF THE “ ROSAURA” EXPEDITION.
By C. C. A. MONRO
Department of Zoology, British Museum (Natural History).
(With 4 text-figures.)
ee Polychaeta of this report were collected by Dr. J. S. Colman during the
1937-38 expedition of Lord Moyne’s M.Y. Rosaura.
They fall into four groups: (1) shallow-water, coastal, benthic forms from
east and south-west Greenland and from Labrador ; (2) similar forms from the
West Indies and off British Honduras ; (3) deep water, benthic forms obtained
off Grenada and off the Canary Islands; (4) pelagic forms obtained at various
positions from the West Greenland Current to the Equator.
There are altogether 71 species belonging to 41 genera. I have established
a new genus of Maldanids on a specimen taken off British Honduras which has
the dorsal surface of several segments provided with a coating of long papillae
which I believe to be branchial. As far as I know, this is the first record of
branchial papillae in the family. I have also described a new EHunice related to
E. cariboa Grube from deep water off Grenada and a new variety of Fauvel’s
Harmothoé joubini from deep water off the Canary Islands.
Much of the ground represented by the present collection has already been
covered in a masterly fashion by Prof. P. Fauvel in his reports on the collections
made from the yachts of Prince Albert I of Monaco, and I have decided, instead
of writing a full report, simply to give a list of the species with their stations and
to restrict myself to the description of the new forms and to commentary only
in very special cases. Some of the material, especially the pelagic forms (exclud-
ing the Tomopterids), is very remarkably preserved and is thoroughly suitable
for commentary, but I repeat that much of the ground has already been covered.
Station 1, 65° 35’ N., 37° 20’ W., Angmagssalik Harbour, gear 2-ft. dredge,
depth 30-50 m. : !
Harmothoé imbricata (L.).
Lagisca extenuata (Grube).
Gattyana cirrosa (Pallas).
? Hteone flava (Fabricius).
Nereis zonata Malmgren.
Onuphis conchylega Sars.
Lumbrinereis fragilis (O. F. Müller).
Scoloplos armiger (O. F. Müller).
Scalibregma inflatum Rathke.
Ammotrypane aulogaster Rathke.
Thelepus cincinnatus (Fabricius).
Pista maculata (Dalyell).
1 Nearly all the species in this list are described with full references by Fauvel in the Faune de
France, vols. V and XVI, 1923 and 1927. Those that are not so described have the date of the
original description placed after the author’s name.
346
NOVITATES ZOOLOGICAE XLI. 1939.
Station 4, 60° 30’ N., 46° 40’ W. to 60° 35’ N., 46° 45’ W., off Julianehaab,
gear 3-ft. dredge, depth 550 m. :
? Pista maculata (Dalyell).
Station 5, 60° 43’N., 46° 02’ W., Julienehaab Harbour, gear 2-ft. dredge,
depth c. 20 m.; cod long line, depth 0-80 m. ; shore coll. :
Pee cirrosa (Pallas).
Harmothoé imbricata (L.).
Eteone longa (Fabricius).
Eulalia viridis (O. F. Müller).
Nereis pelagica L.
Nainereis quadricuspida (Fabricius).
Arenicola marina (L.).
Pectinaria granulata (L.) 1767.
Thelepus cincinnatus (Fabricius).
Spirorbis granulatus (L.).
Station 6, 60° 06’N., 45° 25’ W., off Nanortilik, gear 3-ft. dredge, depth
110 m.:
Gattyana cirrosa (Pallas).
Phyllodoce groenlandica Oersted.
Nephthys caeca Fabricius.
Nephthys ciliata (O. F. Müller).
Onuphis conchylega Sars.
Lumbrinereis fragilis (O. F. Müller).
Stylarioides plumosa (O. F. Müller).
Owenia fusiformis Delle Chiaje.
Myriochele heeri Malmgren.
Amphitrite cirrata (O. F. Müller).
Terebellides stroemi Sars.
Potamilla neglecta (Sars) 1851.
Station 8, 58° 48’N., 46° 11’ W., West Greenland current, gear 2-m.
stramin net, depth 1300-0 m. :
Travisiopsis levinseni Southern.
Tomopteris septentrionalis Quatrefages.
Station 9, 52° 13’N., 55° 45’ W., north-east arm, Horn Bay, Niger Sound,
Labrador, shore coll. :
Nereis virens (Sars).
Station 10, 52° 13’ N., 55° 47’ W., north-west arm, Horn Bay, Labrador,
gear 2-ft. dredge, depth 15 m. :
Lagisca extenuata (Grube).
Nereis pelagica L.
Nephthys ciliata (O. F. Müller).
? Pectinaria granulata (L.) 1767.
Ampharete goesi (Malmgren) 1865.
Terebellides stroemi Sars.
Novrrarrs Zootoarcar XLI. 1939. 347
Station 13, 36° 88’ N., 74° 23’ W., between Gulf Stream and coast ; probably
in counter-current, gear 2-m. stramin net, depth 750-0 m. :
Lagisca hubrechti (McIntosh).
Sagitella kowalewskii Wagner.
Station 15, 18° 21’ N., 75° 25’ W., gear 2-m. stramin net, depth c. 1250-0 m. :
Harmothoé benthophila Ehlers 1913.
Tomopteris planktonis Apstein.
Lopadorhynchus uncinatus Fauvel.
Station 16, 19° 18’ N., 81° 24’ W. to 19° 18’N., 81° 22’ W., Georgetown,
Grand Cayman, shore coll., coral rock ; and North Sound, Grand Cayman, gear
2-ft. dredge, depth 2 m. :
Opisthosyllis nuchalis Verrill 1900.
Eunice vittata (Delle Chiaje) juv.
Sabella melanostigma Schmarda 1861. !
Station 22, 17° 28’ N., 88° 11’ W., Belize Harbour, gear 2-ft. dredge, depth
6m.:
Branchioasychis colmani gen. et sp. nov.
Station 23, 17° 29’ N., 88° 10’ W., Belize Harbour, gear silk tow-net, depth
2-3 m.:
Loimia medusa (Savigny) 1820, larvae.
Station 24, 17° 15’ N., 87° 49’ W. to 17° 16’N., 87° 50’ W., Indian Cay,
Turneffe Is., shore coll. ; Turneffe Is., gear 2-ft. dredge, depth 4 m. :
Hermodice carunculata Pallas.
Syllis spongicola Grube.
Syllis brachychaeta Schmarda 1861.
Opisthosyllis nuchalis Verrill 1900.
Polyophthalmus pictus (Dujardin).
Station 27, 16° 22’ N., 86° 40’ W., off west end of Roatan Is., gear 2-m.
stramin net, depth 600-0 m. :
Harmothoé benthophila Ehlers 1913.
Tomopteris kefersteini Greeff.
Lopadorhynchus uncinatus Fauvel.
Lopadorhynchus brevis Grube.
Station 28, 16° 44’ N., 85° 42’ W., north of Bonacca Is., gear 2-m. stramin
net, depth c. 1100-0 m. :
Harmothoé benthophila Ehlers 1913.
Lopadorhynchus uncinatus Fauvel.
? Tomopteris nisseni Rosa.
Callizona moebii Apstein.
Station 31, 15° 54’N., 82° 13’ W., off Gorda Cay, Mosquito Bank, gear
3-ft. dredge, depth 34 m.:
Syllis corallicola Verrill 1900.
Syllis spongicola Grube.
? Glycera longipinnis Grube 1878.
Nereis trifasciata Grube 1878 juv.
? Pista cristata (Müller) juv.
348 NoviratEs ZooLoGIcAE XLI. 1939.
Station 32, 12° 42’ N., 80° 25’ W., gear 2-m. stramin net, depth 900-0 m. :
Lopadorhynchus uncinatus Fauvel.
Lopadorhynchus brevis Grube.
Callizona angelini (Kinberg).
Tomopteris nisseni Rosa.
Tomopteris ligulata Rosa.
Station 34, 12° 05’ N., 61° 49’ W., off St. George, Grenada, gear Agassiz
trawl, depth 720-800 m. :
Dalhousiella carpenteri McIntosh.
Hunice rosaurae n. sp.
Station 36, 7° 11’ N., 57° 59’ W., off Demerara, gear otter trawl, depth 20 m.:
Onuphis magna (Andrews) 1891.
Station 37, 7° 47’N., 58° 17’ W., off River Essequibo, gear otter trawl,
depth 23 m.:
Loimia medusa (Savigny) 1820.
Station 42, 5° 51'S., 35° 38’ W., off north-east Brazil, gear 2-m. stramin
net, depth 1200-0 m. :
Alciopa cantrainii (Delle Chiaje).
Station 43, 3° 35’8., 32° 20’ W., 12 miles north of Fernando Noronha,
gear 2-m. stramin net, depth 750-0 m. :
Chloeia larva.
Travisiopsis lanceolata Southern.
Sagitella kowalewskir Wagner.
Tomopteris nisseni Rosa.
Spionid larvae.
Station 44, 0° 56’ N., 29° W., St. Paul’s Rocks shoal, gear 3-ft. dredge, depth
50-60 m. ; close to the rocks, gear silk tow-net, depth 2 m. :
Syllis gracilis Grube.
Syllis corallicola Verrill 1900.
Eunice cariboa Grube, var. kinbergii Webster 1884.
2
Station 45, 4° 15’ N., 26° 20’ W., northern edge of north equatorial current,
gear 2-m. stramin net, depth 900-0 m. :
Callizona setosa (Greeff).
Vanadis formosa Claparéde.
Station 46, 7° 27’ N., 23° 08’ W., Guinea current, gear 2-m. stramin net,
depth 1000-0 m. :
Travisiopsis lanceolata Southern.
? Tomopteris nisseni Rosa.
Tomopteris helgolandica Greeff.
Tomopteris sp.
Station 49, 28° 25’ N., 13° 34’ W., between Fuerteventura Is. and Africa,
gear Agassiz trawl, depth c. 1300 m.:
Macellicephala mirabilis McIntosh 1885.
Robertianella synophthalma McIntosh 1885.
Harmothoé joubini Fauvel, var. rosaurae var. nov.
Leocrates atlanticus McIntosh.
Hyalinoecia tubicola (O. F. Müller).
Novirares Zoonoaican XLI. 1939. 349
GENUS HARMOTHOE KINBERG 1855.
1. Harmothoé benthophila Ehlers 1913 (text-fig. 280).
Ehlers, 1913, p. 444, pl. xxvii, figs. 5-11; nec Fauvel, 1923, p. 68, fig. 24, h-o.
Occurrence.—Station 15, 18° 21’ N., 75° 25’ W., gear 2-m. stramin net, depth
1250-0 m. (1).
Station 27, 16° 22’ N., 86° 40’ W., off west end of Roatan Is., gear 2-m. stramin
net, depth 600-0 m. (1).
Station 28, 16° 44’ N., 85°
42’ W., north of Bonaccia Is.,
gear 2-m. stramin net, depth
1100-0 m. (1).
Remarks.—The largest of
these specimens measures c. 8
mm. in length for 28 chaetigers.
The exact number of elytro-
phores is difficult to determine,
but I estimate 11 pairs of scales.
There are no prostomial peaks.
The two pairs of large, con-
tiguous eyes have white lenses.
The palps and tentacles are
smooth. Only asingle detached
scale (text-fig. 280, A) remains.
This is more or less round, and
dotted with short, rather stout
tubercles with truncated tops.
There are in addition a few
slender papillae scattered about
the scale (text-fig. 280, B).
The feet are as figured by
Ehlers. ‘The dorsal bristles are
of two kinds: (1) short, trans-
parent, backwardly curved and
carrying rows of pectinae (text-
fig. 280, c); and (2) very large,
straight, transparent bristles
carrying pectinae arranged ina
spiral and having simple blunt Texı-rıG. 280.—Harmothoé benthophila: A, elytron ;
tips (text-fig. 280, p). The ® tubercles and papillae from scale; c, dorsal bristle; p,
giant dorsal bristle; ®, upper ventral bristle ; r, lower
ventral bristle.
280
0:05mm 0:02 mM
upper and middle ventral
bristles are very long and
slender, carry rows of pectinae and have faintly bidentate tips (text-fig. 280, 2).
The lower ventral bristles are shorter, stouter, and have smooth or very faintly
bidentate tips (text-fig. 280, Fr). The body ends in a small, pygidial appendage.
Except that I find the ventral bristles faintly bidentate, these specimens
agree closely with that described by Ehlers from 18° N., 30° W., taken at a depth
of 3000-0 m. Those seen by Fauvel, on the other hand, from off the Azores
350 Novirates ZooLoGIcAE XLI. 1939.
have both kinds of dorsal bristle with blunt, double-pointed tips, a difference
which caused Fauvel (1916, p. 24) to erect his var. bimucronata. In 1923
(loc. cit.) he abandons the varietal name and gives the varietal characters as
those of Ehlers’s species. The specimens from 10° N., 27° W., attributed
by me (1936, p. 87) to Ehlers’s species, belong in fact to Fauvel’s var.
bimucronata.
Whether H. benthophila and H. benthophila var. bimucronata are stages in
the development of the same species or represent the juvenile phase of different
species there is at present no evidence to decide.
2. Harmothoé joubini Fauvel var. rosaurae var. nov. (text-fig. 281).
Harmothoé joubini, Fauvel, 1914, p. 54, pl. iii, figs. 6-12.
Occurrence.—Station 49, 28° 25’ N., 13° 34’ W., between Fuerteventura Is.
and Africa, gear Agassiz trawl, depth c. 1300 m. (4).
Description.—Only one specimen is complete, and this measures 17 mm. by
3 mm. without the feet for 36 chaetigers. In spirit the head is colourless, but
the back is purplish red, and there is a pair of ventro-lateral purple stripes.
The elytra are colourless. There are 15 pairs and the last 5-6 chaetigers are
without scales.
28 The head has well-developed prostomial peaks, and
two pairs of very small laterally placed eyes, the anterior
pair lying about half-way up the head. There is a
massive median tentaculophore, but the median tentacle
is lost. The lateral tentacles are about as long as the
head. The palps reach back to about the 5th chaetiger.
I see no facial tubercle. The tentacular cirri are lost.
The elytra are large, soft, colourless and completely
smooth except for a patch of small acuminate tubercles
near the scar of attachment. Pseudo-elytrophores are
present in the cirrigerous segments.
The dorsal cirri are all detached, but a few lie loose
among the bristles. They would seem to reach about to
the end of the bristles. They, as also the tentacles and
IN 5 palps, are smooth. Both rami of the feet are extended
by long, sheathed acicula. The ventral cirri reach to the
0:05mm 005" end of the foot. The dorsal bristles are rather short,
sword-like and faintly pectinate. The ventral bristles
Tuxr-rre. 991. Me large, expanded distally, almost smooth and with
Harmathoö joubini var, curved tips. They may be either unidentate or very
rosaurae : A, unidentate faintly bidentate (text-fig. 281, a and B). The lower
ventral bristle ; 8, biden- . -
A rates ventral bristles are short and always unidentate.
ate ventral bristle. nee R
The pygidium is damaged.
Remarks.—These specimens are clearly very close to Fauvel’s H. joubini
taken at a depth of 1473 m. about 60 miles off Cape St. Vincent. They differ
(1) in the absence of a facial tubercle ; (2) in having a different type of ventral
bristle. In Fauvel’s species the ventral bristles are strongly bidentate. These
differences are not, in my opinion, of more than subspecific value. Both Fauvel’s
specimens and mine are deep-water forms, being taken at a depth of over
1000 m.
Novirates ZooLoGIcAE XLI. 1939. 351
GENUS EUNICE CUVIER 1817.
3. Eunice rosaurae n. sp. (text-fig. 282).
Occurrence.—Station 34, 12° 05’ N., 61° 49’ W., off St. George, Grenada,
gear Agassiz trawl, depth 720-800 m. (5).
Description. — The largest specimen which appears almost complete measures
165 mm. by 5 mm. at the widest part without the feet for 135 chaetigers. The
body is of the usual form, more or less cylindrical anteriorly and flattened in
the hinder region. There are traces of brown, transverse dorsal markings.
The head is bilobed and
the median tentacle
reaches back to the 4th—
6th chaetiger. The inner
laterals have about
three-quarters of this My
length and the outer IN
laterals about one-half. —
There is a pair of large
eyes lying just behind N
the gap between the or c
inner and outer laterals. O-lmm.
The tentacular cirri are uvam
equal in length to the
buccal segment. The
tentacles and tentacular
cirri are smooth, but
when contracted they
may give the appear-
ance of being annulated.
The buccal segment is
about equal in length
to the first two chae-
282
tigers. u
The gills begin at
the 3rd chaetiger and ware
continue to about the
35th. They are very
poorly developed and
consist of a single fila- TEXT-FIG. 282.—Hunice rosaurae: A, fifteenth feet; B, comb
bristle; c, ventral bristle; p, subacicular chaeta ; E, upper jaws ;
F, lower jaws.
ment, which is never
more than about one-
third the length of the dorsal cirrus (text-fig. 282, a). The feet are supported
by two or three black acicula. The dorsal bristles consist of capillary bristles
and comb-bristles with rather numerous small teeth and a long lateral filament
(text-fig. 282, B). The ventral bristles (text-fig. 282, c) are compound, bidentate,
hooded hooks. There is a black subacicular chaeta (text-fig. 282, D) which
is bidentate and first appears at about the 32nd chaetiger. The dorsal and
ventral cirri are well developed.
I figure the jaws (text-fig. 282, E). The lower jaws (text-fig. 282, F) have a
calcareous cutting edge. The dental formula is 6 — 7:8 + 7 — 5.
352 Novirates ZOOLOGICAE XLI. 1939.
The anus is terminal and the body ends in a pair of short, ventrally placed,
anal cirri.
The animal builds a parchment-like tube.
Remarks.—This species belongs to the group of Eunicids with single, cirriform
gills, but I know none that has its branchiae so much reduced and confined to
the anterior region. It shows affinities with H. cariboa Grube which differs in
having the single gill filament confined to about 30 segments of the hinder
region instead of the anterior.
Branchioasychis gen. nov.
Head in the form of a rounded plate having a wide border deeply incised
laterally but otherwise entire. Cephalic keel flat, not clearly distinguished.
Nuchal organs short and crescentic. Pygidial region not yet known. No
ventral bristles in the Ist chaetiger. Hooks begin at the 2nd chaetiger. Dorsal
bristles of three kinds. Anterior segments without a collar. Dorsal surface of
6th to 9th chaetigers thickly covered with long branchial papillae. Anterior
region highly glandular.
Genotype: Branchioasychis colmani n. sp.
4. Branchioasychis colmani n. sp. (text-fig. 283).
Occurrence.—Station 22, 17° 28’ N., 88° 11’ W., Belize Harbour, gear 2-ft.
dredge, depth 6 m. (1).
Description.—The specimen is incomplete posteriorly and measures 75 mm.
by 3mm. for 15 chaetigers. The head (text-fig. 283, A) and the first 3 segments are
sprinkled with dark purple markings. Otherwise there is no colour. The
cephalic plate has a wide border, which at the sides merges imperceptibly into
the head. It is notched laterally and forms a deep hood behind. It is quite
smooth at the edge. There is no raised keel and the nuchal organs are short and
erescentic. There are pigment marks on the front border of the head, but I
cannot tell whether these are ocelli, or merely part of the general pigmentation.
The buccal segment is equal in length to the Ist chaetiger. The first three
chaetigers are divided by rings into two unequal parts, the anterior part being
about twice as long as the posterior. At the 4th chaetiger this ring is no longer
apparent. Anteriorly the feet lie near the front border of the segments. Farther
back they seem to move more into the middle of their segments, but, as the
limits of the segments are very indistinct except for the first few chaetigers, their
exact position in their segments in the hinder region remains uncertain.
From the 6th to the 9th chaetigers the dorsal surface is covered with a fur-like
investment of long, filiform papillae which I take to be branchiae (text-fig. 283, B).
They are very different from the regularly arranged vascular caeca found in
Johnstonia. These branchiae decrease in density from before backwards.
They are very thick in the 6th chaetiger and very sparse in the 9th.
There are no ventral bristles in the Ist chaetiger. The dorsal bristles are
of three kinds : (1) simple, plain capillaries (text-fig. 283, c) ; (2) widely bordered
capillaries (text-fig. 283, D) ; (3) barbed bristles (text-fig. 283, m). The uncini begin
at the 2nd chaetiger and are of the usual kind, having a cap of denticles, a main
tooth and subrostral barbules (text-fig. 283, F). The shaft is only slightly
expanded below the head.
Novirates Zoonoarcar XLI. 1939, 353
The Ist and 2nd chaetigers are more or less uniformly glandular. The
3rd, 4th and 5th show a definite glandular pattern dorsally, which is shown in
the figure. Ventrally the body is uniformly glandular up to the 7th chaetiger,
except in the 3rd chaetiger, where the glands are interrupted. From the 7th
c D
0:02mm 0-02mm
aE Mm iy
TExT-rıG. 283.—Branchioasychus colmani: A, anterior end from above, showing pigmen-
tation; 8, dorsal view of 3rd to 9th chaetigers; c, dorsal bristle; p, widely bordered capillary
bristle; e, barbed bristle; F, ventral hook.
chaetiger the glandular areas are confined to the uncinigerous pads and to a series
of lateral spots joining the pads from segment to segment.
The anal region is missing.
Remarks.—In spite of the absence of a hinder region the well-developed
branchiae justify the creation of a new genus, As far as I know, the only mention
26
354 NOVITATES ZooLoGIcAE XLI. 1939.
of a branchiate Maldanid in the literature of the Polychaeta was made by Andrews
(1892, p. 294), who in his paper on the Polychaeta of Beaufort, North Carolina,
under the heading of Maldane elongata Verrill writes: “A fragment belonging,
perhaps, to this genus was peculiar in having the dorsal surface of several somites
thickly set with slender papillae.” It is quite possible that Andrews had an
example of the present species. Branchioasychis, apart from the branchiae,
shows many affinities with Asychis Kinberg 1855.
REFERENCES.
Andrews, E. A., 1892. Report upon the Annelida Polychaeta of Beaufort,
North Carolina. Proc. U.S. Nat. Mus., xiv, 1891, pp. 277-302, pls. xii-xviii.
Ehlers, E., 1913. Die Polychaeten-Sammlungen d. D. Südpolarexped, 1901-
1903. Deutsche Südpolarexped., xiii (4), pp. 397-598, pls. 21.
Fauvel, P., 1914. Annelides Polychetes non pélagiques. Res. Camp. Sci.
Monaco, fase. xlvi, pp. 432, pls. 31.
— 1916, Annélides Polychetes pelagiques. Res. Camp. Sci. Monaco, fasc.
xlviii, pp. 152, pls. 9.
—— 1923. Polychetes Errantes, Faune de France, Paris, v, pp. 488, text-figs.
181.
Monro, C. ©. A., 1936. Polychaete Worms, II. “ Discovery’ Rep., xii, pp. 59-
198, text-figs. 34.
Qu
Novirates ZootocıcAar XLI. 1939. 35
NOTES ON ORIENTAL THECLINAE (LEP. LYCAENIDAE) WITH
DESCRIPTIONS OF NEW SPECIES.
By N. D. RILEY,
Department of Entomology, British Museum (Natural History).
DER Theclids with metallic green males grouped by Seitz (Macrolep. i, p. 269)
under Zephyrus have proved a source of considerable difficulty and, as
Seitz himself remarks, will require very careful study before they can be satis-
factorily determined. The juxtaposition, in the British Museum, however, of
the types of species described by Leech, Butler, Murray, Oberthur and others
has permitted certain conclusions to be reached, purely upon the basis of a
comparison of external features, that may be of value to students of this very
attractive group of species.
In the following notes the species are dealt with, for convenience only,
according to the arrangement in Seitz. To these notes have been added descrip-
tions of several other far eastern Theclids of related genera which appear to have
been hitherto unrecognized. As Hemming has shown (1934, Generic Names
Holarctic Butterflies, p. 115), Zephyrus Dalman (1816) sinks to Thecla Fabricius
(1807), both having the same genotype.
1. Thecla orientalis Murray.’
Thecla diamantina Ob. and T. cognata Staud., here placed as synonyms of
orientalis, have nothing to do with this species (see below).
“ab. suffusa ” Leech appears to be a good species. Except for the narrow
forewing and very wide hindwing black borders, the male upperside is entirely
(not only in outer third) green, of the same tint as in orientalis, but very much
duskier, almost smoky. The underside of the female is dark brown as in the
male ; on the upperside it is the same as the female orientalis. It is well figured
by Leech (Butt. China, pl. 27, fig. 14). With it flies a race of 7’. orientalis which
may be characterized as follows :
la. Thecla orientalis leechi subsp. nov.
3d. Indistinguishable on the upperside from typical orientalis. Recogniz-
able at once by the underside, the ground colour of which, by comparison, is
almost white, i.e. very pale silvery grey, thus rendering the white markings,
which are in fact somewhat reduced, very inconspicuous.
Holotype ¢ “ Ta-chien-lu, 7500 ft., A. E. Pratt collection, July 1889 ” (ex
coll. Leech) ; allotype 2 “ de Ta-tsien-lou & Moupin, chasseurs indigénes, etc.,
1892 ” (ex coll. Oberthur) ; both in British Museum.
This subspecies appears to occur throughout Szechuan, as indicated by Leech
(loc. cit.) and also in Yunnan, whence Oberthur had it from Tsekou, In Central
China (Chang-Yang) it is more strongly marked on the underside, As in many
species of this group a form of the female occurs in which the cell and area la
1 For references to names see Seitz, Macrolep. i, pp. 323-328,
356 NOVITATES ZOOLOGICAE XLI. 1939.
on the upperside of the forewing are blue. This may be known as ® f. coelestina
nov., the type being a female collected by Kriecheldorf at Moupin in June 1890.
2. Thecla taxila Bremer and Grey.
The names “ab. aurorina Ob.” and “ ultramarina Fixsen ” should be
excluded from consideration, as they do not in fact apply to any form of this
species (see below).
This species falls naturally into four subspecies, viz. :
(a) T. taxila taxila. A small form from Manchuria, Hokkaido and the
higher elevations of Honshu.
(b) T. taxila sachalinensis Matsumura.
(c) T. taxila koreana subsp. nov. A larger form, deeper green above and
with wider black borders; intermediate below between T. t. taxila and T. t.
japonica. Korea: Seoul (3 holotype) and Gensan (2 allotype).
(d) T. taxila japonica Murray, occurring apparently throughout Japan
except where 7’. t. taxila occurs, and in Formosa.
The original description of taxila having been based on a female with
unicolorous upperside, Kardakoff’s name unicolor for this form is superfluous ;
similarly © f. bellus Kard. is the same as the older 9 f. regina Butler ; and 9 f.
quercus Kard. corresponds with the 9 f. fasciata Jans. of subspecies japonica,
The assertion (Seitz i, p. 270) that fasciata has a coppery stripe along the inner
margin of the forewing upperside is groundless; the type specimen bears no
such feature, nor does the original description make any reference to it.
3. Thecla aurorina Ob.
Oberthur’s type female of aurorina proves to be a female of the species
dealt with by Seitz (i, 270), a 7. brillantina Staudinger. The Central Chinese
subspecies has been described by Seitz (loc. cit.) as T'. coruscans jankowskii ;
the Japanese as T'hecla aino Matsumura.
4. Thecla smaragdina Bremer.
Oberthur’s type male of diamantina proves to be a male of this species. A
Japanese female is well figured by Matsumura (Ill. Common Insects of Japan,
i, pl. xii, fig. 15) as brillantina Staud. It is doubtful whether Japanese
specimens are subspecifically separable from those from the mainland; I can
find no constant character.
5. Thecla coruscans Leech.
The name is consistently spelt corruscans wrongly by Seitz. The species is
very constant, but in Yunnan a slightly different race occurs which may be
described as :
5a. T. coruscans dubernardi subsp. nov.
d. Transitional between 7’. coruscans coruscans and 7’. souleana (see below).
The upperside resembles the latter, but the underside retains the warm brown
tones of coruscans. The markings on the underside retain the relative positions
of those of coruscans, but the white lines are about half as wide and less regular,
and the orange anal markings are reduced by about one-third in extent. No
discocellular bars.
Holotype 3g, Tse-kou, ex coll. Oberthur.
NOVITATES Zoonoaroarn XLI. 1939. 357
6. Thecla souleana sp. nov.
3g. Closely resembling coruscans on the upperside, but of a duskier colour
suggesting that of 7’. zoa from Darjeeling ; black borders rather broader, apex of
forewing broadly black and costal area mostly black; green of the hindwing
nowhere passing above vein 6; cilia prominently white. On underside ground
colour uniform rather cold grey-brown ; discal white lines prominent but much
narrower and dark edged on inner side ; discocellular lines well developed, dark,
narrow, faintly white edged (this feature is absent from coruscans). Forewing
with much darker wedge-shaped, curved, submarginal band tapering apicad,
very faintly pale-edged ; hindwing with faint exceedingly short white line towards
base, near costa, orange anal markings about half the size of those in coruscans,
and the very faint wavy postdiscal line correspondingly approximated to the
margin, a powdery submarginal band of grey-blue scales ; postdiscal and sub-
marginal bands both parallel to margin, not anteriorly divergent as in coruscans ;
front end of discal line 9 mm. from base of wing. Length of forewing 15 mm.
(18 mm. in coruscans).
Holotype g and 4 paratypes all in British Museum from Yarégong in Talu
district of Szechuan, West China.
7. Thecla desgodinsi Ob.
The Oberthur collection contained two males and one female in addition
to the type. These are from Tien-tsuen, Siao-lou and “ Yunnan.” They
prove to represent a subspecies of the Indian 7’. duma Hew.
8. Thecla ultramarina Fixsen.
Thecla ultramarina Fixsen (= cognata Stgr.) is a species quite distinct from
taxila Brem. and orientalis Murray with both of which it flies in company, both
in Japan and on the neighbouring mainland of East Siberia. The male has the
same upperside coloration as orientalis, but has the marginal band of the hindwing
upperside much broader between the apex of the wing and the shorter of the two
tails, and area 6 of the hindwing (which is blue-green in orientalis) almost com-
pletely filled by the expansion of the costal black border. The female upperside
is like that of orientalis, The underside of both sexes is definitely brownish,
not plain grey, and the female often has orange markings about the tornal area
of the forewing. Japanese examples, which represent a doubtful subspecies,
have been named jezoensis by Matsumura ; small females of this with very brown
undersides and rather curved discal lines are referable to 9 f. jozanus Mats.
9. Thecla lutea adusta subsp. nov.
Smaller than the typical Japanese race and slightly darker beneath, particu-
larly characterized by the proximity of the discal silvery lines to the cell. In
many specimens the inner of those two lines almost touches that at the cell-end.
All the silvery lines also are rather more prominent. On the upperside the
black apical border is rather wider.
19 &, 13 2 in British Museum from Siao-lou (¢ holotype), eastern frontiers
of Tibet (Q allotype), and Tien-tsuen, all ex. coll. Oberthur.
A variety, of which two females are included in the above series, in which
the silver is replaced by pale greyish ochreous may be known as ab. tatsienluica
nov.
358 Novıtares Zootocicar XLI. 1939.
10. Thecla icana setschuanica subsp. nov.
Zephyrus icana Moore, Leech, Butt. China ii, 380, pl. 27, fig. 4.
The race from Szechuan, West China, differs from typical icana in being
blue rather than green-blue on the upperside in the male and much more exten-
sively blue in the basal half of the forewing in the female. The underside shows
no constant difference, though on the average the tranverse bands are narrower,
and the marginal markings much better defined, the marginal lines on the
hindwing grey-brown and never orange-tinted above vein 3.
Holotype ¢ and allotype 9, Ta-tsien-lu, 10,000 ft., A. E. Pratt coll., July
1889, ex Leech coll. A long series of both sexes in British Museum from Ta-
tsien-lu, Omei-shan, Siao-lou and Tibetan borders of Szechuan.
11. Thecla ouvrardi sp. nov.
3. Upperside, dark brown, cilia white; forewing with small triangular
yellow spot on discocellulars, the apex directed towards termen, and an ill-defined
roundish dark-yellow patch beyond, in area 3 ; hindwing with fine blue marginal
line in lc and 2, and the tips of the cilia there black. Underside silvery white,
anteciliar line black, cilia white, a black bar at the discocellulars. Forewing
with a strong black postdiscal line (about 1 mm. wide, and continued on hind-
wing), a faint dusky postdiscal band from costa to vein 4, where it is dislocated
and continued as isolated dark spots (2) in 2 and Ic, a shadowy terminal dusky
line. Hindwing discal line commences at middle of costa (about 1 mm. wide)
and runs, outwardly convex, to cell bar, thence, considerably narrowed, past the
end of cell to reach postdiscal line on vein 2, the subsequent W-mark delicate
but distinct ; black marginal ocellus in 2 placed in a square yellow spot sur-
rounded by a fine silver line, the yellow continued broadly (though divided by
silver along vein 2, and enclosing a wedge of silver in 1c) to anal angle, which
is black, and thence narrowly to end of vein la; tip of cilia in Ic and 2 black.
Length of forewing, 20 mm.
Described from the unique holotype from the Oberthur collection labelled
“ Rahand, Yunnan. Recu du pere Ouvrard, missionaire apostol. & Oui-si ou
Wei-si, Yunnan, Chine, en janvier 1917.”
The species is very closely related to 7. arisanus Wileman, from which it
differs most obviously in the much better development of the markings of the
underside. The upperside is exactly like that of the female of arisanus, the
male of which is devoid of markings on the upperside and was treated by Wileman
as a form of the female having “ a dull unicolorous brown ” upperside (1909,
Ann. Zool. Jap.,7: 92). The 3 genitalia of T. arisanus and 7’. ouvrardi, examined
without dissection, show close similarity, but whereas the median lobe of the
uncus in the former is parallel-sided and somewhat truncate, in owvrardi it is
bluntly Y-shaped. 1’. owvrardi will probably prove to be a mainland representa-
tive of arisanus. T. picquenardi Oberthur also is closely related but has an
additional (submarginal) band on both wings below.
12. Thecla yangi sp. nov.
39. Upperside very pale silvery bluish, the forewing costal and apical
areas broadly dark brown shading in the pale area and extending to tornus
rather broadly in 9, less so in 4, cilia white ; hindwing apex dusky, the veins
Novirates ZooLogIcAaE XLI. 1939. 359
finely black, a fine black anteciliar line preceded by a narrow white line ; cilia
white ; tail black, tipped with white; submarginal black spots in le and 2.
Underside greenish ochreous, both wings crossed by a rather broad (1 mm.)
silvery line and bearing a series of postdiscal inwardly silver-edged black spots, a
white terminal line, a black anteciliary line and white cilia; on the forewing
the transverse line runs from the costa at about three-fourths towards tornus,
but is broken on vein 2 to end on dorsum at three-fourths, and the postdiscal
series consists of one spot each in areas 2-4 (decreasing rapidly in size) and a pair
in 1b ; on hindwing the silver line runs from costa at two-thirds to vein 2, where
it touches the silver edging of the postdiscal spot in 2, thence in a curve to vein
1 and to inner margin at midway ; a complete series of 8 postdiscal spots, of
which those in areas Ic, 2 and 3 are the largest ; anal lobe black, and a conical
marginal spot (outwardly edged or obscured by silver) in le and 2; a broad,
diffuse orange band occupying most of the area between postdiscal series and
margin, but narrowing towards apex. Length of forewing, $ 16 mm.; 2 20 mm.
Described from 1 ¢ labelled “China, Foochow, 1937-38, M.S. Yang” in
the British Museum (holotype); and 1 @ labelled “China, Kwantung, Lung
tao shan, 15.iv.18, R. Mell ” in the collection of Dr. Mell (allotype 2).
13. Satsuma albilinea sp. nov.
Closely allied to S. circe Leech, of which, indeed, it is conceivable that it is
but a seasonal form. Both sexes resemble that species on the upperside, except
that the fringes are entirely unspotted, and the outline of the wings much
smoother. The hindwing in the female has the whole of the basal half blue.
On the underside the pattern is as in circe, but the forewing and the distal third
of the hindwing are rather light ochreous, the dark discal patch on the forewing
and the dark discobasal area on the hindwing being edged by a continuous
narrow white line. It appears to be quite as common as circe, and flies in the
same region, namely Ta-tsien-lu, W. China, from which place the types and a
series of both sexes are in the British Museum ; also 1 & from Yarégong, all ex
coll. Oberthur.
14. Strymon abdominalis persica subsp. nov.
9. Upperside—A prominent white marginal line on hindwing from anal
lobe at least to vein 3, continued inconspicuously to apex. Underside.—Post-
discal transverse white lines prominent, very irregular, internally strongly black-
edged ; orange lunules narrow and pale yellow; hindwing marginal series of
markings well developed and extending fully to apex of wing, the innermost
series consisting of prominent and comparatively large white cones. The spot
in the centre of the ocellar spot in 2 is much larger and more rounded than in
other races of abdominalis, and the same also applies to the black lobe spot.
This subspecies is very well illustrated by the figure of a female (fig. 3) on
the plate accompanying Colonel Peile’s account of the Butterflies of Mesopotamia
and N.W. Persia (J. Bombay N.H. Soc., 28, pp. 135-155 ; 353-370; 1921-2).
The specimen there figured is wrongly referred to subsp. gerhardi Stgr., and it is
obviously distinct from the male of that subspecies illustrated on the same plate
immediately below it (fig. 4). The 9 holotype and two 9 paratypes were all
taken by Lt.-Col. H. D. Peile in the Karind Gorge, N.W. Persia, at an altitude
of 6,000 ft. in July 1918—the holotype on the 14th, the others on the 13th and
17th respectively.
360 Noyirares ZooLoGIcAE XLI. 1939.
15. Strymon persimilis sp. nov.
The upperside of the unique J specimen of this species resembles almost
exactly the fixseni form of the female of S. eximia (Seitz, Macrolep, i, pl. 72i),
but the orange patches are a little less strongly developed—the forewing patch,
for example, extends from vein 2 not quite to vein 6 and does not enter the
cell at all. The androconial patch is relatively very large, being over 2 mm.
long on a forewing 16 mm. long. On the hindwing the red patch extends evenly
from the lobe to vein 3. The underside is best compared with that of percomis
Leech (l.c. pl. 73a), which it strongly resembles both in colour and pattern. On
the forewing, however, the silvery line of narrow lunules is completely missing ;
and on the hindwing this line is composed of much fainter and flatter lunules,
so that the inner edges of the red lunules near the anal angle are almost straight.
Yunnan, R. P. Delavay (ex coll. Oberthur) ; type in the British Museum.
16. Strymon dejeani sp. nov.
In size this species is no larger than prunoides Ster. The upperside is
uniformly very dark brown ; the cilia are white. Both wings are very rounded,
the hindwing having a single narrow tail at vein 2, only the faintest projection
at vein 3, and a very slight lobe. The male patch on the forewing is small,
narrow and very short. The underside is ochreous with a greyish tinge, and a
prominent, interrupted, black-edged white line crosses both wings near their
margins and parallel to them (in the same position as the corresponding line in
Strymon acaciae), not being very sharply angled anywhere. The hindwing has
a prominent pure white marginal line that just extends on to the forewing ;
the lunules are dull yellow and only extend clearly halfway round the margin ;
the black anal spot is very small, the other black spot narrow but moderately
tall, the intervening grey spot broader and shorter than it.
Holotype & Siao-lou, chasseurs indigénes du Pére Dejean, 1903 ; allotype 9,
same data, 1902; 3 & paratypes also Siao-lou, 1 Moupin ; all ex coll. Oberthur,
in British Museum.
17. Strymon ledereri hyrcanica subsp. nov.
The large typical form of ledereri which flies in Asia Minor, the Caucasus,
etc., is replaced in North-East Persia by a much smaller subspecies with a forewing
measurement of about 13 mm. only (16-17 mm. in typical ledereri). In this
subspecies the underside is usually rather lighter and greyer than in typical
ledereri, the black spots are usually a trifle smaller and much less broadly
ringed with white, and the orange marginal spots frequently though not always
relatively larger.
