XI E> RAHY OF THE UN IVER.SITY Of ILLINOIS 590.5 FX v.3L Cop. 5 NATURAL HISTORY SURVEY NATURAL HISTORY SURVEY LIPR*RY »3 FIELDIANA • ZOOLOGY Published by CHICAGO NATURAL HISTORY MUSEUM Volume 31 May 23, 1951 No. 45 ON THE CLAUSILIIDAE OF PALESTINE Georg Haas Department of Zoology, Hebrew University, Jerusalem INTRODUCTION While Pallary (1939) mentions thirty-five species of Clausiliidae from Syria (that is, the entire Levantine region of the Mediterranean Sea), and while more than forty species have been reported from there, this group is only sparsely represented in Palestine as defined today. Pallary also stresses the remarkable fact that the majority of species are confined to very restricted areas, and since the informa- tion about localities in the older literature is frequently inaccurate the rediscovery of a particular form seems practically impossible. This inaccuracy is due partly to arbitrary definitions of the territories known under the names of Palestine, Syria, Lebanon, and Trans- jordania, and partly to the repeated use of certain locality names (for example, the name of Jaffa). Concerning Palestine proper, the literature is full of such mistakes, and our knowledge is furthermore confused by the discontinuous distribution of these snails. I shall characterize the biotopes and deal in full with the six species represented in the collections of the Hebrew University: Laciniaria moesta Ferussac, Albinaria elonensis sp. nov., Cristataria petrboki Pallary, C. genezarethana Tristram, C. davidiana Bourguignat, and C. prophetarum Bourguignat. When collectors are not named, the specimens were collected by myself. According to the literature, Cristataria medlycotti Tristram and Laciniaria hierosolymitana Bourguignat, type locality Jerusalem, should be added to this list. Although I live in Jerusalem and have been searching the environments consistently for years, I have not been able to find this latter species. I am fully aware that the Clausiliidae of the whole of the Levan- tine region should be treated together. I was obliged to restrict myself to Palestine owing to the lack, in our collections, of specimens from the adjacent countries and because I have no personal acquaint- No. 668 479 THEIHWWYOFTK JUL 2 0 1951 -Natural History Survey UNjvertv w *i!ws library © JERUSALEM Map of Palestine, showing localities mentioned in text. Laciniaria (Elia) moesta Ferussac Albinaria (Filumna) elonensis G. Haas Cristataria (Strangulata) petrboki Pallary Cristalaria (Strangulata) genezarethana Tristram \7/\ Cristataria (Strangulata) davidiana Bourguignat O Cristataria (Strangulata) prophetarum Bourguignat 480 © .3 HAAS: CLAUSILIIDAE OF PALESTINE 481 ance with the localities concerned. A full treatment of the Syrian species seems the more desirable, as many details of the morphology of their shells and their clausilia have never been cleared up. The majority of the species of Syrian Clausiliidae that have been described so far are found in the northern and central part of a rather restricted coastal area; we may presume that the surroundings of Beyrouth represent the last crowded outpost on the way to Palestine. Only two species of the genus Cristataria are mentioned as occurring south of Beyrouth: C. medlycotti Tristram (Sarepta, Sidon) and C. gom- baulti Pallary (Vallee du Dahmour, between Saida and Beyrouth). I do not think that these forms enter the region of Palestine proper, but I should like to stress the fact that the region of sparse occurrence of this group begins in Lebanon, rather far from the Palestinian boundaries. If a belt of, say, thirty miles of the coastal region north of Palestine is included in the present survey, the above-mentioned species should be added, as well as some species of Laciniaria. Cristataria gombaulti is not represented in our collections, but we have C. medlycotti from the Litani Valley, about 25 km. from the coast (coll. Moscona). Chicago Natural History Museum has in its collection of mollusks (no. 8063) a paratype of this species. Tristram's original label gives "Galilee" as locality, a very inaccurate geographical conception that may have been used in a very wide sense, so as to include even the locality of Surafend or the Litani gorge. This information has been kindly supplied by Dr. Fritz Haas. KEY TO THE PALESTINIAN CLAUSILIIDAE A. Aperture of shell provided with numerous marginal teeth and several (about 5) clearly visible palatal folds; shell dark brown Laciniaria moesta AA. Aperture of shell without marginal teeth; palatal folds invisible from with- out (with the possible exception of the principal fold). B. Shell smooth (hand-lens enlargement shows a very fine and dense stria- tion), whitish; aperture fused to penultimate whorl. . . .Albinaria elonensis BB. Shell ribbed, gray. C. With complete supraperistomial ridge Cristataria medlycotti CC. Without complete supraperistomial ridge. D. Whorls swollen, ribs very strong and protruding, with large inter- stices; aperture not fused to penultimate whorl. . .Cristataria petrboki DD. Whorls only slightly swollen, ribs densely set, slightly prominent. E. Cervix descending to the ventral border of the aperture without abrupt flexure; mouth gaping, peristomial border expanded, but not dorsally reflected; aperture only slightly detached. Cristataria genezarethana 482 FIELDIANA: ZOOLOGY, VOLUME 31 EE. Cervix abruptly descending to