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Unitat de Botànica, Departament de Biologia Animal, de Biologia Vegetal i d’Ecologia, Facultat de Ciències, Universitat Autònoma de Barcelona, 08193-Bellaterra (Barcelona), Spain
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ABSTRACT |
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 TOP ABSTRACT INTRODUCTIONMATERIALS AND METHODSTAXONOMYKEY TO THE SPECIES...DISCUSSIONLITERATURE CITED |
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Two new genera of Laboulbeniales (Ascomycota) allied to Zodiomyces (Zodiomycetoideae) are described. These are Capillistichus with the single species C. tenellus, parasitic on Laccobius spp. (Coleoptera, Hydrophilidae), and Scepastocarpus with the single species S. peritheciiformis, parasitic on Hydrochus spp. (Coleoptera, Hydrophilidae). Both genera and species are from Spanish collections. Perithecial characters appear to be similar in the two new genera and in the related Zodiomyces, becoming a diagnostic character of the group (subfamily Zodiomycetoideae). Receptacular characters form the basis of separation into genera. To complete the study of all the taxa included in the subfamily, specimens and type slides of all three species of Zodiomyces have been reviewed.
Key words: aquatic beetles, Ascomycota, Hydrochus, Hydrophilidae, Laccobius, Spain, Zodiomycetoideae
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INTRODUCTION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSTAXONOMYKEY TO THE SPECIES...DISCUSSIONLITERATURE CITED |
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. The type species, Z. vorticellarius Thaxt., was collected in the United States on legs of Cymbiodyta lacustris (LeConte) (as Hydrocombus lacustris, Coleoptera, Hydrophilidae). Later, two additional species for this genus were described: Z. subseriatus Thaxt. (Thaxter 1931) and Z. odae T. Majewski & K. Sugiy. (Majewski and Sugiyama 1989). At first sight the most striking feature to distinguish the genus is the pseudoparenchymatous receptacle, consisting of hundreds of cells that support numerous perithecia and appendages in the cup-shaped apex. A pseudoparenchymatous receptacle also is found in other genera, such as Euzodiomyces, Columnomyces and Kainomyces. Several years ago I gathered many collections, including two undescribed species of Laboulbeniales, that seemed to belong to different genera with similarities to the genus Zodiomyces. The perithecial features of these two species agree with those described for the genus Zodiomyces, despite their receptacular features, which appear distinct from those of the three species of Zodiomyces.
Tavares (1985) defined the subfamily Zodiomycetoideae (Thaxt.) I.I. Tav. changing the status of the group for the previous tribe Zodiomyceteae Thaxt. (Thaxter 1908) and family Zodiomycetaceae (Thaxt.) Nann. (Nannizzi 1934). The Zodiomycetoideae containing only the genus Zodiomyces, therefore was defined narrowly by the characters of the genus. The two new genera and species here described should be included in this group and therefore its limits must be emended.
The purpose of this paper is: (i) to describe two new genera and species, (ii) to discuss their apparent relation to Zodiomyces based largely on perithecial characteristics, (iii) to illustrate all the taxa with line drawings and/or photographs, (iv) to provide a key for their identification and (v) to discuss aspects of morphology.
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MATERIALS AND METHODS |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSTAXONOMYKEY TO THE SPECIES...DISCUSSIONLITERATURE CITED |
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) and are kept in BCB (institutional herbarium at author’s address). Most microscopic observations were made with the help of differential interference contrast optics (DIC), and drawings were made with the help of a microscope equipped with a drawing device. Additional observations for Scepastocarpus peritheciiformis were made with scanning electron microscopy (SEM).
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TAXONOMY |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSTAXONOMYKEY TO THE SPECIES...DISCUSSIONLITERATURE CITED |
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Receptacle multiseriate or uniseriate, bearing abundant perithecia and appendages. Spermatia (where known) exogenous. Perithecia bearing two dorsal and two ventral appendages. Outer wall cells of perithecia subequal in height. Aquatic.
Three genera: Capillistichus, Scepastocarpus and Zodiomyces.
Capillistichus Santam., gen. nov.
Thallus uniseriato receptaculo constans, quod multicellulare stratum cellularum subtendit, multas appendices atque perithecia ferentium. Peritheciales cellulae externae parietis difficiles visu in maturite, sed omnes verticales series circa 6–7 subaequalibus cellulis constant. Omnia perithecia duas dorsales et duas ventrales appendices ferentia. Perithecialis apex 4 tenues ligulas ferens.
