Tracheloraphis

Tracheloraphis Dragesco, 1960 (ref. ID; 3690 original paper)

Ciliophora: Kinetofragminophora: Order Karyorelictida: Family Trachelocercidae (ref. ID; 7323)

Synonym Trachelonema Dragesco, 1960 (ref. ID; 2267, 7362)

[ref. ID; 2267]
Tracheloraphis differs from Trachelocerca mainly by the distinct brosse. Furthermore, the glabrous zone is usually considerably wider and the body more distinctly flattened. The brosse of Trachelolophos is unstructured and near the centre of the oral cavity, i.e. within the area bordered by the circumoral kinety, which is not interupted. The second character contained in Foissner et al. diagnosis, viz. the structure of the bristle kinety, is somewhat ambiguous because the minute kineties are inconspicuous in one (T. aragoi) of the four species investigated. Thus, this character should possibly be removed from the diagnosis. However, Foissner et al. prefer it as an additional diagnostic, at least at the present state of knowledge, because it is very conspicuous when compared with the single row found in Trachelocerca, Trachelolophos and Prototrachelocerca. Trachelonema differs from Tracheloraphis in that it is flattened leaf-like and the glabrous stripe extends the whole body width, causing the ciliature to be restricted to the right surface. Foissner rediscovered the type species, T. longicolle, at the locus classicus and can confirm Dragesco's observations. This species is indeed very flat and ciliated only on the right side. However, the somatic and oral infraciliature of T. longicolle is very similar to that of Tracheloraphis phoenicopterus and T. aragoi. Furthermore, there are distinct transitions in body flattening and width of the glabrous stripe, as evident from a comparison of T. aragoi, T. phoenicopterus and T. longicollis. Other species, e.g., Tracheloraphis grisea, are also very flat and have a broad glabrous stripe. Obviously, there is no distinct border between Tracheloraphis and Trachelonema, suggesting synonymy. Both genera were established in the same paper. Foissner suggest preserving Tracheloraphis because it contains more species and has page priority. The following nominal species of Trachelonema are transferred to Tracheloraphis. (ref. ID; 2267)
Etymology; Composite of the Greek nouns trachelos (neck) and raphis (needle). Raphis has feminine gender. Unfortunately, many nominal Tracheloraphis species were supplied with a masculine termination, obviously because most describers assumed a masculine gender of raphis from the termination us in phoenicopterus, the type species. However, phoenicopterus (Flamingo), originally written Phoenicopterus, is a noun in apposition, which retains the same ending, whatever the gender of the generic name with which it is combined (article 31b (ii) of the ICZM 1985). This requires many emendations, undertaken in the last chapter of the results. (ref. ID; 2267)

[ref. ID; 7323]
The main feature separating the genera Tracheloraphis and Trachelocerca is the presence or absence of the non-ciliated glabrous zone. Therefore, every specimen with an extremely narrow glabrous zone (in which case it is a Tracheloraphis) is under suspicion as to whether it indeed has one or it is simply a narrow opening between two neighboring kineties caused by fixation (in which case it is a Trachelocerca). Within the genus, the existing identification keys (Borror 1973; Carey 1992; Uhlig 1964) emphasize several characteristics. These are: general morphological features (organism shape and length; clear versus poor differentiation of the body into regions; rounded versus pointed posterior end; details of cytostome structure); somatic ciliature (number of longitudinal kineties); width of glabrous zone; organization of the nuclear apparatus (single versus multiple groups, and the number of macro- and micronuclei). (ref. ID; 7323)

