Holosticha

Holosticha Kahl, 1932 (ref. ID; 3690) or Wrzesniowski, 1877 (ref. ID; 662, 2014, 3925, 4697, 4905, 4913, 4927)

Phylum Ciliophora Doflein 1901: Subphylum Postciliodesmatophora Gerassimova & Seravin, 1976: Class Spirotrichea Butschli 1889: Subclass Stichotrichia: Order Stichotrichida Faure-Fremiet 1961 (ref. ID; 4778)
Class Polyhymenophora: Subclass Spirotricha: Order Hypotrichida: Suborder Stichotrichina: Family Holostichidae (ref. ID; 2014)
Order Hypotrichida Stein, 1859: Family Urostylidae Butschli, 1889 (ref. ID; 4905)

Synonym Amphisia Sterki, 1878 (ref. ID; 2014)

[ref. ID; 2014]
Elongate, cigar-shaped body. 2 rows of marginal cirri and a midventral series of 2 zigzag rows of cirri extending into the frontal region. 3 to 4 prominent frontal cirri at the anterior end and occasionally a few isolated ones elsewhere. Transverse cirri present. AZM relatively small, less than third of body length. Macronucleus usually in two parts but sometimes numerous. Many species have been described.
Quote; Colin R. Curds, Michael A. Gates and David McL. Roberts "British and other freshwater ciliated protozoa Part II Ciliophora: Oligohymenophora and Polyhymenophora" Cambridge University Press, 1983 (ref. ID; 2014)

[ref. ID; 3925]
One row of right marginal cirri; one or more rows of left marginal cirri; transverse cirri present; midventral cirri in typical zigzag series, 2 cirri per or original oblique ciliary streak, except that 3 or more frontal cirri differentiate from midventral cirri. Holosticha is a large genus (23 species recognized in 1972). (ref. ID; 3925)
Type genus (by monotypy); Holosticha kessleri Wrzesniowski, 1877 (ref. ID; 3925)

[ref. ID; 4391]
The main problem in identifying which ciliates belong to this group is the confusion created by Kahl (1930-1935) between the genera Holosticha and Keronopsis. Kahl (1930-1935) points out that in the genus Holosticha there are several subgenera. The species of subgenus Keronopsis have frontal cirri that are not, or are not clearly, differentiated from the right ventral row. Moreover, the subgenus Holosticha includes ciliates with three distinct spikes at the front end of the frontal field. Despite this clear separation, the same author combines species belonging to both subgenera as was pointed out by Dragesco (1966, 1970), Groliere (1975), Foissner (1982) and Borror & Wicklow (1983). Borror (1979) assigned full generic status to these subgenera and later redefined them. Hemberger and Wilbert (1982) re-examined Penard's description (1922) of Keronopsis and concluded that it was a genus of nonurostyline hypotrichs, closely related to Kerona in the family Keronidae. Because of this, it has been suggested that some species formerly in Keronopsis should be shifted to Holosticha. However, Borror & Wickelow (1983) believe that some of them are distinct at the generic and indeed the familial level from Holosticha. So, they erected Pseudokeronopsis gen. nov. with Pseudokeronopsis rubra (Ehrenberg, 1838) as species type. We agree with Borror & Wicklow (1983) and we think that in order to avoid further nomenclatural problems that ciliates with a clear frontal cirral ciliature should be included in the genus Holosticha while the genus Pseudokeronopsis should be reserved for those ciliates whose frontal cirri are not clearly differentiated from the rest. Furthermore, we agree with Borror & Wicklow that the differences are clear enough to separate sharply the Fam. Urostylidae (with the Subfam. Holostichinae) from the Fam. Pseudokeronopsidae. Foissner (1982) was the first author who, following Hemberger & Wilbert (1982), studied the species Keronopsis muscorum as H. muscorum and suggested that many other species included up to the moment in the old genus Keronopsis to Holosticha should be reallocated. But as a result of our morphological study, we are also interested in clarifying the differences between some species considered by Borror & Wicklow (1983) as belonging to the genus Holosticha. Following the binary key for the suborder Urostylina of Borror & Wicklow (1983), we cannot identify the species we have studied. In fact, the species according to this key that is closest to ours, H. intermedia, possesses several different characteristics: there is a little gap between the posterior end of the double row of mid-ventral cirri at the transverse cirri, the two marginal rows do not cross in the posterior end, there are two buccal cirri, and the mid-ventral cirri are clearly separated in two rows. Later, other species have been described belonging to the genus Holosticha. Holosticha sigmoidea seems to be smaller that H. corlissi n. sp. The left and right rows of the marginal cirri do not reach each other in the posterior end and there are two parabuccal cirri which do not appear in H. corlissi. In addition, there are only three or four transverse cirri and the number of mid-ventral cirri is smaller than in H. corlissi. Holosticha mancoidea does not seem to be a related species to H. sigmoidea or H. corlissi because of the incomplete row of mid-ventral cirri. Holosticha xanthichroma seems to be the most similar species to H. corlissi. The complete and large row of mid-ventral cirri, the shape of the cell and other detains are the same in both species. However, H. xanthichroma has a parabuccal cirrus, the AZM is 1/4 of the body length and the two marginal rows do not join in the posterior end as occurs in H. corlissi. Wirnsberger and Foissner (1987) suggest, after a comparison of the available biometrical data, that the same characters of the infraciliature are particularly useful for identifying Holosticha species: number of dorsal kineties, buccal cirri, parabuccal cirri, mid-ventral cirri, marginal cirri, adoral membranelles, length of adoral zone of membranelles and mid-ventral rows, crossed or open marginal rows. Some of these characters are sufficiently distinct to suggest that H. xanthichroma and H. corlissi are two different species. In order to complete Borror and Wicklow's key we have added the species H. sigmoidea, H. xanthichroma and H. corlissi. Consequently, the key would need to be changed in the following way:

27(26') Ciliates with a nearly complete row of mid-ventral cirri. Holosticha intermedia (Berth, 1889).
27' Ciliates with a row of mid-ventral cirri only about half cell length. Holosticha sphagni nov. comb.
27" Ciliates with a complete row of mid-ventral cirri:
- a) The marginal rows are not superimposed Holosticha sigmoidea (a short mid-ventral row) Holosticha xanthichroma (a large mid-ventral row)
- b) The marginal rows are superimposed: Holosticha corlissi. (ref. ID; 4391)

[ref. ID; 4927]
Holosticha Wrzesniowski, 1877 is a large genus of hypotrichs. It has been a melting pot for all urostylids with three distinct frontal cirri, a midventral complex composed of cirral pairs only, transverse cirri, and one left and open right marginal row. In addition, species with or without caudal cirri are included. Due to this unspecified characterization by plesiomorphies more than 100 species have been originally assinged to Holosticha and many other species have been transferred to and excluded from this genus (Berger 2001). A thorough revision of the urostylids yielded several interesting results showing, inter alia, that Holosticha is a monophyletic group comprising only seven species. The many other species that do not show the newly defined Holosticha groundplan have already been transferred to genera like, for example, Pseudokeronopsis Borror and Wicklow, 1983, Pseudoamphisiella Song, 1996, and Neokeronopsis Warren et al., 2002 or they are assigned to three genera established below. (ref. ID; 4927)
Redefinition (A = supposed apomorphies); Adoral zone of membranelles bipartite in proximal (ventral) and distal (frontal) positions (A). Rearmost membranelles of proximal portion slightly to distinctly wider than remaining (A). Undulating membranes short and in parallel. 3 enlarged frontal cirri. Buccal cirrus distinctly ahead of paroral (A). 2 frontoterminal cirri. Midventral complex composed of midventral pairs only. Number of transverse cirri equal to or only slightly lower than number of midventral pairs (A?). 1 left and 1 right marginal row. Anterior end of left marginal row composed of narrowly spaced cirri and distinctly curved rightwards (A). Caudal cirri lacking. Nuclear apparatus right of or in midline or scattered (A). Frontal-midventral-transverse cirral anlagen originate mainly from right midventral cirri and occur thus basically right of the parental midventral complex (A). Parental adoral zone remains more or less unchanged for proter. Left marginal row anlage for proter originates de novo (A). (ref. ID; 4927)
Brief history of Holosticha; Wrzesniowski (1877) recognized that Oxytricha sensu Stein (1859) is heterogeneous. Thus he suggested it be split into two groups: (i) species with interrupted ventral cirral rows which could keep the name Oxytricha (basically these are the flexible 18-cirri oxytrichids; for review, see Berger 1999); and (ii) species with continuous ventral rows for which he proposed the name Holosticha. Unfortunately, Wrzesniowski did no include the frontal ciliature (three or many frontal cirri) into the definition. Further, he forgot to fix one of the seven species originally included as type species. In spite of these deficiencies, Entz (1884) merged Holosticha and Amphisia which was characterized more precisely by Sterki (1878). Kahl (1932) accepted Entz's synonymy and, in addition, split Holosticha into five subgenera (Holosticha, Paruroleptus, Trichototaxis, Amphisiella, Keronopsis) increasing the number of Holosticha species significantly. Borror (1972) raised the subgenera again to genus rank, and Borror and Wicklow (1983) transferred Holosticha species with a bicorona to Pseudokeronopsis. Recently some further Holosticha species were put into Pseudoamphisiella and Neokeronopsis by Song (1996) and Warren et al. (2002). (ref. ID; 4927)
Type fixation; Wrzesniowski (1877) did not fix one of the seven species originally included as the type. Kahl (1932) assumed that Oxytricha kessleri Wrzesniowki, 1877 was the type species, and likely for that reason Borror (1972) in his revision wrote that this species is the "type by monotypy". Although the phrase "... by monotypy" is incorrect it seems wise to accept Borror (1972) as author who fixed the type species by subsequent designation. Sterki (1878) fixed, although somewhat cryptically. Trichoda gibba Muller, 1786 (incorrectly named Oxytricha gibba Stein by Sterki) as the type of Amphisia. This species was also originally included in Holosticha and is now considered as a senior synonym of Oxytricha kessleri. Synonymy of these two type species was already supposed by Kahl (1932). (ref. ID; 4927)
Brief discussion of apomorphies:

(i) Gap in adoral zone. A distinct gap (break) is not very common in hypotrichs. Within the urostylids a bipartite zone is described basically for Afrothrix Foissner, 1999, Parabirojimia Hu et al., 2002, and Uroleptopsis (unpubl. observations). Since other groups, e.g. euplotids, oxytrichids, amphisiellids, and kahlielids, usually have a continuous adoral zone we have to assume that the gap is a novelty. However, very likely the gap evolved convegently several times within the urostylids because it is unlikely that Holosticha, Afrothrix, Parabirojimia, and Uroleptopsis form a monophylum with the gap as autapomorphy.
(ii) Proximalmost adoral membranelles widened. The widening is different within Holosticha, for example, rather inconspicuous in H. diademata or very pronounced in H. spindleri and H. bradburyae. I do not know another group where such a curious pattern occurs, indicating that the Holosticha state is apomorphic.
(iii) Buccal cirrus distinctly ahead of paroral. Usually the buccal cirrus is right of paroral. By contrast, in all several Holosticha species it is distinctly ahead of the undulating membranes indicating that this is the apomorphic state.
(iv) Number of transverse cirri equals or is only slightly lower than number of midventral pairs. In most taxa with a considerable number of midventral pairs, e.g. Pseudokeronopsis, only the posteriormost anlagen produce transverse cirri (Wirnsberger 1987); by contrast, Holosticha species produce almost the same number of transverse cirri as midventral pairs (e.g. Hu et al. 2002). This is reminiscent of Pseudoamphisiella which, however, has caudal cirri (lacking in Holosticha) and forms a new adoral zone for the proter, whereas the parental zone is retained in Holosticha (Song et al. 1997; Hu et al. 2000). However, polarisation (plesiomorphic/apomorphic state) of this character is difficult. It could also be possible that this Holosticha-state is the plesiomorphic one because in 18-cirri oxytrichids also each frontal-ventral-transverse cirral anlage produces a transverse cirrus. The loss of some transverse cirri in e.g. Urosomoida, is there considered as the apomorphic state (for review, see Berger 1999).
(v) Anterior end of left marginal row composed of narrowly spaced cirri and distinctly curved rightwards. This is likely the best apomorphy of Holosticha because the feature is distinct in all species and does not occur in other groups. Possibly this curve is the reason why the anlage of the left marginal row of the proter originates de novo left of the proximal portion of the adoral zone (e.g. Hu et al. 2000). This is exactly the same region where in species with a normal, that is, straight left marginal row the anlage occurs within the parental row.
(vi) Nuclear apparatus in body midline or right of it. Usually the macronuclear nodules are arranged left of midline. Surprisingly, in all Holosticha species which have two or several nodules they are dislocated distinctly rightwards or at least in the midline.
(vii) Frontal-midventral-transverse cirral anlagen originate basically right of the midventral complex. This feature is described at least for H. diademata, H. pullaster, and H. heterofoissneri (e.g. Hu et al. 2000; Hu and Song 2001). In most (all?) other taxa these anlagen occur left of the midventral complex and usually the left cirri contribute to primordia formation indicating that this is the plesiomorphic state.
(viii) Anlage for left marginal row of proter originates de novo. see feature (v). (ref. ID; 4927)

Species assignable: Holosticha bradburyae Gong et al., 2001; Holosticha diademata (Rees, 1884) Kahl, 1932; Holosticha foissneri Petz et al., 1995; Holosticha gibba (Muller, 1786) Wrzesniowski, 1877; Holosticha heterofoissneri Hu and Song, 2001; Holosticha pullaster (Muller, 1773) Foissner et al., 1991; Holosticha spindleri Petz et al., 1995 (ref. ID; 4927)
Remarks; The list of synonyms contains only the original description of Holosticha and its junior synonym Amphisia, and some major revisions. As already mentioned in the introduction more than 100 species have been originally assigned to Holosticha. At present Holosticha comprises about 49 species. However, only seven of them, including the type species, form a homogeneous group characterized by the features mentioned above. There are two further characters which could be apomorphies of Holosticha, namely (i) the contractile vacuole is usually mid-body or behind it whereas in most other urostylids, oxytrichids, etc. this organelle is usually at least not behind mid-body; and (ii) the undulating membranes are more or less straight and roughly in parallel. In addition, all seven species are marine, except for H. pullaster which occurs in marine and limnetic habitats. Consequently, all species which do not show this Holosticha-groundplan have to be transferred to genera established below. All seven species now included are described in detail either by redescriptions (H. gibba, H. pullaster, H. diademata; e.g. Kahl 1932; Foissner et al. 1991; Petz et al. 1995; Song and Wilbert 2002) or in the original descriptions (other species; Petz et al. 1995; Gong et al. 2001; Hu and Song 2001). Kahl (1932) provided a very good redescription from life of Holosticha kessleri - the junior synonym of the type H. gibba - showing many apomorphies mentioned above. Kahl's redescription should be thus considered as authoritative until the type is defined via a neotype. Unfortunately younger redescriptions of H. kessleri, e.g., those by Borror (1979), Ricci et al. (1982), and Borror and Wicklow (1983), do not show more details than those by Kahl. In spite of this, there is no reasonable doubt that the seven species now assigned form a monophyletic group. (ref. ID; 4927)