Thallus having a uniseriate receptacle subtending a multicellular layer of cells bearing abundant appendages and perithecia. Perithecial outer wall cells difficult to distinguish at maturity, but each vertical row includes about 6–7 subequal cells. Each perithecium bears two dorsal and two ventral appendages. Perithecial apex bearing four thin ligulae.
Etymology. – L., Capilli, from capillus = hair; and Gr., stichus = in a row or line; referring to the uniseriate receptacle bearing abundant appendages ("hairs").
TYPUS generis: Capillistichus tenellus Santam.
Capillistichus tenellus Santam., sp. nov. FIGS. 1–16, 29–46
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Thallus hyaline, 131–191 µm in length (including appendages and perithecia). Receptacle curved to strongly sigmoid (FIG. 5
), consisting of 6–9 superposed cells (including the basal cell), 60–100 µm in length. Basal cell obconical in shape, forming a nearly undifferentiated foot (FIG. 2), which is sometimes faintly yellowish. Above the basal cell, a narrow pedicel consists of 4–7 flattened to isodiametric cells, slightly broadening distally. The two (rarely three) distal cells of the receptacle are more or less globose, broader than long or vice versa, more or less laterally inflated, and always distinctively larger than the receptacular cells forming the pedicel below. The distal cell of the receptacle is always the largest cell, broader (15–42 µm breadth) and longer (often at least twice as long) than the cell below. The upper area of the receptacle, above the largest cell, consists of a layer (FIGS. 2, 4, 5) of many small cells, variable in shape, which are separated by longitudinal septa. These cells give rise to many perithecia or to long, unbranched appendages (up to 123 µm in length). Perithecia 70–95 x18–25 µm, each supported by a pedicel consisting of one or two cells. Stalk cell (VI), secondary stalk cell (VII) and basal cells of perithecia not well defined at maturity. Perithecial outer-wall cells also are difficult to distinguish at maturity, but each vertical row includes about 6–7 cells (FIGS. 39–41, 46). Each perithecium bears two dorsal and two ventral appendages (30–78 µm in length), the four upwardly orientated (FIGS. 14, 16, 40–41, 46). Perithecial apex acute, bearing four thin ligulae (FIGS. 12, 15, 41, 46). Trichogyne simple, downwardly incurved (FIGS. 7, 11, 33). Antheridia unknown.
Etymology. – L., tenellus = referring to the delicate, easily breakable structure of thalli.
Specimens examined. – SPAIN. GIRONA: La Jonquera, Canadal ponds, on Laccobius atrocephalus ytenensis Sharp (Coleoptera, Hydrophilidae, Hydrophilinae), 28 Jul 1990, I. Ribera, BCB-SS1933 (HOLOTYPE). Capmany, Torrelles stream, on undet. Laccobius, 9 Jul 1994 (BCB-SS2212), 10 Sep 1994 (BCB-SS2208a–2208b), I. Ribera. Masarac, Vilarnadal, Llobregat d’Empordà river, on undet. Laccobius, 9 Jun 1998, I. Ribera, BCB-SS2067. MÁ LAGA: Pujerra; Barranco Hondo stream, on L. (Platilaccobius) mulsanti Zaitzev, 13 Jul 1987, C. Hernando, BCB-SS1797. SEVILLA: Villanueva del Río y Minas, Parroso river, on L. mulsanti, 11 Jul 1985, C. Hernando, BCB-SS1776.
Remarks. – The discovery of this species is comparable to that of S. peritheciiformis. Locating specimens under a dissecting microscope is very challenging because they are easily overlooked. Thalli of C. tenellus have been found only on the abdominal sternites of its hosts, an area that very often appears covered by fatty substances and protozoan epibionts concealing the fungi. The fungi are hidden in a whitish, rather unstructured mass. Very thin filaments and white fusiform bodies (the perithecia of the fungi) can be seen when this mass is placed in ethanol. The fungi should be extracted when still soaked with the ethanol because they are almost invisible when dry. Thalli are delicate and break easily.
The description of a new genus appeared to be necessary, although the perithecial characteristics, like those of S. peritheciiformis, fit those described for the species of Zodiomyces. Certainly, perithecia of C. tenellus show the four typical appendages of the genus Zodiomyces (FIGS. 16, 40, 41, 46) and early stages of perithecial development (FIGS. 7, 8, 29–33) are very similar to those seen (FIG. 66) and described for Z. vorticellarius (Thaxter 1896) and those of Z. subseriatus (FIGS. 62, 63). Perithecia of the new species also have four thin ligulae in the perithecial apex (FIGS. 12, 15, 41, 46) as in Z. vorticellarius.