Holotype & and allotype 9 “ N. Persia, ex Staudinger ” in British Museum.
Other specimens in the British Museum are from “ Hyrcania ” (ex Staudinger) ;
Iskander-Kul, Hissar Mountains, 22-25.vi.92; Shahkuh ; Ordubad, 8.v.83;
and Armenia.
One male from Shahkuh is aberrant in that the forewing is devoid of markings
on the underside, except for the submarginal black spots in 2 and 3.
18. Sinthusa rayata sp. nov.
3g. Nearest S. chandrana on the upperside, but with a patch of brilliant
blue at the base of the forewing, of small extent, and the similar blue on the
Novirares ZooLoGIcAE XLT. 1939. 361
hindwing divided up by long black rays along the veins, extending right to the
margin. The abdominal fold is grey. The wnderside is very light bluish-grey
with bright ochreous markings, namely a narrow double stripe at the end of the
cell and an irregular scarcely connected series of narrow discal spots on each
wing. The hindwing in addition has two spots in the cell, a third above them
near the costa, and a fourth against the inner margin ; the black subanal ocellus
is quite small and is surmounted narrowly by an iridescent blue line and then
a yellow one, both of which extend to the black lobe, the yellow one even farther.
The female is unknown.
Habitat: Tien-Tsuen, W. China. Holotype & in British Museum.
19. Rapala caerulea and 20. R. micans (Bremer & Grey) 1853.
Thecla micans Bremer and Grey (p. 8) and Thecla caerulea iidem (p. 9), Schmett., N. China 1853 ;
Menetries, Cat. Mus. Petr. Lep., i, pl. 4, figs. 3 and 4, 1855.
The very clear descriptions given by Bremer and Grey are accompanied by
references to a plate which, however, did not appear with the publication.
Subsequently, on plate 4 of Menetries’ Cat. Mus. Petr. Lep., i, there appeared
figures of these two species unaccompanied by text. Fig. 3 purports to
represent Thecla micans and fig. 4 Thecla caerulea. It is quite obvious, however,
that the names are transposed on the plate, if the figures be compared with the
descriptions where, moreover, “ fig. 3’ is stated to represent 7’. caerulea and
“fig. 4” T. micans. The current usage of these names is clearly based upon
the Ménétriés plate alone and is wrong. Thecla micans Bremer and Grey is
synonymous with Thecla nissa Koll. (1848), or perhaps represents a local race
of that species ; and T'hecla caerulea Bremer and Grey is the oldest name for the
species hitherto known as T'hecla micans. Both species are properly to be
referred to Rapala Moore (1881).
362 NOVITATES ZooLoagicaE XLI. 1939.
SIPHONAPTERA COLLECTED BY DR. GERD HEINRICH IN
BURMA.
By KARL JORDAN.
(With 13 text-figures.)
AS we had hardly any specimens of fleas from Burma the collection made by
Dr. Heinrich was most welcome, and I express here my gratitude to him
and his companions to have taken the great trouble of preserving the external
parasites found on the mammals obtained. The majority were collected on Mt.
Victoria in the Pakokku Chin Hills, a range to the west of the Irawadi R. ;
others were found farther south in western Burma on Mt. Popa and some other
places in the Prome district ; and a small but interesting portion of the material
was obtained in north-eastern Burma near Mandalay and Maymyo. Ten
species altogether are represented in the collection, of which three species and a
subspecies are new to science and nearly all the others new for Burma.
1. Ctenocephalides felis orientis Jord. 1925.
20 km. north of Mesktila, on Felis chaus, 20.1.38, 1 g, 1 2; Maymyo,
800 m., 6.xii.37, on Paradoxurus paradoxus, 1 9.
2. Paraceras hamatus sp. nov. (text-figs. 284-286).
Close to P. melinus Jord. 1925, belonging like that species to the tropical
group of the genus, which differs from the palearctic group in the proboscis
reaching at most to the apex of the
trochanter, in the longer spines of the
pronotal comb and the longer stylet
of the 9.
32. On frons a row of three long
eye-bristles, which are larger in $ than
in 2; farther forward and upward a
second row of smaller bristles, 7 in &
and 2 or 3 in 9, the lowest bristle of
this row slightly above the level of
the upper eye-bristle, the row not
being continued to the oral margin
(text-fig. 284, 2). On occiput, in g, a
single antemedian bristle about the size of the second bristle (from above)
placed at the anterior margin of the antennal groove ; in middle an oblique
row of 3, of which the lowest is large, the second about the size of the
subdorsal one of the posterior row, and the third about half the size ; the posterior
row consists of 5 bristles each side, the lowest being very large, the others as
usual much smaller, interspace between first and second much smaller than
between second and third. In 2 the anterior occipital bristle absent; the
median row consists of 2 bristles, of which the second is small ; posterior row
Novirares Zootoarcarn XLI. 1939. 363
as in g, but the bristles smaller, and there is an additional longish bristle below
the large one. Proboscis reaching to apex of trochanter or nearly.
Dorsal spines of pronotal comb longer than the pronotum, the spines
measured from the anterior margin of the dark vertical band placed at the base
of the spines. Bristles on the two sides together : on pronotum 11 to 14, usually
12; mesonotum 40 odd small ones in a row and scattered over the dorsal and
basal surface, a posterior row of 10 to 12, usually 10; metanotum 13 to 22,
11 to 14; spines on underside of mesonotum 9 to 12. Bristles on mesopleura
ZUR
a
POSS =
\
\\
det mics \)
8 to 10; on metepimerum 8 to 10, usually 4,4, 1. Short apical spines (on the
two sides together) : metanotum 2 to 4; abdominal tergum I 2, II 2 to 4, usually
3, III 2 or 3, usually 2, IV 2, V 1 to 3, usually 2 in g and 1 in 9, VI 0 or 1,
usually 0. Bristles on abdominal terga: on I in g 11 to 13, 10 or 11, in 2 13
to 19,9 or 10; II g 11 to 13, 14 to 16, 2 12 to 14, 13 or 14; III ¢ 10 to 12,
15, 2 12, 14; IV g 9 to 12, 14 or 15, Q 11 to 13, 13 or 14; VII 38 or 9, 12
to 15, 9 7 to 9, 10 or 11. On sterna III to VI in ¢ 6, rarely 7 on one segment,
in © 6 to 8, usually 8. Lower antepygidial bristle of & more than half the
length of the middle one.
In hindtarsus of 9 two slender apical bristles at posterior side of segment
II reach much beyond III, a slender bristle in subapical notch of posterior side
364 Noviratrs ZooLoGIcAE XLI. 1939.
does not quite extend to middle of III; on posterior side of III two slender
apical bristles reach about to middle of V. Measurements: midtarsus ¢ 16 to
18, 22, 14 to 16, 8 or 9, 20 or 21; 9 17, 22 to 24, 14 to 16, 8 to 10, 20 to 22;
hindtarsus ¢ 56 to 66, 32 to 36, 18 or 19, 11 or 12, 22; 2 57 to 63, 30 to 36, 18
to 21, 11 or 12, 23 to 25. The difference in length between segments I and II
of the midtarsus is noteworthy.
Modified Segments. g. On tergum VIII 16 or 17 long bristles, most of
them in the dorso-marginal area ; on inside a spiculose area from anterior dorsal
angle to near the last marginal bristle, the area being nearly six times as long
as broad, the spicules dense at the boundary away from the margin, the boundary
being partly marked by an incrassate stripe and on outer side by a row of long
bristles. Sternum VIII (text-fig. 285) of the usual shape, of nearly even width,
directed obliquely upwards and apically curved distad, bearing a large mem-
branous flap slit into numerous filaments (diagrammatically indicated in the
figure) ; at the ventral margin two longish bristles as in figure, or the lower
bristle accompanied by an additional longish one, or two bristles close together
and the third absent; near apex a very thin ventral bristle. Process P of
clasper touches the anterior margin of the digitoid F at approximately two-
thirds ; digitoid F about as broad as long, distally strongly rounded, in outer
half of ventral margin a row of minute bristles ; at upper end of distal margin
a strong bristle, below it some small slender ones and close to and above it a small
one, which is either short and rather stout or much thinner than in figure; on
inner surface the usual row of bristles, here 7 or 8. Apex of vertical arm of
sternum IX with three projections, two posterior and one anterior ; the most
characteristic feature of the ventral arm is the apical division, which is gradually
curved down like a claw, the pair of claws of the two sides being very con-
spicuous ; the nearest approach to this modification of the distal section of
IX.st. we find in P. crispus Jord. & Roths. 1911. Phallosome of the same
type as in other species of the genus, the ventral paramere being broad in the new
species.
9. Ventral portion of apical margin of sternum VII (text-fig. 286) rounded ;
this part is only a small portion of the margin, the greater part of which is strongly
slanting. Tergum VIII in all 8 99 with a long and a short bristle below the
stigma ; on outer surface of widened area altogether 12 bristles, some of which
small ; on inner side two bristles below apical angle and two or three short stout
ones on the side; apical margin incurved twice, the angle in between 90°.
Stylet and spermatheca as in the other tropical species ; the small projection at
the end of the tail of the spermatheca is not always so distinct as in the figure.
Length: & 2-9-3-3, 2 3-8-4-4 mm.; hindfemur 3 0-59-0-69, 2 0:64-0:67
mm.
Mt. Victoria, Pakokku Chin Hills, 2200 m., on Ratufa gigantea, 21.vi.38,
2 33, 6 99, type 3; also on Martes, 2.v.38,1 g, 2 29.
When I noticed in Dr. Heinrich’s material a species of Paraceras, I had
great hopes that it would turn out to be P. melinus Jord. 1925 ; I am sorry the
type of P. melinus, a 9, does not agree with the series of Heinrich’s 8 99.
When I received the specimen from Oldfield Thomas, who had forgotten locality
and host, it seemed to be worth while to describe it in spite of the absence of
data, because it deviated much from the species known to me in the shorter
proboscis and the long stylet. The specimen is in rather poor condition, but
Novitates ZooLocıcAE XLI. 1939. 365
nevertheless presents clearly the following differences from the new Burmese
Paraceras. In front of the 3 large eye-bristles a row of 5 bristles from the
antennal groove to near the oral margin, the second from above long ; on occiput
a median row of 3 as in ¢ of the new species and on one side also an antemedian
bristle ; posterior row of 6 each side with the second bristle much farther removed
from the long one and the bristle below the large one much smaller (judging
from the smallness of the pit, the bristle broken away on both sides). Sternum
VII apparently not different; I said of P. melinus in Nov. ZooL. xxxviii,
p. 267 (1932), under P. pendleburyi, that “the ventral lobe of VII.st. is obtuse,
rounded-emarginate.” This is true of the left side, but the margin of the segment
is slightly folded over as I now see; on the right side the margin is rounded as
in fig. 286; the row of bristles nearer the margin, distance of ventral bristle
from apical ventral angle only half as long as in the new species. Above stigma
of tergum VIII 10 bristles on the two sides together, in the new species 7 or 8 ;
below the stigma one small and two large bristles, not counting the small one
immediately below stigma; apical margin of widened area more rounded, the
apical angle less projecting. The specimen somewhat larger than our largest 9
of the new species, the tarsi, therefore, longer ; measurements: midtarsus 28,
30, 17, 11, 26; hindtarsus 67, 40, 22, 13, 27. It will be noticed on comparison
with the measurements of the new species that midtarsal I is much longer, being
but slightly shorter than II and a little longer (not shorter) than V. These
differences convince me that P. melinus is specifically distinct.
Macrostylophora.
The genus was “split off from the fimbriatus group (formerly of
Ceratophyllus) on account of the process of the seventh tergite and the
peculiar claspers ” (Ewing, A Manual of External Parasites, London, 1929,
p- 203). As the species allied to fimbriatus are not kept generically separate
366 NOVITATES ZooLoGIcAE XLI. 1939.
from Macrostylophora in the present paper, an explanation is necessary. The
species in question are essentially Oriental squirrel fleas and closely related to
one another. Two of them have in the 3 a long process in between the ante-
pygidial bristles (M. hastatus and M. pilatus J. & R. 1921) and a peculiar knife-
like spiniform at the upper posterior angle of the digitoid (text-fig. 287); in a
third (M. borneensis Jord. 1926) the antepygidial process is quite short, not
projecting beyond the cone bearing the antepygidial bristles, and the spiniform
at the upper posterior angle of the digitoid is not knife-like. These three species
agree with each other, and differ from all the others, in the shape of the manu-
brium of the clasper. I should much have liked to keep these three apart as
Macrostylophora, although “ macro ” does not apply to the short process of M.
borneensis. Unfortunately I have failed to find any character by which the 99
of them are distinguished as a group from the 99 of the gg which have no
antepygidial process and which therefore are not “ stylophores.’” One of the
species, perfectly distinct, M. probatus J. & R. 1922, is known from the @ only ;
its j may be a stylophore or a non-stylophore. What can we do with thisspecies ?
Put it into cold storage until its 5 has become known ? It is the old question of
basing a classification on one sex only. If ancestral characteristics, and therefore
of primary value, are the basis of classification, the results may be correct, but if
secondary characteristics are employed we may be led astray. The aim of
systematics is twofold : (1) to group the species according to blood-relationship,
with which postulate the author of Macrostylophora has complied ; and (2) to
bring the mass of forms into such order by means of definite diagnoses as finger-
posts that we can find out to which squad, company or regiment each individual
belongs, with which postulate the erection of genera on secondary 3-characters
only does not comply. The species with and without antepygidial $-process
here united are undoubtedly all closely related ; there is no difficulty at all in
recognizing the homology of the somatics in which each species differs from the
others. Macrostylophora is an awkward name for this assemblage of species ;
but we should look upon names in Nomenclature as mere symbols and discard
their philological meaning, as we do in the case of Baker, Smith and Taylor, who
may be bankers, lawyers or scientists and not what the name implies. A name
in Nomenclature is a short symbol in place of the diagnosis, and we still labour
under the illogical idea that a name must therefore be diagnostic. Fabricius,
150 years ago, saw the mistake and was quite emphatic : nomina “ optima sunt,
quae omnino nil significant. Characterem essentialem generis generico nomine
indicare impossibile ” (Ent. Syst. 1792, p. x). Let us then at least forget the
literal meaning of the Greek word macrostylophoros, as the Rules do not allow
us to discard the generic term derived from it.
A general description of the genus absolves us from repeating in the descrip-
tions of the new species characters which are found in all.
39. Frons with a small external, usually sharp, tubercle below middle ; in
front of row of 3 long eye-bristles a short row of a few small ones from antennal
groove near to level of upper eye-bristle. Above antennal groove a long median
bristle accompanied by a small one. Proboscis reaching to end of forecoxa or
to middle of trochanter, not beyond. Pronotal comb of 18 spines (17-19).
Abdominal terga with two rows of bristles, anterior row more or less incomplete,
one or two occasional dorsal additional small bristles ; I to IV or V with apical
spines. Sterna III to VI with 2 or 3 bristles each side, on VII one or two more
NOVvITATES ZooLoGIcAE XLI. 1939. 367
in 9. Mid- and hindeoxae without bristles on inner surface ; on outer side of
forefemur 2 to 6 small bristles, on inner surface one ; on mid- and hindfemora
one or two lateral subventral ones on outside in anterior half and none on inner
side (apart from the subapical ventral bristle). Hindtarsi long, I at least as
long as II and III together. Marginal stigma-cavity of tergum VIII short.
&. Bristles of antennal segment II short. One long antepygidial bristle
accompanied by two minute ones. In some species a strongly chitinized narrow
process in between the two sets of antepygidial bristles. Tergum VIII large,
with a row of long bristles along upper half of posterior margin, ventrally one to
three long lateral bristles; on inner surface a narrow dorsal spiculose area.
Sternum VIII narrow, with a very narrow basal dorsad extension ; at apex a long
bristle each side and a long membranous flap divided into numerous filaments.
Tergum IX with large forward extension, acting as a manubrium. Vertical
diameter of clasper much longer than horizontal one, the clasper ventricose ;
two long acetabular bristles above the ventral margin of the digitoid ; manubrium
of clasper broad at base, the sinus between it and the internal extension of
tergum IX very narrow, usually a mere slit. Process P of clasper narrow, several
times as long as broad, its apex angulate posteriorly. Digitoid large, posteriorly
at or below apex with a bristle of varying size, usually modified and above lower
posterior angle several more or less spiniform. Vertical arm of sternum IX dis-
tinctive : approximately its upper half elongate-triangular and pointing upwards.
Anal sternum long and narrow, with long slender bristles at dorsal margin
and apex.
9. Bristles of segment II of antenna not reaching to apex of club. Inter-
space between long ventral bristle of posterior row of occiput and second bristle
twice that between second and third. Upper antepygidial bristle about 4 to 4
the length of middle one, lower one 3 to 4. Tergum VIII with few bristles ;
ventral apical angle projecting, bearing 3 slender marginal bristles, one of them
at apex of angle, the two others above it; on inner surface one short bristle
below the row and 2 or 3 farther forward. On stylet one ventral lateral bristle
and dorsally a very minute one close to long apical bristle. Body of spermatheca
longer than broad, barrel-shaped, with underside concave behind middle ; its
duct much wider than the blind duct, less distinct in a cleared specimen, both
barely longer than the duct of the bursa copulatrix ; this latter duct with a
hump anteriorly above middle.
Species in our collection: hastatus J. & R. 1921, Shan States, Sikkim and
Tonkin, 3 subspecies ; pilatus J. & R. 1922, South Annam ; probatus J. & R.
1922, South Annam ; euteles J. & R. 1911, West China and Yunnan; lupatus
J. & R. 1921, Sikkim ; fimbriatus J. & R. 1921, N.W. India ; phillipsi Jord. 1925,
Ceylon ; levis J. & R. 1921, Selangor; borneensis Jord. 1926, North Borneo.
And the new species and subspecies described below.
3. Macrostylophora hastatus hastatus Jord. & Roths. 1921.
6. Ceratophyllus hastatus Jordan & Roths., Ectoparasites I, p. 168, no. 3, text-figs. 153, 154 (1921)
(Shan States).
The 5 $4 obtained by Dr. Heinrich have a somewhat intermediate position
between our 2 Sikkimese $f and the unique type from the Shan States. The
digitoid is as narrow as in sikkimensis; but its ventral margin is less incurved
(this margin is obscured in type of hastatus) ; the antepygidial process is a trifle
368 Novirares Zoonogicam XLI. 1939.
longer than in sikkimensis, the proportional length being in Tonkin (see below)
37, Shan States 32, Heinrich collection 34-36, Sikkim 29.
In the 99 from Burma the apex of sternum VII is narrower than in sikkim-
ensis, the sinus smaller and the lobe above it shorter.
It is advisable to wait for material from the Shan States before attempting
to solve this problem of geographical variation. The characters mentioned may
be quite unstable in a long series of specimens.
Maymyo, 800 m., on Paradoxurus hermaphroditus, 6.xii, 1 9, and on Tome-
utes, 15.xii.37, 2 $3; Mt. Popa, 600 m., on Tomeutes, 11.v.37, 1 3, 2 99,
and at 1100 m. on Sciurus, 2 3g.
I take this opportunity to describe a subspecies from Tonkin :
Macrostylophora hastatus tonkinensis subsp. nov. (text-figs. 287, 288).
The ¢ differs from M. h. hastatus and M. h. sikkimensis in the digitoid F
being broader and its upper posterior angle drawn out into a nose ; manubrium
narrower; antepygidial median
process a little longer than in
the other sub-species. Sternum
VII of Q asin M.h. sikkimensis.
Tonkin: Boa-Ha, on
Tamiops, 30.x1.23, 1 &, 1 9
(H. Stevens), Perey Sladen-
Godman Expedition.
Approaching M. pilatus a
little in the nose of the digitoid.
4. Macrostylophora heinrichi
sp. nov. (text-figs. 289-291).
Near M. fimbriatus J. & R.
1921 and M. lupatus J. & R.
1921, differing in the tail-ends
of both sexes and apparently in nothing else.
4. At upper margin of tergum VIII 6 long strong bristles, followed farther
down by a submarginal one, nearly on a level with the latter another large
bristle farther forward ; on upper half of side about 8 smaller bristles, on lower
area 2 large ones, one above the other. Sternum VIII (text-fig. 290) as in M.
lupatus, the fringed flap perhaps somewhat different, evidently variable individu-
ally, and of course assuming various unnatural positions in the process of mount-
ing the specimens. Forward extension of IX.t. and manubrium of clasper as
in M. lupatus. Process P of clasper longer, the distance of its tip from the upper
acetabular bristle longer ; the two acetabulars farther apart. Digitoid F much
longer in a dorso-ventral sense, this vertical diameter being about three times as
long as the horizontal diameter measured from the acetabular margin of clasper ;
apical portion of F narrower than in M. lupatus, the twisted large bristle farther
from upper angle, the 3 spiniforms of lower area larger than in M. lupatus, and
the slender bristle between them curved down, not straight (simply curved, or
recurved at apex as in figure). Ventral arm of sternum IX (text-fig. 290) very
distinctive : the postmedian dorsal projection more prominent and the longish
bristle placed at its distal side less modified than in M. lupatus, resembling an
NoviratEs ZooLoGIcAE XLI. 1939. 369
ordinary bristle, not being broad, not spiniform, and not abruptly narrowed to
a point, but as in M. lupatus the apex of this bristle rough with minute teeth ;
the apical lobe of the segment much narrower and longer than in M. lupatus and
with the minute bristles on its lateral surface less close together ; the dorsal
hump between the vertical and ventral arms much larger. Lamina of phallosome
27
370 Novirates ZooLoGIcAE XLI. 1939.
broader (Lam), the ventral apical processes (Par) long and slender, whereas in
M. lupatus this process is broad, barely twice as long as broad, apically slightly
dilated and rounded.
9. Apical margin of sternum VII (text-fig. 291) more or less strongly slanting,
slightly incurved and ventrally excurved, a short rounded lobe being formed,
the outline varying as shown in the figure. Apical margin of widened lower area
of tergum VIII much more deeply incurved than in any of the allied species,
both the upper and lower lobe varying individually, being usually more pointed
---Nlll.st.
than in the specimen figured. Sternum IX (internal) partially sclerified ; this
chitinized area broadly rounded anteriorly and narrowing to a point posteriorly
the ventral margin concave and well defined ; this sclerification not present in
other species of the genus. Bristles of segments VIII, IX and X, as well as the
spermatheca and the bursa copulatrix as in M. lupatus.
Length: & 2:5-2-9, 9 2-6-3-0 mm. ; hindfemur : ¢9 0-45-0-51 mm.
Mt. Victoria, on Tamiops, 13.1i1.38, type g, on Tomeutes, 16. and 18.v and
16.vi.38; a small series of both sexes.
5. Macrostylophora uncinalis sp. nov. (text-figs. 292, 293).
Both sexes are unexpectedly found to differ from the other species in the
striation of the basal abdominal sternite. In all species of the genus except this
new one the striae in the posterior half of the segment are very close together,
there being in the centre from base to posterior margin more than 50 vertical -
striae ; in the new species there are about 30, the interspaces between the posterior
striae being here about half as wide as the interspaces between the anterior
Novirates ZooLoGIcAaE XLI. 1939. 371
striae, in the other species very much narrower. Chaetotaxy as in other species,
apart from the modified segments.
d. Tergum VIII (text-fig. 292) dorsally with 9 long marginal bristles, the
row being preceded by a small bristle, below which there is a still smaller one ;
on side 6 bristles and ventrally a vertical row of 3; on inner side a narrow dorso-
marginal spiculose area. Basal curved-up portion of sternum VIII much
narrower than in the allied species, the ventral arm longer and straighter, the
apical flap with shorter filaments and not so much split up; at the joint where
the pair of bristles are placed (one each side) no additional densely filamentose
flap as in M. lupatus J. & R. 1921 and others. Forward extension of tergum IX
shorter than in M. lupatus and M. heinrichi sp. nov. The bay between this
sclerite and the manubrium (M) of the clasper narrow, its apex rounded. Dorsal
hump of manubrium M nearer base than in the allied species, apex of manubrium
rounded. Body of clasper (Cl) rounded from acetabular bristles ventrally to
‘ manubrium, strongly near the latter, distance of base of manubrium to aceta-
bular bristles much larger than in M. heinrichi, equalling 3 of the length of M in
M. uncinalis and „>, in M. heinrichi; process P reaching to middle of anterior
372 NOVITATES Zoonocicar XLI. 1939.
margin of digitoid F, apex more dilated than in M. heinrichi ; acetabulars on a
short broad projection of the margin of clasper. Digitoid F irregularly triangular,
its dorso-ventral length (measured parallel with posterior margin) not quite
twice the width measured from acetabular margin parallel with ventral margin ;
angle of anterior margin in middle, margin straight from this angle to apical
angle ; ventral margin slightly concave proximally and distally and feebly convex
in between ; posterior ventral angle strongly rounded ; upper (anterior) angle
about 90°; dorsal margin from this angle to large marginal bristle (about 3 of
total margin from anterior apical angle to posterior ventral one) rounded, then
the margin slightly concave to the smaller marginal bristle, and the rest convex ;
4 large spiniforms and 2 smaller bristles, the largest at 2, about as long as F is
broad in middle, the second smaller, a short distance below the large one, the
third above middle away from margin, large, the fourth again small, placed at
beginning of lower convexity of margin, the following 2 placed one above the other
near ventral margin and both as large as the third from above ; none of these
bristles as stout as the large spiniforms of M. lupatus, M. heinrichi, M. phillipsi,
ete., resembling more those of M. levis J. & R. 1922. Upper half of vertical arm
of sternum IX slightly leaning forward ; in angle between vertical and ventral
arms the segment widened, but there is no such prominent hump as in M.
heinrichi (compare text-figs. 285 and 292) ; proximal non-setose section of ventral
arm a little shorter than setose distal portion, abruptly widened ventrad before
middle, the angle rounded off, beyond it a few bristles ; distal setose section of
Novirates ZooLoGIcAE XLI. 1939. 373
nearly the same width throughout, dorsally incurved in middle and apically
rounded, the proximal dorsal prominence of M. heinrichi and others absent,
as is also the long bristle placed in the other species at the distal side of
this dorsal hump. Ventral process of parameres a large hook curved upwards
(Par).
2. Sternum VII (text-fig. 293) truncate, feebly incurved or slightly rounded.
Apical margin of widened apical area of tergum VIII hardly at all incurved, the
ventral angle less than 90°, its tip rounded off, upper angle almost effaced,
replaced by a flat arc. Distance of sensilium from stylet longer than in the
allied species, measured dorsally almost twice the length of the sensilium (6 or
7:11, in M. heinrichi 6 : 8) ; sternum X also longer and narrower. Spermatheca
a little more convex dorsally than in the
allied species.
Length: & 2:4, 2 3-0-3-6 mm. ; hind-
femur: & 0-43, 2 0-45-0-48 mm.
Mt. Victoria, 2600 m., on Tomeutes, 6 and
12.v.28, 13 (type), 3 92; also on Rattus,
NE}
6. Macrostylophora lupatus bamanus
subsp. nov. (text-fig. 294).
9. A little smaller than M. 1. lupatus from
Sikkim (length of hindfemur in lupatus 59 to
66, in bamanus 52 (one specimen !)). Body
of spermatheca narrower and in comparison
with the width longer, being slightly over
twice as long as broad, whereas in M. 1. lupatus
it is at most twice as long as broad. Sinus of sternum VII broader.
Length: 9 2:5 mm. ; hindfemur: 9 0-39 mm.
Maymyo, 800 m., on Funambulus, 6.xii.37, 1 9.
7. Stivalius aporus J. & R. 1922.
Maymyo, 800 m., on Tomeutes, 15.xi1.37, 1 9; forest north of Prome,
24.ix.37, on “Rat,” 1 g; Dudaw-Toung, Pakokku Chin Hills, on T’upaya,
11.11.38, 2 gg, 2 99; Mt. Victoria, 2200 m., on “ Rat,” 19.vi.38, 1 9.
The species is known to us only from South India and Ceylon (and Zululand,
an accidental occurrence). The Burmese specimens do not seem to differ at all.
This may be one of the species, known in other groups of insects, which link the
fauna of Ceylon with that of Burma independently of North India.
8. Neopsylla dispar Jord. 1932 (text-fig. 295).
Mt. Victoria, 2600 m., on “‘ Tree Rat,” 4 gg, 3 29.
Described from a pair obtained in North-East Burma on Epimys. The
ventral arm of sternum IX is slightly broader in these gg than in the type.
Besides, the ventral margin is distinctly rounded-angulate in the type, whereas
there is hardly an indication of the angle in Dr. Heinrich’s gg; the gap in the
row of bristles of the type is present in one of the specimens and less distinctly
374 NovITATEs ZooLoGIcAE XLI. 1939.
in the others. In our figure of the spermatheca (Nov. ZooL. xxxviü, p. 272) the
body is a little too broad and the tail apically slightly too wide.
9. Palaeopsylla remota Jord. 1929 (text-fig. 296).
Mt. Victoria, 1400 m., on Anurosorex, 22.i1ii.38, a small series of both
sexes.
Described from a single 9 collected at Chungking, Upper Yangtse-kiang.
H. Stevens obtained in 1931 a series in Sikkim on Anurosorex assamensis. The
QS
genitalia of the g resemble those of P. sorecis Dale 1878 in the digitoid being
long ; in remota, however, it is straight and projects beyond the apex of the
clasper. Sternum IX quite different, its ventral arm long and almost gradually
narrowed to a point, with a few slender bristles.
10. Thaumapsylla breviceps Roths. 1907.
Lamaing, 30 km. north of Mandalay, 14.1.38, on Rousettus, 1 3. The
usual number of spines on the inner side of the mesonotum is two each side ;
this specimen has two on one side and three on the other. , We have in the
collection also specimens with three each side, a $ from Java and a 9 from
the Cape, together with specimens which have the normal number.
Novirarres Zoonoaicar XLI. 1939.
Fig. 284.
285.
286.
287.
288.
289.
290.
291.
292.
293.
294.
295.
296.
EXPLANATION OF TEXT-FIGURES
The lettering is explained in the text.
Peraceras hamatus, Q . 3
. r genitalia of
” » ” „ Q © 0 0
Macrostylophora hastatus tonkinensis, digitoid of &
= BS es sternum VII of 9
ER heinrichi, genitalia of &, upper sclerites
= a ie „ &, lower sclerites
bP) be) be) 99 2
“3 uncinalis, ,, 5 3
bP) ” 9 99 fe) © 0
5% lupatus bamanus, 2, sternum VII and
spermatheca . ; © : ©
Neopsylla dispar, &, sterna VIII and IX .
Palaeopsylla remota, 3, sternum IX
rg to tS
375
. 362
. 363
. 365
. 368
. 368
. 369
369
. 370
. 371
. 372
. 373
374
. 374
376 NOVITATES ZooLocicaE XLI. 1939.
LARGE AND SMALL FLAME-CELLS IN A CERCARIA
(TREMATODA).
By MIRIAM ROTHSCHILD.
ON two occasions an Allocreadiid cercaria has been found parasitizing Peringia
ulvae (Pennant) 1777 from the mud flats of St. John’s Lake, Plymouth.
Morphologically it shows a fairly general resemblance to the Cercaria of Micro-
creadium parvum Simer, 1929 (Sewell Hopkins 1937). The shape of the body,
closely set eye-spots, alimentary canal, arrangement of penetration glands, and
the ratio of the suckers are similar. The cuticle, however, is unarmed and the
lateral caudal setae (three pairs) are replaced by seven pairs of rudimentary fins.
The size and distribution of the oil droplets in the parenchyma is also different.
The most interesting feature of the cercaria is the excretory system. The
bladder is roughly triangular in shape, lined with giant cells and is of the type
found in the cercaria of Hamacreadium and allied genera, rather than in Micro-
creadium, Crepidistomum, etc. It is filled with large concretionary granules.
The main excretory tubes leave the bladder at the anterior cornu, and divide
into an antero-lateral and postero-lateral branch about the level of the ventral
sucker. Immediately after the division, the postero-lateral tube dwindles in
size until it becomes a mere thread, less than half the diameter of the anterior
collecting tube. The canaliculi and flame cells are correspondingly small. The
latter resemble the early stages of flame cells found in germ balls or very immature
cercariae, but although exceedingly minute they are functional. The antero-
lateral collecting tubes are quite normal and the flame cells of usual size. There
is consequently a very marked contrast between the antero-lateral and postero-
lateral systems.
It is well known that the larval forms of the Allocreadiidae are characterized
by great diversity of form, and there is some evidence that both Stenostoma
and Mesostoma excretory systems occur in the same family. It is perhaps rather
too speculative, but nevertheless tempting, to suggest that the excretory system
of this cercaria from P. ulvae may represent an intermediate stage between these
two types.
REFERENCE
Hopkins, Sewell H., 1937. A new type of Allocreadiid cercaria : The cercariae
of Anallocreadium and Microcreadium.—In Journ. of Parasit., Urbana,
vol. xxiii, No. 1, pp. 94-97, 2 figs.
Novrrares ZooLoGIcaE XLI. 1939. 377
A PRELIMINARY REVISION OF THE PALAEARCTIC SPECIES
AND SUBSPECIES OF THISOICETRUS BR. W. (ORTHOP-
TERA, ACRIDIDAE).
By B. P. UVAROV, D.Se.,
Department of Entomology, British Museum (Natural History).
(With 1 text-figure.)
[eRe of species of this genus affords unusual difficulties owing
to considerable instability of specific characters. A great amount of
detailed taxonomic work would be necessary before a monograph of the genus
may be undertaken, and the following key to species and subspecies, as well as
their annotated list, should be considered merely as an attempt to reach a
temporary arrangement that may be of practical use.
KEY TO SPECIES AND SUBSPECIES.
1 (2). Male subgenital plate bi-tuberculate. Hind femur very slender, about
6 times as long as its maximum width. Hind tibia with 15 external
and 13 internal spines. Dark dorsal stripe terminates abruptly at
the hind margin of pronotum, not extending on to the anal area of
elytra. 6 6 8 1. T. adspersus (Redt.).
2(1). Male subgenital plate not bi-tuberculate. Hind femur moderately
slender or stout, less than 5 times as long as its maximum width.
3 (20). Hind tibia more or less red, or at least pale purplish, with more than
10-11 external spines. Pronotum always with a broad black median
stripe enclosed between two narrow light ones.
4 (5). Hind femur with heavy black markings on the externomedian area, two
of them reaching across it. Face strongly oblique; fastigium of
vertex forming an angle with the frontal ridge, at least in the male.
Very similar in general appearance and coloration to 7’. adspersus.
2. T. rantae Uv.
5 (4). Black markings on the externomedian area of hind femur absent or
small, none of them extending below the median line.
6 (9). Hind tibia with 12-13 (exceptionally 14) external and 10-11 internal
spines.
7 (8). Hind wings colourless or faintly greenish.
3. T. annulosus annulosus (WIk.).
8 (7). Hind wings distinctly bluish. 3a. T. annulosus cyanescens, subsp. n.
9 (6). Hind tibia with 14-17 external and 11-13 internal spines.
10 (15). Male subgenital plate short, not attenuate, very obtuse or even roundly
truncate apically (fig. 297, H).
11 (14). Size medium to quite large (for the genus). Frontal ridge ordinary,
distinctly constricted at fastigium.
12 (13). Size large (S 28-30 mm., 9 40-50 mm.). Lateral pronotal keels diver-
gent backwards. Lateral pronotal lobes and pleurae rugulose; hind
femur punctured . : : : : 4. T. harterti I. Bol.
378 Noviratrs Zoonoaicar XLI. 1939.
13 (12). Size medium (3 22-24 mm., Q 28-30 mm.). Lateral pronotal keels
practically parallel. Lateral pronotal lobes and pleurae less rugulose,
shiny, hind femur smooth, shiny © 0 9 5. T. persa Uv.
14 (11). Size very small ($ 15 mm., 2 unknown). Frontal ridge very wide,
not constricted at fastigium. Lateral pronotal keels practically
obsolete : 6 6 0 5 5 . 6. T. theodori Uv.
15 (10). Male subgenital plate elongate, attenuate, subacute apically (fig. 297, L, S).
16 (19). Size medium to large (3 over 18 mm., 9 over 30 mm.).
ANI
DAY,
Text-ric. 297.—Male subgenital plates, viewed from below and in profile: L, Thisoicetrus
littoralis littoralis ——S, T. littoralis simulis. H, 7. harterti. A, T. annulosus.
17 (18). Hind tibia sanguineous. Male subgenital plate less attenuate (fig. 297,
L). General coloration darker. 7. T. littoralis littoralis (Ramb.).
18 (17). Hind tibia pure light-red. Male subgenital plate more attenuate
(fig. 297, S). General coloration lighter.
7a. T. littoralis similis (Br. W.).
19 (16). Size small (g 16 mm., 2 27 mm.) : 7b. T. littoralis minutus Uv.
20 (3). Hind tibia without red colour, with 10-11 external and not more than
10 internal spines. General coloration buff; pronotum with only a
narrow and imperfectly defined median dark stripe ; elytra without
spots. Size large . 5 : : ; . 8. T. buxtoni Uv.
ANNOTATED LIST OF SPECIES AND SUBSPECIES.
1. Thisoicetrus adspersus (Redtenbacher 1889).
1889. Euprepocnemis adspersa Redtenbacher, Wien. Ent. Zeit., 8: 30.
1898. Thisoicetrus adspersus, I. Bolivar, Ann. Sei. Nat. Porto, 5: 34, 35.
1912. Thisoecetrus adspersus, Uvarov, Horae Soc. Ent. Ross., 40: 33, 34, fig. 3c.
This easily recognizable species has been recorded from Transcaspia (type
locality : Askhabad), E. Transcaucasia, Caspian plains of Northern Caucasus,
Iran, Mesopotamia, Palestine and Egypt, on one hand, and from Spanish Morecco
and S.E. Spain, on the other. Iam now able to record it also from Baluchistan,
Novirates ZooLoGIcAE XLI. 1939. 379
Arabia, Cyprus and Central Algeria, but there still remains a wide gap between
the eastern and the western areas of its occurrence, which cannot be due to
mere lack of records, but represents a real interruption in the present
distribution area.
A long series of specimens from Cyprus are all relatively very small and of
dark general coloration, but it would be premature to separate them subspecifi-
cally, since the species appears to be unstable in these respects elsewhere.
2. Thisoicetrus rantae Uvarov 1936.
1936. Thisoecetrus rantae Uvarov, Linn. Soc. Journ., Zool., 39: 549,
Although somewhat similar in the appearance to 7. adspersus, this species
differs well from it in the structure of the male subgenital plate and in other
characters indicated in the above key and in the original description. A curious
feature of 7’. rantae is the individual variation in the length of elytra which may
be either considerably shorter than abdomen, or extend beyond the hind knees,
while the intermediate condition is less common. This appears to suggest an
incipient dimorphism.
The species is known only from S. Arabia.
3. Thisoicetrus annulosus annulosus (Walker 1870) (text-fig. 297, A).
1870. Heteracris annulosa Walker, Cat. Derm. Salt. Brit. Mus., 4: 673, 674,
1871. Acridium continuum Walker, t.c., 5, Suppl.: 61 (syn. nov.).
1873. Pezotettix (Huprepocnemis) charpentierr Stal, Bih. Sven. Akad. Handl., 4 (5) : 15 (syn. nov.).
1908. Thisoicetrus charpentieri I. Bolivar, Bol. R. Soc. Esp. Hist. Nat., 1908 : 328,
1920. Thisoecetrus charpentieri Uvarov, Nov. Zool., 30: 75, 77,
1923. Thisoecetrus continuus Uvarov, Ent. Mon. Mag. (3) 9: 85.
I. Bolivar (1908) was the first author to define more or less clearly this
species, which was usually confused with 7’. littoralis, but he failed to notice the
important difference between the two in the shape of the male subgenital plate.
