Melanopsis, FERUSSAC, 1824

Heller, Joseph, Mordan, Peter, Ben-Ami, Frida & Sivan, Naomi, 2005, Conchometrics, systematics and distribution of Melanopsis (Mollusca: Gastropoda) in the Levant, Zoological Journal of the Linnean Society 144 (2), pp. 229-260 : 231-256

GENUS MELANOPSIS FÉRUSSAC, 1824

Description

The shell is thick and imperforate, and may be smooth or heavily ribbed. A notch is present at the base of the mouth, where the outer and inner lips meet. The upper part of the mouth is narrowly constricted. On the inner lip, the upper part usually contains a callus; in the lower half, the columella is truncate ( Heller et al., 1999).

The present study reveals ten Melanopsis species in the Levant: three are widespread ( M. buccinoidea , M. costata , M. saulcyi ), two have narrow distributions ( M. dircaena , M. ammonis ), and five are known only from their type localities ( M. khabourensis , M. meiostoma , M. pachya , M. germaini and M. infracincta ).

The ten Melanopsis species of the Levant are described below. For convenience of description, all the smooth-shelled species are presented first, then all the ribbed ones. Within each of these groups, the order of the species follows their year of original description.

MELANOPSIS BUCCINOIDEA ( OLIVIER, 1801)

( FIG. 2A, B; TABLE 1)

Melanie buccinoide Olivier, 1801: pl. 17, fig. 8. Melanopsis ferussaci Roth, 1839: 24 , pl. 2, fig. 10. Melanopsis sesteri Bourguignat, 1884: 119 , from Hula Valley (‘Ain al Bass’) and environs of Aleppo (Sadjour- Sou 70 km NE Aleppo, near ‘Ain Taib’); not sesteri var. diadema Bourguignat from Sadjour-Sou.

Melanopsis praemorsa, Germain, 1921 : pl. 19; Bandel & Salameh, 1981; fig. 23; Heller & Abotbol, 1997.

Melanopsis buccinoidea, Pallary, 1939: 84–85 , pl. 6, figs 1-4, 64, 65; Heller et al., 1999: 56–59, fig. 4A; Bandel, 2000, figs 1, 20-24, 71.

Melanopsis denegabilis, Pallary, 1939: 85–86 , pls. 6, figs 9–12.

Melanopsis prophetarum, Pallary, 1939: 83–84 , pl. 6, figs 16, 17, 23–26, 36, 57.

Melanopsis olivieri, Pallary, 1939 : pl. 4, fig. 11.

Melanopsis praemorsa ferussaci, Schütt, 1983: 40–41 , pl. 2, figs 42, 44, not fig. 43.

Melanopsis praemorsa ferussaci, Ghab form, Schütt, 1983: 41–42, pl. 2, fig. 47.

Melanopsis praemorsa olivieri, Schütt, 1983: 44 .

Types and type locality

‘Melanie buccinoide, M. buccinoidea de Scio ’ ( Olivier, 1801; pl. 17, fig. 8). In the Muséum National d’Histoire Naturelle (Paris) Tillier & Mordan (1983: pl. 5, fig. 11) found ten possible syntypes of M. buccinoidea , labelled ‘ M. praemorsa L., Tripoli, M. Olivier, 1803’. M. buccionoidea was synonymized with M. praemorsa by Germain (1921: 477).

Material examined

SYRIA: Kara Sou ( BMNH 20030348), Nahr Afrin ( SMF 282748), Sadjour-Sou ( BGT 11341), Antiochia ( SMF 290751), Haleb ( NHMW 84539), Orontes ( SMF 282770, 282771, 282772), Akbes ( BMNH 1901.11.26.88–92), Barda River ( BMNH 20030350).

LEBANON: Jeble ( SMF 282739, 282740), Beirut ( SMF 115083, 115084, 290783, 290741, NHMW, 1894, 57575), Nahr el Kabir ( SMF 282729, 282745), Nahr el Kelb ( ZMH), Hermel ( BMNH 20030351), Baalbek ( BMNH 1931.12.30.10345–7, 1933.8.11.72–82, 20030353). JORDAN: Near Irbid, Azraq North, Azraq South, Al Shuna, North Shuna, Wadi Karkara, Hisbon Quelle, Hamam Salomon (all in GPIuMH). ISRAEL: JORDAN VALLEY: Nahal Dan, Nahal Snir, Nahal Gilbon, Nahal Meshoshim, Yehudiya, Enot Huga, En Hanaziv, En Parta, En Sultan (all in HUJ); Ain el Bas ( BGT 11340). COSTAL PLAIN AND ELSEWHERE: Nahal Bezet ( HUJ), En Tamid ( HUJ), Nahal Kishon ( BMNH 20030349), Nahal Taninim ( HUJ), Nahal Yarqon ( HUJ, ZMH), Yafo ( ZMH), Nazareth ( ZMH),

Diagnosis

Melanopsis buccinoidea differs from M. praemorsa L., the type species of Melanopsis , in that it is larger and more cylindrical.

Description

The shell has up to seven whorls. It has a pointed spire, flattened whorls separated by very shallow sutures, and is smooth. In the sperm, the midpiece is exceptionally long ( Hodgson & Heller, 2000); the radula is highly variable ( Mazan-Mamczarz et al., 2002). The variability range of allozymes is exceptionally wide, with populations scattered among clusters of other species, suggesting a broad extent of hybridization with other species ( Falniowski et al., 2002a, b) that has been going on in the Levant since the Lower Pleistocene and is still widespread today ( Heller et al., 1999; Heller & Sivan, 2002a: table 1).

Synonyms

Bourguignat (1884) described M. sesteri from the Hula Valley (‘Ain al Bass’) and Sadjour-Sou. As the type specimens of M. sesteri ( MHNG 11340, 11341) fall within the conchometric (as opposed to geographical) range of M. buccinoidea from the coastal plain ( Table 1) we consider M. sesteri a synonym of M. buccinoidea .

Pallary (1939: 82–83, fig. 9) described M. laevigata Lamarck from the Levant. His assigning of Levantine Melanopsis to laevigata is difficult to investigate, because he illustrated the type shell of laevigata , which is not from the Levant, but his laevigata from the Levant are not illustrated. We assume that laevigata from the Levant are synonyms of buccinoidea , whereas the North African M. episema Bourguignat and M. callichroa Bourguignat , which Pallary considered varieties of laevigata , are not. M. prophetarum described by Pallary (1939) ( BMNH 1901.11.26.88–92) does not differ from typical M. buccinoidea from the Jordan Valley ( Table 1) and thus we consider M. prophetarum a synonym of M. buccinoidea . Pallary (1939) described M. denegabilis de Férussac as a smooth Melanopsis of intermediate dimensions, and illustrated specimens from Lakbe and from Tripoli. His figures of M. denegabilis are all within the range of M. buccinoidea , hence we consider denegabilis a synonym of buccinoidea .

Schütt (1983) considered all Melanopsis of the Levant as subspecies of M. praemorsa (the type species of Melanopsis , from Spain). He assigned the smooth-shelled Melanopsis of the coast and southeastern parts of the Levant to M. p. ferussaci Roth , which differs from the large, narrow, red-brown M. p. buccinoidea Olivier of Greece and the Aegean in being small and dark-coloured. Whether these smooth Melanopsis of the Levant are separate from M. buccinoidea of Greece (as suggested by Schütt, 1983), synonymous with M. buccinoidea (as suggested by Glaubrecht, 1993) or synonymous with M. praemorsa (as suggested by Tchernov, 1975), is beyond the scope of this present study.

Within his M. p. ferussaci, Schütt (1983) placed some Orontes populations as a separate ‘ praemorsa ferussaci Ghab form’, which differs from the coastal M. p. ferussaci in its large size, and in always being completely smooth. The measurements and ratios of his M. p. ferussaci Ghab form ( Table 1) do not differ from M. buccinoidea from the Syrian coast. We therefore consider the M. p. ferussaci Ghab form a synonym of M. buccinoidea .

M. olivieri was described by Bourguignat (1884) from several sites throughout the Levant (the environs of Aleppo, Sajour-Sou, Beirut, Damascus and Jericho); Schütt (1983) placed it as a separate subspecies of north-eastern parts of the Levant, characterized by a shell even larger and wider than the M. p. ferussaci Ghab form. Measurements of two specimens from northern Syria defined by Schütt as M. praemorsa olivieri (NHMW 84539 from Aleppo and SMF 282828 from the Quaik River, near Tell Arsof) are presented in Table 1. The shells fall within the range of M. buccinoidea in all characters. We therefore consider M. praemorsa olivieri a synonym of M. buccinoidea .

Distribution and habitat

M. buccinoidea is widely distributed in the Levant ( Fig. 3 ). In the southern Levant it occurs in a wide variety of aquatic habitats that range from small trickles to springs and streams, where it lives on stones, sometimes also in silty mud; it is absent from the Jordan River and from Lake Kinneret ( Heller et al., 1999, 2002). A generalist feeder, it frequently feeds on dry willow leaves and is an important litter shredder in certain desert oasis ecosystems ( Heller & Abotbol, 1997). Parasites include the eye fluke Philopthalamus lucipetus (Rudolphi, 1819) ( Radev et al., 1999). In the Levant, fossils of M. buccinoidea are known from the mid-Pleistocene, Lower Pleistocene and Pliocene of the Jordan Valley ( Heller & Sivan, 2001, 2002a, b). Both Glaubrecht (1993) and Falniowski et al. (2002b) considered M. buccinoidea a stem species, from which ribbed species of the Levant eventually evolved.

Intraspecific variation

There is considerable variation among M. buccinoidea populations of the Levant. Compared to shells from elsewhere in the Levant, shells from the Jordan catchment area tend to be stouter (shell-diameter/shellheight is 0.50 ± 0.02 vs. 0.46 ± 0.03 elsewhere; the Nahal Taninim population on the coast of Israel is an exception in that it too has stout shells), they have significantly larger mouth-height/shell-height ratio (0.62 ± 0.04, vs. 0.55 ± 0.04 elsewhere) and a lower f/mouth-height ratio (0.16 ± 0.02, vs. 0.21 ± 0.03 elsewhere) ( Table 1, Fig. 4 ).