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Scepastocarpus Santam., gen. nov.
Thallus plus minusve sessili et multiseriato receptaculo constans, quae omnino inclusam cavitatem cingit, multas appendices et perithecia includentem. Peritheciales cellulae externae parietis indistinctae in maturite. Omnia perithecia duas dorsales et duas ventrales appendices ferentia.
Thallus having a more or less sessile multiseriate receptacle which entirely surrounds an enclosed cavity containing abundant appendages and perithecia. Perithecial outer wall cells become indistinct at maturity. Each perithecium bears two dorsal and two ventral appendages.
Etymology. – Gr., Scepasto, from skepastos = covered, and carpus, from carpon = fruit; referring to concealed growth and location of perithecia (fruit bodies) in a cavity of receptacle.
TYPUS generis: Scepastocarpus peritheciiformis Santam.
Scepastocarpus peritheciiformis Santam., sp. nov. FIGS. 17–28, 47–55
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Thallus 100–135 x50–58 µm, yellowish to amber, consisting of an urn-like receptacle having a slightly lateral, uncolored basal cell, which forms a broad, sucker-like foot (FIG. 26). The receptacle consists of many irregularly shaped cells, forming a pseudoparenchymatous tissue enclosing the remaining elements of the thallus, namely the perithecia and appendages, and which opens externally by means of an apical orifice (FIG. 28), resembling the ostiole of a typical "pyrenomycetous" perithecium. At maturity the lateral walls and neck areas of the urn-like receptacle are formed by one layer of cells giving rise internally to slender and very narrow secondary appendages (FIG. 49), which are more abundant in the neck area, where some emerge to the outside through the distal opening (FIG. 22). The base or "floor" inside the receptacle is basically two-layered and gives rise to several stalked perithecia and more appendages (FIG. 49). Perithecia 50–55 x11–13 µm, each supported by a two-celled pedicel, the upper cell of which is laterally extended into a stout upgrowth, reaching approximately half the length of the perithecium (FIGS. 22–24, 47, 48). Stalk cell (VI), secondary stalk cell (VII), and basal cells of perithecia are poorly defined at maturity. Also, the perithecial outer wall cells become indistinct, so that their numbers are unknown. Each perithecium bears two dorsal and two ventral appendages (up to 23 µm in length); the ventral ones are usually downwardly orientated (FIGS. 22–24, 47). Perithecial apex blunt. Trichogyne simple, more or less erect or with a slightly curved tip (FIGS. 53–55). Antheridia unknown.
Etymology. – L., peritheciiformis = referring to similarity in shape of thalli to typical perithecial ascomata.
Specimens examined. – SPAIN. GIRONA: La Jonquera, Canadal ponds, on Hydrochus angustatus Germar (Coleoptera, Hydrophilidae, Hydrochinae), 3 Feb 1990 (BCB-SS1929), 25 Aug 1990 (BCB-SS1928), 25 Nov 1990 (BCB-SS1927, HOLOTYPE), 8 Jan 1994 (BCB-SS2122), I. Ribera; same locality, on H. smaragdineus Fairmaire, 30 Jun 1990 (BCB-SS1932), 25 Aug 1990 (BCB-SS1930a–1930b), 28 Oct 1990 (BCB-SS1931), I. Ribera; same locality, on undet. Hydrochus, 11 May 1994 (BCB-SS2220a), 12 Jun 1994 (BCB-SS2223b), 9 Jul 1994 (BCB-SS2217a), 15 Oct 1994 (BCB-SS2139b, 2147, 2148a), 12 Nov 1994 (BCB-SS2141, 2146, 2163), 17 Dec 1994 (BCB-SS2165), 14 Jan 1995 (BCB-SS2161a), I. Ribera. Sant Climent Sescebes, rec Grimaus, on undet. Hydrochus, 10 Sep 1994, I. Ribera, BCB-SS2207. TERUEL: Tornos, canal lagunica, on undet. Hydrochus, 24 Sep 1994, I. Ribera, BCB-SS2179b, 2181b. Tornos, fuente cañizar, on undet. Hydrochus, 24 Sep 1994, I. Ribera, BCB-SS2186.
Remarks. –
This remarkable species was collected and prepared with no clear concept of its identity beside the possibility of its being a laboulbeniaceous fungus (FIG. 25). It was astonishing to see that, with the magnification of a compound microscope, within these structures a complete set of some typical structures of the Laboulbeniales existed, i.e., perithecia and appendages (FIGS. 18–20, 49). No other member of Laboulbeniales has a comparable habit.
