V

MEMOIRS OF

THE CONNECTICUT ACADEMY

OF ARTS AND SCIENCES

Vol. X

YALE NORTH INDIA EXPEDITION

Article I — Report on Diptera of the Family Ephydridae, by Ezra T. Cresson, Jr.

Article II — Report on Triclad Turbellaria from Indian Tibet, by Libbie H. Hyman.

Article III — Report on Coleoptera of the Family Staphylinidac, by Malcohn Cameron.

Article IV — Report on Coleoptera of the Family Carabidae, by H. E. Andrewes.

Article V — Report on Phyllopod Crustacea (Anostraca, Notostraca and Conchostraca)

Including a Revision of the Anostraca of the Indian Empire, by R. M. Bond.

Article VI — Report on Amphipod Crustacea of the Genus Gammarus, by Masuzo Ueno.

Article VII — Report on Hydracarina, by O. Lundblad.

Article VIII — Report on Terrestrial Families of Hemiptera-Heteroptera, by G. Evelyn

Hutchinson.
\rticlk IX — Report on Rotatoria, by W. T. F.dniondson and G. F. Hutchinson.

September, i 934

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ARTICLE I

REPORT ON DIPTERA OF THE FAMILY EPHYDRIDAE

By Ezra T. Cresson, Jr.

The Academy of Natural Sciences of Philadelphia

(Recei\'ed November 3, 1933)

The Ephydridae submitted to me for study were collected around several saline lakes and
hot springs in the Punjab and Indian Tibet, and are represented by thirteen adults of
three species belonging to the genera Ephydra and Halniopofa, of which two are here
described as new. This did not surprise me as little is known of the Ephydrid fauna of the
Tibetan plateau. Becker's contribution on the Diptera of Central Asia^ is the only report to
my knowledge on the species of this family known to occur there. In this report he records
thirty species, describing several new genera and species, many from localities adjacent to the
saline lakes there. Comparatively few species of insects have adapted themselves to such
severe conditions as are presented by these generally alkaline and saline waters which, in
addition, are often quite hot. Larvae of this f unily have been previously collected in waters
with a temperature as high as 43°C. (109°F.).- In the present collection are specimens of
larvae from a hot spring at Phuga which were obtained up to 49.1°C.

It is interesting to note that while expecting to find a group of species confined to such
a habitat, I was not prepared to encounter such extreme similarity as I did in the species of
the genus Ephydra, here descril^ed, and in one occurring in a similar environment in North
America. It required considerable study of all the material of this genus in my hands to
secure satisfactory characters for specific recognition. I could not believe that we had one and
the same species from such widely separated localities.

This collection also contains some larvae and puparia, but as none of the adults were
bred from any of these, I am unable to determine with certainty the species represented.
However, as far as I can determine, these larvae and puparia seem to represent one and the
same species, although they show a certain degree of variation which I do not think is of
specific value. I append some notes on this material at the end of this paper.

Ephydra glauca Meigen

1830. Ephydra glauca Meigen, Syst. Beschr. Europ. Zweifl., VI, p. 120.

1896. Ephydra ohscuripes Becker, Bed. Ent. Zeit., XLI, p. 222. (nee. Loew. 1866.)

1930. Ephydra glauca Cresson, Trans. Am. Ent. Soc, LVI, p. 115.

Indian Tibet: Tso-kar, 14,850 feet altitude, September 5, 1932. 1 ^ , 19.

The water of Tso-kar is very salt, containing 74,832 parts per million non-volatile solids
in solution.

This species was originally described from Europe, without any citation of definite
locality. I have reported it in 1930 from Lac Sarat, Great Wallacia, Roumania; at the

'Ann. Afus. Zool. Acad. Imp. Sci. St. Petersb., XII, pp. 299-306 (1907).
'Brues in the Proc, Am. Acad, A. & Sc, LXIII, p. 202 (1928).

Mem. Conn. Acad., Vol. X, Art. I. September, 1934.

I DIPTERA OF THE FAMILY EPHYDRIDAE

same time recording a male (not female) labelled "Hdiyka, Indus Phal., 1866." These latter
data I am unable to interpret, but assume that the specimen was collected somewhere along
the Indus River. The present series agrees very satisfactorily with my Roumania specimens,
so I have no doubt of their conspecific status. The species is probably well distributed in the
arid regions of Central Asia. As material I have seen of this species agrees so well with
Becker's description of ohscuripes, I have no doubt of this synonymy. Becker's sjiecies was
originally described from Sarepta, Astrakhan, South Russia, and since recorded from Oren-
burg.^ Popova* also recorded obsciirif^cs from Elton Sea, Astrakhan, Smith Russia.

Ephydra tibetensis n. sj)

This species is so similar to several occurring in the liicrmal waters of North America,
that I rather hesitated to describe it as new without studying a larger series of the North
American forms than I have before me. However, considering their widely separated
habitats and the slight differences that are apparent, I venture to descril)e the present one
as new.

Very similar to pectinulata Cresson" of the United States, but of a more greenish glaucous
tone, not whitish, becoming brownish above. Bristles of the frons stronger; the setulae
assuming bristle-like proportions; the metallic medifrons rough. Mesonotum more uniformly
metallic, less pollinose, cupreous rather than aeneous; the abdomen greenish glaucous. Palpi,
all tarsi and wing veins, black. Differs from glaiica in its more olivaceous tone, strong setulae,
as well as in the flatter, more metallic colored, interfoveal area of the face.

Entirely black except the tawny halteres and squamae.

Head height, length and widths as 15: 18: 22; in profile, facial iinijection l)eyon(l facial
orbits, to distance from occiput to facial orbits, as 8: 7. Eyes oblique witli vertical diameter
slightly in excess of one-half height of head. Frons distinctly broader than long, in profile,
slightly oblicjue; mesofrons sulxjuadrate, broad anteriorly, shining metallic blue, surface hnely
sculptured, bearing two to three pairs of converging, proclinate bristles on anterior portion;
the broad .somewhat metallic parafrons, the narrow frontalia and miular tubercle, opaque
dark brown, with very strong orbital setae in addition to the fimr usual l)ristles. l-'ace
opaque, whitish cinerei ins on the prominent setulose niedifacies; the tlattened horizontal inter-
foveal area, shining metallic blue to green; fovcae, jiarafacials and cheeks olivaceous. Facial
bristles well developed, the lateral ones longest; oral margin slightly retracted, with very long
cilia which are almost as long as the ocellars. Cheeks two-fifths height of head; buccal
bristle slender; lower three to four setae of the postorbital cilia bristle-like.

Thorax olivaceous laterally ; mesonotum shining, metallic, slightly overcast with brown,
s])aringly setulose, humerus and notopleura more grayish, two narrow inter-dorsocentral stripes
and a broad extra-dorsocentral stripe more opaijue and brownish to bluish. Acrostichal setae
much stronger than surrounding setulae, the series closer together than their distance from the
dorsocentral series; presutural acrostichals weak; some inter-acrostichal setulae present and
a few other scattered inter-dorsocentral setulae posteriorly near the dorsocentral bristles.
Scutellum broad as long, pilose apically, flattened, transversely rugulose. Mesopleura very
sparingly setulose.

'Lindner's Flieg. Pal Reg., Fam. 56, p. 75 (1926).
'Russ. Hydrob. Zeit.. VIII, pp. 140-141 (1929).
'Em. News, XXVII, p. 151 (1916).

DIPTERA OF THE FAMILY EPHYDRIDAE 6

Abdomen narrow ; fifth segment longer than fourtli, greenish glaucous ; apical margins
of these segments with few long setae.

Legs olivaceous; post-extensors of fore femora very long. Wings blackish; length to
width as 60: 32; posterior crossvein perpendicular to fifth vein.

9 . Similar to male but with stronger bristles, many of the mesonotal setulae bristle-
like ; the prescutellar convexity bearing some setae and the prescutellar acrostichals stronger
than in the male, jirescutellar acrostichals slightly divergent. Scutellum slightly elongate and
noticeably con\e.x. Postmarginal cilia of mesopleura dense and sternnpleura strongly pilose
above.

Type. — S ; Indian Tibet: Kyam Hot Spring, 15,630 feet altitude, 24 July, 1932.

Paratypes. — 3 <5 , 4 9 ; topotypical.

A female from Phuga hot spring, 14,500 feet altitude, 24 August, 1932, is slightly more
robust, bristling longer on frons, mesonotum and abdomen less metallic, blackish ; face and
pleura also more blackish or brownish than cinereous and olivaceous, but structurally there
seem to be no marked differences.

Halmopota hutchinsoni n. sp.

This is similar to H. c-illosa Becker from northeastern Tibet, but that species is brownish
with ])ale tibiae and tarsi, and the dorsocentrals are not developed. Only the female sex of
z'illosa is known, but I cannot I )elieve the male before me to be of that species.

Entirely black; at most the tarsi are slightly paler distally, and the halteres are pale
yellow. Uniformly opac[ue glaucous gray, with frons, mesonotum and scutellum blackish in
some aspects.

Head 1.2 longer than high, 1.6 wider than high; in profile the frons is slightly convex,
oblique, setting the antennae but slightly above center-line of head and at, or slightly below,
that of eyes; ante-ocellar extension about one-half length of head. Kyes obliquely elongate.
Frons broad as long, sparingly setulose anteriorly; seven to eight strong fronto-orbitals and
a few small setae niesally ; ocellars in line with the anterior ocellus and another pair, as
strong, just behind, some slightly weaker setae behind and between' these, also a strong post-
vertical pair, b'ace with gibbose medifacies less than one-half width of head, strongly sub-
hemispherical, making a distinct sub-horizontal interfoveal hump; four to six facial bristles
in a series on lower half; parafacials very broad, about length of second and third antennal
segments in width. Cheeks almost one-half height of head in width, with five to six strong
dorso-inclinate marginal bristles. Postbucca strongly turgid, setulose. Cilia of posterior
orbits of about six strong blistles. First antennal segment distinctly visible, one-half length
of second ; second segment broader than long, setulose basally with strong dorsal setae ; third
broader than long; arista twice as long as third, with thick pubescent basal three- fourths,
gradually attenuating distally to hair-like tip.

Mesonotum sparingly, strongly, setose, many of the setae may be confused with the
macrochaetae ; the three postsutural acrostichals as strong as the dorso-centrals ; the latter
arranged 2 to 3 : 3 to 4; interalar series of bristles and setae; 1 humeral; 1 presutural ;
2 to 3 notopleurals ; several supralar and postular bristles discernible among the strong setae.
Mesopleura strongly setulose with strong ])ostmarginal cilia; sternopleura setulose above; the
pleural sclerites otherwise bare. Scutellum triangular, as long as broad, flat or slightly convex,
bare, but with two apical and two to four lateral bristles.

4 DIPTERA OF THE FAMILY EPHYDRIDAE

Abdomen sliijhtly broader than thorax, sparingly clothed with appresscd setulae; later-
ally and vciitrally more strongly so; second to fourth segments suhcqual in lengths; fifth
slightly longer.

Bristles of legs strong and numerous but not seriated except on the flexor margins;
setae of posterior margin of fore femora in length equal to diameter of the femora; middle
femora with postflexor comb of closely set spinules on distal three- fourths. Wings translu-
cent, immaculate, with dark veins; venation normal except that the post-crossvein very
strongly undulated; ultimate section of fourth vein not as long as the penultimate; squamae
white with black cilia.

Length, 6 mm.

Type S ; Indian Tibet: Tso-kar, 14,850 feet altitude. September 5, 1932.
(G. E. Hutchinson.)

A^ote on the larvae and puparia

P 10. Punjab: Son Sakasar Kahar, March 13, 1932, from submerged branches of
brush; pH 8.9; 89,306 parts non-volatile solids per million. Three larvae
and six puparia.

L40. Indian Tibet: Panggong Tso, Lagoon II, west end, June 20, 1932; pH 9.3,
alkali reserve 0.0085 N., chloride 0.0007 N., cold water. Two larvae and
one puparium.

The larvae of this lot have two dorsal longitudinal clusters of closely set spinules on seg-
ments one to four, becoming rounded or sometimes coalescing into transverse bands on five
to eight; seventh prolog well developed; post-anal tubercle also distinctly developed. The
puparia have a total length of 6.5 mm. ; greatest thickness at third segment of 2 mm. ;
length of anal tube 1.5 mm., two and a half times as long as its basal diameter; length of
respiratory tube .6 mm. ; dorsal profile of segments one to five practically straight, that of
six to eight convex, causing the prolegs of six and eight to approximate each other to almost
touching; proleg of seven scarcely developed, represented only as a slight swelling bearing a
few curved spinules.

L 58. Indian Tibet: Kyam hot springs, July 19, 1932; 1055 parts non-volatile solids
per million; one larva and several puparia. This form agrees well with Brues description and
figure of Ephydra pcctimdata Cresson, described from the Yellowstone Park, Wyoming," but
I note the following differences: Larva with dorsal patch of spinules as described on the
larvae from Panggong; prolegs with two transverse rows of curved hooks, those in the
anterior series much the stronger and usually four in number. Otherwise the larva and
puparium as described from Panggong except that the anal tube is shorter, which character
seems to vary considerably. This form may be Ephydra tibetetisis here described.

Another larva collected from the above locality, July 20, 1932, in a small pool with tem-
perature of 35.6 C, appears to be of the form described from Panggong, lagoon II.

L80. Indian Tibet: Phuga hot springs, August 25, 1932, temp. 49.1°C.; pH 8.6;
alkali reserve 0.0144N ; chloride 0.01 12N ; very rich in HoS. Eight larvae which appear to Ije
same as those described from Panggong, but their lengths are 4.2 to 6 mm.; anal tube of
largest, .9 mm.

•Proc. Am. Acad. A. & Sci., LIX, pp. 403-405, fig. 5 (1924).

ARTICLE II

REPORT ON TRICLAD TURBELLARIA FROM INDIAN TIBET

By Libbie II. IIyman
(Received November 10, 1933)

The planarian material collected by the Yale North India Expedition was turned over to
me by Mr. G. E. Hutchinson for examination. It consisted of fourteen vials of specimens in
alcohol. These were run up into oil of wintergreen and examined with low power. It was
then seen that the contents of thirteen of the vials are all (presumably) of one species, a
species of Polycelis, while the fourteenth vial contains specimens which are probably to be
assigned to the genus Euplanaria. Fortunately some of the Polycelis are sexually mature and
it has therefore been possible to furnish a diagnostic description. The specimens of the
second species are unfortunately all asexual and consequently cannot be placed generically
with any certainty. All of the material came from Kashmir and Indian Tibet. The sta-
tions labelled K64-K71 are located to the northeast of Srinagar, Kashmir; those marked
K 74-K 83 and L up to number 25 are collecting sites in Indian Tibet on the road between
Srinagar and Leh and in the Indus valley above Leh ; and tlie remainder of the numbers
under L are situated in Indian Tibet north and east of the Ladak range. The following
remarks give the data found on the labels in the vials together with my own notes from
examination of the cleared specimens.

K 64. Gund, rest house, 2080 meters, under stones in a very small stream, temperature
9.4°C., May 17th, 1932. This vial contains three specimens which constitute the second
species mentioned above whose external appearance suggests the genus Euplanaria.

K 66. Small stream with very steep slope, about three miles west of Sonamarg, 2440
meters, temperature 7°C., May 18th, 1932. This vial contains eight good-sized specimens of
Polycelis, none of which appear to have sex organs.

K 71. Stream west of Sonamarg, 2590 meters, temperature 7.0 C., pH. 7.6, May 19th,
1932. Thirteen specimens of Polycelis, mostly rather small. Four of the largest were sec-
tioned but proved to be devoid of sex organs.

K 74. Small stream, temperature 7.3°C., pH. 7.5, mossy, 50 cm. wide, Matayan, 3170
meters, May 20th, 1932. Eight specimens of Polycelis.

K 76. Small stream, temperature 19.0^C., about one mile west of Dras, 3080 meters.
May 21st, 1932. Of the ten,specimens of Polycelis foiind in this vial two were removed and
sectioned but proved asexual.

K77. Stream, Dras, temperature 21.0-22.0 C, 3091 meters. May 21st, 1932. Nine
specimens of Polycelis, five adult and four young ones, all asexual.

K 78. Karbu between Dras and Kargil, spring, temperature 8'C., pH. 7.5, 2819 meters.
May 22nd, 1932. This vial contained four specimens of Polycelis of which two were seen to
be sexually mature and have been sectioned.

K83. Spring, Kargil, temperature 10.2° C., 2679 meters, May 24th, 1932. Two speci-
mens of Polycelis, not sexual.

Mem. Conn. Acad., Vol. X, Art. II. September, 1934.

6 TKU I.AI) TIKHKMAKIA FRDM INDIAN TIBET

L 21.^ Pool just IkIow Ilimis Goiipa, icnipcrature 9.5 C, o500 meters, June lith,
1932. Contains four specimens of Polycelis, asexual.

L25. Stream above Leli, temperature /.S-S.G^C., 3570 meters, June 21st, 1932. Si.x
specimens of Pol\ccIis of which two were obviously in the sexual state. These were removed
and sectioned and furnish the main basis for the taxonomic description.

L 34. Pool, Bao, temperature 20.8 C, 4585 meters, June 2Sth, 1932. This vial con-
tains one Polycelis and three small specimens which are probably rhabdocoels.

L35. Springs, four miles from Bao, temperature 7.2 C, 41 meters, June 26th, 1932,
Five good sized but asexual specimens of Polycelis.

L 60. Stream, under stones, Kyam, 4725 meters, July 21st, 1932. Allhoui^h the sev-
eral specimens of Pulxcclis in tiiis vial are the lar.nest in the cnllection they appear to be in
the asexual state.

L 75. Springs, Tukmuru T.so, temperature IOC, 4385 meters, August 11th, 1932.
Seven rather small specimens of Polycelis.

Search through the literature revealed the fact that this Polycelis had Ijeen seen before.
A number of specimens were taken in the expedition for collecting the aquatic animals of
Tibet made by Captain F. H. Stewart in 1907. The planarians of this collection were turned
over to Meixner and Muth who publishetl their report in 1911. They described and figured
the external features of the specimens, giving several drawings of the arrangement of the
eyes, and placed them in the genus Sorocclis. Owing to variations in the disposition of the
eyes these authors were inclined to think that the material consisted of more than one species.
Their specimens came from streams at Te-ring Gompa, 14,000 feet, and High Hill Gompa
above Gyantse, 14,500 feet. This form was again mentioned by Muth (1912j as similar to
his Sorocclis cbitntca from the region of I^ake Aral. Owing to a lack of sexual specimens,
Meixner and Muth were unable to furnish a diagnosis of the Tibetan material.

The genus Sorocclis was until recently a badly defined genus into which were thrust a
number of many-eyed fresh-water triclads collected chiefly in Asia. In 1930 Kenk, in his
invaluable re-vision of the genera of the fresh-water planarians, proposed to limit this genus
to many-eyed f(n-nis belonging to the family Dendrocoelidae. Those which from the arrange-
ment of the inner muscular layers of the pharyn.x fall into the family Planariidae he has rightly
transferred to the genus Polycelis. The Tibetan specimens at my disposal belong to the family
Planariidae and the sexual apparatus corresponds in all respects to Kenk's definition of the
genus Polycelis. I therefore have no hesitation in placing them in that genus. It is also rea-
sonably certain that my specimens are the same as those recorded by Meixner and Muth. I
do not, however, agree with their supposition that more than one species is concerned. It
is true that the eyes are somewhat variable in different specimens, but these variations are
partly correlated with age and in any case are insufficient to serve as specific distinctions. To
the best of my knowledge, the above references constitute the only records of the Tiljetan
Polycelis. Study of the sexual apparatus shows that the form does not correspond to any
described species of Polycelis, and consecpiently I consider it a new species which fmni its
habitat I name Polvcelis iibetica.

TRICLAD TURBEI.LARIA FROM INDIAN TIBET /

Polycelis tibetica n. sp.

Syti. Sorocclis sp. Meixner und Mutli, 1911.

1. Extcrm^l features. As only preserved specimens are available, the size and shape of
the living animal cannot be stated. In preserved specimens there is always a certain amount
of distortion. The size appears to be moderate, probably not exceeding 15 mm. Judged
from the least distorted specimens, the head has much the same shape as in Polycelis coronata
(see Hyman, 1931a). The anterior margin (Plate I, fig. 5) is rounded in a gentle curve
and provided laterally with two fairly prominent auricles.^ The arrangement of the eyes is
also similar to that of P. coronata, consisting of a semicircular band passing near the anterior
margin, along the base of the auricles, and terminating shortly behind the auricles. Behind
the auricles, the body incurves slightly, then broadens toward the middle regions, and finally
again diminishes towards the rounded posterior end. Plate I, fig. 1, gives the appearance of
one of the less contracted specimens. Presumably in life the auricles would be more promi-
nent, probably still more extended, than in the specimen shown in Plate I, fig. 5, and the body
somewhat more elongated. As is common in the genus, the pharynx is very long and
powerful, terminating near the posterior end and leaving only a short region for the sexual
apparatus.

The eyes in all the specimens form a semicircular band several eyes wide running along
the margin of the head and terminating" shortly behind the auricles. The number and
arrangement of the eyes vary in different specimens, no two worms being exactly alike in
these respects. Frequently the posterior end of the band is at a different level on the two
sides. The number of eyes varies definitely with age, consisting of thirty to forty in the
smallest specimens, sixty to eighty in medium specimens, and eighty to one hundred or more
in the largest worms. Plate I, figs. 2 to 5, inclusive, indicate this relationship with age. In
Plate I, fig. 6, where the number of eyes is quite small although the worm was a large one,
there were various indications that the head was in process of regeneration.

The coloration appears to be a uniform dark brown or black.

2. General histological features. Some of the specimens are in a good state of fixation
but the histology presents nothing in particular. The ventral epithelium is cuboidal and is
ciliated throughout. The dorsal epithelium is slightly taller, contains more rhabdites, and is
ciliated at least in part. The anterior part of the dorsal surface is always ciliated to a greater
or less extent while posterior dorsal regions usually lack cilia. The adhesive zone is very
narrow but so far as could be determined follows the usual course completely encircling the
body margin. The most conspicuous histological elements are the lar;^e gland cells found in
the anterior half of the body distributed thickly below the intestine, slightly less abundant
dorsal to the intestine. They consist chiefly of the large rounded gland cells taking basic
stains which are called Eiweisszellen 1)y German authors. Gland cells of this tyjie and location
are very common in planarians.

The pharynx has the structure typical of the family Planariidae, the circular and longi-
tudinal muscle layers of the inner muscular zone being distinctly separated into two strata.

3. Reproductive system. This system follows the plan typical of the genus Polycelis.
Of the four sexual specimens found in the collection two came from station K 78 and two

'In Plate I, fig. la, I have ventured to add a copy of a sketcli of the anterior end of a living animal, made
from one of the specimens of the series from K 66. — G. E. H.

8 TKK'I.M) I I'KUKI.I.AIv'IA lUnM INlllAN I'lliKT

from station L 25. The two specimens from each station are olniously specifically identical
with each other; bnt at first the penis and penis bulb of the specimens from different stations
appeared to differ. .After some study, however, I decided that the difference was one of
degree of extension and have concluded tiiat all four specimens belong to one species. As
they came from stations rather widely separated, it may Ije considered that all the Polycclis in
the collection are of one species.

The ovaries are a pair of small rounded compact masses in the usual anterior position.
No parovaria were found. The oviducts exit from their lateral surfaces. They could not
be traced very W'ell in any of the specimens but appear to run immediately to the medial side
of the ventral nerve cords. At the level of the penis bulb they curve dorsally and above the
male atrium unite into a common oviduct (fig. 9, co) which immediately turns ventrally and
opens into the male atrium just anterior to the junction of the latter with the bursa stalk
(Plate I, fig. 7, and Plate II, fig. 3, co). This arrangement of course obtains throughout the
genus Polycelis. The oviducts do not embrace the bursa stalk since they unite anterior to it
around the sides of the penis bulb. A few eosinophilous shell glands occur around the point
of union of the oviducts. The yolk glands have the usual appearance and occur as masses of
large granular cells lying between the intestinal diverticula from the level of the ovaries to the
posterior end of the body. In one of the K 78 specimens they are exceedingly numerous and
conspicuous. Presumably the yolk glands behind the copulatory apparatus must connect with
the oviducts by special yolk ducts.

The testes occupy the position typical of the genus Polycelis. They extend near the
ventral surface from the ovaries to the root of the pharynx in a double row, one row to each
side of the midline, lying between the bases of the intestinal diverticula. The testes could be
identified in all of the specimens sectioned consisting in asexual individuals of masses of
rounded cells in the resting state. They were also in this condition in the less mature L 25
specimen but were in active spermatogenesis in the other three sexual specimens in which also
the vasa deferentia contain sperm.

The vasa deferentia form the usual tubular enlargements termed false seminal vesicles
easily seen along the rear part of the pharynx and sides of the bursa copulatrix. At the level
of the penis bulb they curve dorsally and enter the bulb separately one from each side
(Plate I, fig. 8, and Plate II, fig. 4, vd). They penetrate the wall of the penis bulb without
enlargement, each one opening on a papilla (Plate I, fig. 8, and Plate II, fig. 4) which pro-
jects into the cavity of the bulb from its sides.

The penis bulb and penis differ so strikingly in the L 25 and the K 78 specimens that at
first I feared it would be necessary to distinguish two varieties of the species. However, I
finally concluded that the differences rest entirely in the muscular state of these parts. In
the L 25 specimens (Plate I, fig. 7), the copulatory apparatus is withdrawn and in rela.Ka-
tion. There is a large hollow penis bulb and a small conical penis. But in both of the K 78
worms, the bulb and penis are extruded (Plate II, fig. 3). The penis bulb is strongly con-
tracted into a muscular disk and its cavity has been projected into the penis. The latter
organ has thus incorporated the penis bulb and appears as a large muscular elongated organ
with a considerable cavity. If my conclusions are correct, these specimens furnish a striking
example of the rule of the bulb in the protrusion of the penis, its contraction converting
an apparently small weak penis into a large powerful elongated organ. Probably the penis

TRICI.An TURRia.I.ARIA FRIIM INDIAN TIHET y

and bnlb should not be considered as separate parts of the copulatory apparatus but as one
structure.

In the L 25 specimen, whicli seems the most mature of the four, the penis bulb is a
rounded hollow sac of moderate dimensions immediately behind the bursa copulatrix (Plate I,
fig. 7, pb). It is well-defined and plainly marked off from the surrounding tissues but is not
strikingly muscular. The wall is of moderate thickness and composed chiefly of muscle fibers
coursing in several directions. The interior is hollow forming a seminal vesicle (Plate II,
fig. 4, sv), somewhat hourglass-shaped in transverse section owing to the projection into the
lumen of the two lateral papillae which bear the ternn'nations of the vasa deferentia (Plate I,
fig. 8, and Plate II, fig. 4). The lumen is lined by what appears to be a glandular epithelium
densely packed with secretion granules. However, under oil immersions, the epithelial cells
are found to be practically undetectable owing to their penetration by what appear to be tubes
packed with coarse granules (Plate II, fig. 2). Similar granules in groups are found through-
out the subepithelial wall of the bulb as well as in adjacent tissues. It appears probable that
<ve are dealing with very long-stalked unicellular glands which open by ducts through the
lining epithelium of the bulb. These ducts project beyond the epithelial surface and often
a cloud of granules is seen emerging from their open ends (Plate II, fig. 2). These granule-
filled ducts have been found only in the very ripe L 25 specimen and seem to indicate that
some secretion of importance is discharged into the seminal vesicle during the height of
sexual activity.

In the L25 specimens, the penis is a short weak cone-shaped organ projecting into the
male atrium (Plate I, fig. 7) and containing a short duct running from the seminal vesicle
to the tip of the penis. The duct is lined by a columnar epithelium lacking the granular
tubes mentioned above. The male atrium is lined by a cuboidal epithelium encircled by cir-
cular, longitudinal, and radial muscle fillers (Plate II, fig. 1). It is expanded where it con-
tains the penis but immediately beyond this organ narrows at once to a short canal which
receives in its middorsal line first the common oviduct and then the stalk of the bursa
(Plate I, fig. 7). It then immediately opens at the genital pore, the common cavity formed
by its union with the bursa stalk being thus so small as scarcely to merit the name of common
atrium. All of these relations in the atrium are typical of the genus Polycelis.

As already indicated the appearance of penis bulb and penis is very different in the
K 78 specimens (Plate II, fig. 3). Here the penis bulb is apparently contracted and forms a
muscular mass at the base of the penis. The seminal vesicle seems to have been projected
into the penis but its boundaries are still determinable by means of the papillae on which the
vasa deferentia terminate. The penis in consequence of having incorporated most of the penis
bulb appears very much larger, longer and more powerful than when at rest (compare Plate I,
fig. 7, and Plate II, fig. 3j. - The conditions in these specimens seem to prove that the penis
bulb is of great help in the extrusion of the penis and is really an essential part of that organ.

The bursa copulatrix in all the specimens is a large, irregularly rounded sac lying between
the rear end of the pharynx and the penis bulb, and considerably larger than the latter .struc-
ture (Plate I, fig. 7). Its histology is typical, the organ being lined by the usual large
bulbous epithelial cells containing rounded masses. From the center of its posterior surface
the stalk arises and runs posteriorly above the penis bulb. At first the stalk is large and lined
by the same epithelium as the bulb; but at about the level of the union of the oviducts, the
stalk narniws abruptly (Plate I, fig. 7, and Plate II, fig. 3) and its epithelium becomes of an

10 TKICI.AL) TLRISKl.I.ARIA l-KO.M INDIAN TIBET

ordinary coftimnar or cuboidal type. It is, however, lieavily ciliated. This narrow ciliated
stalk turns ventrall}' and joins the male atrium at the genital pore. A very thin layer of
muscles seems to underlie the epithelium of bursa and stalk.

A sas^ittal section of the copulatory apparatus of one of the 1.25 specimens is shown in
Plate I, fig. 7, of one of the K 78 worms in Plate II, fig. 3. Plate I, fig. 8, and Plate II,
fig. 1, are successive transverse sections through the second L 25 specimen, showing in
Plate I, fig. 8, the entrance of the vasa deferentia into the penis l)ulli with the oviducts at the
sides and the wide part of the bursa stalk above; and in Plate II, fig. 1, the male atrium,
union of the oviducts, and narrow part of the bursa stalk above. Plate II, fig. 4, is a diagram
of the sexual apparatus seen from above.

The sexual specimens in the collection were taken (in May 21st and June 21st respec-
tively. The time of sexual maturity is therefore early summer. One of the L25 specimens
taken on May 21st appears to be the ripest of the lot; but presumably the time of sexual
maturity depends in part on altitude, those at higher levels maturing later in the season.

4. Habitat. The worms came from springs, streams, and pools in the high mountains of
Kashmir and Indian Tibet. The altitudes of the collecting sites varied from 2500 to 4700
meters while in the Stewart expedition the species was taken between 4250 and 4400 m.
These are probably the greatest heights at which any fresh-water planarians have ever been
found. The temperatures of the habitats are recorded in some cases and, as might be
expected, are mostly low, ranging from 7 to 22"C. In three cases the acidity is recorded as
pll 7.5-7.6, quite a usual figure for such habitats. The habitat of Polycclis tibetica is very
similar to that oi the only known American representative of the genus, Polycclis coronata,
which also lives in mountain streams and springs. Although the American species has as yet
been taken only at moderate altitudes (below 5000 feet), there is little reason to doubt that
it will be found distributed throughout the high mountain ranges of western North America.
The only other species which seems to be specifically a high mountain form is Polycclis
cormita of Europe which occurs in the streams of the high Alps althougli distributed over a
considerable range of altitude. On the other hand some species of Polycclis are lowland forms
but the entire genus appears to require rapidly flowing water and stony bottom.

Second Species

It seems desirable to make a statement about the secdud species found in the collection.
This is represented by three individuals taken at station K 64 at 2080 meters, in a stream under
stones. As this is the lowest altitude at which planarians were found, it seems probable that
this .species inhabits lower altitudes than does Polycclis tibetica and consecjuently was not
taken at any of the higher stations.

The species is of large size, probably reaching a length of 20 mm., of uniform dark-
brown coloration, and w^ith an evident triangular head with auricles. From the size, uniform
dark coloration, the shape of the head, the appearance of the digestive tract, and in fact, the
general aspect of the form, I am quite sure it is a species of Euplanaria, very close to the
three American memljers of Euplanaria which I have called the dorotoccphala group (Ilynian,
1931 b). However, in the absence of sex organs, the form cannot be placed taxonomically with
any certainty and I therefore forbear to attach a name to it. It has one peculiarity by which
future collectors in this region can probably recognize it. The species tends to Ix; four-eyed.

TklCLAD TURBELI-ARIA FROM INDIAN TIBET 11

Of the three specimens, the smallest (Plate II, fig. 6) has four eyes of ec|ual size, the medium-
sized one (Plate II, fig. 7) has two eyes of regular size in front of which are two small eyes,
and the third, the largest, has one small eye in front of one of the two regular eyes (Plate II,
fig. 5). It is possible that in this form the eyes are regularly replaced and the small eyes of
Plate II, figs. 5 and 7, represent new eyes in process of development. However, the pres-
ence of supernumerary eyes is not at all uncommon in planarians and cannot be used as a
taxonomic character.

Affinities

These species appear to resemble American forms more nearly than they do European
planarians. The European Polycelis species have a single row of eyes along the anterior
margin while the banded arrangement, several eyes wide, occurs only in Asiatic and American
forms. In the eyes as well as in the anatomy of the copulatory apparatus Polycelis tibetica
bears considerable resemblance to Sorocclis sapporo (presumably a Polycelis) of Japan
( Ijima and Kaburaki, 1916). Also the Euplattaria (?) of the present collection is very
like some common Euplanariae of the United States while differing from European Etiplan-
ariae. .Altogether it would seem that the Asiatic species have spread toward North America
(or vice versa) rather than toward Europe.

12 rKUI.AI) Tl-KRF.I.I.AKIA I-K(1M INDIAN TIKK.T

CITATIONS

Hyman, Libbie H. 1931a Studies on the morphology, taxonomy, and distribution of
North American triclad Turbellaria. III. On Polycelis coronata (Girard). Trans.
Amer. Micro. Soc. 50:124.

1931b IV. Recent European revisions of the triclads and their application to the Ameri-
can forms with a key to the latter and new notes on distribution. Same: 316.

IjiMA, I., and Kaburaki, T. 1916 Preliminary description of some Japanese triclads. Anno-
tationes zoologicae japonenses 9: 153.

Kenk, R. 1930 Beitriige zum System der Probursalier (Tricladida paludicola). III. Ver-
such einer natiirlicher Gruppierung der Probursalier. Zool. Anz. 89 : 289.

Meixner, a., and Muth, A. 1911 Report on a collection of aquatic animals made in
Tibet by Captain F. H. Stewart during the year 1907. III. Turbellaria and sum-
mary. Records of the Indian Museum 6: 57.

Muth, A. 1912 Beitrage zur Kenntnis der Gattung Sorocelis Grube. Mitteil. naturwiss.
V'ereins Steicrmark 48:381.

N

Explanation of Plate I.

Fig. 1. General appearance of Polycclis tibetica from a preserved specimen. ;//, mouth;
gp, genital pore.

Fig. la. Sketch of heail of living specimen from lot K 66. (G. E. H.)

Figs. 2, 3. Ej-e number and arrangement of small specimens from lut K 77.

Fig. 4. Eye number antl arrangement of large specimen from lot K 76.

Fig. 5. Large specimen from lot L21 having the best extension of auricles found.

Fig. 6. Large specimen from lut K 66 with head probably regenerating, showing few eyes.

P'ig. 7. Sagittal section of the copulatory apparatus of the ripest specimen, from L 25. be,
bursa copulatri.x ; bs, bursa stalk; (V, common oviduct ; ^^'/i, genital pore ; iini,
male atrium: Z^. i)enis; /ifc, jicnis bulb.

Fig. 8. Transverse section nf second specimen from L 25, taken tln'nugji the center of the
])enis bulb, bs, bursa stalk: o, oviduct; />/'. penis bulli; ;■(/, vas deferens.

MEM. CONN. ACAD., VOL. X.

PLATE

B S

CO

B S

5 <'^'^.'<S»='^'v

'^^y*:.

(?^. \

-V D

\

Explanation of Plate II.

Fig. 1. More posterior section of same series as Plate I, fig. 8, through tlie male atrium.
bs, bursa stalk; ma, male atrium ; o, union of oviducts; p, penis; tviiia, wall of
the male atrium.

Fig. 2. Detail of the wall of the penis bulb of same section as figure 7, showing the granule-
filled ducts projecting through the lining epithelium.

Fig. 3. Sagittal section of the copulatory apparatus of one of the K 78 specimens, recon-
structed from several sections. The penis bulb is contracted forcing the seminal
vesicle into the penis, be, bursa copulatrix; co, common oviduct; gp, genital
pore; p, penis; pb, penis bulb; 7'rf, vas deferens.

Fig. 4. Diagram of the copulatory apparatus seen from the dorsal side, be, bursa ci>pulatrix ;
CO, common oviduct; via, male atrium; p, ])enis; pb. penis bull); .?<', seminal
vesicle; ty/, vas deferens.

Fig. 5. Largest specimen of the second species, ? Euplaiiaria, from lot K 64.

Figs. 6, 7. The two other specimens of lot K 64, showing four eyes.

MEM. CONN. ACAD., VOL. X.

PLATE IL

BS

o

W M A

V D

P B

CO

•1!^..

2

ARTICLE III

REPORT ON COLEOPTERA OF THE FAMILY STAPHYLINIDAE

By Malcolm Cameron, M.B., R.N., F.R.E.S.

(Received January 18, 1934)

The small collection of Staphylinidae, obtained by the Yale North India Expedition,
comprised material of fourteen species enumerated below ; nine species are hitherto unde-
scribed. By arrangement with Yale University the types of these new species have been
incorporated in the collection of the British Museum.

OXYTELINAE

Megarthrus rufomarginatits Cam. Nilgiri Hills : Pykara, altitude circa 7000 feet.
15-XI-32.

Lesteva kargilensis sp. n.

Rather shining, black, the antennae blackish ; femura reddish-yellow, pitchy at apex,
tibiae pitchy, tarsi reddish-yellow. Length 3.75 mm. More robust than fluviata Champ., less
shining, the antennae much longer, thorax more dilated in front and more finely punctured,
elytra more finely punctured. Head bi-impressed between the eyes, closely, moderately
coarsely punctured except on the front where only a few fine punctures are present. Antennae
long and slender, all the joints much longer than broad. Thorax transverse, cordiform, the
sides retracted behind with rectangular posterior angles; l^efore the base with a superficial
impression, closely and more finely punctured than the head. Elytra twice as long as the
thorax, slightly widened behind, as closely and as finely punctured. Abdomen extremely
finely and densely punctured, coriaceous.

Indian Tibet: Kargil. 24-V-32. Mossy stones by spring. Unique.

Geodromicus affinis sp. n.

Rather shining; head and elytra black, thorax and abdomen pitchy (? immature).
Antennae and legs reddish-brown. Length 6 mm. Closely allied to kashiiiircnsis Cam., but
the head is a little narrower and much less punctured, the thorax more finely punctured, the
elytra shorter, more coarsely and less closely punctured and widened behind. Head narrower
than the thorax, deeply impressed on the vertex, the ocelli slightly more apart from each
other than from the eyes, very finely, sparingly punctured near the eyes, almost impunctatc
elsewhere; ground sculpture absent. Antennae long, all the joints much longer than broad.
Thorax strongly cordiform, convex, the sides strongly roundecl and widened in front, retracted
liehiiid, the ]3nsterior angles rectangular, at the middle of the base with a fovea, narrowly

Mem. Conn. Acad., Vol. X, Art. III. September, 1934.

18 COLEOPTF.RA OF TIIF. FAMILY STAPHYLINIDAE

and feebly impressed along the middle, moderately, finely superficially and moderately closely
punctured; ground sculpture absent. Elytra longer (8:5) than the thorax, widened Ijehind,
rather closely and much more coarsely punctured. Abdomen extremely finely, rather
closely punctured and coriaceous.

Indian Tihet: Kargil. 24-V-32. Wet mossy stones near spring. A single specimen.

GeodrO)!iici(s siiiiilis Cam.

A single specimen with the preceding.

Trogophloeus (Taenosoma) porosus sp. n.

Entirely black, the fore-jiarts slightly, the abdomen more shining. Antennae black.
Legs black, the apex of the tibiae and the tarsi brownish-yellow. Length 2.2 mm.

Closely allied to scabrosus Kr. the sculpture scarcely different, but a little smaller and
less robust, the 4th to 6th joints of the antennae a little longer, the penultimate less transverse,
thorax a little longer, less transverse, the sides more retracted behind, the elytra longer.
Head a little narrower than the thorax, feebly bi-impressed in front, the post-ocular region
rounded, a little longer than the eye, the whole surface covered with very coarse, close, rugose,
finely umbilicate punctures, .\ntennae with the 3rd joint shorter than 2nd, 4th and 5th very
slightly longer than broad, 6th as long as broad, 7th to 10th transverse. Thorax a third
broader than long, the sides rounded and dilated in front, almost straight and retracted
Ijehind, the disc without trace of impressions, in the middle with a short, extremely fine
shining line, otherwise covered with coarse sculpture as on the head. Elytra broader and a
third longer than the thora.x, with similar but rather coarser sculpture. Abdomen very finely,
moderateh* closely punctured, finely, moderately closely pubescent.

Nii.GiKi lIii.Ls: Pykara, altitude 7000 feet. 15-X1-32. On wet earth on clilT. Unicjue.

Delopsis consanguinea sp. n.

In colour and opacity similar to glarcusii \Voll. (O.vytcliis) , b\il niucii larger (2.2 mm. I
and more robust, the antennae longer and stouter, the 4th and Stli joints longer than broad,
the penultimate less transverse, the sculpture throughout coarser.

Head very slightly dilated Iiehind the eyes, the impressions as in glarcosa. Thorax very
slightly sinuate before the posterior angles, the sulci deeper. Elytra a third longer than the
thorax.

NiLGiRi Hills : Pykara, altitude 7000 feet. 15-XI-32. Unique.

Stenus (s. str.) pykaranus sp. n.

Shining black. Antennae palpi and legs black, the extreme base of the femora yellowish-
red. Length 4.4 mm. Allied to tortuosus Cam. but smaller, the antennae shorter and like
the palpi entirely black, thorax less uneven, less coarsely punctured. Abdomen much more
finely and sparingly punctured. Head as broad as the base of the elytra, concave Ijetween the
eyes, not elevated along the middle, closely, rather coarsely but not rugosely punctured.

COLEOPTERA OF THE FAMILY STAPHYLINIDAE 19

Antennae rather sliort, the 3rd joint a good deal longer than the 2nd, 4th to 7th gradually
decreasing in length, 8th to 10th slightly transverse. Thorax a little longer than broad
(7: 5.5), widest about the middle, the sides rounded in front, nearly straight and retracted
behind, ali)ng the middle posteriorly with a narrow impunctate line, before the base on each
side with three or four transverse rugae, the rest of the surface closely and more coarsely
punctured than the head and somewhat rugose. Elytra a little longer than the thorax, with
coarse oblique rugae passing from the middle of each disc backwards and inwards to the
suture and sutural angle, a few finer ones also passing forwards and inwards towards the
scutellary region ; shoulders and base closely and moderately finely punctured, postero-exter-
nally coarsely, closely and rugosely punctured. Abdomen gradually narrowed from base to
apex, extremely finely, obsoletely, moderately closely punctured, rather more closely and dis-
tinctly on the 9th segment. Fore-parts almost glabrous. Abdomen sparingly pubescent.

$ : Unknown.

NiLGiRi Hills : Pykara, altitude 7000 feet. 15-XI-32. Unique.

STAPHYLININAE

Actobius basalis Motsch. var. Iiiiiiicralis Cam.

NiLGiRi Hills : Pykara, 15-XI-32. Two specimens.

Type widely distributed in the Oriental region, the variety so far only recorded from
the Nilgiri Hills.

Pliilonthns lidarensis Cam.

Indian Tibet: Kargil, altitude 8790 feet. 24-V-32. On wet mossy stones near spring.
One specimen. Also known from Lidarwat, altitude 9000 feet, and Gulmarg.

ALEOCHARINAE

Athela (Aloconota) iguensis sp. n.

Entirely black, the elytra with very slight metallic reflex, the fore-parts moderately, the
abdomen more shining. Antennae, palpi and legs black, the tarsi brownish-yellow. Length
2.8 muL

Head transverse, suborbicular, nearly as broad as the thorax, the vertex with a fine short
sulcus, extremely finely and very sparingly punctured, strongly coriaceous. Antennae long,
the 3rd joint longer than the 2nd, 4th to 10th all longer than broad, gradually decreasing in
length, the 9th and 10th only a little longer than broad, together as long as the 11th. Thorax
slightly transverse, the sides rounded in front, sinuate and retracted behind, the posterior
angles obtuse, the base on each side obliquely truncate as in iiisecta Thorns along the middle
in the posterior half superficially impressed, very finely, much less sparingly punctured than
the head, the ground sculpture similar. Elytra a little broader and half as long again as the

20 COr.EOPTKRA OF TUF. FAMILY STAPHYLINIDAE

thorax, tlie puncturation similar but mucli closer, the ground sculpture similar. Abdomen
\ery finely, moderately closely punctured on the anterior segments, gradually more sparingly
behind, less strongly coriaceous than the fore-parts. 8th dorsal segiuent very slightly
arcuately emarginatc. The pubescence throughout fine and moderately close. Tibiae without
long setae.

A single e.\ani])le which appears to be a S Alocoiwta frnm the general facies.

Indian Tihf,t: Igu, altitude 11,210 feet. In stream shingle.

Athela (Bessobia) submetallica sp. n.

Moderately shining, black, head, thorax and elytra with .slight metallic reflex. Antennae
black. Legs black, the tarsi yellow. Length 2.2 mm.

In build and size very similar to cxccUciis Kr., the antennae similarly constructed but less
stout. Head large, only slightly narrower than the thorax, the disc with a small impression,
extremely finely, sparingly punctured, distinctly coriaceous, but less strongly than in excellens.
Antennae with the 3rd joint as long as the 2nd, 4th to 10th transverse, the penultimate about
twice as broad as long. Thorax a third broader than long, the sides slightly rounded, more
retracted behind, the posterior angles rounded, feebly and broadly impressed in the middle
behind, the puncturation less fine than that of the head and closer, the ground sculpture
similar. Elytra a little broader and a third longer than the thorax, very finely, closely, asper-
ately punctured, the ground sculpture similar. Abdomen with the first four visible segments
transversely impressed at the base, very finely, asperately, rather sparingly ininctured especi-
ally behind, coriaceous. 8th dorsal segment l)roadly rounded. Tibia without setae.

Indian Tibet: Tsak-Shang, altitude 15,985 feet. 31-VIII-32. Amongst scanty grass
near stream. Unique.

Athela (Microdota) ladakiana sp. n.

Rather shining, entirely black. Antennae black. Legs pitchy black, the knees and tarsi
yellowish. Length 2.2 mm.

Larger, blacker and more shining than indubia Shp. the head larger, the elytra longer,
the thorax, elytra and abdomen much less distinctly punctured. Head large, suborbicular,
narrower than the thorax, the post-ocular region a good deal longer than the eye, practically
impunctate, distinctly coriaceous. Antennae very similar to those of indubia, the 3rd joint
distinctly shorter than the 2nd, clavate, 4th to 10th transverse, the penultimate twice as liroad
as long. Thorax about a half broader than long, the sides gently rounded, more retracted
behind, the posterior angles rounded, extremely finely, very sparingly punctured, the ground
sculpture as on the head. Elytra broader, more than a third longer than the thorax, extremely
finely, rather sparingly punctured, coriaceous. Abdomen a little widened towards the apex,
extremely iinely, very sparingly punctured on the anterior segments, almost impunctate
behind, coriaceous: 8th dorsal segment truncate. The whole insect with a fine yellowish
pubescence, closer on the fore-parts.

Indian Tibet: Tsak-Shang, above Tso-Moriri, altitude 15,985 feet, near a stream
amongst scanty grasses. 31-VIIT-32. A single specimen.

COLEOTTERA OF THE FAMILY STAPIIYLINIDAE 21

Athela (Dimetrota) hutchinsoni sp. n.

Moderately shining', black, the fare-parts with slight metallic reflex. Antennae and legs
black, the tarsi yellow. Length 3.2 mm.

In the broad head resembling cadaz'erina Bris. but differently colored, the antennae a
little stouter, the elytra longer and in this respect resembling marcida Er. Head broad, a
little narrower than the thorax, extremely finely, very sparingly punctured, distinctly coria-
ceous. Antennae with the 3rd joint as long as the 2nd, 4th and 5th slightly longer than broad,
6th as long as broad, 7th to 10th distinctly transverse. Thorax more than a third broader
than long, the sides gently rounded, the posterior angles rounded, along the middle behind
feebly and broadly impressed, very finely, asperately, much more closely punctured than the
head, the ground sculpture similar. Elytra almost twice as long as the thorax, with similar
but rather closer puncturation and similar ground sculpture. Abdomen a little narrowed
before the apex, finely, moderately closely punctured on the anterior segments, more sparingly
behind, less strongly coriaceous, and more shining than the fore-parts. Tibia without long-
setae.

3 : 8th dorsal segment truncate : 6th ventral segment a little produced, narrowed and
rounded.

Indian Tibet: Marsimik La, altitude 18.394 feet. 16-VII-32.

Ororotse Tso, altitude 17,381 feet. 16-VII-32. Two examples under .stones amongst
scanty grasses.

AleocJiara (Coprochara) bilincata Gyll.

Indian Tibet: Tsak-Shang, above Tso-moriri, altitude 15,985 feet. Amongst grasses.
31-VIII-32. A widely distributed species.

ARTICLE IV

REPORT ON COLEOPTERA OF THE FAMILY CARABIDAE

By H. E. Andrewes
(Received February 7, 1934)

The Carabidae collected by Mr. G. Evelyn Hutchinson in the course of the Yale North
India Expedition to Ladak comprised seventeen species, of which four are new ; an enumera-
tion of these will be found below with some notes on distribution, and following this, the
descriptions of the new species. As my Catalogue of Carabidae (part 18) in the series Cata-
logue of Indian Insects, published by the Indian Government, has so recently appeared, in
which full references are given for all the hitherto described Indian species, original refer-
ences only are given here. Mr. Hutchinson has kindly allowed me to retain the type specimens
of the four new species, which will ultimately be placed in the British Museum.

Of the seventeen species collected by the Expedition, six are also found in other parts
of the world, the remainder being endemic. All these six species are found also in Turkestan
and three of them are fairly comnKjnly distributed through the palaearctic region, one extend-
ing its habitat as far- as the western states of North America. While there are at least several
species of Carabidae common to the countries lying within the sandy belt which stretches
from Morocco to Sind, some reaching as far as the North West Frontier Province, there
is no evidence of them in this collection, the region explored being apparently both too distant
and at too great an elevation, but the connexion with the Central Asian fauna is more apparent.

It is interesting to note that the four genera found at high altitudes were also met with
by the Mount Everest Expedition at their base camp on the Rongbuk glacier (16,500 feet),
and one species, Amara brucei Andr., was taken in considerable numbers by both expeditions
up to 17,000 feet.

The enumeration of the species follows:

1. AV^m /'jaMWfO/'Mo Solsky in Fedchenko's Reise in Turkestan ii. pt. 5. 1874, p. 12;
• Andrewes, Fauna of Brit. Ind., Col. Caralx i. 1929, pp. 114 and 118.

Ka.shmir: Kiuhnus, Wular Lake, .SlOO feet, 17-IV-32, 1 ex.

The species has been met with in various parts of Kashmir, but is nol found fartlier
south; it seems to be fairly common in Turkestan.

2. Nebria limbigera Solsky in Fedchenko's Reise in Turkestan ii. pt. 5. 1874, ]>. 1.5;
Andrewes, Fauna of Brit. Ind., Col. Carab. i. 1929, pp. 114 and 120.

Tibet: Tso Nyak region 12-VIII-32 {Tsewang Tashi and Sonam Tergas), 2 ex.

Apparently a more widely spread species than the last, for it is found not only in
Kashmir, Tiljet and the northwestern provinces of India, but also in western China and
rather commonly in Turkestan.

Mem. Conn. Acad., Vol. X, Art. IV. September, 1934.

24 OOKKOPTKKA OF THE FAMILY CA U A IIIIIAK

3. Bciiibuiion phtto Andrewcs in Mi.ssit)n dans li-s rrovinccs Centrales de I'lndc et dans la
region occidentale de I'Himalaya, 1914, jiar (luy Balianlt, Caiab. I''i4, p. 72, t. I. f. o.
Indian Tibet : Tany-yar, 14,300 feet, 241-VI-32, 4 ex. ; Lukung, 14,000 feet, 8-VII-32,
3 ex.; Ign region, 12,000 feet, VIII-32, 3 ex.

First discovered in Rupshu and near Leh. The species seems to be almost confined to
that region, but Mr. II. (i. Champion found specimens in northern Kuniaon near the Tibetan
frontier, and he reports that they were "taken with (icodrouiiciis in running water."

4. Bembidion livens Andrewes, Trans. Ent. Soc. Lond. 1930, pp. 3 and 11.

Indian Tibet: Stream at Khalatse, 9600 feet, 29-V-32, 2 ex.; Iliniis, 21-V1-32,
3 ex. "under leaves or damp moss; dark soil." The two examples from Khalatse are in
poor condition and cannot be identified with certainty.

This species was described from specimens taken in Tibet at 10,000-12,000 feet, by the
Third Mount Everest Expedition, and is confined to the high regions of the Himalayas.

5. Bembidion bracciilaluiii Bates (?), Proc. Zool. Soc. Lond. 1889, p. 212; Andrewes, Ent.
Month. Mag. 1924, p. 194.

Indian Tibet : Matyan, 10,000 feet, 20-V-32, 4 ex.

Known only from Kashmir and Kumaon. The specimens found are a little smaller than
the type and have a greenish instead of a bluish tinge — as have the 'Kumaon examples.
There are some variations too in the colour of the basal joints of the antennae, of the tibiae
and the tarsi so that the identification is not altogether satisfactory.

6. Bembidion ixion sp. nov.

7. Bembidion hutchinsoni sp. nov.

8. Bembidion luntaka iVndrewes in Mission dans les provinces Centrales de I'lnde et dans
la region occidentale de I'Himalaya, 1914, par Guy Babault, Carab. 1924, p. 75.
Indian Tibet: Himis, 21-VI-32, 2 ex.; "under leaves or damp moss; dark soil";
Tang-tse, Mugleb, 14,000 feet, 27-VI-32, 2 ex.

Widely spread throughout the western Himalayas, and very variable in colour. In the
Tang-tse specimens the four pale spots on the elytra are very clearly marked, but those from
llimis are dark, with the pale spots barely visible.

9. Bembidion fuscicrus Motchulsky, Etudes Ent. iv. 1855, p. 79.

Pangur-tso, 14,200 feet, 14-Vni-32, 1 ex.; Tso-nyak region, 14,300 feet, VIII-32, 1 ex.

Found throughout Central Asia and Siberia, also in the western states of North America.
As in the case of so many widely spread species, there is great variability in the coloration;
the two examples in question arc very pale and hardly difi'er from specimens in my collection
from Montana and Oregon.

COLEOPTERA OF TilE FAMILY CARABIDAE 25

10. nniibidion varhim Olivier, Enc. Meth. v. 1790, p. 358.

Kashmir: Lokut Dal Lake, 5200 feet, 28-IV-32, 2 ex.; edge of Phashakuri near
Pampur, 7-V-32, 7 ex.

Widely spread throughout the palaearctic region, but not extending to America. In the
two examples from the Lokut Dal Lake the pale fasciae on the elytra are clearly marked, but
in those from Pampur the elytra are dark aeneous, with the pale markings barely visible.

11. Bembidion eupages sp. nov.

12. C/i/at'MHW /i'K/i(c;w/.«- Bates, Entomologist xxiv. 1891. Suppl, p. 9.
Kashmir: Kiuhnus, Wular Lake, 5100 feet, 17-IV-32, 1 ex.
Confined, so far as I am aware, to Kashmir and Kulu.

13. Stenolophiis cliscoplionts Fischer, Ent. Russ. ii. 1824, p. 141, t. 26. f. 9.
Kashmir: Kiuhnus, Wular Lake, 5100 feet, 17-IVr32, 1 ex.

Central Europe, the Mediterranean basin and western Asia as far as Turkestan; the
only Indian specimens seen were found in Kashmir, between 5000 and 6000 feet.

14. Amara hrucei Andrewes, Ann. Mag. Nat. Hist. (9). xi. 1923, p. 276.

Indian Tibet : Ororotse-tso, 17,400 feet, ll-VII-32, 9 ex. ; Anem La 17,000 feet, 3 ex. ;
Tso Nyak region, 14,300 feet, VIII-32, 1 ex.

First discovered by the Second Mount Everest Expedition in 1922 at the base camp,
16,500 feet, where it was taken in considerable numbers. The Third Mount Everest Expedi-
tion in 1924 met with it again, not only at the base camp, but at various other Tibetan locali-
ties. Mr. H. G. Champion took some specimens in northern Kumaon and again near the
Supi River in Til)et. Tiiis is the first time it has been recorded from so far west. It seems
to be common where it occurs.

15. Cyniindis viannerhehm Gebler, Bull. Ac. Imp. Sci., St. Peters)). 1843, i, p. 36.
Indian Tibet: Tang-yar, 14,300 feet, 24-IV-32, 2 ex.

Not nncommnn in Kashmir, tin- Pamirs and (Jcntral Asia.

16. Cyniindis clianipioiii Andrewes, Ann. Mag. Nat. Mist. (10). ii. 1928, ]>. 589.

Tibet: Tso-nyak region, 14,300 feet, VIII-32, 1 ex. which does not quite agree with
type form, the pattern on the elytra being very indefinite.

Northern Kumaon and Til)et.

17. Cymindis rubriceps sp. nov.

26 COLEOPTERA OF THE 1 AMII.Y CAKAlilDAE

Bembidion ixion ^^l). iici\ .

Lengtli : 4.5 niin.

Piceous beneath, liead and prothorax aeneous, elytra black with a very faint metalhc
tinge and a vague dark red apical spot on each ; joints 1 to 3 of antennae, tibiae, and tarsi
more or less ferruginous.

Head with fairly deep parallel furrows, eyes moderately prominent, antennae reaching
basal third of elytra, surface impunctate. Prothorax convex, subcordate, not (luite a third
wider than head, two-fifths wider than long, base slightly arcuate, as wide as apex, sides
rounded in front, slightly sinuate behind, hind angles sharp, but a little obtuse, each with a
short but fairly sharp carina, bounding the small deep rounded foveae on the outer side;
median line and front transverse impression moderately deep, basal area depressed, finely
but not closely rugose-punctate. Elytra convex, ovate, not quite a half wider than prothorax
and not quite three-quarters longer than wide, bjrder extending inwards a little be\ond stria 5,
shoulders visible but not prominent ; punctate-striate, the inner striae moderately impressed
on disk, much less so at sides, 7 represented by a row of minute punctures, all (except 1)
evanescent towards apex, but 2 is impressed quite close to apex, scutellary striole and apical
stria only moderately developed, intervals a little convex on disk only, dorsal pores fairly dis-
tinct, adjoining stria 3, just before middle and at threc-fnurlhs. Microsculpture of the elytra
consisting of fine transverse lines, which form meshes quite three or four times wider than
long, none on disk of prothorax or head. Metasternal process Ixirdered.

Indian Tibet : Matyan, 10,000 feet, 20-V-32, 1 ex. S.

The species is smaller than bracculatum Bates, but dilYers mainly in the form of the
prothorax, the sides of which are only faintly sinuate behind, while the basal foveae are
small, deep and rounded, adjoining the carina.

Bembidion hutchinsoni sp. nov.

Length : 4.6-5 mm.

Piceous beneath, blue-black abuve ; palpi, joints 1 to 4 of antennae, and legs ferruginous;
basal two-fifths of the elytra dark red, apical fourth pale ferruginous, so that there is a dark
band across them, the outline of which is vague and somewhat variable.

Head with deep uneven furrows, converging very slightly in front, eyes prominent,
antennae slender, reaching basal third of elytra, surface punctate at sides behind, and with
a few scattered punctures on disk. Prothorax convex, cordate, about a sixth wider than
head, a fifth wider than long, base slightly ol)lique at sides, hardly wider than apex, sides
well rounded in front, sinuate a litle before ba.se, hind angles right, fairly sharp and with a
slight carina: metlian line and the small basal foveae moderately deep, transverse impres-
sions shallow, I)asal area a little depressed and finely punctate, disk w ith some slight trans-
verse triae. Elytra subquadrate, moderately convex, a half wider than prothorax, nearly
two-thirds longer than wide, shoulders evident, border reaching stria 5 ; striae clearly punc-
tate, moderately impressed on disk, more lightly at sides and behind, though in some speci-
mens (including type) clearly impressed to apex, scutellary striole and apical stria both
present but rather slight, intervals a little convex on disk, dorsal pores very distinct, on
stria 3, at about a third and two-thirds. Microsculpture of the elytra consisting of very fine

COLEOPTERA OF THE FAMILY CARABIDAE 27

transxcrse lines, w liicli form very wide meshes, none on disk of protliorax or head. Meta-
sternai process bordered and witli a transverse groove behind it.

I.N'DiAN Tibet; Kyam, 15,630 feet, 20-\' 11-32, 6 e.x., taken round the edge of a hot
spring.

The subgenus here is Pcryphus and the species will be readily recognized by the colora-
tion of the elytra.

Bembidion eupages sp. nov.

Length: 2.9-3.1 mm.

Colour Ijlack beneath, upper surface aeneous or blue, and very shiny palpi, joints 1 to 3
of antennae (rest fuscous), apex of elytra and venter, trochanters, tibiae, and tarsi
ferruginous.

Head with fairly deep, nearly parallel furrows, converging a little on clypeus and
diverging behind, eyes prominent, antennae short, submoniliform, surface impunctate. Pro-
thorax convex, cordate, slightly wider than head, nearly a third wider than long, base truncate
at middle, very oblique at sides, narrower than apex, sides strongly rounded in front and
contracted behind, with a rather wide marginal channel, sinuate close to the hind angles
which project on each side as a siuall sharp though slightly obtuse tooth, bounded by a short
oblique carina, within which are the small deep rounded f oveae ; median line very fine, trans-
verse impressions evident, the hind one with a few deep punctures, and one or two generally
transverse punctures along the basal margin, surface otherwise impunctate. Elytra moder-
ately convex, subquadrate, a half wider than prothorax and a little more than a half longer
than wide, shoulders square, border reaching stria 4; striate-punctate, striae 1 and 8 only
impressed, the remaining striae formed by the punctures, which are as clearly marked in the
outer as in the inner striae, but 2 to 7 all disappear behind, scutellary striole formed by rather
large punctures, apical stria wanting and its pore consequently isolated; intervals flat, 1 raised
behind, dorsal pores on interval 3 at about a fourth and three-fifths, a fine clavicular fold
present. No microsculpture. Metasternal process bordered, and with a transverse groove
behind it.

Kashmir: Lokut Dal Lake, 5200 feet, 28-IV-32, 6 ex.; edge of Phashakuri near
Pampur, 7 v. 1932, 3 ex. In the Indian Museum, Calcutta, there is a single specimen labelled
"Kashmir," taken by Mr. H. T. Pease in 1915.

The species fits fairly into the palaearctic subgenus Eniphanes.

Cymindis rubriceps sp. nov.

Length : 7.8 mm.

Colour piceous : head, prothorax, the latter with a vague dark area on each side of disk,
and a small area around scutellum and extending backwards nearly to middle along suture,
dark red; palpi and joints 2 to 11 of antennae ferruginous; joint 1 of antennae, legs,
epipleura, and an elytral pattern more or less flavous. The pattern on the elytra comprises
inter\'al 9, a humeral vitta on intervals 5 to 7, not quite reaching middle, and a small spot
near apex on intervals 3 to 5, extending a little backwards on 5.

Body covered with a fairly long pale pubescence, scanty on the head and prothorax, a
little denser on the elytra, though not concealing- the shiny surface.

28 COI.EOl'TMKA OK THE FA.MILV CAKAr.IDAE

Ifrud wide, convex, with a slight neck constriction, frontal fovcae slight, eyes large Init
liardly prominent, genae sloping gradually to neck, antennae harely reaching basal fourth
of elytra, palpi not dilated, surface moderately punctate along sides and across vertex, disk
and neck smooth. ProtJwrax moderately convex, a sixth wider than head, a fifth wider than
long, base arcuate, a little narrower than apex, sides rounded and reflexed, sinuate quite close
tci the hind angles, which are sharp though a little obtuse, almost dentiform; median line
and front transverse impression moderately deep, basal foveae also fairly deep, rounded,
adjoining the angles, surface moderately punctate, more densely in the foveae, more sparsely
on the disk. Elytra rather fiat, subovate, three-fourths wider than prothorax, about a third
longer than wide, widest behind middle, basal border entire, apex truncate ; striae moderately
impressed and very finely punctate, intervals finely, irregularly, and not very closely punc-
tate, a slight depression on each side on the front of disk. No reticulate microsculpture is
present, but the surface of the elytra is vaguely and micrcscopically rugose-punctate.

A little smaller than C. championi Andr., and somewhat differently coloured. The pro-
thorax is less contracted loehind, with much less conspicuous hind angles ; the elytra are
shorter, w^ith a similar pale humeral vitta, but a slight though quite distinct apical spot
as well, the surface is more finely and more closely punctate, without any reticulate
microsculpture.

Indian Tibet: Anem T,a, 17,000 feet, I-\'IIi-32, 1 ex. S.

ARTICLE V

REPORT ON PHYLLOPOD CRUSTACEA (ANOSTRACA, NOTOSTRACA AND
CONCHOSTRACA) INCLUDING A REVISION OF THE ANOSTRACA

OF THE INDIAN EMPIRE

By Richard M. Bond

(Received February 19, 1934)

INTRODUCTION

The Phyllopoda of the Indian Empire have received sporadic attention since the time of
Baird, who, in 1860, described Streptocephalus diclwtoiiiiis from a single male specimen which
was found swimming in a pail of milk. Since that time Sars, Gurney, Daday and some
others have added to the knowledge of the Phyllopoda of the region. Professor G. E.
Hutchinson has kindly turned over to me for examination the collections of tliese animals
that he made as Biologist of the Yale North India Expedition.