Tracheloraphis aragoi (Dragesco, 1954) Dragesco, 1960 (ref. ID; 2267 redescribed paper, 3690 original paper)
Tracheloraphis binucleata Dragesco, 1960 (ref. ID; 2859)
Tracheloraphis caudatus Dragesco & Raikov (ref. ID; 4858, 4862) reported author and year? (ref. ID; 4743, 4865, 4869)
Tracheloraphis coluber (ref. ID; 4743)
Tracheloraphis conformis Wright, 1982 (ref. ID; 2267, 7323)
See; Trachelocerca sagitta (ref. ID; 2267)
Tracheloraphis crassus Raikov (ref. ID; 2117, 4862) reported author and year? (ref. ID; 4865)
Tracheloraphis discolor Raikov, 1962 (ref. ID; 3119) reported year? (ref. ID; 2117)
Tracheloraphis ditis Wright, 1982 (ref. ID; 2267, 7323)
Tracheloraphis dogieli Raikov, 1957 (ref. ID; 4720) reported year? (ref. ID; 2117) reported author and year? (ref. ID; 4865, 4869)
Tracheloraphis drachi Dragesco, 1960 (ref. ID; 3690 original paper) reported year? (ref. ID; 2117)
Tracheloraphis drachi f. bimicromultinucleata Raikov (ref. ID; 2117)
Tracheloraphis enigmatica Dragesco, 1960 (ref. ID; 2267)
Tracheloraphis enigmaticus Dragesco, 1960 (ref. ID; 3690 original paper)
Tracheloraphis fasciolatus (Sauerbrey, 1928) (ref. ID; 3690)
Syn; Trachelocerca fasciolatus Sauerbrey, 1928 (ref. ID; 3690)
Tracheloraphis flexuosus Raikov & Kovaleva, 1968 (ref. ID; 4880)
Tracheloraphis gracilis Dragesco, 1960 (ref. ID; 2267, 3690 original paper)
Tracheloraphis grassei (Dragesco, 1960) Foissner & Dragesco, 1996 (ref. ID; 2859)
Tracheloraphis griseus (Kahl, 1935) Dragesco, 1960 (ref. ID; 3690 redescribed paper)
Syn; Trachelocerca griseus Kahl, 1935 (ref. ID; 3690
Tracheloraphis incaudatus (Kahl) (ref. ID; 2117, 3690)
Syn; Trachelocerca incaudatus Kahl, 1933 (ref. ID; 3690)
Tracheloraphis kahli Raikov, 1962 (ref. ID; 3119) reported year? (ref. ID; 2117)
Tracheloraphis lacteus Raikov & Kovaleva, 1968 (ref. ID; 7323)
Tracheloraphis longicollis Dragesco, 1960 (ref. ID; 2267 redescribed paper)
Tracheloraphis margaritata (Kahl, 1930) Dragesco, 1963 (ref. ID; 2859)
Tracheloraphis margaritatus (Kahl) (ref. ID; 1879, 2117)
Tracheloraphis monocaryon Dragesco, 1965 (ref. ID; 2267), monokaryon Dragesco, 1965 (ref. ID; 7323)
Tracheloraphis nivea Wright, 1982 (ref. ID; 2267)
Tracheloraphis niveus Wright, 1982 (ref. ID; 7323)
Tracheloraphis oligostriata (Raikov, 1962) (ref. ID; 2267 redescribed paper)
Tracheloraphis phoenicopterus (Cohn, 1866) (ref. ID; 3119) reported year? (ref. ID; 4858, 4862, 7517) or (Cohn, 1866) Dragesco, 1960 (ref. ID; 2117, 2267 redescribed paper, 2316, 3690 redescribed paper) reported author and year? (ref. ID; 4865, 4869)
Syn; Trachelocerca phoenicopterus Cohn, 1866 (ref. ID; 2316, 3690)
Tracheloraphis poljanskyi (Raikov, 1963) Foissner, 1997 (ref. ID; 2859)
Tracheloraphis prenanti Dragesco, 1960 (ref. ID; 2117) reported year? (ref. ID; 4858)
Tracheloraphis prenanti f. multicineta Raikov & Kovaleva, 1968 (ref. ID; 3119)
Tracheloraphis prenanti var. oligocineta Rainkov & Kovaleva, 1968 (ref. ID; 2316)
Tracheloraphis primitarum Epstein, 1994 (ref. ID; 2267, 7323 original paper)
Tracheloraphis remanei Dragesco, 1960 (ref. ID; 3690 original paper) reported year? (ref. ID; 4858)
Tracheloraphis stephani Dragesco, 1965 (ref. ID; 2267)
Tracheloraphis striatus Raikov, 1962 (ref. ID; 2117)
See; Trachelocerca sagitta (ref. ID; 2267)
Tracheloraphis sulcata (ref. ID; 4743)
Tracheloraphis swedmarki(i) Dragesco, 1960 (ref. ID; 1879, 2117, 2267)
Tracheloraphis teissieri Dragesco, 1960 (ref. ID; 3690 original paper) reported year? (ref. ID; 2117)
Tracheloraphis totevi Kovaleva & Golemansky, 1979 (ref. ID; 2117 original paper, 4743) reported year? (ref. ID; 4761, 4858, 4862) reported author and year? (ref. ID; 4865, 4869)
Tracheloraphis vermiformis Raikov (ref. ID; 2117)

Tracheloraphis aragoi (Dragesco, 1954) Dragesco, 1960 (ref. ID; 2267 redescribed paper, 3690 original paper)

