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 0 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
Ar.cocK, A. 1897. Description of a new species of Branciiipus fnim Calcutta. Jour. Asiatic See. Bengal. 65 (II) : 538-539.
Artom, C. 1906. II nuinero dei cromosonii e la maturazione dell' uovo dell' Artemia par- tenogenetica di Capidostria e dell' Arlcmia sessuata di Cagliari. Biologica, vol. 1.
1911a. La sistematica del genere Artciiiia in relazione col nuniero dei cromosomi
delle cellule sessuali e in relazione col numero e colla grandezza delle cellule soma- tiche. Biol. Centralbl. Bd. XXXI.
1911b. Analisi comparativa della sostanza cromatica nelle mitosi di maturazione e
nelle prime mitosi di segmentazione dell' uovo dell' Artemia sessuata di Cagliari (univalens) e dell' uovo dell' Artemia parthenogenetica di Capodistria (bivalens) ; con 3 Tavola. Archiv. f. Zellforschung. Bd. VII.
1912. Le basi citologiche di una nuova sistematica del genere Artemia. Sulla dipend-
enza tra il numero dei cromosomi delle cellule germinative, e la grandezza dei nuclei delle cellule somatiche dell' Artemia salina univalens di Cagliari, e dell' A. s. bivalens di Capo d'Istria. Arch. Zellforschg. Leipzig, 9: 87-113, 2 Taf.
1922. Nuovi dati sulla distribuzione geografica e sulla ])ir)li>gia delle due specie
(micro pircnica e macropirenica) del genere Artemia. Atti R. Accad. Lincei (5) 31: Sem. 2, pp. 225-227.
1926. Tetraploidismo e gigantismo. Esame comparative degli stadi postembrionali
deir Artemia salina diploide e tetraploide. 3 pi. Intern. Rev. Hydrobiol. 16: 51-80.
Baird, W. 1852. Monograph of the Family Branchipodidae, etc. Proc. Zool. Soc. London. 20.
1860. Description of Two New Species of I'jituniostracous Crustaceans from India.
Proc. Zool. Soc. London, 28: 445-446.
B.\KN.\uii, K. LI. 1929. Contributions to the Crustacean Fauna of South .Africa. Nu. 10. A Revision of the South Africa I'ranchiopoda (Phyllopoda). With ?>}> text figures. Annals South y\frican Mu.seum. 29 (I) : 181-272.
I).\i)AY (de Dees), E. 1910. Monographic systematique des phyllopodes anostraces. Ann. des Sc. Nat. (ser. 9), 11: 91-489. 89 fig.
1913. Az eddig ismert kagylos levelFibu rakok attekintese. Math, es Termt. Ert.
Budapest. 31: 559-601.
1915. Monographic systematique de Phyllopodes Conchostraces. Ann. Sci. Nat.
(.ser. 9) 20: 39-330.
• 1925. Monographic systematique de Phyllopodes Conchostraces. Suite. Ann. Sci.
Nat. (ser. 10) 8: 143-184.
1926. Monographic systematique de Phyllopodes Conchostraces. Suite. Ann. Sci.
Nat. (ser. 10) 9: 1-81.
(iuocuuwsKi, M. 1896. Ueber eine neue im Susswasserlebende Species von Artemia. Verhandl. zool. bot. Ges., Wien. 45 : 95.
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, 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, 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
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X\'III.' Bremen.
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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 |
0 |
|
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 |
0 |
|
50 |
330 |
330 |
262 |
0 |
|
58 |
308 |
333 |
242 |
0 |
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 0
|
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 0 |
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 |
|
0 |
25 |
29 |
|
0 |
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. Nat. 38. p. 1.
Thorpe, V.G. 1891. New and Foreign Rotifers. J. R. Micr. Soc. London. 18<)1. p. 301.
TscHiT.UNOFF, N. L. 1921. I'Ikt das Plankton des Nordlichcn Teiles des Kaspisees. Arb. P.iol. W'olga Station. 6. ]>. 107. Russian text, p. 159, German summary.
\'ikii-:lx, j. 1916. Recherches sur le Plancton des Lacs du Jura central. Ann. Biul. Lacustre. 8. p. 5.
Webek, E. F. 1898. b'aune rotatorienne du bassin de Leman. Rev. Suisse Zool, 5. p. 263.
Weber, E. F., and Montet, G. 1918. Rotateurs. Cat. Invert, de la Suisse, fasc. 11. Geneva.
Wesenberg-Luni), C. 1S)30. Contrilnitinns to the Biology of tlie Kotifera. I't. 11. The Periodicity and Sexual Periods. D. Kgl. Danske \'idcns.-selsk. Sl<r. ('•) 2. no. 1.
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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THE TUTTLE, HODEIIOUSt » TAYLOR COMrANY. NEW HAVEN, CONN.
MEMOIRS OF
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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.
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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 : 0 == 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 Diatomeen, und das von 46 m. eine schluffige Feindetritusgyttja.
Die Mikrofossilicnaitalysc der Probe von 31 m. hat einen relativ grossen Artreichtum der Diatomeen gezeigt, die iibrigens zum grossen Teil lebendig waren. Schon die Struktur- analyse gab ja 7% Diatomeen in der Probe von 31 m. Der grosste Teil dieser Prozente wird aus dem relativ grossen Surirella ovalis gebildet, der doch nur 9% des Diatomeenbe- stands macht, oder etwa 200 St. pro mm.^ Am zahlreichsten sind Cyclotclla coiiita: 47% oder etwa 1030 St. pro mm.* Unter Brack- oder Salzwasserformen bemerkt man Aiiiplii- prora paludosa, Aiioiiioconcis poly gramma (weniger als in Pangur Tso), Epithciina tiirgida var. IVestermanni, Navicula pcrcgrina, N. salinarum und Rhoicosphcnia cun'ata. Die meisten sind jedocli in anderen Gegenden auch in siissem Wasser angetroffen worden. In der Prolie von 46 m. gab es keine Diatomeen.
In mikrobiologischer Beziehung gibt es eine sehr grosse Verschiedenheit der beiden Proben: in der Probe von 31 m. leben 22 Diatomcenarten, in derjenigen von 46 lu. ist kein cinziges Fragment davon vorhanden. Es ist jedoch nicht muglich zu sagen, ob die Proben
214 IIOCIIASIATISCHE BINNENSEESEDIMENTE
gleiclizei.tig und also vollkoinmen vergleichbar sind. Von den 22 Arten sind nur 4 typische Siisswasserfornien, 4 sind Brack wasserfonnen und 2 lel>en auch im Salzvvasser. 58% der ganzcn Flora leben niir iin siissen Wasser, die iibrigen ertragen oder verlangen eine stJirkere oder geringere Salzmenge. In Milieul)eziehung ist der grosste Teil der Mora — 47% — Plankton; 23% sind Aufwuchs — und 16% Bodenfornicn."
Zusammcnfassung. Panggong Tso ist ein sehr grosser Brackwassersce, der tlen Salzwas- serseen nahe stelit; das Wasser ist sehr reich an CI, SO4 und Karbonaten; die gefundenen Sedimente gehoren zu zwei Typen : gyttjiger Feinsand und schluffige Feindetritusgyttja, beide sehr kalkreidi, letztere viJllig steril. Nur etvva die lliilfte der Diatoniecn in dem Fein- sand — die zuni grossen Teil kbcndig waren — gehiirt zu reinen Siisswasserfornien; die anderen leben audi luler nur in l)raikigeni oder salzigeni Wasser. Die nieisten sind Planktonfornien.
15. Ororotse Tso (5297 m. {i. M.)
Das Zuflussgebiet ist sehr kkiii und unifasst nur die uingebenden Ilochgebirgsteile. Das Becken ist ein Kar. Der Berggrund besteht aus Granit uiul die Bodenarten sind Moriinen in grossen Mhcngen. Die Vegetation ist sehr armlich ; dominierend sind Griiser und Kruciferen. Das Zufliessen geschieht durch kurze, reissende Alpenbache. Der Abiluss diirfte gegen Norden verlaufen.
Die grosste bekannte Tiefe ist 13.4 m. Die Farbe des Sees ist Forel-Ule XII und die des Wassers 5 mg. Pt/I. Die Sichttiefe ist 5 m. ("hole in ice") und der pTT-Wert 7.1. Der ()kologische Standard ist siiss. Hinsichtlich dieses Sees diirfte hervorgehoben werden, dass die Menge fester Stofife iiberaus gering ist oder nur 78 mg/I, was bei dem Studium der .\nalyscnwerte zu beriicksichtigen ist. Die Summe der Alkalisalze ist 14 mg/1 und der Kar- bonatgehalt betnigt 32 mg/1; daneben bemerkt man einen Uberschuss an Naj CO;j von 5 nig/1. Zuletzt ist eine Cl-Menge von 4 und eine SO^-Menge von 3 mg/1 zu erwahnen. Diese Werte sind unerwartet hoch, weshalb icli an die oben angefiihrte Reservation erinncrn muss. Der allgemeine Typus des Wassers ist dem von Lokut Dal T.ake und Yaye Tso am iilinlichsten.
Der Bodcii ist mit Algenkolnnien l)csetzt. Ini Alikroskop sieht man vereinzelte Chulo- /'/(oro-Zweigc. Mit dicsem Vorkonimen slinimt iibrigens die beobachtete Diatomeenflora gut iiberein. Eine Sediinentprolje ist l>ei 13.4 m. geiiommcn. In getrocknetem Zustand ist sie grau. Die HCl-Probe zeigt nur eine unbetrachtliche Reaktion (0-1).
Striikturanalyse: Feindetritus (griiulich) 85%, Mineralkorner 11%, Diatomeen 3% und Chitin 1%. Der Feindetritus erscheint unter Immersion hyalin-gelblich (in dickeren Khimpen) ; er ist grobkornig und hat ein zerrissencs und si)litterigcs Aussehen. Vx cnthiilt liazillenahnliche, 1 xS-G/^ grosse Korper. Die Tuschproljc zeigt keinen oder hiichstens nur wenig Algenschleim. Die Mineralkorner sind 10-20/*, vereinzelte doch 100/*. Die mei.sten Korner sind splitterig und scharfkantig. Das Sediment ist cine diatomeenreiche Feindetri- tusgyttja.
Die Mikrofossilicnaiialyscn zeigen, dass die Diatomeen die cinzigeii Fossilien sind und
'There is little douI)t tliat by far llic greater part of these diatoms arc derived from an iiitcrglacial lake deposit, V. i. p. 235.— G. E. H.
IIOCIIASIATISCIIE BINNENSEESEDIMENTE 215
dass cs hicr iiur wcnige Arten gibt. Diese sind aber sehr zahlrcich vorhanden. Man bcmerkt hier l>esondcr.s Cocconeis placcntitla var. lincata (42%), Cyclulclla antiqiia (20%), Rhoico- sphcnia curvata (17%) und Ainpliora oralis var. pcdiculus (15%). Daneben gibt es audi z. B. Melosira arenaria in 1 % oder etwa 300 Zellen pro mm.^
Die Mikrobiologie des Sees — der nur etwa 14 Tage im Jahr eisfrei ist (Hutchinson 1933) — ist unerwartet reich entwickelt. 15 Diatomeenarten sind angetroffen worden, von denen die Hiilfte typische Siisswasserformen sind. Von der ganzen Flora bilden diese nur 22%. 77% sind Aufwuclisfornien und 20% fakultative Boden- und Planktonformen (Cyclotella antiqiia).
Zusaiiiiiicnfassung. Ororotse Tso ist der allerhochste der untersuchsten Seen ; er ist von hochalpinem Typus ; das Wasser ist siiss und ohne besonderen Charakter ; das Sediment ist eine schwach kalkige diatomeenreiche Feindetritusgyttja, deren Diatomeenformen zum gn'issten Teil in siissem und brackigem Wasser leben; die nieisten sind Aufwuchsforinen ; eine ungewohnlich grosse Menge sind aber fakultative Boden- oder Planktonformen.
Zusammenfasscnde Bemerknngcn
Die vorstchende Untersuchung der Bodensedimente veranlasst aus verschiedenen Gesicht- spunkten einige zusammenfassende Bemerkungen. Die untersuchten Seen gehoren zu 3 ganz verschiedenen Gebieten: Salt Range, einem salzreichen Becken innerhalb der Bergkette auf der Grenze des nordwestindischen Flachlands, dem Kashmirtal, einem sehr vegetationsreichcn von hohen Bergen umschlossenen Tal, und Ladak, einem stark zerrissenen Alpengebiet. Letzteres gehort zu den Fortsetzungen des Transhimalaya und des Himalaya, wahrend Kashmir zum Himalaya gehort. Die verschiedenartigen Lagen dieser drei Gebiete bestim- men die grossen Ziige der Typen und der Entwicklung der Seen. Einige Einzelheiten darin konnten nicht festgestellt werden konnen, da die Proben gar zu sporadisch genommen warden. Gewisse Schlussfolgerungen konnen ai>er dennoch hervorgehoben werden.
Das Sahproblcin. Seit alters her wusste man, dass Salz- oder Brackwasserseen in diesen Gegenden vorkommen. Schlagintweit (1874, S. 105) sagt im Bezug darauf, dass die Seen immer salziger werden. Sie miissen sich also zu konzentrierten Salzbecken entwickeln. Bei anderen Forschungsreisenden ist die Auffassung hieriiber nicht so ausgepragt. Hedin (1917) gibt eine Darstellung, die deutlich zeigt, wie die Auffassung iiber die betreffenden Seen zu verschiedenen Zeiten wechselte. Er gibt (1917) nach alteren Angaben eine Zusam- menfassung der Abflussanderungen von Manasarovar und Rakastal, wodurch eine Tendenz zu Periodizitat in Erscheinung kommt. Er scheint jedoch der Ansicht zu sein, dass die Seen im allgemeinen austrocknen (1907, S. 603) ; der Salzgehalt kann zwar auch in dem- sclben See jahrlicJi variieren (S. 601). Dasselbe ist iibrigens von Decksbach (1924, S. 275) fiir das Turgaigebiet bewiesen worden. — Nach seiner nachsten Reise hebt Hedin hervnr (1917, S. 173), dass wir nicht wissen, ob das Austrocknen der Seen fortsetzt; er scheint jedoch der Ansicht zu sein (S. 179) : "When it has once reached its minimum it will jirob- ably again return towards a new maximum." Dieses gilt fiir den \\'asscrstand und also indirekt auch fiir den Salzgehalt. — De Terra und Hutchinson (1934) habcn ebenfalls eine Zusammenstellung der Wasserstandsangaben, und zwar besonders fiir Panggong Tso gege- ben. Und dank besonders gliicklicher Umstiinde ist es ihnen gelungen, diese mit einer Kurve darzulegen. Die aus diesem Grunde ausserordentlich wichtige Arbeit, zeigt deutlich, dass
216 IIUCIIASIATISCIIE BINNKNSKKSKDIMKNTE
I'aiijiyoug Tso seit tier Mitte des voris^cn jahrlnindcrts starke und rcgelmiissiijc W^isser- standsvariationen aufweist. In den Jahren 1860-70 war der Wasserstand 5.9 ni. niedriger als jetzt, cin Umstand dcr die Salzkonzcntration betnichtlich l)eeinflusst haljen nuiss. Dieses VerhJiltnis ist auch durcli Vergleichung mil Analyson, die von Henderson und llunu' 1871 ansgefiihrt wurden, nachgewiesen wordcn. Das Resultat wird von de Terra und llutcliinson (1934, S. 316) folgendcnnassen angegeben: "A comparison of the chloride contents of the lake as given by him with that found in 1932, indicates that the lake has been diluted l)y fresh water by 16 per cent, of its former voUune since that date. This increase in volume is com- parable to the increase of 18 per cent, in area of the cross-section at Man indicated alxne. The whole clearly indicates that within 112 years, Tso Pangong has experienced one low and two high levels." Am Ufer von Tso Moriri steht eine Manen-Kiste, die nun vom Eis verschoben worden ist (vgl. Plate 3-4 bei de Terra und Hutchinson). Da die Til:)etaner unter keiner Bedingung' eine solche Kiste links passieren (z. B. Sorlin 1927), kann man nach tier Pliotographie mit Bestimmtheit aussagen, dass die Tage derselben eine ^Vasscrstands- steigerung von wenigstens ein paar m zeigt. Audi in Khyagar Tso, Mitpal Tso und Pangur Tso ist eine solche Steigerung von den erwahnten Verfassern konstatiert worden. Sie halx^n nach meiner IMeinung einleuchtcnd gezeigt, dass mehrerc der Seen von Ladak nach der Mitte des vorigen Jahrhunderts sowoiil Senkungen als Erholiungen des Wasserstands aufweisen, was — nach ihrer Meinung — von A'ariationen der Niederschlagsmenge abhangig ist. Obgleich das fragliche Verhiiltnis ganz wahrscheinlich ist, mochte ich doch zulctzt auch Hedin (1917, S. 192) anfiihren: "But the data we possess regarding the behavior of the lake are too meagre to allow us to draw absolutely reliable conclusions, and at our present state of knowl- edge, it would prol)alily ])e impossible to say whether the monsoon rains or the cold weatlier storms in N. W. India are the most important factor affecting the rise and fall nf tlic lakes, and the volume of waters in the rivers."
Die hier erwahnten Verhaltnisse, also periodisclie Wasserstandsvariationcn und drunit folgende Salzgehaltanderungen, miissen selbstverstandlich den Hintergrund, ausmachcn, gegen den diese indisclien Seen betrachtet werden sollen. Dadurcli knnnen Verschieden- heiten zwischen Seetypus und Mikroflora dieser nicht schr genau gcsammeltcn, friihcr unter- suchten Sedimentproljen, erklart werden.
Das Hauptprinzip des geographischen Vorkommens der Seen I;)etreffs ihres okologischen Standards ist folgender. Die in einem Wassersystem hfichst gelegenen Seen sind Siisswas- serseen, tiefer unter liegen die Brackwasserseen und zu imterst die Salzseen, vorausgesetzt dass das ganze Seesystem in ein abflussloses Becken endet. Beispiele dafiir sind Sta-rtsak- puk Tso und Tso Kar, das grossartigste aber die Seekette, die mit Panggong Tso endet.
Im Ka.shmirgebiet kommen Salz- oder Brackwasserseen nicht vor. Anderseits aljer ist /.u bemerken, dass die Sedimente hier regehnassig Brackwasscrdiatomeen enthalten. Diesen Umstand mochte ich damit erkliiren, dass der jetzige Abfluss zu stark ist, uni eine Salzkon- zcntration zu ermoglichen. Dass Voraussctzungen fiir eine solche Konzentration friiher vorhanden waren, diirfte bewiesen sein. Betreffs der kleineren Seen (Manasbal Lake und der Srinagar-Seen) ist eine derartige Entwickelung nicht so merkwiirdig; dass aber auch W'ular Lake beinahe abfiusslosgewesen sein soil ist unerwartet. Die Losung dieser Fragc ist von klimageschichtlichen Gesichtspunkt aus von grusstem Interesse. Dazu ist aber ein sehr genaues Probenchmen voimotcn.
Ein Problem von grossem Interesse in diesem Zusammenhang ist, woher diese grosse
HOCIIASIATISCHE BINNENSEESEDIMENTE 217
Salzmenge komiiit. Es gibt zwci denkljare Ouellen: den umliegenden Berggrund und das Meer. Inncrhalb des Wassergebietes von Son Sakesar Kahar gibt es Salzlager; Wynne (1878j schreibt aber, dass sie unter dem Wasserspiegel des Sees liegen und also kein salz- reichen Zufluss liefern konnen. Vielleicht miinden jedoch Salzquellen in den Boden des Sees (Fleming 1853 (1854), S. 237). Wynne ist alxM- (S. 47j der Ansicht, dass das Salz aus dem Niederschlag komnien krinnte.
Das Salz wird bei Verdunstung aus dem Meer transportiert und wird von Nebel u. dcrgl. aufgenommen. Die Tropfchen, die den Nebel verursachen, sind oft so minimal, dass sie einen Nebel in gewohnlichem Sinne nicht bilden. Nel^elbildung al:)er setzt die Gegenwart von "hygroskopischen Substanzen" voraus, die als Kondensationszentra dienen konnen. Wahrscheinlich ist es auch eine solche Tropfchenbildung, die nur als eine halbdurchsichtige Triibung iiber dem Meer hervortritt (Kohler 1925, S. 71). Es sei hier aucb an die Angabe von Schlagintweit (1871, S. 467) in Zusammenhang mit dem Klima des Kashmirgebiets erinnert: "Die durch die Besonnung erzeugte Warme ist, bis Schneefall eintritt, auch durch eine sehr starke Triibung der Luft infolge von Suspension fester Korperchen bemerkbar." Nebelanalysen auf Chlor, Magnesium, und Kalzium zeigen so hohe Werte dieser Stoffe, dass sie die Annahme bestiitigen, dass die Meersalze wirklich Kondensationszentra der Nel^el- bildung sind (Kohler 1925, S. 70). Es ist aber die Frage, ob die Salzmengen, die jjihrlich aus dem Meer transportiert werden, so gross sein konnen. Kohler hat aus den Verdunstungs- werten von \\'ust lierechnet, dass die Meersalze "nur fiir 80.2% der Erdol^erflache Konden- sationskerne sein kunnen oder dass nn't anderen ^^''orten die Mg Na — Zone iiber 80.2% der Erde verbreitet ist. Da die Meeroberflache 70.8% der ganzen Oberfljiche betnigt, konnen sie als Kondensationskerne fiir 32.2% des Niederschlags der festen Erdoberflache dienen. Bei diesen Rechnungen habe ich von der Moglichkeit abgesehen, dass Schnee auf verschiedene Weise entsthen kann" (Kohler 1925, S. 73). Das Resultat von Kohlers Berechnungen ist, dass jahrlich 1,969,850,000 Tonnen Salze aus den Meeren transportiert werden. Die Zahl erscheint ungeheuer hoch. Kiirzlich hat aber Meinardus (1934) die von \Viist gegebenen Zahlen korrigiert und gezeigt, dass die Verdunstung des Meeres etwa dreimal grosser ist als die des Landes. Darum muss Kohlers Wert nur als ein Minimum angesehen werden. Obgleich man nichtgar zu grossen Wert auf diese Zahlen legen darf, scheint sie doch die ungefjihre Grossenordnung der Salzmenge zeigen zu konnen. Als einen ferneren Beweis fiir den Salztransport aus dem Meer mochte ich auf die Verteilung des Jodgehalts in den schwedischen Fliissen hinweisen. Die hoheren Jodwerte, die diese im siidwestlichen Schweden enthalten (Eriksson 1929, Fig. 23), zeigen — wenn auch das Analysenmaterial nicht ganz erstklassig ist — eine Tendenz, die mann kaum anders erklaren kann. als durch einen Material- transport mit den Meereswinden bedingt. Kiirzlich hat Cauer gezeigt, dass die Luftmassen von kontinentaler Herkunft sehr jodarm, von mariner Herkunft dagegen sehr jodreich sind (nach Goldschmidt 1934, S. 418, angegeben).
Die Windrichtungen innerhalb der vorliegenden Seegebiete sind also von grosser Ik'deutung. Auf alteren Karten, z. B. in den allgemeinen Handatlassen, wird angegeben, dass die Winde wahrend der Regenzeit regelmassig von SW iiber Indien hinauf wehen. Neuere Untersuchungen haben aber gezeigt (Harwood 1926 (1921) die Karten), erstens dass die Monsunbahnen anders laufen als friiher angenommen wurde, und zweitens, dass eine wesentliche Verschiedenheit zwischen der Windrichtung an der Erdoberfliiche und der in der hoheren Luftschichten herrscht. In- den letzteren kommen ahnliche Verhiiltnisse das
218
I10C11A.S1AT1.SCU1-: BlNNKNSKKSKUIMliNTli
ganze J.llir vor: dcr Wind welit etwa \\'SW in dcin Gcbietc, das iins hier interessicrt. Auf nicdrigcrcn Niveaiis wcht dcr Wind wahrcnd dor kalten Jahreszcit tmgcfahr von W, in Ladak v(»n N (Fig. v3). Wiihrend der Monsunzeit aber komnit dor Wind liaiiptsiichlich voni Bcnga- iisclicn Meerbiisen her, biegt gegen den Himalaya und folgt dein Siidabhang desselben gegen WNW. Die nieisten Rciscbiicher iilicr diese Gegendcn belx;n mehr odor wcnigcr hcr\-or, wie genau der Monsnn dcr Topograpliic folgt und iibcr die Piisse dcs Himalaya aufwilrts- dringt. Ja, der grosste Kenner dieser Gegenden, Iledin, leitct (1907, S. 605) das S-N-liche
Figure 3. Die Wiiidbahncn in Iiidieii uiid im wcstliclicn Tibet; links vviilireiid dcr Kalten Jahreszcit, rechts wiihrend dcr Monsunzeit. Nacli Ilarwood (1926, [1921]) und Bartholomew's Atlas von C. J. Ostnian zusammengestellt.
FrequenzabnclinicMi dcr Seen ini wcstliclicn Tiljet von den Monsunen ab: "This circum.stance is probably in some way connected with the passage of the monsoons across the Tiljetan high- lands and the varying amounts of their precipitation which fall to the lot of the different mountain-ranges." Es gibt also hier grosse I\ir)glichkciten eines Salztran.sports. Die grossten Liiftmassen kommen nach Harwood vom Bengalischen Meerbusen her, woher also auch die grosste Salzmenge kommen diirfte. Nach Schotts Karte (1933) bctragen al^er die Niederschlage hier etwa 2,000 mm., was den Verdunstungswert betriichtlich senken kann. Die geringste Niederschlagsmenge, 100-250 mm., kommt im Arabischen Meer vor, und dort diirfte auch die grosste Verdunstung zu suchen sein. Nach MoUer ( 1933, Taf. 29) ist der
nnciIASIATISCME BINNENSEESEDIMENTE 219
Salzgehalt in diesen Teilen des Indischen Ozeans geringer als weiter gegen O. Die Ursache dieses \^erh;iltnisses diirfte moglicherweise in einem grosseren Salztransport aus dem Meer durcli Verdunstung zu suchen sein. Voransgesetzt dass obige Schlussfolgerungen richtig sind, so waren die West-Winde die wichtigsten fiir den Salztransport zu unseren Gebieten. Da auch die Winde in hoheren Luftschichten diese Richtung haben, mag hervor- gehoben werden, dass ihre viel niedrigere Teniperatur lange Tropfchenbildung weit unter 0' nicht verhindern. Also liegt kein Hindernis vor, dass auch Reif, der ein Subliinations- produkt ist, Meeressalze enthalten kann. Dies ist von Bedeutung fiir kalte Gegenden wie z. B. Ladak. Obiges zusammenfassend scheint es niir glaubhaft, dass das Salz innerhalb unserer Seegebiete hauptsachlich aus dem Indischen Ozean, besonders dem Arabischen Meer, her- riihrt. ScliliessHch diirfte es wohl sel1)stverstandlich sein, dass dann auch die verschiedeuen Salzseen Ursprungsorte eines weiteren Salztransports sein konnen.
Tndessen ist das Salz, einmal in diesen Gegenden angelangt, noch nicht zur Ruhc gekom- mcn. Es ist angefiihrt worden, dass die Seen sakulJire Wasserstandsvariationen erleidcn (de Terra and Hutchinson 1934). Wenn der Wasserspiegel sinkt, vermehrt sich die Salz- konzentration ; grosstenteils bleil)t al>er das Salz auf dem trockengelegten Ufer zuriick (vgl. z. B. Lydekker 1883, S. 337). Bei der spateren Hebung des Wasserspiegels ist es aber nicht wahrscheinlich, dass dieses Salz demsell:)en See aufs neue zu gute kommt. Die meisten Reisebiicher aus diesen Gegenden l^eschreiben malerisch die "Staubmengen," die von den Stiirmen umhergetrieben werden. Es scheint mir darum wahrscheinlich, dass wenigstens ein Tcil der Salzmenge wahrend der Niederwasserperioden in den vorherrschenden Sturmrich- tungen weiterwandert und in die Zirkulation anderer Wassersysteme gelangt.
Die JVasso'typen. Aus obigem geht hervor, dass die Zusammensetzung des Wassers dieser Seen ziemlich schnellen Veriinderungen unterworfen ist. Die vorliegenden chemi- schen Analysen zeigen (Tabelle 1), die jetzigen Verhiiltnisse. Nur einige allgemeine Bemerkungen diirften hier ausgefiihrt werden; von chemischem Gesichtspunkte aus diirfte die Klarlegung dieser Probleme einem Chemiker, der eingehend sowohl die chemischen Eigenschaften des Berggrunds der resp. Wassergebiete sowie auch die chemischen Charak- tere anderer Seen kennt iiberlassen werden.
luncm "gewohnlichen Seewassertypus" am almiichsten sind i.okut Dal Lake, Manasbal Lake, \\'ular Lake, Yaye Tso und Ororotse T.so. Sie weisen alle cine niedrige Alkalisalz- summe auf, danel>en al>er ein unl)etraclitliches, a1)er regelmiissiges Vorkommen von CI und SO4. Die Karbonatgehalte sind einander sehr ahnlich ausser im Manasbal Lake, was wahr- scheinlich davon abhangt, dass das Zufliessen zum grossen Teil aus dem Kalk.steingebiet des Supra Kuling-Systems kommt. Die mikrobiologischen Verschiedenheiten dieser Seen diirfen nicht auf die Ungleichheit der Zusammensetziuig des Wassers zuriickgefiihrt werden. Brack- wasserseen sind Tso Moriri, Khyagar Tso, Mitpal Tso, Pangur Tso und Panggong Tso. Imu gemeinsames Kennzeichen derselben ist die relativ hohe Summe von Alkalisalzen. Unter den Brackwasserseen haben Tso Moriri und Mitpal Tso einen viel niedrigeren Cl-Gehalt als die iibrigen, wahrend Pangong Tso so Cl-reich ist, dass er einen Uljergangstypus zu den Salz- wasserseen biklct. Eine Ahnlichkeit mit diesen ist auch der hohe S04-Gehalt. Von den iibrigen sind auch Khyagar Tso und Pangur Tso C04-reich; sie diirften als Sulf.itsecn bezeichnet werden. Auch das Vorkommen eines relativ grossen Nao CO:,-Uberschusscs ver- ursacht, dass sie einander in chemischer Beziehung ganz nahe stehen. In mikrobiologi.scher Hinsicht sind die ebenfalls deutlich mit einander verwandt; ich mochte beide als Algengyttja-
220
HOCHASIATISCHE BINNENSEESEDIMENTE
seen bezeichncn, trotzdem Khyagar Tso vollig steril ist. Stiitze dieser Ansicht diirfte das Resultat der Tuschprobe sein. Es is moglich, dass in ciiemischer Hinsicht, solche Algen- gyttjaseen sodareiche Sulfatseen sind.
Salzseen sind Son Sakesar Kahar und Tso Kar, zu denen — wie sclion herv'orgehoben wurde — Panggong- Tso einen Ubergangstypus bildet. Dieser unterscheidet sich von den anderen beiden teils durch niedrigere CI- und S04-Gehalte, teils — vor allein — durcli den sehr kleinen Mg CU-Uberschuss. Jene sind einander sehr ahnlich, zeigen aber eine gewisse Ver- schiedenheit darin, dass Son Sakesar Kahar etwas Cl-reicher und Tso Kar etwas SO^-reicher ist. Beide haben einen sehr hohen Karbonatgehalt. Die Salzkonzentration ist zuni Teil eine Iiydrographische Frage. Inwiefern die wesenthche Verschiedenlicit von UnahnHchkeiten des Berggrundes abhiingig ist, entzieht sich nieiner Beurteilung.
TABELLE I Wassert\pcn nach Cheinischcn Analysen von H. Ncvvlands, Z. T. Unigcrechnet von
Dr. A. Assarsson
|
Seen |
Alkali- Salze 2 |
Cl Mg/l |
so. Mg/l |
Cl/SO, in Mol. |
Karbonat- Gehalt* |
Uberschuss an |
|||||
|
Nr. |
Namen |
Standard |
MgCls MgSO, |
NasCOj |
|||||||
|
1 |
Son Sakesar Kahar |
Salz |
730S0 |
34400 |
17176 |
5.42 |
1276 |
7980 |
|||
|
2 |
Lokut Dal Lake . . . |
Siiss |
9 |
1.6 |
5 |
1.0 |
65 |
t |
14 |
||
|
5 |
Manasba! Lake |
Siiss |
23 |
1.8 |
10 |
0.60 |
113 |
||||
|
6 |
Wular Lake |
Siiss |
18 |
1.7 |
11 |
0.40 |
62 |
2.3 |
|||
|
7 |
Tso Moriri |
Brackig |
788 |
22 |
517 |
0.12 |
435 |
30 1 199 |
|||
|
8 |
Khyagar Tso |
Brackig |
3784 |
2S7 |
2069 |
0.34 |
525 |
824 |
|||
|
in |
Tso Kar |
Salz |
61140 |
11662 |
35075 |
0.90 |
1633 |
9960 |
|||
|
11 |
Siiss |
IS |
1 |
7 |
0.4 |
63 |
|
25 |
|||
|
12 |
Mitpal Tso |
Brackig |
1011 |
82 |
625 |
0.35 |
364 |
110 |
|||
|
13 |
Pangur Tso |
Brackig |
3180 |
629 |
1316 |
1.29 |
842 |
.... I |
1936 |
||
|
14 |
Panggong Tso .... |
Brackig |
10039 |
3587 |
2750 |
3.56 |
1553 |
12 1 |
|||
|
15 |
Ororotse Tso |
Siiss |
14 |
4 |
3 |
3.6 |
32 |
5 |
• Mg(Ca)C03 nach Abrechnung von Ca und Mg an Cl und SO4 gebundcn.
Mikrohiolog'ischer Riickblick. Gleichzeitig niit den oben besprochenen Variatinncn iin Wasserstand und Salzgehalt verlaufen die biologischen Veranderungen. Ein zusanimcnfas- sender Bericht dariiber fiir jeden See kann mit dem vorHegenden Material nicht gegeben werden; einige Analogieschliisse diirften aber hervorgehoben werden.
Die Ungleichheit der untersuchten Seen der drei Gebiete ist ganz bedeutend. Son Sakesar Kahar enthiilt ein sehr reiches hakifiles I-eben. Es ist streng spczialisiert, indeni nur zwei Arten vorkommen, daruni aber — wie Thienemann (nach Decksbach 1924, S. 274, angefiihrt) sagt — hochproduktiv. Die Sedimentc sind aber in dieser Beziehung vollstandig nichtssagend.
Die Seen des Kashmirgebiets sind seichte Siisswasserseen niit gulcni .\bfluss und Wasserfarben Forel-Ule X-XVI. Die Sichttiefen sind uncrwartct niedrig, 1.2-4 ni. Mog-
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be S •o XI o u a <u J= (J 15 u |
>^ bo it: 3 c o ™ |
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bo 3 'cj QJ 3 5 |
to 3 OJ c o (5 |
>i be OJ 'S 1 H o 5 |
•o « c ^" fcl-l a; ,60 o |
% bo c _bo < >, 3 u •g 'C |
(t: X o a> .ho |
60 in 3 'C OJ •g X u 'H G |
u u i- H o S |
bt in 3 |
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c g 5 re 'q re JD u ■g i ,bo ?, |
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S N |
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OJ . .SH |
fc.Sf :0 S |
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n! bo H ^ II Ill : 1- o n |
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3 bO |
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3 re _bo |
mX 'aj ^ |
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re^.l |
tn 'oj 3 re L> 60 |
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c t- 6 X |
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I- r C |
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i u o o u 3 to C re |
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HOCIIASIATISCHE BINNENSEESEDIMENTE 227
lich ist die Ursache in Plankton etc. zu suchen. Der pH-Wert ist an den Oberflachen 8.2-9.0, weshalb die Seen also als eutroph angesehen sind. Leider kenne ich die Phytoplanktonpro- duktionen nicht. Die hohere Vegetation, wo man besonders Lotus und Trapa bemerkt, ist oftmals sehr vippig. Audi mikrobiologisch sind diese Seen relativ reichhaltig. An Diato- meenarten kommen insgesamt 118 vor und sie wechseln in den verschiedenen Seen zwischen 28 und 64. Unter diesen sind 79 St. nicht in den Ladakproben angetroffen worden (vgl. Tab. 3). Merkwurdigerweise, gibt es in jedem See 2-3 typische Brackwasser- forinen: Gyrosigma Spenceri, Navicula protracta v. ca pi fat a, N. pygmaca, N. saHnarum und A'^. sp. Prozentuell wechselt ihre Frequenz zwischen < 1% und 3% der Diatunieenflura. Von samtlichen Arten sind die nieisten (17-47 St.) Aufwuchsfornien, 5-11 sind Bodenfor- men und 4-6 Planktonformen. Von der ganzen Diatomeenflora sind 80-90% Aufwuchs- formen. Boden- und Planktonformen sind also ganz untergeordnet. Neben Diatomeen kumnien audi andere Mikrofossilien vor. Desmidieen finden sich in Loknt Dal Lake, Sundar Khun and Manasbal Lake. Alle Ijeobachteten Formen sind klein und unansehnlich. In Lokut Dal Lake gibt es ein paar Pcdiastruui-Arten, 2-4 St. pro min.^ Die Tierreste sind sehr selten. So gibt es von BosininaSdmlen kommen in Lokut Dal Lake und Sundar Khun 4 St. in Manasbal Lake 2 St. pro mm.^ vor. Von Spongiennadeln sind in Lokut Dal Lake 26 St. und in Sundar Khun 16 St. pro mm.^ angetroffen worden.
Die Ladakseen sind in der Regel relativ tief ; am tiefsten ist Tso Moriri, am seich- testen Tso Kar. Die Wasserfarbe ist im allgemeinen Forel Ule V-VIII, aber audi Werte von III bis XII sind beobachtet worden. Die Sichttiefen sind unerwartet verschieden (1.5 — 9ni.); ob die Ursache in Suspensionen oder Plankton zu suchen ist (vgl. doch unten), ist niir unbekannt. Der pH-Wert ist 7.1-9.6, im allgemeinen doch etwa 9 oder niehr. Betrefifs der Phytoplanktonproduktionen weiss ich nur, dass Ororotse Tso "considerable amounts of plankton" besitzt (Hutchinson 1933, S. 136), und dass in Yaye Tso "the water clouded with centric diatom prob. Cyclotella" ist (Hutchinson, briefiiche Mitteilung). Diese beiden Seen sind also eutniph in Naunianns Sinn von 1932. Die hohere Vegetation ist sehr schwach ent- wickelt; gibt es eine solche huhere vie im Sta-rtsak-puk Tso und Pangur Tso, so besteht sie hauptsiichlich aus Potaiiiogcton. In mikrobiologischer Beziehuiig weichen die Ladakseen betrachtlich von den Seen des Kashmirgebietes ab. Die Anzahl Diatomeenformen betragt 72, 30 davon sind nicht in den Kashmirproben gefunden worden. In verschiedenen Seen betragt die Zahl 4-34, ist also durchschnittlich bedeutend niedriger als in Kashmir. Obgleich mehrere dieser Seen brackig (oder salzig) sind, sind typische Brack-oder Salzwasserdiato- nieen nur in Pangur Tso und Panggong Tso angetroffen worden, niimlich : A)iiphiprora paludusa, Anomoconeis polygramma, Epithcmia turgida v. IV estcrmanni, Navicula pygmaca und A'^. salinaruiii. Diese Brack — oder Salzwasserfornien machen in Pangur Tso 79% und in Panggong Tso 16% der ganzen Diatomeenflora; daneben aber gibt es resp. 18% und 23% der fakultativen Siiss- oder Brackwassertypen. In Milieubeziehung sind die Ladak- floren von einander sehr verschieden. In Sta-rtsak-puk Tso, Mitpal Tso und Ororotse Tso liegt der Schwerpunkt auf den Aufwuchsfornien niit resp. 79, 55, und 77%. In Pangur Tso betragen die Bodenformen 81%, 79% sind Anonweoncis polygniinnia. In Mitpal Tso betragen die Bodenformen 33%, selbst dies ist eine ausserordentlich grosse Zahl. In Tso Moriri bilden fakultative Boden- und Planktonformen (= die Siisswasserform C\c!olclla antiqua) 89%. In Vaye Tso und Panggong Tso herrschen die Planktonformen — in beiden Cyclotella coiiita — mit resp. 73% und 47% vor.
228
HOCIIASIATISCIIE BINNENSEESEDIMENTE
Nur Siisswasserformen. Audi Brack-oder Salzwasserformen. Keine Formen gefiindcn. Figure 4. Die Verbreitung der Diatomeen nach Halinitiltstypcn in den rroben der untcrsuchten Seen.
Fakultative Formen nicht mitgerechnet.
Siisswasserseen. Brackwasserseen. Salzwasserseen.
FiGUKE 5. Der okologische Standard der imtersuchten Seen; diirfte niit Figure 4 verglichen werden.
HOCHASIATISCHE BINNENSEESEDIMENTE
229
Diese kurze Ubersiclit zeigt, class oftmals eine sehr grosse Verschiedenheit zwischen dem okologischen Standard der Seen und den Angaben der Bodenproben besteht. Auffallend ist das haufige Vorkonimen von Brackwasserformen in den Siisswasserseen Kashmirs, und ebenso auffallend das Fehlen solcher Formen in den Proben mehrerer Salz- oder Brackwas- serseen Ladaks (Figs. 4 und 5). Ich kann diese Unregelmassigkeiten nicht anders erklaren, als dass der okologische Standard so oft wechselt und die Proben nicht rezent sind. Da Variationen sicherlich ebenso oft wie hier stattfinden (vgl. Panggong Tso) und daneben die Sedimentation ganz sicher unbetraclitlich ist, ist es notwendigger als gewohnlich, die proben mit allergrosster Genauigkeit einzusammeln. Nun sind wahrscheinlich die Proben aus Lokut Dal Lake und Wular Lake mit Rohrlot genommen, weshalb die Gefahr fiir Umriihrung der- selben nicht so gross war. In diesem Zusammenhang mochte icli hervorheben, dass fiir den Alyk-nor dasselbe Verhaltnis zu bestehen scheint (siehe Mereschkovvsky 1906). Der betref- fende See ist abflusslos und die Diatomeenflora siiss. Leider gibt es aber keine Angaben iiber den okologischen Standard des Sees.
Betreffs der Diatomeenfloren der Ladakseen und ihrer okologischen Variationen soil eine andere Moglichkeit in Betracht gezogen werden. Die Seen liegen ganz isoliert und sind wJihrend grosser Zeit des Jahres eisbedeckt. Die Zeit fiir die Einwanderung neuer Formen, ist darum sehr kurz. Durch die Steigerung des Salzgehalts sterben eine grosse Menge Siiss- und Brackwasserarten und neue halofile Formen wandern relativ selten ein. Die Entvvickelung dieser Seen geht also einem "Wiistenstadium" entgegen. — Wenn der Salz- gehalt zufolge der steigenden Niederschlage und durch Wasserstandshebung vermindert wird, kommen die Siisswasserformen, die in Bachen, Siimpfen u. dgl. in der Nahe der Seen leben, zuriick, um die Seen in Besitz zu nehmen.
In mikrobiologischer Beziehung sind die untersuchten Seen nach den Bodenproben zu beurteilen sehr diirftig. Es kann aber nicht genug hervorgehoben werden, dass die Diato- meen, wie andere Algen, zoniert leben, was sowohl fiir Boden — als Aufwuchs formen gilt (Thomasson 1926). Da man nur sporadische Proben bezitzt, ist es selbstverstiindlich nicht anzuraten, die Floren dieser Seen mit denen anderer Seen zu vergleichen. Ganz besonders gilt dies bei den sehr durchsichtigen Seen, wo die Zonen weniger zusammengedrangt vorkom- men. Als Beispiel, wie verschieden die Angaben fiir einen einzigen See sein konnen, will ich die Resultate betr. Panggong Tso nach dem Material dreier Expeditionen mitteilen. Die Expeditionen sind Yale, Trinkler und Hedin und die Bearbeiter Thomasson (hier publiziert), Meister und Hustedt.
Diatomeenarten
Achnanthes lanceolata
" linearis
" microcephala . .
" minutissima . . .
Amphiprora paludosa
Amphora commutata
" ovalis
" " V. pediculus
" Schroederi
|
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Meister |
Hustedt |
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(hier) |
(1932) |
(1920) |
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230
HOCIIASIATl.se HE BINNENSEESEDIMENTE
Diatonieenarten
Anomoeoneis exilis
" polygranima
CaloiK'is Ijacilluni
" fasciata
" silicula
" " V. tnincatiila
sp
Cocconeis pediculus
" placeiitula
Cyclotella conita
Cyml)ella aequalis
affinis
" " V. excisa
" aspera
cistula
" " V. niaculata
" lanceolata
" norvei^ica
" parva
" sinuata v. antiqua
" tuniida
" venlricosa
Denticula knuis v. frigida
Epitheniia argus
" V. alpcstris
" " V. longicornis . . .
" turgida v. Westennanui
" zebra
" " V. proboscidea . . .
Fragilaria construens v. suljsalina .
" leptostauron
" pinnata
Aleridioii circulare
Navicula aiiiphilxila
" cryptocepliala
" cuspidata
sp. (c fr. fortis?)
" oblonga
" peregriiia
" ])yginaea
Rheinhardti
* Massenhaft.
Thomasson (liier)
Meister
(1932
Hustedt (1920)
HOCHASIATISCIIE BINNENSEESEDIMENTE
231
Diatunieeiiarten
Navicula salinarum
" tuscula
Ostrupi
Neidium rectum
Nitzschia capitellata
" communis
" frustulum
" Iningarica
Kittlii
" thermalis
rinnularia appendiculata
" Brebissoiiii
" gracillima
" leptosoma
" major v. linearis
" " V. paludosa
Tibetana
Rhoicosplicnia curvata
Khiipalodia gil)l3a
" " V. ventricosa
" gibberula v. margaritifera
Stauroneis phoenicuteron
Surirella ovalis v. ovata
" var
Synedra ])ulchella ,
" Vauciieriae
" V. capitellata
Thomasson (hier)
1
1
1 1
Meister (1932)
1
1
Hustedt (1920)
1 1 1
rfte die Ursache in d
Die drei Listen sind cinander sehr unabnlich. Zuni Teil di' Vegetationslokalitaten zu suchen sein. So sind die von Hustedt gefundenen Diatomeen a Herbariumpflanzen eingesaminelt worden. Moglich ist aljer auch, dass soich grosse Ver- schiedenheiten von okologischen Veranderungen abhangig sind. Jedenfalls zeigt diese Zusammenstellung von Panggong Tso, wie unzuverlassig nahere Vergleiche zwischen diesen asiatischen Seen und anderen Seen sein miissen.
Nach meiner Meinung ist es notwendfg, die Floren niclit nur qualitativ, sondern auch — und hauptsachlich — quantitativ mit einander zu vergleichen. Leider liegen von friiher keinc solcben Angaben vor. /Ms eine allgenieine Charakteristik kann al)er hervorgehol>en werden, dass vorliegende Floren durch die Abwesenheit von oder unbetrachtliche Mengen Euiwtia-, Fragilaria- und Pinnularia-Arten gekennzeichnet sind. Dies ist nicht sehr erstaunlich, da diese Gattungen zum grossen Teil fiir die humusreicheren Gewasser bezeiclniend sind.
Die Sedivicnttypen. Da die Sedimente das Endresultat des Lebens eines Sees sind, durften sie wenigstens in besonderen Teilen dies Leben wiederspiegeln. Ein Riickblick auf die Sedimente der eruahnten Gebiete in Bezug auf die Strukturanaiysen wird darum hier
232 llOCllASIATISCIIi; binnenseksedimente
gegeben, Zucrst aber niochte ich nochmals betonen, dass das Material in jeder Probe ganz heterogcn ist, uiid dass die Analysenresultate daruni nur als rclativ zu betrachteii sind. Der Grundstoff der Sediniente besteht aus Detritus und Rlineralkomern. V.s sind daruni zu allcr- erst die Variationen in den Mengenverhiiltnissen derselben zu beacliten. Der Detritus ist von gro1)eni oder feinem Typus. Zum Grobdetritus rechne ich Plianeroganienreste, die niclit so destruiert sind, dass die Zellkomplexe ganz aufgclost sind. Sowie die gleichartigcn Gewebe der hoheren Kryptogamen. Der Feindetritus ist das vollstandig destruierte organ- ische Material. Die Mineralkorner sind limnoallochton und bestehen gewohnlich aus Quarz oder Feldspat oder, was besonders angegebcn wird, audi aus dunkeben Mincralien. F.s ist oft schwierig, die limnoautochtonen "Ca-Korner" in der Strukturanalysc von den kleinen Mineralkomern zu unterscheiden, weshalb jene z. T. in der Summe vcrborgen sein kiMinen.