The Notostraca and Conchostraca taken by tlie expedition are few in number, and it
seems wise, in these groups, to limit this treatment to tlie forms in this collection. In the
case of the Anostraca, however, the collections brought back are much more complete, and
for this reason, and because of the ecological and zoogeographical importance of the group,
it seems proper to treat them at greater length.

In this undertaking I was greatly aided by Dr. Hem Singh Prutlii, who secured for me
the loan of all the unidentified Anostraca in the Indian Museum in Calcutta, in addition to
sending me named specimens of certain forms. Records based on this material are marked
with an asteri.'^k (*) throughout tlie present paper. This loan material in addition to tlie
Y. N. I. E. collections has given me an opportunity to compare a larger series of speci-
mens of certain of the species than has probably been assembled hitherto. As a result, I
have raised a "variety" to full specific rank, described 3 new subspecies, and am able to
record for the first time the occurrence of a species in Kashmir hitherto found only in
Mongolia and Manchuria.

In the descriptions of the larger groups, such as families and genera, I have frequently
borrowed, almost verbatim, from the clear, concise paper on the South African Phyllopoda,
by Barnard (1929), to whom I am much indebted.

Subclass BRANCHIOPODA

The classification used in this paper is not only perfectly defensible on purely morpholog-
ical grounds, but has the added recommendation that it follows ecological as well as struc-
tural lines.

Body uniformly segmented, usually elongate, usually ending in a caudal furca; without
carapace, with a dorsal shield-like carapace, or witli a bivalve carapace. Compound eyes

Mem. Conn. Acad,, Vol. X, Art, V. September, 1934.

30 PIIYI.I.OPOD CRUSTACEA

present, and usually a persistent median eye. Five to 19 (in living forms) pairs of trunk
limbs, which are simple foliaceous, modified foliaceous, or (rarely) pediform. Two pairs
of antennae and 2 pairs of maxillae present, the 1st antennae and 2nd maxillae usually much
reduced.

Order PHYLLOPODA (Euphyllopoda)

Branchiopoda with 10 or more pairs of trunk limbs all simple foliaceous, or with the
anterior 1 or 2 pairs somewhat modified for clasping the 2 or as tactile organs. Develop-
ment (with the single exception of Cydcstheria liislopi) always with a metamorphosis from
a free-swimming nauplius or metanauplius stage. The heart has several pairs of ostia.

Suborder I ANOSTRACA

Phyllopoda with an elongate body and without carapace. With 11 to 19 pairs of simple
foliaceous trunk limbs. Paired pedunculate compound eyes, a median ocellus in front. First
antennae small, 2nd antennae large and modified for clasping in $ . Fight or 9 post-
pedigerous (abdominal) segments, the first 2 of which bear the external genital organs and
may be partly fused. Caudal furca when present never segmented. Paired eversible penes
in $ ; ovisac formed by united oviducts in 9 in which ova are retained. Young hatch as
nauplii or metanauplii. (In this group the rami of the caudal furca are usually known as
ccrcopods.)

Key to flic Faiiiilirs and Genera of Anostraca of the Indian Empire

1. 2nd antennae oi S biarliculate

A. Basal joints f)f 2n(l antennae of <? nearly or entirelx' se])arate

i. Basal joint of 2nd antennae of $ with no ])rocesses, or with small

and simple ones only Ikj iimiipah

a. Distal joint of 2nd antennae of $ greatly flattened Irtciiiia

b. Distal joint of 2nd antennae of $ not greatly flattened Branchinecta

ii. Basal joints of 2nd antennae of S bearing 1 or more consiiicuous

fleshy processes Chirocephai.wae

a. (only Indian genus) Pristieephalus

B. Basal joints of 2nd anteimae of $ firmly joined to each other and to

the front of the head to form a clypeus Braxchifodidae

i. Front of head of S bearing 2 long filiform processes which are

connate at base Brancliipiis

ii. Front of head of i without filiform processes Branehipodopsis

2. 2nd antennae of $ triarticulate, cheliform Streptocephai.idae

A. (only genus) Sfreploeephaius

PHYLLOPOD CRUSTACEA 31

Family ARTEMIIDAE Grochowski

1896 Arlciiiiiilae Grochowski. Verh. z(_)ol. bot. Ges. Wien, 45:99

Eleven pedigerous, 8 or 9 postpedigerous segments. Head without frontal process.
Second antennae of 3 biarticulate, not fused at base, or only slightly so. Legs with a single
epite (branchial lamina). Cercopods jointed to last abdominal segment, or fused to it or
absent. Ovisac subglobular or cylindrical. Distribution world wide.

Genus Artemia Leach
1819 Artemia Leach. Diet. Sci. Nat., 14:543

Body slender, abdomen often longer than trunk and head combined. Eight postpediger-
ous segments, the last one longest. Basal joints of 2nd antennae of $ slightly fused; inner
margin with a small round setulose knob. Distal joint of 2nd antennae of i much flattened,
apically acute. Intromittent part of penes without spines. Cercopods movable, fused to last
body segment, or absent. Body form more or less variable according to the salinity of the
environment.

Daday (1910) reduced the many "species" which had l)een described to two, one of
which was from Peru, and was placed in the subgenus CallaoncUa. But the "species"
described by Daday as salina, of wide distribution, has been shown to be heterogeneous by the
work of Artom (1906, 1911a, 1911b, 1912, 1922, 1926, etc.), who found that there are at
least two types, diploid and tetraploid, which he distinguishes at various times as "univalens"
and ''bivalens" (1911b) or as "micropirenica" and "macropirenica" (1922), and these may
be further divided into sexual and parthenogenetic subraces. Hertwig (1931) and Gross
(1932) believe that Artom's "diploid se.xual" Artemia really are diploid, but that his "diploid
parthenogenetic" are tetraploid, and that his "tetraploid parthenogenetic" are octoploid.
Their Ix'licf is based i in tlie fact that tlie chromosomes of the parthenogenetic races are much
larger, each chromosome i)ri)1)alily being bivalent. The most recent review of the situation is
by Stella (1933), who agrees substantially with Artom.

Now, the taxonomic value of these races has never Ijeen properly established, since
Artom did not give formal descriptions, but used his terms rather as conveniences. Daday's
species salina will undoubtedly have to be divided eventually on cytological and grosser struc-
tural grounds, but however the division is made, the name salina will have to be reserved for
the form originally described under that name. This form (of which I have a few specimens
from the type locality) was" found in the salt pans at Lymington, England, and has been
shown to be diploid and sexual — the type called "univalens" and "micropirenica" by Artom.
Fortunately, for the sake of simplicity, this is also the only form that has so far l^een found
in the Indian region.

32

PHYLLOPOD CRUSTACEA

Artemia salina (Linnaeus)

1758 Cancer siiliiuis Syst. Nat. (10th ed.) 1:634

Locality: Indian Tibk.t: Tso Kar. 200+ S S 9 9. 5-1X-32.

Reported from : Nowhere else in the Indian region, but it has been taken in a great numljer of
localities in central and western Asia, as well as in other parts of the world. The Tso
Kar colony may have come from eggs dropped by a caravan carrying salt, though there
seems to be no reason for its not being fnund naturally at several localities in the Indian
region.

Types : Ubi ?

Figure 1. — Artemia salina. A, head of Tso Kar S from above. B, head of Tso Kar 9 from in front. C, end
of abdomen of Tso Kar $ from above. D, end of abdomen of $ raised from Tso Kar eggs in 10% Na CI solution.
(Heads X 23, abdomens X 46.)

PHYLLOPOD CRUSTACEA 33

Always sexual, usually a more or less even distribution of the sexes (in this case
about twice as many $ S as 9 9). Nuclei of the segmenting egg with 42 very small
chromosomes. Nuclei of the ova before emission of polar bodies with 21 diads (observed in
some of these specimens). Differences between specimens raised in brines of different densi-
ties not so marked as in the parthenogenetic Artemia; the caudal furca is never entirely
absent, even in specimens from the strongest brines.

In the Tso Kar^ specimens the abdomen is consistently about 20% longer than the trunk
and head combined; the furca is somewhat reduced and bears from 3 to 10 setae on each
ramus. The mature 9 9 carry from to 40 eggs (average 17.2). The S 5 average
9.96 mm., and the 9 9 11.02 mm. in length.

Professor Hutchinson brought back some viable eggs from Tso Kar which it has been
possible to raise in the laboratory, though so far only in brines more dilute than that of the
lake. As a consequence, the laboratory-raised specimens show better developed furcae and
relatively shorter abdomens than those preserved in the field.

Genus Branchinecta Verrill
1869 Branchinecta Verrill. Am. Jour. Sci. (ser. 2) 48:250

Nine postpedigerous segments, the last usually shortest. Basal joints of 2nd antennae
of <J perfectly separate; unarmed, or bearing 1 or more small processes of spines. Distal
joint of 2nd antennae of $ usually simple, falciform; triangular, oval, or subcircular in
cross section. Cercopods always jointed to last abdominal segment and freely movable.
Ovisac of 9 usually cylindrical, though very short in some species.

About 10 species are known from North and South America, Europe, and Asia. Only 1
species reported from the Indian region, though B. paludosa is found in Siberia to the north,
and B. fcrox is found east to Odessa and Jerusalem.

Branchinecta oriciitalis Ci. O. Sars

1901 Branchinecta orientalis. Sars. Ann. Mus. Zool. Acad. Imp. St. Petersbourg. 6: 144

Localities: Tibet: *Gyantse (coll. Maj. F. M.Bailey) 1 <5 , 2 9 9. 2-VII-23.

Indian Tibet: Chushol, Western Tibet, pond below village. Altitude 4336
meters. About 20 5 5 9 9 . lO-VIII-32.

Lake near Chushol. Altitude 4491 meters. IS. lO-VIII-32.
Togarma Tso. Altitude 5217 meters. 7 <5 3 , 2 9 9 . lO-Vn-32.

Reported from: Hungary, Kecsemet; Russia, Charkov; 4 localities in the Pamir region
(sec. Daday) ; Eastern Mongolia, Chuntu-nor (sec. Sars and Daday) ; Russian Mon-
golia (sec. Smirnov) ; Tibet, Gyantse (sec. Gurney).

Abdomen about the length of the head and tnink or a little longer, in both sexes.
Mandibles with a sharp dentiform process on the posterior comer of the chewing sur-

' An analysis of Tso Kar water sliuvvs the following (figures are mg. per liter) : Total solids 79266; SiO= 25;
Fel.8; Al 5.2; Ca406; Mg2716; Na 16346; K5478; HCO3 2141 ; SO. 35075;' CI 11662.

34

PHYLLOrOn CRUSTACEA

face. The iiul maxillae are pmvided with more setae than is usual in the order. ]'.\nte of the
swimming letjs of jjairs 1-10 creniilate alont^ the hnrder; iL^ill un leijs of pairs 1-10 with
margin entire. Last pair of appendages in both sexes with gill reduced in size and with
a setose end; and with an epite the end of which is deeply notched. In <5 , the basal joint
of the 2nd antennae is stout and cylindrical, with a slight setulose or smooth bulge on the
medial face near the base. Distal joint of 2n(l antennae snidoth, unguifnrni, only slightly
curved. The S genital inmch has 2 pusterinrly directed processes on each side. The dorso-

FiGURE 2. — Brainliinccta oricnlalis. A, head of Chushol i from above. B, head of Chushol 9 from in front.
C, end of abdomen of Chushol i from above. (Heads X H. abdomen X IS.)

lateral process haniifcirm with a ventrally directed side-process. The ventromedial process
cylindrical. The cercopods of the & slender, narrow, and pointed, straight, or slightly curved
outward at the tips; both margins fringed with plumose setae nearly to the l)ase. Tn 9 2nd
antennae Hat and blade-like with a well-marked acute terminal point, with a well-marked notch
on the inner margin lietween the point and the body of the antenna. Ovisac does not reach
beyond the .?rd post-genital (S{\\ post-pedigennis ) segment. Length: <J 12-v38 niiu. ; 9
12-43.5 mm.

Daday (1910) divides this species into a small "forma ^TrmiHs" and a larger "forma
aestivalis," but since all intermediate sizes arc foinid and at all times nf the breeding season,
it seems unnecessary to make this distinction.

This species most closely resembles B. fero.v (Milne-Edwards), and in fact l)oth Daday
(1910) and Smirnov (1932) are a little dinil>tfiil as to whether the two species are really
distinct. It appears to me that they very i)robably are not the same, though they are certainly

I

I'lIYLLOrOD CRUSTACEA 35

closely related. B. fcrox is proportionately much more slender, and averages considerably
longer; the gill of the last pair of limbs appears not to be setiferous (from the descriptions —
I have no specimens at hand) ; the cercopods of the i are always outcurved, and their outer
margins bear setae only near the tips ; in the 2 there is not a well-marked notch between the
body of each 2nd antenna and its apical point; and the ovisac extends beyond the 3rd post-
genital segment. Moreover, B. fcrox has not Ijeen reported from central and eastern Asia,
as it should have been if only environmental variations separate the two. B. oricnfalis has
been reported from Russia and even Hungary, but both these records (especially the latter)
I consider very doubtful. From analogies with the development of other anostracans, it
appears very likely that adults of B. oricnfalis will resemble somewhat juvenile B. fcrox even
more closely than they resemble the adults of that species. From some of Daday's figures
especially it seems possible that he has confused young specimens of B. fcrox with B. oricntalis.

Family Chirocephalidae Daday
1910 Chircoccl^halidac Daday, Ann. Sci. Nat. (ser. 9) 11:175

Eleven pedigerous, 9 postpedigerous segments. S with biarticulate 2nd antennae, with
separate basal joints. In $ basal joints of 2nd antennae bear 1 or more fleshy processes; or
if not, the head bears a median frontal process; or there may be a frontal process as well as
fleshy processes on the basal joints of the 2nd antennae. Legs with 1 or 2 epites. Cercopods
movably articulated with last abdominal segment (except in 'riianiuoccpJntlus) . Ovisac
usually more or less flask-shaped. Distribution world wide.

This is probably the least homogeneous of the families of Anostraca as defined by Daday.
This author further sulidivides it into 3 sub-families (which will not be treated here), but
even with this division certan genera assigned to it by Daday will probably have to be
removed to other families when they are more fully studied. The single Indian genus is close
to Chiroccphaliis, and will certainly remain in the same family, whatever the taxonomic
future of the group as now defined.

Genus Pristiccfluiliis Daday
1910 rristiccphalus Daday, Ann. Sci. Nat. (ser. 9) 11:213

^'Vbdomen without furca,_usually shorter than trunk. Alxlomen of £ unarmed, in ?
bearing various sorts of spines, usually at posterior margins of the segments. Margins of
cercopods setiferous, never spiniferous. Male without frontal process. Basal joint of 2nd
antenna of S often with a subspherical or cylindrical setuliferous process, and always with a
pointed serriform process which is generally carried more or less coiled. Legs wilh 2 epites,
except that last pair may have only 1, or the proximal eijite of last leg may be much reduced.

Four species are known, occurring in parts of North Africa, Europe, Western and Cen-
tral Asia. The species most closely resembling the one found in the Indian region is P. jose-
phinae, which is found in Eastern Russia and in Siljeria, and hence is the nearest geographi-
cally as well.

36

PHVLLOPOD CRUSTACEA

Pristiccpholiis priscu^ Daday
1910 Pristiccphalus prisciis Daday, Ann. Sci. Nat. (ser. 9) 11:224

Localities: Punjab: Sargodhar District, 3 miles South of Nuriwala. Altitude circa 305
meters. 4 $ S . 6-1 11-32.

Simla Hii.i. St.\te.s: *Bet\veen Theog and Matiana. Altitude c/rra 2300 meters.

5$ S , 29 9. Coll. S. Kemp. X-21.

*Below Kupri. Altitude circa 2200 meters. 62 S S 9 9
(all slightly juvenile). Coll. S. W. K. 28-IX-21.

Reported from: Naini Tal, Kumaon; Phagu, Simla Hill States; Suka Tal, above Naini Tal,
Kumaon; Ehovvali Bazar, Kumaon (sec. Daday).

Types: Daday designates no types for any of his species, but he bad specimens of P. priscus
from both the Paris Natural History Museum and the Indian Museum.

Figure 3. — Prisficcl^halits /•ristiis. A, head of Sargodhar i from above (X 15.6;. B, head of Theog 9 from
in front (X 15.6). C, external genitaHa of Sargodhar S from below (X 33). D, egg sac of Theog 9 from below.
E, same from left side. F, end of abdomen of Sargodhar 3 from above ( X 42) .

PIIYU.OPOD CRUSTACEA

37

Tliis cliaracteristically Indian species has not been described nor figured except Ijy
Daday. The specimens tliat 1 have examined agree exactl}' on all important points with
Daday's description and, moreover, the specimens (except those collected by Professor
Hutchinson) are from the same region as Daday's, so that there can be no question of
subspecific or varietal differences. None the less, there are a considerable number of small

Figure 4. — Pristket'halus prisms. A, right 2nd antenna of Sargodhar $ from above (X23). B, right Ist
maxilla of Theog $ (X 38, enlargement X 85). C, right 2nd maxilla of Kupri S, finer setae not shown (X64).
D, E, F, 1st, 6th, and 11th legs of Sargodhar S- Offset from D, flabelkim of same leg of Kupri S. Inset in F,
gill and epite of same leg of Theog $ (all X 22).

points in which these specimens differ from Daday's description, and there are several charac-
ters which Daday seems entirely to have overlooked. His descriptions are in general unnec-
essarily detailed, and to correct all his observation it is needful for me to be very lengthy in
my description also.

Male: Penultimate abdominal segtnent longer than any of the preceding 4. Last
abdominal segment (which is about half as long as the penultimate segment) sometimes
rather deeply notched between the cercopods. Cercopods long, narrow, ensiform; distal
end more or less acutely pointed; fringed all nmnd with moderately long plumose setae
(Figure 3, f).

38 I'lIVl.I.npOn CRUSTACEA

I lead naiiulcd in fmnt. 1st antenna Inarticulate, considerably longer than the basal joint
of ind antennae (bii^ure 3, a), liasal joint of the 2nd antenna roughly 'i as broad as
lonjj, rouyhlv kei;-sha])C{l ( l-'iLTiTe 4, a). A sliijhtly raised area on the outer, distal margin
of the basal joint nf the _'nil antenna may be minutely setulose (not shown in figure); and
there may be a short, ill-defined, transverse ridge, on the lateral side of this joint near the base,
bearing 10-12 slender setae. Distal joint of second antenna with sul>-conical basal portion,
becoming flattened distally. Outer margin of distal portion is a flattened arc; inner margin
sinusdid, and minutely serrate, with the ])oints uf the serrations direi'ted basally. On dorsal
interior surface of basal joint of the 2nd antenna is a pointed process calleil ])y Dada)' the
"serriform process"; it is taeniform with tiie distal end drawn out; the margins are entire;
a row of short digitiform papillae parallels each margin on the ventral surface. In preserved
specimens, the serriform process is usually spirally twisted.

Chewing surface of mandibles in shape of a rough parallelogram, about 30-35 rows of
teeth, the teeth being directed anterior!}'; ;U dorsal edge of chewing surface are a few large,
conical s])ines. 1 have been unal)le to detect any trace of the mandil)ular palp (which in
several other jihyllopod genera is represented by a small papilla). 1st maxilla broad and flat
distally, ending in a row of 15-16 long, plumo.se, biarticulate setae. l>asal portions of the
setae armed with distally directed spines which number 1 or 2 on the lowermost seta and
increase in number uj) to 8-12 on the upi)ermost seta. These spines are on the side of the
setae opposed to the setae of the opposite 1st ma.xilla. Lower than the lowermost seta is a
small spine which appears to be morphologically a much reduced seta, a.s it is supplied with its
iiwn tendon ( h'igure 4, b). 2nd maxilla reduced as is usual in the sub-order, ending in a
large, slightly cur\-ed claw armed with a few minute spines; provided on its medio-anlerior
tidge with three strong, ])iarticulate, jjlumose setae: on ventral surface, lielow bases of setae is
a short, stout spine directed distally (posteriorly). Several patches of extremely fine hairs on
the 2nd maxilla are much too flne to be shown in the figure (JMgure 4, c).

Swimming legs 1-10 with 2 epites with markedly serrate edges. Last pair of legs vari-
al)le in this respect, having 2 sul^equal, narrow, pointed epites, or with the proximal epite much
the smaller, or entirely absent. Last legs of same individual may be unlike in this respect.
Margin of the distal endite of legs 7 and 11 tends to lie bluntly ])ointed ; of legs 2-10 more
evenly rouudetl. (iill with entire margin on all legs. I'^labellnm on legs 2-11 foliaform. with
<lorsal margin flatter than ventral. Mabellum of 1st leg foliaform, or subtriangular in out-
line. (I'igure 4, d, e, f, setae and sjiines shown only for 2 distalmost endites.)

Each side of genital sac with 3 jirocesses directed posteriorly. N'entral ])rocess ends in a
lappet much flattened horizontally, end ol)Ii(niely truncated, sometimes much more so than in
figure, inner dorsal ])rocess digitiform, sometimes slenderer than shown in figure. Outer
dorsal process (penis) longer and nuuli thicker than others, a])proximately cylindrical, ends
in a flat, subtriangular jilate which is eversible and retractable. 'I'lie outline of the plate is
roughly that of a boot viewed from the side, hollowing this analogy, the plate is attached
by the leg, and bears 3 teeth on the top of the toe. These teeth are absent in juvenile
individuals (Figure 3, c).

Total length variable, perhaps depending on season and food sup])ly, averages about
18-1 "J mm. from forehead to end of cercopods.

Female: Second to 11th pedigennis segments with short transverse ridge across median
dorsal line; viewed from the side, the highest point of ridge is towards posterior margin of

PHYLI.OPOD CRUSTACEA 39

segments, especially posteriorly; at each end of each ridge are 1-10 curved spines along the
posterior margin of the segment, the 2nd pedigerovis segment having 1 spine on each side,
and the numl)er increasing- posteriorly ( l-"igure 3, e). According to Daday the 3rd to 7th
ahdominal segments have a girdle of spines around the posterior margin. In the specimens I
have examined, these segments hear from about 6-25 spines on the posterior margin, increas-
ing in numbers per segment posteriorly to a maximum on the 5th or 6th segment. The
spines do not form a girdle, but tend to occur in groups of 2 or 3 with a space between the
groups. The spines are absent, or small and few near the ventral mid line, thus giving an
effect quite unlike Daday's figure (F"igure 3, e). Cercopods as in the male.

Head smoothly rounded in front. First antenna biarticulate and hjuger than 2nil
antenna. Second antenna oval in cross-section, tapering abruptly at distal end, to a pointed
spine-like process (Figure 3, b). Eyes smaller than in male. Mouth parts as in male.

Legs as in the male.

Ovisac short and broad; ventral view something like a beef heart, but with an obtusely
conical process on each side of the posterior end, and slightly dorsal to it (Fig-ure 3, d, e).

Dimensions about the same as in the male or slightly smaller.

All of the specimens that I have examined are heavily infested with epiphytes and
attached protozoa. This suggests that growth is very slow, with long periods lietween
ecdyses.

Family Branchipodidae Daday
1910 Branchipodidae Daday, Ann. Sci. Nat. (ser. 9) 11:287

Eleven pedigerous, 8 or 9 postpedigerous segments. Front part of head of S fused with
basal joints of 2nd antenna to form a clypeus. Front of head of S unarmed, or with a
median process, or with paired processes. Second antenna of $ biarticulate. Legs with
1 epite. Cercopods freely movable, or fused to last body segment. Ovisac generally sub-
gliibular. The familv is absent from North and South America.

Genus Braiichipiis Schaeffer-Daday

1766 BrancJiipus Schaeffer. Elementa Entomologica

1910 Braiichipus Daday. Ann. Sci. Nat. (ser. 9) 11 : 311

Trunk segments smooth, unarmed. I'ostiiedigerous segments unarmed in l)oth sexes, or
with short digitiform processes in i. Cercojjods movably articulated with last body seg-
ment; straight and fringed all round with setae; or curved inward with outer margin setif-
erous, inner spiniferous. Distal joint of 2nd antenna of £ nuuh lunger than basal joint;
falciform, curved inwards. Clypeus of $ with paired short blunt frontal processes. Paired,
long filiform processes with bases connate arise from front of head (dorso-proximal part
of clypeus). Second antenna of 5 flat, blade-like, produced into a sharp a])ical point. Ovisac
short, oval, with a prominent ventral lolse. In the present-day restricted sense, this genus
contains only 2 species, one of which has been found only once, in the French Alps.

40

rnvi-LoroD Crustacea

N. nniihhipiis slai:^niilis (Linnaeus)

1752 Apus piscifoniiis Schaeffer. Abhandl. v. Insecten. vol. 2.

1758 Cancer stagnalis Linnaeus. Syst. Nat. (lOtli ed.), p. 634.

1766 Branchipus pisciformis Schaeffer. Elementa Entomologica.

1906 Branchipus piscifoniiis (iurney. J. and Proc. As. Soc. Bengal (n.s.) 2:275.

1910 Branchipus stagnalis Daday. Ann. Sci. Nat. (ser. 9) 11:312.

Localities: No specimens of this species have come into my liands. It lias been reported only
once from India, by Gurney (1907), who examined specimens in tlie Indian Museum
labeled "J. A. W. Alurray, Sind."

Reported from: The greater part of Europe; North Africa; Palestine. The collections
nearest India were made at Sudak in the Crimea, and Bingol Dagh in Armenia. (Cf.
Daday (1910).)

Types : Ubi ?

Figure S. — Bninchil>iis xla^iuilis. A, lit-ad of $ from above. B, head uf i frcmi Ijelow. C, head of 9 from
above. D, tiid of al)domen of £. !■", cgs sac of 9 from right side. F, end of abdomen cjf 9. (All fnim Daday
(1910) ; magnification unknown.)

Since I have seen no specimens of this species, 1 borrow the descri])tion (much short-
ened) from Daday, as well as some of his figures.

Male: Size very variable according to locality. Abdominal segments unarnied. Cerco-
pods falciform, curved inward, outer margin setiferous, inner margin bearing slender spines.
Clypeus with a short conical process on each side dorsally ; witii paired conical frontal proc-
esses; with a conical tubercle on each side, and a distal digitiform process on each side

PIIYI.LOPOD CRUSTACEA 41

ventrally. Distal pnints uf the 2n(l antenna with l)itul)crculate tips, and witli a digitiforni
process projecting antemlateraliy from the \'entr()lateral niar.t;in at a point slightly distal to
the middle of the joint. Front of head at dorso-posterior margin of clypeus with a pair of
long filiform processes with connate bases. Total length, 8-20 mm.

Female: Cercopods straight, both margins setiferous. Front of head unarmed, gently
rounded. Ovisac short, oval, acutely rounded posteriorly. Total length, 8.5-23 mm.

Genus BrancJiipodopsis G. O. Sars
1898 Branchipodopsis Sars. Arch. Mat. og Naturvid. Krist. 20 (4) : 26.

Nine postpedigerous segments, the last shortest. Cercopods falciform, incurved, mova-
bly jointed to last abdominal segment. No median process from the vertex of head of S , but
there may be a small median, ventral process. Basal joint of 2nd antenna of S (each half
of clypeus) with a conical, subconical or digitiform process on inner anterior side, and a
small setiferous lamelliform process near the distal end. Distal joint strongly curved inward,
often contorted, unarmed. A number of species are found in Africa, one in Asia.

Branchipodopsis affinis G. O. Sars

1901 Branchipodopsis affinis Sars. Ann. Mus. Zool. Acad. Imp. St. Petersbourg. 6: 149

Locality: Kashmir: *Nagmargh. 20-30 5 $9 9. Col. F. Smith. \T-13.

Reported from: Mongolia, Mont Chingan (sec. Sars); Manchuria, near Tyn Chur. (sec.
Daday) ; Russian Mongolia, near Lake Baical (sec. Smirnov).

Types: Museum of Natural History, Leningrad.

These specimens were received in an e.Ktraordinarily damaged condition, apparently
having been completely dried at some time in the past. They were very brittle as received
in alcohol, and not a single specimen had escaped breakage. It was at first impossible even
to make sure of the genus, Init by treating with 5% KOH solution, the horn-like interior
was softened, and the integument resumed something of its former shape. They were then
lightly stained with tetrabromfluorescic acid, and preserved in glycerine to protect the now
very soft specimens. Since these specimens are in such poor condition, the description and
figures are taken largely from Sars (IS^Ol ).

Body somewhat more slender than usual in the genus. The paired median processes on
the dorsal surface of the S clypeus terminate in 2 rounded lobes, having between them a
small spine; digitiform processes on dorsal, distal parts of the clypeus well marked. A small
ventro-median, spinuliferous process on the clypeus. Distal joints of 2nd antenna of $
strongly curved, and somewhat expanded near tips. Second antennae of 2 terminate in an
acute pointed process. The 6th legs of c5 have 5 rounded, tuberculifnrm processes between

* Indian Museum specimens.

42

niVLLOPOD CRUSTACEA

the spinas of the distal endite. (The Kaslimir specimens appear to have only 2 such tuber-
cles.) According- to Daday (in it mentioned by Sars) there are 2 sjiines on the under side of
the last postpedigerous segment. Tiiese cannot be made out nn the Kashmir specimens, though
(piite possibly l^ecause of their pdor preservation. Cercopods of 9 straight, ]i(iinted and
setiferous; those of S longer, strongly curved inward, and setiferous on the greater part of
the outer margin. On the inner margin they are provided with spines which continue to,
and a little around, the tips.

Figure 6. — flraiicliil'odopsis affinis. A, head of $ from in front (Xl4). B, 2nd antenna of 9 (X39).
C, outer endites of 6th leg of 5 (X 32). D, end of abdomen of c5 . E, egg sac of 9 from right side (X 10.5).
(All redrawn from Sars (1901).)

Family Strkptocepiiai.idak Daday
V)\0 Strrplocrphalidac Daday. Ann. .Sci. Nat. (sen 9.) 11:.^.^.=^

Eleven pedigerous, 9 postpedigerous segments, the last always shortest. Head in i sim-
ply rounded in front, or with a frontal process. Second antenna of $ triarticulate, with distal
joint cheliform, a curved, chitenous process projects more or less ventrally from the juncture
of the ba.sal and middle joints. Legs with 1 epite. Cercopods movably articulated with the
last abdominal segment (except in S. scaliif). Ovisac cylindrical, usually elongate. Only
1 genus, which is found in all continents except South America.

PUYLI.OPnn CRTTSTACEA 43

Genus Sircptoccphaliis Baird
1852 Sfn'pfoci'plialus Uaird. I'roc. Zool. Soc. Lniidon 20:20

With the characters of the family.

The species of the genus Strcptoccphaliis liitherto descriljed fmni the Indian region liave
i)een named 5". diclwtomusBa.hd, and S. dichotouiiis var. siinphw (lurney. But an examina-
tion of the specimens of the Yale North India Expedition and the numerous specimens
sent me from the Indian Museum has shown that such a classification is untenable. If it
were to he allowed, a numljer of subvarieties of var. simplex would have to be erected,
some of whicli wmild lie geographically distinct ; and even though the ranges overlap slightly,
there already appears to be a geographical distinction between .S". dichotoiiiiis, and 5'. d. var.
simplex, so that a sub-specific distinction would be proper at the very least. If var. simplex,
the more primitive form, had been described first, it would perhaps be possible to express the
relationships without too much confusion, but under the present conditions it seems much
l)etter to raise (iurney's variety to the rank of a full species, with 3 sub-species. This (to
some perliaps drastic ) step has sound precedent in the suborder, and even witliin the genus,
since S. dregei G. O. Sars, and .S". cirratus Daday are equally close to each other. I may
add that iiu intermediate ftirms have ever l>een recorded between diehotomus and simplex.

Slrcptocepliali(s simplex simplex nov. comb.

1907 Streploeephaliis dichoUimiis var. simplex Gurney. J. and I'roc. Asiatic Soc. Bengal
(New Series) 2:276

Localities: Patiala States *Base of Simla Hills 15, collector for the Indian Museum.
Date?

United Provinces: *Mirihan, Mirzapur, R. B. S. Sewell, coll. 30-XII-12.

Reported from: Cutcli (Gurney); Calcutta (Daday).

Types: Indian Museum, C"alcutta.

Tlie distal chelate joint of the 2nd antenna of the 5 is often spoken of as the "hand,"
the dorsal branch being the "thumb" and the ventral l)ranch the "finger." For tlie sake of
simplicity this terminology will be adopted here. The left hand of the t? from Patiala State
is shfnvn in Figure 7, with the parts to be mentiijned in the descriptions labeled. The terms
"dorsal," "ventral," etc., when a])plied to the 2nd antennae of the c? shall be a])plied as
if these appendages were extendetl out directly forward of the head.

Body ratiier robust for the genus. Abdomen without furca scarcely longer than the
trunk. Head of 9 e\'enly rounded, with the 2nd antennae much folded and crumpled, often
largely obstructing forward vision. Head of <5 produced in front into a short conical pro-
tuberance which is plainly visible from above, lying between the bases of the 2nd antennae
(as in Figure 8, d. d'). Pedigerous and postpedigerous segments simple and unarmed.
Male genital sac of the f<irm usual in the genus, with tiie usual cylindrical, spinous penes.
Ovisac of 9 a slender tapering cylinder, not reaching as far as the last abdominal segment,
tip not bent. Cercopods in both sexes narrow and lanceolate, fringed all round with subequal

44

rilVI.I.Ul'OD CRUSTACEA

plumos^ setae. Cercopods very slightly longer proportionately in S than in 9 . First
antennae of l)oth sexes not showing segmentation or psendoseginentation. Second antennae
of S triarticulate. Basal joints cylindrical, superficially somewhat creased, usually bent more
or less downward. At the juncture of the basal and middle joints is a ventro-laterally directed,
slightly curved smooth process wliich is heavily chitinized. Middle joint of 2nd antennae of

S B

■THUMB

Figure 7. — Siref'tocephahis simplex simplex. Left hand of S from Patiala State (X 13). N = finger notch ;
SB = sickle-shaped branch of finger; BS = basal spine; MB = main I)rancli of finger; G = dorsal groove of
tluiinb ; TN = thnmb notch; VP =; ventral process of thumb; DP = dorsal process of tliunib.

S witii a sigmoid flexure; close to its basal end on tlie dorsal surface are 3 slender fleshy
processes, the innermost one always larger, the other two may nearly equal it, or may be con-,
siderably smaller. All 3 are similar in shape, tapering, curved downwards, pointed, and with
the lower surface provided with a row of small papillae. On the dorsal surface of this same
middle joint is a row of a1)out 10 slender processes, the middle ones usually being shorter.
The whole of the middle segment of the 2nd antennae of the male gives the appearance of
being superficially annulated. The hand, as seen from the outer side, is well shown in
Figure 7. It will be observed that: the distance from the thumb notch to the tip of the

PHYLLOPOD CRUSTACEA

45

tliunil) is aljijut half the length of the main branch of the finger as measured from the basal
spine to its tip. The dorsal process of the thumb is prominent. The dorsal row of spines
tends to run over onto the inner side of the main branch of the finger distally, and the sickle-
shaped branch of the finger is practically smooth along its concave edge. The legs of both
sexes have the epite serrate along the margin, and the gill of the last pair flattened, enlarged,
and finely serrate along the end. Length usually about 20 mm. or more.

FiGURF. 8. — Strclttoccphahis. A, 5". simplex echinus, head from right side. A', same, from above. B, .5". s.
luiigimainis. head from right side. B', same from above. C, 5. j. (iraf'icH.y, head from right side. D,S.diclwtomus,
head frcjin right side. D' same, from above. (All X 8.1.) Side views of heads show right 2nd antenna cut away
to expose the frontal process; the cut surfaces are lined. Many of the differences other than the frontal processes
are the result of the condition of the material. The frontal process of S. s. simplex appears to be exactly like that of
.S'. dicluiliiinus.

Streptocephalus simplex longimanus n. subsp.
Locality: Madras Presidency: Mahabalipuram. 2S, 39, coll. Hutchinson. 4-XI-32.
Types: Peabody Museum of Yale University. Paratypes to Professor Hutchinson.

This subspecies differs sufficiently from S. s. simplex to make it seem worth while
to make the distinction. The terminal joint of the 2nd antennae of the 3 has a much
shallower and less inarked notch on the ventral side of the basal part of the finger. The sickle-
shaped branch of the finger has no basal spine. The main branch of the finger has the spines
on the dorsal edge very few (7-10), short and blunt. The thumb has no dorsal process,
though the dorsal groove is present, and the length of the thumb measured from the notch
on the distal side of the ventral process is actually greater than the length of the main branch
of the finger measured from the dichotomy. The 1st antennae of both sexes are long and

46

I'll VI. I.U POD CKUSTACEA

exhibit a jointed appearance. Tlie cercopods nf both se.xes are unusually wide at the base,
and are much flattened dorso-ventrall}-. In other resix;cts the two sexes appear to be essen-
tiall\- like the t\nical form.

Figure 9. — Sircptoccphahis simplex hiigimaiitis. A, left hand of S from outside (X 13). B, head of 9 from in
front (X 11). C, end uf abdomen of $ from above (X 17.4).

Strcl^tocc/'lialus siiiif^lrx arabicus n. subsj).

Locality: South Ar.mua : Aden. 5 o' , 5 9, coll. G. \i. Hutchinson. 21-11-32.
Aden, 100 or more. Coll. C,. K. Hutchinson. 7-XII-32.

*Aden?, 2 <5 S, 2 9 9, coll.? Date?, poor condition and of doubtful provenance, (|uite
possibly S. s. arabicus.

Types: : Peabody Museum of Vale University. Paratypes in Indian Museum.

Much like the t_\i)ical variety, but for the followint^: First aiUennae of l)(itli sexes
often appearing to be divided into 2 or more segments. Second antenna of 6 with
basal joints so much fused dorsally as almost, or completely to hide the short and ill-
developed frontal process of the head when viewed from above; with the outer 2 fleshy
processes of the 2nd joint nuuh reduced in size (one may be missing), and with the dorsal
row of sj)ines on the main branch of the finger not tending to run over onto the inner face
of the main branch distally. The finger-notch is well marked, and the proximal edge of it is

P11Y],L01'()D CRUSTACEA

47

more or less produced. The 2nd antennae of the 9 not very rarely bent or folded, and nar-
rower than in S. s. simplex; sometimes produced into an obtuse point at the inner side of the
end. The g'ill of the last legs of both sexes is perhaps a little narrower than in the typical
form. The specimens collected by I'rofessor Hutchinson on the different dates differ greatly

Figure 10. — Strcplocephalns simplex arabicns. A, head of Feb. $ from above (X 12). B, head of Dec. 2
from in front (X 7.4). C, left 2nd antenna of Feb. 9 (Xl2). D, end of abdomen of Feb. $ from above.
E, abdomen and egg sac of Feb. 9 from right side (X 12).

in size, those of I-'ebruary measuring al)out 23 mm. f(ir tiie i $ and 20 mm. fur the 2 9.
The specimens taken in December, however, are the smallest se.xually mature specimens I have
ever seen reported for the genus, measuring only about 9 mm. for the largest $ S , and as
little as 6 mm. for 9 9 carrying eggs. In other respects they are precisely like the larger
specimens.

48

PHVLLOPOn CRUSTACEA

Strcptoccplialiis sniiplc.v echinus n. subsp.

Locality : Maduas Prksidkxcy : *(;odaveri ( Town) . 3 <J , 9 9 , coll. N. Aniiaiulale. 28-\M 1 1-18.

Types: Rcturnccl in Indian Mnseum. One £ and one 5, paratypes, retained.

Body and cercopods of Imth sexes relatively slender. First antennae of both sexes
are relatively longer, and appear irregularly segmented. The 2nd antennae of the 5
entirely without fleshy processes of the middle joint; the slender processes on the dorsal side
of the distal part of this joint are reduced in number to 6-8; in the distal joint, the dorsal

KicjIKk U.Slrcf'linrtliiihis siiiiplc.v ahiiius. A, left hand of <^ fmiii (Hitcr .side (X 13). I?, head of 9

from ill front (X H).

row of spines of the main branch of the finger is more regular than in S. s. simplex and does
not run over onto the inner side of the branch, and the spines are much more numerous.
There is also a row of short, conical spines along the outer side of the main branch i>f the
finger. The sickle-shaped branch of the finger is armed along the proximal 4/5 of its con-
cave edge with short close-set spines which become somewhat papilliform distally. The
thumb is a little longer than in 5". j. simplex, its length from thumb-notch to tip being aliout
4/5 of the length of the main branch of the finger as measured from tlie dorsal spine. The
dorsal process of the thumb is lacking, and the dorsal groove of the thumb is scarcely indi-
cated. The finger-notch is obsolescent. The 2nd antennae of the 9 2 are rather narrower
than in the typical form. Length oi $ £ alx)ut 20 mm., of 9 9 about 18. In other respects
this subspecies is very similar to .S". .?. simplex.

PHYLLOPOD CRUSTACEA

49

Strcptoccplialus dichotoinus Baird

1860 Strcploccphalus dicholoinus Baird. Proc. Zool. Soc. London. 28:445

1900 Streptoccphaliis diclwtoiims Sars. Arch. Mat. Naturvid. 22(9) :4

Localities: Madras Presidency: *Madra.s, Spur Tank. 10 c? , 9 5 , N. Annandale. III-ll.
*Tanjore (S. India). 1$ , 1 5 , N. Annandale. 27-X-ll.

United Provinces: *Baraunda Tank, Mirzapur. 1 1 c5 , 5? (juvenile), Mrs.
N. M. Johnstone. 15-VIII-13.

Mysore: ^Bangalore (India). 1 c5 , 1 9 , N. Annandale. 13-X-lO.
Reported from: India (Baird); Calcutta (Alcock ) ; Shevaroy (Stevaroy) Hills (Sars).
Type : Ubi ?

FiciiiRF. l2.Strcl'tuccl^halux diihuloiiiiis. A, left liand of Madras spur tank <? from outer side (X ll.d)-
B, head of Tanjore $ from in front. C, head of Madras 9 from in front. (B and C X9.9).

(There are also about a dozen 9 9 from Mirzapur, collected by Mrs. N. M. Johnstone,
which may be of this species or of S. simplex, as the females are indistinguishable. This is
by no means rare among the Anostraca — for example, no distinguishing marks have been
reported which allow of the separation of the 9 9 of any of the numerous species of
Branch ipodopsis. )

Sars' redescription of this .species is so very complete that it will be unnecessary to dis-
cuss it very fully. A few points of interest have been observed, however, because of the
larger collections from a more wide area that have been available to me.

The species in general is very like .9. simples, differing mainly in the structure of the
2nd antennae of the S , the middle joint of which usually Ijears proximally 4 fleshy processes,
though the S from Bangalore and one of those from the Madras Spur tank have only 3 as in

50

PHYLI.OPOD CRUSTACEA

6". simplex. The finger-notch is generally more deep and open than in the forms described
alxive, and the main branch of the fing'er is always Ijifnrcated for al)Out its last third. The
bifurcation is so constructed that from the outer side the ventral branch appears to be an
enlarged spine, while from the inner face, the dorsal branch gives that appearance. Both
branches may be smooth, or either or both may bear minute spinules. Baird had only the S
(if this species, and Sars says "y\ntenna€ in female simple, blade-like, bluntly rounded at the
tip: . . ." All 16? 9 examined by me had the 2nd antennae folded and wrinkled to a
greater or less degree, even the very immature specimens from Alirzapur showing it plainly

Figure 13. — Strcplocephalns. A, left 2nd antenna of young $ S. simplex from Nundy, seen from outer side.
B, same, somewhat older specimen. C, left 2nd antenna of young S S. dichoiomus from Mirzapur, of about the
same age as 13, D, head of a very young 9 -f. simplex from Nundy, seen from in front. The Nundy specimens
are too young to determine the subspecies.

This agrees with the statement of Alcock (1897) who described the species under the name of
BranchipHs (Strcptoceplialus) bcngalensis, though his figure is almost the precise antithesis
of his description. The 1st antennae of the 9 9 , and to a lesser extent of the S S , are fre-
quently coiled, and often hidden under the 2nd antennae. In neither sex do the 1st antennae
appear segmented.

The bifurcation of the main branch of the 2nd antennae of the S appears very early.
A young stage is shown in Figure 13, c. (The 5". simplex from Nundy, Figure 13, a, b, d, are
not old enougli to place certainly as to subspecies, except to say that they are not echinus.
They are in all probability S. s. simplex.)

All the forms of Streptocephahis here in discussion are quite closely related to each
other, but not to any other forms. If we regard the fiat, unfolded 2nd antenna of the 9
as primitive (it appears nearly universally throughout the genus), it is clear that the oldest
member of the group is that closest to the African center of distribution of the genus. No
Streptocephali are known to occur along the present land route between Arabia and India.
This may be because of inadequate collecting, or the distribution may have taken place before
the present arrangement of the land masses.

PHYLLOPOD CRUSTACEA 51

Key to the Species and Subspecies of the Genus StrcptocepluiJus of the Indian Empire

1. 5 with a complex frontal process, and strong spines on some of the

abdominal segments 6". spinifer

2. S with abdominal segments imarmed

A. Main branch of finger of 2nd antenna of $ bifurcate distally 5". dichotomus

B. Main branch of finger of 2nd antenna of <$ not branched

i. Thumb l^yond thumb-notch at least as long as main branch of $

2nd antenna S". siniphw longiinanus

ii. Thumb not over 4/5 as long as main branch of finger

a. Basal joints of 2nd antennae of $ more or less fused dor-

sally, obscuring frontal process 5". simplex arabicus

b. Basal joints of 2nd antennae of $ not fused dorsally, frontal

process visible from above
b,. Concave edge of sickle-shaped branch of finger sinooth or

nearly so S. simplex simplex

I);. Concave edge of sickle-shaped Ijranch spiniferous 5". simplex echinus

Ecology and Zoogeography of the Indian Anostraca

The Anostraca as a whole are slow swimmers with no sort of protective devices or
behavior. Introduced into an aquarium, they became immediate prey to any sort of fish,
and when they are in company with copepods or even cladocera, the Anostraca are always
the first to be eaten, usually being exterminated l^fore any appreciable inroads have been
made on the (ither forms. I once discovered the skeleton of a single, small fish in the dried
bed of a temporary pond of considerable size which was known to have contained Branchinecta
occidentalis the previous wet season. Outside of this one, rather circumstantial observation,
1 have never seen reported a single case of Anostraca and fish l^eing found in the same
waters.

Possibly in connection with this vulnerability, various devices have arisen which have the
effect of preventing the co-occurrence of these phyllopods and fish. Arteniia will only live
in waters too saline for most fish to inhabit. The other forms may, in the main, be divided
into 2 classes : Those with eggs that require drying to hatch, and those with eggs requiring
freezing. In many temperate -regions the eggs may undergo both processes without detri-
ment. It is not known whether any species require both, or whether in any form drying
can substitute for freezing or vice versa.

As a result it may be said that the Anostraca are ordinarily found in small, shallow
bodies of water, usually of a temporary nature, and that there is usually only 1 generation a
year.

Of the Indian anostracans it may be surmised that both Branchinecta orientalis (the
specimens taken from Togarma Tso, 5,217 m., are from the greatest altitude I have been able

''Found only in Ceylon and not discussed in this paper. Cf. Gurney. Spolia Ceylanica 4 (14-15) : 127, 1906.

pnvLi.nron ciu'stacea

FrcuRE 14. — Map of Sirrfloccf'luihis localities.

l-'i(.LKK 15. — Map of the (listributiuii of tlic Indian .\nostraca except Sln-pluicfilmliis.

PUYTXOPOD CRUSTACEA 53

to tiiul reported fur au}- i'hyllopod) and Branchipodopsis affiiiis have egg-s requiring freez-
ing (though this may not l>e so), and certainly that the eggs of the Bratwhipus and of the
Strcptoccpliali require dr\'ing. Tlie finding of Pristiccplialux in the Sargodhar District seems
to show that these eggs recjuire drying. It is perfectly possible, of course, that this (and
other forms) may have the eggs made ready for hatching by either method indiscriminately.

Since the eggs will (in most species, perhaps all) withstand long periods of drought,
and since ihey are small and light, it might be expected that they would easily and often be
transported by the feet of water birds, by the wind, or by other agencies. As a matter of
fact, however, the ranges of many species are surprisingly circumscribed, even though ponds
and pools offering apparently ideal conditions are to be found a short distance away. Other,
perhaps related, species may be very widely distributed, with, so far as has been determined,
no more efficient method of dispersal.

This curious sort of distribution may perhaps test be explained by the presence or
absence of various necessary, or destructive, factors in the various environments, the different
species differing, of course, in their requirements or sensitivity. Unhappily, very little is
known of the particular ecological factors involved, with the exception of Artemia from cer-
tain regions, and since the environment of brine-pools and salterns is so special, little light
is thrown upon the situation in other genera. Though there is little experimental evidence,
it appears probable that temperature is a very important factor, not only in the freezing of
eggs, but there is reason to suppose that there are both maximum and minimum, sharply
limiting temperatures for many, if not all, species. Thus Heath (1924) has shown that
hatching and the early stages of development in Branchinccta occidcntalis may take place at
lower temperatures, but that sexual maturity is only attained after the water has risen to a
temperature of approximately 22°C., despite abundant food and other suitable conditions.

The Himalayan Mountain system makes a more or less sharp temperature barrier between
North and South, and the higher part of the plateau, even south of the crest is, undoubtedly,
for such species as Branchipodopsis affinis essentially similar to Manchuria and Mongolia.
Whether this form is to be found in many places between the present known sites, or
whether it reached the Himalayan plateau at a period colder than the present, it is impossible
to say. The very occurrence of this species in the cold parts of Asia is at present not easy to
explain, since all the other 11 known species of the genus are found only in the warm and dry
parts of South Africa. The arrival of Branchinccta oricntalis was probably from the North
and West, where it now occurs, and in which direction other members of the genus are to be
found. Pristicephahis prisctis may be surmised to have come from the North and West also,
since its closest relatives are to be found in that direction. Because of its differentiation and
restricted range it wnuld appear to have arrived at an earlier date than the other northern
forms, however. Branchipus stagnalis, like Strcptoccphalus, i)robably arrived from the West,
though it is odd that it has been met with only once, and it appears barely possil^le that it
may have been a chance importation through the agency of modern man.

The localities in which the various forms of Strcptoccphahts are found are shown on the
map, Figure 14. The other species are shown in Figure 15.

54

niYLLOPOD CRrSTACEA

Figure 16. — ■Af'us cancriforniis 9 from Nuriwala.

Suborder 2 NOTOSTRACA
1867 Notostraca Sars. Crust, d'eau douce Norv. : 5

Genus Apiis Schaeffer

1756 Apiis Schaeffer. Der krehsartige Kiefenfu.ss, etc. : 131

1803 Triops Schrank. Fauna Boica 3:251

The correct name for this genus is by no means settled, and though Triops is in general
use at tlie present in continental Europe, I believe that only Apiis has been used by the vari-
ous authors who have treated of the genus in the Indian region. The arguments for this
usage are so well presented by Stebbing (1910) and Gurney (1923,1924), particularly the
former, that I feel constrained to follow their usage in this paper.

PHYLLOPOD CRUSTACEA 55

Apits caiicnfomns Scliaeffer
1756 Apiis cancrifonnis Schaeffer. Der krebsartige Kiefenfuss, etc.
Locality: Punjab: Sargodhar District, 3 mi. South of Nuriwala. 29 9. 6-III-32.
Reported from : Europe, Northern Africa. Kashmir (sec. Barnard).

Carapace oval, shghtly longer than broad. Nuchal organ (between the posterior margins
of the compound eyes) oval as seen from above, conical as seen from the side. Number of
postpedigerous segments in S 6-8, in 9 5-8. Fourth endite of 1st leg longer than carapace.
Rami of caudal furca as long as all the rest of the animal, or longer.

The 2 ? 9 of the collection have 8 postpedigerous segments each — an unusually large
number. The other characters agree so well, however, that there is no doubt of the correct-
ness of the identification. (See Figure 16, drawn by Miss L. Krause.) In nothern Europe
this species is generally parthenogenetic. Whether this is true in this Indian locality the col-
lection is too small to show.

On various bases this species has been divided into a number of subspecies, but as Barnard
(1929) has so clearly shown in his study of the South African forms, the diagnostic char-
acters chosen have no taxonomic value and simply result from individual variation (Cf. also
Gurney (1923)). India may or may not harbor a valid subspecies, but this can only be deter-
mined by a study of some hundreds of specimens.

Suborder 3 CONCHOSTRACA
1 867 Conchostraca Sars. Crust, d'eau douce Norv. : 5

Family Limnadiidae Sars
1896 Limnadiidae (part.) Sars, Fauna Norv. 1 : 84

Shell thin, pellucid, ovate with few and inconspicuous growth lines. Head of adult with
frontal appendage. 18-32 pairs of legs, 1st and 2nd pairs in S prehensile. 9th, 10th and
sometimes 11th pairs of legs in 9 ovigerous. Caudal furca claw-like.

Three not very well defined genera.

Genus Etdimnadia Packard-Daday

1874 EuHiiiiwdia Packard. Rep. Peab. Acad. Sci. Salem. 6:55

1925 EuUmnadia Daday. Ann. Sci. Nat. (ser. 10) 8:145; (9:1-3 (1926))

Hinge line of shell not serrate. 18 or 20 pairs of legs. Lower distal angle of telson
(last abdominal segment) produced into an acute point.

56

niVI.I.OrOD CRUSTACEA

^ Eulimnadia margaretae n. sj).

Locality: South Auabia: Aden, 95 9. 7-X1I-32.

Types: PealKxlv Museum of Yale University. Paratypcs : Indian Museum, liritish Museum,
and retained.

Description of 9 ( <} unknown). Shell tiansparent, oval, highest point just anterior to
tlie middle. Hinge line evenly arcuate. Growth lines 3 in number, the outermost extremely
indistinct. Knstrumiu 9 rounded, or bluntly acute (especially in yi lunger specimens) ; frontal

FicuRK 17.— Eulimnadia margaretae. A, .slid! n{ 9 fnim left side (\ 5.8). B, head of 5 from left side
(X20.5). C, telson of 9 from right side (X20.5). D, distal part of left 1st leg of 9 from I)chiiul. Rases only
of the setae are shown (X22). E, egg (X43).

organ subglobular; posterio-ventral margin of liead nearly straight, sinuous, or wilii a mure
or less sharp notch, hirst antennae with 6 and terminal lobes; 2nd antennae with branches
of 7 and 9 segments. Twenty ])airs of legs, of which jiairs 10 and 11 are ovigerous. I'os-
terior 9-13 segments bearing dorsal setae of variable number and dift'icult to count (.setal
luimliers from k-hind forward on Type: 5?, 5?, "?, 7, 7?, 7?, 7?, 5?, 3, 1, 1. 1, ] ). Telson
much worn in all specimens, with about 35 small, irregular dorsal sjiines, all of which are
smooth; lower distal angle of telson produced into a short, rounded (possibly worn) point.
Furcal claws of all sjiecimens broken, with rounded ends; the bases of about 20 plumose setae
can be made out on each claw, but most of them are bniken off. Dimensions of shell:
10.4x7.4 mm. Ova s])herical, rugose. This form does not very closely resemble any of the
species described hitherto.

PTIYLLOPOD CRUSTACEA 57

Family Cyzicidae Stclihing-Baniard

1910 Cy::icidac (part.) Stebbing. Ann. S. Afr. Mus. 6:486

1929 Cyacidae Barnard. Ann. S. Afr. Mus. 2-J (1) : 253

"Shell thin, pellucid (but often rendered opaque with extraneous matter), laterally com-
pressed, ovate in outline, with numerous and distinct growth-lines and more or less dis-
tinct surface sculpturing. Head without frontal appendage, with distinct fornix on each
side extending to apex of rostrum. Rostrum unarmed, or with a minute apical spinule in the
young wiiich may persist in adult 2 , but not in adult S . Eyes contiguous. First antennae
long with numerous lobes on anterior margin bearing sensory setae. Second antenna strong.
Twenty to twenty-seven pairs of legs; 1st and 2nd pairs in S prehensile, 9th and 10th pairs
in 5 ovigerous. Caudal furca claw-like. Foremost tooth on upper margin of telson larger
and stronger than the following ones." (Barnard.)

Genus Eocycicus Daday

1913 nocy::icus Daday. Math. Termt. Ert. 31 : 567, 574 (sec. Barnard)

1915 Eocyciais Daday. Ann. Sci. Nat. (ser. 9) 29: 190

With occipital angle rounded or rounded-quadrate in bnth sexes. Rostrum of 9 acute,
of S with a less sharp angle (often obtuse).

Eocyzicus hutchinsoni n. sp.

Localities: Punjab: Tahsil of Kushab, Dam between Naushara and Mardwal It? 1$.
12-111-32.

I'uNjAis: Tahsil of Kushab, 3 miles smith on Nuriwala, Kushab-Naushara iviad
29 5. 6-1 11-32.

Types: Peabody Museum of Yale University. Paratypes : British Museum, 1 retained.

Male: (Type only.) Shell ovate, umbone moderately prominent, dorsal margin straight,
passing almost imperceptibly into hind margin; about 15 growth lines, outermost and inner-
most very faint; free margin and outer 2-3 g-rowth lines with minute spines. Pits of shell
sculpture moderately large but so shallow as to be obscure. Rostrum of S acute (extreme
end minutely truncated) ; posterior angle of rostrum rounded; anterior and posterior mar-
gins of rostrum nearly parallel. Supraorbital margin of head sinuous. F^irst antennae with
about 14 lobes; 2nd antennae with both rami of 12-14 joints; spines on anterior margins i>{
joints smooth, or a few slightly ctenate. Twenty-two pairs of legs; inner margin of the
'"hand"' of prehensile legs with strong notch in 1st pair, slightly sinuous in 2n(l i)air; "tlunnb"'
broad; spinous patch long and narrow in both pairs. Last 14 pedigerous segments armed
dorsally (si)ine-formula from behind f (jrward : 1, 1, 1, 1, 3?, 3, 3, 3, 3, 3, 3, 2, 1, 1). Telson
with claws markedly asymmetrical, each preceded by 14 smootii, very unequal denticles, of
A-hich 1 near the middle of the row is about as large as the first. Furcal claw with 7 plumose
setae on the dorsal, inner margin. Dimensions of shell: 10.6 x 6.9 mm.

58

PHVU.OPOD CRI'STACEA

I'Yniiale : Shell as in S , hut with umbones nnidi less pnimineiit; about 12 growth lines,
the outermost very faint. i\(>struni sharply acute. First antennae with about 18 lobes; 2nd
antennae with Ijoth rami with 11-13 joints; spines on joints as in the S . Twenty-two pairs
of legs; 1st pair witli 6tli enclite extending as far as distal end of flabellum or beyond; palp

Figure 18. — Eocyzicus hutchinsoni. A, B, shells of $ and 9 from left side (X S.8). C, D, heads of $ and
9 from left side (X 13.5). E, spines on anterior side of 1st joint of anterior ramus of 2nd antenna of 9 (X 135).
F, left 1st hand of S from behind (X 18). G, distal part of right 1st leg of 9 from behind. The position of some
of the setae is indicated by their bases (X 21). H, left 2nd hand of S from behind. (X 18). J, tclson of S from
left side (X22). K, right ftirral claw of i from left side (X44).

of 5th endite extending nearly to end of 6th endite; 5th endite much less than half as long
as 6th; 4th endite without palp; notches between endites shallow; 9th and 10th pairs of
legs ovigerous. Last 15 or 16 pedigerous segments armed dorsally (forinula of type, from
behind forward: 3, 3, 3, 5, 5, 5, 5, 6?, 5, 5, 4?, 3, 3?, 1, 1, ( 1 ) ). Telson with daws less
asymmetrical than in $ , preceded by alxjut 26 smooth, unequal denticles, with 1 very prom-
inent near the middle of the row. Furcal claw as in 5 . Dimensions of shell: 8.8 x 5.5 mm.
Eggs rugose.

PIIYLLOPOD CRUSTACEA

59

Eocyzicus deterrana n. sp.
Locality: Punjab: Rawalpindi District, Soliawa. About 45 <J cJ 5 2 . 3-III-32.
Types: Peabody Museum, Yale University. Paratypes; Indian Museum; retained.

Figure 19.— Eocyzicus deterrana. A, B, shells of i and 9 from left side (X 7). C, D, heads of 1 and 9
from left side (X 13.5). E, spines on anterior side of 1st joints of anterior ramus of 2nd antenna of 9 (X 135).
F, G, left 1st and 2nd feet of S from behind (X 18). H, left 3d leg of 3 (X 20). J, telson of $ from left side
(X 22). K, enlargement of part of J (X 135). L, right furral claw of c? frmn left side ( :■; 4-4).

Male: Shell ovate, umbone low; dorsal margin straight, often making a definite angle
with hind margin; about 14 growth lines, the outer ones obscure and crowded; free margin
and outer 3-4 grow-th lines with minute spines. Pits of shell sculpture small, very shallow,
difficult to observe. Rostrum of S acute, nearly a right angle; posterior angle of rostrum
very obtuse. Supraorbital margin of head straight. First antennae with about 16 lobes;
2nd antennae with both rami of 11-12 joints; spines on anterior margin of joints smooth or
ctenate. Twenty-two pairs of legs (5 specimens) ; inner margin of "hand" of prehensile legs

60 I'lIVI.I.OI'OD CKl'-STACEA

with slight notch in 1st ])air: nearly straight in 2nd; "tluiinl)" sqnare in 1st pair, slightly
broader in 2nd; in both pairs base of spinous patch of thunil) only slightly longer than the
spines. Last 15 (about) segments armed dorsally (spine- formula of type, from behind for-
ward: 1, 1, 3, 3, 3, 4, 4, 5, 5"^ 5, 5, 4, 2?, 2, 1).

Telson with claws moderately asymmetrical, each precedctl by about 12 very unequal den-
ticles, of which 2 or 3 near the middle of the row are nearly as large as the 1st; 1st (anteri-
ormost) denticle, and some of those following, armed with very fine spinules. Furca'l claw
with 4 plumose setae on the dorsal, inner margin. Dimensions of shell ; 6.6 x 4.0 mm.

Female: Shell as in ^ but with umbones even less prominent; about 11 growth lines,
the outer ones indistinct and crowded and the inner ones very indistinct. Rostrum acute;
supraorbital margin of head sinuous. First antennae with about 14 lubes; 2nd antennae with
both rami with 11-12 joints; spines on anterior margin of these joints strongly ctenate.
Twenty-two pairs of legs (4 specimens) ; 1st pair with 6th endite extending as far as distal
end of fiabellum or beyond; palp of 5th endite extending nearly to end of 6th endite; 5th
enilite much less than half as long as 6th; 4th endite without i^alp; notches l>etween endites
shallow; 9th and 10th legs ovigerous. Last 14 segments of type armed dorsally (formula,
from behind forward; 1, 1, 3, 3, 3, 4, 4, 5, 5, 4?, 3, 3, 1, 1). Telson with claws scarcely
at all asymmetrical, preceded by about 17 imequal, mostly armed denticles (as in the S ) oi
which 2 or more in the middle of the row are about the size of the first (anteriormost).
Furcal claw as in the S . Dimensions of shell : 6.0 x 3.8 mm. Eggs rugose.

No such careful piece of work has ever Ijeen done on the genus Eocycicus as Barnard
(1929) has done for Apus, so that the extent of variation within a natural species is not
known; nor is it known which of the structural details of these Conchostraca are reliable
specific criteria. When such an investigation is made, it may possibly be found that one or
both of the above-described species must be reduced to synonymy with others already known,
l)ut ill the present state of our knowledge of the group it is probably better to describe as new
any specimens about which there is reasonable doubt.

The nearest described relative to E. hntchinsoni is probably the wide-ranging E. oricntalis
Daday, itself very clo.se to E. boiiincri Daday, which differs most conspicuously from the
new species in the shape of the rostrum of the S , and in the presence of a well-marked
palp on endite 4 of the leg 1 of the ?. E. deterrana is most similar to E. perrieri Daday,
from Tobolsk and Buchara, U. S. S. R., but the latter has only 20 pairs of legs, and the
■'hands" of the 5 5 of the two species differ in shape.

• Oshorn Zoological Laboratory,
Yale University.

PIIYLLOPOD CRUSTACEA 61

BIBLIOGRAPHY

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62 PHYI.LOPOD CRUSTACEA

Gross, F. 1932. Untersuchungen iiber die Polyploidie und die Variabilitat bei Artcniia
salina. Naturwiss. 20 : 962-967.

GuRNEY, R. 1906. On two new Entomostraca from Ceylon. Spolia Ceylanica. 4 (14-15) :
126.

1907. On some Freshwater Entomostraca in the Collection of the Indian Museum,

Calcutta. Jour, and Proc. Asiatic Soc. Bengal (new series) 2: 273-281.

1923. Notes on some British and North African Specimens oi .Ipiis cancrifonnis

Schaeffer. Ann. Mag. Nat. Hist. (ser. 9) 11:496-502.

1924. Some notes on the genus Apus (Crustacea Branchiopoda) Ann. Mag. N. II.

London (9) 14: 559-568, 2 figs.

Heath, H. 1924. The external development of certain phyllopods. Jour. Morph. 38 (4) :
453-483.

tlERTwiG, G. 1931. Artemia salina, ein Beispie! fiir die Entstehung einer Gigas-Varietat
durch gieichzeitige Verdoppelung der Chromosomenzahl und des Chromosomen-
volumens. Gegenl)aur's Jahrb. 67: 371-380.

Leach. 1819. Dictionnaire des Sciences Naturelles. 14: (Entomostraces).

LiNNE, C. 1758. Systema Naturae. Editio X, 1758.

Packard, A. S., Jr. 1874. Description of new North American Phyllopoda. Reports
Peabody Acad. Sci., Salem, Mass. 6: 54.

Sars, G. O. 1867. Crust. d"eau douce Norv. (sec. Barnard, 1929).

1896. Fauna Norvegiae 1 (Phyllocarida og Phyllopoda), Cristiania, Aktie Bogtryk-

keriet.