Descriptions

Trunk distinctly, that is 2-3:1 flattened. Greyish in dissecting and bright-field microscope, glabrous stripe appears as light, narrow band. Fully extended specimens filiform with anterior and posterior third gradually tapering, neck and tail thus indistinctly separate from trunk. Glabrous stripe narrow, corresponds to an area occupied by 1-2 kineties, distinctly indented, not tuberculate, underlain by conspicuous, yellowish, discoidal structures, possibly mitochondria. Head claviform to calciform and thus distinctly set off from neck, bright because not filled with ellipsoidal inclusions; oral bulge inconspicuous, distinctly set off from head, about 5 um high, surface flat; brosse cleft distinct. Distal end of tail pointed and curved. Fully contracted live specimens with margins of glabrous stripe distinctly folded and protruding; fixed specimens, however, not banana-shaped but elongate, curved, or sigmoidal. Nuclei in centre of trunk, from distinct, ellipsoid cluster; macronuclei globular to ellipsoid, contain many small nucleoli, some chromatin patches and, in Sete specimens, usually a colourless, oblong protein crystal which does not stain with protargol; micronuclei globular, 2-2.6 um in protargol impregnated, 3.5-5.2 um in methyl green-pyronin stained specimens, scattered within macronuclear group. No contractile vacuole. Cortex very flexible, with spiny projections between ciliary rows, rather distinctly set off from granular endoplasm. Two types of yellowish cortical granules: type 1 ellipsoid, 1.5-2.2x1-1.5 um, forms conspicuous, widely spaced clusters protruding above cell surface between ciliary rows, highly refractile, stains red with methyl green-pyronin, impregnates faintly to intensely with protargol, depending on bleaching conditions, granule clusters in Sete specimens usually so large that somatic kinetids become bulged; type 2 about 0.2-0.5 um across, scattered between ciliary rows, does not stain with methyl green-pyronin and protargol. Cytoplasm conspicuously vacuolated, colourless, packed with brightly shining fat globules and innumerable, 2-4 um sized ellipsoidal (crystalline?) inclusions, becoming inflated and blueish when stained with methyl green-pyronin. Food vacuoles with greenish and brownish content, possibly from ingested algae; a specimen from Roscoff contained a huge (12- x 50 um) vacuole with a decaying ciliate, possibly a Condylostoma. Glides and winds elegantly between sand grains and organic debris. (ref. ID; 2267)

Infraciliature: The somatic and oral infraciliature of T. aragoi is very similar to that of T. phoenicopterus. Most important differences concern morphometric characteristics and features recognizable only in live specimens, as described above. Thus, Foissner refrain from a complete description of the infraciliature and refer to the description of T. phoenicopterus, the detailed figures and figure explanations. Nevertheless, a few features are different and will be thus described in detail. The right somatic ciliation of T. aragoi is much more differentiated than that of T. phoenicopterus. Each kinety commences with a few condensed dikinetids having only the anterior basal body ciliated. The following head and neck dikinetids have both basal bodies of the dikinetids ciliated. All trunk kinetids have only the anterior basal body ciliated. The tail dikinetids have both basal bodies ciliated, except in the distalmost region, where only the anterior basal bodies bear a cilium. The bristle kinety of T. aragoi consists of a single row of loosely and rather irregularly arranged dikinetids, unlike in T. phoenicopterus, T. longicollis and T. oligostriata, which have oblique kineties in the trunk region. Oblique, kinety-like structures are also found in the bristle kinety of T. aragoi, albeit rarely and irregularly scattered. However, only the granules (dikinetids) neighbouring the somatic kineties are ciliated; thus the other granules, which are often slightly smaller and unpaired, are very likely a special sort of cortical granules (extrusomes?). This is supported by the observation that such granules are also scattered between the bristle kinetids in regions where the kinetids form a single line. Nematodesmata-like fibres originated from the anterior arch of the bristle kinety were seen also in this species. The contractile system of T. aragoi is either weakly developed, as indicated by the comparatively weak contractility of the species, or of different chemical composition because myonemes impregnated only very rarely and faintly. (ref. ID; 2267)

Measurements

Extended specimens in vivo about 1,100-2,300 x 50-60 um, usually 1,500-2,000 um long, very flexible but less contractile than many other trachelocericids, size and shape thus comparatively well preserved in protargol slides. (ref. ID; 2267)

Tracheloraphis binucleata Dragesco, 1960 (ref. ID; 2859)

Descriptions

This species has only 2 macronuclear nodules and is flask-shaped and only 200 um long. (ref. ID; 2859)

Tracheloraphis ditis Wright, 1982 (ref. ID; 2267, 7323)

Descriptions

This ciliate is colourless with a slightly swollen apical region and a rounded posterior end. The cytostome is simple, without a slit and is occasionally obscured by inclusions. There are between eighteen and twenty-two kineties; the globerulus zone very narrow and occupying the equivalent of one kinety. The interkinetic spaces and globerulus zone are occupied by plications which have small mucocysts on their surface. The globerulus zone had, on average, eight kineties that terminated against it. There is a loosely associated group of nuclei located centrally. The macronuclei vary in number between four and size, more usually four. They vary in size between 6 and 8 um. They have an irregular outline, contain a large amount of chromatin and may contain one or two small nucleoli. There are two micronuclei, although on one occasion only a single micronucleus was observed. They are large, measuring 5 um across the longest axis and are ovoid in outline. (ref. ID; 2267)

Measurements

Length between 300 and 800 um, average 450 um. (ref. ID; 2267)

Tracheloraphis flexuosus Raikov & Kovaleva, 1968 (ref. ID; 4880)

Type locality

From the Posjet Gulf of the Japan Sea. (ref. ID; 4880)