Holosticha adami Foissner, 1982 (ref. ID; 622 original paper, 4778, 4927) reported year? (ref. ID; 2099, 4842)
See; Anteholosticha adami (ref. ID; 4927)
Holosticha algivora Kahl, 1932 (ref. ID; 4488, 4927) reported year? (ref. ID; 1219, 1621)
See; Caudiholosticha algivora (ref. ID; 4927)
Holosticha alveolata (ref. ID; 1621)
Holosticha arenicola (ref. ID; 1621)
Holosticha arenicola Dragesco, 1963 (ref. ID; 4927)
See; Biholosticha arenicola (ref. ID; 4927)
Holosticha arenicola Kahl, 1932
See; Anteholosticha arenicola (ref. ID; 4927)
Holosticha australis Blatterer & Foissner, 1988 (ref. ID; 4861, 4927)
See; Anteholosticha australis (ref. ID; 4927)
Holosticha begoniensis Fernandez-Leborans, 1990 (ref. ID; 4778 original paper)
Holosticha bergeri Foissner, 1987 (ref. ID; 2128)
See; Anteholosticha bergeri (ref. ID; 4927)
Holosticha brachysticha Foissner, Agatha and Berger, 2002
See; Anteholosticha brachysticha (ref. ID; 4927)
Holosticha bradburyae Gong et al., 2001 (ref. ID; 4913, 4927)
Holosticha brevis Kahl, 1932 (ref. ID; 4927) reported author and year? (ref. ID; 1621)
See; Anteholosticha brevis (ref. ID; 4927)
Holosticha camerounensis Dragesco, 1970 (ref. ID; 4927) reported author and year? (ref. ID; 4778)
See; Anteholosticha camerounensis (ref. ID; 4927)
Holosticha contractilis Dragesco, 1970
See; Uroleptus musculus (ref. ID; 4609)
Holosticha corlissi Fernandez-Galiano & Calvo, 1992 (ref. ID; 4391 original paper)
Holosticha danubialis Kaltenbach, 1960 (ref. ID; 4730)
See; Holosticha pullaster (ref. ID; 4609)
Syn; Holosticha retrovacuolata Tucolesco, 1962 (ref. ID; 4730); Holosticha rhomboedrica Vuxanovici, 1963 (ref. ID; 4730)
Holosticha diademata (Rees, 1883) (ref. ID; 3925) or 1884 (ref. ID; 1621, 3119) reported year? (ref. ID; 3771), (Rees, 1884) Kahl, 1932 (ref. ID; 4927) or (Rees) Kahl, 1932 (ref. ID; 645) reported year? (ref. ID; 5462) reported author and year? (ref. ID; 191)
Holosticha discocephalus Kahl, 1932 (ref. ID; 3119, 3690, 4927) reported author and year? (ref. ID; 1621)
See; Biholosticha discocephalus (ref. ID; 4927)
Holosticha distyla Buitkamp, 1977
See; Anteholosticha distyla (ref. ID; 4927)
Holosticha estuarii Borror & Wicklow, 1983
See; Anteholosticha estuarii (ref. ID; 4927)
Holosticha extensa Kahl, 1932 (ref. ID; 4927) reported author and year? (ref. ID; 1621)
See; Anteholosticha extensa (ref. ID; 4927)
Holosticha fasciola Kahl, 1932 (ref. ID; 4927) reported author and year? (ref. ID; 1621)
See; Anteholosticha fasciola (ref. ID; 4927)
Holosticha flavorubra Entz, 1884
See; Keronopsis rubra (ref. ID; 1621)
Holosticha foissneri Petz et al., 1995 (ref. ID; 4927)
Holosticha geleii (ref. ID; 191)
Holosticha gibba (O.F. Muller, 1786) Stein, 1859 (ref. ID; 1621)
Holosticha gibba (O.F. Muller, 1786) Wrzesniowski, 1877 (ref. ID; 4927)
Holosticha grisea Kahl, 1932 (ref. ID; 4927) reported author and year? (ref. ID; 1621)
See; Anteholosticha grisea (ref. ID; 4927)
Holosticha heterofoissneri Hu & Song, 2001 (ref. ID; 4927)
Holosticha hymenophora Stokes, 1886 (ref. ID; 1621) reported year? (ref. ID; 1618)
Holosticha intermedia (Bergh, 1889) (ref. ID; 1621, 4391) reported year? (ref. ID; 3491)
Holosticha interrupta Dragesco, 1966 (ref. ID; 4927)
See; Caudiholosticha interrupta (ref. ID; 4927)
Holosticha islandica Berger & Foissner, 1989 (ref. ID; 4927)
See; Caudiholosticha islandica (ref. ID; 4927)
Holosticha kessleri (Wrzesniowski, 1877) (ref. ID; 1335, 1621, 1629, 3925, 4488) or (Wrzesniowski, 1877) Wrzesniowski, 1877 (ref. ID; 4609)
Syn; Oxytricha kessleri Wrzesniowski, 1877 (ref. ID; 4609); Oxytricha wrzesniowskii Mereschkowsky (ref. ID; 4609)
Holosticha kessleri var. aquae-dulcis Buchar, 1957
See; Holosticha pullaster (ref. ID; 4609)
Holosticha lacazei Maupas, 1883 (ref. ID; 4889) or 1888 (ref. ID; 1621) reported year? (ref. ID; 5462)
See; Pseudoamphisiella lacazei (ref. ID; 4889)
Holosticha manca Kahl, 1932 (ref. ID; 4778, 4905, 4927) reported author and year? (ref. ID; 1621)
See; Anteholosticha manca (ref. ID; 4927)
Holosticha mancoidea Hemberger, 1985 (ref. ID; 4697 original paper, 4927) reported author and year? (ref. ID; 191)
See; Anteholosticha mancoidea (ref. ID; 4927)
Holosticha milnei (ref. ID; 1621)
Holosticha monilata Kahl, 1928 (ref. ID; 4609, 4813, 4927) reported author and year? (ref. ID; 1629, 4391)
See; Anteholosticha monilata (ref. ID; 4927)