1898. On some South ;\frican l'hy!lop;>ds. .\rch. Mat. og Naturvid. Krist. 20 (4).

1900. On .some Indian Phyllopoda. Arch. Mat. og Naturvid. 22 (9) : 3-30.

1901. On the Crustacean Fauna nf Central Asia. I't. I. Amphipoda and Phyllopoda.

Ann. Mus. zool. Acad. Imi). St. Petersbourg. (>: 130-164.

ScuAEFFEK, J. C. 1752. Der fischfomiige Kicfciifuss. Abhaudl. von Insecten. 2.

1756. Der KreKsartige Kiefenfuss, etc. Regensburg, E. .\. Weiss. (Schaeffer's

Abhandl. v. Insecten, 1 (3).)

1766. Eiementa Entomologica.

ScHR.\NK. 1803. Fauna Boica. 3 : (sec. Barnard, 1929).

Smiknov, S. 1932. Bemerkungen fiber Phyilopoden. Zool. .\nz. Leipzig. 100: 149-155.

Stebbing, T. R. R. 1910. General catalogue of South African Cn.istacea. Ann. S. Afr.
Mus. 6:281-599.

Stella, E. 1933. Phaenotypical characteristics and geographical (hstrilnition of several
biotypes of Artemia salina L. Zeitschr. f. induk. Abst. u. Vererbungsl. 65: 412-446.

Verrh.l, a. E. 1869. Descriptions of some new American Phvllopod Crustacea. .\ni.
Jour. Sci. (ser. 2) 48: 244-254.

ARTICLE VI

REPORT ON AMPHIPOD CRUSTACEA OF THE GENUS GAMMARUS

By Masuzo Ueno

Otsu Hydrobiological Station, Otsu, Japan
(Received February 21, 1934)

The gammarids here discussed were collected by Mr. G. Evelyn Hutchinson during the
Yale North India Expedition in Kashmir, Ladak and the Tibetan borders of North India.
The collection, though consisting of only one common species, Gammanis pulex (Linne), is
highly interesting, as it enables us to ascertain the distribution of gammarids at such unusu-
ally high altitudes as 5000 meters and over, altitudes which are believed to be greater than
those of any previously known localities for this species. I wish herewith to express my best
thanks to Mr. G. E. Hutchinson for kindly permitting me to undertake this work and also
for giving me useful information on many matters. I am greatly indebted to Prof. Kenzo
Kikuchi of Tokyo, for the use of his copy of Sars' monograph and to Dr. A. B. Martynov,
Leningrad, for supplying me with Chevreux's paper.

1. Localities at which Gammarids Were Collected

The collection consisted of twenty-three bottles of specimens in spirit containing over
150 individuals.' As indicated in Table 1, gammarids, though common both in Kashmir
and in the most elevated regions, are quite absent in the intermediate localities lying between
1600-3600 meters, a phennmencm that will be discussed in the last chapter of the present
paper.

2. Description of the Species

Family : Gammaridae

Genus: Gammarus Fabricius

Gauunanis piilcx (Linne, 1758)

In all characters of specific importance, all the specimens from Kashmir and Ladak agree
rather well with the descriptions and figures given by G. O. Sars (1895, pp. 503-505) and
Stebbing (1906, p. 474) for the typical form of Gaiuuianis pulex (Linne). There are, how-
ever, several important features to be noted peculiar to the North Indian specimens, especi-
ally those from Kashmir and the western part of Indian Tibet, north of the Ladak Range,
when compared with material discussed by several authors (Chevreux 1908, Martynov 1930,
Schiiferna 1922, Spandl 1923 and 1924, Tattersall 1914 and 1922) who have studied the
variation of certain characters of this species from different localities.

' Since the material was sent to Dr. Ueno a few more specimens have been t'omid in miscellaneous collec-
tions from several localities. These additional localities are recorded in a note appended to the present paper.
—G. E. H.

Mem. Conn. Ac.^D., Vol. X, Art. VI. September, 1934.

64

AiMrilll'OD CKUSTACKA OF THE GICNUS GAMMARUS

(1) The accessory flag'ellum of tlie first antenna. Tlic number of the joints (^f the
accessory flagelhim of the first antenna is given as four by Sars (op. cit. ), Stebbing (op. cit.),
and Clievreux and Fagc (1925), but the present specimens show considerable variation in the
numl)er of these joints; in most cases 2 or 3 and rarely only one. The specimens with
typical 4-jointed accessory flagella were chiefly collected from the localities in Kashmir at alti-
tudes between 1200-1600 m., wliile the s])ecini?ns collected fnmi the higher Ladak localities
have usually 2-jointed llagella, rarely 3- or 4-jniiiied unes.

TAI5LE 1
List of the Localities in which the Gammarids were Collected

Note: — K . . . the localities in Kashmir; L . . . the localities in Indian Tibet (Ladak) ; the figures
witiiin brackets show the number of specimens examined.

Altitude

No. of

station

Locality

In feet

In meters

Date

Remarks

K 1

Rampur

c.4000

c. 1200

18TII-32

.\n irrigation ditch (5)

K 2

Takht-i-Sulaiman,
Srinagar

c. 5200

c. 1585

20TII-32

Swamp at base of hill (8)

K15

East of Gagirbal
road

c. 5190

c. 1580

30Tn-32

A closed swampy pool ( 5 )

K19

East of Gagirbal
road

c. 5190

c. 1580

31-III-32

A closed swamiiy pool (4)

K23

Nishat Bagh

c. 5200

c. 1585

7-IV-32

A stream (11)

K24

Nishat Bagh

c. 5200

c. 1585

7-IV-32

A pond (16)

K43*

Wular Lake

5160

1573

17-IV-32

Littoral at Kiuhnus spring

(3)
Lagoon at Man by Pang-

Camp 9

Alan (C. G.)

14008

4269

6-VII-32

gong tso ; townetting (2)

L 40(a)

Panggong Tso

13915

4241

29-VH-32

N. W.end (12)

L 40(b)

Panggong Tso

13915

4241

7-VII-32

X. VV. end : littoral of 60
cm. (7 )
N. VV. end; a bottom of

L 40(c)

Panggong Tso

13915

4241

29-VH-32

31 m. collected by Eknian-

Birge dredge (5)

L49

Togom Tso

17506

5334

9-vn-32

(3)

L52

Ororotse Tso

17381

5297

12- VII -32

IVIargin of the lake (4)

L60

Kyam

c. 15500

c. 4725

21-VH-32

Poof (5)

L72

Chushol

14228

4336

9-Vni-32

Pool by pond (6)

L75

Tukmuru Tso

14385

4385

16-MII-32

Smaller part of the lake

(1)
A vertical haul (3 )

L76

Miti)al Tso

15998

4875

23-VHI-32

L 78t

'^'aye Tso

15373

4686

11 -VI 1 1-32

(1)

L81

Khyagar Tso

15330

4672

26-Vni-32

On anchor rojje; mostly
5-10 m. depth (25) -

L 82(a)

Tso-Moriri

14850

4528

30-\TII-32

At N. end, weed in estua-
rine water (17)

L 82(b)

Tso-Moriri

14850

4528

27-Vni-32

Vertical haul. 30-0 m. (1)

L85

Sta-rtsak-puk Tso

14885

4536

2-IX-32

(3)

* Tlie specimens labelled K 42.

t The localities numbered from L 78 to L 85 are located south of tlie Ladak Range.

AMI'HIPOD CRUSTACEA OF THE GENUS GAMMARUS 65

(2) The form of the lower hind corners of the second and third pletjn somites. With
regard to this character Sars (op. cit.) writes that "last pair of epimeral plates of metasome
but very little produced at the lateral comers." Stebbing (op. cit., p. 474) also describes
them as "simply ciuadrate." On the contrary, Chevreux ( 1908, p. 98 ) , who studied an Asiatic
race of this species, pointed out that the lower posterior angle of the third ])leon somite is
prolonged and sharply pointed. y\ similar description is also given by Chevreux and Fage
(1925, p. 254). Tattersall (1914, p. 213) also shows that the specimens of G. pulcx from
the Pamirs at an altitude of 15,600 feet have the third pleon somite with a considerably more
produced and pointed lower hind angle than figured in Sars' monograph.

The present specimens from North India, especially those from the localities nos. K1-L76,
have also the third pleon somites with considerably produced lower hind corners, such as is
seen in those described from high alpine regions in Asia by the above-mentioned authors. In
this feature the present specimens are much more allied to Gaiiunarus occllatus described by
Martynov (1930) from Lake Issyk-koul in Turkestan than to the typical form of G. pule.v.
This prolongation is more marked in the specimens from the lower Kashmir localities
(Plate IV, figs. 1-7) than those from the high Ladak localities (Plate IV, figs. 8-10), and
the specimens from the localities L 78 to L 82 (Plate IV, figs. 11, 12) have the third pleon
somite with much less produced angles which remind us of those of the typical form of
G. pule.v as figured by Sars.

The number of spinules on the lower margins of the third pleon somite is in luost cases
6, l_)ut varies rather considerably from 3 or 4 to 9.

(3) The number of spinules on the last three pleon somites. In most cases 2 median
dorsal and a pair of 2 lateral spinules are present on each side of both first and second pleon
somites. On the last somite the median dorsal spinules are usually absent or with only a
single median. The specimens of Togom Tso (L49, the highest Ideality) and Ororotse Tso
(L 52) have only 1 or 2 median dorsal and one pair of lateral spinules, or even none devel-
oped at all. Some examples of the arrangement of these spinules in the specimens from
various localities are shown in Table 2.

(4) Telson. The telson is usually a little longer than the length of the peduncles of the
third uropods, except a few cases (e.g., L81), in which the telson is nearly as long as the
peduncles of the third uropods. The specimens with the former type of telson, viz., longer
telsons, are very closely allied to Martynov's Gainiuants ocellatiis (loc. cit.). Both lobes of
the telson are more elongated and provided more richly with setae in the specimens from the
lower Kashmir localities (Plate V, figs. 1-4) than in those from the higher Ladak localities,
especially from L 78-L 82 (Plate V, figs. 5-7). Thus the specimens from the last-named locali-
ties have the telson of the typical form of this species, while the form of the telson of those
from the lower localities rathet resembles that of G. ocellatus.

The telson has usually 2 or 3 terminal spines, as seen in Table 2. A single lateral spine
near the base of each \o\x figured by Sars is not seen in the present specimens.

(5) The inner ramus of the third uropod (Plate V, fig. 8) is usually shorter than the
outer ramus, aljout 4/5 of the latter. The arrangement of the spines and fringes of plumose
setae on both rami varies considerably in the specimens from different localities.

(6) The form and size of the second joints of the last three pereiopods are rather differ-
ent not only in both sexes collected in the same locality, but also in the specimens from differ-
ent localities. (lencrally, the s]iecimens from the lower Kashmir localities have the pereiopods

66 AMPHIPOD CRUSTACEA OF THE GENUS GAMMARUS

"^ TABLE 2

Arrangement of Spinules on the Last Three Pleon Somites and the Number of

Apical Spines of Telson-

Locality . . .

Kl .

K15

K19

K23

K24

K24

Sex

?

$

$

?

c5

9

PI

1.4,2

2,0,2
2,2.2

2,2,2
2,2,2

2,2,2
2,2,2

2,2,2
2,2,2

2,2,2
2,2,2

P2

2 2 2

P3

.... 1,0,1

2,0,2

2,0,2

2,0,2

2.0,2

3,0,2

Telson . . . .

.... 2.(0)

2,2

2,2

2,2

2,2

2,2

Locality . . .

L 40*

L40t

L40t

L49

L49

L52

L60

Sex

S

S

$

£

9

<?

S

PI

.... 2,2,2

2,2,2

1,2,2

0,0,0

0, 0,

1,1,1

1(?),1,2

P2

.... 2, 2, 2

2,2,2

2,2,2

0,0,0

0,0,0

1,2.1

2, 2,2

P3

3. 0, 2

2,0,2

2,0,2

0,0,0

2,0,2

1. 1, 1

2, 0,2

Telson . . . .

2, 1

2,2

2,2

1

1,1

2,1

2,3

Locality . . .

L 72

L75

L76

L78

1.81

L82

LBS

Sex

£

9

£

s

£

?

? 9

I'l

.... 0, 0,

2,2,1

1,0,0

2,2,2

1.2.1

2,2,1

1,2.2

P2

.... 2, 2, 2

2.2,1

1,1, 1

3,2,2

1,2. 1

3, 2. 2

2,2,2

P3

.... 2,1.2

1,0,1

1.1,1

2,0,1

1 . 0. 1

3,0,2

1,1,2

Telson . . . .

2,2

3,2

liroken

2,2

S.3

2.2

2,2

* N. W. end.

t Littoral.

i Bottom at 31 m.

3-5 with narrower and more elongated second joints. The iiind margins of the second joints
in these limhs are much more convex in the female (Plate III, figs. 5-7) than in the male
(Plate III, figs. 8, 9), but their lower hind angles are roundcil in Ixith sexes. The second
joints in these limbs show a tendency to become reduced in length in the specimens from iiigher
altitudes (Plate \T). For instance the specimens from L 49 have the last three pereiopods
with the rather short second joint about 2/3 to 4/5 as wide as it is long, so that they resemble
very closely tliose of typical G. pule.v; their lower hind corners are somewhat ])roduced, their
hind margins more convex and the front margins often without spines or setae.

(7) The length and the number of the joints of the fiagellum of the first antenna.

The length of the first antenna is nearly as long as or a little shorter than half of the
body in the specimens from the high Ladak localities, Imt in the specimens from the lower
localities it is often much shorter than half of the body. The number of joints in the first
antenna shows considerable variations, as SpanJl (1923 and 1924) noticed in some luiropean
forms of this species. Some examples are sliown in 'J'able 3. In larger individuals this
fiagellum is composed of 25-28 joints, often reaching as many as 38 joints in some large
specimens from the lower localities, while the specimens from the very high IcKalities (L49,
L 52, etc.) have flagella of a much smaller numlier of joints.

" The figures between two annnias are the number uf median dorsal spinules.

AMPHIPOD CRUSTACEA OF THE GENUS GAMMARUS

67

TABLE 3
Measurements of Gaiuuiarus pulcx (Linne) from Xorth India

Number of

Length

Number of

joints of

Number of

Length

ofL

joints of

accessory

joints of

Altitude

of body

antenna

L antenna

flagellum

II. antenna

Group

Locality

in meters

Sex

in mm.

in mm.

mill. max.

min. 1 max.

min. 1 max.

^Kl

1200

9

20

10

35

4

18

K2

1585

9

19-21

11-12

33-38

2-4

14-17

K2

1585

S

15-17

8- 8.5

26-32

4-4

11-12

<•

K15, 19

1580

S

19-20

10-11

35 - 38

4-4

16-18

K23

c. 1585

$

15

8.5

29

4

15

K23

c. 1585

9

10

6.5

23

3

9

K24

c. 1585

$

13.5

8

29

3

12

K24

c. 1585

9

10

6.5

24

3

11

" ■

'L40a

4241

i

18-20

8-9

25-28

2-3

13-16

L40b

4241

S

15-17

7.5-8

25-28

1-3

13-14

L40c

4241

i

17-17

8-8.5

27-31

2-3

14-16

Camp 9

4269

9

16

8

28

2

11

L49

5450

S

12

?

5

2

9

OQ-

L49

5450

9

14

4

16

2

7

L52

5297

S

16-17.5

6.5 - 8.5

19-25

1-2

12-15

L60

4725

$

16.5

8.5

28

4

?

L72

4336

$

21

9

26-28

3

14-15

L72

4336

9

16

7

■?

p

?

L75

4385

9

c. 10

?

20

2

11

L76

4875

i

15

6

21-22

2

13-16

L78

4686

$

16

7

22

2

12

'-'

L81

4672

$

17-17.5

6

19-21

2

14-15

L82

4528

aS

12

5.5

20

2

8

L82

4528

h$

18

9

28

2

14

L8S

4536

S

12

6

21

2

13

The numljer of the juints of the flagellum in the second antenna also fluctuates greatly,
from 11 to 18, sometimes as few as 7, 8, or 9. The presence of calceoli on this antenna of
the male was not detected in all of the present specimens.

(8) Some other characters. The ecologically most interesting feature is the relative size
of the gill-lamallae in the specimens from different localities, particularly at different alti-
tudes. In the individuals froni the lower Kashmir localities the branchiae are relatively
smaller in size than in the specimens obtained from the higher localities. Several examples
of gills of the specimens of nearly same body length are shown for comparison in Plate VII.
Dodds and Hisaw (1924) demonstrated the comparable fact that in swift and well-oxygen-
ated waters the respiratory organs of some aejuatic insect larvae show a tendency toward
reduction.

The bodies of the specimens from the Kashmir localities and the higher districts l)eyond
the Ladak Range have good deposits of lime and are rather hard and fragile, while those
from the localities south of the Ladak Range have relatively soft bodies with poor deposits

68 AMPIIirOD CRUSTACEA OF THE GENUS GAMMARKS

of liiiK. Tlio former form has a larger body and is less liairy than the latter form, which
is often provided with very rich hairs.

A similar phenomenon was also noticed when comparing the littnral and pelagic forms;
the body of the latter, taken by a vertical haul, is softer and nmch more beautifnl than the
former.

"(^ •!• V "l^ "f.

Examining and comparing many specimens from various localities, I came to the follow-
ing conclusion. In North India, lx)th Kashmir and Ladak, there ajjpear to be distributed at
least two different races of Caiitinarus piilc.v, though there exist some transitional forms
which are difhcult to separate distinctly from one or toth of them. One form, which has
the shortened antennae, the elongated telson, the slenderer second joints in the pereiopods
3, 4 and 5. and the third pleon somite with considerably prolonged and pointed hind corner,
is found in both lower Kashmir localities (Table 3, I A) and the higher localities in
Indian Tibet north of the Ladak Range (Table 3, IB). The other form, which strongly
exhibits the typical features of G. pulex, was collected in the localities south of the Ladak
Range, within and south of the basin of the Upper Indus River (L78 to LBS). In the
intermediate districts in the Ladak Mountains south of I'anggong Tso (L 75, L 76, etc.),
there is found a transitional form between the above-mentioned two forms.

The first-mentioned form (Kashmir localities and L40-L72) is very clo.sely allied to
CJaiiiinanis occllatus Martynov of Lake Issyk-koul in Turkestan, but differs, from that in
having smaller eyes. Although this form has some peculiar features such as the prolonga-
tion of the lower hind corner of the third pleon somite, I believe there is no reason to estab-
lish a new si)ecies for it with such characters as already discussed. In view of the great varia-
tion in some characters which are regarded as of s[)ecific im])ortancc by many authors, we
would be compelled too often to create a new species or subspecies (cf. also Spandl 1923 and
1924, especially the latter ])ai)er, p. 451). Moreover, on the other hand, a comparative study
of numerous specimens from different localities makes it difficult to separate them into one or
more distinct species or suljspecies. .As in some other Crustacea, such as Daplniia and Bosinina
among the Cladocera, especially pelagic forms of them, it seems quite natural that Gammarus
piiJc.Y may also become highly differentiated into numerous races in various localities, and
many forms among the species ascriljed to Gaiiuiiarus with very close relationship to ptile.v
should be regarded as the local differentiates of G. pulcx. With this in mind, I have referred
all the present si^ecimens to one common species, G. pulex, avoiiling the use of a number of
specific names for different forms. Gainmanis occUatus Martynov, G. occllatus minor
Martynov and G. bcrgi Martynov, all descril>ed from Lake Issyk-koul of Turkestan, may be
thus regarded as the different races (probably subspecies) of G. pulex which have been
developed in that locality.

Since the alxive was written, a paper relating to some gannnarids from the western parts
of Asia was published by Karaman (1934). In this paper (pp. 127-129), Karaman has
described a new gammarid, Rwuloga)iii)ianis stolicckae, whose type-specimen was collected
by Stoliczka in 1864 in the vicinity of Lake Tso-Moriri, Prov. Rupschu, Ladak. According
to the original description, this new gammarid is characterized by the 3-jointed accessory
ilagellum of the first antennae, the rather long dactyli of pereiopods, and by the second and
third pleon somites, each with pointed hind C'lrner and provided with long setae on the lower

AM I'll lIMili ( KUSI A( l.A in- I'lll'. CENTS CAMMAKUS 69

margins, and so forth. Karaiiian stated tliat such characters as the longer dactvH and long
setae on the pleon somites show that the animals may have teen originated from a deep lake.
The Yale North India Expedition brought hack two samples of gammarids of Tso-Moriri
(L82), one being captured among the weeds of the estuarine water at the north end of the
lake and the other (only one individual) obtained by a vertical haul (30-0 m. ). These speci-
mens agree rather well with the typical Gammarus pidc.v in various characters, except the
third pleon somite which has a somewhat produced hind corner. The elongation of the
dactylus of each pereiopod as noted by Karaman is also seen in our specimens from Tso-Moriri,
not only in an animal obtained by a vertical haul, but also in many individuals collected among
the littoral weeds. In the other localities, e.g., Mitpal Tso (L 76), Khyagar Tso (L 81), etc.,
relatively longer dactyli were usually observed only in the forms obtained by vertical hauls,
so that such a character seems to be peculiar to the forms of free-swimming life or of deep
water inhabitants, as Karaman suggested. In other words, it must be regarded as an
ecological character due to the peculiar nature of environments; it is not of specific
importance.

With regard to Rivulo gamtnanis stuliczkac, Karaman has pointed out no clear differen-
tial diagnosis from the other allied forms, only writing as ", so jener des R. pulex oder
R. balcanicus mc\\i?, Gemeinsames" (p. 129). In some crustaceans like gammarids, which
show great variations in various body parts according to difference of environment, it is
a very difficult matter whether we are to recognize various forms with slightly different
characters as a separate new species or to treat all different forms as variations of a single
species. In regard to the various forms of gammarids from Kashmir, Ladak and Western
Tibet, as already discussed in the foregoing pages, it is my present opinion that it is best to
distinguish all the related forms as the peculiar local races or subspecies of Gammarus pulex,
thus avoiding the use of one or two new specific names for those forms. As Karaman
(1. c. p. 128) writes as "Oberflachlich einen R. pulex ahnlich," R. stoliczkae may also be a
local race (subspecies) of the pulex-se.v\ts of Gammarus (s. lat.), which has differentiated in
this region of Asia. Karaman, moreover, noted that R. stoliczkae may belong to the oriental
group of Rivulogammarus (1. c. p. 129).

The genus Riznilogammarus has been used by Karaman for certain forms of the pulex-
series of Gammarus, including pulex itself. I have at present only very scant knowledge as to
the validity of this genus, because I have been unfortunately unable to see its original descrip-
tion. If this genus is adopted, all the forms here concerned may be placed under it. As far
as I am able to understand, it seems to me that it may be better to use the generic name
Rivulogammarus as a subgenus of the genus Gammarus. In the present paper, therefore,
with this in mind, I have referred all the forms to the genus Gammarus (s. lat.).

3. DiSTRUujTioN .\N'n Its Limiting Factors
A. GKO(a<ArnicAi. Distriuutkjn

So far as our present knowledge goes, Gammarus pulex (Linne) is the commonest
freshwater amphipod, being distributed over the Palaearctic region, from England to Japan
and some parts of North Africa, but not found in America. In the high altitudes in Asia,
Chevreux (1908) recorded this species from some parts of Turkestan, such as Lake Issyk-koul
(1615 m. above the sea), the Pass of Karacolum (2000 m.), Lake Tchatyr-knul (3200 m.).

70 AMIMIH'on CRl'STACKA OK T 11 K CF.NUS CAM MARTS

etc. From tlie first-n.-iiiR'd lake. Marlyiuiv (l'*3C)j alsn recorded (/. piilc.v t(}t;ether with two
other new species and mie new sul>s])ecies (see foregoing- chapter). Tattersall (1914) first
recorded this species in the region north of India near the western ])art of Tibet. He found
C. pitlcx in a collection made in a pool on the summit of Killik Pass hetw'een the Northern
Hunza Range and the Taghdunkash, Pamir (15,600 feet) and in pcmls near the hanks of the
Killik River. The former locality is the highest hitherto recorded as the habitat of G. piilc.v.
Therefore, the discovery of this species in Togom Tso (5334 meters or 17,506 feet) on the
western border of Tibet by the Yale North India Expedition may lie stated to give a new
record, surpassing the altitude nf the I'aniir locality nunti(}ned ahoNc. Tiie Russo-German
Pamir Expedition in 1928 also collected G. pulex which was examined by I)r. A. B. Martynov.^

In the other parts of Asia, especially east of India, we have only a few records of this
species. Tattersall (op. cit. ) noted its occurrence in Lake Tali V\\ (Erh Hai, Shan-kuan),
Yunan, China, at an altitude of c. 7000 feet. In Japan G. piilcx, first recorded by Tattersall
(1922) from Lake Biwa, is common everywhere, though in the northern parts it is replaced
by G. annandalci Tattersall which is also distrilnited in China ( Tai-Hu, Shanghai).

As described in the foregoing chapter, the (/. pulcx of Lake Issyk-koul has some peculiar
characters compared with the typical form of the species, and the specimens of the same
species collected by* the Yale North India F.xpedition resemble rather closely those from Lake
Issyk-koul. It is supposed from this fact that such races of G. pulcx are rather w'idely s])read
in the vast areas of the high alpine regions in Asia.

6'. ocrllatus, G. occllatus minor and G. bcrgi are the representative races of the North
Indian and Turkistan dift'erentiation of G. pulcx in Asia. The races of the typical form are
distributed in the plain areas from h'urope to Japan. The centre of distribution of the alpine
races is not ascertained at present.

B. Distribution in Kashmir and Ladak, and Its Limiting Factors

As seen from a glance at the list of the localities in which G. pulcx was collected
(Table 1), its distribution in North India is very peculiar and interesting. The species is
common in Kashmir at altitudes up to alx)ut 1600 m. It is however quite absent in the lower
parts of Ladak to the east of Kashmir ( 1600-3800 m. above sea level), Init it reappears again
near the Tiljetan borders at altitudes over 3800 m. It is fpiite common in the lakes of these
high alpine regions, among which Lake Togom Tso is the highest locality, as mentioned
already. I shall discuss below shortly the very interesting problem as to what factors might
cause such a peculiar regional distribution.

1. Physical and Chemical Factors

A number of physical and chemical determinations made on waters from manv locali-
ties during the expedition is summarized in the following table (Table 4).

a. Water temperature. The water temperature does not seem to limit the occurrence of
G. pitlcx in the area investigated, since it ranges from O'C. to as high as 22 ^C. As generally
known, this species is an eurythermal ubiquitous crustacean.

° These specimens from Pamir differ from the typical form of (he species by having somewhat larger eyes
(Dr. Martynov, personal communication dated May 27, 1933).

I

AMPHIPOD CRUSTACEA OF THE GENUS GAMMAKUS

71

TABLE 4
Phj'sical and Chemical Conditions of the Waters In North India

.Alkali reserve

Altitudes

Water temp.

( Methylorange

Locality

in meters

°C.

titration)

pH

Gammarus

1. Kashmir localities

1200-1600

11.9-19.0

0.0014-0.0017 N

8.3-9.3
(diurnal
variation
of K 19)

present

2. Intermediate localities. .

1600-3,SOO

7.0-19.0

0.0007-0.0036 N

7.2-8.6

absent

3. High Ladak localities. .

3800-5400

0.0-22.0

0.0003-0.0610 N

7.1-9.6

present

b. Chemical factors.

i. Dissolved oxygen content. Gammarus pulcx is a mesoxybiont (after Steinniann and
Siirbeck, quoted in Wundsch) which demands water rather rich in dissolved oxygen. Though
I have at hand no data of the dissolved oxygen in the waters of Kashmir and Ladak, it is
difficult to suppose that the dissolved oxygen may play a great part as a limiting factor.
According to Dr. Hutchinson's information, Lake Khyagar Tso in Ladak had an absolute
oxygen deficiency on the bottom. The gammarids in that lake seemed to l)e living as a free-
swimming form in open water, since immense numbers of them were found settling on the
anchor rope while one was making limnological observations.

ii. Salt content of water, especially calcium. The correlation between the distribution
of Gaviniarus and some chemical environmental factors, especially calcium dissolved in water,
is discussed by several authors (cf. also Pia 1933). Thienemann (1912) found that the
gammarids in the torrents of Baumberg districts, rich in calcium, are larger in size than those
found in the torrents of Sauerland, where the water is very poor in calcium. Wundsch
( 1915) observed in the Sieg, a branch of the Rhein, that Gai)imarus pulex entirely disappears
from a biocoenosis when the calcium content of the water Ijecomes less than 9-10 mg. per litre.
Schumann (1930) writes that at least 13 mg. per litre of CaCOg is necessary for the forma-
tion of the shell of Gammarus, because newly ecdysized gammarids with soft bodies take up
the lime for building up their shells in the state of bicarbonate from the water and then make
it monocarbonate in their bodies. Since COo-free water can dissolve only 13 mg. per litre of
CaCOg, below this lime content gammarids cannot use the lime for the formation of their
shells, being obliged to live as soft-bodied animals which have no external protection against
osmotic action. On the contrary, Pentland (1930) concluded that the chemical composition
of the water does not appear to control the distribution of Gammarus, the temperature of
water, vegetation, and the presence of enemies alone limiting the distribution. .Schlagintweit
(cited by Hesse 1924) reported an interesting fact that in Ladak the Puga torrent, rich in
borax, is favourable for crustaceans and fishes.

A study of the physico-chemical data relating to North Indian waters, presented above
in Table 4, makes it very difficult to correlate the non-occurrence of Gammarus with the
alkaline reserve and the |)H-values of the waters in the zone between 1600 m. in Kashmir and
3800 m. in Ladak. The waters containing gammarids at high altitudes often have a
lower alkaline reserve and lower pH-values than those in intermediate region where
Gammarus does not occur. According to Schumann (op. cit.), the optimum pH of the water

/2 jXMl'IIU'Ol) CUrSI AlKA UK Tll|-. CKNl'S CA M M AKUS

for Gamjmints is about 7.i-~.S ami, wlicu tlic \a1uc falls In-low ().3, llic water has a toxic
effect upon Gammarus.

Ik'sides such a chemical relation in the area of the present iinestigation, all of the llow-
ing- water in the intermediate region has run ilowu from hi_i;h valleys at alliiiules at which
the species occurs. It seems therefore very improbable that such a peculiar distribution is to
be explained by chemical factors alone, though in some other parts of the world, as cptoted
above, calcium content of waters would play an important role in this respect. It must be
added that Gaiiiinanis was found in immense numbers in Lakes Panggong Tso and Khyagar
Tso in T-adak, both ha\'ing water of brackish nature.

2. ]->i()lo^ic(il I'ticlors

The types of habitat of G. piilr.v in the area of investigation vary very much, including
swampy pools covered with Lemna, large open-water lakes and streams with a rapid flow.
There niav therefore be no lack of food for Gainiininis. although I have at present no data
relating to this point. Pentland (op. cit.) is of the opini(jn that the food vegetation is one
of the important limiting factors upon the distribution of G"(7;/n;/((/-h.s-. If the source of lime
upon which Gamniants depends should be in some plants containing calcium compounds, such
as Chara or Lcmna, as suggested by Schumann (op. cit.), the ab.sence of sucli |)lants in the
intermediate region in Kashmir must be considered.

Finally, there is a more important biological factor, namely natural enemies, especi-
ally fishes. On this point, Dr. Hutchinson wrote to me that "the presence or absence of fish
cannot be a complete explanation for the distributions, because Gaiiiiiianis may be taken in
streams and lakes where there are many iish, as in the stream between Tangtse and Mugleb,
and in Pangur Tso and Yaye Tso, and there is no very great difference between the fish fauna
of the upper and intermediate zones." However, in the brackish-watcr-lake Khyagar Tso,
Gamniants was found very abundantly, swimming in the open water, and in this locality its
abundance is probably correlated with the complete absence of fish. Such a case was also
observed in some Japanese lakes, where there was an abundance of gamniarids before the
introduction of trout, and Pentland (op. cit. ) also writes that the presence of trout appears to
control the distribution of Gaiiiiiiarus. li lish are in any way a limiting factor for Gammarus
in North Indian localities, it is probable that the presence of genus Schizothorax of the
family Cyprinidae rather than of fish in general is concerned. According to Dr. Hutchinson's
statement, he observed no locality in which Schizothorax occurs with Goiiunanis except
Wular Lake in Kashmir.

Note by the Biologist of the Expedition

Gammunis was found in several samples after the collection had been forwarded to
Dr. Ueno and examined by him. These sam])les are enumerated below:

L i? Between Tangtse and Mugleb, stream, all. e. 417.^ ni. _'(> June, 1932.
L 38 Two to three miles west of Mugleb, alt. c. 4200 ni. 17 June, 1932.
L 47 Chagra, alt. 4636 m., warm spring, temp. 21.7 C. 8 July, 1932.
L 74 Pangur Tso, alt. 4329 m., margin at west end. 12 .\ugust, 1932.
L75 Chushol, spring, alt. c. 4330 m. 12 .\ugust, 1932.

AMi'Mii'dii ('i<i!S'ia<i:a n|- 'rill-, cknus cammauils 73

yVll these speciiiK-ns, i-xcei>t tlinse frdiii the lasl lncahly, apiiear l<i ha\e a pnjlonged
and pointed hind corner to the third pleon segment and an elongate telson : the Gammarus
from the spring at Ciuishol, on the other hand, exhibit indications of the characters of the
specimens from the southern part of Indian Tibet, particularly in their shorter telsons.
These records, therefore, confirm Dr. Ueno's findings based on the major part of the
collections.

The following additional ecological evidence, moreover, supports some of Dr. Ueno's
conclusions.

The oxygen content of the immediate environment of CaiiiiiKinis in the high regions of
Indian Tibet lay between 5.6 and 7.7 mgrms. per litre. In a stream at Dras, in the inter-
mediate zone and so free from Gaminarus, it varied from 6.2 to 6.7 mgrms. per litre; and
in a swamp at Spithug, a very favourable habitat in the intermediate zone, again without
Gamiiuinis, the water in contact with weed was probably supersaturated by day, though no
figures relating to this locality are forthcoming.

Gravimetric analyses of the waters of some of the lakes have been made by Mr. J. A.
Newlands of Idartford, Conn., and it is now possible to state that the calcium content of the
Gammarus localities in Indian Tibet varies from 9 parts per million (Ororotse Tso) to 303
parts per million (Panggong Tso). Since the water of the former locality had the lowest
alkali reserve encountered (0.0003 N.) it is safe to assume that the waters of the intermediate
zone (alk. res. 0.0007 to 0.0036 N.) contained sufficient calcium to support a population of
Gmtimarus.

Potamogeton pcctinatiis Linn, and Ranunculus triclwpliyllum Chaix (kindly determined
by Dr. E. D. Merrill and his staff at the New York Botanic Garden) were the dominant
species in such of the high Gammarus localities as contained higher plants, but more often, as at
Ororotse Tso, Togom Tso, Khyagar Tso, Panggong Tso, etc., no higher vegetation occurred.
Both P. pectinatus and R. trichopliylhim as well as charophytes occurred in suitable localities in
the intermediate zone. It is therefore clear that there is no correlation between the vegetation
and the presence and absence of Gammarus.

Dissection has shown that the pelagic Gammarus of Khyagar Tso is apparently pre-
daceous, feeding on the large, dark, slow-swimming Cladoceran Dapliniopsis, the deep sepia
lirown chitin of this form being easily recognizable in consideralile quantity in the faecal
matter at the posterior end of the alimentary canal of Gammarus from a vertical haul taken
in this lake. Dapliniopsis occurred nowhere in the presence of fish (e.g., Pangur Tso, Yaye
Tso) though some lakes where no fish were found (Ororotse Tso, Tso-Moriri) lacked also
Dapliniopsis. In \ iew of these finflings, taken in conjunction with the results of other
workers who have studied the relation of Gammarus to fish, quoted above by Dr. Ueno,
there can be little doubt that the abundance of Gammarus in the closed and mineralized waters
of these Central Asiatic lakes is due largely to the absence of fish, which leaves Gammarus
unmolested on the nne h.and and on the other introduces no competitor for the most con-
sincuous source of food of these Amijhipncjs. — G. E. II.

74 AMI'llll'iip CKl'STACl-^V or TIIF. GENUS CAMMARtIS

JUIII.IUUK AIMIV

Chevreux, Ed. 1908. fltiules sur la faune de Turkestan hasces sur les materiaux recueillis
par D. D. Pedasclienko (1904-1906). Travaux dc la Societe Imp. des Naturalistcs
de St. Pcter.sburg, T. 17, livr. 2, pp. 106-109 (Russian text); pp. 96-99 (French
text).

Chevreux, Ed., and L. Eac.k. 1925. Amphipodes. l'"aune de France, T. 9. Paris. 1925.
])p. 253-254.

DoDDS, G. S., AND Y . L. HiSAw. 1924. Ecological studies of aquatic insects. II. Size of
respiratory organs in relation to environmental conditions. Ecology, vol. 5, pp.
262-271.

Hesse, R. 1924. Ticrgeograptiie auf okologischer flrundlage. Jena.

Karaman, S. 1934. Ueljer asiatische Siisswasseraniphipoden. Zool. ,\nz., I'.d. 106, 5/6,
pp. 127-134.

Maktvxov, a. B. 1030. .Ampliipoda from tlie Lake Is.syk-koul. .Materials of the Com-
mission of the Expedition; Researches of the Acad. Sci., Pt. 11, Expedition to the
Lake Issyk-koul of 1928. Par! 1. Leningrad. Pp. 51-65 ( Ru.ssian text); 66-70
(English text).

Penti.and, E. S, 1930. Controlling factors in the distrihution of Gammarus. Trans.
Anier. Fish. Soc, vol. 60, 1930, pp. 1-4 (reprint).

I'lA, Juuus. 1933. Die Kalkbildung durch Tiere. Eine Uelx;rsicht der Fragen, vorziiglich
der chemischen. Palaeontol. Zeitschr., Bd. 15, 2/3, pp. 154-195.

Sars, G. O. 1895. An Account of the Crustacea of Norway. Vol. I. Amphipoda. Chris-
tiania and Copenhagen.

Schaferna, Karel. 1922. Amphipoda Balcanica, with notes about other freshwater
Amphipoda. Vesnt. Krai, ceske Spolecnosti Nauk. tr. 11, I'niha, pp. 33-35, p. 98.

ScuLAcuNTWEiT, H. V. (citcd by Hesse, p. 306).

Schumann, E. 1928. Experimentelle Untersuchungen iiber die Bedeutung einiger Salze,
insliesondere des Kohlensauren Kalkes, fvir Gammariden und ihren Einfluss auf
deren Hautungsphysiologie und Lebensmoglichkeit. Zool. Jahrb.. Abt. f. allg. Zool.
und Physiol., Bd. 44, pp. 623-704.

Standi. , II. V^'llt. Bcobachtungen an Gammariden (\(irl;iuligc Alitteilung.). Verhandl. d.
nalurforsch. \'er. in Briinn, Bd. 58, ]ip. 1-3.

1924. .Studien iiljer Siisswasseramphipoden. 1. .SitzungslK'r. d. Akad. d. Wiss. in

Wien, Alathem.-naturwiss. Klasse, Abt. I, Bd. 133, Heft 9, pp. 444-452.

Stebbing, T. R. R. 1906. Amphipoda. I. C/animaridea. Das Tierreich. Lfg. 21. Berlin.
Pp. 460-462 and p. 474.

AMl'Iill'OI) CRUSTACEA OF TIIK GENUS GAMMARUS 75

'I'atteksai.l, W. M. \'^H4. Notes on some Amphipods collected on llie I 'amir at an altitude
of 15,600 feet. Rec. Ind. Mus., vol. 10, pp. 213-14.

1922. Amphipoda with notes on an additional species of Isopoda. Mem. Asiat. Soc.

of Bengal, vol. 6, pp. 451-52. Calcutta.

Thienemann, a. 1912. Der Bergbach des Sauerlandes. Faunistisch-biologische Unter-
suchungen. Internat. Rev. d. ges. Hydrobiol. u. Hydrogr., Biol. Suppl. Bd. 4.
1912. Stuttgart.

WuNDSCH, H. 1922. Beitriige zur Biologic von Gainiiiarus piilcx. Archiv. f. Hydro-
biologie, Bd. 13, pp. 478-531.

Explanation of Plate III.

Gammanis pidex (Linne) from K 2, Takht-i-sulainian, Srinagar, Kashmir.

Fig. 1. Gnathopod 1. of female.
Fig. 2. Gnathopod 2. of female.
Figs. 3-7. Pereiopods 1 to 5 of female.
Fig. 8. Pereiopod 3 of male.
Fig. 9. Periopod 5 of male.

MEM. CONN. ACAD., VOL. X.

PLATE III.

5 mm.

Explanation of Plate IV.

Ganunants pulex (Linne)
Epimeral plates of pleon somites

Figs. 1-3. Ep. pi. of pleon 1-3 of female, K 2. Taklit-i-sulainian, Sriiiagar, Kashmir.

Figs. 4, 5. Ep. pi. iif pleon 2-3 of male from the same locality.

Fig. 6. Ep. pi. of pleon 3 of male, K 15, east of Gagirhal Ixoad, Sriiiagar, Kashmir.

Fig. 7. Ep. pl. of pleon 3 of male, K 23, Nishat Bagh, Kashmir.

Fig. 8. l".p, pl. i'i pleon 3 df male, L 40, Panggong Tso.

Fig. 9. Ep. pl. (if pleon 3 of male, L 49, Togom Tso.

i''ig. 10. I4). pl. of pleon 3 of male, L 76, Mitpal Tso.

Fig. 11. Ep. pl. of pleon 3 of male, L Id^, Yaye Tso.

Fig. 12. Ep. pl. of pleon 3 of male, L81, Khyagar Tso.

MEM. CONN. ACAD., VOL. X.

PLATE IV.

^nm.

Explanation of Plate V.

Gaiiiiiianis pidcx (Linne)

Fig. 1. Dorsal part of three pleon somites and telsoii, K 1, Tal<ht-i-sulaiinan, Srinagar,
Kashmir.

Fig. 2. Telson, dorsal view, K 19, Gagirl)al, .Srinagar, Kashmir.

Fig. 3. Telson, dorsal view, K J3, Nishat Bagh, Kashmir, male.

Fig. 4. The same, female.

I'ig. 5. Telsiin, dorsal view, 1,78. Yaye Tso.

I'^igs. 6, 7. Telson, dorsal \iew, LSI, Kliyagar Tso, male.

l""ig. 8. Third ur(j])od, K 2, Takht-i-sulaim;in, Srinagar, Kashmir.

MEM. CONN. ACAD., VOL. X.

PLATE V.

1 TTITTI.

Explanation of Plate VI.

Gammarus pulex (Linne)

Fig. 1. Last pcrciopod of male, L 49, Togom Tso.

Figs. 2-4. Pereiopods 3-5 of female, L 49, Togom Tso.

Figs. 5-7. Pereiopods 3-5 of female, T. 71, Ciuishol.

Figs. 8-9. Pcrci(>|)i)ds 3 and 4 of male, 1.81. Kliyagar Tso.

l'"ig. 10. Pereio|)()d 5 of leinale, LSI, Kliyagar Tso.

I'ig. 11. Pcreiopod 5 of male, L 72, C hushol.

MEM. CONN. ACAD., VOL X.

PLATE VI.

-> 5 m:

vn.i.

Explanation of Plate VII.

Gammariis pulex (Linne)

Fig. 1. Pereiopod 1 $ and 9, L 49, Togom Tso.

Fig. 2. Gnathopod 2, inner, L 49, Togom Tso.

Fig. 3. Terciopod 1 of male, K 23, Ni.siiat Ragli, Kashmir.

Fig. 4. Pereiopod 1 of female, K 1. Uampur, Kashmir.

g. gill-lamella, ac. accessory gill. o. oostegitc.

MEM. CONN. ACAD., VOL. X.

PLATE VII.

ARTICLE VII

REPORT ON HYDRACARINA

By O. Lundblad, Stockholm
(Received March 1, 1934)

During the course of the Yale North India Expedition, Mr. G. E. Hutchinson collected
some water-mites, which he has asked me to work out. Because of the high altitudes at
which the main part of the material was collected, only some few species are represented.
Nevertheless the collection is a highly interesting one, forming a very valuable contribution
to our poor knowledge of the water-mite fauna of India, and it is a great pleasure to me to
offer Mr. Hutchinson my best thanks for giving me the opportunity of studying his material,
which contains quite a number of forms new to science.

Family EYLAIDAE

Subfamily Eylainae

1. Eylais liaiitata Koen.

Eylais hamata Koenike, 1897, pp. 282-83

Piersig, 1897-1900, p. 427

" georgei Soar, 1901, pp. 68-69

" longipons Daday, 1901, pp. 94-96

Eidais hamata Piersig, 1901, p. 22

Eylais longipons Daday, 1903, pp. 359-62

" hamata Halbert, 1903, p. 506
Eulais marenzelleri Thon, 1905, pp. 158-62

" hamata Thon, 1906, pp. 15, 44-45
Eylais " Koenike, 1909, p. 16

Koenike, 1910, pp. 152-53
Eulais " Lundblad, 1912, pp. 59-60
Eylais " Lundblad, 1912 a, p. 222
Koenike, 1919, pp. 521-24
Soar and Williamson, 1920, pp. 110-11
V. alpina Walter, 1922, pp. 247-49
Soar and Williamson, 1925, pp. 59-61
Viets, 1921, p. 342
Viets, 1928, p. 10

The collection contains some few, more or less mutilated, specimens, nioslly without legs
and palps.

In all characters still available for study the specimens agree well with European material.
The species was found for the first time in Palestine in 1895 and described from German
material in 1897. Since then it has been found in many other countries, as will be seen from
the foregoing list, which, however, does not complete the synonymy. Earlier authors, who

Mem. Conn. Acad., Vol. X, Art. VII. September, 1934.

86

HYDRACARINA

(lid nut possess our present knowledge of the great variability in the genus Ilylais, liave estab-
lished new species for some slightly aberrant specimens of E. liamata. From Asia Minor
Thon (1905) described E. marcnselleri, which later was withdrawn l)y Koenike (1910) as a

Figure 1. — Eylais luiinata Koen. 9. A, B, eyeplates; C, maxillary organ from beneath; D, from the right side;
E, right palp from tlie imier side: F. cpimeral striutiire: G, female genital field.

synonym of Iiaiiiata. E. marcnzeUeri is characterized by its very long and thin eyebridge.
Sul)sequently, in 1912, I descril^ed a nymph-form of liamata, having an equally thin and
slender, though not especially long, eyebridge (Lundblad, 1912 a, p. 222, fig. 1) and mentioned
also some adults with comparable structure. Soar in 1901 described E. gcorgei, charac-
terized by having the intcrcapsular bridge curved backwards in the middle. Soar and

IIYDRACARINA 87

Williamson (1920, p. Ill; 1925, p. 59) now regard this form as the same species as
E. hamata. Lastly Uaday, in the same year (1901) in his monograph on the Hungarian
Eylais-specics, published the description of a supposed new species, E. longipons, which
almost exactly agrees with E. gcorgei Soar, having an eyeliridge of the same shape. I do not
hesitate in considering this form a real hamata. The broad and short maxillary plate, the
immense overlapping pharynx with its distal hooks and the great mouth-disc indicate that
both forms are identical. The third palp-segment is more conspicuously protruded than in
hamata, but 1 think this character is referable to a mistake, like so many others in Daday's
drawings. Walter's var. alpina, finally, is merely a dwarfed form of hamata and in all other
respects quite typical.

E. hamata is an easily recognizable though rather varialjle species. The specific features
are to be found mainly in the eyeplate, the maxillary organ, the last palp-segment and, as
pointed out by Thon (1905, p. 158; 1906, p. 15, p. 45), in the structure of the epimera. The
eyebridge is always very long, but length, as w-ell as width, is variable. The maxillary
organ is always broad, with a short maxillary plate, a broad and immense mouth-plate and an
overlapping, hook-bearing, distally widened and rounded pharyn.x. The end-nails of the last
palp-segment are very blunt. The epimera present a close mesh-work of chitinous balks of
various thickness, quite different from those of most other species. A drawing may serve to
illustrate this peculiar structure. The thicker main balks always connect the front and the
hind margins {i.e. the longer sides) of the epimera, whereas the thinner balks run in dif-
ferent directions Ijetween the main balks. Lately I have drawn the attention to a somewhat
similar structure in E. iiintila Koen. (Lundblad, 1929, pp. 5-6.) In this species, however,
the epimera are still more chitinized, the meshes being reduced to narrow pores, so that no
balks can he distinguished. In E. hamata the structure of the epimera varies greatly, in some
examples no distinction can be made between main balks and secondary balks, the meshes
sometimes lacing more or less pore-like, though not so narrow as in E. mutila. Also the
nymphs of the two species in question are distinguished by the same respective characters.
In other species of Eylais there is no real meshwork, only main balks with some few connect-
ing secondary balks between them being developed (cf., for instance, Thon, 1906, p. 69:
E lati pons and p. 72: E. ineridionalis. Lundblad, 1929, Plate III, fig. 19: E. discreta).

Regarding all characteristics mentioned above, the present specimens are quite typical
hamata. All examples are females. The male possesses two semicircular genital plates like
most other species of the genus, whereas the female is destitute of real plates. Her genital
opening is surrounded in front by a great numlaer of long bristles. Originating from the
opening there is a subdermal, suture-like, chitinous rod directed backwards. Some short and
strong bristles are inserted in the skin on each side of the rod.

Locality. Indian Tibet: near Chu.shol, altitude ca. 4,340 m. 14 Julv, 1932.

Distribution. Most European countries, Palestine, Asia Minor, .Siberia. In Switzerland
it is fijuiul at 2450 m. above sea level (Walter, 1922, p. 247).

88 IIVDKACARIMA

2. Eylais degenerata K(3cii.

Eviais degenerata Koenike, 1897, pp. 292-93

Koenike, 1898, pp. 307-09
liiiliiis ■i'liriabilis Sig Thor, 1902, pp. 450-51

" " var. iiiagtm Sig Thor, 1902, p. 451

" intermedia Sig Thor, 1902, p. 451
" degenerata Nordcnskiohl, 1905, p. 2
" pscudoriinosa Piersig. 1906, pp. 380-82
" degenerata Daday, 1910, p. 239
Eylais angulata Viets, 1911, pp. 155-56
" " galcata Viets, 1911, pp. 156-57
" angulata Yiets, 1911a, pp. 351-54
" degenerata galeata Viets, 1911 a, pp. 354-57
" consors Szalay, 1912, pp. 70-73, 81
" eregliensis Szalay, 1912, pp. 73-77, 81-82
" stagnalis Szalay (non lialbcrt!), 1912, pp. 77-80, 82
" degenerata Viets, 1914, p. 83
" taurica Viets, 1914 a, p. 560
" degenerata hispanica Viets, 1918, pp. 19-23

microstoma Viets, 1921, pp. 419-20

Walter, 1922, pp. 64-65

galeata Walter, 1922, pp. 65-66

microstoma Walter, 1922, p. 66

galeata Szalay, 1926, pp. 21 1-12, 215
" consors Szalay, 1926, pp. 212-13, 216
" taurica Szalay, 1926, pp. 213-14, 216
" degenerata sumatrensis W&is, 1926, pp. 101-02

asiatica Viets, 1926 a, pp. Z7Q-72
" asiatica Marshall, 1928, pp. 602-03
" galeata Marshall, 1928, p. 603
" degenerata Viets, 1930, pp. 208-09

angidata Viets, 1930, pp. 209-10

This species was originally described from Madagascar, Egypt and East Africa. Later
Nordenskiold and Walter reported it from Soudan, Viets from the Cape Province. More
or less aberrant forms were described by Viets from East Africa (angidata, galeata, micro-
stoma), Spain (hispanica), Sumatra (siimatrensis) , and India (asiatica). Some of these
later have been met with in other places (angidata in Spain by Viets, galeata in llungar}'
by Szalay and in China by Marshall, and asiatica in China by Marshall).^ The forms angidata
and galeata at first were looked upon as distinct species, but after some time degraded by the
author himself to the rank of varieties.

Eylais degenerata is a very variable and widely distributed form that has often been
misinterpreted. There seems to be no doubt that Daday and Viets are quite right in identify-
ing Thor's E. variabilis, variabilis magna and z-ariabUis intermedia from the Cape Province
with the species of Koenike. The figure of the maxillary organ seems to me undisputably to
confirm this opinion. For the same reason Piersig's E. psciidorimosa from Sumatra prob-
ably also belongs to degenerata, as pointed out already by Viets. Another form, consors,
which Szalay describes from Asia Minor and which he afterwards refound in Hungary, is

' It seems somewhat uncertain whether the specimens mentioned by Marshall from China have been rightly
identified.

IIYDKACARINA

89

regarded I)y Viets, witli some hesitation, as being conspecific witli dcj^i'iirralti. 1 think Viets
is right in this identification and I may add that I regard also the other forms described in the
same paper by Szalay as merely synonyms of degcnerata, viz., eregUcnsis Szalay and taiirica
Viets {=stagnaUs Sz3\a.y)} Both these forms are also distinguished l)y their characteristic

Figure 2. — Eyiais degcnerata Koen. A, B, eyeplates ( 9 ) ; C, maxillary organ from the left side ( $ ) ;
D, from beneath ( 9 ) ; E, right palp from the inner side ( 9 ) ; F, right mandible from the inner side { $) ;
G, epimcra and genital field (9).

maxillary plate and pharynx. Tlie latter form, taiirica, was later refound by Szalay in
Hungary.

There is no other character to he relied upon in identifying E. degeiicrala but tlie struc-
ture of pharynx and maxillary plate (Viets, 1930, p. 209). vMl other characters vary, for
instance, shape of eyebridge, shape and chaetotaxy of palpi, shape and width of oral disc, and
length of posterior maxillary processes.

^ Concerning the change of name, see Viets, 1914, p. 560.

90 IIYDRACARINA

III Mnler to demonstrate the variability in the characters just mentioned the following
talile may l>e submitted of Eylais degencrata s. sir. and allied forms.

( 1 ) Oral disc :

wide (degencrata)

medium width (censors, taurica)

narrow {pscudorbiiosa, eregliensis, inicrostoma)

(2) Posterior maxillary processes:

shorter than pharynx (degencrata, variabilis, taurica. hispaiiica, asialica)

as long as pharynx (pseudoriinosa, consors, eregliensis, lulcrostoiiia. siniiatrensis)

(3) Intercapsular bridge:

V-shaped (degencrata, angtilata, sumatrciisis)

not V-shaped (all other forms)

narrow (iiiter)iiedia)

fairly narrow (microstoma)

fairly wide (magna, galeata, consors)

wide (variabilis, pseudoriniosa, eregliensis, taurica, hispanica, asiatica)

(4) Projection of 3rd palp-segment:

about 4 setae (pseudoriniosa)

" 6 " (sitinatrensis)

" 7 " (angulata, taurica)

" 8 " (cotisors, microstoma)

" 9 " (degencrata, variabilis, hispanica)

" 10 " (asiatica)

" 14 " (eregliensis)

(5) Inside of 4th palp-segment :

about 8 setae (degencrata: 3 spiniform, 5 pectinate)

10 " (pseudoriniosa: 3 spiniform, 7 pectinate; eregliensis, microstoma:

5 spiniform, 5 pectinate; taurica: 6 spiniform, 4 pectinate; siima-

trensis: 4 spiniform, 6 pectinate)
13 " .(angulata: 10 spiniform, 3 pectinate)

18 " (z-ariabilis: 10 spiniform, 8 pectinate; consors: 14 .spiniform, 4

pectinate)

19 " (asiatica: 5 spiniform, 14 pectinate)

20 " (hispanica: 4 spiniform, 16 pectinate)

(6) Out.side of 4th palp-segment:

alxjut 5 setae (pseudoriniosa: 3 spiniform, 2 pectinate)

9 " (angulata: 4 spiniform, 5 pectinate)

10 " (taurica: 6 spiniform, 4 pectinate)

11 " (hispanica: 6 .spiniform, 5 pectinate; variabilis: 11 spiniform)

12 " (consors: G .spiniform, r> pectinate; degencrata: 4 spiniform, 8

pectinate)

13 " (eregliensis: 6 spiniform, 7 pectinate; microstoma: 5 spiniform,

8 pectinate)
19 " (asiatica: 5 spiniform, 14 pectinate)

IIYnRACARINA 91

(7) 4tli palp-segment :

slender {degcncrata, consors)

rather thick {variabilis, pseicdoriinosa, crcgiicnsis, taurica. liispanica, asiatic. suina-

trensis)
thick {microstoma)
slender ( 5 ) or thick ( 9 ) {angulata)

From the above it will be seen that especially the palpi vary considerably. We find all
stages from few to numerous bristles. It must be remembered, however, that it is often diffi-
cult not only to distinguish and to count exactly all the bristles, but especially to decide whether
a bristle is feathered or not, depending upon the point of observation, i.e., the position and
direction of the bristle. It is obvious, therefore, that not t0(j much stress must be laid upon
statements concerning the structure of the l)ristles. It must also be kept in mind that it is
quite an exception to find two specimens of the same species of an Eyiais, in which the position
and shape of the palp-bristles are exactly the same, at any rate in a limited collection. There
is no doubt, therefore, that most species are extremely variable and that it is inappropriate to
separate species by means of minute differences in the armament of the palpi, the right and
left of which are often differently shaped in the same specimen. Not seldom the eyebridge
is asymmetric or deformed, sometimes entirely wanting, and specimens having but one well-
developed eye-capsule also have been met with (Lundblad, 1929, Plate III, fig. 26). In one
of the specimens of degenerata, here figured, there is an unpaired extra bristle in the middle
of the ey bridge.

The main feature in degenerata is the very short maxillary plate, by which the greater
part of the pharyn.x is laid bare. The maxillary plate is described as being coalesced with the
pharynx, without leaving any suture. However, in the specimens studied by me, there seems to
be a very fine, nearly invisible suture, which separates pharynx and maxillary plate from one
another. This suture runs not far away from the outer oral circle, so that anyhow the
maxillary plate is very short.

In the specimens before me the projection, of 3rd palp-segment bears 13-14 bristles, the
inner side of 4th seginent 17-22, and the outer side 10-13. In the 3rd segment the specimens
thus come nearest to eregliensis, whereas the inner side of the 4th segment resembles that of
variabilis, consors, asiatica, or hispanica, the outer side that of taurica, hispanica, variabilis,
consors, degenerata, or eregliensis. In the female the numl^er of bristles on the inner side
of 4th palp-segment is somewhat higher than in the male, corresponding to the condition in
some other species of Eyiais (cf. Lundblad, 1929, concerning E. infnndibidifcra and E. dis-
creta). The oral disc is of medium size and the intercapsular bridge varies from narrow to
wide, in both cases being rather straight.

The Kashmir specimens tlius present a remarkable intermixing of characters, making it
impossible to refer them to one of the "species" already described. This indicates, as far as
I can see, that there is no meaning in describing forms, separated by such slight differences
in number and situation of palp-bristles or in shape of intercapsular Ijridge, as distinct
species. Such a proceeding totally neglects the great variability prevailing in the genus
Eyiais. And I think it is not even worth while to give all the different forms the rank of
varieties, since it is obviously difficult to find another specime;i exactly satisfying the original
description in all structural details. Therefore, I regard the Kashmir specimens as being real
E. degenerata.

92 IIYDRACARINA

'riic iiKiIe genital opening is siirroim(U<l l>y two soinilunar, liristlc-bcaring genital valves.
The female has no genital plates, but some bristles are inserted in the skin lietween genital
opening and epiniera; at the anterior end of the opening tlie l)ristles are crowded, forming a
group of 4-5 bristles on each side.

Localities. Kashmir: Phashakuri (K33) altitude c. 1585 m., in a ditch, 10 April,
1932; Phashakuri swamp (K35), 10 April, 1932; Gagirbal Pond, Srinagar (K36), alti-
tude c. 1580 m., 11 April, 1932; Bakh Hajan, Jhil (K46), altitude c. 1575 m., 19 April,
1932; Anchar Lake, S. of Bandipur, marginal swamp, altitude c. 1580 m., 6 May, 1932;
Punjab: Sohawa, Rawalpindi dist. (P2-3), altitude c. 528 m., 3 March, 1932; Gungrila,
Rawalpindi dist. (PI), edge of shallow, weedy pool, altitude c. 525 m., 2 March, 1932.

Distribution. North, Middle and South Africa, Madagascar, Spain, Bulgaria, Ilungaria,
Asia Minor, Kashmir, India, Sumatra, China.

Family PROTZIIDAE

Subfamily Protziinae

3. Protziella hutchinsoni gen. et sp. n.

Generic diagnosis. Skin papillated. Eyes in capsules. Frontal organ lying in a shield,
the latter consisting of frontale, prae- and postfrontalia, dorsocentralia 1, and postocularia.
Frontal organ rudimentary, but clearly visible and redoubled, divided in two lateral parts,
each with a small spot of pigment. Palp chelate. Legs without swimming hairs. Claws
simple, as in Partnunia, not split up into a number of teeth. Genital plates present, situated
inside the acetabula.

At present there are four genera known within the family ProtzHdae,^ viz., Protzia, Part-
nunia, Calony.v, and Neocalonyx. One of these, Partnunia, has simple, all the others com-
posite claws. Protzia differs from Ccdonyx and Neocalonyx by the lack of genital plates.
Neocalonyx differs from all other genera in the palpi not being chelate and in tlie skin, tend-
ing to a development of chitinous plates. The new genus comes nearest to Partnunia, the
claws being simple, but differs remarkably from all genera by the structure of the frontal
shield and the fairly well developed frontal organ, which is composed of two distinct parts.
In the structure of the skin the new genus somewhat resembles Neocalonyx, a number of
small dorsal chitinized plates or punctures being developed. 1 f there is any closer affinity
in the genital area it is impossible to say so far, since it is difficult to ascertain the structure of
the genital organ in Neocalonyx from Walter's description, for instance, if the genital plates
are situated inside or outside the acetabula.

Description of species. Length of body 983ju. ( 5 ) - 1500iU ( S ). A detailed study of
the structure of the skin, the presence and situation of chitinized plates and dermal glands,
etc., in the family Protziidac has never been performed. Some years ago, however. I shortly
drew attention to the fact that Protzia cxiuiia has the dermal glands and dors.il bristles dis-
tributed over the skin quite after the same plan as the Thyasinae, for which I have drawn up
a special terminology (Lundblad, 1927, pp. 210 and 221-23). According to this we find on

' Or five, if also Wandesia Schecht., known hitherto as a nymph only, belongs to the family in question.

HYDRACARINA

93

each side an outer row of 4 lateroglandularia and an inner row of 7 dorsoglandularia. The
first dorsoglandulare is also termed antennifonn. Between the just-mentioned two rows
there is a row of 4 chitinized shields, the dorsolateralia, and on the medial side of the dorso-
glandularia we find a row of 5 other shields, the dorsocentralia. Near the frontal organ
there can sometimes be distinguished two small shields on each side, viz., the prae- and post-

FiGURE 3. — Protziella hutchinsoni gen. et sp. n. A, animal from above ( ? ) ;
C, frontal organ ( 9 ) ; D, maxillary organ from the left side ( $ ) ; E, mandible ( S).
side (9).

B, frontal shield ( 9 ) ;
F, palp ( 9 ) ; G, ventral

frontalia, and, outside of these, two bristles on each side, the prae- and postocularia. In the
Thyasinae there is a marked tendency to develop more or less strong and voluminous chitin-
ized shields, which either remain isolated from one another or coalesce in a different way in
different genera.

It is of considerable interest to find that the family Protsiidae presents the same chitin-
ized skin-elements and dermal glands as just described in the Thyasinae, and this fact indi-
cates a rather close relationship between tiic two families. In Profsia, Calonyx and Partnuiiia
there are no skin-plates developed, but according to Walter (1919) there are some in the

94

IIYDRACARINA

genus •A^eocalonvx. Beyond doubt these plates are arranged after the same scheme, though
\Vaher does not give any details. In the new genus Protciclla the chitinized elements around
the frontal organ are united into a frontal shield, which is built up from the prae- and post-
frontalia, the frontale, and the dorsocentralia 1. As in so many 7'hyasiiiar, there are two
bristles arising from the shield. These bristles are homologous with the postocularia in the
Thxasinae. The praeocularia are inserted free in the soft skin in front of the shield. In the
construction of the dorsal surface of the body the genus Protcuila thus puts one in mind of
the genera Paiiisoides and Paiiisopsis among the Thyasinae (cf. Lundblad. 1''33. Figure 12 a
and c), though in Panisoidcs and certain Panisopsis species some of the dorsocentralia and
dorsolateralia are much more enlarged.

The shape of the frontal shield in Protsiella hutchitisoiii will ]x seen from the drawing.
The shield is somewhat incised in front as well as behind, and the posterior margin is thick-
ened to fonn a callosity, whereas the rest of the shield is fairly weakly chitinized. The post-
ocular bristles are inserted in the lateral corners. The frontal organ is well developed and
divided into two regularly shaped lateral parts, with a pigment spot in each. Such a structure
we again meet with in some of the Thyasinae, viz., Euthyas and Thyasides (Lundblad, 1927).

S . The maxillary organ is 278/1 long. The upper side of the rostrum is convex, and
the rostrum is l>ent downwards. The mandible (including the clawj measures 286/i in length.
The lengtli ol the extensor and flexor sides of the palp-segments are (in /n) :

I

II

III

IV

V

Extensor side

Flexor side

71
46

111

2')

54
54

1Q3
125

30
32

The penultimate segment ends sharply ptMUted. and the second segment is ])r()\ided
dorsally with rather numerous bristles, whereas otherwise the i)alp is only poorly l)eset with
bristles.

The anterior two pairs of legs are short and robust and, like the two posterior ones,
provided with a great many strong spines, the longest of which are arranged in whorls at the
distal ends of the segments 3-5. The 4th leg dilYers by its considerable length from the other
ones, the 4th segment especially being prolonged. This segment increases in length from
the first to the fourth leg; the measurements for this segment in tlie difYerent legs are as
follows (in /^) :

1st leg: 147; 2nd leg : 182; 3rd leg: 220; 4th leg: 392.

The indifferently shaped epimera are widely separated into four groups. The anterior,
projecting corners bear tufts of long, backwardly directed bristles or hairs.

