Corresponding author: William D. Hummon (
Academic editor: Antonio Todaro
The near eastern geographical region is almost devoid of reports of macrodasyidan gastrotrichs, the exceptions themselves being part of this study. Here, as Part 1 are described fourteen new Macrodasyida from countries of the Near East (Cyprus, Egypt and Israel, representing both the Mediterranean and the Red Seas), and a redescription of the previously described
As of December 2010 there are 305 described species of Macrodasyida, of which 142 species belong in the family
Basically, work in this geographical region began with
Littoral collections were made by whole-beach transects, with 8–10 sites spaced more or less equally from lowest to highest water levels. Locations are presented in a table (
Locations in the Mid East that are referred to in the text, along with the alphabetic symbols that are used in Figure 1.
MEDITERRANEAN SEA | |||
---|---|---|---|
CYPRUS | CB – Coral Bay | ||
EGYPT | MM - Marsa Matruh | ||
AR - Sidi Abd al-Rahman | |||
G - Green Beach | |||
H - Hannoville | |||
A - Alexandria {Betash ’Agami, Bir Mesud, Cleopatra, Mamura} | |||
ISRAEL | P - Palmachim N | ||
RED SEA | |||
EGYPT | ES - Ein Sukhna | ||
16 km south of Ein Sukhna, | |||
23km south of Ein Sukhna, | |||
WA - Wadi ’Araba | |||
U - ’Uyun Musa | |||
Ras Sudr | |||
D - Daghashland | |||
MB - Moon Beach | |||
HP - Hammam Pharoan | |||
H - Hurghada {Moon Valley, Giftun Village, Princess Village, Mugawish} | |||
SA - Sharm el-Arab | |||
SN - Sharm el-Naga | |||
S - Safaga | |||
G - Giftun Island S | |||
AR - Abu Ramada 1, 2, 3 | |||
S - Sharm el-Sheikh {Ras Mohamed National Park: Main Beach, Marsa Bareika, Tip RM, West Gate}, {Far Garden, Middle Garden, Na’ama Bay, Nabq, Ras Nasrani, Ras Qanti} | |||
TR - Tareef el-Reeh | |||
N - Nuweiba | |||
ISRAEL | E - Eilat {Princess Hotel, Snuba Dive Shop, North Beach, Coral Beach: S, M2, M3, M4, M5, N1} |
Map, with the collection locations listed in Table 1 indicated
Analytic methods are more fully given in
Drawings of holotype specimens were made using a drawing tube on the compound microscope. My finished drawings involve both dorsal and ventral views as mirror images of one another, occasionally with a composite dorsal/ventral view of juveniles and/or subadults, and always with drawings of cuticular armature, when appropriate, all with appropriate scale bars. Several of the species described herein are based on subadults, which already show characters of the adults that distinguish them from other species. In addition to the species name, for each is included a unique ‘DOS name’ based on a three-letter designation for genus and a four-letter designation for the specific epithet, giving the eight-letter ‘title’ that originated when the MS-DOS (Microsoft Disk Operating System) would only allow eight letters for a file-name.