Although at first appearance this fungus seems unique, careful examination revealed the similarities of this species with those species described in the genus Zodiomyces. The characteristics of the perithecium agree well those described for the perithecia of species of Zodiomyces, such as Z. vorticellarius. Mature perithecia of S. peritheciiformis bear four appendages on the wall cells (FIGS. 22–24, 47): two adjacent appendages from cells in dorsal rows and two from ventral rows, precisely the same as in Z. vorticellarius. In addition, the cell under the typical stalk cell (VI) shows a comparable prolongation (FIGS. 22–24, 47, 48). Furthermore, the development of this species and Z. vorticellarius (and it also may be assumed for Z. odae and Z. subseriatus) is similar. As indicated by Thaxter (1896), a shallower cavity is produced in young thalli of Z. vorticellarius and appendages and perithecia begin to grow from its floor; later the cavity forms an enlarged cup-shaped structure, filled with numerous perithecia, appendages and antheridia. In S. peritheciiformis, an enclosed cavity is formed in young individuals (FIG. 18), and only later an orifice is opened above (FIGS. 27, 28, 49). Scepastocarpus peritheciiformis obviously may be considered an extremely reduced form in comparison with Z. vorticellarius and, based on these receptacle features, it is described as a new genus, allied to Zodiomyces, that should be included with it and with Capillistichus in the subfamily Zodiomycetoideae (Tavares 1985).
The almost enclosed structure of the new species needs some mechanism for ascospore dispersal. We have observed that entire mature perithecia of several thalli emerged through the upper orifice of the thalli (FIGS. 27, 47) and remained attached in this position for a period of time (FIG. 47), as could be seen under the dissecting microscope. With manipulation in slide preparation, the perithecia become dislodged easily (FIG. 22). It may be assumed that the dispersion of ascospores occurs when the perithecium is attached apically, until it later is freed and disappears. Perithecia seem to be unattached even when they emerge through the orifice, because no elongation of a supporting pedicellar cell was observed. Even some immature perithecia are released (FIGS. 23, 24) if a slight pressure is applied to the cover glass over the thalli. If this is a natural occurrence, such a mechanism for release of the entire perithecia is unknown in any other members of the Laboulbeniales.
As stated in the description, the antheridia have not been seen (see discussion). When some thalli were squashed, young perithecia could be seen with long, straight or slightly curved trichogynes (FIGS. 53–55), supporting the probability of the presence of antheridia and, judging from the length and direction of trichogynes in S. peritheciiformis, antheridia might be located in the neck area of the receptacle.
This species coexists, although rarely, on the same host with Autoicomyces aquatilis (F. Picard) I.I. Tav. (for Spanish records of A. aquatilis, see Santamaria 2001).
Zodiomyces Thaxt., Proc. Am. Acad. Arts 25:263. 1891.
Thallus having a large, multiseriate, pseudoparenchymatous receptacle, at the top of which a cavity is formed and broadens into a shallow depression enclosing appendages and perithecia. Perithecial outer wall cells poorly defined at maturity, but each vertical row includes about eight subequal cells. Each perithecium bears two dorsal and two ventral appendages. Perithecial apex bearing four thin ligulae.
TYPUS generis: Zodiomyces vorticellarius Thaxt.
Zodiomyces odae T. Majewski & K. Sugiy., Trans. Mycol. Soc. Japan 30:83. 1989. FIGS. 56, 60
Known distribution and hosts. –
Known only from the collections included in the original description (Majewski and Sugiyama 1989) on Sternolophus rufipes Fabricius (Coleoptera, Hydrophilidae) from Japan (Iriomote Island).
Specimens examined. – JAPAN. IRIOMOTE ISLAND: Sonai, on S. rufipes, 4 Aug. 1985, K. Sugiyama et al, KS 4541 (PARATYPE).
Remarks. –
According to Majewski and Sugiyama (1989), this species differs from Z. subseriatus and Z. vorticellarius in lacking projections near the base of the receptacle and in having more cell layers in its receptacle. These authors also mentioned the impossibility of determining perithecial characters from the material available. The slide that I had the opportunity to study included two thalli (FIG. 56), in which perithecia in the apical cup of the thallus were concealed and impossible to describe without squashing the material. As will be discussed later in relation to Z. subseriatus and Z. vorticellarius, the number of buffer projections in the base of the thallus is not a sufficient character to distinguish the species of Zodiomyces. Nevertheless, it is true that the thalli of Z. subseriatus or Z. vorticellarius all bear at least one projection. However, the receptacle of Z. odae appears to be different in cell structure and cell arrangement when compared with the other two species mentioned. The pseudoparenchymatous tissue is formed by an outer layer of nearly cubical and relatively large, slightly inflated cells, whereas the inner cells are very narrow and rectangular (FIG. 60). This organization was not observed in the other species, where all the cells are very irregular in shape, and therefore, I take the conservative view of maintaining Z. odae as a separate species.