ventral border of aperture, forming an abrupt flexure; rim of aperture expanded, funnel-shaped; dorsal border of aperture free. F. Whorls of last two-thirds of shell flattened, the suture at the bottom of a narrow furrow; cervix strongly contracted; ribs of lower whorls delicate; aperture free, strongly deflected from penultimate whorl Cristataria davidiana FF. Whorls of lower two-thirds of shell rather swollen, the suture at bottom of a gradually deepening furrow; cervix moderately con- tracted; ribs of lower whorls coarser than in davidiana; aperture free but less deflected from penultimate whorl. Cristataria prophetarum Laciniaria (Elia) moesta Fe>ussac. Figures 90 and 96, A. Clausilia moesta Ferussac, 1821, Tableaux Syst., No. 539; Rossmaessler, 1839, Icon., 2, fasc. 3-4, p. 23, pi. 48, fig. 634. Clausila (Euxina) moesta 0. Boettger, 1877, Clausilienstudien, p. 86. Laciniaria (Elia) moesta Pallary, 1939, Mem. Inst. Egypt., 39: 66. Clausilia saulcyi Bourguignat, 1853, Catal. rais. moll. Saulcy, pp. 50-51, pi. 2, figs. 7-9. Laciniaria (Elia) saulcyi Pallary, 1939, Mem. Inst. Egypt., 39: 65. All specimens at my disposal are distinctly ribbed and are provided with the small teeth at the lower edge of the aperture and with an angular flexure at its base; the aperture is adnate to the penultimate whorl. All of these features are reported by Pallary as characteristic of saulcyi. In some specimens the aperture appears more roundish, but there is always the basal flexure at the border of the aperture. The color is brown to brownish-violet; the initial whorls, the mouth rim and the interior of the aperture are whitish; by reflected light the shell appears iridescent, in bronze hues. The morphological details of the aperture and the internal folds in this species are particularly complicated and rather variable. The aperture carries, to the left of the upper lamella, a rather narrow sinus extending upward; the lower margin is angular. The palatal rim is beset with a row of eight to ten marginal teeth, some of which may shift a little inward. There are five very marked palatal folds. The upper ones, which begin at some distance from the row of marginal teeth, run close together and are parallel in their external portion. The third fold starts as a thickened ridge, but is soon reduced to a thin ledge resembling the upper two. The fourth fold is situated in the left-hand margin of the angular lower peristomial corner; it is directly connected with a marginal tooth and rises, S-shaped, ending just before the inner end of the third HAAS: CLAUSILIIDAE OF PALESTINE 483 fold (cf. below). The fifth fold turns inward and upward and becomes confluent with the columellar lamella, constituting, so to speak, a continuation of the latter into the lowermost palatal corner, Fig. 90. Laciniaria (Elia) moesta Ferussac, from Mount Carmel. A, Front view of shell; X 5. B, Clausilium; about X 20. C, Aperture; about X 20. where it describes a short arc to the right of the lower peristomial angle. On the inner end of the marginal teeth there may be a variable number of short crests, interposed between the constant palatal folds (e.g., in one specimen, folds 1, 2, crest, 3, crest, 4 and 5). From eight to eleven wrinkled ridges extend inward from the inter- lamellar space, exhibiting many individual variations. The upper ones converge toward the obliquely descending lateral (outer) crest of the lower lamella, which bifurcates internally, the massive main portion running left (internally) of the axis of the shell and overlying the fusion between the fifth palatal fold and the columellar lamella. 484 FIELDIANA: ZOOLOGY, VOLUME 31 The upper lamella rises abruptly to its maximum height and slowly falls off internally, turning through half a whorl. From its middle, downward, it is accompanied laterally by the sharp-edged low colu- mellar fold, which also describes half a whorl. The spiral line of the upper lamella is apically continued as a detached crest of variable development. In one case the inner end of the spiral lamella projects about a third of its length beyond this accessory upper lamella. In another case I found (proximally of the axis) a tiny portion of the accessory upper lamella, the inner portion of the true upper lamella being reduced over a certain length to a low calliform crest, rising farther inward to its former height; it is thus divided into three portions. The spiral lamella becomes higher toward the interior and falls off steeply in its terminal portion. The first palatal (principal) fold runs approximately opposite the columellar fold, a relatively broad space being left between the two. The inner end of the fourth palatal fold rises in a shallow S-shaped curve toward the end of the third fold. Judged by its relation to the clausilium, this portion of the fourth fold represents the lunar fold; it ends just before and below the end of the third palatal fold. The ends of the four upper palatal folds are so arranged that the uppermost projects farthest inward, the second somewhat less, the third again less, etc. All terminal portions converge. The margin of the clausilium is entire, the sides of the broad terminal wing being almost parallel. The right margin forms a vertical lobe, projecting inward. The lower end tapers symmet- rically, both margins being pronouncedly thickened. The terminal portion of the clausilium is especially