The Jordan catchment area M. buccinoidea may perhaps be a separate subspecies. We have previously referred to this ( Heller, Sivan & Ben-Ami, 2002), noting that on the coast of the Levant, M. buccinoidea tends to be more elongate than in the Jordan Valley. We return to this point below, when discussing M. ammonis . It is noteworthy in the dendrogram of Figure 7 that, within the Jordan Valley, samples of M. buccinoidea from the northern region of the valley (e.g. Azraq, Al Shuna II) cluster separately from those of the southern region (e.g. N. Gilbon, Yehudiya).

We are undecided as to the taxonomic status of three samples of short, eroded, smooth shells from Ba’alBek (BMNH 1931.12.30.10345–47, 1933.8.11.72– 82 and 20030353, labelled M. brevis ). With a shelldiameter/shell-height of 0.54 ± 0.03 these shells are significantly stouter, not only than those of Lebanon, but even than those from the Jordan Valley. Further, mouth-height relative to shell-diameter in these specimens is only 1.02 ± 0.04, smaller than that of all other populations of M. buccinoidea , and the mouth is more rounded (mouth-diameter/mouth-height 0.49 ± 0.04). The type locality of Melanopsis brevis Mousson 1854 , to which these shells are referred, is the Litani ( Germain, 1921: 463). Pallary (1939) discussed the taxonomy of brevis and concluded that it is a synonym of laevigata .

Shell-diameter (mm) Shell-height (mm) Mouth-height (mm)

Melanopsis sharhabili was recently described by Bandel (2000: 156, figs 119–123, type no. 4267 in GPIuMH) from a ‘spring and creek next to Wadi Raiyan Plantation in the Jordan Valley near the town of Wadi Raiyan and close to the mosque of Sharhabil Ibn Haddanna’. The typical M. sharhabili differs from M. buccinoidea of the Jordan Valley in that its four last whorls are more expanded, as compared to the previous whorls. The shell consists of nine whorls. The first five to six increase regularly in diameter; the next increases rapidly, both in diameter and in height, so that it is two-thirds of total shell height. Bandel (2000) noted that M. sharhabili grades in shape (not in size) into local M. buccinoidea , and suggested that it is probably a hybrid of M. buccinoidea and one of the ribbed species, M. saulcyi or M. bandeli , that live in the same creek and spring. As M. sharhabili grades into M. buccinoidea in the only locality from which it is known, we are undecided as to its subspecific position.

MELANOPSIS AMMONIS ( TRISTRAM, 1865)

( FIG. 2C, TABLE 2)

Melanopsis ammonis Tristram, 1865: 542 , no. 102.

Melanopsis eremita Tristram, 1865: 542–3 , no. 107.

Melanopsis ammanensis, Pallary, 1939: 88–89 , pl. 4, fig. 3.

Melanopsis praemorsa eremita, Mienis & Ortal, 1994 : II.

Melanopsis doriae, Bandel, 2000: 152 , figs 25–29. Not Melanopsis doriae Issel, 1886 .

Melanopsis ammonis, Bandel, 2000: 177 , figs 117, 118.

Material examined

Types: four syntypes ( BMNH 1968665 ) from ‘streams at Heshbon and Ammon, east of Jordan’ .

Additional material: JORDAN: Rabat Ammon ( BMNH 1936.3.10.7–11), W. Walla, Hamam Yarmouk, Rumeimin, opposite Enot Qane (all in GPIuMH). ISRAEL: En Boqeq ( BMNH 1968666, 1936.3.10.12– 18), En Amatziahu, En Tamar, En Namer (all in HUJ). Gaza ( BMNH 1937.12.30.10402–16).

Diagnosis

Melanopsis ammonis differs from M. buccinoidea of the Jordan Valley in its higher penultimate whorl, as expressed in its higher values of the ratio f/mouthheight.

Description

The shell is smooth, narrow, tall and elongate with a pointed spire. It consists of up to nine whorls, which are high, very flattened and separated by very shallow sutures. Shell colour is almost always uniform black, or reddish brown; sometimes it is greyish yellow. The callus is white. Conchometrics of three of the four syntypes, BMNH 1968665, together with BMNH 1936.3.0.7–11, are in Table 2.

Synonyms

Tristram (1865: 542–3) described M. eremita from ‘the little stream of the Wady Um Baghek, between Sebbeh and Jebel Usdum, at the south-west corner of the Dead Sea’ (today this stream is named En Boqeq). He described it as spindle-shaped, semitransparent, smooth, glossy, with very faint nonregular bands, and eight whorls that grow regularly. He commented that in this region ‘which supplies the smallest of its group, the common Melanopsis praeorsa [ buccinoidea of this study] attains its greatest magnitude’. Measurements and ratios of eight of the 12 types ( BMNH 1968666) and seven additional specimens ( BMNH 1936.3.10.12–18) of M. eremita ( Table 2) illustrate that M. eremita is broadly similar to M. ammonis , differing only in its slightly higher ratios of shelldiameter/shell-height and mouth-height/shell-height. These differences are not diagnostic, whereby we consider M. eremita a synonym of M. ammonis . In a recent visit to En Boqeq, the type locality of M. eremita , no M. eremita but only M. buccinoidea were found.

Bandel (2000: figs 25–29) described Melanopsis doriae Issel from northern regions in the Kingdom of Jordan (a thermal spring at Hamma, in the Yarmouk Valley). From his figures we could not determine any differences between his M. doriae on the one hand and our M. ammonis from nearby sites of the Yarmouk and Jordan valleys on the other. We therefore consider Bandel’s M. doriae a synonym of M. ammonis .

Distribution

All records are from the southern Levant ( Fig. 5 ), mostly from the Jordan Valley: (Heshbon, Wadi Mujib near its outlet to the Dead Sea, Wadi Walla and Yarmouk in the Kingdom of Jordan; En Boqeq and En Amatziahu in Israel). A sample from Gaza (coastal plain of the Palestinian Authority, conchometrics in Table 1; originally labelled Melanopsis ferussaci Roth ) is the single record beyond the Jordan Valley.

Comparisons

M. ammonis differs almost diagnostically from M. buccinoidea of the Jordan Valley in its higher ratio of f/mouth-height (0.25 ± 0.03 vs. 0.16 ± 0.02) ( Table 2, Fig. 6A ). In addition it differs significantly in its ratios shell-diameter/shell-height (0.44 ± 0.02 vs. 0.50 ± 0.02), mouth-height/shell-height (0.50 ± 0.03 vs. 0.62 ± 0.04), mouth-height/shell-diameter (1.14 ± 0.05 vs. 1.24 ± 0.06), e/mouth-height (0.38 ± 0.05 vs. 0.28 ± 0.04), mouth-diameter/mouth-height and shouldering. Figure 7 presents relationships among the samples of smooth shells from the Jordan Valley catchment area. In this dendrogram the samples from the Yarmuk, Wadi Walla, En Boqeq, En Amatziahu and Heshbon, fall into one group ( M. ammonis ), whereas those of Nahal Gilabon, Yehudiya, Nahal Snir, En Hanatziv, En Sultan and En Huga fall into another ( M. buccinoidea ).

M. ammonis of the Jordan Valley catchment area is clearly distinguished from buccinoidea of the same area. However, it is less well-separated from

Mouth-height (mm) Mouth-height (mm)

M. buccinoidea of elsewhere in the Levant ( Fig. 6B ). In separating M. ammonis as a distinct species we place more weight upon comparisons between close populations (in geographical terms) than between distant ones. To conclude, in the Jordan Valley region, M. ammonis can be clearly separated from M. buccinoidea ; elsewhere in the Levant this separation is more difficult.

MELANOPSIS DIRCAENA PALLARY, 1939

( FIG. 2D, TABLE 3)

Melanopsis dircaena Pallary, 1939: 87–88 , pl. 6, figs 31–35.

Not Melanopsis dircaena var. ex-colore luctuosa Pallary, 1939: 88, pl. 4, fig. 19.

Standardized Euclidean

Material examined

Types: six syntypes in BMNH (1937.12.30.10.383–88); all from ‘lac de Homs’ ( Pallary, 1939: 140).

Additional material: SYRIA: vicinity of Homs ( SMF 282785 , 283565 , 290938 , 290954 ) .

Homs N = 6 (pooled) M.dircaena Max. shell-height 16.8 mm

Max. shell-diameter 9.2 mm

Max. mouth-height 10.0 mm Shell-diameter/shell-height 0.59 ± 0.05/0.56–0.66 Mouth-height/shell-height 0.60 ± 0.04/0.54–0.66 Mouth-diameter/mouth-height 0.44 ± 0.02/0.42–0.46 Mouth-height/shell-diameter 1.04 ± 0.04/1.00–1.09 Notch-width/notch-depth 0.85 ± 0.08/0.75–0.91 Notch-width/mouth-diameter 0.51 ± 0.10/0.39– 0.63 e /mouth-height 0.36 ± 0.09/0.28–0.51 f/mouth-height 0.18 ± 0.03/0.14–0.21 Conicality (a/b) 0.81 ± 0.02/0.78–0.84 Figurativity (c/d) 0.90 ± 0.01/0.89–0.92 Shouldering (b/c) 0.73 ± 0.03/0.69–0.76

Shell–diameter / Shell–height Axis 1

Diagnosis

Melanopsis dircaena differs from M. buccinoidea in that it is almost always more stepped.

Description

The shell is stout, smooth and has up to seven whorls. It has a short spire, and heavily shouldered whorls separated by deep sutures. The mouth-height is large (relative to shell-height). Shell colour is either uniform black or pale brown with three dark bands. The ground colour of the columella and (well-developed) callus is whitish; the callus is tinted brown. Conchometrics (N = 6) are presented in Table 3.