Holosticha mononucleata Gelei, 1954 (ref. ID; 2129)
Holosticha multicaudicirrus Song & Wilbert, 1989
See; Caudiholosticha multicaudicirrus (ref. ID; 4927)
Holosticha multinucleata var. decolor Wallengren, 1900
See; Keronopsis decolor (ref. ID; 1621)
Holosticha multistilata Kahl, 1928 (ref. ID; 1629, 2128, 3925, 4609, 4861, 4927) or 1932 (ref. ID; 662, 4778) reported author and year? (ref. ID; 191)
See; Anteholosticha multistilata (ref. ID; 4927)
Holosticha muscicola Gellert, 1956
See; Anteholosticha muscicola (ref. ID; 4927)
Holosticha muscorum Kahl, 1932 (ref. ID; 662, 4778, 4927) reported year? (ref. ID; 2099)
See; Anteholosticha muscorum (ref. ID; 4927)
Holosticha (Paruroleptus) muscorum Kahl, 1932 (ref. ID; 7585)
Holosticha musculus (ref. ID; 191)
Holosticha mystacea Stein, 1859 (ref. ID; 1621)
Holosticha navicularum Kahl, 1932 (ref. ID; 4927) reported author and year? (ref. ID; 1621)
See; Caudiholosticha navicularum (ref. ID; 4927)
Holosticha novitas Horvath (ref. ID; 1335)
Holosticha oculata (Mereschkowski, 1877) (ref. ID; 1621) or Mereschlowsky, 1877 (ref. ID; 4927)
See; Anteholosticha oculata (ref. ID; 4927)
Holosticha pseudorubra (Kaltenbach, 1960) (ref. ID; 4730)
Syn; Keronopsis pseudorubra Kaltenbach, 1960 (ref. ID; 4730)
Holosticha pulchra Kahl, 1932
See; Anteholosticha pulchra (ref. ID; 4927)
Holosticha pullaster (Mueller, 1773) Foissner et al., 1991 (ref. ID; 4488, 4609, 4927) reported author and year? (ref. ID; 1629)
Syn; Holosticha danubialis Kaltenbach, 1960 (ref. ID; 4609); Holosticha kessleri var. aquae-dulcis Buchar, 1957 (ref. ID; 4609); Holosticha retrovacuolata Tucolesco, 1962 (ref. ID; 4609); Holosticha rhomboedrica Vuxanovici, 1963 (ref. ID; 4609); Holosticha simplicis Wang & Nie (ref. ID; 4609); Oxytricha alba Fromental, 1876 (ref. ID; 4609); Trichoda pullaster Mueller, 1773 (ref. ID; 4609)
Holosticha punctata Rees (ref. ID; 1621)
Syn; Holosticha wrzesniowskii var. punctata Rees, 1884 (ref. ID; 1621)
Holosticha randani Groliere, 1975
See; Anteholosticha randani (ref. ID; 4927)
Holosticha retrovacuolata Tucolesco, 1962 (ref. ID; 4609, 4730) reported year? (ref. ID; 3698)
See; Holosticha danubialis (ref. ID; 4730), Holosticha pullaster (ref. ID; 4609)
Holosticha rhomboedrica Vuxanovici, 1963
See; Holosticha danubialis (ref. ID; 4730), Holosticha pullaster (ref. ID; 4609)
Holosticha rubra Ehrenberg, 1838
See; Keronopsis rubra
Holosticha scutellum Cohn, 1866 (ref. ID; 1621, 3925)
See; Anteholosticha scutellum (ref. ID; 4927)
Holosticha setifera Kahl, 1932 (ref. ID; 4927) reported author and year? (ref. ID; 1621)
See; Caudiholosticha setifera (ref. ID; 4927)
Syn; Holosticha obliqua Kahl, 1928 (ref. ID; 1621)
Holosticha sigmoidea Foissner, 1982 (ref. ID; 662 original paper, 4778, 4927) reported year? (ref. ID; 4842) reported author and year? (ref. ID; 4391)
See; Anteholosticha sigmoidea (ref. ID; 4927)
Holosticha similis Stokes, 1886 (ref. ID; 4778) reported author and year? (ref. ID; 191)
Holosticha simplicis Wang & Nie, 1932 (ref. ID; 1620)
See; Holosticha pullaster (ref. ID; 4609)
Holosticha sphagni Groliere, 1975 (ref. ID; 4927) nov. comb. (ref. ID; 4391)
See; Anteholosticha sphagnia (ref. ID; 4927)
Holosticha spindleri Petz et al., 1995 (ref. ID; 4927)
Holosticha stueberi Foissner, 1987
See; Caudiholosticha stueberi (ref. ID; 4927)
Holosticha sylvatica Foissner, 1982 (ref. ID; 662 original paper, 4778, 4861)
See; Caudiholosticha sylvatica (ref. ID; 4927)
Holosticha tetracirrata Buitkamp & Wilbert, 1974 (ref. ID; 662, 2345, 4778)
See; Caudiholosticha tetracirrata (ref. ID; 4927)
Holosticha thononensis Dragesco, 1966
See; Anteholosticha thononensis (ref. ID; 4927)
Holosticha vernalis Stokes, 1887 (ref. ID; 1621) reported year? (ref. ID; 1618)
Holosticha violacea Kahl, 1928 (ref. ID; 1621)
See; Anteholosticha violacea (ref. ID; 4927)
Holosticha viridis Kahl, 1932 (ref. ID; 4927) reported author and year? (ref. ID; 1621)
See; Caudiholosticha viridis (ref. ID; 4927)
Holosticha warreni Song & Wilbert, 1997 (ref. ID; 4905)
See; Anteholosticha warreni (ref. ID; 4927)
Holosticha wrzesniowskii (Mereschkowski, 1877) (ref. ID; 1621)
Holosticha wrzesniowskii var. punctata Rees, 1884
See; Holosticha punctata Rees (ref. ID; 1621)
Holosticha xanthichroma Wirnsberger & Foissner, 1987 (ref. ID; 4927) reported author and year? (ref. ID; 4391)
See; Anteholostica xanthichroma (ref. ID; 4927)