Tracheloraphis grassei (Dragesco, 1960) Foissner & Dragesco, 1996 (ref. ID; 2859)

Descriptions

It has large brown granules only on the left side. Furthermore, it has a distinct tail and only about 14 ciliary rows. (ref. ID; 2859)

Tracheloraphis longicollis Dragesco, 1960 (ref. ID; 2267 redescription paper)

Descriptions

Filiform and flattened ribbon-like, neck and tail indistinctly separate from trunk, head claviform and dark, distal end of tail curved. 4 macronuclei and 2 micronuclei forming tight cluster in centre of trunk. 11-13 somatic ciliary rows; glabrous stripe as wide as body, left side thus barren. Bristle kinety at margins of left side, of usual structure in anterior and posterior third of cell, composed of many minute kineties each comprising 2-4 dikinetids in trunk region. 2 oblique brosse kineties. Cortical granules inconspicuous, about 1 um across, colourless, loosely spaced. (ref. ID; 2267)

Somatic infraciliature: Tracheloraphis longicollis has only the right surface ciliated, the left is barren, i.e. occupied by the glabrous stripe, at the margins of which, the bristle kinety extends. The cilia are arranged in longitudinal rows which are distinctly separate from the circumoral kinety and extend between flat cortical crests. The anterior end of the ciliary rows has condensed, i.e. more narrowly spaced dikinetids and is slightly curved to the right. Usually, the condensation is inconspicuous or even lacking in some kineties. One to four ciliary rows are gradually shortened in the neck region left of the glabrous stripe and posteriorly, where the body narrows to the tail, on both sides of the stripe. In other words, and anterior and posterior secant system are formed at the margins of the cell where some kineties abut to the bristle kinety. Thus, the head, neck and tail have about one quarter fewer kineties than the trunk. The ciliary rows neighbouring the right branch of the bristle kinety are unshortened anteriorly and thus run alongside the glabrous stripe. The distance between the ciliary rows decrease slightly from right to left, i.e. those forming the anterior secant system are more narrowly spaced than those right of the glabrous stripe. The entire infraciliature consists of dikinetids whose ciliation and fibrillar system highly resemble those of T. phoenicopterus, with, however, some differences. First, oralized somatic dikinetids are very likely lacking, at least were not recognizable despite the high quality of the preparation. Second, the ciliation of T. longicollis is extremely variable. In some specimens most dikinetids have both basal bodies ciliated, while in others only the anterior basal bodies bear a cilium. Usually, the condensed dikinetids at the anterior end of the ciliary rows and the kineties neighbouring the glabrous stripe have barren posterior basal bodies. The contractile apparatus of T. longicollis consists of a myoneme close to the left of each kinety. As in the other species investigated, the distinctiveness of the myonemes varies highly, depending on preparation condition. The myonemes are flattened ribbon-like and extend the whole length of the kinety, but are wider (thicker, i.e. about 0.6 um) in the trunk than in the head and tail region (<0.2 um). No myonemes impregnated in the glabrous stripe. The glabrous stripe extends along the whole body length and width, except on the neck and head, where it gradually narrows, occupying only about one third of the head's width. The glabrous stripe is bordered by the bristle kinety which is very similar to that of T. phoenicopterus, especially in having small, oblique kineties in the trunk region. Tracheloraphis longicollis possibly lacks the nematodesmata-like fibres associated with the dikinetids forming the anterior arch of the bristle kinety in T. phoenicopterus. (ref. ID; 2267)

Oral infraciliature: The oral infraciliature of T. longicollis consists of a circumoral kinety and a small brosse difficult to recognize in living specimens. The circumoral kinety extends in the flat furrow separating the oral bulge from the head and is composed of a single row of vertically oriented dikinetids having only the posterior basal body ciliated. Each circumoral dikenetid is associated with a distinct nematodesma obliquely extending into the head. The nematodesmata of neighbouring dikinetids unite to conspicuous bundles, forming a cone-shaped oral basket. As in the other species investigated, the circumoral kinety of T. longicollis is very likely composed of several fragments, as indicated small gaps, 1-2 dikinetids wide, and the bundled arrangement of the nematodesmata. The circumoral kinety is interrupted where the brosse kineties are inserted. Its left end simply abuts to the margin of the brosse cleft, i.e. to the left lip of the oral bulge. The right end is more complicated. It extends along the oblique margin of the brosse cleft and curves back at the cleft vertex in such a steep angle that a loop-like structure, or a small oblique segment, is formed paralleling the descending portion of the circumoral kinety and the anterior end of the neighbouring somatic kinety. The brosse is located in flat cavity, the brosse pocket, just above the arch of the bristle kinety, and intersects the circumoral kinety. The brosse invariably consists of 2 oblique kineties, arranged in parallel and one behind the other, composed of closely spaced dikinetids having only the posterior basal body ciliated. Brosse kinety 1 is invariably smaller than kinety 2. (ref. ID; 2267)

Measurements

Fully extended cells in vivo 600-900 um long. (ref. ID; 2267)

