R E I N W A R D T I A Published by Herbarium Bogoriense, Bogor, Indonesia Volume 7, Part 5, pp. 561 — 578 (1969) SARAWAKUS LLOYD, A GENUS OF THE PYRENOMYCETE FAMILY HYPOCREACEAE MIEN A. RIFAI * SUMMARY The scope of the monotypic genus Sarawakus is enlarged to enable it to accommodate the newly described species Sarawakus succius Rifai. A complete taxonomic revision of the genus is presented, in which it is shown that Sarawakus belongs to the Hypocreaceae and not to the Xyla- riaceae as some authors have suggested. Hypocrea gelcutinosa (Tode ex Fr.) Fr. subsp. oligotheca Penz. & Sacc. is accorded specific status as Hypo- crea oligotheca (Penz. & Sacc.) Rifai and Phaeocreopsis pezizaeformds Boedijn is transferred to Hypocreopsis. INTRODUCTION The discovery of an undescribed species of Sarawakus Lloyd has made it necessary to propose an emendation to the scope of this so far monotypic genus. This is mainly due to the fact that the characters of the new species deviate in such a way from the type species that the need for a new generic delimitation is inevitable: it is reflected by the fact that it has been found justifiable to classify each species in a subgenus of its own. The discovery of this new species has also induced me to restudy the taxonomic position of Sarawakus. As is well known the type species of this genus, Sarawakus lycogaloides (Berk. & Br.) Lloyd, was twice described as new and had been referred to five genera — i.e. Hypoxylom, Bull, ex Fr., Hypocrea Fr., Sarcoxylon Cooke, Penzigia Sacc. apud Sacc. & Paolet and Clintoniella (Sacc.) Rehm — which in, turn are usually distributed among two families (the Xylariaceae and the Hypocreaceae), which some mycologists placed in one order (the Sphaeriales) while others would classify them in two orders (the Sphaeriales and the Hypocreales). Much of our understanding of Sarawakus comes from the extensive study of Boedijn (1934), who tentatively considered this genus as a transitional form between the xylariaceous and the hypocreaceous fungi. In contrast * Herbarium Bogoriense, Bogor, Indonesia. — 561 — 562 R E 1 N W A R D T I A [VOL. 7 von Arx & Miiller (1954) definitely included the genus in the Xylariaceae, whereas Petch & Bisby (1950) and Ainsworth (1961) listed it as a member of the Hypocreales. More recently Boedijn (1964) stated that Sarawakus " is most probably related to Xylariaceae " It will be shown below that the place of Sarawakus should be amongst the hypocreaceous alliance. THE TAXONOMY OF SARAWAKUS SARAWAKUS Lloyd emend. Rifai Sarawakus Lloyd, Mycol. Writ. 7(71): 1258. 1924; Boedijn in Bull. Jard. bot. Buitenz. Ill, 13: 263. 1934; von Arx & Miiller in Beitr. Kryptogamenfl. Schweiz 11 (1) : 340. 1954; Ainsworth in Ainsworth & Bisby's Diet. Fung. : 359. 1961. Stromata superficial, gregariously seated on an extensive and compact tissue-like subiculum or the latter very poorly developed to almost absent, subglobose, globose-depressed or sometimes pulvinate, sessile or contracted below into a short and broad stalk-like base, smooth surfaced, yellow or brown coloured but at maturity dotted with darker and ultimately greenish ostioles, with fleshy to firm fleshy or fleshy corky consistency. Stromal tissue indistinctly prosenchymatous but often almost simulating a pseudo- parenchymatous tissue and consisting of hyaline or subhyaline to yellowish, subglobose, angular, polygonal elongated or lobsd celled hyphae interspaced by more distinct filamentous hyphal elements, surrounded on the outside by a distinct, darker-coloured cortex layer made up of smaller, sometimes more isodiametric and much thicker-walled cells. Perithecia one-layered, completely immersed and confined to the upper part of the stroma, ovoidal or subglobose, their yellowish walls quite distinct and composed of a few layers of slightly flattened angular cells, provided with slightly prominent ostioles lined with a dense layer of periphyses. Asci eight-spored, cylindrical, thin-walled, unitunicate and having a simple apical apparatus with pores not turning blue in Melzer's reagent. Asco'spores uniseriate, one-celled, ellipsoidal to rarely ovoidal, at first hyaline but soon turning dark green or olive-green, becoming brown in preserved material, without germ-slit or germ-pore, but with surface studded with numerous large