Morphological symbols and conventions are as follows: Lt: Length, total: from anterior tip of head to posterior tip of caudum and its adhesive tubes; LPh: Length, pharynx from anterior tip of head to PhJIn; PhJIn: Junction between pharynx and intestine; WHd /WNk /WTr /WFrBs /WFrTp: Width of head, neck, trunk, furcal base and furcal tips; TbA /TbL /TbD /TbV /TbP /TbS: adhesive tubes of the anterior, lateral, dorsal, ventral, posterior (caudal), and “Seitenfüsschen” series; U: Percentage units of Lt from anterior to posterior × 100; the ‘Adult’ condition is attained when evidence of reproductive maturity is attained: testes, ovules or accessory sex organs. Columns: longitudinal in orientation; Rows: transverse in orientation; the caret ^ refers to a type locality. Latitude and longitude are given for type localities (Ordinance Survey references are given for Israeli locations); those for all other localities can be found in the Global Data Base (
Adult Lt to 772 µm; PhJIn at U28. Head small, pyriform, with a broad circumcephalic band of cilia at U02–U05, separated from the rest of the body by a long gradual neck constriction; trunk narrow, bowed outward along the mid-gut, caudum slightly flaired. Epidermis finely granular, without glands. TbA 4 per side, borne on fleshy hands; TbVL 6 per side, with 5 inserted regularly from U27 to U51 and the 6th at U72, one in the rear pharyngeal region, the others in the fore- and midgut region; TbP 10–12, inserting on the rounded caudum. Locomotor ciliature: 2 longitudinal bands, separate from one another, but uniting with cilia of the transverse cephalic band in front, and joining together behind the anus. Mouth terminal, of medium diameter; buccal cavity cylindrical; pharyngeal pores basal; intestine broadest at the rear mid-gut, narrowing behind; anus ventral at U95. Hermaphroditic; testes begin at the PhJIn, vasa deferentia run rearward, but termini not seen; a column of eggs lies along the mid-gut, developing front to rear; frontal and caudal organs not seen.
Adult Lt 615–772 µm; LPh 205–215 µm to PhJIn at U33–U28 (
There are nine video sequences of
Dolichosoma (Greek:
When mature,
Adult Lt to 517 µm (perhaps reaching 600 µm); PhJIn at U38. Head pyriform, with a broad circumcephalic band of cilia at U02–U07, separated from the rest of the body by a long gradual neck constriction; trunk medium, broadest at the mid-body, caudum slightly flaired. Epidermis finely granular, without glands. TbA 4 per side, borne on fleshy hands; TbVL 10 per side, inserted regularly from U24 to U66, three along the rear pharynx, the others along the fore- and midgut; TbV 2 per side amid the TbVL series at U44 and U49; TbP 18 of varying lengths (12 longer, 6 shorter), inserting on the rear of the rounded caudum. Locomotor ciliature: 2 longitudinal bands, separate from one another, but uniting with cilia of the transverse cephalic band in front and joining together behind the anus. Mouth terminal, of medium diameter; buccal cavity broadly cylindrical; pharyngeal pores basal; intestine broadest at the rear mid-body and narrowing behind; anus ventral at U92. Hermaphroditic; possibly protandric, testes begin at the PhJIn, vasa deferentia run rearward, but termini not seen; female system not seen, nor were frontal or caudal organs.
Adult Lt 492–517 µm; LPh 191–196 µm to PhJIn at U39–U38 (
There are four video sequences of
Saegailus (Greek:
Adult Lt 390 µm; PhJIn at U32. Head bluntly rounded, but without ocelli, neck constriction extended but slight; trunk slender, with two broad caudal lobes that incise medially to U92, without a peduncle. Glands not seen; protonephridia 3 per side, at U32, U78 and U87; longitudinal muscles are striated. TbA 3 per side (L=6, 8, 11 µm) insert in parallel, protruding obliquely to the rear; TbVL 6 per side, arising in groups of 3/2/1 (L=14, 8, 6 / 17, 7 / 14 µm) at U36–U38 /U46–U49 /U57, all protruding obliquely to the rear, proceding rearward from longer to shorter in each group; TbP 6 per caudal lobe (L=8–10 µm), longest medially on each lobe. Mouth terminal, of medium breadth; buccal cavity goblet-shaped; pharynx width follows the head/neck contours, with inconspicuous basal pores that open well behind the neck constriction; intestine narrows fore to aft, anus ventral at U91. Ventral ciliation: a unified field beneath the head splits into a pair of longitudinal bands, each narrow in breadth, that continues rearward to the level of the anus, and a second pair of longitudinal bands that lie medially from U12 to U34, with a an isolated patch lying medially behind the anus. Probably parthenogenic; male system not seen; ovaries paired in hindgut region, with oocytes on both sides behind the predominant ovum that develops medially forward toward the midgut; caudal and frontal organs not seen.