Comments

The Senckenberg Museum has four samples from the vicinity of Homs (290938, labelled M. costata bullio ; 290954, labelled M. costata jordanica ; 282785 and 283565, both labelled M. praemorsa costata ). Samples 290938 and 282785 contain only smooth shells; 290954 and 283565 contain a mixture of smooth and ribbed shells, and perhaps also hybrids. We assign all of these samples to M. dircaena .

Distribution

Localities available to us include, in addition to that of the type samples, also samples from Lake Homs, from south-west of Rastane (above the Stan Lake ) and from Sheizar ( Fig. 5 ) .

Comparisons

M. dircaena differs from M. buccinoidea (to which it is closer than to other smooth species of the Levant) in that it is diagnostically more stepped (shouldering 0.73 ± 0.03 vs. 0.82 ± 0.03) and stout (shell-diameter/ shell-height 0.59 ± 0.05 vs. 0.46 ± 0.03) ( Tables 1, 2 and Fig. 8A ). It further differs in that the mouth is wider as compared to mouth-height, mouth-height is lower relative to shell-diameter, and the shell is less conic (as expressed by higher values of conicality and figurativity). These differences contribute to the separate clustering of the M. dircaena samples in Figure 8B .

Pallary (1939) suggested that M. dircaena was the stem species, from which all ribbed species of the Orontes eventually evolved. Pérès (1946) however, suggested that it is a degenerate species.

MELANOPSIS KHABOURENSIS PALLARY, 1939

( FIG. 2E, TABLE 4)

Melanopsis (Mesopotamia) khabourensis Pallary, 1939: 102–104 , pl. 5, figs 13-15, 21.

? Melanopsis (Mesopotamia) khabourensis var. elongata Pallary, 1939: 103 , pl. 5, fig. 16.

Not Melanopsis (Mesopotamia) khabourensis var. dolichosoma Pallary, 1939: 103 , pl. 5, fig. 18.

Not Melanopsis (Mesopotamia) khabourensis var. plicata Pallary, 1939: 103 , pl. 5, fig. 19.

Not Melanopsis (Mesopotamia) khabourensis var. gradata Pallary, 1939: 103 , pl. 5, fig. 120.

Ras al ‘Ayn’ N = 6 M. khabourensis Max. shell-height 29.1 mm

Max. shell-diameter 15.3

Max. mouth-height 16.0 Shell-diameter/shell-height 0.53 ± 0.02/0.51–0.56 Mouth-height/shell-height 0.54 ± 0.03/0.42–0.58 Mouth-diameter/mouth-height 0.43 ± 0.02/0.40–0.45 Mouth-height/shell-diameter 1.01 ± 0.04/0.97–1.07 Notch-width/notch-depth 0.70 ± 0.03/0.64–0.74 Notch-width/mouth-diameter 0.38 ± 0.04/0.32– 0.42 e /mouth-height 0.39 ± 0.05/0.32–0.42 f/mouth-height 0.21 ± 0.02/0.19–0.24 Conicality (a/b) 0.79 ± 0.04/0.75–0.83 Figurativity (c/d) 0.91 ± 0.01/0.88–0.93 Shouldering (b/c) 0.79 ± 0.02/0.76–0.81 Material examined

Types: six syntypes ( BMNH 1937.12.30.293–302); all from ‘ Sources du Khabour, dites Ras el “Ain”, northern Syria’ ( Pallary, 1939: 102) .

In the type series, together with M. khabourensis we also found another species, which we assign to M. infracincta (see below); shells intermediate between these two species suggest some degree of hybridization.

Diagnosis

Melanopsis khabourensis differs from M. buccinoidea in that mouth-height is small relative to shelldiameter.

Description

The shell is smooth, stout and consists of seven whorls separated by shallow sutures. The spire is short, the callus weak and the columella broad. The notch is deep and from it a hollow ridge extends upwards, winding around and above the columella and ending in the mouth. This ridge separates the columella from the parietal lip with its callus. Shell colour is greyishbrown, usually with two dark bands; the columella is white. Conchometrics (N = 6) are in Table 4.

Distribution

Ras al ‘Ayn’ ( Fig. 5 ), a series of Karst springs in northern Syria which together form the main spring of the river Khabur, a tributary of the Euphrates, is on the boundary between two different landscapes, the mountains of Turkey in the north and the semidesert flats of Syria in the south ( Schütt & Sesen, 1989).

Mouth-height / Shell-diameter

Comparisons

A major difference between M. khabourensis and other smooth-shelled species is that the hollow ridge that extends from the notch upwards is very pronounced, and the columella is very wide. We could not conveniently measure and express this character in quantitative terms. In the conchometrics used in this study, M. khabourensis further differs from buccinoidea in that it has almost diagnostically lower ratios mouth-height/shell-diameter (1.01 ± 0.04 vs. 1.19 ± 0.07) and notch-width/notch-depth (0.70 ± 0.03 vs. 0.87 ± 0.08) ( Fig. 9 ). M. khabourensis differs from M. d ircaena in its diagnostically lower ratio notchwidth/mouth-diameter (0.38 ± 0.04 vs. 0.51 ± 0.10) and notch-width/notch-depth (0.70 ± 0.03 vs. 0.85 ± 0.08), and almost diagnostically higher ratio b/c (0.79 ± 0.02 vs. 0.73 ± 0.03) ( Tables 3, 4). It further differs from dircaena in its lower mouth-height relative to shell-height.

Within Melanopsis, Pallary (1939) placed M. khabourensis in Mesopotamia, a separate subgenus that he characterized by a shell with two distinct cords at the base of the lower whorl. Whether or not Mesopotamia should be a separate group is beyond the scope of this study. We note however, that M. khabourensis has only one ridge. Within Mesopotamia, Pallary (1939) described M. khabourensis as differing from other species in that it has only two dark bands on the shell, whereas other species have three; the limited material available to us supports this difference.

MELANOPSIS MEIOSTOMA HELLER & SIVAN, 2000

( FIG. 2F )

Melanopsis meiostoma Heller & Sivan, 2000 .

Material examined

Types: Holotype HUJ 7966 and 39 paratypes ( HUJ 7967 , all from ‘En Haruv’, a small spring in the southern parts of the Golan Heights.

Diagnosis

Melanopsis meiostoma differs from M. buccinoidea in that it almost always has a smaller mouth-height relative to shell-height.

Description

The shell is small, narrow, and has up to eight whorls. It has a pointed spire, flattened whorls separated by very shallow sutures, and is smooth. The mouth is small (relative to shell-height) and rather round, due to an outward flaring of the outer lip. Shell colour is uniform, dark-brown to black; also the columella and the (weakly developed) callus are blackish. Conchometrics of M. meiostoma are in Heller & Sivan (2000).

Distribution and habitat

M. meiostoma is recorded only from its type locality ‘En Haruv’ ( Fig. 5 ) a small brackish spring that pours into a cement pool, on the walls of which the snails were found.

Comparisons

M. meiostoma differs from M. buccinoidea in having a more narrow shell and smaller, more rounded mouth (lower ratios of shell-diameter/shell-height, mouthheight/shell-height and mouth-height/shell-diameter, higher ratio of mouth-diameter/mouth-height; Heller & Sivan, 2000: fig. 2). It also has a larger notch relative to mouth size (higher ratios notch-width/mouthdiameter and notch-depth/mouth-diameter; Heller & Sivan, 2000: fig. 3).

The sperm of M. meiostoma differs from that of M. buccinoidea in its larger nucleus, larger acrosome, and shorter midpiece ( Hodgson & Heller, 2000). The radula of M. meiostoma differs from that of M. buccinoidea in the long central cusp of its rachidian tooth ( Mazan-Mamczarz et al., 2002). In allozyme studies however, M. meiostoma invariably adjoins the cluster of M. saulcyi ( Falniowski et al., 2002a, b).

Conchometric differences between M. meiostoma and M. ammonis are slight. M. meiostoma has a smaller mouth-height, relative to shell-height and shell-diameter (0.47 ± 0.02 vs. 0.50 ± 0.03, 1.05 ± 0.05 vs. 1.14 ± 0.05), a larger mouth-diameter relative to mouth-height, a less conic shape, a wider notch (in relation to both notch-depth and to shell-diameter) and higher whorls.

Evidence concerning the separate identity of M. meiostoma is thus conflicting. It is placed as a species separate from M. buccinoidea on grounds of the shell, sperm and radula, but not so on grounds of electrophoresis; and as separate from M. ammonis on grounds of conchometric differences that are significant but not diagnostic.

MELANOPSIS COSTATA ( OLIVIER, 1804)

( FIG. 10 , TABLE 5)

Melania costata Olivier, 1804 : pl. 31, fig. 3.

Melanopsis costata, Pallary, 1939: 90–91 ; Heller et al., 1999: 59–61, fig. 4C; Bandel, 2000: 166, figs 72–75.

Melanopsis praemorsa jordanica, Mienis & Ortal, 1994 : II.

Type and type locality

‘Melanie a cotes, Melania costata ; de Orontes’ ( Olivier, 1804; pl. 31, fig. 3). Olivier’s type material is not found in the Muséum National d’Histoire Naturelle (Paris) and has probably been lost ( Tillier & Mordan, 1983).

Diagnosis

Melanopsis costata differs from M. buccinoidea in its ribbed shell.

Description

The shell is ribbed, except for its uppermost whorls. The ribs almost always extend the entire height of each whorl and most of them descend vertically; sometimes however, the ribs above the mouth may bend leftwards, towards the columella. The ribs are almost always uniform rather than bumpy and the upper section of each rib (near the suture) is almost always rounded. Shell colour, which varies from greyish yellow to reddish brown or black, may be uniform or banded. In the sperm of M. costata the nucleus and acrosome are longer and the midpiece shorter than in M. buccinoidea ( Hodgson & Heller, 2000) .