massive, as compared with the other Palestinian species. Distribution. — Syria, Palestine, Asia Minor (Westerlund) ; Brou- mana (600-800 meters), Jaffa, Beyrouth, Amchit in Lebanon, Asia Minor (Germain); Beyrouth Lighthouse, Tartous, mouth of Nahr el Kelb (Pallary, 1929); Jerusalem, Carmel, Beyrouth (Pallary, 1939). Specimens. — The specimens in the Hebrew University Collection are from the environment of Haifa, slopes of the Carmel; wadis in the area of Carmel Cape (Wadi Tartar); wadis opening into Haifa Bay (Wadi Rushmieh) ; wadis descending to the sea, south of Carmel Cape; Elon and Hanita, near the northern frontier of Palestine (the most easterly Palestinian locality of this species). Habitat. — Both the Carmel and the northern frontier are situated in Mediterranean oak forest, on rugged Cenomanian limestone, and HAAS: CLAUSILIIDAE OF PALESTINE 485 belong to the Palestinian regions of maximal rainfall (700-1,000 mm. per year). The animals are usually found under larger stones or at the foot of well-shaded rocky precipices, either crawling up the wall or under stones beneath it. This species appears to be negatively phototropic and favors moist places. Certain localities reported from Lebanon are from 600 to 800 meters above sea level (Broumana), while our highest locality, Elon, is about 450 meters above sea level, and others, as Wadi Rushmieh, are hardly more than 100 meters. These localities probably represent the southernmost outposts of a rather wide area of distribution, unless the locality Jerusalem should prove correct after all. Germain figured specimens said to be from Jaffa. If the well-known town in the coastal plain, and not another place with this frequently used name, was meant, this would appear a most unlikely and even impossible site. All the neighborhood is characterized by sand dunes, which cannot constitute a suitable biotope for a species bound to shade and moisture, and there are no limestone rocks. Pallary (1939) reported specimens from the sur- roundings of Jerusalem, the supposed original locality. The speci- mens may be referable to hierosolymitana Bourguignat, but this does not follow clearly from the context. The nature of the soil and the quantity of annual rainfall around Jerusalem would not exclude the species. However, in spite of searching for years, with the co-operation of many helpers, I have not been able to find any clausiliid near Jerusalem. Westerlund's reference to this variety (Jerusalem and Antiochia) only adds to my doubts. Discussion. — Pallary (1929) figures a specimen of hierosolymitana that differs from my specimens by the smaller size and by the very faint ribs, which become somewhat more distinct near the sutures only. The heights of the three specimens measured from the photos are 12, 10.6, and 9.1 mm., respectively. Since our collection contains a very similar specimen measuring 11.4 mm. from Sidon (Lebanon), it does not appear unlikely that by a mistake specimens from the north were wrongly labelled. According to Germain, this species may reach a length of 17 mm.; his figured specimen from Jaffa measures 14.5 mm. and is not very distinctly ribbed, resembling hierosolymitana and the specimen from Sidon. (This fact also seems to speak in favor of a Jaffa in Lebanon or Syria.) The fifty-one specimens from the Carmel area measure 12-16.6 mm., and twenty-four specimens from the northern frontier area measure 12.8-16.5 mm. They belong therefore to the same size group. Populations of small individuals were found at Wadi Tartar 486 FIELDIANA: ZOOLOGY, VOLUME 31 (12-14 mm.) ; Wadi Rushmieh (13.2-15.7 mm.) ; and eastern Carmel, Haifa (12.5-16.6 mm.). These smaller specimens are ribbed like the largest forms, thus differing distinctly from hierosolymitana, although they correspond in size. For this reason I am inclined to consider Laciniaria hiero- solymitana a separate species and not only a forma minor of moesta, as suggested by Pallary (1929, p. 32). The locality Jerusalem, how- ever, remains doubtful. According to 0. Boettger (1877), Clausilia saulcyi Bourguignat is a synonym of moesta. Westerlund (1884, p. 36; 1901, p. 201) and Germain (1921, p. 346) adhere to this viewpoint, whereas Tristram (1884, p. 201) and Pallary (1939, p. 65) consider the two as distinct. From my own experience, I share the opinion of Boettger and Westerlund and see in saulcyi only a slimmer race of moesta. Albinaria (Filumna) elonensis sp. nov. Figures 91 and 96, B. Type and paratypes in the collection of the Hebrew University; 20 paratypes in Chicago Natural History Museum No. 32110; type locality Elon, near the northern border of Palestine. Description. — Shell fusiform, with twelve whorls. The cylindrical narrowing of the last whorl as compared with the penultimate is striking, especially in small specimens. Viewed from above, the last whorl appears bent to the right. The last but two and the last but three whorls are as a rule the broadest. Beyond these the shell narrows, resulting in an elongated turnip shape. The first two or three whorls are smooth, but all the others bear delicate regular stripes that are most marked in the cervical region, but never amount to wrinkles. There are two not very prominent cervical folds separated by a shallow groove. The right is more prominent, more rounded and narrower, and particularly protruding and sharp- edged in the umbilical region; the left one passes toward the lateral peristomial rim and gradually vanishes. There is no trace of a peristomial collar ("collerette suprapenstomiale"). The shell is calcareous, somewhat bluish white, and the five upper whorls are light brown, passing into chestnut near the apex. The free peristomial margin is strongly thickened. The upper portion of the border of the aperture is completely fused with the penultimate whorl dorsally; the unattached portion of the aperture forms a spoon-shaped expansion bent on both sides toward the fused dorsal rim, thus forming a very characteristic curvature. Therefore the A Fig. 91. Albinaria (Filumna) elonensis sp. nov. A, Front view of shell; X 5 B, C, D, Lower part of shell in three different views. E, Clausilium; about X 20 F, Aperture; about X 20. 