He was also unable to know of the existence of two earlier names given to the
same species by Walker, annulosa and continuum, but my examination of the
types shows them to be synonymous with Stäl’s charpentieri. The name T.
annulosus (W1k.), unfortunately, takes precedence of the others, although it is
based on a female from unknown locality.
When redescribing 7’. continuus in 1923, I had before me a limited series of
that insect, and had no means of comparing it with the true charpentieri. There-
fore, my redescription was made comparative with 7. littoralis, which is, of
course, not even closely related to the insect in question. Now I have before
me very abundant material from Tunis, Egypt (Siwa, Helouan, etc.), Arabia,
Palestine, etc., and am able to record its extreme variability as regards size,
coloration and the degree of rugosity of integument. It appears that, being
essentially an oasis insect, this species splits up into numerous local population
groups, fairly uniform within themselves, but always slightly different from
other populations. The Palestine (Jordan Valley, especially) group might even
be separated subspecifically (see my redescription of continuus), as its members
are usually much more roughly sculptured and have relatively shorter and
stouter hind femora than North African forms. I believe, however, that it would
be wiser not to attempt subspecific division without more detailed studies, and
380 Noyrrates ZooLoGIcAE XLI. 1939.
the only subspecies of 7’. annulosus which I propose to recognize is that from
the Somaliland, differing in wing colour (see below).
The known distribution of 7’. annulosus comprises Spanish Morocco, Algeria,
Tunis, Libya, Egypt, Anglo-Egyptian Sudan (Kordofan ; British Museum),
Sinai, Palestine, Arabia.
3a. Thisoicetrus annulosus cyanescens subsp. n.
One male (type) and three females in the British Museum, from Buran,
British Somaliland, 3000 ft. (©. L. Collenette), are strikingly different from the
typical form in the light-blue coloration of hind wings, while quite similar to it
morphologically. Since no case of blue coloration of hind wings is known to me
amongst very long series from other localities, I regard the Somalian specimens
as representing a distinct subspecies.
4. Thisoicetrus harterti I. Bolivar 1913 (text-fig. 297, H).
1913. Thisoicetrus harterti I. Bolivar, Nov. Zool., 20: 614.
1923. Thisoecetrus bolivari Uvarov, op. cit., 80: 76 (syn. nov.).
1923. Thisoecetrus harterti, Uvarov, l.c.
In my 1923 paper I have made a mistake in identifying the Biskra insect
with that redescribed by I. Bolivar as 7’. charpentieri St. (now T. annulosus, see
above) and believed that author incorrect in his determination, while I can see
now that he was right.
The number of tibial spines in this insect, now known to me by a long
series from Southern Algeria (Bou-Saada, Djelfa, Colomb-Bechar), French
Sudan, Northern Nigeria and Kordofan, is greater than in 7’. annulosus and by
this character the species must be referred to the group allied to 7’. littoralis.
On the other hand, the male subgenital plate of 7’. harterti is of the type approach-
ing that in 7’. annulosus, but clearly different in the truncate apex. In Kordofan,
T. harterti was found actually together with 7. annulosus, from which it differs
strikingly even in the size, and there can be no doubt of its independent specific
status.
T. harterti is easily recognizable at first sight by its large size and large
dark spots on elytra. I. Bolivar’s type from Biskra is a bleached female, prob-
ably because it hibernated in the adult stage (taken in April), but in fresh speci-
mens the coloration is very contrasting. The Kordofan specimens are paler
than those from Algerian oases, and they were also taken at the end of winter.
The distribution area of this species embraces the Sudanese zone of dry
grasslands south of the Sahara, while in the Sahara itself it occurs only in oases,
as a relic of the former northward extension of tropical savannas.
5. Thisoicetrus persa Uvarov 1933.
1933. Thisoecetrus charpentieri persa Uvarov, Trav. Inst. Zool. Acad. Sci. U.R.S.S., 1: 231.
On re-examining some paratypes still available to me, as well as additional
series of specimens from British Baluchistan, I am now convinced that this
insect deserves specific recognition, the more so that in Baluchistan it occurs
together with T. annulosus (former charpentieri). By the number of tibial
spines it belongs to the same group as 7’. littoralis and 7’. harterti, sharing with
the latter the shape of the male subgenital plate. It is easily recognized by the
strongly polished shiny surface and practically parallel lateral pronotal carinae.
Novirates ZooLoGIcAE XLI. 1939. 381
6. Thisoicetrus theodori Uvarov 1929.
1929. Thisoecetrus theodori Uvarov in: Bodenheimer u. Theodor, Ergebnisse der Sinai-Expedition,
1929: 102.
A curious diminutive species with abnormally expanded frontal ridge.
Unfortunately, the type, a male from Sinai, remains the only specimen known.
7. Thisoicetrus littoralis littoralis (Rambur 1838) (text-fig. 297, L).
1838. Gryllus littoralis, Rambur, Faune Andal., 2:78.
1908. Thisoicetrus littoralis, I. Bolivar, Bol. R. Soc. Esp. Hist. Nat., 1908 : 329.
1923. Thisoicetrus littoralis littoralis, Uvarov, Nov. Zool., 30: 77.
This subspecific name is restricted to the Spanish representatives of the
species, while two more subspecies are recognized below.
7a. Thisoicetrus littoralis similis (Brunner 1861) (text-fig. 297, S).
1861. Caloptenus similis Brunner von Wattennwyl, Verh. zool. bot. Ges. Wien., 11: 224.
1870. Cyrtacanthacris notata Walker, Cat. Derm. Salt. Brit. Mus., 3: 574 (syn. nov.).
1912. Thisoecetrus similis Uvarov, Horae Soc. Entom. Ross., 40 (3) : 32, fig. 3b.
1931. Thisoicetrus littoralis asiaticus Uvarov, Trav. Inst. Zool. Acad. Sci. U.R.S.S., 1: 230 (syn.
nov.).
Lack of sufficient material on this insect led me into unnecessary confusion
in the past, but I have before me now very extensive series from the whole of
south-western Asia and north-eastern Africa. This enables me to establish
what I hope is the final synonymy of this subspecies of 7’. littoralis.
Brunner in 1861 has described a Caloptenus similis from Egypt (which I
designate as the type locality) and Syria, giving only a very brief diagnosis that
may be applied to several different species of T’hisoicetrus. In 1912 I have
suggested that his description fits a Transcaspian insect, and I then gave a
figure of its male genital appendages. Later (in 1931), however, I became doubt-
ful in my interpretation of similis, mainly because I had no sufficient Egyptian
material before me and thought that Egyptian specimens may be different from
Asiatic ones. At present, I am able to say definitely that the genus T'hisoicetrus
is represented in Egypt by T. adspersus, T. annulosus and a subspecies of 7’.
littoralis which extends its area into Palestine, Arabia, Iraq, Iran, Central Asia
and Baluchistan, without exhibiting any appreciable geographical variation.
Since the type of Caloptenus similis appears to have been lost (or possibly, has
never been labelled as such), the question arises which of the three Egyptian
members of the genus answers best the brief original description. T. adspersus
is at once ruled out because Brunner would not have failed to notice the bi-
tuberculate male subgenital plate characteristic of this species. The description
of the lateral pronotal carinae, which are said to be “rectissimae ’” in ©. similis,
can never be applied to 7’. annulosus, while it is very apt in the case of 7’. littoralis.
A useful supplementary character may be seen in the presence of three black
spots on the external face of hind femur, since these spots are usually well deve-
loped in 7’. littoralis, but mostly quite obsolete, or very indistinct, in 7’. annulosus.
Some uncertainty is introduced only by the description of the male subgenital
plate which is said to be short and obtuse, while the cerci do not exceed the
plate. The subgenital plate in 7’. annulosus is certainly shorter and more obtuse
than in 7’. littoralis, but the cerci in the former are definitely (if a little) projecting
beyond the plate, while in 7’. littoralis they are relatively shorter and, therefore,
382 NOVITATES ZootocicaE XLI. 1939.
correspond better to the description of ©. similis. I think, therefore, that my
1912 view was correct and that Brunner’s name should be applied to the sub-
species of littoralis, which has been later described by Walker as Cyrtacanthacris
notata (from “ Upper Egypt ”) and by myself as T. littoralis asiaticus (from
Iraq, Iran and Transcaspia).
7b. Thisoicetrus littoralis minutus Uvarov 1921.
1921. Thisoecetrus littoralis var. minuta Uvarov, Trans. Ent. Soc. London, 1921: 123.
1923. Thisoecetrus littoralis minuta Uvarov, Nov. Zool., 30: 77.
This is a very distinctive small race of T. littoralis described from Bone on
the Algerian coast and now known to me also from Bau-Saada, where it occurs
together with 7’. harterti, as well as from St. Germain in Tunis.
8. Thisoicetrus buxtoni Uvarov 1921.
1921. Thisoecetrus buxtoni Uvarov, J. Bombay Nat. Hist. Soc., 27: 65.
A remarkable large species, described originally from Amara in lower
Mesopotamia, but now known to me also from Baghdad.
Note 1.—When studying Palaearctic species of T’'hisoicetrus, I had to consider
Euprepocnemis caerulescens Stal 1876, referred by some authors to T'hisoicetrus.
An examination of a J and a @ types, kindly sent by Dr. M. Beier from the
Brunner collection in Vienna, proved them to belong to the genus Bibulus I.
Bolivar 1914, based on T'ylotropidius brunni Giglio-Tos. This latter species was
considered by its author to be identical with Huprepocnemis caerulescens as
re-described by Brunn in 1901 from East African specimens, but not with
Stal’s original H. caerulescens from Massowa on the Red Sea. Actually, however,
no specific differences can be found between specimens from the Red Sea coasts
and those from the interior of East Africa, as I am able to say after having
examined good series from various localities. Therefore the following synonymy
should be established :
Bibulus caerulescens (Stal 1876).
1876. Euprepocnemis caerulescens Stal, Bih. Sven. Avad. Handl., 4 (5): 16.
1901. Euprepocnemis caerulescens, Brunn, Mitt. Naturhist. Mus. Hamburg, 18 : 84, 132
1907. Tylotropidius brunni, Giglio-Tos, Boll. Mus. Torino, 22: 31 (syn. nov.).
1914. Bibulus brunni, I. Bolivar, Trab. Mus. Hist. Nat. Madrid, 20: 31.
Note 2.— The name Thisoicetrus similis Ramme 1931 (Mitt. Zool. Mus. Berlin,
16 : 942) is preoccupied by Th. similis Brunner 1861, and the species is here
renamed Th. oxyurus nom. nov.
Noviratres ZooLogIcAE XLI. 1939. 383
ON SOME TROPICAL POLYCHAETA IN THE BRITISH MUSEUM,
MOSTLY COLLECTED BY DR. C. CROSSLAND AT ZANZIBAR,
TAHITI AND THE MARQUESAS.
By C. C. A. MONRO,
Department of Zoology, British Museum (Natural History).
(With 4 text-figures.)
II. FAMILIES SYLLIDAE AND HESIONIDAE.
Note.—The first part of this paper containing an account of the Families Amphino-
midae and Phyllodocidae was published in the August number of the Annals and
Magazine of Natural History for the present year.
FAMILY SYLLIDAE.
Genus Syllis Savigny 1816.
1. Syllis variegata Grube 1860.
Fauvel, 1923, p. 262, fig. 97, h-n ; and 1932, p. 76.
Occurrence—Red Sea (3); Marquesas (5); Taunoa Reef, Tahiti (3) ;
Fautaua Reef, Tahiti (5); Papeete, Tahiti (6) ; Hava Hevane Reef, Tahiti (3) ;
Cape Verde Islands (3).
Distribution.—Atlantic, Mediterranean, Indian Ocean, Pacific.
Remarks.—I find the discrimination of these Syllids extremely difficult.
A number show rather long and slender blades to the upper chaetae in the front
region, and I was tempted to ascribe them to S. alternata Moore. According
to the Berkeleys, the latter has the palps fused at the base, a character which
is often very difficult to verify on the material, and examples of S. variegata
often have rather long and slender blades to the bristles in the anterior region.
Unfortunately I have no examples of S. alternata for comparison. Syllis hartı
E. and C. Berkeley, which the Berkeleys claim to show affinities with S. alternata
especially in the linear dorsal bristles, is quite distinct from my material and has
altogether longer blades to the bristles.
I can find no eyes in the Cape Verde specimens which bear some resemblance
to S. caeca mihi from the Panama region, but otherwise they agree with 8.
variegata.
2. Syllis gracilis Grube 1840.
Fauvel, 1923, p. 259, fig. 96, f-i ; and 1932, p. 76.
Occurrence.—Marquesas, from various washings (numerous); Tahiti ;
Papeete Harbour, 15 ms. (5); and from washings near Papeete (numerous) ;
Suez (1); Sherm Sheikh, Red Sea (1).
Distribution.—Atlantic, Mediterranean, Red Sea, Indian Ocean, Pacific.
Remarks.—This is probably the most abundant Syllid in the washings.
In some of the young specimens from Papeete Harbour the fusion of the shaft
with the blade in the large crutch-like chaetae is not yet complete. According
to E, and C. Berkeley (1938, p. 41), the simple chaetae in Syllis elongata (Johnson)
384 NOVITATES ZoonocicaE XLI. 1939.
are not formed, as Moore believed, by the fusion of the blade with the shaft as
in S. gracilis, but by the shedding of the blade as in 8. amica Quatrefages.
3. Syllis hyalina Grube 1863.
Fauvel, 1923, p. 262, fig. 98, a—b.
Occurrence.—Chuaka, Zanzibar, 2-3 ms. (2); Red Sea (4); Marquesas (2) ;
Tahiti (3).
Distribution.—Atlantic, Mediterranean, Indian Ocean, Pacific.
Remarks.—I have referred a number of small Syllids to this species on the
ground of the short and slender cirri and the bidentate character of the bristles
throughout. I have provisionally separated this species from S. closterobranchia
Schmarda (v. Fauvel, 1932, p. 77) because in Schmarda’s species the chaetae
are usually unidentate in the middle region of the body and the dorsal cirri are
relatively stout and fusiform. Practically it is often very difficult to decide
to which of the two species a specimen belongs, and the distinction between
them is doubtfully valid.
4. Syllis spongicola Grube 1856.
Fauvel, 1923, p. 257, fig. 95, a-d; and 1932, p. 76.
Occurrence.—Miludumadulu, Maldives (coll. J. Stanley Gardiner) (1); S.
Nilandu, Maldives (coll. J. Stanley Gardiner) (1); Torres Straits (coll. F. A.
Potts) (1); Red Sea (5); Zanzibar, 10 ms. (1); Suez Bay, 44 ms. (5); Tahiti
(10) ; Marquesas, from blue sponge (numerous).
Distribution.— Atlantic, Pacific and Indian Oceans. .
Remarks.—There is much variation in size and shape of body in these
specimens, but they all agree in having the main axis of the second tooth of the
bristles at right angles to the shaft. In S. depressa Augener and in Fauvel’s
var. dollfusi the second tooth is either more or less parallel with the main tooth
or else it is reduced to a small boss.
On the specimens from Tahiti Augener has the following note: “ Living
in sponge which is green outside and yellow within. The Syllids are the same
colour, obviously no protective significance. No details recorded, as only seen
by naked eye alive. Dead when washed out but colour seems to be uniform.
Body slender, feet well developed, but cirri and tentacles small.”
5. Syllis zonata (Haswell) 1886.
Gnathosyllis zonata Haswell, 1886, p. 746, pl. 52, figs. 4-6.
Augener, 1913, p. 195, pl. 3, fig. 22, and text-fig. 21 a-c.
Occurrence.—Tahiti (numerous) ; Marquesas (numerous).
Distribution. —South and South-west Australia, Galapagos Islands, Mar-
quesas.
Remarks.—One of the larger specimens measures about 20 by 1 mm. for
about 80 chaetigers. As far as can be seen on preserved material, the colour
pattern is very variable. There are typically three purplish brown bands in
each segment, but these may be fused so that the whole dorsum is a more or
less uniform purplish brown. Dark spots at the base of the dorsal cirri may
be present or absent, and the dorsal cirri may or may not be banded with brown.
In some of the smaller and presumably younger specimens the more or less
Noviratrs Zoonoaroan XLI. 1939. 385
uniformly brown dorsum carries dorso-lateral pairs of white spots in some but
not in all the segments of the front region.
The dorsal cirri are rather massive, of moderate length and with large
clearly separated articles. In the larger specimens the number of articles is
alternately about 25 and 12.
The bristles are all clearly bidentate, but show nothing characteristic.
Crossland has the following note on some specimens from Tahiti: “ Not
seen alive. Only found after long narcotization of material. Large with long
dorsal cirri over the back, conspicuous palps. The body is almost uniformly
dark brown dorsally, white ventrally, and dorsal cirri are banded brown and
white. Small white spots dorso-laterally in first few segments, but not on all,
may be alternate or on every third. This brown colour breaks into patches and
dies out before half the body length, and before the brown on the body begins
to go the dorsal cirri have but one brown band or none.”
Another note on some specimens from Tahiti is as follows: ‘“ Very distinctly
striped brown. May be a variety of 229 (the note given above), but not
noted there whether the dorsal cirri are annulated or not. These are smooth
and have brown dots throughout the length of the body, numerous on
anterior cirri, one or two on posterior. Ground colour whitish. Palps small.
Ground colour of body anterior third yellowish, after which sudden change to
dark, dull, green. Brown lines, three to the segment, conspicuous on both
ground colours, but naturally more so anteriorly. Underside without marks.
“8.vi.25. Coralline weed. A deeply pigmented specimen, brown bands
confluent on first 12 setigerous segments, so leaving intersegmental lines con-
spicuous.
“ There are two species of Syllid with brown bands: this one with 2 long
and 1 shorter band per anterior segment, 2 dots on side of body below the banded
area and dots on dorsal cirri which are smooth.”
Crossland is wrong in saying that the dorsal cirri are smooth. The other
banded species to which he refers is T’rypanosyllis zebra, under which heading
the note is given.
6. Syllis corallicola Verrill 1900.
Monro, 1933, p. 249, fig. 3, a-b.
Occurrence.—Praslin, Seychelles (J. Stanley Gardiner) (1).
Distribution.—West Indies, Seychelles.
Remarks.—An anterior fragment measuring 7 by 1 mm. for 26 chaetigers. I
have compared it with some specimens from Dry Tortugas attributed by me to
Verrill’s species, and I believe them to be conspecific. The bristles in the front
region have rather long and slender blades, and the dorsal cirri have alternately
between 60 and 70 and between 30 and 40 articles. The bristles are bidentate
throughout.
7. Syllis armillaris (Malmgren) 1867.
Fauvel, 1923, p. 264, fig. 99, a-f.
Occurrence.—Agig Bay, Red Sea, 44 fins (1).
Distribution.—Arctic, Atlantic, Mediterranean, Pacific.
Remarks.—I had some hesitation in reporting this species from the Red
Sea, but except that the eyes cannot be seen, the specimen shows all the character-
istic features of the species. Apart from northern Europe and the Mediterranean,
28
386 NOVITATES ZooLoGIcAE XLI. 1939.
this species has been reported from Alaska by Moore and from Vancouver Island
by Berkeley. I believe this to be the first record from tropical waters.
8. Syllis verruculosa Augener 1913 (text-fig. 298).
Augener, 1913, p. 203, pl. iii, fig. 39, and text-fig. 24, a-c.
Occurrence.—Wasin, Zanzibar, 7-10 fins (1).
Distribution. —South-west Australia, Zanzibar.
Remarks.—The single specimen is colourless and measures 17 by 1 mm. for
62 chaetigers. There is a very broad and more or less rectangular prostomium,
such as Augener figures. The pharynx reaches to the 6th chaetiger and the
proventriculus to the 11th. The tooth is anterior. The body is rather sparsely
dotted with large cylindrical or conical papillae (text-fig. 298, A), which become
smaller towards the hinder end. The dorsal cirri have 15 to 20 articles. The
bristles are all unidentate, and in the hinder region the ends of the shafts are
much expanded and the tips bent outwards (text-fig. 298, B). The blades
consist of small curved hooks, which in shape to some extent resemble the
hooks in the hinder feet of Syllis exilis. They
are smaller than the latter and not so much
298 modified. This appears to be an uncommon
species, for I know of no record of its occurrence
other than the original.
B 9. Syllis exilis Gravier 1900.
0-OImm Gravier, 1900, p. 160, pl. x, fig. 9.
Fauvel, 1932, p. 77.
Syllis fuscosuturata Augener, 1927, p. 52.
TEXT-FIG. 298.—Syllis verruculosa: Monro, 19334, p. 32, fig. 14, a-e.
dy HEDIS (OS 4) 8 > lowisiild strom Occurrence.—Zanzibar, 7-10 ms. (7);
hinder region (x 2). on
Tahiti (2).
Distribution.—West Indies, Red Sea, Indian Ocean, Pacific.
Remarks.—This is a fairly common tropical species inhabiting dead coral,
and it is less well represented in the Tahiti and Marquesas washings than I
should have expected.
On a specimen from Tahiti Crossland has the following note: “ Among
fucoid weed with the preceding (Syllis variegata) and Nereis dwmerilii. White
with bright green-grey markings, as sketch, as far as the end of the buccal appa-
ratus, after which the connexion between the pair of marks is only visible some-
times.” (Crossland’s sketch shows in the anterior part of the segment a pair
of rectangular marks joined by a line at the inner and hinder corners. This is
followed by a single rectangular mark in the hinder part of the segment.) ‘“ After
this a small black dot in the middle of each segment over the opaque white gut.
Palps large, four small bright red eyes. Dorsal cirri long and very slender,
colourless but marked with minute dark dots, visible under a lens. Dorsal
cirri appear smooth.”
10. Syllis vittata Grube 1840.
Fauvel, 1923, p. 263, fig. 98, i-1.
Occurrence.—Marquesas, Hiva Oa (3) ; Moorea shore, Papeete (4).
Distribution.—Atlantic, Mediterranean, Pacific (?).
Remarks.—With some hesitation I have assigned these specimens to Grube’s
NOVITATES ZooLoGIcAE XLI. 1939. 387
species. The colour has faded, but otherwise I see no ground for separation.
The cirri in the front region are about twice as long as those near the hinder
end. The longer anterior dorsal cirri show about 40 articles and those of the
posterior region about 20. The dorsal cirri are less stout and fusiform than in
S. krohnii and the heads of the chaetal shafts less enlarged. The specimens
from Moorea show a slight second tooth on the chaetal blades. The present
examples may belong to S. fasciata Malmgren, an arctic species, which Fauvel
has recorded from Japan and the Gulf of Petchili.
11. Syllis krohnii Ehlers 1864.
Fauvel, 1923, p. 259, fig. 96, a-e; and 1934, p. 303.
Occurrence—Marquesas, Hiva Oa (numerous) ; Red Sea (1) ; Zanzibar (2) ;
Seychelles (1) ; Cape Verde Islands (2).
Distribution.—Atlantic, Mediterranean, Indian Ocean, Pacific.
Remarks.—The markings on the specimens from the Cape Verde Islands are
different from those on the material from the Marquesas. The latter have the
dorsum coloured reddish brown in varying degrees, the colour showing a tendency
to break up into segmental markings ; the Cape Verde specimens have a single
clearly defined purple band in each segment.
In some specimens the dorsal cirri show traces of banding.
12. Syllis (Ehlersia) cornuta Rathke 1843.
Fauvel, 1923, p. 267, fig. 100, g-7.
Occurrence.—Marquesas, from various washings (15) ; Tahiti (1).
Distribution. —-Atlantie, Mediterranean, Indian Ocean, Pacific.
Remarks.—A number of these specimens have very short dorsal cirri with
alternately about 14 and 10 articles and each article encloses one or two brown,
refringent bodies. These examples usually have only one #hlersia bristle in
each foot, and these bristles are absent from the first few chaetigers. I was
tempted to separate them from cornuta and attribute them to Grube’s cerina
with which they agree well, but there are other specimens which grade into the
typical cornuta with a higher number of articles to the dorsal cirri and two or
three Hhlersia bristles in the feet.
The specimen from Tahiti has with it a reference to the following note by
Crossland: “5.vi.25. Salmon pink from end to end without markings, rather
darker at head end. Long palps, eyes red and very small. Dorsal cirri colour-
less, smooth, fairly long. 8.vi.25. Mossy weed on stones exposed to surf
Fautaua Reef. Colour much fainter, just visible in body wall, conspicuous on
head, palps and walls of pharynx. Rest of body coloured brown by gut only.”
The European representatives of this species are yellowish white or colour-
less. Crossland’s note may refer to some specimens of S. variegata which were
with the example of the present species, but in my opinion his account of the
colour is more likely to refer to the S. (H.) cornuta.
Genus Pionosyllis Malmgren 1867.
13. Pionosyllis marquesensis n.sp. (text-fig. 299).
Occurrence.—‘‘ Among weed, etc., lining pool on the shore shelf at Tai o Hae,
Marquesas, common in this situation and under stones in the pool. Its colour
makes it conspicuous, though it is small,” (Crossland) (8) ; Tahiti (1).
388 Novirarrs ZooLoGICAE XLI. 1939.
Description.—One of the larger specimens measures 37 by 1 mm. for 118
chaetigers. The head (text-fig. 299, A) is broader than long, divided by slight
grooves into two lobes separated by the median tentacle. There are two pairs
of eyes set in a widely open are. The palps are broad and fused at their base.
The median and lateral tentacles are subequal, about twice the length of the
head, and slightly shorter than the tentacular and anterior dorsal cirri. Both
tentacles and dorsal cirri are vaguely and irregularly constricted. The pharynx
reaches to the 7th chaetiger and the proventriculus to the 12th. The rim of
the pharynx is smooth and surrounded by a circlet of about 16 papillae. The
tooth is not quite anterior. The dorsal cirri are alternately longer and shorter,
the longer extending a little more than halfway across the back, and the shorter
having about half this length. The feet (text-fig. 299, B) are supported by two
299
Olmm
Trext-FIG. 299.—Pionosyllis marquesensis: A, head from above (x 4);
B, foot from anterior region (X 4); C, upper bristle (x 4); D, lower bristle
(x 4); E, lower bristle from hinder foot (x 4).
or three dorsal acicula. They are rather long and narrow and at the apex the
lips are continued into a pair of cirriform processes. The ventral cirrus is
slender and reaches to the end of the foot. About the four uppermost bristles in
each foot are narrow and linear (text-fig. 299, C) and end in delicate, clearly
bidentate tips. The lower bristles (text-fig. 299, D) have shorter and more curved
blades also with clearly bidentate tips. There is little change in the feet through-
out the body. In the hinder region the blades (text-fig. 299, E) of the more
ventral bristles are shorter and more curved than in the front region. The body
ends with a pair of pygidial cirri.
Remarks.—This species shows affinities with the Antarctic P. stylifera
Ehlers, but is distinguished by the presence of the slender linear bristles and by
the absence of the enlargement of the dorsal cirrus of the first chaetiger. Augener
has described from South-west Australia two species, P. ehlersiaeformis and
P. weissmannioides, both with linear bristles of the Ehlersia type, but these
bristles are unidentate in both and of a different form from the linear bristles in
Novitatrs ZOOLOGICAR XLI. 1939. 389
the present species. P. divaricata (Keferstein) is also an allied species, but
differs in having unidentate bristles.
In spirit the colour is pale yellow, but of its colour in life Crossland gives
the following account: “ Colour yellow, deeper posteriorly, one specimen full
of greenish eggs posteriorly. The body is thick and soft, thicker than the
prostomium, feet small, with long slender dorsal cirri. Prostomium deep in
colour, orange, with red eyes. Behind this is a conspicuous opaque white spot,
rather irregular, occupying most of the dorsum of the peristomium. From about
the 5th segment a pair of clear white spots on each segment, one either side of
the middle line. The translucence of the body makes these opaque spots the
more conspicuous.”
Of the two species of Pionosyllis described by Moore from the north Pacific,
P. gigantea has altogether longer dorsal cirri and flattened, foliaceous ventral
cirri and P. magnifica also has flattened ventral cirri partly fused with the feet
and lacks the slender, linear bristles of the present species.
Of the specimen from Tahiti Crossland writes: “ One specimen from
the coralline weed of Fautaua Reef. In spite of small size it is easily
identified by the naked eye by the white spot on the peristomium.”
Genus Opisthosyllis Langerhans
1879.
14. Opisthosyllis brunnea
Langerhans. A A
Fauvel, 1930, p. 15, fig. 2, a-k. 25mm
Occurrence.—Tai o Hae, Mar-
quesas (1) ; Seychelles (J. Stanley
Gardiner) (2).
Distribution —Kastern Medi-
terranean, Atlantic, Indian Ocean,
Pacific.
Remarks.—The specimen from
the Marquesas has a long re-
generating area at the hinder
end. There are no papillae. The
number of articles of the longer
dorsal cirri is about 30. The
bristles are unidentate through-
out. In spirit the colour is a
Cc
brownish yellow dorsally. ora
15. Opisthosyllis longicirrata TEXT-FIG. 300.—Opisthocyllis longicirrata: A, head
n.sp. (text-fig. 300). from above (x 3); B, middle foot (x 3); C, anterior
bristle (3); D, large bristle from hinder region (x 3).
Occurrence—Hululu Male
Atoll, Maldives (J. Stanley Gardiner) (2); Red Sea (2); Suez (1); Tahiti,
Taunoa Reef (2).
Description —The specimens from the Maldives are the best preserved,
and the larger of these measures 19 by 1 mm. for 74 chaetigers. All colour
markings have disappeared. The head (text-fig. 300, A) is more or less bilobed,
390 Novitatrs ZooLoGIcAE XLI. 1939.
grooved posteriorly and carries two pairs of eyes set in a rectangle. There is a
large nuchal flap or gibbosity. The median tentacle is about three times as
long as the palps and has between 50 and 60 articles. The lateral tentacles are —
all incomplete. The pharynx reaches to about the 10th chaetiger and the
proventriculus to the 18th. The tooth lies in about the 8th chaetiger.
The feet (text-fig. 300, B) are triangular in outline and are supported by two
or three dorsal acicula. The dorsal cirri in the front region are very long and
have about 120 articles. They are shorter in the hinder region, one of the longer
type having about 60 articles. In the hinder region the apexes of the pedal
lips are prolonged into a pair of small papilliform processes. The ventral cirri
are short, not reaching to the end of the feet. The bristles are all clearly biden-
tate ; the blades in the front region (text-fig. 300, C) are rather slender and
elongate, and they become shorter and broader from before backwards. In the
hinder region there are at the base of the foot two or three chaetae larger than
the rest, but of similar shape (text-fig. 300, D). There is no papillation.
Remarks.—Of the species of Opisthosyllis known to me, O. brunnea, O.
nuchalis, O. viridis, O. australis and O. ankylochaeta the present species is nearest
to Augener’s O. australis. It differs in having clearly bidentate
bristles throughout, in its much longer dorsal cirri and in the
301 absence of all papillation. The presence of two or three
bristles considerably larger than the rest in the feet of the
hinder region recalls Fauvel’s O. ankylochaeta, but in
Fauvel’s species these bristles are simple, the blade being
fused with the shaft. Moreover, in O. ankylochaeta the
bristles are unidentate.
16. Opisthosyllis viridis Langerhans 1879 (text-fig. 301).
Langerhans, 1879, p. 543, pl. xxxi, fig. 9, a-c.
Occurrence—Taa Hu Ku Bay, Hiva Oa, Marquesas,
from three washings, W. 39, W. 41 and W. 44 (3).
Distribution.—Madeira, Marquesas.
Remarks.—The best preserved of these specimens is from
W. 44 and measures 7 by 0-5 mm. for about 60 chaetigers.
In spirit there is no colour. As in Langerhans’s figure,
there are two pairs of eyes in a widely open are and a
papillated nuchal flap reaches forward almost to the eyes.
Text-rıc. 301 The median tentacle reaches just beyond the palps and has
Opisthosyllis viridis: about 15 articles. The laterals are a little shorter. The
fe ee tus body is papillated. The pharynx reaches to the 10th
middle region. chaetiger and the proventriculus to the 17th. The tooth
lies in the 9th. In the larger but poorly preserved specimen
from W. 39, measuring 13 by 1 mm. for 67 chaetigers, the pharynx reaches
to the 15th chaetiger, the proventriculus to the 22nd, and the tooth lies in the
13th. In the smaller specimen the number of articles of the longer dorsal
cirri is about 15; in the larger a few of the longer cirri in the front region
have as many as 25 articles.
The bristles (text-fig. 301, A and B) are unidentate throughout and except
for a slight expansion of the head of the shaft in the hinder region show little
A B
O0-0lmm 0-Olmm
Novirarrs Zoonoarcar XLI. 1939. 391
change from before backwards. In the hinder region there is a single needle-like
simple bristle in the feet.
Remarks.—These specimens agree well with Langerhans’s description
except that he states that the bristles are bidentate. He also figures two bristles
(fig. 9, b and c), and of these fig. 9, c shows a very minute second tooth, but fig. 9, b
appears to be unidentate, as are the bristles of my specimens. In the Syllids it is
often difficult to decide whether a bristle is bidentate or not because the hairs at
the top of the concave surface of the blade suggest the presence of a minute second
tooth.
Augener’s O. australis is an allied species, but according to his description
the longer dorsal cirri have about 40 articles, the bristles are clearly bidentate
anteriorly and unidentate posteriorly and the papillation is apparently inconstant.
Genus Trypanosyllis Claparéde 1864.
17. Trypanosyllis zebra Grube.
Fauvel, 1923, p. 269, fig. 101, a-e; and 1932, p. 78.
Occurrence.—Suez, mud flat (coll. Potts) (1); Zanzibar (4); Red Sea (6) ;
Tahiti (1).
Distribution.—Atlantic, Mediterranean, Indian Ocean, Pacific.
Remarks.—The specimen from Tahiti is unfortunately much damaged, and
although I believe it to belong to this species its determination remains uncertain.
Crossland has the following note on it: “ Has a long body tinged pink like the
dorsal cirri which are finely annulated, banding confined to the pharyngeal area,
two bands of equal length per segment over a pink or yellow, not white, ground.
One specimen has a sexual bud full of pink eggs.” There is also a female sexual
bud which from its highly flattened shape and the character of its bristles may
very well belong to this species. Crossland has the following note: “ Size
15 mm. by 3 mm. when dead, not seen alive. Washed out of coral from shore
reef at Taunoa. Colour creamy, apparently full of sexual products, with
rather vaguely outlined red-brown intersegmental lines and specks of same
colour between the feet. These lines also on the ventral surface. Dorsal
cirri fairly long, slender, moniliform, pink. Head appears scarcely to be made
out, but 4 very large red eyes. Dorsal cirri of first two segments long and
thicker than ordinary. Feet uniramous, setae directed upwards at angle
of about 45 degrees.”
18. Trypanosyllis misakiensis Izuka 1912.
Izuka, 1912, p. 185, pl. xx, figs. 2-6.
Fauvel, 1932, p. 79.
Occurrence.—Red Sea, Crossland (2).
Distribution. Japan, Madras Coast, Red Sea.
Remarks.—Iwo milk-white specimens without any buds. The absence of
pigment together with the separation of the apical from the second tooth of the
chaetal blades suggests that these specimens belong to this species. They also
agree with Izuka’s account in the shape of the head and the form of the dorsal
cirri.
392 NoVvITATES ZooLoGICAE XLI. 1939.
FAMILY HESIONIDAE.
Genus Leocrates Kinberg 1864.
19. Leocrates claparedii (Costa).
Fauvel, 1923, p. 237, fig. 88, i-n ; and 1932, p. 61.
Occurrence.—Red Sea (1) ; Tahiti (7).
Distribution. —Mediterranean, Red Sea, Indian Ocean, Pacific.
Remarks.—Of the specimens from Tahiti Crossland writes as follows:
““ Among coral Taunoa shore reef. No pigmentation or markings except a dark
brown line on the peristomium. General colour a dull red-brown, one specimen
dark purple (?) and containing eggs. Iridescence very brilliant, coppery and
purple. ‘Cushions’ at sides of body little developed anteriorly. Feet with
stiffer, slightly greenish ventral setae and fan of delicate ones above. Feet
colourless and base of cirri colourless and inconspicuous. Dorsal cirri brown,
long and slender. No visible segmentation, fine wrinkles across body about 10
to the segment. Size largest specimen 32 mm., rest rather smaller.”
Fauvel gives the colour in life as rose-chair irisé.
Genus Hesione Savigny 1809.
20. Hesione genetta Grube 1878.
Horst, 1924, p. 193.
Monro, 1931, p. 10, text-fig. 5.
Occurrence.—Tahiti (1) ; Barrier edge, Faa, Tahiti (1).
Distribution.—Philippines, Malay Archipelago, Great Barrier Reef, Tahiti.
Remarks.—These specimens have the back covered with brownish purple
spots, confluent in places and forming bars. In the bristles the accessory tooth
of the chaetal guard approaches the subapical tooth. Crossland writes as
follows: “ Among coral Taunoa shore reef. General colour the usual dull pink,
but this specimen is mottled with brown on a pinkish white ground throughout
its length. The mottlings are irregularly rounded, fewest but larger on the
“side cushions’ and aggregated in each segment to form a median irregular
triangular blotch in front of which is a space with less colour. These brown
markings are least regular in the anterior segments where the colour is rather
more developed. Bases of dorsal cirri not conspicuous, about the same colour
as the feet, transparent whitish. Dorsal cirri very slender, tinged brownish
yellow. Setae greenish yellow. Acicula conspicuous. Underside marked with
pink spots segmentally (? ganglia). Prostomium also pink. One pair of eyes
prominent. Spotted character of the coloration is conspicuous to naked eye.
Size alive 54 by 5 mm. or across feet (not setae) 9 mm.”
REFERENCES.
Augener, H. 1913. Polychaeta—I. Errantia: Michaelsen, W., and Hart-
meyer, R. Die Fauna Südwest-Australiens, vol. iv, pp. 65-304, pls. 1
and iii, 42 text-figs.
— 1927. Polychaeten von Curacao. Bijdragen tot de Kennis der Fauna van
Curacao. Resultaten eener Reis van Dr. C. J. van der Horst in 1920. Pp.
38-82, 9 text-figs.
Novirares Zoonoarcan XLI. 1939. 393
Berkeley, E. and ©. 1938. Notes on Polychaeta from the coast of Western
Canada. Ann. Mag. Nat. Hist. (11), i, pp. 33-49, 12 text-figs.
Fauvel, P. 1923. Polychetes errantes. Faune de France, v, pp. 488, 181 figs.
1930. Supplement to the littoral Fauna of Krusadai Island in the Gulf of
Manaar. Annelida Polychaeta. Bull. Madras Govt. Mus., New Series,
Nat. Hist. Section, vol. 1, no. 2, pt. 1, pp. 72, 18 text-figs.
—— 1932. Annelida Polychaeta of the Indian Museum, Calcutta. Mem.
Indian Mus., xii (1), pp. 262, pls. 9, 18 text-figs.
Gravier, C. 1900. Contribution a l’etude des Annelides Polychetes de la Mer
Rouge. Nowy. Arch. Mus. Hist. Nat. Paris (4), ii, pp. 137-182, pls. ix-
xiv.
Haswell, W. A. 1886. Observations on some Australian Polychaeta. Proc.
Linn. Soc. N.S.W., vol. x, 1885, pp. 733-756, pls. 50-55.
Izuka, A. 1912. The Errantia Polychaeta of Japan. Tokyo J. Coll. Sci.,
XXX, pp. 262, pls. 24.
Langerhans, P. 1879. Die Wurmfauna von Madeira. Zeits. Wiss. Zool.,
Xxxii, pp. 513-591, pls. 31-33.
Monro, C. C. A. 1933. On a collection of Polychaeta from Dry Tortugas,
Florida. Ann. Mag. Nat. Hist. (10), xii, pp. 244-269, 12 figs.
—— 1933a. The Polychaeta Errantia collected by Dr. C. Crossland at Colon,
in the Panama Region, and the Galapagos Islands during the Expedition
of the S.Y. St. George. Proc. Zool. Soc. London, i, 1933, pp. 1-96, 36
figs.