The genital area occupies the region I^etween the pcsterior pairs of epimera. The genital
opening is 260/^ long and bounded at the anterior end by a supporting, chitinized body.
Another chitinous piece is to l)e found farther back. The genital plates are much shorter
than the opening, but 121/^ long, tapering" towards Ixith ends and sup])lied with rather few,
thinly inserted bristles, most of which arise from the median border. The genital lips inside
the plates are papi Hated, the papillae being very small and scattered. Lateral to the plates
there are many acetabula, about 24 on each side. They var}- somewhat in size and are not
stipitate like those in most other Prolziidac.

HYDRACARINA

95

9 . The female does not differ essentially from the male. It is larger and the genital
area has not quite the same appearance, owing to the genital lips, which are convexly
swollen and much more projecting and conspicuous than in the male. They are covered by
papillae, the size of which equals that of the other papillae of the skin. This sexual diifer-
ence is similar to that in Protaia (Lundblad, 1927, p. 212), though the male in Protziella

Figure 4.— Protziella hutchinsoni gen. et sp. n. A, first leg ( 5 ) ; B, fourth leg C 5 ) ; C, genital organ {$)

D, genital organ ( 9 ).

has no bristles on the genital lips. The genital plates resemble those of the other sex, as do
also the acetabula, the number and shape of which also agree. As in the male there is only
an anterior supporting body : another chitinous spot lies at some distance behind the genital
field, free in the skin. Posterior to the genit:d plates there is a very small supplementary
genital plate on each side, carrying some few bristles. The genital opening measures 318/*
in length.*

' In the figure the male genital organ is more magnified than tliat of the female and thus seems to be larger,
but in reality it is rather smaller.

96

HYDRACARINA

In the male the epiineral groups are more crowded and the space between them is nearly
filled up by the genital organ, whereas they are more distant in the female, leaving plenty of
space between each other and the genital organ.

In both sexes some few bristles are scattered between the acetabula in the anterior part
of the genital field, and the excretory opening is surrounded by a ring of chitin.

Nymph. Like the adult the nymph is recognized by the prolonged last leg. In the only
specimen examined the genital organ consists of four acetabula on the right and three on the
left side. Judging from the situation of the acetabula probably three acetabula on each side
is the rule.

Figure 5. — Protziella hutchinsoni gcMi. ct sp. n. Genital organ of nymph.

Locality. Indian Tibkt: Shimsha Karbu between Dras and Kargil (K78), 22 May,
1932, in a spring (temp. + 8°C., pH. 7.8), altitude ca. 2819 m. ; springs (temp. + 7.2='C.)
4 miles from Bao, between Bao and Drugup (L35), altitude c. 4100 m., 26 June, 1932.

Systematic affinities. The genus Protziella differs from all other Frotziidac in a series of
characters, viz. : (1) the strong chitinization of the dorsal surface, a well-developed frontal
shield (and small dorsocentralia and dorsolateralia) being present. (2) the well-developed
frontal organ, with its double spot of pigment; (3) the non-stipitate acetabula, situated out-
side the genital plates.

In some of these characters the genus more resembles certain ineni])ers of the subfamily
Thyasinae than the Protsiidae. Very often the dorsal surface of the body is more or less
chitinized in the Tliyasinae, and in some genera there is, as already mentioned, a double pig-
ment spot. The non-stipitate acetabula also gives the animal an appearance somewhat
unfamiliar to a Protziid and more in correspondence with that of a Thyasin."' The situation
of the acetabula outside instead of inside the genital plates is, however, a character unknown
in both groups.

Like Eutliyas and Thyasides among the Tliya.<:iiiac — both with two spots of pigment
in the frontal organ — Protziella seems to occupy quite an isolated position among the
Protziidae. It has been already mentioned that in some respects — chitinization of dorsal tody
surface, frontal organ, non-stipitate acetabula — Protziella resembles certain Tliya.<;iiiae, but
probably this agreement is to be explained by convergence. However, it is impossible to
deny the affinity of the Protziidae and the Thyasinae (or 1 1 yd rypJwntidae) . W'c find another
similarity between the two groups in the complicated claws which Viets recently (1929) has
described in the aberrant genus Tcratothyas, belonging to a special subfamily, Teratothya-
sinae, among the liydryphantidae. The claws in that genus are said to be like those of

'^ It must be remembered, however, that according to Walter tlic acetabula in Catonyx hilus arc non-stipitate.

HYDRACARINA

97

Protzia or Calonyx, i.e. being dilated at the apex and consisting of a main claw and some
lateral teeth. It is interesting to find that we are able to draw parallel lines of develop-
ment within both families (claws, frontal organ, chitinization of body). Even if some of
the characters are to be regarded as being independently acquired by the two families, some
others may well be of common origin. In this connection it is of a great interest to draw atten-
tion to a paper of Motas (1929) in which he has described the larva of Calonyx brcvipalpis.
His drawings, compared with the figures given by other authors and the present writer ( 1927),
of the larvae of some Thyasinae and Hydryphantinac again reveal the fact that all these
three groups undoubtedly are nearly allied to one another. However, the declaration of Motas
(1. c, p. 261) that the Protsiidac occupy an intermediate position between the Thyasinae and
Hydryphantinae seems to me to require further evidence. I think it is better to join, as
hitherto, the subfamilies Thyasinae and Hydryphantinae into the same family, Hydryphan-
tidac, and to place the Protsiidac in the vicinity of the Hydryphantidae.

4. Calonyx montanus sp. n.

(5 . Length of body about 965/it. Skin without chitinous plates. Frontal organ very
small, hardly bigger than a skin-papilla and lying some distance behind a line connecting the
composite eyes. The organ looks like a rounded, circular papilla, which projects a little more
over the skin-surface than do the ordinary papillae. The row of dorsoglandularia is char-
acterized by the third dorsoglandulare having been much displaced in lateral direction, so
that the distance between the two glands of that pair is distinctly longer than that between
the eyes.

Length of maxillary organ 275/i. Seen from the side the rostrum is rather flat. The
ventral surface projects angularly over the rest of the rostrum and the dorsal side is undu-
lated. The mandible is 264/<. long from the base to the tip of the membrane, which tapers and
is sharply pointed at the end. The claw is 89/* long. The bristles nn the palpi are few in
number and the lengths of the palp-segments are (in ft) :

I

II

III

IV

V

Extensor side

29
39

93
35

50
46

150
79

42

Flexor side

28

The projection at the end of the fourth segment is long and slender, with the tip bent
ventrally.

The bristles on the epimera are few and principally restricted to the lateral, anterior
corners of the first three pairs. The first pair has a forward protruding corner, bearing
about 10 rather short, stout, more or less spine-like bristles; on the second and third pairs there
are about 4-5 bristles. The suture between the first and second epimera vanishes before reach-
ing the median border. The greatest length of the anterior group of epimera is 268/u, that of
the posterior group 246^. In the third epinieron the anterior and median borders form
together a continuously curved arc, not being angularly bent so as to constitute well-
distinguished anterior and median borders, as in the following species.

The last segment in all legs is considerably thickened toward the distal end, which is pro-
vided with two strong claws of different lengths. Each claw consists of a central tooth of

98

IIYDRACARINA

considsrable length and thickness and a great number of shorter and thinner lateral teeth.
These are so arranged that about 15 stand on one side, about 5 on the other side of the long
tooth. Seen from the side, the last segment of the leg shows a deep incision and the upper
side of the segment, which limits the incision from above, forming a long, triangular projection
(see the figures for the following species). Especially the segments 3 and 4 exhibit a row of
stiff bristles of middle length along the extensor side. The number of these bristles does not
exceed 7. On the lirst leg these bristles seem to be a little shorter than on the others.

Figure 6. — Calonyx montanus sp. n. A, animal from beneath (S) ; B, maxillary organ from the left side ( $) ;
C, palp ( 5 ) ; D, E, claws ( 9 ) ; F, genital organ (S) ; G, genital organ ( 2 ) .

The structure of the genital organ shows that the species in question is a true Calonyx,
genital plates being present. These plates, however, are extremely poorly developed, consist-
ing only of a narrow strip of chitin, wide enough just to support the bristles, which are
inserted to the number of about 12, forming a single row. The strips are so short that only the
anterior half of the genital organ is enclosed by them. Three pairs of acetabula are situated
here, whilst the rest lie behind the strips. These latter acetabula are more numerous and
more elongated than the others. The genital lips inside the acetabula are furnished with
some very small, but rather high and well-marked papillae. Posterior to the genital organ
the skin is chitinized as a small shield.

9 . The female diiifers, except in its larger size, very little from tlic male. The antero-
medial border of the third epimeron is perhaps somewhat more prominently arched, forming

IIYDRACARINA

99

a more distinctly marked medial border than in the other sex. There is a well-developed
anterior supporting chitinous piece for the genital organ, and the genital lips are projecting and
provided with big papillae all over. The posterior acetabula are more elongated than in the
male and the posterior part of the genital plates seems to be bent out laterally.

Locality. Indian Tibet: c. 1 mile W. of Dras (K76), altitude c. 3081 m., 21 May,
1932 (temp. 19.0°C.).

The present species differs from all hitherto described Calony.v species, except C. latus,
by the claws being split up into a very large numljer of teeth.

5. Calonyx flagellum sp. n.

9. A rather large species, measuring about 1550-1640^1 in length. The body is broad,
attaining a breadth of about 1320j«. The frontal organ is bigger than that of the foregoing
species and has the shape of a distinctly marked circle. Of the dorsoglandularia only the first
pair (=antenniformia) is situated anterior to the eyes, whilst the second pair lies on a line
connecting the posterior margins of the eyes. In this respect the species differs from the
above-described Calonyx species and the Frotziella species as well, whereas the concordance
with Calonyx montanus as to the situation of the third pair of dorsoglandularia is complete.
The glandularia are characteristic, supported as they are by a very conspicuous, subcutaneous
framework of chitin. This consists of three rings of different shape. In the drawing the
lowest circle is black, the middle one is dotted, and the highest one, which lies immediately
under the skin, is not marked in any special way. The latter embraces the fissure-shaped
opening of the gland.

The maxillary organ is differently shaped, compared with that of the previous species.
The rostrum especially is dissimiliar, being bent ventrally and abruptly truncated at apex,
without the ventral, projecting tip, so well marked in montanus. The upper dorsal margin
differs also in being quite straight, not undulated. The organ is 362/*. long. Seen from above
it is more alike in both species, though a little wider in the present one. The lengths of the
palp-segments are (in ii) :

I

II

III

IV

V

Extensor side

Flexor side

46
46

132
36

75
64

196
96

46
36

The length of the mandible from the base to the tip of the mandibular membrane is
362/x and that of the claw 153/*.

As in C. montanus the anterior corners of the first three pairs of epimera bear bristles
about equal in number to those described above. Most of those on the first pair are devel-
oped into strong, short spines. The third epimeron projects inwardly at the antero-medial
corner, thus dift'ering very much from the preceding species. The greatest lengths of the
anterior and posterior groups of epimera respectively are 332 and 357/*.

The bristles, mentioned in the foregoing species as bordering in particular the extensor
surface of the segments 3 and 4 in the legs are longer and more conspicuous in the present
one. The claws are shaped as in C. montanus, but the main claw is shorter and thinner, not

100

IIYDKACARINA

overlajjping the side-claws so nuuh.'' Tlic dorsal end of the last leg-segment is protruded as in
C. montanus.

The genital organ resenihlcs that of the latter species, the acetabula, however, being
more numerous, about 30 in nunil>er. and unich more elongated. Likewise the genital
plates form a narrow strij) on each side of the anterior acetabula. Tn some .specimens there

Figure /.— Calonyx flagellum sp. n. 2 • A, supporting skeleton for a dermal gland ; B, maxillary organ from
the left side ; C, mandible ; D, palp ; E, part of a leg ; I*", end segment of a leg seen from the right side ; G, end
segment of a leg seen from above (claws omitted) ; H, I, claws ; K, genital organ.

is a small, roimded, posterior plate, bearing some bristles, which is distinctly separated fnMU
the anterior strips. Becau.se of the bad state of preservation of the present species it is impos-
sible to decide whether the structure mentioned holds good for all specimens. In some the
spot seems to be connected, or nearly connected, with the strip, but more, well-preserved
material is needed to settle this question. There are no chitinous sup])orting pieces before or
behind the genital organ.

The male is not represented in the collection.

Locality. Indian Tibkt: Shimsha Karbu, between Dras and Kargil (K 79), on stones
in rapid stream, altitude 2819 m., 22 May, 1932.

' In the figures tlie ilaws arc seen from a somewhat different position in the two species and thus not exactly
comparable.

IIYDUACAKINA

101

Family HYDRYPHANTIDAE

Subfamily Thyasinae
6. Parathyas primitiva sp. n.

S . Length of body 896m, breadth 672(1. The chitinization of the skin is less developed
and more primitive than in P. thoracata, the sole hitherto known species of the genus. In
the latter species all dorsolateral ia and the last pair of the dorsocentralia are very large; in

Figure 8. — Parathyas primitiva sp. n. A, animal from above (^ ) ; B. nia.\illary organ and right palp seen
from the left side ( <i ) ; C, mandible {$) ; D, end segments of third leg ( 9 ) ; E, genital organ (S) ; F,
genital organ ( $ ) .

the present species, on the contrary, they are not larger than the dorsocentralia 1-4. The
frontal shield has a characteristic shape ; it is somewhat prolonged and truncated posteriorly.
The hole for the frontal organ is well marked and fairly large. As in many Tliyas species
the fourth and fifth pairs of the dorsocentralia arc widely separated from one another.

102

riYnRACAKINA

The maxillary organ measures 186/1 in lengtli. The rostrum is high, with a curved
dorsal margin. The length of the mandible from the base to the end of the membrane is
232/i, that of the claw 71^. The dorsal margin of the mandible is evenly curbed. The meas-
urements for the palp-segments (in /*) are as follows :

I

II

III

IV

V

Pvtensor side

54
36

86
36

52
52

146
95

46

PIfxor side

39

Between the second and third pairs of epiincru Uic .^kin projects laterally in the shape of
a triangular fold. The legs hardly display any specific characters. The spines at the distal
end of the segments are finely dentated.

Genital organ of normal shape. The medial margins of the genital plates carry about 8
short and, behind these, alxnit 8 long bristles. In front the plates are obliquely cut off, as is
best seen when they are closed. Anterior and posterior to the genital organ, at the same
distance, there is a chitinous, circular plate. The excretory opening is bordered at both ends
by a chitinous knob.

9. As usually, the female is larger, attaining a length of about 1170/^. Otherwise it
differs from the male only in the genital organ, the anterior chitinous piece of which lies close
to the genital lips, not at some distance in front of the organ. The eggs measure alx)ut 241/^
in diameter.

Nymph. Length about 690/^. In the skin-plates and certain other characters it resem-
bles the adult. The shields, however, are smaller, only the frontal shield being of correspond-
ing size. There is no fold betw-een the two epimeral groups. The genital organ consists of
four acetabula, the two on either side being separated by a medially directed chitinized flap,
broadly rounded at the tip, which carries some few hairs.

Locality. Indian Tibet: Dras (K77), in a stream (temp. + 21.0-24.3°C.), altitude
3091 m., 21 May, 1932; CI mile W. of Dras (K76), altitude 3080 m., 21 May, 1932
(temp. + 19.0°C.).

7. Kashmirothyas hutchinsoni gen. et sp. n.

Generic diagnosis. Colour red. Skin armoured with large shields. Right and left
shields of dorsocentralia 3 and 4 united respectively into two shields. Frontal shield large,
composed of frontale, prae- and postfrontalia, postocularia, dorsocentralia 1 and 2. Frontal
organ non-pigmented. Eyes stalked, projecting over the sides of the body, attached to a
chitinous plate, from which also the praeocular bristle arises. First pair of epimera differ-
entiated sexually, the anterior corner being blunt in the male, acutely protruded in the female,
in both sexes inwardly set with a row of long, pectinate hairs. First pair of legs with
sexual dift'erences. Genital organ with more than 3 pairs of acetabula.

Description of species. 9. Length of body 1017/^. Skin-papillae sharply pointed,
spine-like. Dorsal surface covered with large shields, the shape of which varies, as is usual
in the Thyasinac. The frontal shield is largest. It is tnmcated in front and tapers gradually
toward the posterior end, which is cither truncated or more or less rounded. The frontal

TIYDRACARTNA

103

organ is not pigmented; laterally and a little posterior to it we find the two postociilar
bristles. The frontal shield reaches backwardly to behind the fourth pair of dorsoglandularia,
thus indicating that the first and second pairs of the dorsocentralia have been absorbed. The
third pair of dorsocentralia is united into a single, medial shield, as is also the fourth pair,
whereas the shields of the last, fifth pair, are isolated and very large. Moreover, the dorso-
lateralia of the pairs 2-4 are large and easily visible, whilst the first pair is small and invisi-

FiGURE 9. — Kashmirothyas hutchinsoni gen. et sp. n. A, animal from above { 6 ) ; B, eyeplate ( $ ) ; C, maxillary
organ from above (_S) ; D, from the left .side ( $ ) ; E, mandible ( 3 ) ; F, palp (S).

ble from above, lying on the sides and even partly on the underside of the body.'' The first
and third pairs of dorsoglandularia consist of a bigger plate than the others. As in Lund-
bladia the bristles of pairs 1 and 3-5 are much thicker, those of 2 and 6-7 thinner. The
hair of the second pair is especially very long and delicate. The two antenniformia (= first
[lair of dorsoglandularia) are free, not united, but lying close up to each other. As in
Javathyas, Lundbladia, and Trichothyas the ocularia have coalesced with the praeocularia
to constitute a transverse plate. I have been unable to ascertain whether the praeocular
bristle is bifid, as in Lundbladia, or not. I should not be surpri.sed if it actually were, but the
preparation is not clear enough to settle this. The epimeroglandulare 1 is large and triangular,
as in the female of Lundbladia.

' In Figure 9 A the outline of this pair is stippled.

104

llYDRACAklNA

The shields on the wntnil surface resemble those in LundbJadia. There is a postgenital
shield of various shape, munded or elongated, a more or less longitudinal excretalc, two very
large, medially incised ventralia 2, two elongated ventralia 1 and snuie smaller, accessory,
lateral plates.

Figure 10. — Kashmirothyas hutchinsoni gen. ct sp. a. \, animal from beneath ( ^ ) ; B, epiniera and genital
field ( 9 ) ; C, genital organ {&) ; D, genital organ ( 9 ).

Seen from the side the ma.xillary organ is rather llat, the mstrum heing straight, very
little bent downwards. The organ is 307/^ long. The posterior margin of the maxillary plate
is straight or a little concave because of the slightly projecting posterior lateral angles. The
outer wall of the articular socket for the palp is angularly protruding laterally. The mandi-
bles are elongated and narrow and the claw is fairly straight. The measurements for the
palp-segments (in i^) are:

r

II

TIT

IV

V

Extensor side

Flexor side

4,^
54

80
30

56
61

146
111

50

48

II YUU A CARINA

105

The palp is lony and slender and the process of the penultimate segment thin, spine-like
and much shorter than the prolonged end segment.

Epimera of usual shape. The first pair is prolonged in front, forming a very conspicu-
ous, pointed projection. The epimera of the first pair are widely separated from each other,
their inner margins being parallel, so that the maxillary bay retains the same width through-
out. From the inner margins a single row of about 18 feathered bristles arises. The
posterior inner corner of the first epimeron is acutely protruded inwardly. The three first

Figure 11. — Kashmirothyas hutchinsoni gen. et sp. n. A, first leg of male; B, of female, from the inner side.

legs especially are armed with strong spines, arranged in whirls at the distal end of the
segments. Besides, the first leg has a long, backwardly curved, strong bristle on the ventral
side of the second .segment. There is also a thinner bristle on the ventral side, arising in
some distance behind the spine. Such a bristle stands also at the same place on the second leg.

The genital organ is characteristic. It reaches up to the inner end of the maxillary bay
and in structure it is reminiscent of that of Tricliothyas or Liiiulbladia. However, it differs
considerably from the organ of all known genera. The genital plates are long and narrow,
posteriorly connected with the^ posterior acetabula and accordingly probaI)ly but little mova-
ble. Their interior border carries about 10, the exterior 2 bristles. The first pair of acetabula
lies in fnmt of the organ, the second is situated much farther back, posterior to the middle
of the plates. Both these pairs of ace(;il)ula are much elongated and attached to low socles,
liehind the plates but coalesced with tlu-ni and resting upon their hindpart we find some other
acetabula, more rounded in shape. Their number varies from two to four on each side.
In front of the genital aperture is a chitinous transverse bolt.

S . The male differs in many respects from the other sex, the following characters
especially being worth mentioning. It is smaller, about 930ai long. The maxillary organ is

106 TiYi)K.\r.\r<iNA

somewhat narrower in the posterior part. The epimeral groups lie closer together, the first
epimeron is rounded apically and the bristles along the medial border have longer feathers.
Also at the base the first epimeron is broadly rounded, without the slightest trace of the hook-
like projection of the female. The maxillary bay decreases in width posteriorly, and the
epimera of the first pair are brought closer together than in the other sex, pressing the
maxillary organ out of position in a dorsal direction. The first epimeroglandulare is hardly
smaller than in the female (cf. Lundbladia).

The anterior pair of legs presents very striking differences. They are distinctly short-
ened, particularly in the basal segments and resemble very much the same leg in Lundbladia.
Whilst the female possesses a long and strong bristle on the ventral side of the second seg-
ment, the male has this bristle developed into a thick spine. On the dorsal margin of the
following segment the male has two curved, strong spines, just as in Lundbladia. On the
other hand, I am unable to find the angular swelling, described in Lundbladia, on the outer side
of the second segment of the first leg. Consequently the leg in question is not distorted as in
the genus just mentioned.

In the genital organ the sexual differences are very conspicuous. It fills completely the
space between the four groups of epimera, which allow the genital structures very little space.
The first pair of acetabula, for instance, overlaps the posterior ends of the first pair of
epimera. Obviously, through the pressure against the epimera, these acetabula have altered
their shape from elongated to more rounded, being even bluntly distended in front, as if they
had been pressed together. As in the other sex, anterior and median acetabula are attached
to socles. The second pair is not placed posterior to the middle of the genital plates as in the
female, but has been pressed forward until somewhat in front of the suture between the third
and fourth pairs of epimera. The posterior acetabula, however, have retained the same posi-
tion. The genital plates are narrower and have thinner bristles. Their lateral margins are
partly overlapping the borders of the epimera.

Locality. Kashmir: Stream W. of Sonamarg (K71), altitude 2590 m. (temp.
+ 7.0°C.), 19 May, 1932. Indian Tibet: Shimsha Karbu, between Dras and Kargil
(K78), altitude 2819 m., in a spring (temp. + 8 C, pH. 7.8), 22 May, 1932.

Systematic affinities. The above described, very interesting genus is nearest related to
Tricliotliyas and Lundbladia, with which it has many characters in common.

Such characters are :

(1) composition of frontal shield

(2) non-pigmented frontal organ

(3) right and left elements of third and fourth pairs of doscocentralia united to

form two medial shields

(4) eyes stalked and lying in a shield together with the praeocularia

(5) the same sexual differences in the first pair of epimera

(6) the same sexual differences in the maxillary organ

(7) the same sexual differences in the situation of the second pair of acetabula

(8) principally the same sexual differences in the first pair of legs

The most important difference is to be found in the genital organ, which comes nearest
to that of Lu}uJbladia. Especially the males of the two genera are very much alike, whilst the

INI)l;Ar'AIUNA

107

females differ particularly in the shape of the posterior part of the genital plates. Moreover,
both sexes differ in having a greater number of acetabula.

In some of the characters mentioned Kashmirothyas also resembles the genus Javathyas
and belongs undoubtedly to the same group of genera as Javathyas, Trichothyas, and Lund-
hladia. It has, therefore, to be looked upon as a ramification from the same tribe as these
genera, mainly differing in the multiplication of the acetabula (cf. Lundblad, 1933).

Family HYDRACHNIDAE

Subfamily Hydrachninae

8. Hydrachna {Diplohydrachna) conjccta Koen.

Hydrachna conjecta Koenike, 1895, pp. 145-46
Koenikei Sig Thor, 1898, pp. 7-8

Sig Thor, 1899, pp. 17-18
conjecta Piersig. 1897-1900, Plate L, fig. 177 a-f

Piersig, 1901, pp. 47-48

Koenike, 1904, pp. 29-33
" -f- Koenikei Koenike, 1908, p. 263
Hydrarachna " Koenike, 1909, p. 43

dissecta Viets, 1911, a, pp. 343-46

Halbert, 1911, pp. 12-13
Hydrachna koenikei Sig Thor, 1916, pp. 14-18, figs. 15-16
Hydrarachna conjecta Lundblad, 1920, pp. 169-70

Soar and Williamson, 1925, pp. 175-78
Hydrachna " Viets, 1928, p. 15

Lundblad, 1929, pp. 18-23

Figure 12. — Hydrachna i-.)/i,Vi7(i Koen. A, eyes and dor.-^al plates ( 9 ) ; B, left palp from the outer side ( 9 ) ;
C, stigma ( 9 ) ; D, epimera and genital organ {$) ; E, epimera and genital organ ( 9 ).

lOS

iivi)i;Ar.\uiNA

This species, whicli Koenike described from a nymph, and nl whicli \'iets later described
another nympli under the name of H. conjccta dissecta, is a very variable species, as already
shown by me in two papers (1920 and 1929), to which I may refer here. For the sake of
completeness I give some drawings here from the India Expedition material, including a
figure showing the shape of the stigma. In all details the specimens agree well with European
material of this well-known species.

Localities. Kashmir: Srinagar, Gagirbal, closed swamp (K 19), altitude c. 1580 m.,
9-11 April, 1932; Phashakuri (K34) altitude c. 1585 m., 7 May, 1932; Shadipur (K40),
altitude c. 1582 m., 13 April, 1932 (nymphs).

Distribution. Many European countries, Palestine, Kashmir.

Family HYGROBATIDAE

Subfamily Hygrobatinae

9. Megapus proximalis sp. n.

9 . Length of body 880/*. Skin without any distinct structure, except two poorly devel-
oped chitinous plates far back on the dorsal side. The glandularia are also chitinized, as
usual, and can be seen as small, circular spots.

The palps constitute the most striking character of the species. .'Xs always in this genus
the penultimate segment is supplied with a strong, lateral spine on the inner side. In the
new species, however, this spine is placed near the base instead of in or beyond the middle of
the segment. Characteristic of the species are also two very long bristles, attached to the
ventral side near the base. At the point from vvhere these bristles arise the segment is dis-
tinctly swollen and the whole segment is curved, turning the concave side ventrally. The last
segment is unusually short. The penultimate segment carries a number of fine hairs near the
dorsal side. The measurements for the palp-segments are (in /*) :

I

II

III*

IV

V

Extensor side

Flexor side

43
32

107
53

93
61

134
113

32
32

The mandible ( iiirluding the claw) is 286/* long. The rostrum of thi' maxillary organ
is short.

The epimera are not very characteristic. The posterior group projects triangularly
toward the middle of the ventral l>ody surface. The first leg' again shows remarkable features,
the last segment being unusually short, measuring but 114/* in length. It is strongly curved.
The penultimate segment is supplied with the usual two spines of different sha])e and size and
the long, curved bristle.

* The measurements of the third segment are from its inner side.

HYDRACARINA

109

The genital opening is 243/n long and equipped at both ends with chitinized supporting-
bodies. The acetabula, three in number on each side, are arranged in a curve. The diameter
of the egg is \75i^.

Locality. Indian Tibet: Shimsha Karbu, between Dras and Kargil (K78), altitude
2819 m., in a spring (temp. + 8°C., pH. 7.8), 22 May, 1932.

FioUKE 13. — Megapus proximalis sp. n. 9 . A, animal from beneath ; B, ma.xillary organ from the left side ;
C, mandible; D, right palp from the inner side; E, first leg; F, end segments of first leg; G, genital organ.

Subfamily Acercinae

10. Accrcits ornatus C. L. Koch

Tipliys onuitus C. L. Koch, 1835, 5, fig. 20
Acercus " C. L. Koch, 1842, p. 24
Piona oniata Piersig, 1897-1900, pp. 143-48
Laiiiiniprs ornaliis Piersig, 1901, p. 202
Acercus " Koenike, 1909, p. 106

Soar and Williamson, 1929, pp. 16-19

no

HYDRACARINA

Unfortunately the male sex is not represented in the collection, so the determination is
not quite to be relied upon. The females, however, agree exactly with European females of
Acercxts ornatns.

Localitx. Kashmir: Phashakuri, S. of Pampur (K35), altitude 1585 m., 10 May,

1932.

Distribution. Most European countries, Algeria. Kamschatka, Kashmir.

COMl'AKATIVE NoTES ON THE DISTRIBUTION OF THE WaTERMITE FaUNA AT

Higher Elevation

The watcrmite fauna occurring at high altitudes is very little known, except in the Alps.
In spite of this it may be of some interest to compare here the faunae of certain elevated
districts.

If we begin with the Alps and consider only the species living over a height of 1800
meters w'e can group them in the following way. The distributional data are taken from
Walter (1922 b).

List of the IVatermites of the Alps {Above 1800 m.Y

1800-2000 m.

Protsia distincla
Partmiiiia angusta
Limnocliarcs holosericea
HydrypJumtes spinipes
Dartia borncri
Mega pus nod i pal pis

" loricatus
Lebertia cognata

" " sevocata

" lineata

" dubia

Lebertia aspera
" extendens
" euneifera
" giardinai
Gnaphiscus setosus
Pionacercus leuckarti
Feltria zschokkci
Aturits clinitiis
Arrhenurus conicus
" inaeulator

" neuniani

2000-2200 m.

Hydrovolzia placophora

Protzia alpina

Colony X rotund us

Sperchon mtitilus
" squaiiiosus
" longirostris

Panisus bazettae

Panisopsis curvifrons

Zschokkea oblonga

Megapiis -i'aginalis
Lebertia subtilis
" robust a
Feltria inenzeli

" setigera

" rubra
Pion<i cornea
Brachypoda versicolor

'Of course most of the species occur also beneath 1800 ni. For instance, the figures 1800-2000 mean that
the highest level at which the species licre enumerated are found lies s'lmewherc between 1800 and 2000 m.

HYDRACARINA

111

2200-2400 m.

Partnunia steinamanni
Spcrchon hrevirostris
" glandulosiis
" dcnticulatits
Panisus niicJiacU
Limnesia fulgida

Protzia invalvaris
Eylais hamata

" extend ens
Lehertia rufipes

2400-2600 m.

Hygrobates longipalpis
Rivobates norvegicus
Lebertia rufipes impcnnata
" pavesii
" maglioi
" zermattensis

Lebertia tuberosa

" zschokkei
Feltria minula
" nussbaumi

2600-2800 m.
Lebertia porosa

The above list shows us that the watermite fauna of the Alps is a rich one. Above 1800
meters there still live no less than 60 species and varieties.

In Norway Thor (1901) has studied the watermite fauna in the mountains. He was
unable to find watermites above 1200 meters. Thor mentions the following species:

List of the Watermites of Norway {Above 900 m.)

900-1000 m.
Teiitonia priniaria

Lebertia insignis

S perch on brcrirostris
lineatus
" mnltiplicatus
" squamosus

1200 m.

Piona coccinoides

Acerciis liitcscens

Hygrobates foreli
Mcgapus nodipalpis
Aturtis scaber
Feltria mimita

In Sweden the watermites reach no higher than in Nonvay. In the moimtains of North
Sweden I have collected a great many mites but most of them have not been determined as
yet. I have not found any mites there at higher elevation than 1112 meters. At present I
am al)le to list the following species only:

List of the JVatcrmites of Siveden {Above 900 m.)
900-1000 m.

Sperchon squamosus

Zschokkea oblonga
Piona coccinoides
Pionaccrcus Icuckarti
Hygrobates foreli

1000- About 1100 m.

Tcutonia subalpina

Giuipliiscus setosus
Neobrachypoda ekmani
Arrhcmirus subarcticus

112

IIYDRACARINA

La'Stly we shall list the watermites of Mount Elgon in equatorial Africa. Mt. Elgon is
the only tropical mountain of which the waterinite fauna has been studied. The following-
species have In^en found (Lundblad, 1927a) :

List of the ll'alcrniitcs of Mt. Elgon {Above 1800 m.)

1800-1900 m.

Spcrchon fcnestratus

1900-2400 m.

Hygrobates loveni
Hygrobatnf^sis !ei ipalpis
Hxgrohatuiiicgapiis spatliulifertis
Megapus linearis
" tigandcnsis

Hydrachna cldorctica
Hygrobates elgoncnsis

Hygrobates laceratus

Spcrchon elgoncnsis

2400-3200 in.

3200-3300 m.

3300-4200 m.

Megapus splendidus superbus
Octoniegapus viinutissiinus
A trad ides juciindiis
bryki

Megapus splendidus
Atractidcs leniniiis

Megapus affinis
Fiona angiilala

Unfortunately we can by no means say that the watcrniitc fauna is well knuwu at higher
altitudes either in Sweden, Norway, or on Mt. Elgon. Probably, however, there are but few-
species to lie added to the Scandinavian fauna. Our highest mountains do not rise much
above 2000 meters, and they are always isolated summits. Our highest plateaus arc
hardly higher than 1000 m. Their area is very extended in Sweden, but is much split uj)
into smaller, isolated districts, the area of each measuring at the utmost some 10 square
kilometers. When following a certain level — for instance, that of 700 m. — from South to
North, we would find, however, that the number of species to be enumerated as living above
or at this level would diminish very rapidly. This indicates that the mites are not restricted
to a special elevation but that the temperature of the waters — or other circumstances, for
instance, the presence of food or hosts — determines their distribution. It is, therefore, of
little interest to state the vertical distribution of a species unless we at the same time report
the latitude at which the species was found.

Consequently we find quite a numlier of watermites on Mt. Elgon even above 2000
meters, viz., 16 species, whilst there are no species in Sweden above 1100 meters. Undoubtedly
the fauna of Mt. Elgon, being at present superficially known, contains at least the same
number of undiscovered species as those already described.

HYDRACARINA 113

The species of Kasliniir may Ijc arranged in the following way:

List of the IVatcnnitcs of Kashmir (Above 1000 m.)
1500-1600 m.

Eylais degenerata Accniis ornatits

Hydrachna conjccta

1600-2800 m.

Calonyx flagcllum Megapus pro.viiiialis

Kaslniiirofhyas hiitchinsoni

2800-3000 m.
Calonyx montanus ParatJiyas priitiith'a

3000-4100 m.
Pro taicUa hutch iiisoni

4100-4300 m.
Eylais Iiauiata

It is difficult to decide at what height the corresponding zoogeographical limits run in
North Sweden, in the Alps and on Mt. Elgon, so far as the watermite fauna is concerned.
A personal knowledge of all three districts would be necessary in order to settle this. How-
ever, I think we can use preliminarily the uppermost limit of the forest as an indicator of the
climate." On Mt. Rlgon this limit runs at an altitude of ahout 3400 meters, in North
Sweden at 400-900 meters, sinking cnnsiderahly from South tn N<irth. \\\ the Aljis the
forest-limit is situated at about 1900 meters (1800 m. along the northern and 2000 m. at
the southern border). In the part of the Kashmir \'alley studied by the Yale Expedition
the forest-limit, according to information in writing from Dr. Hutchinson, is situated between
3000 and 3500 meters. Within this depression, i.e., southwest of the main Himalayan range,
in the Srinagar region, at a height of about 1500-1700 meters, the winter is not severe at
all, but most localities there are likely frozen in parts of January and February, whereas the
summer temperature must be high. The species found in stagnant waters there are Eylais
degenerata, Hydrachjui conjccta, and .Icrrcus oruaftis. The water temperature varied between
13.0-23.2°C.

The expedition crossed the, range at the Zoji-La pass at 3528 meters, where many kilo-
meters of snow were found along the road in the middle of May. On the southwestern
slopes of the range before the expedition reached the pass only one species, Kashniirothyas
liutchinsoni, was collected, in streams at Sonamarg at 2590 meters. The climate here is
undoubtedlv more rigorous, with lower temperatures, than in the Kashmir basin. Once over

" It is clear, however, that this limit is a very rough one, owing to the fact that distribution of water animals
depends upon the temperature of the water, which does not always correspond to that of the air nor is it a simple
exponent of the height above the sea level. At the same level different bodies of water often present ciuite difTerent
temperatures.

114- IIYDRACAKINA

the Da5s the forest disappears entirely even below 3000 m., the whole of Indian Tibet being
arid or semi-arid. The winters must be very severe, though with little snow, but certain
springs remain open all through the winter. This may be true of the spring at Shimsha
Karbu at 2S19 meters, where Prut del la hutchinsoni, Kashniirotliyas liutchinsoni and Megapus
proximalis live at a temperature of 8.0"C. However, some of the small streams in this dis-
trict are heated up relatively high by day, to over 20^0., and cool off to around 8.0X. at
night, owing to the rarity of the atmosphere. In such a stream Parathyas primitiva was
collected at Dras at about 3100 meters in a temperature of 19.0-24.3C., and Calonyx
tnontainis in 19.0" C. At Bao, at an altitude <>i 4100 meters, ProtzicUa hutchinsoni was
found in 7.2^ C. in a spring. For Calonyx flagclhun, found only in a rapid stream at
Shimsha Karbu at an altitude of 2819 meters, there are no temperature records. It may be
mentioned also that Kashniirotliyas hutchinsoni, l)esides in die above reported spring, was col-
lected in a stream at Sonamarg, altitude 2590 meters, in 7.0 C. Lastly Eylais haiiiata was
taken in ponds at Chushol, near the Tibet frontier, at an altitude of about 4340 meters.
Unfortunately there are no precise temperature data availal)le in this case, but the ponds in
question very likely freeze comj)letely solid during the winter. It is rather strange to find
that this mite, which lives even at the level of the sea, is able to thri\c at such ;i height. But
we have noticed already that Eylais haiiiata is one of those species which forces its way far-
thest up the Alps, where it lives above the forest limit. The ])onds at Chushol are the highest
records for mites in this region, and in the world, and though there were a number of ponds
at still much higher altitudes, no mites were found therein. It is a matter of interest that
the species from the highest locality is a widely distributed one and not an exclusively mountain
form. Thus only two species, Eylais haiiiata and ProtcicUa hutchinsoni, extend their range
above the forest limit as defined on the eastern slopes of the Kashmir \'alley, while two more
species, Calonyx iiinntanus and Parathyas primitiva, ju.st reach this limit.'" It is likely that
still mnre species do so though they have hitherto escaped discovery.

The watermite fauna of the Western Himalayas appears, in the present state of our
knowledge, to l)e much less rich than that of other districts enjoying a somewhat similar
climate, either in the far North or at high elevations in temperate or tropical countries. It is
difificult to decide with absolute certainty whether this difference is due to the \\'estern I lima-
layan fauna l)eing less well known, or to actual poverty of species. One would expect not
only a greater assemlilage of endemic cold-water forms, but also a greater number of lowland
species in the Kashmir Valley.

It is not advisable to give any detailed opinion as to the composition and immigration
of the watermite fauna of Kashmir, owing to the poverty of the data at present available,
founded as they are on but a few months' collecting. But even if we postulate the occur-
rence of more species, the fauna seems to be very poor. The reason fcir this is jjrobably that
mountain barriers have prevented the mites fr.)m .s])reading. The most characteristic high-
land forms, such as Protziclla and Kashinirolhyas, are probably endemic and of ancient,
perhaps preglacial, origin, while the other element in the fauna comprises such easily dis-
tributed species as Eylais haiiuita and E. degencrata; the immigration of this latter element is
presumably .still occurring, though more slowlv than in m^st other parts of the world.

'"But in ttie Dras basin where they occur the country is too arid to support forest, though at this altitude on
the opposite side of the Zoji-La about 35 miles to the west there are numerous trees.— C E. II.

lIVnRACARINA 115

If we compare the Scandinavian mountain watermite fauna, living above the forest hmit,
with that of the Alps, we see that the Alpine fauna is much richer, consisting of 60 species,
as against 18 in Scandinavia. This, moreover, is true not only of the watermites but of other
groups, such as the insects. One of the reasons for this richness is that the Alps are sur-
rounded by a much more abundant fauna and have received immigrants from many different
directions. Tropical mountains also will probably be found to have a rich fauna, though
experience on Mt. Elgon hardly supports this contention, for but two species were found there
above the forest belt. It must, however, be remembered that the fauna at the top of this
mountain has not been at all thoroughly studied.

Considering the favorable situation of Kashmir, in the middle of a large continent,
bounded to the North by the enormous palaearctic area and to the South by the tropical
Indian region with its luxuriant fauna, one should expect to meet a rather rich assemblage of
watermites there, comparable to that of the Alps. This is not the case, however, and conse-
quently there must be some special causes preventing the development of such a fauna. Proba-
bly the main obstacle is to be found in the high Himalayan ranges, which most of the mites
are unable to force and by which the Kashmir upland is isolated from the surrounding
districts.

116 IIYDRACARINA

REFERENCES

Uaday, I-^ 1901. A magyorszagi Eylais-fajok. — Math, es Termeszcltucloni. crtes. XIX.

Biulapest.
1903. Die F.ylaisarten Ungarns. — IMatli. uiul naturw. Ber. aus Ungarn. X\'III.

Leipzig.

1910. Untersuchungen iilier die Siisswasser-Mikro fauna Deutsch-Ost-Afrikas. —

Zoologica. Heft. LIX. Stuttgart.

Halbert, J. N. 1903. Notes on Irish Species of Eylais. — Annals aiul Mag. Nat. Hist.
Ser. 7, Xil. London.

1911. llydracarina. Clare Lsland Survey. 39.— Proc. Royal Irish Acad. XXXI.

Dublin.

KoENiKE, F. 1895. Liste des Hydrachnides recueillies ])ar le Docteur Th. Barrels en Pales-
tine, en Syrie et en figypte avec la description de quelques especes nouvelles. —
Rev. Biol, du Nord de la France. \'II. Lille.

1897. Zur Systematik der Gattung Eylais Latr. — Abh. Naturw. \'er. Breni. XIV.

Bremen.

1898. Hydrachniden-Fauna von Madagaskar und Nossi-Be. — Abh. Senckenb. naturf.

Ges. XXI. Frankfurt a. M.

1904. Hydrachniden aus dor nordwestdcutschen Fauna. — Abh. Naturw. \'er. Brem.

X\'III.' Bremen.

1908. Beitrag zur Kenntnis der llydraclini(k'n. Iliid. XIX.

1909. Acarina. — iJraucr: Siisswasser fauna ncutschlands. XII. Una.

-1910. Fin Acarinen-insl)esondere Hydracarinen-Systeni nel)st hydracarino-logischen

Berichtigungen. — Abh. Naturw. \'er. lircm. XX. liremen.

1919. Fine Wassermilbe als Cast bei cineni Wasserkafer. — Ibid. XXIV.

Koch, C. L. 1835. Deutschlands Crustaceen, Myriopoden und Arachniden. Regensburg.

1942. Ubersicht des Arachnidensystems. III. Niirnberg.

Ll'Ndbl.M), O. 1912. Nagra bidrag till kannedomen om vara hydracarincr och dcras utbred-
ning inom Upland. — Entomolog. tidskr. XXXIII. Stockholm.

1912a. Hydracarinologiska notiser. — ibid.

1920. Siissvvasseracarinen aus Diinemark. — Mem. de I'.Vcad. Royale Sci. Lettr.

Danemark. Sect. Sci. 8 me serie. VI. Copenhague.

1927. Die Hydracarinen Schwedens. I. — Zoolog. bidr. fr. Upsala. XI. Upsala.

1927 a. Zur Kenntnis der Hydracarinen fauna des Mount Elgongebiets im britischcn

Ostafrika. — Arch. Hydrobiol. XVIII. Stuttgart.

1929. Die Hydracarinen des Sees Takern. — Sji'ni Takcrns fauna och Hora. V.

Stockholm.

HYDRAfARINA 117

1933. Zur Kenntnis von Lundbladia petrophila (Michael) und der verschiedenen

Entwicklungsrichtungen bei den Thyasinen. — Zoolog. bidr. fr. Upsala. XIV.
Upsala.

Maglio, C. 1909. Idracarini del Trentino (Contributo alia conoscenza dell' idracnofauna
alpina.)— Atti della Soc. Ital. di Sc. Nat. XLVIII. Pavia.

Marshall, R. 1928. Watermites from China. — Trans. Wisconsin Ac. Sci., Arts and
Lett. XXIII.

MoTAS, C. 1929. Sur le developpement posteinbryonnaire de Calony.x brevipalpis (Maglio)
et sur les affinites des Protziidae. — Annales scientif. de I'univers. de Jassy. Jassy.

NoRDENSKiOLD, E. 1905. Hydrachniden aus dem Sudan. — Results Swed. Zool. Exp. to
Egypt and the White Nile, 1901. XX A. Upsala.

PiERSiG, R. 1897-1900. Deutschlands Hydrachniden. — Zoologica. Heft XXII. Stuttgart.

1901. Hydrachnidae. — Das Tierreich. Lief. XII. Berlin.

1906. Uber Siisswasser-Acarinen von Hinterindien, Sumatra, Java und tlen Sand-

wich-Inseln. — Zool. Jahrb. Abt. Syst. XXIII. Jena.

Soar, C. D. 1901. British Freshwater mites. Genus Eylais. — Science-Gossip. New ser.
VIII, No. 87, p. 68. London.

and Williamson, W. 1920. Hydracarina : The Genus I'-ylais Latr. — Journ. Ouekett

Micr. Club. Ser. 2. XIV. London.

1925. The British Hydracarina. I. — The Ray Society. London.

1929. The British Hydracarina. III.— Ibid.

Szalay, L. 1912. Kis-Azsiai Hydracarin.nk. — Allattani Kozlemenyek. XI. Budapest.

1926. Hydracarinr'ik a Balaton knmyekerol. — Ibid. XXII.

Thon, K. 1905. Hydrachniden. — Analen k. k. Naturh. Hofmus. XX. Wien.

1906. Monographic der Hydrachniden Bohmens. I. Teil. — Arch, naturw. Landes-

durchf. Bohmen. XII. Prag.

Thor, S. 1898. Nye Hydrachnide former fundne i Norge sonimeren 1898. — Arch. math.
og naturvid. XX. Kristiania.

1899. Tredie bidrag til kundskaben cm Norges hydrachniden — Ibid. XXI.

1901. Fjerde bidrag til kundskal^en cm Norges hydrachnider. — Ibid. XXIII.

1902. South African Hydrachnids. — Annals of the South Afr. Mus. II. London.

1916. Sur le genre Hydrachna Miill. et sur des nouvelles especes provenant princi-

palement de la Russie. — Rev. Russe d'Entomol. X\T. St. Petersburg.

ViETS, K. 1911. Neue afrikanische Hydracarinen. — Zool. Anzeig. XXX\^II. Leipzig.

1911a. Plydracarinologische Beitrage. IV-V. — Abh. naturw. Ver. Brem. XX.

Bremen.

1914. Hydracarinen aus Siidafrika. — Deutsche Siid])(>lar-Kxp. I'101-lf103. X\T.

Berlin.

118 llVnUACAKINA

1914a. Die Fortschrittc in <1ct KciiiUiiis dcr 1 lydiacaiincn. (l'H)l-1912.) — Arcli.

Hydrobiol. u. Planktonk. IX. Stuttgart.

1918. llyilracarinologische Beitnige. IX-X.— Abh. Natiirw. Vcr. I'.reni. XXIX.

Bremen.
1921. Hydracarina.— Wiss. Ergehn. Dcutsch. Zentral-Afrika-l-lxj). 1907-1908. V.

Leipzig.

1921. Neiiere englische Hydracarinen-Litcratur. — Arcli. Ilydmli. XI II. Stuttgart.

1926. Fauna sumatrensis. Hydracarina. — Entomol. Mitteil. XV. Ikrlin-Dahlem.

1926a. Indische W'assermilhen. — Zool. Jahrl). Abt. Syst. 1. 11. k-na.

— 1928. WassermilbLMi, Hydracarina. — Die Ticrwelt Aiiltclcuropa.s. iU. Licb. 4.

Leipzig.

1929. Dritte Mitteilung iilx'r neue Ilydracarinen vim dcu Sunda-Inseln. — Zool. Anz.

LXXXIH. Leipzig.

1930. Zur Kenntnis dcr' Hydracarincti-l'"auna von .S]ianifn. — Arcli. llydrob. XXI.

Stuttgart.

W.M.TER, C. 1919. Ilydracarinen aus den peruani.^chen .\nden mid ans I'.ra.silien. — Rev.
Suisse de Zool. XXVIL Geneve.

1922. Hydracarinen aus den vMpen. — -Ibid. XXIX.

1922 a. Zoologische Resultate der Reise von Dr. P. A. Chappuis an den oberen Nil.

Hydracarina.- — Ibid. XXX.

1922 b. Die Hydracarinen der Alpengewiisser. — Denksclir. Scbweizer. Xaturf. Ges.

LVIII. Basel.

ARTICLE VIII

REPORT ON TERRESTRIAL FAMILIES OF HEMIPTERA-HETEROPTERA

By G. Evelyn Hutchinson
Biologist, Yale North India Expedition

The present paper is based on the collectiim of terrestrial Heteroptera made during the
course of the Yale North India Expedition in Indian Tibet and the borders of Tibet proper
in 1932. My very best thanks are due to Dr. Hellmut de Terra for the opportunity to make col-
lections and observations in the little known territory traversed l>y the expedition and for his
continued interest in the progress of the work after the return uf the expedition. In a later
paper I hope to discuss in detail the ecology and zoogeography of the various elements which
compose the fauna of the highest inhabited zones of the Himalaya and Karakorum. I believe
that it will be possible to correlate many of Dr. de Terra's geological findings with the results
of such zoogeographic studies. Meanwhile a short zoogeographical account of the fauna of
the highest localities is appended to the present contribution.

The taxonomic work here reported was begun at the British Museum in January, 1934.
While working in London I received invaluable help from Mr. W. E. China, who is in charge
of the unrivalled collections of Hemiptera at South Kensington. Mr. China spared himself
no trouble in assisting me, and any merit that the present paper may possess is largely due to
him. My thanks are also due to my friend Prof. A. Petrunkevitch for help with the Russian
Hterature, and to Dr. E. D. Merrill and the staff of the New York Botanical Garden for deter-
mining specimens of food-plants.

The only previous work dealing with the Heteroptera of the region under discussion is
Distant's report (1879) on the collections made by Stoliczka during the Second Yarkand Mis-
sion. Most of the Heteroptera in these collections were obtained at Murree and in the vicinity
of Yarkand, but among terrestrial species Lamprodema hrevicolle Fieb. is recorded from
between Tangtse and Chagra (altitude c. 4, 200 m.) in Indian Tibet. The specimen was
determined by Edward Saunders and is presumably correctly named. The species is not repre-
sented in the present collection.

In Oshanin's catalogue (1912) several Heteroptera are recorded from Ladak, apparently
on the authority of Horvath (1889), who enumerated a number of species collected by Pauli
"in itinere suo e provincia Ladak in provinciam Pendshab." Since this collection contained a
numl:)er of large brightly colored forms, some of which are known from other i)arts of the
western Himalayas at comparatively low altitudes, it .'^eems reasonable from the available data
to sujipiisc that the collection was made either in Kulu or in the Kashmir depression. As
the present material consists exclusively of specimens from consi(leral)le altitudes, and contains
no species present in Horvath's collection, the latter is not further discussed.

The material collected by the Yale North India Expedition comprises 76 specimens, repre-
senting thirteen species, of which one, a species of Stictoplcura, is represented only by a
female and a nymph; in the absence of a male it seems unwise to attempt a specific determi-

Mem. Conn. Acad., Vol. X, Art. VIII. September, 1934.

120 TF.RRF.STiu \i I'AMiMES OF I ii-M 11' ^Kl^\- 1 1 irri:R(ii"n:R.\

nation. "^Of the remaining twelve species, four appear to lie already known, while twn new
genera, eight new species and one new subspecies are here described for the first time. It lias
also been necessary in the course of the work to re-examine some of the criteria used in tlie sep-
aration of tlie sul)families and tril>es of the Lygacidac, and to study rather closely certain
members of the genus Xysiits allied tn A', cricac (Schiil. ). The results of these studies are
set out in the appropriate places below. .\.ll species in any way associated with a(|uatic
localities will Ix; described in a later paper.

In general Oshanin's catalogue (I'Jlii has been followed as to nomenclature and classi-
fication. Bibliographic references are given for all si)eciiic names not included in that work,
but otherwise only to papers to which actual reference is made in the te.xt. A representative
set of all species, including the types of those here descriljed, has been incorporated in the
collections of the Peabody Museiun, Yale University: a first set of duplicates has been sent
to Mr. China for the collections of the llritish .Museum.

b'aniily I'ENT.XTOM IDAE

Subfamily ScHTKi.i.KKlN.\K

Tribe Odontotarsaria

1. Phimodera rupshuensis sp. n.

\\'idel\- oval ( I'late \ III, fig. 1 I, holotype 1.44 times as long as wide; moderately conve.x,
opaque, covered with short, sparse pale jnibescence; head, save for the raised parts of the
jugae and clypeus and the regions at the bases of the eyes, pronotum, scutellum and exposed
portions of the elytra, punctured, the distance between the punctures very irregular, averag-
ing rather more than their diameter, under surface punctured but more sparsely so.

Color. I'ale greyish yellow, puncturation black, head black, a narrow area round the
eyes, jugae particularly in their raised part, and base of the clypeus largely yellow, a large spot
on the anterior margin of the pronotum, and the base of the scutellum black; basal joints of
antennae brown, the first and second narrowly yellow apically, terminal two joints black ;
anterior femora with a black ventral stripe and a less w-ell defined posterior stripe fusing with
the black puncturation dorsally, intermediate and posterior femora with black postero-ventral
stripes, tibiae black a])ically and basally, on the dorsal surface the dark markings forming a
.stripe interrui)ted in its middle third, first and di.stal half of third tarsal joints dark brown.

Head. In front view (Plate \TII, fig. 2), about one-sixth longer than intemcular
width, subparallel and hardly constricted in front of the eyes, juga with outer angle widely
rounded, anterior margin lightly curved from the external to the .sub-obtuse inner angle, inner
part of the juga in its basal half .somewhat raised and encroaching on the clypeus; clypeus
anteriorly subacute and jirojecting Ijeyond juga, carinate in its anterior part, carina depressed
posteriorly, Ijecoming obsolete opposite the elevated part of the juga, vertex behind clypeus
somewhat elevated. Ocelli .separated from eye by a space sulKjqual to the maximum diameter
of the eye and rather greater than its width. F)ucculae subprominent below clypeus, obtusely
rounded behind (Plate VIII, fig. 3).

TKRRF.STRIAI, FAMILIES OF HEM I I'TF.K A- H F.TKKOl'TF.R A IJl

y\ntennae with first and second jdints sub.-tiual, half as loiiy again as third, tourtli twice
as long as the latter, fifth just over half as long again as fourth (0.27, 0.27, 0.18, 0.36,
0.58 mm.).

Rostrum reaching to posterior coxae.

Pronotum just over twice as wide as long (3.35, 1.53 mm.), with anterior margin evenly
concave when seen from in front and practically straight when viewed from above, posterior
margin almost straight centrally, laterally bent forward to the rounded posterior angles, lat-
eral margins emarginate behind the subrectangular anterior angles, disc with a central
impunctate carina, which in its posterior third is fragmented to form two irregular tubercles,
and with a transverse depression, obsolete centrally, in front of which are two raised areas
divided by transverse V-shaped depressions (probably apodenie bases) and falling off abruptly
toward the lateral margins.

Scutellum with a longitudinal central carina, reaching to just beyond its centre. (The
specimen also shows two folds running obliquely from the anterior angles to behind the middle
of the disc, but these appear to be due to an injury that has also removed the right elytron
and so buckled the scutellum on that side as to make measurement impossible. )

Marginal abdominal tubercles but moderately prominent (Plate VIII, fig. 4).

Length 4.8 m. ; breadth ?>.?>S m.

Indian Tibet: 1 $ (type) I'eldo-le, near N. end of Tso Moriri, altitude 4529 m. (14,855
ft.), among roots of short grass.

This species, as is indicated by its subparallel head, subrectangular anterior pronotal
angles and the coloration of its antennae, clearly belongs in the fourth cohort of the key in
Reuter's monograjjh of the genus (1908). It differs from the species placed in that group in
its small size, unarmed trochanters, less conspicuous marginal tubercles, and apparently in the
somewhat elevated center of the vertex behind the clypeus.

P. rcnlcri Kiritshenko (1910), the only species described since Reuter's monograph.
belongs to first cohort. The present species appears to be the smallest member of the genus.

Family COREIDAE

Subfamily Corizinae

2. Stictoplciira sp.

Indian Tibet : 1 9 antl 1 nymph, between Tsak-shang and Tsak-ra, road from Tso
Moriri to Tso Kar, altitude 4570 m. ( c. 15,000 ft.), 1 Sept., 1932.

The single adult before me is a female in not very good condition. It is most closely
allied to n\sioidcs Kiritshenko, but since the genus contains several very similar species and
since these probably cannot be satisfactorily determined without a study of the S genitalia the
present specimen is l)est left unnamed.

\22 TKKRKSTKTAI. IWMII.IF.S (11" HFAJ I I'T1:r \-H KTKROl'TERA

Family LYGAEIDAE

Subfamily Lygaeinae

Tribe Orsillaria

3. A^ysiiis ericac (Scbill.)

The sixteen specimens of Xy-^'us in the collection present so much diversity that at first it
seemed as though several rather distinct species were represented. Before attempting to
elucidate the present collection it ap])cared advisable to examine rather minutely certain of the
described Palaearctic species. In particular, since Evans (1929) had shown thai in discrimi-
nating between certain Australian species, the parameres of the male provide valuai)le ciiar-
acters, special attention was paid to these structures. As a result of these studies it l^ecame
clear that all the Yale North India Expedition material was referable to A'', ericac (Schill.).
tliough it has seemed desira1)le to descrilje as a subspecies a rather distinct form fmm very
high altitudes.

The Palaearctic species of Nysins have been studied by Horvath (1890) whose valuable
key provides a satisfactory basis for further work. In this key a group of species of the
restricted sub-genus Nysius (now to be regarded as a genus, cf. Evans, 1929) are character-
ised by having no well-marked pale longitudinal ruga on the scutellum and by the bucculae
being distinctly lowered posteriorly and not quite reaching the posterior margin of the ventral
surface of the head. This group includes thymi (Wolff), ericac (Schill.) and its var.
ohscuratus Yiorw, cymoiJcs Spin., i^raiiiiiticola (Klti.), and groenlandicus (Zell.), the latter
form, which Lindroth (1931) regards as a synonym of ohscuralus Horv., being excluded by
Horvath on geographical grounds. As pointed out below, groenlandicus, which is found
in the N. of Europe and Iceland as well as in. the Nearctic region, though undoubtedly a
subspecies of ericae, differs in several characters from obscuratus. This group, which may
be known as the thymi-group, appears to include most of the species described from the
tropical regions of the world, but with the exception of the .Vustralasian and African species
described by Evans very few of these species can l:»e recognised from descriptions alone. It
seems therefore desirable to put on record the following notes, which, though they relate
only to three of the most closely allied Palaearctic forms, may hel]) to stabilise our con-
ception of this difificult group of species and provide a point of reference for workers studying
tropical and sub-tropical species.

a. A'^. tliymi (Wolff). This species is distinguished externally by its oblong-ovate
shape, the posterior corial margin being rounded and ampliate (Plate \'I11, fig. 7). The
genital segment of the male is black and the longitudinal veins of the corium are brown or
Ijlackish. According to Horvath the vertex is destitute of a pale immaculate basal spot, but
this is actually often very feebly developed. Horvath also states that the ante-apical black
line on the pronotum is nblii|ue, curved forward and interni[)te(l centrally. This refers to
a pair of marks, of essentially the same f(jrm in all the .species, presumably th« bases of
thoracic apodemes, which are black and surrounded by a dark suffusion. In ericac, how-
ever, this suffusion generally forms a straight uninterrupted transverse band so that the
forward curve of the apodeme bases is less easily distinguishable.

TERRESTRIAL FAMILIES nv IIEMirTERA-IIETEROPTERA 123

The genitalia of two specimens from I'ritain were examined, one from Polzeath, Corn-
wall, the other from Kidw'elly, Carmarthen. The parameres in lateral view (Plate VIII,
fig. 14) are distinctly angulate dorsally, the angnlation not being emarginate, and the ven-
tral margin distinctly flanged. In dorsal view (Plate VIII, fig. 15) the angulate prominence
hardly projects over the inner margin of the base of the shaft.

b. A'^. ericac cricac (Schill.). The typical subspecies of cricac is much narrower than
thymi, the corial margins less ampliate, though very slightly curved from the widest point
towards the membrane (Plate VIII, fig. 8). The genital segment is black and the corial
nerves infuscated, but the basal immaculate spot on the vertex is much more strongly devel-
oped and the antiapical apodeme bases of the pronotum are normally included in a straight
unbroken transverse band. Material from North America (North Haven, Conn.) appears to
differ in no respect from a S from Marburg, Germany, determined by Horvath and in the
Britisii Museum collection.

The genitalia were studied in two specimens from North Haven. The dorsal angle of
the parameres is very prominent, setose, and distinctly emarginate, the ventral flange is
obsolete (Plate VIII, fig. 16). In dorsal view the angular prominence projects over the
inner margin of the base of the shaft (Plate VIII, fig. 17).

c. A'^. e. obscuratus Horvath. I have been unable to examine an authenticated specimen
of this form. Horvath's (1899) description is as- follows: Articulo primo antennarum,
saepe etiam basi articuli secundi, femoribusque nigris, femoribus feminae interdum pallidis,
nigro-maculatis; pronoto posterius fusco, angulis posticis maculaque parva media postica
pallidioribus; hemelytris griseo-fuscibus, interstitiis vernarum corii f usconebulosis ; ventre
feminae magnam partem nigro ; statura sexuum conformi. c5 . 2. Long, 4^-4^ mill.

Apart from its size the first male from Renka-le appears to agree with this form but its
smallness indicates a transition to cricac cn'cae. The genitalia are quite typical.

A^. e. obscuratus was originally recorded from Turkestan, Siberia and China ; in
Ekblom's map it is indicated as co-occurring with the typical subspecies throughout its entire
Central Asiatic range, but it is clear from Horvath (1904) and Kiritshenko (1931a) that it
is the only form found in the Tian-shan and in the Pamirs so that it may justifial)ly be
given subspecific status.

d. A^. c. groenlandicus (Zett). Lindroth (1931) synonymises this form with obscuratus.
In groenlandicus, however, the corial margin has a peculiar shape well marked in a series
of ? 9 in the British Museum collection and also in a 5 from Kugsuk, Godthaab Fjord,
West Greenland, collected by Major Hingston and kindly sent me by Professor G. D. H.
Carpenter of Oxford (Plate VIII, fig. 9). In N. e. ericac and in the Renka-le specimen,
discussed above under N. c. obscuratus, the corial margin is slightly and very gently rounded
from the straight basal part to the region of maximum dilatation, while in groenlandicus the
dilatation is more sudden so that in this region the corial margin appears almost obtusely
angulate. Moreover, in grociihnnlicus the ])ale purtion of the elytra is more transparent
than in the other forms so that when compared with obscuratus the color pattern of the
former shows much more contrast than that of the latter, likblom (1931) records the
Lapland form of cricac as obscuratus without description, and without indicating any Ice-
landic or Greenlandic records on his map. In the absence of specimens from this region it

124 TKRRKSTKlAi. K\MII.I1",S dl" 1 1 l-M I I'll'.U A- 1 1 F.TI:K(>PTI".R A

is not possible to settle the matter finally Init it seems more reasonable at present to refer
all these toreal forms to grocnlandicus. It is clear from l-^kblom's map that the latter sub-
species, as here understood, is separated from the other forms by a wide intervening
subboreal zone in which the species is absent.

The parameres of the West (ireenland specimen are iclentieal in shajie with those of the
North Haven specimens, though the angular prominence is a little more setose, a character
that varies in parameres of insects from the same locality in Indian Tibet. There can be
no doubt therefore that ^^rocnlaitdicus is rightly referred to this species.

e. A^. gniDiinicohi ( Klti. I . This species is easily distinguished by its coloration from the
preceding, for the longitudinal veins of the corium are hardly, if at all, infuscated and the
general coloration is paler. In shape gni)}iinicola is more elongate than thyiiii, but the corial
margins are posteriorly more strongly and more regularly rounded than in cricac. The pro-
notum is without a transverse black band obscuring the bases of the apodemes, which are
at most surrounded with an intcrru])ted black suffusion. The vertical margin sjjot is very
feebly developed.

The genitalia of a sjiccimcn from I'orto d' Ischia, on the island of Ischia. ltal_\-, were
studied. The dorsal angle is very feebly emarginate and '.be ventral keel moderately devel-
oped (Plate VIII, figs. 20, 21).

It is clear from the above that lliyiiii. cricac, and gniiiiinicola, three very closely allied
but adequately defined species, all show differences in their genitalia, while the various forms
here grouped under cricac show no such differences, thus justifying the present arrangement.
I have not been able to examine the genitalia of cyiiioidcs, a most distinct species with very
long subparallel elytra.

The material collected l)y the Yale North India E.xpeditioii was obtained from five
localities, as enumerated below. Measurements and notes on the individual specimens are
also set out in Table I. It will be seen that the material from the lowest locality is prac-
tically identical in form and color with typical A', c. cricac, while from the highest a rather
distinct new form w'as obtained which is described below as alticola subsp. n. From the
intermediate localities series were obtained which appear to combine the characters of all the
Central Asiatic forms known, viz., cricac s. str., obscuratu^ and alticola.

A. Leh. 1 $ Residency Garden. 19 Sept., 1932, altitude 3506 m. Parameres typical of
species. This specimen may lae considered as a very slightly atypical member of A^ c. cricac
(PlateVTII, fig. 10).

B. Tsak-shang, N. of Tso Moriri. 2 9 9. 31 Aug., 1932. altitude 4872 m. These
specimens are comparable to some of the 9 9 from the next locality; they are ])robablv
nearer to A', c. cricac than any other form.