487 488 FIELDIANA: ZOOLOGY, VOLUME 31 profile figure shows that each peristomial side forms an angle with the dorsal, adnate portion, only the anterior cervical edges lying in one plane. The border turns markedly outward, but hardly upward. The adnate upper portion of the peristome is marked off against the penultimate whorl as a smooth reflecting surface, into which some of the fine striae of the former penetrate (or through which they may be discerned). In a few specimens the lateral peristomial edge passes into the deeply sunken lower lamella, which is joined from above by a shallow, accessory fold, constituting the right-hand border of the aperture. The deeper main branch of the lower lamella projects in surface view as a small deflection above this border. The peristomial rim of the columellar side can hardly be traced beyond the umbilicus, since above the recess it is already completely adnate. The upper lamella is a short crest and remains distinct from the spiral lamella. The latter begins internally and left of the upper lamella, as is easily seen in oblique view of the aperture from below. The principal fold is sharper and thinner than the latter and its beginning in the depth of the left upper angle is just visible; this fold is rather long and ends internally in a blunt thickening; below this runs a short parallel ledge ending at the same level — the rudiment of the second palatal fold. The third palatal fold is very short; beginning far inside, it rises in an arc toward the tiny second palatal fold and in its lower portion is almost invariably fused with the broad calliform lunar fold, which thus appears as the thickened end of the third fold. In a minority of cases there is a small furrow separating the third fold from the calliform portion (representing no doubt the primitive condition) . The columellar fold begins deep inside. The clausilium is rounded in front and devoid of a recess. Its longitudinal edges are somewhat thickened and bent up, so that it forms a channel. With a higher magnification it reveals a finely granulated surface structure, in contrast to the smooth polished clausilia of the genus Cristataria. Size. — Height rather variable, 10.5-15 mm., mostly 13.5-14.5 mm.; breadth 2.8-3.2 mm.; height of aperture 3 mm.; breadth of aperture 2.5 mm. Distribution. — Elon, Wadi Kerkara near Elon, a cave between Elon and Hanita, Wadi Qurn. All these localities are near the northern frontier of Palestine, from 200 to 460 meters above sea level, in Mediterranean scrub- wood, on Cenomanian limestone; yearly rainfall more than 800 mm. HAAS: CLAUSILIIDAE OF PALESTINE 489 Remarks. — Albinaria elonensis is markedly smaller than the nearest relative, namely A. hedenborgi Pfeiffer (according to Pallary, 1929, p. 28, this species includes ehrenbergi Roth, filumna Pfeiffer and maronitica Westerlund). It differs from this species in that the peristomial rim is sharply set off and fused on and by the spoon- shaped expansion of the lower peristomial portion. Moreover there are no wrinkled cervical folds. In the localities mentioned, the species is rather common on rocky slopes and under smaller stones. This is the first instance of the occurrence of an Albinaria in Palestine, A. hedenborgi being recorded from Tannourine and from the Nahr el Kelb Valley, A. Veprevieri Pallary from the Ansarieh Mountains. Cristataria (Strangulata) petrboki Pallary. Figures 92 and 96, C. Clausilia (Strangulata) petrboki Pallary, 1939, Mem. Inst. Egypt., 39: 63, text fig. 4, pi. 3, figs. 10-12. Description. — Cristataria petrboki bears strong and broad ribs, and its whorls are distinctly set off against one another by deep furrows. The last half-whorl is detached from the remainder of the spire and by a deep contraction, which is not ribbed, becomes thinned out to form an upper sharp rim, running to the left upper sinus of the aperture and continuing as a deep fold toward the columella. Two cervical folds are almost completely fused to form a prominence, which bears (as seen from above) several low sharp folds on the right side, spaced like the ribs; on the left side a number of fine wrinkles are spaced twice as densely. The principal and the lunar folds are distinctly seen through the shell. Along the border line between the dense wrinkles and the double-spaced sharp ribs runs an indistinct depression situated between the two cervical folds. The border of the aperture is widened and lies exactly in one plane running obliquely to the axis of the spire. There is no trace of a supraperistomial collar ("collerette"). The color is dirty yellowish-white, grayish-brown