Zodiomyces subseriatus Thaxt., Mem. Am. Acad. Arts 16:331. 1931. FIGS. 57, 61–64
Known distribution and hosts. –
This species was described from a small undetermined hydrophilid collected in China, and from Enochrus esuriens Wlk. (as E. exuriens) and Berosus vitticollis Boh. collected in the Philippines (Thaxter 1931). Later, it was reported on Helochares anchoralis Sharp from Taiwan (Sugiyama 1981), on Helochares lentus Sharp from Thailand (Sugiyama and Phanichapol 1984; their FIG. 11 shows what appears to be Z. vorticellarius, instead, judging from shape of perithecia), on Helochares pallens (MacLeay) from Malaysia (Sugiyama and Majewski 1985), on H. lentus from Korea (Lee and Choi 1992, reiterating remarks and description of previous Asian reports), on Helochares sp. from Sudan (Weir 1993), on undetermined Helochares and Berosus from Sierra Leone (Rossi 1994; his FIGS. 20–21 show Z. vorticellarius, instead, with one receptacular basal cell), and on Helochares obscurus from Poland (Majewski 1999). All hosts belong to the Hydrophilidae (Coleoptera).
Specimens examined. – CHINA. Amoy, on a small hydrophilid of undet. genus, Chung, FH7167 [No. 3425 Type 1925] (HOLOTYPE).
Remarks. –
This species was described without an illustration (Thaxter 1931), a circumstance which hampered its easy recognition and cast doubt upon its distinctiveness from Z. vorticellarius. Sugiyama (1981) illustrated this species for the first time. Two primary characters have been used to separate Z. subseriatus and Z. vorticellarius: (i) Z. subseriatus has only one buffer projection at the base of the receptacle, whereas Z. vorticellarius has two or more of such projections; and (ii) Z. subseriatus has 4–9 undivided, superposed cells in the base of the receptacle, whereas Z. vorticellarius has only one undivided cell. After the examination of the type slide, the study of many slides (including the type) of Z. vorticellarius and review of the literature, I have come to no conclusion concerning the distinctiveness of the two species. It is rather evident that the number of buffer projections is a variable feature, and in Z. vorticellarius thalli occur with only one projection (FIG. 59). The presence of several undivided cells at the base of the thallus seems to be the best character to distinguish these species (FIGS. 57, 61).
Zodiomyces vorticellarius Thaxt., Mem. Am. Acad. Arts 12:371. 1896. FIGS. 58, 59, 65–74
Known distribution and hosts. –
Zodiomyces vorticellarius was described on several species of Cymbiodyta (as Hydrocombus) from USA (Thaxter 1896). It subsequently has been reported on undetermined Hydrophilidae from USA, Cuba, Trinidad, Venezuela and Argentina, on Hydrobiomorpha from Argentina and Cameroon (as Hydrophilus and Neohydrophilus wehnkei P., respectively), on Sternolophus angolensis Erichson from West Africa (Thaxter 1931), on Enochrus sp. (as Philhydrus) from France (Picard 1913), on Helochares lividus (Forster) from Poland (Siemaszko and Siemaszko 1928), from Germany (Scheloske and Matthes 1966), and Italy (Rossi and Cesari Rossi 1980), on Helochares lentus Sharp from Korea (Lee and Choi 1992), and on H. obscurus (O.F. Müller) (Majewski 1994) from Poland. Here, it is reported for the first time from Mexico and Spain. Any doubtful or corrected record from Z. subseriatus must be added (see above, under Z. subseriatus section). All hosts belong to the Hydrophilidae in the Coleoptera.