Distribution

Throughout the Levant, from Kara Sou in the north via Orontes River, coastal plain of northern Israel, Jordan River, Lake Kinneret, Yarmouk River and Azraq Oasis, down to the environs of the Dead Sea in the south ( Fig. 11 ). In the Levant, fossils of M. costata are known from the Mid- and Lower Pleistocene of the Jordan Valley ( Heller & Sivan, 2001, 2002a). Bandel (2000) suggested that M. costata , together with M. saulcyi and M. bandeli , forms a group that originated in the Balkan, during the Late Miocene or Early Pliocene.

M. costata - M. buccinoidea intermediates Intermediates between M. costata and M. buccinoidea are frequently reported in the literature. Germain (1921; figs 51–56) described and mentioned such intermediates from Qusayr (on the Orontes south of Homs), Jayrud (north-east of Damascus) and Baalbek; Schütt (1983) noted that intermediates occur at Cebelib (40 km north of Iskenderun), Samandagi (= Suwaidiya) on the Orontes and Qala’at al Mudiq.

Glaubrecht (1993: figs 5–9) described in detail three sites in the Kingdom of Jordan at which both smooth, ribbed and intermediate shells were found. However, as Glaubrecht assigned all ribbed specimens to M. costata (not being aware that another ribbed species, M. saulcyi , also occurs in this region; see below) his analysis of these populations should be approached with some caution. His fig. 5 suggests that at one of these sites, Ash Shunah, ribbed and smooth shells are well separated (unfortunately Glaubrecht blurred the picture by adding to this figure also his data from a nearby site, that of North Shuna). Further intermediates were found in the Jordan Valley, in three streams that flow into the Kinneret, and also in the lower Hasbani (a tributary of the Jordan; Heller et al., 1999); and in the Azraq oasis ( Bandel, 2000).

At least some of these intermediates are hybrids. Evidence for hybridization between M. costata and M. buccinoidea , but without merging of their genetic pools, was found also at the level of allozymes, in a study that suggested to Falniowski et al. (2002a, b) that M. costata speciated from M. buccinoidea by a mild founder effect. Further, M. costata - M. buccinoidea hybrids have been found at a Lower Pleistocene site of the Jordan Valley, in the southern Levant ( Heller & Sivan, 2002a).

M. costata - M. dircaena intermediates

Five samples from the Orontes (from the environs of Hamma) contain shells that seem intermediate between M. costata and M. dircaena : the shell is stout and the ribs are poorly developed. They may be hybrids between these two species. Transitions, of possible hybrid origin, between M. dircaena and M. costata were noted also by Pérès (1946).

Germain (1921; perhaps in the footsteps of Dautzenberg, 1894) described M. bullio from the Lake of Homs (without illustrating it) as having ribs that are pronounced near the suture, where they sometimes form tubercles, and gradually become shorter, until they disappear from the basal part of the last whorl. Pérès (1946) considered M. bullio a synonym of M. costata . He placed it as a separate variety only to mark a step in the scale of variation of costata , but noted a completely gradual transition between a typical M. costata and a typical M. bullio . The specimens determined as M. bullio by Germain are illustrated by Pérès (1946; pl. 2, figs 4–9). They may perhaps be M. costata - M. dircaena intermediates .

Intraspecific variation

Within M. costata we distinguish four groups that differ significantly (not always diagnostically) in their conchometrics and distribution. These groups are hereby described as four subspecies: M. c. costata , M. c. lampra , M. c. jordanica and M. c. obliqua .

MELANOPSIS COSTATA COSTATA

( FIG. 10A )

Melania costata Olivier, 1804 : pl. 31, fig. 3.? Melanopsis callista Bourguignat, 1884: 118 .? Melanopsis sesteri Bourguignat, 1884: 119 .? Melanopsis alepi Bourguignat, 1884: 119–120 . Melanopsis hiera Bourguignat, 1884: 121–122 .? Melanopsis hiera Bourguignat, 1884: 121 , from Lake Homs; not Melanopsis hiera Bourguignat, 1884: 121 , from Hula Valley (‘Ain Mallaha’).

Melanopsis costata var. hörnesi Blanckenhorn, 1897: 130 , pl. 10, figs 9, 10.

Melanopsis costata Olivier , cotypes: Pallary, 1939: pl. 5, figs 26-28.

Melanopsis costata var. strigosa Pallary, 1939: 90 , pl. 5, figs 35, 36.

Melanopsis costata var. subnodata Pallary, 1939: 90 , pl. 5, figs 30-32.

Melanopsis costata var. obliquata Pallary, 1939: 91 , pl. 5, fig. 34.

Melanopsis orontis var. elevata Pallary, 1939: 92 , pl. 6, fig. 74.

Not Melanopsis costata costata Heller et al., 1999: 61– 64 .

Types and type locality

‘Melanie a cotes, Melania costata ; de Orontes’ ( Olivier, 1804: pl. 31, fig. 3).

Material examined

Types: the six syntypes of Melanopsis costata , as determined by Pallary (1939) (BMNH 1937.12.30.10361– 66) are from the ‘R. Orontes’. Another lot, determined by Pallary as ‘locality original’ (BMNH 1933.8.11.352– 59) are from the Orontes at ‘Djishr ech Cheghour’, between Lattaquie and Aleppo; Pallary noted that this is where Olivier originally collected his M. costata .

Additional material: SYRIA: Orontes River and tributaries ( BMNH 1933.8.11.14–21, 1933.8.11.33–42, 20030346; NHMW 1838.4.16, 55113; SMF 167427, 282787, 282792, 282805, 282806, 282807, 282812, 282813, 282816, 282824, 283565, 290956, 290984); Aleppo ( SMF 271604 ) .

Description

The shell is elongate, with stepped whorls. The uppermost whorl is smooth, the six lower whorls are ribbed. Ribs extend almost the entire height of each whorl and most of them descend vertically, or in a slightly diagonal direction. The upper section of each rib (near the suture) is rounded. Shell colour is greyish yellow to almost black. Conchometrics (of 14 specimens, those labelled cotypes and those labelled as from Djisr ech Cheghour) are given in Table 5.

Synonyms

Blanckenhorn (1897) described Melanopsis costata var. hörnesi from Aleppo. The type specimen of M. c. hörnesi (NHMW 1838.4.16; Table 4) has slightly higher values of conicality and figurativity than in M. c. costata (conicality index 1.02 as compared to 0.90–0.99; figurativity index 1.06 as compared to 0.97–1.05); all its other conchometric characters fall within the range of c. costata . We therefore consider M. c. hörnesi a synonym of M. c. costata .

From the type locality of M. c. costata (Djishr ech Chegour), Pallary (1939) described six varieties of M. c. costata : inflexa, strigosa, subnodata, obliquata, inodata and lineata. These taxa are all so similar that we could not find conchometric gaps between them, whereby we consider them as synonyms of M. c. costata . Also listed as synonyms are M. orontis var. elevata Pallary from Djishr ech Cheghour, and a sample in the Pallary collection (BMNH 1933.8.11.14–21) labelled M. stephanota Bourguinat from Kara Sou, 80 km north of Djishr ech Cheghour. These taxa are all so similar that, again, we could not find conchometric gaps between them.

Distribution

Kara Sou, Sadjour Sou, Aleppo and the Orontes River ( Fig. 11 ).

Comments

In addition to the shells of M. c. costata that are typically characterized by well-developed ribs, many samples also contain shells with weak ribs, which are frequently not traceable on the last whorl. Such are the type specimens of M. alepi from the environs of Aleppo, M. callista from Sadjour Sou, and some of the M. sesteri from Sadjour Sou. These weak-ribbed shells may perhaps lie within the normal range of variation of M. c. costata , or they may perhaps represent an eastern form of M. c. costata , or they may even be hybrids between M. c. costata and some smooth species.

The M. costata of the upper Jordan Valley were previously considered as M. c. costata ( Heller et al., 1999) . However, the additional material available in this study reveals a conchometric gap between M. costata of the type locality on the one hand, and those of the upper Jordan Valley, some 300 km further south, on the other. We therefore reassign the upper Jordan Valley M. costata to a separate subspecies, M. c. lampra .

MELANOPSIS COSTATA LAMPRA BOURGUIGNAT

( FIG. 10B )

Melanopsis lampra Bourguignat, 1884: 132–133 . Melanopsis phoeneciaca Bourguignat, 1884: 133–134 . Melanopsis belusi Bourguignat, 1884: 134 . Melanopsis tanousi Leternoux , in Bourguignat, 1884: 137–138.

Diagnosis

M. c. lampra differs from M. c. costata in that it is almost always stouter and its whorls are almost always more conic.

Description

The shell is conic, the ribs are mildly pronounced and their shoulders are rounded. M. c. lampra further differs from M. c. costata in that it has a larger mouth (higher mouth-height/shell-height), higher number of ribs, and these are denser. Conchometrics are in Table 5 (from Heller et al., 1999; with additional data). The radula of M. c. lampra is described by Mazan- Mamczarz et al. (2002) and allozyme variation by Falniowski et al. (2002b).

Melanopsis jordanica Roth var. parvula Bourguignat, 1884: 141–142 .

Melanopsis eumorphia Bourguignat, 1884: 146 .

Melanopsis praemorsa costata, Schütt, 1983: 42–44 , pl. 2, figs 48–56.

Melanopsis costata costata, Heller et al.,1999: 61–64 , fig. 4B.

Melanopsis bovieri, Germain, 1921: 504–505 , pl. 20, figs 7, 8.

Material examined

Types: The syntypes of M. c. lampra ( MHNG unnumbered; N = 102) are from ‘Bélus, prés de Saint- Jean-d’Acre’ (environs of the Na’aman of today).

Additional material: SYRIA: Orontes ( SMF 282816). LEBANON: Hamik ( BMNH 1937.12.30.10341–44). JORDAN: Azraq ( ZMH). ISRAEL: seven sites along the upper Jordan River, as given by Heller et al. (1999).