Die Sediinente von Son Sakesar Kahar weichen, was zu erwarten, war ganz von dencn der anderen Seen ab, in struktureller Beziehung zwar nicht so auffallcnd. Die Lagenfolge ist aus schwarzen Fe S-reichen und weissen Kalziunikarbonatreichen Schicliten aufgebaut worden. Die Verschiedenheiten ihrcr Zusamniensetzur\g werden durch die Strukturan- alysen (Tabel 2) deutlich gezeigt. Der grosse Gehalt an Fe S (und Ho S) ist von der Zerlegung der betrachtlichen Mengen Microcystis und Diaptonms abhangig. Speziell aus dem letzteren stamnien die karotingefjirbten Reste, die jedoch nur in der Oberlliichenlage vorkommen und darum nioglicherweise aufgelost worden sind. Als Ursache dieses Unter- schiedes im Sedinienttypus diirfte nicht die Jahreszciten angenommen werden. Dagegen spricht die relativ grosse Miichtigkeit der Schichten. Ehcr diirften diese das Pulsieren in den sakulJiren Wasserstandsvariationen und die davon bedingten biologischen Veninderungen wiederspiegeln, also dasselbe Phanomen beleuchten, das von de Terra und Hutchinson betr. Panggong Tso beschrieben wnrde. Fiir eine niihere P>esprechung der Sediniente und Natur- vcrhaltnisse wenn die letzteren abgelagert wurden, scheinen niir chemisclie Analysen selir wertvoll zu sein. Solclie stehen mir aber leider nicht zu Gebote.
Die Kashniirsedimcnte sind in trockenein Zustand klar grau und frei von Kalziumkar- bonat. In diesen seichten Seen konnte man einen durchgehend hohen Grobdetritusgehalt erw arten. Das Vorkommen eincs solchen ist aber ganz unregelmiissig. Nur betr. Lokut Dal Lake und Sundar Khun niaclit sich dieser Gehalt in den Analysen geltend. Damit sei jedoch nicht gesagt, dass es den iibrigen Proben an diesem Gehalt mangelt. In den ganz und gar von Litoraltypus l)estehenden Kashmirsedimenten, sind dagegen die Mineralkorner sclir zahlreich (20-30%). Was die Grosse betrifft sind sie gewohnlich < \Q) f-, also hauptsiich- lich Schluff; cinzelne grosscre Korner konnen aber selbstverstJindHch vorkommen. In diesem Zusanmienhang sei bemerkt, dass nach den Feldnotizen die oberfiiichlichen Sediniente von Lokut Dal Lake reichhaltige Muschelschalcn enthalten, wahrend tiefcr in den Lagen- folge ihrer viel weniger sind. Im Mikroskop kann man zwar in den untersuchten Proben einige Fragmente davnn l)cmerken; die HCl-Probe gab aber gar keine Reaktion. Obgleich sich der Muschelgchalt gegen die Gcgenwart hier vermchrt haben kann, ist cs wahrschcin- licher, anzunehmen dass die Schalen infolgc des unbctnichllichen Karbonatgehalts des Wassers und des relativ starken Hunmszuschusses (Versumpfung der Umgebungen) sehr schnell aufgelost werden. So verhalt es sich oft in den dystrophen Seen. Der Feindetritus bildet im allgemeinen den Hauptbestandteil dieser Sedimente. Er enthiilt iminer etwas Algenschleim, der jedoch zum Teil von den Schleimstielen der Aufwuchsdiatomeen hcr- riihren kann. Bezeichnend fiir die Kashmirsedimente sind schliesslich die Diatomeenmen-
IIOCHASIATISCHE BINNENSEESEDIMENTE 233
gen, die sich audi in struktnreller Beziehung in so hoheni Grade geltend machen, dass sie fiir das Benennen der Sediinente bestimmend geworden sind. Ziisammenfassend, kann man iiber die Kashmirsedimente sagen, dass sie, von den Variationen im Grobdetritusgehalt in Lokut Dal Lake und Sundar Khun abgesehen, einander sehr ahnlich sind. Sie gehoren zum Typus der seichten Seen, die sich dem Zinvachsstadium nJihern. Von den ini folgenden besproclienen Sechmenten sind diese in vieler Beziehung stark verschieden.
Die Sediniente des Ladakgeliietes sind viel mannigfaltiger als die von Kashmir; sie sind hauptsachHch von den sehr verschiedenen Seen typen abhangig. Auffallend bei der Mikro- skopierung ist also hier das minerogene Material, die Mineralkorner. Beziiglich der Fre- quenz wechseln sie allerdings sehr, 2-63%, durchgehend aber sind die Korner viel grosser als die des Kashmirgebiets. Gewohnlich sind sie grosser als 15 z^, oftmals 20-40 /i*; Korner von 100-200 i". sind aber nicht so selten in den Proben bemerkt worden. Weiter sind die Korner hier oft splitterig und scharfkantig. Dies diirfte damit in Zusammenhang stehen. dass die Zufliisse der Seen so kurz sind : die Mineralkorner haben darum nicht Zeit genug, abgerundet zu werden. Ich vermute, dass auch Frostsprengung und Wind beitragende Ursachen zu dem betreffenden Verhaltnis sein konnen : kleine SpHtter werden weggesprengt und von Winde beinahe unmittelbar in die Seen getragen, wo sie schnell sedimentieren. Auch Kalziunikarbonat gibt es nach der HCl-Probe in mehreren dieser Seen; es kommt aber hauptsJichlich als limnoallochtones Korn aus den Kolkformationen des Panjalsystems vor und wird zu den "Mineralkornern" gerechnet. Der Grobdetritus ist am reichhaltigsten (jedoch nur 7%) in dem seichten Sta-rtsak-puk Tso, von dem besonders berichtet wird, dass er mit Potamogcton bewachsen ist. Die Chitinreste sind sehr ungleichartig ; sie konnen nicht inuner bestimmt werden. Die zahlreichen Reste in Tso Kar diirften zu Arteiiiia gehoren, in Pangur Tso aber sind sie Ostracodenschalen. Diese bestehen grosstenteils aus Kalziuni- karbonat und diirften dannn in hohem Grade zu der starken Reaktion der HCl-Probe beige- tragen haben. Bemerkenswert genug sind keine Bosmina-Keste in den Ladakproben ange- troffen worden. Diatomeenreich sind Yaye Tso, Panggong Tso (in 31 m.) und Ororotse Tso. Beachtenswert scheint mir, dass auch dieser letztere, in einem wirklich arktischen Milieu gelegene See ein so reiches Leben beherbergt. Prinzipiell erinnert dies beinahe an den Planktonreichtum der antarktischen Meere. Der Feindetritus ist quantitativ wechselnd, bildet aber in den meisten Fallen den Hauptteil der Sediniente; Ausnahmen sind Tso Moriri, Sta-rtsak-puk Tso und Panggong Tso, 31 m. Algenschleim fehlt oder seine Ouantitat ist sehr untergeordnet, ausser in zwei Seen, Khyagar Tso und Pangur Tso, wo er > 90% des Feindetritus ausmacht. Diese Gyttjen kann man also Algengyttjen nennen. Der Sediment- typus in Pangur Tso ist durch seinen Gehalt an Algenschleim und durch den hohen Gehalt von AnoDwconeis polygraiiniia gewissen Seen der Stockholnier Gegend zu der Zeit, da sie von der Ostsee isoliert wurden, sehr ahnlich. Man war friiher der Meinung, dass das Algen- gyttjenstadium der Seen sich nur unter warmeren Klimaverhaltnissen ausbildete. Da dieses Stadium unter besonderen Bedingungen im Ilochland Tibets angetroffen worden ist, diirfte wohl darin diese erwahnte Meinung keineswegs eine Stiitze erhalten haben. Daraus ware auch der Schluss zu ziehen, dass das Leben der Seen zum grossen Teil von den Variationen der Kliniatypen nicht unmittelbar unabhangig ist; sehr eigentiimlich scheint mir, dass die Temperatur von ganzlich untergeordneter Bedeutung ist. Von grundlegender Bedeutung dagegen sind die cheniisclien Verhaltnisse des Berggrundes des Wassergebietes. Besondere Umstiinde treffen aber selbstverstJindlich ein, wenn ein Zuschuss von Meeres-
234 IIOIIIASIATISCIIIC BINNENSEESEDIMENTE
salzcn liinziikdiiiint. Dies liiulet statt (lebicteii, die (lurch NivcauvcriliKkTun^t'ii odt-r ahnliclic \'crli;iltiiissc von deiu Mecr isdliert wcrdcn, sowie audi Ikm \\'iisten,s;el)i(.'ten. Die X'oraussetzung al^er, dass sich die auf \erscliiedenc W'eise in den See eino'efiihrten I'.kktni- ivten sjeltend niachen kiinnen, ist ein gewisser Ziisaninienhang zwiselun Zu- und Abfliessen. Sclilagintwcit (1872, S. 139) iiusscrt sich ul)er eine illinliche Frage : . . . "sDwie Crosse des Oiiellengehietes der Zufliisse im Verhiiltnisse zum Wasservolumcn und zur Oberlliiche, dies siiid dabei die wichtigsten Momente."
Ein ]\iickl)Hck auf die Mikrofossilien- und Scdinienttypen der I)esprochenen Seen gibt folgende .\uffassung vnn ik-n allgenieinen I'rinzipicn des luitwicklungsvcrl.aufs. In den tiefen Seen herrschen die Plankton formen vor, hauiitsiichlich Diatonieen und Tierclien (Copepodcn, die jedoch niclit resistent sind). Die Sedimentation ist ini allgenK'inen sehr schwacli, kann aber ortlich dnrch kriiftige Zufuhr von minerogenem Material in den litoralen Teilen verstJirkt werden. Wenn das Verhiiltnis zwischen Zu- und Ablliessen unveriuidert !)leil)t, verniehren sich die Bodenforinen vvenigstens ortlich, bald. Sie sind in den tieferen Seen des Gebietes die relativ wichtigsten. bis die \'erschlaniniung und andere Milieuverbiilt- nisse so entwickelt werden, dass eine h(>here X'^egetation (z. ]]. J'ataiiioi^cton) einwandern kann. Dann verniehren sich auch die Aufwuchsfornien und nehmcn iiberhand. Wenn der See innerhalb eines kalkreichen Gebietes liegt und das Wasser deshall), oder anderer Unistiuide h.ilber, von eineni kalk- und sulfatreicheren Typus ist, so wandeni die Myxophycceninassen ein. Der Sediuientzuwachs gcschieht dann dank dieser starken Materialzufuhr viel schnellcr. Innerhalb (lebietc mit geringen Niederschliigen und starker Verdunstung maclieu sich ;uich schnell kk'ine \'ariationcn in Niederschliigen und \^erdunstung gcltend und der Konzentra- tiousgrad in dem Wasser der Seen verandert sich relativ leicht. W^enn sich die Nieder- schliige vermindern, nimmt das Zufliessen al). Kommen dann Meeressalze hinzu, kann sich der okologische Standard des Sees verandeni und brackig oder salzig werden. (ileichzeitig sterben dann zuni Teil die Siisswasserfornien aus und ISrack- oder Salzwasserfornien wan- dern ein. Innerhalb des wiirmeren Kashmirgebictes findet dieser Austausch relativ leicht statt: aber in ariden Gebieten wie Ladak, wo die Seen den grossten Teil des Jahres eislK'- deckt liegen, sind die Aussichten auf eine Neueinwanderung stark vermindert. Daruni sind die llrack- und Salzwasserfloren in diesen Gebieten sehr arm. Innerhalb Gegenden, die nicht so lange Zcit eisbedeckt sind, z. 15. weiter in Tibet hincin, ist der Reichtiun der Formen viel gn")sser (vgl. Mereschkowsky 1006 und Oestrup 1909 j. XX'enn das Wasser der Seen steigt und der Salzgehalt also abnimmt, k(")nnen die Siisswasser- und besonders die fakulta- tiven Siiss- und P>rackwasserformen sehr leicht einwandern, da sie noch in I'fiitzen u. dgl. in miiuittelbarer Nahe der Seen le1)en.
Jiisblick
Nacb vorstehender Linlersuchung dieser asiatisi-hen i'linnengewassersedimi'ule muchte ich einige daraus gezogene Konsequenzen darlegen. Obgleicb die I'mben nur kleine llruch- stiicke der Geschichte jedes Sees bilden und ausserdeni nur von cinem einzigen Platz des l)etrefifen(len Sees stammen, kann man bier, gestiitzt auf friihere Untersuchungen anderer Art M(')glichkeiten fiir fortgesetzte Arbeiten wahrnehmen. Dnrch die Arbeit von de Terra und Ilutchinsf)!! wird gezeigt, dass die Seen relativ schnell und regelmiissig zwischen Siiss und Brackwasserstadien iiendeln. Vorstehende l^ntersuchung zeigt, dass die Sedimente deutlich
HOCHASIATISCHE BINNENSEESEDIMENTE 235
diese Stadieii w iederspiegeln. Wcitcrhin siiul, liauptsiiclilicli in Kashniirsedinienten, Pollen verschiedener Arten so reichlich angetroffen wurden, dass es nicht unin(")glich ware, niit Bei- stand dersell>en die Variationen in der Zusammensetzung der \\'aldcr zu studiereii. Dadurch wiirde man auf einen festen Boden kommen betreffs der Klimagescliichte Kashniirs. Schwie- riger ist es, diese Erfahrungen fiir die Hochgebirgsketten und die waldlosen Gegenden Ladaks zu auszuniitzen. Als Ausgangspunkte scheinen inir die pollenreicheren Zoncn von Kashmir geeignet. Mit grosseren I'ollenmengen diirften die Moglichkeiten fiir Ver- l)reitung der leichtfliegenden NadelwaldpolIenkfJrncr zunehmen. Durch Niveaukombina- tionen mit weithergeflogenen Pollen (z. B. audi Tanuiri.v-PoWen?) und den Wechsel zwis- chen Siiss- und Brackwasserstadien der Seen also nacli den Mikrofossilien diirftc man eine relative Chronologie auf die Sedimente aufbauen konnen.
Rein theoretisch ist die hier vorgeschlagene Untersuchung relativ einfach auszufiilircn. Die Schwierigkeit Hegt darin, ein geeignetes Material aufzuspiiren und einzusammeln, aber sicher wiirde sich die Miihe reichlich lohnen, da durch eine solche Untersuchung einer der Schliissel zur Klimageschichte Zentralasiens gefunden werden Konate.
Sveriges Geologiska Undersokning, Stockholm, im Januar 1935.
Note by Biiilogisf of the Expcditiaii
After Dr. Lundcjvist's paper was received for publicati(_)n, 1 heard from Dr. de Terra tliat lie had forwarded a sample of diatomaceous deposit from the interglacial lacustrine beds of Pang-gong Tso to Mr. Iv. E. Lohnian for determination. Mr. Lohman most kindly for- warded the names of the species present; as he did not think the report worth publishing as a separate paper, his list and comments are transcribed below. I have added indications of the present distribution in Indian Tibet of the sjiecies encountered. It is probable that the greater part of the diatoms in the shallow water deposit in Pang-gong Tso are derived from this interglacial l)etl and not from the living flora of the lake. — G. E. H.
236 JIOCHASlATISCllE BIN NENSEESKDIMENTE
Diatoms from a frcsh-watcr intcrglacial lake hcd in tlic hoiiIktii Iliinalayas. India. Sample No. K-108, U. S. G. S. Dial.. 111 L.xality No. 2274.
Cyclotclla comta (EhrenlK-rt;) Kiitziu!-- (F) A Ka. Yy. M. P.31 .
StephanoclLsciis astraca niimitula (Kiitzing) Griinow . . Cold R not rec.
Fragilaria leptostaiiron (Ehrenberg) Hustedt (F) R Ka. P31.
Eunotia suecica A. Cleve Cold K not ixc.
Cocconeis placentula lineata (Ehreiiberg) Clevc (FBj A Ka. AI. Or.
Rlioico.sphenia ciirvata ( Kiitzing) Grunow (FB) F Ka. Tm. St. Yy. P31 Ov.
Mastogloia elliptica Agardh R not rec.
Calonei.s silicula (EhrenlxTg) Cleve (F15) F .St. Yy.
Stauroneis montana Krasske Cold R not rec.
Anomoeoneis polygramma (Ehrenberg) Cleve (FB) C i'31, I'a.
Navicnla reinhardtii Grnnow ( F) Cold R I'a. P31.
Navicula oblonga Kiitzing A not rec.
Pinnnlaria niicrostauron (Ehrenberg) Cleve Cold R not rec.
Amphora ovalis Kiitzing (FB) F Ka. Tm. Yy. Or.
Cymbella ventricosa Kiitzing (F) C Ka. Or.
Cymbella lanceolata (Ehrenberg) Van Heurck (F) C Ka.
Cymbella cistula (TTemprich) Grunow (FB) F Ka. St. Yy.
Gomphonema intricatum Kiitzing (F) R Ka. St. M.
Gomphonema cf. parvnlum (Kiitzing) Grunow R not rec.
F.pithcmia turgida (Ehrenberg) Kiitzing (FB) A Ka. St. M.
Epithemia zebra porcellus (Kiitzing) Grunow (FB) F Ka. Pa.
Epithemia sorex Kiitzing (FB) C Ka. Yy.
RhoiKdo<lia gibba (F.hrenberg) Miiller (FB) F Ka.
Nitzschia denticula Cjrunow F not rec.
Ka. = Kashmir Valley, Tm. = Tso-moriri, St. = Sta-rtsak-puk Tso, Yy. = Yaye Tso, M. z= Mitpal Tso, Pa. = Pangiir Tso, P31 ^ Panggong mud 31 m., Or. ^ Ororotse Tso, not rec. =: not recorded by Lundqvist from recent deposits, F. = fresh, FB. = Fresh and Brackish.
A = abundant, C := common, F =: few, R = rare.
"Ail uf the above species are living today in freshwater to somewhat saline lakes, ( )iic species, Anomoeoneis polygraiitma, is living at present in the (ireat Salt i-ake in Utah and in the Gulf of Mexico. Its connnon occurrence in the Indian material suggests that the water was at least somewhat saline, and this is further indicated by the fact that half of the species live in both fresh and brackish water environments.
Stephanocfisctis astraca iiiiiiiihila, Eunotia suecica, Stauroneis niontaiM, Navicula rein- hardtii, and Pinnnlaria niicrostauron are now living in cool water lakes and suggest that the Indian material was deposited from a cool water lake. These species all occur rarely, how- ever, so thcv ofifer no verv conclusive evidence."
HOCHASIATISCIIE BINNENSEESEDIMENTE 237
LITTERATUR
Antevs, Ernst. 1929. Maps of the I'leistoceiie Glaciations. Bull. Geol. Soc. of America,
Vol. 40, p. 631-720. Dainelli, Giotto. 1922. Studi sul Glaciale. Spedizione Italiana de Filippi, Ser. II, \^ol.
Ill, Bologna. Decksbach, N. 1924. Seen und Fliisse des Turgai-Gebietes (Kirgisen-Steppeii). Verh.
Intern. Verein. Limnol., Bd. II, S. 252-288. DE Terra, H. 1933. Preliminary Report on the Yale North India Expedition. Science,
Vol. 77 (N. S.), p. 497-500, 1933. ■ 1934. Physiographic Results of a Recent Survey in Little Tibet. Geographical
Review, Vol. XXIV, No. 1, p. 12, 1934. and Hutchinson, G. Evelyn. 1934. Evidence of recent climatic changes shown
by Tibetan Highland lakes. Geogr. Joum., vol. LXXXIV, No. 4, p. 311-320. Eriksson, J. V. 1929. Den kemiska denudationen i Sverige. Medd. fr. Statens Meteorol.-
Hydrogr. Anst., Bd. 5, No. 3, 4:o. Fleming, A. 1853 (1854). Report on the Geological Structure and Mineral Wealth of
the Salt Range in the Punjaub. Joum. of Asiatic Soc. of Bengal, Vol. XXII, p.
229, 333, and 444, Calcutta 1854. Geological Map of India and Adjacent Countries. Fifth Ed. 1928. Geological Sur- vey of India. GoLDSCHMiDT, V. M. 1934. Drei Vortrage iiber Geocheniie. Geol. Fi'iren. Fcirh., Bd. 56. Harwood, W. a. 1926 (1921). The free atmosphere in India. 8. Upper air movement
in the Indian Monsoons and its relation to the general circulation of the atmos- phere. Mem. of the India Meteorological Department, Vol. XXIV, 1922-25,
Calcutta. Hedin, Sven. 1907. Scientific Results of a journey in Central Asia 1899-1902. Vol. IV,
Central and West Tibet, Stockholm.
1909. Transhimalaya. Upptackter och afvcntyr i Til)et, Bd. I och II, Stockholm.
1917. Southern Tibet, Vol. II, Stockholm.
Holland, Thomas. 1926. Indian Geological Terminology. Mem. Geol. Surv. India,
Vol. LI, part 1. Huntington, Ellsworth. 1906. Pangong: a glacial lake in the Tibetan plateau. Journ.
of Geol., Vol. XIV, p. 599-617. Hustedt, Friedrich. 1920. Bacillariales aus Inner-Asien gesammelt von Dr. Sven Hedin.
Southern Til>et by -Sven Hedin, Vol. VI, part IV, Stockholm. 1925. Bacillariales aus den SalzgewJissem bei Oldesloe in Holstein. Mitt. Geogr.
Ges. Naturhist. Mus. Liibeck, 2. Reihe, H. 30, S. 84-121, 1925. Hutchinson, G. E. 1933. Limnological Studies at High Altitudes in Ladak. Nature,
Vol. 132, p. 136. KoLBE, R. W. 1932. Grundlinien einer allgemeinen Okologie der Diatomeen. Ergebn. d.
Biologic, Bd. 8, 1932. KoHLER, Hilding. 1925. Untersuchungen iiber die Elemente des Nebels und der Wolken.
Medd. fr. Statens Meteorol.-Hydrogr. Anst., Bd. 2, No. 5.
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Leightom, Marshai.i. O. 1905. Field assay of water. U. S. Cienl. Survey: Water Supply
and Irrit;atit)n Paper No. 151 (Series L, Quality of Water, 11), Washington,
1905. LuNDQViST, Ci. 1927. Bodenablagerungcn und i'.ntw iiklungstypen der Seen. Die llinnen-
gewiisscr, Bd. II, Stuttgart. LvDEKKER, RiciiAKD. 1883. The geology ui tlie Kashmir and L'haniba Tcrrittiries, and the
British District of Khagan. Mem. Geol. Surv. India, Vol. XXII, Calcutta. Meinardus, W'. 1934. Die Nieder.schlagsverteilung auf dcr I'.rde. Mutenrdl. Zcilscln-.,
Bd. 51, H. 9, S. 345-350. Meister, Fr. 1932. Kieseiaigen aus Asien. \'erl. Gelir. iMirntniger, iierhn. Mereschkow.sky, K. S. 1906. Diatdniowyja vordomsli 'ril)cta, in Kozlov. P. K., Mon-
golija i Kain. Trudy ekspedicii Imp. Russkago geograficeskago obscestva, sover-
senoj V 1899-1901 gg. T. 8. S.-Peterh. 1907-08, 4 : o. MiH.LER, LoTTE. 1933. Zur Frage der Tiefencirkulation im Indi.schen Dzean. Ann. d.
Hydrograpliio und Marit. Metcorologie, Jahrg. 61. Naumann, Einar. 1918. Uhcr das Nachweisen gewisser Gallertstrukturcn hci Algen mit
gewohnlichen Fari)stiften. Zcitschr. f. wis.sensch. Mikmskopie n. f. mikrosk.
Techn., Bd. 35, S. 243-244, 1918.
1932. Grundziige der regionalen Limnologie. Die Binnengewasser, I'd. XI, Stuttgart.
NoRDisK Vari.ds.\tlas. Stockliolm, 1926.
Oestrup, Ern.st. 1909. Beitriige zur Kenntnis der Diatonicenllora des Kossogoilx-ckcns
in der nordwestlichen Mongolei. Pledwigia, Bd. 48, S. 74-100. PoRECKij (Poretzky), V. S., JocsE, A. P., et Sesukova, V. S. (SniEscinKovA, W. S.).
1934. Sur les Diatomacees de la prescju'ile de Kola c't la composition microsciipicjue
de ses diatomites. Geomorfologiceskij Institut, Trudy, V'yp. 8, Leningrad.
(Russisch.) Schlagintweit-Sakunlunski, Hermann, von. 1869-80. Reisen in Indien und Ilm-lia-
sien, Bd. I, 1869; Bd. 11, 1871; Bd. Ill, 1872; Bd. IV, 1880. Jena. (1871) 1874. Untersuchungen iiljer die Salzseen im westliclien Tibet und in 'i'urkis-
tan. 1. 'ilu-il : Rupclni und Pangkong; das ( iebict der Salzseen im westliclien
Til)et. .\l)h. mathcni.']>hysik. CI. d. krMiigl. haycrischcn Akad. d. Wisscnsch.,
Miinchen. ScuoTT, Gerhard. 1933. Die jidirlichen Nicderschlagsmengcn ;uif dcni Indischcn und
Stillen Ozean. .\nn. d. I lydrographie u. Marit. Meteorologie, Jahrg. 61. SuPAN, A., und Obst, [•'.. 1927. Grundziige der physischen F.rdkundc, l!d. II, Berlin und
Leipzig. Sorlin, Anton. 1927. Till Indi.-iland . . . , .Stuckhdlm. Thomasson, 11. 1''26. Methodcn zur Cntersuchung der Mikniphyten der linmischen
Literal- und I'mfundalzone. Ahdcrhaldcn : llandbuch der liinl. ArheitsnKth' 'den.
IX : 2. Ule, Willi. 1892. Die l>estimmung der Wasserfarbe in den Seen. Petermanns Mitteil-
ungen, Bd. 38, S. 70-71, 1892. Wynne, A. B. 1878. On the Geology of the Salt Range in the Punjab. Mem. Geol. Surv.
India, Vol. 14.
Explanation of Plate XI.
Fig. 1. Aspect of Vegetation around Dal Lake. Note: the natives are collecting water-plants (Trapa?). G. Jarring, July, 1935 (Copyright).
Fig. 2. Lsland with luxuriant vegetation, Dal Lake. Figures 1 and 2 give a good idea of the richness of the Kashmir I>akcs in contrast to the Ladak Lakes. G. Jarring, July, 19,35 (Copyright).
Fig. 3. The flat, flooded shore of W'ular Lake in the midst of the great Kashmir-Basin. G. Jarring, July, 1935 (Copyright).
Fig. 4. Mitpal Tso. G. E. Hutchinson, August, 1932.
MEM. CONN. ACAD., VOL. X.
PLATE XI.
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ARTICLE XIII
REPORT ON MYRIAPODS By F. Silvestri (Portici)
(Received February 12, 1935)
The collection of Myriapods made by Mr. G. E. Hutchinson as biologist of the Yale North India Expedition contains specimens of 3 species of Chilognatha, 2 of S\'inpli\la and 8 species and varieties of Chilopoda, of which last 6 are described as new.
Having before us the results of both this expedition and those of the "Niederlandischen Expedition in den Karakorum" we might deduce that the Chilognatha are almost absent from high altitudes, only a single species of Polydesniidae {Kashmiriosoma contortipes Schub.) having been collected at an altitude of about 3200 ni. ; but it is necessary to search more, in very special ground, before definitely drawing such a conclusion.
Figure 1. Kashmiriosoma contortipes, specimens of the Y. N. I. E. : 1, copulatory leg seen from the internal side; 2, the second joint of the same without the apical large and contorted branch, which is represented isolated in two positions in 3. and 4.; 5, anterior view of inferior part of -Ith segment of the male: A, first joint of the fourth pair of legs ; Ci, lower lateral carinae ; Pe, external median process of the external branch of the copulatory legs ; Pi, internal median process of the copulatory legs ; Pr, sternal processes between the 4th pair of legs; Ps, internal branch (pseuduflagellum), St, sternum.
Both the above-named expeditions collected a number of Chilopods of the family Litho- biidae, which family appears to attain higher altitudes than any of the other Myriopods. This can l)e understood when we bear in mind the predacious habits of the Lithobiidae, which prey on other Arthropods, while the Chilognatha need decomposing vegetable matter, and a quite high humidity. wSpeciinens of Lithobiidae were collected up to altitudes of alxmt 5100 m.i
' The highest locah'ty from which Myriopods were obtained was L Z2, on the eastern slope of the Shakya-la, at about 5250 m. Immature and so indeterminable specimens of Lithohius here occurred among tufts of sparse grass and withered herbaceous vegetation.
Mem. Conn. Acad., Vol. X, Art. XIII, June, 1936.
242 RF.rORT ON MYRTAPODS
W'itji regard to the species of Litliohiiis here recorded, it must I:)e noted that tlie speci- mens beinjj few and nut all in adult condition, it was not possible to appreciate the value of the range of variability of some characters, as for instance that of spines on first joint of the legs of the posterior part of the body; therefore I have, fur the present, preferred to retain specimens sliowing such variation as varieties. It will he necessary, later, when more numerous specimens are available, to establish the range of variability of the characters under consideration.
As to the generic reference of the greater part of these IJthohiidac, notwithstanding the characters of the antennal joints and of the teeth of maxillipedes, I have referred them to the genus Lithobius s. 1. and not to Porubiiis proposed for forms possessing such characters by Attems, attending a more scientific revision of the old genus.
CIIILOGNArilA
Family POLYXENIDAE
Polyxcnus sp.
One young specimen of but 8 segments was collected at K 2. Takht-i-.Sulaiman, Srinagar, 20 March 1932, at c. 1585 m.
Family I'OLYDESMIDAE
Subfamily Strongylosominae
Kashmiriflsoina contortipcs Schubart
Through the kindness of Dr. J. B. Corporaal 1 have Ix'cn able to compare the type material (3 specimens) of Kashmiriosoma contortipcs Schubart, collected by the Dutch Expedition in Kashmir (Sind Valley and Matayan near Dras ) with the few specimens (1 male, 2 females) of the Yale North India Expedition collected also at K 7i, Matayan (under stones) at about 3170 m., 20 May 1932. The only appreciable difference I have found is in the copulatory legs, which in the typical specimens have not the external spine (Text-figure 1, 1-2, S) on the second joint well developed, and have a shorter process at the base of the pseudoflagelkun. As the typical and paratypical material is too scanty and does not permit us to ascertain the range of the variation of the characters in (piestion, I refer the specimens of the Yale Expedition to the same species.
Subfamily Polydesminae
Opisthoporodesmiis sp.
One young female specimen with only 18 segments, which is too immature to refer to any of the descril)ed species or to descril)e as new. Localitv: Niknri Hills: Pvkara at c. 2133 m.
REPORT ON MYRIAPODS
243
SYMPHYLA
Family SCOLOPENDRELLIDAE
Hanseniclla sp.
In tlie collection of the Expedition there are 2 specimens : 1 young- and 1 in moult, which very probably belong to H. subunguidulata (Immsj, but the state of the specimens does not permit a certain determination.
Locality: Nilqiri Hills: Ootacamund at 2316m., 8 Nov. 1932.
Symphylella sp.
One adult .specimen from Takht-i-Sulaiman, Srinagar, at about 1.t85 m., 20 March 1932, has characters related to those of 5". simplex (Hansen) Bagnall, described from the Island Koh Chang, Gulf of Siam, but being poorly preserved, is not fitted for a complete study.
Figure 2. Gcophihis iiilcniiissus var. crenulata: 1, corporis pars aiitica prona; 2, eadem stipi'na ; 3, labruni ; 4, pedum niaxillariuin unguis ; 5, segmentum decimum pronum ; 6, idem supiinim ; 7, segnieiituin quiu- quagesimum supimim ; 8, pes paris decimi ; 9, corporis pars postica prona ; 10, eadem supina.
244 RF.rORT ox MVRIAI'ODS
CIIILOPODA
Family GEUl'lllLlUAE
Mccistoccplialiis ccf^lialotcs Mein
One specimen from (N17) Nilt;iri Hills: Ootacamund, dam above Dho1)i's quarter c. 2.M6m., 15 Nov. 1932, and a very _viiun_<4' specimen from (K55) Kans^an (Rest House) c. 1766 m., 16 May 1932.
GcophHiis iiifrniiissux Silv.
var. crenulata nov.
Text-figure 2
S Corpus stramineum capite fiavo-ferrugineo.
Lamina cephalica paullum latior quam longior setis brevibus modice numerosis instructa, antennae articulis elongatis, decimo exempli gratia c. duplo longiore quam apice latiore, pedes inaxillares fiexi capite retracto laminam cephalicani tractu brevissinio haud superantes, subcoxis et articulis ceteris inermibus, articulo ultimo tantum tuberculo interno basali per- parvo aucto, ungue l^ene arcuato margine interno maxima pro parte tenuissime crenulato.
Tergita setis brevissimis parce numerosis instructa.
Sternituni priinuni area subpostica mediana poris 8 instructa, stemita 2'"" ad 18"'" fovea antica transversali et area mediana subpostica porosa ut Text-figure 2, 6 demonstrat, nee non setis nonnullis brevissimis instructa ; sternita cetera areis duabus parvis sublateralibus subposticis porosis, in segmento 19"^ poris 10+5, in segmento jienultimo 5 + 5.
Pedes breves parum setosi, ungue seta basili antica breviore et postica brevissima.
Scgmentum ultimuin pedifcrum scuto sulitrapcznideutn, subcoxis ])oris 4, (juorum duo interni magni partim obtecti et duo sublaterales externi parum niinures, sternito trapeze mleum c. 1/3 ad basim latiore (juain longiore, pedibus crassiusculis, articulis 1-5 sul)tus setis brevi- oribus numerosis instruct!, ungue terminali parvo.
Pedum paria 57; long, corporis mm. 22; long, antennaruni 3; long, pedum paris decimi 0.52, ultimi 0.70.
Habitat. Exemplum typicum ad "L 20 Ravine above i I imis (lonpa at 3525 m. 12 June 1930," lectum est.
Observ'atio. Varietas haec a forma typica ("Matayan near Dras" Xicdcrl. Exp. Kara- koriiiii) labri parte mediana denticulis sat distinctis, pedum maxillarium ungue margine interno minutissiine crenulato salteni jim teniimre distinguenda est.
REPORT ON MYRIAPODS
243
Family LITHOBIIDAE
Litliohiiis (Arcliilithobiiis) clcctiis Silv.
var. imminuta nov. Text-figure 3 S Corpus sublatericiuni.
Lamina cephalica paruiii latior quam longior setis sparsis brevioril)us (in exempio typico maxima pru parte al^ruptisj instructa ; antennae 20-articulatae (iaeva in exempio typico 19-articulata) articnlis elongatis, decimo duplo longiore quam latiore omnibus setis numerosis brevibus instructis; oculi ocellis 11 compositi subtriseriatis, ocello postico a ceteris aliquan- tum remoto; pedes maxillares antice aliquantum angustatis margine externo ad setae anticae latus externum paullum latiore, margine antico medio profunde inciso, utrimque dentibus 2 + 2 et seta externa breviore, superficie pone dentes setis paucis brevibus et brevioribus instructa.
Figure 3. Lilhohius (Arcltilitlwbiits) elcctus var. imminuata: 1, capitis regio ocularis; 2, pedes maxillares; 3, pedum maxillarium subcoxarum pars antica magis ampliata; 4, pes paris decimi.
Tergita omnia angulis posticis rotundatis, margine postico tergiti segmenti penultimi ])editeri vix sinuato ; tergitum segmenti ultimi pediferi minus quam 1/6 longius quam latius postice vix sinuatum; sternita setis sparsis brevioribus a segmento 12° postice aliquantum
magis numerosis mstructa.
Pedes spinis primi paris 1,0,3, 1, 1
0,0,2,2.0
paris ultimi
0, 0, 2, 3, 1 1,0.3, 1,0
paris decimi
0, 0, 2, 2, 2
paris penultimi
0, 0, 2, 3, 2 ' , subcoxis segmenti 13' etiani spina supera antica
0, 1,3,2, 1 ' ' 0, 1,3,2, 1
arniatis, pedes 14' et 15' quam praecedentes aliquantum longiores et cressiores, facie interna ab articulo 4'^ ad tarsum inclusum poris glandularibus numerosis instructa, practarsi ungui- cula interna breviore. Poris subcoxalcs 4, 4, 4, 4 (in parte laeva anormaliter 3, 5, 4, 4), cir- culares, uniseriati.
246 REPORT ON .MVKIAPODS
Apjiendices geiiitales niininiae, tuberculiforines, convcxae, setis trilnis instriictae.
Lung, corporis mm. 13; lat. laminae cephalicae 1.56; long, antennarmn 5; long, pedum paris decimi 2.80, paris ultimi 5.
Habitat. Exemplum descri])tum ad "(K 7?>) Matayan m. .il70 under stones near K 72) 10 May 1932" lectum est.
Observatio. X'arietas haec a /.///(. (./.) clccliis fnrina t)pica (Sanju, Karakasli X'alley, Niederl. Exp. Karakorum) , statura minora ocellis minus numerosis, poris glandularilius sul)- co.xabbus minus numerosis distineta est.
Lilhoh'nis (.Ircltililliobiits) iirclus Si! v.
var. secessa nov. Text-figure 4
Corpus avelianeum.
Lamina cephalica paullum latior quam longior; setis pauciorilnis l)revissimis sparsis instructa; ocuii ocellis 13-16 longitudinaliter 4-.seriatis in exempio uno oculus iaevus anor- maliter ocellis tantum 9 (3-seriatis) ; antennae breves, 20-articulatae, articulis longioribus quam latioribus (decimo c. 2/5 longiore quaui latiore), omnibus setis brevissimis numerosis instructis; pedes maxillares subcoxis antice angustatis angulo externo rotundato, margine externo obliquo, margine antico medio sat profunde sinuato, cetero dentibus perparvis 2^2 et seta brevi laterali aucto, superficie infera postdentali setis tantum brevioribus instructa.
Tergita niarginata angulis posticis plus minusve rotundatis (hand acute pruductis) ; ter- gitum segmenti ultimi pediferi jiarum longius quam latins, margine postico ]iaullum sinuato; sternita setis brevissimis sat numerosis instructa.
00322 . 00 3 22
Pedes spinis primi paris ' ' J J .y , paris decimi ' ' V -,' ^ . spinis minoribus apice
\Jy \Jj ^f tjj ^ \Jy \Jj v), O y t^
denticulato, spina apicali ventrali mediana articuli decimi mm. 0.31 longa, tarso articulo ])rimo quam secundus fere dupio longiore ct ambobas setis brevioribus mnnerosis instructis,
, . . 1.0, 3, 2, 2 ... 1,0,3, 1,0 ,. , , . ,
pans penultnni ^ ^ ? ' I^'""'^ nltnni (m exempio uno pes Iaevus articulo
secundo supra ctiam spina breviore aucto), ungue tcrminali seta basali sat longa aucto, articulorum 4-7 facie interna poris glandularibus ob.sessa, articulo sexto quam cjuintus paul- lum breviore; pedum j)aris 12' et 13' etiam subcoxa spina supera antica brevi robusta apice denticulato aucta.
Pori glandulares subcoxales 6-7, 8, 8, 8 subuniseriati, rotundi.
Feminae appendices genitales calcarilms 2 + 2 robustis, ungue terminali simplici.
Long, corporis ad mm. 18, lat. laminae cephalicae 2.10; long, antennarum 6.5; long, pedum paris decimi 4.5, paris ultimi 8.5.
Mas juvenilis quam femina minor (corporis long. mm. 13) oculis ocellis 10 compositis, poris subcoxalibus 5, 6, 6, 6.
Mas adultus. Feminae statura subsimilis, oculis ocellis 12-13 cf)mpositis, poris sub- coxalibus ul in femina, corporis parte postica steriiaruni ct ]>cdum setis ut mas tyi)icum (cfr. fig. Niederl. Exp. Karakorum, Zoologie, p. 196, fig. II).
Habitat. Exempla nonnulla vidi ad L 32a Zung-Lung, Ladak (.Typi) c. 4400 m. under stones in stream bed, wet place, 24 June 1932.
REPORT ON MYRIAPODS
247
Figure 4. Litlwbius {ArchUithobius) eleclus var. secessa: 1, capitis regio ocularis; 2, capitis regio ocularis exempli alii; 3, pedes maxillares; 4, . pedum maxillarium suhcoxarum pars antica; 5, pes paris primi ab articulo secundo ; 6, pedum paris decimi, minus quam primus ampliatus; 7, ejusdem tarsi apex et praetarsus mafjis ampliatrs ; 8, pes paris penultimi a facie externa inspcctus ; 9, pes paris ultinii a facie externa inspectus ; 10, spina dorsualis articuli tertii pedis ultimi magis ampliata; 11, pedis ultimi tarsi apex et praetarsus extcrne inspecti ; 12, feminac pars postica supina.
Paratypi ad Ladak L 62 Nyin,L;-ri, alt. 51 14 in., 26 July 1932; excnipla tria juvenilia ad L 68 An-zunna (camp below Anem La) 4890 ni. 1 Aui,aist 1932; niareni juvcnileni ad L 53 Sachuk-kont^ma 5160 m^ 15 July 1932; exempla mascuiina 3 plu.s niinu.sve juvenilia ad Tso Nyak region (western end) c. 4245 m., 12 August 1932; 1 9 e 1 c5 juvencs ad L 20, Ravine above Himis gonpa, 3525 m., 12 June I'*32; 1 pullus L 68a between An-zin-nia and Dambu-guru, under stone, c. 4725 m., 1 August 1932.
Observatio. Varietas haec a forma typica Lith. clcctiis Silv. femina adulta ocellis magis niunerosis pedibus posticis magis elongatis, subco.xis segmenti pediferi 12' etiam spina dor- suali armatis, ungue pedum paris ultinii seta basali sat longa instructa distinguenda est.
248
Klil'OKT ON MVKIAI'ODS
■^ Lithobius (Archilithobius) bispinosus sp. n.
Text-figure 5
Corpus pallide testaceum.
Lamina ccplialica vix latior quani l(in>;ior, setis taiilum paucissimis brcvissiniis instructa; oculi ocellis 16, quaruin 15 inter sese adiacentes, subtriseriatis et ocello singulo postico majore a ceteris paruin remote; antennae 20-21 articulatae, articulis elongatis, decimo c. 34 longiore quam latiorc, ultimo c. duplo longiore quam latiore, omnibus setis brevissimis numerosis instructis; pedes niaxillares subcoxis antice angustatis angiilo extenio et margine cxtcrno convexiusculis, margine antico medio profunde sinuato, cetero deiitilms 2-1-2 par\is arniatn et seto brevi externa, superficie infera (postdt-ntali ) setis nonnullis brevilius et lirevioribus instructa.
Tergita angulis posticis subrectis vcl late mtundatis, margine postico etiam tergiti seg- ment! pcnultinii ])cdifcri vix sinuato; tergitum segmenti ultimi pediferi c. j4 longius quam latins, longius quam in specicbus praccedentibus ct scgmentuni anale fere omniud obtegens; sternita setis paucis sparsis brevissimis instructa.
Figure 5. Lithobius (Archilithobius) bispinosus: 1, capitis regio ocularis; 2, pedes niaxillares; 3. pedum maxillarium subcoxarum pars antica magis ampliata ; 4, corporis pars postica prona ; 5, idem supina; 6, scgmenta genitale et analc supina ; 7, pes paris ultimi a facie interna inspectus ; 8, cjusdcm tarsi apex et prae- tarsus subtus inspecti.
REPORT ON MYRIAPODS 249
T3 , 0,0, 2, 1, 1 . , . . 0,0, 2, 2, 2
Pedes spims prinii pans q q ^ ^ :, , paris decinii q q ^ ^ '^ ' P^""'^ penultiini
2, 0, 2, 1, 1 . . 1, 3, 3, 1, 0
o' 1 3 3^' "'*''"' q' i' 3' 9 1 ' pedes pariuni 12", 13', ut 14', supra antice spinis brevi-
oribus robustis duabus armati ; pedes omnes tarso in articulinis duobus bene diviso, articu- lino prime quam secundus parum longiore, secundo, praesertim infra, setis numerosis I)revi- bus instructo, praetarsi ungue mediano rol)usto, unguicola postica quam antica pauilum breviore et robustiore; pedes paris 14' (juani praecedentes aliquantum crassiores et longiores, articuli 4' usque ad tarsum inclusum facie postica (vel interna) poris glandularibus mimimis pernumerosis instructa ; pedes paris ultimi quam praecedentes, praesertim articulis quarto et quinto, crassiores, poris glandularibus praecedenti similes, praetarso ungue seta externa brevi et unguicola interna breviore composite.
Pori glandulares sulxoxales 6, 7, 7, 6.
Appendices genitales ebsoletae, minimae, forma tuljercoli convexi manifestae.
Long, corporis mm. 17; lat. laminae cephalicae 2.30; long, antennarum 6.50; long. pedum paris decimi 4 paris ultimi 3.20.
Habitat. Exemplum typicum tantum vidi ad "L 27-28, Digar Polu valley" lectum at 4050 m., 21 June 1932.
Observatio. Species haec a Lithohius (P.) clcctus Silv. pedum parium 12', 13', et 14' subcoxis, spinis duabus superis armato et tergito segmenti ultimi pediferi majore, nee non pedibus ultimis crassioribus facile distinguenda est.
Lithohius {Archilithobius) crraticitlus Silv. var. plurispinata nov.
Text-figure 6 Corpus testaceum.
Lamina cephalica subaeque longa atque lata setis sparsis I)revibus et brevioribus non- nullis instructa; ocuii ocelli 6-7, longitudinaliter subtriseriatis, ocello postico a ceteris ali- quantum remoto; antennae 20-articulatae, articulis elongatis, decinio c. 1/3 longiore quam ad apicem latiore, omnibus setis numerosis brevibus instructis; pedes maxillares subcoxis antrorsum lateribus convergentibus, margine extemo obliquo baud vel vix convexo, margine antico medio profunde sinuato, utrimque dentibus 2 armatis et seta externa quam dentes pauilum longiore, superficie infera pone dentes setis paucis Iirevibus et lirevinribus instructa.
Tergita omnia angulis posticis subrectis vel rotundatis, setis paucis lireviljus et l)revi()ri-
bus instructa; tergitum penultimum margine postico pauilum sinuato, tergitum ultimuni
c. 1/5 longius cjuam latius, margine postico parum sinuato; sternita setis paucis brevil)us et
setis brevioribus vel l)revissimis parum magis numerosis instructa.
^ , 0,0. 1,2, 2 . , . . 0, 0, 3, 2, 2 . , . .
Pedes spnns pans prnni „ ,. ^ - „ , pans decuni n n -2 -i o < P^^s penultuni
1,0, 3, 1, 1 . , . . 1,0,3, 1,0
r^ ■. ^ ? 1 ■ P^'ii'is ultuni -— — - ; pedum 11-15 articulo primo (subcoxa) supra spina
brevi ad apicem armato, pedes penultimi et ultimi aliquantum longiores et crassiores et facie interna ab articulo quarto ad tarsum inclusum poris glandularibus obsessa, pori ultimi ])rae- tarso unguicula interna breviore, seta proximali externa brevi. Pori glandulares suljcoxales 3,4, 4, 3 vel 3, 4, 4, 4.
;5o
REPORT ON MYKIAPODS
Figure 6. Lilhohlus (Architilhohius) crraticuUis var. plurispinata: 1, capitis regio ocularis; 2, pedes maxillares; 3, pedum niaxillarium subcoxaruiii pars antica; 4. pes priini paris (semper ab articulo secuudo) ; 5, pes paris decimi ; 6, pes paris peiuiltimi a facie externa inspect! ; 7, pes paris ultimi etiam a facie externa inspecti ; 8, feminae corporis pars postica supina ; 9, feminae appendix genitalis magis ampliata supina; 10, fcminae appendicis genitalis articuli secundus et tertius proni.
l'"eininac appendices geiiitales calcariuni pariijus lUiulms, uiiyiie termiiiali apice iiitej^m, sed iiicisione iiiarginali externa infera, ali(|iiantnm longe a1> apice, affecta, articulo pemiltimo supra externe setis I^rcviorilms subspinifonnil)tis 6-7 et articuli i ultinu) supra etiain setis siinilibus duobus.