Holosticha begoniensis Fernandez-Leborans, 1990 (ref. ID; 4778 original paper)

Diagnosis

Elongate ciliates which are flat on their dorsal and ventral sides, measuring 187-198 um in length and 48-67 um in width, having a narrow caudal zone (42-44x13-15 um). Numerous macronuclear nodes (55-64). Adoral area with 49-55 membranelles and paroral formation with two parts: a diplostichomonad on the right and a polystichomonad anteriorly combined with a posterior stichomonad on the left, on the right edge of the oral cavity. 12-20 frontal cirri in two rows. 35-45 left marginal cirri and 46-50 right marginal cirri. 48-54 ventral cirri in two rows. 5 transverse cirri arranged in a "J". 2-3 ventral cirri directly above the transverse cirri. Habitat in marine areas. (ref. ID; 4778)

Descriptions

The specimens are elongate, flat on the dorsal and ventral sides, and more or less oval when considered from the dorsal or ventral side. The body is 187.5-198.6 um long and 48.1-67.5 um wide. The posterior zone of the body has a narrow caudal area that is 42-44.2 um long and 13.5-15 um wide. There are 55-64 macronuclear nodes distributed over the entire body. Each of these is 5.4-8.4 um long and 3-5.1 um wide. Each of the numerous micronuclei has diameter of 1.2-2 um. The ciliature forms two groups of structures on the ventral surface: the oral ciliature and the cirri. (ref. ID; 4778)

Oral ciliature: This is located on the anterior left side of the ventral surface and comprises two components: the zone of membranelles or adoral organelles (AM) and the paroral formation (PF). The zone of membranelles (AM) is 72-85.8 um long and 6.7-11.4 um wide. It consists of a group of 49-55 membranelles. These all have a similar structure except the posterior and anterior ones, which have fewer rows of kinetosome containing less kinetosomes. At the posterior end, there is usually one membranelle with two kinetosome rows that are shorter than in the other membranelles. On the anterior part of the body, the AM surrounds the upper margin, and extends to the right side of the ciliate, although it is developed to a greater degree on the left side. Most of the membranelles have four parallel rows of kinetosomes arranged perpendicular to the main axis of the AM. The two middle rows are of the same length and have an average of 32 kinetosomes each. The anterior row is the shorter row and has some 20 kinetosomes. The front row is much shorter and has an average of three kinetosomes. The cytostome of the ciliate is located near the posterior end of the AM, which is situated 44.4-46.8 um from the anterior pole and 143.9-148.5 um from the posterior pole of the ciliate. The paroral formation (PF) is on the right of the zone of membranelles, runs lengthwise and borders the right side of the oral region. It consists of two structures: paroral formation 1 (PF1) and paroral formation 2 (PF2). The two formations often cross one anther more or less at the median point of their course. PF1 is furthest to the right, at least at its anterior end. It is median point of their course. PF1 is furthest to the right, at least at its anterior end. It is 25.8-30.6 um long and comprises two parallel rows of 37-46 kinetosomes each (the kinetosomes are sometimes staggered and form a stichodyad). PF2 has its anterior end to the left of PF1 and is longer (31.2-40.8 um). The arrangement of kinetosomes in this structure is variable. In general, it is divided into two parts: an anterior one consisting of an elongate group of kinetosomes arranged in a triple (polystichomonad) or double row (diplostichomonad), and a posterior part invariably consisting of a single row of kinetosomes (stichomonad). In all, there are 68-78 kinetosomes in PF2. The anterior end of the paroral formation is 16.8-21 um from the anterior pole and the posterior end 147-151.6 um from the posterior pole of the ciliate. (ref. ID; 4778)
Cirri: There are four types of cirri: frontal, marginal, ventral and transverse. The frontal cirri are anterior, and on the right of the paroral formation. They are arranged in two rows (F1 and F2) that curve in the anterior area and run lengthwise toward the midsection of the silicate. The F1 cirri are further to the right and occupy an area with a total length of 21-34.8 um. There are 7-10 of them and they are usually arranged in three rows with three kinetosomes each. Their bases are 1.2-1.25 um long and 1.26-1.56 um wide and exhibit no appreciable derivatives. However, in some specimens a short bundle has been observed to run from the cirrus toward the posterior right side of the ciliate in the two or three anterior-most cirri. This is the posterior right bundle (prb). The F2 cirri close to the paroral formation occupy an area that is 27.6-33 um long. Numbering 5-10, they generally have the same kinetosomic composition as the F1 cirri, but in some specimens one or two DWS with 4 kinetosomes have been observed in each cirrus. In other cases, some of these cirri possess only two rows with three kinetosomes each. The derivatives of these cirri (F2) are of two types: there is a bundle stemming from each cirrus in an anterior right direction (the anterior right bundle, arb) and having a length of 0.84-1 um; the other derivative is a long, thick fibrillar bundle running from the cirrus in a posterior left direction (posterior left bundle, plb) to a point near the paroral formation and having a length of 10.8-13.2 um. The posterior left bundles join together in the area adjacent to the paroral formation and curve posteriorly towards the inferior end of the oral cavity. The marginal cirri occupy a large part of the right and left hand edges of the ventral zone of the ciliate. There is a row of right marginal cirri (M1) and a row of left marginal cirri (M2). The right marginal cirri (M1) run from a point near the right anterior end of the zone of membranelles to the narrow caudal area at the posterior end of the ciliate. This group occupies an area of 125.6-131.7 um. It comprises 35-45 cirri consisting in most specimens of two rows with 4 kinetosomes each. However, cirri of this type having two rows with three kinetosomes each have been observed in some cases and three rows with 5 kinetosomes each in others. There is a bundle running in an anterior right direction (anterior right bundle, arb) from each cirrus. This bundle is 1.92-2.4 um long. In some specimens and in the cirri located most anteriorly in this group, there are two such bundles per cirrus, i.e. there is supplementary bundle to the left of the one described. The left marginal cirri (M2) run from an area near the posterior left part of the zone of the membranelles to points near the narrow caudal area. The area they occupy is 93.4-98 um long. There are 46-50 of these cirri, and they have the same derivatives and kinetosomic composition as the right marginal cirri. The ventral cirri (V) are located midway along the ventral surface and extend from the posterior end of the frontal cirri to the beginning of the narrow posterior end of the ciliate. They occupy an area 87.3-96.5 um long. The posterior end of this group is located 117.3-121 um from the anterior pole and 69.2-71.4 um from the posterior pole of the ciliate. In all, there are 48-54 ventral cirri grouped in pairs. The members of each pair are morphologically different, and it is thus possible to distinguish between right and left ventral cirri. The right ventral cirri (24-27) each consist of three rows having four kinetosomes each. These generally possess no derivatives. In some specimens, however, the 2-3 anterior-most cirri have a derivative of similar length and position to that of the marginal cirri. The left ventral cirri (24-26) each have two rows with four kinetosomes per row. Each of these cirri has two derivatives running beside each other from the cirrus to the posterior zone of the ciliate (posterior bundles: pb). Both are 0.28-0.36 um long. Directly above the transverse cirri there are 2-3 cirri with a kinetosomic composition similar to that of the left ventral ones and without appreciable derivatives. There are considered to be of taxonomic significance. There are five transverse cirri (T) arranged in the shape of a "J" as in many other similar ciliates. They are located on the median left zone of the narrow caudal area. Each cirrus consists of three rows with 4-5 kinetosomes each. (ref. ID; 4778)