Specimens examined. – FRANCE. CORSICA: Mignataja, on Helochares lividus (Forster), 28 Jun 1989, C. Hernando, BCB-SS E264. Saleccia, Santo Pietro di Tenda, on H. lividus, 22 Jun 1989, C. Hernando, BCB-SS E268. Sarténe, Fiume Rizzanése, on H. lividus, 30 Jun 1989, C. Hernando, BCB-SS E267. MEXICO. BAJA CALIFORNIA: Hamilton Ranch, Río Santo Domingo, on Helochares sp., 22 Apr 1963, R.K. Benjamin, BCB-SS E256. SPAIN. ALBACETE: Robledo, El Arquillo, on H. lividus, 1 Mar 1997 (BCB-SS2052), 22 Apr 1997 (BCB-SS2046), I. Ribera. ÁVILA: Navalmoral, on H. lividus, 13 Nov 1978, I. Ribera, BCB-SS1958. BADAJOZ: No-gales, Rivera de Nogales, on H. lividus, 23 Apr 1992, R.M. Cros, BCB-SS1465. BALEARIC ISLANDS: Menorca, Alaior, Son Bou, on H. lividus, 16 Nov 1990, S. Santamaria, BCB-SS1007. BARCELONA: Castellet i La Gornal, Foix damp, on H. lividus, 23 Nov 1985, I. Ribera, BCB-SS1985a–1985b. Gualba, Gualba stream, on H. lividus, 12 Apr 1993, S. Santamaria, BCB-SS1641. Mura, Nespres stream, on H. lividus, 3 Jun 1993, S. Santamaria, BCB-SS1675. GIRONA: Capmany, Torrelles stream, on Enochrus sp. and H. lividus, 9 Jul 1994, I. Ribera, BCB-SS2213–2214. La Jonquera, Canadal ponds, on H. lividus, 25 Nov 1990 (BCB-SS1983), 12 Jun 1994 (BCB-SS2221), I. Ribera. Masarac, Vilarnadal, Llobregat d’Empordà river, on H. lividus, 9 Jun 1998, I. Ribera, BCB-SS2066. GUADALAJARA: El Pobo de Dueñas, on Helochares sp., 20 May 1990, C. Hernando, BCB-SS1784. HUEL-VA: Almonte, on Helochares sp., 16 Aug 1985, C. Hernando, BCB-SS1798, 1807. Laguna de la Mujer, on Helochares sp., 18 Aug. 1985, C. Hernando, BCB-SS1786. HUESCA: Barbastro, Cregenzán, on H. lividus, 12 Aug 1990, I. Ribera, BCB-SS1957. Candasnos, on H. lividus, 5 Mar 1994, I. Ribera, BCB-SS2081. Ontiñena, Hospital gully, on H. lividus, 9 Apr 1995, I. Ribera, BCB-SS2091. Sariñena, Pallaruelo de Monegros, Lafarda gully, on Enochrus sp., 5 Mar 1994, I. Ribera, BCB-SS1965. Villanueva de Sigena, Hospital gully, on Enochrus sp., 9 Apr 1995, I. Ribera, BCB-SS1966. LLEI-DA: Llastarri, on Helochares sp., 7 Apr 1984, C. Hernando, BCB-SS1795. Oliana, Segre river, on H. lividus, 26 May 1990, I. Ribera, BCB-SS1960. Pinell de Solsonès, Madrona, on H. lividus, 23 Apr 1990, I. Ribera, BCB-SS1984. Sanaüja, on H. lividus, 23 Jun 1990, I. Ribera, BCB-SS1959. SORIA: Ucero, Cañp ón río Lobos, on Helochares sp., 14 Jun 1990, C. Hernando, BCB-SS1804. TARRAGONA: Amposta, Delta de l’Ebre, on H. lividus, 24 Jun 1995, I. Ribera, BCB-SS1946. Santa Perpètua de Gaià, Gaià river, on H. lividus, 2 Jul 1994, I. Ribera, BCB-SS2156. TERUEL: Alcañiz, on Helochares sp., Apr 1984, C. Hernando, BCB-SS1790. Tornos, canal lagunica, on H. lividus, 23 Jul 1994, I. Ribera, BCB-SS2133. ZA-RAGOZA: Berrueco, on H. lividus, 24 Jul 1994, I. Ribera, BCB-SS2129. Monegrillo, San Benito pool, on H. lividus, 25 Jun 1994, I. Ribera, BCB-SS2152. USA. CONNECTICUT: West Haven, on Hydrocombus lacustris LeConte, May 1890, R. Thaxter, FH7117 (HOLOTYPE).
Remarks. –
This is the type species of the genus, and it was exhaustively studied by Thaxter (1896). Its pseudoparenchymatous, large thallus is variable in size and is symmetric to asymmetric. The buffer projections also are variable in number (FIGS. 58, 59), correlated with the position on the host surface.