Maris appendices genitales perparvae tuberculi formes.
Long, corporis ad luni, 10; lat. laiuinac cephalicae 1.20; long, anteiinaruin 3.25; long, pedum paris decimi 2.10 paris ultimi 3.40.
Habitat. L 32, Zung-Lung, Ladak (2 9 9 and 1 $) 4224 m.
Oljservatio. Varietas haec a forma typica (N. Picngal, Kashmir) spina apicali supera etiani a pedum pare decimn primi) incipiente, saltern diversa est.
Lithobius (Archilithobius) materiatus sp. n.
Text-figure 7 9 Corpus pallide castaneum.
Lamina cephalica subaecpie longa atque lata setis paucis brevioribus instructa; oculi
(icellis 10 longitudinaliter triseriatis; antennae 19-articulatae, articulis longioribus (|uam
latioribus, articulo decimo, exempli gratia, fere 1/3 longiore quam latiore, articulis omnibus
setis numerosis brevioribus instructis, articulo ultimo (|uam penultimus ])annii longiore; pedes
maxillares margine laterali alicpiantiun pone marginem anticum sinuatinn, siiui subrecto,
margine antico ipso parum pone li!)ellam marginis antici interni ])edum maxillarium articidi
secundi pertimente, medio profunde sinuato et utrimque dentibus duobus et seta externa brevi
aucto.
REPORT ON MYRIAPODS
251
Figure 7. Lithobius (Archilithobius) materiatus: 1, capitis regio ocularis; 2, pedes inaxillares; 3, pedum niaxillarium subcoxarum pars antica ; 4, pes primi paris (ab articulo secundo) ; 5, pes paris decimi ; 6, pes paris penultitni a facie externa ; 7, pes paris ultimi a facie externa ; 8, paris ultimi tarsi apex et praetarsus a facie externa inspect! ; 9, feminae corporis pars postica supina ; 10, feminae appendix genitalis, supina, magis ampliata.
Tergita angulis posticis plus minusve rotimdatis setis nonnullis brevissitnis instructa;
tergitum segmenti ultimi pediieri paullum longius quam latins, niargine postico vix sinuate;
sternita setis nonnullis brevioribus instructa.
0, 0, 2, 2, 1 0, 0, 2, 2, 2
Pedes primi paris spinis n a' 9' ■;' 7 ■ P^ris decimi ' „' ' "' ^ , tarso divisione obsoleta
articulum praecedentem longitudine aequante, paris penultimi ' ' ' ' , paris ultimi
1,0,3,1,0
Q 2 3 2 1 ' ""b"s termmah submtegro (seta ventrali vix distincta), pedum parium 14' et
15' facie interna poris numerosis obsessa.
2B2 REPORT ON MYRIAPODS
Fori sulicoxales 4, 4, 4, 4 iiarvi, rotundi.
Appendices genitales calcarilnis robiistis 2 + 2, uiigue termiiiali paullum trilnliatM.
Long, corporis mm. l,i; lal. laminae ccphalicac 1.38; long, antcnnarum 4..^(); long. pedum paris decinii 1.80, paris ullimi ,v"0.
Hahitat. I'^xcmphmi typieum (antum vidi ad "Pass to Gya at base, Ladak lectum, at about 4203 m., Septeml)er 1932."
Observatio. Species haec pedimi maxillariuTn suhcoxarum forma, pedum brevitatc, tarso brevi divisione obsoleta et spinarum numero Ijene distincta est.
/.(iiiiyclcs albipcs (Pock.)
2 females collected at "Kandan (Kashmir), 1800 m., 16 May 1''32, under rotten wmkI." These specimens have antenn;c of 28 and of 29 joints; and in the remainder of their
characters are identical with a specimen from Srinagar previously figured by me (Niederl.
Exp. Karakorum, Zoologie p. 202, fig. VII).
ARTICLE XIV
REPORT ON DJPLURA AND THYSANURA
By F. Silvestri
(Reckivkd Fedruary 16, 1935)
The Diplura of the Yale Nnrth linha Kxpeditiun were not cnlkcted at very high alti- tudes, the highest locality being at 2159 m. in the Nilgiri Hills, where Indjapyx Harrisoni was obtained. Regarding the vertical distribution of this order, Mr. Hutchinson kindly wrote to me that "No Caiiipodeidac could be found in Ladak, in spite of considerable search being made." Some of the Thysanura on the other hand, such as Ctenolepisma sp. and Machilanus Hutchinsoni, found at 5315 m., were collected at very high altitudes in Indian Tibet.
DIPLURA
Family CAMPODEIDAE
Lcpidocampa Weberi Oud.
A few specimens of this species, which is very widely distributed throughout central and south Asia, and in the Oriental Islands, such as Japan and the Philippines, were col- lected at (K 55) Kangan, Kashnn'r, 1810 m., under rotten wood and earth, 16 May 1932, and a single specimen at (K2) Takht-i-Sulaiman, Srinagar, 1645 ni., 20 March 1932.
Family JAl'YdlDAE
Indjapyx Harrisoni sp. n. Te.\t-figures 1 and 2
Corpus stramineinn aliili>iniiR- a scgniciiln se])tini(i ocraceu, segmcntu deciino ferrugineo, fcrcipis niarginibus nigrescentibus.
Caput supra setis sat longis (mm. 0.28 longis) c. 15+ 15 et setis nonnullis brevioribus et brevissimis instructum; antennae 34-articulatae (anormaliter in e.xemplo uno antenna lacva 33-articulata el in cxeinpln alin antenii.i laeva 33-, dextera 32-articulata), articulis primo el secundo setis brevibus et brevioribus paucis, articulis tertio et quarto etiani setis paucis sat longis (tcrtii externa mm. 0.26 longa), articulis ceteris, decimo sexto exempli gratia, dimidia parte distali setis brevioribus sat numerosis et setis nonnullis proximalibus brevibus et non- nullis apicalibus parum brevroribus, articulorum 4-6 trichobothris consuetis, articuli quarti tricho1>othrio supero, fore basali, longo. Maxillae primi paris lobus internus laniinis pecti- natis 5.
Thorax. I'mnuliiin setis sat Inngis (ad mm. 0.28) 4 + 4, brevibus 3 H- 3 et 6 4 7 brevis- simis; mesiiniitum praescuto setis submedianis 1 + 1, scuto setis sat longis (lateralibus ad mm. 0.33) 5 + 5 et setis nonnullis brevioribus et brevissimis instructum, metanotum me.sanoto simile. Prosternum antice setis 3 + 3 sat longis, setis submedianis et sublateralibus, 4 + 4 sat longis et setis nonnullis brevioribus et brevissimis.
Mem. Conn. Acad.. Vol. X, Art. XIV, June, 1936.
254
REPORT ON DIPLURA AND TIIYSANURA
Figure 1. Indjapyx Harrisoni: 1, antennae dexterae pars proximalis prona; 2, cjusdem antennae articulus decimus sextus; 3, pes paris tertii a trochantero; 4, ejusdem pedis praetarsi apex et tarsus; 5, urotcrgiti scxti dimidia pars postica; 6, urotergiti septimi dimidia pars postica; 7, urotergiti octavi pars postica lateralis; 8, urosternitoruni primi et secundi dimidia pars; 9, urosterni primi dimidia pars postica magis ampliata; 10, maris regionis genitalis dimidia pars.
Figure 2. Indjapyx Harrisoni : abdominis pars postica a segmento sexto prona.
REPORT ON DIPLURA AND THYSANURA 255
Pedes setis nonnullis brevibus et minus nunierosis brevioribus et brevissimis instruct!, tertii paris tibiae seta apicali mm. 0.13 longa, tarsi setis inferis etiam mm. 0.10-0.14, praetarso quam tarsus parum magis quam dimidium breviore, ungue postico quam anticus aliquantum longiore, unguicula mediana attenuata.
Abdomen. Tergitum prinniin praescuto eidem metanoti simili, scuto setis duabus sub- medianis subanticis brevioribus, duabus submedianis subposticis brevibus et setis nonnullis brevissimis, tergitum secundum etiam seta submediana sublaterali brevi et seta postica sub- laterali brevi instructum; tergita 3-7 setis brevibus 5 + 5, setis brevioribus paucioribus et non- nullis brevissimis; tergitum sextum angulis posticis rotundatis; tergitum septimum angulis posticis retrorsum mm. 0.28 productis et parte apicali angustiore acuta; tergitum octavum lateribus postice acute aliquantum productis.
Urostemum primum organis subcoxalibus inter sese parum magis quam unius latitudo sitis, serie transverse setarum glandularium c. 32, longitudine inaequalium (mm. 0.06-0.08 longis) et serie postica setarum minimarum instructis, organo glandulari median© parvo dis- culis 13 instructo, superficie pone organa subcoxalia setis brevibus (mm. 0.06-0.08), numer- osis, transverse 3-5 inordinatim seriatis, pone organum glandulare medianum uniseriatis et setis minimis submedianis 1 + 1 subposticis, superficie cetera setis paucis brevibus transverse 4-subseriatis et setis brevioribus et brevissimis sat numerosis instructa; urosterna cetera setis paucis brevibus 4-subseriatis et setis brevioribus et brevissimis instructa, vesiculis minimis, stilis conicis consuetis.
Segmentum decimum supra mensum subaeque longum atque latum, acropygio parvo transverse rugoso, carinis lateralibus parum abbreviatis, superficie supera setis 3 + 3 sat longis, 4 + 4 brevibus et nonnullis brevissimis instructum. Forceps brachiis subsimilibus, dente submediano sat magno, margine praedentali tuberculis superis duobus et tuberculis inferis 4-6 (in exemplo uno tribus sed majoribus quam tubercula exemplorum ceterorum), margine postdentali denticulato vel crenulato.
Femina mari similis sed urostemi primi organis subcoxalibus setis glandularibus parum minus numerosis (ad c. 20) instructis.
Long, corporis ad mm. 10.5; lat. urotergiti septimi 1.24; long, antennarum 2.5; long, pedum paris tertii 1.80.
Habitat. Exempla typica tria vidi ad "Ponds beyond Cemetery, Pykara Rd., near Ootacamund, under leaves and moss, alt. c. 2195 m., 6 November 1932," lecta et tria ad "N 17 Ootacamund, by dam above Dhobi's quarter, alt. c. 2316 m., 15 November 1932."
Indjapyx Petrunkevitchi sp. n.
Text-figures 3 and 4
Corpus stramineum abdomine a segmento septimo ochroleuco forcipis marginibus fuscis.
Caput supra setis sat longis (ad mm. 0.13) c. 15 + 15 et setis minus numerosis breviori- bus et brevissimis instructum. Antennae 30-articulatae, articulis primo et secundo setis non- nullis brevibus et brevioribus, articulis a tertio (praesertim tertio et quarto) setis nonnullis sat longis (tertii supera externa mm. 0.12 longa), articulis ceteris decimo quinto exempli gratia dimidia parte distali setis proximalibus brevibus et ceteris 1-2 transverse seriatis ali- quantum brevioribus instructis, articulis 4-6 trichobothriis consuetis, quarti supero dorsuali subbasali longiore; maxillae primi paris laminis pectinatis 5.
256
REPORT ON DIPI.URA AND TIIYSANURA
>^ft^
FiGURK 3. Indjapyx Petrunkevitchi ; I, anlciiiiae doxtcrac pars proxiiiialis prona; 2, aiitciiiiariiin arlioiihis decimus quintus niagis anipliatus; 3, pes paris tertii a trochaiitcro ; 4, ejusdcni pedis tarsi apex et practarsus; 5, urotergiti sexti dimidia pars postica; 6, urotergiti septinii dimidia pars postica; 7, urotergiti octavi pars postica lateralis ; 8, urosternitoruni primi et secundi dimidia pars ; 9, urosteniiti priini partis posticae dimidia pars.
Figure 4. Indjapyx Petrunkevitchi: abdominis pars postica a segiiiento sexto proiia.
REPORT ON DIPLURA AND THYSANURA 257
Thorax. Pronotum setis sat longis (ad mm. 0.18) 5 f 5, setis brevioribus 4 + 4, et nonnulla brevissima; mesonotum praescuto setis submediaiiis sat longis 1 + 1, scuto setis longis (lateralibus mm. 0.23) 5 + 5 et setis c. 15 + 15 brevibus et brevioribus, metanotum mesonoto simile. Prosternum antice setis 3 + 3 sat longis, setis 4 + 4 brevibus vel sat longis (lateralibus subposticis) et setis nonnullis brevissimi instructum; sterna cetera similia.
Pedes setis sat longis, brevibus et brevioribus nonnullis instructi, tibiae seta apicali infera mm. 0.09 longa, tarsi setis inferis apicaliljus 0.07 longis, praetarso quam tarsus ali- (juantum magis quam dimidium breviore, ungue postico quam anticus aliquantum breviore, unguicula mediana breviore.
Abdomen. Tergitum prinium praescuto eidem metanoti simili, scuto setis submedianis subanticis brevibus, duabus submedianis subposticis sat longis et setis nonnullis brevissimis ; tergitum secundum etiam seta submediana-sublaterali subantica et seta postica laterali sat longis instructum; tergita 3-7 setis brevibus vel sat longis T -^7, quarum subantica et sub- mediana laterali, basi marginali infera, et setis brevissimis c. 20 + 20 instructis; tergitum sextum angulis posticis rotundatis, septimum angulis posticis retrorsum mm. 0.32 productis et parte distali angustiore acuta; tergitum nonum lateribus postice baud productis.
Urosternum primum organis subcoxalibus inter sese panun magis quam unius latitudo sitis, setis glandularibus 14 inter sese inaequalibus (mm. 0.20-0.28 longis) uniseriatis et setis posticis minimis etiam uniseriatis, organo glandulari mediano parvo, disculis c. 10 instructo, superficie ante organa subcoxalia setis numerosis brevibus transverse inordinatim 3-4 seriatis, ante organum glandulare medianum setis brevibus uniseriatis et setis minimis duabus sub- medianis subposticis, superficie cetera setis nonnullis brevibus 4-seriatis et setis nonnullis brevioribus vel brevissimis; urosterna cetera setis nonnullis brevilms transverse 4-seriatis et setis paucioribus brevissimis, vesiculis perparvis, stilis consuetis.
Segmentum decimum supra mensum suliaeque longum atque latum, acropygio breviore, late rotundato, carinis parum abbreviatis superficie supera setis 10-10 sat longis et setis non- nullis brevioribus et brevissimis instructa. Forceps brachiis subsimilibus, dente submediano sat magno, margine praedentali tubercula supero et tuberculis 2-3 inferis parvis aucto, margine postdentali denticulato.
Long\ corporis mm. 5; lat. urutergiti septimi 0.()7; long, untennarum 1.60; long, pedum paris tertii 0.90.
Habitat. Exemplum descriptum ad "Takht-i-Sulaiman, Srinagar, 20 March 1''32, alt. c. 1585 m.," lectum est.
Observatio. Species haec. clar. Prof. A. Petrunkevitch dicata, a specie praecedente abdomine parum magis setoso, antennarum artioulnruni numero et forcipis forma distinctis- sima est.
TUYSANVRA
Family LFPISMTDAE
Ctenolepisma longicaudata Esch.
Few specimens from (K86) Cliff alx)ve Wakka Chu River, 2 miles West of Kargil at 2740 m. This species has a wide distribution, at least in Africa and Asia.
258
REPORT ON DIPLURA AND THYSANURA
^ Ctenolepisma sp.
Specimens from (L68), Camp below Anem La at 5315 m., 1 August 1932, are in poor condition and it can only be said that they are very similar to Ct. mauritanica (Lucos).
Ctenolepisma sp.
A few specimens, also in such a poor condition that a description cannot be given, were collected at (L22) between Leh and Spitok (Spithug) at about 3350 m. The distribution of combs of setae of urotergites is as in the group Ct. Itncata.
Lepidospora ceylonica Silv.
One female collected at (N 17) Ootacamund (Nilgiri Hills) 15 November 1932, has the same characters as are shown by the typical specimens, which I described from Ceylon. This species was also collected later at Rotuna and Kumaon so that it appears to have a wide distribution, from North India to Ceylon.
Figure S. Machilanus Hutchinsoni: 1, oculi ct ocelli laterales supra inspect!; 2, oculus ct ocellus laevi supra parum oblique inspecti ; 3, antennae pars proximalis; 4, ejusdem antennae partis distalis articuli duo; 5, ejusdem partis distalis articulini duo magis ampliati; 6, feniinae palpus maxillaris; 7, feminae palpus labiallis; 8, feminae pes primi paris; 9, ejusdem pedis tarsus et praetarsus magis ampliati; 10, urosterni quinti dimidia pars; 11, ejusdem urosterni stilus magis ampliatus; 12, feminae urosterna 6"°' ad 9""' cum ovipositore ; 13, ovipositoris valvulae inferae laevae pars distalis; 14, ovipositoris valvuiae stiperae lacvae pars distalis; IS, maris pes paris primi a femore ; 16, maris urosterna octavum et nonum cum pcne et paramcris.
REPORT ON DIPLURA AND THYSANURA 259
Family MACHILIDAE
Machilanus Hutchinsoni sp. n.
Text-figure 5
Color ? (exempla typica squamis denudata sunt).
Oculi magni inter sese tractu quam oculi dimidia longitudine paullum longiore tangentes, oculus singulus c. 1/S latior quam longior; ocelli laterales transverse subrectangulares oculi dimidiam latitudinem fere aequantes.
Antennae in exemplis typicis baud integrae, parte sistente mm. 8 longa, articulo primo fere duplo longiore quam latiore, flagello gradatim attenuato articulis distalibus partis sis- tentis (mm. 7.4 longa) articulinis 12-14 compositis, serie transversali setarum brevium et seta alia nonnulla breviore instructis; palpi maxillares gradatim attenuati, forma et arma- tura consuetis ; palpi labiales articulo ultimo apicem versus gradatim parum latiore ut Text- figure 5, 7 demonstrat.
Thorax. Arcus thoracicus parvus late convexus ; pedes primi paris femore et tiljia quam ceteri parum crassioribus, secundi et tertii paris processu coxali longo, omnibus squamis paucis setis pernumerosis brevibus et tibia tarsoque infra setis brevibus robustis sub- spiniformibus biseriatis instructis.
Abdomen. Urosternum 2-6 pars mediana proximali lata, urosterni quinti exempli gratia c. duplo ad basim latior quam longior, subcoxarum angulo externo late subrotundato, angulo interne baud producto rotundato; vesiculae urosternorum 2 + 5 ulrimque duae; stili uros- terni quinti quam urostenium aliquantum breviores squamis paucis et setis brevibus numer- osis, a basi ad apicem gradatim robustioribus, instructi.
Subcoxae urosterni septimi angulo intemo retrorsum mm. 0.26 producto et margine rotundato, subcoxae segmenti noni angulo externo obtuso, angulo interno retrorsum acute aliquantum producto; stili segmenti noni quam subcoxae paullum breviores.
Ovipositor robustus plurianulatus, subcoxarum IX marginem posticum c. mm. 1.30 superans, anulis brevioribus, valvulis inferis anulorum superficie media setis nonnullis brevis- simis, laterali setis brevibus et anulorum 6 posticorum setis brevissimis obtusis robustioribus ut Text-figure 5, 13-14 demonstrant instructis, valvulis superis magis attenuatis setis sub- similibus. Cercus medianus quam corpus aliquantum et quam cerci laterales fere triple longior, setis ? (in exemplis tipicis abruptis).
Long, corporis ad mm. 13; lat. thoracis 3; long, antennarum ?; long, pedum paris tertii 5.5 stilorum IX-1.90, cerci mediani 14.
Mas. Palpi maxillares quam feminae paullum crassiores, setis? (in exemplo typico abruptis) ; pedes paris primi femore parte infera preapicale paullum inflata et setis brevibus numerosis sat robustis aucta.,
Subcoxae segmenti octavi angulo intemo oblique truncate, sul^coxae segmenti noni angulo interno acute retrorsum aliquantum producti, stili subcoxarum longitudine aequantes.
Paramera apice attenuato ad stilorum basis libellam pertinentia, setis brevissimis numer- osis instructa, penis quam paramera paullum longior.
Habitat. Exempla typica ad "Camp below Anem-la, L 68 at 5315 m., 1 August 1932," lecta vidi et exemplus paratypicum ad "Slope opposite Mulbe Gonpa, coll. G. E. Lewis, 27 May 1932."
ARTICLE XV
REPORT ON COLLEMBOLA
By J. R. Denis (Dijon) (Received August 2, 1935)
Les difficultes de recolte, principalenient aux hautes altitudes, ont fait que bon nombre d'echantillons ne sont pas en etat satisfaisant et que certains d'entre eux ne sont pas etudi- ables. J'ai cependant fait en sorte — tout en sauvegardant leurs restes precieux — que les echantillons a peu pres etudiables soient decrits aussi completenient que possible. Et si certaines de nies descriptions sont encore incompletes, j'ai cru devoir les publier quaiid menie, laissant a mes successeurs le soin de les parfaire.
Hypogastrnra aniiala (Nic.) Station: Edge of swamp, Phashakuri, near Pampur, 7-V-1932; 4 exempl.
Notes. Rapport Dens: niucron : 2.1 a 2.5. Rapport epine anale/g3 : 1.2 a 1.6. Rapport epine anale/mucron : 1.7 a 2.1. Rapport g3/mucron : 1.20 a 1.45. Rapport mucron/post- antennal : 1.40 a 1.60. Caracteres habituels, griffes avec une dent interne; ergots nets on indistincts.
L'espece pent etre consideree comnie a peu pres cosmopolite.
Hypogastrnra coiinimnis (Folsoni)
Station: Gagirbal, swampy pond, 1580 m. alt., l-IV-1932; 2 exemplaires.
Notes. Ces deux exemplaires concordent exactement avec ceux que j'ai pu voir jusqu' a present, du Japon, de Chine et de Formose.
Friesea excelsa n. sp.
Text-figures 1-4
Station: Ororotse La, top, 5500 m. alt., 11-VII T932; 4 exemplaires, qui me sont i)arvenus a sec et que j'ai du traiter par la potasse.
Diagnose. Taille 1 mm. environ, couleur noire. Pas d'epines anales. Revetement tres incomplet, mai.s, a en juger d'apres les insertions, il n'est pas impossible qu'il y ait eu des .soies ca])itees sur le tins. S yeux par cote, avec deux soies sur chaque champ oculaire. Organites autennaires mm rludiables, mais les ])iiils (ilfactifs d'aul. I \' sniil nombreux, longs, courbes et tres differencies. I 'ieces buccaies (Text-ligures 1 et 2) indubitabieinenl du genre Fricsca. (Iriffes sans dent interne ni ai)pendice enipodial. Sur I'ape.v des lil)ias, il y a au ninins (leu.\ ccrcles de soies eftilees, courbees a I'apex et legerenient eapitees. 'i'enaculum a deux dents bien nettes. Furca ayaiit uii developpement comparable a celui (|ue montrent les especes typiques d'Europe. Dens plus courte que la crete interne de g3, avec trois soies posterieures, sans mucron net — celui ci tout au plus represente par iin petit tuljercule (Text- figures 3-4).
Mem. Conn. Acad., Vol. X, Art. XV, Junk, 19.36.
262
REPORT ON COLLEMBOLA
Figures 1^. Friesea excelsa n. sp. (Obj. imm. 90Xoc.lO), 1, mandibule. 2, maxille. 3-4, mucro-dens
(assez defornie), tenaculum et g3.
Justification. Autant que je sache, la seule Friesea a 8 yeux par cote, connue jusqu'a present est F. grisea (Schaffer), forme antarctique. Ses epines anales la separent radicale- ment de la presente espece.
Proisotoma ladaki n. sp.
Text-figures 5-8
Station: Mitpal Tso, N. edge., 16-VIII-1932; 6 exemplaires de 1 a 1.4 mm.
Diagnose. Taille maximum observee: 1.45 mm.; couleur gris uni forme clair, pouvant se foncer sur le dos. Yeux noirs, 8 omnia, subegales, par cote avec 6-7 soies sur la plaque oculaire. Entre les antennes, une petite bande longitudinale pigmentee en noir occupe la place d'un ocelle. Postantennal de I'ordre de longueur de deux diametrcs d' omnia. Rapport post-
$^<s<^^^^^^
Figures S-8. Proisotoma ladaki n. sp. S, prolil. 6, postantennal avec la longueur de I'appcndice empodial (ae) et du niucron (m) a mcme echelle, pour le meme exemplairc, 7:g3. 8, mucrodens, profil, face cxterne. 8A, un mucron a plus fort grossissement.
antennal/niucron : de 0.95 a 1.35, niais le plus souvent au voisinage de 1.0. Rapport g3/post- antcnnal:de 1.00 a 1.50, mais, semble-t-il, le plus souvent au voisinage de 1.40. Organitcs antennaires habituels, soies olfactives d'ant. IV, greles et tubules d'org. ant. Ill, caches on decouverts, inclines ou droits selon I'etat de turgescence de I'antenne. Antennes plus courtes que la diagonale de la tete, .segments II et III subegaux, IV environ 1.5 fois le precedent. Tibias a subsegment distal incomplet mais trcs net, avec un certain nombre de soies greles, effilees et un pen courbes a I'apex, mais qui ne sauraient passer pour des ergots differencies (Text-figure 7), griffes minces et sans dent interne, appendice empodial effile. Tenaculum
REPORT ON COLLEMBOLA 263
a 3 dents et une forte soie au corpus. Furca de taille moyenne. Rapport Manubrium/dens- mitcron : 1.0-1.2. Manubrium n'ayant, sur la face antcrieure, que deux soies apicales. Dens (Text-figure 8) a face antcrieure a 3 rangs de soies et munie de rides obliques, a face pos- terieure plutot tuberculeuse qu'annelee, a deux rangs de soies. Mucron (Text-figures 8, 8A) plutot court par rapport a la dens, a cote tres courbe, mince et a trois dents, apicale plus petite, subapicale plus forte et externe sensiblement aussi forte que la subapicale; ces deux dernieres sont reliees par une lamelle, les autres lamelles sont sensibles; la base du mucron est renflee sur sa face posterieure: il s'agit d'un prolongement du bord dental.
Justification. Cette Proisoloma (s. str.) semble bien separee des especes voisines de par son mucron. Une forme de mucron a peu pres semblable se trouve chez P. bayoiiensis Mills (1931, Amer. Mus. novit., n°464, p. 2-3, figs. 1-6), trouvee en Louisiane. Ces deux especes me semblent tres voisines I'une de I'autre. Les seules differences portent sur la conformation des ergots (un scul, fort bien differencie dans I'espece americaine)^ — cette difference est faible ; sur le revetement dental qui n'a, dans I'espece americaine, de soies posterieures que dans la seule moitie apicale; enfin sur le nombre (4 centre 3) des dents du tenaculum. (Je n'ai pas observe le nombre 3 un nombre de fois suffisant pour afiSrmer que ce caractere signifie quelque chose.)
Ainsi, I'espece la plus proche parente de notre forme du Ladak serait une forme de I'Amerique du Nord. II est bien evident qu'on doit s'interdire, pour le moment, toute con- sideration d'ordre biogeographique.
Isotoma spinicanda Bonet
1930. Bonet F., Eos, 6, p. 249-51, fig. 1.
Stations: Nyag-tzu, lower camp, Ladak, dry bank, l-VIII-1932; 1 exempl. Ororotse-la, S. side, 5300 m. alt., ll-VII-1932; 2 exempl.
Mes cxemplaires concordent fort bien avec les donnees de M. Bonet et il n'est pas douteux qu'on ait bien a faire a I'espece indiquee. Celle ci n'est connue, pour le moment, que du Waziristan, pres de la frontiere de I'Afganistan. Comme I'a note M. Bonet, elle est tres voisine de 1'/. viridis et risque fort d'etre confondue avec cette derniere si on ne prend pas garde aux epines manubriales. M. Bonet indique encore d'autres afifinites. A mon sens, il est permis d'admettre qn'I. spinicauda est une forme paiearctique, mais il est encore plus raisonnable de ne rien dire.
f Orchesellides boraoi Bonet Text-figures 9-12
1930. Bonet F., Eos, 6, p. 253-5, fig. 2.
Stations: E. of Shakya-la, 5200m. alt., 25-VI-1932.
Ororotse-la, S. side, 5300 m. alt., ll-VII-1932.
Marsimik La, under stones, 5600 m. alt., 16-VII-1932.
Remarques. Mes exemplaires sont en mauvais etat et je ne puis etudicr correctement leur coloration. Bonet donne une figure de I'omementation mais signale que les dessins plus sombres peuvent s'estomper. Ce serait le cas de mes exemplaires. Meme certains sont tres pales et presque sans pigment.
2(A
REPORT ON COLI.EMBOLA
La -taillc pent atteimlre 2.6 mm.
Te rclcve, eiitrc la description cle IIonet et mes exemplaires 1) les re.ssem!)lanccs suivantes: Rapports Ant. II/I, III/II, Ics divers rappnrts tires des ti^riffes, en particulior A/1, qui varie de 1.2? a 1.45 et qui, d'apres la fiy. 2C dc I'A., est de 1.45 et le rappurt ae/d (3.10 dapres I'A.), i^-i 2.95 i 3.75.
Figures 9-12. Orchcsellidcs Boraoi Bonet. 9-10, deux e-xemplaires, I'liii hien colore, I'autrc pale. 11-12, griffes
et miicrou (irappartciiaiit pas an iiicme p .xemplaire) .
2") les differences suivantes: a) les yeux : je ne trcnive ([ue 6 yeu.K par cote (mais I'etat des exemplaires ne permet pas une etude appro fnndic) ct ce qui est plus jirave, les yeux C et D me seml)lent subegaux aux yeux .\ et W et nun ])as |)lus ])etits cumme I'indiqut' l;i fii^nre 2B de I'A. !>) la rci^ion dentale non anneleee est plus courte cliez mes exemplaires (|uc sur ceux de Bonet. c) Ant. I Will est, selon I'A. 1.7 environ, tandis que je ne trouve que 1.2 a 1.4.
II est aussi vain d'affirmer (|ue ces differences sont specifi(|ues (|ue de supposcr le con- traire. Seule I'expcrience prouvera ce qu'elles valcnt. Pour le mumcnt je me contente de mettre un ? devant ma determination; mais ce (jui importe surtout, c'est la trcs proche parente entre mes exemplaires et ceux de Bonet.
Les exemplaires de M. Bonet proviennent du W'asiristan, pres de la fmntiere aft^anc, et d'altitudes superieures a 1000 m.
Sur le genre Orclicscllidcs Bonet. — J'adopte ce genre pour de simples raisons de com- modite, mais je crois devoir faire, a son sujet, les reserves suivantes.
REPORT ON COLLEMBOLA 265
1 ) II lie se distingue pas d'Onhcsclld par It- iii)iiil>i-e iryt'iix, car nn sail (pie les Onltc- sclla peuvent avoir 8 yeux (G. et II. reduits) on seuleniciit (i. On sait aussi epic, dans la praticjue, le nonibre des yeux est souveiit indeterminable.
2) II ne se distingue pas d'Orcliesclla par la presence d'une sole lisse en face de I'ergot de p. 3, car cette soie lisse existe chez Orchesella.
3) L'aljsence de suhsegnientation d'ant. II le distingue assurenicnt des Orchesella eurfipe- ennes. En sera t-il toujours ainsi cpiand on connaitra inieux des 0)xhcscUa exotiques tclles que mon O. sinensis ou la subsegmentation en question est fort pen nette ? On sait ce que vaut ce caractere chez Dicranoccntrus et Heteromunis: pas grand chose.
4) La massue trilobee de I'apex d'ant. IV constitue, jusqu' a nouvel ordrc, le meilleur caractere distinctif qu'ait invoque M. Bonet. Ce n'est pas qu Orchesella soit depour\'u d'un lobe plus ou moins saillant a I'apex d'ant. IV, niais Orchesellidcs exagcre la saillie et la trilobe. II ne me semble pas que I'organite en question puisse etre dit retractile. En tout cas, il n'est aucunement comparable a la massue retractile des Entomobrya.
On pourra penser que les caracteres justificatifs du genre Orchesellidcs sont plut<'it sus- pects et qu'on ne manquera pas de trouver des formes telles qu'on ne saura si elles appartien- nent a Orchesella ou Orchesellidcs. O. sinensis m. en est une sans doute (mais je n'en ai pas vu assez d'exemplaires pour I'affirmer).
Mais, proinsoirement, Y Orchesellidcs de M. Bonet est assez distinct des Orcheselles connues jusqu'a ce jour pour meriter d'en ctre separe, afin d'alleger la classification, si pen claire, des Orchesella.
Sira brahamides n. sp.
Text-figures 13-16
Stations: Karpet, S. shore of Panggong Tso c. 4250 m., damp grassy place, 8-\'II-1932; 2 exempl. Tang-yar. Ladak. 4400 m. alt. env., under stones in .stream, 24— VI-1932; plusieurs excmplaires mais si deterinres que leur elude est impossible. Rapporles avec doute a I'espece ci-dessus.
Description. Taille : 1.6 mm. env., presquc incolore; sur la tete, seules les plaques oculaires et I'ocelle sont noirs. Aucune bande ne relie les yeu.x a I'ocelle (Text-figure 13). Avec les caracteres haliiluels du genre (Ecailles pointues, costulees; pas d'ccailles denlales cl mucron complet) on notcra la tres grande minceur des ecailles et la largeur relativement grande de I'appendice empodial qui est du type aigu. Les antennes sont cassees mais devaient avoir la taille baliituelle. Une elude statistique des elements des griffes et de la furca est impossible el on devra se contenter des figures ci-joinles.
Justification. Le genre Sira, tel que le comprennent la jjlupart des autcurs, pent se decomposer en deux groupes A et B avec un groupe AB (intermediaire?).
A. Appendice empodial tronque: .S". foniiosaua m. et S. hyolina Ilandscbin
AB. Appendice empodial paraissant tronque ("schrag abgestutzt" dit I'A.), sans doute efYectivement tronque, mais sans que les figures ou les textes permettent de I'affirmer formellemcnl : Sira abnipfa Schott, 5". binoculata C. B.
B. Appendice empodial aigu : les autres especes.
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Notce S. brahamides ajjpartient a cc groupe B. A Tintericiir dii groiipe B, on est force de s'en tenir a dcs caracteres d'omementation et de repartition geographique : cri- tcriums detestables.
Laissant de cote le groupe S. platani Nic, revise par Handsciiin et fait dc formes d'Eiirope et d'Amcrique du Nord, nous avons outre une serie d'espcces colorees, toutes carac- terisees par la tendance a former des bandcs transverses: 5". intcniicdia Schott (de Juan Fernandez), 5. Jacobsoni C. B. (de I'Archipel Malais, de I'lnde et des iles Hawai), S. japonica Folsom (du Japon), 5". parajacobsoni m. (de Madagascar) ; 5". purpurea Schott
Figures 13-16. Sira brahamides n. sp. 13, profil. 14, ^cailles d' abd. II. 15, gl(I) et g3(III). 16, mucrons.
(de Californie — d'ailleurs douteuse en tant que Sira), S. fricincta Schott (du N. Queensland) et S. variabilis Schiiffer (de la Terre de Feu). Je laisse de cote la forme hongroise: S- pal- lidipes Renter, que M. Stach signale dans son travail sur les Collemboles de Hongrie, mais qu'il ne doit pas avoir revue.
La seule Sira incolore ou a pigment unifprmement reparti, done comparable a la notre, est 5". brahama Imms (Proc. zool. soc. London, 1912, p. 99, PI. 8, figs. 43-44). Mais les descriptions de Imms sent tellement imprecises, qu'on ne pent tabler sur elles. Ainsi 1 'A. indique une griffe a deux dents, mais je me garderai bien de tabler sur cette particularite. Cependant 1 'A. figure un appendice empodial si allonge que je crois pouvoir justifier mon espcce en me fondant sur la difference de forme des appendices empodiaux. II s'agit d'une justification tres provisoire et je ne m'illusionne pas sur sa valeur. ^. brahama vient d'AlIa- hal)ad. Imms la decore du nom d'espece orientale. Je me garde de toute opinion sur les afiinites des Sira.
Sira nilgiri n. sp. Text-figures 17-20
Station: Hill south of Ootacamund, Rhododendron, env. 2378m. alt., 13-XI-1932; 1 exempl.
Description. Taille 1.5-1.6 mm., aspect general et dessins cf. Text-figure 17. La plupart des ecailles sont pointues, mais certaines sont arrondies au bout libre (Text-figure 18) ; toutefois leur striation est celle des Sira typiques. Antennes environ 1.5 fois la diago-
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nale de la tete, avec anneaux apicaux relativement nets et ant. Ill plus court que II. Grififes assez etroites (Text-figure 19), appenclice empodial etroit avec aile postero-interne nettement crenelee. Get appendice est du type aigu encore que, sous certaines incidences, les deux ailes externes puissant sembler confondues sur leur trajet apical (Nota: L'existence d'une telle
Figures 17-20. Sira nilgiri n. sp. 17, profil. 18, ecailles. 19-20, g3 et mucron au mcme grossissement (Obj.
90XOC.10).
disposition in'enipeche de considerer les formes du groupe AB comme veritablement tron- quees). Rapport manubrium/dens-mucron : 0.85 a 0.90. Region non annelee de la dens et mucron du type habituel. Rapport ae3/mucr. : 1.70.
Justification. L'appendice empodial est etroit et long comme chez 5". brahaiiia Imms, niais le type d'ornementation est different, cette derniere ayant une pigmentation uniforme. Je m'abstiens de toute consideration de [larcnte.
Parasira subornata n. sp. Text-figures 21-26
Stations: Dambu-guru, Ladak, 4603m. alt., dry bank, l-VIII-1932; 1 cxempl. Takht-i-Sulaiman, Srinagar, 1585 m. alt., 20-III-1932; 3 exempl.
Description. Tailie: 2.4— 2.8 mm. Aspect general ct dessins: cf. Text-figures 21, 22. Ecailles allongees, aigues du type Sira ; face anterieure de la furca sans ecailles. Yeux conformes a la figure 7C de Bonet pour P. ornata. Antcnnes sans ecailles Rapport: Ant./diag. ceph. : 2.20 a 2.55. Rapport: tronc/ant. : 1.4 a 1.6 Ant. II aussi long ou un pen plus long que III. Ant. IV/III:1.05 a 1.15. Massue apicale presente a ant. IV. Pas
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d'ecaille^ sur les pattcs. Tibias II ncttcment plus lon<;s que tiliias I. Griffe confornie a celle de P. ornata Bonet et n'cn different que par la position plus distale des dents laterales (et de la dent externa — que Bonet ne figure pas). L'appendice empodial, du type aigu, montre sur I'aile habituelle de tres fines crenelures (Text-figure 25). Kapport niucro-dens/manu- brium: 1.15 a 1.30. Region dentalc non annclce finissant graduellcment. Mucron (Text- figure) en crochet a epine basale.
Figures 21-26. Parasira subornata n. sp. 21-22, deux types d'orncmciitation. 23, corncules eii vue oblique. 24, ecailles d'abd. IV (obj. 60Xoc.lO). 25, gl(I) et g3(III) et 26, mucron au mtinc grossissement (Obj. 90XOC.10).
Justification. Cette espcce est extrcniemcnt voisinc de celle de Bonet. Le type d'orne- mentation est sensiblenient de ineme. Dans I'un de mes echantillons, on voit nieme une bande transverse sombre sur le dos d'abd. III. Cependant mes echantillons ne montrent pas la fine bande mediane figuree par Bonet (Text-figure 7A). Cette legcre difference ne m'em- pecherait certes pas de faire rentrer mes exemplaires dans le cadre de P. ornata et, si j'ctablis P. subornata, c'est principalement a cause de la position, nettement plus distale, chcz celle ci, des dents laterales de la griffe. Je ne serai d'ailleurs pas surpris qu'on vicnne a montrer que ce caractere est variable et qu'on reunisse P. subornata a P. ornata.
Notes sur le genre Parasira Bonet.
M. F. Bonet (1930, Eos, 6, p. 263 & seq.) a etabli le genre Parasira pour separer des Sira les formes a mucron falci forme et pourvu d'une epine basale.
Jusqu'a present je n'ai pas montre beaucoup d'ardeur a adopter les diverses classifica- tions du groupe — Sira-Lepidocyrtiis et je reste encore tres sceptique sur les genres Pseudo-
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sira, Mesira et Lepidocyrtinus — tout en reconnaissant que M. Bonet ait beaucoup clarifie les choses. En particulier je connais des formes (telles que Lepidocyrtus vexans m.) qu'on ne saurait actuellement classer. De plus j'estime qu'on doit laisser tomber le mot Mesira, car personne ne saura jamais ce que fut la Mesira squaiiwornata de Scherbokow — qui, par son ornementation se rapprochait sans doute du Lepidocyrtinus DoUfusi Carl : forme rare que pen d'auteurs ont vue. Cependant je souscris pleinement au genre Parasira de M. Bonkt et j'y inscris les especes suivantes :
Seira frigida Imms 1912, P. ornata Bonct 1930, Scira pallida Brown 1926, P. subor- nata m. et Sira villosa Borner 1903.
Toutes ces formes me semblent bien presenter les particularites requises pour faire partie du genre Parasira. On remarquera que I'aire de dispersion geographique du genre (tel qu'il est actuellement compose), comprend la Mesopotamie, le Wasiristan, I'Himalaya (Garhwal), le Kashmir, le" Thibet occidental et le Japon. En dehors de cette aire, le groupe est encore inconnu.
Lepidocyrtinus unifasciatus n. sp.
Text-figures 27-30
Station: Edge of I'hashakuri, near Pampur, env. 1585 m. alt., 7-V-1932; 3 exempl.
Description. Taille: 1.4 mm. env. Blanc avec une fascie noire sur le dos d'abd. III. Yeux noirs et individualises ou a pen prcs (Text-figure 28), pas de bande les reliant a I'ocelle. Antennes de 0.6 fois la longueur du tronc, a segments II et III subegaux, a seg- ment IV env. 1/3 plus long que III et non annele — ce qui devrait faire placer I'espcce dans
Figures 27-31). Lepidocyrtinus unifasciatus n. sp. 27, profil. 28, tache oculaire. 29, ecaille. 30, niucroii.
le genre (ou sous-genre) Pseudosira et je I'y placerais si j'etais convaincu que les jcunes Pseudosira ne deviennent jamais des Lepidocyrtinus en vieillissant et si j'etais certain que mes exemplaires soient bien des adultes. II y a naturellement une massue apicale a ant. IV et les antennes sont trcs pales. Les ecailles ovales ont I'ornementation habituelle et il y a naturellement des ecailles sur la face anterieure des dentes. Griffes du type habituel, appen- dice empodial aigu (non tronque) avec une ciliation longue et fine sur I'aile postero-interne. A titre d'indication, voici quelques rapports tires d'un seul exemplaire: A/B de 1.90 a 2.05; A/D: 1.30 a 1.35; A/D' : 1.10 a 1.15; A/e : 1.95 a 2.10; A/1 : 1.95 a 2.35; A/ae: 1.50
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a 1.60; A/E: 1.05 (p. I), 1.00 a 1.05 (p. II), 0.95 (p. Ill) ; ae3/m: 2.30; riia/m: 1.5 a 1.8. Mucro-dens nettenient plus long que le manubrium, region dentale annelce finissant brus- quemment, mucron en crochet simple, sans epine basale.
Justification. En systematique de Lcpidocyrtinus, nous en sommes reduits a comparer priiicipalemcnt des ornementations. Les deux formes qui me seml)lent les plus prochcs de la presente sont: Lcpidocyrtinus Schaffcri (Schott), forme de Nouvellc Guinee, de I'ile Selco et des Philippines, decrite par Sciiott (1901, Ten Fuz., p. 323, figs. 31-9) et revue par Handschin (1930, Philipp. Journ. sci., 42, p. 417, figs. 24-7). Cette espece est plus coloree que la presente, ant. IV y est annele (encore que les exemplaires d'll. n'aient qu'cnviron 2 mm.). Au sujet de I'appcndice empodial Sciiott ecrit : "Untere Klaue lancetteualmlich, nicht schrag abgestutzt, unbewaffnet." et Handschin : "Empodialanhang schmal, nach ausscn, schrag zugespitzt." C'est dire que les auteurs ne sont pas precisement d'accord et qu'il est bien probable que 1' on doive considcrer deux especes la ou Handschin n'en fait qu'une seule. Je puis affirmer que notre unifasciatus n'est pas celle de M. Handschin. Je serais moins affirmatif touchant celle de Sciiott, encore que Fornementation puisse seml>ler m'autoriser a I'etre.
La seconde espece a prendre en consideration est celle que Ritter (1910, Ann. k. k. naturh. Hofmus., 24, p. 391-4, figs. 44-6) a decrite sous le nom de Calistocyrtiis iitdicus. et dont BoNET (1930, Eos, 6, p. 260-1, fig. 5) fait, a juste titre, une Pseiidosira. Cette espece est plus coloree que la notre ; en particulier, la bande somljre transverse occupe le dos d'abd. Ill et II et il y a une bande transverse au bord posterieur d'abdomcn IV. Je crois pouvoir considerer L. unifasciatus comme tres voisin de L. indictis (Ritt.).
L'afiinite de L. unifasciatus avec deux formes appartenant a la faune dite orientale ne doit tromper personne car personne n'oserait dire la parcnte des Lcpidocyrtinus.
Lcpidocyrltis oricntalis E. Handschin Text-figures 31-33
Station: Ootacamund, under leaves and moss, 2195m. alt., environ, Nilgiri Hills, 6-XI-
1932; 1 exempl.
Description. Taille : 2.4 mm. Coloration pale, pratiquement sans pigment, alors que la fig. 17 de Handschin (1929, Rev. suisse Zool., 36 p. 242) en indique sur les coxae; ant. Ill et IV violet fonce, moitie apicale d'ant. II un pen coloree (concordance avec les donnees de 1 'A) ; ecailles du type Lepidocyrtus. Taches oculaires noires a 8 yeux dont deux plus petits; ant. II, III, IV subegaux et IV pounai d'une mas.sue apicale retractile (non notee par 1 'A). Rapport tronc/ant. : 2.15; rapp. ant./diagonale cephalique: 1.65 Mesonotum un peu debord- ant. Tibia III/I: 1.85. Ergots non etudiables. Griffes a 4 dents internes, dents latcrales l>ien ecartees et dent externe en pointe trcs detachee. Appendice empodial nettement du type tronque a p. I et II, mais, a p. Ill, la troncature est peu nette et il scmble bien que les deux lamelles externes se poursuivent jusqu'a I'apex, tres rapprochees mais non soudees. Aile postero-interne a denticulations presque indistinctes. Rapports: A/B : 3-3.4 (p. I), 2.85- 3.15 (p. II), 2.5-2.6 (p. HI) ; A/D: 1.75-1.90 (p. I), 1.70-1.75 (p. II), 1.55-1.60 (p. HI) ; A/D': 1.25-1.30 (p. I), 1.25 (p .H, HI); A/1: 1.0-1.1 (p. I), 1.1 (p. II), 1.2 (p. HI); A/e:0.9 (p. I, II), 0.95 (p. HI); A/ae: 1.7-1.8 (p. I), 1.4-1.5 (p. II), 1.5 (p. HI). Manu-
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brium subegal an mucro-dens ; pas de lobe particulier a la racine des dentes ; region dentale non annelee 2.05 fois le mucron. Mucron complet a dent apicale longue et epine basale. Rapport: ea3/mucron : 1.25.
Justification. La diagnose de L. oricntalis est forte courte. L'A. ne signale pas la massue apicale d'ant. IV et ne figure pas completement la griffe. Ma determination doit done comporter certaines reserves.
Figures 31-33. Lcpidocyrtus orientalis E. Handschin. 31, profil. Z2, gl(I), g2(II) et g3(III) (Obj. 90XOC.10). 33, mucron avec longueur de g3 et d'ae3 a la meme echelle (Obj. 60 X oc- 10).