toward the apex. The aperture is white. In many living specimens the last whorl appears much lighter than the others, probably because the blackish body is seen through the upper whorls of the shell. In a front view, only the shallow upper lamella is visible, the others being more deeply situated. Externally (i.e., to the left) at a short distance from the inner end of the upper lamella the relatively shallow spiral lamella begins. The interlamellar space is smooth. The lower lamella begins at a very deep level, as a simple broad fold that sharpens upward and inward toward the free margin. The 490 FIELDIANA: ZOOLOGY, VOLUME 31 lower end of the columellar fold arches toward the lower lamella; upward, it rises approximately parallel to the lower lamella, being half as broad as the latter. A deep broad furrow runs between the two lamellae below. The principal fold has a sharp margin, exactly Fig. 92. Cristataria (Strangulata) petrboki Pallary. A, Front view of shell; X 5. B, Back view of lower part of shell; X 5. C, Clausilium; about X 20. D, Aperture; about X 20. opposite the spiral lamella, describing somewhat less than half a whorl upward. A slit runs between the principal fold and the spiral lamella, gradually tapering upward. The second palatal fold is very short and thread-like, running at an angle of 30 degrees toward the inner end of the principal fold but ending at a distance below the latter, though at the same vertical level. The lunar fold is a bean-shaped callus running at an angle of about 70 degrees toward the external (anterior) end of the second palatal fold, but stopping HAAS: CLAUSILIIDAE OF PALESTINE 491 just before it. The concavity of the lunar fold, which is hardly longer than the second palatal fold, is directed upward toward the latter. The distal end of the clausilium is broad and shovel-shaped, forming a groove (in cross section it describes an arc of about 120 degrees). The lower margin is somewhat notched and forms angles of about 80 degrees with both longitudinal borders. Thus the end appears as though clipped off, as a marked difference from the elongated, rather pointed ends of the other clausilia described in this paper. Size. — According to Pallary (1939) the maximum height reached by this snail is 15 mm., but some of our specimens reach a height of 16.2 mm. As in the other Palestinian Clausiliidae, I could not find any relation between the size of the shell and the altitude of the habitat of the specimens, as stated by Pallary (1939, p. 55). According to this author, the highest places in the area of occurrence are inhabited by forma major, the lowest by forma minor. Distribution. — The type locality is Gilboa Mountain, near Beth Alpha, in the Valley of Esdraelon (Emek Jesreel); Pallary (1939) writes erroneously "Gibboa Beith Alfa." Walking southward uphill from the settlement Beth Alpha, I found the snail at 50-60 meters above the bottom of the valley; it was equally numerous toward the east on limestone rocks exposed to the north, and also over a distance of 3-4 km. before the descent into the same valley. At lower levels, on rocks exposed to the northwest, the animals are found within a moist lime crust (Nari) covering the hard rock, while higher up they live in deeply carved crevices and hollows on the very surface of the rock, which is covered by lithophagous lichens. The yearly rainfall in this area is approximately 500 mm. Cristataria (Strangulata) genezarethana Tristram. Figures 93 and 96, D. Clausilia genezarethana Tristram, 1869, Proc. Zool. Soc. London, 1869: 539. Clausilia (Cristataria) genezarethana Westerlund, 1901, Mem. Acad. Sci. St. Petersb., (8), 11: 47. Description. — Pallary (1929) figures genezarethana, emphasizing differences from the Syrian C. praegracilis. C. genezarethana is the largest species found in Palestine. The color is a light brownish-gray, the last whorl light gray, the aperture inside pure white; the upper whorls are often light brown. There are 123^ to 133^ whorls. The border of the aperture is complete and not fused with the preceding 492 FIELDIANA: ZOOLOGY, VOLUME 31 whorl ; there are two very unequal cervical ridges, the left one only a third as broad as the other, and set off distinctly from the latter and from the umbilicus. Toward the other side the right ridge is set off by a shallow notch in the last whorl. There is no peristomial collar. The apertural margin is moderately expanded, but not reflexed upward. It lies exactly in one plane with the exception of the sinus, which is somewhat bent upward, left of the upper lamella. In the aperture the upper lamella is seen, starting directly from the margin; the simple, unforked lower lamella at a relatively high level passes obliquely into the columellar margin. The principal fold appears drawn far forward; the external depression between the two cervical folds protrudes distinctly inside as a low, ascending ridge (the lowest palatal fold) . The columellar fold runs, in its lowest portion, nearly parallel to the lower lamella and does not approach it. The upper lamella has a broad, blunt crest and is strikingly low in its inner portion. Directly left and external of its inner end the spiral lamella begins, as if it were a faulty continuation of the former. A crescent-shaped fold opening apically may well represent the second palatal fold in its sharp-edged upper portion rising toward