394 NOVITATES Zo0LOGICAE XLI. 1939.
ON SOME TROPICAL POLYCHAETA IN THE BRITISH MUSEUM,
MOSTLY COLLECTED BY DR. C. CROSSLAND AT ZANZIBAR,
TAHITI AND THE MARQUESAS.
By C. C. A. MONRO,
Department of Zoology, British Museum (Natural History).
(With 4 text-figures.)
III. FAMILY NEREIDAE.
Genus Nereis L. 1758.
1. Nereis jacksoni Kinberg 1865.
Fauvel, 1932, p. 97, with eitations.
Occurrence.—Suez (1).
Distribution.—Red Sea, Indian Ocean, Pacific.
Remarks.—A single specimen in poor condition that apparently belongs to
this species. It is characterized by the presence in the hinder feet of one or
two homogomph falcigers with bi- or tridentate blades and also by having only
a single row of paragnaths in VII and VIII.
2. Nereis coutierei Gravier, 1901.
Gravier, 1901, p. 167, pl. xi, figs. 38-41 ; text-figs. 166-172.
Fauvel, 1932, p. 96.
Occurrence.—Suez (9).
Distribution.—Red Sea, Indian Ocean.
Remarks.—This small species is characterized by a single row of paragnaths
in VII and VIII, smooth, homogomph falcigers only in the dorsal ramus of the
posterior feet and the enlargement of the upper dorsal lamella in the hinder
region.
3. Nereis gisserana Horst 1924 (text-fig. 302).
Horst, 1924, p. 151, pl. xxx, figs. 6-7.
Occurrence.—Coll. J. Stanley Gardiner, Suvadiva, Maldives (4); Saya de
Malha, Maldives (1) ; Amirante Ids., 30 ms. (1).
Distribution.—Malay Archipelago, Maldives, Amirantes.
Description.—These specimens are all incomplete and in rather poor condition.
The body is swollen anteriorly with genital products, but there is no modifi-
cation of the feet. The largest measures 45 by 2 mm. at the widest part without
the feet for about 90 chaetigers. The palps are short and stout. The tentacular
cirri are unusually long, the longest reaching back to about the 15th chaetiger.
The arrangement of the paragnaths (text-fig. 302, A and B) is as follows: I, 0;
II, a transverse row of about 5 paragnaths, in front of which a second rudimen-
tary row can sometimes be with difficulty seen ; III, 3 groups: a middle group
consisting of about 3 transverse rows of small paragnaths, and two lateral groups
each consisting of 2 paragnaths ; IV, a small patch of rather larger paragnaths
with distally 3 or 4 large transverse bars; V, 0; VI, 3 in a transverse row ;
Novirarrs Zoonoaicar XLI. 1939. 395
VII and VIII, a single row of 5 or 6 widely separated paragnaths. The armature
of the proboscis agrees well with Horst’s description except that Horst gives 2 to
5 paragnaths in a longitudinal row in Group I. These are absent in my material.
In regard to the feet, in the anterior region the languets are short and blunt,
but they gradually lengthen out from before backwards. A foot from the middle
region (text-fig. 302, C) has a long dorsal cirrus extending well beyond the end
TEXT-FIG. 302.—Nereis gisserana : A, head from above; B, head from
below; C, middle foot; D, posterior foot; E, large falciger from middle
foot ; F, large faleiger from hinder foot.
of the foot, a conical upper dorsal languet, a shorter lower dorsal languet that is
fused for part of its length with a slightly smaller middle dorsal languet. The
ventral chaeta sac has two equal digitiform lips, and there is a rather small
ventral languet, below which is a short ventral cirrus. In the hinder region
(text-fig. 302, D) the middle dorsal languet disappears, and the dorsal ramus
greatly overshadows the ventral, but there is no special development of the
upper dorsal languet.
The dorsal ramus carries homogomph spinigers, the upper ventral ramus
396 NoviratEes ZooLoGIcAE XLI. 1939.
carries homogomph spinigers and two or three rather large heterogomph falcigers
(text-fig. 302, Eand F). The lower ventral ramus carries one or two heterogomph
spinigers and a number of falcigers considerably smaller than those in the upper
ventral ramus.
Remarks.—The most remarkable character of this species is the large
transverse bars in Group IV of the proboscis. I know no other nereid with a
similar structure. Moreover, the fusion basally of the lower and middle languets
of the feet is very unusual.
4. Nereis semperiana Grube 1878 (text-fig. 303).
Grube, 1878, p. 77, pl. iv, fig. 6.
Fauvel, 1935, p. 303.
Occurrence —Tahiti (2).
Distribution. —Philippines, Java, Indo-China, Tahiti.
Remarks.—Two very small specimens in which the arrangement of the
paragnaths beyond the fact that there is a single row in VII and VIII cannot
be made out. There is a dark brown
ea dorsal band covering the buccal and first
303 ja chaetigerous segments. The feet (text-
fig. 303, A) are unusual. The dorsal
cirrus is swollen basally and tapers to a
point distally. The two dorsal languets
are unequal, the upper finger-shaped and
considerably longer than the blunt and
conical lower dorsal languet. The ventral
languet is very small and papilliform and
the ventral cirrus is also very small.
The dorsal bristles are homogomph
spinigers, the upper ventral bundle con-
tains homogomph spinigers and one or
two rather large heterogomph falcigers. The bristles of the lower ventral
bundle are all heterogomph falcigers with extremely small end-pieces (text-fig.
303, B) recalling those of the Syllids and of the type figured by Fauvel for his
Leptonereis foli. "This species is allied to N. unifasciata Willey, but the shape of
the feet and especially that of the lower ventral languet is different. Crossland
has the following note: “ Broad brown bands, of a red chocolate tint, particu-
larly on peristomium and next segment. These are from weed on basal parts of
the ‘stag’s horn’ coral Acropora (?) pulchra on the shore and lagoon reefs of
Taunoa Passage, in which it seems to be the common nereid. Large examples
of 234 (Perinereis cultrifera var. obsfuscata) are also present, and this species is
found in other washings as well. The worm is small and slender, yellow in
colour owing to the gut, feet transparent, sometimes opaque white, apparently
owing to sexual products. The two broad bands on the peristomium and the
next segment are the main masses of colour. The same colour may be found
on the prostomium in two longitudinal lines. All head appendages colourless
and rather short. Segments 2-3 colourless, 4, 5, 6 have broad bands of light
brown, but all have a couple of dark brown blotches laterally, backed by a white
line across the whole back. These are faint on segments 2-3 and died out at
12-15. Underside colourless.”
Text-Fia. 303.—Nereis semperiana :
A, middle foot; B, small falciger.
Novirarres Zoonocicarn XLI. 1939. 397
5. Nereis picteti Malaquin and Dehorne 1907 (text-fig. 304).
Malaquin and Dehorne, 1907, p. 336, figs. 1-4.
Occurrence.—Seychelles, 34 ms. (1).
Distribution.—Amboina, Seychelles.
Remarks.—An anterior fragment measuring 18 by 4 mm. without the feet
for 23 chaetigers. The head is deeply embedded in the buccal segment, which
at the sides extends beyond the anterior pair of eyes. The tentacles are about half
as long as the head, and the tentacular cirri reach back to the 5th chaetiger.
The arrangement of the paragnaths, which are heavy and boldly defined, agrees
fairly well with Malaquin and Dehorne’s figures. They are as follows: Group I,
3 in a line; II, a curved row of about 6; III, a straggling row of about 7, some
large, some small; IV, 4 in an irregular line; V,0; VI, 3 ina triangle; VII
and VIII, a single row of about 5 paragnaths. Malaquin and Dehorne give
Group III as 3 in a triangle, and their
number of paragnaths in VII and VIII is 04mm 304 N
low 0:02mm \\
ower. \
In the feet the dorsal cirrus is equal A Bl 7.
to the upper dorsal languet. There are
three dorsal languets. The upper and
lower are rather large, conical and equal,
and the middle is slender and about half
the size of the other two. There is a
ventral languet about equal to the lower
dorsal languet and a small ventral cirrus
(text-fig. 304, A).
The dorsal bristles consist of homo-
gomph spinigers only. The upper ventral
bristles are homogomph spinigers and ne A Nee
about half a dozen large heterogomph A, mel fois 3, lm len.
faleigers (text-fig. 304, B) in which shaft
and blade show signs of fusion. In my very small material I have not found
one in which the articulation has disappeared, but they are clearly of the type
found in N. anchylochaeta Horst and Ceratonereis pachychaeta Fauvel.
The lower ventral bundle consists of heterogomph spinigers and small
heterogomph falcigers.
As far as can be seen on the present material this species is closely allied
to N. anchylochaeta Horst. It differs in having a small patch of paragnaths in
VI instead of 1 or none ; in having the middle dorsal languet much smaller than
the rest instead of equal and in having the large heterogomph falcigers confined
to the upper ventral bundle. More material is required to decide whether the
joint in the latter actually disappears as in N. anchylochacta, but I suspect that
it does.
Genus Perinereis Kinberg 1865.
6. Perinereis cultrifera (Grube) 1840.
Fauvel, 1923, p. 352, fig. 137, a-1; and 1932, p. 104.
Occurrence.—St. Vincent, shore, Cape Verde Islands (3); Tahiti, from
mossy green brown weed on vertical face of Fautaua Reef. Stones exposed to
surf just W. of Fautaua Reef (v. numerous juv.) ; Hiva Oa, Marquesas (1).
398 Novirates ZooLoGIcAE XLI. 1939.
Distribution. —Cosmopolitan.
Remarks.—These specimens have the arrangement of paragnaths proper to
the typical form, ie. I, 1 or 2ina line; V, 3inatriangle. The tentacular cirri
are usually described as reaching to the 5th or 6th chaetigers. In most of these
specimens they are rather shorter, reaching to the 3rd or 4th chaetigers. The
common Nereid in the washings from Tahiti is P. cultrifera var. obfuscata, of
which very large numbers were obtained in the immature stage. It is remark-
able to find a patch of the typical cultrifera taken by Crossland from the face of
Fautaua Reef, Tahiti, surrounded, so to speak, by the obfuscata variety. The
slight differences between the various forms of this species may be correlated
with differences in habitat, but in our present state of knowledge we cannot so
correlate them. It should also be noted that the same washing produced a
number of var. obfuscata, but these appear on the whole to be at a slightly later
stage of development than the examples of the typical cultrifera.
7. Perinereis cultrifera (Grube) var. obfuscata Grube 1878.
Perinereis obfuscata Grube, 1878, p. 86.
Perinereis obfuscata Monro, 1931, p. 16, fig. 10, a-d.
Occurrence.—Tahiti, numerous young specimens from various washings.
Distribution.—Philippines, Malay Archipelago, Great Barrier Reef, Tahiti.
Remarks.—Group I, a small patch of 4 or 5 little paragnaths ; V, a single
paragnath. The tentacular cirri reach back to about the 5th chaetiger and not
to the 3rd as Grube states. In this they approach Grube’s var. striolata,
distinguished only by the fact that the tentacular cirri reach back to the
9th chaetiger.
Crossland has the following notes: ‘ Inner reefs of Taunoa Passage. Never
seen when taken from the sea, but very abundant on dead sheets of the umbrella-
like Acropora hyacinthus when broken up and put into water with magnesium
sulphate. The specimens include a ripe heteronereid and a ripe male not changed
in form. Feet and sides of anterior half of body transparent, but posteriorly
speckled whitish or with light yellow marks. Back strongly marked with
greenish brown spots, which extend to hind end, though becoming smaller and
less conspicuous. Prostomium to base of tentacles and between eyes coloured,
palps white, white line left between prostomium and peristomium. Mid-dorsal
part of peristomium and Ist and 3rd setigerous segments nearly uniformly
coloured with broad intersegmental colourless lines. Deepest colour on anterior
and posterior parts of each block (of spots) and there are whitish diagonal lines
latero-posteriorly, two on each side. Tentacles and tentacular cirri short.
The worms are small, the larger among them and of much the same colour are
Eunice antennata (?). In larger specimens the blotches seem to run into one
another and pattern seems less distinctive.
“ From caulerpa on the Fautaua Reef edge. Probably the same species but
colour greenish grey with white intersegmental lines and white line diagonally
laterally. Grey colour forms a deeper band just behind anterior margin and in
small specimens in posterior part of the segment also. Palps as well as prosto-
mium are here brown. This isthe worm characteristic of the bare lithothamnium
outer slope of the Fautaua Reef edge, just as N. dumerilii is of the coralline weed
band. Compare the outer reef edge which provided hosts of E. antennata here
apparently absent. Variations of the markings are mostly due to age. Again
Novirates ZooLoGIcAE XLI. 1939. 399
dominant in mossy, green brown weed of Fautaua Reef edge in which the
Opheliid is also abundant.”
8. Perinereis cultrifera (Grube) var. perspicillata Grube 1878.
Perinereis perspicillata Grube, 1878, p. 90, pl. iv, fig. 10.
Perinereis cultrifera (Grube) var. perspicillat1 Fauvel, 1932, p. 105.
Occurrence.—Red Sea (4) ; Suez (4) ; Zanzibar (numerous heteronereids).
Distribution. —Philippines, Singapore, Red Sea, Persian Gulf.
Remarks.—Group I, a small patch of 4 to 8 paragnaths ; V, 3 in a triangle.
According to Fauvel this variety is also found on the French coast.
9. Perinereis cultrifera (Grube) var. helleri Grube 1878.
Perinereis helleri Grube, 1878, p. 81.
Perinereis helleri Monro, 1931, p. 14, fig. 8, a-c.
Perinereis camiguina Grube, 1878, p. 87.
Perinereis cultrifera (Grube) var. helleri, Fauvel 1932, p. 105, with citations.
Occurrence.—Coetivy (J. S. Gardiner) (3) ; Zanzibar, Chuaka Bay (2).
Distribution.—Philippines, Indian Ocean, Red Sea, Great Barrier Reef.
Remarks.—Group I, 3 in a triangle; II, 1 to 3 in a line. Tentacular cirri
reach to 6th-9th chaetigers.
10. Perinereis vancaurica (Ehlers) 1864.
Monro, 1931, p. 14, with citations.
Perinereis vancaurica Fauvel, 1932, p. 103, with citations.
Occurrence.—Suakim Harbour (3).
Distribution. —Red Sea, Indian Ocean, Pacific.
Remarks.—The specimens have been allowed to dry and are in very poor
condition. They are recognizable only by the distinctive arrangement of the
paragnaths. The maxillae often have no secondary teeth. In these specimens
they are faintly serrate.
11. Perinereis nuntia (Savigny) 1820.
Fauvel, 1919, p. 415, with citations.
Occurrence.—Suez, mud flats (6).
Distribution.—Red Sea, Indian Ocean.
Remarks.—This is the typical form with long tentacular cirri and narrow,
elongated pedal languets. Group I consists of 1 or 2 small paragnaths and V
of 3 in a triangle.
12. Perinereis nuntia (Savigny) var. heterodonta Gravier 1901.
Fauvel, 1919, p. 419, with citations.
Occurrence.—Suez (numerous).
Distribution.—Red Sea, Persian Gulf.
Remarks.—A large form with all the paragnaths of Group VI of the trans-
versely elongated, cutting type. Group I has 1 or 2 paragnaths, and V none.
The tentacular cirri are short.
400 NOVITATES ZOOLOGICAE XLI. 1939.
13. Perinereis nuntia (Savigny) var. brevicirris Grube 1867.
Fauvel, 1919, p. 417, with citations.
Occurrence.—Red Sea (10) ; Zanzibar (numerous heteronereids).
Distribution.—Red Sea, Japan, Australia.
Remarks.—This is the form with short tentacular cirri and blunt pedal
languets. The arrangement of the paragnaths is the same as in the stem-form.
The heteronereids from Zanzibar were partly this variety and partly var. vallata.
The only reliable way to distinguish the two varieties was by the presence or
absence of the triangular group of 3 paragnaths in V.
In my opinion they scarcely deserve two varietal names.
14. Perinereis nuntia (Savigny) var. vallata Grube 1857.
Fauvel, 1919, p. 418, with citations.
Occurrence.—Chuaka Bay, Zanzibar (numerous heteronereids).
Distribution.—Red Sea, Cape of Good Hope, Australia, Philippines, Chile.
Remarks.—This is the variety with short tentacular cirri, 1 to 3 paragnaths in
I, and a single paragnath in V. I have given my opinion of its relation to var.
brevicirris under the heading of the latter.
Genus Ceratonereis Kinberg 1865.
15. Ceratonereis mirabilis Kinberg 1865.
Fauvel, 1932, p. 98, with citations.
Occurrence—Amirante (1 juv.); Wasin, Zanzibar (3); Chuaka shore,
Zanzibar (1).
Distribution. Atlantic, Red Sea, Indian Ocean, Pacific.
Remarks.—This common circumtropical species is represented by very poor
material.
16. Ceratonereis costae (Grube) 1840.
Fauvel, 1919, p. 402, pl. xvii, figs. 87-88 ; and 1923, p. 349, fig. 136, a-f.
Occurrence.—Wasin, Zanzibar (2); floating dock, Suez (5); Suez (3); Red
Sea, from coral (1).
Distribution.—Atlantic, Mediterranean, Red Sea, Indian Ocean, Pacific.
Remarks.—The specimens from the floating dock, Suez, are remarkable for
having a paragnath in Group I, where no paragnath is normally found. This
species is identical with both C. fasciata Grube and C. lapinigensis Grube.
C. fasciata is also described as having a paragnath in I.
17. Ceratonereis pachychaeta Fauvel 1919.
Fauvel, 1919, p. 403, pl. xv, figs. 22-25 ; fig. viii, a-g; and 1933, p. 57.
Occurrence—Hululu Male, Maldives (2) and Coetivy (coll. J. Stanley Gardiner)
(1); Suez, quayside (4) ; Dongonaab Harbour, Red Sea (3); Tahiti (2); Papeete
Harbour, Tahiti, 15 fins (1).
Distribution.—Red Sea, Madagascar, Gambier Islands, Indian Ocean,
Pacific.
Remarks.—This is a well-characterized and easily recognizable species.
The paragnaths of the various groups often appear to be fused into solid, toothed
bars like Eunicid jaw-plates. Crossland has the following note: “ Washings of
Novirares ZooLoGIcAE XLI. 1939. 401
“ coralline ’ weed from Fautaua Reef edge. Conspicuous bright red with yellow
tinge, making its colour resemble that of Oenone but for the worms being short.
This colour extends the length of the body without diminution, and feet with
large dorsal lobes strongly coloured increase the resemblance to Oenone. Colour
remarkably uniform. Cephalic appendages all short, nearly colourless, as is the
middle part of the prostomium. Head end the broadest part of the body, giving
it a truncated appearance. Larger than any preceding Tahitian Nereid.”
Genus Platynereis Kinberg 1865.
18. Platynereis coccinea (Delle Chiaje) 1827.
Fauvel, 1923, p. 360, fig. 141, g-n.
Occurrence—Wasin, Zanzibar (4) ; Zanzibar (2).
Distribution.—Azores, Sargasso Sea, Mediterranean, ? Indian Ocean.
Remarks.—Having failed to find any paragnaths except in Group IV, I am
reluctantly forced to attribute these Zanzibar specimens to this species, which
has hitherto to the best of my knowledge not been recorded outside Atlantic or
Mediterranean waters.
19. Platynereis pulchella Gravier 1901.
Gravier, 1901, p. 202, pl. xii, figs. 55-56, text-figs. 210-213.
Platynereis dumerilit var. pulchella Fauvel, 1911, p. 402, pl. xx, figs. 30-32.
Platynereis festai Holly, 1935, p. 27, pl. i, figs. 7, k, and text-figs. 13-15.
Platynereis pulchella Monro, 1937, text-fig. 10, a-n.
Occurrence.—Red Sea (1 epitocous and 2 atocous) ; Dongonaab Harbour,
Red Sea (2 epitocous and 4 atocous).
Distribution —Red Sea, Persian Gulf, Arabian Coast, Hawaii.
Remarks.—I have already (loc. cit.) published an account of the curious
heteronereid of this species which in its atocous stage is practically inseparable
from P. dumerilii, and the present damaged material does not permit me to add
anything to it. It is very possible that some of the young specimens that I have
attributed to P. dumerilii do in fact belong here.
20. Platynereis dumerilii (Audouin and M.-Edwards) 1834.
Fauvel, 1923, p. 359, fig. 141, a-f; and 1932, p. 113.
Occurrence—Egmont Reef (1 juv.) and Coetivy (coll. J. Stanley Gardiner)
(3 juv.); Wasin, Zanzibar, 10 fins (10 juv.); Tahiti from various washings
(v. numerous juv.) ; Marquesas, from blue sponge (5), and from washing (2).
Distribution. — Cosmopolitan.
Remarks—Young specimens of this widely known species are extremely
abundant in the Tahiti washings, but relatively rare in those from the Marquesas.
When the material is immature and in the absence of heteronereids I cannot
confidently distinguish this species from Gravier’s P. pulchella. Crossland has
the following note: “ Reddish throughout, due to pink specks on a light green
ground, as well as the blood-vessels. Bases of the long tentacular cirri red,
themselves all translucent green colour. White dots middle each segment
dorsally and specks scattered over the back; these specially conspicuous
posteriorly. Notopodia particularly pink, no yellow spots. Eyes large, brown
with a small black pupil. Body broadest just behind head, Builds a trans-
29
402 NOVITATES ZooLoGIcAE XLI. 1939.
parent mucus tube in which body undulates (?) to draw water through. The
worm can turn in the tube. Very active when out of the tube.
“ I think this is the universal P. dumerilii, and there is absolutely nothing
new in all this, but copy in case.”
Genus Pseudonereis Kinberg 1865.
21. Pseudonereis anomala Gravier 1901.
Gravier, 1901, p. 191, pl. xii, figs. 50-52, figs. 194-202.
Fauvel, 1932, p. 112.
Occurrence.—Red Sea (8) ; Suez (3) ; from buoy in Canal entrance, Suez (3).
Distribution. —Red Sea, Indian Ocean, Malay Archipelago, Australia.
Remarks.—Most of these are young specimens. In one or two of those from
Suez the dissected proboscis appeared to show the paragnaths of VI in clusters
instead of a single row, but the presence of dorsal homogomph falcigers dis-
tinguishes them from Ps. rottnestiana Augener and the enlargement of the dorsal
ramus of the foot in the hinder region from Ps. masalacensis Fauvel (? non Grube).
22. Pseudonereis variegata (Grube) 1857.
Pseudonereis gallapagensis Kinberg, Fauvel 1932, p. 111, with citations.
Occurrence.—Tahiti (3) ; Funufuti, outer reef (numerous juv.).
Distribution.—Tropical and southern temperate regions of the Atlantic,
Indian and Pacific Oceans.
Remarks.—Fauvel, following Gravier, has recently adopted Kinberg’s name
for this species. As Fauvel (loc. cit.) points out, Kinberg’s figure of gallapagensis
and Grube’s description of variegata both appeared in 1857, and the priority is
doubtful. I prefer to retain Grube’s name, which has been more generally used.
Crossland writes of the Tahiti specimens as follows: “ From ‘ mossy’ green
brown weed on stones exposed to surf W. of the Fautaua Reef. Of a fair size.
Head is the broadest part of the body. Tentacles and tentacular cirri short.
No obvious structural difference from large specimens of 245-234 (Perinereis
cultrifera var. obfuscata Grube), but head seems broader and palps fatter and
shorter. Body coloured a deep green-blue, darkest on the prostomium. Anterior
half of palps is coloured, the rest white. This colour extends to tail but becomes
fainter after first third of the body length. Feet, tentacular and dorsal cirri
white, as are end pieces of the palps. Intersegmental grooves white, throwing
up darker band of pigment across anterior part of each segment. Faint diagonal
lines laterally.”
23. Pseudonereis masalacensis Fauvel 1923 (? non-Grube) 1878.
Fauvel, 19234, p. 39; and 1930a, p. 532, fig. vii, a-k.
Occurrence.—Praslin, Seychelles, J. Stanley Gardiner (3).
Distribution.—Gambier Islands, New Caledonia, Seychelles, ? Philippines.
Remarks.—These small specimens agree closely with Fauvel’s descriptions.
The proboscis is withdrawn, and as far as I can see the paragnaths in VII and
VIII are more numerous than he describes them. Otherwise in the feet, the long
dorsal cirri, the dorsal homogomph, posterior falcigers, ete., they are indistin-
guishable. The species is close to Gravier’s Ps. anomala, but differs in the
paragnaths of the oral ring and in the absence of change of the dorsal ramus in
the hinder region.
Noviratres ZOoLOGICAE XLI. 1939. 403
I have already (1933, p. 43) expressed a doubt as to whether this is in fact
Grube’s species, and Fauvel himself in a later publication (1935, p. 314) shows
a similar uncertainty.
Genus Leonnates Kinberg 1865.
24. Leonnates jousseaumei Gravier 1901.
Gravier, 1901, p. 160, pl. xi, figs. 34-37.
Fauvel, 1930, fig. 5, a-e.
Occurrence.—Diego Garcia, J. Stanley Gardiner (1) ; Suez (numerous juv.) ;
Red Sea (1); Wasin, Zanzibar, from cavities of a sponge (2).
Distribution.—Red Sea, Indian Ocean. i
Remarks.—In young and ill-preserved specimens I find it often impossible
to see any paragnaths on the maxillary ring. The homogomph falcigers in the
feet are unlike those of any other species.
25. Leonnates simplex n. sp. (text-fig. 305).
Occurrence.—Aldabra coast (1).
Description. —The specimen is more or less complete and measures 53 by
2 mm. without the feet for 110 chaetigers. The palps are long and slender,
TEXT-FIG. 305.—Leonnates simplex: A, head from above; B, head
from below ; C, middle foot; D, hinder foot ; E, faleiger from middle foot.
and the tentacular cirri rather short, the longest reaching back to the 6th chaetiger.
There is no colour.
The maxillary ring of the proboscis carries paragnaths and the oral ring only
papillae, The arrangements is as follows (text-fig. 305,.A and B): Group I,
404 NOVITATES ZOOLOGICAE XLI. 1939.
2 in line; II, a small rounded patch ; III, a small transverse patch ; IV, rather
long oblique patches; V, 3 papillae in a triangle; VI, a transverse ridge of
several small papillae ; VII and VIII, a band of papillae two of three deep.
Both feet and bristles are very simple. There is a dorsal cirrus (text-fig.
305, C) about equal to the upper dorsal languet. In the dorsal ramus the upper
and lower dorsal languets are about equal. There is a rounded ventral chaetal
lobe ; the ventral languet is a little smaller than the dorsal and the ventral
cirrus is shorter than the ventral languet. In the hinder region (text-fig. 305, D)
the dorsal ramus dominates the ventral, there is a large black gland at the base
of the dorsal cirrus, and both dorsal and ventral cirri are longer relatively to the
languets.
The dorsal ramus carries homogomph spinigers, the upper ventral ramus
homogomph spinigers and heterogomph falcigers, the lower ventral ramus one
or two heterogomph spinigers and about half a dozen heterogomph falcigers
(text-fig. 305, E). I see nothing distinctive about either feet or bristles. The
pygidium is damaged.
Remarks.—L. jousseaumei Gravier and L. decipiens Fauvel both have very
distinctive falcigers and differ in several other characters. L. nierstraszi Horst
has three dorsal languets. JL. ehlersi Augener is a highly modified species of a
quite different character. L. indicus Kinberg I find unrecognizable. L. virgata
Grube has three dorsal languets and is figured with a group of papillae in the
maxillary ring.
REFERENCES.
Fauvel, P. 1911. Annélides Polychétes du Golfe Persique. Arch. Zool. exp.
gen., Vi, pp. 353-439, pls. xix—xxi.
1919. Annélides Polychétes de Madagascar de Djibouti et du Golfe Per-
sique. Arch. Zool. exp. gén., viii, pp. 315-473, pls. xv—xvii, 12 text-figs.
— 1923. Polychétes errantes. Faune de France, v, pp. 488, 181 figs.
—— 19234. Annelides Polychetes des Iles Gambier et de la Guyane. Mem.
Accad. Nuovi Lincei Roma, Ser. 2, vi, pp. 89-147, 8 text-figs.
—— 1930. Supplement to the Littoral Fauna of Krusadai Island in the Gulf of
Manaar. Annelida Polychaeta. Bull. Madras Govt. Mus., New Series,
Nat. Hist. Section, i (2), i, pp. 72, 18 figs.
—— 19304. Annélides Polychétes de Nouvelle-Calédonie. Arch. Zool. exp.
gen., \xix (5), pp. 501-562, 9 figs.
—— 1932. Annelida Polychaeta of the Indian Museum, Calcutta. Mem.
Indian Mus., xii (1), pp. 262, pls. 9, 18 text-figs.
—— 1933. Mission Robert Ph. Dollfus en Egypte: Annelides Polychétes.
Mem. Inst. Egypte, xxi, pp. 31-83, 4 text-figs.
—— 1935. Annélides Polychétes de l’Annam. Mem. Accad. Nuovi Lincei
Roma, Ser. 3, ii, pp. 279—354, 8 text-figs.
Gravier, ©. 1901. Contribution 4 l’étude des Annelides Polychetes de la Mer
Rouge. Nouv. Arch. Mus. Hist. Nat., Paris (4), iii, pp. 147-268, pls. vii-x.
Grube, E. 1878. Annulata Semperiana. Beiträge zur Kenntniss der Anne-
liden Fauna der Philippinen. Mém. Acad. sci. St. Pétersb. (7), xxv, No. 8,
ix, pp. 300, pls. 15.
Holly, M. 1935. Polychaeta from Hawaii. Bishop Mus. Bull., cxxix, pp. 33,
pl. 1, 15 text-figs.
Noviratres ZOOLOGICAE XLI. 1939. 405
Horst, R. 1924. Polychaeta errantia. “ Siboga” Haped. Mon., xxiv (c),
pp. 145-198, pls. xxx—xxxvi.
Malaquin, A., and Dehorne, A. 1907. Les Annelides Polychetes de la Baie
d’Amboine. Rev. Suisse Zool., pp. 335-400, pls. 51-58, 20 text-figs.
Monro, C. €. A. Polychaeta, Oligochaeta, etc. Great Barrier Reef Exped.,
1928-29. Sci. Reps., iv (1), pp. 37, 15 figs.
1933. The Polychaeta Errantia collected by Dr. C. Crossland at Colon,
in the Panama Region and the Galapagos Islands during the Expedition of
the S.Y. St. George. Proc. Zool. Soc. London, i, pp. 96, 36 figs.
—— 1937. Polychaeta. Murray Exped., 1933-34. Sci. Reps., iv (8), pp.
243-321, 28 figs.
406 Novirates ZOoOLOGICAE XLI. 1939.
THE RACES OF JAGUAR (PANTHERA ONCA).
By R. I. POCOCK
Department of Zoology, British Museum (Natural History).
(With 5 text-figures.)
SSNS and skulls, particularly skins, of Jaguars appear to be much less
plentiful in most of the museums of Europe and America than those of
the three old-world species of the genus, the Lion, Tiger and Leopard. This, at
all events, is the case in the British Museum, where the Jaguars have never
hitherto been systematically studied since the time of Gray. A good deal of
work upon the species has, however, been done in America, notably by Mearns,
Hollister, Nelson and Goldman, and Cabrera. References to their papers are
given below.
The most comprehensive contribution to the subject was the monograph of
the species in 1933 by Nelson and Goldman who supplied a valuable revision of
the bibliography from 1758 onwards. This portion of their paper has been
particularly helpful ; but the systematic portion, containing the descriptions of
the subspecies, of which many were new, would have been greatly enhanced in
value by the supply of a number of regrettable omissions regarding the actual
dimensions of the comparatively large number of skulls, borrowed from various
sources, which the authors had in their hands. At most a single example of
each sex of the claimed different kinds was selected for measurement, practically
no detailed information on the variations in size within subspecific limits being
given. In the discrimination of the subspecies, moreover, importance was
attached to a number of cranial differences known in the case of Lions, Tigers
and Leopards, let alone some other species of Carnivora, to fall within the range
of individual variation and to be, therefore, of no systematic significance. Similar
individual variation is shown by the skulls of the Jaguars I have seen, no two of
which are alike, even from the same locality. I follow these authors in admitting
only one species.
INDIVIDUAL VARIATION IN THE COLOUR AND PATTERN OF SKINS FROM THE
SAME Locatity.
Since both colour and pattern have been used as characters for distinguishing
various named forms, the variations in these features shown by two skins from
the same locality, the Upper Guapore River, Matto Grosso, presented by the
Marquis de Wavrin, are of special interest. One (no. 22.7.7.3) is a bright
tinted skin with the principal rosettes large and mostly entire, but only one of
them, close to the spine behind the shoulder, encircles a supplementary spot ;
on the forequarters, however, only a few rosettes are indicated dorsally, those
on the sides of the shoulder and outer surface of the foreleg being broken up
into solid spots like those of the Hunting Leopard. The other (no. 22.7.7. 2) is
noticeably duller in the tint of the ground colour, nearly all the principal rosettes
contain from one to four supplementary spots, and the pattern on the shoulders,
both above and laterally to the base of the foreleg, consists of well-defined
Novirarrs Zoonoaican XLI. 1939. 407
rosettes ; and rosettes are traceable up the nape to the occiput. A third skin
from Matto Grosso, without special locality (no. 99.3.2.1) is nearly intermediate
in its pattern between the other two.
Three skins from Buena Vista, Santa Cruz, Bolivia, also differ individually.
An adult & (no. 26.1.12.1) is ochreous buff in ground colour, with the edges
of the rosettes thinner and
more broken up. This skin
is merely a shade darker
than the brighter of the two
from the Guapore River,
Matto Grosso (no. 22.7.7.3).
An adult 2 from Buena Vista,
Santa Cruz (no. 26.12.4.29),
is decidedly duller, more
sandy buff in ground colour
than the g and has the rims
of the rosettes better defined,
thicker and less broken up.
The ground colour in this
exactly matches that of
the Matto Grosso skin no.
99.3.2.1. The skin of a
half-grown specimen from
Buena Vista, Santa Cruz, is
a still paler sandy buff in
ground colour than the 9
from that locality, with the
centres of the rosettes dus-
kier and more contrasted
with the ground colour.
Theindividualdifference
between theskinsfrom Matto
Grosso and Bolivia, above
pointed out, show that no
systematic importance can
be attached to such varia-
tions in the colour and TEXT-FIG. 306.—A, occipital area of adult g P. onca
onca from Teperinha, Santarem. B, the same of ad. 3g P.
If onca onca from Para (45.8.25.22).
pattern as they present.
the localities of the 5 adult
skins were unknown, it would be impossible to sort those from Matto Grosso
as distinct from those from Bolivia.
INDIVIDUAL VARIATIONS IN SKULLS FROM THE SAME LOCALITY.
Three adult skulls from Parad, 2 gg, 1 9, show that certain features that
have been cited in support of the admission of local races may fall within the
limits of individual variation. Their dimensions, entered in the table below,
show a difference in the condylobasal length in the two 3 specimens. The larger
of the two is just adult. It would not have grown appreciably in length, but in
zygomatic width would probably have increased considerably with age. Despite
408 Noyiratrs ZooLoGIcaE XLI. 1939.
its being younger, it has a higher sagittal crest, 16 mm. just behind the fronto-
parietal suture, whereas in the much older, smaller specimen the crest is only
10 mm. The two also differ greatly in the shape of the nasals. In the older
skull they are broad behind, 30 mm. between the points of the frontals ; in the
younger that width is 25 mm. (text-fig. 309). In the latter skull also the frontals
are appreciably less inflated with air-cells. The bullae of the smaller 3 are
actually and relatively larger than in the larger 3, and much larger than in the
Q, their length and width in the 3 being 35 and 26 mm., in the 9 26 and 22 mm.
In the ? also they are much flatter, projecting in profile view less deeply below
the auditory orifice (text-fig. 307, A—D).
Again, in two skulls, 3, 9, from Buena Vista, Santa Cruz, Bolivia, the
bullae differ somewhat
similarly. In the &
their length and width
are 35 and 30 or 32
mm.—they vary in
width on the two
sides—and in the 9
those dimensions are
29and 27mm. In the
6 also the mesoptery-
goid fossa, measured
from its inner edge,
is actually and rela-
tively considerably
wider, being 28 mm.,
whereas in the 9 it is
23 mm. There is no
Text-rig. 307.—A, lower view of right auditory bulla of ad. ¢ reason to suppose that
P. once onca from Para (45.8.25.23). B, lateral view of the same. the above recorded
C, lower view of right bulla of ad. 2 P. onca onca from Para diff intheball
(45.8.25.21). D, lateral view of the same. ifferencesin the bullae
and the mesopterygoid
fossa have anything to do with sex; and this is proved by the occurrence of
even greater difference in the two & skulls next described.
Two adult & skulls from the Magdalena Valley, Colombia (Mrs. Vaughan),
also differ a good deal in many details. They have approximately the same
condylobasal length, but one of them (no. 35.3.6.1) is an older skull than the
other (no. 35.3.6.2). Their principal measurements are given below. Setting
aside such differences as the wider zygomata, interorbital and mastoid areas,
attributable to age in the older skull, the frontal bones just behind the
postorbital processes are more inflated with air-cells affecting the shape of the
profile of this area of which the width immediately behind these processes is
53 mm. as compared with 47 in the younger skull. The older skull also has a
much wider mesopterygoid fossa and the bullae noticeably smaller in their three
dimensions, their narrowness making the basioccipal wider between them. The
actual measurements of these parts in the two skulls are as follows, those of
the older coming first : width of postpalate at interior margin of fossa 36 : 29 ;
inner width of fossa at pterygoid suture 27:21, greatest width of basiocei-
pital between bullae 29:23; length and width, at stylomastoid foramen, of
Noviratres Zoonoarcan XLI. 1939. 409
expanded part of bullae 31 and 22:32 and 24; depth of bulla below auditory
orifice 20 : 23.
These data attest that the inflation of the frontals with air-cells, the shape
of the nasals, the size of the bullae, the width of the mesopterygoid fossa and the
height of the sagittal crest may vary considerably in adult skulls of the same
TEXT-FIG. 308.—A, Mesopterygoid fossa of ad. g. P. onca centralis from Magdalena Valley,
Colombia (35.3.6. 2). B, the same of another ad. 3 from the same locality (35.3.6.1.) C, the
same of yg. ad. ¢ identified as P. onca goldmani (39.537) from Mexico.
sex from the same locality. Some other features that have been used for dis-
tinguishing the subspecies are also quite unreliable. It is, for instance, well
known that in a great many genera of Carnivora, parts of the skull, other than
those mentioned, to which powerful muscles, especially those concerned with
mastication, are attached, are liable to vary considerably, apparently in
response to the action of the muscles, during the life of the individual. The
Wry frre
A
Text-ric. 309.—A, Posterior end of nasals and maxille of ad. 3 of P. onca onca from Para
(45.8.25.23). B, the same of a second ad. & from Para (45.8.25.22). C, the same of ad. 2 from
Tocantins (36.5.26.2).
postorbital area, for example, may become narrower and the zygomatic arches,
mastoid area and occipital crest may continue to expand after the skull has
attained its fulllength. All such variations in Jaguars’ skulls are under suspicion
of being systematically valueless.
From an examination of the skulls of P. onca in the British Museum, it
appears to me that several of the described subspecies are untenable on the
evidence supplied for their validity, some being based on single specimens pre-
senting characters which may fall within the limits of individual variation. By
410 Noyiratrs ZooLOGICAE XLI. 1939.
adopting the view that such features are worth nominal distinction, the number
of named forms could be greatly increased on the material available for examina-
tion; and it is no exaggeration to say that I found myself unable to identify
subspecifically with certainty any of the specimens in the British Museum from
the characters furnished by the describers. The localities were the only guide.
DESCRIPTION OF THE RACES OF P. ONCA.