Synonyms

From the swamp of Hamik (in Lebanon, near the Nahr ez Zair bridge on the route to Damascus; 33∞43¢N, 35∞35¢E) Pallary (in Germain, 1921: vol. 1: 504–505, vol. 2, pl. 20, figs 7, 8) described Melanopsis bovieri , in which the upper whorls are smooth and the lower ones sparsely ribbed, each with a nodosity on its upper part. Germain commented (p. 505) that this species differs from M. costata and M. saulcyi mainly in its ‘trochiform shape with a broad base’. Pallary (1939) illustrated M. bovieri (pl. 5, figs 40–43) and mentioned (p. 93) that on visiting the site in 1934, he found that this species had become rare; Pérès (1946) considered it a separate taxon, close to M. saulcyi . We examined four specimens of M. bovieri in the Pallary collection ( BMNH 1937.12.30.10341–44). They fall within the range of M. c. lampra ( Table 5), whereby we consider M. bovieri a synonym of M. c. lampra .

Distribution and habitat

M. c. lampra is distributed in the upper Jordan River (Hula Valley), coastal plain of northern Israel and, in the Kingdom of Jordan, the Azraq Oasis ( Fig. 11 ). The northern range of M. c. lampra is difficult to assess. We assign to M. c. lampra some specimens available to us from the region of Hamma, and to M. c. costata some others, from the same sample. We note that Pallary (1939: 91–92, pl. 5, fig. 29) described M. atramentatria from Acharne (on the Orontes, between Hamma and Qala’at al Madiq) and commented that this species (or variety) that differs from M. c. costata in its more solid, stout shape, lives in the Orontes together with M. c. costata . We consider M. atramentatria a synonym of M. c. lampra ; this extends the distribution of M. c. lampra to the north of Hamma.

In the Jordan Valley, the dominant habitats of M. c. lampra used to be Lake Hula (surface area 14 km 2, depth 1.5–2.5 m), its several tributaries and its outlet (the upper Jordan River). This lake was drained in 1958. Today this subspecies has a narrow, linear distribution along the upper Jordan River and its tributaries, where it is found near overhanging vegetation, especially of Salix ( Heller et al., 1999) . In the coastal plain of the southern Levant it was recently described from ‘En Afeq (biblical Na’aman, later named Belus) on muddy substrates, submerged plants and bits of vegetation (Heller et al., 2002).

Comparisons and intraspecific variation

As compared to M. c. costata , in M. c. lampra the shell is almost always stouter (shell diameter/shell height 0.48 ± 0.03 vs. 0.44 ± 0.03), has a larger mouth-height compared to shell-height (0.54 ± 0.03 vs. 0.50 ± 0.03), is more conic (0.86 ± 0.05 vs. 0.93 ± 0.05), and has significantly more ribs (11.0 ± 1.14 vs. 9.4 ± 1.1) ( Table 5). Differences between M. c. lampra and M. c. costata are best seen in shells of above 19 mm shell-height and therefore, in Figure 12 , only such shells are presented. In addition, the ribs of M. c. lampra are narrower. Consequently, M. c. lampra clusters separately from M. c. costata ( Fig. 13 ).

M. c. lampra from the coastal plain of the southern Levant differs from that of the Jordan Valley in that it has fewer ribs, a more shallow notch, a higher rib index and it is more conic (Heller et al., 2002). However, the extent of overlap between the coastal plain and Jordan Valley populations is so considerable that we consider them conspecific.

MELANOPSIS COSTATA JORDANICA ROTH

( FIG. 10C )

Melanopsis costata var. jordanica Roth, 1839: 25 , pl. 2, figs 12, 13.

Melanopsis jordanica, Rossmässler, 1854: 17 , 905; Bandel, 2000: 176, figs 127, 128.

Melanopsis ovum Bourguignat, 1884: 143 .

Melanopsis costata jordanica Roth, Heller et al., 1999: 64–67 , fig. 4D.

Material examined

Holotype: Melanopsis costata var. jordanica from ‘mari Galilaeo’ in ZSM.

Additional material: ISRAEL: Six sites along the shoreline of Lake Kinneret, as in Heller et al. (1999).

Diagnosis

Melanopsis costata jordanica differs diagnostically from M. c. costata in that its shell is stouter and its

Shell-height (mm) Axis 1

mouth larger. In these characters it differs also from M. c. lampra , though in statistically significant, not diagnostic, terms.

Description

The shell is stout. The ribs are mildly pronounced, with rounded shoulders. Shell colour varies: some are black but many others are banded, the dark bands being broad and black, the pale bands narrow and pale grey to dark brown. The last whorl of each shell has three dark bands and two pale ones. Conchometrics are in Table 5 (from Heller et al., 1999 and from additional data).

Distribution and habitat

The many samples of M. c. jordanica in the various collections we inspected are all from Lake Kinneret. One exceptional sample from Damascus (coll. Capt. Burton, BMNH) is the only record beyond Lake Kinneret. As there is no further evidence of M. c. jordanica in the well-collected region of Damascus, we reject Damascus as a mislabelling.

Lake Kinneret is about 21 by 12 km and frequent daily winds cause it to become wavy, choppy or stormy. Its natural salinity was once almost 400 mg / L (chlorinity) but recent diversion of haline springs from the lake has decreased salinity levels to about 200 mg /L. Within the Kinneret M. c. jordanica occurs along rocky shores consisting of gravel, cobble, stones and boulders, down to a depth of 5 m; it is not found on shores consisting of mud, sand and silt. Two snail-predating fish, Barbus longiceps (Valenciennes, 1842) and Blennius fluviatilis Asso, 1801 , are found in Israel, mainly in Lake Kinneret, and they may perhaps exert selection on shell colours ( Heller, 1979).

Synonyms

From Lake Kinneret, Bourguignat (1884) described Melanopsis ovum . Upon inspecting and measuring the type series of M. ovum ( BGT 11395, N = 11) we find no differences between it and M. c. jordanica ; hence we consider M. ovum a synonym.

Comments

As suggested by Figure 14 , M. c. jordanica is well separated from M. c. lampra . Falniowski et al. (2002a, b), in an analysis of allozyme data, found that populations of M. c. jordanica cluster close to but separately from populations of M. c. lampra . Also, the radula of M. c. jordanica clusters separately from that of M. c. lampra ( Mazan-Mamczarz et al., 2002) .

Axis 1

MELANOPSIS COSTATA OBLIQUA BOURGUIGNAT

( FIG. 10D )

Melanopsis obliqua Bourguignat, 1884: 138 .

Melanopsis eumorphia Bourguignat, 1884: 146 .

Melanopsis egregia Bourguignat, 1884: 146 .

Melanopsis noetlingi Bourguignat in Noetling, 1886: 817, pl. 23, fig. 6; Bandel, 2000: 171–2, figs 105, 106.

Melanopsis lampra, Pallary, 1939: 93–94 , pl. 6, figs 45, 46.

Melanopsis costata noetlingi Bourguignat, Heller et al., 1999: 67–68 , fig. 4E.

Material examined

Holotype: Melanopsis obliqua from ‘ Le Belus pres St. Jean d’Acre’ , BGT 11394.

Additional material: lower Jordan River: ( BMNH 1937.12.30.10366–77, SMF 111879, 168956, NHMW 52309, Allenby Bridge ( HUJ); Yarmouk River ( HUJ).

Diagnosis

Melanopsis costata obliqua differs from M. c. costata in that in all specimens the ribs on the last whorl extend the entire length of the whorl. Each rib consists of a very prominent upper tubercle, fused to a straight, prominent, lower ridge.

Description

The shell is conic to elongate. The upper three whorls are smooth whereas the lower ones are ribbed. The ribs on the last whorl are very high and prominent, and extend the entire height of each whorl. The upper section of each rib (near the suture) consists of a very prominent upper tubercle, fused to a straight, prominent, lower ridge. The tubercle and the ridge are either continuous, or separated by a depression. Shell colour is greyish yellow to almost black. Conchometrics are in Table 5 (all material: Yarmouk and lower Jordan; N = 63).

Synonyms

M. eumorphia , M. egregia and M. noetlingi (and also the specimen from the Yarmouk named M. lampra by Pallary, 1939; pl. 6, figs 45, 46) all have ribs in which there is a very prominent upper tubercle, fused to a prominent, lower ridge, and on this basis we consider them synonyms of M. c. obliqua .

Distribution and habitat

M. c. obliqua is known from the lower Jordan Valley and the coastal plain of the southern Levant ( Fig. 11 ). In the lower Jordan Valley, the current in the outlet of the Yarmouk is torrential and we found M. c. obliqua on boulders; in the lower Jordan River the current is slow and we found it on submerged vegetation. The coastal plain (Belus, today named Na’aman) is the type locality of M. c. obliqua ; in a recent survey we did not find M. c. obliqua in this region (Heller et al., 2002).

Comparisons

General conchometrics of M. c. obliqua are similar to those of M. c. costata , but its tubercles, combined with its longer ribs, place it in a separate position. The tubercles also separate it from M. c. lampra and M. c. jordanica , which are closer to M. c. obliqua in geographical terms. A sample recently collected from the lower Jordan is of smaller size and is less tuberculose than specimens collected in the prepollution era of the nineteenth and early twentieth centuries.

MELANOPSIS SAULCYI BOURGUIGNAT, 1853

( FIG. 15A, TABLE 6)

Melanopsis saulcyi Bourguignat, 1853: 66 , pl. 2, figs 53, 54.

Melanopsis hiera Bourguignat, 1884: 121–122 .

Melanopsis jebusitica Bourguignat, 1884: 126 .

Melanopsis saulcyi Bourguignat, 1853, 1884: 127 ; Mienis & Ortal (1994: II; Heller et al., 1999: 68–72, fig. 4F, G; Bandel, 2000: 176, figs 76-79, 84; Heller & Sivan, 2002b: 601–603, fig. 3E.

Melanopsis aterrima Bourguignat, 1884: 127–128 .