the end of the principal fold, while the central calliform portion may be the homologue of the lunar fold. The clausilium is elongated, forming a groove; its lower margin is entire, with an obtusely angled bend at the lower longitudinal border, its ultimate tip rounded. Size. — The height of the shell is rather variable, whereas the breadth is comparatively constant. There appears to exist a rather fixed relation between the height on one hand and height and breadth of aperture on the other. According to Tristram the measurements are 20.5 mm. and 3 mm., respectively. Our specimens are smaller. The following are the measurements of ten specimens from Tiberias, in the collection of Chicago Natural History Museum. Height Breadth Height Breadth No. of shell of shell of aperture of aperture 1 15.8 2.9 3.2 2.1 2 17.0 3.0 3.4 2.4 3 17.4 3.0 3.4 2.5 4 16.3 2.9 3.0 2.3 5 17.6 3.0 3.3 2.5 6 17.0 3.1 3.5 2.5 7 17.4 3.0 3.2 2.4 8 15.9 2.9 3.1 2.5 9 16.3 3.2 3.2 2.4 10 17.7 3.1 3.2 2.4 Mean 16.8 3.0 3.3 2.4 HAAS: CLAUSILIIDAE OF PALESTINE 493 Distribution. — Tristram (1884) found C. genezarethana "only on rocks near the plain of Gennesaret." The specimens in the Uni- versity collection were found in two very limited areas on the A m Fig. 93. Cristataria (Strangulata) genezarethana Tristram. A, Front view of shell; X 5. B, Clausilium; about X 20. C, Aperture; about X 20. western shore of Lake Galilee, in the scattered limestone enclaves of a territory characterized by basalt. The first locality is a steep precipice facing north and northeast, situated south of Tiberias, opposite the hot springs; the shells are found in straight deep rock crevices. The second locality (probably identical with Tristram's) is the Wadi Chamam, opening toward the lake near Migdal (plain 494 FIELDIANA: ZOOLOGY, VOLUME 31 of Gennesareth), north of the other locality; here also the snails were found on steep rocks facing north. The annual rainfall is approximately 500 mm. The character of the flora is Mediterranean, but devoid of the original forest (at present ruderal flora). Cristataria (Strangulata) davidiana Bourguignat. Figures 94 and 96, E. Clausilia davidiana Bourguignat, 1868, Rev. Mag. Zool., (2), 20: 376, pi. 16, figs. 12-15. Clausilia (Cristataria) davidiana Westerlund, 1901, Mem. Acad. Sci. St. Petersb., 11, no. 11: 46; Germain, 1921, Moll. terr. fluv. Syrie, p. 338, text figs. 33-35. Cristataria (Strangulata) davidiana Pallary, 1939, Mem. Inst. Egypt., 39: 56. Description. — The coloration of C. davidiana is the same as that of C. genezarethana, which appears much more elongated and slender, partly on account of the relatively narrower aperture and the smaller left dorsal sinus. In davidiana, the aperture is more rounded and passes gradually into the dorsal sinus, which is wide open ventrally; the border of the aperture is more broadly expanded and reflected. Moreover, davidiana is more densely ribbed. The lower whorls in genezarethana are rather cylindrical and are separated by a narrow, impressed suture; in davidiana the whorls are more convex and the suture is more rounded and less accentuated. In a dorsal view, the last whorl of genezarethana appears more compressed than that of the other species; viewed from the right side, davidiana exhibits the suggestion of cervical protrusion that does not merit the name of a "collerette supraperistomiale" ; at any rate, in this view the right outline falls off toward the straight aperture profile more steeply and is somewhat swollen. The aperture in front view is similar in both species, but the upper lamella in davidiana is broader and shorter and the lower lamella not so much protracted toward the left, thus exhibiting a smaller anterior face. The columellar fold appears below only after turning the aperture through 90 degrees to the right; it converges in an arc toward the lower lamella, the two together describing the outline of an upright biconvex lens. The principal fold begins rather low, opposite the spiral fold. The latter is only half as high as the upper lamella (which reaches its greatest height in its middle portion) and starts behind and left of the inner end of the upper lamella. The lunar fold (plus the second palatal fold) arches up toward the inner end of the principal fold, pad-shaped and flat below (but not in the form of a callosity), sharpening above. It ends before the principal fold, running parallel HAAS: CLAUSILIIDAE OF PALESTINE 495 to the latter in its terminal portion. There is no trace of the lower palatal fold. The lower portion of the clausilium has parallel long sides and between these a groove-shaped depression. The tapering distal Fig. 94. Cristataria (Strangulata) davidiana Bourguignat. A, Front view of shell; X 5. B, Clausilium; about X 20. C, Aperture; about X 20. part bears an inconspicuously pointed corner below. It differs markedly from the distally more twisted and differently tapering end of the clausilium of genezarethana and from the truncated one of petrboki. Size. — The measurements (in millimeters) of twenty specimens from Wadi Ghurab near Artuf are as follows: 496 FIELDIANA: ZOOLOGY, VOLUME 31 Length Height Height Breadth No. of shell of shell of aperture of aperture 1 15.1 3.0 3.0 2.3 2 18.2 3.1 3.4 2.9 3 17.3 3.2 3.3 2.5 4 17.8 3.0 3.3 2.6 5 17.9 2.8 2.9 2.4 6 16.0 3.0 3.0 2.4 7 17.4 3.1 3.1 2.4 8 15.3 2.9 3.0 2.5 9 14.0 2.9 3.0 2.5 10 14.6 