1. Panthera onca onca Linn. (text-figs. 306, 307, 309, 310 A).
Felis onca Linnaeus, Syst. Nat., ed. 10, 1, p. 42, 1758.
Felis onca onca, Nelson and Goldman, Journ. Mamm., 14, p. 224, 1933.
Felis onca var. major Fischer, Addend. Syn. Mamm., p. 366, 1830; Nelson and Goldman, op. cit.,
p. 231 (subsp. of onca).
? Felis onca mexianae Hagmann, Archiv. Russen.- und Gesellsch.-Biol. 5, p. 10, 1908; Nelson and
Goldman, op. cit., p. 233.
? Felis onca coxi Nelson and Goldman, op. cit., p. 225.
? Felis onca boliviensis Nelson and Goldman, op. cit., p. 229.
Felis onca ucayalae Nelson and Goldman, op cit., p. 230.
Felis onca madeirae Nelson and Goldman, op. cit., p. 233.4
Locality of the type of onca, Pernambuco (settled by Thomas, 1911); of
major, Surinam ; of mexianae, Mexiana Island; of coxi, Espiritu Santo; of
boliviensis, Santa Cruz, Bolivia ; of ucayalae, Sarayacu on the Rio Ucayali; of
madeirae, Auara Igarapé on the Rio Madeira.
Distribution (suggested) : From Pernambuco southwards to Espiritu Santo
and Rio Grande do Sul and westwards along the Amazon and its tributaries, the
Tocantins, Tapajos, Madeira, Ucayali and Rio Negro, thence northwards up the
Rio Branco to the Guianas and Venezuela.
No zoologist seems to have described a topotype of the typical race of P.
onca. I follow Nelson and Goldman in considering that the name may be
adopted in a subspecific sense for specimens from the Lower Amazon. They had
3 skulls, one from Parad and two from the Tocantins River Valley. One of the
latter, an adult 4, was selected for measurement, presumably on account of
its exceptional size, its condylobasal length of 253-5 mm. greatly exceeding that
of any skulls in the British Museum, including specimens from Para and
Tocantins.
Three adult skulls from Parad (R. Graham, 45.8.25.21-23) were briefly
discussed above to illustrate the unreliability for subspecific distinction of
certain features which have been used for that purpose (text-fig. 307). They
were not sexed by the collector ; but I have very little doubt that my determina-
tion of the two largest as $ and the smallest as 9 is correct, the last, apart from
its smaller size, having a very low sagittal crest. Their dimensions are given in
the table below and they call for no further comment at the moment, except the
remark that the smaller and older of the two ¢ skulls is about 27 mm., over
1 inch shorter than the $ from Tocantins, measured by Nelson and Goldman to
demonstrate the large size of typical onca. Unfortunately they gave no measure-
ments of their skulls from Para.
A 3 skull from Marajo Island, on loan from Col. H. W. Murray, differs
merely in minor details of no systematic importance from the skulls from Para.
1 The suggested synonymy of this race is based upon an examination of specimens referred to
typical onca, to major and to boliviensis ; and, in the case of the other forms, upon the inadequacy
of the evidence of their distinctness supplied by the descriptions.
Novirarrs Zoonoarcarn XLI. 1939. 411
It is a little longer than the larger of the two Para skulls, but has the zygomatic
width narrower, because it is a little younger, being only just adult.
A & skull from Taperinha, close to Santarem, collected by Hagmann and
kindly lent to me by Sir Frank Colyer, is a very close match in size and other
features of the skull from Marajo Island, but has the sagittal crest lower, the
bullae rather flatter and the mesopterygoid fossa a trifle wider, features of no
systematic significance. Santarem is at the mouth of the Tapajos River, whence
Hagmann in 1908 (see below) recorded a ¢ skull which he identified as typical
onca. It had a basilar
length of 225 mm., which, 310
judging from other
skulls, would indicate a
condylobasal length of
about 248 mm., a little
longer than the skull
from Taperinha, San-
tarem. Hollister also
measured a ¢ skull from
Santarem which had
the condylobasal length
shorter than in the
Taperinha skull but the
same as in my larger
skull from Para.
Of @ skulls from the
Amazons assigned to
typical onca I have seen
only two, the one from
Para, already mentioned,
which is like the ¢
skulls from that district Trxt-Fic. 310.—A side view of skull of ad. 9 P. onca onca
except for its smaller from Tocantins (36.5.26.3) with the forehead greatly inflated with
o air-cells. B, side view of ad. skull from Ecuador (34.9.10.64)
size, smaller bullae, low marked ¢ by the collector, but treated as 2 and provisionally
sagittal crest, relatively identified as P. onca peruviana. This shows the normal form of
wider postorbital area the skull in P. onca. (These two skulls are drawn approximately
to the same scale ; but the forehead in A is a little too flat behind
the postorbital processes.)
and slightly smaller teeth;
and one from Tocantins
(Col. H. W. Murray, 36.5.26.3) which is much smaller. It may be noted that it
came from the same locality as the huge & skull of onca measured by Nelson and
Goldman, but is about 57 mm., nearly 24 inches, shorter in condylobasal length.
It differs remarkably from all the other skulls of P. onca I have seen in the pro-
nounced inflation of the forehead between the nasals and parietals with air-cells
(text-fig. 310). This, with a scarcely developed sagittal crest, makes the skull
dome-shaped in profile view and steeply sloped from the brow to the occipital
erest which is comparatively poorly developed. This feature is, no doubt,
merely an individual peculiarity. A very similar frontal inflation occurs in an
adult g skull of P. o. palustris (see p. 417) from Chaco, Paraguay, figured by
Cabrera. In this case the character is clearly individual, since the rest of the
skulls from Chaco have normally shaped foreheads. This skull from Tocantins
412 Novirates ZooLoGIcAE XLI. 1939.
also differs from those from Para in the slight overlap of the maxillae by the
nasals. It amounts to about 2 mm., whereas in the Para skulls it is as much
as 9 mm. This also I regard as an individual peculiarity.
MEASUREMENTS OF SKULLS ASSIGNED TO TYPICAL ONCA FROM THE LOWER
AMAZON AND ITS TRIBUTARIES FROM PARA TO SANTAREM.
j
|
Locality and sex. rote, | Oped | Ayas Boa tak | Al | am | m.
Tocantins Valley (N. and G.) . ad. &| 296-3 | 2534 | 205 — 573 764 | 284 | —
Tapajos River (Hagmann) .ad. & — (248+ ) 180
Marajo Island (Murray in B.M. ) ad. &| 271 241 173 47 50 73 28 21
Taperinha, Santarem (Colyer) . ad. g | 2714 242 180 44 45 74 29 20
Santarem (Hollister) 6 .ad. g| 269 234 183 — — _ 264 | —
Para (B.M. no. 45.8.25.22) .ad. g) 265 234 179 49 49 74 28 21
a » 45.8.25.23) .ad. g| 258 227 178 50 49 73 27 —
( » ..45.8.25.21), „ad. 9 | 238 216 172 49 45 64 26 19
Tocantins (B.M.no. 36.5.26.3) ad. 9 | 222 197 150 51 43 63 25 19
The great individual variation in size between the skulls of supposedly
typical onca from the Amazons is graphically shown by the table of skull-measure-
ments. All that need be added is the evidence supplied that the average condylo-
basal length of the 3 skulls from Parad to Santarem, including the one from
Tapajos recorded by Hagmann, is, as nearly as may be, 240 mm. In the two 9
skulls from Para and Tocantins the average of the condylobase is about 206 mm.
These averages have, in my opinion, an important bearing on the status of
the described subspecies of P. onca cited in the synonymy.
The Jaguar from Mexiana Island, in the estuary of the Amazon just north
of Marajo Island, was described as Felis onca mexianae by Hagmann, who claimed
that it was smaller than specimens of typical onca from the Rio Tapajos and
Southern Brazil. Nelson and Goldman admitted the race on Hagmann’s
authority, quoting the measurements of the largest g and 2 he gave. The
basilar length of Hagmann’s three adult & skulls was respectively 210, 194 and
190 mm. The basilar length in my three 3 skulls from Marajo Island and Para
is 218, 216 and 207 mm. respectively, the last being smaller than Hagmann’s
biggest ; but the average of the Mexiana specimens is less. In Hagmann’s two
9 skulls the basilar length was 186 and 176 mm. In my two @ skulls from
Para and Tocantins that length is 200 and 179 mm. Here, again, there is an
overlap, but the average of the Mexiana skulls is less. The data are most
unsatisfactory ; but on the evidence it is impossible to turn down mexianae as
an invalid race.
Panthera onca madeirae, based by Nelson and Goldman on a single 9 speci-
men from Auara Igarapé on the Rio Madeira, was distinguished from P. 0. onca
by having the skull less arched in its upper outline and decidedly smaller. No
measurements of Q skulls of onca were cited; but the condylobasal length of
the skull of the type of madeirae was 203 mm., intermediate in length between
1 In this and other tables the total length is taken with callipers from the front edge of the
premaxilla to the tip of the occipital crest ; the condylobasal length from the same point in front
to the tip of the occipital condyles; the zygomatic width is the greatest spread of those arches ;
the postorbital and interorbital widths are the least widths of those areas; the maxillary width is
taken just above the insertion of the canines; under pm‘ and m, are included the lengths of the
crown of the upper and lower carnassial teeth respectively. Figures in brackets are estimated
measurements. The letters N. and G. stand for Nelson and Goldman.
Novirares ZooLoGIcAE XLI. 1939. 413
my two 2 skulls from Para and Tocantins respectively ; and considering the
great difference in the convexity of the dorsal profile in these two skulls, it does
not appear to me that madeirae can be admitted as a valid race on the evidence
supplied.
The race P. onca ucayalae Nelson and Goldman has for its type an adult &
from Sarayacu on the Rio Ucayale, a tributary of the Upper Amazon, there
known as the Marafion. Nelson and Goldman also had specimens from Moyo-
bamba and Iquitos. The race was distinguished from typical onca by certain
details in the skull, especially the narrower zygomata. Since all the skulls of onca
differ from each other in details and the zygomatic width is known to be an
entirely untrustworthy systematic character, this information is not very helpful.
From the measurements of the skull, moreover, the zygomatic width is almost
exactly the same as in the skull of onca from Marajo Island, its narrowness in this
case being due to the skull being only just adult, obviously younger than the skull
from Taperinha, near Santarem. In the matter of size, the skull of the type
of ucayalae is certainly shorter in its condylobasal length than the skull from
Tocantins chosen for measurement by Nelson and Goldman, but it is longer in
that dimension than any ¢ skull of typical onca I have seen, coming between
the skull from Taperinha, Santarem, and the one from the Tapajos River recorded
by Hagmann. It is about 5 mm. longer than the average of skulls of typical
onca ; but on the available evidence cannot be regarded as representing a distinct
subspecies.
P. o. ucayalae was also said to be like boliviensis but larger, with the rosettes
heavily outlined and black spots on the head, limbs and underparts larger than
usual. But since the distinctness of boliviensis from typical onca is also in
doubt and the size of the spots on the underside and elsewhere varies considerably
in two of my skins from Matto Grosso, clearly varying within subspecific limits,
the diagnosis is here also unsatisfactory.
Felis onca var. major Fisher was based on Hamilton Smith’s figure and
description of a ‘ great variety ’ of F.onca from Surinam, in Griffith’s An. King.
Mamm., 2, p. 455, 1827. It was admitted as a valid race by Nelson and Gold-
man, who stated that it is distinguished from typical onca by its smaller skull.
In support of this they gave the dimensions of a 3 skull from Surinam as 242 mm.,
and of a 9 skull from the Caura Valley, Venezuela, as 2144 mm.
It is quite true that the g skull is 11 mm. shorter in its condylobasal
length than the exceptionally large & skull of onca from Tocantins Nelson and
Goldman selected for measurement ; but it is practically the same as in my
two ¢ skulls from Marajo Island and Taperinha respectively, and considerably
_ larger than both the ¢ skulls from Para, its condylobasal length being almost
exactly the average of the six ¢ skulls recorded above which I assign to typical
onca. Similarly, the 9 skull from Venezuela with a condylobasal length of
2141 mm. is the merest trifle smaller than my @ skull of typical onca from Para
and larger than the 9 from Tocantins.
On the available evidence there do not appear to be any reasons for regarding
the Jaguars from Surinam and Venezuela as racially distinct from those from
the Amazons.
A young & skull in the British Museum from the Orinoco (Parzudaki,
no. 51.8.25.12), with the condylobasal length 214 mm., resembles the Amazonian
skulls in the size of the teeth but differs in characters due to immaturity and in
414 Novirarres ZooLoGIcAE XLI. 1939.
having the auditory bullae a little more swollen than the most swollen of the
Amazonians.
Nelson and Goldman also recorded major from Frechal, Rio Surumu, and
from Serra da Lua, Amazonas, both in Brazil. I am indebted to Sir Frank
Colyer for the loan of a ¢ skull from “ the forest of Boa Vista, Rio Branco,” a
tributary of the Rio Negro, which joins the Amazon at Manaos, where the skull
- was purchased. Boa Vista is high up, just south of the point where the tributaries
of the Branco rise in Venezuela and British Guiana. The skull is not quite full-
sized and would probably have reached about the length of my larger & skull
from Para. Its bullae are almost exactly the same size as in my smaller ¢ from
Para and the nasals are similarly broad posteriorly but only overlap the maxillae
by about 2 mm., instead of 9 mm. in the Para skull ; but the overlap is exactly
the same as in my @ skull of onca from Tocantins and is about half that of my
skull from the Orinoco, which also differs in having the bullae noticeably more
inflated.
Felis onca boliviensis Nelson and Goldman. For the original record of this
race its authors had 3 skins and 2 skulls, ¢ and 9, from Buena Vista, Santa
Cruz, Bolivia (Steinbach coll.). They did not compare it with onca, but with a
Jaguar from Matto Grosso they had previously described as milleri (see below,
p- 416) and with the little-known form named peruviana by Blainville (see
below, p. 417).
In the British Museum there are two adult skulls, 3, 9, also collected by
Steinbach at Buena Vista. The four known skulls attest considerable individual
variation in size. Inthe Sand 2 recorded by Nelson and Goldman, the condylo-
basal length was 2364 and 212 mm. respectively. In the $ and @ skulls in
the British Museum, the same dimension is 225 and 203 mm. respectively. The
two 3 skulls are respectively a trifle larger and a trifle smaller than the two adult
¢ skulls recorded above from Para which I assign to typical onca ; but the two
pairs from the different localities have the same average length of the condylobase.
Similarly, the two 9 skulls are intermediate in size between my two ® skulls
assigned to onca from Para and Tocantins ; but here again the average condylo-
basal length of the Bolivian and Amazonian pairs is as nearly as may be the
same. It may be noted, moreover, that my Bolivian 9 has exactly the same
condylobasal length as the 2 type of madeirae. The available data clearly leave
the status of boliviensis unsettled. They are unsatisfactory, at all events so far
as Bolivia is concerned. There is very little justification for the conclusion that
Sg skulls from this country are on the average smaller than those from the
Amazons. It seems as likely as not that future collecting there will bring to light
skulls as large as the larger known Amazonian skulls.
Felis onca coxi Nelson and Goldman, based upon an adult 2 from north of
the Rio Doce in Espiritu Santo, was also recorded from the north-eastern part of
Minas Geraes, near the Rio Doce. The skull was described as small with a low
sagittal and a well-developed lambdoid crest, much smaller than the skull of
onca with the frontal region more constricted behind the postorbital processes
and the bullae relatively broader and somewhat flatter posteriorly. It is not clear
how many skulls were available. Only the measurements of the type were given;
but no measurements of 9 skulls of onca were supplied to show the difference.
The skull is actually a trifle longer in its condylobase than my skull assigned to
typical onca from Tocantins, its zygomatic and interorbital widths are the same
NOVITATES Zoonoaicar XLI. 1939. 415
and in both the sagittal crest is low. Its lambdoid crest is said to be well deve-
loped, but since the difference between the total and condylobasal lengths is less
than in the Tocantins skull, that crest in the type of coxi is presumably a little
less well developed, a character of no systematic value. The only other differ-
ences the measurements show are a narrower muzzle and a slightly smaller pm‘.
The diagnosis is obviously quite insufficient to justify more than provisional
acceptance of cowi as a valid subspecies. It must be remembered, moreover,
that the distributional area of coxi is considerably closer to Pernambuco, the
type-locality of onca, than is Para. Hence, the Jaguars of Minas Geraes and
Espiritu Santo are at least as likely to be referable to typical onca as those from
Para and other places near the estuary of the Amazon.
The skull of an adult 9 from Taquara, Rio Grande do Sul (H. von Jhering,
B.M. no. 84.2.8.1), is a good deal larger than the type of coxi, but only slightly
shorter in condylobasal length than my @ skull assigned to typical onca from
Para. It differs from the latter chiefly in having the sagittal and occipital crests
a little better developed and the zygomata much narrower ; but the differences
between it and the Para skull are much less than the differences between the
© skulls from Para and Tocantins. Also it differs less from the type of coxi than
those two Amazonian skulls differ from each other. Geographically, perhaps, it
should belong to the next described race palustris ; but, with our present know-
ledge, it is too small for that determination. It must be left unidentified,
although if it had had no locality I should have assigned it to typical onca.
In the following table are included the skull measurements of the so-called
races regarded as distinct by Nelson and Goldman but entered above as synonyms
of typical onca. The only omission is mexianae Hagmann.
Post.
j
Total Cond. bas. | Zygom. Int. orb.| Max.
Rema, Wcecelltay emal Se Length. | Length. | Width. | yon, | Width. | width. | 27 | 7%
(major) |
Surinam (N. and G.) .ad. d| 264 242 182-3 — 544 | 74 28-7 | —
Boa Vista, Rio Branco (Colyer) |
subad. g| 254 225 170 46 51 72 29 20
Orinoco (B.M. no. 51.8.25.12).yge d | 237 214 159 43 43 67 29 20
Caura Valley, Venezuela (N. and |
G.) 2 : A .ad.Q | 239 214-3 163-3 — 47 655 | 27-7 | —
(ucayalae)
Sarayacu, Rio Ucayali (N. and
G.) ! 5 3 .ad.d| 275 | 2454 | 1738| — | 493 | 74 1299| —
(madeirae)
Auara ee Rio Madeira (N.
and G.) .ad.Q | 224 203 1504 — 44-7 | 62 25-7 | —
(boliviensis) |
Santa Cruz, Bolivia(N. and G.)ad. {| 266 2364 1754 — 47 68:2 | 284 | —
29 » (B.M. no.
26.1.12.1).ad. | 250 225 174 42 47 67 27 22
60 » (N. and G.)ad. 2 242 212 160 — 47 65:8 | 284 | —
” ” (B .M. no.
26.12.4.29)ad. 9? 224 203 155 41 39 57 25 184
(coxi) | |
Espiritu Santo (N. and G.) . ad. 9 219 1984 150 | 43 | 584 | 244+) —
(? race) |
Taquaraé, Rio Grande do Sul
(B.M. no. 84.2.8.1) . .ad.Q | 241 | 213 160 47 | 42 | 64 |27 | 19
416 NOVvITATES ZOOLOGICAE XLI. 1939.
Of the four ¢ skulls entered in this table, namely the one from Surinam,
the one from Ucayali River and the two from Bolivia, all, except the second
from Bolivia, which is 2 mm. shorter, fit into the series of seven skulls of typical
onca from the Lower Amazons in their condylobasal lengths ; and the average
of that dimension in the four is 237 mm., only 3 mm. less than the average in
the Lower Amazonian skulls. The average of the eleven skulls combined is
239 mm., as nearly as may be, 1 mm. less than in the series from the Lower
Amazon.
Similarly, the five 9 skulls from Venezuela, Rio Madeira, Bolivia and
Espiritu Santo fall in condylobasal length between the two @ skulls of typical
onca from Para and Tocantins ; and their average condylobasal length is practi-
cally the same, namely 206 mm.
2. Panthera onca palustris Amegh.
Felis palustris Ameghino, Rapidas diagnosis Mam. Fos., p. 6, 1888.
Panthera onca palustris Cabrera, Not. Prelim. Mus. La Plata, 2, pp. 9-39, 1934.
Felis paraguensis Hollister, Proc. U.S. Nat. Mus. 48, p. 170, 1915; Nelson and Goldman, Journ.
Mamm., 14, p. 227, 1933.
Felis notialis Hollister, Proc. U.S. Nat. Mus., 48, p. 170, 1915.
Felis ramsayi Miller, Journ. Mamm., 11, p. 14, 1930 (substitute for paraguensis).
Felis onca millert Nelson and Goldman, Journ. Mamm., 14, p. 226, 1933.
Felis onca paulensis Nelson and Goldman, Journ. Mamm., 14, p. 228, 1933.
Locality of the type of: palustris, Lujan (fossil form) ; of paraguensis,
Paraguay ; of notialis, San José, Entre Rios, N. Argentine ; of ramsayi, Para-
guay ; of milleri, Descalvados in Matto Grosso ; of paulensis, Sao Paulo.
Distribution (of existing forms) : Matto Grosso, Paraguay and the adjoining
parts of the Parana River Valley, Entre Rios, north-eastern Argentine.
This mainly more southern race intergrades with typical onca, but may be
distinguished by its larger size, the average condylobasal length of the ten adult
6 skulls entered in the table being about 254 mm. and of the four adult 2 skulls
about 231 mm., exceeding that dimension in onca by about 14 and 15 mm.
respectively.
From the synonymy it may be seen that Hollister thought two distinct
species, paraguensis and notialis, were indicated by the cranial differences. But
Nelson and Goldman detected that notialis represented the 9 of the ¢ previously
described as paraguensis. Cabrera independently came to the conclusion that
the two names had been applied by Hollister to the same form, although he left
the sex undetermined. Also it may be seen that Nelson and Goldman con-
sidered that three races, paraguensis, milleri and paulensis, were involved, the
three being distinguishable by cranial differences which appear to have no
systematic significance.
Cabrera, without being acquainted with the monograph by Nelson and
Goldman, studied the Jaguars of Southern Brazil and Paraguay and compared a
number of skulls of recent specimens with those of late Pleistocene skulls described
and named by Ameghino. He saw examples from Matto Grosso which Nelson
and Goldman had named milleri and came to the conclusion that they belonged
to the same race as Hollister’s paraguensis. For this name he substituted the
older name palustris given by Ameghino to a Pleistocene skull. Since the
numerous photographs and measurements he published of recent skulls from
Novirares Zoonocicarm XLI. 1939. 417
Chaco, Paraguay, and of subfossil skulls from localities near by bear out his view,
there seems to be no escape from the acceptance of palustris as the name for this
large southern Jaguar.
Cabrera’s measurements, reprinted in my table, show the impossibility of
distinguishing the Matto Grosso Jaguars from those found in Paraguay and also
the great range in size between adult 3 skulls from the same locality, Chaco in
Paraguay, amounting to 30 mm. in condylobasal length, which exceeds by a few
millimetres the range of variation of that dimension in typical onca.
No less clearly do his admirable photographs show the great differences
between adult ¢ skulls from Chaco in minor variations affecting the general
shape, proving that such variations have no systematic importance. They are
similar to those pointed out above in connection with skulls of typical onca from
Para and of the skulls from the Magdalena Valley, Colombia (P. o. centralis).
Of particular interest is one ¢ skull from Chaco which has the frontal region
greatly inflated with air-cells, apparently almost exactly as in the 9 skull of onca
from Tocantins, above described.
Cabrera did not sex his skulls from Chaco; but from his photographs it
may be inferred that they are ¢.
I have added to the synonymy of palustris the name paulensis based on a
single © skull, of which the characters appear to me to be systematically valueless.
SKULL MEASUREMENTS OF P. 0. PALUSTRIS, MOSTLY TAKEN FROM CABRERA’S
PAPER.
Locality and Sex. Total | Cond. bas. | Zygom. Host: Int. orb.| Max. ms om
zi Length. | Length. Width. Width. Width. | Width. | ?”- 22
Corumbä, Matto Grosso (milleri, s
N. and G.) .ad. d| 302 277 207 — 57 803 | 28-2 | —
(The same measured by Cabrera) 305 280 208 — — — |28 | —
Chaco, Paraguay (Cabrera) .ad. d| 304 272 213 — 61 88 29 —
Paraguay (paraguensis type, N.
and G.) . „ad. | 2954 | 264 195 — | Be | we |e | =
(The same measured by Hollister) 303 265 196 — — — |288| —
Corumba, Matto Grosso (Cabrera)
ad. & | 300 260 196 — — — 128 —
Chaco, Paraguay en . ad. & | 280 252 198 — 47-7 | 773 | 32 —
.ad. & | 285 250 197 — 54 80 29 —
Matto Grosso (B.M. no. 99. 8.4.1)ad. g| 276 245 181 42 49 73 28 21
Chaco, Paraguay (Cabrera) .ad. &| 275 242 192 — 614 803 | 29 —
Chaco Santiagueno (Cabrera) . ad. 3 | 274 242 1654 | — 50 74 30 —
Misiones (Cabrera) . 2 ad. &| 265 242 184 — 46 74 31 a
Descalvados, Matto Grosso (mil-
leri type, N. and G.) . ad. 2 | 267-8 238 1754 a 514 | 70:3 | 27-8) —
Sao Paulo (paulensis type, N.
and G.) . .ad.Q | 266-2 | 2354 | 1753 | — | 47-7 | 704 | 263 | —
San José, Entre Rios (notialis
type, N. and G.) & .ad. Q | 261 232-3 175% — 43-8 | 72-3 | 30:8 | —
Paraguay (Hollister) : .ad. 9 | 248 220 179
3. Panthera onca peruviana Blainv. (text-fig. 310, B).
Felis onca peruviana Blainville, Osteogr. Mamm., 2, pt. 12, p. 186, Atlas, pl. 8, 1843; Nelson and
Goldman, op. cit., p. 230.
This race, as stated by Nelson and Goldman, cannot at present be placed.
The name was printed on the plate, without diagnosis, under the figure of a skull
of a specimen known to haye been brought alive from Peru and exhibited in a
30
418 Novirates ZooLogicaE XLI. 1939.
menagerie. It was said to be a 9 skull; but on account of its size, its well-
developed sagittal crest and the constriction of the postorbital area, making it
narrower than the interorbital area, I believe it to be a ¢ skull, although the
slight prominence of the occipital crest resulting in a comparatively small differ-
ence between the total and condylobasal lengths (see below) is a more likely 9
character in adult skulls. Judging from the figures there are, however, certain
peculiarities about the skull, and I incline to the suggestion of Nelson and
Goldman that it may have been affected by the conditions that captivity imposes.
Evidence, although not proof, of this is supplied by the coronoid process of the
mandible, which is less robust and more backwardly inclined than in normal
skulls, its posterior edge at the summit projecting well beyond the condyle.t
The mastoid and paroccipital processes, as Nelson and Goldman detected, are
also abnormal, and this peculiarity may be attributable to the same factor,
although these processes are subject to a great deal of individual variation in
the skulls of wild killed Jaguars.
Except in one respect the figures, said to be half natural size, are in close
agreement. The discrepancy lies in the condylobasal length as represented in
the profile and ventral views. In the ventral view this dimension is 116 mm. ;
in the profile view itis 111mm. _ I believe the former figure to be incorrect because
when doubled it makes the condylobase 232 mm., only 12 mm. less than the total
length of the skull, a much smaller difference than in any adult Jaguar skull of
either sex that I have seen. The acceptance of the smaller figure, 111 mm.,
doubled, makes the difference between the two lengths 22 mm., exactly the
same as in the 9 skull of typical onca from Para. But this, as already said, is a
very small difference for a g skull. The measurements of this skull, taken from
the figures, show that it is a little smaller in all its important dimensions than
my smallest 3 skull of typical onca from Para and comes very close to the smaller
of the two ¢ skulls from Santa Cruz in Bolivia.
In the British Museum there is a skull of a young ¢ labelled “‘ Cerro Azul on
the Upper Ucayali, Peru” (R. W. Hendee, no. 28.5.2.139). Since that locality
is in the Peruvian Andes it must be close to the source of the Ucayali, but on the
eastern slope of the watershed. No doubt this skull represents peruviana. It
has the nasals rather exceptionally narrow between the frontals as in Blainville’s
figure, and considering its immaturity a tolerably well-developed sagittal crest,
about 6 mm. high, just behind the fronto-parietal suture ; the occipital crest is
poorly developed and the postorbital area is wider than the interorbital, both
characters attributable to youth ; the frontal is more inflated with air-cells than
usual ; the teeth are large but quite unworn ; the mastoid and paroccipital pro-
cesses are normal and the bullae are larger than depicted in Blainville’s specimen.
This skull shows no feature suggesting that, when full grown, it would have
differed from that of typical onca.
On the evidence the status of the Peruvian Jaguar must be left
unsettled.
A specimen in the British Museum from Santa Domingo de los Colorados,
2000 ft.,in Ecuador (B.M. no. 34.9.10.64) is of some interest. By its collector,
Söderström, it was marked 3. If correctly sexed it probably represents an
undescribed local race, because the skull, which is adult, is a good deal smaller
than the adult 3 skull of any other subspecies. But it has the characters
1 I have seen this modification in many skulls of Carnivora reared from cubhood in captivity.
Novirates ZooLocicaAE XLI. 1939. 419
of a 9, the sagittal crest being merely a low ridge, although the postorbital
area is narrower as compared with the interorbital than is usual in that sex
(Fig. 310, B).
The richly tinted skin is ornamented with bold rosettes, mostly with entire
rims enclosing one or two small spots.
I provisionally refer this specimen to peruviana on account of its locality
and its measurements are entered in the table under that name. Its bullae are
tolerably large for so small a skull, 32 by 25 mm.
4. Panthera onca centralis Mearns (text-figs. 308, A, B).
Felis centralis Mearns, Proc. Biol. Soc. Wash., 14, p. 138, 1901.
Felis onca centralis, Goldman, Smiths. Misc. Coll., 69, no. 5, p. 166, 1920; Nelson and Goldman,
Journ. Mamm., 14, p. 234, 1933.
Locality of the type : Talamanca in Costa Rica.
Distribution : From Colombia throush Panama to Costa Rica, Honduras,
Nicaragua and Salvador.
Distinguished by Mearns from hernandesii by its smaller size and more normal
pattern of rosettes; but according to Nelson and Goldman the rims of the
rosettes are considerably broken. These two authors state that the skull is
very similar to that of hernandesii but has the nasals less depressed, more highly
arched from the front. The three skulls that I have seen differ individually in
this particular.
Mearns gave the basilar length of two adult & skulls from Talamanca as
200 and 212 mm. respectively. The condylobase of these would be approxi-
mately 220 and 232 mm. Nelson and Goldman chose the former of these,
citing the condylobase as 2195 mm. They also recorded the race as passing
from Panama into Colombia as far south as Guadas, Cundinamarca.
There are four skulls in the British Museum which I identify as centralis.
An adult g, labelled “ Nouvelle Grenade, Central America, Dr. Bonnet ”’
(Lataste’s coll., B.M. no. 19.7.7.3689) is intermediate in its condylobasal
length between the two Talamanca skulls measured by Mearns; but it
apparently has bigger teeth and may belong to the race, veraecrucis, mentioned
below.
Two adult & skulls from the Magdalena Valley (no. 35.3.6.1-2) were
noticed above in some detail (p. 408). They differ as much from each other as
either differs from Lataste’s skull except that in both the occipital crest is more
prominent, the sagittal crest a little higher, and the nasals hardly overlapping
the maxillae, whereas in Lataste’s skull the overlap amounts to about 7 mm.
In the latter also the teeth are larger. But in a young adult, apparently ¢ skull
from Medellin Colombia (Salmon, B.M. no. 74.4.8.2), which differs in details
from both the other Colombian skulls, the nasal overlap is about 5 mm., and
pm’ is practically as long as in Lataste’s skull, although m, is smaller.
The average condylobasal length of the 5 adult & skulls entered in my
table is 223 mm., considerably less than in typical onca, although individuals
may overlap. But the 2 of centralis from Salvador is intermediate in size
between my two Q skulls of typical onca. From the little that has been recorded
of the skull of hernandesii, it appears that centralis has not a smaller skull despite
Mearns’ statement to that effect,
420 NOVITATES ZOoOLOGICAE XLI. 1939.
SKULL MEASUREMENTS OF JAGUARS FROM PERU, ECUADOR, COLOMBIA AND
SOUTHERN CENTRAL AMERICA.
cr . 2 nd. bas. | Zygom. Eos: ns Max.
anna ocaltyianllsen- resem, | | Width, | OF. | orb. | wiath, | amt | ™-
. | |
peruviana | |
Peru (Blainville) . o ad.? g| 244 Do 72 A 46 68 28 18
Cerro Azul, Peru (B.M. | |
28.5.2.139) . 6 .yg. &| 232 217 | 159 | 45 43 69 30 22
Ecuador, 2000 ft. (B.M. | |
34.9.10.64) . e .ad.Q | 224 202 | 156 | 40 41 | 65 27 20
|
| |
centralis |
Talamanca, Costa Rica (Mearns) ad. & — (2325) | 175 | 43 45 — | 264 | —
> » (N.andG.) ad. $&| 250 219% | 169% IN 414 | 63:8 | 254 | —
Central America (Lataste coll.). ad. &| 250 226 | 176 42 44 12 29 21
Magdalena Valley (B.M. no. | |
35.3.6.1) 5 3 .ad. g| 249 219 | 175 | 47 48 70 264 | 20
Magdalena Valley (B.M. no. | |
35.3.6.2) ° 5 .ad. d | 249 218 | 168 47 + 65 254 | 19
Medellin (B.M. no. 74.4.8.2) yg.ad. d& | 232 214 | 161 | 45 44 63 29 19
Salvador (N. and G.) : .ad.?Q | 2314 | 207-7 | 156-2 ie 43-8 | 634 | 25-9) 20
Of the following races described from Central America and the Southern
United States there is very little material in the British Museum. Since all that
is available is the skin of the type of hernandesii, the skull having been mislaid,
and a couple of skulls and a skin provisionally identified as goldmani, the informa-
tion about them that follows has been taken from the papers by Mearns and
Nelson and Goldman.
5. Panthera onca hernandesii Gray.
Leopardus hernandesii Gray, Proc. Zool. Soc., 1857, p. 278, pl. 58.
Felis hernandesii, Mearns, Proc. Biol. Soc., Wash. 14, p. 141, 1901.
Felis onca hernandesii, Goldman, Proc. Biol. Soc. Wash., 45, p. 144, 1982; Nelson and Goldman,
Journ. Mamm., 14, p. 235, 1933.
Locality of the type : Mazatlan in Sinaloa, Mexico.
Distribution : Mexico west of the Central Plateau from Sinaloa to Tehuantepec,
additional localities mentioned being Colima, Guerrero, Nayarit and Oaxaca.
Gray distinguished this Jaguar from typical onca on the evidence of a skin
from Mazatlan distinguished from the latter by its disruptive pattern, the edges
of the rosettes being broken up into spots so that they are at most indistinctly
traceable. According to Mearns, it is a comparatively dry zone form, with
pallid ochraceous buff ground colour.
Very few skulls showing the range in size have been recorded, the largest
apparently being that of an adult ¢ from Tehuantepec stated by Mearns to
have a basilar length of 217 mm., whence the condylobasal length may be esti-
mated to be about 237 mm. This skull was referred by Mearns to his race
goldmani ; but I infer from Nelson and -Goldman’s paper that it represents
hernandesii, although they did not mention it (see below under goldmani, p. 421).
6. Panthera onca arizonensis Goldman. \
Felis onca arizonensis Goldman, Proc. Biol. Soc. Wash., 45, p. 144, 1932; Nelson and Goldman,
Journ. Mamm., 14, p. 237, 1933.
Locality of the type : Cibecue in Arizona.
Distribution : Arizona, Sonora and New Mexico.
Novitarrs ZooLoGIcAR XLI. 1939. 421
Said to be closely allied to hernandesii, with the general colour and pattern
much the same, but distinguished by being larger and having a more massive
skull with a broader rostrum, flatter nasals, wider anterior nares and narrower
mesopterygoid fossa, the flatness of the nasals distinguishing it from all other
races of the species.
Although claimed to be larger than hernandesii, the skulls of the adult g and
2 of arizonensis are respectively about the same size as those of hernandesii
entered in my table. Whether the differences in details of the skulls have any
systematic value may remain an open question.
7. Panthera onca veraecrucis Nelson and Goldman.
Felis hernandesii goldmani Mearns, Proc. Biol. Soc. Wash., 14, p. 142, 1901 (in part, skulls but not
skin).
Felis = veraecrucis Nelson and Goldman, Journ. Mamm., 14, p. 236, 1933.
Locality of the type: San Andres, Tuxtla, Vera Cruz.
Distribution : Central America east of the Mexican plateau from Tabasco
through Vera Cruz and Tamaulipas to Central Texas.
This race, based upon an adult & skull assigned by Mearns to goldmani, was
described as the largest of the North American Jaguars, but similarly coloured,
with rather large rosettes. The skull was said to be like that of hernandesii, but
decidedly larger and more massive, with the anterior nares higher, nasals more
arched and canines larger. The skull of the type has the condylobasal length
only about 10 mm. longer than in the skull of hernandesit from Tehuantepec,
whereas the ¢ skull of veraecrucis from Palenque, Chiapa, is smaller than that skull
of hernandesii. The measurements also show that the claim that the skull of
veraecrucis is bigger than that of arizonensis conveys very little, seeing that the
length of the condylobase of the type skull of arizonensis is almost exactly the
same as the average of the two & skulls of veraecrucis.
The published evidence that these three races differ in the size of their skulls
amounts to nothing. The claimed detailed differences may be left for what
they are worth; but there is evidence that those races are larger than goldmani.
8. Panthera onca goldmani Mearns (text-fig. 308, C).
Felis hernandesii goldmani Mearns, Proc. Biol. Soc. Wash., 14, p. 142, 1901 (in part, skin not skulls),
Felis onca goldmani Goldman, Proc. Biol. Soc. Wash., 45, p. 144, 1932; Nelson and Goldman
Journ. Mamm., 14, p. 235, 1933.
Locality of the type: Yohattan in Campeche, Mexico.
Distribution : Campeche and Guatemala in Mexico.
Mearns distinguished this race from hernandesit by its more intense coloration
and bolder pattern, responding to a more humid environment. His statement
that the skull is decidedly larger than in centralis was, judging from Nelson
and Goldman’s paper, based upon his erroneous association of one skull of
hernandesii and two of veraecrucis with the type skin of goldmani. According to
Nelson and Goldman, the race is most nearly allied to centralis, with the colour
and pattern much the same, but has the skull smaller and less angular, with the
sagittal and lambdoid crests less developed. Of the two skulls they measured,
the ¢ from Guatemala is almost exactly the same size as my smaller & of centralis
422 Novirates ZOOLOGICAE XLI. 1939.
from the Magdalena Valley; but on the published evidence the race may be
provisionally admitted.
In the table of measurements I have provisionally entered under this race
a couple of skulls in the British Museum. One is that of a subadult & labelled
“ Mexico, J.B.” (no. 39.537). It is a small skull with a well-developed occipital
crest, but has a great peculiarity in the extreme narrowness of the mesopterygoid
fossa which is only 15 mm. wide from edge to edge, from 5 to 6 or so mm. less
than in the narrowest of other skulls (Text-fig. 308, C). The other skull, that
of an old 9, clearly menagerie reared, but without locality, comes into goldmani
by size, and is associated with a skin in which the margins of practically all the
rosettes on the body are broken up into from 3 to 6 larger spots.
SKULL MEASUREMENTS OF THE RACES OF P. ONCA, EXCLUDING CENTRALIS,
DESCRIBED FROM CENTRAL AMERICA AND FROM THE SOUTHERN UNITED
STATES.