Tracheloraphis margaritata (Kahl, 1930) Dragesco, 1963 (ref. ID; 2859)

Descriptions

It is only 350 um long when extended, has a narrow glabrous stripe, and possesses only 5 macronuclear nodules forming a short strand. (ref. ID; 2859)

Tracheloraphis oligostriata (Raikov, 1962) (ref. ID; 2267 redescribed paper)

Descriptions

Trunk distinctly, that is about 3:1 flattened, cells thus ribbon-like. Greyish and rather transparent in dissecting and bright-field microscope. Fully extended specimens filiform with anterior and posterior region gradually tapering, neck and tail thus indistinctly set off from trunk. Glabrous stripe about as wide as body in trunk region, slightly convex, without groove in midline. Head calciform and thus distinctly set off from neck, bright because studded with ellipsoid inclusions; oral bulge indistinctly set off from head, surface flat, contains 1.5 x 1 um sized granules, possibly extrusomes; brosse cleft distinct. Distal end of tail pointed and curved. Fully contracted specimens about 150-300 um long, banana-shaped with many transverse and oblique folds, left side distinctly protruding and tuberculate. Nuclei form distinct strand in trunk, number and arrangement highly variable, frequently 2-3 macronuclei with 1-2 micronuclei in between unite to a small cluster; macronuclei and micronuclei globular to slightly ellipsoid, macronuclei with one, rarely two large nucleoli and some inconspicuous chromatin condensations. No contractile vacuole. Cortex very flexible, in contracted specimens conspicuously tuberculate, indistinctly set off from granular endoplasm, contains numerous small, colourless granules. Cytoplasm with some food vacuoles containing unidentifiable debris and many 2-4 um sized ellipsoid (crystalline?) inclusions, which become inflated and reddish, respectively, blue after prolonged supravital action of methyl green-pyronin and cresyl blue. (ref. ID; 2267)

Infraciliature: The somatic and oral infraciliature of T. oligostriata is very similar to that of T. longicollis. Most important differences concern morphometric characteristics. Thus, Foissner et al. refrain from a complete description of the infraciliature and refer to the description of T. longicollis, the detailed figures figure explanations. Nevertheless, a few features are different or were seen clearly only in this species, and are thus described in some detail. The somatic ciliation of T. oligostriata is as variable as that of T. longicollis, i.e. some specimens have both basal bodies of the dikinetids ciliated in the main portion of the cell, while others mainly have only the anterior basal bodies ciliated. The two rightmost kineties have condensed, i.e. more narrowly spaced dikinetids at the tail end, highly reminiscent of the condensation found in loxodids; however, the condensed dikinetids of T. oligostriata are not associated with special fibres, as in loxodids, and could thus simply be caused by a strong or special mode of contraction of the tail during fixation. The bristle kinety is structured as described in T. longicollis, i.e. consists of short, oblique kineties in the trunk region, each composed of ciliated and unciliated argyrophilic granules; usually ciliated and unciliated granules alternate within a row, the latter being slightly smaller and often unpaired, but sometimes they are distinctly paired or triplicate. Furthermore, a special fibrillar system, highly reminiscent of that described in Trachelocerca ditis, was recognizable in a few excellently prepared specimens. It consists of a comparatively thick fibre extending from the ciliated basal body to the somatic kineties and of a very fine fibre extending from each basal body of a pair into the glabrous stripe. The unpaired granules or paired granules without a cilium lack these fibrillar associates, indicating that they are not kinetids but a special type of cortical granule, possibly extrusomes. The brosse of T. oligostriata invariably consists of a single kinety slightly obliquely implanted at the right wall of the brosse pocket. (ref. ID; 2267)

Measurements

Extended specimens in vivo about 300-500 um long, highly flexible and contractile, size and shape thus poorly preserved in protargol slides. (ref. ID; 2267)

Tracheloraphis phoenicopterus (Cohn, 1866) (ref. ID; 3119) reported year? (ref. ID; 4858, 4862, 7517) or (Cohn, 1866) Dragesco, 1960 (ref. ID; 2117, 2267 redescribed paper, 2316, 3690 redescribed paper) reported author and year? (ref. ID; 4865, 4869)

Synonym

Trachelocerca phoenicopterus Cohn, 1866 (ref. ID; 2316, 3690)