Adult Lt 322–390 µm; LPh 103–126 µm to PhJIn at U32–U33 (
I found this species in the Red Sea sites while on a Fulbright Senior Research Scholarship during 1994; thus my drawings and videos antedate by seven years its formal description by Hochberg, 2003. He had found it in eastern Australia during 2001, the species now having a much broader biogeographical range than just Austalia or the upper Red Sea. His recent report of the species from Panama (
The reason for redescribing this species is that Hochberg’s drawing (
Agadasys (Greek:
Adult 272 µm; PhJIn at U20. Body slender; head has crescent-shaped anterior, with protruding mouth and laterally directed lobes that have rounded tips, with a knob-shaped pestle organ on each side lying largely exposed beneath the rear of the head lobes; neck constriction slight, marking the pharyngeal pore openings; trunk parallel-sided, narrowing gradually in the rear, ending in a long, narrow caudal peduncle, with a bifurcate apex that indents medially to U90; pharynx short, pharyngeal pores basal; intestine narrows fore to aft; anus ventral at U68. Glands 7 per side, with another medially in the caudal peduncle; longitudinal muscles are striated. TbA 1 per side, with a duo-gland tube extending forward from a tapering base; TbL absent; TbP 3 per side, the longer one, arising from the caudal base, and the shorter two, arising from the bifurcate tip of the caudum. Mouth diameter narrow, protruding forward from anterior head curvature; small goblet-shaped buccal cavity moderately cuticularized; pharyngial pores located at the level of the neck constriction, and only detected with maturity; intestine broader in front, narrower behind, its lumen fringed by actively-beating cilia; anus ventral at U68. Locomotor ciliation forms paired longitudinal bands from the TbA rearward, joining behind the anus and continuing as a unified band onto the caudal peduncle, with a patch lying just before the bifurca. Hermaphroditic; paired testes lie along the fore-gut and paired ovaries along the rear mid-gut, eggs maturing rear to front; an ovoid frontal organ bearing active sperm occurs opposite the largest ovum on the right side.
Adult Lt 244–272 µm; LPh 48–53 µm to PhJIn at U20 (
Occasional (10–30% of samples) in frequency of occurrence, scarce to prevalent (3% to more than 30% of a sample, the latter sometimes a co-dominant
There are four video sequences of
Rubomarinus (Latin:
Adult being described Lt 544 µm; PhJIn at U43. Head stepped, narrowing toward the mouth, pestle organs in the step at U03; trunk broadest in the pharyngeal region, narrowing gradually to the long caudum. Glands inconspicuous. TbA 5–6 per side, in transverse rows that insert directly on the body; TbL/TbV 35 per side, the series beginning as TbL, then rotating to TbV and finally along the caudal base the two series duplicating one another; TbL 7 per side, with 0 along the fore and and 4 along the rear pharynx, 2 along the rear intestine, and 1 behind the anus; TbV 28 per side, uneven in size and location, but symmetrical in placement, with 1 along the rear pharynx, and the remainder in the intestinal region, the final tubes duplicating those of the rear of the TbL series; TbP 11 per side, surrounding the elongate caudum. Locomotor ciliature: a single field runs from the oral opening to the tip of the caudum, unciliated in spots surrounding ventral openings. Mouth terminal, of medium-broad width; buccal cavity lightly cuticularized, expanding with depth; pharyngeal pores para-basal; intestine narrows gradually to the rear; anus ventral at U80. Hermaphroditic; testes begin just before the PhJIn, vasa differentia join beneath the frontal organ; ova probably develop rear to front; frontal organ tubular, with cuticular nozzle, lies behind the foremost ovum; long caudal organ with muscular/glandular construction in a half to half ratio, extensively overlapping with the frontal organ.