= : S Total | Cond. bas. | Zygom. TUS Tnt. orb.| Max. ‘
Das SEEM Oe Se Length. | Length. | Width. | yon; | Width. | width. | PR | ™-
hernandesii
Tehuantepec . : ö . ad. & u (237+) | 188 47 51 25 —
Agua Brava, Sinaloa (N. and G.) ad. 3 — — 1774 — 534 72 26-8 | —
Colima (N. and G.) 5 . ad. 9 | 218-8 | 1994 154-8 — 44-8 | 62-6 |25-8 | —
San Blas (Mearns) . a . ad. 2 — a 159 50 46 — | 24 —
arizonensis :
Cibecue, Arizona (N. and G.) .ad. 3} 273 2374 187 — 51 744 | 28 —
Greaterville, Arizona (N.and G.) ad. 2 | 2174 1924 155:3 — 44-3 | 644 | 254) —
veraecrucis
Tuxtla, Vera Cruz (N. and G.) . ad. g| 279 2474 180 — 50 72:3 | 28 —
Palenque (Mearns) . 6 .ad. J — (230+) | 178 44 49 — |27 —
goldmani
Peten, Guatemala . B .ad. &| 237 217-8 1664 — 44 64-9 | 234 | —
Mexico (B.M. no. 39.537) yg.ad. & | 237 209 156 44 44 65 26 20
Campeche (N.and G.) . .ad. 9 | 204 187-8 143-8 — 38-8 | 55 24-4 | —
? loc. (B.M. no. 45.5.23.5) .oldQ | 207 186 140 42 36 61 24 18
Novitarrs Zoonoarcan XLI. 1939. 423
DESCRIPTIONS OF ANTHRIBIDAE, MOSTLY FROM FIJI.
By KARL JORDAN.
(With 1 text-figure.)
1. Tribotropis limodes sp. nov.
&. Very near to T. conicollis Jekel 1855; more robust, the green colour more
or less replaced by olive. Median carina of frons and rostrum rather more
prominent ; between dorso-lateral carina and raised margin of antennal groove
a shallow channel continued backwards to near basal third. Antenna short, not
reaching to middle of elytra, segment II one-third shorter than III, this one-
tenth longer than IV, IV = V, VI and VII a little longer than V, VIII and tip
of VII white, VIII a trifle shorter than VI, IX = VII, X very little longer than
II, XI nearly twice X, longer than III, in ¢ of T. conicollis VIII longer than IX.
Median swelling of pronotum less high than in 7. conicollis, being a little
lower than the lateral swellings, not higher. Elytra as in 7. conicollis.
Intercoxal process of mesosternum apically slightly narrower. Abdominal
segments distinctly flattened-impressed in middle, the last external segment
bearing a semicircular flattened central area; apical margin of this segment
without distinct indication of lateral angles. Tarsi broader and shorter, foretarsal
segment I 17: 7-5, II 9:10 (length measured to middle of apical margin, breadth
across the same.point), in 7’. conicollis I 24:6, IL 10: 7-5; hindtarsus in 7’.
limodes 1 17: 6-5, IL 9:7; in T. conicollis I 24: 6, II 12: 6-5.
Length (head excl.) 15 mm., breadth 5 mm.
Upper Amazon, 1 ¢ received from Messrs. Staudinger & Bang-Haas.
2. Tribotropis apollinaris sp. nov.
®. In colouring halfway between 7. compressicornis Jord. 1895 and T.
conicollis, nearest to the latter, but the pronotal carina continued forward to
middle of side. Rostrum broadly and deeply depressed in middle from base to
apex, with the median carina higher than in the preceding new species, especially
on frons, where the carina bears a shallow longitudinal groove; margin of
antennal groove cariniform, high, the channel between it and dorso-lateral
carina of rostrum long, extending farther apicad than the antennal groove and
fading away on the widened apex of rostrum. Antenna compressed (III to
VII), segment VIII woolly, white, extreme base of IX also white, III the longest.
Measurements: II 8, III 23, IV 18, V 17, VI 17, VII 18, VIII 21, IX 14, X 9,
XI 21.
Prothorax less regularly conical than in 7’. conicollis and less convex, apex two-
thirds as broad as middle, which is the broadest point, disc with the usual two
impressions, median swelling a little higher than sides, from near apical margin
to carina a thin median raised line ; dorsal carina nearly straight in middle, with
a narrow median interruption, gradually flexed forward at side to meral suture
and then continued a short distance by a slight swelling.
Elytra flattened at suture, interspace IV convex, but not sharply cariniform,
interspaces I, VII, and IX tessellated with brown, II and IV with fewer but
424 Novirates ZooLoGIcAE XLI. 1939.
larger brown spots ; subapical swelling almost evenly convex, not tuberculiform.
Pygidium flat, with the apex rounded-truncate.
Transverse grooves of pro- and metasterna deep; intercoxal process of
mesosternum flat; metasternum swollen, more strongly convex than in 7’.
conicollis. Tarsi long, measurements: hindtibia 70, hindtarsal I 29, II 11,
IV 20.
Colombia : Bogota, received from Hermafio Apolinar-Maria, in whose
honour the species is named ; type at Tring, paratype in the collection of the
donor.
3. Mecocerus dolosus Jord. 1914.
In the specimens of a series from the Ogové R., Gaboon, the upperside of
segment II of the tarsi is black, the grey pubescens conspicuous in type and
paratype being absent or barely indicated. The division of the African Mecoceri
with median channel on frons into two groups according to the colouring of the
pubescence of the upperside of tarsal segment II can evidently not be relied on.
A grouping according to the width of the frontal channel ($9) and the size of
the velvety metasternal groove (3) is more natural.
4. Rhinotropis evansi sp. nov.
The Anthribid described by me as Notiana buxtoni in 1928 from Samoa is
near to or the same as Rhinotropis cristiferus Fairm. 1881 from Fiji. I was
misled by Fairmaire placing the species near Phloeops Lacord. 1866 (= Phlaeops,
Fairm., Ann. Soc. Ent. France, 1881, p. 467). The genus is nearest to Sintor
Schénh. 1839. The name cristiferus does not seem to have been amended by
anybody, and we may look upon it, leniently, as meaning figuratively a “ wild
coxcomb.” Dr. Evans has discovered a second species in Fiji which I have
much pleasure in dedicating to him. Notiana Jord. 1928 is a synonym of
Rhinotropis Fairm. 1881.
It resembles Phloeops platypennis Montr. 1857 even more than does Rh.
cristuferus. Underside of rostrum with median carina, and antennal scrobe ovate.
On upperside of rostrum a strong carina each side beginning at upper margin of
eye and disappearing above antennal groove ; between the two carinae the surface
deeply depressed, the depression divided into two broad channels running along
the carinae, curving sidewards and not quite reaching apical margin ; the apical
area between the two channels triangular, the apex of the triangle bearing a low
interrupted carina which extends backwards to near base of rostrum and reappears
on frons, where it is double ; apical margin with median sinus; on occiput a
very shallow depression each side, the two meeting on frons; pubescence of
head and rostrum clayish hazel mixed with grey, occiput blackish, the two
shallow depressions variegated with grey; from antennal groove backwards
three short lateral carinae, the uppermost visible from above. Antenna entirely
pale rufous, similar to that of Rh. cristiferus (cf. description of N. buxtoni Jord.
1928), segment II shorter than III (Fairmaire says II nearly as long as III).
Pronotum shorter and broader than in Rh. cristiferus, sides more strongly
rounded posteriorly, one-third broader than long, obscurely clayish hazel, mixed
with grey, centre blackish for the greater part, at each side of middle a fairly
deep impression which does not extend to apical margin, the two impressions
united posteriorly ; dorsal carina broadly concave, close to base in middle,
Noviratrs Zoonoaican XLI. 1939. 425
slightly convex laterally and then flexed forward in a round arc, lateral carina
more curved than in Rh. cristiferus. Scutellum greyish white, a little broader
than long, rounded.
Elytra strongly flattened together, punctate-striate, basal margin slightly
incurved from shoulder to shoulder, subbasal callosity raised as a large tubercle
bounded posteriorly by the antemedian depression, which is laterally continued
around the tubercle to basal margin, before apical declivity a second tubercle,
and in VII another, small, slightly in advance of the large subapical one ; pubes-
cence a mixture of black and dark hazel, with very little grey, depressed area
somewhat paler than sides, particularly behind subbasal tubercle, sutural inter-
space tessellated with black, as are more or less feebly also the alternate inter-
stices, apex hazel shaded with grey except a dark transverse line. Pygidium
dark hazel mixed with grey, as long as broad, with round apex.
Underside dark hazel and black, a lateral stripe on prosternite, the mesepime-
rum and a lateral stripe on abdomen cream colour; median area of thorax and
abdomen more or less covered with creamy pubescence ; anal sternum truncate,
in middle somewhat longer than IV, the lateral angles somewhat projecting ;
intercoxal process of mesosternum broader than in Rh. cristiferus, as broad as
coxa, apically rounded (Fairmaire calls the process subquadrate). Femora
variegated with grey, tibiae barred with grey and brown on upperside, apical
two-fifths black, as are base and apex of tarsal segment I.
Length 7 mm., breadth 2-3 mm.
Fiji: Taveuni, Oilai, x.24 (Dr. H. S. Evans), 3 3.
In Fairmaire’s description of Rh. cristiferus there is an evident slip on
p- 468, line 3 from above: “impressionis latioribus obtuse elevatis”’ has no
sense ; it should read “ lateribus.” The 2 or 3 tubercles at the apex of the
elytra said to be present in between the compressed elevations in R. cristiferus
are absent from buxtoni and evansi.
Dinosaphis gen. nov.
Q. Closely related to Rhinotropis Fairm. 1881; as in that genus the labium
entire, the dorsal carina of pronotum medianly close to base and with the lateral
angle broadly rounded, and the elytra tuberculate. But the rostrum broader
than long, flattened, without distinct carinae above and below, little narrower
at base than at apex. Antennal groove foveiform. Genotype: sp. nov.
here described.
4. Dinosaphis vitiana sp. nov.
@. A small species. Black, partly rufescent, pubescence black, hazel, and
grey, the only conspicuous markings being two white spots close together in
interspace III of elytrum.
Rostrum one-third broader than long, very little dilated at apex, flattened-
depressed, middle of apical area slightly convex, in basal depression a vestigial
carina, the whole surface rugate-reticulate, as is the head ; in front of eye the
ridges indicate a dorsal and lateral carina ; apical margin with shallow median
sinus ; no median carina on underside.
Frons five-eighths the width of apex of rostrum. Eye elliptical ; interspace
between eye and antennal groove as broad as segment II of antenna is long,
being very much narrower than in Rhinotropis. Antenna rufous, segments IX
426 Novirates Zootocicar XLI. 1939.
and X brown, XI pale, length of III to XI: 12, 10, 9, 8, 7, 6, 12, 6, 8, XI a little
broader than long.
Pronotum one-half broader than long, with central tubercle, at the side of
which a slight depression, sidewards of this a slight swelling ; a very thin buff
median line enlarged into a short streak at frontal side of tubercle, pubescence in
submedian depression grey, at posterior side of dorso-lateral swelling a grey dot,
at anterior margin behind eye a larger grey spot, basal and apical areas more or
less buff; dorsal carina shallowly concave, practically basal, somewhat convex
towards side, lateral angle rounded, lateral carina oblique, undulate, the side of
pronotum rounded-dilated behind middle and between this point and basal angle
incurved, the lateral carina reaching to middle and continued to near apex by
a longitudinal swelling ; apex rather abruptly constricted.
Elytra black, almost rectangular, one-third longer than broad, truncate at
base, rounded-subtruncate at apex (in dorsal aspect), rather coarsely punctate-
striate, interstices somewhat convex, subbasal swelling raised as a large tubercle,
which is longer than broad and crowned with brownish black hair, in interstice
III a small postmedian tubercle ; pubescence dark grey, suture and alternate
interstices dotted with black, in front of postmedian tubercle two white spots
one behind the other, shoulder angle and a transverse apical marginal band
hazel. Pygidium broader than long, rounded, hazel.
Underside sparsely pubescent grey ; sides of thorax punctate. Tibiae with
a narrow grey subbasal ring and a much broader median one, apex also grey,
the derm of these rings rufescent ; tarsi black, segment I rufescent and grey in
middle, IV more rufescent than black, I somewhat shorter than II to IV together.
Length 3-8 mm, width 2-1 mm.
Fiji: Taveuni, Waiyevo, ii.24 (Dr. H. S. Evans), 1 9.
Proscopus Jord. 1924.
This genus was based on a single species from Fiji, P. veitchi Jord. 1924.
I have now seen several other species which differ from P. veitchi in the size of
the eye and the structure of the rostrum. Instead of erecting for these species
another genus, which on receipt of further material might have to be sunk, it is
advisable to widen the diagnosis of Proscopus so as to embrace all the species of
this group with the proboscis short, vertical, and the lateral carina of pronotum
reaching to or to near apex.
5. Proscopus rudicollis sp. nov.
9. A distinctive feature of this species is the surface sculpture of the prono-
tum, the whole surface being densely covered with large deep pits the edges of
which are raised and form a net.
Pale rufescent buff. Rostrum rugate-reticulate, two-fifths broader at apex
than long, one-fifth narrower in middle than at apex, median apical area convex,
bearing a rather irregular carina which extends to base, apical edge centrally
forming an obtuse angle, base of rostrum tranversely depressed, the lateral
carina, which is so sharply marked in P. veitchi, is here vestigial. Frons a little
narrower between eyes than half the rostrum at its narrowest point, reticulate-
foveate, a swelling extending from tubercle of antennal groove to behind basal
depression of rostrum. Eye different from that of P. veitchi, its lower lobe
Noviratres Zoonoatcan XLI. 1939. 427
narrower than upper lobe, which extends well above tubercle of anntenal groove
and a short distance forward, the eye being about semicircular. Segment I of
antenna thin at base, widest beyond middle and slightly narrowing to apex, II
claviform, length of segments I 16, II 10, III 23, IV 25, V 25, VI 20, VII 21,
VIII 19, IX 16, X 13, XI 16, the last very pale.
Pronotum one-fourth broader than long, strongly rounded at sides, coarsely
reticulate-foveate, the margins of the pits raised into granules, before centre of
dise an indication of a transverse depression ; lateral carina not quite reaching
apical margin, situated in a blackish longitudinal smear. Scutellum transverse.
Elytra nearly three-fourths longer than broad, dorsally somewhat flattened,
subbasal swelling vestigial, behind it a slight transverse depression ; pubescence
scattered as on pronotum, greyish white, not concealing the derm ; strongly
punctate-striate, interstices convex, subcarinate, III and V posteriorly united
and continued as a single ridge ; lateral margin from below shoulder to middle
blackish, this colour occupying in middle the margin and interstices IX and X.
Pygidium broader than long, rounded, somewhat convex, with large punctures
at base, rest smooth.
Underside with large punctures. Segments II and III of tarsi dilated,
being especially broad in foretarsus.
Length 3-7 mm., width 1-5 mm.
Fiji: Suva, iv.1924 (Dr. H. S. Evans), 1 9.
6. Proscopus gentilis sp. nov.
Q. Like P. rudicollis, but rostrum broader, not appreciably narrowed in
middle, with a distinct dorso-lateral carina, the lateral surface below this carina
broader and more impressed, and the median sinus of apical margin rounded.
Eyes rounded, space between them at narrowest point two-thirds as wide as
middle of rostrum, lower lobe much broader than upper, which does not extend
above tubercle of antennal groove. Segment I of antenna rather more suddenly
widened beyond middle and more distinctly narrowed to apex, II less claviform
than in P. rudicollis ; length of segments: I 17, II 11, III 25, IV and V 26,
VI 21, VIII 17, IX 15, X 10, XI 13, the last slightly paler than X.
Pronotum less coarsely sculptured, the pits about as numerous, but their
margins not raised, not granulate; side without black smear, which is also
absent from side of elytrum. Interspaces of elytrum less cariniform. Basal
median groove of pygidium continued a short distance down.
Length 3-8 mm., width 1-7 mm.
Fiji: Rewa, Naduruloulou, i.1924, at light (Dr. H. S. Evans), 1 9.
Plesiobasis gen. nov.
$2. Near Basitropis Jekel 1855, but the lateral carina of pronotum extending
only to or to near middle. Rostrum short, slightly narrowed to base. Club of
antenna consisting in both sexes of three segments, IX triangular, very little
longer than broad, X shorter than IX, XI somewhat broader than IX, ovate
or nearly circular. Labium entire, apart from a small shallow median sinus.
Antecoxal portion of prosternite somewhat shorter than the coxa is broad.
Genotype: P. monera sp. nov.
The specimens resemble in appearance exceptionally small individuals of
428 Novirates ZOOLOGICAE XLI. 1939.
Basitropis and are so similar to each other in facies that they might be taken for
specimens of one variable species. According to peculiar secondary $-characters,
however, they fall into three groups, each group consisting of one or two (possibly
more) species. In one category of 3 the hindleg is simple, in a second it bears
at the end of the tibia a process, and in the third the process is found on the
hindfemur. Such distinctions of species which otherwise agree very closely are
of great interest. The shifting of a secondary j-character from one part of the
insect to another part is known in several genera of Anthribidae, for instance in
Phloeotragus such an organ occurs in some species on the metasternum, in others
on the hindcoxa.
In all species of Plesiobasis before me the rostrum is almost exactly twice as
broad as long, conspicuously rugulose, the ridges more or less longitudinal at
sides, no carina, but
one or two of the
irregular ridges in
middle of base more
prominent (as also on
frons) ; apical margin
with shallow median
sinus. Frons from
one-half to less than
one-third as broad as
rostrum, rugulose.
Eye elliptical, with
very small sinus to-
wards antennal
groove; interspace be-
tween eye and this
groove very narrow ;
the groove triangular,
ventral, not visible
fromabove. Antennal
segment I (in ventral
aspect) longer and
thinner than II, this shorter and much thicker than III, IV nearly like III,
III to VIII together about one-half longer than foretibia. Pronotum very finely
coriaceous, almost smooth, with traces of puncturation laterally at some distance
from lateral and dorsal carinae ; lateral angle of carina about 90°. Elytra
subeylindrical, strongly punctate-striate, interstices slightly convex, more
strongly at sides than dorsally ; the stripes quite distinct on apical declivity.
Pygidium smooth, evenly rounded.
Underside smooth, i.e. very minutely coriaceous ; on prosternite some large
shallow punctures laterally in front of meral suture and close to hindmargin in
lateral basal depression, others in the grooves behind meso- and metacoxae, on
metepisternum and on side of metasternum. Intercoxal process of mesosternite
truncate, broader than long. Abdomen medianly short in 3, (external) last
segment not wider at sides than in middle and here as narrow as the preceding
one, the pygidium therefore inclining frontad ; in 9 the last segment medianly
broader than the preceding one. Midtibia of 3 with apical tooth on inner side.
Novirarrs ZooLoGIcAE XLI. 1939. 429
Metasternum and abdomen flattened in ¢ medianly and here covered with a
silky pubescence.
Derm for the greater part testaceous, more or less pale, with blackish brown
areas on upperside. Markings evidently individually variable. All specimens
about twice as long as broad.
Four species are here described, based on differences in the males. I am
not certain in every case which females belong to these males and therefore do
not mention them. The receipt of further material must be awaited to settle
this doubtful point.
7. Plesiobasis monera sp. nov. (text-fig. 311, A).
SP. Hindleg simple. Upperside whitish grey ; on pronotum four well-
defined elongate brown patches, which reach neither apical margin nor carina,
all narrowed at both ends, the two dorsal ones rounded-dilated in middle and
here merged together, usually each connected with apical margin by a diffuse
brown line, lateral patch with or without grey spot near its middle. On elytra
a square brown patch on suture before middle, its anterior angles connected with
an irregular brown ring on subbasal swelling, on apical declivity an irregular
transverse brown band, which is connected in type with sutural patch by a brown
sutural line. Middle of femora, extreme tips of tibiae and tarsal segments IT
to IV brown.
Frons of ¢ less than one-third as broad as rostrum (6 or 7 : 22 or 23), rather
coarsely rugulate. In both sexes IX of antenna only a trifle longer than broad,
X broader than long. Lateral area of pronotum with the vestigial puncturation
fairly distinct.
Length 2-2 to 2-4 mm., width 1-1 to 1-2 mm.
Fiji: Taveuni, Waiyevo, ii, iv, and x.1924 (Dr. H. S. Evans), 3 3g, 1 .
8. Plesiobasis phelos sp. nov. (text-fig. 311, B).
dg. Hindleg simple. Upperside grey; the brown patches of pronotum
extending to base, the two middle ones continued to apex, their anterior portions
narrower than posterior, the antemedian connection between them incomplete,
rudimentary, these two stripes enclosing between them two grey patches one
behind the other, the anterior one elliptical, the posterior one nearly straight at
sides, pointed in front and truncate behind ; lateral brown patch a laterally open
ring with a forward projection and a connection with the brown submedian
stripe, the grey lateral area being divided into four patches. On elytra an
elliptical spot on subbasal swelling, the shoulder angle, a broad transverse ante-
median band from margin to a little beyond punctate line II and extending
along margin to shoulder brown, an inconspicuous line on suture before middle
pale brown, apical area marmorated with very pale rufescent irregular lines
(probably dark brown in more mature specimens), two of these lines longi-
tudinal on each elytrum and joining the transverse band. Underside and legs
very pale.
Frons one-third as wide as rostrum. Club of antenna longer than in the other
species before me, IX, X and XI longer than broad. Lateral carina of pronotum
curved, its dorsal side being convex and the angle of the carina therefore a little
430 NOVITATES ZooLoGICAE XLI. 1939.
smaller than in other species, the lateral carina anteriorly pale and not well
developed, not reaching to meral suture.
Length 2-2 mm., width 1-1 mm.
Fiji: Natova, vii.1918 (R. Veitch), 1 3.
9. Plesiobasis grallina sp. nov. (text-fig. 311, C).
$. Hindtibia with apical process. On pronotum a large brown median
area from apical margin close to carina, occupying a little less than half the
surface, divided by a narrow grey median stripe which widens in middle into a
transverse patch ; no brown lateral patch, but a stripe curved posteriorly as a
halfring bare of grey pubescence and perhaps due to abrasion. On elytra a
square patch on subbasal swelling, and before apical declivity a transverse band
from line VIII across suture to VIII brown, suture from this band to middle and
interstice IV from band to base rufous brown, before middle a brown spot in
III joined to line in IV, interstices V to VIII irregularly and indefinitely spotted
with rufous brown. Legs entirely pale, only segments II to IV of tarsi slightly
brownish.
Frons a very little over two-fifths the width of proboscis. Antenna as
in P. monera, X being broader than long. Process of hindtarsus less than
half the length of tibia, its apical two-thirds flattened and glossy on side
towards tarsus, the underside convex, being in lateral aspect incurved from
near base to near apex on side towards body, apex slightly curved towards
tarsus.
Length 2-3 mm., width 1-1 mm.
Fiji: Taveuni, Waiyevo, x.1924 (Dr. H. 8. Evans), 1 £.
10. Plesiobasis charax sp. nov. (text-fig. 311, D).
3. Hindfemur with long process projecting downwards. Pronotum with
four brown stripes, the two upper ones united anteriorly and continued on to
head as a triangular patch, which ends in a point on frons, lateral stripe narrow,
extending from base to apex, narrowing apically, being here narrower than the
grey stripe above it, whereas posteriorly it is broader than it. The two dorsal
stripes continued on to elytra, where they are somewhat narrower than on
pronotum ; they are first straight, then slightly oblique and beyond middle
curve round to outer margin, extending here forward to base, from the curved
portion of this large irregular halfring a stripe of the same colour runs backwards,
occupying interstices VII and VIII, and joins a transverse brown band which is
placed at the beginning of the apical declivity and is narrow at the suture and
lateral margin. Femora brown except base and apex, tarsal segments II to IV
very faintly brown.
Frons and antenna as in P. grallina. The process of hindfemur emanates
near apex from the inner side, lying against the abdomen ; it is slightly inclined
towards base of femur, and its extreme apex is curved towards the tibia, measured
on side towards body the process about half the length of the tibia (in this
specimen).
Length as in P. grallina.
Fiji: Lantoka, v.1919 (W. Greenwood), 1 4.
Novirarrs Zoonoagrcan XLI. 1939. 431
11. Stenorhis psomus sp. nov.
9. Much shorter than St. ampedus Jord. 1928 and lateral carina of pronotum
nearly extending to middle.
Cylindrical, twice as long as broad, derm dark brown and partially pale
rufous. Frons narrower than the eye is broad transversely. Antenna pale
rufous, segment III shorter than II, IV to VIII almost gradually shorter, VIII
twice as long as broad, a little longer than X and a little shorter than IX, IX and
X triangular in outline, with the sides somewhat rounded, XI elongate-elliptical.
Pronotum very densely punctate-reticulate, a transverse median area
blackish, sinuous behind, extending forward at side, behind apical margin a
blackish transverse line, rest of surface pale rufous, pubescence of the rufous
parts grey with a golden sheen in certain lights; angle of carina larger than
90°, not rounded off.
Elytrum punctate-striate, the punctures large, especially in stripes VI to
IX, the interspace between adjacent punctures of neighbouring rows not wider
than the punctures, row III containing 12 punctures, rows II to VIII obsolete
on apical declivity ; derm of elytrum rufous, with some darker clouds, pubescence
of base grey, extending along suture and as a line from above shoulder obliquely
to suture, which the line reaches before middle, in front of apical declivity an
indication of a transverse grey band. Pygidium rounded, broader than long,
with sparse grey pubescence. Legs pale rufous, middle of femora darker.
Length 1-8 mm., width 0-9 mm.
Fiji: Taveuni, Waiyevo, x.1924 (Dr. H. S. Evans), 1 9.
12. Melanopsacus stilbus sp. nov.
9. Pitchy brown, elliptical, glossy, with sparse grey pubescence. Antennal
segments I and II, mouth-parts and legs pale rufescent buff. Pronotum with
minute dispersed punctures ; lateral carina straight, angle with dorsal carina
90°, not produced laterad. Elytra not striated apart from sutural stripe, rows
of punctures distinct in basal half, obsolescent in apical half. Pygidium broader
than long, rounded, minutely and dispersedly punctate. Prosternite with large
punctures, in lateral depression posterior to meral suture four transverse rows
of punctures, along lower side of lateral carina a space bare of punctures. Abdomen
slightly coriaceous, without large punctures.
Length 1-1 mm., width 0-6 mm.
Fiji: Taveuni, Waiyevo, x.1924 (Dr. H. S. Evans), 1 9.
13. Doticus sentus sp. nov.
$@. Similar to the Australian D. palmaris Pasc. 1862; coloration of head
and pronotum different, subbasal tubercle of elytrum higher, proportional length
of foretarsal segments of & different, etc.
On head three buff (4) or white (9) lines, median one straight, the others
oblique, the three uniting anteriorly close to base of rostrum ; from this point a
broadish diffuse band of the same colour runs laterad and slightly forward,
enclosing antennal groove ; posteriorly the lines extend near to middle of pro-
notum, terminating rather abruptly ; usually the lines sharply defined, rarely
somewhat diffuse. Eye bordered with buff (¢) or white (2) above. Apical
margin of rostrum slightly curved forward in middle, not incurved. Antenna
432 NOVITATES ZooLoGIcAE XLI. 1939.
pale testaceous, club slightly darker, segment II a little longer than in D. palmaris,
proportional length of segments III to XI in largest & 15, 14, 13, 11, 11, 9, 10,
11, 11, in largest 2 14, 12, 11, 10, 9, 8, 10, 10-5, 12.
Pronotum convex each side of middle, centrally more flattened than in D.
palmaris, basal area more depressed, medianly less convex ; lateral carina less
incurved, usually straight or rounded in dorsal aspect.
Shoulder of elytrum elevate as in D. palmaris, but more evenly convex, less
compressed ; subbasal tubercle posteriorly higher and here more abrupt; in &
a black halfring across middle of suture, open in front, the horns not reaching
the tubercles, the halfring bordered with buff laterally and posteriorly and
placed in a buff patch, which extends to base, but is usually much obscured
anteriorly, the tubercles more or less within the ellipse and usually some black
spots in between them ; in 2 these markings absent, the elytra being blackish,
the anterior half of sutural area darker. Legs testaceous; tibiae with four
brown spots, the first of them basal and usually indistinct. Foretarsus of &
more or less black, segments I and II shorter and slenderer than in D. palmaris,
length (measured in centre) in type I 15, II 8, width I 7-5, II 6-5.
Length ¢ 4:7 to 5-0 mm., width 2-9 to 3-1 mm.; 2 5-0 to 5:3 mm., width
3-0 to 3-1 mm.
Mandated New Guinea: Rawlinson Mts., a series received from Messrs.
Staudinger and Bang-Haas.
Novirarrs Zoonoaican XLI. 1939. 433
ON SOME OLD-WORLD LEPIDOPTERA HETEROCERA.
By KARL JORDAN.
(With 8 text-figures.)
I. Two small North African species of Procris were described by me and
figured in Seitz, Macrolep. ii, pp. 7 and 9 (1909) as Procris cirtana Lucas 1849
and P. orana Austaut 1880. The latter identification was erroneous, there
being nothing in the original description to guide me. The name orana was
based by Austaut on a single ©. The specimen is in the Oberthür collection
of Zygaenids acquired by the late Lord Rothschild. It differs externally from
what I called orana in Seitz in the large tuft placed in between the antennae
and hanging on to the frons consisting to a large extent of long thin hairs and
in the pronotum, coxae and underside of the femora being likewise hairy.
In this the type of orana Aust. agrees perfectly with cirtana, of which orana
is a synonym.
The P. orana Jordan nec Austaut (error of identification) is P. algirica Roths-
child 1917. In this species the head is much more smoothly scaled, the long
narrow scales between the antennae being truncate and not drawn out into a
long point as in the case of a hair-scale, and the pronotum, coxae and femora are
less hairy. The three segments of the g-antenna preceding the last segment are
more or less formed as in P. statices L. 1758 and not pectinate as in P. cirtana.
The type locality of P. algirica is Batna.
II. The genotypical species of the Oriental Saturnid genus Cricula Walker
1855 is C. trifenestrata Helfer 1837, a species which extends in a number of sub-
species from Ceylon and North India to Lombock, Ceram and Halmaheira. In
North-East India a second species is found, C. andrei Jordan 1909, which differs
in the early stages as well as in the imago. Among some duplicates of Cricula
received from the Joicey collection there is a Sumatran ¢ that is similar to
small North Indian examples of C. andrei, but stands apart in structure and less
conspicuously in pattern, representing a new subspecies :
Cricula andrei sumatrensis subsp. nov. (text-fig. 313).
g. In colour of upperside like pale ochraceous buff C. a. andrei, subcostal
brown spot 3 mm. wide, brown line from near apex of forewing to abdominal
margin of hindwing somewhat broader than in C. a. andrei. Underside for the
greater part as brown as distal marginal border. Penis-sheath (Pen) much
narrower than in C. a. andrei (compare text-fig. 312), its internal armature
reduced, consisting of two very narrow scythe-like sclerites and a small triangle ;
ventral portion of penis-funnel representing sternum IX two-horned as in
C. a. andrei, but the horns longer and the undivided proximal portion
narrower.
West Sumatra: Mt. Korintji, 7,300 ft., Aug.-Sept. 1921 (Messrs, Pratt),
i &
31
434 \ Noviratrs ZooLoGIcAE XLI. 1939.
On Mt. Korintji, but at a somewhat lower altitude, also occurs a Cricula
which I described in 1909 as C. andrei elaezia. I had at that time only a single
.6, from Java, of which the characteristic sternum IX was accidentally lost. We
have now a series of both sexes, differing constantly from C. andrei. It is, in my
opinion, the original Malayan Cricula, the commoner C. trifenestrata being a
later arrival in the archipelago and C. andrei a still later immigrant from the
North. The relationship will be better understood if we treat this Malayan
insect as a distinct species, Cricula elaezia, the range of which overlaps that of
C. andrei in Sumatra. Sternum IX (text-fig. 314) is much smaller than in C.
andrei, its horns are always short (though variable to some extent, as they also
are in C. andrei), the penis-sheath remarkably narrow and without armature;
the specimen from which our figure is drawn is somewhat larger than those to
which text-figs. 312 and 313 belong ; the apical margin has minute traces of teeth,
but there are no internal sclerites. In the 2 the postvaginal plate (sternum VIII)
is the same as in C. andrei, the margin bearing a shallow median sinus of some-
what variable breadth (text-fig. 316), whereas in C. trifenestrata there is always a
narrow and deep median sinus (text-fig. 315), an incision rather than an excision.
Both C. andrei and C. elaezia differ from C. trifenestrata also in the j-antennae
the branches being longer. This is particularly noticeable in the distal segments,
as shown in text-figs. 318 (trifenestrata) and 319 (elaezia) (both sketched from
Javan examples, dorsal side, the cilia and sensory cones omitted).
C. elaezia is dichromatic, as is C. andrei, the specimens being dark brown,
with or without a vinaceous tint, or ochraceous buff, the dark individuals con-
trasting with C. trifenestrata from the same district. The species varies also
geographically. The subspecies from Celebes will be described by Professor W.
Roepke and is here omitted.
Novirarrs ZOOoLOGICAE XLI. 1939. 435
(a) C. elaezia elaezia Jordan 1909 (text-fig. 318).
&. Circula andrei elaezia Jordan, Nov. Zoou. xvi, p. 303 (1909) (Preanger, Java)
In the Rothschild collection from : Sumatra, Mt. Korintji, 5,000 ft., Aug.—
Sept. 1921 (Messrs. Pratt), 3 ¢ 4g. Western Java, Preanger, 1 &; Eastern
Java, Nongkodjadjar, 4,000 ft., all
months Dec. to June (J. P. A. Ralis,
A.M.R.Wegner), 9 SI, 7 98,
Tengger, 5,000 and 6,000 ft., April
and June (Kalis), 6 34, Ardjoeno,
4,500 ft., June (Kalis), 1 9, Water-
fall Baoeng, 1,200 ft., July (Kalis),
1 Q. Bali, Batoeriti, 3,500 ft.,
June (Kalis), 4 $4, Gilgit, 5,000 ft.,
April (Kalis), 2 ¢¢.
The usual 3 transparent spots
on forewing of @ larger, at their
basal side 1, 2 or 3 small trans-
parent spots. 7 99 are more or
less orange-ochraceous, 1 9 nearly
tawny and 1 @ dark chestnut ;
none of the SG are as brightly
coloured as the ¢¢ of C. andrei
andrei. 3 of the Bali dd are tinted
with vinaceous and 3 are orange
buff, the vinaceous colour being
more pronounced than in any of our
specimens from Java and Sumatra.
(6) C. elaezia buruensis
subsp. nov. (text-fig. 317). 318 319
&. The single specimen
from Buru agrees with
dark examples from Java,
differing only in the apex WY
of the claspers: as in
many Heterocera the clas-
per of Cricula is divided
at the apex into two lobes,
the two halves of the
clasper being homologous
with the two pleura of the
abdominal segments. The
apical lobes are more or
less strongly rounded in
Cricula; they lie closer
together in buruensis than elsewhere, the ventral lobe (v) is broader than the
dorsal one (d), and the gap between them smaller than in other subspecies ; the
3 figures give a ventral (A), dorsal (B) and lateral (C) view of the two lobes.
Central Buru ; Kako Tagalago, 2,700 ft., May 1922 (Messrs. Pratt), 1 3.
436 Novirares ZOoOLOGICAE XLI. 1939.
EXPLANATION OF TEXT-FIGURES.
Fig. 312. Cricula andrei andrei 3, penis-sheath and sternum IX . p. 434
313. Cricula trifenestrata 3, the same p. 434
314. Cricula andrei sumatrensis 3, the same p. 434
315. Cricula trifenestrata 9, sternum VIII . p. 435
316. Cricula andrei sumatrensis 9, the same : : . p. 435
317. Cricula elaezia buruensis 3, ventral (A), dorsal (B) and lateral
(C) aspects of clasper ; d, dorsal lobe ; v, ventral lobe p- 435
318. Cricula trifenestrata 3, distal segments of antenna, Java p. 435
319. Cricula elaezia elaezia 3, the same, Java . ; ; . p. 435
Novitatrs ZOOLOGICAE XLI. 1939. 437
ON SOME ANTHRIBIDAE FROM AFRICA AND THE
SOLOMON ISLANDS.
BY KARL JORDAN.
(With 3 text-figures.)
1. Litocerus phygus sp. nov.
6. Similar to L. insignis Jord. 1894, which we have from the Kuilu R. and
Camerun. Club of antenna shorter, vittae of pronotum narrower and inter-
rupted, and markings of elytra somewhat different.
Rostrum grey, with three dorsal carinae, median one extending to middle,
lateral ones disappearing on widened apical area; genal groove very distinct.
Frons as in ¢ of L. insignis about one-fourth as wide as the narrowest part of
the rostrum. Proportional length of the segments of the antenna: III 27,
IV 23, V 27, VI 29, VII 34, VIII 29, IX 15, X 12, XI 18; segment VIII about
twice as long as IX, whereas in L. insignis it is practically as long as IX, the
distal segments measuring in L. insignis VIL 30, VIII 25, IX 23, X 15, XI 22.
The broad grey lateral vitta of the pronotum of L. insignis replaced in the
new species by a narrow dorso-lateral stripe which does not quite reach to the
dorsal carina and by a narrow stripe placed at the lateral carina, this second
stripe ending in front of and a little above this carina and extending posteriorly
for some distance along the dorsal carina as a faint border ; median vitta not
reaching apical margin and interrupted (or nearly) between transverse groove
and dorsal carina; behind the carina three spots corresponding to the three
vittae.
On elytra a grey stripe in sutural interspace from before middle to apex,
bearing on suture itself an inconspicuous brown streak before middle ; interstice
II with a short grey dash at base and another at apex and a longer one in middle
on a level with the brown sutural streak and contiguous with short dashes and
spots in interstices III, IV and V; in basal fourth a number of more or less
distinct grey dashes and above and below shoulder a grey patch, the subbasal
swelling remaining brown for the greater part; in apical area a distinct dash
in interstice III and between this dash and the outer margin four or five short
ones forming a transverse band ; laterally at apex a small patch. Pygidium
very little longer than broad, laterally bordered with creamy grey.
Length 5-5-6-0 mm.
Congo Belge : Rwindi, 1,000 m., 20-24 Nov. 1934 (G. F. de Witte), 2 gd;
type returned to the Institut des Parcs Nationaux du Congo Belge, Bruxelles,
paratype retained at Tring.
"2. Gulamentus nephelus sp. nov.
32. Nearest to G. laetus Jord. 1922 from Natal; as in that species the
antennal grooves a little nearer together than are the eyes posteriorly. Markings
different.
Derm black, pubescence greyish white ; in type and some paratypes the
central area of pronotum and the sides of elytra black, this elytral area extending
438 NoviratEs ZooLOGICAE XLI. 1939.
upwards to near suture in middle and again behind it, the black areas irregular
in outline and variable ; they bear some white irregular spots, while there are
black spots on the white areas ; in the other paratypes the dorsal surface greyish
white, the black areas indicated more or less distinctly by diffuse olivaceous
clouds ; in all specimens the scutellum and its immediate surroundings black,
sometimes a few white hairs on it. Underside greyish white; near apex of
prosternum below carina a black spot or a trace of it.
Antenna rufescent, club darker except tip of XI; in g reaching well beyond
base of elytra, in large 3 to near apex, all segments of shaft much longer than
broad, variable in length according to size of specimen, IX longer than VIII,
in large ¢ half as long again as broad and X one-fifth broader than long, in small
3 IX a little longer (one-fourth) than broad, X about twice as broad as long and
XI at least one-half longer than broad. In @ the antenna shorter, VIII less
than twice as long as broad, XI a very little (one-sixth)
broader than long, X more than one-half broader than
long (8:5) and XI twice as long as broad. Pygidium
round at apex in both sexes.
Length 2-8-5-1 mm. ; width 1-5-2-5 mm.