Redescription

Trunk distinctly, i.e. 2-3:1 flattened. Greyish and opaque in dissecting and bright-field microscope. Fully extended specimens filiform with anterior and posterior third gradually tapering, neck and tail thus indistinctly separate from trunk. Head slenderly claviform, dark to black at low magnification (< /_ x100) due to many about 4x2 um sized, refractile (crystalline?) inclusions; oral bulge inconspicuous but easy to recognize because distinctly set off from head, about 3 um high, surface slightly depressed in centre, contains, like the head, ellipsoid inclusions; brosse cleft difficult to recognize. Distal end of tail pointed and distal curved. Fully contracted specimens banana-shaped in vivo, convex side with glabrous stripe distinctly protruding and tuberculate; partially contracted cells sometimes spiral and with glabrous stripe indistinctly tuberculate. Nuclear apparatus (capsule) in centre of trunk, surrounded by voluminous layer of pale, 2-3 um sized granules faintly stained with protargol; nuclei form tight, distinctly ellipsoid cluster, possibly a capsule, their number thus difficult to recognize, 6-12 macronuclei and 2 micronuclei are common. Macronuclei 6-8 um in vivo, with small and medium-sized nucleoli, two of them contain a cubiform protein crystal which does not stain with protargol. Micronuclei about 4 um in vivo, in centre of macronuclear cluster. See Raikov & Kovaleva (1978) for a detailed transmission electron microscope account of the nuclear apparatus of T. phoenicopterus. No contractile vacuole. Cortex very flexible, about 1 um thick, forms tubercles in contracted specimens. Cortical granules ellipsoid to fusiform, minute (about 1.2x0.6 um), yellowish, form narrow stripes between ciliary rows and rather dense layer in cortex of glabrous stripe. Glides and winds elegantly between sand grains and organic debris. (ref. ID; 2267)

Somatic infraciliature: The surface of T. phoenicopterus is densely ciliated, leaving blank a rather wide zone, the glabrous stripe, extending the whole body length in the midline of the left side. The cilia, which are rather stiff and can be spread, are about 10 um long and arranged in longitudinal rows which are distinctly separate from the circumoral ciliature and extend between flat cortical crests. The anterior end of ciliary rows has condensed, i.e. more narrowly spaced dikinetids and is curved to the right. Usually, the condensation is inconspicuous or even lacking in some kineties; rarely, it is absent in most kineties. The ciliary rows are gradually shortened anteriorly in the neck region left of the glabrous stripe and posteriorly, where the body narrows to the tail, on both sides of the stripe. In other words, an anterior and posterior secant system are formed on the left surface of the neck and tail where some kineties abut to the bristle kinety. Thus, the head, neck, and tail have about one third less kineties than the trunk. The ciliary rows neighbouring the right branch of the bristle kinety are unshortened anteriorly and thus extend alongside the glabrous stripe. The distance between the ciliary rows decrease slightly from right to left, i.e. those forming the anterior secant system are more narrowly spaced than those right to the glabrous stripe. The entire infraciliature consists of dikinetids which have, however, a highly specialized ciliation and fibrillar system. The dikinetids are rotated about 20-30 degrees counter-clockwise to the kinety axis and have both basal bodies ciliated, except the condensed kinetids at the anterior and the widely spaced kinetids at the posterior end of the somatic kineties, where only the anterior basal bodies are ciliated. Likewise, the kinetids at the ends of the secant kineties lack the posterior cilium. The dikinetids are associated with various distinct fibres, all very likely originating from the posterior basal bodies. Foissner et al. observations largely agree with the transmission electron microscopic investigations of Raikov & Kovaleva (1995) and Raikov et al. (1975) who, however, did not recognize the oralized somatic kinetids and some site-specific differences. On the other hand, the transverse microtubule ribbons and kinetodesmal fibres did not stain in Foissner preparations. The most conspicuous fibres are the postciliary microtubule ribbons, several of which overlap to form a distinct bundle (postciliodesma) right of each kinety. The postciliodesmata are thinner in the head and neck region than in the trunk and tail. The subkinetal microtubules form a very thin, but sharply impregnated bundle underneath or close to the left of the kineties. They do not or hardly overlap in the head, neck and tail region, where this comma-like shape can thus be recognized. All head and neck dikinetids have associated a thin, rather irregular fibre, very likely a nematodesma, extending obliquely to the centre. Thus, they are oralized somatic kinetids as defined by Foissner & Foissner (1988). The contractile apparatus of T. phoenicopterus is very similar to that described in Trachelocerca sagitta and T. ditis. However, the myonemes appear thinner and string-like and are lacking or unstained in the head and neck region. The glabrous stripe, which extends the whole length of the body, is narrow in the head region and widens, respectively narrows, gradually on the neck and tail. Its full width on the trunk corresponds to an area occupied by about 10 kineties, i.e. approximately two thirds of body width. The glabrous stripe is bordered by the bristle kinety which consists, like the ordinary ciliary rows, of dikinetids having about 12 um long, rather stiff cilia. However, the bristle kinety is easily distinguished from ordinary somatic ciliary row because its dikinetids are more widely spaced and more irregularly arranged and either lack or have very inconspicuous postciliary microtubule ribbons, too small to be recognized with the light microscope; there is, however, a very faintly stained fibre along its left branch. The bristle kinety is continuous at the posterior end of the cell, whereas its anterior end appears covered by a short, oblique kinety ("anterior arch of bristle kinety") composed of about 10-15 rather irregularly arrange dikinetids. The dikinetids of this segment might belong to the oral ciliature because they have, like the circumoral kinety and the brosse kineties, associated nematodesmata-like fibres extending posteriorly near the cell surface. The ciliation of the bristle dikinetids is the same as described in Trachelocerca sagitta, i.e. those along the right margin of the glabrous stripe have the posterior basal bodies ciliated, whereas the dikinetids along left stripe margin have the anterior basal bodies ciliated. The bristle kinety of T. phoenicopterus and some other species mentioned below is unique in being composed of many minute, oblique kineties, consisting of 2-5 dikinetids, in the trunk region. The proximal (inner) granule (rarely two) of the oblique kineties frequently appears unciliated and unpaired, indicating that it is not a kinetid but a special type of cortical granule, possibly an extrusome. The oblique kineties become gradually shorter and more vertically arranged towards the ends of the cell, where the bristle kinety is of usual structure, i.e. composed of single, rather widely spaced dikinetids. Foissner et al cannot exclude that this peculiar pattern is caused by a particular mode of contraction of the cell during fixation. However, Tracheloraphis aragoi and Trachelocerca sagitta, which are also highly contractile, lack such kineties. On the other hand, the glabrous stripe is much more narrow in T. aragoi and T. sagitta than in the other species, which certainly influences its shape in contracted cells. This could also explain the lack of minute kineties in the neck and tail region of T. phoenicopterus, T. longicollis and T. oligostriata, where the glabrous stripe is as narrow as in the trunk region of T. aragoi. Furthermore, the bristle kinetids are more widely spaced in the neck and tail than in the turn region, i.e. have enough space to arrange one behind the other when the cell contracts. (ref. ID; 2267)