Adult being described Lt 544 µm (others Lt 541–625); LPh 234 µm (others LPh 200–205) to PhJIn at U43 (others PhJIn at U32–U38) (
Occasional in frequency of occurrence (10–30% of samples), scarce to prevalent in abundance (3 to greater than 30% of a sample; occasionally a co-
There are ten video sequences of
Imbricatus (Latin:
Adult Lt being described 590 µm; PhJIn at U36. Head ovoid, with a narrow band of circumcephalic cilia at U02 and pestle organs at U03; trunk broad throughout, narrowing gradually along the rear intestinal region to the long caudum. Glands inconspicuous. TbA 11 per side, in arcs that insert directly on the body; TbL 4–5 per side, with 1–2 along the fore and 0 along the rear pharynx, 3 in the rear intestinal region, and 0 behind the anus; TbD 4 per side, of similar size and spacing, in the rear half of the body from U53 to U75; TbV 17 per side, nearly even in size and spacing, with 3 along the rear pharynx, the remainder in the intestinal region, and 0 behind the anus; TbP 7 per side symmetrically along the long caudum. Locomotor ciliature: a single field from the TbA to the tip of the caudum, unciliated in spots surrounding ventral openings. Mouth terminal, of narrow width; shallow buccal cavity is lightly cuticularized, expanding with depth; pharyngeal pores sub-basal; intestine broadest in front, narrowing quickly along the mid- to hind-gut, then bending around the caudal organ; anus ventral at U83. Hermaphroditic; testes begin just before the PhJIn, vasa differentia join beneath the frontal organ; ova develop rear to front; frontal organ elongate ovate, its nozzle in close contact with the ovum, with a beak-like cavity lying on the left that has a ventral pore and an accessory chamber behind; short, thick caudal organ has spiral muscles throughout, except for the rearmost glandular sac, the fore half appearing glandular, the rear half bearing an internal canal; caudal organ barely overlapping the accessory cell of the frontal organ.
Adult being described Lt 590 µm (others Lt 477–620); LPh 192 µm (others LPh 195–203) to PhJIn at U36 (others PhJIn at U33–U41) (
Occasional in frequency of occurrence (10–30% of samples), scarce to prevalent in abundance (3 to greater than 30% of a sample; often a sub-
There are seven video sequences of
Macrurus (Greek:
Adult being described Lt 524 µm; PhJIn at U31. Head stepped, narrowing toward the mouth, with a band of circumcephalic cilia at U01–U02, pestle organs in the step at U03 and black ocelli borne just behind pestle organs at U05; trunk broader in the pharyngeal than in the fore-gut region, broadest in the mid-body region, narrowing quickly in the hind-gut region to the long caudum. Glands 8 per side. TbA 7–8 per side, in arcs that insert directly on the postoral body surface at U02–U03 and project forward to obliquely outward; TbL 14, of similar size, with 3 at U55, U64 and U73, and 12 at U80 and behind (7 along the hind-gut, and 4 behind the anal aperture); TbV 14 per side, even in size, but uneven in spacing, with 1 along the rear pharynx, 1 just behind the PhJIn and the remainder in the intestinal region from U41 to U77, mostly concentrated in the rear, and 0 behind the anus; TbP 10–11 per side symmetrically along the caudum. Locomotor ciliature: a single field lies between the TbV series from the TbA back onto the caudal base, with small bare spots surrounding the ventral openings. Mouth terminal, narrow; buccal cavity shallow, lightly cuticularized, expanding with depth; pharyngeal pores sub-basal; intestine narrows gradually front to rear; anus ventral at U84. Hermaphroditic; testes begin just before the PhJIn, vasa differentia join beneath the frontal organ; ova develop rear to front, with two large developing ova and several smaller bilaterally to the rear; frontal organ Y-shaped, its nozzle in close contact with the most developed ova, with an elongate cavity having a ventral pore lying on the left; caudal organ of medium length, its fore half appearing glandular, its rear half having an internal canal and a spiral muscular covering, except for the rearmost glandular sac, the caudal organ does not overlap the frontal organ.