C. Renka4e. Ijetween AIit])aI Tso and ^'aye Tso. 3 $ S, S 9 9. 18 .\ug.. l'>32,
altitude ^\Mi in. 'ihe specimens numbered 1 and 2 are very close to (ihsciiraliis. The third

i is practically typical ('. rr/cat', though very small. Specimen 2 (Plate \lll. lig. 11) is
slightly wider than the others, so approaching alticola. The females are rather variable in
width, but none .show the coloration of obscitratits.

D. Kyang-La, Koh Lungpa valley. 2 S $. 9 July, ].':>:^2. altitude .5100-5200 m.
These two specimens are both here referred to alticola; the darker one is rather similar to

TERRESTRIAL FAMILIES OF IIEMIPTERA-IIETEROPTERA

125

TABLE I
Dimensions of and remarks on specimens of A^ysiiis examined.

Locality

Sex

Length

Breadth

^Ratio

Departure from
Normal Coloration

Posterior
Pronotal Angles

A", ericae

Leh
alt. 3506 m.

15

3.45

1.11

3.12

Femora l)Iack with tes-
taceous apices

Very slif^htly
reduced

TSAK-SHANG

alt. 4872 m.

1?

29

3.75

4.25

1.38
1.56

2.71

2.72

Femoral spots confluent
Femoral spots confluent

Slightly reduced
Slightly reduced

Renka-i.e

alt. 5156 m.

\$

3.28

1.09

3.00

F'emora save apices, pro-
notum largely, and
inner part of elytra,
very dark

Almost normal

25

3.52

L27

2.86

As above

Almost normal

35

3.28

1.11

2.96

Femoral spots confluent

Slightly reduced

49

3.88

1.34

2.89

Femoral spots strongly
confluent

Slightly reduced

5 9

3.82

1.40

2.73

Practically typical

Slightly reduced

69

3.96

1.54

2.60

Femora black with pale
apices

Slightly reduced

79

4.00

1.42

2.82

Femoral spots some-
what confluent

Slightly reduced

89

3.96

1.49

2.66

As above

Slightly reduced

Kyang-La

alt. 5100-5200 m.

15

3.0' :i

1.13

2.74

Femora black save at

Considerably

sulisp. allirola

aj^ex, pronotum very
dark, elytra suffused
with brown

reduced

25

2.97

1.13

2.64

As above but lighter

As above

Ororotse Tso

alt. 5297 m.

15

3.20

1,24

2.58

Femora black save apic-

Much reduced

subsp. alticola

ally, elytra and pro-

notum suffused with
brown

2 9 3.38 1.42 2.38 As above but some pale As above

maculation on femora

3 9 3.65 1.45 2.50 .As above but rather paler As above

North FIaven,

Conn., U.S.A.
subsp. ericae

1 5 3.48

1.16

3.00

East Greenland 1 5
subsp. grocniandicus

4.15 1.42 2.92 Femora, inner part of

elytra and most of
pronotum black, outer
part of elytra hyaline

Unreduced

N. thyini

Kidwelly, Wales

15

4.07

1.49

2.72

A', graiuinicola

Ischia

15

4.62

1.49

3.10

126 TERRESTRIAL 1- AMIJ-IES OF IIE.MIPTERA-IIETEROPTERA

the wide. (lark specimen from the previous locality, but the latter is larger, prnpdrtionately a
little narrower, and has more prominent pron :)tal angles.

E. Ororotse Tso. 13,229. 11 July, 1932, altitude 52^^7 m. These specimens
(Plate \'lll, figs. 12, 13) are very broad and have the posterior pmnntal angles much reduced
so that the posterior border is but little reflexed and the sides are straight. In color they
are less dark than the obsctiratiis form from Renka-le. The present specimens constitute
the typical series of alticola subsp. n., primarily characterised by its small size, wide form
which is comparable to that of tliyiiii rather than to cricac s. str., and straight lateral pronotal

Nysitis ericac alticola subsp. n.

S Robust (Plate VIIT, fig. 12); dorsal surface covered wiih fine short adpressed hairs.

Color. Head black, mottled with testaceous yellow on the clypeus and juga, and w itli a
conspicuous smooth spot on the posterior margin yellow; antennae dark brown, ventral surface
of first joint and proximal half of first joint, .save the extreme base, yellowi.sh ; bucculae
grey. Pronotum yellowish-grey, with heavily black pnnctnration, save in the posterior angles
and a spot on the posterior margin; transverse black line on the anterior part of the pronotal
disc complete centrally and turned forward laterally; scutellum black. Dorsal surfaces of all
femora Ijlack, save at their a])ices which are testaceous; anterior femora black ventrally save
for the testaceous apices, intcriuediatc and posterior femora testaceous ventrally, lieaxily
spotted with black, tibiae testaceous with black spots apically, first tarsal joints testace(jus
darkening distally to brown, second joints brown, third joints black. Elytra opaque, yellowish-
grey, with the inner margin of the clavus obscurely darkened, lateral margins of curium wvy
narrowly black, indefinitely mottled with black along the outer corial vein (snb-costa) and
less consi)icuously on the disc, membrane hyaline with a large I)lack sj)ot on its corial border
fading to brown at the edges and just invading the posterior angle of the corium. Thorax
lx;neath black, posterior borders of pleurae and edges of articulations of legs yellowish-grey,
outer part of lip of scent-gland yellowish. Ab.lomen black, with yellowish mottling on the
edge of the connexivum.

Head about one-sixth narrower than the pronotum postericjrly; eyes relatively small,
vertex moderately flat in profile, bucculae not quite reaching the posterior margin of the ven-
tral surface of the head, slightly lowered in their posterior half and more al)ruptly termi-
nated opposite the apex of the first rostral joint. Eirst joint of antenna \n\\. little surpassing
the apex of the head; second joint twice as long as first and very slightly shorter than the
pronotum ; third joint three-fourths the length of the second : and fourth joint slightly longer
than the third (0.29, 0.58, 0.44, 0.51 mm.).

Pronotum trapeziform and moderately transverse, sides straight, posterior angles not
prominent, posterior margin but little deflexed, transverse black lines on anterior part of
disc incomplete centrally and turned forward laterally. Scutellum sul)-c(|ual in length to
pronotum and about as long as its basal breadth.

Apex of alxlomen not covered by elytra. Wings developed.

Genitalia as in the typical subspecies (Plate VIII, figs. 18, 19).

9 Somewhat broader than the male (Plate VIII. fig. 13 l. ^'ellow mottling of the clypeus

TERRESTRIAL FAMILIES OF HEMIPTERA-IIETEROPTERA 127

extending back throughout the central region of the vertex. Antennae entirely black. Elytra
just surpassing the apex of the abdomen. Otherwise as male in non-sexual characters.

Length $, 3.20 mm. (holotype) ; 2, 3.38 mm. (allotype), 3.65 mm. (paratype).

Indian Tibet. S (holotype), 2 5 9 (allotype and paratype) L 32, Ororntse Tso,
altitude 5297 m. (17,381 ft.), near margin of lake, among short sparse grass in company
with Chlaiuydatus pachycerus Kiritsh. 11 July, 1932; 2 & S L48, Kyang-La, altitude
5100-5200 m. (16,700-17,100 ft.), among short sparse grass with Pcgaeophyton prob.
scapifolhim Marq. and Skan., in company with C. pachycerus, 9 July, 1932.

In form this subspecies in its most extreme facies differs very widely from A^. e. ericae,
departing as much from the latter in its proportions as does A'', tliymi (cf. Table I). Were
it not for the existence of intermediate specimens and the identity of the genitalia throughout
the entire series it would have been regarded as a very distinct species. Though the feeble
development of the posterior pronotal angles suggests brachyptery, the wings appear to be
as well developed as in the North Haven specimens of the typical subspecies.

Key to the Subspecies of Nysius ericae (ScJiill.), applicable primarily to Male Specimens

1. About three times ( S 2.92-3.12) as long as wide, lateral margins of the pro-
notum sinuate, posterior angles subprominent 2

Less than two and three-quarters ( S 2.58-2.74) times as long as wide, lateral margins

of pronotum straight, posterior angles reduced N. e. alticola subsp. n.

Koh massif, Chang Chenmo Range, from over 5100 m.

2. Posterior part of promitum, femora and intcrvenal spaces of corium widely suffused
with l)lackish-l)rown 3

Posterior part of pronotum and intervenal spaces of corium testaceous, femora
testaceous with black spots A', e. ericae (Schill.)

Palaearctic from France to Sil^eria but absent in Britain,
Scandinavia, Northern Germany and Northern Russia ; Nearc-
tic throughout U. S. A. and Southern Canada ; locally wholly
or in part replaced by other subspecies.

3. Corium evenly rounded to its ma.ximum width, pale parts of elytra opaque

A'', e. obsciiratus Horv.

Central Asia from the Caspian to China, in part replacing
A'^. e. ericae.

Corium suddenly expanded to its maximum width, pale parts of elytra hyaline

A'', e. groenlandicus (Zett.)

Lapland, Greenland, Iceland and Arctic and Sub-arctic
America.

128 TF.RKKSTNIAI. FAMILIES OF IIEMIPTKRA-IIKTFROrTF.R A

^ Subfamily Oxycarenixae

4. Microplax hissarietisis Iviritshenko
.1/. hissaricnsis Kiritsheiiku ( 1^'13\

Indian Tibet 1 9 . Between Tsak-shani^' and Isak-ra, mad trum Tso Moriri li> Tsu Kar,
altitude c. 4570 m. (c. 15,000 ft.), 1 Septcniher, 1"32.

. The single specimen which is here identified witli .1/. Iiissaricnsis appears to aj^ree in all
essential points with the original description of this very distinct species. The only slight
differences concern the coloration of the elytra, which seems to be more intense in the speci-
men now before me, the dark marks on the corial nerves appearing to extend ontward
farther onto the disc of the corium than is indicated in the original descrijnion, while the
black base of the clavus fades to brown with black punctures apically. Since but a single
specimen is known it seems unwise to describe the present form as a subspecies.

}f. hissariensis is, as Kiritshenko points out, sharply distinguished from its congeners
!)y its larger size (4.0-4.2 mm. in typical series, 3.94 mm. in the jjresent specimen ), the entirely
black antennae, and the brown-black apical corial angle ( J'lale \T1I, fig. 5). The typical
series was taken in northern Buchara.

5. Bianchiclla udchnii^i Renter

Indian Tibet. 1 9 Igu, in the Indus V'alley above I.eh, on the bark on Populu^ sp.,
ahitude 3417 m. (11,210 ft.).

The single brachypterous specimen obtained was one of several observed, l)ut extremely
difficult to capture owing to the rapidity of their movements. It has been comp.-ircd with
material determined by Kiritshenko in Ihe llritish Muscmn and ajipears to be identical.
Since this remarkable form has not been figured, a drawing is given in I'late \T11. lig. (i.
The species is known from .Siberia, Mongolia and iVorlhern China ( Oshanin, 1"12) but
curioush' enough a])pe;u's to be unrecorded trom l\ussian Turkestan: a second sjjecies
( />'. saninilica Kiritshenko, 1"26) is, however, known from l'"uropcan Russia.

Subfamily Aphaninae

Tribe Gonionotaria

6. Emblethis horvathiana sp. n.

Ovate subparallel and rather robust, 2.25 times as long as wide.

Color, dark greyish-yellow heavily punctured with black, antennae and legs darker than
head pronotum scutellum and elytra, eyes brown, apical joint of antennae, ocular margin of
head, base of scutellum showing through the pronotum and some irregular spots joining
punctures on the disc of the pronotum and the scutellum. black ; thorax below black, margins
of coxal articulations greyish-yellow, abdomen beneath brownish, darkening to piceous along
the midline.

Head with eyes, seen from above, twice as wide as long (1.20, 0.58 mm. ), antenniferous
tubercles acutely rounded in lateral view, antennae 1.11 times as long as the maximum width

TERRESTRIAL FAMILIES OF HEMIPTERA-HETEROPTEKA 129

of the pronotum, basal jtjiiit sub-cylindrical, just over twice as long as wide (0.15 mm.),
second joint just over twice as long as the first, third joint just over two-thirds as long as
the second, fourth suljequal to the latter (0.33, 0.76, 0.55, 0.74 mm. ) ; basal three joints
richly setose, the setae being slightly shorter than the maximum diameter of the first joint,
apical joint with a few setae basally and with fine short hairs throughout; rostrum long,
reaching almost to the center of the posterior coxae, second joint very slightly longer than
the first, third subequal to second, fourth subecjual to first.

Pronotum trapeziform (Plate X, fig. 1 ), not greatly narrowed anteriorly, rather under
twice as wide as long (2.15, 1.16 mm.), sides moderately explanate, anterior margin slightly,
evenly and roundly excavate, lateral margins neither reflexed or marginated, slightly con-
verging anteriorly from just before the posterior angles, slightly emarginate behind middle,
with about eight setae on their anterior portion, including the anterior angles.

Scutelluni equal in length to the pronotum and basally slightly wider than its length
(1.34, 1.16 mm.). Mesosternum with well-developed and closely appro.ximated tubercles,
disc of metasternum not very conspicuously impressed.

Elytra nearly reaching the apex of the abdomen.

Posterior tibia a little shorter than the posterior width of the pronotum, and just over
twice as long as the basal tarsal joint, the latter two and a half times as long as the subequal
second and third joints together, claws two-fifths as long as one of the latter (2.00, 0.91,
0.18, 0.1 S, 0.07 mm.).

Length 5 {type) 5.45, breadth 2.43 mm.

Indian Tibet. 2 9? {^yps and paratype). L 77a. Renka-le, altitude 5136 m.
(16,917 ft.), between Mitpal Tso and Yaye Tso, on grassy bank in valley. 18 Aug., 1932.

In the paratype the anterif)r margin of the pronotum is practically straight centrally and
the elytra reach to the apex of the abdomen, the two specimens otherwise agree and are
undoubtedly conspecific.

E. Iion'athiana is perhaps more closely allied to E. vcrbasci, than tt> any other species of
the genus known to me. It differs conspicuously in having much more setose antennae, the
fourth joints of which are subeejual to the second, rather less explanate lateral pronotal mar-
gins, in being narrower and in its dark greyish coloration. At first I believed my material
was to be referred to brezicornis Horv., but, on seeing a drawing of one of the present
specimens, Dr. Horvath pointed out to me that in my species the form of the pronotum
and antennae are very different. In brevicornis the lateral margins of the former are quite
straight and converge more markedly anteriorly, while the fourth joint of the antenna is
very much shorter than in the present species. I am greatly indebted to Dr. Horvath for
calling my attention to these points and have much pleasure in associating this high-altitude
species with his name.

Dolmacoris^ gen. n.

Head bearing conspicuous bristles, ocelli set well on vertex, close to the inner margins
of the eyes; antennae with first three joints and the extreme base of the fourth with well-

' Tibetan sGrot-via. pronounced Dolnia, the most popular goddess of the lamaistic pantheon, better known by
the Sanskrit name of Tara. The specific name is in honor of my friend Dr. Helhnut de Terra, leader of the Yale
Nortli India E.xpeditiun.

130 TERRESTRIAL FAMILIES OF IIEMIPTERA-IIETEROPTERA

developed bristles; bucculae well developed; rostrum short reaching Init to the posterior
margin of the prosternum; apex of second joint reaching but to the base of the head, the first
joint the longest; sides of pronotum slightly explanate, pronotal disc with two large raised
circular areas, and an ill-defined longitudinal carina; sutures between thirtl and fourth and
Ijetween fourth and fifth abdominal stcrnites almost straight, reaching almost to the con-
nexivum where they become fragmented and obscure ; glandular patches on fourth sternite
apparently absent; abdominal spiracles all ventral save that of the fourth segment which
is situated dorsally on the connexivuni ; anterior femora incrassated but unarmed, well-devel-
oped tarsal aroliae absent. Genotype : — D. deterrana sj). n.

7. Dolmacoris deterrana sp. n.

Color. Dull greyish-yellow, somewhat suffused with orange, eyes and ocelli reddish
brown, punctures and bases of bristles very dark brown or black, posterior smooth part of
vertex and central carina of pronotum slightly paler, elytra with two large tubercles reddish,
abdomen dorsally obscurely mottled with brown, anterior margin of connexival portion of
tergites and a small transverse stripe on the same on tcrgites five, six, and seven, black;
abdomen ventrally black mottled with greyish-}-ell<)w laterally, antennae and legs greyish-
yellow with large black spots at the bristle bases, fourth antennal joint uniformly blackish
brown, femora with heavy black puncturation, apices of tarsi somewhat darkened.

Head. Dorsal surface, save for two areas immediately around the ocelli, a median
area nn the extreme jjostcrior i)art of the vertex and the anterior three-quarters of the
\entral surface lateral to the bucculae, coarsely :im\ irregularly ])uncturc(l: anterior and
postero-central part of head dorsally with conspicuous sparsely set bristles, two being set
on the labrum; width of head with eyes greater than length seen from above (0.98, 0.76
mm.) ; clypeus very distinctly separated l)y furrows from the jugae; antenniferous tubercles
well developed, downwardly directed in lateral view (Plate IX, fig. 3), and giving the pre-
ocular part of the head a very slightly constricted outline in front of the eyes; eyes very
large and subpedunculate, situated behind the middle of the head; bucculae well developed
and elevated, somewhat divergent, reaching practically to the posterior margin of the head,
suddenly and obliquely lowered in the posterior eighth of the latter; rostrum short reaching
but to the posterior margin of the ])rosternutn, the first joint the longest, about one and
one-half times as long as the second, which reaches to about the posterior margin of the
head, the third subequal to the second and slightly longer than the fourth (Plate IX, fig.
5). Antennae rather short, basal joint cylindrical, reaching alx)ut to the apex of the head
and stouter than the others, three basal joints and base of the fourth with strong bristles
which are a little longer than the diameter of the second joint, the latter joint twice as long
as the first, and just under twice as long as the third which is just under half the length
of the fourth (0.25, 0.47, 0.22, 0.45 mm.V

Thorax. Pronotum (Plate IX, fig. 1) subequal in length to the head and one and
two-thirds times as wide as long (1.18, 0.72 mm.), anteriorly narrower and posteriorly
wider than the head, subtrapeziform, with all sides slightly and widely emarginate, anteri-
orly with an ill-defined collar behind which the lateral margins are slightly explanate, forming
a cariniform expansion which is impunctate above and bears a row of five short bristles

TERRESTRIAL FAMILIES OF HEMIPTERA-IIETEROPTERA 131

just within the margin; disc with a few short bristles and two large circular raised areas
with central depressions, behind which are a pair of ill-defined tubercles, between each
raised area and continued behind between the tubercles a very ill-defined longitudinal carina.
Propleuron punctured, its posterior margin distally bent back towards the posterior angle of
the prothorax. Prosternum with a wide well-defined longitudinal rostral depression, the
sides of which are raised posteriorly against the articulation of the anterior coxae, anterior
part of prosternum forming a distinct collar which is coarsely punctured. Apertures of
metathoracic scent-glands small and set a little obliquely, their margins hardly elevated. Legs
with numerous well-developed bristles throughout. Anterior coxa with an inwardly project-
ing lamelliform tooth; anterior femur moderately incrassated, its maximum diameter being
about twice that of the femora of the other legs, subequal in length to the anterior tibia;
the latter slightly expanded apically, twice as long as the tarsus (0.84, 0.90 mm.), first tarsal
joint twice the second which is about two-thirds the length of the third and equal in length
to the claws (0.16, 0.09, 0.13, 0.09 mm.). Intermediate coxa acutely angulate interno-
posteriorly but not produced into a definite tooth, femur subequal in length to tibia; the
latter twice as long as the tarsi (0.84, 0.44 mm.), first tarsal joint three times as long as
second, second about two-thirds as long as third and equal in length to the claws (0.18, 0.09,
0.16, 0.07 mm.). Posterior coxa obtusely angulate interno-posteriorly, femur very slightly
shorter than tibia; the latter twice as long as the tarsus (1.24, 0.62 mm.), first tarsal joint
equal in length to the others together, third one and two-thirds as long as second, claws a
little shorter than the latter (0.29, 0.11, 0.18, 0.07 mm.). No aroliae can l)e made out on
any tarsi.

Elytra (brachypterous) covering the proximal half of the abdomen ; widely expanded
in their proximal quarter so as to cover the base of the connexivum, in their distal three-
fourths slightly narrowed exposing the connexivum; posteriorly obliquely truncate; claval
vein (cubitus) well developed and tuberculate, inner corial vein {media) more or less obsolete,
represented by a feebly developed carina bearing a single minute tutercle; subcosta -}-
radius well developed, dividing behind the middle of the elytra to form two large tubercles,
with the inner, more anterior, one of which, the inner corial vein appears to fuse, three longi-
tudinal tuberculate carinae behind the tubercles apparently re])resent the subcosta, radius and
media freely approaching the posterior margin of the elytron.

Abdomen. Broad, depressed centrally, coarsely and irregularly punctate, sutures between
sternites two and three, three and four, and four and five, almost straight, very slightly
turned forward at their distal ends, especially in the case of that between three and four,
the latter antl that between four and five not quite reaching the connexivum and irregularly
fragnnented at the ends; all spiracles, small, those of the fourth segment dorsal, the rest
ventral (Plate IX, fig. 2) ; opaque glandular patches not developed on the fourth or any
other sternite ; fourth and fifth tergites with their posterior margins produced backwards as
obtuse angle, each angle enclosing a well-marked tubercle, the two tubercles sul^equal in size
and rugose.

S Seventh abdominal tergite evenly rounded behind, posterior margin of sternite
straight (Plate IX, fig. 8) ; genitalia as in most Aphaninae, with a long spiral vesica (cf.
Singh-Pruthi 1925), ba.sal plates moderately large (Plate IX, fig. 6), parameres dilated
sub-hasally, narrower and slightly bent apically (Plate IX, fig. 7).

132 TERRESTRIAL FAMILIES OF IIEMIPTERA-HETEROPTERA

9^Seveiith abdominal teryitc with a wide, deep semicircular emaryination in its posterior
border, seventh stemite cleft throughout. Eighth tcrgite with posterior margin sharply
emarginate in the extreme centre. Gonapophyses unarmed (Plate IX, fig. 4).

Length 6 (liolotypc) 4.13 mm., l)readth 2.05 mm.

Length 2 (allotype) 4.55 mm., Ineadth 2.15 mm.

Indian Tibet. 2 S S (holotype and parol ypc), 5 5 9 (allotype and paratypcs).
L63, l)etween Nying-ri and Chungang La, altitude 5100-5300 m. (16,800-17,400 ft.), under
and between sparsely distributed j)lants of Artemisia minor J^icq., in company with Tibct-
ocoris margaretae gen. n., sp. n., and Psyllids, on which furm.s it probably feeds, 18-19 July,
1932. The male i)aratype was taken on a slope just above the summit of the Chungang La,
altitude 17,397 ft., on the boundary between Indian Tibet and Tibet proper.

In spite of its unarmoured femora and straight abdominal sternal sutures this remark-
able insect is referred to the .\phanine tribe Gonionotaria on account of the disposition of
the bristles on its head and antennae, and the position of the abdominal spiracles. Doluiacoris
is clearly allied to Dioinphalus Lieb., which also has straight abdominal sternal sutures, very
similar brachypterous elytra, no conspicuous tarsal aroliae (Fieljer, 1864, T. 1., iig. ]\^ f.)
and a small spur on the anterior coxa. The structure of the rostrum, which in Diomphalus
reaches to beyond the middle of the mesosternum, with a basal joint reaching almost to the
base of the head, constitutes the most striking generic character of Ihiliiiacoris. The short
antennae and trapezoidal pronotum suggest comparison rather with the little known 'i'rans-
baikalian Dioinphalus anmilicornis Jak., than with 1). hispidulus I'ieb., but Jakovleff (1889)
makes no mention of his species differing from hispidulus in the structure of its rostrum so
that it is presumably correctly placed in Dioinphahis. Doluiacoris ajipears to have larger
eyes than either species of Dioiuphalus and in the latter genus the anterior femora appear to
bear spurs; the shape of the pronotal bosses prol)ai)ly furnishes a further generic character.

In the cour.se of examining DoluMcoris it became apparent that no adequate informa-
tion was available as to the position of the abdominal spiracles in the \arious tribes of the
.Iphauiuae as well as in certain of the other subfamilies of the Lygaeidae. Mr. \V. E. China
most kindly offered to make ])reparations from representative species of each tribe of the
Aphauiuac, using as far as possible the typical genera and also of representatives of a numl)er
of other subfamilies. The results of these studies Mr. China most generously asked me to
incorporate in the present paper (Table II). A few words may therefore be appropriately
devoted to the problems of the classification of the Lygaeidae raised by these data. Omitting
the Aphaninae it is clear that while there is a general progression from a dorsal to a ventral
position when the subfamilies are considered in the order currently used in systematic works,
3'et this [jrogression is not as regular as would appear from the keys that have been published,
as, for instance, those given by Stal (1872) or in the excellent work of Barbour (1917,
1918). The following points require comment:

1. The Lyga-einae and Cyminae are generally stated to have entirely dorsal spiracles;
this appears to be essentially correct, though the spiracles on the seventh segment of Cvuius
are almost lateral, being situated dorsally on the conjunctival membrane between the con-
nexivum and the sternite.

2. The Hcnestarinae are omitted from Barbour's key as the subfamily is unrepre-
sented in the Nearctic Region. If it is to he included with the Blissiuac and Geocorinac, as
is done by Stal (1872), the key character defining this group of subfamilies must be emended

TERRESTKIAI. FAMTI.IKS OF HF.MT PTERA-IIF.TF.ROPTERA
TABLE II

Position of Abdominal Spiracles in Lygaeidae

Species Segment

2 3 4 5

Lygaeinae

Lygacus pandunis D D D D

Chauliopinae

Chauliops bisantula D D D D

Cyminae

Cyinus claviculaliis D D D D

Metrarginae

Metrarga (Ncsocryptias) villosa D D D D

Geocorinae

Gcocoris limbatus D D D D

Henestarinae

Hcncstaris laficcps V D D D

Blissinae

Blissus leucoptcnts D D D V

Artheneinae

ChUacis typhae D V V V

Oxycareninae

Oxycarcnus hyalinipennis D V V V

Heterogastrinae

Hctcrogastcr urticac V V V V

Pachygronthinae

Pachygrontha antennata V V V V

Aphaninae-Cleradaria

Clerada apicicornis , V-L V V V

Myodocharia

Orthaca paUicornis D D D V

Rhyparochromaria

Rhyparochroimis chiragra V D-L D V

Plinthisus hrcvipcnnis V V V V

Aphanaria

Aphanus vulgaris V D D V

Gonionotaria

Gonionotus niarginipunctaliis V V D V

Ischnopcza pallipes \' V I ) V

Doluiacoris Jclcrrana V V I) \''

Lethaearia

Lethacus longiroslris V V V V

D := dorsal, \" = ventral, D-L = dorsal on conjunctival nienihrane, V-L

conjunctival membrane.

L33

D

D

D

D

D

D-L

D

D

D

V

V

V

V

V

V

V

V

V

V

V

V

V

V

V

V

V

V

V

V

V

V

V

V

V

V

V

V

V

V

V

= ventral on

134 TERRESTRIAL FAMILIES OF II KMl PTEKA-ll KTKROI'TERA

to "allxjf the abdiiminal spiracles not situated ventraliy, at least those of the third and fourth
segments dorsal"; for, as is indicated in the table, the spiracles on the second sei^inent of
Hcncstaris are ventral, so that the original statement that at most only the last three
spiracles are ventral is incorrect.

3. In the Hetcrogastrinae, Pachygronthinac, Arthcncinae and Oxycareninae all the
spiracles are usually said to be ventral. In Oxycarcnus and Chilacis, however, those of the
second segment are dorsal. The emended key character dehning this group of subfamilies
should therefore run "'all or at least the live posterior abdominal spiracles situated ventrally."

4. In the Chouliopimic and Mctrarghuie, subfamilies not examined by Stal or Barbour,
all the abdominal spiracles are dorsal. The Mctrargiuac were stated by Kirkaldy (1902),
in erecting the subfamily, to be allied to the Cyiiiinac, but to have the last three abdominal
spiracles placed ventrally. Later he concluded (1908) that the subfamily was more probably
allied to the Ox\careninac. It is probable that Kirkaldy mistook three prominent pairs of
trichobothria for ventrally placed spiracles in uncleared material. In reality the aflinilies of
this peculiar Hawaiian subfamily with the Cyiiihuic are great, the chief differential character
being that in the Metrarginac, unlike any other member of the family, the hamus of the alar
areole is "continuous, extending from the vena subtensa upwards In the ujjper vein" (Kirkaldy,
1902).

5. The Aplia)iiiuu (Rhyparocliroiniiiae) have never been separated on spiracular char-
acters and show great diversity in this respect. The Myodocharian arrangement as exempli-
fied by Orthaca is similar to that obtaining in the Blissinae, while in the Lcthacaria and in
Plinthisns an entirely ventral arrangement is found as in the Hclcrogastrinae and Pachy-
gronthinac, and an almost identical pattern is found in Clerada. On the other hand the
arrangements with the spiracles of the third and fourth segments alone dorsal or dorso-lateral
as in Aphanus and RhyparocliromiiS, or with only those of the fourth segment dorsal as
exemplified by the Gonionotaria, are not found outside the Aplmninae. In conclusion it
would seem that although the position of the spiracles may be of great value in the construction
of artificial keys and in determining the relationships of individual genera and tribes, too
much stress must not be laid on so variable a character in determining the natural subdivisons
of the family.

Family ANTHOCORIDAE

Tribe Anthocoraria

8. Ectemnus paradoxus sp. n.

Color. Head, pronntum, scutellum and ventral surface l)iack; eyes and (xelli dark
vinous; first antennal joint black, second yellow with a little black ba.sally and the extreme
apex greyish-black, third yellow, narrowly black apically, fourth black, slightly paler basally.
Elytra with fine sparse pale golden pulx?scence, clavus brown, its inner margin ])aler and outer
margin darker than the disc, corium and embolium basally lacteous, apically i)iceous, extreme
apex of corio-embolial suture, in the neightourhood of the anterior margin, hyaline, cuneus
piceous black, membrane opaque lacteous with a large central and a still larger sub-apical
spot greyish-black, the areas around and Iietween the spots luteous; femora black, tibiae ])ale
testaceous, their apices somewhat darker, tarsi greyish-black.

TERRESTRIAT. FAMILIES OF UEMIPTERA-I I ETKUOPTERA 135

Head elongate (I'late X, fig. 3), just under one and a half times as long (0.53 mm.)
as width, with eyes (0.36 mm.), anterior margin of eye inserted very slightly behind middle
of lateral margin, head somewhat constricted in front of insertion of antennae and also before
the posterior margin; rostrum reaching to posterior margin of anterior coxae; first antennal
joint reaching just to apex of head, second three and a third times the length of the first,
third twice the length of the first and very slightly shorter than fourth (0.11, 0.40, 0.24,
0.27 mm.), second joint slightly thickened baso-apically, third and fourth hardly narrower
than the middle of the third.

Pronolum twice as wide posteriorly (0.73 mm.) as long (0.36 mm.) with a very distinct
apical collar less than half the posterior width (0.31 mm.) and marked transverse impression,
disc finely rugose and covered with very fine short pale hairs, lateral margins sinuate, some-
what raised and marginate, posterior angles sub-acute, directed backward and not projecting
laterally.

Scutellum slightly shorter than pronotum, and one and a half times as wide (0.44 mm.)
as long (0.29 mm.), disc with sparse, very short fine pale hairs, little raised anteriorly,
remotely punctate and nitid, slightly depressed before apex which is rugulose.

Presternum rugose, its disc flattened centrally, posterior margin produced to form an
acute xyphus between the anterior coxae.

Mesosternum nitid, very finely and regularly rugulose, posterior margin narrowly emargi-
nate, disc with a fine groove running forward from the emargination and becoming obsolete
anteriorly.

Metasternum transverse, between the widely separated posterior coxae, but little raiseil,
coarsely and irregularly rugose, posterior margin truncate.

Orifice of metathoracic scent-gland straight, produced rather prominently at the outer
end (Plate X, fig. 7).

Tibiae of all legs but little longer than femora (ant. 0.51, 0.55; inter. 0.51, 0.55; post.
0.80,0.87 mm.).

Elytra and wings macropterous, the latter without a hamus (Plate X, fig. 6).

Abdomen distinctly surpassed by the elytra.

S Left paramere short, broad and semicircular (Plate X, fig. 8), right paramere vestigial.
Length 2.55 mm., breadth 0.80 mm,

Indian Tibet. 3 5 5 (holofype and paratypes) Igu, in the Indus \'alley above Leh, on
the bark of Populus sp. ; altitude 3417 m. (11,210 ft.), Sept., 1932.

The present species is anomalous in that it lacks the hamus of the wing cell, a character
which would remove it from the Anthocoraria and place it in the Lyctocoraria as defined by
Poppius (1909). Eetemnus paradoxus, however, runs down perfectly to its genus in the key
to the Anthocoraria given by this author, if once its membership in that tribe be admitted.
Apart from the absence of the hamus it appears tn lie an entirely normal member of its genus.
If, therefore, it is to be removed from the Anthocoraria, a new genus of the Lyctocoraria
must be defined, isolated from all the other memlwrs of that tribe, and differing only from
the Anthocorarian genus Eetemnus in the single character under discussion. This is clearly an
unsatisfactory proceeding and the present species is therefore described as an Eetemnus. It
is clear that the value of the presence and position of a hamus as a major taxonomic character
is dubious, but I am not in a position to revise the tribal characters of the Anthocoridae,

Llf) TKKKKSTKIAl. FAMTI.IES OF TTKMTPTF.R A-II KTRRDI'TF.KA

nor to pFOvide any new distinction Ijetween tlie two tribes. It may be pointed (lut tliat Cliina
(1933) also appears to be soniewliat doubtful of the value of liainal characters for this
purpose.

The genus Ectoimus at present contains four species. E. loiigirostris 1 lorv. from the
Balkans is sharply distinguished by its rostrum which reaches to the intermediate coxae. Of
the remaining species the widespread Palaearctic E. rcdmimts (H.-Sch.) is an insect of very
different facies from paradoxus; it is usually brachypterous and the head and pronotuni are
ferrugineous brown. E.. parilis llorv. is known only in the brachypterous state, the head
and anterior part of the pronotum are black as in paradoxus, but the posterior part of the
latter fades to ferrugineous, and the whole of the fourth, the a])ical half of the third and all
of the second antennal joint save a yellow ring are black. /:'. pictiprniiis Esaki (1931) a
macropterous species from Japan, in which, as in paradoxus, the head .and pronotum are
entirely black, differs from the latter, as is clear from Esaki's excellent description and
figure, in having the fourth antennal jc^nt yellow, the sides of the pronotum straight and the
head unconstricted l^ehind the eyes.

The species nearest geographically to paradoxus is rcduvinus, which is recorded by
Oshanin (1889, 1912) from Russian Turkestan, but it is possible that Galchana Distant
(1910) is a synonym of Ecfeiunus, though the type and only species, G. Iiuiiicralis from
Simla, is clearly distinguished by its pointed posterior pronotal angles from E paradoxus.

9. Anthocoris gyalpo" sp. n.

Moderately broad and robust, 2.85 times as long as wide.

Head, antennae, basal half of rostnnn, pronotum, scutellum, dorsum abdominis and
ventral surface black; ape.K of penultimate joint of rostrum testaceous, ultimate joint brown,
posterior part of metapleuron liehind scent-gland and apex of abdomen beneatii, obscurely
testaceous; legs testaceous, the bases of the coxae piceous, extreme bases of femora and tibiae
slightly darkened, dorsal surface of anterior femora slightly infuscated subapically, posterior
femora darkened along the posterior margin, tarsi brown, all these markings obscure, the
legs l)eing without any definite spots or annulations; elytra pale testaceous brow^n, practically
unmarked, the base and internal margin of the clavus, the corial veins and the apex of the
cuneus being very slighdy darker, membrane grey, infuscated subapically.

Head with a few pale hairs anteriorly, one and a sixth times as long as wide (0.60,
0.51 nun.), suddenly and then more gradually narrowed in front of the eyes, postocular region
constricted, antennae longer (1.44 mm.) than the length of the head and pronotum together
(1.07 mm.), first joint not reaching the apex of the head, second joint subecpial in length to
tiie width of the head and eyes, and half as long again as the third which is subequal in
length to the fourth (0.15, 0.55, 0.36, 0.38 mm.); second joint about half as thick basally
as subapically, where it is very slightly thicker than the first joint, all joints clothed with fine
])ale hairs which are subequal in length to or a little shorter than the niaximum thickness of
the .second joint, hairs niore abundant and more closely adjjres.sed on the apical half of the
fourth joint ; rostrum reaching to just beyond the centre of the anterior coxae, its first visible
joint (damaged in unique type) apparently not quite reaching to the insertion of the antennae;
second joint about twice as long as third (0.25, 0.47 mm.).

' Tibetar. rGyal-f'o, a king ; the garden in which the unique holotype was taken formerly surrounded a pavilion
or summer residence of the Gyal-po of Leh.

TICRRKSTRIAT, KAMIT.IES OF 1 1 KM I I'TKKA-IIF.TKRI 1I'T|-.UA 137

ProiiotiiDi (Plate X, fig. 2) covered with fine slmrt jiale hairs, its niaxinuiin wicUh aliout
two and a quarter times the median length (1.06, 0.47 mm.), anterior collar moderately well
developed, its width (0.38 mm.) just over one-third the maximum width of the pronotum,
lateral margins immarginate feebly rounded from the collar and quite straight throughout the
greater part of their length, posterior angles obtusely pointed, disc strongly rugose, save a
longitudinally impressed raised transverse area immediately in front of the transverse fovea,
which is set in the middle of the mid-line of the pronotum and occupies more than one-
third of the width of the pronotum at that level, posterior part of disc with traces of a longi-
tudinal central depression, posterior margin widely emarginate before the base of the scutellum.
Scutellum covered with fine short pale hairs, longer (0.62 mm.) than the pronotum and
about one and a cpiarter times as wide (0.77 mm.) as long, with a well-marked transverse
fovea behind the middle, anteriorly somewhat swollen and remotely punctate, apex somewhat
rugose.

Prosternum with its posterior margin somewhat marginated, except centrally where it
is produced backwards as a short xyphus betwesn the anterior coxae, its disc somewhat rugose,
with an indistinct transverse carina behind the middle.

Mesosternum smooth, its posterior margin rounded and centrally a little emarginate,
disc with a fine longitudinal groove running forward from the emargination almost to the
anterior coxae.

Metasternum rounded posteriorly and elevated.

Orifices of metathoracic scent-glands curved slightly forward externally and \\ith a very
fine carina running forward from the outer end of the orifice (Plate X, fig. 4).

Legs with fine pale hairs on all joints, slightly sparser than those of the antennae,
tibiae slightly incrassated apically, anterior femur very slightly shorter than the tibia, which
is just over three times the length of the tarsus, the latter just under three times the length
of the curved claws (0.73, 0.80, 0.25, 0.09 mm.), intermediate femur very slightly shorter
than the tibia, which is just under three times the length of the tarsus, the latter just over
three times the length of the curved claws (0.76, 0.84, 0.31, 0.09 mm.), posterior femur
about five-sixths the length of the tibia, which is a little over three and a half times the length
of the tarsus, the later about three times the length of the straight claws (1.02, 1.20, 0.33,
0.11), last tarsal joint of each leg just over half the length of the tarsus.

Elytra surpassing the apex of the abdomen, covered with fine short pale hairs, coarsely
but obscurely punctured and sub-nitid throughout, cuneus entirel}' Ijehind the apex of clavus,
its marginal length (0.62 mm.) about three-fifths that of the embolium (1.16 nun.), embolial
margins straight and subparallel, all membranal veins save the outer one more or less obsolete.

S . Left paramere narrow, sickle-shaped, and angulate (Plate X, fig. 5).

Length 3.52 mm., maximum breadth 1.24 mm.

Indian Tibet. S (holotypc) Leh, Residency Garden, apparently IjIdwu from Pupitlics
sp., 19 September, 1932.

The present species, in its rostruiu, metasternum, odoriferous glands, runeus and pro-
notum agrees sufficiently well with Antliocoris to be included in that genus. It differs from
its previously described congenus in its almost unicolorous elytra which are perhaps more
clearly punctate than usual in Antliocoris. A. gyalpo appears to belong to that section of the
genus in which the antennae are longer than the head and pronotum together, of which
A. syh'cstris (Linn.) is the best known member, but is easily distinguished from var. nigri-

138 TEKRESTKIAI, FAMII.IKS Ol' IIF.M ll'TKKA-l I KTKUOITr.UA

cornis (.Fieb.) of this species by its almost uniform clytral C(jloration and curved orifice of
the odoriferous gland. From the other species included in this section it is also distinguished
by the different proportions of the antennal joints. Poppius (1909) has described two
members of this group, viz., anmdipcs and indicus from Darjeeling, l)ut these appear to be
normally coloured memljers of the genus witli black or annulatcd femora {vide Distant, 1909,
figs. 166, 167). The almost complete suppression of all but the outer membranal veins
suggests Conipsobiella Poppius (1909) but the present species shows none of the other char-
acters of this Central .\frican insect and a somewhat similar rcductinn is found in the species
of the sylvcstris group.

Family MIRIDAE

Subfamily Duvi'Iiinak

Triljc Dicypluiria

10. Dicyphus physochlaenae sp. n.

Plead black, tlic inner Ijorder of the eyes margined witii }-eIl(iw which spreads out
towards the central black area of the vertex from tlie postero-internal angle of the eye, centre
of frons with a longitudinal yellow stripe w^hich spreads anteriorly to the bases of the
antennae, vertex with two submarginal yellow spots posteriorly (Plate X, fig. 9). Pronotum
grey, with a transverse stripe across the calli and subapical fossa piceous lihick, tiie stripe
interrupted l)y a longitudinally elongated yellow spot between the calli but uninterru])ted more
anteriorly, grey part behind median fossa with an anterior median yellow sjiot narrowly
connected with the spot between the calli, outer part of calli marked with brownish yellow.
Scutellum black, basal angles narrowly orange, apical half of margins w^ith greyish yellow
vittae wliich become obsolete towards the posterior angle. Ventral surface black. Antennae
witii first joint black, very narrowly greyish yellow at the extreme base and apex, second
joint black, very narrowly greyish yellow at the base and witii a conspicuous yellow band
occupying its central quarter, third joint black, a little paler basally, fourth joint brown.
Rostrum yellow, base of second and third and whole of fourth joint black. Coxae yellow
with black bases, femora yellow heavily spotted with black, dorsally tibiae yellow with tiie
extreme base brown and with al)out five brown (anterior) or black (intermediate and pos-
terior) spots on the postero-dorsal part of the proximal third, first and second tarsal joints
yellow and the third black ; all tiljiae armed with fine black spines. Elytra hyaline, greyish,
with the apex of the cuneus piceous, membrane very transparent, greyish, its nerves yellowish
grey suffused with brown.

Head transverse, rather less than one and a half times as wide as long (0.60 mm.,
0.44 mm.), evenly rounded and little produced anteriorly alxive clypeus, the latter in lateral
view with its anterior margin straight ventrally, in its dorsal third rather suddenly rounded
to meet its dorsal suture with the frons, gula longer than bucculae and slightly sinuate.

Antennae fairly thick, first joint surpassing a|)cx of head 1iy about three-quarters of
its length, and about two-thirds the length of the head from above, second joint twice as long
as the first, third joint just over two-thirds the length of the second, fourth about two-thirds
the length of the third, second joint distally about twice as thick as proximally, but through-
out slightly narrower than the first (head length 0.44 mm., antennae 0.25, 0.51, 0.36,
0.25 mm.).

TERRESTRIAL FAMILIES OF IIEMIPTEKA-IIETEROPTF.RA 139

Rostrum not quite reaching" the uiiddle of the intermediate coxae, first joint hardly
surpassing the base of the head.

Pronotum anteriorly about three-quarters of the width of the head, posteriorly about
one and three-cjuarters times as wide as head, two and one-third times as wide as the anterior
breadth and twice as wide as long (head width 0.60 mm., pronotuiu, anterior width 0.45
mm., posterior width 1.02 mm., length 0.1 mm.), anterior collar well marked, its anterior
luargin very slightly sinuate, calli well marked, subconfluent centrally, posterior transverse
fossa central, sides but little sinuate, posterior margin widely and deeply eniarginate, posterior
part of disc rather feebly rugose.

Elytra long, their length from the insertion to the level of the apex teing five times the
median length of the pronotum, and two and one-third times their greatest width, outer
margin slightly explanate centrally. Anterior coxae reaching to the middle of the meso-
stemum, posterior tibia (1.58 mm.) 2.66 times as long as the width of the head and eyes,
third tarsal joints of all legs slightly shorter than second.

3 Left paramere as in Plate X, figs. 11, 12.

$ (holotypc) length 3.53 mm., breadth 1.13 mm.

5 (allotype) length 3.75 mm., breadth 1.20 mm.

Indian Tibet. 2 S S , 4 9 $ (Iwlotype, allotype and paratypes) L 67. Dambu-
guru, altitude 4603 m. (15,100 ft.), on Physochlaena pracalta Hook. (Solenaceae), 31
July, 1932.

In the coloration of the head and legs this species closely resembles D. orientalis Rent,
from Turkestan ; it may ultimately have to l)e treated as a sub.species or form of that species.
The posterior tibia is, however, proportionately shorter than is indicated in Renter's descrip-
tion (1884) of orientalis and the coloration of the antenna is comparable to that of the
widespread western Palaearctic species D. annulatus (Wolff.). In the latter species the black
spots extend throughout the intermediate and posterior tibiae at the bases of the black spines.
D. montanus Poppius (1912) from the Alexander Mts. is another closely allied species which,
however, appears to have a longer basal antennal joint ("nur wenig kiirzer als der Kopf von
ober gesehen") and to have a rather different color pattern on the vertex.

These forms are all clearly closely allied and the coloration, \\hich has been chiefly used
in separating them, is undoubtedly variable. It is hoped that the present figures of the
parameres of D. physochlactiae will make it possible for other workers to decide whether the
present form is specifically distinct. There can meanwhile be little doubt of its Central
Asiatic affinities.

11. Dicyphus sengge* sp. n.

Head yellow, posterior margin black, centre of vertex with a large V-shaped black
mark which tends to become somewhat diffuse at its posterior apical end (Plate X, fig. 10).
Pronotum grey, with a transverse stripe across the calli black, centrally interrupted by a
longitudinal yellow vitta, outer margin of calli yellowish, posterior part of pronotum grey.
Scutellum black, basal angles dull orange, apical two-thirds of margins with broad greyish
yellow vittae which do not quite reach the apical angle. \^entral surface brownish.
Antennae with basal joint black, its apex very narrowly whitish, second joint yellow with the
extreme base and apical third black, third joint black, fourth joint piceous black. Ro.strum

' Tibetan Seng-ge, a lion, tlie setose angle of the left paramere being suggestive of a mane.

140 TF.RKKSTKIAI. FAMIl-IF.S OF 1 1 KM 1 1'TI-.KA-TI ICTI-.Km^TFUA

lirownisK yellow, fourtli joint black. Legs very pale greyish yellow, bases of coxae blackish,
femora with small brown spots, tibiae immaculate, third tarsal joints black; tibiae with
numerous fine black spines which do not arise from spots. Elytra hyaline t;rey throughout,
apices of corial nerves suffused with l)lack, covered throughout, but most strongly in the
lateral (anterior) region with fine black hairs.

Head transverse, rather more than one and a halt limes as wide as lung (0.()0 mm.,
0.37 mm.), evenly rounded and hardly produced anteriorly over the clypeus, the latter in
lateral view with its anterior margin straight ventrally; l)ucculae shorter than gula.

Antennae moderately thick, the first joint suqiassing the apex of the head by al)OUt
three-quarters of its length, and about two-thirds the length of the vertex seen from above,
second joint two and two-thirds times as long as first, third just under three-fourths as long
as the second and fourth about three-fifths as long as third (0.25, 0.65, 0.45, 0.27 mm.).

Rostrum apparently reaching just beyond middle of intermediate coxae (somewhat
damaged in tmique type), basal joint distinctly shorter than head.

Pronotuiii anteriorly about four-fifths as wide as head with eyes, posteriorly about one
and four-fifths as wide as the head, just over twice the anterior breadth and just over twice
as wide as long (head width 0.60 mm., pronolum, anterior width 0.49 mm., posterior width
1.03 mm., length 0.4'* mm.), anterior collar well marked, its anterior margin very slightly
sinuate, calli well marked, subconfluent centrally, posterior transverse fossa lying just
anterior to center, sides rather sinuate, posterior margin widely emarginate, posterior part
of disc feebly rugose.

Elytra about .six times as long as pronotum, and just over twice their greatest width,
outer margin very slightly explanate. Anterior coxae reaching to middle of mesosternum,
posterior tibiae (1.64 mm.) 2.73 times as long as width of head and eyes, third tarsal joints
suljequal to second.

S Left paramere as in Plate X, fig. 13, with very long hairs on the basal part of the
shaft, and a small triangular projection, situated more apically tlian the corresponding pro-
jection in the preceding species and directed upwards.

Length S (type) 3.69 mm., breadth 1.23 mm.

Indian Tibet. 1 S (type) L 37, Ijetween Tangtse and Mugleb, altitude c. 4175 m.,
among grasses, 27 June, 1932.

This species is very close to the preceding, differing in the different coloration of the
head, the more sinuate lateral margins of the pronotum and in the left paramere. Both
species are allied to the above-mentioned Central Asiatic species and to anniilatns. From the
latter species D. scns_i^e differs in the coloration of the head and tibiae, from oriciilalis in the
coloration of the head and antennae, from montanus in the short basal joint of the latter.

Subfamily ri.AGIOGNATniNAE

Tribe Plagiognatharia

12. C/ilaiiiydiiliis piicliycenis Kiritsh.

C. pachycenis Kiritshenko. 1931.

Indian Tibf.t. 1 $ macr., 3 9 5 brachypi. L 33, Shakya La, .south slope c. 5200 m.
(c. 17,000 ft.), air temp, in shade 7.2 C. "jumping about in sun around moss and grass," 25

TERRESTRIAL FAMILIES OF HEMIPTERA-IIETEROPTERA 141

June, 1932. 3 99 brachypt. L 48 Kyang La, among sparse grass and Pcgm-o/'M'/oH prob.
scapifoUiim Marq. and Skan., altitude 5100-5334 m. (16,800-17,500 ft.), 9 July, 1932.

1 $ macropt., 1 9 brachypt. Ororotse Tso, altitude 5297 m. (17,381 ft.). 11 July, 1932.

2 $ $ macropt., 3 9 2 brachypt. L 54a, north side of Marsimik La, altitude c. 5300 m.
(17,400 ft.), grassy place, 16 July, 1932. 1 $ macropt., 1 5 brachypt. Kyam, altitude
4733 m. (15,530 ft.), grassy place, 20 July, 1932. 3 5 5 , 1 9 all macropt. Nyagtzu, alti-
tude 4671 in. (15,324 ft.), grassy place, 30 July, 1932; 1 <5 macropt. Peldo-le, north end
of Tso Moriri, altitude 4529 m. (14,835 ft.), mixed vegetation with grasses dominant, 31
Aug., 1932. 1 9 brachypt. Tsak-shang, 31 Aug.; 1932.

This species was described from material taken between 13,500 and 16,500 feet in
southern Tibet by Major R. W. G. Hingston on the Third Mount Everest Expedition.
According to Kiritshenko both sexes may be brachypterous ("Hemelytra — magis minusve
abbreviata"), while only males may be macropterous. In the present collection all the males
and a single female appear to be macropterous. Hingston, at his highest locality, notes that
the species was "common at the entrance to tunnels of mouse-hares"; in spite of much
observation on this point I never found the slightest trace of such an association. Since
short grass is the only plant, common to every locality, on the vegetation of \\hich I have
notes, there can be little doubt that this species is graminivorous.

Tibetocoris gen. n.

Elongate, cluthed abi)\-c with lung irregular sparse, ])alc pubescence, which is somewhat
tomentose on the head.

Head (Plate X, figs. 14, 15, 1()) from above but little produced anteriorly, facial angle
subrectangular, clypeus moderately prominent, wide, very slightly depressed dorsally, slightly
compressed ventrally, subparallel in lateral \iew, dorsal suture indistinct, lying just above a
line drawn across the insertions of the antennae, bucculae moderately well developed, gula
distinct, rostrum reaching almost to the apex of the intermediate coxae, anterior joint short
and thick, but little surpassing the posterior margin of the head, vertex unimpressed, its
posterior margin convexly rounded centrally and feebly marginate laterally, eyes large,
ommatidia granuliform, interocular distance less than twice the dorsal width of an eye, in
lateral view eye elongate; loro-genal suture distinct; frons and anterior part of vertex feebly
striate on each side. First antennal joint surpassing the head by about half its length.
Pronotum very transverse, just over twice as wide as long, anterior border centrally emargi-
nate, posterior margin very widely and lightly sinuate, lateral margins straight, anterior
callosities poorly developed. Proxyphus flat, its margins obscurely marginate ; mesostemum
reaching a little beyond the apex of the anterior coxae, its posterior border emarginate
centrally.

Tibiae with fine Ijlack spinous bristles; pseudarolia narrow, connate throughout its
entire length, reaching about to the centre of the evenly and lightly curved claw, basal tooth'
obtuse, aroliae very fine, bristle-like and subparallel (Plate X, fig. 17). Hamus of wing
cell arising opposite the base of the vena decurrcns. Gciiol\'/^c : T. margaretae sp. n.

142 TERRESTRIAL FAMILIES OF II F.MirTERA-nF.TEROPTERA

13. Tibetocoris margaretae sp. n.

Head, pronotum, scutelliim and elytra clothed above with long irregular sparse pale
pubescence, which is somewhat tonientose on the head ; and with a few black hairs on the
elytra. I'ale greyish white, tinged with yellowish green, vertex near eyes minutely trans-
versely striate with brown, pronotum. scutellum, elytra, and distal third of femora with
minute brown spots, antennae greyish brown, liasal joint paler, tarsi brown becoming almost
black apically, inesosternum and apical joint of rostrum black, alxlomen greenish grey. First
joint of antenna with two subapical black bristles, second joint narrow proximally, somewhat
widened apically but throughout narrower than the first, third and fourth subequal in
width, and slightly narrower than the proximal end of the second; second joint just under
three times as long as the first ; third joint about three-fifths as long as the second and fourth
three-fifths the third (0.33, 0.95, 0.58, 0.33 mm.).

Anterior femur with three conspicuous subapical and three small apical bristles, tibia
with about nine black liristles; the tibia one and one-third times as long as the femur and
twice the length of the tarsus (0.80, 0.98, 0..^1 mm.), the tarsal joints overlapping at their
articulations, the second a little longer than the first and a little slmrler than the third which is
twice as long as the claws (0.15, 0.18, 0.22, 0.11 mm.).

Intermediate femur with two apical and one conspicuous subapical bristle tibia with
about 16 bristles, tibia just over one and one-third times the length of the femur and two
and a half times the tarsus (0.90, 1.24, 0.51 mm.), second tarsal joint twice as long as first,
third just longer than .second and more than twice as long as claws (0.12, 0.24, 0.25, 0.11
mm. I. Posterior femur with two conspicuous subapical bristles, tibia with about 16 bristles,
tibia about one and a half times as long as femur and three times as long as the tarsus
(1.34, 2.04, 0.69 mm.), second tarsal joint two and a half times as long as the first and
slightly longer than the third which is just over twice as long as the claws (0.15,0.36,0.33,
0.15 mm.).

S Right paramere hook-.shaped ( Plate X, fig. 18), left jjaranicrc styliform (Plate X,
fig. 19).

1-ength 3.75 mm., breadth 1.27 mm.

Indian Tibet. 5 ^ ^ , L 57, Slope of mountain on south side of the valley of the
Chang-chenmo River, near Pamzal, altitude c. 5220-5270 m. ( c. 17,000-17,300 ft.), on
Artemisia minor Jacq., 18 July, 1932. 3 3 <J {type and paratypes) L 62a, Nying-ri c.
5120 m. (c. 16,800 ft.), on Artemisia minor Jacq., 26 July, 1932. 6 $ S L 63 Chungang
La, just alx)ve top of pass which constitutes tha boundary of India and the independent terri-
tories of Tibet, altitude 5305 m. (17,400 ft.), on Artemisia minor Jacq., 19 July, 1932. 1 5 , 77
Kakstet La, altitude c. 5360 m. (c. 17,600 ft. ), on Artemisia minor Jacq., 18 August, 1932.

The present genus is probably most clossly allied to Tuptonia Renter, from which it
differs mainly in the longer pseudarolia. This character, if the feeble markings on the
femora be neglected, would bring the genus into the neighbourhood of Asciodema in Renter's
key (1884); Asciodema, however, differs m:irkedly from V'/fcc/orwrn in the structure of the
head and legs. The specimens are all somewhat teneral and the prosternuin therefore tends
to be distorted, but in the l:)est preserved of them it shows no trace whatever of being convex,
in this rather resembling the series of genera originally separated by Renter as the flivision
Oncot\Iaria.

TERRESTRIAL FAMILIES OF IIEMII'TERA-IIETEROPTERA 143

ZnOGEOGRAPIlICAL NOTi:s ON THE HeTEROPTEROUS FaUNA OF HiGU ALTITUDES

The zoogeographical pruljlems raised by the present collection center around the exist-
ence of a number of endemic species, some belonging even to endemic genera, at liigli
altitudes in a mountainous region which was undoubtedly subjected to intense glaciation
during the Quaternary Ice Age.

Prior to the present investigation the highest recorded locality at which Heteropterous
bugs had been collected was Rongbuk in South Tibet, where at an altitude of 16,500 ft.
Kingston obtained much of his material of Chlamydatus pachycenis (Kiritshenko, 1931b).
A hitherto unrecorded nymph of Nysiits was also obtained by the same investigator at an
altitude of 17,000 ft. In the Yale North India Expedition collections the following four
species are represented from still greater elevations :

Nysiiis cricar alticola siibsp. n., u\) to 5297 m. (17,381 ft.)
Dohnacoris dctcrrana gen. n., sp. n., up to 5300 m. ( 17,400 ft.)
Chlamydatus pachycenis Kiritsh., up to 5334 m. (17,500 ft.)
Tibctocoris margarcfae gen. n., sp. n., up to c. 5360 m. (17,600 ft. )

These four species fall into two ecological and zoogeographical groups. A^^. c. alticola
and C. pachycenis occur among grasses and small herbaceous plants, mostly Cruciferae, the
dominant members of the mesophj-tic high-altitude vegetation. Both are probably widely
distributed in the Himalayan and Karakorum ranges wherever the appropriate flora is devel-
oped, for, as has been pointed out, C. pachycenis and an undetermined species of Nysius
are known from but slightly lf)wer localities in the Everest region. It is also of interest to
note, that of the three free-living species of Heteroptera inhabiting Greenland, two (cf.
Qiina, 1934) belong to the genera Nysius and Chlamydatus. It is therefore very probable
that, during the Quaternary Glaciation, both these genera, and perhaps no others, could
survive in the highest zone of vegetation, in the immediate vicinity of the ice. D. deterraiia
and T. niari^arctae, on the other hand, are apparently confined to a drier type of hal)itat,
where the dominant plant is Artemisia minor Jacq. Both species belong to monotypic genera
most closely related to groups that reach their highest development in Central Asia. The
well-defined generic characters of Dolmacoris make it most improbable that it is a recent
immigrant to the western Tibetan Plateau and strongly suggests that the fauna of the present
region of xerophytic vegetation in this region has survived the Quaternary Glaciation in
unglaciated parts of Western Central Tibet under semi-arid conditions.

The Pamirs are the only mountains of Central Asia of which the Heteropterous fauna
is at all well known. Here from heights of over 4,000 m. Kiritshenko (1931a) records 27
species of which three belong to the Acanthiidae, not treated in the present paper. Of the
24 truly terrestrial Heteroptera of the High Pamir only two are endemic species. Omitting
three forms only recorded from the Indus valley from localities lying below 4,000 m. and
in each case associated with poplar trees which do not grow above this height, the num])er of
species at present known from Indian Tibet is 11. If to these are added two extremely
doubtful records, a nymph of Psallus sp. (Dambu-guru) and Teratocoris sp. (Tukung, S.
of the Panggong Tso), which I noted but of which no specimens were found when the
collections were unpacked, the total number of species is increased to thirteen, just over half
the number recorded fnmi tlic High Pamir. Of these, however, si.x appear to be endemic

144 TERRESTRIAI, TAMIUES OV II KMIITKRA-Ii KTKROl'TKRA

to Indian Tihi't and one to Indian Tibet and tlie Southern ilinialaya. Moreover, it is
probable that were a male of the species of Sticto/^lnira c)l)tained availaljle, tliis tno would
be found to l)e an endemic species. It tlierefore appears that at least liaif tlie species of
the region are peculiar to tlie I liinalayan and Karal<i)rum ranges, and that the High Taniir
though richer in species is much poorer in ]ieculiar forms. This is ])robably to l)e explained
by the fact that, while in the Pamirs after the Quaternary Glaciation a number of routes
for recolonisation were open (Reinig, 1932), putting the high regions into easy communi-
cation with the richest Ilcteropterous fauna in the Palaearctic region (cf. Kiritshenko,
1931 a), in Indian Tibet the only migration nnites were from the North over extensive
mountain ranges and deserts, from the humid south and west where the Sub- Himalayan and
Kashmirian forest fauna is apparently ecologically unsuited to penetrate into very elevated
and semi-arid regions, and from the east where a restricted pre-glacial high-altitude fauna may
have survived in the less glaciated parts of western Tibet jimper. The material available
suggests that certain forms such as Nysitis cricac obsciinitus and Microplax hissaricusis
belong to a Central Asiatic element that has entered by the northern route, while the endemic
genera and perhaps some or all of the endemic species represent a migration from hypothet-
ically unglaciated regions of the Tibetan plateau, where a fauna of undoubted Central Asiatic
origin survived and differentiated at a time when the greater ])art of the Karakorum and
Western Himalaya were heavily glaciated and quite uninhaljitable. Without some such
hypothesis it seems impossible to explain the large proportion of endemic forms in a region
that has suffered so much glaciation in relatively recent times.

Osborn Zoological Laboratory,
Yale University,
July 18, 1934.

TERRESTRIAL FAMILIES OF HEMIPTERA-lIETEROrTER A 145

BIBLIOGRAPHY

Barber, H. G. 1917-1918. Synoptic Keys to the Lygaeidae ( Ilemiptera ) of the United
States. Part I. Psyche, XXIV, p. 128, 1917; Part II. ibid., XXV, p. 71, 1918.

1923. Family Lygaeidae. Guide to the Insects of Connecticut. Part IV. The

Hemiptera or Sucking Insects of Connecticut, p. 708. Hartford, C(jnn.

China, W. E. 1933. A New Genus and Species of Anthocoridac (Hemiptera) from
New Zealand. Ann. Mag. Nat. Hist. ser. 10. XI, p. 514.

1934. Hemiptera collected by the Oxford University Expedition to West Greenland,

1928. Ann. Mag. Nat. Hist. ser. 10. XIII, p. 330.

Distant, W. L. 1879. Rhynchota. Scientific Results of the second Varkand Mission,
based on the collections of the late Ferdinand Stoliczka. vol. II. Calcutta.

1910. Fauna of British India. Rhynchota. vol. V, p. 297. London.

Ekblom, T. 1931. Hemipteren aus dem Sarekgebiet. Naturwiss. Untersuch. des Sarek-
gebirges in Schwedeisch-Lappland. Bd. IV, Zool. Lief. 10. p. 939. Stockholm
and Berlin.

EsAKi, T. 1931. Undescribed Hemiptera from Japan and Formosa. Annot. zool. japon.
XIII, p. 264. Tokyo.

Evans, J. W. 1929. A new species of Nysiiis (Hem., Lygaeidae) from x\ustralia. Bull.
Entom. Research. 1929. p. 351.

HoRVATH, G. 1889. .\nalecta ail Cognitionem Heteropterorum Himalayensiunr Term.
Fuzetek, Xil, p. 29.

1890. Synopsis des Nysiiis palearctiques. Rev. d'l^ntonL IX, p. 185.

1904. Insecta Heptapotamica. I. Hemiptera. Ann. Mus. Nat. Hung. II, p. 574.

Jakowleff, B. E. 1889. Zur Hemipteren Fauna Russlands und des angrenzenden Lander.
Hor. Soc. Ent. Ross. XXIV, p. 332.

KiRiTSHENKO, A. 1910. Espcce nouvelle du genre Fhiviodcni Latr. trouvee dans I'Altai.
Rev. russ. entom. X, p. 21.

1913. Hemiptera-Heteroptera turanica nova. Rev. russ. entom. XIII, p. 412.

1926. BeitrJige zur Kehntnis palaearktischer Hemipteren. Konowia. V, p. 218.

1931a. Hemiptera-Heteroptera. Abhandlung. der Pamir-Expedition 1928. VIII,

p. 77. (Russian and Latin text) p. 117 (German zoogeographical summary).

1931b. Hemiptera-Heteroptera of the Third Mount Everest Expedition, 1924. I.

Ann. Mag. Nat. Hist. ser. 10, VII, p. 362.

KiRKALDY, G. W. 1902. Plemiptera. Fauna Hawaiensis. III. p. 164. Cambridge,
England.

146 TF.RKESTKIAK FAMILIES OK 11 EMIl'TERA-IIETEROrTKR A

KiRKALDY, G. W. 1908. A List of the Described Heiiiiptera (excludinfj Aleyrodidac and
Coccidae) of the Hawaiian Islands. Proc. Hawaiian luit. Soc. I. p. 185.

LiNUROTH, C. H. 1931. Die Inscktfiifauna islands nnd ilirc I'roblcnu'. j). 1.^0. Inansj.-
Diss. Uppsala.

OsHANiN, B. 1891. The Zoogeographical Character ul the llcniipterous I'auna of Turke-
stan. Zapiski Russk. Geogr. Obsch. XXIII, p. 56 (in Russian).

1912. Katalog der Palaearktischen Hsniijiteren. P«erlin.

PoppR-s, B. 1909. Beitriige zur Kcnntnis iler Anthocoriden. .Act. Soc. Sci. Penn.
XXXVII, No. 9, p.' 1.