Melanopsis faseolaria, Bourguignat, 1884: 128 .

Melanopsis sancta, Bourguignat, 1884: 129 .

Melanopsis cerithiopsis Bourguignat, 1884: 130–131 ; Pallary, 1939: 95, pl. 4, figs 18, 20, pl. 6, figs 47–50, 53, 54, 73.

Melanopsis cerithiopsis var. curta Bourguignat, 1884: 131 , from Hula Valley (‘Ain el Mellaha).

Melanopsis desertorum Bourguignat, 1884: 134–135 , from Tarsous ( Anatolia, the type) and Belus.

Melanopsis costata var. luteopsis Germain, 1921: 494 , pl. 20, figs 9, 10.

Melanopsis palmyrensis Pallary, 1939: 96 , pl. 6, figs 60-62.

Melanopsis praemorsa obsolete, Schütt, 1987: 66 , fig. 5(3).

Not Melanopsis var. obsoleta Martens 1874: 32 , pl. 5, fig. 39.

Melanopsis praemorsa bandeli Schütt, 1988: 216 , fig. 7.

Melanopsis cerithiopsis Bourguignat, 1884 , Mienis & Ortal, 1994: II.

Melanopsis bandeli, Bandel, 2000: 176–77 , figs 80–83, 85-93, 95.

Material examined

Types: four syntypes ( BGT 11358) from ‘Artouse, en Syrie’ ( Bourguignat, 1853: 66). The type locality of M. saulcyi , given by Bourguignat, 1853: 66) as ‘Artouse, en Syrie’, is problematic, because Pallary (1939) argued that there is no such site in Syria; he suggested that the type locality should be Tartous, on the Syrian coast ( Fig. 16 ). An inscription on the label of the type specimen also mentions Artous as being near Jericho. However, Artous (Artuz in modern maps) is a site south-west of Damascus. To conclude, within the southern Levant we could not determine the precise whereabouts of Artous, the type locality of M. saulcyi . Additional material: SYRIA: vicinity of Homs ( SMF 290940, 290952, 290954, 29059, NHMW 29382, 30407); Palmyra ( SMF 282826, 282827, NHMW 84546). ISRAEL: coastal plain: Nahal Kishon ( BMNH 20030249, NHMW 52219), Nahal Taninim ( HUJ); Jordan Valley: Enot Huga, En Hanaziv, Hamat Gader (all HUJ).

Diagnosis

Melanopsis saulcyi differs from Melanopsis c. costata in that it has more ribs, and these are usually bumpier.

Description

The shell is usually elongate, and its mouth-height small in relation to shell-height. The 2–4 upper whorls are smooth, the 3–6 lower ones are ribbed. In most shells, the ribs descend from the suture to about the middle of the lowest whorl; in others they descend further, sometimes exceptionally so, even extending to the base of the whorl. Each rib usually consists of an upper tubercle, fused to a lower ridge. Consequently the rib normally has a waist between the tubercle and ridge; this waist is present in all ribbed whorls (not only in the ultimate whorl, as is sometimes the case in M. costata ). When the ribs extend the entire height of the lower whorl this is due to the appearance of an additional ridge, beneath the previous (upper) one; eventually the two ridges fuse to a varying degree. Usually, the overall appearance of the shell is bumpy. Shell colour varies from uniform pale brown via dark brown to uniform black. On the inner lip the pale callus of the upper parietal wall merges with the white columella, so that the entire inner lip is pale. Conchometrics of M. saulcyi are in Table 6.

Synonyms

Under his heading saulcyana, Bourguignat (1884: 75) placed six species: saulcyi from Artouse, Ain Soultan and Ain el Bass; jebusitica from near Jericho; aterrima from Ain Soultan; faseolaria from Ain Mallaha and Belus; sancta from Jordan 4 km from the Dead Sea, Ain el Placa (Hula Valley) and Ain Soultan; and cerithiopsis from Ain el Mallaha. We inspected Bourguignat’s types of these species and as we found no significant conchometric differences among them, we consider them all as synonyms of M. saulcyi (as suggested also by Germain, 1921).

Melanopsis costata var. curta , a small shell (height 16 mm) with ribs that are not well developed and that fade out at the base of the lower whorl, was described (but not illustrated) from Lake Homs and Lake Antioch by Locard (1883) and Germain (1921). From these descriptions we find no differences between M. c. var. curta , with its small shell and ribs that fade out in the lower part of the last whorl, and a typical M. saulcy i; hence we consider M. c. var. curta a synonym of M. saulcyi . From Lake Homs and also from Jayrud, Germain (1921) described and illustrated M. c. var. luteopsis . We see no differences between M. c. var. luteopsis , with its slim shell and ribs that fade out in the lower part of the last whorl, and a typical M. saulcy i, so we consider it a synonym of M. saulcyi .

From Palmyra, Dautzenberg (1894) and Germain (1921) described M. saulcyi var. obsoleta as having a small size, slender shell and faint ribs. Schütt (1987) removed this variety from M. saulcyi and placed it as a separate subspecies, M. praemorsa obsoleta . Schütt suggested that M. p. obsoleta is an endemic relict of the Pliocene Melanopsis inhabiting the Pliocene connection between the Euphrates and the Jordan Valley via the basins of Palmyra, Qaryatain and Damascus, and that M. cerithiopsis Bourguignat of the Jordan Valley is its closest relative.

In a previous study ( Heller et al., 1999) we presented evidence that M. cerithiopsis is a synonym of M. saulcyi . Here we present conchometrics of three samples (eight specimens) of Melanopsis from Palmyra described as M. p. obsoleta by Schütt ( Table 6). The shells have ribs similar to those of M. saulcyi , but in some of them the ribs are diffuse. The shells from Palmyra have a slightly smaller mouth than those of M. saulcyi from the southern Levant, giving small differences in all the ratios that include mouth-height; these differences are not diagnostic and the overlap is so great that we consider M. p. obsoleta a synonym of M. saulcyi .

From Syria (Palmyra) Pallary (1939: pl. 6, figs 62- 64) described M. palmyrensis , which was considered by Schütt (1983, 1987, 1988: fig. 6) as M. praemorsa obsoleta . M. p. obsoleta falls within the range of M. saulcyi , so we consider it a synonym.

From the Kingdom of Jordan, Schütt (1988; fig. 7) described Melanopsis praemorsa bandeli as having a narrow shell with dense ribs that do not reach down to the base of the whorl. This description closely follows

Axis 1

that Bandel assigned to M. bandeli ; so too are the 13– 16 mm M. saulcyi measured by Germain (1921: 503). A similar size range is found in M. saulcyi of the Jordan Valley ( Heller et al., 1999: fig. 12). These data do not suggest a separate taxon within the shell-height range of 13 mm. We see no character with which to distinguish M. bandeli from M. saulcyi , and thus we consider it a synonym.

that of M. saulcyi . M. bandeli was re-described by Bandel (2000: figs 80–83, 85-93, 95), from the Kingdom of Jordan (both Recent and from fossil layers ‘just before the Brackish Lisan Lake’. Bandel (2000: 177) described M. saulcyi as smaller and shorter than M. bandeli without giving any further differences between these two species. From one of his sites in the Jordan Valley, he (p.177, figs 94, 96) noted transitions between M. bandeli and M. saulcyi but observed that the majority of individuals belong to saulcyi . As concerns size, for M. saulcyi Bandel gave a shell-height of 13 mm (his figs 78, 84), whereas for M. bandeli he gave shell-heights of 13–20 mm (his figs 80–83, 85–93).

It is noteworthy, however, that Bourguignat (1853) described the type of M. saulcyi as reaching 15 mm and Roth (1855) described M. saulcyi as having a shell-height of 18.5 mm, both well within the range Distribution and habitat

M. saulcyi is widely distributed in the Levant ( Fig. 16 ) in springs and streams, where it occurs on boulders and also on aquatic and submerged vegetation. It is our impression that, up to the beginning of the twentieth century, the original habitat of M. saulcyi used to be streams that border swamps; today, no swamps remain ( Heller et al., 1999, 2002).

Comparisons

In Figure 17 M . saulcyi clusters separately from M. costata . M. saulcyi differs from M. costata mainly in its more numerous, dense and short ribs, and also in that each rib, on the penultimate whorl, has a waist (for the southern Levant the number of ribs is 13.1 ± 2.2, rib density 4.15 ± 0.90, rib length 2.46 ± 0.40). Within M. costata , it differs from M. c. costata also in that it is more conic (a/b 0.84 ± 0.05 vs. 0.93 ± 0.05), less stepped (b/c 0.80 ± 0.03 vs. 0.74 ± 0.04), has lower figurativity index (c/d 0.93 ± 0.02 vs. 1.00 ± 0.04), and a smaller ratio 2-rib width/shell-diameter. It differs from M. c. lampra , to which it is close in geographical terms, in its slimmer shell and smaller mouth (see Heller et al., 1999). All of these differences are significant, not diagnostic.

M. saulcyi falls well away from M. costata also in allozyme investigations, with M. buccinoidea scattered among both clusters ( Falniowski et al., 2002a, b). Mazan-Mamczarz et al. (2002) described differences in the radula between M. saulcyi and M. costata . The sperm of M. saulcyi differs from that of M. costata in having a larger acrosome and shorter midpiece ( Hodgson & Heller, 2000).

Intraspecific variability

Within M. saulcyi , the type specimens of M. saulcyi (BGT 11358) are not typical of the species: they do have numerous dense, short ribs, but they lack the bumpy appearance, and their mouth heights are large. In the Bourguignat collection, the M. saulcyi from Ain Sultan (BGT 11357) are more typical of M. saulcyi .