Adult being described Lt 524 µm (another Lt 558); LPh 161 µm (another LPh 230) to PhJIn at U31 (another to PhJIn at U41) (
Occasional in frequency of occurrence (10–30% of samples), rare to scarce in abundance (fewer than 1% to 5% of a sample);
There are seven video sequences of
Nigrocellus (Latin:
Adult being described Lt 635 µm; PhJIn at U46. Head stepped, narrowing toward the mouth, with a narrow band of circumcephalic cilia at U02 and pestle organs in the step at U03; trunk of similar breadth throughout, narrowing quickly in the hind-gut region to the medium caudum. Glands inconspicuous. TbA 7–9 per side, in arcs that insert directly on the postoral body surface at U02–U04 and project forward to obliquely outward; TbL 11, of similar size, with 0 along the fore half and 2 along the rear half of the pharynx, 9 in the intestinal region, and 0 behind the anal aperture; TbV 14 per side, similar in size, but giving the impression of linear pairs, all in the intestinal region, with 0 behind the anus; TbP 11–12 per side symmetrically around the caudum. Locomotor ciliature: paired lateral bands lie between the TbV series back to the tip of the tail, with additional sparse cilia running medially to U73. Mouth terminal, of medium width; buccal cavity lightly cuticularized, expanding with depth; pharyngeal pores sub-basal; intestine narrows gradually front to rear; anus ventral at U91. Hermaphroditic; testes begin before the PhJIn, vasa differentia join beneath the frontal organ; ova develop rear to front, with a large ovum above the fore-gut and smaller ovules to the rear; frontal organ pyriform, its nozzle, having a thick refractive cuticular cap, and lying in close contact with the developing ova, bearing sperm internally; caudal organ of medium length, its fore half appearing glandular, its rear half having an internal canal and a spiral muscular covering, except for the rearmost sac; the caudal organ barely overlaps the rear of the frontal organ, and not at all in younger specimens.
Adult being described Lt 635 µm (others 394–800); LPh 290 µm (others 159–300) to PhJIn at U46 (others to PhJIn at U29–U47) (
There are 25 video sequences of
Scleracrus (Greek: scleros + akros = meaning ‘hard, tough summit, peak’) refers to the refringent cuticular nozzle cap on the frontal organ.
Several species have frontal organs that are tipped by a small cuticular nozzle:
Trunk of adult specimen being described Lt 440 µm, Lt tail 1100 µm; PhJIn at U40 of the trunk. Head bluntly ovate, with a narrow band of circumcephalic cilia and broad pestle organs at U05; neck constriction slight; trunk broadest along the mid-gut, narrowing gradually in the hind-gut region to the elongate tail. Glands 15–16 per side. TbA 7 per side, 3 in transverse rows at U07, which project obliquely forward, and 4 in longitudinal columns at U08–U17, which project obliquely outward, all inserting directly on the postoral body surface; TbL 4 per side, similar in size, all in the intestinal region; TbVL 4 per side, similar in size, 2 along the fore and 0 along the rear pharynx, and 2 in the rear intestinal region; TbD 5 per side, similar in size, 1 along the fore and 0 along the rear pharynx, and 4 in the intestinal region; TbV absent; TbP 10 or more per side, asymmetrical along the tail, depending on its length. Locomotor ciliature: paired lateral bands run from the TbA back to the rear of the caudal organ, with more sparse cilia medially from the TbA to U49. Mouth terminal, narrow; buccal cavity lightly cuticularized, expanding with depth; pharyngeal pores sub-basal; intestine narrowing gradually front to rear, lacking an anus. Probably hermaphroditic, though testis was not seen; ova develop from rear to front, with two bilateral ova in the mid-gut region of recently mature specimens, much larger in fully mature specimens (video #1904); frontal organ not seen; caudal organ, appearing enclosed in an oval capsule, has a hyaline bulblet in the rear, with an internal canal, that cycles around to include a darkish mass on the left and a stylet on the right, which is curved in the rear, widening and straightening toward the front, where it has an asymmetrical bulb that has a symmetrical opening at the end.