Holosticha corlissi Fernandez-Galiano & Calvo, 1992 (ref. ID; 4391 original paper)

Descriptions

Ellipsoid to nearly parallel sided, both ends rounded and slightly narrowed at the level of the AZM. Average of 45 adoral membranelles formed by four kineties. Two paroral kineties, the endoral one being double. Three frontal cirri, the one situated on the right is placed at the beginning of the double midventral row. Two cirri placed behind the last one described, called fronto-terminal cirri. There is a single buccal cirrus. Generally six transverse cirri (range, 5-7). The two rows of marginal cirri cross in the posterior end. (ref. ID; 4391)

Comments

Body shape is nearly the same as that described for Holosticha intermedia and Holosticha monilata: ellipsoid with parallel margins, anterior and posterior ends rounded (with the posterior end a little more pointed than the anterior end), and slightly narrowed at the level of the adoral zone of membranelles. It is dorsoventral flattened. The endoplasma contains numerous subpellicular granules that may be mucocysts. The food vacuoles are relatively large, often containing green unicellular algae and diatoms. The movement is moderately rapid at the bottom of the Petri dishes. With respect to comparable morphological structures of H. corlissi n. sp., the infraciliature of this species is very stable in our populations as opposed to observations of H. monilata made by Dragesco (1966, 1970), Dragesco & Dragesco-Kerneis (1986) and Groliere (1975). The adoral zone of membranelles (AZM) is about one third of the body length (~/_72 um). It is formed by 38-55 paramembranelles with four kineties each: two are equal in length, one is shorter with 15 kinetosomes, and the other, the most anterior one, is the shortest with only 3-6 kinetosomes. At the right side of the buccal area, there is a double endoral membrane, a diplostychomonad, and a single paroral one, a stychomonad. They are clearly superimposed. All the ciliate have the three enlarged frontal cirri (FC): two are located closely behind the coronal paramembranelles and the third is situated where the AZM begins. Occasionally in some ciliate, another cirrus occurs to the left of the others. All frontal cirri are formed by six kineties. A short row of fronto-terminal cirri (FTC) with only 3-4 kineties is at the upper right corner. The two rows of mid-ventral cirri (MVC), characteristics of the order Urostylina, are in this species so close together that they seem to form a single row. In fact, it is a double row with 25 elements: each one has two cirri near to each other but separated by an argentophilic material (possibly a kinetodesmal fiber). The right cirrus has three kineties. When it is at the level of the cytostome, we can see the kinetodesmal fiber at the anterior corner as it appears in the cirri of the right marginal row. The left cirrus has 3-5 kineties, but it begins to have two when the right one shows the kinetodesmal fiber. It is placed at a 60 degrees angle to the transverse axis of the cell. This double row begins on the anterior AZM side and it is obliquely arranged with the last cirrus terminating very close to the transverse cirri (TC). There are six TC (occasionally five or seven can be seen). One of them is followed by the MVC and the rest are parallel and to the left of the MVC. They have four kineties. The left marginal row (LMR) begins below the level of the cytostome (under the 4-5 paramembranelles) with all the cirri situated near the body margin. Each cirrus has two kineties with a very long kinetodesmal fiber in it anterior right corner. The row of right marginal cirri begins at the level of the fronto-terminal cirri. Each of these cirri has two kineties. Its kinetodesmal fiber does not appear in the former cirri and it is shorter than the kinetodesmal fiber of the left row of marginal cirri. The two marginal rows never come together at the posterior end. They cross each other slightly with the last 2-3 right marginal cirri in front of the left ones. Six dorsal kineties are always present. The nuclear apparatus is formed by 8-16 macronuclei (Ma). They are ovoid with several small spherical nucleoli. They are located in the left side, between the mid-ventral and the left marginal rows. There are 2-8 micronuclei (Mi) located near the macronuclear segments. (ref. ID; 4391)

Type locality

Holostica corlissi n. sp. was isolated from the moss Calliergonella cuspidata taken from the beech wood of Montejo de la Sierra (Madrid, Spain). (ref. ID; 4391)

Type specimen

The holotype was deposited as Holosticha corlissi n. sp. at the Non Insect Invertebrate Collection, National Museum of Natural Sciences of Madrid, Spain, accession number MNCN-39.03/1. (ref. ID; 4391)

Measurements

Size in vivo 160-220x40-70 um. (ref. ID; 4391)