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KEY TO THE SPECIES AND GENERA OF ZODIOMYCETOIDEAE |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSTAXONOMYKEY TO THE SPECIES...DISCUSSIONLITERATURE CITED |
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DISCUSSION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSTAXONOMYKEY TO THE SPECIES...DISCUSSIONLITERATURE CITED |
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). A similar large thallus was known in only one another genus, Euzodiomyces, which Thaxter (1908) placed in Zodiomyceteae, together with Kainomyces. However, Benjamin and Shanor (1951) demonstrated many important differences, including antheridial and perithecial structure and development. Columnomyces ptomaphagi R.K. Benj. (Benjamin 1955) and the three known species of Kainomyces (Thaxter 1908, 1931; Terada 1978) have also a more or less large receptacle, showing both longitudinal and intercalary transverse divisions in their receptacles, but there are many differences and these genera are unrelated (Tavares 1985). Now the description of C. tenellus and S. peritheciiformis calls for a reassessment of the use of characters in addition to the large receptacle to distinguish the Zodiomycetoideae. Another distinctive feature, the buffer outgrowths, are present in several genera of Laboulbeniales, were so named by Thaxter (1931:331–332) presumably because of their function of maintaining the position of the thalli during the movements of the host. Unicellular buffer organs are present in Hydrophilomyces and Rhizopodomyces, whereas those of Scelephoromyces and Osoriomyces are uniseriate and multicellular; multiseriate buffer organs are present only in Zodiomyces. These buffer organs are lacking in the two new genera.
Perithecial features, which are important in Laboulbeniales systematics, indicate that the two new genera described here are closely related to Zodiomyces, belonging in the same group, currently the Zodiomycetoideae. As in some other genera, the perithecia in Capillistichus, Scepastocarpus and Zodiomyces, have indistinct cell walls. The failure to develop distinct cell walls is a highly modified characteristic and often is accompanied by a reduction in the number of cells, according to Tavares (1985:69). However, Capillistichus and Zodiomyces may be considered primitive because each of the four vertical rows of wall cells consists of many (about 6–8) cells approximately equal in height (FIGS. 9, 10, 69, 71). These cells usually may be seen only in young perithecia except in C. tenellus where some wall cells may be more or less clearly seen even in mature perithecia (FIGS. 40, 41, 46).
Perithecium development in all species studied is similar, as can be observed in figures representing young stages (FIGS. 7, 8, 29–34, 53–55, 62, 63, 65–68). Zodiomyces vorticellarius, Z. subseriatus, and S. peritheciiformis show a similar curious outgrowth arising from one cell below the perithecium (FIGS. 22–24, 47, 48, 64, 71, 74). The significance of this outgrowth is unknown. Capillistichus tenellus lacks this outgrowth, but in young stages a similar outgrowth is seen above cell VII (FIGS. 11, 30–32), but later this prolongation breaks off (FIGS. 7, 8).
The perithecial apices of Z. vorticellarius and C. tenellus are similar in having four thin, ligula-like projections (FIGS. 12, 15, 41, 46). All species of Zodiomycetoideae (Z. odae is excluded from this discussion because no perithecial characteristics were included in the original description and those of the paratype could not be seen clearly) show four appendages arising from the perithecial walls. The significance of these appendages is unknown. They begin to develop when perithecia are slightly immature (FIGS.13, 23, 24, 39, 48). Two appendages develop from dorsal wall cell rows, and the other two from ventral rows. The two appendages originating from ventral rows are lower in position. The two pairs of appendages may be different, especially in Z. vorticellarius and Z. subseriatus where the two dorsal appendages are upwardly orientated, curved, or even falcate, and broader than ventral appendages, which are longer as well (FIG. 74). In C. tenellus and S. peritheciiformis no significant differences are noticeable between the two pairs of appendages. Those wall cells producing these appendages may be distinguished earlier in C. tenellus and Z. vorticellarius, where they may appear slightly inflated (FIGS. 37–39, 71).
Receptacular appendages are abundant in all species of Zodiomycetoideae. They arise close to the perithecia and from receptacular cells. Among the very abundant sterile appendages it might be assumed that appendages bearing antheridia must be present. The existence of trichogynes suggests fertilization, and the direction of their growth indicates where the antheridia may be found. In three of the five species in Zodiomycetoideae, trichogynes curve downward (FIGS. 7, 11, 33, 62, 66), in S. peritheciiformis they are long and more or less erect (FIGS. 53–55), whereas in Z. odae, there is no information about them. Additional information about antheridia in the genus Zodiomyces has been published by Tavares (1985). Thaxter (1896) defined and drew what he called exogenous antheridia. Tavares (personal communication) re-examined Thaxter’s slides to search for antheridia and found only one pair in situ; possible detached spermatia could not be positively identified as such. I have examined the type of Z. vorticellarius without seeing any antheridia. According to Tavares (personal communication), antheridia can be seen clearly only in well-stained specimens, acetocarmine being the best stain (Tavares 1985: Pl. 25a). The spermatia are borne on short stalks, hidden among short appendages and very young perithecia clustered at the bases of the larger perithecia, and there seem to be very few of them (Tavares pers comm). Unfortunately, I have not seen any evidence of antheridia or spermatia among the many slides of the species of Zodiomyces, Capillistichus or Scepastocarpus studied.