L. orioitalis nie semble tres voisin de L. cacndeicornis Bonet (1930, Eos, 6, p. 262, fig. 6), des environs de Bombay. Mais I'A. dit formellement (p. 263) "appendice empodial lanceole, non tronque." Pour cette raison je ne puis songer a une synonymie. II y aurait, entre L. oricntalis et L. caeruleicornis les memes differences qu'entre nos L. ruber et laimgi- nosiis. II y a aussi, dans Ritter (1. c, p. 390) un L. scaber, de Peradenya, qui pourrait bien entrer en ligne de compte, s'il etait mieux connu.
Nota. M. Bonet inscrit parnii les Lcpidocyrtus son L. caeruleicornis, pourvu d'une massue apicale a ant. IV. Jusqu'a nouvel ordre, je suis I'exemple de M. Bonet.
Repartion geographique. Handschin indique, pour L. orientalis, plusieurs stations des Nilgiris, de 1600 m. a 2400 m. d'altitude. Autant qu'on puisse juger, L. orientalis presente de proches parents dans la faune orientale, inais il n'cst pas impossible qu'on lui en trouve d'aussi procbcs ailleurs (L. ruber par exemple).
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Paronella (Aphysa) longicornis (Oudemans) Text-figures 34-35
Station: Pykara, cnv. 2133 m. alt., on wet, eartliy cliff, 16-XI-1932; 4 exenipl.
Reniarques. En rapportant, avec quelque doute, mes exeinplaires a I'espece d'OuDE- MANS, je dois dire que j'adopte I'idee que M. Handsciiin (1925, Treubia, p. 261) se fait de ladite espece. En particulicr j'admets que "Typisch ist die Aniage eines kleinen Aussen- zahnes, sodass das Gebielde (Mucro) 7-zahnig wird." On sait que SciKirr ne signale pas cette petite dent.
Figures 34-35. Aphysa longicornis (Oud.).
34, deux types de coloration. 35, deux profils de mucrons en vue externe.
Je range mes exemplaires dans Ic genre (ou sous-genre) Aphysa ericore qu'ils present- ent, entre I'apex de la dens et la base du mucron, une petite saillie — sans dmite articulaire — pouvant rappeler I'appendice dental des Microphysa.
Paronella longicornis, selon Handsciiin (1930, Philippine j. of Sci., 42, p. 424) est largement repandue sur toute la region orientale (I'A. la note aux Philippines) ct (I. c, p. 421) fort pen variable, quant a la coloration, dans toute son aire.
II n'est peut-etre pas inadmissible que nos exemplaires des Nilgiri-hills apparticnncnt a cette espece si repandue. Cependant, afin d'eclairer ceux qui reprendront la question, je donnerai la description desdits exemplaires.
Trois exemplaires out de 3.2 a 3.8 mm., le quatricme a env. 5 mm. Les Text-figures 34-35 donnent une idee de leur coloration et j'ajoute que les tibias sont toujours violaces. Ant. I/II varie de 1.00 a 1.15; tibia I/II varie de 1.10 a 1.15; tibia III/I : de 1.05 a 1.25; mucro-dens/manubrium varie de 1.35 a 1.50. Voici, pour les trois premiers exemplaires les rapports couQernant les griffes: A/B : 3.8-4.1 (p. I), Z.2,-Z.7 (p. II), 2.6-3.0 (p. Ill); A/D: 1.5-1.6 (p. I), 1.4-1.5 (p. II), 1.4-1.5 (p. Ill) ; D' manque toujours, de meme: impos- sible de voir la dent externe (e) ; A/1: 1.2-1.4 (p. 1), 1.4-1.5 (p. II), 1.3-1.7 (p. Ill);
REPORT ON COLLEMBOLA 273
A/ae: 1.6-1.8 (p. I), 1.4-1.5 (p. II), 1.2-1.3 (p. Ill); appendice empodial tronque avec denticule et aile postero-interne crenelee; rapport ae/d (d etant la longueur de la tronca- ture): 1.8-2.1 (p. I), 1.9-2.0 (p. II), 1.9-2.1 (p. Ill); E/A: 1.1-1.2 (p. I), 1.2-1.3 (p. II), 1.1-1.4 (p. III). Les figures ci-jointes donnent una idee de la structure du niucron — tres comparable a celle qu'admet Handschin — et niontrent le lobe articulaire dento- nuicronal. II n'est pas impossible que cette formation soit homologue a la vesicule des Mkropliysa. Rapport mucron/ae3 : de 1.3-1.6.
Le grand exemplaire de 5 mm. doit sans doute etre traite a part — les coordonnees logarithmiques seraient pent etre indiquees ici, si j'avais un plus grand nombre de donnees ma disposition.— A/B : 3.0-3.2 (p. I), 3.3 (p. II), 2.5-2.8 (p. Ill); A/D : 1.4 (p. I, II, III) ; D'et e manquent; A/1 : 1.8(p. I), 1.6(p. II), 1.6-1.7(p. Ill) ; A/ae: 1.4-1. 5(p. I et II), 1.2(p. Ill); ae/d: 1.8(p. I, II), 1.9-2.0(p. Ill); E/A: 1.0-1.1 (p. I, II, III); mucro/ ae3: 1.1-1.2.
S'il ne s'agit pas — ce qu'on saura plus tard — exactement de I'espece d'OuDEMANS- Handsciiin, il s'agit a coup sur d'une forme tres voisine. En tout cas, on devait s'attendre a trouver, dans le materiel de la Yale North India expedition, des Apliysa et des Micro- physa. II s'agit de types caracteristiques de la faune orientale.
Salina celebensis (Schaffer) Te.xt-figures 36a, b, c
Stations. Ootacamund (Hill, south of.), 2378m. alt. environ, Rhododendron, 13-XI- 1932; 6 exempl. Ootacamund, dam above Dhobi's quarter, 2316 m. alt. env., 15-XI- 1932; 2 exempl.
Description. Tailles : 2.3-3.3 mm. En ce qui conqerne la coloration, les Text-figures 36a, b, c montreront les types notes. II in'est absolunient impossible de les separer de ceux que j'ai pu etudier d'Indo-Chine et dent j'ai dunne la description (1935, Bull. soc. entom. Fr., p. 138-42).
En ce qui conqerne les caracteres morphologiques, je ne vois aucune possibilite de dis- tinguer specifiquement ces exemplaires de ceux d'Indo-Chine. On comparera les chiffres suivants a ceux que je donne dans le travail ci-dessus indique.
(Nota) les chiffres suivis d'un "■' se rapportent a la forme dubiosa nov. (ut species) dont je parlerai plus loin.
Rapport A/Ba-p: 2^(3.0), 3(3.25), 1(3.50), 7(3.75), 9(4.00), 2(4.25), 5(4.50) pour p. I; 3(3.00), 1(3.25), 5(3.50), 5(3.75), 3(4.00) pour p. II; 1(2.75), 5(3.00) et 2°(id.), 9(3.25), 7(3.50), 2(3.75) pour p. III. Tableau tres semljJable au correspondant des formes indochinoises. Rapport A/D: 1(1.3), 2(1.35), 5(1.40), 3(1.45), 2(1.50) a p. I; 3(1.35), 4(1.40), 2(1.45) .\ . 1(1.55) a p. II; 1(1.30), 2(1.35), 7(1.40) et r(id.), 2(1.45) a p. Ill; meme conclusion que ci-dessus. Rapport A/D' : 2(1.15), 7(1.20), 3(1.25) a p. I; 4(1.15), 4(1.20), 2(1.25) a p. II; 1°(1.10), 2(1.15), 8(1.20) ; meme conclusion. Rapport A/e: 7(0.9) a p. I; 6(0.9) a p. II; 7(0.9) et l°(id.), 2(1.0) a p. Ill; mcmes conclusions. Rapport A/1: 1(0.9), 3(1.0), 6(1.1), 3(1.2) a p. I; 4(1.0), 7(1.1) a p. II; 1(0.9), 9(1.0) et l°(idem), 5(1.1), 1(1.2) a p. Ill; memes conclusions. Rapport A/ae: 1(1.4), 1(1.5), 3(1.6), 5(1.7) a p. I; 5(1.4), 3(1.5), 1(1.6) a p. II; 5(1.4) et ^(idem), 4(1.5) a p. Ill; memes conclusions, Rapport ae/d: 1(1.6), 2(1.7), 3(1.8), 4(1.9), 1(2.0)
274
REPORT ON COLLEMBOLA
... 1(2.3) a p. I; 3(1.8) . . . 5(2.0) . . . 2(2.6) a p. II; 1(2.0) . . . 2(2.2), 3(2.3), 2(2.4) . . . 1(2.6), 1(2.7) et l°(ideni) . . . 1(2.9) memes conclusions. Rapport E/A: 1 (1.4), 1(1.6), 3(1.7), 4(1.8), 3(1.9) a p. I; 7(1.8) 2(1.9) a p. II; 1(1.6) et P(idem), 1(1.7), 7(1.8),
Figures 36a, b, c. Trois types de coloration de Saliiia celebeitsis (Schaffer). Figure 37. Salina dubiosa n. sp., profil.
1(1.9), 1(2.0) a p. Ill; memes conclusions. Rapport nmcron/g3 : 4(0.95), 6(1.0), et l°(id.), 1(1.05), 2(1.10); encore que les maximum soient deportes vers la gauche du tableau, on ne saurait fonder une distinction specifique sur ce caractere. Rapport mucron/ap- pendice dental: 1(1.3), 2(1.4), 6(1.5), 3(1.6) ; ce qui rentre dans les cadres donnes pour les formes d'lndochine. Rapport Dens/manubrium : 3( 1.2), 3(1.3), 1(1.4) et l'(id.); meme conclusion.
REPORT ON COLLEMBOLA 275
En resume : ni I'etude du dessin ni celle des principaux caracteres morphologiques liabituellement utilises en determination ne permet de distinguer ces exemplaires de ceux d'Indo-Chine. C'est pourquoi je leur attribue le meme nom.
II n'est pas impossible que le Cremastocephalus indiciis de Imms (1912, p. 104-5, figs. 58-9) doive rentrer dans le groupe dont le chef de file serait I'espece de Schaffer. Le Cremastocephalus montanus Imms prescnte un rapport mucrou/'appendice dental voisin de 1.0. J'ai trouve ce rapport chez 5. celebensis d'Indo-chine.
Salina dubiosa nov. sp.
Text-figure 37
Station : meme localite que les precedents et recolte en meme temps que les exemplaires du 15-XI-1932; 1 exemplaire.
Description. Caracteres morphologiques compris au tableau precedent ; memes aspects des griffes et du mucron. Coloration comme I'indique la Text-figure Z7.
Justification. Le type de coloration differe de tons ceux dont j'ai pu me procurer la figure et notamment de tons ceux que figure Handschin (1928, Treubia, 10) dans sa revision du genre. C'est done uniquement faute d'intermediaries entre les formes pales, les formes a bandes ou series de taches longitudinales et ce type a bande transverse, que jene puis faire autement que de donner un nom au present exemplaire.
Sminthurides aquaticus (Bourlet) Text-figures 38-41
Station: Kyani. Small spring surface at 35.5°, 4763m. alt. env., 21-VII-1932; qq. exempl.
Justification. Je ne donne cette determination que sous la reserve suivante : il s'agit d'une forme qui peut rentrer dans la comprehension actuelle de Sminthurides aqtiaticiis (Bourl.). II est possible que, quand cette derniere sera mieux connue, Ton doive renommer cette forme du Thibet. Afin de justifier ma determination — en tant que premiere approxi- mation— je crois devoir donner les Text-figures 38-41 qui serviront de points de comparaison.
La meilleure description que nous possedions de 5". aquaticus est celle de Borner (1901. Abh. naturwiss. Ver. Bremen, 17, p. 96 & seq.). En comparant mes figures a celles de Burner, on trouvera sans doute de minimes differences, notamment sur I'antenne $ . Cependant on ne saurait se defendre d'y trouver de grandes similitudes. De telles struc- tures n'ont pas ete etudiees systematiquement chez 5". aquaticus. II convient done d'attendre qu'on sache leur valeur taxonomique. Pour le moment, afin de ne pas encombrer la nomen- clature, je nomme mes exemplaires de Kyam du nom de S. aquaticus.
Repartition geographique. Que I'avenir demontre I'identite de mes exemplaires avec I'espece d'Europe ou bien qu'il me contraigne a etablir un noveau nom, il restera toujours ceci : on aura a faire a des formes excessivement voisines et on ne pourra se defendre de leur attribucr une origine commune.
.V. aquaticus (Bourl.) existe certainenient en Europe et on ne saurait douter que Folsom (1928, Mem. 101, Cornell. Univ. agr. exp. sta., p. 16) n'ait parfaitement raison quand il
276
RF.rORT ON COI.I.EMBOI.A
admet spn identite avec la forme americaine S. amicus Fols. D'autre part, Handschin (1928, Arch. Natiirg., 92, p. 8) le note d'Algerie. Womersley (1932, Cone. sci. ind. research, painphl. n°34, p. 17) le signale d'Australie. Cependant je n'ose tenir pour assuree cette determination (en particulier la fig. 4d de 1 'A., representant un nuKron, ne semble
Figures 38-41. Smmthurides aquaticus (Bourlet). 38, ant. II et III du $. 39, g2(II) et g3(III). 40, chetotaxie dentale en vue anterieure, cote medial a droite. 41, niucron, profil en vue e.\terne.
pas s'accorder avec ce qu'on sait de 5". aquaticus). Je crois done, jusqu'a plus ample informe, laisser dans le doute la presence de ^. aquaticus en Australie.
Dans I'etat actuel de nos connais.'^ances, on pent considerer S. aquaticus comme une forme paiearctique et nearctique. Sa presence dans le mediterraneen doit etre affirmee par des stations plus nombreuses que celles qu'on possede actuellement. La presence de 5". aqua- ticus au Thibet (ou d'une forme tres voisine) indique, sans doute, une affinite paiearctique de la faune thil)etaine.
Sminthurides (Stenacidia) violaceus (Renter) Text-figures 42-48
Station: Bao, East of Shakya la. surface of pool, 4616 m. alt. environ, 25-VII-1932; 1 exemplaire.
Justification. Je fais, pour cette determination, les memes reserves que pour la pre- cedente et, pour la justifier, dans le mesure du possible, je dunne les figures ci-jointes ainsi
REPORT ON Cnr.I.EMBOLA
277
que la description suivante : 2 : taille : 0.6 mm., violet pale. Revetemcnt assez dense, fait de soies assez longues et greles; pas de soies en epine sur la tete; yeux habituels sur taches noires. Antennes : cf. Text-figure 42 ; ant. IV avec tres faible ebauche d'annulation. Griffes a dent interne tres nette a p. I et II et nulle a p. III. Je ne trouve rien de comparable a une "tunica" — Stach (1919-21, Mull. acad. polon. .sci. & lett., p. 197) en indique une, con-
FlGURES 42^8. Sminthuridcs (Stcnacidia) vlolacciis (Reuter). 42, profil de I'antenne (on ne saurait considerer ant. IV comme subsegmente). 43, gl et 44, g3 avec, en dessus, I'organe tibio-tarsal a plus fort grossissenient et le tenaculum. 45, dens et mucrons en vue posterieure. 46, chetotaxie dentale en vue anterieure, cote medial a gauche. 47, mucron. 48, profil du petit segment abdominal 9 .
trairement a la plupart des auteurs. Dents laterales tres nettes; appendice empodial a sole subapicale et plus large a p. Ill qu'aux autres paires. Organe tibio-tarsal fait de deux capuchons a pointe courbe et d'une sole elargie, assez courte (Text-figures 43-44). Get organe est tout a fait con forme aux figures donnees par Linnaniemi. Tenaculum avec trois soies au pars anterior. (Linnaniemi en indique 4.) Mucron et chaetotaxie dentale: cf. Text-figures 45, 46, 47. Mucron conforme a la description de Stach (1. c).
Repartition geographique. II s'agit d'une forme europeenne — d'ailleurs rare. Je ne pense pas trop aventurer en ecrivant que le S. (St.) violacea de Womersley (1. c, p. 14-6, figs. 2b-g) n'est pas I'espece de Reuter et n'ai pas scnipule de ne pas etendre a I'Australie la repartition geographique de cette derniere. La presence de S. z'iolaceus au Thiljet indique une affinite palearctique.
278
RErORT ON COLLEMBOLA
^ Bourletiella arvalis (Fitch)
Text-figures 49-59
Station: Edge of Phashakuri near Pampur, env. 1585 ni. alt., 7-V-1932; 3 exempl.
Description. Le plus grand exemplaire a 1.2 mm. (S), le plus petit est encore un jeune dont on ne pent determiner le sexe. Ses antennes ont leur IV article indistinctement segmente. Les figures ci jointes tiendront lieu de description et justifieront la determination que j'adniets ici.
Figures 49-55. Bourletiella arvalis (Fitch). 49, 9. 50, soies anterieures de la tete. 51, profit d'ant. IV. 52, p. et g: soies du petit et du grand abdomen comparecs a la longueur du mucron, menic ccliclle que fig. 50. 53, tenaculum. 54, mucron en vue externe. 55, abd. VI en vuc postcrieure ( 9 ) .
11 faut (lire que j'admets la systematique proposee par Folsom (1934, Iowa st. coll. sci., 8, p. 466, figs. 52-58) selon laquelle B. an>alis (Fitch, 1863) est synonyme de B. lutca (LublxDck, 1868) et de la plupart des auteurs curopeens. Je renvoie done a la synonymic donnee par M. Folsom. Cependant, je n'admets pas la synonymic avec la forme nominee an'alis par Womersley, 1932 (Pap. r. soc. Tasmania, 193, p. 9, PI. I, fig. 5 et text-fig. IV, 1-5), principalement a cause de la forme des appendices anales 9 . Par suite, je dois rejeter la determination du meme auteur (1932, Cone. sci. ind. res., n°43, p. 26-7, fig. 8a). Ces formes d'Australie ne sont certaineinent pas les memes que celles du palearctique et du nearctique.
REPORT ON COLLEMBOLA
279
Repartition geographique. Dans I'etat actuel de nos connaissances — il n'est pas dit que I'etude approfondie de B. an'alis (au sens acluel) n'apportera pas des surprises analogues
Figures S6-58. B. arvalis (Fitch). 56, abd. V et VI en profil (2). 57, a gauche: appendix anahs 9 en vue oblique et, a droite: le meme a peu pres de face, striation exageree a gauche, non figuree a droite. 58, chetotaxie dentale en vue anterieure.
Figure 59. B. anatis (Fitch) ; p. I et p. III.
a celles dont j'ai pu fairc soup(;onner I'existence a propos de Bourlcticlla hortensis (Fitch) (Denis, 1931, Bol. soc. entom. Espan., p. 85-9, fig. 10-3) — il n'est pas illicite de penser que la presence de B. arvalis au Kashmir denote une affinitc faunistique vers I'Europe et rAmerique du Nord.
280 REPORT ON COLLEMBOLA
Addendum
II existe encore, dans Ic materiel qui m'a etc confie;
1°) un excmplaire d'un genre indeterminable — la furca maiiquant — de la station L 20, Ravine above Himis, 3360 m. alt.
2°) au moins deux especes A'Entonwhrya.
T.'une provient du Mt. Pamzal, Ladak., par 5240 m. d'altitude et resscnible bcaucoup a notre Eiitoiiiobrya lanuginosa (Nic.).
L'autre, de la station L 20, prescnte une ornenientation rappelant celic de notre R. dis- jiiitcta (Nic.), mais sans serie mediane de taches.
Je ne puis prendre la responsabilitc de ces determinations, n'ayant a ma disposition qu'un seul exeinplaire des deux Entoiiiobrya en question, mais je crois devoir signaler leur presence a reux (jui reprendront nion travail quand la systematique des Entomobrya europcens sera plus assurce qu'elle ne Test actuellement.
\'uc d'ensemble sur le materiel recolte par la YNIE. 17 especes ont pu ctre etudiees. On en drcssera le tableau suivant
I .'^ffinites N
Noms Altitudes Palearctiqucs Orientates Douteuses
Hypogastrura aniiafa (Nic.) 1585 x
H. couumtnis (Fols.) 1580 x
Friesea excelsa n. sp 5500 x
Proisotoma ladaki n. sp 4753 x
Isotoma spinicauda Bonet 5300 ?
OrclicscUidcs Boraoi Bon 5200 ?
Sira brahamides n. sp 4400 x
S. nilgiri n. sp 2378 (Nilgiri Hills) x
Parasira subornata n. sp 1585-4664 x
Lepidocyrtinus unifasciatus n. sp 1 585 ?
Lcpidocyrtus oricntalis H 2195 (Nilgiri 1 1 ills ) x
Aphysa longicornis (Oud.) 2133 (Nilgiri Hills) x
.S-,///;;rt cclcbcnsis (Scliaff.) 2316-78 (Nilgiri Hills) x
S. dubiosa n. sp 2316 (Nilgiri Hills) x
SmintJiiirides aqiialiciis (Bourl.) 4763 x
S. (St.) znolaceus (Renter) 4616 x
Bourletiella. arvalis (Fitch) 1585 x
Je ne crois [jas devoir epiloguer sur la (piestiun ties altitudes, le nunibre des stations etant notoirement insuffisant. II ne faut jjas oublier qu' Hypogastrura armata est pratiquement cosmopolite et que les Siniitlhuridcs snnt difficiles a capturer. II n'est peut-etre pas interdit de remarqucr (|ue les formes veritablenicnt orientales {.Ipliysa et Salinn) provienncnt d'alti- tudes rclativenient faibles et deja Lmms (1912) avait fait une constatation analogue. Mais, a mon sens, cette sim[)le remarque ne saurait sufifire ])our etablir quelque conclusion fernie.
Deux auteurs ont etudie des regions analogues a celles qui ont occu])e la '\'NI1£. Les
REPORT ON COLLEMBOLA 281
resultats de run d'eux: M. Bonet (1930, 1. c.) sont fort interessants en ce qui conqerne les constituants palearctiques de la faunule du Waziristan. Voici la liste de M. Bonet:
Isotoma spinicauda Bonet
Orchescllides Boraoi Bonet
Drepanura falcifcra Bonet
Paranura ornata Bonet
On ne manquera pas de souligner la grande similitude de la liste de Bonet et de la precedente.
L'autre auteur: Imms (1912 I.e.) admet, dans sa liste, 5 formes palearctiques et ecrit qu'elles proviennent d'au dessus de la limite des forets de I'Hymalaya. Ce sont : Isotoma siva Imms (Garhwal, 10,300 ft.) Tomoccrus vulgaris Tullb. (id.) Entomobrya crassa Imms (id. 12,500 ft.) Sinella montana Imms (id. 10,300 ft.) Seira frigida Imms (id.)
Mais les biogeographes feront bien de ne pas accepter sans controle le contenu de cette
liste, car la systematique de 1 'A. appelle les remarques suivantes.
1°, Seira frigida Imms est une Parasira.
2°, L'A. donne avec un certain doute sa determination de Tomoccrus vulgaris Tulb. II est vrai que les Tomoccrus sont pratiquement inconnus dans la region orientale et qu'on est en droit de les considerer comme palearctiques. Je ferai noter qu'il y a un Tomoccrus en Indo-Chine (encore inedit). Cependant il reste fort probable que le Tomoccrus de Imms denote des afifinites palearctiques de la faune des Garhwal.
3°, Isotoma sii'a Imms n'a certainement rien a voir avec le genre Isotoma ni meme avec la super famille des Isotomiens. Elle n'en a aucun caractere et I'A. ecrit lui meme que le quatrieme segment de I'abdomen est "triple longius quam tertium." Les descriptions et figures de I'A. ne permettent pas de savoir ce qu'est /. si-va. Pent ctrc un tres ordi- naire Entomobrya? Pent etre quelque chose de beaucoup plus interessant pour le systematicien?
4°, Entomobrya crassa Imms presente un "segmentuni abdominis tertium quartum longi- tudine fere aequans" et, selon I'A. ressemble assez a une OrchcscUa. II ne s'agit cer- tainement pas d'un Entomobrya, mais ni la description ni les figures (fig. 30-1) ne peuvent permettre une supposition soutenable. On peut soupconner que Imms est passe a cote de quelque Orchescllides.
En resume, on fera prudemincnt de s'en tenir a la liste suivante : Un Tomoccrus ^^ni-eirt voisin de vulgaris Tullb. Parasira frigida (Imms) Sinella montana Imms
Encore n'est-il pas prouve que la Sinella denote des affinites palearcticjues.
II me reste a attirer I'attention sur les Parasira dont I'aire de repartition va de la Meso- potamie au Japon et sur 1' Hypogastnira conimiaiis (Fols.) qu'on trouve au Japon, en Chine et en Indo-Chine. J'ose esperer qu'on ne tirera pas, de ces repartitions, de conclusion exagerement generale.
282
REPORT ON COI.LKMBOLA
^■lildciidtoii n"2 (September 21, 1935)
Proisotoma ladaki in. — Station: Togarnia Tso, lO-VlI-1932; 7 exemplaircs confornies a
ceux dccrits plus hant. OrchescUidcs Boraoi Bonet. — Station: Ororotse-Tso, 5513 m. alt., Indian W. Tibet, 11-
VII-1932; 2 exemplaires a bandes transverses bien nettes et presqiic unifornies. Salitia cclcbcnsis (Schaff.). — Station; Ootacanuind, S. of town by I)eating Rbododendron,
ll-XI-1932; 4exempl.
Entomobrya Hutchinsoni n. sp. Text-figures 60, MI I Station: Shakya La, Indian Tiiiot, E. side, talus slope, 5440 ni. alt., 25 -VI-1932; 1 ex.
Description. Taille 1 mm. 34 cnv., Jaunatre, sans pigment sur le corps; yeux noirs, unc ombre tient lieu de bande frontale; ant. II un pcu teinte de gris a I'apex, ant. Ill bien tcintc, ant. IV idem, mais plus clair; pattes et furca pfdcs. Rapport tronc/antenncs : 1.40; ant. II, III et IV subcgaux; Ics tubules de I'org. antennaire III sont particulierement volumineux; apex d'ant. IV a massue retractile comme chez la plupart des especes du genre. Revetement
Figure 60. Entomobrya Hutchinsoni n. sp. I, p. I, a cote: la longueur dc Tergot. II, p. III. Ill, uuicruii.
Tout (lu memc excmplaire au meme grossisscment.
general trcs abimc mais certaincmcnt sans ccailles. Griffcs trcs particulicres: dent externc probablement presente, dents latcralcs tres distales (Text-figure 60, I-II), une paire de dents internes dans la moitie distale de la griffe dont I'apex est un peu courljc; impossil)le de dccouvrir les dents impaires. Appendice empodial du type acutus allonge; impossil)le d'y decouvrir des crenelures. Ergot beaucoup plus long que la crete interne de la griffe. Rap- port Manubrium/mucro-dens : 0.95 ; mucron (Text-figure 60, III) a dent anteapicale trcs nettement plus faible que I'apicale; pas trace d'epine basale.
Justification. Les Entomobrya — j'entends ceux qui sont suffisamment connus — a mucron sans epine basale sont rares, de plus, je ne connais pas d'Enloinobrya dont la griffe presente les caractcres que je viens de dire. II est done infiniment probable que la nouvelle espcce que je propose — d'apres un seul exemplaire — sera facile a reconnaitrc. Je nc saurais lui trouver de proche parentc parmi toutcs celles qui sont suffisamment connues, encore que la recente tentative de revision du genre par M. Bonet (1934, Eos, 9, p. 152 & seq.) facilite grandement la recherche. Les quelques especes dont M. Bonet dit ne pouvoir tenir compte, ne sauraient non plus etre comparees ;i E. Hutchinsoni.
ARTICLE XVI
REPORT ON CLADOCERA
By Dr. V. Brehm
Wnii 8 Text-Figures
(Received August 16, 1935)
Die Yale North India Expedition untersuchte in crster Linie die Hochgebirge von Kaschmir und Ladak, doch warden audi in Punjab und in den Nilgiri Hills Aufsammlun- gen gewonnen.
Bei dem kosniopolitischen Charakter der Cladoceren konzentriert sich aber das Interesse auf die im Plochgebirge vorgenommencn Untersuchungen, die okologisch bemerkenswerte Resultate erwarten liessen. Doch erwies sich audi die Untersuchung der nicht im Hoch- gebirge gewonnenen Proben als dankenswert, da sie zur Auffindung einer neuen Moina fiihrte, von der spater die Rede sein wird.
Zunachst sei an der Hand einer Tabelle das Augenmerk auf einen Vergleich mit der Cladocerenfauna bcnachbarter Gebiete gelenkt. Als solche komnien Pamir und das angren- zende Tibet in Betracht, woriiber folgende Berichte vorliegen :
Daday E. v. : Entomostraca et Hydrachnidae e Tibet. Erschienen in Report on a collection
of aquatic animals made in Tibet by Caj^tain F. H. Stewart, I. M. S., during the year
1907. Werestschagin, G. : Notiz iiber die Siisswasserfauna des Pamirs. Deutsches Resume
einer russischen Arbeit, deren Erscheinungsort aus dem Separatum nicht zu entnehmen
ist. Rylov, M. : Cladocera et Copepoda in Abhandlungen der Pamir — Expedition 1928. H.
Zoologie. 1930. Sars, G. O. : On the Crustacean Fauna of Central Asia. Part II. Cladocera. Annuaire
du Musee Zoologique de I'Academie Imperiale des Sciences de St. Petersbourg. T.
VIII. 1903. Sars, G. O. : Local Faunae of Central Asia. Ibidem.
In der nebenstehenden Tabelle sind in der ersten Spalte niit R die von Rylov gefundenen Arten bezeichnt, die von Werestschagin gefundenen mit W, in der zweiten mit S die von Sars angegel^enen, in der dritten die von Daday gefundenen mit D, wahrend die letzte Kolunme die von der Yale — Expedition mitgebrachten Arten enthiilt, wobei die Signatur des Fundortes angegeben ist.
Vergleichen wir die hier gegeniibergestellten Artenlisten, so sehen wir im allgemeinen eine weitgehende Ubereinstimmung. Wir begegnen unter den Yale-Cladoceren nur sechs, die von den Untersuchern der Nachbargebiete nicht namhaft gemacht werden : Sida spec, Drep- anomacrotlirix sp., Alonclla cxigiia Leptodora hyalitia, Camptocercus rectirostris und Polyphemus pediculus.
Mem. Conn, .'^cad., Vol. X, Art. XVI, June, 1936.
284 REPORT ON CLADOCERA
Moina esau sclieidet als Vergleichsobjckt aus, da sie deiii Nilgiri-Clel)irg:e anqehort. Drcl'iiiioiiuicrothnx alx^r ist von IjesoiukTcni Intcrcssc, da dicse l)isht'r als kaukasisilier ImuIc- mismus gegolten hat. Es scheint sicli da urn cine ausgesprochene Hochgebirgsforni zu Iiaii- deln, da ihre Fundstcllc ini Kaukasus bei rund 3000 ni. liegt uiid die von der \'ale Expedition n)itgcl)racliten I-'.xeniplare gar aus 5217 ni. Sccholic staninicn. Dass .lloiuila cxigua in den Nachbargebieten nicht gefunden wurde, mag lx"i dieser kk-inen, weit verbreitetcn /\rt wohl nur einem Zufall zuzuschreiben scin nnd sie wind kiinftig wohl noch dort gefunden werden. Dass Lcptodora von der Yale F.xpedition gefunden wurde, verdient Reachtung, da diese Form wohl mehr tiefer gelcgene, wiirniere Gewiisser bewohnt, wenn man nach den Verbrei- tungsverhilltnissen in den eur()]);iischen Alpen schliessen darf. Sie felilt daher wohl in alien oben zitierten Arl)eitcn. Abcr ihr Vorkommen in 1548 ni. Seehohe, wie es von der ^'ale Expedition gefunden wurde, steht nicht ohne Beispiel da, weil Sars die Art — allerdings ohne Ilohenangabe — aus dein Altai zitiert nnd Rylov aus dem Kaukasus, wo sie sogar noch in 2080 m. Ilcihe angetroffen wurde. Merkwiirdig ist schliesslich, dass in keiner der Ver- gleichsarbeiten Rosniinen erwJihnt werden, die ja auch im Kaukasus vollig zu fehlen scheinen, wiihrend sie (lurch die Art B. longirostris in Kaschinir von vier Fundstellen ver- treten sind.
Haben wir damit einige Formen namhaft gemacht, die in den benachbarten I lochge- birgsseen fehlen oder zum mindesten bisher nicht gefunden wurden, so sei andererseits auch bemerkt, dass im Yale-Material die aus dem Pamirgebiet lieschrieljene Gattung Cor- niicUa nicht gefunden werden konnte.
In den Travaux de la Station Biologique du Caucase du Nord (Vol. III. — I'KW) veroffentlichte K. Decksbach einc Arbeit: ,,Zur Cladoceren fauna von Kaukasus imd Nord- Persien." In dicser Arl)eit werden als Kaukasus-Endemismen crwahnt : Ih-cpunoiuacro- thrix Stschelkamnvaewi, Ceriflddphma reticulata vnr. dubia inid Alona ariilrata. in negativer Hinsicht wird abgesehen von dem bereits crwahnten Mangel der Hosminen noch auf das Fehlen der Gattungen Holopcdiimi und Bytlwt replies aufinerksam gemacht, die in den Gebirgen Nord und Mitteleuropas verbreitet sind, wiihrend die Gattung Pnlyf^heiiiiis dem Kaukasus ebenso zukommt, wie den genannten europiiischcn (iebirgen. Den innerasiatischen Gebirgen .scheinen nun diese Gattungen mit Ausnahme des Polypliciiitis pcdiculus zu fehlen, da sie auch von der Yale Expedition nicht erbeutet wurden.
Njichst der geographischen Verbreitung interessiert an dem vorliegenden Material besonders die vertikale \'erbreitnng der einzelnen Arfen, die mm in einzclncn h'iillen den analogen Angalx;n frithercr Reobachter gegeniibergestellt werden mugen. Es sei aber vorher darauf aufmerksam gemacht, dass die Angal)e der Seehohe des Fundortes noch kein geniigender Anhaltspunkt zur Beurteilung der cikologischen Verhaltnisse ist. Ich verwei.se diesgeziiglich auf die Mitteilungen die F. H. Stewart der ()1>en zitierten Arbeit von Daday iil)er die X'erhaltnisse des in 14,700 Fuss Hfihe gelegenen Gyan Tse vorausschickt. Es heisst da: ,,b"rom .\pril to October this lake is entirely free from ice. During the summer no climate could be more delightful. In the daytime the temperature rises to aljout that of an I'.nglish summer's day and even at night remains moderate, so ringed in is the jilain by l)are hills which store up the sun's heat."
Immerhin geht aus den I'bercinstimmungen der von verschiedenen Autoren mitge- teilten Tabellen zur Geniige liervor, dass gewisse Arten in ansserordentlich unwirtliche
REPORT ON CLADOCERA 285
Regionen (.'inpurzust.-iyen vcninincn uiid dass andere yeradezu als Charakterformen sehr hoch gelegener Gewiisser zu gelten haben.
Merkwiirdig sind die Befunde hinsiclidicli dcr ]'"ort])flaiizungsverli;iltnissc, wenn wir sie den im Paiiiirgehiet gemachtcn licohaclitungen gegeiiuherstclkn : W'erestschagin scliliesst seine Arbeit niit (k'n Worten: ,,iii den I inchgebirgsseen des Pamirs sind die Cladoceren wahrscbeiniicb nionoeyclisch nnd die Zeit der parthenogenetisehen Fortpflanzung ist sehr rednziert, so dass es fiir einige Arten wahrscheinlich ist, dass die sexuelle Fortpflanznng in der ersten Generation nach dem lirscheinen ans den Ephippien eintritt." Dieser Anffas- sung pflichtet anch Kylnv 1)ei, der in seiner oben zitierten Arbeit sagt : ,,Unsere Befnnde bestatigen dnrcliaus die Angaben Werestscliagins. Das Zuriicktreten der parthenogeneti- sehen Verniehrung der Pamircladoceren nnd cHe vijrwiegende Bedeutung der Verniehrung auf sexuellem Wege wird durch folgende Talielle illustriert" und hier schaltet Rylov eine Tabelle ein, ans (k'r hervorgeht, dass von 14 Arten 7 im miinnbchen Geschlecht vertreten warcn nnd von 6 Arten l^phippialweibchen vorlagen.
Demgegeniiber mnss fiir das Yale-Material festgestellt werden, dass iiberwiegend par- thenogenetische Exemplare angetroffen wnrden. Es erinnert dieser Fall an eincn iihnlichen, der die Cladoceren f anna Spitzl>ergens lx;trilift. Anch fiir diese, wie iiljerhaupt fiir die ark- tische Cladoceren f anna, wnrde lange Zeit das Fehlen bzw. Zuriicktreten der Parthenogenese behauptet, bis es Olofsson durch langere, personlich an Ort und Stelle vorgenommene Unter- suchungen nachzuweisen gelang, dass anch auf Spitzbergen Parthenogenese zu den regel- mjissig auftretenden Erscheinungen gehort. Wenn nicht iiber einen liingeren Zeitraum sich erstreckende FJinge vorliegen, kann es leicht passieren, dass die bisexuelle Periode der Beo- bachtung entgeht. Bei der Untersuchung der in 4000 m. Seehohe am Mount Elgon lel^enden Cladoceren (Cladocera und Phyllopoda in ,, Mission scientifique de I'Omo. — Tom. II. Fasc. 8. — Paris, ]f'35) konnte ich bei keiner Art das Mannchen nnd niu' bei zweien E])hippial\veib- chen beobachten. W'aren die Proben 14 Tage spiiter entnommcn worden, hiitte das Bild jedesfalls ganz anders aussehen kdnnen. Jedesfalls — um auf das Yale-Material zuriick- zukommen — sprechen alle hier gemachten Beobachtungen gegen die Annahme, dass aus den iiberwinternden Ephippien sofort wieder eine bisexuelle Generation ausschliipfen wiirde.
Audi Daday erwiihnt fiir den Rham Tso das Vorkommen eines einzigen MJinnchens einer einzigen Cladocere, namlich \-on Moina rcctiroslris.
DIE UNTERSUCHTEN LOKALITATEN, DEREN HOHENLAGE UND FAUNA
I.
Puiijah and ad jnci-ul pari of I lie Xortli-ll'rsI Frontier Province
P 2 Sohawa (.528 m.): P 2 (4) Sinioccplialiis rrlnla mit rundeni Augenflcck. P2(7)
Pleiiroxus aduncits, Sinioccplialus vctuloidcs und .llona cf. btikobcnsis. P3(2) Sohawa: Siiiioccphalus zrtuloidcs. P 6 Sohawa : Sivwccplialus vctida mit rundem Augen fleck. P 13 Haripur: Macrotliri.v Iiirsuticornis.
285 REPORT ON CLADOCERA
II.
Stations in Kashmir
K21 Lokut Dal Lake (1582 m.): Campfoccrcns rectirostris, Bosniina longirostris. K21a: Polxphcimis pcdiadns, SimoccphaJus vetula, Scapholcbcris Kingi, Bosinina longi- rostris, Diaphanosoma hrachyunim, Acropcrus liarpac, Graptolcberis tcstudinaria, AloncUa c.vigua, Chydorus sphaericus.
K 8 Gagirbal: Srinagar. Open swamp. 1580 m.: Brudistiickc eiiier Sididc. Bosinina longirostris, Acropcrus Iiarpac, Graptolehcris tcstudinaria, .llonclla c.vigua.
K 19 Gagirbal: Srinagar. Closed swamp. 1580 m.: Daphnia pulcx, mit EY>h'\\i[)m\, Sca- pliolcberis Kingi, Ccriodaphnia quadrangula, AloncUa cxcisa und nana.
K 26 Sundar Khun (1582m.): Einige junge nicht n.'iher bestimmbarc I''xcniplarc von Diaphanosoma und Ccriodaphnia.
K31 Pampur (1585m.): Moina rectirostris.
K 24 Nishat Bagh, pond (1585m.): Alonella cxcisa.
K 33 Phashakuri (1585m.): Daphnia magna.
K 34 Ebenda : Ccriodaphnia sp. Acropcrus harpac, Graptolcberis tcsludiiuvia, Duiihc- vedia crassa var. interrupta.
K 34 Ebenda: Simoccphahis zrfula, mit rundem Augenficck, Ccriodaphnia qtiadrangula, in Menge, vereinzelt die Formen der vorigen Probe.
K 46 Bakh Hajan, Jhil (1575 m.) : Simoccplialus z'ctula mit rundem Augenfleck, Ccrio- daphnia quadrangtda.
K48 Manasbal Lake (1584m.): Lcptodora Iiyalina.
K51 Bod Dal Lake: Bosmina longirostris, Sida cristallina, Diaphanosoma juw Scapholc- bcris Kingi, Acropcrus harpae, Ccriodaphnia pulchella, Graptolcberis tcstudinaria, Alonella cxcisa. Ccriodaphnia und Acropcrus dominierend.
K 54 y\nchar Lake (1580m.): Polyphemus pcdiculus.
K 68 Pond, Sonamarg (2620 m.): Moina rectirostris.
K 69 Ebenda : Moina rectirostris.
III. Stations in Indian Tibet (Ladak and Rupshu)
L 1 Shargola (3050 m.): Chydorus sphaericus.
L 13 Leh. Pool (3506 m.): Chydorus sphaericus.
L 14 Spitok (3270 m.) : Simocephalus vctida, Chydorus cf. sphaericus.
L 16 Spitok. Small muddy hole: Simocephalus juv., Alona rectangula, Chydorus sphaericus.
L 18 Gulam Bagh, Chu.shod, pools in swamp (3230 m.): Simoccplialus z'Ctuloidcs, Chy- dorus sphaericus.
L 47 Lung-yun (4977 m.): Chydorus sphaericus.
L 50 Togarma Tso (5217 m.): Drei Proben mit Daphniden, in Probe L 50 (2) aus.ser- dem Drepanomacrothrix und Alona guttata.
L 60 Kyam, largest pool (um 5000 m.) : Chydorus sphaericus.
L61 Ebenda: Chydonis sphaericus.
Diaphanoioina brachyurum
Sida crystallina
Daphnia pulcx*
" magna
" pamirensis . . . .
" longispina
Daphniopsis tihetana* . .
Scapholcbcris aurita ... .
" mucronala
" Kingi . . . .
Ccriodaphnia quadrangida
" reticulata
" pulchcUa
Simocephalus vctida . . .
" " init ruiidem Augeii-
fleck
" vctnloidcs Moiiia rcclirostris
" dubia
" esau
Bosinina longiroslris
Macrothri.x rcctiroslris
" hirsuticornis
" laticornis . . .
Drepanomacrolhri.t sp. . .
Camptocercus rectirostris
Acroperus harpae
Euryccrcus lamcUalns Euryalona aimaiulalci Alona afflnis
" quadrangularis
" ladacensis ....
" coslala
" cf. bukcbcnsis .
" guttata
rectangula
Rhynchotalona falcata . . . Graptolchcris tc.ittidinaria
CornncUa pamirensis . . .
Pleuro.vits cf. aduncus ..
" cf. irigoncHus
Alonella c.veisa
nana .. e.xigua
" karua Dunhcvedia crassa
Chydorus sphacricus
var. interrupta
Polyphemus pcdiculus Leptodora hyatina . . . .
Friihere Beobachter
Pamir
R Rylov.
W Werest-
schagin
R,W K,W R,W
R,W
R
W
W
R.W
W
R,W
\v
w
"\v"
R
"r"
w
R R, W
W R
R,W
Iiinera- siens Sars
s s
Tibet Daday
D
D D
D
D
D
D D
Yale North India Expedition
Pendschab Kashmir
Ladak
P2,3
P2(4),6 P3(2)
K 21a. 26,
51
K.51
Kl'J
K3.3
P13
P2(7)
P2(7)
P2(7)
Kl'J, 21a,
51 K 19, 34,
46
K51 K21a
K 34, 46
K31,68 69
K8,21,
21a, 51
K31,68
69
K21 K8, 21a,
34,51
K8.21a, 34, 51
K 19, 24.
51
K19
K 8, 21a
K34 K8,21
K 21a, 54 K48
L80
L73
L 14, 16, 71,73
L71a L50(2^
L82
L 50, 71a,
73,82
L 16. 82,
82a
r. 1.13, 14, 16, 18, 47, 60,61, 71,72
Nilgiri
NIS NIS
L 18, 73 (Mysore)
N5,6 N14 N19
NS
NIS
N5, IS
N ?. 6, 13, 15
* The species of the geiuis Daphnia arc being studied by Professor R. Woltercclc and consequently are not treated in the present contribution. — G. E. Hutchinson.
288
REPORT ON CLADOCF.RA
L71a Tso Nyak (4241m.): Siinoccplialus vctula, Ccriodaphnia juv., Macrothrix laticor- nis, Alona guttata mit starker Langsstrcifunq' und viel Cliydonts sphacriciis.
L 72 Chushol, Large pond: Daphnien.
L 72a Small pool by Pond: Chushol (4491 m.) : Cliydonis spliacriciis.
L 73 Chushol Pond: Viel Ccriodoplniia ijiiaih-diigiilti. i.-'nuv;c Siiiioccplialu.'! ^'ctitloiilcs und stark gestreifte Alona guttata.
L76 Mitpal Tso (4875 m.) : Daphnia.
L 80 Khyagar Tso (4672 m.): Daphniopsis tilntana.
L 82 h'.stuary, Tso Moriri (4.^2''' in. i : . //lUd/ i^a/Zi;/,/, . I. /•(■(7(;;(.i,';(/i; und Alona ladacensis.
IV.
Nilgbi mils
N 5 Ootacamund Lake (2201 m.) : Moina dubia dominiercnd CItydurus cf. spluwricus sehr hiiufig. Vcreinzelt : Macrothrix laticornis, Pleuroxus cf. trigoncllus und cf. adun- cus. Ein einzelnes Stiick einer schlecht crlialtiMicu und daluT niclit iklfrniinicr- baren 1 lelnidaiihnie.
Pool near niik'stune 4. Moina dubia, Cliydorus spec.
Ootacanumd, small grassy pool: C/(y(/<)r)M-fragmente.
Umbrella Tree (2316m.): Chydorus sphacricus.
Ponds. Pykara Road near Ootacamund Moina esau.
]"".benda: Ccriodaphnia quadrangula, Scapholchcris Kiiigi ?, Plcuroxus si)ec. ( vgl. Seite 296) . Chydorus sphacricus. N 19 Pykara: Bosinina longirostris I. cornula.
Yanni Hole Mysore : Siiuoccphalus vctuloidcs.
|
N 6 |
|
N 9 |
|
N13 |
|
N14 |
|
N 15 |
Diese Tabelle mag noch durcli einige Angaben iiber Cladocerenfunde in den Ilochgebir- gen des siitllichen China ergimzt werden. Diese stammen von O Pe.sta — Wien und finden sich veri)ffentlicht in nieiner ;\rl)eit ..iilxr siidasiatische Diaptomiden" (Archiv f. Idydrobiologie Bd. XX I I. 1''30.) — Das Material stainnite mmi den I'orsrhuiigsreisen llandel-Mazettis. Seite 158 ft.
Setschwan. Sumpfgewiisser am Tscheschajoch. 4100
Yiinnan. Yiinnau. Yunnan.
Yiinnan. Yunnan.
I'ongatong. See 4075 m. Tsukne. See 3825 m. See bfi Likiang. 2820 m.
See l>ei Mahaidse. 3675 m. Wahaschimi. See 4325 m.
Alona costata^tubcrculala Chydorus sphacricus Chyilorus sphacricus L li vdnrus sphacricus Chydorus sphacricus Graptolcbcris tcstudinaria . Ilonclla c.vcisa Hosiuina Unigiroslris Ccriodaphnia spec. Chydorus sphacricus Alotui guttata Alona costata
REPORT ON CLADOCERA
289
Man sieht, class die faunistischc Zusamnieiisetzung ganz der entspricht, die aus unserer Taljelle fiir die weiter westlich gelegenen Hochgebirge zusammengestellt wurde, nur dass hier gar keine spezifischen Faunenbestandteil zu verzeichnen sind.