Oral infraciliature: The oral infraciliature of T. phoenicopterus consists of a circumoral kinety and a distinct brosse, both associated with conspicuous fibres, very likely nematodesmata, clearly recognizable, however, only in perfectly impregnated specimens. The circumoral kinety extends in the flat furrow separating the oral bulge from the head and is composed of a single row of vertically orientated dikinetids having, very likely, only the posterior basal body ciliated. Each circumoral dikinetid is associated with a distinct fibre (nematodesma) obliquely extending into the head. The nemotodesmata of neighbouring dikinetids unite to conspicuous bundles, forming a cone-shaped oral basket. The circumoral kinety is very likely composed of rather many fragments, as indicated by small gaps, 1-2 dikinetids wide, and the bundled arrangement of the nematodesmata. The circumoral kinety is interrupted where the brosse kineties are inserted. Its left end simply abuts to the margin of the brosse cleft, i.e. to the left lip of the oral bulge. The right end is more complicated. It extends along the oblique margin of the brosse cleft and curves back at the cleft vertex in such a steep angle that a loop-like structure, or a small oblique segment, is formed paralleling the descending portion of the circumoral kinety and the anterior end of the neighbouring somatic kinety. These peculiar configuration becomes evident, as in the bristle kinety, from the ciliation of the dikinetids: those in the bulge furrow have the posterior basal body ciliated, whereas the anterior basal bodies are ciliated in the curved segment. The brosse is located in a rather deep cavity, the brosse pocket, just above the arch of the bristle kinety, and intersects the circumoral kinety. The cilia of the brosse emerge through the brosse cleft, which divides the oral bulge and the circumoral ciliature into a right and left half. The brosse consists of 2-4, usually 3, oblique kineties composed of closely spaced dikinetids having only the posterior basal body ciliated. The variation in the number of brosse kineties is not caused by a mixture of different species because the specimens with 2 or 4 kineties match those with 3 kineties very well in all other characteristics. Furthermore, a similar variation has been observed in T. aragoi and Prototrachelocerca. The brosse kineties are arranged in parallel one behind the other and their dekinetids are associated with distinct fibres extending, like the circumoral nematodesmal bundles, into the head. (ref. ID; 2267)

Identified by median cleft in the anterior mouth region and 6-8 oval macronuclei. (ref. ID; 2316)

Measurements

Size of fully extended specimens in vivo about 1,000-1,500x30-50 um, highly flexible and contractile, size and shape thus poorly preserved, but better than in Trachelocerca spp., in protargol slides. (ref. ID; 2267)
Mean length 310 um. (ref. ID; 2316)

Tracheloraphis poljanskyi (Raikov, 1963) Foissner, 1997 (ref. ID; 2859)

Descriptions

This species has only about 12 ciliary rows and a long, narrow neck. (ref. ID; 2859)

Tracheloraphis prenanti var. oligocineta Rainkov & Kovaleva, 1968 (ref. ID; 2316)

Descriptions

Body spindle shaped. Anterior head region slightly inflated and on contraction organism displayed a distinctive tail. Nuclear apparatus in large ovoid group, individual components could not be identified without staining. Whether this organism differed significantly to be assigned to a new species could not be ascertained from the observations made here. Borror (1973) in his review of the genus Tracheloraphis placed this species into the 'phoenicopterus' complex and Wright (1983) retained the variety. This species must therefore be viewed with caution until the varieties included in this species are reviewed. (ref. ID; 2316)