2.8 2.8 2.3 11 17.2 3.0 3.1 2.5 12 14.6 2.9 2.9 ca. 2.4 13 14.9 ' 2.8 3.0 2.4 14 17.5 3.1 3.2 2.8 15 16.3 2.9 3.0 2.5 16 17.0 3.0 3.2 2.6 17 17.9 3.0 3.1 2.6 18 16.2 2.8 2.8 2.4 19 16.4 2.9 2.9 2.4 20 17.0 3.1 3.0 2.6 Mean 16.4 3.0 3.0 2.5 This species is on the average 4 mm. shorter than genezarethana, its width being the same. The aperture is lower (3 against 3.3 mm.), and the breadth is nearly identical. The ratio of height to breadth is 5.6 as against 5.5 in davidiana. Distribution. — Germain reports "Beyrouth, rochers sur les bords de la riviere du Chien; environs de Jaffa; Amchit, dans le Liban." These localities, however, are given for both this and the following species (see p. 501). I mention again my doubts concerning the possibility of an occurrence of this shell in a Palestinian Jaffa. The specimens in Chicago Natural History Museum and in the Hebrew University collection were found in the Wadi Ghurab, between Artuf (Hartow) and Beth Mahzin, at the western margin of the Central Judean Hills; some of the specimens were collected from holes of a soft limestone (nearer to Artuf), others from the gorge- like wadi on hard limestone. The annual rainfall averages 600-700 mm. Remains of the Mediterranean evergreen forest may be found in this area, which is 45 km. from Jaffa as the crow flies, at the Mediterranean shore. Cristataria (Strangulata) prophetarum Bourguignat. Figures 95 and 96, F. Clausilia prophetarum Bourguignat, 1868, Rev. Mag. Zool., (2), 20: 378, pi. 16, figs. 8-11. HAAS: CLAUSILIIDAE OF PALESTINE 497 Clausilia (Cristataria) prophetarum Westerlund, 1901, Mem. Acad. Sci. St. Petersb., (8), 11, no. 11: 47; Germain, 1921, Moll. terr. fluv. Syrie, p. 338, text figs. 36-38 (as synonym of davidiana). Cristataria (Strangulata) prophetarum Pallary, 1939, Mem. Inst. Egypt., 39: 56. A Fig. 95. Cristataria (Strangulata) prophetarum Bourguignat. A, Front view of shell; X 5. B, C, Lowest whorl in side and back views; X 5. D, Clausilium; about X 20. E, F, Aperture; X 20. Description. — The shell of C. prophetarum is elongate-conical rather than fusiform. The ribs are not as dense as those of davidiana and resemble those of genezarethana, in which the intercostal spaces are at least twice as broad as the costae, the latter being more massive 498 FIELDIANA: ZOOLOGY, VOLUME 31 and more bluntly edged than in davidiana. The shallow intercostal field of prophetarum is about one and one-half times as broad as the rib, and the ribs are also appreciably stouter than in genezarethana; these facts apply, however, only to the lowermost whorls. The last whorl is elongated and is subcylindrical as in genezarethana. The suture between the whorls is deeply impressed. The aperture is notched dorso-ventrally in the cervical region, but less so than in davidiana, whereas there is no notch at all in genezarethana. The peristomial border is most strongly expanded and reflected upward in this species. The boundary between the two cervical folds is only just indicated by a flexure of the ribs that cross this region. A supraperistomial fold is more distinct than in davidiana (especially on the left as viewed from the dorsal or, better, the cervical side of the shell), but this fold is incomplete and cannot be compared with the sharp and prominent ridge of medlycotti. The relatively narrow aperture contrasts with the broad peristomial border. The upper and lower lamellae are closely approximated, more than in the other two species. The lower lamella descends but little along the columellar margin. The left upper marginal sinus is more distinctly marked than in genezarethana by the thickening and contraction of the left margin, which runs down to the lower end of the lower lamella, narrowing the aperture, and which is most thickened opposite the upper lamella. The lower end of the columellar fold diverges by an angle of 45 degrees from the lower margin of the lower lamella, which passes into the marginal protrusion below, while above it thins out to form a sharp crest. The upper lamella arches outward in a semicircle to its maximal height and falls off inward into a shallow, long, drawn-out fold. A very long longitudinal fold runs in the same direction toward the inner end of the upper lamella; it may join the latter, thus constituting its prolongation, in which case the border between the two folds is marked by a slight depression; or it may bend left of the inner end of the upper lamella and ascend as a low ridge toward the columellar margin of the lower lamella, stopping short before it. This accessory fold may be absent. The spiral lamella begins near the inner end of the upper lamella. Toward the inner end of the normal principal fold, an arched lunar fold (plus the second palatal fold?) is developed. The ascending branch is hardly indicated as a low crest; the part that may represent the palatal fold and that runs opposite the principal lamella is a short, blunt ridge as high as the latter, which broadens perceptibly in this region. The clausilium is similar to that of davidiana, but its lower part is rounded, not drawn out into an angular shape. B D E Fig. 96. A, Laciniaria (Elia) moesta F6russac. B, Albinaria (Filumna) elonensis sp. nov. C, Cristataria (Strangulata) petrboki Pallary. D, Crislataria (Strangulata) genezarethana Tristram. E, Cristataria (Strangulata) davidiana Bourguignat. F, Cristataria (Strangulata) prophetarum Bourguignat. All X 2. 