BEMERKUNGEN ZU EINZELNEN ARTEN
1. Moina esau n. sp.
Text-figure 1, a-f
In der Bearbeitung der Cladoceren der Deutschen Limnologischen Sundaexpedition habe ich liei einem provisorischen Versuch, die mir bekannten Moina — arten iitersichtlich zu gruppieren jene Arten als besondere Gruppe herausgegriffen, die durch einen Haarbesatz
Figure 1. Moina esau n. sp. A, Ephippialweibchen ; B, Kontur des § Postabdomens ; C, Distaler Teil des Postabdomens, Bewiniperung der Winiperzahne nicbt eingezeichnet ; D, Anteiiiiula des ? ; E, Antennula des c5 ; F, I Fuss des £ . Eiidteil vvegeii Rauminangel separat gezeichiiet.
am Kopfe sich von den anderen unterscheiden und die man nach der am langsten liekannten zugehorigen Form als Banff\i-C,i-u\)\>e 1)ezeichen konnte. Diese Gruppe umfasste bisher die Arten Banff yi aus luiropa, Belli ( iurney aus Siidafrika und tonsitrafa Hrehm aus W'estafrika. Zu diesen Arten kumnit nun als vierte die hier zu l)eschreilx'n(le neue Art esau ans dem Nil- girigebirge. Bevor ich deren Beschreibung gebe, will ich nochmals wie in der Arbeit iiljer die Sundacladoceren und wie bei der Beschreibung der tonsnrata betonen, dass es mir sehr zweifelhaft scheint, ob diesen Arten eine wirkliche Verwandtschaft zuzuschreiljen ist. Ihre
290 REPORT ON CI.ADOCERA
Zusamnltnfassung erfolgte ja nur mil Riicksicht auf die fiir einen Bestimmungsschliissel sehr bequeine Abtrennung von den iibrigen Arten.
Ini Korperuniriss unterscheidet sich unsere Art von den Vergleichsarten durch eine hinter dem Auge gelegene leichte Impression. Die Kopfhaltung entspricht der zwischen ,,semierect" und „semidepressed" gelegenen Stellung der dubia — Form nach dem Schema, das Penelope Jenkin auf Seite 148 ihrer Arbeit Cladocera from the Rift Valley Lakes in Kenya (Ann. & Mag. Nat. ist., 1934. Ser. X. Vol. XIII.) niitgcteilt hat. Die antennula des Weibchens zeigt nicht das schuppige Bild wie bei Banffyi, doch ist die Kontur fein wellig und den kleinen \Vellenbergen sitzen kleine Stacheln auf. Die seitliche Sinnesborste ist etwa in der Mitte inseriert. Die Endkralle des Ncbenkamnies l)esitzt im Gegensatz zu Banffyi und Belli, aljer in Ubereinstimniung mit tonsuratn einen Nebenkamni. Die Zahl der Wimper- ziihne betragt 10 oder 11.
I-lphippialwcibchen lag leider nur ein schlecht erhaltenes vor, das keine verlasslichen Angaben iiber die Struktur des liphippiums erlaubte. Doch enthielt das Ephippium jedes- falls nur ein Dauerei. Auch Miinnchen fand sich nur eines in dem Material, dessen erstes Beinpaar in Figur 1 abgebildet ist. Die antennula tragt am Ende 4 Ilaken,^ wie totisurata, wiihrend Belli 6 Ilaken haben soli und fiir Banffyi niir keine Angal)en hieriil)cr bekannt sind. Die Knickung, an der die seitliche Sinnesborste inseriert ist, befindct sich \k\ esau etwa in der Mitte der antennula, wiihrend bei toiisurata diese Stelle etwa im proxinialcn Drittel liegt und l)ei Belli eher noch vor diesem.
Fassen wir also die mit behaartem Kopf versehenen Moinaarten der BccjuemHchkeit halber als eine Gruppe auf, so ergiibe sich folgende Trennung der Ijetreffenden Arten :
I. Endkralle mit Nebenkamni :
1. Mannliche antennula in der Mitte geknickt: esaii Brehm
Mannliche antennula im ersten Drittel geknickt : totisurata Brehm II. Endkralle ohnc Nebenkamni:
1. Scliale hinten in eine Spitze ausgezogen Belli (Jurncy
Scliale hinlen abgerundet Banffyi Daday
Nach dieser Tabelle scheint die Trennung von Belli und Banffyi recht gekiinstelt. Leider liegen mir die beiden Orginalbeschreibungen nicht vor, die vielleicht eine sicherere Trennung moglich machen wiirden.
2. Dunhez'edia crassa var. interrupta n. Text-figure 2, c Von der Fundstelle K 34 in Kaschniir lag in mehreren parthenogenetischen Weibchen die genannte Art in einer Form vor, die neu zu sein scheint. \\';ihrciid namlich nach den Abl>ildungen, die Stingelin von dieser Art sowohl nach Schweizer (Text-figure 2a), wie auch nach indischen Exemplaren (Text-figure 2b), gibt, die Lippe niehr oder wcniger zugespitzt ist und der Vorderrand dersellxin eine gleichmassig fortlaufende Kontur zeigt, war liei den
' Nachtriiglich fand ich ein Miinnchen, das an der einen antennula 5, an der andcren 6 Haken triig, sich also in diesem Punkte der Art Belli naherte.
REPORT ON CLADOCERA
291
Figure 2. IJppe von Diiiihcvedia crassa King. A, Form von Basel, nach Stingelin ; B, Form von Indien, nach
Stingelin ; C, Form interrupta aus Kaschmir.
Tieren aus Kaschmir die Lippe stunipf und der Vorderrand zeigte eine Knickung der Kon- tur (Text-figure 2c), die vielleicht als Anlauf zu einer Zahnbildung gedeutet warden kann, wie eine seiche bei anderen Arten dieser Gattung vorliegt.
3. Drepanomacrothrix sp. Text-figure 3, a-c
Die Probe L50(2) von Toganna-Tso enthielt ein vollstiindiges und zwei defekte Weibchen einer Macrotrichide, die schon ohne nahere Untersuchung als zu der von Werest- schagin aufgestellten Gattung Drepanomacrothrix gehorig erkenntlich war. Da ich an der
r^
FiGiTRF, 3. Dref'a)iomacrothnx sp. von Togarma Tso. A, Antennula ; B, Antenna ;
Postabdomens.
C, Disfaler Teil des
292 REPORT ON CLADOCERA
Existenzbereclitigung- dieser Gattung Zweifel liege, ware es natiirlich erwiinscht gewesen, wenn ich durcli Untersuchung der vorliegenden Form die Berechtigung nieincr Zweifel hatte erweisen konneii. Ich woUte aber das einzige unverselirte Exemplar nichl opfern und die i^iden defekten Exemplare gestatteten keine so eingehenden Ergebnisse, wie sie zur Begriindung meiner Meinung notwendig gewesen wiiren. Ich habe daher vorliiufig die Bezeichiuing Drcpanoiitacrotliri.r beibehalten und lasse auch die Speciesfrage offen, obwohl selbst die unvollstandige Beschreibung erkennen litsst, dass in mancher llinsicht unsere Eorni von lien zwci oder drei l)islier bekannteii und als Prcpanoviacrothrix zusanimengefassten Arten verschieden sein diirfte; freilich bliebe auch die Moglichkeit dlTen, dass alle diese Arten Fornien einer einzigen, variablen Art seien.
Bevor wir auf die Erorterung dieser Angelegenheiten (.■ingclun, wird es gul sein, sich iiber die Aufstellung der Gattung Drcpanonuurothrix und ii])er die zu dieser Gattung gerech- neten Arten zu orientieren, zumal da die Aufstellung dieser ( lattung in einer niclit leiclit erreichbaren Zeitschrift erfulgte.
Uber die Gattung Drcpiuioiiiacrolhrix
Die Aufstellung der fraglichen Gattung erfolgte durcli G. Werestschagin in seiner Arbeit ,,Nutiz iiber die Cladocerenfauna des Karsischcn I'lateau und des Batumischen Strandes" (Arb. d. Hydrobiol. Station am See fJlubokoje. Bd. \'. Lief. 1. 1913). Da mir diese Arbeit nicht zugiinglich w^ar, hatte Ilerr Dr. Smirnov — Leningrad die Lieljenswiirdig- keit, mir nicht nur eine Abschrift des Deutschen Resumes dieser Arl)eit — soweit unsere Gat- tung in Frage kommt — sondern audi Kopieii der dnrl niitgeleilten l'"iguren zu iibermitteln, wofiir ich deni Geiiannten an dieser Stclle iiuclimals herzlichst dauken niuclite. LIberdies al)er teilte mir Herr Dr. Smirnov etliche weitere, nur aus dem russischen Text ersichtliche Daten niit, die ich hier einzuschalten fiir gut linde, da auch den ineisten anderen Cladocer- enuntersuchern diese Mitteilungen schwer erreichbar sein diirften.
Gefunden wurde diese neue Form von Werestscliagiu am 2. VII. 1907 ini See Chant- schanly (Kreis Aclialkalaki, Gouv. Titlis) in 4 weiblichen I^xeniiilaren. Dieser See liegt 6385 Fuss ii.d.M. ist 6.5 km. lang, 2.5 km. breit und 1 bis 1 ' _■ m. tief. Die Ufer sind stark bewachsen. Decksbach ist der zweite Autor, der diese l'"urm gefunden hat und zwar ini kleinen Kabarda ini Nordkaukasus. (Zur GladcKereu fauna von Kaukasus und Nurd — I'ersien. Trav. de la Stat. Jiiol. du Caucase du Nurd. v. III. fasc. 1-3. 1930.)
Beschreibung der Gattung Prrptiiidiiiacrnlhri.v und ilirer Art
StscliclkanozvccTi'i nach Werestschagin
IhYpiiiKiiiuhrdtlii'XX (sic !)
„Bei der Seitenansicht befindet sich am J\iickenrand des Ki'irpers cin zalinartiger l-'ort- satz; er ist vi>n einem kragenarligen Fortsatz iles 1 liuterraudes der Kopfschildes gebildet. Der Kopf ist konusartig zugespitzt; das Auge i.st gn'isser als der i'igmentlleck ; die ersten Antennen bestehen aus einem Glied ; der Aussenast der 2. Antenne ist 3 gliedrig init 5 Schwimmborstcn ; der Innenast viergliedrig init 4 Schwimmborstcn ; Kiemenfiisse, die denen der Drcpanothrix dcntata sehr ahnlich siiul. Die Cauda ist breit und kurz; ihr Riickenrand besteht aus zwei konvexeu Teiien. Der .\nus belindet sich an dem, der den
RKPORT ON n.ADOCERA 293
caudalen Krallen am niichsten liegt. Der Darmkanal ist ohne Schling-e und Blindsacke. Die Abdominalborsten sind zweigliedrig, die Abdominal fortsJitze fehlen.
Zum neuen Genus gehort nicht nur die neubeschriebene Fomi, sondern auch die im Jahre 1903 von Daday beschriebene Macrotlirix cornuta, der die neubeschriebene Form am niichsten steht. Beide stehen zwischen dem Genus Drcpanotliri.v und Macrothrix und miissen in eine besondere Gattung gestellt werden, da sie durch das Fehlen der Darmschlinge den Arten von Macrothrix sich nahern und durch die Form der Kiemenfiisse und den Riicken- fortsatz der (jattung Drepanotlirix.
Drcpaiwiitacrotlirix Stsclielkanowzewi n. g., n. sp.
Die (iesammtform des Korpers ist oval. Am Riickenrand befindet sich am Ende des Kopfschildes ein zahnartiger Fortsatz; am Hinterende des Riickenrandes befindet sich ein gut ausgesprochener Kiel, der am Aussenrand seiir kurze und seltene Dornen triigt. Der Hinterrand ist mit unregelmassigen Tuberkeln, die am Ventralrande allmalilich zu sagear- tigen Durnen werden, versehen. Der Vorderrand tier Schalenklappen entl)ehrt aller Haare und Borsten. Das Auge ist zweimal grosser als der Pigmentfleck, die Entfemung vom Auge bis zum Pigmentfleck ist beinahe ebenso gross, wie die vom Pigmentfleck zur Basis der antennula. Die Fomices sind nicht stark entwickelt. Die antennulae sind eingliedrig, schwach kolbenformig und konkav. Ihr Vorderrand ist wellenformig, in 9 bis lo Telle geteilt, an deren iuide lange und starke sehr durchsichtige Haare sich befmden. Am Ende l)elindet sich eine 'i'astborste und ein Biindel kurzer, diinner Flarchen.
Bei der zwciten Antenne trJigt das erste Glied des Aussenastes am Ende eine 2 gliedrige, an einer Seite bewiinperte Schwimmborste, das 2. Glied triigt die liingste Schwimmborste der Antenne, die dreimal liinger als das Aussenglied der Antenne ist. Diese Borste ist zwei- gliedrig und triigt am Ende des ersten Gliedes cinen .starken Dorn. Das dritte Glied triigt drei gleich lange, zweigliedrige, einseitig be\vim[)erte Schwimmborsten und ausserdem am Ende einen starken Dorn. Das erste Glied des Innenastes triigt keine Borsten, das 2. ist an der Innenseite am Ende mit einem stumpfen Fortsatz versehen, an der Hinterseite befindet sich ein starker, langer, krummer Dorn. Das dritte Glied triigt an der Innenseite einen eben- solcJR-n I'ortsatz wie das zweite, und am luule der Aussenseite eine Schw imnijjorste ; diese ist (lie kiirzestc Sclnvimmbnrste. Das 4. Glied triigt am l-jide drei Schwimmborsten wie beim Aussenast. Das Labrum ist zugespitzt. Die Maxillen bestehen aus drei schwach gekriimmten, befiederten Ziihnen. Der Darmkanal ist ohne Schlinge. Der Riicken der Cauda jjesteht aus 2 konkaven Teilen, von denen der, iler den Abdominalborsten zuniichst steht, zweimal liinger als der andere ist. An den Seiten der Analfurche finden sich 6 bis 7 .schwach narh binten gekriimmte Dornen. Der pro.ximale Teil der cauda isl am ivande mit zahl- reichcn diiniieu kurzen Dorneu vcrscjieu. Die Gaudalkrallen sind glalt und (.hiie Piasal- stachel. AbdiiniinaIfi)rts;Uze fehlen. In der iirutkammer befanden sich 1 bis 2 Embfyonen. Die Dimensionen sind f olgende : Eiinge = 0.42 jjis 0.43; grosste Breite ^= 0.28 bis 0.29. Die Liinge der antemuda= 0.12 mm. Die Liinge der Cauda von der Basis der Abdominal- lx)rsten bis zur i'.asis der Candalkrallen 0.14 nun."'
Werestschagin charakterisiert also die Drcpaiuniiacrothrix als Zwischen form zwischen Macrothrix untl Drcpanothrix und jjegriindet seine Ansiciit damit, dass das b'ehlen einer Darmschlinge seine Gattung in Beziehung zur Gattung Macrothrix setze, wiihrend der zahn-
294 RErORT ON CLADOCERA
artige ^^iickenfortsatz sowie der Ban der Beine Beziehungen zur Gattung Drcpanothrix bediiige. Es scheint niir nun, dass die Gattung Drcpaiwiiiacrothrix zur Gattung Macro- thrix gehore, mit der sie u. a. in folgenden Punkten iibereinstimnit : Die von Werestschagin als Tastborste uiid Sinneshaare bezeichneten Sinnesschliiuche entsprechen ganz den Verhalt- nissen bei Macrothrix, nicht aber denen von Drepaiiofhrix; die zweite Antcnne ist cine typisclie Macrothrix antenne, da sie einen init 4 Borsten versehenen Aussenast liat, wiihrend bei Drepanothrix das vorletzte Glied dieess Astes keine Schwimniborste triigt, also der ganze Ast nur dreiborstig ist. Der von Werestschagin erwahnte an der Gliederungsstelie der liing- sten Schwimniborste befindliche starke Dorn ist bei viclen Macrotlirix artcn in der glciclien Form entwickelt. Der Hintcrkcirper der zu DrcpanoDiacrothrix gestellten Formen erinnert sehr stark an den mancher Macrotlirix artcn, z.B. den \nn Macrothrix spinosa und zeigt die TeiUnig in zvvei konvexe — Werestschagin spricht von kt)nkaven — Abschnitle, ein W'rlialtcn, das fiir Drcpanothrix nicht zutrifft.
Stimmt somit Drepanomacrothrix weitgehend mit Macrothrix iiberein, so zeigt sich andererseists, dassdie Ubereinstimmungen mit Drcpanothrix nur sciieinbare sind. Denn der Riickenzahn der Gattung Drcpanothrix scheint mir etwas ganz andcrcs zu scin als der zahnartige Vorsprung des Hinterrandes des Kopfschildes bei Drcpanoinacrotlirix. Und was die Ubereinstimmung der Gliedmassen anbelangt, sei zuerst erwiihnt, dass Werestschagin gar keine spezielle Beschreibung derselben gibt und dass Daday zwar Bilder und Beschrei- ung seiner heiher geh("irigen Art mitteilt, dass aber diese kaum bcweisend gcnug" sind, da ja nach Lilljeborg iibcrhaupt kein fundamentalcr Unterschied zwischen den l^xtrcmitiiten dieser beiden Gattungen besteht, was nicht verwunderlich ist, da j;i inncrhalb tier Gattung Macro- thrix weitgehende Verschiedenhciten im Extremitiitenbau vorlicgen. Lilljeborg sagt diesbe- ziiglich von Drcpanothrix in seiner Monographic der schwedischen Cladoceren (pag. 368) : "Es sind fimf z. Th. denjenigen der vorigen Gattung und der Gattung Macrothrix ahnliche Fusspaare vorhanden." So diirfte die Abtrennung der Gattung Drepanomacrothrix von Macrothrix kaum haltbar sein.
Eine andere Frage ist nun die, wie sich die von der Yale-Expedition gclundene Art zu den bereits bekannten hieher gehorigen Arten verhiilt. Vergleichen wir unsere Art mit der kaukasischen Stschclkanowzewi, so finden wir folgendc Differenzen : Bei der kaukasischen Art ist die Entfernung des Auges voni Pigmentfleck etwa eben so gross wie die Entfernung von diesem zur Insertionsstelle der antennulae, bei unserer Form ist die erste Distanz ctwa doppelt so gross als die zweite. Die Stacheln am ventralen Schalenrand sind Ijei uusercr Art viel langer. Fiir die antennulae der Art Stsclielkanoivzeiifi gibt der Entdecker eine Teilung des Vorderrandcs in 9 bis 10 Abschnitte an, wahrcnd sich bei den Tieren der Yale expedition deren nur 6 bzw. 7 finden liessen.
Bei einem Vergleich mit der turkestanischen Art cornuta ergeben sich folgende Differ- enzen : Die Art cornuta trJigt an der antennula zwei lange Sinnesschlauche, unsere Art hat nur einen ; unsere Form zeigt an den Gliedern der zweiten Antenne drei fliichenstandige Haarsiiume, wiihrend fiir cornuta von Daday eine unregelmiissige Behaarung abgebildet wird. Die Art cornuta zeigt am distalen Teil des Postabdomens ,,5 Querreihen feiner Borsten," wahrend unsere und die kaukasische Art bier ,,nach hinten gekriimmte Dornen aufweisen. Es giibe vielleicht noch manche andere Differenzen, aber wenn man auf einen Vergleich mit Bildermaterial angewiesen ist, konnen hier leicht Irrtiimer unterlaufen, weshalb ich nicht wciter auf einen solchen Vergleich eingehe.
REPORT ON CLADOCERA
295
Resumieren wir das gesagte, so ergibt sicli, dass die als Stschelkanowzewi, cornuta heschrielienen Arten sowie unsere Form wolil eine zusaninien gehorige iniierasiatische Gruppe von Macrotlirix formen l^ilden, iiljer dereii Sell)standigkeit sich olme Nachpriifung von Originaltypen zur Zeit nichts sagen lasst. Ob, wie Daday meint, auch odontocephala dazugehort, bleiljt nocli unentschieden. Fraglich ist es, ob man in diesen Formen einen Uhergang zur Gattung Drepanothrix sehen darf. Nach meinen oben gemachten Mitteil- ungen zweifel ich daraii. Es sclicint niir eher, dass Drepanoflirix zu den altersstarren Mono- typen gehort, die zusammenhanglos neben den anderen Gattungen ihrer Verwandtschaft stehen.
4. Uljer eine vermutlich ncue Alona aus deni Tso Moriri ; A. ladacensis
Text-figure 4, a-c
Unter den ziemlich vielen Exemplaren von Alona guttata, die allerdings durch langs- gestreifte Schalenklappen vom Typiis aljwicben, fand sich neben einem vereinzelten Stiick der Alona rcctangula ein Weibchen einer Alona, das auf den ersten Blick ein guttata — Exemplar zusein schien, sich bei niilicrem Ziiselien, aber als ziemlich abweichend gebaut erwies.
]''u;uRK 4. A, Alona ladacensis- n. sp. 13, Vordereiide unter starkem Deckglasdruck ; C, Postabdomen ; D, Alalia sp. aus dem Tso-Nyak; E, Postabdomen.
Die undeutlich polygonal gefelderten aber stark punktierten Schalenklappen dieses 500 /* grossen Tieres zeigten den durch unsere Figur gezeigten Umriss. Auch die Form der anten- nulae und der breiten gerundeten Lippe ist aus der Abbildung ersichtlich. Was nun dieser Form ein besonders auffallendes Aussehcn gibt ist die Bewehrung des Abdomens, das im Umriss dem von guttata ziemlich nahe kommt. Die distale Halfte tragt am Rand 6 Gruppen
296 REl'OKT ON C1.ADUCERA
von niei^t zu dreieii lieisainiiKMistcliciukMi s^toIm.'!! Staclicln und olR-rlialli ilit-sor an der I-'lankc mehrcre TIaari;Tui)i)cn. I-ls ist mir kcine Alona hekaniit dit' dicsi' lui^fntiiiidiclikeit zcigte ausser ciiicr l'\inn. die iiioi^licherweise einen ahnlichen Ban hal und die von Schiklejew in seiner Arl)eit ..Die Cladocercnfauna der kaukasischen llegegewasscr," etc. I Archiv. fiir llvdrohiologie. I'.d. XXI. I'l^O) aiif Seitc 34,3 hc.^cliriehcn und auf Seite M4 ahgehiklet wurde. Da iliin audi nnr lin cinziges l^.xeniphir vorlag. unterliess Schikk^jew eine llenen- nung. Leider sind die Figuren (k's genannlcn .Vutors so niangellialt, dass man u\>vv ukukIrmi I'unkt Zwcifel hegen muss und keiuen einwandfreien X'ergleich (hn-clifiiliren k.iun. Zutkni hig (k'ln gcuannten I'iologen ein mannlichcs Tier vor, so dass die Unterscliieck' ini Uniriss sowolil cks Schaknl)ildes als audi des Posta1)doincns auf (iesdikclitsdifferenzen zuruckgelicn konnten. P'rappant ist jeck's falls das Vorhandenscin von ,,sedis IWindel Orn-ndien" am dis- talen Rand des Postahdomens. Im weitercn \'crlauf spridit Schiklejew von 5 bis 6 W'im- pern, aus denen sicli jedes soldics Biindcl zusaminensctzt und audi seine Ahhiidung spridit dafiir, dass es sich uni feine Borsten handle, wiihrcnd in unscreni I'all grohe lOornen oder Zahne vorliegen. jcdesfalls ist das Vorhandenscin von seclis (iruppen soldier .Anliiinge eine selir aufl'allcnde Uhereinstimmung. RTerkwiirdig ist es. dass sowohl bei <ler aus deni kauka- sischen See Inkrit beschriebenen Form wie bei luiserer, nur je ein I'.xemplar gefunden wurde, was den Verdacht aufkonimen lasst, es handle sich um cine vereinzelte Mutation cincr bereits bekannten Art. Andererseits modite ich al^er doch zu Ijedenken geljen, dass diese I'orm gcrade aus zwei Geljieten vorliegt, die tiergcographisch cnge Beziehungen aufweisen, wenn wir an Drcpanotitacrothrix denken und daher eine Benennung dieser Form in Vorsclilag- bringen : Alona ladacensis.
5. Alona sp. .''
Text-fisure 4, d-e
In der f'robe aus dem Tso Nyak, T.71;i, dcr in cincr Scchohc \'on 4241 m. gelegen ist, fand -ich ein einzelnes Rxeniplar cincr .lloiiti, vim dcr ich abcr nur das .Sdialcnbdd und den Ban des I'ostabdomens erniittcln konnte. Lipjie und antcnnnla warcn nicht gut crhaltcn. Ich bin im Zwcifel. ob ich diese .\rt niit eiiier bereits Ijckannlcn idcntitizicren kann und niit weldier und gcbc dahcr bios die beidcn Figuren, aus denen ersichtlidi ist, dass die am unteren hinteren Schalenwinkel al^erundete, nicht gezidinte Sciialenklappe niit weit iiber 20 recht niarkanten Langslinien versehen ist und flass das kurze, breite Postabdonien ctwa 8 kurzc Ixandstadieln zeigt, liinter denen noch einige Borstchengrupi)en folgcn. snwie fcrner, dass die Flanken des Postabd' miens mil klcincn Brirstchcngruppcn vcrschcn sind. Die luid- kraile triigt einen Basaldorn.
6. Die /'/(•»r().r/(j-— Artcii
Plciir('xiis ailiiiuiis Jurine. Sowohl die wenigcn I'.xcmplarc \dn P 2, .SoJiawa als audi das einzelne ]''xem])lar von Oot.icunnnd Lake stimmtcn mit dcm 'lyims so weit iiljerein, dass man sic ohiie Bedenken als adiincits bezeichnen kann. Anders liegt der I'.ill bei den PIcuro.vus exeniplaren, die sich in einigen Proben aus dem Nilgiri Gebirge iK'fandcn. Ilier liegt die Schwierigkeit darin, dass innerhalb derselben Population sich nicht nur .selir l>etrachtliche Verschiedenheiten bei verschiedenen .Mterstufen zeigen. sondern, dass auch etwa gleichalterige Stiicke eine starkc individuelle \^ariation aufweisen. Die von Birge
REPORT ON CLADOCERA 297
vorgenoiiinienc SclifiduiiL; in lani;.s,rfstreckte und Imcli^cw i illile l'"(iriiK-ii liissl liii-r iiisofcrne im Sticli, als junt^e I'iere niclir dein ersten T3-i)us zuiieigen, iiltcre luit I'.mljryoncn versclienc al)er deni zweiten. Nach solchen iiltcren Typcn nn'iclitc icli alle dicso Tierc dt-in tri}^onclliis — Kreis zurechncn, docli zeigen vielc Individuen cine Iciclitc, lioim lypisclien trii:;onr!liix niclit vorliandene Konkavitiit des Hiiiterrandcs des rostahdomens, die als Kennzcichen der anieri- kanischen Art dcnticiilatits angesehen wird. In dieseni Punkte kiimen auch die Arten assiiiiilis Brady aus Ostafrika sowie die leider nicht gut ahgebildete Art australis Henry von Aiistra- lien als Vergleichsarten in Retracht. Rci deni gegenwiirtigcn Stand der Systeniatik dieser (iattung- ist es wohl am Ijesten die Tiere als dcnticiilatus-\)7.\x (7.s\v/;;;/7/.s-;ihnliche Fornien aus dem trigoncUus Kreis zu bezeichnen. Die Plcuroxus arten scheinen sehr varial^el zu sein und ich vermute, dass die grosse Menge der bisher beschriebenen Arten sich l)ei genauerer Analyse dieser Formi-nkrcisc- auf selir wenige Arten rcduzicren wird, in welcheni Falle wolil die Nilgiri-Tiere zur Species trigoiwllus fallen werden.
7. Chydonis n. sp.? Text-figure 5, a-b
Die I'robc N 15 aus den Nilgiri Hills entliielt eincn Chydanis aus der spliacriciis — Gruppe ini weiteren Sinn, der wohl ncu sein diirfte, den ich aber — da nur parthenogene- tische Weibchen vorlagen — nicht als neu beschreil:)en will, da gerade bei einer so vielgestal- tigen und nur durch minutiose Merkmale die Artentrennung ennoglichenden Gruppe zur Charakterisierung einer Art die MJinnchen und womoglich auch die Ephippialweibchen bckannt sein snllten.
A
Figure 5. Chydonis n. sp. ? der C. spluwiuiis Ciriippe. A, Vurdereiide ; B, Postabdumeii.
Unserc l'"iirin stcht wohl den ;ds riirxiinl ii.'i uwd (/r;;//(7//(////.v beschriebenen [■"(irnien durch die breite nicht zugespitzte Ltppe am niichsten, docli zeichnet sich diese Lippe durch einen kleinen knopfformigen Vorsprung aus. Ob diese Kleinigkcit konstant ist und als Merkmal einer kleinen Art vcrwendet werden kann, ist bei der Variabilitat, die die Lippenformen gerade in der sf'liacriciis — Gruppe zeigen, noch unsicher. Das rnstabdimien und die i'.nd- klauc zeigen in Form und Bewehrung grosse Ahnlichkeit mit der von Roljert Gurney fiir Cliydorus dcnticidatus gegebenen Abbildung, wie unsere Figur zeigt (Vgl. dessen Arbeit ,,Some Australian Freshwater Fntomostraca reared from dried mud." Proc. Zool. Soc. London. 1927. Fig. 11 auf Seite 77.)
ARTICLE X\II
REPORT ON FISHES. PART I: COBITIDAE'
By Sunder Lal Hora, D.Sc, F.R.S.E., F.A.S.B.
Zoological Survey of India, Calcutta
(Received August 16, 1935)
CONTENTS
Page Introduction 299
Ecology and Structural Modifications 300
Air-bladder 300
Caudal fin 303
Geographical Distribution and Origin of the Fish Fauna 304
Systematic Account 306
Nemachilns stoUc"kac (Steindachner) 306
Ncmachihts gracilis Day 309
Nemachilus microps ( Steindachner 310
Ncmachilus tcnuicauda (Steindachner) 311
Nemachilus vittatus (tieckel) 311
Nemachilus deTerrai, sp. iiov 311
Nemachilus hutchinsoni, sp. nov 314
Nemachilus panguri, sp. nov 318
Botia hinti Chaudhuri 321
introduction
Mr. G. E. Hutchinson, biologist to the Yale North India Expedition, entrusted to me an extensive and interesting collection of fishes for study and report in October 1933, but it was not possible to take up the work till the later part of 1934. The collection was made in the Kashmir Valley and Ladakh, and comprises representatives of Sisoridac (Glyplo- fhora.v and Glyplosfcnmin), Cypr'umme (Labco and Crossochcilus), Schisothoracinae {Schiso- tliora.v, Ptycliobarbus, Schicopygopsis, Diptycliiis and Orcinus) and Cohitidac (Bolia and Nemachilus). As was to l)e expected, Glyptotliorax, Labco, Crossocliciliis and Botia were found only in the Kashmir Valley, while Nemachilus and the Schicothoracinae were cciivdUy abundant at high altitudes and in the Valley. In this article, I propose to deal with the Cobittdae, especially the genus Nemachilus, which in the lakes and torrential streams of Central Asia has proliferated into many species showing diverse structural, adaptive modifi- cations. The present collection contains one species of i?of/a and eight species of A'^r/;;ar//(7».?.
1 take this opportunity to offer my sincere thanks to Mr. Hutchinson for affording nie an npportunity to investigate this interesting material and for his invaluable field notes. I am indebted to the authorities of the Yale North India Expedition for a grant towards tiie cost of drawings which were executed by Balm R. Bagchi under my supervision.
' Published witli permission of the Director, Zoological Survey of India.
Mem. Conn, .-\c.\u., Vol. X, .Art. XVII, June, 1936.
300 REPORT ON FISHES. PART i: COBITIDAE
ECOLOGY AND STRUCTURAL MOPtFICATIONS
Ecologically, the eight species nf tlie .i;cinis Xi'iiunliihts represented in tlu' cullectiMU nf the Yale North India Expedition may I)e divided into three "associations": (i) llottmn- dwelling species of the lakes, snch as A^ 7'ittati(s, which live in 4 tn 6 feet of water Init lirohably rise from the Ixjttom occasionally and swim al)out ; ( ii ) lK)ttoni-(lwelling species in torrential streams, such as A', stolicckae, N. gracilis, N. niicrops and X. tcnuicaiida. wliicli haliitually live atlliering to rocks and stones in swift currents though at times, especially dur- ing the i)reeding season, mav enter into sjirings, pools and lakes; (iii) free-swimming lake species, such as N. deTerrai, N. hutchinsoni and N. panguri. wiiicli swim alxnit freely in still waters luit for feeding purposes have to cling to rocks and otlicr ohjects, usually at the lK>ttom. ("orreiated witli the al>ove differences in hahits and habitats, the species have undergone remarkable modifications in the structure of certain organs.
Air-bladder. In 1930, P referred to the modifications of the air-bladder in species of Nemachilus from several localities and indicated the close relation I)etwcen its structure and the type of habitat in which the species lived. On account of the occurrence of gradations Ix'tween the Ncmacliiius-typc of bladder and the Dif^lophysa-iype of bladder, it was indicated tliat the differences in the structure of the bladder could not be used for taxonomic pur|)Oses. Rendahl'^ has, however, used this character in proposing several sub-genera for the species of A^cjiiarliilus obtained by Dr. Sven Hedin in Central Asia, and has given a detailed morpho- logical account of the modifications observed by him. The accompanying figure shows some of his illustrations and an attempt is made below to explain the possible significance of these modifications, as I interpret them.
The structure of the air-bladder of A', barbutiila ( Text-Hgure 11'") is characteristic of the species that live in swift currents and, though they may dart from place to place, are rarely seen to swim. In these circumstances, the air-bladder has lost ils biioy.uit function and its anterior portion is represented by two small lateral chambers {a) enclosed in bony capsules and the posterior chamber by a small, thick-walled bag (c). This type of struc- ture is found in A'^. stoliczkae, N. gracilis, N. niicrops and A'^. tcnuicauda. In N. yarkandcn- sis (Text-figure ID) the fonn of the bladder remains the same but the lateral chamljers are greatly enlarged so that they come in close contact with the skin. Tn the areas of contact, the Ixiny capsules are incomplete so that the bladder can react to the surrounding changes in pressure. The structure of the bladder in .Y. yarkandcnsis is more or less similar to that of A^ vittatus (Hora 1930, Text-figure 6c) and probably the habits and habitats of the former are similar to those of the latter. A'^. vittatus is a lake species and the Yale North India I'-xjic- dition obtained several specimens, mostly from the weedy marginal areas of the Kashmir lakes. Probably the species lives at or near the bottom and does not swim about niucli. The Netherlands Karakorum Expedition obtained specimens of A'^. yarkandcnsis from pools in the neighbourlioo<l of extensive marshes at Rabat-Utsang. The real lake forms that move al)ont in all possible directions are characterised by a bladder of the type found in N. stnvarli (Hora 1<'*30. Text-figure 8), ^V. hiitjertjuensis (Text-figure lA) and three new species of Nemachilus described hero from Western Til>et (Text-figures 5/), //;, '^t). The posterior
'Hora, Jount. Homlmy Nat. Hisl. Soc. XXXIV, pp. 379-385 (1930). "Rendahl, Arkiv for Zoologi XXV, No. 11, pp. 1-51 (1933).
REPORT ON FISHES. PART I : COBITIDAE
301
chaniLer of the I)la(](ler lias assumed the form of the typical Cyprinid bladder and probably functions in exactly the same way.
The type of bladder found in A^. kiingcssaniis (Text-figure IB) shows that the spec
les
Fkuire 1. Various types of air-bladder found in Nnnailtiliis (after Rendalil). Various magnified. A: .V. Iiiiljcrljucnsis Rend.; B: N. kiingcs.minis (Kessl.) ; C: A'. pafillo-Iabiatus (Kessl.) ; D : A^. yarkandnuis Day; E: A^. hsulschonrnsix Rend.; F: A', barhatnia (Liiui.). a = portion of bladder enclosed in bone; 6 = duct con- necting; the enclosed and the free portions of the air-bladder; r = anterior chamber of the free portion of the bladder; rf = posterior chamber of the free portion of the bladilcr ; c = pneumatic duct connecting the air-bladder with oesophagus ; g ■= gut.
has reverted again to a ground habit of life in comparatively swift currents. Its long, nar- row and thick-walled posterior portion shows that the bladder is losing its utility as a hydro- static organ. In this connection reference may be made to the modification of the swim-
302
REPORT ON FISHES. I'ART i: COBITIDAE
bladder 411 species of the genus Garra* in which the torrential species possess a similar type of posterior chamber. This process of retrogression is much more pronounced in A'^. hsiit- schoiicnsis (Text-figure IE).
The type of bladder found in N. pal>illo-labiatus (Text-figure IC), N. strainiiii (Hora 1*^130, Te.xt-figure 7) and A', actiticcplialus (Ilora 1930, Text-figure 9) is rciuarkable in so far as the free bladder is situated very far back in the abdominal cavity and is connected with the bilobed anterior portion and the oesophagus ])y means of a long tul^e. In the case of A'^. acuticcphalits I surnu'scd tliat the pfisition of the liladder was ])riil)ably due to its burrow-
Fir.uRE 2. Form of caudal fin in llie torrential species of A^c/iiac/ii'/jii from Western Tibet, a: Ncmachilus gracilis Day X2; b: Nentachilus stolicskae (Steind.). X2J^; c: Nemachilus tcnuicauda (Steind.). X3/^; d: Nemachilus microps (Steind.). X 2}/i.
ing habits. This hypothesis receives sujjport frimi tlic fad tliat in Psi'udapocryplrs luncco- Itiliis, an eel-like burrowing Gobioid fish, the bladder has shifted backwards to the anal region, though in the young stages, when the fish leads a pelagic life, the bladder occupies almost the whole of the abdominal cavity.'
In air-breathing fishes of the families Anabantidac and Aphiccphalidac, the air-bladder extends into the caudal region as far as the base of the caudal fin. The utility of this remark- able modification has been explained l)y me in another place (Hora, Cur. Sci. Ill, p]). 336- 338, 1935). It seems probable, however, that the backward position of the bladder in A^. papillo-IabJatus, N. strauchii and N. acuticcphalus enables them to lie horizontally at the bottom and obviates any tendency of the anterior part to rise. This is merely a tentative sug- gestion as no observations have yet Ijeen made on the mode of life oi these fishes.
'Hora, Rcc. hid. Mus. XXII, p. 646 (1921). 'Hora, Current Science III, p. 336 (1935).
REPORT ON FISHES. PART I : COBITTDAE
303
From the alxive it is clear lliat tl
lie lonii ami structure of the air-liiaddcr is liable to considerable variation and that the modifications noted above are definitely correlated with the diverse types of habitats. In view of these considerations, and also on account of the fact that all possible gradations exist between different types of bladders, I am of the opinion that the character of the air-bladder should not be used for splitting up species into genera. All the same, it is a good index of the type of habitat of a particular species.
Caudal Fin: It is well known that in most of the hill-streani fishes the lower lobe of the caudal fin is distinctly longer and better developed than the upper. A powerful stroke from
Figure 3. Form of caudal fin in the lake inhabiting species of Ncmachilus from Western Tibet, a. Nemachilus vittatus (Heckel). X3; b: Nemachilus panguri, sp. nov. X3; c: Nemachilus hutchinsoni, sp. nov. X; d: Nemachilus deTerrai, sp. nov. X 3.
such a fin would not only result in the forward movement of the fish but the differential dis- placement of water by the two lolx;s would tend to rotate the anterior end of the fish upwards. This is probably advantageous in the case of torrential fishes when darting from rock to rock in shallow rapid-running waters. By the operation of this device the fish actually moves with its head pointing towards the surface and thus obviates encountering any obstructions in its path. In the torrential species of Nemachilus (Text-figure 2), the caudal loljes are either ecjual or the lower is slightly longer than the upper. I'ut in the case of till- free-swimming "lake" species (Text-fig"ure 3), the u]>pi'r Inbc is Idngcr and better develojjed than the lower. This modification no doubt enables these buoyant fishes to go to the hiittiiui more easily and to keep the head-end directed towards the bottom. The dift'erence in the form of the caudal fin is so marked in the species of the two habitats that it is usually easy to separate, with its help, the specimens with a free air-bladder in the al)dominal cavity. Attention may be directed to the modifications of the air-bladder and the structure of the caudal fin in the species of Nciiuithilus obtained by the Netherland Karakorum Expedi-
.>04 REPORT ON FISHES. PART I: COBTTinAE
til 111.'' 'Hic four t'lrrciitial species, N. stolicckac, .\ . i^racilis. W li-iniiaiitilii and A'. iiiiiT(>ps, witli i;Tcatl\- reduced I)ladder ami witli the lower iol)e of the caudal liii better developed than the upper, were found either in small, rapidly flowing streams or in springs and pools to which they resort for breeding purposes. In the specimens of A^. ladaccnsis from Alinazar- Kurghan, the bladder is like that of A^. /'0/>i7/o-/a6i(;/!<j (Text-figure IC) and it is likely that the fi.sh leads a bottom life in still or slowly Howing waters. The two loljes of its caudal tin arc almost symmetrical, suggesting that the lish does not perform any regular vertical movements.
The three species collected from the plains of Turkestan near Varkand were obtained from marshes, lakes or canals and, in consequence, the bladder is considerably modilied. The structure of the bladder of A^. yarkaudcusis is referred to above (]>. 300) ; and that of Neiiiailiiliis sj). prox. tariiiiriisis is similar. Tiie liladder of A'c nun hi I us sp. from Kaliat Utsang is like that of A^ papillo-hibiatus. Jt is thus seen that all the three species of Ne)iia<liiii(s from Turkestan are adapted to live at the bottom in the marshy areas and do not swim alx)ut much. In A', yarkaiidetisis and A^. iariiiicnsis the upper lolx; of the caudal fin is longer as is the case in A'. I'itlatus {ride supra, \). 300), while that of Xriiunliilus sp. is almost symmetrical. It is thus seen that the study of the material obtained by the Nether- land Karakorum Expedition supports the hypotheses advanced regarding the ecology and bionomics of the species collected by the Yale North India I'^xpedition.
GEOGRAPHICAL DISTRIBUTION AND OKICIN OK T UK KISll FAUNA
r have often remarked that the i'lsh fauna of the high altitudes of Central ;\sia is derived from the fauna of the low-lying lands of the neighbouring countries, and this hypothesis is supported 1)\' the geographical distribution of the species and the modilicatioiis undergone by the air-bladder of the forms living in stationary waters at great heights, .\linost all the species of Nciiuichilus found along the slopes of the llimalay.'i are characterised by the great reduction of their air-bladder, but when they enter lakes, etc., a functional bladder is developed once again. The three new .species of the Panggong complex are no doubt descendants of forms once living in torrential streams. When acknowledging the prelimi- nary determinations of the fish collected by the Expedition Mr. liutrhinsoii made the ful- lowing observations regarding the distribution of the new s[)eeies :
"As you will sec from tlie map, one species ( N. hutchinsoni) whieh now seems to occur in small ponds and the very small lake Tsar Tso occui)ies a regitjii formerly filled by the great freshwater lake which represented Panggong in the late glacial and which extended far to the west. The species from Man (N. deTerrai) occurred only in a .small lagoon, cut otT from the edge of the lake. I saw one specimen actually in the lake, almost certainly of this species, and think that it may have been washed out from the lagoon during a rather heavy storm the nighl
before It is quite clear that all fish are extremely rare in lake Panggong
itself at the ])re.sent time. This is due no doubt ])artly to its high salt content and still more to its com])lete lack of higher vegetation which is abiiiidarit in the
"Hora & Mukerji, Visser's Karakorum I, pp. 426-445 (193.S).
'Hora, Rec. hid. Miis. XXIV, p. .S8 (1922) ; Phi!. Trans, h'oy. .Sor. Loudon (P.) CCXVIII, p. 268 (1930) ; Rec. hid. Mus. XXXVI, p. 281 (1934).
bEPORt ON FISHES. PART 1: COBITIDAfi 305
lagoons. No doubt tlie latter are far richer in food stuffs than the lake itself. In the case of the third species ( N. panguri), from Tso Nyak and Pangur Tso, there is no doubt that the s])ecies actually lives at present in the lakes which are ' fairly rich in vegetation, though it also enters the streams running into them. To my mind there can be no doubt that all three species developed in the late glacial lake but only the third one has remained common in lacustrine environments, the others hanging on as best they can chieHy in small pools in the basin."
It is clear from the above that at the present time the Panggong Lake acts as an effective barrier for the distribution of various species. Further it seems probable that this hal)itudi- nal segregation may have induced the development of different species. It is likely that in the late glacial lake there was only one species derived from a torrential stock and that when the environments became restricted, it developed along different lines in different localities and resulted in the production of several new species. ** This supposition supports the hypothesis of Regan^ "that as a rule the first step in the origin of a new species is the for- mation of a community with a new and restricted environment, or with new habits; in other words, that some form of isolation, either localization or habitudinal segregation, is the condition of the development of a new species."
From the modifications of the bladder described above, it does not follow that the lake species cannot enter into brooks or vice versa. All species of Nemachilus are flattened and adapted for clinging to foreign objects and if ponds, pools and lakes are in communica- tion with brooks it is possible that the species uf one habitat may enter the habitat of the other set of species. For instance, it often happens that torrential forms enter springs and pools for breeding purposes. The occurrence of a species (A^. panguri) in both types of habitats, therefore, does not in any way help to fix its evolutionary status.
A'^. x'ittatus seems to have developed in the Kashmir lakes and is endemic in them. Neiiicichiliis sttilicckae, N. iiiicrtipx. N. touiicauda and N. gracilis are widely distributed species. The first three are, however, restricted to the high altituiles, whereas A', gracilis is found as far down in the Indus as Attock.^" Almost all the species were obtained by the Netherland Karakorum Expedition not only from the Nubra Valley but also from the Kara- kash river, which now drains into the Tarim river system. Mukerji and I referred to this discontinuous distributidu of the species, but the difficulty has now disappeared for "On the basis cjf his geomorphological studies. Dr. de Terra has reconstructed the Tertiary drainage pattern of the western part of the Tibetan plateau. A nunilx-r of rivers ran from west to east, one of them occupying the present valley of the Upper Indus."'' The close similarity between the torrential fish fauna of the Karakash river and of western Tibet suggests, at any rate, a common drainage for the waters of these two areas at no great distant date and lends great support to the hypothesis advanced by de Terra. Reference may also i)e made to the occurrence of A', ladacensis in Ladakh and the Karakash \'alley, but it has to be remembered that only a few specimens of this species are known so far, and, in consequence, its specific limits have not yet been precisely defined.
' Dr. A. W. C. T. Herre liad a similar problem in the evolution of the seventeen species of fishes of Lake Lanao in the Philippines {Amer. Nat. XLVII, pp. l.S4-l()2, 1933). •Regan, Nature. CXIII, p. 569 (1924). '"Hora, Rcc. hid. Mus. XXXV, p. 189 (1933). "Hutchinson, Naliire, CXXXIV, p. 87 (1934).
306 REPORT (IN KISHKS. PART T ! rORITIDAE
SYSTEMATIC ACCOUNT
NcmacJiilus stoliczkae (Steindachner) 1866. Cobitis slolicckac, Steindachner, Vcrh. Zool.-bot. Gcs. JVicii, p. 793, pi. xiv, fig. 2. 1868. Ncmachiliis stoliczkae, Giinther, Cat. Fish. Brit. Mus. VII, p. 360. 1876. N eviachcilus stoliczkae, Day (in part), Proc. Zool. Soc. London, p. 795. 1878. Ncmachcilus stolicckae, Day (in part), Sci. Res. 2nd Yarkand Miss. ]chth\oL, p. 14, pi. v,
fig. 2. 1878. Ncuiacheihis stoliczkae, Day (in ])art), I-isli. India, p. 620, pi. civ, fig. 10. 1889. Ncmacliihis stoliczkae, Day (in part), faun. Brit. Ind. Fish. I, p. 235, fig. 84. 1922. Nemachihts stoliczkae, Hora, Kcc. Ind. Mus. XXIV, p. 78. 1935. Nemachilus stoliczkae, Hora & Mukerji, Visser's Karakoruni I, p. 429, ])!. iv, fig. 4.