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ACKNOWLEDGMENTS |
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FOOTNOTES |
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E-mail: sergi.santamaria{at}uab.es
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LITERATURE CITED |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSTAXONOMYKEY TO THE SPECIES...DISCUSSIONLITERATURE CITED |
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———. 1971. Introduction and supplement to Roland Thaxter’s contribution towards a monograph of the Laboulbeniaceae. Biblioth Mycol 30:1–155.
———, Shanor L. 1951. Morphology of immature stages of Euzodiomyces lathrobii Thaxter and the taxonomic position of the genus Euzodiomyces. Amer J Bot 38:555–560.
Lee YB, Choi DS. 1992. Studies on Taxonomy and Distribution of the Laboulbeniales collected in Korea (species from Cheonham Province). Korean J Mycol 20:183–194.
Majewski T. 1994. The Laboulbeniales of Poland. Polish Bot Stud 7:1–466.
———.1999. New and rare Laboulbeniales (Ascomycetes) from the Bialowieza Forest (NE Poland). Acta Mycol 34:7–39.
———, Sugiyama K. 1989. Some Laboulbeniales (Ascomycotina) collected in Japan, IV. Additional species on coleopterous insects from Iriomote Island. Trans Mycol Soc Japan 30:77–88.
Nannizzi A. 1934. Repertorio Sistematico dei Miceti dell’Uomo e degli Animali. Vol. IV. In: Pollacci G, ed. Trattato di Micopatologia Umana. Siena, Italy: S. A. Poligrafica Meini. p 1–557.
Picard F. 1913. Contribution à l’étude des Laboulbéniacées d’Europe et du nord de l’Afrique. Bull Soc Mycol France 29:503–571. Pls. XXIX–XXXII.
Rossi W. 1994. A new contribution to the knowledge of the Laboulbeniales (Ascomycetes) from Sierra Leone. Accad Naz Lincei Quaderni 267:5–17.
———, Cesari Rossi G. 1980. Su alcune Laboulbeniali (Ascomycetes) parassite di insetti acquatici italiani. Rivista Idrobiol 19:147–154.
Santamaria S. 2001. New and interesting Laboulbeniales (Fungi, Ascomycota) from Spain, IV. Nova Hedwigia 72:375–389.
Scheloske HW, Matthes D. 1966. Ein parasitischer Pilz. Mikrokosmos 55:314–315.
Siemaszko J, Siemaszko W. 1928. Owadorosty polskie i palearktyczne. Polsk Pismo Entomol 6:188–211.
Sugiyama K. 1981. Notes on Laboulbeniomycetes of Formosa III. Trans Mycol Soc Japan 22:311–319.
———, Majewski T. 1985. The Laboulbeniomycetes (Ascomycotina) of Peninsular Malaysia. II. Trans Mycol Soc Japan 26:449–462.
———, Phanichapol D. 1984. Laboulbeniomycetes (Ascomycotina) in Thailand, I. Nat Hist Bull Siam Soc 32:47–88.
Tavares II. 1985. Laboulbeniales (Fungi, Ascomycetes). Mycol Mem 9:1–627.
Terada K. 1978. Additions to the Laboulbeniales of Taiwan, with descriptions of two new species. Trans Mycol Soc Japan 19:55–64.
Thaxter R. 1891. Supplementary note on North American Laboulbeniaceae. Proc Am Acad Arts 25:261–270.
———.1896. Contribution towards a monograph of the Laboulbeniaceae. Mem Am Acad Arts 12:187–429. Pls. I–XXVI..
———.1908. Contribution toward a monograph of the Laboulbeniaceae. Part II. Mem Am Acad Arts 13:217–469. Pls. XXVIII–LXXI.
———.1931. Contribution towards a monograph of the Laboulbeniaceae. Part V. Mem Am Acad Arts 16:1–435. Pls. I–LX.
Weir A. 1993. Rare and interesting Laboulbeniales on Sudanese Coleoptera. Mycol Res 97:509–512.
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