Trunk of adult specimen being described Lt 440 µm (others Lt 413–480); Lt tail 1100 µm (others Lt 588–1620); LPh 174 µm (others LPh 161–173) to PhJIn at U40 (others PhJIn at U34–U40) (
Occasional in frequency of occurrence (10–30% of samples), scarce to prevalent in abundance (3% to greater than 20% of a sample, sometimes a sub- dominant
There are eight video sequences of
Toxostylis (Greek:
Adult Lt 332 µm; PhJIn at U29. Body short, robust; head end truncated, without pestle organs, tentacles or lobes; lacking any neck; trunk broadening through out the pharyngeal region, then even more along the fore-gut, before narrowing gradually to the caudal base; cirrata 6 per side of nearly similar lengths, dorsolateral at U10, U30, U45, U62, U79 and U96; caudal pedicles medium, borne on fleshy lobes, with a broad concave margin separating the two lobes, incising medially to U94. Glands 6 per side (3–8 µm diameter) in lateral columns at U15–U79. Epidermis covered with pentancres twice as long as wide, with the center tine 20% longer than the others, though ancres are smaller fore and aft; ancres occur in 55–60 rows of 15–17 ancres each, extending onto the rear of the oral hood and onto the caudal lobes. TbA 7 per side forming 3 rows of 2, 3 and 2 tubes, all projecting obliquely forward, tubes inserting directly on the postoral body surface at U05–U07; TbVL 18 per side, 1 at U08, a group of 11 at U25–U71, 1 at U80, and a group of 5 at U86–U95; TbV 3 per side, 1 at U65 and 2 at U72; TbDL 4 per side (L 17–19 µm) at at U13, U47, U64 and U80; TbL/D
Adult Lt 280–332 µm; LPh 85–80 µm to PhJIn at U30–U29 (
Sparse in frequency of occurrence (fewer than 10% of samples), rare to scarce in abundance (fewer than 1% to 5% of a sample);
There are three sequences of
The species is named with reference to the coralline-calcareous sediment in which it was always found.
Adult Lt 344 µm; PhJIn at U30. Head end bluntly rounded, bearing club-shaped, ear-like pestle organs at U04; body narrows along the hind-pharynx; trunk broadens along the mid-gut, narrowing gently to the caudal base; caudal pedicles medium, with a sharply concave margin, indenting medially to U94. Glands 37 per side in medial and lateral columns. Epidermis covered with tetrancres of similar size, but smaller fore and aft; ancres occur in 36–38 rows of 13–15 ancres each, excluding the oral hood, but covering the caudal lobes. TbA 5 per side forming a transverse arc, tubes inserting directly on the postoral body surface; TbVL 12 per side, 1 along the fore pharynx, behind the TbA, 4 along the fore-gut, and 7 closely spaced from anus onto the caudal lobes; TbV 3 per side in a transverse row at U87; TbDL 2 per side at U51 (L 26 µm) and U90 (L 17 µm); TbP 3 per side on the caudal pedicles, forming the fused ‘two fingers and a thumb’ typical of the family, without supplemental cirrata-like structure projecting from between the ‘fingers’ or additional tubes in the space between the peduncles. Locomotor ciliature: a single field covers the entire ventral surface from TbA to anus, with a narrower tract continuing beneath the caudum. Mouth subterminal, as broad as the fore end of the body, buccal cavity lightly cuticularized; pharynx narrows to inconspicuous basal pharyngeal pores; intestine narrows fore to aft, anus ventral at U87. Hermaphroditic; testis on left side as seen from below; vas deferens opens near the anus; developing ovum occurs above the midgut, with oocytes bilaterally to the rear; caudal organ small, ovoid and thick-walled, with an interior of refractive material and a central canal; frontal organ broadly oval and hyaline, bearing active sperm.