1912. Neue Miriden aus deni russischcn Reiche. Ofv. h'insk. \'etcrsk.-Soc. Forh.

LIV, A. N:o 29, p. 11.

Reinig. W'. F. 1932. Beitriige zur Faunistik des Pamir-Gehietcs Wiss. Frgeb. der Alai-
Paniir Expedit. Tl. III. Band 1. Berlin.

Reuter, O. M. 1884. ITenii])tera Gymnocerata luiropac. Act. Soc. Sci. Fenn. XIII, p. 1.

1885. Monographia .\nthocoridaruni orbis terrestris. Act. Soc. Sci. Fenn. XIV,

p. 555.

— -1908. Monographia generis Heteropterorum Pliiinodcra Germ. Act. Soc. Sci. Fenn.

XXXIII. No. 8, p. 1.

SiNGii-PRmii, H. 1925. The Morphology of the Male Genitalia in Rhynchota. Trans.
I'.nt. Soc. Lond. 1925, p. 127.

St.\l, C. 1872. Genera Lygaeidanini I'.uropae. Ofvcrs. Kimg. X'etensk. .\kad. Forh.
Stdckhnltn. 1872. N :o. 7. p. 37.

Explanation of Plate VIII.

Fig. 1. Phimodera rupshuensis sp. n.

Fig. 2. P. rupshuensis, anterior aspect of head.

Fig. 3. P. rupshuensis, ventral aspect of head.

Fig. 4. P. rupshuensis, anterior part of connexi\un).

Fig. 5. Micropla.v hissaricnsis Kiritsh.

Fig. 6. BiaiichieHa adelungi Reut.

Fig. 7. Nysiits thyini (Wolft'j. S , Kidwelly, Britain.

Fig. 8. A'', cricae ericae (Schill.). $, North Haven, Connecticut.

Fig. 9. A'^. cricae groenlandicus (Zett.). $, East Greenland.

Fig. 10. A'^. ericae ericae (Schill.). 5, Leh.

Fig. 11. A^. ericae afif. ohscuralus Horv. $, Renka-le.

Fig. 12. A'^. cricae alticola suhsp. n. $ , holotypc, Ororotse Tso.

Fig. 13. A'. cr/a7c alticola subs]), n. 9 , allotype. Ororotse Tso.

Fig. 14. A'. Iliyuii (Wolff). <5 , lateral aspect of paraniere, Kidwelly.

Fig. 15. The same, dorsal view.

Fig. 16. N. cricae cricae (Schill.). $, lateral aspect of paramere, North Haven.

Fig. 17. The same, dorsal view.

Fig. 18. A', cricae alticola suhsp. n. S , holotype, lateral aspect of paramere.

Fig. 19. The same, dorsal view.

Fig. 20. N. graininicola (Klti.). S, lateral aspect of paramere, Porto dTschia, Italy.

Fig. 21. The same, dorsal view.

MEM. CONN. ACAD., VOL. X.

PLATE VIIL

Explanation of Plate IX.

Dolmacoris deterrana gen. n., sp. ii.

Fig. 1. Dorsal aspect of $ .

Fig. 2. Lateral margin of abdominal sternites 2-7 and connexivum of tergites 2-4, partially
detached, to show arrangement of spiracles.

Fig. 3. Lateral aspect of head.

Fig. 4. Gonapophyses of 9 .

Fig. 5. \'entral aspect of head and prothorax.

Fig. 6. Genitalia of S . dorsal aspect. (W. F, China del.) hp. hr. hasal plate bridge, bp. basal
plates, ejy. ejaculatory reservoirs, phs. ]iliallosuma, ();;'. conjunctiva, I's. vesica, gnp.
gonopore, pr. ])aramerc.

Fig. 7. Paramere of $ .

Fig. 8. Seventh abdominal segment of o ■ ventral aspect.

PLATE IX.

Explanation of Plate X.

Fig. 1. Emblethis horvathiana sp. ii. Head and ]iro!i()tniii of type.

Fig. 2. Anthocoris gyalpo .sp. n. Head and pronutuni of type.

Fig. 3. Ectemnus paradoxus sp. n. Head and pronotuin of type.

Fig. 4. Anthocoris gyalpo sp. n. Aperture of inetathcjraeic scent-gland.

Fig. 5. A. gyalpo sp. n. Left paraniere.

Fig. 6. Ectemnus paradoxus sp. n. Wing cell.

Fig. 7. Ectemnus paradoxus sp. n, .\perture of inctathoracic .scent-gland.

Fig. 8. E. paradoxus sp. n. Left paramere.

Fig. 9. Dicyphus physochlaenae sp. n. Head of holotype.

Fig. 10. D. sengge sji. n. Head of tyi)e.

Figs. 11 and ]2. D. physochlaenae sp. n. Left paraniere.

Fig. 13. D. sengge sp. n. Left paramere.

Fig. 14. Tibetocoris margaretae gen. n., sp. n. Dorsal asjiect of head and ])ronotuni.

Fig. 15. T. margaretae gen. n., sp. n. Anterior aspect of head.

Fig. 16. T. margaretae gen. u., sp. n. Lateral asjjcct of head.

Fig. 17. T. margaretae gen. n., ,sp. n. Claws, aroliae and psendaroliae.

Fig. 18. T. margaretae gen. n., sp. n. Right jiaraniere.

Fig. 19. T. margaretae gen. n., sp. n. Left ])araniere.

MEM. CONN. ACAD., VOL. X.

PLATE X.

ARTICLE IX

REPORT ON ROTATORIA

By \V. T. Edmondson and G. E. EIutciiinson

I. INTRODUCTION

Tow-nettings and uther samples containing rotifers were obtained by the Yale North
India Expedition in fifty-two localities in the Punjab and Northwest Frontier Province (4),
the Kashmir basin (15 ), Indian Tibet and the extreme western part of Tibet proper (24) and
the Nilgiri Hills (9). From these collections we have succeeded in determining ninety-nine
species excluding a few doubtful forms. No new species were discovered, though it has been
necessary to bestow one specific and two varietal names. This paucity of new forms is prob-
ably due to the fact that the collections were made for the most part in alkaline waters
at moderate or low temperatures. The great number of new species recently obtained from
acid waters by Myers (1931-4) is well known; moreover it is probal)le that truly tropical
waters will be found to yield a number of remarkable new species when they are more
adecjuately known. At the same time as the work reported in this paper was in progress
one of us had the opportunity to examine a small collection from somewhat alkaline waters
at temperatures between 25.9-29.2°C. from the Island of Hispaniola, in which, though but
thirty-six forms were obtained, three of these represented very distinct new species
(Edmondson, 1934).

In preparing this report we have had invaluable assistance from Mr. Frank J. Myers,
who has confirmed a number of doulitful identifications and placed his great knowledge of
the group at our disposal whenever difficulties have arisen. Mr. Elbert H. Ahlstrom has
most kindly examined our material of the genus Brachionus and his determinations have been
included in the lists of species. He has also made several useful comments on species of other
genera. Dr. J. Wiszniewski has most kindly studied our material of the very interesting
species that he has recently described as Pcdalia biilgarica Wiszn. To these investigators
our sincerest thanks are due. We are extremely grateful to Professor Alexander
Petrunkevitch and Professor George Vernadsky for translating various descriptions in
Russian and Polish. We are also indebted to Dr. E. D. Merrill and the staff of the New
York Botanical Garden for determinations of water plants.

The types of the new forms described, and as far as possible a representative set of
slides of the known species encountered, will be deposited in the Peabody Museum of Natural
History of Yale University. ~ A second set of slides will be forwarded to Mr. M\'ers for
incorporatif)n in the collection of the American Museum of Natural History.

II. LIST OF I.nCAI.ITTKS

P. Localities in the Punjab and Nortlizvcst Frontier I'roinnce.

P 2. Sohawa, Jhelum district, altitude c. 528 m.

Large dam near village. Marginal vegetation chiefly a narrow-leaved Potaniogcton

Mem. Conn. Acad., Vol. X, Art. EX. September, 1934.

154 ROTATORIA

^ and cliarophytes. Temp. 10.45 a.m. (nvercast) 16.2"C., 4.30 p.m. (bright sun-
light) 21.2°"C. 3 March, 1932. P 2. (6) frum open water hetween weeds, P 2.
(7) from among weeds.

P 3 Sohawa, Jhelum district, altitude c. 528 m.

Small pond, about 2X5 m. and 50 cms. deep, in swampy stream-bed just below
P 2. Temp. 11.25 a. m. U'C. 3 March, 1932.

P 6. Sohawa, Jhelum district, altitude c. 528 m.

Pond about 18 X 18 m., shallow, very turbid, Jkiiciis in centre suggests that it is
temporary. Temp. 3.45 p. m. 26.5°C. 3 March, 1932.

P 13. llaripur, Abbotabad district. Northwest Frontier Province, altitude c. 550 m.

Small and presumal)ly very transient puddle by roadside, about 2-3 X 1 m., and
10 cms. deep. Masses of filamentous algae. 17 March, 1932.

K. Localities in the Kashmir Valley.

K 4. Srinagar, altitude c. 1585 m.

Small pond by office of I'^ish and Game Commission. \'ery tnrliid and tending to
polysaprobic condition. 21 March, 1932.

K 8. Swamp at Gagirbal, Srinagar, at lower end of Lokut Dal lake, altitude c. 1580 m.

Limnanthcmutn nyiiiphaeoidcs dominant plant. Diurnal temp, range, 6 Ajjril, 1932,
15.3-19.0'C.; pH range, 8.0-8.4. 5 April, 3 May, 1932.

K 19. Swamp at Gagirbal, Srinagar, altitude c. 1580 m.

Cut off from lower end of Lokut Dal by an embankment.

Dominant plant Ranunculus tricho phyllurn. Temp, range 6 April, 1932, 13.2-

23.0°C., pH range 8.3-9.3. 5 April, 3 May, 1932.

K21. Lokut Dal Lake, Srinagar, altitude 1582 m.

Shallow lake, about 1.3 m. deep, entirely carpeted with vegetation, Potamogcton
luccns being dominant, with P. amphibiioii, and Liimmntheiiiuiii nyi)ipliaeoidcs at
the margin. Temp, range, 6 April, 15.85-18.12°C. ; pH range 8.2-8.5. 5 April,
1932.

K 21a. Floating garden at side of west side of Lokut Dal, altitude 1582 m.

These are artificially constructed swamps, formed by anchoring masses of decaying
vegetation on sticks at the edge of the lake. Tomatoes and other vegetables are
cultivated on the islands so obtained. Small pools are cut off from the edge of the
main lake in this way and in one of these collections were ni.ade S May, 1932. A
full ecological account of stations K 8, K 19, K 21 and K 21a is in jneparation anti
will appear in a later paper in this series.

K 24. Nishat Bagh, altitude c. 1535 m.

Small artificial pond, 8 m. in diameter and about 50 cms. deep, muddy and with
undetermined Myriophy Hum-like plant. 7 April, 1932.

ROTATORIA 155

K 26. Suiidar Khun, altitude 1582 m.

A small lake flowing into the swamps in which lie the two Dal Lakes. Max. depth
5 m. Potaiiiogcton luceiis, P. pcctinatus, Ceratophyllitm demersum and charophytes,
the whole floor of the lake being carpeted with vegetation. Surface temp. 19.4,
pH 8.5. 2 May, 1932.

K 35. Phashakuri, near Pampur, altitude c. 1585 m.

A very large swamp in which the dominant plants are Potatiwgeton crispus and
Liniiianthemum nympliaeoidcs. Temp. 7 May, 1932, 19. IC.; pH, 10 April, 1932,
9.3, 7 May, 1932, 9.6. Collections 10 April, 7 May, 1932.

K43. Wular Lake, altitude 1573 m.

Large shallow lake, max. depth 6 m. but mostly under 3 m. Collections are from
plankton samples taken on the western side of the lake, April, 1932. Surface temp.
15.5-19.OX., pH 8.9-9.0.

K 46. Bakh Hajan, altitude 1584 m.

Large swamp of "jhil," Ijetween K 43 and K 48, mostly about 30 cms. deep, with
much RamiHculiis trichophyllum and Marsilia sp. pH 8.5. 17 April, 1932.

K48. Manasbal Lake, altitude 1584 m.

Lake with a maximum depth of 12.7 m. Surface temj). 19.85°C., pl^ 8.8. Collec-
tions 21 April, 1932, K 48 (1) epilimnetic plankton 7-0 m., K 48 (2) hypolimnetic
plankton 10-7 m.

K 51. Bod Dal Lake, altitude 1582 m.

Shallow lake; max. depth 4.0 m., but mostly under 2 m. deep with beds of
Potaiiiogcton crispus, etc. Temp, surface 16.4; pH 8.5. 1 May, 1932.

K 68. Sonamarg, altitude c. 2620 m.

Pond about a mile west of rest house, 20 x 10 m. and about 50 cms. deep. Masses
of filamentous algae. Temp. 9.00 a. m. 10 'C, pH 7.0. 19 May, 1932.

K 69. Sonamarg, altitude c. 2620 m.

Similar pond near K 68 with fresher looking algae. Temp. 9.30 a. m., 13.2, pH
7.6-f . 19 May, 1932.

L. Localities in Indian Tibet (Ladak, Rupshu, etc.) and Tibet Proper.

L 2. Mulbe, road from Kargil to Leli, altitude 3200 m.

Small ]>i")l behind Gonpa rock. About 4 m. in diameter, 50 cm. deep. I'"ilanienti>us
algae. Temp. 8.30 p. m. 7.0''C. ; pH 8.0. 27 May, l'>32.

L4. Spring beliiw I'hotho-la, pass on mad to Leh, between Mulbe and Lamayuru, altilude
c. 3960 ni. 28 May, 1932.

L 10. Khalatse, on road to Leh, altitude 2957 m.
Small pond. 30 May, 1932.

156 KOTATORIA

L 14. Si)itliu-, "U ihe Indus south of l-ch, altitude .i_'70 m.

Pool in a swampy meadow, Potamogcton pi-cliiiatus, IJlncithina sp. cliarophytes, etc.
pH 8.0. 4 June^ 1932.

I. 1(). .Spithug, altitude 3270 m.

Pool a little south of L 16, vegetation similar but l\'tiiiiinculi(s tn'cliopliylliiiii also
present. Temp. 1 p. m., 28.7°C. 9 June, 1932.

T. 32. Zung--lung, Ix-low Shakya-la, altitude 4224 m.

Algal growth in stream, temp. 2.30 p.m., 5.2, pil 7.V. 24 June, 1932.

L 3*^. Tsar Tso, IxHween Mugieb and Panggong Tso, altitude 4252.

Small shallow lake, lacking outlet, about 400 m. in diameter, almost cunipletely full
of a water-plant, probably P. pectimtus. Temp. 5 cms. below surface at edge,

10 a.m., 12.5''C.; pH ".3, 28 June. l')32.

L 40b. Lagoon 2 at west end of Panggong Tso, altitude 4241 m.

A long narrow pond shut off from the lake by a bar of sanil and shingle. Brackish,
chloride, 0.249 grms. per litre. 30 June, 1932.

L47. Lnng-yun, above Chagra, Northwest of Panggong Tso, altitude 4977 m.
Pools in swampy ground. 9 July, 1932.

L49. Togom Tso, northwest of Panggong Tso, altitude 5334 m.

9 July, 1932. This and the ne.xt two localities are discussed in detail Ijclow.

T, 50. Togarma Tso, northwest of Panggong Tso in the Ko-luiigpa valley, altitude 5217 m.

10 July, 1932.

L 52. Ororotse Tso, just south of Chang-chenmo River, altitude 5297 m. 12 July, 1932.

L 60. Kyam, Chang-chenmo valley, altitude c. 4725 m.

Largest of several pools fed by non-therinal .spring below terrace. 21 July, 1932.

L61. Kyam, Chang-chenmo valley, 24 July, 1932.

Pond in swampy hummock ground, about 5 m. in diameter, with Ranunculus
trichophyUiim.

L71a. T.so Nyak, Tibet, altitude c. 4250 m.

Visited l)y two Ladakis, Tzewang Tashi and .^niiam Tergas, wlm made littoral tow-
nettings in the western part of the lake. Though the water of the lake drains into
Panggong Tso it is probably slightly mineralised; a sample lirought back to camp
had a ])1 i value of 8.9. 12 August, 1932.

L 74. Tangur Tso, Tibet, altitude 4329 m.

Large lake lacking outlet studied at the west end, where the maximum tlepth was
9.5 ni. Littoral vegetation abundant, not flowering but probably P. pcctinatus.
Temp. 14.1-15.1, pH 9.6-|-. Collections 14 August, 1932.
L 74 (1) marginal, L 74 (2) open-water plankton.

ROTATORIA 157

L 72. Chushol, S. of Panggong Tso, altitude 4491 m.
Large pond south of village. 9 August, 1932.

L 72a. Chushol, S. of Panggong Tso, altitude c. 4491 m.
Small pool by L 72. 9 August, 1932.

L 7Z. Chushol, S. of Panggong Tso, altitude c. 4330.

Pond with large spherical algal colonies, N. of village. 10 August, 1932.

L 76. Mitpal Tso, Ijetween Chushol and the Indus Valley, altitude 4875 m.

Small lake lacking outlet, max. depth 24 m. Plankton from middle of lake, surface
temp. 12.49^C., pH 9.1. 17 August, 1932.

L 78. Yaye Tso, draining into Indus, altitude 4686 m.

Small lake, max. depth 18.1 m. Planktim from middle of lake, surface temp.
14.19 C, pH 8.7. 19 August, 1932.

L 82. Tso Moriri, Rupshu, altitude 4528 m.

Very large lake, with a max. depth probalily in excess of 50 m. Plankton from
northern part of lake, temp, surface 11.73, pH 9.0. 28 August, 1932.

L 82a. Estuary at Peldo-le, northern end of Tso Moriri, altitude 4528 m.

Temp. 14.9^C. Water much fresher than in lake. Consideral)le amounts of water-
weed, presumably P. pecfinatiis.

L 85. Sta-rtsak-puk Tso, Rupshu, altitude 4536 m.

Small very shallnw freshwater lake flowing into the salt lake Tso Kar. 4 Septem-
ber, 1932."

L 86. Tso Kar. Rupshu.

Shallow salt lake, abcjut 2 m. deep, chloride 11.66 gms. per litre, pH 8.9. 5 Septem-
ber, 1932.

L 86a. Pool by Tso Kar, Rupshu. altitude 4527 m.

Slightly brackish pool by salt lake, chloride .078 gms. per litre, pH 9.2. 5 September,
1932. '

N. Localities in the Nilgiri Hills, Madras Presidency.

N 2. Ootacamund, altitude c. 2255 m.

Pond on Marimund Rd., about 50 m. in diameter, blue waterlilies and Utrieularia.
Temp. 16.1-17.0 C, pH 6.6. 7 November, 1932.

N 3. Ootacamund, aUitude c. 2255 m.

Small pond in swamp above N 2, about 13 m. across, temp. 16.5. 7 November, 1932.

N 5. Ootacamund, altitude c. 2201 m.

Lake near the town, very turbid owing to recent rains, small amount of l)lue water-
lily. Temp. 17.5, pH 6.6. 8 November, 1932.

1 38 ROTATORIA

N 6. botacaiiiunil, altitude c. 2280 m.

I'und at tiiurth milestone on Connemara Rd.. about 2 x 3 ni. and 20 cms. deep, in
swampy ground, temp. 22.7'C., pH 6.3. 9 Novemlier, 1"32.

N 8. Ootacamund, altitude c. 2195 m.

Pond about 15 X 6 m. and 1 m. deep, nn Pykara Ud., bc-yond cemetery, temp.

21.3 C, pll 7.3. 10 Noveml)er, 1932.
N 13. "Umbrella Tree" near Ootacamund, aliitude c. 2316 m. Small pool in swamp,

2 X 1.5 m. and c. 60 cms. deep. Iltricularia .-ind I'lIamcntDUs algae. Temp, midday,

22.1X., pli (1.1. 18 November, 1932.

N 15. Ootacamund, altitude c. 2195 m.

Pool 3X8 m. and 1 ni. deep, full of a lanccolate-lcavcd Potamogcton. pll 6.4. 13
November. 1932.

N 17. Ootacamund, altitude c. 2316 m.

Dam towards Government House, c. 30 X 70 in., Icmj). 13.7 C, pll 6.3. 15 Novem-
ber, 1932.

N20. Pykara, altitude c. 2133 m.

Littoral plankton from large artificial lake. Temp. 19.5^C., ])TT 6.8. 16 November,
1932.

III. ENUMI'.KATION OF SPRCIES

In general Ilarring's Synopsis has been accepted as indicating the standard nomenclature,
save in a few cases where later investigators have found it necessary to modify his
terminology.

The symbol c. after ;i statinn number indicates that the species in question was common,
vc. that it was very coinnidu ; in all other cases it ma}' be assumed th;it but a lew individuals
were found.

Actinunis ncptunius Ehrenberg, P2 (7) c, P 3, K 48.
Anuraeopsis fissa (Gosse), P2 (6, 7).
Ascomorpha cucaudis Perty, K 35, K 51.
A. sal'atts Bartsch, L 72.
A. sp. L 13.
Asplanchna brightzvelli Gosse, K 26, K 35, K 43 vc, K 51. L 14, L 16, N 20.

A. priodonta Gosse, K 8, K 21, K 43, K 48 vc., K 51.
Asplanchnopus iiiulticeps Schrank, L 73.

Bdelloids, indet., P 2 (7), K 8, K 19, K 21a, K 24, K 35, K 46. K 51. I, 2. L 4, L 13, L 14.
L 32, L 47, L 49, L 60, L 71a, L 72, L 72a. L 73, L 82a. L 86a, N 2, N 3, N 5, N 8, N 15.
Bracluoims annularis Gosse, P 2, K 43.

B. calyciflorus Pallas, P 2, K 19, K 35, K 43 vc, K51c

B. capsidiflorus Pallas, P 2 f. quadridcntatus Hermann, L 71a f. rhcnaniis Lauterborn, L 74
f. clitniorbicidaris Skorikov, f. ciizi'i Franc: and transitional forms, I. 86a f. cnzii France.
B. furculatus Thorpe, K 4 (form).

ROTATORIA 159

B. plicatilis Miiller, L 40b vc, L 71a vc, L 86 c. L 86a.
B. quadratus Rousselet (== ? leydigii Colin), P 6.

B. sericus Rousselet, K 4.

Cephdoddla catdlina (Miiller), ? K 35, L 74 (1) vc. f. iahlstromin.

C. cxigna (Gosse), ? P 2 (7), ? P 6, K 35.
C. forfictda Ehrenberg, P 2 (6), P 6.

C. gibba (Ehrenberg), K 8, K 21, K 21a, K 35, K 44, K 51, L 16, L 72, L 73, L 74, L 86a.
C. panarista Myers, P2 (6).
Cephalodella wiszniewskii n. n. L 86a.
C. sp. P 2, K 4, K 35, K 44, K 51, L 74, N 13.
Collotheca campanidata (Dobie) K 8.

Colurella bicuspidata (Ehrenberg) P 2 (6, 7), P 3, K 8, K 21, K 21a, K 35, L 14, L 74, N 5,
N8.

C. colurus (Ehrenberg), K 35.
Conochilus hippocrcpis (Schrank), K21.
Cupclopagis z'orax (Leicly) L21.
Dicranophorus myriophylli (Harring) L 72.
Diurclla brachyura (Gosse), P2 (7).

D. cavia (Gosse), L 14, L 16.
D. stylata Eyferth, N 2, N 19.

D. zueberi Jennings P 2 (7), K 8, K 19, K 21, K 21a, K 34, L 14, L 16, L 73.
Eosplwra najas Ehrenberg, L 60.

Epiphancs brachionits (Ehrenberg), L51.

E. senta (Miiller) K 4, K 8, K 34.
Enchlanis alata Voronkov ? P 2 (7).

E. (Dapidia) dcflexa Gosse, K 8, K 34, K 51.

E. dilatata Ehrenberg P 2 (6, 7), P 13, K 8, K 19, K 21a, K 34c, K 46c, K 51c, K (.8, K 69,

L 14c, L16c, L39, L 73.
E. mcncta Myers P 3, K 21a, K 24, K 35, K 51, L 14, L 16, L 60, L 72.
E. pan'a Rousselet, P2 (6, 7), P3, K21a, K 24, K 34, K 46, K 51c, L 14c, Lol, L 73, L 82a.

E. triquetra Ehrenberg P2 (7).

Filinia longiseta (Ehrenberg) P 2, K 43, K 46, K 48 (2) vc, K 51, L 16, L 49c, N 2, N 5.

F. terminalis (Plate) N 2, N 5.
Floscidaria conifera (Hudson) ? K 8.
Flosculariid, indet. P2 (7).
Gastropus Iiyptopus (Ehrenberg) K 4.

Itura aurita (Ehrenberg) P 2 ,(7), P 3, L 86a.

Keratella cochlcaris (Gosse) K 8, K 24, (f. cochlcaris c, f. carinata (Levander) one speci-
men) K48 (1, 2) vc, K51, L 52, L 76 vc, L 78.

K. qimdrata (Miiller) f. divergens ( Voigt) w^ith transitions to f. qiiadrafa K 8, K 24, K 35c
46c, K 48, N 5, N 6, N 10, N 17, N 20, f. fre7t::cli (Eckstein) K 26 vc, f. quadrata and
f. tcstiido L 76c, f. quadrata, short spines trans, ad divergens L 82, the same with f.
valgoides n. L 71a.

K. z>alga (Ehrenberg) f. valga, P2 (6), f. tropica Apstein P2 (6), N 5, f. tropica-
asyiiimctrica Barrois and Daday K43, f. tropica-moiistrosa I5arrois and Daday K 43c.
Lccaiie aciilcata Jakubski, P 3.

160 ROTATORIA

A. flcxiirs (Gosse), N 5.

L. horncvianni (Ehrenljerg) K 19.

L. lima (Miillcr) P2 (6, 7), P 3, K 8. K21, K 26, K ,U, L 14, L Ibc, L71a, L 73c.

L. ohionisis (Ilerrick) P2 (7).

L. papuana (Murray), P2 (6, 7)c.

Lepadclla acuminata (lihrenberg) N 5, N 15.

L. oralis (Miilier) K 24, L 73.

L. patella (Miiller), P 2 (6, 7), vc, I' 3, P 6, K 19, K 21, K 21a, K 35 vc, K 43, K 51, L 13,

L14, L16, L61, L74(l), L 86a.
L. rhomboidcs (Gosse), P 2, N 5.
L. triptcra Ehrcnberg, P2 (6), K 24, K 35, N 13.
Lophocaris oxystcrnon (Gosse), P2 (6, 7), L 86a.
L. salpina (Ehrcnberg) P2 (6).
Monostvia bulla (Gosse) P 2 (6, 7) vc, P 3 vc, K 8, K 19, K 21a, K 35, L 14, L 16, N 2, N 5,

N 13, N 15.
.1/. clostcrocerca Sdnmirda. P2 (6), P2 (7) vc, P 3c, P 6, K 8, K l'>, K21, K21a, K 24,

K 35, K 43, K 46, K 51c, L 14, L 72, N 5, N 8, N 15.
.1/. crcnata Harring, P2 (7), K 8, K21a, K 35, N 6.
.1/. hamata Stokes, P2 (7), K 19, K21a, K 35, N 5.
M. lunaris (Ebreiiberg) K21, 1.71.

M. quadridentata (Ehrenlierg) K 1*', K2!, K 35 vc, K 43, K 51.
M. stenroosi Meissner P 3.

Mytilim mucrormta (Muller) P2 (7), K 8, K 19, K21, K 35, vc, K43, K 51.
M. trigona (Gosse) L61, L 72 (long form).
M. vcntralis (Ehrenlx^rg) P2 (6, 7) f. 7'cnfrali.'! and f. Iirrvispiiui l*".lircnb(.-ri;'. I\ 8 lung imst.

spines, K 21a, long post, sjiincs, K 34 c. f. rv»/r(//L\- ami f. fcn-r/.v/'/Hcr, K 48, f, brcri.'^piihi.

L 16 f. brcvispina.
Notholca striata (Miilkri, i\51, 1\ 60, L71a, L 73, L 74, L 86a, all reduced f. striata

(= scaph ula Stewart ) .
Notomniata copcns Ehrenl)erg L 16.
N. epaxia Harring and flyers, P 2 (7), L 86a.
A^. trifnis Ehrcnberg, P 2 (7).

Notommatids, indet. K 24, K 35, K 51, L 10, L 14, L 72, L 72a.
Pcdalia bulgarica Wiszniewski L 50, L 73, L 74 (2).
P. mira (Hudson) K 19, N 5.
Polyarthra euryptera Wierzejski K 48 vc.

P. trigla Ehrcnberg P 2 (6), K 8, K 1^, K 21 c, K 24 vc, K 43 vc. K 7^. K 85 vc
Platyias patulus (Muller), K 34, N 5.
P. quadricornis (Ehrcnberg), K 8, L 14.
Pompholyx sulcata Hudson, L 82a vc.
Proalcs decipicns (Ehrcnberg), P2 (6).
Ptygura sp., K46.

Scaridium longicandtim (Miiller), P2 (6, 7), P 3.
Squatinella mutica (Ehrcnberg), P2 (7). L 76.
Synchaeta littoralis Rousselet, ? K 51.
5". oblonga Ehrcnberg, K 51.

ROTATORIA 161

S. pcctiiiata Ehrenberg, K 8, K 19, K 21, K 21a, K 26, K 43, K 46, K 48 ( 1) vc, K 51 c, N 5.

5". stylata Wierzejski, K 43 c, N 5.

S. tazina Hood, K 43, ? L 72.

S. tremida (Muller), P2 (6, 7), K 43 c, K51, ? N 19.

5-. sp., K 35, L 14.

Tcstudinclla incisa (Ternetz), N 2, N 8, N 13.

T. viucronata (Gosse), K 35.

T. patina (Hermann), P2 (6, 7), P 3, K 8, f. patina and f. intcnncdiata Anderson, K 19,

K 21, K 24, f. intcnncdia, K 35 c, K 51.
Trichocerca cristafa Harring, P2 (7).
T. cylindrica (Imhof), K 43.

T. elongata (Gosse), K 8, K21, K21a, K 26, K51.
T. iernis (Gosse), K 8.

T. longiscta (Schrank), K 19, K 35, L 14, L 16, L 72.
r. rattus (Muller), P 2 (6, 7) c, P 3, K 8, K 21, K 35, K 69, L 14, L 16.
T. scipio (Gosse), P2 (7).

Trichotria pocilliim (Muller), K 19, K 21, K 51, L 14, L 16, L 72, L 73.
T. tctractis (Ehreberg), P2 (7), K 8, K 19, K21, K 21a, K 46, L 72a, L 73, N 5, N 8.

IV. TAXONOMIC AND ZOOGEOGRAPHICAL NOTES

1. Ct'plialodcUa catcUina ( Muller ).■* Several allies of this species, distinguished pri-
marily by the relative length and shape of the toes, appear to exist. There can be little doubt
that the form figured by Ehrenberg, who gives the first recognisable illustration of the species
(1838, T. LV, fig. iii) is the same as that figured ^by Hudson and Gosse (1889, Plate
XIX, fig. 10a) and by Harring and Myers (1924, Plate XXVII, fig. 3). If Ehrenberg's
reference of his Diglcna catclUna to Ccrcaria catcUina Miiller determines the significance of
the latter, which otherwise would be quite unrecognisable, it is clear that the form of the
species with long slender straight toes about one-sixth of the tutal length must be regarded
as typical.

On the other hand Weber has figured, as Diglcna catellitia a form which is not only
somewhat larger than the typical form as figured by Harring and Myers, Init has a propor-
tionately shorter and basally much broader toe. Though this form is perhaps less widespread
than the true C. catcUina, it apparently is found in the New World as well as in the Old,
for Mr. Myers informs us (in Hit.) "after our paper was published, . . . Harring found l)oth
forms of C. catcUina and it was at that time decided that we should have described both."

' After the present paper had gone to press, we saw, through the kindness of Mr. Myers, part of the proof of
a paper by Dr. Wiszniewski, in which this species is discussed. Dr. Wiszniewski has shown that calclVina Weber
differs from catcUina Harring and Myers, not only in sh:ipe and in the form of the toe, but in possessing salivary
glands. These structures appear to be present in our volvocicolous form and absent in f. ahlstroini. We have
attempted, with a minimum alteration of our text, to bring our arrangement into line with that of Dr. Wiszniewski.
We find it impossible, however, to agree with him that the true catcUina is the species figured by Weber, but
have much pleasure in associating the name of this distinguished Polish investigator with such an interesting
species. Were it not for the fact that catellina is the genitype of Cephalodella, it would probably be better to dis-
card the name and use for the form figured by Harring and Myers, the new name introduced by Dr. Wiszniewski.
The posterior position of the foot in Miiller's figure makes it certain that whatever species he may have had
before him, it was not either of the present forms.

162 ROTATUKIA

We have uhtained a single specimen of this form from sample L 86a, and I)elieve it to lie
sufficiently distinct to merit recognition; we therefore propose the name C. wiszniewskii n. n.

In addition to these forms, two other very close allies of the species have l>een described.

Dii^lcna rok-ocicola Zavadovski (1916), for which the author proposed, should it prove
to be but a variety, the unnecessary alternative D. catcUina parasitica (ncc Plciirotrocha para-
sitica Jennings = C. parasitica Ilarring and Myers), is a form living in colonies of J'oli'ox.
We have unfortunately been unable to obtain the original description, but according to
Harring and Myers the figures that accompany it do not show "any differences that luight
be considered of specific value. Some physiological dissimilarities are described, the most
striking one being the parasitism of the animal in Wilvox colonies." Hutchinson, Pickford
and Schuurman (1932) record from inside colonies of Vol^-ox in two shallow fresh-water
pans a form determined by them as C. catclliiia. Re-e.xamination of some of this material
shows that it represents a form of Cephaloddla in some respects intermediate between C.
catellitui and C. ri'iszniewskii, resembling tlie former in the narrow l)ase of the toe in proportion
to its length, and the latter on the other hand in the concavity f)f its anterior margin, though
this is more marked in the volvocicolous form.

Recently de Beauchamp (1932) has described a very large form with a short curved toe
from the plankton of Lake Elmenteita in Kenya. This form he regards as a species Cepha-
lodella elmenteita because the preserved specimens on which it was based lacked the character-
istic eye-spots of C. catclliiia. We have, however, been unable to oliserve eye-spots in any
specimens of forms of C. catellina preserved in formalin and then cleared in glycerine, and
therefore believe the difference to be illusionary. Moreover, in the marginal sample from the
alkaline Pangur Tso we have o!)tained a form of C. catcUina which has toes \vhich though
straight show a basal constriction as in C. elmenteita. Our Pangur Tso form, therefore,
seems in one of its characters to be transitional to de Beauchamp's species. For this Pangur
Tso form with its basally constricted toe, -wt propose the name f. ahlstromi, Mr. Ahlstrom
who observed some speciiuens of the form in a tube of Brachioniis material having pointed
out to us the desirability of examining the form more closely. Mr. Ahlstrom believes the
Pangur Tso form to have a shorter and in preserved specimens more plicated head than
is normal, but we are not convinced that these differences are not entirely due to presers^a-
tion. Intergrades l^etween f. catellina and f. ahlstromi appear to occur, for of throe s])eci-
mens of catellina from Los Angeles, California (det. Myers), kindly lent us for study by the
American Museum of Natural History, two specimens have a typical toe, and the third a
definitely constricted toe as in ahlstromi, the toe in this specimen is however slightly longer
(body 120/i, toe 2U'.) than in the latter form, in this resemlding the specimens of f. catellina
with which it occurred.

The various forms of C. catellina may be tabulated as follows :
C. catellina (Miiller). Length 110-135^, to.' 20-25^, about one-fifth or one-sixth of the
total length, long, narrow, anterior margin practically straight, widest at the practically
unconstricted base. (Figttre 1 c.)

f. ahlstromi n. Length 95-112/u, toe 15-18'/, just under one-sixth nf the total length.
long, narrow, slightly concave both anteriorly and posteriorly, straight, basally ctmstrictcd
and widest distinctly distally to base. (Figure Id.)

ROTATORIA

163

C. wiszniewskii n. n. Length 1S)0 (type), 100-140/x (Weber), toe 21/^, about one-ninth of the
total length, short, wide basally, anterior margin very slightly concave, widest at the uncon-
stricted base. (Figure la.)

f. volvocicola (Zavadovski). Length 126/*, toe 19/t, about one-seventh of the total length,
moderately narrow basally, anterior border strongly concave, widest at the practically uncon-
stricted base. (Figure lb.)

C. clmcntcita de Beauchamp. Length 210/*, toe (from figure) 18/t, short, narrow, strongly
curved, the anterior border concave, the posterior convex, basally constricted and widest
distinctly distally to base. (Figure 1 e.)

Figure 1. — E.xternal aspect of left toe of A, C. wiszniewskii n. n. ; B, C. iv. f. volvocicola (Zavadovski) ;
C, C. catclUna (Miiller) ; D, C. c. f. ahlstromi n. ; E, C. clmcntcita de Beauchamp, redrawn from
de Beauchamp.

The tropin are essentially similar in all these forms, the characteristic asymmetrical
development of a tooth on the distal edge of the right ramus is particularly well marked in
South African specimens of f. z'olz'ocicola and in the East African C. litiicntcita.

2. Filinia loiigiscfa (Ehrenberg) and F. terminalis (Plate). In plankton samples
from the lake at Ootacamund two forms of Filinia are not uncommon. One of these is a
typical limnetic form of F. longiseta, the other is a form with a small very narrow spindle-
shaped body and completely terminal insertion of the posterior seta. No intermediates occur
between the two forms in these samples. The form with the terminal insertion of the pos-
terior seta was met with in South Africa by Hutchinson, Pickford and Schuurman. Some
of their material was kindly examined by Mr. D. Bryce, who pointed out that this form is
referable to F. terniinalis (Plate), a species which has been rarely found in Europe and
which was placed in the synonymy of F. loiigisctahy Harring. Unfortunately Plate (1886)
gives no figures but there is no reason to doul)t that Mr. Brycc's determination is correct and
as the two forms are frequently found together in South .Africa as well as in the Ootacamund
locality, without any intermediate forms occurring, there seems no reason why F. feniiinalis
should not be accorded specific rank. The speci;s may be easily recognised by its small narrow
spindle-shaped body and by the insertion of the posterior seta ( Figure 2c) as pointed out above.

It is probable that the chief reason for the almost universal failure to recognise
F. tcrmitialis is to be fmuid in the great varialjility of the distance l.)etween the posterior end

164

ROTATORIA

i)f the T)ody and the insertion of the posterior seta in /■". loii^i^isi-ta. Sloniiuski ( 1926) has
shown that in Europe tliis species exhibits a considerable amount of cyclomorphosis and
that, in the autumn and winter, forms occur in which the distance Iwtween the posterior end
and the insertion of the seta is reduced to 7/*, these forms are called tcniiiuaiis by Slominski
who consequently regards the latter as Init a form of longiseta. It is however to be noted
that his measurements show that these winter "fcniiinalis" forms are actually shorter and
wider than the summer forms in which the insertion of the seta is markedly ventral. It is

B

I'^iiiURE 2. — A, Filinia longisela (Ehrenberg), Ootacamund Lake, NS; B, /-'. loiigiscia Togom Tso, 1-49; C,

F. tcrmwalis (Plate), Ootacamuiul Lake, N 5.

clear therefore that our form, which we lielieve to be the true Icniiiualis. is a quite different
organi.sm to Slominski"s form with a sul)-terminal insertion.

Our material from Kashmir ( W'ular Lake and the hypi>lininion of L. Manasijalj and
from the ice-covered Togom Tso in Indian Tibet has a terminal or sub-terminal insertion
(Figure 2 b), while from the Ootacamund Lake the specimens have a typical ventral insertion.
Measurements of specimens taken at random from these localities are given in Table I. It
will be seen that normally longiseta has a lx>dy al)out twice as long as deep, while in fcniiiiiaiis
the body is about two and a half times as long as deep. The material of the former from
Togom Tso however is about as proportionately long as fcniiinalis but comparison of the
figures of the two forms will show (Figure 2 1), c) that even such elongate specimens of
longiseta retain a more giblwus dorsal profile than is found in tenninalis; moreover Slomin-
ski's data, as pointed out alx)ve, indicate that elongate forms of longiseta are only found in
populations of very large specimens, having a much greater total length than terniinalis, so
that they do not constitute a real transition between the two species. On the basis of the

ROTATORIA

165

relative length of the anterior setae our Togoni Tso specimens may be considered as typical
longiscta, while our other specimens are transitional to f. lUnnctica (Zacharias) :

Length
F Hi Ilia longisela

ToGOM Tso 230

ToGOM Tso 240

WuLAR Lake 138

WuLAR Lake 133

WuLAR Lake 146

WuLAR Lake 142

L. AIanasbal 150

L. Manasbal 150

OOTACAMUND 171

OOTACAMUND 208

Filinia tcrininalis

OoTACAMUND 138

OoTACAMUND 138

OoTACAMUND 142

TABLE I

dimensions

in fi)

Distance between

Dorso-

insertion of post.

ventral

R. Ant.

L. Ant.

Post.

seta and apex of

Depth

Seta

Seta

Seta

body

90

480

455

360

8

100

475

490

395

4

79

400

380

262

1

71

410

410

290

71

420

< • .

237

8

71

382

390

233

4

83

420

380

325

4

83

450

450

325

4

83

590

590

341

21

100

595

595

305

25

54

342

300

242

50

330

330

262

58

308

333

242

3. KcratcUa qiiadrata (MuUerj and Kcmtclla vdga (Ehrenberg). In spite of the
valuable contributions of Hartmann (1918) anil others, there still appears to be considerable
uncertainty as to the status and correct designation of the various forms included by Harring
(1913) under K. qiiadrata (Miiller). After examining extensive material from South
Africa and India we are fully convinced of the correctness of the contention that two species,
A', qiiadrata (=aculeata Ehrenb.) and K. valga, are to l)e recognised. Moreover, the
nomenclature of these two species in their typical forms raises no difficulties.

Amiraca aculeata Ehrenberg (1832) is expressly stated by its describer (1838) to be
identical with Brachionus quadratus Miiller. Ehrenberg's best figure (1838, T. LXII,
fig. xiv, 1) may therefore be taken in conjunction with those of Miiller (1786, T. XLIX,
figs. 12, 13) as defining the typical form of qiiadrata. The posterior margin of the lorica
in Ehrenl>erg's figure, measured across the outer margins of the bases of the posterior
spines, is wider than the anterior margin similarly measured across the bases of the antero-
lateral spines. The posterior spines are parallel (Miiller) or very slightly divergent, al)out
two-fifths as long (Ehrenberg) or half as long (Miiller) as the lorica. Both reticular and
punctate sculpture are clearly indicated by Ehrenberg. Miiller's figure is so unsatisfactory
that in fixing the typical form it is best to abide by the excellent illustration given by
Ehrenberg, save that it is convenient to regard typical qiiadrata as having practically parallel
posterior spines, as is indicated by Miiller.

166 ROTATORIA

Ariuraea Z'olga Ehrenherg (1834) maybe regarded as typically represented by Ebren-
berg's best figure (1838, T. LXII, fig. xv, 1), which shows a form in which the posterior
width of the lorica is distinctly narrower than the anterior. The right spine is alx)ut one-
third, the left one-quarter of the length of the lorica. Reticulate and punctate sculpture are
1)1 >th clearly indicated.

In examining as much material and as many illustrations as are available to us we have
rarely had any difficulty in assigning individuals or illustrations to one or the other species.
It is true that in a few figures of European specimens (e.g. Virieux 1916, fig. 46, Wesenberg-
Lunil, 1930, Plate VH, figs. 8, 26, and 46) the aiitcrinr and posterior borders are equal
or the former very slightly longer than the latter. But such specimens do not apjiear to
show the pronounced narrowing of z'alga and it must be remembered that a very minute
error in drawing on the part of an observer not studying the dimensions in question would
produce just such differences as found in some of these figures. Fadeev (1927) used the
relation between the anterior and posterior widths of the lorica to separate tropica Apstein
from the other forms of K. qiiadrata (s. lat.), apparently understanding b}' Apstein's name
what we here regard as K. ralga. Within these species the use of misapplied varietal terms
appears to have led to considerable confusion. Thus in his monumental work on the cyclical
phenomena in rotifers Wesenberg-Lund (1930) objects to certain of Hartmann's conclusions
as to the production of nalga forms from resting eggs. But it is clear from a study of
Wesenberg-Lund's figures that what this author calls 7'alga is a form of qiiadrata (s. s.)
while Hartmann's observations on this point referred primarily to the true /v. t'a/T'a (Ehren-
berg). Since such confusion is bound to result from the present unsatisfactory state of
the varietal nomenclature of these two species we have attempted to standardise as far as
possible the names of the various forms, introducing a minimum of new names and adhering
to the principle of priority. Though the latter is not binding in such cases, it would ajipear
to provide the best method of determining the relative merits of two synonyms, and the
neglect of the princijjle in the past has led authors to create new varietal names without
an adequate study of the literature so that the form of 7'alga with a single posterior spine
has been provided with at least three and probably four names.

Kcratclla qiiadrata (Miiller).

Kratzschmar (1908, 1913), Hartniann (1918) and Wesenberg-Lund (1930) have
studied the morphological cycle in this species. Normally the first generations from se.xual
eggs are composed of forms with long divergent spines (f. diz'crgcns Voigt) ; these later give
place to forms with shorter more parallel spines (f. qiiadrata Miiller). Later one (f.
valgoides n.) or both spines (f. curvicornis Ehrenberg) are lost and at such periods sexual
reproduction is stated to occur. The amount of cyclomorphosis is very variable in different
localities but in general this species exhibits a cycle of reduction. In a few cases an initial
elongation of the spines has been observed and in a series called by Hartmann "A. aculcata-
valga," but consisting of apparently morphologically normal reduced forms of K. qiiadrata,
the cycle begins with ciiri'icornis, to which the addition of one minute papilliform spine pro-
duces the form named by Jakubski (1915) irregularis, and two minute spines tcsludo
Ehrenberg.

In general the two posterior spines of qiiadrata are subequal in length, but in certain
forms unequal spines apparently indicate a transition from diz'crgens (right spine) to qiiadrata
(left spine). Fadeev has figured as valga a specimen of qiiadrata with unequal subparallel

ROTATORIA

167

Figure 3. — KcralcUa quadrala (Miiller), dorsal aspect of lorica of A, B, f. divcrgcns (Voigt) Phashakuri,
K35; C, f. divcrgcns Ootacamund Lake NS; D, f. jrcnzcli (Eckstein), Sundar Khun, K26; E, f. quadraia
trans, ad divcrgcns, Tso Nyak, L71a; F, f. valgoides n, Tso Nyak, L71a; G, f. quadrata, Mitpal Tso, L76; H,
f. tcstudo (Khrenberg), Mitpal Tso, L 76.

Spines but this may best be regarded as a transition to valgoides, though Wesenberg-Lund's
figures and our collection from Tso Nyak indicate that the latter form may probably appear
in a quadrata population without intermediate forms with unequal spines. The following
names are probably sufficient to describe the chief members of the cycle and to designate
the varieties present in single collections.

KcratcUa quadrala (Miiller).

Diagnosis : Six anterior spines, reticulate sculpture forming a median series of undivided
hexagons, maxinmiu breadth of lorica slightly greater than the posterior breadth, the latter

168 RdTATORTA

greater^than the aiitrrinr Ijrcacltli, twn posterior spines wliich arc iisualiy suliequal, or (Hie
asyninietrically i)lace(l spine, or without spines.

f. platci ( J;igerskir)kl). Posterior spines long, strongly divergent basally, bent ruund so
thai apically thev lie at right angles to the long axis of the body. An e.Kiiberant development
of f. divergetus from the Baltic.

f. dh'crgcns (Voigt). Posterior spines long, more than half the length of the lorica
(excluding anterior spines), subequal and strongly divergent, reticulate and punctate sculp-
ture well developed. Syn. var. longispina Thiebaud (1911).

f. frcnzcH (Eckstein). Posterior spines more than lialt' the length of the lorica. Sub-
equal and somewhat sinuate, but with parallel axes. Sculpture practically absent.

f. qiiadrata (Miiller). Posterior spines alxmt half the length of the lorica, su1)equal,
and subparallel, reticulate and punctate sculptare well developed.

f. testudo (Ehrenberg). Posterior spines short, about one-seventh of the length of the
lorica, often somewhat divergent. Punctate and reticulate sculpture both present.

f. bmnspina (Gosse). Posterior spines short and equal as in testudo, often somewhat
divergent. Punctate sculpture absent.

f. valgoides f. n. Right posterior spine well developed, left absent. Punctate sculpture
absent. Syn. z'alga Fadeev 1927, p. p. Wesen1>erg-Lund 1930 etc., nee. Ehrenberg.

f. irrcgtilaris (Jakubski). One posterior spine (right in tj'pical figure) absent, the other
represented by a minute papilla.

f. curvicornis (Ehrenberg). No posterior spines, punctate sculpture absent.

Of the other forms described and probably referalile to this species, Auuraca squaunda
Ehrenberg is clearly a very reduced curuicornis without reticulate sculpture. However, it is
very doubtful that this is really the same as Brachionus squariuda Miiller, the anterior
spines of which suggest a very round reduced form of Nothoka striata, in all pr(jbal)ility the
use of this name is superfluous. A. falculata Ehrenberg and -i. qiiadridentata Ehrenberg
are doubtful forms which probably need not be recognised.

In the Nilgiri Hills and in most localities in Kashmir (Figure 3 a, b) moderately devel-
oped forms that have rather divergent spines and may be regarded as transitional between
the typical form and f. (/;'trr^r«.y are common. Well-developed examples (Figure 3 c) of
the latter are found in the Ootacamund Lake (N 5). In Sundar Khun (K26), however, a
very well developed unsculptured form with long parallel posterior spines, which may be
referred to f. frcnzeli, is abundant (Figure 3d). No reduced fonns were found in any
samples from these regions.

In the high lakes of Indian Tibet qiiadrata forms with moderately long spines, which
in the specimens from Tso Moriri and Tso Nyak in 'ril)et are fairly divergent ( l'"igure 3 e),
are found. In Tso Nyak the reduced f. valgoides (Figure 3 f) also occurs sporadically, while
in Mitpal Tso a form (Figure 3 h) referable to f. testudo is found though less commonly
than is f. quadrata. The specimens from Tso Nyak, and to a less degree from Tso Moriri,
are feebly sculptured, while the Mitpal Tso form which is rather wide and gibbous dorsally
has well developed sculpture and slightly convergent posterior spines. It is probably unwise
to attempt to correlate the occurrence of reduced forms in these cold elevated lakes with
ecological conditions without more knowledge of the seasonal cycle, but their occurrence may
be of significance.

ROTATIIIUA 169

Tlie occurrence (if /\'. quadrata in the Nilyiris is of interest for the species is probably
absent in most truly tropical waters, occurring only within the tropics in elevated localities
under temperate cimditions such as are afforded iiy the aquatic iiabitats arnmid Ootacamund.

Kcratella valga (Ehrenberg).

In true K. valga the cycle (Klausener 1908, Hartmann 1918) appears to involve pri-
marily the addition of spines. The initial phase in the most complete cases is the form
called by Klausener K. curviconiis f. brchiiii. This form is figured as being somewhat longer
than K. quadrata f. cun-'icornis. In certain irregularities in the line of the posterior margin
of this figure there is, moreover, perhaps a hint of the position of the spine bases, demark-
ating a typical valga posterior margin, though this may be accidental. Fadeev (1927) has
described and figured as f. aspina a comparable form which is certainly clearly referable
to valga on the shape of the posterior margin alone without having to take into account the
other members of the cycle. An essentially similar elongate form (Figure 4 a) was recorded
from South Africa by Hutchinson and by Hutchinson, Pickford and Schuurman ( 1932)
as curvicornis. Klausener distinguished f. brehtiii from curvicornis by the fact that the
antero-median spines do not diverge in the former. This character is probably too vari-
able and in general does not separate valga from quadrata. It must be admitted that the
two species must often Ije hard to separate in their most reduced forms.

In Hartmann's studies of this species the most exuberant form was one in which the
two posterior spines are subequal. This form seems to be the one figured by Schmarda
( 1850, Plate IV, fig. Ill) as A. longicornis but it has doubtless been regarded by other authors
as actileata (i.e. quadrata s. s.).

Apstein has described another exuberant form from Ceylon as tropica. This form is
characterised by the very long and unequal posterior spines, the left being about two-fifths
the length of the lorica without the anterior spines, the right about six-sevenths of this
length. This form, in spite of Hartmann's statement that it agrees well with liis form of
May, 1915 (f. 7'alga), differs from his figure of the latter in which the right spine is liardly
more than one-half the length of tlie lorica. It is clear from the work of Tschugunoff
(1922) and Fadeev (1927) that forms essentially similar to tropica are common in South
Russia, and Skorikov as long ago as 1896 figured (T. VIII, fig. 29) such a form from
near Kharkov. Jaku1:iski (1915) has figured a monospinous tropica form from Poland and
in South Africa, Hutchinson (1930) and Hutchinson, Pickford and Schuurman (1932)
fcnind tropica to be the only common form of ixilga.

In all probability the most developed form of I'alga varies from place to place, and if
f. valga and f. longicornis represent the high;st development in some European localities,
tropica probably does the same in Ceylon, S. Russia, South Africa and Kashmir. The names
proposed for forms of I'alga in which the left spine is very reduced or al)sent may lie used in
conjunction with the name tropica in localities where the right spine is very elongate, and
where tiiese reductions occur, e.g., K. valga f. tropica-asyininctrica and K. 7'olga f. tropica-
motistrosa.

The following terms may then be applied to designate the forms of A', zxilga.

Keratella valga (Ehrenberg).

Diagnosis: six anterior spines, reticulate sculpture forming a medial series of undixided
hexagons, maximum breadth of lorica very distinctly greater than the posterior breadth, the

170

ROTATORIA

latter niHch less than the anterior breadtli, two posterior spines which are usually unequal, or
one asyniinetrically placed spine, or without spines.

f. c'iilga (Ehrenberg). Posterior spines well developed, rather sliort, unequal, the riyht
typically one-third and the left one-fourth of the length of the lorica. Punctate and reticulate
sculpture both present.

f. asx)iiiiictric'a (P)arrois and Daday). Right posterior spine well developed, typically
about half as long as the lorica, left spine rudimentary. Punctate and reticulate sculpture
both present. Syn. f. heterospiiia Klauscner (1908) p. p.

lOOy.

Figure 4. — Kcralclla ralga (Ehrenberg), dorsal aspect of lorica of A, f. aspina (Fadecv) Weltevreden
West, Tvl. S. Africa; B, f. valga, Sohawa, P2; C, f. tropica (.-Xpstein) Ootacamund, NS; D, f. iropica-
asymmctrica Apstein-Barrois and Daday, Wular Lake, K43; E, f. tropica-iiwiistrosa Apstein-Barrois and Daday,
Wular Lake, K43.

f. monstrosa (Barrois and Daday). Right posterior spine well developed, typically alxjut
half as long as the lorica, left spine absent. Reticulate and punctate sculpture well developed.
Syn. ? var. duinasi Richard (no figure or indication if this is really a form of valga)
var. asymmetrica Daday nee. Barrois and Daday
f. nionospina Klausener (1908)
i.monospUia Fadeev {\927 )^ tropica-motistrosa

i. longieornis (Schtnarda). Both posterior spines well developed and practically equal,
typically about three-fifths as long as the lorica, reticulate and punctate sculpture both well
developed, as indicated in Schmarda's typical figure, or the latter reduced (Hartmann, 1918,
Figure 76).

f. tropica Apstcin. Rigiit spine very long, typically about si.x-sevenths of the lengtli of
the lorica, left spine much shorter but well developed, about two-fifths as long as the
lorica. Reticulate sculpture developed, punctate often reduced.

f. reducta Fadeev. Right posterior spine alone developed, short, about one-fourth of tlie
length of the lorica (typical figure may be taken as Fadeev, 1927, T. 2, fig. 12), punctate
sculpture absent, reticulate very reduced.

ROTATORIA 171

f. brchmi Klausener. Both posterior spines absent, reticulate sculpture present (Brehm
and Zederbauer, 1904, fig. 1 ), punctate apparently absent.

f. aspina Fadeev. Both posterior spines absent, reticulate sculpture almost and punctate
entirely absent.

If Fadeev is correct, in South Russia aspina is to be regarded as the extreme reduced
phase of tropica, i.e. tropica-hreluni, and rediicta an intermediate between tropica-mofisfrosa
and this extreme reduced phase. More knowledge is clearly needed of the cyclomorphosis of
tropica forms before it is certain that the extreme reduction in sculpture noticed in aspina is
really characteristic of this series of forms. In the only South African locality that has been
followed throughout an entire year, Florida Lake, near Johannesburg (Schuurman, 1932),
the species appears stable, but a spineless unsculptured form which we would refer to aspiiia is
recorded from certain shallow pans, as has been already indicated. In the present collec-
tions a single specimen which may be referrd to tropica (Figure 4) occurred in the plankton of
the lake at Ootacamund, in which K. quadrata f . diz'crgens was abundant. A few very similar
specimens occurred in the pond at Sohawa designated as P 2, where a single short subequal
spined form with a pentagonal posterior polygon (Figure 4) here referred to f. z'alga was
also obtained. Both the Sohawa and Ootacamund K. valga f. tropica have well-developed
punctate as well as reticulate sculpture. In Wular Lake a very long spined form which may
be referred to tropica-nwnstrosa (Figure 4) was not uncommon and in the same locality a
very few specimens of f. tropica-asymmctrica were also obtained (Figure 4). The punctate
sculpture in these Kashmir specimens is considerably reduced.

It is clear from Jakubski's (1916) record and from some of Klausener's data that the
asymmetric condition does not always involve reduction of the left, but sometimes also of
the right spine. It is highly probable that truly dextral and sinistral forms may occur, with
a concomitant difference in the sense of their spiral swimming movements. Such a possi-
bility is of great biological interest and would merit close attention on the part of any worker
to whom living material is available.

Certain names applied to forms of K. quadrata and K. valga or included under the
former species by Harring remain to be considered.

Anuraca scutata Thorpe (1891) from Brisbane appears to be allied to K. valga f. asym-
mctrica but such sculpture as is indicated seems to show the median cariniform arrange-
ment of the mid-dorsal reticular partitions characteristic of K. cochlcaris (Gosse). The
dorsal surface is said to be markedly gibbous in lateral view. In view of the doubt raised
by the sculpturing the name scutata Thorpe is iiest suppressed unless an animal identical with
the figure remains to be rediscovered.

Anuraea procurva Thorpe (1891) from the Island of Ascension is in form nearer to
quadrata than zvlga but is distinctly asymmetric in its posterior spines. In side view the
lorica is seen to be bent forward, particularly in its ventral part, in a most peculiar manner.
This form is probably best retained as a somewhat doubtful species under the name of
Keratella procurva (Thorpe).

Anuraca stipitata var. Warlnianni Asper and Heuscher (1889) is another problematic-
form but there seems no reason to treat it as a synonym of K. quadrata f. curvicornis, as
is done by Weber (1898).

172

ROTATORIA

TABLE II
(All dimensions in /i)

Max. Ant. Tost. R. Post. L. Post. Median Dorsolat.

Lcngtli Breadth Breadth Breadth Spine Spine Ant. Spine Spine

A', quadrata
SuNDAR Khun

f. frenccli 134 96 77 92 125

121 100 75 92 100

129 104 79 96 133

142 98 79 94 108

Phashakuri
trans, ad

i. divergais 132 92 75 90 83

138 96 75 92 75

OOTACAMUND

f . diver gens 116 79 58 7?, 75

121 83 60 77 79

TSO MORIRI

trans, ad

f. divcrgens 121 92 67 79 46

121 94 69 83 42

Tso Nyak
trans, ad

f. divergens 125 90 67 77 48

i. valgoid es 118 90 69 77 54

MiTPAL Tso

f. quadrata 121 96 67 75 42

125 96 69 75 42

f. festudo 108 90 63 71 17

K. valga

Ootacamund

f. tropica 112 71 58 46 85

So HAW A

f. vali^a .*.... 100 67 58 46 23

i. tropica 116 75 67 50 75

Wl'lar Lake

i.tropica-asyimuctrica .. 129 71 60 50 108

i.tropica-monstrosa 138 77 58 54 120

.... 131 68 54 46 108

Banagiier Pan 2,
Transvaal.
{.tropica 104 79 68 54 96

Frisc h gew a agd,
Transvaal.
{.tropica 102 90 75 50 108

Weltevreden

West, Transvaal.
f . aspina 106 73 60 c. 42

108

83

134

117

38
46
33
42

29
29
27

25

79
58

38
38

25
25

67
71

?,2,
29

21
25

42
40

25
33

21
21

46

38
33

29

25

38
38
17

25
25
25

17
21

10

42

54

50

25

37

46

31

21

19

23

17

21

27

23

25

29

25

25

29

29

29

21

31

17

ROTATORIA

173

Anuraca aculcata var. cochlcaris Voigt is, according tu Carlin-Nilsson (1934), identical
with K. paludosa (Lucks).

Two forms recently described by Athanassopoulos (1930) as Anuraca aculcata var.
graeca and var. conica, both appear to belong to the genus Brachionus, in the figures of both
forms what appears to be a foot-shield is indicated; the former is apparently B. capsuliflonis
i. quadridcntatiis Hermann, the latter bears some resemblance to B. sataniciis Rousselet.

FiGUUE 5.— Map of the world showing the distribution of Lccanc f^apiiaua (Murray) in relation to tlie mean

annual isotherms for 15° C.

In Table II measurements of specimens of the two species under discussion, from vari-
ous localities in India, Tibet and in South Africa are given.

4. Lecane papuana (Murray). This species was originally described from New
Guinea. Harring and Myers ( 1926) record it from Panama, Guatemala and Polk County,
Florida, and Ahlstrom (Myers in litt.) has also taken it at Miami in Florida. Tarnogradsky
(1930) reports the species from the North Caucasus and Wiszniewski (1931) from near
\'alencia, Spain. Hutchinson, Pickford and Schuurman (1932) met with it rather fre-
quently in a number of locahties in the Transvaal. In the present collection it occurred at
Sohawa in the Punjab. If these records are plotted on a map of the world (Figure 5) on
which the mean annual isotherms for 15"C. are drawn, it is seen that they fall on or within
these isotherms, while the countries whose rotatorian fauna is best known lie for the most
part outside them. There can be little doubt, therefore, that the present species is a sub-
tropicopolitan form, providing what appears to be the most conspicuous case of such a
distribution yet recorded among the Rotatoria.

1 74 ROTATORIA

\'. ROTATORIA I'RKVIOUSLY RECORDED EROM INDIA AXD TIBET

In the present section we have collected tog;ether all the previous Indian and Tibetan
records of Rotatoria, and have revised these lists in accordance with the now generally
accepted nomenclature of Harring ( 1913). All species of doubtful validity have teen omitted.

The earliest Indian list is that of Anderson (1889) who studied the rotifers in the
vicinity of Calcutta. The following list of thirty-seven species cnm])riscs all those recorded
by him, the nomenclature being standardized as indicated above.

CoUotlicca oniata (Ehrenberg)

C. caiiipauulata (Dobie)

C. aiiihigua (Hudson)

C. tcnuilohata (Anderson")
*FlosciiIaria rini^ciis (Schrank)

Lhnnias ccratophylli Schrank

L. vicliccrta Weisse

Beauchampia crucigera (Dutrochet)

Ptygura stcphanion (.Anderson)

Simiittcrina socialis (Linnaeus)

PhilodUia citrina Ehrenberg

Rotaria rotatoria (Pallas)

R. macroccros (Gosse)

R. mcnto (Anderson)

Actinurm ovatiis Anderson
*Notommata tripus Ehrenberg
*CcphalodcHa forficula (Ehrenljerg)

Monoiiunafa orbis (Miiller)

Dicranophorus forcipatus Miiller

DhircUa tigris (Miiller)
*Scaridiiuit longkaudiini Ehrenberg

SqiiatincUa tridcnfata (Fresenius)
*Mytilina vcntralis (Ehrenberg)
*M. ventraJis breiispina (lihrenberg)

Etichlanis iiiacnira Ehrenberg
*Lecanc lima (l-^hrenberg)

Monostyla cornuta (Miiller)
*ilf. quadridentata Ehrenberg
*M. bulla Gosse

Cohirclla caitdata ( I'.hrcnberg)
*LcpadcUa ozvlis (Ehrenberg)
*L. triptcra (Ehrenberg)

L. chrcbcrgii (Perty)
*Tcstiidiiu'lla patina form iii/cniicdia (Anderson)
*Brachioniis capsidiflorus Pallas

Brachionus urceolaris Ehrenberg
*Platyias patitlus (Miiller)
*P. quadricontis (Ehrenberg)

The species preceded by an asterisk are to be regarded as wide spread, having occurred
in our collections also.

ROTATORIA 175

Murray (1906) has listed the following rotifers from the slopes of the Himalayas
between altitudes of 2000 and 8000 feet (610 and 2440 m.). Most of these rotifers are
Bdelloids, which is to be expected since the collections were made in moss.

Philodina indica Murray

P. sqiiaiiiosa Murray

P. citrina Ehrenljerg

P. brevipcs Murray

P. flaz'iccps Bryce

P. vora.v Janson

P. laticeps Murray

Habroirocha perforata (Murray)

H. angiisticollis (Murray)

H. angiisticollis attcmtata (Murray)

H. nodosa (Murray)

H. aspcra Bryce

H. lata Bryce

H. Icitgcbii (Zelinka)

H. inicroccphala (Murray)

Macrotrachcla forinosa (Murray)

M. qiiadricornifera Milne

M. papulosa Thompson

M. tmdtispinosa Thompson

M. plicata (Bryce)

M. habit a (Bryce)

M. bidlata (Murray)

M. muscnlosa Milne

Rot aria sordid a (Western)

R. sordida fimbriata (Western)

R. rotatoria (Pallas)

Adincta vaga (Davis)

Proales quadrangularis (Glasscott)

Squat inclla tcnclla (Bryce)

Colurclla adriatica (Ehrenberg)

Monostyhi litnaris (Ehrenberg)

BracliidiiKs urccolaris Ehrenberg

Our Kashmir stations are comparable in altitude to Murray's localities, but since our
collections were made with a tow-net, his list is of no value for comparison.