The species was originally described from 12 specimens obtained from Tsho Mararai (Tso Moriri), a lake in the Rupshu Province of Western Til)et at an altitude of 15,500 ft. Day^' assigned a very wide range of distribution to this species and recorded it from Leh, Snima, Lukong, Cliagra, Yarkand, Sarikol and Akta.sh. Since then it has been reported fmni widely different places in Central Asia and several varieties of it have Ijeen described by Russian ichthyologists" from Eastern Turkestan (Tarim River System) and Tm-kestan (Oxus River System). In 1922, it was pointed out by me that of the large miniber of specimens referred to A^. stoliczkae by Day and nnw preserved in the cnjk-ctinn uf the Indian Museum, only those that came from Rupshu, Lukong and Chagra could Ik; definitely assigned to this species. A specimen from "Kashmir" was also referred to A^ stoliczkae, but there seems to have been some mistake alx)Ut the locality of this example, as the species ap])ears to be restricted to high altitudes. Mukerji and I have recorded this species from Leh, I'anamik, Nungstet, Suget-Karaul and Alenazar-Kurghan after studying the material obtained I)y the Netherland Karakorum Expedition. The first three localities arc on the Iieadwaters of the Indus, while the last two are in the Karakash Valley whence the waters flow into the Tarim River. All these places are situated at fairly high altitudes.
The Yale North India Expedition made collections in Western Tilx!t and ttbtained specimens from several places to the north-east of the type-locality. Though originally descril)ed from a lake," it appears to be a torrential form as it is devoid of a functional air-bladder. It seems likel)- that the species enters lakes for breeding purposes. Several yomig" si)ecimens were obtained by the Expedition liom \'ay(' Tso; wliilo fnllv grown specimens were obtained from several streams.
In view of the great confusion that prevails regarding this species, I take this oppor- tunity to give a detailed description with figures from freshly preserved material.
D.3/8; A.3/5: P. 13; V.8; C.19 Neiinuliiliis stoliczkae is a long and slender s])ecies in which tlir luad and the anterior ])art of the body are depressed; while the tail region is compressed and whip-like. The
"Day, I'rvc. /mil. Sue. Lnmlvii, p. 595 (18/6); .SVi". h't-s. Jml Yail:,nul Mission, hhtliyol., p. 14, pi. v, Hr. 2 (1878).
" Herzensteiii, Wiss. Res. Pr=nmtski Ccntrut As. Kris. /.ool. Ill (>), |i. H (1888); Hcfk, I'oiss ili-s luiii.v Douces dc L'U. R. S. S., pt. ii, p. 559 (1933).
"Mr. Hutchinson informs me that he used a trawl on very favourable ground at the north end (estuary of Pcldo-le stream) of Tso Moriri, and found no fish. He is of opinion that the types of A', stoliccltae must have come from a stream flowing into the lake. (This is stated to be the case in the original description. G. E. H.)
REPORT ON FISHES. PART T: COBITIDAE 307
dorsal profile is gently, luit sli^litly, arclicd ami the ventral |)rolile is straight and hori- zontal throughout. The head is lung and narrow and broadly pointed ; its length is contained from 5.6-5.9 times in the total length and from 4.2-4.8 times in the length without the caudal. The head is relatively longer in the female specimens. The greatest width of the head is contained from 1.4-1.6 times and its height at occiput from 1.7-2.1 times in its length. The eye is almost in the middle of the head in female specimens, while in the males the snout is sometimes considerably longer than the postorbital part of the head. The diameter of the eye is contained from 5.2-6.5 times in the length of the head, from 2.3-3.2 times in the length of the snout and from 1.3-1.6 times in the interorbital width. The supraorbital margin of the eye projects slightly beyond the profile and the eyes arc not visible from the ventral surface. The mouth is on the ventral surface considerably behind the tip of the snout; it is lunate and horizontal. The lips are thick, continuous and greatly papillated or striated. The posterior lip is reflected backwards so that a portion of the jaw is left bare. The post-laliial groove is interrupted in the middle by a slight ridge. The posterior jaw has a sharp, evenly rounded edge. The form of the lips is a very characteristic feature (jf the species. The barbels are short and stumj)y; they are as long as or slightly longer than the diameter of the eye.
The greatest height of the body is above the pectoral fins; the dejjth of the body is con- tained from 8.2-10.4 times in the total length and from 6.8-8.6 times in the length without the caudal. The body is scaleless. The lateral line is complete; anteriorly it is continued over the head and divides into two branches Ijehind the eyes. The caudal i)eduncle is long and narrow; its least height is contained from 3.1-3.6 times in its length.
The dorsal fin is inserted somewhat in advance of the ventral and its commencement is distinctly nearer to the base of caudal than to the tip of snout. The longest ray of the dor.sal is considerably higher than the depth of the lK)dy below it; its anterior margin is rounded near the ti]) and the free border is concave. The paired fins are broad, rounded and horizontally ])laeed. The jjectoral is somewhat shorter than the head and extends almost half the way to the ventral. The ventral fin extends beyond the anal opening and in some cases almost reaches the anal fin. The anal fin is separated from the caudal by a distance equal to its own length. The caudal fin is almost as long as or slightly longer than the head in males, while in the females it is .shorter th.m the head; its length is contained from 5-5.8 times in the total length. It is slightly emarginate and has two nnnnled lobes; the lower lol)e is Ix-tter develojied and longer than the U])per.
Nemachilus stolicdcac exhibits sexual dimorphism. The secondary sexual characters of the male are well developed and of the tyjie described l)y nie^''^' for A', tibctanus. In the males the head is relatively shorter and the snout is longer than the postorbital part of the head.
In si)irit specimens, the general colour is dark above and on the sides and nuich lighter l)elow. The head and bo<ly are mottled with numerous black spots, and in some specimens short, .saddle-.sha|)e(l, black bands are distinguishable along the dorsal surface, especially in the tail region. The dorsal and the candrd tins are sjjotted. The anteriormost ray of the dorsal fin is provided with ;i series of consjjicuous sjxits. The dorsal surface of the outer rays ui the paired fins is sometimes spotted.
In young specimens, the whole of the body is grayish in eolotu", though somewhat
■"Hora, Rcc. hid. Mus. XXIV, p. 81 (1922).
,l08 REI'ORT ON FISHF.S. PART I: rnBITIDAE
lii^litt-r 1*41 tlic ventral surface. 'I'liere is a series of spots aloni; tlie lateral line and also along tile dorsal surface. The fms are witlumt any colour niarkins.;s.
Pistribiitioii. Reference has been made above to the .general distributimi nf the s])ccies. Its precise range is, however, difficult to assign till the limits of the forms referred to .V. stolicchae from all over Central Asia are properly elucidated. The \'ale North India ICxpedition obtained specimens from the following localities in June-August, \'^^32:
Between Tangtse and Mugleb, ca 13,700 ft. (L37). 1 .specimen (S).
Mig])al-kongma, ca 16,082 ft. (L64). 5 specimens (i).
Between Chuine-sang and Nyagtsu, ca 15,500 ft. (L()5). 4 specimens (39 +13).
Nyagtsn, ca 15.324 ft. (L65). 1 s]K'cimen (S).
Tso-skani, ca 15.800 ft. (L77b). 1 specimen (9).
Yaye Tso, ca 15,373 ft. (L78). 18 specimens (young).
Rciimrks. A'cDiachiliis stolicskar can be readily distinguished by the fullowing combina- tion of characters :
(i) The \entrals extend considerably bexcuid the anal opening.
(ii) The comiiieiiceiiicut of the dorsal is nearer {<> the base of the caudal than to the tip of the snout.
(iii) The least height of the caudal jieduiicle is abnut 3-4 times in its length.
(iv) The lijis are papillated and conliiUK ms ; the posterior li|) is broad and rellecled backwards.
Bionomics, b'roni its general buikl, position and form of the jjaired fms and the struc- ture of the lips and jaws, N. stolicchac appears to l)e a torrential species. The absence of a functional swim-bladder indicates that it is a stream form and li\i's at the bottom. An examination of the stomach contents has shown that it feeds on insect larvae and algal growths that encrust rocks and stones. Caddis- worms, dipterous larvae, eggs (probably of Trichoptera) and slimy matter have been found in the stomachs of specimens dissected from dififerent localities. The length of the alimentary canal is slightly greater than the length of the fish. Some of tlu- s]ieciniens opened ha\e been found to harbour wdrnis.
M casiirciiwnls in iiiilliiiuircs
6 £ S 9 9 9
Total length including caudal 121 .0 106.0 100.0 102.5 96.9 84.5
Length of caudal 21 .2 18.S 17.0 18.0 16.2 14.8
Length of head 21.0 18.0 17.8 18.5 19.0 16.4
Width of head 15.0 11.5 11.3 12.5 12.1 9.3
Height of head 11.8 9.2 9.8 10.4 9.0 7.8
Depth of body 14.0 10.3 10.7 9.8 10.2 10.2
Lengdi of snout 10.0 8.8 7.8 7.6 8.2 7.0
Diameter of eye 4.0 2.9 2.8 3.0 . 3.0 2.8
liiterorbital width 5.5 4.0 4.0 3.9 5.0 3.6
Length of caudal peduncle 23.6 20.0 20.0 21.0 20.0 18.0
Least height of caudal peduncle 7.0 6.4 5.5 6.2 5.8 5.0
Longest ray of dorsal 18.1) 15.1 14.0 15.0 14,8 11.0
Longest ray of anal 1 5.5 1 3.6 12.0 1 4.0 1 4.0 10.5
Length of pectoral 18.5 16.0 14.5 1 5.8 15.0 12.0
Length of ventral 15.6 14.5 13.0 12.5 13.0 11.0
L66 L64 L6S
REPORT ON FISHES. PART 1 : COBITIDAE 309
Ncmachiliis gracilis Day
1876. Neiuachcilus gracilis, Day, Proc. Zool. Soc. London, p. 798.
1878. Neiuachcilus gracilis, Day, Sci. Res. 2nd Yarkand Miss. Ichthyology, p. 16, pi. iv, fig. 5.
1878. Nemacheilus gracilis, Day, Fish. India, p. 621.
1889. Ncmachilus gracilis. Day, Faun. Brit. hid. Fisli. I, p. 257.
1898. Neniachilus stoUczkae, Alcock {ncc Steindachner), Rep. Nat. Hist. Pamir Bound. Comui.,
p. 38.
1922. Ncmachilus gracilis, Hora, Rec. Ind. Mus. XXIV, p. 74.
1933. Ncmachilus gracilis, Hora, Rcc. Ind. Mus. XXXV, p. 189.
1935. Ncmachilus gracilis, Hora & Mukerji, in \'isser's Karakornni, I. p. 430, pi. iv, fig. 2.
Ncmachilus gracilis appears to be one of the commonest loach of the Indus River and its range extends from very high altitudes to as low down as Attock in the North-Western Frontier Province. Few specimens of the species were also obtained by tlie Netherland Karakorum Expedition from the Karakash Valley. In the collection of the Yale North India Expedition, A^. gracilis is represented from the following localities. The specimens were collected during May to July, 1932.
Stream 1 mile of IJras, ca 10,100 ft. (K76). 6 specimens (young).
Dras, (-(( 10,144 ft. (K 77). 22 specimens (9 3 — 13 9 ).
Spring below Kargil, ca 8,790 ft. (K81). 1 specimen ( <5 ).
Above Leh, ca 15,000 ft. (L25). 3 specimens (young).
Between Tangt.se and Mugleh, ca 13,700 ft. (L37). 1 specimen ( S ).
Kyam rivulet, ca 15,500 ft. (L59). 1 specimen (young).
Kyam, a pool below camp, ca 15,500 ft. (L60). 1 specimen { S )■
Yalapuk, ca 13,521 ft. (L79). 2 specimens (young).
Sta-rtsak-puk Tso, ca 1-1,889 ft. 4 specimens (young).
The above distribution shows that the species frecjuents ptiols in the course of streams, springs and lakes for breeding purposes as young specimens were collected from such localities. It is essentially a torrential species of wide range.
N. gracilis is readily distinguisiied by the fact that, as a rule, the ventrals do not extend as far as the anal opening, the eye is almost in the middle of the head and the ventrals cnnnnence in advance of the dorsal. The structure nf the biwer lip is also characteristic of the species.
The air-bladder is of the usual reduced type, consisting of two lateral chambers enclosed in bone. The alimentary canal is simple and not much convoluted; its length is about three- fifths of the total length of the fish. The food consists of insect larvae, mostly free-living Dijitera and Trichi>])tera, and of the slime encrusting rocks and stones in rajjid current. The small, fan-shaped, horizimtal ])aired fins, reduced air-bladder and its food .strongly suggest that the lish lives in very fast currents. Young specimens were collected from underneath slimes in a ris'ulet.
Ill the mature females, the ovaries occu])y almost the whole of the abdonu'nal cavity and even the alimentary canal is flattened out. The eggs are of a fairly large size (diameter about 1.25 mm.).
310 REPORT ox 1I>HI.S. I'ART I : CORTTIOAE
V NciiHichiliis niicrops (^Stcindachner)
1866. Cobilis microps, Stciiuladiner, Fcr/i. Zool.-bot. Gcs. Jl'irii, XVI, p. 794, pi. xii, fig. 3. 1868. Nciiiachiltis viicrops, Giinther, Vat. Fish. Brit. Mtis. VII, p. 357. 1878. Ncmachihts microps. Day, Sci. Res. 2nd Yarkand Miss., Ichthyology, \\ 17. 1922. Newachiliis microps, Hora, Rcc. Ind. Mits. XXIV, p. SO. 1935. A^cwachilus microps, Hora & Mukerji, in Visscr's Karakoniiii, I, p. 430, pi. iv, fig. 3.
I refer to Ncmachilus microps 4 specimens, from 30 to 86 mm. in total length, collected by the Yale North India Expedition on the i/th of August 1932 from underneath large stones in the bed of a stream flowing into the west end of the Tso-Moriri lake about 14,853 feet above sea level. The two larger specimens are females with fully developed ovaries. The eggs are minute and the ovaries do not extend forwards l)eyond the midille of the abdominal cavity. The air-bladder is reduced and enclosed in Ixmy capsules, as is char- acteristic of the stream-dwelling forms. The alimentary canal is about as long as the length of the fish and the stomach contents show tliat the fish feeds on white, slimy stuff tliat is found encrusting rocks and stones.
In my key to the species of NciiiacliUiis from Central .\sia in tjie collection of llu' Indian Museum (1922, p. 7i) N. viicrops was scfjarated fmrn the 9 of A^. yasinciisis by the rela- tive lengths of the anal fin and of the caudal ]KHluncIe. in tlie specimens now before nie the portion of the caudal peduncle is much less than the iengtli of the anal lin, l)ut in all other respects they agree with tlie other specimens in tlic Indian Museum collection. In A^. yasiiiciisis the caudal peduncle is low, while in the four specimens from Western Tiljet it is two-fifths as high as long. In the earlier s])ccimens, the caudal peduncle is one-third as high as long. These differences do not seem to me suflicient to justify the erection of a new species in such a variable genus. For future reference, however, 1 give below measurements of two mature female specivnens.
The species was originally described from 10 specimens, 4 from I.eli and (> from "I'hirse- Bach in einer llohc von circa 16000 Fuss bei .Manecban in Rupsbu i juli 1(%5)." The specimens in the collection of the Yale North Intlia ]'".xpe(lition were also collected in the Rupshu Province, Western Tibet.
.]Icasurciiiciils in millimclrcs
2 9
Total length including caudal 86.0 86.0
Length of caudal 1 5.6 1 5.5
Length of bead 15.0 16.0
Width of head 9.5 11.2
Height of head 7.5 7.0
Depth of hotly 8.8 9.5
Length of snout 5.5 6.2
Diameter of eye 2.8 2.6
Interorbital width 3.8 3.8
Length of caudal ])eduncle 13.2 12.8
Least height of caudal peduncle 5.3 5.0
Longest ray of dor.sal 1 1.6 12.5
Longest ray of anal 10.5 10.0
Length of pectoral 12.2 12.5
Length of ventral 1 1.5 1 1.0
REPORT ON F1?HES. PART i: COBITIDAE 311
Nciiiacliiliis tcniticaiida (Steindachner )
1866. Cobitis tciuiicauda, Steimlachner, I'crli. ZattL-hot. Gcs. IJ'icii. W'l. p. 792, pi. .wii, fig. 3.
1868. Ah^iinn-hiliis tcmiiamda, Giiiither, Cat. Fish. Brit. Miis. VII, p. 357.
1922. Neutachihts tcnuicauda, Hora, Rcc. Ind. Mtts. XXIV, p. 79.
1935. Nemachilus tcnuicauda, Hora & Miikerji, in Visser'.s Karakoniiii, I, \). 430.
There is a single, mature, female specimen of Ne)itachilHS tcnuicauda, alx)ut 62 mm. in total length ; it was collected by the Expedition from a pool in a swamp by Sta-rtsak-i)uk Tso at an altitude of 14,885 feet. It is a small species and was originally collected from a small brill ik in Western Tibet. It is also known from Leh and the Nubra Valley.
The ovaries occupy only the posterior half of the abdominal cavity. The air-bladder is reduced and enclosed in two bony capsules. The length of the alimentary canal is about seven-tenths of the total length of the fish. The food consists of Dipterous and Trichoj)- terous larvae and of insect eggs. The long and narrow caudal peduncle indicates that the species lives in turbulent waters.^"
A'Ciinicliilits 7'ittiifiis ( Meckel )
1838. Cobitis vittata, Ilcckel, Fischc Kaschni., p. 80, pi. xii, figs. 3 and 4. 1844. Cobitis vittata, Heckel, in Hugel's Kashmir IV, p. 382, fig. 1922. Nemachilus vittatus, Hora, Rcc. Ind. Mits. XXIV, p. 74.
1930. NcmachUus vittatus, Hora, Journ. Bombay- Nat. Hist. Soc. XXXIV, p. 379 (air-bladder structure).
Nemachilus vittatus is represented by 40 speiimens in the cullection of the Yale North India Expeditinn; of these 33 are from the Wular Lake and 7 from a small lake at Shad- ipur. The Wular Lake specimens were dredged at Kiuhiuis. Though a lake form, N. vittatus appears tn be a bottom-living species and, in cunsequence, is fievnid of a free air-bladder in the abdominal cavity. The two lateral chambers are large and lie next to the skin as the Ixiny capsule is incomplete in that region. The alimentary canal is four- fifths of the total length of the fish. The food consists of algae, mud, insect larvae, leeches, etc.
The species exhibits well marked sexual dimorphism and the secondary sexual char- acters of the males are similar to those of the other species described in this paper. The gonads were ripe towards the end of April when the specimens were collected. The ovaries occupy almost the whole of the abdominal cavity and the eggs are of a relatively large size.
Of the 40 specimens obtained by the Expedition, 24 are females and 16 are males, giving a percentage of 60 females and 40 males. L^sually the males predominate in collec- tions as the females are of rather secretive habits, but in this case the dredge used seems to have made a considerabfe difference in the projiortional representation of sexes in the collection.
Nemachilus deTerrai, sp. nov.
D./38; A.2/5; P.IO; V.7; C.16
Nemachilus deTerrai is a long and slender species in which the head and the anterior part of the liody are slightly depressed, while the posterior part, especially the tail region, is compressed and whip-like. The head is moderately long and broadly pointed ; its length
"Hora, Phil Trans. Roy. Soc. London (B) CCXVIII, pp. 250-25-4 (1930).
312
REPORT ON FISHES. PART I : COlilTIDAE
is contained from 5.7-6.2 times in the total Icns^lh and from 4.6-5 times in the length without the caudal. The width of the head is contained from 1.4-1.8 times and the height of the head from l.()-l.S times in the length of the Iiead. iiie eye is situated somewhat nearer
Figure 4. Lateral vitu of a female spcciiiieii of Nemachilus deTerrai, sp. iiuv. X Ij^-
the tip of the .-^noul tliaii to tlie o])eri'ular margin; its superior horder |iiMJeets slightly heyond tile dorsal profile of tiic head, imt it is slightly visible I'mni llie \eiitral surface. The diam-
<Z.
Figure S. Alimeiilary canal and air-bladder nf a male ."ipeciiiieii of Nemachilus deTerrai, sp. nov. X 2J^.
a : alimentary canal ; b : air-bladder.
eter of the eye is contained from 4-4.7 times in the lengtii of tiir lieail. Iri'in 1.4-l.S times in the length of the snout and from 1-1.4 times in the interorhital width. The mouth is lunate and transverse; it is situated on the ventral surface somewhat behind the tip of the snout and is bordered by fleshy lips. The lips are striated; the posterior lip is interrupted in the middle and reflected towards the sides so that a small, triangular portion of the
REPORT ON FISHES. PART I : COBITIDAE 313
posterior jaw is left bare. The posterior jaw is sharp and shovcl-lil^e and the anterior jaw lies as a Imod in front of it. The harlx-ls are thin and long; tlie inner rostrais are as long as the diameter of tlie eye while the other two pairs are mnch longer.
Some of the specimens are heavily parasitised l)y worms, so that the depth of the body is liable to considerable variation. The depth of the body is contained from 8.5-9.9 times in the total length and from 7-7.6 times in the length without the caudal. Behind the gill- opening and above the base of the pectoral fin, the lateral line is represented by a thin-walled, broad tube beyond which it is faintly marked, thriugh it is continued to the base of the caudal fm. The caudal peduncle is long and narrow; its least height is contained from 4.8-6.4 times in its length. In the male specimens the least height is either erjual to or greater than the diameter of the eye while in the females it is considerably less.
The dorsal fin is inserted slightly in advance of the ventrals and its commencement is considerably nearer to the tip of the snout than to the base of the caudal; it is longer than the head ; its posterior edge is truncate or slightly crenulate. The paired fins are horizontally placed ; the pectoral fin is broad but pointed in the middle ; it is somewhat shorter than the head and separated from the ventral by a distance equal to half of its length. The ventrals e.xtend beyond the anal opening and in some cases even beyond the commencement of the anal fin which e.xtends about half the way to the base of the caudal. The caudal fin is longer than the head; its posterior border is concave with the upper rays considerably longer than the lower.
Nemachilus deTerrai exhibits sexual dimorphism. The secondary sexual characters of the male are similar to those described above for N. stoliczkac. The difference in the height of caudal peduncle is also well marked in the two sexes.
Air-bladder: The air-bladder is divided into two parts, (i) the anterior part consisting of two round, lateral chambers enclosed in b(jny capsules and connected by a short, trans- verse tube, and (ii) a large posterior part lying free in the abdonn'nal cavity and connected with the trans\erse tube bv a short tul)e. By another short, liut broader, tube it is connected with the oesophagus. The posterior part is slightly constricted in the middle so that it con- sists of two chambers. In a specimen about 110 mm. in total length, the measurements of the bladder are as follows :
Total length of bladder 25.00 mm.
Length of posterior part 20.00 mm.
Width of anterior part 7.50 mm.
Width of ])osterior part 7.50 mm.
Transverse diameter of each anterior chamber .... 3.25 mm.
Length of tube between two anterior chamljcrs .... 1.00 mm
Length of tul)e between anterior and posterior ])arts 1.70 mm. The ai)()ve measurements are of the bladder after its removal from the bony capsules.
The bony cajisules of tlu' air-liladder lie just I)eneath the skin and are distinctly visible from the external surface.
In spirit specimens the general colour of the body is pale-olivaceous. There is usually a lilack, fairly broad streak along the lateral line which is composed of a series of darker blotches. In some the dorsal surface is gray so that there is a lighter stripe between the dorsal band and the lateral line. The dorsal and the caudal lins are provided
314 REPORT (IN FISHKS. TAKT i: COKITIDAE
witli 2 tji 4 series of spots ami the anterior ray >>i tlie (lorsal fin is provided witli 3 nr 4 l)iaci< splits aliinj^ the front niarj^in. Tlie dorsal surface of tlie paired fins and thi- anal iin are sometimes jirovided willi Mack jialches.
Localitv: Nine specimens of N. deTerrai were cilitaiiicd l>y the ^■ale North India Ivxpeditioii from the Man I.a^oon on the 4tii and 3th of July. It is an isolated !a.L;oon in the drowned valley at an altitude of 14,008 ft.
Rt'iiuirks: The most distinijuishing- feature of N. deTerrai is the iL^real len.s;th of its dorsal fin. The other fins are also eloni^atcd. 1"he form of the caudal Iin is very charac- teristic of the species.
Bionomics: From the extensive air-hladder in the abdominal cavity, and from the nature of the fins, it is clear that the fisli is adajited to live in stationary waters. The gen- eral facies, especially the \\hi])-like caudal peduncle, suggests that the fish is a fast swimmer. For feeding purposes, the fish proliahly adheres to rocks with the help of the jjaired fins and .scrapes off animal and vegetable matter. In the ca.se of two .specimens di.s.sected the stomach was found to be full of a whitish, pidi)y material \vithout any sand or small bits of stones. The alimentary canal is not much convoluted: its length is about three-fifths of the total length. It wiiuld thus seem to be a flesh-eating species.
Measurements in niilinnetrcs
$$$99
Total lenglh including caudal 104.0 95.0 95.0 83.0 68.0
Length of caudal 21.0 18.5 19.0 14.8 11.5
Length of head 17.9 16.5 15.2 14.0 12.0
Width of head 11.5 9.2 8.8 9.8 7.0
I leight of head .' 10.') 10.0 9.5 8.2 6.5
Depth of body 12.0 10.5 9.0 12.0'^ 8.0
Length of snout 6.5 6.5 6.5 5.0 4.5
Diameter of eye 4.2 3.5 3.8 3.5 2.8
Intennbital width 5.2 5.0 4.0 3.5 3.2
Length of caudal peduncle 23.0 ■ 21.6 20.0 20.5 16.0
Least height of caudal peduncle 4.8 4.3 4.0 3.2 2.5
Longest ray of dorsal 19.8 17.8 17.0 15.0 14.2
Longest ray of anal 15.0 14.0 13.2 12.5 9.0
Length of pectoral 17.2 15.0 15.0 13.2 11.5
Length of ventral 15.5 13.8 14.0 1 1.5 10.0
Nemachilus hutchinsoni, sp. nov.
D.3/8; y\.2/5; P.9; V.7; C.16
In Nemachilus hutchinsoni the head and the anterior ])art of the body are depressed so that the ventral surface is somewhat flattened. In the tail region the body is compressed and whip-like, d'he head is short, high and broadly pointed ; its length is contained from 5.4-5.6 times in the total length and from 4.4-4.6 times in the length without the caudal. The width of the head is contained from 1.4-1.6 times and the height of the head at the occiput 1.6 times in its length. The eyes are situated nearer to the tip of the snout than to the posterior
" The abdominal portion is greatly swollen due to heavy parasitisation by worms.
REPORT ON FISHES. TART I : COBITIDAE " 315
margin of the ojjerculuni ; tliey are dorsolateral in position and invisible from the ventral sur- face. The diameter of the eye is contained from 4.1-4.9 times in tlie lens^th of the head, from 1.5-1.7 times in the length of the snout and from 1-1.5 times in the interorbital width. The month is lunate and transverse; it is on the ventral surface not very far behind the tip of the snout and is bortlered by ileshy lips which are continuous at the angles of the mouth. The ventral lip is divided in the middle almost imperceptibly. The lips are folded, fimbriated and covered with minute papillae. The lower jaw is sharp and liorizontal, while the upper jaw is vertical and lies in front of the lower. The barbels are thin and long; the inner rostrals arc almost as long as the diameter of the eye, while the other two pairs are much longer. The only mature female specimen is heavily parasitised and, in conserjuence, the depth of its body is relatively greater. In the male specimens the depth of the body is contained from 7.7-8 tiiues in the total length and from 6.2-0.6 times in the length without the caudal. The lateral line is well developed above the base of the pectoral fin beyond which it is incon-
Figure 6. Lateral view (if a female .specimen (jf Nemachilus hutchinsoni, sp. iiov. Nat. .size.
spicuous. The caudal peduncle is long but fleshy; its least height is contained from 4.4-5.1 times in its length. The least height is either greater than or etpial to the diameter of the eye; in the female specimen the least height is considerably greater than the diameter of the eye.
The dorsal liu is inserted in advance of the ventrals and its coiumencement is consid- erably nearer tn the ti[) of the snout than to the base of the caudal fin; it is somewhat longer than the head but this character is more marked in the female specimen. The posterior margin of the fin is almost truncate. The paired fins are horizontally placed and are broadly jiointed in the middle. The pectoral is shorter than the head and is separated from the ventral by a considerable distance. The ventral extends beyond the anal opening and almost reaches the anal fin which extends half way to the base of the caudal fin. The caudal fin is somewhat longer than the head ; it is slightly emarginate with the two lobes broadly rounded. The upper lobe is better developed and longer than the lower.
Nemachilus hutchinsoni exhibits sexual dimorphism and the secondary sexual charac- ters of the male are similar to those of the other species discussed here. The mature male and female specimens are from two different localities so one cannot be certain that they beldug to the same species. A male specimen has been selected as the type of the species. .\ttention luay 1^ directed to the fact that in the female specimen the caudal peduncle is relatively deeper and the dorsal fin longer than is the case in the males.
Air-bladder. The air-bladder is of the usual Diplophysid tyjie. The anterior ])art is dumbbell-shaped and is enclosed in two bony capsules \\hile the posterior part, which is deeply
ol6
KKl'OKT ON FISHES. PART I : CUBITIDAE
6.
a..
Figure 7. Alimentary canal and air-bladder of a male specimen of Nemachilus hutchinsoni, sp. nov. X 4.
a: alimentary canal ; h: air-bladder.
cdiistricted to form two cliainl)ers, lies free in tlic alidnniinal cavity. The two anterior chambers are connected by a short tube and the anterior and jjosterior parts of tlie bladder are connected by a short tube. In a male specimen about 90 mm. in total length the measurements of the bladder are as follows :
Total lcn!,nli of bladder 21.60 mm.
Length of posterior part 18.40 mm.
Width of anterior ])art 7.00 mm.
Width of ])ostcrior ])art 4.80 mm.
Transverse diameter of each anterior chamber .... 3.20 mm.
I^ength of tube between two anterior chambers .... 0.66 mm.
Length of tube between two parts of bladder 0.90 mm.
REPORT ON FISHES. PART i: COBITIDAE 317
The above measurements are of the bladder after its removal from the Ixmy capsules.
The bony capsules are incomplete in the part where they touch the skin so that their position can be readily made out from the external surface.
In spirit specimens the general colour of the body is pale-olivaceous. A series of fairly broad blotches is present along the lateral line and in some specimens they unite to form a longitudinal band. Along the dorsal surface, especially in the tail region, there are a number of saddle-shaped bands. The dorsal surface and the sides are further irrorated with small black dots. The dorsal and the caudal fins are provided with two to three broad bands. The anal and the ventral fins are provided with one or two I)ands each.
Localities: In all six specimens were collected by the Yale North India Expedition during June 1932 from the following localities :
About 3 miles west of Mugleb, ca. 13,525 ft. (L3S). 1 specimen (young). Pond between Durlnik and Tangtse, ca. 13,000 ft. (L36). 3 specimens (S ). Tsar Tso, ca. 13,950 ft. (L39). 1 specimen (young). Pool isolated from the river at Lukung, ca. 14,164 ft. (L40). 1 specimen ( 9 ).
It is seen from the alxjve that the species occurs in pools, and small lakes.
Bioiioiiiics: The species is adapted for life in stationary waters of lakes where its well developed air-liladder enal)les it to swim about freely at different depths. It feeds on insect larvae and pupae that encrust rocks and stones. Its horizontal, paired fins enable it to adhere to rocks and its lower jaw appears to lae capable of acting as a shovel for rasping off encrusting organisms. The alimentary canal is a simple tube without many convi)lutioiis and its length is about three-fifths of the total length. The eggs are small and the o\'aries extend right up U> the anterior end of the abdominal cavity.
Measurcvicnts in niiHiinclrcs
Total length including caudal . .
Length of caudal
Length of head
Width of head
Height of head
Depth of body
Length of snout
Diameter of eye
Interorbital width ^
Length of caudal peduncle . . . . Least height of caudal peduncle
Longest ray of dorsal
Longest ray of anal
Length of jjcctoral
Length of ventral
|
2 |
S |
S |
$ |
|
110.0 |
94.0 |
90.0 |
89.0 |
|
20.5 |
18.5 |
17.0 |
15.5 |
|
19.5 |
17.0 |
16.5 |
16.0 |
|
12.2 |
11.0 |
11.0 |
11.0 |
|
11.9 |
10.5 |
10.5 |
10.0 |
|
17.0 |
12.2 |
11.5 |
11.0 |
|
6.9 |
6.5 |
6.0 |
6.0 |
|
4.0 |
4.0 |
3.6 |
3.6 |
|
5.0 |
4.2 |
4.0 |
5.5 |
|
22.0 |
20.5 |
19.5 |
18.0 |
|
5.0 |
4.0 |
4.0 |
3.6 |
|
21.5 |
17.0 |
16.0 |
17.5 |
|
16.5 |
12.0 |
12.5 |
12.6 |
|
17.5 |
15.5 |
15.0 |
15.0 |
|
15.5 |
13.2 |
13.2 |
13.5 |
L 40 L 36
318
REPORT (IN FISHKS. PART 1: rdlUTIDAE
Nemachilus panguri, sp. nov.
D.3/8; A.2/5; \\9; \'.7; C'.U.
The liuilil of Nemachilus panguri is mure or less nf tlie same type as in the twu pre- ceding species. i'lie iiead ami the anterior ])art •<( tlie IhkIv are sumewliat deprcsseil wliile tlie tail region is siigiitly compressed and wiiip-likc. The head is moderately long and broadly pointed anteriorly; its length is contained from 5-5.6 times in the total length and from 4-4.6 times in the length of the caudal. Tlie width of the head is contained from 1.78-1.95 times and the heiglit of tlie head from 1.73-1.7() times in its length. The jiosition of the eye in the length of the head is varialile: the upjier margin of the orbit is slightly raised above the dorsal profile of the head and the eyes are not visible from the ventral sur- face. The diameter of the eye is contained from 4-5 times in the length of the head, from 1.5-2 times in the length of the snout and frnm 1-1.1 times in the interorbital width. The mouth is small, lunate, transverse and horizontal; it is situated on the ventral sm^face slightK'
Figure 8. Lateral view uf a female siiecimen nf Nemachilus panguri, sp. iiuv. fruiii Tso Nyak. X 1^2.
behind the tip of the snout and is bordered by llesiiy and papillated li])s. i'he lower lip is interrupted in the middle. The posterior iaw is sharp, truncate and horizontal. Ibe three ])airs of barl)els are fairly well de\elo]ied : the inner mstrals are as long as or slightly longer than the diameter of the eye while the other two ])airs are much longer.
The depth of the body is cf)ntained from 9-10.5 times in the total leui^th and from 7.3-8.6 times in the length witliout the caudal, liehind the gill-opening and abo\e the base <if the pectoral tin, the lateral line is represented In- a thin-walled, broad tulx; beyond whicli it is faintl}' marked to the l)ase of the caudal flu. The caudal ]icdunclc is long and narrow; its least height is contained from 5.3-6.5 times in its length. In both the sexes the least height of the caudal peduiK'le is usualh' le>s ilian the diameter ol the eye, but in some female specimens it is greater than the diameter of the eye.
The dorsal fin is inserted slighth' in advance of the vcntrals and its coniuKiuement is either equidistant l>etween the tip of the snout and the base of the caudal or nearer to the tip of the snout than to the base of the caudal; it is almost as long as the head; its jiosterior border is slightly arched. The ])aired fins arc broad and horizontal ; the pectorals arc ])ointed in the middle, especially in the males. The pectoral fin is considerably shorter than the head and is .separated from the ventral by a distance almost equal to half of its length. The ven- trals extend beyond the anal ojjcning and reach the base of the anal fin. The anal fin is similar in shape to the dorsal. The caudal fin is almost as long as the head; it is slightly eniarginate with the upper lobe considerably longer and better developed than the lower.
REPORT ON FISHES. PART I : COBITIDAE
319
6.
\
a
Figure 9. Alimentary canal and air-bladder of Nemachilus panguri, sp. nov. X 5. a: alimentary canal of a
male specimen ; h : air-bladder of a female specimen.
Nemachilus panguri exhibits well-marked sexual dimorphism. The secondary sexual characters of the male are similar to those of other Nemachiloid fi.shes of Central y\sia.
.■iir-b!a<Idcr: The air-bladder of N. panguri is similar to that of N. deTerrai. In a female specimen about 95 mm. in total length, the measurements of its various parts were as follows :
Total length of l)l;ul(ler 26.6 mm.
Length of jjosterior jiart 19.0 mm.
Width of anterior part 6.7 mm.
Width of posteri<ir ])art 5.8 mm.
Transverse diami-ter of each anterior chamber .1.0 mm.
Lenj^th of tube between two anterior chambers 0.9 mm.
Length of tube bi-lween anterior and posterior parts uf bladder 1.4 mm.
The above nieasm-ements are of the bl:idder after its removal from the bony capsules.
320 REPORT ON FISHES. PART I : COBITIDAE
Tliejx5ny capsules of tlie air-bladder lie just Ijeneath the skin and are distinctly visible Iruiii outside.
The ground color of the spirit specimens is pale-brown, the dorsal surface lieing some- what darker than the ventral. There are jiatches of dark culnur along the lateral line and saddle-shaped l)anils along the dorsal surface. These colour markings are more pronounced in the younger .specimens. The head is grayish above and pale-yellow Ixdow. The dorsal fin is jirox'ided with 4-5 dark bands in the adult while in the smaller individuals there may be only one or two bands. Ihe \entral and the anal iins are also siniilarb' niarketl. The caudal fin is ])rovided with 3 broad Ijands, Init in young specimens only one broad, prominent band is present in the middle of the fin.
Localities: Several specimens in N. panguri were collected by the ^'ale North India Expedition in August 1932 from the fLillowing localities:
I'angur Tso. n;. 14.203 ft. (L74). Several )i)ung, half-grown an. 1 adult specimens. Tso Nyak (L 71a). Several young, half-grown and adult specimens.
Remarks: Nemachilus panguri seems to be a very close ally of N. deTerrai from which it differs in i)r<i])ortions, esi)ccially of the eye and the dorsal fin. Though the differ- ences do not seem to be markedly .specific, it is better to regard the two .species as distinct in the present state of our knowledge of the fish fauna of Western Tiljet.
Bionoiiiics: Like the two preceding species, N. panguri is also adapted to live in sta- tionary waters where it can dart froni place to ])lace with the help of the whip-like caudal peduncle or make vertical movements with the help of the large swim-bladder. The stomach contents of a male specimen consisted of Chironomid larvae with their sandy and calcareous cases. The alimentary canal is only slightly convolute; its length being seven-tenths of the total length of the fish.
Mcasiirciiioils in iiiilliiitctrrs
Total length including caudal . .
Length of caudal
Length of head
Width of head
Height of head
Depth of body
Length of snout
Diameter of eye
Interorbital width
Length of caudal peduncle . . . . Least height of caudal peduncle
Longest ray of dorsal
Longest ray of anal
Length of pectoral
Length of ventral
|
£ |
S |
S |
9 |
|
73.0 |
84.0 |
98.0 |
58.0 |
|
12.6 |
15.5 |
18.6 |
11.0 |
|
13.0 |
16.0 |
19.6 |
11.5 |
|
7.1 |
8.2 |
11.0 |
6.2 |
|
7.5 |
9.1 |
11.5 |
6.5 |
|
7.4 |
8.0 |
10.9 |
5.8 |
|
4.8 |
5.9 |
8.0 |
3.8 |
|
3.2 |
3.2 |
4.0 |
2.5 |
|
3.5 |
3.5 |
4.2 |
2.5 |
|
15.8 |
18.0 |
20.2 |
13.0 |
|
2.5 |
3.0 |
3.8 |
2.0 |
|
14.0 |
16.1 |
19.2 |
11.0 |
|
10.5 |
12.0 |
12.5 |
12.2 |
|
12.9 |
14.5 |
14.5 |
9.8 |
|
11.2 |
12.9 |
14.1 |
7.8 |
REPORT ON FISHES. PART I: COBITIDAE 321
Botia birdi Chaudhuri
1909. Botia birdi, Chaudhuri, Rcc. hid. Miis. Ill, p. 339. 1922. Both birdi, Hora, Rec. Ind. Mies. XXIV, p. 319.
The Yale North India Expedition collected a dozen specimens of Botia birdi at Srin- agar during March 1932. The specimens vary from 86 mm. to 138 mm. in tt)tal length. The colour pattern on the body is subject to consideraljle variation.
In 1922, I assigned Day's B. gcto to the synonymy of B. birdi. but nn an examina- tiiin of the material from the Eastern Himalayas it Ijecame clear that Day's form repre- sented a new .species,'* differing from B. birdi mainly in the nature and form of its head.
"Hora, Rcc. Ind. Mus. XXXIV, p. 571 (1932).
Explanation of Plate XII.
Neniachiliis fn>m Western Tibet
Nemachilus hutchinsoni, sp. imv.
Fig. 1. I-atcral view of a male speeinieii. x j/j.
Fig. 2. W'utral surface of iiead ami anterior pari of l>M<ly of same, x IVj.
Nemachilus panguri, sp. nov.
Fig. 3. Lateral view of a male specimen. X yi.
Fig. 4. N'eiitral surface (if head ami anterior jiart of l)o(ly of same. X 1)4.
Nemachilus deTerrai, sp. nov.
]Mg. 5. Lateral view of a male s])ecimcn. x ^A.
Fig". 6. Ventral Mu-face nf head and .anterior [lart of body (if same, x lj<2.
A'c-iiiacliiliis stoliczkac (Steindachncr)
Fig. 7. Lateral view of a male specimen, x l^.j.
Fig. 8. \'entral surface of head and anterior part of body of same, x 1>4-
MEM. CONN. ACAD., VOL. X.
PLATE Xn.
ARTICLE XVIII
REPORT ON FISHES. PART II: SISORIDAE AND CYPRINIDAE
By Dev Dev Mukerji, M.Sc.
Zoological Survey of India, Calcutta
(Published with permission uf tlie Director, Zoological Survey of In(Ha)
CONTENTS
Page
Introduction 323
Systematic Account 326
Sisoridae :
Glyptothorax kashinircnsis Hora 326
Glyptostcrnum rcticiihituin McClelland 328
Cyprinidae :
Cyprininae :
Laheo diplosfoimis (Heckel) 329
Crossocliiliis piDijabciisis Mukerji 331
Schizothoracinae :
Scliicothorax labiatus (McClelland) 333
Scliicotliorax longipiimis Heckel 334
Schiaothorax csociniis Heckel 335
Schiaothonix planifrons Heckel 340
Schicothorax iincropoi^on Heckel 343
Scliiaothorax curvifrons Heckel 346
Orcinus sinuatus (Heckel) 347
? Schisothorax labiatus x Orcinus sinuatus 350
? Scliiaothomx planifrons x Orcinus sinuatus 351
Schiaopygopsis stoliczkae Steindacliner 351
Diptychus niaculatus Steindacliner 354
Ptyclwbarbus conirostris Steindacliner 356
INTRODUCTION
Professor G. Evelyn Hutchinson, Biologist to the Yale North India Expedition, entrusted to Dr. S. L. Hora of the Zoological Survey of India a representative collection of fisiies made l)y the Ivxpedition in 1932 in various localities in the Kashmir valley and Indian Til)et ( Ladak ) fur taxnnoniic study and rcpnrt. The collection comprises repre- sentatives of the families Sisoriihw. Cypriniihic and Cobitidac. A report on the Cohitidac is presented by Dr. Hora as Part I above, antl the collection of the Sisoridae and the Cyprinidae was after a jjreliminary determination by him turned over to me for detailed stud}- and report.
Though the material came in my hands towards the latter ])art of 1934, it was not possible for me tn take up the work l>efore the middle of 1935. I'urthermore, in studying
Mf.m. Conn. Acah., Vol. X, Art. XVIII, June, 1936.
324 REPORT ON FISHES. PART II: SISORIDAE AND CVrRINIDAE
the Schi::(itlioracinac, particularly the different species of Schizothorax, more time had to be devoted than I anticipated. Though all the sjjecies of the genus brought back by the I'Lxpe- dition are referable to the already known forms from the Kashmir basin'), the specific limits and the ranges of variation ;uni)ng these fishes are by no means easy to define. The difitlculty is further accentuated by hybridisation in the Sclii::ollioraciiiac, as a result of which even the limits of genera sijmetimes become obscure.
The collection under report consists of 119 specimens," of which 0 belong to the laniily Sisoriihw and the rest to the Cyprinidnc. Among the ("yiirinids, 12 belong to the subtaniily Cypriniiiac and 101 to the Schizothoracinac. This clearly indicates the great preijonderance of the ScliicoHioracinac over other fishes in the Kashmir valley and in the portions of Indian Tibet traversed by the lixpedition.
So far as the present collection is concerned, the Kxpedition obtained specimens mainly from (i) the lakes, rivers and channels in the Kashmir valley and (ii) from several rapid running and torrential streams of the Upper Indus system in Indian Tibet at altitudes vary- ing from 8,790 feet to 15,215 feet. 1 give below a complete list of the specimens with their localities:
Kashmir Valley: Sisoridae:
''*Glyptothoni.v kasluiiircnsis llora JlRJuni River, Sha(lii)ur (Sta. K 40).
Cyprinidae:
I.abco diplostoiiiHx (Heckel) Srinagar (Sta. K 7) ; Jlu-knn River, Pampur
(Sta. K 10).
Crossochihfs piiiijiibciisis Mukerji Srinagar (Sta. K7).
Schizothorax longipinnis Heckel Channel, Manasbal Lake, Srinagar (Sta. K 48).
Schizotlwrax csoc'mus Ifeckel Srinagar (Sta. K 7) ; Jhchiin Kiver, Srinagar
Sta. K 14) ; Channel, Manasbal Lake, Srina- gar (Sta. K48).
*Scliiz(>lli()rtix plauifrons Heckel Canal to Dal Lake, Srinagar (Sta. K6) ; Dal
Lake, Srinagar (Sta. Kll); Channel to Manasbal Lake, Srinagar (Sta. K48).
*ScIi!zolliorax inicropogon Ueckel Canal to Dal Lake (Sta. K6); Dal Lake,
-Srinagar (Sta. Kll); Manasbal Lake, Srinagar (Sta. K48) ; Kashmir.
*Srliizothorax cttrvifrnns Heckel Srinagar (Sta. 7) ; Channel to Manasbal Lake,
Srinagar (Sta. K48).
Orcimis sinualns (Heckel) Jheliim River. Srinagar (.Sta. K 14) ; Jliehini
River, Pampur (Sta. K 16) ; Rivers and canals in Srinagar.
Schizothonix Jahialns / Orchuts sinuatus Jheluni River, Srinagar (Sta. K14); Rivers
and canals in Srinagar. Schizothorax plauifrons :< Orciniis simtalns ..Main canal, Srinagar (Sta. 12) ; Jheluni River
Srinagar (Sta. K 14) ; Channel to Manasbal Lake, Srinagar (Sta. K48).
'Heckel, J. J.: Fische aus Caschimr (Wien, 1838).
" For keeping down the cost of transport some of the specimens of easily determined species were not sent for report.
'^Aii asterisk (*) against a species denotes that it is endemic in the Kashmir Valley.
REPORT ON FISHES. PART II : SISORIDAE AND CYPRINIDAE 325
Indian Tibet (Ladak) : Sisoridae :
Glyptostcrnnm rcticiilatum McClelland A small rapid stream at Kalatse (Sta. L 10) ;
Leh (Sta. L 13). Cyprinidae :
Schizothorax labialus (McClelland) Spitok, Upper Indus (Sta. L 17).
Schisothorax csocinus Heckel Spitok, Upper Indus (Sta. L 17).
Orcinns sinuatus {WeckeX) Kargil (Sta. K 88) ; Spitok, Upper Indus
(Sta. L 17).
Schizotliorax hdnatus :< Orciniis siiiitulus . . . .Lake near Chushol (Sta. L 73).
Schizopygopsis stoliczkac .Steind Spitok, Upper Indus (Sta. L 17) ; Large rapid
stream between Tangtse and Mugleb (Sta. L37) ; Stream above Lukong (Sta. L45a) ; Stream at Chagra (Sta. L46) ; Stream into Pangur Tso (Sta. K 74) ; Yalapuk near Nyoma ; Upper Indus (Sta. L46); stream at Leh.