Kenya: Ngong, Dec. 1924 (Dr. van Someren), a short
series, received from the Director of the Imperial Institute
of Entomology, Sir Guy A. K. Marshall, C.M.G., D.Se.,
F.R.S., to whom I am indebted for the majority of the
species here described.
3. Phloeobiopsis farinosus sp. nov. (text-fig. 320).
3g. Close to Ph. plagifer Jord. 1913, differing much in
the antenna.
Black ; upperside of head and rostrum entirely buffish
white, this pubescence very dense, completely concealing
the derm ; pronotum variegated with black, white and
ochraceous nearly as in Ph. plagifer; on elytrum a large
patch above shoulder and another behind middle white,
the latter rather well defined, particularly posteriorly, the
rest of the elytra shaded with white except the following
markings: an ochraceous streak on subbasal swelling and some black dots each
side of the streak, in middle of interstices III and V a white dot each encircled
with black, behind postmedian white oblong a streak each in III and V (and less
distinct in VII) consisting of black and white spots, the adjacent portions of
interstices II, IV and VI ochraceous ; black and white markings shining through
the white powdering, these half-suppressed white spots in median lateral area
transverse. Pygidium densely creamy white. Underside shaded with creamy
white, submaculate at sides, abdomen more densely creamy white each side of
middle. Legs brown shaded and diffusely ringed with white.
Rostrum depressed in middle, width (at widest point) 35, length 22 (meas-
ured from eye to submedian angle of apical margin). Width of frons 22, of
head inclusive of eyes 44. Antenna short, reaching to base of elytra, length of
segments II to XI: 6, 9, 13, 9, 6, 5, 4, 8, 5, 9, IV and V widened on anterior
side (text-fig. 320), not quite so broad as IX, this segment as broad as long,
triangular, asymmetrical, X wider than long, XI with very distinct notch,
NOVITATES ZooLoGIcAE XLI. 1939. 439
III, IV and V covered with long hair on anterior side. Transverse diameter of
eye 16, longitudinal 12.
Pronotum a little more than one-third broader than long (75 : 55), irregularly
rugate-reticulate, dorsally flattened from carina to about middle, side incurved
before basal angle, which is somewhat less than 90°, and convex from apex of
lateral carina forward, the anterior margin not projecting above the level of the
neck ; dorsal carina distinctly, though slightly, convex each side. Scutellum
longer than broad, white.
Length of elytra 135, width 75, without tufts; interstices III, V and VII
slightly raised, apex flattened-depressed, especially interstice II. Pygidium
rounded, one-third broader than long (32 : 24).
Prosternum narrow in front of coxa, slanting, without ridge, except the
elevate anterior margin. Mesosternal intercoxal process narrower than coxa,
slanting, apex rounded, not abruptly curved back. Abdomen somewhat uneven
at side of middle. Hindfemur reaching to apex of abdominal segment IV;
hindtibia on underside with a median swelling bearing long pubescence, the
under surface from this swelling to the apex flattened and hairy. Segments II
and III of tarsi broader than in Ph. plagifer.
Length 12 mm.; width 5 mm.
Kenya: Kitui, 1 ¢.
4, Aulodina bifax sp. nov.
9. A narrow species with parallel sides, nearly two and one-half times as
long as broad (head excluded from measurement), bearing a circular patch on
pronotum and a smaller one on suture before apex of elytra.
Rostrum very short, in front of antenna nearly four times as broad as long,
apical margin with median sinus, of which the edge is marginate and therefore
well defined, the sinus occupying one-third of the entire apical margin ; rostrum
and head black, with a very small white tuft in middle of frons, the whole surface
reticulate, the meshes of the net shallow and more or less hexagonal. Eye more
coarsely granulate than in A. unicolor Jord. 1903. Antenna rufescent, segment
II claviform, as long as III, but broader, half as long as the frons is wide, III to
V gradually shorter, VI to VIII about equal in length, a little shorter than V,
club a little longer than II + III + IV, nearly four times as long as broad,
proportional length of the three segments: 6, 5, 9, XI being elongate-ovate and
slightly wider than IX and X.
Pronotum as long as broad, strongly convex, the central area covered with
rather sharp ridges (in A. unicolor the ridges flattened), sides coriaceous, pubes-
cence grey mixed with white, brown and luteous, the grey more concentrated at
apical angle, appearing almost white, on disk a short distance from apical margin
a black ring the external diameter of which is as long as half the width of the
pronotum, the ring diffusely bordered on outside with grey, white and luteous
and its interior filled with the same mixture of pubescence ; lateral angle of
carina over 90°, lateral carina extending to apical margin, ending here with a
distinct tooth. Scutellum white.
Elytra with grey and luteous pubescence, diffusely spotted with white and
bearing a few blackish dots; on apical declivous area, but not touching apical
margin, a deep black circular spot, the diameter of which is two-thirds that of
the black ring of pronotum, the black spot diffusely encircled by grey and luteous
pubescence ; the rows of punctures distinct.
440 NOVITATES ZOOLOGICAE XLI. 1939.
Underside blackish, the grey pubescence not concealing the derm. Tibiae
rufescent, with grey pubescence and three incomplete brown rings.
Length 3-7 mm. ; width 1-5 mm.
Kenya: Rabai, Aug. 1937 (Dr. van Someren), 1 9.
5. Apatenia phaeura salomonis subsp. nov.
d. Like A. phaeura Jord. 1903 from New Guinea, with grey apex to the
elytra. Frons broader, the rostrum being 2-75 times as broad as frons in A. ph.
phaeura and 2-40 times in A. ph. salomonis; angle of pronotal carina 90° in
salomonis and the lateral carina very slightly curved, in phaeura the angle obtuse
and the carina distinctly curved.
On pronotum a buffish grey lateral stripe parallel with margin, nearer to
the latter than to middle, its anterior half broader than posterior half, being
almost interrupted, disk with central depression bearing an indefinite silky
buffish grey patch, a median line of the same colour indicated by a short thin
stripe at apex and before carina, ending in front of scutellum with an ovate
spot. On elytra numerous black dots, some of them raised as pustules, the
most conspicuous one of them behind middle of interstice III ; grey apical patch
conspicuous, the suture and apical margin remaining black. Legs less rufescent
than in A. ph. phaeura.
Length 7-3 mm., width 3-2 mm.
Solomon Islands : Guadalcanar, 15 Nov. 1935 (R. A. Lever), 1 3.
6. Plintheria leveri sp. nov.
9. Near P. woodlarkiana Jord. 1898 ; the rostrum and antenna shorter and
the elytra entirely striped with grey.
Rostrum as broad at apex as long, coarsely rugate, with three carinae,
which disappear on apical dilated area. Frons half the width of apex of rostrum.
Antenna rufescent, club black, length of segments I to XI: 5, 5, 9, 7, 6, 6, 5:5,
5-5, 7, 5, 6; VIII distinctly broader than VII, but not forming part of the club
as it does in Phaeochrotes Pasc. 1860, IX as broad as long (length measured
along middle), X broader than long, XI short, ovate, strongly rounded at base
and sides, apex pointed.
Pronotum one-half broader than long, with dispersed grey pubescence,
which is denser laterally at apex and before dorsal carina; the middle of this
carina less deeply concave than in P. woodlarkiana (and P. luctuosa Pasc. 1859)
and its lateral portion less curved forward. Elytra more strongly punctate-
striate than in the allied species, all the interstices grey, the pubescence forming
regular stripes from base to apex (some united subapically), those in sutural
and alternate interstices broader than the others.
Tibiae grey except apex, which is black-brown. Tarsal segment I shorter
than in P. woodlarkiana, in hindleg a little over one-half the length of the tibia
(in P. woodlarkiana two-thirds),
Length 2-8 mm ; width 1-4 mm.
Solomon Islands: Lunga, Guadalcanar, May-June 1935 (R. A. Lever),
3 99. Ihave much pleasure in associating the name of the discoverer with this
distinct little species.
NOVITATES ZoonoaicaArn XLT. 1939. 44]
7. Proscopus aper sp. nov. (text-figs. 321, 322).
3g. Rufescent clay-colour, cheek grey, elytra with numerous short grey
lineoles, femora rufous, dark brown at apex. Face long, measured from vertex
to tip of mandibles nearly thrice as long as it is broad at the narrowest point,
sides strongly incurved; upper surface coarsely granulose-rugose ; rostrum
with short smooth median line at base, from tubercle bearing the antenna down-
wards a vestige of a thin lateral carina ; beyond constricted median area a horn
each side curving forward-upward-inward and like the surface of the rostrum
rough with small tubercles. The two horns alike, but appearing different in
text-fig. 321, because this figure presents a view taken obliquely from the side
and behind. Text-fig. 322 is drawn from above and behind, the face being
foreshortened in this aspect ; apical margin of rostrum with a rather prominent
tubercle each side of labrum, the sinus between the tubercles shallow and but
feebly rounded. Eye bilobate, its frontal lobe narrower than the lateral one
and only a little wider than the interspace between the eyes. Antenna nearly
three times as long as pronotum and elytra together, length of segments I to XI:
27, 23, 75, 56, 55, 51, 49, 42, 29, 20, 24 (globular base of I not included), last
segment pale rufous. Length of face 70 (inclusive of mandibles).
Pronotum one-third broader than long, granulose-rugulose, sides rounded,
lateral carina reaching to near apical margin. Elytra punctate-striate. Abdo-
men punctate-coriaceous. Foretarsal segments II and III strongly dilated,
III being twice as broad as the tip of the tibia.
Length 3-8 mm.
Solomon Islands : Bukum, Guadalcanar, 8 Sept. 1935 (R. A. Lever), 1 ¢.
8. Proscopus vellanus sp. nov.
6. Like the preceding species; smaller; face very slightly narrower ;
rostrum without horns, apical tubercles somewhat higher, interspace between
dorsal lobes of eyes a little more concave. Antenna shorter and the length of
442 NoviratEs ZooLoGIcAE XLI. 1939.
the segments different : 20, 15, 40, 31, 28, 27, 27, 25, 19, 14, 18, last segment
pale as in the previous species ; taking the length of X as 1, the proportional
length is as follows :
I II III IV Vv VI | VIL | VIIL}| IX x XI
P. aper 0 .| 1:35 | 1:15 | 3:75 | 2:80 | 2:75 | 2:55 | 2:45 | 2-10 | 1-45 | 1-00 | 1-20
P. vellanus . . | 1-43 | 1:07 | 2:86 | 2:22 | 2:00 | 1-93 | 1-93 | 1-79 | 1:36 | 1-00 | 1-29
I is one-sixth longer than II in P. aper and one-third in P. vellanus, IV about
one-third longer than V in P. aper and about one-ninth in P. vellanus, ete.
There do not appear to be any other differences.
Length 3-1 mm.
Solomon Islands: Lambulambu, Vella, 18 Sept. 1935 (R. A. Lever), 1 &
(according to genitalia).
Novirarres ZooLOGIcAE XLI. 1939, 443
ON RHOPALOPSYLLUS BAKER 1905 (SIPHONAPTERA).
BY KARL JORDAN.
(With 10 text-figures.)
Swe the publication of the survey of Rhopalopsyllus Baker 1905 in Ecto-
parasites, pp. 320-351 (1923) a considerable number of new species have
been described, and it is to be expected that many more will be discovered. In
these circumstances it is advisable to treat the three natural groups into which
we divided the genus in 1923, l.c., as different genera, our expectation expressed
l.c. that the discovery of new species would fill the gaps between the three
sections not having come true.
1. Rhopalopsyllus Baker 1905; genotype: Pulex lutzi Baker 1904.
Rhopalopsyllus Baker, Proc. U.S. Nat. Mus., xxix, pp. 128-130 (1905) (partim ; lutzi designated
genotype).
Rothschildella Enderlein, Zoolog. Anzeiger, xl., p. 72 (1912) (genotype: cryptoctenes = lugubris) ;
Jord. & Roths., Ectoparasites, p. 320 (1923) (Rothschildella = Rhopalopsyllus).
The diagnosis given in Hctoparasites for Section A reads as follows : “ Bristles
of anterior row on frons weak, except upper bristle. Prosternum anteriorly
extending downwards as a rounded projection. Mesosternum as high as long.
Fourth hindtarsal segment short, very strongly dilated apically, fifth of all tarsi
long, in hindtarsus longer than second midtarsal segment. Anterior dorsal
margin of pygidium
strongly project-
ing.”
In none of the
species is the body
of the spermatheca
humped. The
phallosome is
characteristic, as ex-
plained below.
The most widely
distributed species is
Rh. australis Roths.
1904, occurring in
several subspecies
from Mexico to South-East Brazil. A series of Brazilian specimens of this
species lately received for determination from Dr. Fred L. Sloper, Rockefeller
Foundation, Rio de Janeiro, calls for a remark on the variability of a minute
subspecific distinction. In the males from Anapolis, Goyaz (Rockefeller
Foundation) and from Sao Paulo (Tring collection) the lower internal incrassation
of the posterior margin of the clasper (text-fig. 323, lo; specimen from S.
Paulo) is smaller than the upper incrassation (up). In typical Rh. australis
tomoyus J. & R. 1923 from Paraguay the lower incrassation is at least as large
444 NOVITATES ZoOLOGICAE XLI. 1939.
as the upper. The material from Paraguay is not extensive enough for
ascertaining whether this is always the case in that country (and Argentina,
whence we have no examples of Rh. australis). The males from Santa Cruz
de la Sierra, Bolivia, submitted by the Rockefeller Foundation, are more or
less intermediate. For the present I look upon all these specimens as being
Rh. australis tomoyus. Inthe S. Paulo specimen from which fig. 323 is taken the
groove (g) of the clasper which corresponds to the angle of the anterior margin
of the digitoid F is exceptionally large ; as a rule the groove is about half the
size as in the figure.
2. Polygenis gen. nov.; genotype: Pulex roberti Roths. 1905.
Prosternum not humped anteriorly between the coxae. Mesosternite
(measured from anterior margin of sternum to apex of mesepimerum) longer
than high. Tarsal segment V much smaller than in Rhopalopsyllus, that of
hindtarsus not longer than midtarsal segment II.
Here belong the majority of the species now standing as Rhopalopsyllus.
In most of them the body of the spermatheca is distinctly humped. We have
selected roberti as genotype, because the species is easy to recognize ; one of its
main recognition characters, however, varies to some extent, which must be
borne in mind: the subapical dorsal notch of the hindtibia usually bears three
heavy bristles, but in one of our males there are only two in one tibia ; in another
specimen there is a strong dorsal bristle between the second and third pairs
instead of a thin hair.
3. Tiamastus gen. nov.; genotype: Pulex cavicola Weyenb. 1881.
Proboscis extending to the apex of trochanter or beyond, whereas in Poly-
genis it reaches at most to the base of trochanter. Prosternum not humped
324
marg. =—<--- hg.
anteriorly between the coxae. Mesosternite longer than high. In & posterior
margin of clasper with subapical tooth below which there is a long submarginal
bristle. In @ sternum VII with lateral sinus, and spermatheca with long
nose.
Fr
Novırares Zoonogican XLI. 1939. 445
Only a few species are known, in all of which the longest bristle of hind-
tarsal segment II reaches to or beyond apex of IV: T. cavicola Weyenb. 1881,
T. callens J. & R. 1923, T. subtilis J. & R. 1923 and T. palpalis J. & R. 1923.
The species of the three genera have an interesting characteristic in common
which does not seem to have been mentioned in the descriptions of these
fleas : the distinction
is found in the
sclerites which are
accessory to the duc-
tus ejaculatorius and
form a compact body
for which we accept
the term phallosome
introduced in the
systematics of Dip-
tera. The phallosome
of Rhopalopsyllus re-
sembles in dorsal
aspect (text-fig. 324,
Rh. lutzi) an elongate
(very pale) caverni-
cole beetle with long
prothorax, its outer
covering, the mantle
(mtl.), being divided
into the apical hood,
the median neck and
the larger, elongate-
ovate, anterior rump-
cover. This rump-
cover is open below
and its margin, some-
what more strongly
chitinized than the
rest of the mantle,
curves down and back
and unites with the
margin of the other
side in a sort of heel
(hmg.), similar to the
heel of sternum IX, the heel terminating the ventral mantle of the neck (text-
fig. 325), but being continued forward by a tendon. The elongate rump-cover,
which resembles the elytra of a beetle, is longitudinally marked with a suture.
The cavity within this portion of the mantle is centrally divided by a lamina
which is suspended from the suture and ends distally with a transverse bar
(dorsal aspect), the bar appearing as a kind of condylus (cond.) in lateral aspect.
The neck contains the strongly chitinized and long tube through which the
ejaculatory duct passes distad. The proximal portion of the ejaculatory duct
446 NOVITATES ZOOLOGICAE XLI. 1939.
which lies below the body of the mantle is supported by a long sclerite and
forms a vesicle (ves.) before entering the tube. It is an interesting fact that
the penis-tube consists in the three genera of two portions : a strongly chitinized
basal segment (bpt.) and a narrow distal one (apt.), separated from each other
Marg. ves . jam.
by a joint (articulation 2); the basal tube is also articulated (art. 1) with the
condylus of the lamina, there being a cavity (caps.) between the condylus and
penis-tube variously filled with ligaments and ossicles. There are some specific
differences in the phallosomes of the four species represented by fig. 325 (Rh.
lutzi), fig. 326 (Rh. lugubris), fig. 327 (Rh. australis) and fig. 328 (Rh. cacicus) ;
Novirares Zootogıcan XLI. 1939. 447
in Rh. cacicus, for instance, the basal segment of the penis-tube is longer than
in the other species, the capsule is wider, the free distal portion of the duct is
for the greater part enclosed in muscles and the heel (hmg.) is not thickened ;
in Rh. lutzi a portion of the apical duct also lies in a muscle, whereas in Rh.
lugubris and Rh. australis the duct appears to lie free in the cavity of the neck,
the dorsal membrane of which is thin and flexible. All four species have this
point in common that the duct, on leaving the tube, turns dorsad in a lateral
view as depicted. The free duct is more or less rolled up as a spiral except in
australis and the position of the coil is variable. If the duct lies partly at the
side of the tube before the specimen is cleared, the pressure of the coverslip may
cause the beginning of the duct first to turn towards the ventral side instead of
the dorsal one.
In Polygenis (text-figs. 329-331) the two segments of the penis-tube are
usually longer than in Rhopalopsyllus,
the basal segment being particularly
long in P. tripus (text-figs. 330, 331).
The pale space in the dorsal wall of the
basal segment is very much longer in
P. tripus (and P. axius J. & R. 1923)
than in other species, as shown in the
figures. The duct protruding from the
tube turns ventrad and is coiled near
the apex of the tube. Fig. 331 (less
enlarged) represents a specimen of
P. tripus in which the basal tube is
turned ventrad. The heel is not
inbrassate. Only P. klagesi is an
exception as regards the free duct,
agreeing in this particular with the
next genus.
The phallosomes of the species of
all Tiamastus resemble that of T. cavicola (text-fig. 332). The free duct turns
dorsad as in Rhopalopsyllus, but is not rolled up, being long and extending
forward to the base of the penis-tube, more or less as in the figure. The
ventral heel (hmg.) of the mantle has a distinct incrassation, as in Rh. australis,
Rh. lugubris and Rh. lutzi, more or less.
Although P. klagesi agrees best with Polygenis, it has much in common
with T. cavicola. The long bristles of the hindtarsus are longer than in other
species of Polygenis, as is the case with all Tiamastus, segment V of foretarsus
is short and broad as in T. cavicola, and the ejaculatory duct turns dorsad on
emerging from the tube (as in Rhopalopsyllus and Tiamastus) and extends
forward to the basal segment of the tube, making a loop and a coil or two, but
not being rolled up as in other Polygenis. "There is no distinct heel to the mantle.
The basal penis-tube is exceptionally short for a species of Polygenis. We
observe in P. klagesi a mixture of somatics which suggests that we have here
a side-line of development of which so far only this one representative is
known.
With the exception of fig. 323, the illustrations are drawn from specimens
mounted in balsam. The appearance of the detail in the composition of the
448
NOVITATES ZOOLOGICAE XLI. 1939.
phallosome depends much on the method of clearing and mounting and the
depth of the focus, particularly as regards the condylus and the contents of
the capsule.
EXPLANATION OF THE TEXT-FIGURES.
Fig. 323. Rhopalopsyllus australis tomoyus J. & R. 1923, Sao Paulo. Cl,
clasper ; up, upper incrassation ; g, groove ; lo, lower incrassation.
Fig. 324. Rhopalopsyllus lutzi lutzi Baker 1904, phallosome extracted, dorsal
aspect; 100 x mtl., mantle; lam., lamina; art. 1, first
articulation ; art. 2, second articulation ; caps., capsule; marg.,
ventral margin of mantle; hmg., heel of mantle; ves., vesicle ;
bpt., basal segment of penis-tube ; apt., apical segment of penis-
tube ; ej. d., ejaculatory duct.
Fig. 325. Rhopalopsyllus lutzi lutzi, the same, lateral aspect as in all the
following figures ; 180 x.
Fig. 326. Rhopalopsyllus lugubris J. & R. 1908.
Fig. 327. Rhopalopsyllus australis tupinus J. & R. 1923.
Fig. 328. Rhopalopsyllus cacicus saevus J. & R. 1923.
Fig. 329. Polygenis roberti Roths. 1905.
Fig. 330. Polygenis tripus Jord. 1933; 180 x
Fig. 331. Polygenis tripus ; 105 x.
Fig. 332. Tiamastus cavicola Weyenb. 1881.
abdominalis (Strymon),359.
Abtrichia, 217, 224, 228.
acaciae (Strymon), 360.
Accipiter, 67.
Acherontia, 261.
Acomys, 191.
Acorynus, 140.
Acostatrichia, 228-230.
Acraea, 260.
Acrobates, 58-60.
Acryllium, 311.
Actinote, 304.
acuticauda (Poephila), 71.
adana (Nycteris), 204.
Adoratopsylla, 164-166.
adscitus (Platycercus), 69.
adspersa (Euprepocnemis), 378.
adspersus (Thisoicetrus), 377, 378, 381.
adusta (Thecla), 357.
aegyptiaca (Vulpes), 200.
aegyptius (Pluvianus), 311.
aequalis (Argyrolepidia), 249, 250.
aequinoctialis (Procellaria), 134, 137.
aesalon (Falco), 310.
aestiva (Amazona), 312.
aethiopicus (Paraechinus), 202.
Aethomys, 112.
afer (Stivalius), 115, 116.
africana (Chapinia), 63, 70.
Agapornis, 69, 312, 313.
Aglais, 125.
agyrtes (Ctenophthalmus), 103-105, 109.
Ailanthus, 130.
ailanti (Daphnusa), 130.
— (Smerinthus), 130.
aino (Thecla), 356.
airensis (Jaculus), 191.
Aix, 66, 309.
alba (Chionis), 68.
— (Kgretta), 308.
albatus (Paraechinus), 202.
albertensis (Foxella), 124.
albescens (Rhea), 308.
albicauda (Ichneumia), 198.
albicaudus (Herpestes), 198.
albilinea (Satsuma), 359.
albinasus (Gulamentus), 144,
32
INDEX.
albior (Paraechinus), 202.
albocristatus (Tropicranus), 70, 313.
albogularis (Garrulax), 314.
albolineata (Mortoniella), 217, 218.
Alciopa, 348.
aldabrae (Utetheisa), 271.
aldabrensis (Utetheisa), 271.
alexandrinus (Rattus), 192.
algirica (Procris), 433.
Allocanthochasmus, 159.
alternata (Syllis), 383.
alternatus (Kurodaia), 306.
alveatus (Gulamentus), 145.
amabilis (Lithosia), 273.
Amadina, 314.
Amaurornis, 311.
Amazona, 312.
americana (Rhea), 308.
amhara (Utetheisa), 253-256, 261, 263-266, 268,
272, 274.
amir (Paraechinus), 203.
Ammotrypane, 345.
Ampharete, 346.
Amphitrite, 346.
Amyrsidea, 306.
anahua (Exitrichia), 233.
Anas, 66.
Anaticola, 307.
Anatoecus, 67, 307.
anchylochaeta (Nereis), 397.
andrei (Cricula), 433-436.
andricus (Parapsyllus), 300.
angulata (Misthosima), 147.
ankylochaeta (Opisthosyllis), 390.
annulata (Caulibugula), 340, 342.
annulosus (Thisoicetrus), 377, 381.
anomala (Pseudonereis), 402.
antennata (Abtrichia), 226, 227.
— (Deiopeia), 278, 279.
— (Eunice), 398.
— (Utetheisa), 253, 261-263, 266, 268, 278.
Anthropoides, 311.
Antilope, 185.
antiquorum (Adoratopsylla), 165-169.
— (Phoenicopterus), 66, 309.
Antoptila, 219, 220.
Aunrosorex, 374,
449
450
Apatenia, 440.
aper (Proscopus), 441, 442.
aphanis (Utetheisa), 281, 282.
Aplodontia, 319.
Apodemus, 103, 105, 107, 108, 110.
apollinaris (Tribotropis), 423.
Apophallus, 155.
aporus (Stivalius), 373.
aquilus (Lophuromys), 114.
aquitanius (Microtus), 106, 108.
Ara, 313.
arabica (Antilope), 185.
— (Gazella), 184-188.
— (Vulpes), 199-210.
arabicus (Lepus), 189, 190, 201.
arabium (Dipodillus), 195.
arabs (Canis), 201.
arcuata (Deiopeia), 274.
Ardea, 66, 308.
Ardeicola, 62, 65, 68.
arduus (Gerbillus), 195,
Arenicola, 346.
argentatus (Larus), 312.
Argina, 252.
Argyrolepidia, 133, 249.
arida (Ludia), 132.
arietans (Bitis), 304.
— (Vipera), 304.
arimalius (Meriones), 193.
arisanus (Thecla), 358.
aristotelis (Phalacrocorax), 308.
arizonensis (Panthera), 420, 422.
— (Sciurus), 317.
armata (Caulibugula), 342.
armatum (Stamnosoma), 155.
armiger (Scoloplos), 345.
armillaris (Syllis), 385.
arquata (Numenius), 311.
arvernus (Ctenophthalmus), 105.
Asellia, 203.
asiatica (Cyanops), 65, 70.
— (Zenaida), 69.
asiaticus (Thisoicetrus), 381.
asio (Megabothris), 122.
assumptionis (Utetheisa), 271.
astrild (Estrilda), 314.
asunicus (Typhloceras), 111.
Asychis, 354.
asymmetrica (Dahlemhornia), 64, 65.
Athene, 70.
atlanticus (Leocrates), 348.
atrata (Chenopis), 66.
atropos (Acherontia), 261.
attenuatus (Notocotylus), 75, 78.
audax (Uroaétus), 310.
aulaster (Ammotrypane), 345.
Aulodina, 439,
aura (Cathartes), 309.
aurita (Cercaria), 78.
aurorina (Thecla), 356.
australiacus (Parapsyllus), 134, 137.
australis (Microcavia), 293, 294.
— (Opisthosyllis), 390, 391.
— (Rhopalopsyllus), 443, 444, 446, 448.
austrinum (Hippotion), 130.
Austrogoniodes, 62, 64-66.
aviatica (Zenaida), 312.
avocetta (Recurvirostra), 68.
axius (Polygenis), 447.
baeticus (Ctenophthalmus), 108.
bailwardi (Tatera), 196.
Balearica, 311.
bamanus (Macrostylophora), 373.
bantorum (Xenopsylla), 112, 114.
Baritius, 255.
barrerai (Parapsyllus), 297.
Basitropis, 427.
Beania, 321, 327, 329.
bella (Utetheisa), 251, 252, 255, 260, 263, 266,
268.
bengalus (Uraeginthus), 71, 314.
benthophila (Harmothoé), 347, 349, 350.
bergylta (Labrus), 179.
bernicla (Branta), 309.
Betrichia, 230.
Bibulus, 382.
Bicellaria, 339.
bicolor (Deiopeia), 274.
— (Dejopeja), 274.
— (Hucus), 140.
— (Utetheisa), 288.
bifax (Aulodina), 439.
bifida (Canoptila), 218.
bilaminata (Beania), 329.
bimucronata (Harmothoé), 349.
bisetosa (Adoratopsylla), 164-168.
Bitis, 304.
blanfordi (Paraechinus), 203.
Blennius, 179.
bolivari (Thisoecetrus), 380.
boliviensis (Panthera), 410, 414, 415.
Bombyx, 268, 278.
bonana (Icterus), 72.
borneensis (Macrostylophora), 366, 367.
brachychaeta (Syllis), 347.
Branchioasychis, 347, 352-354.
Branta, 309.
brasiliana (Antoptila), 219, 220.
brasiliensis (Xenopsylla), 112, 115.
breviceps (Thaumapsylla), 374.
brevipalpe (Eidmaniella), 306.
brevis (Gulamentus), 146.
— (Lopadorhynchus), 347.
brillantina (Thecla), 356.
broscus (Hectopsylla), 303.
Bruélia, 62, 64, 70-73, 307.
brunnea (Opisthosyllis), 389.
brunni (Bibulus), 382.
— (Tylotropidius), 382.
Bubo, 70, 313.
bubo (Bubo), 313.
Bubuleus, 66.
Bucephala, 309.
Bucorvus, 70.
budensis (Procris), 213.
budini (Parapsyllus), 294, 299.
Bugula, 329-331.
Capra, 184.
Caracal, 197.
caracal (Caracal), 197.
— (Felis), 197.
caraibica (Caulibugula), 342.
cardinalis (Richmondena), 72, 315.
Cariama, 305, 311.
cariboa (Eunice), 345, 348, 352.
carpenteri (Dalhousiella), 348,
carunculata (Hermodice), 347.
cascadensis (Orchopeas), 317.
castanea (Anas), 66.
castanotis (Taeniopygia), 314.
451
burtoni (Procavia), 182, 183. catalina (Sciurus), 317.
buruensis (Cricula), 435. Catallagia, 124.
— (Mimeusemia), 249. Cathartes, 309.
buryi (Meriones), 193. Caulibugula, 321, 339-342.
buski (Emma), 327. cavicola (Pulex), 444, 445.
buskii (Emma), 326, 327. — (Tiamastus), 445, 448.
— (Menipea), 327. ceblebrachys (Eustrigiphilus), 307.
Buteo, 67, 310. celticus (Ctenophthalmus), 104.
buteo (Buteo), 67, 310. centralis (Panthera), 417, 419, 420.
butleri (Hippotion), 131. Ceratonereis, 397, 400.
— (Panacra), 131. Ceratophyllus, 63, 67, 365.
— (Procavia), 183. Cercaria, 42-50, 52-56, 75-79, 81, 93, 148, 155,
buxtoni (Notiana), 424. 178, 180.
— (Thisoicetrus), 378, 382. Cereopsis, 309.
cervicornis (Emma), 321, 326.
— (Goniodes), 64, 67.
cethis (Nessiara), 142.
Chalccophaps, 69.
cacicus (Rhopalopsyllus), 447, 448.
caeca (Nephthys), 346.
— (Syllis), 383. chalcoptera (Phaps), 69.
caerulea (Rapala), 361. chalcospilus (Turtur), 312.
— (Thecla), 361. chalybeata (Hypochaera), 72.
caerulescens (Euprepocnemis), 382, chandrana (Synthusa), 360,
cafer (Bucorvus), 70. Chaperia, 327.
c-album (Polygonia), 125. Chapinia, 63, 70.
Calidris, 312. charax (Plesiobasis), 430.
callens (Tiamastus), 445. Charaxes, 130.
callima (Utetheisa), 262, 275. charpentieri (Thisoecetrus), 379, 380.
Callizona, 347, 348. chaus (Felis), 362.
cameratus (Nirmus), 175. cheesemani (Lepus), 190, 201.
camiguina (Perinereis), 399. Chenopis, 66.
canagica (Philacte), 309. cheopis (Xenopsylla), 112, 116.
candida (Mauia), 146. Cheyletiella, 63, 65, 71, 73.
Canis, 201. chinensis (Cissa), 71.
canis (Ctenocephalides), 116. — (Ptolyptychus), 127.
Canoptila, 218. Chloeia, 348.
canora (Tiaris), 72. Chloris, 72.
chloris (Chloris), 72.
chloros (Procris), 213, 215.
canorus (Cuculus), 69.
canutus (Calidris), 312.
capense (Bugula), 330. Chrysolampis, 73.
capensis (Bubo), 70, 337. chrysolophus (Eudiptes), 65.
— (Bugula), 329-332. Chrysopa, 304.
— (Chaperia), 327. ciliata (Caulibugula), 342.
capistrata (Lioptila), 313. — (Nephthys), 346.
capitulatus (Onychomys), 121. ciliatus (Monopsyllus), 120-122.
caponis (Lipeurus), 62, 64, 67. cincinnatus (Thelepus), 345, 346.
452
cinerea (Ardea), 66, 308.
— (Neotoma), 317.
Circaetus, 310.
cirrata (Amphitrite), 346.
cirrosa (Gattyana), 345, 346.
cirtana (Procris), 433.
cisandinus (Rhopalopsyllus), 293.
Cissa, 71.
Citellophilus, 106, 107.
citrinellae (Philopterus), 64, 72.
clivosus (Rhinolophus), 204.
claparedii (Leocrates), 392.
Clonorchis, 148, 155.
closterobranchia (Syllis), 384.
coelebs (Fringilla), 315.
cognata (Procris), 213, 216.
— (Thecla), 355, 357.
colmani (Branchioasychis), 345, 352, 353.
Coloceras, 307.
Colpocephalum, 63, 66, 67, 176, 306.
columbae (Columbicola), 65, 69.
columbarius (Falco), 310.
Columbicola, 65, 69, 307.
communis (Turritella), 84.
completa (Utetheisa), 262, 275.
compressicornis (Tribotropis), 423.
conchylega (Onuphis), 345, 346.
conicollis (Tribotropis), 423.
constantina (Felis), 197.
continuum (Acridium), 379.
continuus (Thisoecetrus), 379.
coomani (Acorynus), 140.
contrainii (Alciopa), 348.
corallicola (Syllis), 347, 348, 385.
coromandus (Bubulcus), 66.
coronanda (Cercaria), 148-163.
cornuta (Ehlersia), 387.
— (Syllis), 387.
corticea (Ludia), 132.
coruscans (Thecla), 356, 357.
Cossypha, 313.
costae (Ceratonereis), 400.
costalis (Polyptychus), 126.
Costatrichia, 228.
coutierei (Nereis), 394.
coxi (Panthera), 410, 414, 415.
Craneopsylla, 164.
Craspedorrhynchus, 307.
crassus (Meriones), 193.
Crepidistomum, 376.
crestatus (Eudiptes), 66.
Cricula, 433-436.
crispus (Paraceras), 364.
cristata (Cariama), 305, 311.
— (Paroaria), 73, 315.
— (Pista), 347.
cristatus (Eudyptes), 66, 134, 135.
— (Pavo), 67, 310.
cristatus (Podiceps), 66.
cristiferus (Rhinotropis), 424, 425.
Crocidura, 108, 202.
crosslandi (Bugula), 329, 330, 334-338.
cruentata (Deiopeia), 270, 271.
— (Utetheisa), 257, 258, 261, 263, 266, 267,
270.
Cryptocotyle, 102, 148, 151, 152, 178.
erystallina (Emma), 321-326.
Ctenocephalides, 116, 362.
Ctenolabrus, 179.
Ctenophthalmus, 103-109, 114, 124, 165.
Cuculus, 69.
cultrifera (Perinereis), 396-399, 402.
Cummingsiella, 307.
cuneata (Geopalia), 312.
cunhai (Adoratopsylla), 166.
cursitans (Philopterus), 64, 70.
curvirostra (Loxia), 72.
cuvieri (Gazella), 185.
cyanea (Pitta), 70.
cyanescens (Thisoicetrus), 377, 380.
Cyanops, 65, 70.
cyanoptera (Anas), 66.
cyathus (Emma), 321, 322.
cylindricus (Gulamentus), 144, 145.
Cyrtacanthacris, 381.
Dahlemhornia, 64, 65.
Dalhousiella, 348.
danubianus (Citellophilus), 106, 107.
Daphnusa, 130.
darwini (Utetheisa), 283, 284.
dasyurus (Dipodillus), 194.
decimfasciatum (Colpocephalum), 63, 66.
Degeeriella, 62, 64, 67, 68, 70, 307.
Deilemera, 270.
Deilephila, 261.
Deiopeia, 269-287.
Deiopia, 282.
dejeani (Strymon), 360.
Dejopeja, 274.
Delias, 130.
demersus (Spheniscus), 65.
Dendronessa, 67.
depictus (Gulamentus), 143, 146.
depressa (Syllis), 384.
deserti (Paraechinus), 202, 203.
desgodinsi (Thecla), 357.
Diadema, 304.
diadematus (Stephanophorus), 71.
diamantina (Thecla), 355.
dianae (Psammomys), 192.
Diardigallus, 67, 310.
diardi (Diardigallus), 67, 310.
Diaulus, 218.
Didelphis, 166.
dilecta (Adoratopsylla), 168.
dilutior (Utetheisa), 262, 263, 275, 276.
dimidiatus (Acomys), 191.
— (Mus), 191.
Dimorphozoum, 321, 327, 329.
Dinosaphis, 425.
Diodosida, 131.
diplogonus (Goniocotes), 64, 67.
Dipodillus, 194, 195.
Dipodomys, 121, 122.
discors (Anas), 66.
discreta (Adoratopsylla), 165-168.
dispar (Neopsylla), 373.
dissimilis (Malaraeus), 119.
diva (Deiopeia), 269, 270.
divaricata (Pionosyllis), 389.
dollfusi (Syllis), 384.
dolosus (Mecocerus), 424.
Dolotrichia, 235.
domesticus (Glyciphagus), 308.
dominicana (Paroaria), 73, 315.
dominicanus (Larus), 68.
donicum (Rossicotrema), 152.
Doratopsylla, 165, 169.
dorsalis (Paraechinus), 202, 203.
dorsifusa (Utetheisa), 283.
Doticus, 431.
Dromiceius, 65.
Dromicia, 59.
Drosophila, 251.
dubernardi (Thecla), 356.
dubitans (Dolotrichia), 235.
— (Protoptila), 221.
duma (Thecla), 357.
dumerilii (Nereis), 386, 398.
— (Platynereis), 401.
ealanus (Gulamentus), 143, 145.
ebraeum (Esthiopterum), 176.
Echidna, 304.
echinata (Flustra), 327.
Echinometra, 304.
econia (Mimeusemia), 248.
edithae (Meriones), 193, 194.
edwardsii (Herpestes), 198.
Egretta, 308, 309.
Ehlersia, 387, 388.
ehlersi (Leonnates), 404. 5
ehlersiaeformis (Pionosyllis), 388.
Eidmaniella, 306.
elaezia (Cricula), 434, 435,
elata (Bombyx), 268.
elata (Utetheisa), 257, 258, 261, 266, 268, 270,
271.
elatus (Chrysolampis), 73.
elgonensis (Ctenophthalmus), 114,
453
Eliomys, 106.
elongata (Maldane), 354.
— (Syllis), 383.
Elzerina, 329.
Emma, 321.
engis (Temnora), 132.
Enodice, 314.
ephemera (Cercaria), 45, 75-79.
Epifregata, 176.
Epipelicanus, 176.
Episbates, 176,
Epitedia, 124.
epops (Upupa), 70.
erithacus (Psittacus), 70.
erlangeri (Gazella), 185.
erythroura (Gerbillus), 193.
— (Meriones), 193.
Erythrura, 6, 72.
eskeyi (Hectopsylla), 292, 293.
Esthiopterella, 65, 68, 176.
Esthiopterum, 176, 306, 307.
Estrilda, 71, 314.
Eteone, 345, 346.
Euchelia, 269, 273.
Eudyptes, 66, 134, 135.
Eulalia, 346.
eumelas (Mimeusemia), 249.
Eunice, 345, 347, 348, 351, 398.
Eupetomena, 73.
Euplectes, 71, 314.
Euprepocnemis, 378, 379, 382.
europaea (Talpa), 105, 110.
eurygaster (Goniodes), 307.