499 500 FIELDIANA: ZOOLOGY, VOLUME 31 The numerous differences between C. davidiana and C. prophet- arum are in my opinion fully sufficient to warrant the adherence to Bourguignat's conception and to disprove the validity of Ger- main's arrangement of this species. Size. — Measurements of ten specimens (nos. 3 and 5 in Chicago Natural History Museum, the others in the collection of the Hebrew University); specimen 6 is broken. Height Breadth Height Breadth No. of shell of shell of aperture of aperture 1 17.0 2.8 2.8 2.4 2 16.8 2.8 3.0 2.4 3 16.9 2.9 3.0 2.4 4 16.5 2.6 2.8 2.3 5 17.4 2.6 3.0 2.4 6 17.4 2.7 7 16.3 2.7 2.5 2.2 8 15.1 2.5 2.8 2.2 9 15.8 ca. 2.9 2.9 2.4 10 16.2 2.9 2.8 2.3 Mean: prophetarum 16.5 2.7 2.8 2.3 davidiana 16.4 3.0 3.0 2.5 genezarethana 16.8 3.0 3.3 2.4 The ratio of height to breadth is 6.1 as compared with 5.6 in genezarethana and 5.5 in davidiana. This should be compared with Bourguignat's figures (quoted from Germain, 1921, p. 339), giving for davidiana 57/12 (=4.75), and for prophetarum 72/12 (=6.0). There is good agreement in the ratios of prophetarum, while Bourgui- gnat seems to have dealt with a different, stouter population of davidiana (perhaps from the Lebanon?). Comparing the mean measurements of these three species, which are, no doubt, closely related, it is seen that genezarethana is the longest form with the highest aperture, and that in the ratio of height to height of aperture, prophetarum is first (5.9), davidiana second (5.5) and genezarethana last (5.0). A superficial comparison gives the impression that davidiana is the stoutest of the three species, whereas the two others may easily be distinguished by the gaping mouth of genezare- thana and the thick-bordered contracted one of prophetarum. Distribution. — Syria (Westerlund) ; the only known locality from Palestine (collected by Kushnir) lies between Khurbeta ibn Harith and Midie, 10 km. east of Lydda, on the western border of the Judaean hills, about 20 km. northwest of the locality of davidiana. Only empty shells were found on rocks in relatively open land. The HAAS: CLAUSILIIDAE OF PALESTINE 501 author has no personal knowledge of this place. Annual rainfall almost 700 mm. Germain's localities are confused with those for davidiana. GENERAL REMARKS The distribution of the Clausiliidae in Palestine, as in many other regions, is discontinuous and the range of certain species may be very limited. Laciniaria moesta seems to constitute the only exception. A number of small areas of occurrence may still have escaped the notice of the numerous collectors. All Cenomanian limestone regions, namely, the northern mountain area and the range of the central hills, wherever the annual rainfall is not below 500 mm. average, still should be searched for these snails. Moun- tain ranges in the west or northwest, situated relatively near the sea, appear to be most favorable. The localities in the Jordan Valley (genezarethana) and at the transition from the plain of Esdraelon to the extremely dry and hot Jordan Valley (petrboki), both with relatively low rainfall, face partly to the northeast and are exposed to the scorching desert storms; thus they do not seem to offer ideal biotopes for a clausiliid. The mean temperature in the coldest month (January) does not sink below 10° C. in all these areas. At all events, the sparsely distributed Clausiliidae of Palestine must be considered as relicts of a moister and possibly cooler era. The occurrence in the Jordan Valley may well date from the post- glacial period, when a fresh- water lake of considerable dimensions covered the valley as well as the eastern part of the plain of Esdraelon, producing a rather high atmospheric humidity in the neighborhood. A more comprehensive picture of this highly interesting distribu- tion area (which ends in Palestine) might be gained by investigating the occurrence of Clausiliidae in the more humid region of northern Transjordania, where no representative of this group has ever been found. This area is partly covered by woods to this day. Further searching of the whole of northern Palestine, and the coastal slopes from Beyrouth southward to about halfway between Gaza and Jaffa is needed. I am very much indebted to Dr. Fritz Haas, Curator of Lower Invertebrates, Chicago Natural History Museum, for useful and indispensable advice and for his kind help in the preparation of this paper. 502 FIELDIANA: ZOOLOGY, VOLUME 31 REFERENCES BOETTGER, O. 1877. Clausilienstudien. Palaeontographica, Suppl. 3, pts. 6 and 7, pp. 1-122, pis. 1-4. Germain, Louis 1921. Mollusques terrestres et fluviatiles de Syrie. Voy. Zool. Henri Gadeau de Kherville en Syrie, 2: 1-523, 56 figs., 23 pis. Pallary, Paul 1929. Premiere addition a la faune malacologicfue de la Syrie. Mem. Inst. Egypt., 12: 1-43, pis. 1-3. 1939. Deuxieme addition a la faune malacologique de la Syrie. Mem. Inst. Egypt., 39: 1-141, pis. 1-7. Tristram, Henry Baker 1884. The survey of western Palestine. The fauna and flora of Palestine. London, 455 pp., 20 pis. Westerlund, Carl Agardh 1884. Fauna der in der palaarktischen Region (Europa, Kaukasien, etc.) lebenden Binnenconchylien. IV. Gen. Balea Prid. u. Clausilia Dr. Karskro- na, Lansboktrykkeri, 212 pp. 1901. Synopsis molluscorum in regione palaearctica viventium ex typo Clausilia Drap. Mem. Acad. Sci. St. Petersb., (8), 11, no. 11: xxxvii+203.