Dypliclnis niaciilatus A rapid stream at Nimu (Sta. L 12) ; Torren- tial stream at Lhabaps ( Sta. L 3 1 ) ; Large rapid stream between Tangste and Mugleb (Sta. L 37) ; Stream at Chagra (Sta. L 46) ; Stream at Leh.
Plycliobarbns conirostris Steindachner Kargil (Sta. K 88) ; Kalatse, Upper Indus
(Sta. L 17) ; Yalapuk, Lljiper Indus (Sta. L79).
As is evident from the aljove list, the family Sisoridae is represented in the collection by two genera and only two species, 7'/"., Glyptothorax kashniircnsis, and Glyptostcrnnm reticulatuin. G. kasJimirensis is an endemic species in the Kashmir Valley and is a river form mainly restricted to sluggish water. G. rcticulaluiii is essentially a torrential species and is widely distributed in the liead-waters of the ])rincipal river systems in High Centra! Asia and Eastern Turkestan, and, as is to \x expected, specimens of the species were obtained by the Expedition only from the rapid streams in Indian Tibet. Of the two representatives of the sub-family Cyprininae, vis., Labeo diplostomiis and Crossochiliis punjahensis, special attention may l)e directed to the latter species which had hitherto been known from the Punjab and Baluchistan. The present record of the fish from the Kashmir valley greatly extends its range in the Western Himalayan territory. The subfamily Schizotlwracinae is represented in the collection by six species of Scliizothorax and one species each of Oreimis, Schizopygopsis, Diptyclius and Ptycliobarbus. It may l)e noted that of the eight species of Schizotliorax descril)ed by Heckel {op. cit.) from tiie Kashmir basin, only five were obtained by the Expedition; three of these (S. planifrons, S. uiicropogon and S. cnrvifrons) are endemic in the valley. .S". hibiatiis is essentially an Afghanistan species and its occurrence in the head waters of the Indus is reported here for the first time. Orciniis sinuatns, Schizo- pygopsis stoliczkac, Diptychus inactilatus and Ptychobarbus conirostris were found in their usual habitats and do not call for any special remarks. The two hybrid forms between Schizolhorax and Orciniis are worthy of interest, but similar hybridisation among fishes in nature, particularly at high altitudes in Central Asia, is by no means a rare phenomenon
326 REPORT ON FISHES. TART 11: SISORIDAE AND CYl'RINIDAE
A detailed account (if tlie ecolnt^y, Iiinnomics and znnrrcnorraphical distribution of the fishes of the I'^xpedition is heinj; ])uhlished hy Mr. (!. F.velyn Hutchinson, and in tlic present report no attempt has, therefore, l)een made to refer to these pro])lems. In the syslenuilic account, besides discussing various taxonomic pro1)lems, and giving necessary descriptive notes, I have, however, added, wherever possiljle, certain l^iological observations which, though made by me independently, are meant only to corroborate the observations of Mr. Hutchinson. Such information as I have been able to collect on the breeding habits of the different species of Schicothoraclnac from a study of the material is also included here. OI)servations on the parasitism in several species of fish^ under investigation have also l)een made.
Before concluding T must mention that but for the excellent preservation of the material and the invaluable I'leld-notes by Mr. Hutchinson this rejiort would have Ijeen of much less value, and I take this opportunity to record here my ajipreciation of the care with which Mr. Hutchinson dealt with the collection in the field. I am grateful to the authorities of the Yale North India Mxjiedition for a grant towards the cost of illustrations which have been executed b\- Mr. U. Bagchi under my supervision. 1 am indelited to Dr. B. Prashad, Director, Zoological Survey of India, and Dr. S. L. Ilora for affording me every facility and lielp in tlu' course of ]irepar.'Uion of the re])ort. To 1 'r. Hura ! am ]),'irticnlarly thankful for tiie op|i(irtunity to investigate this interesting materi.al an<l fur jiis \aluable suggestions.
SYSTEMATIC ACCOUNT Family SISORIDAE
Genus Glyptotlwrax Blyth (1860) Glyptothora.v kashniirotsis Hora 1923. Glyptothorax kaslniiirnisis, Hora, Rcc. hid. Afiis. XXV, pp. 22-24, figs. 2, a, h and c.
Glyptothora.v kaslniiirciisis belongs to the group of species of the genus in win'ib the pectoral spine and the ventral rays are not plaited below. It is provided with a strong dorsal spine and a moderately developed adhesive ap[)aratus on the chest with a circular depression in the middle.
'l"he Yale North India I'^xpedition obtained 2 specimens of the species in April, 1''32, from the following localitv:
Jhelum River, Shadipur:
(Sta. K 40) ; ra. 5,192 ft 2 (98.85 mm.)
The eyes are situated almost in the middle of the head. The pectorals arc slightly shorter than the he;id and are sejiaratcd from the \entrals by a considerable distance. The ventrals extend as far as the anal papilla. The caudal tin is much shorter (ban the head: its upper lobe is longer than the lower.
The colouration of the specimens in alcohol is light to dusky brown with a lighter under
'At tlie instance of IJr. S. L. Hura, my colleague, Mr. Al. N. IJatta, has undertaken a systematic study of the intestinal parasites of the fi.shes of the Yale North India Expedition and a separate report on this material will he puhlishcd in another place.
REPORT ON FISHES. TART II: SISORIDAE AND CYPRINIDAE 327
surface, lilack dcits are irre!L;"uIarly scattered all over the body. All the fins are marked with faint black sputs and bands.
Pistribiitioii: Glyptothora.v l<:(isliiiiirciisis is an endemic species of the Kashmir Valley and is fairly common in rivers and sluggish waters.
[Attention has already been directed to the two ecological groups into which the species of the genus Glyptothora-v can be divided {Rcc. hid. Mns. XXV, pp. 4-8, 1923). The first group comprises less specialized forms which live in larger streams at the bases of hills and are liable to be carried into slow currents, while the members of the second group are more specialized and are invarialjly found in torrential streams. In the former case, the body is cylindrical and the adhesive apparatus is restricted to the chest and the belly as an elongated, coin])act structure. The paired fins are not horizontally placed and play only a small part in the adhesion of the fish. The species of the second group are depressed and possess a flattened ventral surface ; the paired fins are horizontally placed and the function of adhesion is transferred from the chest region to the outer, flattened rays of the paired fins. The thoracic, adhesive apparatus is considerably reduced and is restricted to the region behind the isthmus in a semilunar patch. The probable evolution of Glyptotliorax from forms like Erctliistcs and Laguvia and their ultimate development into forms like Glyptostcrnuin have already l)cen explained {Phil. Tran.<;. Roy. Soc. London, B, CCXVIII, p. 236,1930).
The form of the body and the structure of the adhesive apparatus of Glyptotliorax kashinir- cnsis are dififcrent from the species assigned to the above groups. The Ijody, though cylindrical, is much deeper and the adhesive apparatus is restricted to the chest region only {Rcc. Ind. Mus. XXV, p. 23, fig. 2 a-c, 1923) ; it is almost circular in outline and possesses a deep pit in the centre. Not knowing the precise habitat of the fish, it was not possilile to explain on ecological grounds the differences in the structures referred to above. Mr. Hutchinson now informs me that "the s])ecies is moderately common in the Jbelum near Srinagar. and also inhabits some of the sluggish canals around the city." The species would thus appear to be restricted to dee]) and flowing waters. Init a rapid current is not essential for its existence. The deep form of the body can be assrK'iated with lift- in sluggish waters where, it seems prohable, that tlie fish feeds l>y scraping ofl' algal matter from hard objects. During the feecling process the fish pnibablv adheres to the substratum before bringing into action its shar]). lower jaw. This would explain the utility of the adhesive ajiparatus in this species even when living in sluggish waters.
Attention may here be directed to the mode of life of the fishes of the genus Garra (Mukerji, Acliiiryya Sir P. C. Ray Goiinii. I'ol.. Calcutta, pp. 477-482, 1932) which possess a suctorial disc behind the lower lip. The disc ma\' ha\e originally develojxjd for stemming rapid currents, but some of the species now live in lakes and deeper streams where the sucker is used for holding on to hard f)bjects while the fish scrajies off algal matter from them. There is thus a parallelism between the mode of life of Glyplolhorax kasliiiiircit.\'i.\- and the lake-inhabiting sjiecies of Garra.
The adhesive apparatus of Glyptotliorax is not a sucker device in the same sen.se as the mental disc of Garra. but is a mechanical device for increasing frictii>n {Phil. Trans. Roy. Soc. London, 1'. CCXN'lll, p. 262, 1930). The ridges are providetl with sharp, curved spines {Rcc. [lid. Mn.<:. XXIV, pp. 55-58, 1922) which fi.x themselves into the tmevennesses of the rocks. When the fish applies its adhesive apparatus to a r(x;k, it is likely that the water flows out of the grooves by adpression and a series of partial vacua is created. The central de])ression in the adhesive apparatus of G. kaslunircn.Kis is probably a device for the production of a partial vacuum. The circular form (tf the ajiparatus and the fact that it is incomplete posteriorly shows that the species is iirobably derived from a member of the highly sjieciali/.ed .group. This again show's how at high altitudes torrential forms revert to primitive mode of life on finding sluggish waters, but it is significant that structures acquired for stemming currents in torrential streams are retained
328 KICrdRT ON FISHES. PART II: SISORIDAt; AND CYI'KIN IDAK
in coiiiiectuMi with their feechiit; habits. Thus the iitihty of a structure plaNs an important ])art from the he,u;inning of its evolution to its ultimate fate.
The mechanism of adhesion of the highly specialized menihers of (ilyptothora.v is similar to that of the Glyptosternoid fishes. The ridges and grooves on the lips and the outer rays of the paired fins are used for adhesion which is greatly facilitated by the inner rays of the pectoral fins which, by their vigorous movements, shoot out any water that may enter below the fish when adhering (Phil. Trans. Roy. Soc. London, P., CC.WIII, p. 258, 1930). Sunder I.al IIora.\
Genus Clypto.'ftcrniiiii McClelland (1842) Glypfostcnniin rcticulafuiii McClelland
1842. Glyptostcrnum rcticidatum, McClelland, Calcutta Joiirn. Nat. Hist., II, p. 584.
1860. Glyptostcrnum rcticulatxwi, Blyth, Journ. Asiat. Soc. Bengal, XXIX, p. 153.
1876. Exostoma Stoliczkac, Day, Proc. Zool. Soc. London, p. 7S2.
1877. Exostoma Stolicskae, Day, Fish. India, p. 502, pi. cxvii, fig. 3.
1878. Exostoma stoUc::kac, Day, Sci. Res. 2nd Yarkand Miss., Ichthyology, p. 1, \i\. i, figs. 1.
1889. Exostoma Oschanini, Herzenstein, Mel. Biol., XIII, p. 69.
1890. Exostoma Oschanini, Herzenstein, Bull. Acad. St. Pelersh., XXX III, p. 120. 1905. Exostoma stoliczkac. Berg, Ryhy Turkestan, p. 211, fig. 31.
1905. Parcxosioma stoliczkac, Regan, Ann. Mag. Nat. Hist. (7), XV, p. 183. 1907. Exostoma gracilc, Grazianov, Trudy Otdela Icluhyologic, IV, p. 58.
1907. Exostoma labrax, Grazianov, ibid., p. 59.
1908. Exostoma stoliczkac. Berg, Ezhegodnik Zoologischeskago Muxcya Akadcniii N auk, WW, p. 450.
1916. Parcxostoma stoliczkac. Berg, Poiss. des Eaux Douces de Russic, \). 371, figs. 280, 290.
1923. Glyptostcrnum stoliczkac, Hora, Rcc. Ind. Mus., XXV, p. 37.
1925. Parcxostoma stoliczkac, Norman, Am. Mag. Nat. Hist. (9), XV, p. 572.
1932. Glyptostcrnum rcticulatum, Hora, Ami. Mag. Nat. Hist. (10), X, p. 179, fig.
1932. Glyptostcrnum rcticulatum, Hora, Current Sci., I, p. 130.
1933. Glyptostcrnum rcticulatum, Hora, Journ. Bombay Nat. Hist. Soc, XXXVI, p. 697.
1933. Glvptostcrnum rcticulatum, Berg, Poiss. dcs Eaux Douces de I'U. R. S. S. (3r<l ed.), jH. 2, p. "S97, figs. 549-551.
1934. Glyptostcrnum rcticulatum. Hora, Journ. Bombay Nat. Hist. Soc, XXX\'II. \\ 1934. Glyptostcrnum rcticulatum, Hora, Rcc Ind. Mus., XXXV, pp. 287-292.
From the above list of synonymy it is clear that the true identity and the systematic position of Glyptostcrnum rcticulatum have long l:)een obscure, while the wide range of varia- tion that it exhibits in regard to relative proportions of the principal parts of the body has led to its being described under several names from different areas of its extensive range. Recently, Hora (op. cit., 1932, 1933, 1934) has succeeded in rediscovering this interesting species and made its position abundantly clear; he also pul)lished a detailed study of the species both frnm systematic and biological jjoints of view.
The Yale North India Expedition collected 4 specimens of the species in May-June, 1932, from the following localities:
A small rapid stream near Kalatse : Ladak (Sta. L 10), ai., 9,700 ft. .3 (10o-128mni. : 1 ,i and 2 9 2 ) Leh: Ladak (Sta. L13), 11,500 ft 1 (130mm.; 9)
REPORT ON FISHES. PART II: SISORIDAE AND CYI'RINIDAE
329
Of tliese 4 specimens, 2 from Kalatse and 1 from Leh are females, while the remain- iiit^- 1 from Kalatse, 128 mm. long, is a male. Hora (op.cif., 1934) ol)served that the species exhibits sexual dimorphism. In the males there is a sharp, conical and well-defined papilla projecting behind the anal opening. In the females there is no anal papilla but the vent is bordered by prominent, fleshy lips. The secondary sexual characters noted above are well marked in the specimens under report (Text-figure 1, a-b).
Figure 1. Glyploslcrnuin rclkulatum McClelland. Anal region of a male specimen and a female specimen from rapid streams at Kalatse, Indian Tibet, showing sexual diflferences and form and position of anal opening and structures of anal papilla in male. X ca. 2.
Distribution: Glyptoslcnuiin rcticnlatiiiii is "widely distrilmted in the head waters of the Indus (Basgo, Sneema, Leh, Ladak and the Kashmir valley), of the Kabul River (Sri-i- Chusma, Tulraiz, Paghman and the Chitral valley), of the Syr-Darya and the Amu-Darya in the Eastern Turkestan (Oxus system)."
Family CYPRINIDAE
Subfamily CYPRININAE
Genus Labco Cuvier (1817)
Labeo diplostomiis (Heckel)
18.38. I'aricorliimis diplosluiims, Heckel, Fischc ans Cascliniir, jip. 67-75, i)l. xi.
1839. Gobio ricnorhynchus, McClelland, Asiat. Res. (Ind. Cyprin.), XIX, pt. ii, pp. 279, 363-
364, pi. Iv, fig. 1.
1844. Tylognathiis valcncicmicsii, Heckel, Fischc Kasclunir's in Hugd's Rcise, p. 378, fig. xiii.
1868. Labco diplostomus, Gunther, Cat. Fish. Brit. Mns., VII, p. 57.
1877. Labco diplosloiints. Day, Fi.<;h. India, p. 540. pi. cxxix, fig. 2.
1889. Labco diplostomus, Day, Faun. Brit. Ind., Fisli., I, pp. 265, 266.
1913. Labco diplostomus, Zugmayer, Die Fische von Balutschistan (Munchcn), p. 26.
330 REPORT ON FISHES. PART 11: SISORIDAE AND rvrRINinAE
Labco diplostoiiius belongs to the group of species characterised by a tliick and fleshy iiead, sh<irt dorsal and anal fins, a thick, prominent and pendulous snout, a wide mouth, homy jaws, and thick and loose lips. The species is distributed along both the l'"astern and the Western slopes of the Himalayas, and presents a certain .-nndunl of \ariatinn in biuly proportions, etc., which, as far as I can judge from the nialcrial bclMrc nie. is UKirc indixidual than geographical.
In the collections nf the Indian Museum /.. (lif^liislniiiiis is rciirescntcd li\- 11 specimens. The localities of 10 are definitely known, 5 are fmm Siml;i, 1 fnmi ilardwar, 2 frnm Assam, 1 from Chuniba and 1 from Baluchistan. The lirst nine specimens are the orig- inals of Day's description of the species, while the one from LJalnchistan is a representative of the series collected by Dr. l^rich Zugmayer fnim the \'indar l\i\er at Snnnnani during his travels in that ciunitry in l*'!!. The lltli s])ecinien, labelled "/.iilh\) riciinrlixiiflnis" (Keg. No. Cat. 687), is fmm the old collections of the .\siatic Society of llengal, but unfortunately its history is completely effaced from the original label which is still attached to the specimen. It appears probable, however, that the specimen comes from "Northern part of Bengal" and is the original of McClelland's descrijition of Gobio ricnorhynchus. for, in reference to the only specimen of the species that he examined, he mentioned: "The .specimen here described was found by Mr. Hodgson by whom it was presented to the Asiatic Society" (r'/'. cit. p. 364). The only other Asiatic Society of Bengal specimen of the species preserved in the Indian Museum collections is Day's original from Ilardwar.
The 'S'ale North India Expedition obtained 2 specimens f)f the species in March, 1932, from the following localities in Kashmir:
Srina,gar (Sta. K 7) ; ca. 5,200 ft 1 ( 16.S mm.)
jheluni River. l'ain|inr ( Sta. K 10) ; ca. 5,200 ft 1 ( 140 mm.)
The s|)ecimens agree in all respects with the above mentioned series of specimens from the different localities as alst) with lleckel's description of the species, and do not call for any special remarks, except for the fact that there are only 41 scales along the lateral line and 13 rows of scales in a transverse series bet\\(.'en the bases of tlu' dorsal and the \entral fins. The scales on the chest region are considerably reduced in size and are jiartly imbedded in the skin, thereby producing a more or less smooth under surface. In most of the speci- mens that I have examined, the snout is studded with well defined horny tul)ercles which apparently develop quite early in life. In the specimens from Kashmir under report the snout is, however, perforated with a few fine mucous pores instead. It appears that the tuberculate condition of the snout is a secondary sexual character of the males, but from tlu' material before me it is not jiossible to give a derniite opinion.
The colouration of the specimens in alcohol is uniformly reddish-brown with a slightly- darker upper surface. The outer edges of the dorsal and the caudal fins are dusky.
Ihslrihutioii: Labco diplostoiiius tKcurs in Kashmir (Type-locality), "along the Sind hills and Himalayas, also in lirabmaputra in Assam" (Day). In regard to the li;ibitat and the size of the .species in the Assamese waters McClelland (op. cil. p. 364) observed: "Tiie Nepura of the .'\s.samese I found as low as Bishenath, where the current is slow, and the bottom is .sandy; here its colour is deep blue on the back. It is small, and very rarely met with in Lower Assam; Init above the rajjids Mr. Criffith says it is very connnon. and attains a large size, and that the fins and tail arc dusky, the body below white, above
REPORT ON FLSMES. PART 11: SISORIDAE AND CVPRINIDAE 331
Attention may here be directed t<i the fact that in 1930 and later in 1931 Tchang'' recorded a species under the name Labco diplostoimis Heckel from Kiating in the Szechwan Province in China. In 1932 Rendahl" reported the same fish from Clningking in the said Province, Imt douhted its identity with Ileclxel's species, in view of tlie differences in the form of tlie mouth, body proportions, squamation, etc. In 1934 Kimura' recorded the form from Yangtze-kiang and definitely assigned it to a new species, "Labco {\'aricorli\iicluis) Rciiddlili.''
As Labco diplostoiiiits presents variations in different body proportions, lengtli of the fins, etc., I give below tlie measurements of the two specimens under report:
Mcasiirciiiciils in iiiilliiiictrcs
Total length without caudal 165.0 140.0
Length of head 37.0 32.0
Width of head 24.0 20.0
Height of head 28.0 23.0
Diameter of eye 6.0 6.0
Length of snout 13.0 12.0
Interorbital width 14.0 12.0
Deptli of body 40.0 30.0
Longest ray of dorsal 33.0 31.0
Longest ray of anal 25.0 22.0
Length of pectoral 31.0 26.0
Length of ventral 27.0 23.0
Length of caudal 38.0
Least height of caudal peduncle 19.0 15.0
Genus Crossnchilns \'an Hasselt (1823)
Crossocliilus piiniabciisis Mukerji
1934. Crnssochiliis laliits piiiiiabciisis. Muk-erji, Jotirii. Bombay. Nat. Hist. Soc, XXXVIl, p. 53, fig. 7.
broni an examination of large series of specimens of "Crossocliilii.': hiliiis" from dif- ferent parts of India and Uurnia, preserved in tlie collections of the Indian Museum, I came to the conclusion that tlie species is a comjiosite one and that the dwarf form from the Punjab and I5a1ucbistan, at any rate, sliould be regarded as distinct from the typical form from iKirtbcni I'lengal. At that time 1 had no specimens at mv disposal from the Kashmir waters for comparison with the subspecies pnnjabcnsis and it was, therefore, not possible to ju^lge the affinitives of the former with the latter. On an e.\aminati(in of the ten specimens collected by the "N'ale North India b'.xpedition in March-May, 1''32, from parts of Kashmir, 1 tind that in all essential char.acters they correspond to the form piDiju- bcnsi.'; whicli 1 now consider on re-e\aniinatioii to be sufficiently distinct Iroin C'. hitiiis lo merit specific rank.
"^^TchaiiR, I..: Sim-nsUi, I, No. 7, pp. 87-94 (1930); Bull. Pan. Mew. In.':!. Biol.. II, Ni>. 11, p. 227 (1931).
"Rendahl, II.: Arl,iv. /. Zoologi, XXIV (A), No. 16, pp. 74-79 (1932).
"Kimura, S. : Journ. .Shmiglmi .S"ci'. huit.. Sec. 3, I, pi), 125-12S, pi. iii. fiR. 2 (1934).
332
REPORT ON FISHES. PART II: SISORIDAE AND CYPRINIDAE
The Yale North India Expedition obtained 10 specimens from the following localities:
Dal Lake. Lokut (Sta. K 3), ca. 5.190 ft 3 (55- 62 mm.)
Srinagar (Sta. K 7), ca. 5,200 ft 7 (62-108 mm.)
Of these 10 specimens, 3 taken at I.nkut Dal Lake, are l)adly desiccated and shrivelled. The majority are females either with ripe eggs or fairly matnre gonads. The species seUlom exceeds 150 mm. in length, and apparently attains se.xual maturity at 30 tn 45 mm. stage (Mukerji, o/'. cit.). The tish is of a stout and thick huild with a snmewliat deep body and a liroad and blunt snout. The position of the vent is from 3 tci 4 scales in advance of the in.'iertion of the anal fin. In small specimens the \cntrals extend as far ;is the vent, Imt in larger full-grown individuals they just miss it.
The colouration of the specimens in alcuhcil \;iries tmni dark hi'dwii {n dusky with a darker upper half. In medium sized specimens of 30 to 70 mm. in length, the scales are sometimes infuscatcd with fine black dots. All the fins are diajjlianous.
Bionomics: The peritoneum is black. The length of the air-bladder is contained about 3.5 times in the length of the body excluding the caudal iin. The posterior chanil)er is long and narrow with a sharp end; it is of uniform thickness and its length is almost twice that of the anterior chanil)er which is somewhat of a knob-like structure (Text-figure 2).
2
Figure 2. Crossochiliis /'uiijabeiisis Mukerji. Air-bladder of (a) an adult specimen, 108 mm. long from Kashmir valley and (6) a young specimen, 65 mm. 'X.Syj.
REPORT ON FISHES. PART II: SISORIDAE AND CYPRINIDAE 333
The intestine is of considerable length, narrow with many convolutions. In specimens over 100 mm. in length, the intestine is about 6 times as long as the body excluding the caudal fin, while in smaller specimens it is about 6.5 times the same length. The gut contents include lumps of slime and algae mixed with sand particles, bits of leaves and twigs.
Parasites: The specimens obtained by the Yale North India Expedition are heavily infected with Trematode parasites the cysts of which appear as small black nodules all over the body and the fins. In a gravid female, 108 mm. long, six specimens of a species of an Acanthocephalan worm have been found.
Distribution: As far as I can judge, C. pimjabensis is distributed in the mountainous rivers, streams and lakes in the Western Himalayan territory through Kashmir and the Punjab along the North Western Frontier to Baluchistan in the Western territory. In the Eastern sub-Himalayan and Burmo-Malayan territories the species is replaced by C. latins (Ham. Buch.).
Subfamily Schizothoracinac Genus Schizothorax Heckel (1838)^ Schizotlwrax labiatus (McClelland)
1842. Racoma labiatus. McClelland, Catculta Jonrn. Nat. Ilisl., II, ]i. 578, pi. xv, fig. 1.
1842. Schiznihorav Ritchicana, McClelland, Hiid., p. 580.
1868. Racoma labiatus, Giinther, Cat. Fish. Brit. Mus., VII, ]>. lf>2 (footnote).
1868. Schizolliora.v rilcliiauus, Giinther, Cat. Fish. Brit. Mus., VII, p. 162 (footnote).
1877. Schizothorax Ritchianus, Day, Fish. India, p. 531 (footnote).
1877. Schizothorax labiatus. Day, ibid., p. 532 (footnote).
1934. Schizothorax labiatus. Mora, Rcc. Ind. Mus., XXXVI, pp. 292-297, figs. 2 and 3.
The narrow, suljcylindrical body, the large and pi->intcd head, the well developed and reflected posterior margin of the lower lip which is invariably triloljed and the sharp and shovel-shaped lower jaw are some of the principal features of S. labiatus originally described from the Kunar river near Jalallabad. Recently, Hora (o/>. cit.) published a detailed account of this hitherto little-known species from a series of specimens obtained from the Chitral Valley. He has shown that in the species the structures of the lower lip and the air-bladder are considerably variable. Further, according to him .S". ritchicana McClelland, a species descrilied also from Afghanistan, is synonymous with 5. labiatus.
The Yale North India Expedition obtained a single specimen of the species in June, 1932, from the following locality:
Upper Indus, Spitok, Ladak (Sta. L 17), 10,730 ft (260 mm.)
" Just before this report went to tlie press. Dr. Yiianting T. Chu's paper on "Comparative studies on the scales and on the pharj-npeals and their teeth in Chinese Cyprinids, with particular reference to taxonomy and evolution" (Biol. Bull. St. John's Vniv., Shanghui, China, pp. 1-22S, pis. i-.xxx, No. 2, 1935) was available for reference. In regard to the taxonomic positions of the genera {Schizothorax Heckel and Orcinns McClelland the author has pointed out that according to Bleeker's restriction {Nat. Tijd. Dicrk, I, p. 196, 1863) the name Scln::opygc Heckel should be used in place of "Schisolhora.v" of authors and Schizothorax Heckel should replace the name Orcinns of McClelland. Although I thoroughly agree with this statement I am in favor, in view of the general application, to conserve the names Schizothorax and Orcinns in tlieir hithertofore accepted sense.
334 REPORT ON FISHES. PART II: SISORIPAE ANI1 CYPRIMDAK
In tlie single specimen be lore nic the snout is sparsely studded with fairly sharp, warty tubercles. The eves are situated slij;htly nearer to the tip of the snout than to the posterior margin of the operculum. The lower lip is tril'ibed: llie lateral li^bes are broad and llat, while the central one is narrower and somewhat elongated. The dorsal spine is fairly strong and sharply denticulated posteriorly; its length is equal tn the head behind the nostrils.
The ground colouration of the specimen in alcohol is reddish brnwn with a darker dorso-lateral surface.
Pistrihiilion: Sclii::otlioro.v hihintiis is essentially confined to Afghanistan and is com- mon in the Kunar or the Chitral river and in its tributaries in the ("hitral X'alley. Its occur- rence in the Upper Indus is rejiorted here fur the tirst time. Hybrids Utween this sjiecies and Orriinis siiniatiis occur in the Kashmir \alley (<•. ;'. ]). 349) lint d" nut necessarily indi- cate that Ixith parent species are present in that region. The species is usually found in rapid-flowing waters.
Schicolhorax longipinnis TIeckcl
1S38. Schicollwra.v loiii:;il'iiiiiis, Hcckel, /■'isclic aiis Cascliiiiir, |)p. 27-29, pi. iv.
b'^44. Sclii"olhora.v longif'iiinis, Meckel, Pisclic Kasclniilr's in [{ui^cl's Rcisc.
1868. Schizothorax loni:;i/^l)i)iis. Giinther, Cat. Fish. I'rit. Mii.<:.. \'II. j). U)().
1877. Schicnthom.v hitgipiiuii.';. Day, F/.v/;. India, p. 532 (f(K)tn()te).
1889. Scliiaotliord.x- !ongil>inni.s. Day, Paitii. Brit. hid. l'i.<;h.. I, p. 2.^2 (footnote).
1916. Schizotliora.v long{fiiniii.<:. \'inci.guerra, ./;/». .I/h.v. Civ. Star. Wit. Ciciuiva. .\l.\ II. ]i. 141.
The species can Ix? readiU distinguished troni others by its narrow and liigli anal tin. which when laid l)ack, extends to the root of the caudal fin (Text-figure 3) and also by its .sharp lower jaw. The body is somewhat narrow and cylindrical with a moderately com- pressed caudal peduncle. The contour of the body as well as the development and structure ">{ the tins highl}' suggest that the fish is an inhabitant of the rapid-running stream-, and rivers.
The N ale North India I'.xpcdition obtainetl a single gravid female in April, 1932, from the follow ing locality :
Channel : :Manasbal Lake ( Sta. K 48) ; 5,196 ft (3(K)nini.)
The mouth is inferior, horse-shoe-sha])ed and \erv slighlK ol)lii|uc; it is nnich broader than long and its .gape is almost ecpial to the length of the head in front of the nostrils. The upper jaw is longer than the lower and is ])rovided with a fairly developed and partly protrusible fleshy lip: the lower jaw has a free and sharp margin, practically denuded of a true lip. The lower lip seems to have undergone atrojihy Inith in its structure and func- tion, and is represented only laterally as two short and thin loose l],i])s (Text-figure 4).
The osseous ray of the dorsal fin is moderately strong and almost as long as the head behind the nostrils; its posterior serrations are fine and close-set. The length of the head is cont.ained nearly 4.25 times in the length of the body without the caudal. The diameter of the eye is contained 5.5 times in the length of the head. The interorbital space is flat and wide and more than twice the orbital width. 'Jhe snout is broadly rounded anteriorlv and almost twice as long as the diameter of the eve.
The colouration of the specimen in alcohol is reddish brown with a somewhat darker dorsal surface. All the tins are ti|)i)ed with black.
REPORT ON FISHES. PART II: SISORIDAE AND CYPRINIDAE
335
Figure 3. Lateral view of posterior part of body of Sclihollwrax Iniigil^iiinis Meckel from Afanasbal Lake, showing form of anal and caudal fins and nature of tiled rows of anal scales. X Vi- The arrow indicates the extent of the anal fin in relation to the root of the caudal.
Figure 4. Yentro-Iateral view of head of Schizollunax loiigipiiinis Ileckcl from Manasbal Lake, showing sharpness of free margin of lower jaw and specially niodilied lower lip. X i'/i-
Distnbulinii: Schicothorux lon'^i[^iimis is fduml in the Kashmir Valley and the Indus at Skardii. It presumably inhabits rapid streams and rivers, occasionally entering lakes.
|
1838. |
Schicutliora.v |
|
1842. |
Schiaothorax |
|
1844. |
Schiaotlwrax |
|
1868. |
ScJiizotlionix |
|
1876. |
Schizolhonix |
|
1876. |
Schisothonix |
|
1877. |
Scluzotliovax |
|
1877. |
Schisothorax |
|
1878. |
Schizothonix |
|
1878. |
Schicothonix |
|
1889. |
Scliiaatlinnix |
|
1889. |
Siliizothorax |
Schi:!othorax csocinus Meckel
esocinus, Heckel, Fischc aits Caschniir, p. 48, pi. ix.
esocinus, McClelland, Calcutta Joiini. Nat. Hist., II, p. 579.
csocinus, Heckel, Fisclir Kashmir's in Iliii^cl's k'risc. p. 372, 3 figs.
esocinus^ Gihither. Cat. I-'isli. I'rit. Mus., \'\\. p. 166.
csocinus, Day, Proc. Zool. .Sac. I.oihlon, p. 78.s.
[•iindalus. Day, il>iil.. ]t. 785.
csocinus. Day, Fi.sii. India, p. 533, pi. cxxiii, fig. 4.
punctatus. Day, ibid.. ]>. 532 (footnote), ]il. cxxiii, fig. 3.
csocinus. Day, .S"c/. Res. 2nd Varkaiid Miss., Iclitlivoloiiy, ji. 4, j)l. i, fig. 4.
punctatus. Day, //'/(/., p. 4. pi. i, lig. 3.
csocinus. Day, IhitDi. liril. hid. I'isli., T. ji. 254.
puiictalus. Day, ildd.. \> 252 (footnote).
336 REPORT ON FISHES. PART II: SISORIDAE AND CYI'KINIDAE
1910. Scliicothorax csociiiiis, Zuginayer, Zool. Jalirb. {.Ibih. Sysl.), XXIX, p. 277.
1916. Scl\izothorax csociiius, Vinciguerra, Ann. Miis. Civ. Stor. Nat. Genova (3) VJI, p. 142.
1934. Schizothora.v csocimis. Ilora, Kcc. Ind.Mus., XXXVl, pj.. 297-300.
The Yale Nortli India Expeditidn iil>taitie(l 12 specimens of the species in i\prii-June, 1032, fnmi the following localities:
Sriiiagar (.Sta. K 7) ; ca. 5,200 ft 5 (270, 250 nun., 9 9 ; 102-105 mm.)
Main Canal. Srinayar (Sta. K 12) ; ca. 5,200 ft 1 (235 mm., 9 )
Jhelum River, Srinagar (Sta. K 14) ; ca. 5200 ft 3 (270, 235 mm., $ $ ; 410 mm., 9 )
Channel : Manashal Lake (Sta. K 48) ; 5,196 ft 1 (260 mm., 9 )
Upper Indns, Spitok, Ladak (Sta. L 17) ; 10,730 ft 3 (290 mm., ,5 ; 425 mm., 9 )
Scliicotliora.v esociniis is, as far as one can judge from a study of series of specimens from ilifferent localities, a very variable species in regard to the different body proportions, structure of the jaws and the lips, the position of the eye and the relative positions and lengths of the fins and its colouratitm. Among an assemblage of specimens of the species there may l>e found forms which show a remarkable combination of variations from the typical r.Torm!/,?-characters. Such aberrant forms, if judged by themselves, are very baffling and tend to assert claims to distinct si)ecinc ranks. Day's .Schizollun-a.v puiictatits is one of such forms, as I lind from a study of Day's originals of the figures of S. c.^ocinii.'; and .v. pioictatiis, as also from a careful examination of a fair series of si)ecimens from various localities in Kashmir. Zugiuayer, Vinciguerra and recently Mora referred at some length to the discrepancies in Day's descriptions of the two species and to the inaccuracies in his drawings and considered the two species as itlentical. Opportunity is here taken to sub- stantiate the views of these authors by a detailed analysis of more e.xtensive material before me.
Condensing Day's loose descriptions of the two species and the differences exhibited by his original specimens I find that S. csocinii.^ and .S". puiictatKs differ in the following three principal characters only :
.S". csociiius S. piinctattis
1. Upper jaw slightly longer tli;m lower. 1. Lower jaw slightly longer than upper.
2. Serrated dorsal spine equal to length of 2. Serrated dorsal spine e<|ual to length of head behind nostrils. head behind middle of eye.
3. Anal fin laid flat almost reaches root of 3. Anal fin laid flat does not reach root of caudal or just misses it. caudal or widely separated from it.
With a view to testing the \alidity and studying the nature and the range of variation, if any, of these three distinguishing features, I have made observations in details, which fcjr convenience of reference, are given in a tabular form (Table 1 ). It is clearly seen froiu the table that of a series of 20 specimens examined by me, including Day's 2 original specimens, in 17 cases, of which 6 are adult females and 4 adult males, the upper jaw is slightly longer than the lower, similar to the typical ('.vorfH;/,s--condition (Text-figure 5, a), while only in 3 cases, one of which is a full-grown female and two half-grown, the lower jaw is slightly longer than the upjier and corresponds to that of the typical piinctatus (Text- figure 5, b). In regard to the length of the serrated dorsal spine, it was found that in 14
REPORT ON FISHES. PART II: SISORIDAE AND CYPRINIDAE
337
cases, the majority of which are full grown and of which at least 7 are females and 4 males, the spine is like that of typical pitnctatus, equal to the length of the head behind the middle of the eye, whereas only in 6 individuals, which are either half-grown or young, the length of the spine is nearly equal to that of the head behind the nostrils. In reference to the third character, c. g., the length of the anal fin, the table shows that in 12 specimens, all of which are full grown and of which at least 6 are females and 4 males, the anal fin laid flat does
TAl'.LK I Variation of characters in Scliicothorax csociiuis Heckel
|
LOCALITY |
c 5 |
u a.— |
1- u tic = 1 |
C n •b-5 Us oSg C u >« |
HI o^ o O-a re |
Length of dorsal spine equ,tls lenjjth of head behind nostrils |
11 CO rt |
cs'ra <E |
< u |
||||
|
Y.N |
.I.E. (Sta. K;) Srinagar |
2 |
270.0 |
X |
X |
X |
X |
||||||
|
U t< it (( |
9 |
250.0 |
X |
X |
X |
X |
|||||||
|
(( <t (1 (( |
? |
2 |
* 105.0 105.0 |
X X |
X X |
X |
X |
X X |
|||||
|
(( (( (( tl |
? |
102.0 |
X |
X |
X |
X |
|||||||
|
(Sta. K 12) Srinagar |
9 |
235.0 |
X |
X |
X |
X |
|||||||
|
(Sta. K14) Srinagar |
$ |
270.0 |
X |
X |
X |
X |
|||||||
|
(1 (1 (1 11 |
$ |
235.0 |
X |
X |
X |
X |
|||||||
|
(1 H ft It |
9 |
t410.0 |
X |
X |
X |
X |
|||||||
|
(Sta. K48) Srinagar |
5 |
260.0 |
X |
X |
X |
X |
|||||||
|
(Sta. L17) Spitok.. |
5 |
425.0 290.0 |
.X X |
X X |
X X |
X X |
|||||||
|
(I.^ |
I.) Chitral R |
s |
230.0 |
X |
X |
X |
X |
||||||
|
Jhelum R it H |
? |
*11S.0 *108.0 |
X X |
X X |
X |
X |
X |
||||||
|
tt H |
? |
* 97.0 |
X |
X |
X |
X |
|||||||
|
Wular L |
9 |
200,0 |
X |
X |
X |
X |
|||||||
|
Gandarbal |
? |
280.0 |
X |
X |
X |
X |
|||||||
|
Leh (Day's S. esocinus) |
? |
*170.5 |
X |
X |
X |
X |
|||||||
|
Kashmir L. |
- |
||||||||||||
|
(Day's .S". piinctatus) . . |
? |
1 |
1 1 92.0 |
X |
X |
X |
X |
||||||
|
7 99 |
20 |
17 |
3 |
16 |
4 |
14 |
6 |
12 |
7 |
1 |
|||
|
4 Si |
Y. N. I. E. — Denotes specimens collected by the Yale North India Expedition. I. M. — Denotes specimens preserved in the collection of the Indian Museum. * Denotes typical esocijiits characters, t Denotes typical PHttctatiis characters.
338
Ki.i'oKT <)\ KisiiKS. r.vkr ii: sis<ii<ii> ai-: and tvi'iuxidak
not rcadi ^he caudal and is rather widely separated from its root, a condition that corrc- sjjontls to the pinutntiis-tyiic (Text-figure 6, </ ) ; in 7 cases, all of which are, with the excep- tion of an adult female, young or half-grown, the anal hn laid llat just misses the root of the caudal, similar to the typical «Of/;nYi--condition (Text-figure 6, b), while imly in a single s])ecinien (115 mm.) from the Jhcluni River does the anal I'm actu.illy reach the caud;il (Text-figure 6, c).
l'"roni a stud_\- of the tahle it is moreover apparent that of the IS s]iccimens examined by me, excluding Day's 2 originals, .all the three ty|)ical (•.s()(/«;f.s--characters are present in half-.grown specimens only (m;irkcd with an ;i>tcrisk in the tahle"), while only a single
5
Fir.URF. 5. Lateral view of anterior parts of body of two specimens of Schholhora.v csociitits Hcckcl sliowing "esociniis" and "punclaliis" types of head and jaws, (a) Day's original specimen of "Schicolhorax csociitiis" from Leli (upper jaw longer). X l/<3- (6) Day's original specimen of "Schicotlwrax puintalKs" from "Kaslimir Lake" i.e., Wukir Lake (lower jaw longer). Nat. size.
s])ecimen, a large femrde |41() mm.) from the Jhclum Uiver (marked with a ilagger in the table) represents all the true /vnn/a/H.s-characters. The rest of the specimens exhibit nn'xed variations.
Summarizing the above analysis of characters the following facts may now be definiteU' established :
(i) Tn the majorit\' (d" full-grown specimens the njij)er jaw is, irrespective of sex
and age, longer than the l^wer, while in a lew cases the lower jaw is longer
than the up])er.
fii) Tn the majority of full-grown specimens, the length of the serrated dtirsal spine
is, irrespective of sex, almost ecpial to the length of the head Ix-hind the middle
REPORT ON FISHES. PART II: SISORIDAE AND CYPRINIDAE
339
of the eye, while as a rule, in yduiig or half-grown individuals, the spine is comparatively long, equalling- the length of the head Ijehind the nostrils, (iii ) In the majority of full-grown individuals, irrespective of sex, the anal fin laid flat is widely separated from the root of the caudal, while invariably in young and half-grown specimens it just misses or only exceptionally it reaches the root of the caudal.
(iv) Combination of all the three csociniis-characters, is found alnmst invarial)ly in young or half-grown specimens, and
(v) Combination of all the />;»;< /(///(.s-characters is rarely met with.
6
FinuRF. 6. Lateral views of anterior parts of body of three specimens of Schicolhorax rsocinus Heckel sliowing different forms of anal and candal fins. (a) Day's original specimen of "Schholluira.v pnnctaliis" from Wular Lake (anal widely separated from caudal). X H- 0') Day's original specimen of "Schizolhorax esocimis" from Leh (anal almost reaching caudal). X H- ('') A young specimen, 115mm. long (Kashmir Survey coll.), from Jhelum River, Kashmir valley (anal reaching caudal). X 1'4. The arrow indicates the extent of the anal tin in relation to the root of the caudal.
From the foregoing account it is, T believe, abundantly clear that specific differentiation between A", rsocinus and S. pimctatiis is impossible and that Day's .S". pimctatiis is only an .aberrant \ariation t)f .S\ csdciiius Heckel.
Hiniidiiilcs: The ])eritiinetuii is bhick. The length of the intestine is approximately 1.5 times the length of the body excluding the caudal fin. In adult individuals the length of the anterior chaml>er of the air-])ladder, in propnrtion to that of the posterior chamber, is ciiuipjiratively linger than it is in yinnig specimens. In a sjiecimen, 270 mm. long, the leiiglb (il the anterior ch.'imber is 33 mm. and that of the posterior chamber fiO mm., while in a specimen about 105 mm. long, the corresponding measurements are 16 mm. and 22 mm. The gut contents of several .specimens from the Jhelum I'iiver and other localities in Srinagar were ex;nnined. In some cases the iiilesline was pr;icticall\' empty, whereas in idbers frag-
340 REPORT ON FISIIKS. PART II: SISORIDAF. AND CYPKINIUAE
ments of semi-digested insect larvae and a kind of suft, pulpy ori^anic sul)stance mixed with sand and gravel and bits of twigs and weeds were fonnd. I'rom a large gravid female. 425 mm. from Spitnk Inmps of semi-digested fleshy substance and large (luantity of frag- ments of fish bones were also found. Judging from the nature of the stomach contents it appears that 5". csocinus is .somewhat a dirty, nonselective and mixed feeder, but its short intestine indicates that it feeds chiefly on animal matter. It is believed to be a scavenger lish, feeding on dead fish and other organisms at the Ixittoni of pools, etc.
Breeding: Nothing is definitely known about the breeding habits of the species, but it is significant that all the fenialc specimens c()llected by the \'ale North India ivxpedition during the months of April and June Uar mature eggs.
Parasites: A species of Acanthocej)halan worm lias been found in moderate numbers in the intestine of certain specimens from Srinagar and Spitok.
Distribution: Scliizotliorax csocinus is distributed in I.eh ;ni(l I.adak, in the head waters of the Indus, in the Kashmir and in the ("hitral valleys and in Afghanistan. In the Kashmir valle\' the species is popularly known ;is ''cliiruh" ( Ilutchinson).
Sclii.Z()tli(>rax plunifroiis IJeckel
1838. Schi::ulhorax planifroiis. lleckel, /•'/.scZ/r aus Kiuchmir, p. 48, pi. viii, fig. 2.
1844. Schisothorax plamfrons. Heekel, Fisclic Kaschinir's in /7((;,'(7'.v h'cisc, p. 370, 3 figs.
18()8. Scliizollwrax phiiiifrcnis, (jiinther, Cat. Fish. Brit. Mus., \'II, ]>. lo3.
1877. Schizothorax plamfrons, Day, Fish. India, p. 532 (footnote).
1889. Schizothorax planifrons, Day, Faun. Brit. Ind. Fish., I, p. 252 (footnote).
191(3. .Srhizothorax planifrons, Zugmayer, Zool. Jahrh. .Ihlh. Syst., XXIX, ]>. 278.
The Yale North liulia Expedition obtained 9 specimens of the species in March-May, 1932, from the following localities:
Canal to Dal Lake; Srinagar (Sta. K 6) ; ca. 5.200 ft 2 (212 mm., £ ; 200mni., 9 )
Dal. Lake: Srinagar (Sta. K6) ; ca. 5.200 ft 3 (218nini., 9 ; 178 mm., 9 ; 165 mm.)
Channel, Manasbal Lake: Srinagar (Sta. K 48) ; 5,196 ft 4 (235 mm., i ; 178 mm., 9 ;
175 mm., £ : 168 mm., S )
Schizothora.Y phinifrons is a narrow and elongated form; the ma.xiinum depth of the body is contained from 4.5 to 5.3 times in the total length excluding the caudal and the least height of the caudal peduncle is about twice in its length. The mouth is somewhat anterior, oblique and wide, and the chin is sharply ascending forwards. The jaws are almost of ecpial length. The margin of the lower jaw is not sharp, but its inside is covered with a thin deciduous cartilaginous layer" (Text-figure 7), which in some cases may be absent. The lips are thick, the lower one l>eing broadly interrupted in the middle. The barbels are nearly equal to or slightly longer than the di;imeter of the eye. The head is elongated, flat
"The presence of this diaracter may induce an impression that the species is a hybrid between .Schizolliora.x and Orciitu.'!, but in such hybrids the manifestation of tliis diaracter is always more pronounced and attended witli several other Orciiius characters. I am inclined to think that the development of a thin cartilaginous layer on the inner margin of S. planijrous is an adaptive modification which can be correlated with the scraping and scooping mode of feeding of the species. Furthermore, it is suggested by Dr. Hora that the mouth parts of .V. plauijroits indicate the probable mode of evolution of the Oreinus type of structure which in this case may have developed in sluggish waters for scraping food from hard objects and later becomes accentuated in swift currents.
REPORT ON FISHES. PART II: SISORIDAE AND CYPRINIDAE
341
above and arched below; its length is contained from 4 to 4.5 times in the length of the body excluding the caudal. It is slightly higher than broad. The snout is thick and prominent with broadly rounded anterior edge; its length is contained about 3.5 times in the length of the head. The interorbital space is flat and wide, with a prominent muchal region; it is nearly twice as broad as the orbital width. The eyes are moderate and their diameter is contained from 5.4 to 5.7 times in the length of the head and about 1.5 times in the length of the snout.
Figure 7. Lateral view of anterior part of body of a specimen of Schizothorax planifrons Heckel from Manasbal Lake, showing contour of head, nature of mouth, jaws, Hps and barbels. Nat. size. The dotted portion of the lower jaw indicates the position of the cartilaginous layer on its inner margin.