Adult Lt 271–344 µm; LPh 102 µm to PhJIn at U39–U30 (
Occasional in frequency of occurrence (10–30% of samples), scarce to prevalent in abundance (3% to greater than 30% of a sample, occasionally a co-dominant
There are eight video sequences of
Rhopalotum (Greek:
Adult Lt 423 µm; PhJIn at U34. Body of medium length, narrow; head end rounded, without pestle organs, tentacles or lobes; neck indistinct along the rear pharynx, trunk broadening slightly in the mid-body, before narrowing gently to the caudal base; caudal pedicles short, borne on short lobes, with a wide concave margin separating the two lobes, incising medially to U97. Glands 8 per side, small, scattered in lateral columns. Epidermis covered with tetrancres of similar size, slightly smaller fore and aft; ancres occur in 45–50 rows of 9–10 ancres each, excluding the entire oral hood, but extending onto the base of the caudal lobes. TbA 7 per side forming arcs radiating from forward to outward, tubes inserting directly on the postoral body surface at U03–U05; TbVL 43 per side, 1 at U06, a group of 36 at U25–U80, 1 at U89, and a group of 5 at U93–U97; TbV 5 per side in a transverse row at U85; TbDL 9 per side, of two sizes, 7 (short) subequally spaced at U10–U90, and 2 (long) at U86 and U93; TbL/D
Adult Lt 423 µm (others Lt 241–410); LPh 146 µm (others LPh 95–141) to PhJIn at U34 (others PhJIn at U39–U34) (
Common in frequency of occurrence (30–60% of samples), scarce to prevalent in abundance (3% to greater than 30% of a sample, occasionally a co-
There are 12 video sequences of
Sinaiensis (pronounced ‘sinaiënsis’ with a dieresis over the ‘e’ to indicate that it is to be pronounced separately from the diphthong ‘ai’) is named after the geographical region, the Sinai Peninsula, in which it was first found.
Adult being described Lt 246 µm; PhJIn at U35. Body short, robust; head end rounded, without pestle organs, tentacles or lobes; neck a slight narrowing at the PhJIn; trunk broadening in the mid-gut, before narrowing gently then quickly to the caudal base; cirrata dorsolateral, 3 per side; caudal pedicles medium, naked, with a narrow concave margin separating the pedicles, incising medially to U92. Glands 9 per side scattered in lateral columns at U24–U85. A strange sub-cylindrical finger-like structure occurs laterally at U34. Epidermis covered with curved pentancres three times as long as wide, slightly smaller fore and aft; ancres occur in 46 rows of 14–15 ancres each, on dorsal and lateral surfaces, extending onto the middle of the oral hood and onto the caudal base. TbA 4 per side, 1 medially and 3 laterally, all projecting forward or obliquely outward, tubes inserting directly on the postoral body surface at U09–U10; TbVL 11 per side, 1 along the fore half of the pharynx at U14, 7 along the intestine at U39–U75, and 3 at and behind the anus at U88–U92; TbV 3 per side in a transverse row at U80; TbL
Adult being described Lt 246 µm (others 197–322); LPh 87 µm (others 79–128) to PhJIn at U35 (others to PhJIn at U40) (
Sparse in frequency of occurrence (fewer than 10% of samples), scarce to numerous in abundance (3% to 20% of a sample, sometimes a sub-dominant
There are three sequences of
Xenodactylum (Greek:
Adult Lt 657 µm; PhJIn at U23. Body elongate, slender; mouth a narrow outwardly rolled protrusion, head with a band of circumcephalic cilia at U03 and prominent pestle organs at U04, but lacking lateral lobes; neck constriction lacking, body sides parallel over most of their length, thinning gradually to the caudal base; caudum is slightly cleft, incised from its tips to U97; medial cone usually absent. Glands inconspicuous, ca. 28 per side. TbA 8 per side, the shortest one inserting on the medial edge, occur on fleshy hands that insert at U11; TbL absent; TbD 7 per side at U27–U84 and TbV 14 per side at U29–U88, all in the intestinal region, of similar size and spacing; ‘dohrni’ [Seitenfüsschen] tubes 2 per side, posteriolaterally directed (L longer tube =20 µm, shorter =15 µm), inserting ventrolaterally just behind the fleshy hands at U12; TbP 8 per side, the outermost being the longest and thickest, the others being shorter, with none occurring on the lateral or leading edges of the lobes. Locomotor ciliature: 2 longitudinal bands run from the pestle organs back and join behind the level of the anus. Mouth terminal, breadth narrow; buccal cavity small, deep, vaseshaped; walls of medium cuticularization; basal pharyngeal pores large and conspicuous; intestine narrows gradually front to rear; anus is ventral at U93. Hermaphroditic, protandrous to simultaneous; paired testes extend rearward from just behind the PhJIn, their vasa deferentia recurving to the fore and exiting at about U31; small developing ova occur bilaterally in the mid-gut region; frontal and caudal organs not seen.