Samples from Homs differ from those of the Jordan Valley in that they almost always have a higher figurativity index (0.97 ± 0.03 vs. 0.93 ± 0.02), often have less ribs (10.2 ± 1.33 vs. 13.1 ± 2.2), and lower rib density (2.92 ± 0.90 vs. 4.15 ± 0.90). They further differ in that they are significantly more stepped, the ribs are wider as compared to shell-diameter (2-rib width/ shell-diameter 0.41 ± 0.07 vs. 0.30 ± 0.04) ( Table 6). Consequently, within M. saulcyi the Homs region samples cluster separately from those from the Jordan Valley ( Fig. 18 ).

Within M. saulcyi of the Jordan Valley there is considerable variation in rib length; in one population (Sheluhot) they were exceptionally long ( Heller et al., 1999). Similar long ribs were found in the Bourguignat collection also in shells from Ain Mallaha, and named by Bourguignat as M. cerithiopsis (BGT 11349–50) and M. faseolaria (BGT 11351).

In the southern Levant hybrids were found between M. saulcyi and M. buccinoidea . As this hybridization has not resulted in the merging of the genetic pools of the two species, its occurrence does not reject the species distinctness of M. saulcyi and M. buccinoidea ( Heller et al., 1999, 2002; Falniowski et al., 2002a, b).

Comments

Falniowski et al. (2002b) suggested that M. buccinoidea , with its high genetic variability, may be the ancestral species from which M. saulcyi eventually speciated, perhaps by a founder effect and the allopatric speciation of peripheral isolates.

MELANOPSIS PACHYA PALLARY, 1939

( FIG. 15B, TABLE 7)

Melanopsis pachya Pallary, 1939: 86 , pl. 5, figs 10,12.

Material examined

Types: eight syntypes ( BMNH 1933.8.11.372–79) from ‘ Dans les Sources de Mézérib , au N. O. de Derâa (Syrie méridionale)’. Deraa is in southern Syria, on the bor-

Axis 1

Deraa N = 8 M. pachya

Max. shell-height 29.3 mm

Max. shell-diameter 16.3 mm

Max. mouth-height 14.4 mm Shell-diameter/shell-height 0.50 ± 0.01/0.49–0.52 Mouth-height/shell-height 0.59 ± 0.02/0.59–0.62 Mouth-diameter/mouth-height 0.41 ± 0.02/0.39–0.44 Mouth-height/shell-diameter 1.15 ± 0.05/1.06–0.20 Notch-width/notch-depth 0.83 ± 0.05/0.77–0.88 Notch-width/mouth-diameter 0.51 ± 0.04/0.46– 0.56 e /mouth-height 0.30 ± 0.04/0.24–0.35 f/mouth-height 0.18 ± 0.02/0.14–0.20 Conicality (a/b) 0.74 ± 0.04/0.69–0.81 Figurativity (c/d) 0.92 ± 0.02/0.87–0.95 Shouldering (b/c) 0.79 ± 0.01/0.77–0.80 Number of ribs 15.9 ± 1.57/13–17 Rib length 1.70 ± 0.67/1–2.5

Rib density 2.67 ± 0.29/2.5–3

2 rib width/shell- diameter 0.24 ± 0.03/0.22–0.28 der with the kingdom of Jordan; Mezerib is a small spring some 5 km further north-west.

Diagnosis

Melanopsis pachya differs from M. costata in that it almost always has more ribs; these usually reach only half the length of the whorl.

Description

The shell is thick and heavy. The upper whorls are eroded in all specimens available, the three lower whorls are weakly ribbed. The ribs, which descend vertically, may extend the entire height of each whorl; or, in other specimens, they may reach to about the middle of the last whorl. The upper section of each rib (near the suture) is rounded. The inner lip is whitish. Conchometrics are in Table 7. As all shells are eroded, shellheight was calculated by adding an estimate of about 0.8 mm to the measured shell-height (this estimate was reached by comparing the pachya shells with noneroded shells of other Melanopsis taxa of similar form).

Distribution

M. pachya is known only from the type locality (Deraa) ( Fig. 16 ).

Comparisons

In geographical terms, M. pachya is close to M. c. lampra of the upper Jordan Valley. M. pachya differs from M. c. lampra in its rib parameters: it has more ribs (15.9 ± 1.57 vs. 11.0 ± 1.14), that are usually shorter (1.70 ± 0.67 vs. 3.55 ± 0.43) ( Fig. 19A ) and it has significantly lower values of the ratio 2-rib width/shelldiameter (0.24 ± 0.03 vs. 0.35 ± 0.05). It further differs from M. c. lampra in its significantly higher ratios of shell-diameter/shell-height, and mouth-height/shellheight; and lower mouth-diameter/mouth-height ratio and conicality index ( Tables 5, 7). Consequently, M. pachya falls separately from M. c. lampra , in the PCO ( Fig. 19B ). M. pachya differs from M. saulcyi in that its ribs do not consist of an upper tubercle fused to a lower ridge. Consequently, the ribs do not have a waist between the tubercle and ridge.

Axis 1

MELANOPSIS GERMAINI PALLARY, 1939

( FIG. 15C, TABLE 8)

Melanopsis germaini Pallary, 1939: 93 , pl. 6, figs 5-8, 14.

Material examined

Types: three syntypes in MNHN (unnumbered) from ‘ Sources du Nahr es Sine, au Sud de Lattaquie, sur la route de Beyrouth’ .

Additional material: SYRIA: Nahr Senne: Paratype ( BMNH 20030571 ) , and five specimens ( BMNH 1933.8.11.51–55) determined as M. germaini by Pallary; five specimens collected by Pallary, and four additional specimens in MNHN; five topotypes ( NMW 1955.158 . 1727); Nahr as Zirud ( SMF 282739 ) .

Diagnosis

Melanopsis germaini differs from M. costata in that it has at least 15 and usually 17 ribs. It differs from M. pachya in that it has longer ribs and is not so stout.

Description

The shell is conic. The upper whorls are eroded in all specimens, to such an extent that it is not possible to determine whether they were smooth or ribbed; the five lower whorls have well-defined, regularly spaced ribs which descend, usually vertically but sometimes with a slight curve to the right, almost the entire Nahr es Sine N = 23 (pooled) M. germaini Max. shell-height 29.7 mm

Max. shell-diameter 13.3 mm

Max. Mouth-height 14.7 mm Shell-diameter/shell-height 0.47 ± 0.02/0.44–0.51 Mouth-height/shell-height 0.53 ± 0.03/0.46–0.57 Mouth-diameter/mouth-height 0.44 ± 0.02/0.40–0.48 Mouth-height/shell-diameter 1.13 ± 0.04/1.05–1.21 Notch-width/notch-depth 0.90 ± 0.07/0.77–1.00 Notch-width/mouth-diameter 0.50 ± 0.06/0.35– 0.57 e /mouth-height 0.36 ± 0.04/0.28–0.42 f/mouth-height 0.23 ± 0.04/0.17–0.33 Conicality (a/b) 0.77 ± 0.05/0.66–0.84 Figurativity (c/d) 0.93 ± 0.02/0.90–0.95 Shouldering (b/c) 0.81 ± 0.02/0.78–0.84 Number of ribs 17.6 ± 2.01/15–21 Rib length 3.6 ± 0.30/3–4

Rib density 3.4 ± 0.55/2–4.5

2 rib width/shell- diameter 0.24 ± 0.04/0.18–0.31 height of the last whorl. The upper section of each rib (near the suture) is rounded. The inner lip is pale brown. Conchometrics are in Table 8.

Distribution

M. germaini is known only from its type locality, Nahr Senne (on the Syrian coast, north of Banias) ( Fig. 16 ), and the near environment.

Comparisons

M. germaini differs diagnostically from M. c. costata of northern Syria (to which it is closer than other subspecies of M. costata , in geographical terms) in its number of ribs (17.6 ± 2.01 vs. 9.4 ± 1.1), relative rib width (2 rib width/shell-diameter 0.24 ± 0.04 vs. 0.48 ± 0.07) and conicality index (0.77 ± 0.05 vs. 0.93 ± 0.05) ( Tables 5, 8 and Fig. 20A, B ). It further differs significantly from M. c. costata in the ratios shell-diameter/ shell-height, mouth-height/shell-height, figurativity, shouldering, e/mouth height, rib density and rib length. Consequently, M. germaini is clearly separated from M. c. costata in the PCO ( Fig. 20C ).

MELANOPSIS INFRACINCTA MARTENS, 1874

( FIG. 15D, TABLE 9)

Melanopsis costata var. infracincta Martens, 1874: 32 , pl. 5, fig. 38.

Melanopsis (Mesopotamia) nodosa, Pallary, 1939: 98– 99 , pl. 4, fig. 21.

Melanopsis (Mesopotamia) mesopotamica Pallary, 1939: 99–100 , pl. 5, figs 1–7.

Melanopsis (Mesopotamia) infracincta, Pallary, 1939: 103–104 , pl. 4, figs 13–17.

Melanopsis (Mesopotamia) khabourensis var. plicata Pallary, 1939: 103–104 , pl. 5, fig. 19.

Melanopsis (Mesopotamia) cheragragensis Pallary, 1939: 104–105 , pl. 5, figs 8, 9, 11.

Types and type locality

‘ Quellen des Chabour bei Ras-el-ain’. The Khabour , in north-eastern Syria, is a tributary of the Euphrates and Ras al ‘ Ain’ is its major source. We did not find the original types .

Ras al ‘Ayn’ N = 8 M. infracincta Max. shell-height 24.3 mm

Max. shell-diameter 12.1 mm

Max. Mouth-height 11.1 mm Shell-diameter/shell-height 0.52 ± 0.02/0.49–0.54 Mouth-height/shell-height 0.49 ± 0.03/0.45–0.53 Mouth-diameter/mouth-height 0.48 ± 0.03/0.45–0.54 Mouth-height/shell-diameter 0.95 ± 0.03/0.90–0.98 Notch-width/notch-depth 0.78 ± 0.16/0.50–0.97 Notch-width/mouth-diameter 0.48 ± 0.12/0.25– 0.61 e /mouth-height 0.48 ± 0.04/0.44–0.55 f/mouth-height 0.30 ± 0.04/0.26–0.36 Conicality (a/b) 0.90 ± 0.11/0.76–1.06 Figurativity (c/d) 1.01 ± 0.05/0.94–1.08 Shouldering (b/c) 0.72 ± 0.06/0.66–0.82 Number of ribs 9.0 ± 0.93/8–11

Rib length –

Rib density 1.63 ± 0.35/1–2

2 rib width/shell- diameter 0.44 ± 0.06/0.35–0.51 Material examined

Five specimens ( BMNH 1937.12.30.293–302) and three specimens ( SMF 290976 ), all labelled M. infracincta from Ras al ‘ Ain’, Syria .