Figure 8. Dorsal fin of a specimen of Schhothorux I'laiiifrons Heckel from Manasbal Lake, showing struc- ture of spine and nature of serration on its posterior border. X ^'A.
The origin of the dorsal fin is opposite that of the ventrals and is much nearer the root of the caudal than the tip of the snout. The dorsal spine is long and strong and has from 18 to 21 coarse serrations posteriorly (Te.xt-figure 8) ; it is slightly shorter than the head f)r equal to the length of the head behind the nostrils which are situated much nearer the anterior margin of the eye than the tip of the snout. The anal fin is somewhat narrow and its longest ray is almost as long as that of the dorsal; in the grown-up specimens it almost reaches the root of the caudal (Text-figure 9), while in young and half-grown indi- viduals it is separated by a little distance. The pectorals are much shorter than the head, alxjut as long as the head l)ehind the anterior margin of the eyes; they are separated from the ventrals by a wide space. The ventrals are a little shorter than the pectorals and are separated from the insertion of the anal by a considerable distance. The caudal is deeply
342
REPORT ON FISHES. PART 11: SISORIDAE AN'O CYPRIXIDAK
furcate with pointed lobes, the upper of which is appreciably lonyer tiian the lower. It is siiglitly shorter than the head; the lengtii of its central rays is contained nearly 2.5 times in that of the outer ones. The lateral line is uniformly and moderately concave, and runs to the middle of the root of the caudal; in some specimens it is irre,L;ularl\' wavy in the anteriur half of the iKxly. The vent is in front of the origin of the anal; in female specimens it is provided with a raised Heshy area. The scales on the body are very small. The tiled row of anal scales are moderately developed, the largest ones being about half as broad as the orbit.
The coluuratiiin of the majority of specimens in alcohol is reddish brown with a much darker upper surface, while a few are somewhat paler.'" The entire body, except for the ventral surface is powdered with black pigments. All the llns are dusky.
9
Figure 9. Lateral view of posterior part of a specimen of Schholhorax phnijrons Heckel from Manasbal Lake, showing form of anal and caudal fins. Nat. size. The arrow indicates the e.\tent of tlie anal fin in relation to the root of the caudal.
Bionomics: The peritoueimi is black. The structme of the air-bladder is more or less similar to the other memljers of the genus. Its posterior chaml)er is elongated and spindle- sha])ed, while its semi-rounded and wider anterior chamber is much shorter. In a specimen, 212 mm. long, the lengths of the anterior and the posterior chambers are 28 mm. and 45 mm. respectively. The alimentary canal is long, wide and moderately convoluted. Its length in grown-up individuals is nearly 3.5 times the length of the body excluding the caudal. The gut contents of 4 specimens from the Dal and the Manasbal lakes were examined. I.timps of a kind ui soft pulpy substance mixed with sand and mud, vegetable debris and huge amount of filamentous algae were found. There was no trace of any insect larvae or of any other animal matter. In the case of a gravid female the intestine was partially empty. The gut ci>ntent^ and also the- length of tlu- iule>line suggest that the species is a vegetable teeder.
'"Ill one of his letters Mr. (i. K. Hutchinson informed us that the Kashmiri fishermen observe difference between the paler specimens and the darker ones by using the name "Cluilla" and "Chush" respectively. There is not the least doubt, however, that "Clialla" is a colour variation of "Chush." Dr. S. L. Hora kindly informs me that in the Punjab a similar word "Chitia" means white, while "Challa" would denote a licking habit on the part of the (ish. Evidently reference is made to the colour of the species in this case.
REPORT ON FISHES. PART II: SISORIDAE AND CYPRINIDAE 343
Breeding: There are im data available so far about the breeding of the species, but all the female specimens collected by the Yale North India l^xpedition during the months of March to June are full of mature eggs, the approximate diameter of which is 1.5 mm.
Parasites: Large numbers of a species of Acanthocephalan worm were found in the intestine of both male and female specimens.
Distrihutinn and Remarks: Schicoflwrax planifrnns. known amongst the Kashmiri fish- ermen and anglers as "Cliusli." is one of the commonest endemic species of the Kashmir val- ley, usually inhabiting the principal lakes and the adjoining channels. It co-occurs with two closely allied species, e.g. S. inieropogon and S. niger. S. inicropngon is a somewhat smaller species and is known amongst the Kashmiris as "Ranigliurdi," while 5". niger, as the name indicates, is a much darker species with somewhat larger eyes, a shorter anal and shorter barlK-ls; it is locally known as "Alghard" or "Algliad." It is probable that -S". plani- frons and S. niger represent one and the same species, but in the absence of more extensive material it is difficult to judge the range of variation and to come to any definite conclusion. The specimens obtained by the Yale North India Expedition, however, correspond to the planifrons-type.
MeasiirciiiciJis in iiiilliiiictres
Total length without caudal
Length of head
Width of head
Heiglit of head
Diameter of eye
Length of snout
Interorbital width
Depth of body
Longest ray of dorsal
Longest ray of anal
Length of pectoral
Length of ventral
Length of caudal
Distance between pcctnral and base of ventral
Distance between ventral and base of anal
Length of caudal jieduncle
Least height of caudal peduncle
Seliirjclhorax niicropitgan Ileckel
1838. Scliicolhora.v inieropogon, Hcckel. Piselie aits Casehniir. p. 41.
1844. Schicothorax micropogon, Meckel, Fisclic Kaschmir's in Hiigel's Reise. p. 369, 3 figs.
1868. Selticotliorax inieropogon, Giinthcr, Cat. Fish. Brit. Mus.. \TI, ]ip. 163, 164.
1877. Sehicothorax inieropogon, Day, Fisli. India, p. 532 (fnotuutc).
1889. .Selii::otIiorax inieropogon. Day, rami. Urif. Ind., fi.sli.. I, p. 252 (footnote).
Ill the collection of the ^'ale North India l'"xpedition the species is represented by 10 female specimens obtained in March-May, 1932, from the following localities:
|
£ |
9 |
9 |
|
235.0 |
218.0 |
200.0 |
|
57.0 |
52.0 |
51.0 |
|
30.0 |
30.0 |
30.0 |
|
35.0 |
34.0 |
34.0 |
|
10.0 |
9.0 |
9.0 |
|
16.0 |
15.0 |
15.0 |
|
20.0 |
19.5 |
18.0 |
|
44.0 |
44.0 |
42.0 |
|
42.0 |
42.0 |
36.0 |
|
41.0 |
40.0 |
36.0 |
|
40.0 |
39.0 |
36.0 |
|
35.0 |
35.0 |
31.0 |
|
52.0 |
51.0 |
44.0 |
|
26.0 |
26.0 |
25.0 |
|
24.0 |
24.0 |
24.0 |
|
40.0 |
40.0 |
34.0 |
|
19.0 |
19.0 |
17.0 |
344 REPORT ON FISHES. PART II: SISORIDAE AND CYPRINIDAE
Canal to DaJ Lake : Srinagar (Sta. K 6) ; ca. 5,200 ft 4 ( 105-165 mm.)
Dal Lake : Srinagar (Sta. K 1 1 ) ; ca. 5,200 ft 1 (112 mm.)
iManasbal Lake: Srinagar (Sta. K 48) ; 5,196 ft 2 (135, 132 mm.)
Kashmir 3 (165, 160, l.Wmm.)
Schizothorax micropogon is one of tlie smallest species of the Kashmir valley, rarely exceeding 7 inches in length excluding tlic caudal fin. It lias a sonicwhat dwarfed ajjpcar- ance with a thick and broad head and a deep Ixniy. The depth (if the Ixxly is contained from 4 to 4.5 times in the total length excluding tlic caudal and tlie least height of the caudal peduncle about \.5 times in its length. Tlie cleft of the mouth is small, crescentic and obliquely
Figure 10. Lateral view of anterior part of body of a specimen of SchizolJwrax microt>ogon Heckel from Dal
Lake, .showing contour of head, nature of mouth, jaws, hps and barbels. X 2. Figure 11. Dorsal fin of a specimen of Schholhorax micropogon Heckel from Dal Lake, showing structure of spine and nature of serration on its posterior border. X 2.
ascending forwards. The jaws are sub-equal in length, the upper one l^eing the longer. The margin of the lower jaw is rounded and devoid of horny covering. The lips are well developed (Text-figure 10) ; the lower one is interrupted in the middle. The l)arl)els are always shorter than the diameter of the eyes. The head is sul>-triangular in shape, with a flat and trenchant upper surface; it is slightly higher than l>ro;id and its length is contained about 3.7 times in the total length excluding the caudal. The snout is thick with ol)tuse anterior margin; its length is contained from 3.1 to 3.3 times in the length of the head. The interorbital space is flat and is about 1.5 times as broad as the orbit. The eyes are
I
REPORT ON FISHES. PART II : SISORIDAE AND CYPRINIDAE
345
pruminent and their diameter is contained from 4.2 to nearly 5 times in the length of the snout.
The commencement of the dorsal fin is opposite that of the ventrals and considerably nearer the root of the caudal than the tip of the snout ; in certain specimens it is almost equidistant l)et\veen these two points. The osseous dorsal spine is more or less narrow and slender towards the distal end, slightly curved and armed posteriorly with 17 to 20 promi- nent teeth (Text-figure 11). It is as long as the head behind the anterior margin of the eyes or the nostrils. The anal fin is short and its longest ray is shorter than that of the dorsal ; it rarely reaches the root of the caudal. The pectorals are considerably shorter than the head, nnt ni<jre than the length of the head from behind the middle of the eyes; they are sepa-
12
Figure 12. Lateral view of posterior part of body of a specimen of Schicothorax micropogon Heckel from Dal
Lake, showing form of anal and caudal fins. X 1/4.
rated from the origin of the ventrals by a distance equalling more than half their own length. The ventrals are slightly shorter than the pectorals and are separated from the insertion of the anal by a distance equalling almost half their own length. The caudal fin is somewhat shorter than the head, deeply furcate and appreciably higher than long with more or less equal lobes; the length of the central rays is contained nearly 2.5 times in the length of the outer rays (Text-figure 12). The vent is situated in front of the anal and in female speci- mens is provided with a prominent fleshy papilla. The scales on the body are minute. The tiled row of anal scales are small, the largest one being less than half as broad as the orbit.
The colouration of the specimens in alcohol is reddish brown with a darker upper sur- face. As in vS". planifrons some of the specimens are paler. The entire Iwdy excluding the ventral surface is dusted with fine black pigments. The dorsal and the caudal fins are dusky.
Bionomics: The peritoneum is black. The structure of the air-bladder is similar to that of S. planifrons. In specimen, 132 mm. long, the lengths of the anterior and the posterior chambers are 19 mm. and 31 mm. respectively. The alimentary canal is short and of mod- erate width with a few convolutions. Its length varies from 1.5 to about 2 times the length of the body excluding the caudal fin. The gut contents of 5 specimens from the Dal and the Manasbal lakes were examined, and the intestines were found to be partially empty, but small
346 REPORT ON FISHES. TART 11: SIS()Rin.\r. AND CYPRIN' IDAl'.
aiimuiit I if a snft liulp}' .substance lui.xcd willi IllamciitDUS alL;ac and frat;nii;nts ut in.scct larvae were found. The species is apparently a mixed feeder and the short length of the intestine suggests that the lish feeds chielly nn animal matter.
Brccdini^: So far as 1 can gather, nothing is known alxuit the hrecding habits of the species. Tt is, however, significant that all the female specimens obtained by the b'-xjiedition during the months of March to May bear mature eggs, the diameter of which is about 1 mm.
Parasites: No Acanthocephalan parasites were found in the intestine of the specimens examined, Init in a sjieeimen, 13J mm. long, from the Manashal lake several specimens of a species of Neniatode worm were found.
Pislrlhiition: Silii.::i>tlitini.v iiiicml^di^dii is one nf tin- smallest endemic s])ecies of the Kashmir valley, co-occurring with .S". planifrons in the laki's and the adjoining channels. It is known by the Kashmiris as "RaingJuirdi."
Mcasiirciiiciils in niilliiiichcs
Total length without caudal
Length of head
Width of head
Height of head
Diameter of eye
Length of snout
Intcrorhital width
Depth of body
Longest ray of dorsal
Longest ray of anal
Length of pectoral
Length of ventral
Length of caudal
Distance between ])ectt)ral and base of ventral
Distance between ventral and base of anal
Length of caudal ]ieduncle
Least height of caudal peduncle
SiiiirjatliDrdX curvifrons Ileckel
1838. Schizolhoiax curvifrons. Ileckel. l-isilic aits Cuscliiiiir, p. 25. pi. iii.
1844. Schizothorax rurrifroits, Ileckel. I'isclic Kasclniiir's in lliii^rrs Rrisc. p. ,^ol. 3 figs.
1868. Scliicotliorax cur'i'ifrous. (liinther. Cut. I'isli. Brit. Mux., VII, ]>. 1()4.
1877. Schicothorax curvifrons. Day, Fisli. India, p. 532 (footnote). .
1889. Schizothorax curvifrons, Day, faun. Hrit. Ind., I'ish., I, p. 252 (footnote).
The species is represented in the collection of the Yale North India Expedition by five young and half-grown specimens collected in March-May, 1932, from the following localities :
Srinagar (Sta. K7), ca. 5,200 ft 4 (110-178 mm.)
rhannel: Manashal Lake: Srinagar (Sta. K48), 5,197 ft 1 (122mm.)
|
? |
9 |
9 |
|
165.0 |
132.0 |
112.0 |
|
44.0 |
35.0 |
,30.0 |
|
27.0 |
21.0 |
17.0 |
|
31.0 |
21.0 |
20.0 |
|
9.0 |
8.0 |
7.0 |
|
14.0 |
11.0 |
9.0 |
|
14.0 |
11.0 |
9.0 |
|
40.0 |
32.0 |
25.0 |
|
32.0 |
26.0 |
21.0 |
|
28.0 |
21.0 |
16.0 |
|
28.0 |
21.0 |
21.0 |
|
25.0 |
20.0 |
20.0 |
|
34.0 |
28.0 |
25.0 |
|
21.0 |
16.0 |
11.5 |
|
15.0 |
11.0 |
6.0 |
|
25.0 |
19.0 |
19.0 |
|
14.0 |
12.0 |
11.0 |
REPORT ON FISHES. PART II : SISORIDAE AND CYPRINIDAE 347
Tn the speciim-iis before me the length of the head is coiitainetl ahout 4.5 times or a little over in the total length excluiling the caudal fin. The eyes are large and their diam- eter is contained from 4.5 to nearly 5 times in the length of the head. The snout is short and stump\- and 1.5 times longer than the orhit. The interorhital space is nearly 1.5 times the orbital width. 1"he barbels are very shtjrt and not more than half as long as the diameter of the eyes. The nmuth is somewhat narrow and horse-shoe-shaped. The lips are mod- erately developed and more or less tough. In some specimens the lower jaw is provided with a sharp margin.
The dorsal fin is inserted slightly nearer the root of the caudal than the tip of the snout ; its osseous spine is straight and fairly strong and has rather fine serrature on the posterior Ixirder. In the smaller specimens the anal fin laid flat does not reach the root of the caudal, whereas in the larger ones it almost reaches it.
The colouration of the specimens in alcohol is reddish brown with a slightly darker upper surface. The inner margin of the caudal fin is dusky.
Bionomics: The air-bladder is of the normal Sclii.'^otliora.v-type. It appears that in young specimens the posterior chamber is proportionately shorter than the anterior cham- ber. In a specimen, 120 mm. long, the length of the anterior chamber is 15 mm., while that of the posterior one is 20 mm. In a specimen, 178 mm. long, however, the correspond- ing measurements are 25 mm. and 50 mm. The length of the intestine is almost 3 times the length of the body excluding the caudal fin. The gut contents of 2 specimens (120, 178 mm.) were found to consist of fragments of some insect larvae mixed with a brownish pulpy substance.
Parasilrs: In the intestine of a specimen ( 178 mm. ) a few specimens of a species of .\canthoeephalan worm were found.
fh'.stribittidii: Sclii::i>thora.v curvifrons is one of the endemic species of the Kashmir vallew It grows to a fairly large size, weighing about 31bs. and is known by the Kashmiri fishermen as "Sattir."
Genus Orciniis IMcClelland (1839)
Oreinus sinuatus (Heckcl)
1838. Schizothorax sinuatus, Heckel, Fische aus Caschmir, \). 21, pi. ii.
1839. Oreinus maculatus, McClelland, Asiat. Res. XIX, pt. ii, pp. 274, 345, pi, Ivii, fig. 6. 1844. Scliizntlwrax sinuatus, Heckel, Fische Kascliniir in flu^i^cl's Rcisc, \>. 259, 3 figs. 1868. Oreinus sinuatus, Giinther, Cat. I'isli. Frit. Mus., VII, ]). 161.
1876. Oreinus sinuatus. Day, Proc. Zooi. Sac. London, p. 783.
1877. Oreinus sinmitus. Day, Fish. India, p. 529, pi. cxxiv, fig. 4.
1878. Oreinus sinuatus. Day, Sci. Res. 2nd Yarlcand Miss., Ichthyoloi:,v. p. 3. 1889. Oreinus .Kinuatus, Day, luiiiii. Urit. Iiid., Fish.. I, ]>. 248.
The Yale North India Expedition obtained 14 specimens of the species in March-June, 1''32, from the following localities:
Jheluni River, Srinagar (Sta. K 14) ; ca. 5.200 ft 1 (208 mm.)
"jhelum River, Pampur (Sta. K 16) ; ca. 5,200 ft 3 (270, 220, 220mm.)
Rivers and Canals, Srinagar 2 (280, 5 . 184 mm.)
Kargil, Ladak (Sta. K 88) ; 8,790 ft 5 (495. 270. 260, 180, 145 mm.)
Upper Indus, Spitok, Ladak (Sta. L 17) ; 10,730 ft 3 (280, 275, 180 mm.)
348
REPORT ON FISHES. PART II: SISORIDAE AND CYPRINIDAE
The speciiufiis under report agree tairly well with the (lescriptimi nf tiie speeies and exhibit weTl marked differences from the variety griffithi McClelland from Afghanistan, whirh has been recently redescrilied by Ilora." O. siitualus is very closely allied to O. pla- giostoiuiis, but differs from it chiefly in the structure of the serrated dorsal spine, which is long and strong with moderate serrations on its posterior border in sinuatiis and consid- erably weak with feeble or obsolete serrations in plagiostoiiius (Text-figure 13, o, b).
13
Figure 13. Dorsal spines of nearly equal-sized specimens of (a) Oreimis plagiostcmus (Heckel) and (6) Oreinus sinuatus (Heckel), showing difference in structure of spines and nature of serration on their posterior borders. X ca. 2.
Figure 14. Ventral views of head of nearly (■(|iial-sized specimens of (<i) Orciiiiis f'liif;i,isli'iiiiis (Heckel) and (b) Oreinus siiiiialus (Heckel), showing difference in gape of mouth and structure of lower lip. X 1>4-
Further, siuimhis has invariably a deeply concave margin of the lower li]) which is more or less straight or slightly concave in plagiostoiiius (Text -figure 14, a, b). Ihe anal scales in siniMtus are not well developed and are often olxsolete, the largest lx;ing nearly half as broad as the orbit, whereas in plagtostonius they are well defined and almost as broad as the orbit. From the Expedition material as also from the Indian Museum speciriiens from dif- ferent localities T find that in O. siiniafus the dorsal spine is somewhat variable in length.
" Hora, S. L.: Rcc. Ind. .Uxs., XXX\I. pp. 300-306, figs. 4 and 5 (1934).
REPORT ON FISHES. PART II: SISORIDAE AND CYPRINIDAE
349
It is nornially as Imii; as tlie head l)eliiii(l tlie nostrils, l^ut it may be as long as or a little longer than the entire head.
Bionomics: The perituneuni is iilack. The air-bladder, although bipartite is peculiarly modified. The anterior chaml)er is roughly knob-like with a broader anterior portion, while the jjosterior chamber is narmw in the lieginning, IjecDming broadest in the middle ami narnnver again towards the free purtinn and has thus a somewhat bulbular appearance. The posterior chamber is moderately thick-walled, specially near and behind the constric- tion between the two chamljers (Text-figure 15). The posterior chamber is nearly twice as long as the anterior one. The alimentary canal is capacious, long and much convoluted ;
16
FiGiiRK 15. Oicimis sinmilns (Heckcl). .\ir-liladder uf (a) a specimen, 275 mm. Idiig. from Spitok, Indian Tibet and (/>) a specimen, 255 mm. lung, fmm Kargil, Indian Tibet, showing structural variation. Nat. size.
it is about 4 to 5 times as long as the body e.xduding the caudal fin. The intestines of sjjeci- mens from Kargil and Spitok were examined and found to contain lumps of gravel and mud mixed with pulpy vegetable matter. A few chironomid larv-ae were also found.
Br(\'iiiii<i: No delinitc information is available in regard to the breeding habits of the species. Day, however, observed: "In May (at ("humba) the ova of these fish inhab- iting the main stream, were almost fully develo])ed, Ix'ing numerous and of large size, whilst there were a considerable number of fry in the side streams of the Ravi." Of the few Kxpedition specimens opened by me, I find in one case, a female 280 mm. Imig, that the eggs are fully developed and have a diameter of about 1.5 mm.
Distribution: Orcinus siniiatiis occurs in the rivers of Kashmir and the runjab, and prefers to live among rocks. In Afghanistan the species is replaced by the variety griffitlii
350
KErUKT UN FISHES. PART II: SISUKIUAE AiNlJ C\rRIMDAE
McClellatKl, O. iiuuiilaltis McClelland from the Kalml river'" (nrc Simla), as indicatt-d by Hora (op.cit.), is a young form of the var. griffitlii.
? Schizothorax labiatus >- Orriniis siiiiiatus
Tlybridisation among fishes in nature is not a rare occnrrenee and usually takes place between closely related genera of the same ancestral stock or l)et\vccn two congeners that co-occur in specialized and restricted areas, such as the high altitudes of Central Asia, etc.. and live under more or less similar cniiiliiiniis of life. Intermediate forms between Schico- tliora.v and Orciiiiis ha\c often been observed by various authors and recently flora''' has recorded a very interesting series from the Chitral valley which he has designated as ScJii::(>- tliom.v labiatus x Orcinus sitiuatus var. griffitlii. Among the fishes l)rought by the Yale
Figure 16, A scries (a-e) of vt-iitral biirfaco of head uf five specimens, intermediate lictwicn Schicnlhorax and Orcinus, riniglily showing progressive modifications from Schizoihorax lalnalns towards Orcinus si)ni(ilus type of month parts. (Figures variously magnified.)
North India i'.xpeditiou there is a series of specimens detailed below which undoubfedly re])- resent intermediate forms l)etween Schicotliorax and Orcinus and as far as can be judged friiui the nature <if the bead, the jaws and the lips, they seem to be hybrids U'tween .S". hibi- (ilits and (). siiiualus: they agree fairlv well with the series describee! li\- Ibira.
Srinagar (Sta. K 14) ; ca. 5,200 ft 1 (1X0 mm.)
Rivers and Canals, Srinagar 3 (223, lOO, 150 mm.)
Lake near Chushol (Sta. L 73) ; 14,735 ft 1 (215 mm.)
•= McClelland, J.: Calculla Jouni. Xat. Ilisl., II, p. 580 (184.'). "Ilora, S. L.: Kcc. Ind. Mus., XXXVI, pp. 3U7-J10, figs, / and 8 (1934).
REPORT ON FISHES. PART II: SISORIDAE AND CYPRINIDAE 351
These 5 specimens can roughly Ije arranged in a series showing progressive modifica- tions towards the formation of an Orcimis type of mouth ])arts (Text-figure 16, a-c). The specimen from Srinagar (Sta. K 14) represents true Schizotliorax lab'uitus type, while the one from the lake near Chushol (Sta. L73) almost approaches typical Oreimis simiatus type. The three specimens from the rivers and the canals show the successive intermediate stages.
It is a matter of congratulation to mention here that the striking peculiarities of these five specimens did not escape the notice of Mr. G. E. Hutchinson, who had rightly labelled tlu-ni in the fields as "Intermediates."
? Scliicolliorax planifrons x Oreiniis siniiafiis
Besides the five specimens of an intermediate type which I have designated as Scliico- thorax labiatus x Oreinus siniuihts, there are, in the collection of the Yale North India Expedition, six specimens collected from the following localities which correspond to S. planifrons in all essential characters except for the sharp, horny tubercles on the snout and the sharp, horny covering of the lower jaw. In one of his letters Mr. G. E. Hutch- inson informed us that this intermediate form "co-occurs with Scliizothorax csocinns and the Schizotliorax species called in Kashmir "Chiisli." The "Chiisli" of the Kashmiris is .S". planifrons and I am inclined to believe that the six specimens under report are hybrids between S. planifrons ("Chush" ) and U. sinuatiis with tlominant ''Chiisir characters. This statement finds sujjport in the fact that, according to Mr. Hutchinson, these intermediate forms are also called "Chnsh" by the Kashmiri fishermen.
The specimens have been collected from the following localities:
Main Canal : Srinagar (Sta. K 12) ; ca. 5,200 ft 2 (228, 215 mm.)
Jhelum River : Srinagar (Sta. K 14) ; ca. 5,200 ft 2 (290, 275 mm.)
Channel to Manasbal L^ke : Srinagar (Sta. K 48) ; 5,196 ft 2 (370, 150 mm.)
Scliizopygopsis stoliczkac Steinclachner
1866. Schisopygopsis sloUczkac, Steindachner, Vcrh. Zool.-hot. Gcs. IVicn, XVI, p. 786, pi. xvi,
fig. 2. 1868. Schizopygopsis stoliczkae, Giinther, Cat. Fish. Brit. Mus., VII, p. 170. 1876. .Schicopygopsis stoliczkae, Day, Proc. Zool. Soc. London, p. 791. 1878. Schizopvi^op.'ds stoliczkae, Day, Sci. Res. 2nd Yarkand Miss., Ichthyology, p. 9, pi. ii,
fig. 2. "' 1878. Schizopygopsis stoliczkae, Day, Fish. India, p. 531, pi. cxxiv, fig. 2. 1889. Schizopygopsis stoliczkae. Day, Faun. Brit. Ind., Fish., I, p. 251, fig. 89. 1801. .Schizopygopsis stoliczkae, Herzenstein, IViss. Result. Przc'a'alski Cenlral-Asicn uuter
Rei.s-en, Fi.uhe, p. 191, pi. xvi, fig. 3. 1898. .Schizopygopsis stoliczkae, Alcock, Report J\at. Hist. Res. Pamir Bound. Coiuiii., j). 14. 1907. Schizopygopsis stoliczkae, Berg, Ann. Mus. Zool. Petersb., X (1905), p. 323.
1910. .Schizopygopsis stoliczkae, Zugmayer, Zool. Jalirb. Abth. Syst., XXIX, j). 290.
1911. Schizopyi^opsis stoliczkae, Stewart (in part), Rec. Ind. Mus., VI, ]). 73, ])1. iii, figs. 1-3. 1914. Schizopygopsis stoliczkae. Berg, Faune dc la Russic. I'oissnns. HI, \). 702.
1916. SchizopVij^of^sis stoliczkae. Berg, Poi.ss. des Faux Douces dc la h'lissie, ]>. 290.
352 REPdkT ON FISHES. I'ART II: SISORIDAE AND CYPRINIUAE
1916. Sclii:iopyi^of^sis slolicchac, N'incitvuerra, ./)/)). ^[us. Civ. Slor. \'al. Geneva, XLVIT, p. 143. 1932. ScUhot>yi^opsts stolkckiie, Berg (in part), Poiss. dcs liaii.v Doiici'S dc !'U. R. S. S. (3rcl
ed.), pt. i, p. 473. 1935. Scliicopy};of^sis slolicskae, Hora and Wukerji, in J'isscr's Karahontiii, I, j). 434, pi. iii,
figs. 1-4."
Aniont^ the Schizothdracinae of Central Asia Sclii.zopyi^opsis stolicckac is one of the most variable forms, particularly in regard to size and the proportions of the height of the body, and of the length, breadth and the height of the liead in relation to the length of the body. The diameter of the eyes in proportion to tlie length of the head is also considerably variable. The species is generally uniform in colouration, but spotted and l)lotched forms are not rare. In view of considerable variations of a nmnber of characters — which again are not infrecpiently inconstant even in series of si)ecimens from any ])articular area — a very wide interpretation has been given to i". stolic::ktu' by certain authors, while others form restricted groups of e.xtremc and intermediate variations which may almost be regarded as distinct taxonoiuic entities.
In I'Ul, Stewart {op.cit.), while reporting the species from various localities (from 11,500 ft. to 15,000 ft.) lietween the Chumbi valley and the town of CJyantse in the TiU-tan province of Tsang,'^ took opportunity of e.xamining the extensive collection of S. stolic.^kur preserved in the Indian Museum. Liesides s])ecimens from various localities, the Indian Milium colifction includes tlie topotyijcs of tiie species wiiich were obtained by Dr. Stoliczka from the Pamirs and Ladak during Sir ]>niglas Forsyth's Mission to \'ark:ni(l in 1873-74, as also those of i". sc-^'crcowi from the I'amris.
Uerzenstein's species, A\ sc2'cr::o7in, may be superficially separated from S. stt'lic::kae l)y its somewhat smaller size (dwarf), flattened lower surface of tiie head instead of Ijeing curved, and by the position of its mouth which is somewhat overhung by the snout and is more terminal than ventral. iUit Stewart has shown, after a \ery detailed analysis of the so-called characteristics of the two forms that all i)rol)al)le gradations exist between the typical form of stolicckae and the sei'cr::oiin type, so much so that specific distinction seems hardly justifiable.
In this connection mention may also Ik- made of the Seistan form of stulircknr reported by .Annandale and I lora.'"' In all probability this is yet anotluT dwarf race, distinct Iroin both the typical slolicskac and scz'crcoun. Recently, while ilealing with the material obtained by the Netherland Karakorum b'xpedition, Hora and Mukerji'" have intlicated, after thor- oughly examining the entire collection of S. stolicckar in the Indian Museum, that l)oth severzowi and the Seistan form should Ik.- considered at least as separate subspecific forms. But unless further material from tin- Pamirs and .Seistan l)ecome available, no definite conclusions are possible.
The Yale North India Expedition obtained 17 specimens from the fnlldwing localities in June-August, 1932, which, in my o])inion, are .'dl referable to the typical form of stoliczkae. The specimens are of variable sizes, the largest one from .S])itnk being about 300 mm. long excluding the caudal fin.
" "The province of Tsang lies roughly iiortli of Sikhim and Nepal and imiudi-s the di.strirt from Tang-la to Gyantse and Shigatse."
"Annandale, N., and Hora, S. L.: Rec. Intl. Mus., XVIII, pp. 173, 174, figs. 7a, 7h, 7c (1920). "Hora, S. L., and Mukerji, D. D. : in Visser's Karakorum, I, pp. 434, 435, pi. iii.
Report on fishes, part ii : sisoridae and cyprinidae 353
Spitok, Upper Indus : Ladak (Sta. L 17) ; ca. 10,730 ft 2 (300 mm. ; 1 10 mm.)
A large rapid stream between Tangtse and Mugleb: Ladak (Sta. L37) ; ca. 13,700 ft 6
Stream above Lukong; Ladak (Sta. L 54a) ; ca. 14,164 ft 3 (50-72 mm.)
Stream into Pangur Tso; Ladak (Sta. L 74) ; ca. 14,203 ft 1 (77 mm.)
Yalapuk, Upper Indus near Nyoma ; Ladak (Sta. L 79) ; ca. 13,521 ft 2
It is apparent from the a!jove detailed distribution that only the young and small adult specimens were obtained by the Expedition from the shallow streams and pools in the vicinity of Pang-gong and Pangur lakes, while the larger individuals were taken from the Indus river itself. This suggests that either the species breeds in the shallower waters of the streams and pools that are associated with the lakes and the river, or the young forms migrate to the former habitat for the opportunity of having better food supply and other more favoural>le conditions of life that naturally prevail there. The former view finds support in the fact that the Yale North India Expedition collected mature eggs of 5". stoHczkae from the streams between Tangtse and Mugleb." The eggs are comparatively large, their average diameter being 2 mm. Further, the Second Yarkand Mission also obtained "fry and small fish from Lukong and Chagra" (Day).
Breeding: Steivart {op. cit.) has observed that the "breeding season (of S. stoUczkae) in tiie neighbourhood of Gyantse appears to occur about June. In the less favoured waters near watershed it is probably later." The mature eggs of the species referred to above were collected by the Expedition on the 27th June, and indicate that the breeding of the species in Western Tibet probably occurs at almost the same time as in Eastern Tibet.
Bionomics: The peritoneum is black. The air-bladder of ^. stolicckae, as is character- istic of the cyprinoid fishes, is free in the abdominal cavity, extensive, and bipartite ; the posterior chamber is longer and somewhat narrower than the anterior one which is short and rounded and marked off from the former by a deep constriction. A well developed pneu- matic duct opens at the junction of the two chaml^rs (Text-figure 17). It is interesting to niite that unlike the cyprinoid fishes of the typical calm waters, such as lakes, ponds, etc., both the chambers of the air-bladder of S. stolicckoe are very thick-walled. This type of air-bladder is to be expected in this species in so far as "its characteristic habitat is in the streams and small rivers of the (jpen, tree-less, flat, grassy uplands at an elevation of 11-16,000 feet; broad valleys in the centre of which a river runs witli moderate rapidity, while on either side are marshy pools fed by springs, shallow ponds or irrigated land." (Stewart, op. cit. ). i\ stolicchac, owing to its being a denizen of rapid waters and in con- sequence of ground habitat, does not use the air-bladder as a hydrostatic organ, and its wails, therefore, become more and more thick.
The alimentary canal is much convoluted; in young and half-grown specimens its length is a little less than twice the length of the body including the caudal fin, while in adult individuals it is just a little more than twice. The stoiuach contents of some of the young specimens obtained by the Vale North India Expedition show that, in young and immature individuals at any rate, S. stulic::kcie, like the young of most of its allies, feeds on nymphs and lar\ae of insects, such as I'lecoptera, Diptera, etc. The examination of the gut con-
" The piipulaticiii in tlie Chagra stream is isolated, ami it is clear that its largest members never reach the size of the largest specimens from the Indus. An operculuiii froiu the shore of Tso-Nyak suggests this species, and indicates a very large fish. — G. E. H.
354
RT.rORT dX FISHES. PART II: SISORIDAF. AND CY I'RI NIUAK
tents (if tlii^ adult specimens, Ikiwcvct, slmus tliat tin- siK-ries, wlicn full s^niwn, feeds exclusively on vegetable matter.
Parasites: Day already observed in ctinnection with the specimens obtained by the Second Yarkand Mission that "these fishes ai)i)ear to be much attacked by parasites, which occasion yellowish elevated tubercles, not only on the head and body but also on the dorsal fin." The specimens under report not only show similar parasitism but in the intestine of certain si)ecimcns taken at the streams l^etween Tangtse and Mugleb a species of Acantho- cephalaii worm has also been fmuid. I'^rdUi a single specimen as many as eleven worms were collected.
t^.
/:
17
Figure 17. Schhopygo/'sis stolicckac Steiiidaii.iicr. Air-ljladdcr of (a) a young specimen, 72 mm. lung, from a stream above Lukong, Indian Tibet, and (/)) an adult specimen, 300 nun. long from .Spitol<, Indian Tiliet. Nat. size.
Distributii'ii: The species is widely distributed "from I'.adak.shan and the Pamirs to the Eastern Ilim.ilaya including the u])pcr waters of the O.xus, Indus, Sutlej and Urahmaputra ( Tsauj^-iio 1. (_)n the south face of the llimahiya it has hitherto lieen found in the Chumbi Valley."
(lenus r>!/^lychiis .Steind.achner (1866) Pipl yell IIS iiKiciihiliis Steindaclmer
1866. Diptychtis maculatus, Steindachner, I'l-iii. Zool.-hul. (,'cs. W'icii. X\'l. y. 7i:^'f>. fig. 6. 1868. Diptychus macitlalus, Giinthcr, Cat. l-'isli. Brit. Mtis., VII, \>. 171. 1876. Diptychus imiculalus, Day, Proc. Zool. Soc. London, p. 272.
REPORT ON FISHES. PART II : SISORIDAE AND CYPRINIDAE 355
1878. Diplyclnis iiuicitlatiis. Day, Sci. Res. 2nd Varkand Miss., Ichthyology, ]t. 10, pi. ii, fig. 3.
1878. Viplyiliiis iiiaculahts, Day, Pish. India, p. 534, pi. cxxiv, fig. 3.
1889. Diptyclius inaculatiis. Day, Faun. Brit. Ind., Fish., I, p. 255, fig. 92.
1910. Diptychns maculatus, Zugmayer, Zoo!. Jahrb., Ahth. Syst., XXIX, p. 292.
1914. Diptyclius inacuhitus. Berg, Faitnc de la Russie, Poissons, III, p. 677, figs. 133-135.
1916. Diptyclius viaciilatus, Berg, Poissons des Eau.v Donees de la Russie, p. 286.
1916. Diptyclius maculatus, Vinciguerra, Ann. Mus. Civ. Stor. Nat. Genova, XLVII, p. 145.
1931. Diptychns maculatus. Berg, Zool. Anz., XCVI, p. 311.
1932. Diptychns maculatus, Berg, Poiss. des Faux Donees de I'U. R. S. S. (3rd ed.), pt. i, p. 466.
1935. Diptyclius iiiaciilalus, TTora and Mukcrji, in Visscr's Karakoruin. I, p. 435.
Dipl\clius maculatus is a very widely distributed species in Central Asia, usually inhab- iting rocky, niuuntainiius streams. It exiiibits considerable variations in different body pro- portions, nature of the barl)els, tiie diameter ui the eyes and tlie colouration. The Yale North India Expedition obtained 16 specimens of the species in May-July, 1932, from the following localities:
A rapid stream at Nimu: Ladak (Sta. L 12) ; ca. 10,250 ft 4 (123 mm. ; 33-60 mm.)
A torrential .stream at Lhabaps: Ladak (Sta. L31); ca. 11,855 ft 4 (108 mm. ; 27-32 mm.)
A large rapid stream between Taiigtse and Mugleb: Ladak (Sta. L 37) ; ca. 13,700 ft.
2 (118 mm.; 107 mm.)
Stream at Chagra: Ladak (Sta. L46); ca. 15,215 ft 2 (270mm.; 225 mm.)
Stream at Leh 5 (95-135 mm.)
From the al)ove list of localities it will be seen that the majority of the specimens collected by the Expedition are either young t)r half-grown and they all come from rapid to toi-renlial streams. The two specimens taken at Chagra are large; the larger one l>eing 27U mm. in length excluding the caudal fin.
Preeding: The breeding season of I>. inaculatus is not definitely known so far,''* but from an examination of the nature of the gonads of a number of specimens brought back by the Yale North India Expedition, it appears that the spawning of the species takes place, like Schicopygopsis stoliczkac, during the months of May to August. Both the specimens taken at C'hagra are gravid females.
lHonoiiiics: The peritoneum is black. The air-bladder is more or less similar in struc- ture to that of .Schi::opygopsis stoliczkac: (Text-figure 18). The alimentary canal is much convoluted. In young and half-grown specimens its length is alxiut twice the length of the body including the caudal fin, while in larger individuals it is more than twice. The gut contents of a numlK'r of specimens under report indicate that the young immature stages fi'ed on nyni]>bs and l;irvae of b.pbimeroiiterous, I 'lecopterous and r)i])terons insects, while llu' aduhs li\e cbietly on slime, filamentous algae, and other vegetable matter.
I'lirasitcs: Like .Sclii.':iipyg(ipsis slalic.ckar this species is also attacked with parasites which cause brownish tubercK's on the head, cheeks, different parts of the body and the fins. No intestinal jiarasitcs were found in the specimens under report.
'" V. .\. .Aiiikin'.s work, "Die Fische dcr Gattimg Diplychus Steincl., ilire Sy.stematic und biologisclie BedcutuiiK" {Tonuli, 1900), seems tu contain interesting biological accounts of Diptyclius. hut unfortunately it is nut available in India.
356
REPORT ON FISHES. PART 11: SISORIDAE AND CYPRINIDAE
18
Figure 18. Diptychus maculalus Steindachner. Air-bladder in different stages of growth. X l/^- (a) A specimen, 220 mm. long, from Cliagra, Indian Tibet. (/)) .'\ specimen, 138 mm. long, from Leh, Indian Tibet, (f) .K specimen, 72 mm. lung, from Lukong, Indian Tibet. ((/) A si)ccimen, 60 mm. lung, frum Nimu, Indian Tibet.
Distribution: D. imiciilalus is one of tlie iimst coniinoii species foiiiul in the Iiulus, Tarim and the Yarkand river systems. It has also l)een fcuind in otlier parts of TilxH and Nepal. It is an inhabitant of the rapids.
Roiiarks: Reference may here be made of a species, "Diptyclius aiinumlalci" described by i\euan''' from Katamundu in Nepal. I have shown elsewhere"" that the form is con- generic with Siliiciilhoni.v and that tiie description is based on juvenile specimens.
Genus Ptyclwbarbus Steindachner
Ptychobarbus conirostris Steindachner
1866. Plycliobarbiis conirostris. Steindachner, Verb. Zool.-bot. Gfs. Jl'ioi, X\'I, ]). 790, pi. xvii,
fig. 4.
1868. Ptychobarbus conirostris, Giinther, Cat. Pish. Prit. .Mas., \ll, p. 169.
1876. Ptychobarbus conirostris, Day, Proc. Zool. Soc. London, p. 789.
1878. PtycliolHirbits conirostris. Day, Sci. Res. 2nd Yarkand Miss., Ichthyology, p. 7, pi. ii, fig. 3.
1878. Ptychobarbus conirostris, Day, Pish. India, p. 5^3. \A. c.xxv, fig. 3.
1889. Ptychobarbus conirostris, Day, Faun. Brit. Ind., Fish.. 1, ]>. 253, fig. 91.
'Regan, C. T.: Rec. Ind. Mhs.. I, p. 158 (1907).
'Mukerji, D. D. : Rcc. Ind. Mus., XXXIII, pp. 63-65, figs. 1-4 (1931).
REPORT ON FISHES. PART II : SISORIDAE AND CYPRINIDAE 357
1898. Ptychobarhiis coniroshis, .'Mcuck, Rep. Proc. Pamir HohihI. Coiiiiit., p. 37, pi. i, figs. 2
and 3.
1910. Ptycbobarhus conirostris, Ziigniayer, Zool. Jahrh., Abtli. Sysl.. X.\IX, pp. 291, 292.
1935. PtycJiobarbiis conirostris, Hora and Mukerji, in Visscr's Karakonmi. I, p. 436.
The species is represented in the collection of the Yale North India Expedition by five specimens collected in June-August, 1932, from the following localities:
Kargil : Ladak (Sta. K 88) ; ca. 8,790 ft 1 (262 mm. ; 9 )
Kalatse, Upper Indus : Ladak (Sta. L 6-9) ; 9,700 ft 1 (174 mm. ; 5 )
Spitok, Upi>er Indus: Ladak (Sta. L 17) ; ca. 10,730 ft 2 (270mm. 9 ; 250mm. S)
Yalapuk, Upper Indus near Nyoma: Ladak (Sta. L79); ca. 13,521 ft 1 (100mm.)
In the report on tlie fishes obtained I)y the Netherland Karakorum Expedition Hora and Mukerji (up.cit.), on the authority of Mr. G. E. Ilutcliinson, stated tliat rtychobarbiis is usually found in large, deep and rapid-flowing rivers and seldom in small, clear streams or springs overgrown with vegetation. The above list of localities corroborates this view inasmuch as all the specimens of P. conirostris collected by the Expedition come from rivers. Aicock ( op. cit. ), during the Pamir Boundary Commission, collected specimens of the species, both males and females, from the Yasin river at an altitude of about 8,500 feet between Kashmir and the Pamirs. This adds further support to the statement that P. conirostris is essentially a mountainous river form.
The species is variable specially in regard to the size and proportions of the head and the eyes. In young and half-grown specimens the diameter of the eyes is contained nearly four times in the length of the head and 1.5 times in the length uf the snout; in fairly grown up individuals, on tlie contrary, tiie proportions are 6.5 times and 2.5 times respec- tively. The interorbital space is equal to the diameter of the eyes in young specimens, l)ut in adults it is much wider, being from 1.5 to 2 times the orbital width.
The head is slightly higher than or as high as broad. Its width is equal to tiie length f tiie head behind the middle of the eyes. The length of the head is contained nearly five times in the length of the body including the caudal fin. In young and half-grown indi- viduals the caudal fin is almost as long as the head, but in adults it is much shorter, being e(|ual to the length of the heatl behind the nostrils. In young stage the barljels are equal to the orbital width, while in full-grown specimens they are twice the same.
.\ well-developed thick fleshy appentlage is present in the axillae of the ventral fins. In Scliizopygopsis stoUcsIcae this structure is poorly developed, while in Diptyclnis nuiciilatus it is hardly present. The .scales on the chest are considerably reducetl.
The colouration of the specimens in alcohol is pale brown with a silvery sheen. The upper half of the body, the head and the back are comparatively dark. The upper surface of the head is spotted with black. The back and the upper half of the body are somewhat irregularly marked with black patches which give the fish a characteristic maculated appear- ance. In young and half-grown sinH'imens these markings show a tendency to form reticu- lations, while in adult iixlividuals the)' represent series of small stars. All the lins are marked with black sjjots and small blotches.
Sc.viKil J>iiiiorpltis)n: h'rom an examination of the specimens of P. coniroslris taken b\' him ,it the ^'asin river, Alcock {i)p.cit.) observed that the females "are singular in having
o
358
REPORT ON FISHES. PART II: SISORIDAE AND CYPRINIDAE
J
20
Figure 19. Ptychobarhus coniroslris Steindachner. Lateral views of anterior portion of body of a male specimen, 250 mm. long, from Spitok, Indian Tibet, and a female specimeti, 262 mm. long, from Kargil, Indian Tibet, showing sc.xnal differences in structure of upper lip and curvature of snout. X H-
Figure 20. Air-bladder of a female specimen of Ptychobarhus coniroslris Steind., 262 mm. long, from Kari;il, Indian Tibet. X fi-
the iipi)er lip L;rcatly l>r(ia(lciied anil thickened and the pnifilc of the snout is conspicuously concave." Similar secondary sexual characters are al.so to he found in the female specimens of the species hrought back by the Yale North India I'^xpedition (Text-figure 19, a, b).
Bionomics: The peritoneum is black. The air-bladder is more or less similar to that of Diptychiis viacidalits. Its ]iosterior chamber is greatly elongated and extends as far back as the rectal end of the intestine. In a female specimen. 26J mm. long, excluding the caudal fin, the length of the anterior chamber of the air-bladder is 25 mm. and that of the posterior chamber 88 mm. (Text-figure 20). The alimentary canal is much convoluted: its length in grown-up specimens is 1.2 to 1.5 times the length of the body including the caudal fin. In the gut contents lumps of slime and algae mixed with mud and sand, as also large ninnbers of Chirononiid larvae were found. It appi'ars probalile that /', Cdniroslris is a mixed feeder.
Lhstribiition: I', cdiiirdslris has so far been rejiorted from the headwaters ni the Indus and the western border of Tibet.
Rriiiarks: Herg'"' regards Ptycliobdrhiis as a subgenus of nifitxrlnis and gives the following synopsis for the separation of the various subgenera:
""Berg, L. S.: Fainic tic hi Russic, Poissoiis, III. p. 6/7 (1914).
I
REPORT ON FISHES. PART II : SISORIDAE AND CYPRINIDAE 359
A. Lower jaw provided with sliarp, Imi-iiy coverins;'. liody densely covered with scales above the lateral line Piplyt'lnis (s. str.)
B. Lower jaw without sharp, horny coverins;'.
a. Body almost naked; scales present alon.^' lateral line and hase of pectoral fin
(iyiiniodiplycliits
b. Body covered with scales Ptychobarbus
In the character of the inoutli, the lower jaw, the lips and the squamation, however, Diplychus differs so markedly from Ptychobarbus that I am unable to agree with Berg, and consider Ptychobarbus to be a distinct genus.
rilE TUTTLE, MOKEIIOUSe & TAVLOR COMFANY. NEW HAVEN, CONN.
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