Measurements

Length 500 um. (ref. ID; 2316)

Tracheloraphis primitarum Epstein, 1994 (ref. ID; 2267, 7323 original paper)

Descriptions

Living ciliates were vermiform, 0.4-0.9 mm in length, narrow (length-to-width ratio 10:1 to 15:1), and elliptical in cross section. In both living and fixed specimens, the body was divided into three clearly defined regions conditionally called the "head" (bulbous anterior end with cytostome), the flexible "neck", and the body proper with rounded posterior end ("tail"). Under incident light, live specimens appeared gray and were not transparent except for the "neck" region. There appeared to be no contractile vacuole. Several specimens contained large unidentified diatoms. Upon fixation, the ciliates contracted; the length and width of fixed and stained specimens were 88-176 um (128 um average of 21 measurements) and 26-49 um (36 um average of 21 measurements), respectively. Somatic ciliature consisted of 18-29 (most often 24-28) kineties of dikinetids. The glabrous zone was exteremly narrow (6-7 um) and corresponded to the area occupied by one kinety and the two neighboring interkinetal spaces. Both sides of the glabrous zone were bordered by rows of disordered (not placed in straight kineties) monokinetids; a very similar pattern was observed in Tracheloraphis beninensis Dragesco and Dragesco-Kernies, 1987. Ten to fifteen kineties terminated against the glabrous zone, while the remaining 8-14 kineties ran the entire length of the cell. Interkinetal extrusomes (mucocysts?) were approximately 1 um in diameter; their number varied widely from a few to several hundred per interkinetal space. The cytostome was simple a cleft on the lip was not observed. One contiguous circumoral kinety surrounded the cytostome; corresponding kinetids appeared to be monokinetids. Extrusomes were present around the cytostome; their number varied from a few to several hundred. The nuclear apparatus was always positioned in the middle of the cell and consisted of 3 irregular macronuclei (Ma) and 2-3 micronuclei (Mi). The nuclear group was spherical or slightly oval in shape with a large axis 8-15 um (11 um average of 19 measurements) and small axis 6-13 um (10 um average of 19 measurements). The texture of macro- and micronuclei was different among specimens. In one quarter of the specimens, macronuclei were semi-transparent with distinct chromatin patches numbering five per macronucleus. In this case, parts of the nuclear apparatus were closely clustered but not encapsulated. In another quarter of the stained specimens, the macronuclei were smaller, dense and not transparent; the nuclear apparatus appeared to be encapsulated. In the remaining half of the specimens, the nuclear apparatus was seen in intermediate forms between the two extremes. Such variation in nuclear morphology are common within the genus Tracheloraphis and have been by others. (ref. ID; 7323)

Remarks

This species is sufficiently different from all the other Tracheloraphis species. Several of them (T. monokaryon Dragesco, 1965; T. lacteus Raikov and Kovaleva, 1968; T. conformis Wright, 1982; T. ditis Wright, 1982; T. niveus Wright, 1982), however, share with T. primitarum such key characteristics as rounded posterior, nuclear apparatus in the form of a single cluster and very narrow globrous zone. This necessitates detailed comparison of the above species. Compared to T. monokaryon Dragesco, 1965, T. primitarum sp. n. differed in the number of kineties (~~40 versus 18-29), width of the glabrous zone (2-3 versus 1 kinety) and the structure of nuclear apparatus (4Ma/2Mi versus 3Ma/2-3Mi). Compared to T. lacteus Raikov and Kovaleva, 1968, T. primitarum sp. n. differed in size (~~1.6 mm versus ~~0.65 mm), color of living specimens (white versus gray under incident light), the number of kineties (~~40 versus 18-29), and the structure of nuclear apparatus (8Ma/4Mi versus 3Ma/2-3Mi). Compared to T. conformis Wright, 1982 and T. ditis Wright, 1982, T. primitarum sp. n. differed in that the latter had three quite distinctive body regions, in the number of kineties (11-14 and 18-22 versus 18-29), and the structure of nuclear apparatus (4Ma/2-3Mi and 4-6Ma/1-2Mi versus 3Ma/2-3Mi). Compared to T. niveus Wright, 1982, T. primitarum sp. n. differed in color of living specimens (white versus gray under incident light), the number of kineties (36-47 versus 18-29), and the structure of nuclear apparatus (4Ma/2Mi in a loose group versus 3Ma/2-3Mi in a tight cluster or encapsulated form). The above differences indicate that the specimens indeed represent a distinct species. (ref. ID; 7323)

Etymology

The species name, primitarum (firstcoming), reflects the fact that this species was the very first that I met when I began working in this area of the Northwest Atlantic coast. (ref. ID; 7323)

Type material

The protargol stained type preparations are stored at the Marine Science Center, Northeastern University, Nahant, MA 01908. (ref. ID; 7323)