Diagnosis

Melanopsis infracincta differs from M. c. costata in that it has diagnostically higher values of the ratio shell-diameter/shell-height and a significantly higher mouth-height/shell-diameter.

Description

The shell is elongate and its overall appearance is knobby. The upper whorls are eroded in all specimens, to such an extent that it is not possible to determine whether they were smooth or ribbed; the four to five lower whorls have elaborate ribs which descend, either vertically or diagonally to the left. The upper section of each rib (near the suture) consists of a huge tubercle that may reach gigantic size. Beneath it and fused to it, a narrow ridge extends downwards. This ridge may expand considerably in its lower part to form a lower tubercle, which may be subequal in size to the upper tubercle. On the lowermost whorl a pronounced ridge starts above the columella, curves

Shell-height (mm) Shell-diameter (mm)

downward flanking the columella to the left and terminates as the notch. A second ridge starts near the callus, curves downward around the whorl and terminates on the lower part of the upper lip. Shell colour is black or dark brown to greyish yellow. Conchometrics are in Table 9.

Synonyms

Melanopsis arrousiana and M. stolliana , two further species described by Pallary (1939) from ‘Ain Arous’ (his pl. 5, figs 22-25, 44–48) are probably synonyms of M. infracincta .

Melanopsis (Mesopotamia) khabourensis var. dolichosoma Pallary is characterized by upper whorls with vertical ribs, and lower whorls that are smooth. We consider dolichosoma an intermediate, perhaps a hybrid, between M. khabourensis and M. infracincta .

Schütt & Sesen (1989) suggested that M. infracincta and M. khabourensis may ‘belong to Melanopsis nodosa Férussac’ which they considered a highly variable species. M. infracincta is indeed similar to M. nodosa in its complexity of rib structure; the precise relations between these two species are beyond the scope of this study.

Distribution

M. infracincta is known from its type locality, Ras al ‘Ayn on the Khabur’. It is also known from ‘Ain ar Arous’, the type locality of M. mesopotamica , and from Tell Abiad near Nahr Balik, the type locality of M. cheragragensis . All of these localities are from tributaries of the Euphrates, in north-eastern Syria.

Comparisons

In its bumpy appearance, M. infracincta differs from all other ribbed species here described. However, bumpiness is difficult to express in quantitative terms by the tools available to us. In the conchometric terms of this study, M. infracincta differs from M. c. costata in that it is almost diagnostically stouter (shelldiameter/shell-height 0.52 ± 0.02 vs. 0.44 ± 0.03) and the mouth is smaller relative to shell diameter (0.95 ± 0.03 vs. 1.12 ± 0.07) ( Tables 5, 9 and Fig. 21A, B ). Further, M. infracincta has significantly higher ratios of mouth-diameter/mouth-height, e/mouth-height, f/ mouth-height and its ribs are less dense.

For the Melanopsis of Mesopotamia, Pallary (1939) established a separate section Mesopotamia, characterized by the presence of two distinct ridges at the base of the lowermost whorl that terminate on the left of the columella. He commented that Mesopotamia seems very close to a fossil group of melanopsids, Melanoptychya.

MELANOPSIS SP.

( FIG. 15E, TABLE 10)

A sample of six shells (NHMW 15826) are labelled ‘ costata , Ras-el-Ain, near Baalbek, Syria. Leg. Dr F. Käufel; Walter Klemm collection’. These shells differ from Melanopsis costata in their larger mouth-height, and in that each rib has an upper tubercle, which tends to fuse with neighbouring tubercles, so as to form a pronounced ring. The three upper whorls are smooth, the three lower whorls are ribbed. Ribs extend the entire height of each whorl and descend vertically. The upper section of each rib (near the suture) consists of a pronounced tubercle, from which a low ridge extends vertically, until it reaches a lower tubercle; from here a thick, well-developed ridge extends downward, to the mouth. Strong ridges extend from one upper tubercle to the next, thereby forming a thick, ‘articulate’ ring, immediately beneath the suture. The overall appearance of the body whorl is as consisting of a ring, beneath which there is an indentation, beneath which ribs emerge from prominent tubercles and extend downward to the mouth. In the mouth, the anal slot is exceptionally long and narrow, expanding somewhat in the upper part. Shell colour is horny brown to black, the callus and inner lip are white. Conchometrics (N = 6) are in Table 10.

Melanopsis sp. is so closely similar to M. letourneuxi Bourguignat from northern Africa (BMNH 1937.12.30.10.113–18, 1937.12.30.10.28–32 and 1937.12.30.10.35–40) as to suggest that it may perhaps be conspecific and mislabelled as to its locality.

Baalbek N = 6 M. sp.?

Max. shell-height 18.0 mm

Max. shell-diameter 9.6 mm

Max. Mouth-height 12.5 mm Shell-diameter/shell-height 0.50 ± 0.02/0.48–0.53 Mouth-height/shell-height 0.67 ± 0.04/0.61–0.71 Mouth-diameter/mouth-height 0.39 ± 0.01/0.38–0.41 Mouth-height/shell-diameter 1.33 ± 0.05/1.26–1.40 Notch-width/notch-depth 0.76 ± 0.04/0.69–0.81 Notch-width/mouth-diameter 0.58 ± 0.03/0.55– 0.60 e /mouth-height 0.29 ± 0.05/0.21–0.37 f/mouth-height 0.15 ± 0.02/0.11–0.18 Conicality (a/b) 0.89 ± 0.06/0.83–0.97 Figurativity (c/d) 0.95 ± 0.05/0.88–1.02 Shouldering (b/c) 0.68 ± 0.07/0.59–0.76 Number of ribs 8.2 ± 0.41/8–9

Rib length 3.92 ± 0.20/3.5–4

Rib density 2.50 ± 0.32/2.5–3

2 rib width/shell-diameter 0.44 ± 0.03/0.39–0.46

Figures

Figure 2. Smooth-shelled Melanopsis of the Levant. A & B, M. buccinoidea (elongate and Jordan Valley forms). C, M. ammonis. D, M. dircaena. E, M. khabourensis. F, M. meiostoma.

Figure 3. Distribution of Melanopsis buccinoidea in the Levant (inspected records).

Figure 4. Melanopsis buccinoidea from the Jordan Valley (excluding Dan and ‘En Bass’) (Δ) compared with M. buccinoidea from elsewhere in the Levant (O). A, shellheight vs. shell-diameter; B, mouth-height vs. shell-height; C, whorl-height vs. mouth-height.

Figure 5. Distribution of Melanopsis ammonis (), M. dircaena (•), M. khabourensis (Δ) and M. meiostoma (-) in the Levant (inspected records).

Figure 6. Whorl-height vs. mouth-height. A, Melanopsis ammonis of the Jordan Valley (Δ) compared with M. buccinoidea of the Jordan Valley (O); B, M. ammonis of the Jordan Valley (Δ) compared with M. buccinoidea of elsewhere in the Levant (O).

Figure 7. Melanopsis ammonis and M. buccinoidea of the Jordan Valley catchment area, UPGMA dendrogram.

Figure 8. Melanopsis dircaena (Δ) and M. buccinoidea (O). A, shouldering vs. the ratio shell-diameter/shellheight, B, PCO scores.

Figure 9. Melanopsis khabourensis (Δ) and M. buccinoidea (O). The notch-width/notch-depth vs. mouth-height/ shell-diameter ratio.

Figure 10. Melanopsis costata of the Levant. A, M. c. costata. B, M. c. lampra. C, M. c. jordani. D, M. c. obliqua.

Figure 11. Distribution of Melanopsis costata costata in the Levant (inspected records). M. c. costata (-), M. c. lampra (•), M. c. jordani (Δ) and M. c. obliqua ().

Figure 12. Melanopsis costata lampra from upper Jordan Valley (Δ) and M. c. costata from Antiochia and northwards (O) (shells of at least 19.0 mm shell height). A, shelldiameter vs. shell-height. B, mouth-height vs. shell-height. C, conicality. D, rib number.

Figure 13. Melanopsis costata lampra (Δ) and M. c. costata (O): PCO scores.

Figure 14. Melanopsis costata jordanica (Δ) and M. c. lampra (O): PCO scores.

Figure 15. Ribbed Melanopsis of the Levant. A, M. saulcyi. B, M. pachya. C, M. germaini. D, M. infracincta. E, M. sp.

Figure 16. Distribution of Melanopsis saulcyi (•), M. pachya (-), M. germaini (), M. infracincta (Δ) and M. sp. (') in the Levant (inspected records).?, type locality of M. saulcyi as suggested by Pallary.

Figure 17. Melanopsis saulcyi (Homs and Jordan Valley pooled) (Δ) and M. costata (c. costata and c. lampra, pooled) (O): PCO scores.

Figure 18. Melanopsis saulcyi from the Homs region (Δ) vs. M. saulcyi from the Jordan Valley (O): PCO scores.

Figure 19. Melanopsis pachya (Δ) and M. costata lampra (O). A, rib length vs. rib number. B, PCO scores.

Figure 20. Melanopsis germaini (Δ) and M. c. costata (O). A, 2 rib width/shell-diameter vs. number of ribs. B, conicality (a vs. b); C, PCO scores.

Figure 21. Melanopsis infracincta (Δ) and M. c. costata (O). A, shell-diameter vs. shell-height. B, mouth-height vs. shell-diameter.