Holosticha diademata (Rees, 1883) (ref. ID; 3925) or 1884 (ref. ID; 1621, 3119) reported year? (ref. ID; 3771), (Rees, 1884) Kahl, 1932 (ref. ID; 4927) or (Rees) Kahl, 1932 (ref. ID; 645) reported year? (ref. ID; 5462) reported author and year? (ref. ID; 191)

Descriptions

Being 50 to 70 um in length, the members of the population studied appear to be the degeneration forms described by Kahl (1932): normal animals are between 70 to 140 um long. Kahl states that he found the species in all the saline waters he investigated. Pastsch (1974) found them in fresh water, and gives a detailed description of the infraciliature. Comparison of this with our samples made it possible to identify the species unequivocally. Characteristics of the genus are two rows of ventral cirri passing from one end of the body to the other, with 10 to 12 cirri in each row. In front of the hind end there are usually 7, rarely 8 transverse cirri, all of which project beyond the hind end. The row of marginal cirri on the left, numbering 9 to 13, begins below adoral membranelle zone with three cirri close together, at a right angle to the other cirri in this row. The right marginal row is composed of 12 to 16 cirri. The AZM, about 1/3 the length of the body, is unusual in shape, having a wide gap at the front end of the body that is apparently caused by the absence of membranelles. The AZM can comprise 22 to 25 membranelles. Usually 7 of these are isolated form the others, at the anterior end. (ref. ID; 645)

Holosticha hymenophora Stokes, 1886 (ref. ID; 1621) reported year? (ref. ID; 1618)

Descriptions

5 anals; 2 contractile vacuoles; shallow pools. (ref. ID; 1618)

Measurements

160-200 um long. (ref. ID; 1618)

Holosticha manca Kahl, 1932 (ref. ID; 4778, 4905, 4927) reported author and year? (ref. ID; 1621)

See

Anteholosticha manca (ref. ID; 4927)

Redescription

Cell flexible, shape usually elongate or fusiform with both ends more or less narrowly rounded, margins almost parallel, only slightly convex on right. Dorsa-ventrally about 2:1 flattened. Pellicle thin, cortical granules about 1 um in size, often several together and irregularly arranged in rows. Cytoplasm colourless to greyish, contains numerous shining globules (2-5 um in diameter). Contractile vacuole (CV) small, near left border and behind 1/3 of body length, with two conspicuous collecting canals during CV formation. Pulsation interval up to 5 minutes. About 50-70 ellipsoid macronuclear segments, each ca. 3-5 um long containing several large nucleoli. Micronuclei lenticular to globular, only recognized after protargol staining. Movement slowly to moderately fast crawling on substrate, sometimes jerking back and forth. Buccal field medium-wide, adoral zone of membanelles about 2/3 of cell length with distal end only slightly curved to right border. Bases of membranelles ca. 7-8 um in length. Cilia of membranelles 15-20 um long. Undulating membranes almost parallel and straight, somehow crossed. Three slightly enlarged and one smaller frontal cirri arranged in most anterior portion of buccal area, cirri about 20-25 um long. One single buccal cirrus near anterior end of undulating membranes. Frontoterminal cirri relatively fine, 10-15 um long, located between right end of adoral zone of membranelles and anterior terminus of right marginal row. Midventral row composed of oblique pairs of cirri, extending to about posterior 1/3 of cell length. Almost always 5 transverse cirri terminally positioned with relatively short cilia (ca. 12-15 um long), which extrude in vivo only inconspicuously over posterior margin of cell. Marginal rows separated posteriorly, no caudal cirri. Constant 3 dorsal kineties over entire length of cell, cilia about 2-3 um long. (ref. ID; 4905)

Comparison

The identification of this species is sound since the body size shape, biotope, general morphology and ciliature of our population correspond pretty well with previous studies (Kahl 1932; Agamaliev 1972, 1974). It differs from the similar form, Keronopsis rubra (Ehrenberg, 1838) sense Agamaliev, 1974 (possibly misidentified) in having markedly shortened mid-ventral rows (Agamaliev 1974). In contrast to Holostica warreni -a new species described by the same authors (Song & Wilbert 1997)- the form studied here is characterized by different body shape, form and arrangement of cortical granules, absence of contractile vacuole and evidently lower number of transverse cirri. (ref. ID; 4905)

Neotype specimens

1 neotype as a slide of protargol impregnated specimens has been deposited in the Protozoological Laboratory, College of Fishereis, Ocean University of Qingdao, Qingdao, China. (ref. ID; 4905)

Holosticha

Holosticha begoniensis Fernandez-Leborans, 1990 (ref. ID; 4778 original paper)

Diagnosis

Descriptions

Holosticha corlissi Fernandez-Galiano & Calvo, 1992 (ref. ID; 4391 original paper)

Descriptions

Comments

Type locality

Type specimen

Measurements

Holosticha diademata (Rees, 1883) (ref. ID; 3925) or 1884 (ref. ID; 1621, 3119) reported year? (ref. ID; 3771), (Rees, 1884) Kahl, 1932 (ref. ID; 4927) or (Rees) Kahl, 1932 (ref. ID; 645) reported year? (ref. ID; 5462) reported author and year? (ref. ID; 191)

Descriptions

Holosticha hymenophora Stokes, 1886 (ref. ID; 1621) reported year? (ref. ID; 1618)

Descriptions

Measurements

Holosticha manca Kahl, 1932 (ref. ID; 4778, 4905, 4927) reported author and year? (ref. ID; 1621)

See

Redescription

Comparison

Neotype specimens

Holosticha (Paruroleptus) muscorum Kahl, 1932 (ref. ID; 7585)

Descriptions

Holosticha vernalis Stokes, 1887 (ref. ID; 1621) reported year? (ref. ID; 1618)

Descriptions

Measurements