Adult Lt 635–657 µm; L to PhJIn 155–163 µm at U26–U23 (
Sparse (less than 10% of samples) in frequency of occurrence, rare to scarce (fewer than 1 to 3–5% of a sample) in abundance;
There are eight video sequences of
Levantia is named after the eastern Mediterranean region in which it was first found.
Adult Lt 440–486 µm; PhJIn at U33. Body medium-short, slender; head sculptured, with a narrow band of circumcephalic cilia and shallow lateral lobes at U05, but no tentacles; neck constriction greatest along the mid-pharynx; trunk broadest along the mid-body, thining gradually to the caudal base; caudum is slightly cleft, incised from its tips to U98, bearing a medial cone. Glands 36–37 per side, small, inconspicuous. TbA 7 per side, inner-most shorter than the others, on fleshy hands that insert at U11; TbVL 9 per side, with 1 along the fore half and 0 in the rear half of the pharynx, and the other 8 evenly spaced and symmetrically arranged along the intestine, and none behind the anal opening; ‘cirrata’ [Seitenfüsschen] tubes are short, but present at U35; TbP 7 per side, lengthening medial to lateral, inserting along the trailing edge of each lobe. Locomotor ciliature forms 2 longitudinal bands from buccal cavity to anus. Mouth terminal, of medium width; buccal cavity shallow, mugshaped; walls lightly cuticularized; pharynx broad, with conspicuous basal pharyngeal pores; intestine narrows fore to aft, with a slight bulge at the ventral anus, U92. Hermaphroditic; paired testes extend back from the PhJIn, its vasa deferentia recurving to the fore and probably exiting just behind the PhJIn, sperm sometimes descending in clusters to the base of the vasa deferentia; the bilateral ova occur in the fore-gut region and develop rear to fore; frontal and caudal organs not seen.
Adult Lt 440–486 µm; L to PhJIn 162–166 µm at U38–U33 (
Sparse in frequency of occurrence (fewer than 10% of samples), scarce to prevalent in abundance (less than 3–5% to greater than 30% of a sample, sometimes a dominant
There are five video sequences of
Erythrothalassia (Greek:
I am greatly indebted to colleagues who facilitated my collections for this study: These were part of the fauna that were encountered in Egypt and Cyprus during an eight month Fulbright Research Fellowship from April through November 1994 with the Department of Oceanography in the University of Alexandria, Dr. Sharif el Din, sponsor and Dr. Heshem Mustafa, host. The work in the South Sinai twice was facilitated by Dr. Michael Pearson, Park Manager, and the Ras Mohammed National Park staff. Three trips were made to Israel, one in February 1992, a second in February 1999, and the third for the month of September 1999 was based at the Department of Zoology at Hebrew University, Jerusalem, Dr. Francis Por, sponsor and Dr. Nachama Ben Eliahu, host. The work in Eilat twice was facilitated by the Steinitz Marine Biological Laboratory staff. The Fulbright Committee in Cairo, Egypt was of inestimable value. I am indebted to two reviewers, Dr. Antonio Todaro and one anonymous reviewer, for helpful suggestions in revising the manuscript, the former important in his challenge of some of my assumptions.