The rotifers of Southern Tibet have been studied by Stewart (1908), who collected 17
species, including five that he described as new, from the neighbourhood of Gyantse, at alti-
tudes between 13,000 ft. and 14,000 ft., i.e. approximately 4000 m. and 4270 m. Of the
five new species Mastigocerca auchinlcckii Stewart is synonymised by Harring (1913) with
Trichocerca longiseta (Schrank) and Salpina shape Stewart with Mytilina centralis brezispina
(Ehrenl). ). Rotifer tridentatiis Stewart is considered unrecognisable by Harring as is
Cathypna auibaii Stewart by Harring and Myers ( 1926) . Notholca seaphida Stewart, omitted
through an oversight by Harring, is an obvious synonym of A^. striata (Miiller). The Tibetan

176 KOTATORIA

list is further rt'clucctl l>y tlie union of Proalcs gibba Ehrenh. and Jh'aschica sciiiiapcrta Gosse
under the name of Cephalodella aHriculata (Miiller) (Harring and Myers, 1924). The
following;" list, tlierefore, gives all the valid species recorflcd 1)y Stewart:

J'liiludiiia crytlioplilhalnia l'",lirenl).

1'. roseola Ehrenh.

P. citrina Ehrenl).

Notoimiiata aiirifa (JMuller )
*A'^. copeus Ehrenh.

Cephalodella auricidala (Miiller)
*C. catcllina (Miiller)

C. exigna (Gosse)

Scaridiuin longicaiiditin ( IMiiiler)
*'rricJwcerca longiscta (Schrank)
*Trichotria pocilluin (Miiller)
*Mytilina z'entralis brei'ispina (Ehrenherg)
*Euchlaiiis dilatata l^hrenherg
^Xotholca striata (Miiller)

An asterisk indicates that the species in question was obtained also from our collections
from W'estern Tibet.

.\ndersou's and Murray's lists alone refer to territory which is within the boundaries of
the Indian Empire. Taken together sixty-seven species are recorded by these two authors, of
these, sixteen were found in our collections. In the latter, therefore, eighty-three species are
fnund, nut hitherto recurded fr(im India, and the total Indian list is brnught up tu une hundred
and fifty. When it is remembered that Ahlstrom (1933 J has recorxled one hundred
and nineteen species from a single embayment of Lake Isrie, it becomes clear that the
Rotatoria t)f India are still extremely little known and ctTer a prumising field for furtiier
investigation.

VI. THE ROTATORIA^ FAUNA OF HIGH AFTITUDES

The only previous collection of Rotatoria from the higher parts of the Himalaya is that
described by Stewart, whose recognisable species are enumerated above. No other collection
from over 4000 m. appears to have been hitherto reported. Smirnov (1930) has enumerated
14 species and 2 varieties frnni the Pamirs but his collection was made apparently between
3700 and 3900 m.

In our material 42 determinable species are recorded from 18 separate localities in
Indian Til)et between 3500 and 5334 m., while from 12 localities in Kashmir, lying between
1580 and 2667 m. we record 58 species. It would aiijiear, therefore, that a slight decrease
in number of species occurs in ])assing from the lower to the higher localities. Further
analysis brings this out much more clearly. Of the 42 species recorded from Indian Tibet
33 were fountl in 9 localities (of which 3 were alkaline) below 4500 m. and 22 species were

ROTATORIA 1/7

confined to this zone. The 9 locahties lying alcove 4500 m. may best be considered in three
groups.

4500-4600 (3 locahties)

Brachionus plicatilis

B. capsuliflorus f. ensii
Ceplialodclla gibba

C. wiszniewskii
Ifiira aurita
Kcratella quadrata
Lophocaris oxysternon
Notholca striata
Notonitnata epaxia
Polyarthra trigla
Pompholyx sulcata

4600-5000 m. (3 localities)
Eosphora najas
Euchlanis mcneta
E. parva

Kcratella quadrata
K. cochlcaris
Lepadella patella
Mytilina trigona
Notholca striata
Polyarthra trigla
Squatinella niutica

5000-5334 m. (3 localities)
Filinia longiseta
Kcratella cochlcaris
Pcdalia bidgarica

It would seem that the increasing rigor of the environment with increasing altitude
plays a considerable part in reducing the rotatorian fauna but that in the region studied the
limiting factors do not become very intense until an altitude of about 5000 m. is attained.
The localities above that altitude may therefore be profitably examined more closely.

Togarma Tso, altitude 5217 m. (Figure 6).

Three small ponds lie close together in the wide valley that ends to the North in the
pass called Ororotse La, above the Ororotse Tso. The smallest pond is about 30 m. long,
15 ni. wide and 30 cms. deep. Both the largest and smallest ponds yielded Pcdalia bidgarica,
but no other rotifers were obtained. The water at 11.00 a. m., 10 July, 1932, had a temper-
ature of 16.2°C. but during the night probably fell to about freezing point. The chloride con-
tent was less than 0.0005 N., the alkali reserve (methyl orange titration) 0.0012 N. and the
pH 8.9. In the largest of these ponds an abundant copepod and cladoceran fauna occurred,
and much Spirogyra, forming brick-red masses, but no other rotifers were obtained in
tow-nettings. The poverty of the rotatorian fauna is emphasized by the fact that these
ponds superficially resemble the ponds at Chushol from which a considerable number of
species are recorded.

Ororotse Tso, altitude 5297 m.

178

ROTATORIA

Figure 6. — The group of ponds known collectively as Togarma Tso. The localities for Pcdalia bulgarica are the
small pond indicated by the arrow, and the large central pond.

Figure 7. — Togom Tso.

ROTATORIA 179

This lake is a small fresh-water lake in a "kar" at the head of a small tributary of
the Chany-chenmo River. The lake has a maximum determined depth of 14 m. When
visited 11-13 July, 1932, it was covered by a sheet of ice with a maximum thickness of
about 1 111. which was melting around the edges and from below. Most of the water was
at about 4.0°C., falling to 1.25 below the surface of the ice. A single specimen of Kcratdla
cochlcaris was noted in a vertical haul made from 13.5 to the surface. An extended study
of the lake will be given in a later paper.

Togom Tso, altitude 5334 m. (Figure 7).

This very small lake, lying between Togarma Tso and Chagra, was visited on the
afternoon of 9 July, 1932, when it was found to be almost entirely covered with ice. A
small belt of free water at the edge varied in temperature from 0°C. against the ice to
9.5°C. at the extreme margin. A few specimens of undeterminable bdellofds, one perhaps
Dissotrocha aculcata var. tiiberculata, and of FiUnia longiseta. were obtained in this marginal
water. The chloride content of the latter was less than 0.00005 N., the alkali reserve 0.0003
N., the pH 7.3.

Although many lakes in the western part of Tibet apparently lie at about the altitude
of these three it is doubtful if any habitats capable of supporting planktonic or other swim-
ming rotifers exist much above 5500 m. On the other hand, Heinis (1910) has shown
that in the Alps the muscicolous fauna extends to 4000 m. so that it is reasonable to sup-
pose that bdelloid rotifers e.xist in the Himalaya at altitudes of over 6000 m.

In considering the limiting factors determining the existence of organisms at very high
altitudes, it is clear that many which apply to terrestrial plants and animals cannot affect
aquatic forms. Thus terrestrial organisms may theoretically be limited by low temperature,
low oxygen tension, perhaps intense ultra-violet radiation, low CO^ tension in the case of
plants and some animals with a complex respiratory mechanism, and in the case of animals
deficiency in fuud supply. As will be pointed out in a later paper the oxygen tensions in the
high-altitude lakes examined, owing to their coldness, lie within the values frequently found
in surface waters in low-lying temperate countries. The penetration of ultra-violet light
into water is slight (cf. Carter and Beadle, 1930, and some unpul^lished observations made
on this expedition) . It may be of importance in the surface layers of water at high altitudes,
but much less so than in the case of terrestrial habitats. The available COo content of natural
waters is largely regulated by the quantity of alkali carbonate in the water, while so far as
the food supply of animals is concerned, evidence available as to the productivity of Ororotse
Tso suggests that this lake compares favourably with lakes at much lower altitudes (Hutchin-
son 1933). It seems, therefore, that temperature is the most important limiting factor in
the ecology of the high-altitude members of such a group as the Rotatoria.

Sufficient is known of the high-altitude rotatorian fauna of Europe to justify some
comparison between that fauna and the present collection. From the monumental work of
Zschokke (1900), the papers of Brehni and Zederbauer (1904) on the Tyrol and Monti
(1906) on the Italian Alps and from the catalogue of Swiss Rotatoria by Weber and Montet
(1918), it is possil)le to prepare lists of the rotifers of the Central European Alps. Zschokke
indeed gives such a list of 65 valid, fully determined species from over 1450 m., and by
inclusion of later records this list is raised to 108. The limits chosen by Zschokke, however,
include the upper part of the forested zone ; if only the region above the forest line, from
1700 ni. upward, be considered, the list is reduced to 89 species. Such a list has indeed

180 ROTATORIA

been given by Pesta (1929) in his valuable work on the high mountain lakes of the Alps,
but since it is in need of a few minor corrections, is not according to the standard nomen-
clature now universally used and is not arranged by zones, it seems desirable to present the
data critically in full. The altitude after the name of each species gives the highest station
recorded. No species is given in a lower zone if it also occurs in a higher zone.

2700-4000 m. (nival zone)
Adincta vam 3800
Macrolrachcla chrcnbcrgi 300C
Mniohia magna 4000
]\I. scarlatina 4000
Plcurctra alpiuni 4000 m.

2300-2700 m. (subnival zone)

Asplanclina priodonta 2453 ni.

Bracliionus iircciis 2350

Ccphalodclla gibba 2340

Clironiogastcr oralis 2306

Conochilus unicornis 2359

Dicranophorus forcipatus 2600

( Diitrclla sp. 2375)

Euchlanis dilatata 2630

Pilinia longiscta 2400

riabrotrocha torquata 2686

K. q. Volga 2350

LepadcUa patella 2400

Macrotrai-hcla plicata 2440

Monommata longiscta 2344

Monostyla htnaris 2600

Notliolca longispina 2640

N. striata 2600

Fcdalia bvlgarica ? 2630 (2200 in Ihilgaria, Wis/nicwski, 1933)

Pliilodina citrina 2600

P. erythophthalma 2445

Poly art lira trigla 2600

Proalinopsis caiidatiis 2313

Rotaria citrina 2445

7?. rotatoria 2550

Syncliaeta pcctinata 2307

Trichoccrca carinata 2350

(Trichocerca sp. 2640)

1700-2300 m. (alpine zone)
Adincta gracilis 2028
Bracliionus calyciflorus 2000
Cephalodclla aiiriculata c. 2000
C. ez'a 1938
Collotheca ornata 1810
C. deflcxa 2000
C. uiicinata 2144
C. 7Jinssnie7vskii 1825
Dicranophorus uncinatns 2028

KUTATORIA • jgj

DissotrocJia aculcata 1796
D. macrostyla 2048
DiiircUa sejiinctipcs 1874

D. tigris c. 2000
Embata parasitica 2189
Epi plumes bracliioiius 1725

E. senta 2093
Eosphora najas 2102
Eothinia elongata 2102
Euchlanis niacnira 2144
£. triquctra 2048
Floscularia mcUccrta 2100

F. ringcns 2000

Gas tr opus stylifcr 1920

Habrotrocha angu-sticollis 2000

/:/. bidetis 2100

H. munda 2087

Keratella cochlearis 2189

7v. quadrata f. quadrata 2270

K. serrulata 2189

Lecane luna 2189

Lc pad ell a ovalis 2000

Maerotrachela multispinosa 1900

Mniobia symbiotica 1950

Mytilina mucronata spinigera 1782

iW. j»wfica c. 2000

M. ventralis breznspina 2200

Notholca foliacea 2102

Noiommata aurita 2189

A'', pachyura 2102

TV. /n>?Zf 2000

Philodina roseola 2200

P. wm.r 2000

Pleurotroclia petromyzon 2000

Proalcs decipieiis 2000

Ptygura crystallina 2000

Rhiiioglcna frontalis 1800

Rotaria macrura c. 2000

i?. jorfl^iWa 20000

7?. tardigrada 1938

Squatinclla inutica 2000

Stcphanoceros fimbriatus 2144

Tcstudinclla patina 2000

Trichotria poeilliiin 1815

Trichoccrca hui'^iseta ]<134

r. ra.//;/.?2189 '

Zschokke (1900) in his analysis of his earlier list brings out clearly a limitation of
tlic number of species with increase in altitude, which limitation is supported by the later
list, given above. In the nival zone only muscicolous bdelloids occur, no rotifers being
recorded from the few lakes above 2700 m. that have been studied biologically in the Alps.
Nine species are, however, known to occur between 2600 and 2700 m. Zschokke gives a

\g2 KOTATOUIA

little evidence that the liniitatit)ii of the algal llora in the higher lakes is one of the factors
involved, hut considerahly more data are required to suhstantiate tliis.

It appears clear from the results of all investigators that the conmionot pelagic and
semipelagic rotifers of the high Aljjine ponds and lakes are Conocliiliis unkoniis, Euchlanis
dilatafa. Xofholca longispina and Polyarthra frigla. Zschokke concludes that the Alpine
rotatorian fauna is composed of widely distributed common species, a fact further empha-
sized by the list reproduced above. Pcdalia bulgarica appears to be the onl)- exception to this
generalisation.

In no other part of the worUl have any relevant data been assembled. All the lakes
studied at high altitudes in N. America appear to lie lielow the timberline. Bryce (1931)
records 13 species from the sacred lake on Mt. Za(|uala in .\byssiiiia at an altitude of 2700 m.
(9000 ft.), but this locality clearly enjoys a temperate climate, and the same is true of Lake
Titicaca at an altitude of 3800 m. (12,500 ft.), from which Murray (1913) obtained ii
species.

In comparing our list with that of the Rotatoria of the central European Alps it is
necessary to establish some sort of correlation between the ecological zonation in the two
areas. This raises considerable difficulties owing to the fact that the whole of Indian Tibet
enjoys a semi-arid climate and within the llimalaxan front-range no true forest occurs.

The nival zone begins in the region studied at about 6000 m. Above 5000 m. larger
lakes such as Ororotse Tso and Togom Tso a])parently remain frozen for almost, if not
quite, the entire year. It is probably correct to regard the zone lying between 5000 and
6000 m. as the equivalent of the upper part of the subnival zone of the Alps. Thickets of
Salix sp. exist in sheltered valleys, such as the Nyagtsu valley North of the Panggong Tso,
up to 4600 m., but such rare and isolated thickets can hardly be regarded as representing the
upper limits of the forest zone of the Alps. Cultivation is carried on up to 4524 m. at
Phobrang, northwest of Panggong Tso and up to aljout 4540 ni. at Korzok on the shore
of Tso Moriri. The fields at the latter settlement are probably the highest cultivated land
in the world, but Francke (1914) gives evidence of former higher cultivation in this region.
It must, however, be remembered that owing to the poverty of the country and the isolation
of its communities agriculture is carried on under circumstances that would be economically
unremunerative in luirope. It seems, therefore, reasonable to suppose that the Al])ine zone
of the Alps is represented by the zone above 4500 m. though its upper limit is uncertain.

In comparing oiu' list with that of the .\lps the data from above 1700 in. may therefore
be legitimately used, but the bdelloids and the attached forms (except Conocliiliis) must be
omitted, as our methods of study inevitably preclude their recognition. If this omission lie
made, the Alpine list is found to consist of 58 species, or just under three times the numl)er
recorded from the supposedly corresponding zones in Indian Tibet. When it i< rcmemljered
that the iVlpine list is the result of over fifty years' work by several investigators while our
list represents but a single season's collecting by one individual, it becomes prolialile that the
rotatorian fauna of Indian TilK't is at least as rich as that of the higher zones of the
mountains of Central Eurojic.

Of the 21 species recorded in our list, 11 species arc found in the Alps above 1700 m.
and 7 species alx)ve 2300 m. Unfortunately these numbers are small ; an attempt was made
to determine if there is any correlation between the maximum altitude records of these
species in the Alps and in Indian Tiljet; the vahie of the correlation coefificient obtained, viz..

KOTA-IOUIA 183

0.19, is (juite without signilicaiice wlien derived Irmn eleven pairs of ol)servations. While
it is clear that there are considerable similarities between the two faunae, evidence not sus-
ceptible to statistical treatment strongly indicates that there are also differences l^etween
them, lor it is to be noted that of the four commonest Alpine free-swimming rotifers only
one, Polyarthra trigla, occurs in our list for over 4500 m., and that Notholca longispina,
perhaps the commonest Alpine species, and Coiiocliilus uiiicoiiiis, were found nowhere in the
regions studied.

With the exception of Pcdalia bitlgarica, all the species that we record have previously
l)een found at about sea-level in widely distant countries, and in general our list supports
Zschokke's conclusion that the .Vlpine rotatoria are predominantly eurytopic species of
immense vertical and horizontal distribution. Bearing in mind the differences that we have
just noted Ijetween the fauna of the Alps and that of Indian Tibet, it is safe to conclude
that the rotifer fauna of very elevated waters represents, with the single exception of P.
bulgarica, a remnant of the common, and in general eurytopic, fauna widely distributed
throughout the earth, but that the composition of this remnant differs from place to place,
such differences l^eing perhaps in part due to chance and in part due to ecological factors,
of which latter the occurrence of Brachionus plicatilis provides an extreme example.

Pcdalia bulgarica has been mentioned so often throughout this discussion that a few words
as to its ecology may be appropriate. The species was met with in three localities.

As indicated above, a number of well-preserved specimens were obtained from two of
the Togarma Tso ponds; these were determined by Dr. W^iszniewski. A single specimen
was obtained in one of a similar series of ponds at Chushol (altitude 4336 m.) and several
very poorly preserved specimens from the open water of Pangur Tso (altitude 4329 m. )
were found to be undoubtedly referable to this species on account of the absence of posterior
appendices and the six teeth of the uncus. None of these waters were very cold at the time
when the material was collected. At Togarma Tso there was undoubtedly a great diurnal
variation in temperature, but in Pangur Tso, the temperature lay between 15.1°C.,
(surface) and 14.2'C. (9 meters) on 13 August and 14.1°C. (surface) and 13.8"C. (9
meters) on 14 August, so that in this locality the species must l)e continually exposed to a
moderate temperature of about 14°C. in the summer. The record of the species in Pangur
Tso is also of interest as indicating that it is tolerant of considerable alkalinity (alkali reserve
0.0610 N., chloride 0.022 N., pH c. 9.6). These facts are of interest as indicating that
this species, the only recorded alpoliiont rotifer, is more tolerant of diverse conditions than
might be expected.

Osborn Zoological Laboratory of Yale University,
23 July, 1934.

1^4 UOTATORIA

BlllLlUGKAPIlY

AuLSTROM, E. II. 1933. A Quantitative Study of Rotatoria in Terwilligcr's Pond, Put-
in-]'.ay, Oliio. Ohio Biol. Surv. Bull. 30, in Ohio State Univ. lUill. 38 (5). pi.

Anderson, H. li. 1889. Notes on Indian Rotifers. J. Asiat. Soc. Beni;-. 58. j). 345.

Apstein, C. 1907. Das Plankton ini Colombo-See auf Ceylon. Zool. Jahrb. Abt. Syst. 25.
p. 201.

AsPER, G. and Heuschkr, J. 1889. Zur Naturgeschichte dur Alpenseen. Ber. St. Cal-
lischen Naturwiss. ' Ges., St. Gallen (for 1887-1888), p. 246.

Athanassopoulos, G. 1930. Sur deux formes nouvelles de Anuraea aculeata Ehrb.
variete groeca et var. conica. Bull. Soc. Zool. Fr. 1930. p. 476.

Beauchamp, p. de. 1932. Report on the Percy Sladen Expedition to some Rift Lakes in
Kenya in 1929. iii. Rotiferes des Lacs de la Vallec du Rift. Ann. Mag. Nat. Hist,
(sen 10), 9. p. 158.

Bakkois, T. C, and Daday, E. 1894. Adatok az Aegyptonii, Palaestinai es Syriai Rota-
toriak ismeretehez. Math. Termesz. firtes. Budapest. 12. p. 222.

Brehm, V. and Zederbauer, E. 1904. Beitr.ige zur Planktonuntersuchung alpiner Seen.
Verb, zool.-bot. Ges. Wien. 54. p. 48.

Bryce, D. L. 1931. Report on the Rotifcra: Mr. Omer Cooper's Investigation of the
Abyssinian Fresh Waters. (Dr. Hugh Scott's l'"xpedition.) Proc. Zool. Soc. Lon-
don, 1930, p. 865.

Cari.in-Nilsson, B. 1934. Uber einige fi'ir Schweden neue Rotatorian. Ark. Zool. 26A,
n: o 22. p. 1.

Carter, G. S., and Beadle, L. C. 1930. Reports of an Expedition to Paraguay and lirazil
in 1926-27. The Fauna of the Swamps of the Paraguayan Chaco in Relation to
its Environment. I. Physico-Chemical Nature of the luivironnient. J. Linn. Soc.
(Zool.). London. 37. p. 205.

Daday, E. von. Az Anuracidac Rotatoria-csalad revisioja. Math. Tcrniesz. firtes. Buda-
pest. 12. p. 364.

lu)MONnsoN', W. T. 1934. Investigations of .some Ilispaniolan Lakes. (Dr. R. M. Bond's
Expedition.) I. The Rotatoria. Arch. Hydrobiol. 26. p. 465.

EiiRENBERG, C. G. 1838. Die Infusionsthierchcn als vollk>ininiL-ne Organismcn. Leipzig.

I-'adkev, N. N. 1927. Materials for the Study of the Rotatorian l'"atuia of U. S. S. R.
Proc. Nat. Ili.st. Soc. Kharkov. 15. ])art 2. (Reprint .separately paginated.)

Francke, A. H. 1914. Antiquities of Indian Tibet. Archaeol. Surv. India, New Imp.
Ser. 38. Part I, p. 54. Calcutta.

Harring, II. K. 1913. Synopsis of the Rotatoria. Bull. V. S. Nat. Mus. 81. p. 1.

ROTATORIA 185

Harking, H. K., and Myers, F. J. 1924. The Rotifer Fauna of Wisconsin. II. A Revision
of the Notominatid Rotifers, exclusive of the Dicranophorinae. Trans. Wise. Acad.
Sci. Arts Lett. 21. p. 415.

1926. III. A Revision of the genera Lecane and Monostyla. ibid. 22. p. 315.

Heinis, F. 1910. Systematik und Biologic der moosbewohnenden Rhizopoden, Rotatorien
und Tardigraden von Basel mit berucksichtigung der iibrigen Scweiz. Arch.
Hydrobiol. 5. pp. 89, 217.

Hutchinson, G. E. 1931. New and Little-known Rotatoria from South Africa. Ann.

Mag. Nat. Hist. (ser. 10). 7. p. 561.
1933. Lininological Studies at High Altitudes in Ladak. Nature. 1932. p. 136.

Hutchinson, G. E., Pickford, G. E., and Schuurman, J. F. M. 1932. A Contribution
to the Hydrobiology of Pans and other Inland Waters of South Africa. Arch.
Hydrobiol. 24. p. 1.

Jakubski, a. W. 1915. Apis fauny Wrotkow powiate Sokalskiego. Rosprany i Wiado-
mosci z Museum im Dzieduszychkick 1. p. 1.

Klausener, C. 1908. Die Bkitseen der Hochalpen. Int. Rev. Hydrobiol. 1. p. 359.

Kratzschmar, H. 1908. Ueber den Polymorphismus von Anuraea aculeata Iihrbg. Int

Rev. Hydrobiol. 1. p. 623.
■ 1913. Neue untersuchungen iiber den Polymorphismus von Anuraea aculeata Ehrbg.

ibid. 6. p. 44.

Monti, R. 1906. Rcchcrchcs sur quelques lacs du Massif du Ruitor. Ann. Biol. Lacustre.
1. p. 120.

Mui.LER, O. b\ \7H(). /Nniniacula Infusiiria. Ilauniae.

Murray, J. 1906. Some Rotifera of the Sikkim Himalaya. |. Iv. Micr. Soc. London.
(Ser. 2.) 9. p. 259.

Murray, J. 1913. Quoted in Bryce, 1931.

Myers, F. J. 1931-34. The Distribution of Rotifera on Mount Desert Island. Amer. Mus.
Nov. nos. 494, 659, 660.

Pesta, O. 1929. Der Hochgebirgssee der Alpen. Die Binnengewiisser. 8. Stuttgart.

Plate, L. H. 1886. Beitrage zur Naturgeschichte der Rotatorien. Jena Z. Naturw. 12.
p. 1.

ScHMARDA, L. K. 1850. Neue Formen von Infusorien. Denkschr. Akad. wiss. Wien 1.
pt. 2. p. 1.

Schuurman, J. F. M. A Seasonal Study of the Microflora and Microfauna of Florida Lake,
Johannesburg, Transvaal. Trans. Roy. Soc. S. Afr. 20. p. 333.

Skorikov, a. S. 1896. Rotateurs des environs de Kharkow. Trav. Soc. Nat. Kharkov.
30. p. 207. (In Russian.)

186 ROTATORIA

•s. ^

Slominski, p. 1926. Sur la Variation Saisonnicre chez Triarliira (iMlinia) longiseta E.
C. R. Soc. Biol. 1926. p. 543.

Smu^xov, N. 1930. Rotatoria. .Xhliandlungen der I'aniir-Expedition. 2. Zool. p. 87.

Stkwart, F. II. 1908. Rotifers and (lastrotricha from Tiliet. l\cc. Ind. Mus. 2. ]). 316.

T.\KNOGRADSKY, D. 1''30. /ur Iv ilatnricu fauiia ck-s XMrd-Kaukasus aus dcm ( iciicra
Lecaiie, Moiiastyla und CultircUa. Trav. Stat, liinl, du ( aucast- du Xurd. 3 pt. Yi.
(Abstract only seen.j

TriiKB.M'D, M. 1911. Les Rotateurs du Canton de Ncuchatel. Neuchatel r.uil. Soc. Sci.
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Thorpe, V.G. 1891. New and Foreign Rotifers. J. R. Micr. Soc. London. 18<)1. p. 301.

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8. p. 5.

Webek, E. F. 1898. b'aune rotatorienne du bassin de Leman. Rev. Suisse Zool, 5. p. 263.

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Geneva.

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WiSZNlEWSKi, J. 1931. Sur (piel((ucs Untifcres trouves en J-'.spagne. Arrh. d'l lydrnliinl.
et d'lchthyol. 6. p. 41.

1933. Un nouveau Rotifere du genre Pedalia habitant les lacs des hautes montagnes.

Int. Rev. Hydrobiol. 29. p. 229.

Zavadov.ski. 1916. Quoted in Harring and Myers, 1924.

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Naturw. Zurich, i?. p. 1.

THE TUTTLE, HODEIIOUSt » TAYLOR COMrANY. NEW HAVEN, CONN.

MEMOIRS OF

THE CONNECTICUT ACADEMY

OF ARTS AND SCIENCES

Vol. X

YALE NORTH INDIA EXPEDITION

Article X — Ueter Einen Land-Isopoden aus Latlak, by K. \V. Verhoeff.

Article XI — Report on Hirudinea, by J. Percy Moore.

Article XII — Hochasiatische Binnenseesedimente, by G. Lundqvist.

Article XIII — Report on Myriapods, Ijy F. Silvestri.

Article XIV — Report on Diplura and Thysanura, by F. Silvestri.

Article XV — Report on Collembola, by J. R. Denis.

Article ■ XVI — Report on Cladocera, by Dr. V. Brehm.

Article XVII — Report on Fishes. Part I: Cobitidae, by Sunder Lai Hora.

Article XVIII — Report on Fishes. Part II; Sisoridae and Cyprinidae, bv Dev Dev
Mukerji.

June, 1936

Price, $4.75

NEW HAVEN, CONNECTICUT

published by the

CONNECTICUT ACADEMY OF ARTS AND SCIENCES

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ARTICLE X

UEBER EINEN LAND-ISOPODEN AUS LADAK
(55. Isopoden-Aufsatz)

By K. W. Verhoeff (Pasing bei Munchcn)

With 7 Text-Figures
(Received November 8, 1934)

Die niir in sechs Tuhcn von 6 I'^nndplatzen durcii Herrn Prof. G. E. Hutchinson
freudlichst iibennittelten Isopoda-tcrrcstria aus deni nordwestlichen Indien geln'iren alle zu
der einzigen, ini Folgenden besprochenen Protracheonisciis-Art.

Hinsichtlich dieser Gattung Protracliconiscus niochte ich al)er zunachst Folgendes her-
vorheljen : In meinem 22. Isopoden-Aufsatz "Zur Kenntnis der Entwickelung der Tracheal-
systeme und iiber die Gattungen PorceUio und Trachconiscus'' (SitzBer. Ges. Naturf. Fr.,
Berlin, 1917, N.3 S. 195-223.) habe ich nicht nur eine neue Umschreibung dieser Gattun-
gen besonders auf (irund der Atniungsorgane gegeben, sondern audi mehrere Untergat-
tungen beider begriindet. Unter den Untergattungen von Tracheoniscus befindet sich auch
Protmchcomsctis, eine Gruppe, welche jedoch spater ebenfalls um so mehr als eigene Gat-
tung betrachtet werden musste, da sich die Notwendigkeit ergab sie selbst wieder in Unter-
gattungen zu teilen.

Dies geschah in meinem 42 Isopoden-Aufsatz, iiber Isopoden aus Turkestan (Zool.
Anzeiger, Leipzig 1930, Bd 91. H. 5/8 S. 101-125.) wo ich nicht nur die Beziehungen von
Protracheoniscus und A''a^ara behandelt, sondern auch auf S. 105 die lieiden Untergattungen
ProtracJiconiscits s. str. und Mongoloniscus unterschieden lial)e. Wahrend Mongolonisciis
fiir Ostasien charakteristisch ist, stellt Protracliconiscus die artcnrcichstc imd iihcrhaiipt
hervorstechendste Isopoden-Gruppe der mittlcrcn Ldndcr Asicns vor. Aus Turkestan habe
ich allein 7 Arten nachgewiesen. Aus luehreren im letzten Jahrzehnt erschienenen Aufsatzen
wissen wir jetzt auch, dass Protracheoniscus fiir den Siidosten Europas bezeichnend ist und
dass die tccstlichstcn Vorposten der Gattung fast genau in der Mitte Deutschlands stehen,
so namentlicli der bekannte politiis Koch, \'erh. Die oek(jlogischen Anspriiche der einzchien
.\rten sind ausserordenthch verschieden und ebenso die Grossen der Areale. Es giel)t wiirnic-
bediirftige, mediterrane Arten von geringer Verbreitung einerseits, aber andererseits auch
weiter verbreitete und weniger empfindliche Arten, wie z. B. den asiaticiis Ulj.

Jedenfalls kennen wir aber in Europa keine Art, welche in den Gebirgen besonders hocli
ansteigt, z. B. ol)erhallj 1500 m. ist meines Wissens in Europa nie cin Protracheoniscus beo-
Iiaclitet worden. Aber auch aus Asien waren bisher Vorkommnisse von bedeutender II(")lic
nicht bekannt. Uin so mehr hat es mich iil)errascht, dass die im Folgenden beschriebene Art in
Nordindien in Hohen lebt, in welchen es in Europa iiberhaupt gar keine Isopoden giebt. In
den niitteleuropjiischen Alpen sind schon l)ei 2000 ni. I lohe in vielen Gebirgen keine Lsopoden
mehr anzutreffen, im Gegensatz zu den klimatisch viel widerstandsfahigeren Chilopoden und
Diplopoden.

Die Starke Vertretung der Gattung Protracheoniscus in Asien, woljei aber zweifellos

Mem. Conn. Acad., Vol. X, Art. X, June, 1936.

188 UEBER EINEN LAND-ISOPODEN AUS LADAK

die weitiuis incistcn Arten noch unbekannt sind, ferner die Tatsache, dass sowohl die Untcr-
tjattung Moniiolaiiisciis als audi die nahe verwandte Gattung' Dcscrtoniscus \'erh. (cljen-
falls ini 42. Ausatz beschriel)cn ) in Asien lieimatcn, wiilircml cntsprcfheiidc Aerwandte (irup-
])cn in Europa (und aiidern Continenten) nicht bekannt sind und schliesslich das extreme hcjlie
Vurkommen des Pr. nivalis, sind in liinklang stehcncle Erschcinungen, wclcli dafiir sprcchcn,
dass Asicn die Urliciiiiat z'on Protracheoniscus isl.

Protracheoniscus (Protracheoniscus) nivalis n. sp.

9 11-14 nun., i 12 mm. Jang. Iviickcn gran l)is hrauiiscliwarz.

Von alien hekannten Protracln'oiiiscus — Arten nntersciieidet sieli die vorliegende schon
iinsscrlich.

1. Durcli die Stirnleisten, welche zwischen deni Mitteiteil derselhen und den Seitenlapjien

fa.st einen shiiiipfni Jl'inlcrl hilden ( wiihrend sie sonst liier einfacli gehogen sind),

2. Durch den Hinterrand an den 1. rereion-l'"])imeren, welclu-r ganz i:;crailc streicht

(wahrend er sonst bogig verliiuft),
.V Zeigen die 1. I'leopoden des S (A1)I). 1 und 3) naeii ]''.\(ipiKlil und iMKJDpDdit eine recht
eigentiimliche Beschaffenlieit.

Riicken luelir oder weniger gliinzend, .\n(ennen vtm tx'pischer I.iinge, die l)ei(len fieissel-
glieder gleich lang, oder das terminale etwas (bis 1/3) kiirzer, 3 (ilied am l'".nde xuvu und
hinten mit kleinem Zahn, 4 und 5 deutlicli gefurcht. An den 3 gliedrigen, seiir kleinen
Antennulen (Abb. 6) das mit mehreren Sinnesstiibehen besetzte Englied nur halb .so lang wie
das mittlere und dieses etwas schmiiler mid weiiig kiirzer wie das Grundglied. Letzteres
springt am Ende innen etwas gerundet vor.

Ocellen in vier Reihen stehend. Die Seitenlappen des Kopfes gross, fast halljkreisffirmig,
ihr Endrand uni etwa ^ der Liinge des Ocellenhaufens von diesem entfernt. Stirnleiste in
der Mitte mit stunipfen Winkel vorragend.

E.xopodite der 1 Maxillen mit 4 derlieren und 5 feineren Zannchen, von den 4 derljeren
einer viel kiirzer als die andern. 2 Maxillen am luide tief eingesschnitten in zwei fast
gleich breite Lappen, deren ausserer nackt und deren innerer fein behaart und gestrcift ist.

Die Kieferfiisse (Abb. 7) sind durchaus iiacli dem Ijekannten Porcc!!ionidcii-Tyin\f.
gebaut, ihre 3-gliedrigen Taster, wie iiberjiaupt die ganzen Kieferfiisse zeigen kaum etwas
Besonderes gegeniiber denen der \"erwandten, stimmen auch fast ganz iiberein mit denen
von Dcscrtoniscus (Abb. 17 in nieinem 42. Aufsatze) doch ist hier bei Dcscrtoniscus das 3
Tasterglied schlanker und zugleich nicht deutlich abgegrenzt.

Der Riicken erscheint (unter der Lupe) feinpunktirt. Auf den h'pinieren des Pereion
zeigt sich eine schwache aber deutliche Korncliiui^. Noduli laterales am 1.-4. Pereioiitergit
viel hoher stehend als am 5.-7. .\m. 1. Tergit, wo die Noduli zugleich etwas grul)ig vertieft
liegen, sind sie vom Seitenrand 1 ' j mal weiter als vom Hinterrand entfernt, am 3 Tergit vom
Seitenrand doppelt so weit wie vom Hinterrand entfernt, am 4 Tergit 3 mal so weit. Am 5-7
Tergite stehen also die Nnduli dem Seitenrand viel niilier und zwar sind sie am 5. und 6.
Tergit vom Seiteii- und I iinlerrand i^leich weit entfernt, am 7 vom Seitenrand I'/, mal weiter
als vom Hinterrand.

In den Punkten der genannten Punktirung sitzen iil)er den ganzen Riicken zcrstrent
kurze und sehr feine Borsten, die sich (mikroskopisch) als einfach erweisen.

UEBER EINEN LAND-ISOPODEN AUS LADAK 189

Pleon ohne Kornelung. Das im dreieckigen Spitzenteil grubig eingedriickte Telson reicht bis
zum Hinterrand der Uropoden-Propodite. An den Rjindern der Epimeren miinden Driisen
iind zwar stehen an den 1 Pereion-]*2pimeren die Driisenporen in einer Langsreihe hinter
den al)gerundeten V^irderecken. An den 7. Epimeren sali ich 5-7 Driisenporen in einer Langs-
reihe im mittleren Gebiet, dicht nelien dem Seitenrande.

Am 7 Beinpaar des $ ist das Ischiopcidit unten bogig ausgelnililt, kurz beborstet, Ijesitzt
oben in der J'^ndhiilfte eine im Bogen angeordnete Reihe vnn 7 Stacheliiorsten. Meropodit
am Ende olien und unten mit je drei Stachelborsten, Carpopudit unten mit drei stufigen Abset-
zungen und 7-8 Stachelljorsten.

Von sehr charakteristichem Ban sind die 1. Pleopoden des S . Die gerade nach hinten
gestreckten und allmahlig verschmalerten 1. Endopodite (Abb. 3) lanfen in einen fast
dreieckigen, am Ende abgerundcten Endzipfel ans. Die aussere Basis dieses Endzipfels
tritt nach aussen cckig vor und vor dieser Ecke miindet die Spermarinne (.r. ). An der
immeren Basis ist der Innerrand unterbrochen (at.). Hinter dieser Unterbrechung zeigt
sich eine kurze Wimperreihe und vor ihr ist der Rand mit ausserst feinen und kurzen
Spitzchen Besetzt.

Trachealsysteme treten, wie bei alien Protrachconisciis, an den 1.-5. Exopoditen auf. Die
1. und 2. Exopodite sind pigmentlos, die 3.-5. von zahlreichen verzweigten Pigmentzellen
durchsetzt, (Abb. 4.).

Die 1. Exopodite des S (Abb. 1.) sind hinten breit abgerundet, innen gerundet, aussen
hinten gerade abgeschragt, wahrend vom der Aussenrand in einen abgerundeten happen
vorspringt, an (lessen hinterer Basis das Trachealsystem miindet. Plinter dieser Miindung
bemerkt man innen neben deni Schrrigrande eine Tracheal feldleiste, deren Hinterende
undeutlich.

An den 2. Exopoditen des & , welche am Ilinterrande aussen eine Reihe kriiftiger
Borsten tragen, befindet sich mitten im Trachelfeld eine stumpfwinkelige, ziemlich tiefe
Einbuchtung (tf, Abb. 2.) und auch die Tracheal feldleiste ist stump fwinkelig eingebuchtet.
Die 2. Exopodite werden von ihreni Endopodit, welches im Endteil sehr diinn ist und spitz
auslauft, nur wenig iiberragt.

Den starksten Borstenbesatz und zwar am Ilinterrande besitzen die 5. Exopodite des
<J (Abb. 4) welche vorn quer abgestutzt, aussen und hinten zugerundet und innen fast
gerade nach hinten streichen. Ueber dem Innenrande zeigt sich die bekannte, taschenartige
Einsenkung. Die Trachealysteme (Abb. 5.) miinden am vorderen Aussenrande etwas vor
der Mitte und sind an den 5. Exopoditen am schwiichsten entwickelt.

Vorkoiiiincn. Die in etvva 14 Stiick vorliegende Art ist auch in zwei Jugendlichen vertreten
und im obersten Indusgebiet an folgenden Orten gesammelt worden :

L. 20 Hemis Gonpa, unter Steinen, 12-VI-1932. c. 3660 m.

L. 31 Lhabaps, unter Steinen, 22-VI-1932. 3614 m.

L. 32 Zung-Lung, Tangyartal unter Steinen, 24-VI-1932. 4224 m.

L. 68 Zwischen Anzurma und Dambu-guru, l-VIII-1932. c. 4725 m.

L. 72 Tokung, bei Panggong Tso, 8-VIII-1932. c. 4250 m.

L. 68 und L. 72 befinden sich zu beiden Seiten des Panggong Tso. Ausserdem liegen
nncli vor von Tso Nyak in Tibet Tzewang Tashi und Sonam Tergas 12-viii-1932 durch
auffallende Weichheit ausgezeicimet 2 9 9. Ein 9 mit Marsupium wurde nicht beobachtet.

100

UEBER EINEN LAND-ISOHOUEN AUS I.AIIAK

^liiiiirrk'un}^: V'icllciclit liaiulelt cs sich bei tlicseu Ticren uin zwci Ivasscn, dercn cine
scliniiilcr uiid licller iiiul dcren andere l)reiter und dunkler ist. Dies iJisst sich jedoch
mir an zahireidicrcn Objecten entschcidcn als niir vorj^elejjen liabcii. Wie cs scheint
ist dcr Traihconiscus nk'alis die am hochstni icbcndc /siipiii!cn-.\r{. wckiic bislicr auf
nnscrc I'-rdc l)C()l)aciitet wordcn ist.

2. ex

Figures 1-7. Protracheoniscus (Protracheoniscus) nivalis n. sp. 1, Eiii I. PIcnpoclcncxnpiKlit ties S,
voii uiitcn gcsclicn, X 56. 2, Aiisscror .\hsclinitt cincs 2. Plcdpiidciicxopodit des S, I, Tracliccii, //, Trachcalfeld.
X 125. 3, Endteil eines 1. Plcopodciicndopodit des S, Ansicht von unten, i, Innen- a, Aussenrand, .r, MiinduiiK
der Spermarinne, X 125. 4, Ein 5. Pleopodcnexopodit des S von unten geschen, pr, Propodit, X 56. 5, Ausserer
Teil desselben mit dem Trachealfeld (tf), X 125. 6, Linke Antcnnule (an) und der angrenzende Teil der
Gelcnkgrube der linkcn .Antcnne, t', Vorder- //, Hintcrrand, X 125. 7, Endteilc des reehtcn Kicferfusses, Ansicht
von unten, X 125.

ARTICLE XI

REPORT ON HIRUDINEA

By J. Percy Moore
(Received December 15, 1934)

The collection of leeches is very small, consisting of ten lots representing three
species. One of these came from the Nilgiri Hills in Madras Presidency and two from
Kashmir, one being a new record for that state. All are from moderate elevations and
the absence of land leeches is worthy of note.

THEROMYZON SEXOCULATA (Moorc)

Protoclcpsinc scxoculata Moore, 1898.
Thcromyzon sexoculata Moore, 1924.

Harding, 1927.
? Protoclcpsis mcyeri Livanow, 1902.

This species is new to Kashmir, though it was previously known from India, having
been recorded from Manipur (Moore, 1924) ; the type is from Bering Island (Moore,
1898 j. Livanow considered T. scxoculata (unfortunately so named) identical with a
species known from Russia, Sweden and France, to which he gave the new name of T. mcyeri.
Externally the resemblance, in respect to annulation, eyes, genital orifices and color, is close.
The only difference noted on the Manipur specimens was that the second annulus (ai) of
somite XXVI was differentiated only at the margins, as is also the case in the Kashmir
specimens. Somite III is triannulate on all specimens. But a series of sections which this
material permitted definitely establishes the distinction between the Indian form and T.
mcyeri. The gonopores are similar in position ( S XI/XII, 5 XII a2/a3) but the two
oviducts are united beneath the nerve cord into a slender, tubular vagina which runs vertically
to the ? orifice, exactly as in T. tcsscUata, as contrasted with T. mcyeri in which there is no
such common vagina. The condition of the oviducts in the original T. scxoculata is
unknown, as it could not be determined from the single very poorly preserved type specimen.
It is not improbable that the south Asiatic species will eventually prove to be distinct from
T. sc.vociilata but present evidence gives no ground for separation.

The color of the present specimens is largely faded but all retain at least traces of the
six series of metameric yellow spots and in addition a moderate number of similar but unseg-
mental spots. One specimen has a dark band on each side of the buccal ring, wide laterally
and tapering to a point medially. The dark green, contracted chromatophores are con-
spicuous and on the venter of one specimen form dark rings about the sensillae which
appear to the naked eye as black dots. On complete somites a3 is constantly somewhat
larger than the other annuli Ijut there is no indication of subdivision. Ventrally throughout
the length, and dorsally at both ends, the intersegmental furmws are conspicuously deeper
than the others and a2/a3 is deeper than til/aZ.

Mem. Conn. Ac/ml, Vol. X, Art. XI, June, 1936.

192 RKPORT OX TTIRT-niNKA

AlPfour specimens (K34 Nos. 691 and 698) were taken at Pliashakuri, near Pampur,
Kashmir, May 7, 1932, at an altitnde of 5200 ft.

Erpobdella octoculata (Linn.)

Ilintdo octoculata Linnaeus, 1758.
Erfobdella octoculata Moore, 1924.
Moore, 1927.

This widely distributed Eurasiatic species is very common in tiie lakes and ponds of
Kashmir. It was fully discussed in my 1924 paper. It is the best represented species in this
collection but all specimens are small. The gonopores are normally separated by 3 annuli,
the $ in or immediately behind the furrow XII bl/b2 and the 9 constantly in XII b5/b6
but the d may shift caudad as far as the middle of b2 in which case it is only 2^^
annuli anterior to the ? pore. One specimen has two <5 pores in XII b2 and XIII b2
respectively.

K 15 and 19 Gagirbal, Srinagar, Kashmir, swampy pond east of ruad, altitude 5190 ft.,
No. 703, 707; K 24, Nishat Bagh, pond, April 7, altitude 5200 ft.. No. 651 ; K 42, W'ular
Lake, April 18, alt. 5180 ft., dredged. No. 763, depth 1.5-2 m., No. 765, depth 1.0 m.;
K61, (iund, Sind Valley, May 17, altitude 6824 ft., under stones on nnfddy bottom of
small stream.

FORAMINOBDELLA IlEPTAMERATA Kaburaki

Foraminobdella hcptamcrata Kaburaki, 1921.

Moore, 1927.

This interesting and little known species is represented by four poorly preserved speci-
mens N. 8 (No. 834, 849) taken in ponds on Pykara Road near Ootacamund, Nilgiri Hills,
Madras, November 10, 1932, altitude 7200 ft. The specimens measure from 23. x 4. to
42. X 5.3 mm. All have the gastropore of large size: this structure, the gonopores and the
annulation are as described in 1927 Irom the tyjie, at that time unique, which also came
from the Nilgiri District. In January, 1931, I was fortunate in tinding several populous
colonies of this species in the lake at Ootacamund and was able to study its mode of life
and to secure material from which an anatomical description will be pul)lished in another con-
nection. Its favorite habitat is in the little gravelly deltas at the niduths i>i streams emptying
into the lake. Here it is found under stones especially just above water level and when
exposed disappears quickly into the gravel and silt. It is as muscular, hard and slippery as
an eel, very difficult to hold and with its pointed head and slender form an adept burrower.
When placed in water a current may be seen to issue periodically from the gastropore and
when removed to the air a fine jet of water was sometimes ejected from it a distance of
several inches. In life the color is a dull or bright red or pink according to size and contents
of stomach. The food consists chiefly of tubificid oligochaetes.

Full references to all of the papers cited in the synonymy appear in the bibliographies
of Harding and Moore, Fauna of P.ritish India. Ilirudinea. L(indon. 1927.

ARTICLE XII

HOCHASIATISCHE BINNENSEESEDIMENTE
By G. Lundqvist

With 1 Plate and 5 Text-Figures
(Received February 8, 1935)

Fonvort

In einigen iilteren Arbeiten, besonders von 1927, habe ich die Auffassung verfochten,
dass die Einsammlung von Bodenproljen in Seen niit der grossten Genauigkeit und von dem
Forscher, der dieselben bearbeiten wird (Lundqvist 1927), ausgefiihrt werden muss.
Weiterhin ninss man, um den Tj-pus eines Sees feststellen zu konnen, eine pers("inliche Erfab-
rung desselben baben, denn in dem Typus ist eine Mannigfaltigkeit unbedeutender und
unbeschreibbarer Faktoren vorhanden. Es scheint deshalb inkonsequent, dass ich die Bear-
beitung der Bodenproben von "The Yale North India Expedition, 1932," die mir von Dr. H.
de Terra und Dr. G. E. Hutchinson freundhchst angeljoten wurde, iiljernonimen halje. Ander-
seits aber war die Moglichkeit, Sediniente aus den hochst gelegenen limnologisch untersuch-
ten Binnengewassern der Erde zu sehen, gar zu verlockend. Ich mochte darum den erwahn-
ten Forschern fiir ihr freundhches Anerbieten meinen besten Dank aussprechen. Ganz
besonders mochte ich Dr. Hutchinson fiir die ortHchen Observationen iiber die verschiedenen
Seen, die er mir bereitwilHg geliefert hat, danken. Ich l)in ihm und Dr. de Terra auch
dankbar fiir das Durchlesen der Korrcktur, das ich infolge drucktechnischer Umstiinde leider
nicht selbst ausfiihren konnte.

Von denen, die mir im iibrigen geholfen haben, mochte ich Dr. H. Thomasson, der die
Diatomeenbestimmungen au.sgefiihrt hat, zuerst nennen. Weiter hat Dr. S. Thunmark das
Desmidieenmaterial durchmustert. Dr. G. Assarsson hat die W'asseranalysen, die mir von
Dr. Hutchinson zur Disposition gestellt wurden, durchgerechnet und mit mir diskutiert.
SchHesshch habe ich den Vorteil gehabt, mit Dr. R. Mehn und Dr. C. J. Ostman iiber die
Klimafragen sprechen zu konnen. Besonderen Dank schulde ich deshalb auch diesen fiinf
Forschern.

Schon hier diirfte erwahnt werden, dass die Lokalitatsnamen dieser Gegenden in ganz
verschiedener Weise geschrieben werden. Ich habe selbstverstandiich die Namen der Berichte
der Yale-Expedition benutzt (de Terra 1934). Daneben habe ich aljer die Seenanien ange-
fiihrt, die auf solchen bekannten Karten wie z. B. denjenigen von Hedin (1909) und Dainelli
(1922) gebraucht worden sind. Eine mehr systematische Priifung und Vergleichung von
Karten der vorliegenden Gebiete von ortographischen Gesichtspunkten aus habe ich selbstver-
stiindlicli nicht versuciit.

Um ein Missverstandnis zu vermeiden, hat jeder See seine Nummer erhaiten, und (Hese
ist sowohl in den Text als auf die Karten eingesetzt worden.

Uber die Disposition der folgenden Arbeit sei nur angefiilirt, dass einerseits das Mate-
rial (also die deskriptiven Kapitel ) andererseits, die mehr tiieoretischen Kapitel und die

Mem. Conn. Acad., Vol. X, Art. XII, June, 1936.

194 imCTrASIATISCHE BINNENSEESEDIMENTE

Riickblicke fiir sich stehen. Das Ilauptgewicht ist auf das crstere gelcgt, da die Sedimcnte
dieser Gegenden bisjetzt ganz iinbekannt wareii. Darum diirfte hier jedt-s Detail von Inter-
esse sein kuiincii.

Mcthodischc Bcmcrkungcn

Die Probeii siiid mit deiu llknian-r>irge-Bodengroifer eingesammelt wnnk'n. Nur in
einigen Seen (Son Sakesar Kabar, Lokut Dal Lake und W'ular Lake) wurde das Ivohrlut
von Nauinann benutzt. Die rro1)en, die mit deni IJudengreifer genonimen warden, sind ja
betrcffs ibrer Lage in Verhiiltnis zu der Sediment obertlache nicbt so gut Lockalisiert wie die
ul)rigen, und das Jtlaterial ist auch zusammengeriihrt und beterogen geworden. Man kann
darum in solcben Proben Klumpen aus Gyttja mit ganz vcrschiedener mikrobiologischer
Zusammensetzung finden. Die iiusserst genauen Metboden, mit denen die Proben I)earl)eitet
wurden, sind desbalb oft nicbt notig gewesen. Ich niocbte daber die Aufmerksamkeit darauf
ricbten, dass hier in hohem Grade der Ausspruch von Hagen gilt : "Der Mangel an matbe-
matiscbcr Bildung gibt sich tlurcb nichts so auffallend zu erkennen, wie durcb masslose
Schiirfe im Zablenrecbnen." Damit mocbte ich also audi bcrvorbelx'n, dass die I'"requenz-
zahlen, die fiir die Diatomeen oder Strukturelemente angefiibrt werden, nicbt so exakt,
wie sie im Druck erscheinen, aufzufassen sind.

Ul)er die Observationen an Ort und Stelle ist auch zu Ijemerken, dass die Farlie des Sees
nacb dem Masstab von Forel-Ule (Ule 1892) und dicjenige des Wasscrs nacb dem Platina-
C'blorid-Masstab der U. S. (leological Survey (Lcigbton 1905) bestimnit worden sind. Die
natiirlicbcn L'arlK'n der Sedimente sind scbwer festzustellen, da .samtlicbc Proben, die nicbt
getrocknet sind, in Spiritus oder Formalin fixiert sind.

Die Bearljeitung der Prol)cn im Laboratorium ist mit einigen Erweiterungcn nacb Lund-
qvist, 1927, ausgefiibrt worden. Icb babe mich also nicbt mit eingetrockneten Prol^en
beschaftigt. Die Eintrocknung verursacbt In-i den Kullniden, feinereni Detritus u. a. den
W'rlust sowobl ibres Aussehens als audi ilires \ oluniens. Dadurcb wird also das X'olumen-
verbiiltnis zwiscben organogenem und minerogenem Material venindert. Die Strukturan-
alyse ist durch Rechnung der verschiedenen Bestandteile uuter dem Netzokular von Leitz in
1 mm.'' von der gefeuchteten Probe ausgefiibrt worden. Bei dieser Analyse kann es ganz
scbwierig sein, das feink("irnige minerogene Material und I'eindetritus von einandcr zu unter-
scheiden. Ich babe darum verscbiedene Fiirbungsmittel prol)iert, um eines zu finden, das das
minerogene Material unbedeckt und farblos lasst. Das l)este der jirobierten Mittel ist ein
gewohnlicher Anilinfarljenstift (vgl. Naumann 1918). Nacb cbemiscber Metbode kann man
dieses feinkornige Material, besonders den Tonschlamm, erreicben. Die metbotliscbe
Schwiiche bestebt darin, dass man hierbei in der Sunime des unorganischen Materials auch
die Diatomeen und die anderen Kieselskelette bekommt. Es ist daber meines Erachtens vom
mikrobiologiscbem Gesichtsj)unkte aus mebr anzuraten, die Proben mikroskopisch zu unter-
suchcn. W^iinscht man aber den Charakter der Sedimente von rein cbemischem Gesichtspunkt
aus berauszufinden, ist es eine ganz andere Frage.

Die mikrobiologischen Untersucbungen sind ebenfalls auf 1 nim.^-Prol)en ausgefiibrt
wc)rden. In den mebr eingebenden Diatomeenanalysen, von II. Thomasson, werden die
Resullate in Prozentzahlen vorgelegt, wol>ei die Prozente der Summe der nicbt kolonienbil-
denden Individuen au.sgerechnet sind. Ausnahmen bilden z. B. Mclosira armaria und Cyclo-
tella, dagegen weder Mclosira '^ninuhila etc. nndi l-nt;^ilaria.

IIOCHASIATISCIIE BINNENSEESEDIMENTE 195

Auf diese jetzt olien beschriebene Weise siiul audi die Prol3en niit der "Trockenprobe,"
der "HCl-Probe" und der "Tiischprobe" iintersucht worden. Die Trockenprobe ist ein ein-
faches Hilfsmittel, um eine ungefahre Vorstellung des Dygehalts in den Sediinenten zu ver-
mitteln : je dyreicber dieselben sind, um so mehr braunschwarz werden sie in trockenem
Zustand. In den vorUegenden bunuisarnien Gebieten ist die Prul^e jedoch von vnitergeord-
netein Interesse. Die HCl-Probe beabsichtigt die Angabe, einer ungefiibren Auffassung des
relativen Karbonatgehalts. Auf etwa 1 mm.^ des Sediments wird ein Tropfen HCl (etvva
10%-ig) getropft und das Schaumen wird beobachtet. Fiir die Bestimmung desselben babe
ich eine 5-gradige Skala benutzt nach dem Prinzip : == kein Schaumen, 5 =^ sehr starkes
Schaumen. Die Anwendung ist sehr einfach und weitere Besclireibung unnotig. Die Tusch-
pr(i1)e gil)t eine recbt gute Vorstelhuig des Gebalts an Algensciileim iin Feindetritus. Diese
Probe ist schon friiher, besonders in der algologischen Literatur, mehrmals beschrieben
worden.

Schhesshch mochte ich bier auch hervorheljen, dass ich auf das Zusammenbringen von
Data iiber die Naturverhiiltnisse viel Zeit verwendet hal^e. Denn fiir das V'erstehen der
Biologie eines Sees ist es viel wichtiger, dass man die Umgebungen und das Milieu des
Sees beherrscht, als dass man z. B. samtliche Arten einer besonderen Tier-oder Pflanzen-
gruppe, die darin lebt, kennt.

Das Material, das iiber die Naturverhaltnisse dieser Gegenden vorliegt, ist ebenso gross
als auch schwer zu iiberblicken. Diese Gebiete sind klassischer Boden sowohl von dem
Gesichtspuiikte der indischen Naturforschung aus als auch aus limnologisch klassischer
Boden. Hier waiiderten die Briider Schlagintweit schon Mitte des vorigen Jahrhunderts und
untersucbten auch die Seen. Selbstverstandlich waren ihre Methoden ausserst primitiv ; in
einer Hinsicht al^er waren diese Forscher ihrer Zeit voran : sie machten relativ genaue Obser-
vationen iiber die Transparenz und Farbe der Seen. Es diirfte lohnen aus der Vergessenheit
gerettet zu werden, dass sie im Ladak- und Kashmir.see die Sichttiefe durch Messen der Tiefe
Ijestimmten, bei der ein Zylinder aus Carrara-Marmor unsichtbar wurde. Fiir die Farben-
bestimiuungen benutzten sie eben falls ein grosses Glasprisma, das man unter der Wasserflache
drehte, bis die stiirkste Farlje erhalten wurde (Schlagintweit 1871 [74] S. 170).

Einige Bodenproben der asiatischen Seen von reinem Sediment-gesichtspunkt aus
scheinen jedoch vorher nicbt genommen worden zu sein, obgleich einige "Schlammproben"
auf Diatomeen bin bearl>eitet worden sind. Ich mochte in dieseni Zusammenhang Meister
(1932, S. 2) anfiihren: "Es fehlten also bis jetzt Proben vom Grunde stehender Gewiisser,
die erfahrung.sgemass die reichste Ausl^eute liefern. Wenn einmal richtiges Benthos zur
Untersuchung gelangt, werden sicherlicb nocb viclc neue Formen bekannt." Aus den in
vorliegender Arbeit untersucbten Pmbcn scbeint es jedoch, als ob Meisters Pnjphezeiung
nicht Stich halten wiirde.

Die hier untersucbten Seen liegen in drei Geljieten : Salt Range im Punjab, dem Kash-
niirtal und Ladak im westlichen Tiljet (Text-figure 1 und 2). Ich werde versuchen, ii])er
jedes flicser drei Gebiete eine kurze Naturbescbreil)ung zu liefern.

Salt Range

Die Salt Pange ist ein O-W'-licber Komple.x von Gebirgsketten zwischen (lulus luitl
Jhelaiu im Punjab. Gegen S Ijesitzt das Gebiet einen starken AI)fall ; gegen N ist es aber
nicht so scharf abgegrenzl. Das Seegebiet liegt auf einer schwach kupierten Hochebene

196

HOCHASIATISCIIE BINNENSEESEDIMENTE

Figure 1. Die Seen der drei Untersuchungsgebiete: Nr. 1 Salt Range (Sun Sakesar Kaliar), Nr. 2-6 Kashmir

und Nr. 7-15 Ladak.

Figure 2. Die untersuchten Seen: 1 = Son Sakesar Kahar, 2 = Lokut Dal Lake, 3 = Bod Dal Lake, 4 =
Sundar Khun, 5 = Manasbal Lake, 6 = Wular Lake, 7 = Tso Moriri, 8 =; Khyagar Tso, 9 = Startak-puk Tso,
10 = Tso Kar, ll = Yaye Tso, 12 =: Witpal Tso, 13 = Pangur Tso, 14 = Panggong Tso und IS^Ororotse Tso.

IIOCIIASIATISCIIE BINNENSEESEDIMENTE 197

zwischen zwei der Gebirgsriicken. Der hochste Punkt ist Sakesar, LSOOm. u. M. Der Berg-
grund besteht hauptsachlich aus Nummuliten-Kalkstein. Gegen S gibt es auch Karbon-
Kalksteine und kleine Partien von Jura-Gesteinen. Die Kalksteine liegen gewohnlich unbe-
deckt und kleinhiigelig in O-W Richtung ausgestreckt. Die Bodenarten nehmen hauptsach-
lich die Senken zwischen den Riicken ein und sind in grosser Ausdehnung kultiviert. Die
Waldvegetation dieser Zentraipartie der Salt Range ist ganz unl)etrachtlich und nur aus
einigen verkiimmerten H\pcranthcra und Bouibax heptaphyllum zusammengesetzt (Fleming
1853, S. 237). iJbrigens wird die hohere Vegetation von Fleming (S. 238) als "a low
bush jungle, formed in great part of Dodoiiaca Biiniuumiana ( Sunhetta) and Adhatoda
2'assica (Behikkur)" bezeichnet. Die letzteren sind fiir das Gebiet sehr charakteristisch. Das
Klima ist denijenigen von Kashmir ahnlich. Wiihrend der warmen Zeit ist die Hitze ganz
driickend; aus einer Angabe von Fleming (1853, S. 229) geht hervor, dass er darum Mitte
iXpril wegen der Hitze mit seiner Untersuchung aufhoren musste, wodurch die Arbeit ein
halbes Jahr verz<")gert wurde. Die Winde blasen in der Salt Range wahrend dieser Zeit von
OSO und wahrend der kalten Zeit von W. (Harwood 1926). Die Niederschlagsmenge
erreicht 250-500 mm. (Schott 1933).

1. Son Sakesar Kahar
(ca 750 m. ii. M.)

Wynne (1878) auf der Karte : Son Sukesur Kahur, im Text auch Samandar genannt.

Das Zuflussgebiet umfasst einen Teil der Hocheliene Son in den hijchsten Partien der
Salt Range. Die Topographie ist teilweise stark zerschnitten; die Hohenunterschiede zwis-
chen Gipfeln und Talboden erreichen hochstens 700-800 m. Der Berggrund besteht am See
aus Kalksteinkonglomeraten und Sandsteinen, oberem und unterem Pleistozan angehorend.
Die Hochgebirgsabhange bestehen hauptsachlich aus flinsteinhaltigen eozjinen Kalksteinen.
Die Bodenarten sind in der Niihe des Sees vor allem alte Salzablagerungen. Die Vegetation
der Umgebung des Sees ist als Macchien charakterisiert worden, ein Begriff der jedoch
heterogen ist. In der Nahe des Sees liegen die kleinen Dorfer Chitta, Uchhali u. a. Das
Zufliessen scheint nicht so stark zu sein, einen Abfluss gibt es nicht.

Das Seebecken ist seicht, deshalb wechseln Gnisse und Wassertiefe mit den Nieder-
schlJigen (Wynne 1878 S. 46). Die grosste mir bekannte Tiefe ist 8.8 m. Die Farlie des
Sees, die man nicht nach Furel-Ule I)estinimen konnte, ist, auf Grund einer grossen Menge
von Microcystis rosco-pcrsicinits graulich-hellrot. Ul^er die Farbe spricht mir Hutchinson in
einem Briefe: "Later in the year, according to Dr. Pruthi, this alga disappears and the only
plankton that he discovered consisted of Diaptoiiiiis salinus. The lake was still pink, as this
crustacean is a reddish species. The inhabitants maintain that the lake is always pink, except
in the late autumn when it may be whitish. I am not very clear whether this is true, but it is
possible that the calcium carbonate deposits represent a whitish phase occurring annually at a
time when a good deal of sediment is washed into the lake."

Das Wasser ist in okologischer Beziehung extrem salzig. Die Cl-Menge ist 34400 mg/1.
Die Summe von Alkalisalzen 73050 und SO4 17176 mg/1. Auch die Karbonatmenge ist
ungewohnlich hoch : 1276 mg/1 (vgl. die Tabelle).

Der Boden. Hieriiber liegen folgende Ijriefliche Angaben von Dr. Hutchinson vor:
"The bottom of Son Sakesar Kahar is very peculiar, consisting, I suspect, largely of ferrous
sulphide in which there seem to be thin bands of white material, probably calcium carbo-

198 iiociiAsiATisciir: binnenseksedimente

nate." Drei Bodcnproben, von 8.8 m., aus einer etwa 6 cm. langen Schlammwurst, mit clem
Rohriot von Naiimann heraufgeholt, sincl untersucht worden. Zwei von ilmcn gehiircn dem
sclnvarzcn sultidreichen Sediment, die dritte einer wcis.sen dazwischen liegenden 2 cm. unter
(icr Sedimentflache befindlichen Schiclit an. Ciewtiluilich sclieint es, als ob die .'iclnvarzen Sedi-
mentschichten etwa 2 cm. und die weis.sen < ^j cm. dick waren. In trocknem Znstand ist
die Farbe der Ol^erflachenprobe dunkelgrau nnd die der unteren i'nil>e geil)lich-dunkel-
grau. Die weissen Schiclitcn veriindern sich I)eim Eintrocknen niclit. Die iiCi-l'rt)lje galj
eine starke Reaktion (2 und 3) von siimtliciien I'roben.

Stnikturanalysc.