Eustrigiphilus, 307.
euteles (Macrostylophora), 367.
evansi (Rhinotropis), 424.
evelynae (Meriones), 193, 194.
eversum (Macravestibulum), 78.
Evotomys, 103, 105, 107, 108, 119.
exclamationis (Hippotion), 130, 131.
exilis (Caulibugula), 342.
— (Monopsyllus), 120-122,
— (Syllis), 386.
Exitrichia, 232-235.
expansa (Bugula), 321, 329, 330, 338.
extenuata (Lagisca), 345, 346.
Eyprepia, 274.
f-album (Polygonia), 125.
faceta (Neopsylla), 124.
falcata (Anas), 66.
Falco, 67, 310.
Falcolipeurus, 308.
falklandica (Cathartes), 309.
famulus (Dipodillus), 194, 195.
farinae (Tyroglyphus), 308.
454
farinosus (Phloeobiopsis), 438.
fasciata (Amadina), 314.
— (Syllis), 387.
— (Thecla), 356.
— (Utetheisa), 273.
fasciatus (Gulamentus), 145.
— (Nosopsyllus), 107.
fatela (Utetheisa), 270.
fatua (Deilemera), 270.
— (Utetheisa), 270.
favosus (Typhloceras), 111.
felineus (Opisthorchis), 148, 152, 155, 157, 159,
170.
Felis, 197, 362.
felis (Ctenocephalides), 116, 362.
Fennecus, 199.
ferrugineus (Herpestes), 198.
ferulis (Hypoaspis), 308.
festai (Platynereis), 401.
fimbriatus (Macrostylophora), 365, 367, 368.
fisheri (Agapornis), 312.
flava (Deiopeia), 274.
— (Eteone), 345.
— (Utetheisa, 288.
flavescens (Colpocephalum), 176.
— (Vulpes), 200.
flavirostris (Lophoceros), 313.
flaviventris (Rattus), 192.
florentiae (Jaculus), 191.
floridensis (Cercaria), 151, 152, 155, 157.
Flustra, 327.
Flustrella, 329.
oli (Leptonereis), 396.
formosa (Anas), 66.
— (Euchelia), 269.
— (Vanadis), 348.
Foxella, 123, 124.
fragilis (Lumbrinereis), 345, 346.
franciscana (Euplectes), 71, 314.
franciscanus (Foxella), 123.
Fringilla, 315.
fringillae (Ricinus), 307.
frontalis (Acrobates), 58-60.
frugivorus (Rattus), 192.
fulvoculata (Cercaria), 77.
Funambulus, 373.
furcatoides (Thalurania), 315.
fusca (Degeeriella), 62, 64, 67, 307.
fuscipes (Neotoma), 317.
fuscosuturata (Syllis), 386.
fuscus (Ornithobius), 64, 66.
fusiformis (Owenia), 346.
galapagensis (Utetheisa), 251, 259, 261, 268.
Galea, 297.
galeanus (Parapsyllus), 294.
galericulata (Dendronessa), 67.
gallapagensis (Pseudonereis), 402.
Gallicolumba, 69.
gallicus (Circaetus), 310.
gallinae (Ceratophyllus), 63, 67.
— (Menopon), 306.
Gallus, 311.
Garrulax, 314.
Garrulus, 71.
garzetta (Egretta), 309.
Gattyana, 345, 346.
gattorugine (Blennius), 179.
Gazella, 184-189.
gemina (Hectopsylla), 292.
genetta (Hesione), 392.
Gennaeus, 67.
gentilis (Proscopus), 427.
Geomys, 123.
Geopelia, 312.
Geophaps, 69.
Gerbillus, 193, 195.
gerhardi (Strymon), 359.
gigantea (Pionosyllis), 389.
— (Ratufa), 364.
gisserana (Nereis), 394.
glabra (Bicellaria), 339.
— (Caulibugula), 340.
— (Stirparia), 339.
glandarii (Brüelia), 64, 71.
glandarius (Garrulus), 71.
glandulosa (Ailanthus), 130.
glareolus (Evotomys), 103, 105, 107, 108.
globulariae (Procris), 212-216.
Glycera, 347, 348.
Glyciphagus, 308.
Gnathosyllis, 384.
Gobius, 157, 179.
goesi (Ampharete), 346.
goldmani (Panthera), 420-422,
Goniocotes, 64, 67, 305, 307.
Goniodes, 307.
gracilis (Syllis), 348, 383.
Gracula, 72.
grallina (Plesiobasis), 430.
grandidieri (Temnora), 131.
grandis (Niltava), 70.
— (Typhlopsylla), 124.
granulata (Pectinaria), 346.
granulatus (Spirorbis, 346.
Graomys, 294.
griseoflavus (Graomys), 294.
groenlandica (Phyllodoce), 346.
gruis (Esthiopterella), 65, 68.
— (Esthiopterum), 306, 307.
grunnellus (Pholis), 179.
Guira, 69.
guira (Guira), 69.
Gulamentus, 143-146, 437.
guttatus (Gulamentus), 145.
habessinica (Procavia), 182, 183.
hadramauticus (Canis), 201.
haedinus (Taphozous), 204.
haematodus (Trichoglossus), 69.
hageni (Rhyacopsyche), 218.
Haliaétus, 310.
Haliastur, 310.
Halipegus, 49, 54.
Halipeurus, 176.
Hamacreadium, 376.
hamatus (Paraceras), 362.
hamifer (Leptopsylla), 319, 320.
hamiltoni (Gennaeus), 67.
hansali (Ludia), 132.
haplogonus (Goniocotes), 307.
Haplorchis, 155.
Harmothoé, 345-350.
harterti (Thisoicetrus), 377, 380.
hastatus (Macrostylophora), 366-368.
Hectopsylla, 292, 303.
hedya (Mimeusemia), 248.
heeri (Myriochele), 346.
heinrichi (Macrostylophora), 368, 371, 373.
Heleonomus, 307.
helgolandica (Tomopteris), 348.
helleri (Perinereis), 399.
helvetica (Cercaria), 75, 78.
Hemerobius, 304.
hemprichi (Otonycteris), 204.
hermaphroditus (Paradoxurus), 368.
Hermodice, 347.
hernandesii (Panthera), 419-422.
Herpestes, 198.
Hesione, 392.
heterodonta (Perinereis), 399.
Heterophasia, 71.
Heterophyes, 152.
heterophyes (Heterophyes), 152.
heterotypus (Lagopoecus), 307.
himalayensis (Oxylipeurus), 64, 67.
hipposideros (Rhinolophus), 204.
Hippotion, 130, 131.
hircus (Capra), 184.
hispanicus (Ctenophthalmus), 108.
hollandicus (Leptolophus), 69.
homericus (Acomys), 191, 192.
hospes (Degeeriella), 64, 68.
hubbardi (Phaneris), 318.
hubrechti (Lagisca), 347.
Hucus, 141.
humulinus (Hemerobius), 304.
hyalina (Syllis), 384.
Hyalinoecia, 348.
hyalocauda (Cercaria), 78.
Hydrobia, 75, 79-81.
Hydroptila, 217, 230.
hyemalis (Junco), 315.
Hypoaspis, 308.
Hypochaera, 72.
hypomelas (Paraechinus), 203.
hyrcanica (Strymon), 360.
icana (Thecla), 358.
Ichneumia, 198.
Icterus, 72.
ignotus (Foxella), 123.
ikhwanius (Pipistrellus), 203.
illinoisensis (Geomys), 123.
imbricata (Cercaria), 77, 81.
— (Harmothoé), 345.
impejanus (Lophophorus), 310.
inkermanni (Cercaria), 77.
indica (Chalcophaps), 69.
— (Kittacincla), 70, 313.
indicae (Cercaria), 78, 155, 170.
indicus (Leonnates), 404.
indus (Haliastur), 310.
inflatum (Scalibregma), 345.
insignis (Litocerus), 437.
intermedia (Tritopsylla), 164, 169.
intermedius (Metorchis), 155.
irritans (Pulex), 116.
isacanthus (Rhadinopsylla), 110.
islandica (Bucephala), 309.
ismahelis (Meriones), 193, 194.
jacksoni (Nereis), 394.
— (Procavia), 183.
Jaculus, 191.
jaculus (Jaculus), 191.
— (Mus), 191.
j-album (Polygonia), 125.
jankowskii (Thecla), 356.
japonica (Thecla), 356.
javanicus (Herpestes), 198.
jayakari (Procavia), 182, 183.
jejuna (Cryptocotyle), 102, 151, 155, 158, 179.
jobiensis (Gallicolumba), 69.
Johnstonia, 352.
Joubertia, 308.
joubini (Harmothoé), 345, 348, 349.
jousseaumei (Leonnates), 403, 404.
jubata (Neochen), 309.
Junco, 315.
Kakatoe, 313.
kalisi (Polyptychus), 126-129.
kallima (Utetheisa), 254, 262, 263, 272-276.
kefersteini (Tomopteris), 347.
kelanus (Polyptychus), 126, 127.
kelloggi (Colpocephalum), 306.
kinbergii (Eunice), 348.
Kittacincla, 70, 313.
456
kohli (Gulamentus), 146.
konadensis (Cercaria), 78.
koreana (Thecla), 356.
kowalewskii (Sagitella), 347, 348.
krohnii (Syllis), 387.
kuhli (Pipistrellus), 203.
Kurodaia, 306.
Labrus, 179.
lactea"(Deiopeia), 271.
— (Utetheisa), 257, 258, 261-263, 266, 267, 269,
271, 272, 277.
lacteus (Bubo), 313.
ladislavii (Diaulus), 218.
Laelaps, 63, 65, 68.
laenatus (Acorynus), 140.
laetus (Gulamentus), 145, 437.
lafayetii (Gallus), 311.
Lagisca, 345-347.
Lagopoecus, 307.
lanceolata (Travisiopsis), 348.
laqueator (Cercaria), 54.
Larus, 68, 312.
lebouri (Cercaria), 78, 81.
leclancheri (Passerina), 315.
ledereri (Strymon), 360.
leechi (Thecla), 355.
Leocrates, 348, 392.
Leonnates, 403.
lepida (Deiopeia), 286.
— (Utetheisa), 253, 256, 276, 278, 286-288.
leptoceros (Gazella), 185.
Leptolophus, 69.
Leptonereis, 396.
Leptopsylla, 319.
Lepus, 189, 190, 201.
leucoblephara (Galea), 297, 300, 303.
leucocephalus (Haliaétus), 310.
leucomels (Lophoceros), 313.
Leucophaser, 72.
leucopus (Vulpes), 199.
leucotis (Turacus), 70.
leveri (Plintheria), 440.
levinseni (Travisiopsis), 346.
levis (Macrostylophora), 367, 372.
lidthi (Garrulus), 71.
ligulata (Tomopteris), 348.
lilloi (Maxilliopsylla), 303.
limbata (Brüelia), 64, 72.
— (Mimeusemia), 248.
Limnaea, 241.
limodes (Tribotropis), 423.
lingua (Cryptocotyle), 148, 151, 152, 155, 156,
178-180.
Lioptila, 315.
Liotheum, 176.
Liothrix, 315.
Lipeurus, 62, 64, 67, 176.
Liponycteris, 204.
Liponyssus, 63, 65, 70, 72, 308.
Lithosia, 273.
Litocerus, 140, 141, 437.
littoralis (Gryllus), 381.
— (Littorina), 156.
— (Thisoicetrus), 378-381.
littorea (Littorina), 85, 87, 156.
Littorina, 78, 85, 87, 156.
lixa (Dipodillus), 195.
loderi (Gazella), 185.
loftusi (Jaculus), 191.
Loimia, 347, 348.
longa (Eteone), 346.
londiniensis (Nosopsyllus), 117, 118.
longicirrata (Opisthosyllis), 389.
longicornis (Parapsyllus), 134-138.
longifrons (Meriones), 193.
longiloba (Leptopsylla), 319.
longipinnis (Glycera), 347.
Lopadorhynchus, 347, 348.
lophocerca (Cercaria), 155, 176.
Lophoceros, 313.
Lophophorus, 310.
lophotes (Oxyphaps), 306, 312.
Lophura, 310.
Lophuromys, 114.
lotrix (Phalaena), 286-288.
— (Utetheisa), 252-286, 288-291.
Loxia, 72.
Loxotrichia, 238.
lucania (Cercaria), 78.
luctuosa (Delias), 130.
— (Plintheria), 440.
Ludia, 132, 133.
ludlowi (Paraechinus), 202.
lugens (Rallicola), 68, 307.
lugubris (Rhopalopsyllus), 443, 446, 448.
— (Rothschildella), 443.
lujai (Gulamentus), 145.
Lumbrinereis, 345, 346.
lunula (Protolichus), 308.
lupaster (Canis), 201.
lupatus (Macrostylophora), 367, 373.
lupus (Canis), 201.
lutea (Liothryx), 315.
— (Thecla), 357.
luteatus (Polyptychus), 126, 127.
lutzi (Pulex), 443.
— (Rhopalopsyllus), 443-448.
lybica (Felis), 197.
Macellicephala, 348.
maclachlani (Peltopsyche), 218.
macracanthus (Paraechinus), 203.
Macravestibulum, 78.
Macrostylophora, 365.
macroura (Eupetomena), 73.
macrura (Cercaria), 45.
maculata (Pista), 345, 346.
madagarensis (Margaroperdix), 68.
madeirae (Panthera), 410, 412, 413, 414, 415.
magellanica (Beania), 329.
magellanicus (Spheniscus), 134-137.
— (Parapsyllus), 135-138.
magna (Onuphis), 348.
magnifica (Pionosyllis), 389.
magniovatus (Notocotylus), 78.
major (Gulamentus), 144.
— (Panthera), 410, 413, 414.
malabarica (Enodice), 314.
— (Kittacincla), 313.
malacca (Munia), 72.
Malaraeus, 119, 120.
Maldane, 354.
Mangusta, 198.
maniculata (Felis), 197.
maracana (Ara), 313.
Mareca, 67, 309.
Margaroperdix, 68.
marginalis (Falcolipeurus), 308.
marica (Gazella), 184, 185, 188.
marina (Arenicola), 346.
Maritrema, 95.
Marmosa, 168.
marquesensis (Pionosyllis), 387.
marshallarum (Utetheisa), 282.
marshallorum (Utetheisa), 278, 282, 283.
marsupialis (Didelphis), 166.
Martes, 364.
masalacensis (Pseudonereis), 402.
Mauia, 146.
Maxilliopsylla, 303.
mazzai (Hectopsylla), 303.
Mecocerus, 424.
medus (Nosopsyllus), 117.
Megabothris, 120, 122.
Megninia, 65, 71.
melampyga (Utetheisa), 273.
Melanopsacus, 431.
melanostigma (Sabella), 347.
melanota (Sarkidiornis), 309.
meleagridis (Goniodes), 62, 64, 67.
melinus (Gulamentus, 145
— (Paraceras), 362, 364, 365.
Melopsittacus, 63, 312.
melpoda (Estrilda), 71, 314.
Menacanthus, 63, 306.
mengesi (Capra), 184.
Menipea, 327.
Menopon, 306.
Meriones, 192-194,
mesa (Rhadinopsylla), 110.
Metagonemus, 155.
Metopiana, 67, 309.
Metorchis, 155.
mexianae (Panthera), 410, 412, 415.
mexicana (Sialia), 313.
Mexitrichia, 222.
medusa (Loimia), 347, 348.
micans (Rapala), 361.
— (Thecla), 361.
Microcavia, 293, 294.
Microcreadium, 376.
microphylla (Joubertia), 308.
Mierotus, 106, 108, 119, 320.
milleri (Panthera), 414, 416, 417.
Mimeusemia, 248.
mimulus (Dipodillus), 195.
mincopicus (Polyptychus), 126, 127.
minima (Bugula), 332, 334-337.
minimus (Rhinolophus), 204.
minuta (Thisoecetrus), 382.
minutus (Nirmus), 175.
— (Thisoicetrus), 378, 382.
mirabilis (Ceratonereis), 400.
— (Macellicephala), 348.
misakiensis (Tripanosyllis), 391.
Misthosima, 147.
Mitu, 310.
mitu (Mitu), 310.
modicus (Ctenophthalmus), 114.
moebii (Callizona), 347.
molops (Sintor), 142.
monera (Plesiobasis), 427, 429, 430.
mongolicus (Phasianus), 67.
monostomi (Cercaria), 77.
Monopsyllus, 120-122.
monroei (Ludia), 132.
moorei (Hippotion), 130.
Mortoniella, 217-220.
mosonica (Nessiara), 142.
Munia, 72, 314.
murex (Phascogale), 58.
Murexia, 58.
murina (Crocidura), 202.
— (Marmosa), 168.
Mus, 107, 108, 191.
muscatansis (Gazella), 184, 185.
musculus (Mus), 107, 108.
mustela (Onos), 179.
muticus (Pavo), 310.
Myriochele, 346.
Myrsidea, 63, 73, 306.
Nainereis, 346.
nakagawai (Cercaria), 78.
nanus (Gulamentus), 146.
neglecta (Potamilla), 346.
Neochasmus, 159.
Neochen, 309.
457
458
Neopsylla, 124, 373. occidualis (Halipegus), 49, 54.
Neotoma, 317. ocellaris (Daphnusa), 130.
nephelus (Gulamentus), 437. ocreata (Felis), 197.
Nephthys, 346. olii (Dipodomys), 121.
Nereis, 345-347, 386, 394-396. omanensis (Lepus), 190.
nerii (Deilephila), 261. onca (Panthera), 406-422.
neritina (Bugula), 329, 330, 334, 336. oniscus (Paraechinus), 202.
neritinoides (Bugula), 329, 330, 334. Onos, 179.
neritoides (Littorina), 78. Onuphis, 345, 346, 348.
Nessiara, 142, onychius (Parapsyllus), 297.
nicobarica (Cercaria), 155. Onychomys, 121, 122.
nierstraszi (Leonnates), 404. opadus (Monopsyllus), 121.
niger (Paraschinus), 203. Opisodasys, 316.
nigeriensis (Stivalius), 116. Opisthorchis, 152, 155.
nigricauda (Taterona), 196. Opisthosyllis, 347, 348, 389-391.
nigrosignata (Utetheisa), 271, 272. oralis (Jaculus), 191.
nilotica (Vulpes), 199-201. orana (Procris), 433.
Niltava, 70. Orchopeas, 317.
Nirmus, 175. orectus (Megabothris), 122.
nissa (Thecla), 361. Oreortyx, 67.
nisseni (Tomopteris), 347, 348. orientalis (Thecla), 355.
nisus (Accipiter), 67. orientis (Ctenocephalides), 362.
nivalis (Ctenophthalmus), 108. orinoptena (Ludia), 133.
niveicapilla (Cossypha), 313. ormina (Mexitrichia), 222.
nobilis (Ctenophthalmus), 104, 105, 108, 109. ornatrix (Utetheisa), 251, 255, 260, 263, 266,
— (Flustra), 327. 268.
Noctua, 273. Ornithobius, 64, 66.
noctua (Athene), 70. Oryzivora, 314.
norwegicus (Rattus), 113, 116, 117. oryzivora (Oryzivora), 314.
Nosopsyllus, 107, 116-118. — (Paddo), 72.
notata (Cyrtacanthacris), 381. osafunei (Cercaria), 78.
— (Procris), 213. Otilipeurus, 68, 307.
noteuna (Exitrichia), 232. Otis, 68, 311.
notialis (Panthera), 416, 417. Otonycteris, 204.
Notiana, 424, Otyphantes, 71.
Notocotylus, 75, 77, 78. ouvrardi (Thecla), 358.
novaehollandiae (Cereopsis), 309. ovona (Exitrichia), 233.
novae-hollandiae (Dromiceius), 65. Owenia, 346.
— (Larus), 68. Oxylipeurus, 64, 67.
novara (Exitrichia), 232. Oxyphaps, 306, 312.
nubiana (Capra), 184. oxyurus (Thisoicetrus), 382.
nuchalis (Opisthosyllis), 347, 390.
nudiventris (Liponycteris), 204.
numatus (Hucus), 140. ; pachychaeta (Ceratonereis), 397, 400.
numenii (Degeeriella), 307. Pachydota, 255.
Numenius, 311. Paddo, 72.
nuntia (Perinereis), 399, 400. pagodorum (Temenuchus), 314.
Nyctea, 313. Palaeopsylla, 374.
nyctea (Nyctea), 313. palaestina (Vulpes), 200, 201.
Nycteris, 204. pallida (Utetheisa), 273.
Nyroca, 309. pallipes (Canis), 201.
nyroca (Nyroca), 309. — (Herpestes), 198.
— (Mangusta), 198.
palmaris (Doticus), 431, 432.
obfuscata (Perinereis), 396, 398, 402. palpalis (Tiamastus), 444, 445.
obtusicaudum (Macravestibulum), 78. palustris (Panthera), 411, 415-417.
occidentalis (Neotoma), 317. Panacra, 131.
occidentis (Citellophilus), 106. Panthera, 406.
Papilio, 265.
papualis (Pygoscelis), 135.
papuana (Utetheisa), 278, 283-285.
Paraceras, 362.
paradisea (Anthropoides), 311.
— (Steganura), 314.
— (Tetrapteryx), 68.
Paradoxurus, 362, 368.
paradoxus (Paradoxurus), 362.
Paraechinus, 202, 203.
paraguensis (Panthera), 416, 417.
Parapleurolophocerca, 148, 152, 154, 170-173.
Parapsyllus, 134-138, 294-302.
pardalis (Gulamentus), 145.
Paroaria, 64, 73, 315.
parviceps (Goniodes), 307.
parvomelaniae (Cercaria), 155.
parvum (Microcreadium), 376.
Passerina, 315.
paulensis (Panthera), 416, 417.
Pavo, 67, 310.
pavonia (Balearica), 311.
pavonis (Goniodes), 64, 67, 307.
Pectinaria, 346.
pectinata (Utetheisa), 262, 277-281, 285.
pelagica (Nereis), 346.
pelerinus (Meriones), 193.
pellucida (Cercaria), 77.
Peltopsyche, 218.
pendlburyi (Paraceras), 365.
penelope (Mareca), 309.
penicilliger (Malaraeus), 120.
peposaca (Metopiana), 67, 309.
percomis (Strymon), 360.
perforatus (Taphozous), 204.
Perinaeus, 176.
Perinereis, 396-402.
Peringia, 42, 44, 75, 78-89, 84-88, 91-96, 101,
240-249, 376.
perla (Hemerobius), 304.
perotis (Pronops), 303.
persa (Thisoicetrus), 378, 380.
persica (Strymon), 359.
persicus (Herpestes), 198.
persimilis (Strymon), 360.
perspicillata (Perinereis), 399.
peruviana (Panthera), 414, 417-420.
Pezotettix, 379.
phaeura (Apatenia), 440.
Phalacrocorax, 308.
Phalaena, 272, 286.
Phaneris, 317.
Phaps, 69.
Phascogale, 58.
Phasianus, 67.
phelos (Plesiobasis), 429.
Philacte, 309.
philbyi (Tatera), 196.
459
Philichthyophaga, 176.
philippinensis (Polyptychus), 126-129.
phillipsi (Macrostylophora), 367, 372.
Philopterus, 64, 70, 72, 73.
Phlaeops, 424.
Phloeobiopsis, 438.
Phloeops, 424.
Phoenicopterus, 66, 309.
phoenicurus (Amaurornis), 311.
Pholis, 179.
pholis (Blennius), 179.
phygus (Litocerus), 437.
Phyllodoce, 346.
Physconelloides, 64, 69.
picaoides (Heterophasia), 71.
picatus (Gulamentus), 146.
picta (Oreortyx), 67.
picteti (Nereis), 397.
pictus (Polyophthalmus), 347.
pilatus (Macrostylophora), 366-368.
Pionosyllis, 387-389.
Pipistrellus, 203.
piquenardi (Thecla), 358.
Pista, 345-347.
Pitta, 70.
Pitymys, 108.
plagifer (Phloeobiopsis), 438, 439.
plana (Cercaria), 77.
planirostris (Litocerus), 141.
planktonis (Tomopteris), 347.
platensis (Rhopalopsyllus), 292, 293.
Platycercus, 69.
Platynereis, 401.
platypennis (Phloeops), 424.
platystomus (Craspedorrhynchus), 307.
plaumanni (Acostatrichia), 228-230.
Plesiobasis, 427-430.
Pleurolophocerca, 148, 152-155, 170, 172.
pleurophocerca (Cercaria), 155.
Plintheria, 440.
plumosa (Stylarioides), 346.
Pluvianus, 311.
Podiceps, 66.
poecilops (Dipodillus), 194, 195.
Poephila, 71.
poliocephalus (Porphyrio), 68, 311.
Polygenis, 444-448.
Polygonia, 125.
Polyophthalmus, 347.
Polyptychus, 126-129.
poppei (Typhloceras), 110, 111.
Porphyrio, 68, 311.
Potamilla, 346.
probatus (Macrostylophora), 366, 367.
Procavia, 182, 183.
Procellaria, 134, 137.
Procris, 212-216, 433.
producta (Hydroptila), 217, 236.
460
projecta (Cercaria), 49.
Pronops, 303.
Proscopus, 426, 441, 442.
Protolichus, 308.
Protomolis, 255.
Protoptila, 221.
provincialis (Ctenophthalmus), 104.
prunoides (Strymon), 360.
Psammomys, 192.
Pseudonereis, 402.
Pseudonirmus, 176.
Psittacus, 70.
psittacea (Erythrura), 72.
psomus (Stenorhis), 431.
Pterolichus, 65, 68-71.
pulchella (Bombyx), 273.
— (Deiopeia), 273, 274, 282, 287.
— (Deiopia), 273.
— (Dejopeja), 274.
— (Lithosia), 273.
— (Phalaena), 272.
— (Platynereis), 401.
— (Utetheisa), 252-291.
— (Tinea), 272.
pulchelloides (Utetheisa), 252-286,
pulchellus (Acrobates), 58-60.
pulchra (Deiopeia), 273.
— (Euchelia), 273.
— (Eyprepia), 273.
— (Noctua), 273.
— (Utetheisa), 273,
Pulex, 116, 443-445.
pulex (Rhynchopsylla), 303.
punctata (Degeeriella), 307.
pusilla (Vulpes), 199, 200.
putris (Succinea), 84.
Pygidiopsis, 155, 159.
pygmaeus (Acrobates), 58-60.
Pygoscelis, 135.
pyrenaicus (Pitymys), 108.
quadricuspida (Nainereis), 346.
quercinus (Eliomys), 106.
quercus (Thecla), 356.
querquedulae (Trinoton), 63, 306.
Quiscalus, 72.
quiscula (Quiscalus), 72.
Rallicola, 68, 307.
ramosa (Bugula), 334.
ramsayi (Panthera), 416.
rantae (Thisoicetrus), 377, 379.
Rapala, 361.
Ratufa, 364.
Rattus, 113, 114, 116, 117, 192, 373.
rattus (Rattus), 113, 114, 192.
rayata (Synthusa), 360.
rebeli (Hippotion), 131.
recula (Foxella), 123, 124.
Recurvirostra, 68.
regularis (Beania), 329.
reichenowi (Otyphantes), 71.
religiosa (Gracula), 72.
remota (Palaeopsylla), 374.
repens (Argyrolepidia), 249.
Rhadinopsylla, 110.
Rhea, 308.
rheae (Struthiolipeurus), 308.
Rhinolophus, 204.
Rhinopoma, 204.
Rhinotropis, 424, 425.
Rhodogastria, 250, 255.
Rhopalopsyllus, 292, 293, 443-448,
Rhyacopsyche, 218.
Rhynchopsylla, 303.
Richmondena, 72, 315.
Ricinus, 62, 64, 73, 307.
ridibundus (Larus), 312.
ridibundum (Menopon), 306.
roberti (Polygenis), 444, 448.
— (Pulex), 444.
Robertianella, 348,
robertsianus (Parapsyllus), 137, 138
robur (Hippotion), 130.
robusta (Bugula), 329-337.
— (Cercaria), 78, 81.
— (Tatera), 196.
robustus (Opisodasys), 316.
rolandi (Typhloceras), 110.
rosaurae (Eunice), 348, 351.
— (Harmothoé), 348, 349.
roseicapilla (Kakatoe), 313.
roseicollis (Agapornis), 69.
Rossicotrema, 152.
Rothschildella, 443.
rothschildi (Leucophaser), 72.
— (Phascogale), 58.
rotunda (Emma), 322-326.
Rousettus, 374.
ruberrima (Utetheisa), 277, 278, 280.
rubra (Utetheisa), 289, 290.
rubritorquis (Trichoglossus), 69.
rubrosignata (Utetheisa), 271, 272.
rudicollis (Proscopus), 426.
rufa (Aplodontia), 319.
— (Degeeriella), 64, 67.
— (Lophura), 310.
ruficeps (Procavia), 182.
rupestris (Ctenolabris), 179.
ruppelli (Vulpes), 201.
russatus (Acomys), 191.
— (Mus), 191.
russula (Crocidura), 108.
ruthensparri (Gobius), 157, 160, 179.
sabaea (Vulpes), 199, 201.
sabaeus (Paraechinus), 203,
Sabella, 347.
sachalinensis (Thecla), 356.
Saemundssonia, 307.
saevus (Rhopalopsyllus), 448.
Sagitella, 347, 348.
sagittarius (Cercaria), 42, 44, 46, 48, 50, 52,
53.
salomonis (Apatenia), 440.
— (Utetheisa), 254, 256, 263-266, 268, 273,
277, 279.
sanctaehelenae (Estrilda), 71, 314.
Sarkidiornis, 309.
Satsuma, 359.
satyra (Tragopan), 67.
saudiya (Gazella), 184-188.
Scalibregma, 345.
scaphoides (Bugula), 329, 330, 333, 334.
Scardafella, 69.
Sciurus, 317, 368.
Scoloplos, 345.
seineti (Notocotylus), 77.
sellatus (Gulamentus), 146.
— (Nirmus), 175.
selliger (Nirmus), 175.
semara (Utetheisa), 257, 259, 260, 261, 263, 266-
268, 290.
semisignata (Utetheisa), 273.
semperiana (Nereis), 396.
seniculus (Paraechinus), 203.
sentus (Doticus), 431.
septentrionalis (Tomopteris), 346.
setosa (Callizona), 348.
— (Echinometra), 304.
setschuanica (Thecla), 358.
sexdentatus (Orchopeas), 317.
shoana (Procavia), 183.
Sialia, 313.
sibilatrix (Mareca), 67.
sieboldi (Peltopsyche), 218.
signatus (Gulamentus), 143, 145.
sikkimensis (Macrostylophora), 367, 368.
similis (Caloptenus), 381.
— (Thisoicetrus), 378, 381, 382.
simplex (Citellophilus), 106.
— (Leonnates), 403.
— (Mimeusemia), 249.
simulans (Acostatrichia), 229.
— (Hydroptila), 217.
sinaitica (Capra), 184.
sinensis (Clonorchis), 148, 155.
sinitzini (Cercaria), 42, 43, 48, 52, 54-56.
Sinthusa, 360.
Sintor, 142.
smaragdina (Thecla), 356.
Smerinthus, 130.
smithi (Geophaps), 69.
socotrensis (Utetheisa), 287, 288.
sonantis (Polyptychus), 126, 127, 129,
souleana (Thecla), 356, 357.
sparsa (Hydroptila), 236.
spatula (Cercaria), 78.
Spirorbis, 346.
Spheniscus, 65, 134, 135, 137.
spongicola (Syllis), 347, 384.
sponsa (Aix), 66, 309.
Spreo, 72, 121.
squammata (Scardafella), 69.
squamosa (Abtrichia), 217, 226, 228.
Squatarola, 68.
squatarola (Squatarola), 68.
Stamnosoma, 155.
statices (Procris), 213, 433.
Steganura, 314.
Stenopsylla, 164, 165.
Stenorhis, 431.
Stephanophorus, 71.
stevensi (Argyrolepidia), 133.
stigmata (Utetheisa), 254, 256, 289.
stilbus (Melanopsacus), 431.
Stirparia, 339.
Stirpariella, 339.
Stivalius, 115, 116, 373.
stomis (Hectopsylla), 292, 293.
striata (Munia), 314.
striolata (Perinereis), 398.
stroemi (Terebellides), 346.
strutheus (Austrogonioides), 64, 66.
Struthiolipeurus, 308.
Strymon, 359, 360.
Sturnus, 314.
Stylarioides, 346.
stylifera (Pionosyllis), 388.
subflava (Estrilda), 71, 314.
subglobosa (Bugula), 329, 330, 332.
subnotus (Mauia), 146.
subsellatus (Lipeurus), 64, 67.
subsignatum (Anaticola), 307.
subtilis (Tiamastus), 445.
subtuta (Misthosima), 147.
Succinea, 84.
suffusa (Thecla), 355.
sumatrana (Utetheisa), 252, 266, 268, 289.
sumatrensis (Cricula), 433.
summus (Pygidiopsis), 155.
Suncus, 202.
suturalis (Sintor), 142.
Syllis, 347, 348, 383-387.
sylvarum (Liponyssus), 308.
sylvaticus (Apodemus), 103, 105,
110.
synophthalma (Robertianella), 348.
syriaca (Felis), 197.
— (Procavia), 182, 183.
syrius (Meriones), 193.
107,
461
108,
462
Tadorna, 309.
tadorna (Tadorna), 309.
taeniatus (Gulamentus), 145.
Taeniopygia, 314.
taeniura (Tatera), 196.
taichui (Haplorchis), 155.
taihokui (Haplorchis), 155.
Talpa, 105, 108, 110.
Tamiophila, 124.
Tamiops, 370.
Taphozous, 204.
tarda (Otis), 68, 311.
Tatera, 182, 196.
Taterona, 196.
tatsienluica (Thecla), 357.
taxila (Thecla), 356.
tectonis (Charaxes), 130.
Temenuchus, 314.
Temnora, 131, 132.
tenuata (Bugula), 330, 332.
tenuella (Utetheisa), 288.
tenuicornis (Procris), 213-216.
tephras (Gulamentus), 146.
Terebellides, 346.
terrae-sanctae (Psammomys), 192.
tertia (Exitrichia), 235.
testor (Neopsylla), 124.
testudinarius (Cummingsiella), 307.
Tetrapteryx, 68.
teutona (Maxitrichia), 222, 223.
thalia (Papilio), 304.
Thalurania, 315.
Thaumapsylla, 374.
thebaica (Nycteris), 204.
Thecla, 355-358.
Thelepus, 345, 346.
theodori (Thisoicetrus), 378, 381.
Thisoecetrus, 379-382.
Thisoicetrus, 377-382.
thyter (Deiopeia), 273.
Tiamastus, 444, 445, 447, 448,
Tiaris, 72.
timanus (Stivalius), 115, 116.
timius (Litocerus), 141.
tinnuneulus (Falco), 67.
tonkinensis (Macrostylophora), 368.
Tomeutes, 368, 370, 373.
Tomopteris, 346-348.
tomoyus (Rhopalopsyllus), 444.
torvus (Stivalius), 116.
trabeculata (Cercaria), 77.
Tragopan, 9, 67.
translucens (Cercaria), 155.
Travisiopsis, 346, 348.
triangula (Emma), 322, 323.
Tribotropis, 423.
tricellata (Emma), 322, 326, 327.
Trichoglossus, 69.
trieirratus (Onos), 179.
trieolor (Gulamentus), 145.
tridens (Asellia), 203.
trifasciata (Nereis), 347.
trifenestrata (Cricula), 433-436.
trilineatus (Polyptychus), 126-129.
Trinoton, 63, 66, 306.
triophthalmia (Cercaria), 78.
triptus (Monopsyllus), 122.
tripus (Tiamastus), 447.
trithorax (Briielia), 64, 73.
Tritopsylla, 164, 166, 169.
troglodytes (Estrilda), 71.
Tropicranus, 70, 313.
Tropideres, 140.
Trypanosyllis, 385, 391.
tuberosa (Caulibugula), 340, 342.
tubicola (Hyalinoecia), 348.
tumidum (Menopon), 306.
Tupaya, 373. _
tupinus (Rhopalopsyllus), 448.
Turacus, 70.
turatii (Procris), 216.
turmalis (Otilipeurus), 64, 68, 307.
Turritella, 84.
Turtur, 312.
Tylotrepidius, 382.
Tympanistria, 312.
tympanistria (Tympanistria), 312.
Typhloceras, 110.
Typhlopsylla, 124.
Tyria, 255.
Tyroglyphus, 308.
tyto (Spreo), 121.
ucayalae (Panthera), 410, 413, 415.
ultramarina (Thecla), 356, 357.
ulvae (Peringia), 42, 44, 75-96, 240-247,
376.
umata (Utetheisa), 279, 281, 284.
uncinalis (Macrostylophora), 370.
uncinatus (Lopadorhynchus), 347, 348.
undatus (Polyptychus), 126-129.
undulatus (Melopsittacus), 69, 312.
unicolor (Aulodina), 439.
— (Thecla), 356.
unifasciata (Nereis), 396.
uniserialis (Bugula), 329.
unota (Mexitrichia), 222-223.
Upupa, 70.
upupae (Degeeriella), 62, 64, 70.
Uraeginthus, 71, 314.
urbanensis (Cercaria), 77.
Uroaétus, 310.
urticae (Aglais), 125.
Utetheisa, 251-291.
vaga (Cercaria), 77.
—(Utetheisa), 279, 284,
vallata (Perinereis), 400.
Vanadis, 348.
vancaurica (Perinereis), 399.
variegata (Pseudonereis), 402.
— (Syllis), 383, 386.
vatia (Nessiara), 142.
vectifera (Bugula), 329, 330.
veitchi (Proscopus), 426.
vellanus (Proscopus), 441, 442.
ventrosa (Hydrobia), 75, 79-81.
venusta (Deiopeia), 269, 270.
— (Utetheisa), 269.
venustus (Apophallus), 155.
veraecrucis (Panthera), 419, 421, 422.
verbanus (Ctenophthalmus), 104.
verruculosa (Syllis), 386.
vexillaris (Psammomys), 192.
vigens (Leptopsylla), 319, 320.
Vipera, 304.
virens (Nereis), 346.
virgata (Leonnates), 404.
virgatus (Gulamentus), 145.
viridis (Eulalia), 346.
— (Opisthosyllis), 390.
virilis (Misthosima), 147.
vitiana (Dinosaphis), 425.
vittata (Eunice), 347.
— (Syllis), 386.
vogeli (Cercaria), 155.
vulgaris (Sturnus), 314.
Vulpes, 182, 189, 199-201.
vulpes (Vulpes), 182, 189, 199, 200.
vulturinum (Acryllium), 67, 311,
watersi (Emma), 322, 326.
weissmannioides (Pionosyllis), 388.
wenmanni (Ctenophthalmus), 124.
woodlarkiana (Plintheria), 440,
Xenopsylla, 112-116,
Xenopsyllus, 115.
yangi (Thecla), 358.
yenchingensis (Cercaria), 78, 81.
yokogawai (Metagonemus), 155.
zamboangae (Sintor), 142,
zanzibariensis (Caulibugula), 339, 340.
zebra (Trypanosyllis), 385, 391.
Zenaida, 69, 312.
Zephyrus, 355.
zilaba (Loxotrichia), 238,
zilbra (Betrichia), 230, 231.
zonata (Gnathosyllis), 384,
zonata (Nereis), 345.
— (Syllis), 384.
zostera (Cercaria), 77.
Zumatrichia, 217, 226,
463
LEPIDOPTERA
COLLECTED BY THE
British Ornithologists’ Union and Wollaston Expeditions in
the Snow Mountains, Southern Dutch New Guinea
WITH TWO COLOURED PLATES
By THE Hon. WALTER ROTHSCHILD, Pu.D.
(LORD ROTHSCHILD)
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A REVISION OF THE LEPIDOPTEROUS FAMILY
SPHINGIDAE
By THE Hon. WALTER ROTHSCHILD, PH.D,,
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