Probe Grobdetritus Feindetritiis MiiKralkunicr C.i-.Sclil.iiiini I'.vrit Chiliii

Ca 1 cm. u. P. <17o 71% 12% 12% 2% Z%

" 2 " " " 2% 20% 6% 64% ... 87o

" 5 " " " 67% 18% 13% <1% 1%

Der Feindetritus ist in den scliwarzen Scliicbten hyalin, in den oberen Schicbten dagegen
reicb an granen oder karotingefarl)ten KUnnpen; er ist kornig nnd n(x:kig und entbiilt eine
reichbche Menge kleiner l)azillen;ibnlicher Korper, etwa }<• x 1 /'. Die Tnschprobe zeigt
10-20% von Algenscbleim, mit bcstimmtem j\leln-gc\viciit in dtr unteren Probe. Die Mineral-
k("irner sind gut abgernndct oder stark splitterig, 15 m oder weniger. Der l^'eindetritus ist in
dan weissen Schicbten bellgelb und reicli an abgerundeten KcJrnern, die zum Teil eventuell
minerogen sind. Die Mineralkorner sind bier 15-25 /^, sebr gut abgenmdet, oder 5-15 f, gut
abgerundet und stark liclitbrecbend. Sie besteben mit Gewissheit aus Karlx)nat, nacli Dr.
Assarsson gewc'ibniicb aus Kalziumkarlionat. Auch Eisenkarbonatki'inier kommen vor,
obwohl es nicbt nKiglich war, die Proportion der.seiben festzustclicn. Die i\arbonatki"irner
untersclieiden sicb von den "Alineralki'irnern" (Ouarz, Feldspat, (ib'inmer und aiideren niclit
ausgefiillten Mineraben) durcb die bobe Licbtbrechung und den Gbmz und die lClx;nenbeit
olme Splitterigkeit der Flacbe. Die Karlxjnatkorner sitzen wie Trauljen auf den Feindetri-
tuskUunpclien. Auf dem Grobdetritus sind die KarbonatkcJrner grosser und etwas kantig,
sind aber docli reciit gut abgerundet. Das Pyrit ist gewobnlicb kugelformig und khunpen-
weise angesamnieit. Die scbwarzen Setbniente sind kalkreicbe I'eindetritusgyttja, die weissen
cliitinreicbe Kalkgyttja.

MikrofossUicnanalysc. Unter ik'n Diatomeen gibt es nur ein I'ragment von Cyiiibclla
aspera in der oljeren Prolje. Danel)en sind Cosiiiariuiii sj^. und andcre ( Idoropbyceenreste
angezeicbnet. An Pollen findet man "Ficca," unter denen ein ISO/j. grosses l^xemijlar,
"Sali.v," "Cbenopodiaceen-pollen," einen Querctts-tihnUchcn Pollen und einige, die einen bellen
y^agiw-Pollen ahnlicb sind. Was die Quantitat betrifft, so kommen nur ein paar von jedem
P'ollen auf den mm.'' Dies alles gilt nur fiir die dunklen Scbicbten. Die weissen Scliicbten
sind beinabe steril, nur ein C7(///»r«- fragment, mi'iglicbcrweise von eincm Rotatorien-lu, ist
angetroffen worden.

Mikrobiologiscb sind die Angabcn dieser Sedimente ziemlicli diirftig; sie deuten viel-
leicbt an, dass der See biologiscb zu einer Wiiste umgewandell wnrde, ein Scbluss, der jedocb
von den I""eldlx^obacbtungen widerlegt wird.

/^usaiiiiiii'iifiissung: Son Sakesar Kabar ist ein seicbter See mit einem sebr (1- und .SO^-
reichen Wasser, die Lagenfolge i.st gescbicbtet, die zwei Sedinunttypen der Scbicbtenreibe

IIOCirASIATISCriE BINNENSEESEDIMENTE 199

sind kalkreiche Feindetritusgyttja und cliitinreiche Kalkgyttja ohne MikrofossiliL-n. Mikro-
biolugisch herrschcn ^[icl■(lcysl!s nisco-prrsifiniis und Dial^toiiiiis saliiius vor.

Kashmir

Das Kashmir-Geljiet ist ein ctwa 7 :■: 11 Mcileii brcites Tal, scliarf al>gcgrt'nzt vmi den
Abhiingen der umgebenden Gel)irgszuge. Das Gebiet war friiher von einem Siisswassersee
eingenommen, wovon noch zerstrent liegende Sediniente mit Siisswassermuscheln zeugen.
Der Berggrund besteht aus Saiidsteinen, Otiarziten u. a des Panjal-Systems, Kalksteinen des
Karbon, Perm und Trias und im Norden aus andesitiscbeni Trapp (Lydekker 1883). Die
P>odenarten der Talebene sind muschelreiche Karewa-Sedimente, diejenigen der Abhiingc
Iiesonders gegen unten — Morane oder warwiger Ton (Sorlin 1927). Das Flachland ist mit
Reisfeldern und Obstgarten schon bewachsen. Der Boden ist so stark ausgeniitzt worden,
dass man im "Srinagar-See" (ich I)in nicht ganz sicher, welcber dem Srinagar naheliegende
See damit bezeicbnet wird) Garten auf Fltissen angelegt hat. Diese bestehen aus Schilf,
Zweigen usw., sind mit Erde bedeckt und fliessen auf ledernen Luftsacken, das ganze wird
durch in den Seeboden eingetriebene Stamme befestigt (Schlagintweit 1871, S. 411). An
Baunien stellt man Platanen, Pappeln und Walnussbiiume fest. Die umgelienden Abhiinge
sind mit Nadelwiildem von z. B. Abies Jl^cbbiaita, Picca Morinda, Finns lotif^ifnlia. P. Pence
und Taxus baccafa (Sorlin, briefliche Mitteilung) bewachsen. Es sind dalier hauptsachlich
die gegen N exponierten, also die nicht von der Sonne trockengelegten Seiten, die so
bewachsen sind. Hinter den Waldern erheben sich in der Feme die schneebekleideten Alpen-
gipfel. Das Klima des Kashmirgebiets ist ein warm gemassigtes Regenklima von warmem
und wintertrockenem Typus (Koppen auf der Karte in Supan-Obst 1927). Die heftigen
Sommerregen im Juli verursachen ein kraftiges Schneeschmelzen im Hochgebirge, also eine
Alpenfluss, das die Fliisse sehr schwer zu passieren macht (Schlagintweit 1871, S. 466). Im
Winter wird das Klima durch heftige, kalte Luftstromungen von den Hochgebirgen her ver-
schlechtert. Die jahrliche Niederschlagsmenge betragt etwa 1000 mm. (Schott 1933). Die
W'inde scheinen wahrend dieser kalten Jahreszeit von WN\\\ wahrend der Rlonsunzeit
al)er von OSO und auch etwa von NNO zu kommen (Harwood 1926). Dnch sei hier
bemerkt, dass die Topographic einen grossen Einfluss auf die ortlichcn Windrichtungen
ausiibt.

2. Lokut Dal Lake
(1582 m. ii. M.)

Die Grosse des Wassergebiets ist schwierig zu bestimmen ; es scheint jedocli recht
gering zu sein. Der Berggrund, der zwar nicht entblosst vorkommt, l^esteht aus einem
andesitischen Trapp, der dem Panjalsystem angehort (Lydekker 1883). Die Bodenartcn
sind pleistozener Ton aus der Karewa-Serie (de Terra, Isrieflicli). Der See liegt auf dem
ebenen, zum Teil versumpften Talboden; im Osten steigen die Gebirge steil empor. I^ie
Vegetation rings um den See Ijesteht aus Salts und Populus und im Osten aus Nadelwiiidern.
Das Wassergebiet zum grossen Teil Ijel^aut. Das Zufliessen ist aus Bod Dal; der AbOuss
geht siidwest warts l)ei Srinagar vorI>ei.

Die Tiefe des Sees ist gering, 1-2 m. Die Farbe des Sees ist nach Forel-Ule etwa XIV
und diejenige des Wassers am Abfiuss 15 mg. Pt/1. Die Secchi-Scheibe ist am Boden gut
sichtbar; der pH-Wert ist 8.2-8.5. Der okologische Standard ist siiss wie in den nahelie-

200 IIOCHASIATISCIIE BINNENSEESEDIMENTE

genden Seen. Cheniische Analysen licgen von "Rod Dal" vor. Jcdcn falls sind die Analysen
sichcr fiir samtliche Seen der Srinagar-Gegend repnisentativ. Die Sunimc der Alkaiisalze
betrrigt 9 nig/1. Die Cl-Menge ist 1.6 und der i\arlx)natgeliait 65 nig/1. Die einzige Bezie-
Ining, in der sicli das \Vasser dieses Sees von dcni der iil)rigen uutersucliten unttTsclieidet,
ist der relativ holie Uherschuss an Na^ CO;,: 14 nigv 1.

Der Bodcn ist zuni grossen Teil niit PotaDic'^cton l)e\vaclisen iind cnthidt einc arnie
/./»;;;(rra-Fauna. Ubrigens wachsen hier sowolil rnte /.('//(.v-iilnnicn zu Tausenden als audi
Trapa (Sorlin 1927). Die Oljerllachenschiclit ist hraun-grau und enthiilt Schneckenschalen ;
das konsolidierte Sediment ist helll)raun-grau luid enth.'ilt nur wenige Schalenreste. Die
(irenze zwisclien den erwiihnten Scliicliten wire! dnrch eine diinne graue Scliicht niarkiert, cs
war jedocli nicht nn'igjich, sie von (k'u unigL-l)cuden zu separieren. Nur die ( Jherlliiclicn-
sehiclit ist untersuclit worden. Die Pniln.* ist in trockeneni /ustand duukclgrau. Die- IK'l-
J'robe gal) keine Reaktion (0).

Slniktitranalysc. Grohdetritus SA'/( , {'"eiudL'tritus ?>?>'/<, Mineralki'irner 2^^ , Chitinreste
10%, Kalkscldaniin < \'/( und Diatomcen und i'yritkugeln < 1%. Der Gmlidctrilus
stamint event, aus rotaiiioi^cton lier. Der J'"eindetritus ist hyaliugelblich und relativ grni);
mit Immersion sieht man rundliche Kr)rner und vereinzelte hazilleniihnliche Korper 1 x 4 /i.
Die Tusehprobe zeigt < 10% Algenschleim. Die ]\Iineralk(")rncr sind aljgerundet und etwa
25/' oder 5-10 /'. Die ("bitinreste Ijcstelien ini allgenKMUun aus luisiiiiiui. Das Sediment ist
eine cliitinreiclie Grobdetritusgyttja.

.]filcrnf(>ssilicijaitalysc. Die Diatomcen sind die wiclitigsten der Alikr^fossiben. Unter
iluien douiiniert Goiiiplionciiia iiilricatiiin, 31% oder beinahe 50 St. pro mm.'' Danach
folgen L'yiiibclla /'arr'c/ \5'/r und Rpitlicmia zebra \'ar. porccllus 14'?, d. i. nieln- als 20 St.
pro mm.'' Unter den iibrigen MikrofossiHen bemerkt man l)esonders Spongicn-nadein, 22
St. pro mm.'' Es kommen audi I'fdiastnini duplex (4 St.), Eiiastrum sp.,Cosiimriiiiii sp.,
Gl<n(ilrieliia u. a. vor. Die Giiitinreste sind zuni grossen Teil 5oj;»/wa.-Schilde. Unter den
Pollen gibt es "Piniis," "Pieea" und "Oitereiis," von dem ersten 3 St., von den iibrigen 2 St.
pro mm.'' i'",in Riicklilick zeigt eine ausserordenttich reiche Mikrobiologie. \^on den Dia-
tonieen gil)t es 28 Formen ; 77'/f dieser Flora geh(")ren zu den Siisswasscrarten und 23%
sind fakuitativ Siiss- und ISrackwasserfnrnien. Die meisten, 90%, sind .\ul'\vudisli.irmen.
Typische Plankton formen kommen nicht vor.

Zusaniiiicnfassung. Lokut Dal Lake ist ein seichter, sebr vegetationsreidier See mit
siissem Wasser und ohne besonderen C'haraktcr; das Sediment ist eine cliitinreiclie Grobde-
tritusg)'ttja mit zahlreichen Diatomeen, besonders Susswasser-.Xufwudis formen.

3. Bod Dal Lake
(1582 m. ii. M.)

Daindli (1922) auf der ixarte: Dal.

Das Wassergebiet umfasst ein grosseres, gegeii NO 3-4000 m. hoch gelegenes Pergland
und den darunter liegenden Tallioden. Der Rerggrund ini TIodigebirgsgeI)iet bestelit aus
andesitischem Trapp (Lydekker 1883). Die Uodeiiarten sind nadi de Terra Sand und Ton
aus der Karewa-Serie ; zum grossen Teil sind sie versumpft. Die luihere N'egetation der
Umgebungen bestdit aus ,S"(;//.r und Popidiis und auf den ibilien ini O aus Nadelwaldem.
Grosse .\reale rings um den See sind bebaut. Das ZuHiesseii vnu dem Gebirge ini T) diirfte
betriichtlich sein ; das Abtliessen ist audi recht effektiv.

HOCHASIATISCHE BINNENSEESEDIMENTE 201

Der See hat wenigstens im W. eine Tiefe von etwa 4 m. FJie Farbe des Sees ist Forel-
Ule XV unci die des Wassers 10 mg. Pt/1. Die Sichttiefe ist 1.2 m. und der pH-Wert 8.5.
Der okologische Standard ist siiss; chemisch diirfte derselbe mit demjenigen von Lokut Dal
Lake iibereinstininien.

Der Bodcn ist mit einem dunkeigrauen, in trockenem Zustand grauen Schlamm bedeckt.
Eine Probe davon (von etwa 4 m.) ist untersucht worden. Das Material, mit dem Ekman-
Birge-Bodengreifer heraufgeholt, ist jedoch sehr heterogen. So kommen in einer Probe
wenige Diatomeen, in einer anderen zahlreiche vor. Die Produktionsziffern, die hier unten
angefiihrt werden, haben chiruni mir einen zienilich relativen Wert. Die HCl-Prol)e gab
keine Reaktion (0).

Struktiinmalysc: Feindetritus 68%, Mineralkurner 29% und Diatomeen 3%. Der
Feindetritus ist hyalin-gelblich und recht grobkornig; die Bestandteile sind abgerundet oder
bilden zahlreiche bazillenahnliche Korper, 1 x 3 /«. Die Tuschprobe zeigt nur sehr w'enig
Algenschleim. Die Mineralkorner sind gewiihnlich < 10/^; es gibt aber auch solche bis 25 /-t;
siimtliche sind aligerundet. Das Sediment ist eine diatomeenreiche Scliluffgyttja.

Mikrof()ssilicnanal\sc. Dominierend sind die Diatomeen, obleich, wie schon oben
ervvahnt wurde, die Mengen verschiedener Priiparate ziemlich variieren. Am zahlreichsten
Scheint Mclnsira granulata mit var. angustissiuta, zusammen 9% oder beinahe 500 St. pro
mm.^ vorkonimen. Dann folgen Cyiiibdla pan'a 12%), C. z'cntricosa 11% und Navicida
rhynchocephala var. rostcllafa 12%'. Im iilirigen wird auf die Tabelle 3 hingewiesen. Nur
ein "Picea" -PoWen pro mm.'"* ist festgesteUt worden.

Eine Zusammenfassung der MikrofossiHen zeigt 64 Diatomeen formen, die hijcliste
Anzahl in den untersuchten Seen. Die meisten sind Siisswasserformen, danel)en eine Brack-
wasserform, Nazncula pygmaca, und eine Art, die liauptsachlich in Salzwasser leljt, Navicida
sidinaruui, sind auch angetroffen worden. Prozentuell sind die l)eiden letzteren untergeordnet,
die typischen Siisswasserformen sind aljer nur 62%. Unter den Milieutypen dominieren
Aufwuchsformen mit 85%; daneben kommen audi u. a. 3% (+16%) Plankton formen vor.

Zusammenfassung: Bod Dal Lake ist ein seichter, vegetationsreicher See mit siissem
Wasser; das Sediment ist eine diatomeenreiche Schluffgyttja mit zahlreichen Siisswasser-
Aufwuchsdiatomeen und auch ein paar Brackwassertypen.

4. Sundar Khun
(1582 111. ii. M.)

Das Wassergebiet ist dasselbe wie das fiir Bod Dal Lake Ijeschriebene, da der See in
umiiittelliarer Nahe davon liegt. Die Sand- und Tonflachliinder sind mit Gebiischen von
Salix und von Populus und Plafanus bewachsen. Der Zufluss diirfte zum grossten Tcii (lurch
den Abfluss aus liod Dal Lake gebildet werden.

Der See ist etwa 5 m. tief und der Buden ist mit eiuL-ni fur den Bodengreifer undurch-
l.'issigen C/iara-Teppich bewachsen. Die I'^arbe des Sees ist Forel-Ule XII-XIII. Die
Secchi-Scheibe ist unter Boden vegetation auf 3 m. sichtbar; der pH-Wert ist 8.5. Der okolo-
gische Standard ist siiss; in chemischer Beziehung diirfte das Wasser dem von Bod Dal
Lake ziemlich iihulich sein.

Der Bodcn besteht aus einem bniunlich-grauen, in trockenem Zustand grauen Sediment
von sehr heterogener Beschaffenheit. Die HCl-Probe gab keine Reaktion (0).

202 IIOCMASIATISCIIE HINNENSEESEDIMENTE

SlruhtKniiuilysc: Cmlxletritus 237r, Fciiidftritus 56%, I\Iiiicr;ilk(')nKT W/v, Pyrit 2'/( ,
Chitinrestc 4'/v und Diatomccn 5%. Der (irolxlctritus ist aus Chara iiiul aus Phaiu'r<is;anK'n
(Ccratophyliuin ?) gebildet. Dcr I'ciiulctritus ist liyalini^xlhlicli, feinkornig-flockig und
enthalt nur vereinzclte hazillenformige K(')ri)er, 1 ■: 2 a*. Die Tuschprol)e zeigt etwa 10-20%
Algenschlciiii. Die i\Iincra!k<")mcr sind 2-10 /i und ahgenindct. Die Chitinrestc sind zuni
gnissten Teil BosiitinaSdvdK'n. Das Sediment ist cine diatoniccnrciclic Clrobdetritusgytt ja.

Mikrofossiliciianalysc. Die Diatunieen dciminieren, al>cr aurli die Desniidieen, und
Spongiennadeln konimen rclativ zahlreich vor. Unter den ersteren benicrkt man Cyiiibcllo-
Artcn, zum grossen Tcil C. pan'a 9%, Goinplinnciiia intricatnm 18%, Nitzschia ainpliibia
12% und Fragilaria construcns 20%-. Epithciiiia wird von /;. sorcx 5%, E. turgida 7%> und
E. zebra van porccUits 4% repriisenticrt. In einigen Praparaten sind diese Arten so zahl-
reich, dass es z. B. von E. zebra > 60 und von E. sorex 20 St. pro nun.'' konstatiert wurden.
Das Sediment ist al)er, wie schon hervorgeholK'n wurde, sehr lieterogen, was selbstverstand-
iicli audi in den /.allien ausgedriickt werdcn kaiin. X'oii den Spongien-nadehi kann man
15 St. und von den Kopfscliilden von Basiiiiini 4-5 St. i)ro nim.'' finden. Pollen von "Salix,"
"Picea," "Piiitis" und "Oiierciis" komnien niit 1-2 St. i)ro iiini.''' vor.

lune Zusaninienfassung der Diatonieen giht 44 Arten usw., wovon die nieisten Siisswas-
scrformcn sind. 12 St. lel)en auch ini lirackwasser. Von der ganzen 1 )iatiinicen-FIora
niachen die typischen Susswasserformen 71 ^f- aus. 88%. sind Aufwiu-lisfdrinen und 3%
Plankton fomieii.

Ziisaiiuiiciifassung. Sundar Khun ist ein kleiner, seicliter, vegetationsreicher See niit
siissem Wasser; das Sediment ist eine diatonieenreiche Grohdetritusgyttja init Siisswasser-
.\uf\vuclisdiatoinecn.

5. Manasbal Lake
(1584 m. ii. M.)

Das Zullussgebiet ist reclil scliwer zu bestininien, da der See auf deni I'lachlande am
P'uss der Ilochgebirge liegt. Das I-'lachland ist aus Sand und Ton der Karevva-Serie aufge-
baut; im Hochgebirge im Norden steht andesitischer Trapp an (T.ydekker 1883). Die
hi'ihere Vegetation der Umgebung des Sees besteht aus .V(//(.r und fapiiliis, wo der Boden
nicht bebaut ist; an den Abhiingen im Norden wachsen Nadelwalder. Das Zufliesscn scluint
nach der Karte von Dainelli (Taf. CL XTT) unterirdisch zu gelien ; dcr Abfluss geiit in den
Jlielum und dann in den \\'ular Lake.

Der See ist etwa 15 m. tief (Lydekker 1883).' Die b^arbc desselben ist l'"orel-Ule X
und diejenige des Wassers 10 nig. Pt/1. Die Sichttiefe ist 4 m. und der ]ill-\\'ert 8.5. Der
okologische Standard ist siiss. Die Cl-Menge ist 1.8 nig/l und die Summe der .Mkalisalze
23 nig/1. Der Karbonatgehalt ist 113 mg/1.

Der Boden ist von einem 1)einahe schwarzen, in trockeneni Zustand grauen Sediment
mit heilbrauneni Oberflachenlager 1>edeckt, das nach den Angaben reich an grol)em Pllanzen-
detritus ist. In der sehr heterogencn Prol^e von 12 m. Tiefe, die ich untersucht babe, ist
jedoch dieser ( irobdetritus sehr untcrgcordnet. Die IlCi-Prolje gab keine Reaktion (0).
Ubrigens konnten 13 'rubitRiden ])ro m." festgestellt werden.

Die Stniktiiraiialy.s'i- eines dunklen und festen dyttjaklumps gab; drobdctritus < 1%,

'Eighty-eight soundings in all parts of the lake failed tii disclose any depth greater than 12.8m. — G. E. H.

IIOCHASIATISCHE lilNNENSEESEDIMENTE 203

I'^eindetrilus 75%, Mineralkorner 18% und Diatomeen 7%. Der Feindetritus ist hyalingel-
blich, flockig, nur ein wenig kornig und enthiilt mir vereinzelte bazillcnformige Korper,
1 X 3 /i. Die Tuschprobe zeigt etwa 10% Algenschleim. Die Mineralkorner sind splitterig
oder etwas abgerundet, gewohnlich < 10 ft, aber auch einzelne bis 50 ;« sind wahrgenonimen
worden. Das Sediment ist eine diatomeenreich Feindetritusgyttja.

MikrofossiUcnanalyse. Die Diatomeen dominieren und unter ihnen besonders Cyclo-
tclla coiiita 14%, Cocconeis placcntula mit var. lincata 15% und CymbcUa panu 13%. Am
zahlreichsten kommt jedoch Synedra acus v. angnstissima, 300% (ausserhalb der Summe
berechnet) vor. Dieser Wert entspricht 1140 St. pro mm.^ Die Epitheima-Artcn machen
zusammen 10% oder I^einahe 40 St. pro mm.^ aus. Ausser Diatomeen gibt es kleine Des-
midieen der Gattungen Cosmar'mm, Spondylosmm und Staurastrum. Zwei Pollen von
"Picea" und "Quercus" pro mm.^ sind auch konstatiert worden.

Die Diatomeen bestehen aus 46 Formen, von denen 26 typische Siisswasser- und 3 Brack-
wasserformen sind. 80% sind Aufwuchsformen und 14% Planktonformen, dazu kommen
aber 300% Synedra acus v. angusHsshiia, die hier in der Summe nicht mitgerechnet sind, um
die anderen Formen nicht ganz zu iiberglanzen.

Zusanuiicnfassiing. Manasbal Lake ist ein mitteltiefer Siisswassersee ; das Sediment
ist eine diatomeenreiche Feindetritusgyttja mit Siisswasser-diatomeen von Aufwuchs-und
Planktontypus ; auch einige Brackwasserformen kommen vor.

6. IVular Lake
(1573 m. u. M.)

Das Wassereinzugsgebiet besteht aus dem grossten Teil des Kashmirbeckens und ist also
ausserordentlich gross. Im S und O von dem See ist das Land sehr niedrig und eljen, im N
und W aber reichen die hohen Gebirge (> 3000 m. ii. M.) beinahe an den See. Der Berg-
grund besteht hauptsachlich aus Sedimenten des Panjal-Systems (Sandsteinen, Konglomer-
aten und Ouarziten). In der Nahe des Flachlands steht andersitischer Trapp an. Die
Ouellen der nordostlichen Zufliisse finden sich in Formationen alt- und jung-palaozoischen
Alters bei denen Schiefer und Kalke vorwiegen. Das Flachland rings um den See besteht
aus einem sandigen Karewa-Ton. In der unmittelbaren Niihe des Sees gibt es Torfablager-
ungen (Lydekker 1883 S. 332). Die Umgebungen sind mit Salix und Popuhis bewachsen
oder bebaut. An den Abhangen im N und O wachsen Nadelwalder. Das Zufliessen muss
ganz bedeutend sein, da der See ein Sammelljecken fiir einen grossen Teil des Kashmirtals
ist. Bemerkenswert ist, dass viele der fliessenden GewJisser nicht bis im den See reichen,
sondern in den umgebenden Torfboden enden." Dies deutet darauf hin, dass sie nur wahrend
der Hochwasserzeit des Jahres wasserfiihrend sind (vgl. die Karte, Taf. CL XII von Dainelli
1922). Der Abfluss geht gegen SW durch den Jhelum.

Die Tiefe des Sees betrJigt wenigstens 5 m. Die Farhc des Sees ist Forel-UIe XVI uiul
die des Wassers 30 mg. Pt/1. Die Sichttiefe ist 1.5 m. und pH-Wert 9.0. Der ()kologische
Standard ist siiss. Die CI-Menge ist 1.7 mg/I, der Uberschuss an Mg CI2 2.3 mg/1. Die
Summe der Alkalisalze Ijetrrigt 18 mg/1. In chemischer Beziehung schliesst sich dieses

'Two well-dcfinod anil probably inimcrous smaller channels now conduct the Jhekini water into the lake.

G. E. H.

204 not II ASIA risen K iunnenseeskdimentf.

Wasscr^clir i;ut don aiulereii, im \\'assergel)ict von Jlielaiii wcitcr aufuiirts liegfiulcii
Seen an.

Dcr Bodcn ist niit I.oliis lunl Trapn bewachsen (Sorlin, brielliclic Mittciluu!;- ), uiul im
Sclilamm ist Tuhifcx allgemein. Das Sediment ist ol)crfl.-ulilicli luckcr, hraun-^raii uiul vdu
einer dunkelgraucn, festen Gyttja nnterlagert. Die rrul)en, von 4 und 5 ni. sind niit deni
Rohrlot von Nauniann aufgenommcn. Die untersuchten Proben von 5 m. sind aus dcr
Schlamniwurst von 0-1 cm. und 6-12 cm. hcrausgenommen. Beide sind in trockeneni Zustand
grau und die IlCl-Proben derselben geljen keinc Rcaktion (0).

Striikturanalyscn zeigen, dass die Zusanimensctzungen dcr [)eiden Proben mit cinandcr
anniiliernd uljereinstiminen; unter folgenden Ziffern beziehen sich die crstcn auf die Ober-
flachcnprobe : Feindetritus 70-69%, Mineralkorner 28-29% und Diatomeen 2-2%. Der
Feindetritus ist liyalin, ganz grob und cnthalt abgerundcte Korner, 2-5 /* in Diam., und bazil-
lenaluiliclie Korper, 1 x 2 m. Die Tuscbprolx^ zcigt sehr wenig Algcnscblcim. Die Mineral-
ki'irner sind klcine Splitter und grosserc, ganz abgerundcte Korner; gcwohnlich sind sie
< 10/*. Das Sediment ist eine diatomeenreiche Schluffgyttja.

Mikrofosstlienanalyscn hal^en eine sehr grosse Verschiedenheit der lieiden Prolx;n
gczeigt, liesonders hinsichtlich dcr absoluten Mcngen der .1/tVo,s-/;'(7-Zellen. In dcr ol)cren
Probe gibt es anniiliernd 500 St., in der unteren dagcgcn ctwa 1150 St. pro nini.^ Die eiit-
sprechenden Prozentwcrte sind 105 und 126% (ausser der Summe). Sie gehoren Mclosira
distmis V. alpigaia, M. gramilata mit v. angush'ssinia. Ubrigens gibt es in der oljeren Prol^e
16% Goiiiplioncnia infricafuni v. dicJiofoma. was in der unteren Probe fehlt. Dagegen sind
die Epithemien der unteren Prol^e zahlrcieh. Die Prozentsumme ist bier 20%, in dcr oliercn
Probe 4%. Ausser Diatomeen gibt es Pollen von "Picca," "Piniis" und "Sali.v," von jedcm
ein oder zwei pro mm.'

Die mikrobiologische Verandcrung, die die beidcn Proben angeben, enthidt cincn \\'ider-
spruch. In dcr unteren Probe sind die Siisswas.serformcn 48% (+129%), in der oberen
73% (+107%). Die fakultativen Siiss- und P.raekwassertypen sind resp. 43% und 20%
(+6%) und die reinen Brackwasser oder Salzwasserformen 2% und 4%. Trotz dcs frag-
lichcn Verhaltnisses zeigen die Proben aufwarts meiner Meinung nach ein Abnehmen des
Salzgehalts. Die Aufwuclisformen sind zu einer Zunahmc aufwarts — von 75% bis 82% —
geneigt. Dieses kann auf eine N'enneliruiig der Vegetation des Sees hindenten, kanii aber
audi nur l)losser Zufall seiii.

ZusaiiDiicnfassung. W'ular Lake i.st ein relative grosser, al)er seicliter, vegetations-
reicher Siisswassersee. Das Sediment ist eine diatomeenreiche Schluffgyttja mit Siisswasser-
diatomeen, besonders von Aufwuchstypus; aucli Brackwasserfornien kdinnien vor.

Ladak

(Plate XI, Figure 4)

Ladak ist ein sehr stark zersclinittenes Hochgebirgsbebiet in der Fortsetzung des Trans-
himalaya und Himalaya und wird von dem oberen Indus durchflossen. Das siidliche Ladak
setzt sich geologische aus zwei Ilaupt-Granitmassiven zusamen, die von nietamorphen
palaozoischen Koniplexen uniraiimt sind. In der umgegend des Panggong-Sees stehen verg-
neisste Schiefer, Griinsteine. niarmorisicrte Kalke und Granite an. Die Secufer sind ortlich
von quartaren Moranen und interglazialen Seetonen eingeraiimt. Am Siidabhang der Ladak-

IIOCIIASIATISCIIE BINNENSEESEDIMENTE 205

Kcttc, in (Icr Uniyeljunt;' der Seen Yaye- unci Mitpal-Tso erscheinen eozrinc l)is oberkreta-
zische Sandsteine und altere Ouarzite (de Terra, brieflich). Die Vegetationsverhilltnisse des
Gebiets sind sehr arm selig: im allgemeinen gibt es nur Taniarisken und Xerophyten oder
nicht einmal die geringste Vegetation. Das Kiinia ist ein Tnndren-Klinia (K(")ppen auf der
Karte in Supan — Obst 1927) mit < 250 mm. Niedenschlag per Jahr (Nordisk Varldsatla,s
1926).^ Die Stiirme sind zeitweise unerhort heftig und reissen grosse Mengen des Bodcn-
materials mit sich in die Luft. Die Temperatur ist sehr niedrig, und die Seen sind darum
nur in einem kurzen Teil des Jahres eisfrei. Ul)er Ororotse Tso z. B. schreibt Hutchinson
(1933) : "When visited on July 11, 1932, Ororotse Tso was still covered witli ice save at
the extreme edge, and it seems doubtful if it ever becomes entirely clear." Nach diesen
kurzen Angalien iil:)er die Seen konnte man vermuten, dass sie vollstandig wiistenahnlich
wjiren, eine Annahme, die im folgenden jedoch widerlegt wird.

7. Tso Morii'i
(4528 m. (i. M.)

Hedin (1909): Tschamomeril Lake.

Das Zuflussgebiet ist relativ gross unregelmassig zerschnitten und umfasst auch Geljirge
von > 6300 m. Meeresliohe. Der Berggrund bestelit aus Graniten und Kalkschiefern; die
Bodenarten sind pleistozener Ton, Sand und Schotter. Die Umgebung des Sees ist beinahe
steril, nur mit einigen Grasem und Xerophyten bewachsen. Unter anderm konstatiert man
hier Caragana versicolor (Schlagintweit 1874, S. 126). Ein Dorfchen mit kleinen, bebauten
Feldern liegt am westlichen Ufer. Die Zufliisse sind wasserreich ; der grosste Zufluss ist der
Yan, der — wie mehrere andere — von Schnee feldern und kleinen Gletschern im Hochgebirge
im Westen kommt. Auf der Karte Taf. CXXV von Dainelli (1922) weist der Sec auch
einen grossen Zufluss von dem grossen Haupttal im Siiden auf. Aljfluss fchlt.

Der See ist im allgemeinen 30-60 m. tief; die grosste Ijekannte Tiefe ist 74.7 m. in der
Nahe des Ufers, in einer Bucht im SO. Die Farbe des Sees ist Forel-Ule V und die des
Wassers < 5 mg. Pt/1. Die Sichttiefe ist 9 m. und der pH-Wert 9.0. Der okologische
Standard ist brackig. Die Cl-Menge ist 22 mg/1 und ein Uberscliuss an IMgCU von 30 nig/i
kommt vor. Die SOj-Menge ist f iir die Brackwasserseen von Ladak relativ gering :
517 mg/1; Mg SO, ist 199 mg/1 (Uberschuss). Auch die Summc der Alkalisalze ist niedrig:
788 mg/1.

Der Boden. Zwci Prolx^n, 1)cide von derselben Stelie, 48 m. u. Wfl., sind untcrsucht
wcirden. Das Sediment ist liell graugriin, in getnjcknetem Zustand hellgrau. Die HCi-
I'robe hat eine sehr starke Reaktion (4) gezeigt.

Die Sfnikfuranalysc der Proben gab: Feindetritus (hyalin-graulich) 43% und mineral-
korner 57%. Mikrofossilien u. s. w. also < 1%. Der Feindetritus scheint unter Immersion
flockig, allgemein hyalin, in dickercn Klumpen aber gell)licli ; darin finden sich einzelne bazil-
lenahnliche Korper, 5^2 x 2 /i, die sicher kcine Mineralkiirner sind. Die Mineralkorner sind
allgemein 10-40 jn, am haufigsten sehr scharfkantig und splitterig. 10-15% der Korner — ein
ungewohnlich grosser Teil — bestehen aus dunklen Mineralien. Das Sediment ist ein gyttjiger
Feinsand.

"At Leh, the nearest station to the lakes, the mean anin-al precipitation is 81 mm. (Smithsonian Misc. Coll.
79, p. 271).— G. E. H.

206 IIOCIIASIATISCHE lilNNENSEESEDlMENTE

Miki-ofossilicitanal\sc. Die Mikrofossilien Ijcstelien iiur aus Diatoineen, von denen 89%
Cyclotclla antiqiia (178 St. pro niin.^) sind. Es fulgt Diploncis cllipliai niit 6% (12 St.
pro mm.*)

An Diatomcenformen unterscheidct man 10 St., von dciicii () St. fakiillativ in siissem
mid brackigem Wasser leben. Von der ganzcn Flora sind jcdoch S9'/o typische Siisswasser-
formen, die zu den Plankton-oder Bodentypen gchoren.

Zusa>ii>itcnfassung. Tso Moriri ist ein grosser, vegetationsarnicr lirackwassersee; die
CI- und SOj-Mengen sind rclativ klein; das Sediment ist ein kalkreiclier, gyttjiger Fein-
sand mit besonders planktischen Siisswasserdiatomeen ; Drackwassertypen wurden nicht
angetroffen.

8. Khyagar Tso
(4672 m. u. M.)

Dainelli (1922): Tso Tasancuru.

Das Zuflussgebiet ist klein, da der See in einer Berggrul^e liegt, die im Siidcn an Tso
Moriri grenzt. Das Gebiet aus Graniten und Schiefern bestcht, die mit pleistozJinem Ton
und Sand bedeckt sind. Die Topographic ist stark zerschnitten, da der Hohenunterschicd
dcs kleinen Gebiets etwa 1350 m. betragt. Der Boden ist fast steril, nur cinige Xerophyten
kommen vor. Die Zuflusse sind kurz und steil; der grosste kommt von den Schneefeldern im
westlichen Teil des Gebietes. Al)fluss fehlt.

Die grosste bekanntc Tiefe des Sees ist 20.2 m.; hohere Vegetation kommt nicht vor.
Die Farbe des Sees ist Forel-Ule VIII und die dcs Wassers 5 mg. Pt/1. Die Sichttiefe ist
3 in. und der pH-Wert 9.5. Der okologische Standard ist brackig. Die CI-Menge ist 257
und die SO^-Menge 2069 mg/i. Der Karbonatgchalt ist relativ hoch; 525 mg/i. Der
Ulicrschuss an Na^, CO.., betragt 824 mg/I. Der Bcrggrund des Wassersystems scheint a1x?r
kalkfrei zu sein. Die Summe der Alkalisalze ist 3784 mg/1. Dieser Wert triigt ebenfalls
dazu l)ei, die Ahnlichkeit dieses Wassers mit dem von Pangur Tso zu erhohen.

Der Bodcn. Das Sediment ist auf 21 m. grau.schwarz ; nach Ilcraufholen wird cs ein
wenig rotlich, in getrocknetem Zustand gniulicli-hcllrot. Die HCI-Probe hat keine Rcaktion
gezeigt (0). Der Siebrest ist beinahe nur Gain}iiarns-Ch\t\n. Das Tiefwasser ist reich an
freiem HoS.

Sfniktjiranalysc: Feindctritus (graulich-hyalin) 93%, Mincralkorncr 6%, Chitinen-
reste 1%. Der Feindetritu^ scheint untcr Immersion komig-flockig, gelblich-hyahn und
enthalt vereinzelte, J^ x 1-2 /j- grosse Partien. Die Mineralkorner sind gewohnlich etwa 20/*;
man findet aber auch einzelne Korner bis 100 m gross. Sie sind abgenmdet, zuweilen aber
scharfkantig. Die Chitinreste sind graubraun, quadratfiinnig liniiert, Ijeinahe sicher aus
Cainnianis. Andere limnische MikrofossiHen, auch Diatomeen, sind nicht angetroffen
worden. Dagegen sind 'Tjcm"-Pollen 1 St. und "Sa/iV-Pollen 2 St. pro mm.* beobachtet
worden. Diese Pollenkomer sind genau von demselben Typus wie die in den Kaslunir-
Prol)en gefundenen. Das Sediment ist eine Feindetritusgyttja oder — besser gesagt — Algen-
gyttja, trotz der Abwesenheit deuthcher Algenstruktur passt dieser Name Ix^sser.

Die MikrofossiHenaimlysen deuten darauf hin, dass der See vollig steril ist. Die Gyttja
stammt aber, wie gesagt, zum grossen Teil aus Algenschleim, weshalb es recht wahrscheinlich

HOCHASIATISCHE BINNENSEESEDIMENTE 207

ist, dass der See reich an Myxophyceen ist oder — richtiger — gewesen ist.* Die Sediment-
probe ist ganz sicher nicht rezent.

Znsammcnfassung. Khyagar Tso ist ein kleiner Brackwassersee niit relativ hohen Cl-
und S04-Mengen und auch relativ hohen Karbonatgehalt ; das Sediment ist eine Feindetri-
tusgyttja mit reichlich Algenschleim ; Mikrofossilien sind nicht angetroffen worden.

9. Sla-rtsak-pitl: Tso
(4536 m. ii.M.)

Dainelh (1922) : Tso-Ciiim.

Das Zuflussgebiet ist im Verhjiltnis zum Areal des Sees relativ gross. Es weist grosse
Ilohenunterschiede auf und zwar besonders im NO (etwa 1460 m.) ; der grosste Teil aber
ist aussergewohnlich flach. Die Bergarten sind schieferige Granite und die Bodenarten
Schotter und Ton. Das Zufliessen ist relativ reichlich, der Abfluss im N gelit in den Tso
Kar. Die grosste Tiefe ist etwa lyi m. Der Boden ist mit einem dichten Teppich von
Bliitenpflanzen bewachsen. Die Sichttiefe ist 1.5 m. und der pH-Wert 9.6. Der okologische
Standard des Wassers ist siiss. Chemische Analysen fehlen.

Der Boden. Eine Probe von 1.5 m. ist untersucht worden. In getrocknetem Zustand
ist das Sediment gran. Die HCl-Probe zeigte keine Reaktion (0).

Strukturanalyse: Grobdetritus 7%, Feindetritus 36%, Mineralkorner 55%, Diatomeen
1%, Merismopedia etwa 1%. Der Grobdetritus ist unbestimmbar, diirfte aber aus den
Bliitenpflanzen des Sees stammen. Der Feindetritus scheint unter Immersion grobflockig,
gelblich und enthalt vereinzelte kleine, bazillenformige Korper, y^ xl p-. Die Tuschprobe
zeigte keinen Algenschleim. Die Mineralkorner sind 5-20 /*, aber auch Korner 50-60 /i
gross erscheinen; sie sind imnier sehr splitterig und scharfkantig. Das Sediment ist ein
gyttjiger Schluff.

Die Mikrofossilien. Quantitativ und qualitativ dominieren die Diatomeen in den Mikro-
fossilien. Am zahlreichsten sind Epithcmia zebra (34%) und Navicula rhynchocephala van
rostellata (14%). Daneben kommen Fragilaria brevistriata (9%) und F. pinnata (6%)
vor, beide nicht in der Summe mitgerechnet. Uljerhaupt dominieren die Aufwuchsformen.
Ubrigens sei bemerkt, dass Rhoicosphenia in < 1% vorkommt.

Die Anzahl pro mm.* ist z. B. fiir: Epitliemia zebra 75 St., Caloneis silicula 20 St.,
Cymatopleura solca 4 St., und Mensmopedia-ls.o\o\\\(t\\ sind etwa 20 pro mm.''' Ausserdem
bemerkt man vereinzelte "P/t"rfl"-Pollen.

Die Diatomeenformen umfassen hier 21 St. Unter diesen leben 12 St. in sowohl
sussem als auch brackigem Wasser. Sie machen 71% (+ 15%) der ganzen Flora aus. 79%
sind Aufwuchsformen und 11% (+9%) Bodenformen.

Zusaiiiiiiciifassung. Sta-rtsak-puk Tso ist ein kleiner, relativ vegetationsreicher Siiss-
wassensee; das Sediment ist ein gyttjiger Schluff mit recht zahlreichen Aufwuchsdiatomeen,
von denen die meisten fakultative Siiss- und Brackwasserformen sind ; typische Brackwas-
serformen wurden nicht angetroffen.

'Myxophyccae and Chlorophyceae are very abundant in tlic plankton. M 12 in. the water contained 5000 cells
and colonies per cc, tlie highest number encountered in Indian Tibet.

208 imciIASIATISCITE BINNENSEESEDIMENTE

^ 10. Tso Kar

(4527 m. ii. M.)

riedin (1909): Tsokr Tschumo Lake ; Dainelli (1922): Tso Cemnio.

Das Ziillus.sgebiet ist gross uiid umfasst audi dasjenigc von Sta-rtsak-puk Tso. Die
I)eiden Seen liegen in einem ausgedehnten und flachen Gebiet, von hohcn und slciicn Bergen
umrandet. Nur wenige Schneefelder, z. B. die auf Rukchen ini W und W vmi Shing-lnik
].a ini NW, sind bestiindig. Die Bcrgarten sind schieferige Granite und die Pxuk-narten Ton
und Schotter, die letzterer in Terrassen. Der See ist von salzigeni l-dadiland unigebcn, das
iin S grasbewachsen ist, sonst aber nur Xerophytenvegetation aufweist. Der /uduss sclieint
ganz nnl)etr:ichtlidi zu sein. Am wichtigsten ist das Wasser aus deni Sta-rtsak-puk Tso und
aus eineni Badi aus Shing-lnik La. Die meisten fiiessenden Gewiisscr trod<nen sdion aus,
wenn sie in das Flachland erreidicn.

Die Tiefe des Sees ist etwa 2 ni. und die Farbe Forel-Ule VIII. Die Sidittiefe ist nielir
als 2 ni. und der pH-Wert 8.9. Der okologische Standard ist sehr salzig. Die Cl-Mengc ist
11662 nig/1 und ein Uberschuss an Mg CI, von 9960 nig/1 komnit vor. Daneben beinerkt
man einen extremen Gehalt an SO4 : 35075 mg/I, es ist dies der hochste Wert in diesen Seen.
Der Karbonatgehalt ist 1633 mg/1 und die Summe von Alkalisalzen 61140 mg/L Ilinsicht-
lifli (k'S Wassertypus steht also dieser See Son Sakesar Kahar am nJichsten.

Der Bodcn. Line Probe von 2 m. ist untersuclit \vi^rden. In gelrockncteui Zusland ist
sie grauweiss. Die HCl-Probe gab eine starke Reaktion (3).

Sfrulduranalysc: Grobdetritus 1%, Feindetritus 70%, Mineralkorner 19%, Chitin
10%. Der Feindetritus ist hyalin, kornig-flockig, aber reidi an kleinen bazillenfc'irmigen
Korpem, Yi-l h- >- 1-4/* gross. Die Tusch prolje zeigte nur wcnig Algensdileim. Im
Priiparat scheint aber mehr cntbalten zu sein, da die KalkUimpcben l)ciuabe denselben
farblosen Eindruck geben. Die Mineralkorner sind 10-60/*, gerundet und nur selten etwas
scharfkantig. Die Chitinreste bestehen aus byalinen Stiickchen, miiglicherweise von Raupen-
liaut und Cladocerenextremitaten herstammend. W'ahrsdieinlich staminen sie von Artciiiia
lier, (k'r wichtigsten Komponente der Fauna. Andere Mikrofossilien, auch Diatomeen, sind
nicht angetroffen worden. Das Sediment ist cine chitinreiciic Feindctritu.sgytt ja. IMikro-
biologisch ist der See eine Wiiste.

Ziisaiimicnfassting. Tso Kar ist ein steriler, relativ grosser See niit eincni cxtrcmcn
Gehalt an CI und SO4; auch der Karbonatgehalt ist hodi ; das Sediment ist eine diitinrcidie
Feindctrilusgyttja, die im iibrigen steril und kalkreich ist.

11. Yayc Tso
(4686 111. (i. M.)

Daindli (1922) : Ichi T.so.

Das Zuflussgebiet ist im Verhaltnis zum Areal des Sees ziemlich gross. Im N grcnzt
es an das des Mitpal Tso. Es ist sehr stark zerschnitten, und die hohen Gebirgsabhiinge
fallen beinahe an das Seeufer. Die Bergarten sind Sandstein, Mergel, Konglomerate und
Granite. Die Gegend ist ausscrordentlich steril ; im Norden findet sich eine spar.same Vege-
tation von Ciriisern und Xerophyten. Im Talstrich, etwa 10 km. NNO von dem See, licgt das
Dorf Pialung. Das Zufliessen ist im Veriiiiltnis zum /\real des Sees sehr riirhlich, wt^halb

IIOCIIASIATISCHE BINNENSEESEDIMENTE 209

der Wasseraustaiisch sehr schnell von statten gehen muss. Der See fliesst in den nahgeleg-
enen Indus ab.

Die grosste Tiefe des Sees ist 18 m. und die Farbe Forel-Ule X. Die Farbe des
Wassers ist < 5 mg. Pt/1. Die Sichttiefe ist 4 m. und der pH-Wert 8.2. Der okologisclie
Standard ist siiss. Die CI-Menge ist nur 1 mg/1, es ist dies der niedrigste Wert in den vor-
liegenden Seen. Die Summe der Alkalisalze ist nur 15 mg/1. Der Karbonatgehalt ist
6v^ mg/1, wobei noch ein Uberschuss an Na^ CO3 von 25 mg/I vorkommt. Der allgemeine
Typus dieses Wassers ist dem der Srinagar-Seen am ahnlichsten.

Der Boden. Eine Sedimentprobe von 18 m. ist untersucht worden. In getrocknetem
Zustand ist diesell^e grau. Die HCl-Probe gab kein Reaktion (0).

Struktiiranalysc: Grobdetritus 1%, Feindetritus (graugrun) 88%, Mineralkorner 5%,
Diatomeen 6%, Pyrit und Chitin < 1%. Der Feindetritus scheint unter Immersion gelblich,
grobkornig und sehr reicli and bazillenahnlichen, 5^-1 ft x2-4a' grossen Korpern. Die
Tuschprobe zeigt keinen yVlgenschleim. Die MineralkcJrner sind 10-20 /*, gerundet oder
ebenso hjiufig splitterig. Das Sediment ist eine diatomeenreiche Feindetritusgyttja.

Bei den Mikrofossilicn dominieren die Diatomeen und unter diesen ihrerseits CyclotcUa
comfa, 73%. Daneben finden sich Campylodiscus noricus 3%, Gyrosigma attennatum und
G. kiifzingi, beide 2%, zuletzt Melosira arcnaria 1%. Von CycloteUa coiiifa kommen etwa
1960 St. pro mm.^ vor; die Anzahl variiert jedoch etwas in der heterogenen Probe. Von
den tibrigen Mikrofossilien sind "Ficca-" und "Chenopodiace"-PoIlen als etwa 2 St. pro mm.^
angezeichnet.

In mikrobiologischer Beziehung weist dieser See einen fiir die Ladak-Seen ausserordent-
lichen Reichtum auf. An Diatomeenformen wurden 34 St. gerechnet und unter diesen sind
12 St. fakultative Siiss- und Brackwasserformen. Von der ganzen Flora machen diese
Formen 9% aus. Die Aufwucbsformen sind 11 (+1)% und die Bodenformen 13 (4-3)%
aus.

Zttsaiiinicnfiissung. Yaye Tso ist ein kleiner hochalpiner Siisswassersee mit etwas
Karbonatgehalt; das Sediment ist eine diatomeenreiche Feindetritusgyttja mit Siisswasser-
Planktondiatomeen. Brackwasserformen sind nicht angetroffen worden. i\Iikrnl)iologisch
scheint der See sehr iippig zu sein.

12. Miip(d Tso
(4875 m. u. M.)

Dainelli (1922) : Mirpa Tso.

Das kleine Zuflussgebiet liegt zwischen dem des Yaye Tso und Panggong Tso. Es
ist ein einheitliches Becken, yon metir als 6000 m. hohen Gipfeln umgeben. Die Gesteine
sind Granite und kristalline Schiefer, die Bodenarten pleistoziiner Sand und Schotter. Das
Gebiet ist fast ganz steril ; nur eine arme Xerophytenvegetation gedeiht. Die Zufliisse sind
kurze, reissende Bache von den Ilnhen rings umber, zum Teil Abfliisse der Schneefelder im
SO. Ein Abfluss des Sees fehlt.

Die gn'Jsste Tiefe ist 23 m. die Farbe des Sees Forel-Ule XI und die des Wassers
< 5 mg. Pt/1. Die Sichttiefe ist 7 m. der pH-Wert ist 9.1. Der okologische Standard i.st
brackig. Die Cl-Menge ist 82 mg/1 und ein Uberschuss an Mg Cl.^ von 1 10 mg/1 kommt vor.
Die SO^-Menge betriigt 625 und die Summe der Alkalisalze 1011 mg/1, der Karbonatgehalt

210 IIOCIIASIATISCHE BINNENSEESEDIMENTE

364 mg/i. Trotz grosser Verschiedenheiten scheint dieses Wasser Ijetr. dcs allgemeinen
Typus deni von Tso Moriri am nachsten zu stehen.

Der Bodcn. Eine Probe aus der grossten Tiefe, 23 in. ist uiitersucht worden. In
getrockneteni Zustand ist sie hellgrau. Die HCl-Probe gab cine scliuaclie Reaktion (1).

Stniktiiranalysc: Grobdetritus 3%, Feindetritus (h3-alinliellgrau ) 90% und Mineral-
konier 7%. Der Feindetritus scheint unter Immersion hyalin und grobkornig. Er ist reich
an hyalinen, unrcgelmiissigen, abgerundetcn, 4 /j- grossen Kornern, und ein wenig stachelig.
Unsicher ist jedoch, oi) diese Kurner organisciien oder minerogenen Ursprungs sind. Bazil-
lenahnliche Korper sind nicht beoI)achtet worden. Die Tu.scliprobe zeigt keinen y\lgcn-
schleim. Die Mineralkorner sind etwas abgcrundet und nieistens 15/* gross; vereinzclte
grossere sind indessen konstatiert worden. Das Sediment ist eine Feindetritusgyttja.

Mikrofossilicnaiuilyse: Die Mikrofossilien werden von Cocconeis placenlula var. liiieata
52%, Diploims elHptica 22%, Amphora ovalis 11% und Mclosira arcnana 10% dominiert.
Die Cocconcis-An7.i\\\\ pro nim.^ betrilgt etwa 50 St. ; unter diesen sind aber mchrere so stark
destruiert, dass nur die Randpartie zuriickgeblieben ist und wie ein schwarzer Ring hervor-
fritt. Dasselbe gilt iibrigens audi fiir Diploneis. Von Melosira arenaria kommcii 10 St.
pro nim.^ vor.

Von ubrigen Mikrofossilien kommen nur 2 Pollenarten : Clienopodiace-Pollen unci eine
Art, die dem Quercus ahnlich ist, aber wahrsclieinlich von cincm Poptdus stammt, vo.

An Diatomeenformcn kommen 8 St. vor, von denen 4 typische Siisswasserformen sind.
87% der Flora sind fakultative Siiss- imd Brackwassertypen ; es ist dieser iiochste Wert in
den untersuchten Seen. Die Aufwuchsformen betragen 55% und die Jjodenfurmen 33%.
Ausserdem kommen hier 1% Plankton formen (Cyclotella comta) vor.

ZusiDiimcnfassiing. Mitpal Tso ist ein kleiner Brackwassersee mit einer durclisclinitt-
lichen Menge von CI, SOj und Karbonaten; das Sediment ist eine schwach kalkige Fein-
detritusgyttja mit sehr zahlreichen Diatomeen, von denen die meisten in Siiss — resp. Brack-
wasser als Anfwuclis — oder Bodenfnnnen lelien ; es wurdcn kcine Brackwasscrfurmfii
angetroffen.

13. Pangur Tso
(4329 m. ii. M.)

Es ist mir nicht moglich gewesen das Zuflussgebiet zu bestininicn: auf der Karte der
Yale-Expedition ist das Gebiet im O nicht kartiert, auf den Karten von Hedin (1909) und
Dainelii (1922) ist die Abgrenzung sehr unbestimmt. Das fragliche Gebiet scheint jedoch
relativ gross und stark zerschnitten zu sein. Im Westen liegt ein Flachland gegcn Tsaka-
iungpa. Der Berggrund besteht in den bekaimten Teilen aus kristalHnen Scliiefern und
Kalksteinen ; die Bodenarten sind pleistocane, schneckenhaitige Tone. Die Vegetation
bestelit aus spiirlichen Griisern an den Ufem und einigen xerophytischen Kriiutern. Die
Zufliisse kommen in den bekannten Teilen aus dem Ilochgebirge im S. Die Biiche von den
n(")rdlichen Hohen erreichen den See nicht. Der Abfluss ist ganz unbetriichtlich.

Die grosste bekannte Tiefe ist im NW etwa 10 m. Uber den ganzen untersuchten Boden
verbreitet sich eine hohere Vegetation (Pota)uogcfon). Die Farl^e des Sees ist Forel-Uie VI
und die des Wassers < 5 nig. Pt/1. Die Siclittiefe ist 8.5 ni. und der pll-Wcrt 9.6. Der
okologische Standard ist l>rackig. Die C!-Menge betragt 629 und die von SO4 1316 mg/1.

IIOCHASIATISCHE BINNENSEESEDIMENTE 211

Der Karbonatgehalt ist 842 mg/1; Na2 CO;, zeigt einen Uberschuss vou 1936 nig/1. Die
Summe der Alkalisalze betriigt 3180 mg/1. Der allgemeine Typus des Wassers ist dem von
Khyagar Tso am ahnlichsten.

Der Boden. Es gibt nur zwei Proben dieses grossen Sees und beide stammen von der-
selben Stelle, 9.5 ni. u. Wfl. Ob sie aus demselben Niveau in der Lagerfolge herriihren oder
nicht, ist mir unljekannt. Es scbeint mir aber sehr glaublich, dass die eine (Feld-Nr. 58)
mehr oberflachlich ist, die andere dagegen unmittelbar unter der Sedinientflache genommen
wurde. Beide Proben sind nun von einer hellen rotgelben Farbe; es ist aber unmoglich, die
natiirlicbe Farbe zu bestimmen. In getrocknetem Zustand sind die Proben graulich rotweiss.
Die HCl-Probe gab eine recht starke Reaktion (3). Die Feldnoten geljen nur Siebreste von
Potaiiwgcton an. Die Konsistenz ist stark elastisch und makroskopisch scheint das Sedi-
ment cine Algengyttja zu sein.

Strukturanalyse der beiden Proben

Obere Probe Fertiges Sediment

Grobdetritus < 1% . . . .

Feindetritus 95% 87%

Mineralkorner 2% 6%

Oscillatoria 2% 1%

Diatomeen 1% 1%

Ostrakodensclialen .... 5%

Cbitinreste < 1%

Charakteristisch ist also der Feindetritus. Er scheint ganz iiomogen zu sein. Die
Tuschprobe aber zeigt, dass etwa 90% der Detritusmenge Algenschleim sind. Der Feinde-
tritus erscheint unter Immersion vollkommen hyalin, sehr feinkornig und besteht aus 1 it-
grossen rundlichen und auch etwa 1 x 4 /^ grossen bazillenalinlichen Korpern. Die Mineral-
korner sind If- oder etwa 15/*; vereinzelte sind doch 100 /*. Daneben sieht man 2x4/*
grosse eirunde Korper. Mineralogisch besteht das Material aus Quarz. Die erwahnten
kleinen Korner, die eine starke Lichtbrechung besitzen und durch Milchsiiure leicht zu losen
sind, bestehen wahrsclieinlich aus Kalziunikarbonat. Ihre Frequenz ist — auf Grund ihrer
unbetrachtlichen Grosse und ihrer AhnHclikeit mit kornigeren Partien von Feindetritus —
schwierig festzustellen. Oscillatoria bilden allerdings nicht so grosse Volumenprozente, ihre
Frequenz ist aljer 200-250 St. pro mm.-'', was sehr bedeutend ist. Das Sediment ist eine
typische Algengyttja.

Die Mikrofossilienaiialy^e hat gezeigt, dass unter den Diatomeen Anoinoconcis poly-
gramma (79%) dominieren und danach Epitlieinia zebra var. porccllus (18%), I:)eide in dem
konsolidierten Sediment. Die absoluten Werte betragen 34-40 St. Anomoeoneis und
15-25 St. Epithcmia pro mm.^ Es ist bemerkenswert, dass so hohe Frequenzen nicht besser
mit der Strukturanalyse ausgedrvickt werden. Dieses berulit darauf an, dass oftmals von den
ganz grossen Diatomeenschalen vielfach nicht mehr als die Raphe iibrig geblieben ist, die
anderen Teile sind ganz aufgelost worden. Dieses Verhaltnis ist betreffs gewisser kalkreicli-
erer Sedimente nicht seiten. Von den iibrigen Mikrofossilien ist nur Oiiadnila subglobosa
(20 St. pro nim.^) in dem konsolidierten Sedimente hervorzuhelien.

212 IIOCIIASIATISCIIE BIXXENSEIiSEDIMENTE

In raikrobiologischer Beziehung wird der See durch die ausserordentlich starke Spezial-
isieruiig cliarakterisiert : 2 Arten — Anomoeoncis polygraiiuua und Oscillatoria — dominieren
vollstiindig. Die letztere Art diirfte der wichtigste SediiiK-ntproduzent sein, denn beinahe
die ganze Feindetritusmenge Ijesteht aus Algenschkim. I'nter Diatomeen gibt es nur 4
Arten, von denen cine oder 79% der ganzen Diatomeenflora eine Brackwasserforni (und
Bodenform) ist.

Zusaniiiienfassung. Pangur Tso ist ein ziemlicii grosser Brackwassersee niit ctwas
Potauiogeten; das Wasser ist besonders reich an CI, SO^ und Ivarbonaten; charakteristisch
ist ein holier Naa COs-Uberschuss ; das Sediment ist eine sehr kalkreiche Algengyttja init
einer stark spezialisierten Mikrollora von hanptsachlich Anomoeoncis poIygra)nina und
Oscillatoria, die erstere Art ist eine Brackwasser-Bodenform.

14. Fanggong Tso

(4241 m. ii. M.)

Schlagintweit (1874): Tschomagnalari ; Dainelli (1922): Pancong-Tso.

Das Zuflussgebiet ist auf der Karte der Yale-Expedition im Osten nicht abgeschlossen ;
nach jilteren Untersuchungen (Huntington 1906, Hedin 1909) wissen wir al>cr, dass der See
der k-tzte einer Reihe ist, die etwa unter 81 (ireenwioli-Lange (in der Nalie von Jai-
Tonghok auf der Karte Hedins) beginnt. Das Gebiet, das jedcnfalls sehr gross ist, ist ein
tektonisches Becken (de Terra 1934) iu:d zeigt grosse topographische \''erschiedenheiten.
Die Gipfel erreiclien > 6600 m. ; l:)esonders im Norden sind sie mit Schnee oder Gietschern
Ijedeckt. Der Berggrund besteht aus paliiozoischen Schiefern, auch Mergelschiefern, niit
Griinsteingebieten. Die Bodenarten in den TJilern sind pleistoziine schalenfiihrende Tone
uder Sand. In kleinen Gebieten ist der Schotter aus ausgefalltem Kalk zusannnengesintert.
Wanne Quellen springen einer Verwerfungslinie entlang, in der Niiiie des nordlichen Ufers,
liervor. Die \'egetation bestelit liauiitsiiclilicli aus xerophytisclien Kniutern. Ausserdem
komnit ortlich Salices, Rosa und Myricaria vor. An dem siiillichen Ufer lit-gcn einigc kleine
Dorfer (Spangmik, Man und Alirak), von unbetnichtlichen Gerstenfekk-rn umranik-t.
Weiter oben in dem Tsaka-Lungpa-Tal im Siiden liegt das Dorf Chushul, von kleinen
l)el3auten Feldern umgeben. Das Zufliessen muss wegen der Grosse des Zuflussgebiets ziem-
lich bedeutend sein, sofern nicht das Verdampfen sehr gross ist. Wahrscheinlich ist jcdoch
dies der Fall, denn der See ist heute ohne Abfluss. Friiher ist er gegen NVV in den Shayok
(Shejok auf der Karte Hedins) abgeflossen.

Die grosste bekannte Tiefe des Sees ist 51.8 m. und liegt an der cistlichcn Seite in der
Nahe des Ufers. Nordlich von Spangmik ist 47.7 m. gelotet. Nach niir zur Verfiigung
stehenden Karten, be.sonders der Yale-Karte, zu beurteilen, ist die Tiefe der verschiedenen
Teile des Sees ganz regelmiissig; sie wird aber ortlich von kleinen jiihen .Vbhangen (vgl.
I)esonders in der nordwestlichen Bucht) abgebrochen. Die h'arbe des Sees ist Forei-L'le Ii
und die des Wassers < 5 mg. Pt/1. Die Sichttiefe ist 11 m. und der pH-Wert 9.3. Der
okologische Standard ist allerdings brackig, zeigt aber eine ausserordentlich hohe Cl-Menge :
3587 mg/1. Auch der S04-Gehalt, der 1553 mg/1 betriigt, ist der niichst hiJchste dieser
Seen. Die Sumnie der Alkalisalze ist 10039 mg/1. AIs allgemeine Charakteristik des
Wassers diirfte man sagen konnen, dass es einen Zwischentypus zwischen den brackigcn
Gewassern und den extremen Salzseen Son Sakesar Kahar und Tso Kar bildet.

nOCIIASIATISCIIE BINNENSEESEDIMENTE 213

Dcr Bodcn. Aus diesem grossen See gibt es nur 2 Proben : von resp. 31 m. in dem
nordwestlichen Teil und 46 m. vor dem Dorf Man am siidliclien Ufer. Das Sediment von
Panggong- Tso ist an der Oberfljiche hellbraun und etwa 3 cm. darunter grau. In getrock-
netem Zustand ist die Prol^e von 31 m. gelljlich-hellgrau und die von 46 m. grauweiss. Die
HCl-Pro1:)e zeigt eine recht starke Reaktion, 1-2 fiir die Prolje von 31 m. und etwas starker
fiir diejenige von 46 m. : 2. Der Siebrest enthalt Fragmente von Gaiiniiaru^, der frei
schwinimend "in tlie middle water" (Hutchinson, brieflich) lebt.

Sinikturtinalysc dcr hcidcn Proben

31 m. 46 m.

GroI)detritus ....

Feindetritus 30% 81%

Mineralkorner 63% 17%

Diatomeen 7% ....

Chitin 2%

Pyrit < 1% < 1%

Der Feindetritus ist in der Prol^e von 31 m. hyalingraulich und in derjenigen von 46 m.
grau. Die Verschiedenheit der Feindetritusmenge pro Volumen ist ja eine natiirliche Folge
der litoralen Zunahme des minerogenen Materials. Dasselbe Gesetz reguliert iibrigens die
Verteilung der Diatomeen. Unter mimersion scheint die Feindetritusprobe von 31 m. hyabn,
teils sehr feinkornig und teils recht grobkornig und daneben ganz reich an bazillenahnlichen
Korpern (1 x5/^). Die Feindetritusprobe von 46 m. scheint hyalin, feinktirnig und enthalt
reichlich von rundlichen, etwa 1 /^ grosse Korner und vereinzelte bazilienahnliche Korper
(3^x2 /a). Die Tuschprol^e von 31m. zeigt nur wenig Algenschleim und diejenige von
46 m. gar keinen. Die Mineralkorner sind in der Probe von 31 m. gewohnlich 25-50 /a; dock
kommen auch zahlreiche solche von 100-200 ft vor. Dunkle Mineralien sind sehr zahlreich.
iibrigens sind die Mineralkorner sehr scharfkantig und splitterig. In der Probe von 46 m.
sind sie dagegen nicht so auffallig scharfkantig: die Grosse ist etwa 10-20 m. Die Chitin-
reste in der Probe von 46 m. diirften aus Gaiiuiiarus sein. Das Sediment von 31m.
liesteht aus gyttjiger Feinsand, reich an Diato