tubercles. Pseudoparaphyses present at young stage, partly or wholly deli- quescing at maturity. HABITAT: on bark of living trees and culm sheaths of bamboo shoots. TYPE SPECIES: Hypoxylon lycogaloid.es Berk. & Br. SCOPE AND DISTRIBUTION: two species from Ceylon, Borneo and Java. The generic name Sarawakus was derived from the geographical name Sarawak (Malaysian Borneo). The latter is the type locality of Hypocrea rhytidospora Ces., which is a later synonym of Hypoxylon lycogaloides; the latter species was originally described from Ceylon. For the purpose of euphony and to comply with the Recommendation 75A of the Inter- 1969] RIFAI: The pyrenomyeete genus Sarawakus 563 national Rules of Botanical Nomenclature I take this opportunity to desig- nate the gender of Sarawakus as masculine. In proposing this genus Lloyd (1924) did not give a formal generic diagnosis or description, so that for about ten years its existence was ignored (Clements & Shear, 1931). It was left to Boedijn (1934) to for- mulate the concept of the genus, in spite of Lloyd's (1924) expressed reluctance for the need of creating this genus. The characters stressed by Boedijn (1934, 1964) in delimiting Sara- wakus, and later in distinguishing this genus from Thuemenella Penz. & Sacc. emend. Boedijn, are the more or less corky stromata which " originate in large numbers from extensive subiculum. The stromal tissue consists of cells with thickened cell-walls. The cortical layer is distinct and is made up of very thick-walled cells of which the lumina are nearly obliterated. The dark spores are ellipsoidal " (Boedijn, 1964: 2). The poorly developed subiculum of the new species is markedly different from the extensive and tissue-like subiculum of Sarawakus lycogaloides, and if compared with the latter species its stromal tissue and cortex layer have thinner-walled cells so that on the whole its stromata have also a softer consistency. The stromata of Thuemenella britannica Rifai & Webster (1965) have even a much softer consistency than those of the new species because their tissue is also made up of more delicate cells. Although in this respect the new species would appear to occupy an intermediate position between Sarawakus and Thuemenella, the idea to merge the two genera is unwarranted because of the closer relationship between the new species and Sarawakus lycogaloides. Nevertheless it is obvious that the inclusion of the new species in Sarawakus will bring the latter to a position close to Thuemenella, and invalidates much of the taxonomic evidence enu- merated by Boedijn (1964) to distinguish the two genera. The separation of Sarawakus and Thuemenella based on the last character mentioned by Boedijn — that of the ascospore shape — is fully justified, because the genera of the hypocreoid fungi such as Hypocrea, Hypocreopsis Karst., Thuemenella, Thyronectria Sacc, Calonectria de Not., Nectria Fr., Pseudo- nectria Seaver and others are often distinguished from each other solely by their ascospore characters. The ellipsoidal and tuberculate ascospores of the new species make it a stranger amongst species of Thuemenella. The ascospore shapes of the two genera indicate further that Thuemenella is closer to Hypocrea than Sarawakus is: the globose, subglobose or elon- gated ellipsoidal, smooth-walled or minutely echinulate or warted asco- spores of the former two genera have broadly rounded to almost truncated 564 [VOL. 7 ends so that in optical sectiori they often appear subcuboidal, subangular to almost oblong (compare the descriptions and illustrations of ascospores of various species of Thuemenella and Hypocrea given by Seaver, 1910; Boedijn, 1964; Webster, 1964; Rifai & Webster, 1965, 1966, 1966a). The creation of another genus based on this new species and distin- guished from Sarawakus by the absence of a distinct subiculum and by the softer consistency would seem to be unwarranted, because by them- selves these two characters do not appear to be of diagnostic value at the generic level. In other genera of Hypocreaceae one finds that Podostroma Karst., for example, contains species with stromata having a fleshy to horny consistency (Boedijn, 1934a, 1938), whereas species of Hypocrea may vary from the soft fleshed Hypocrea gelatinosa (Tode ex Fr.) Fr. to the firm fleshed Hypocrea aureo-viridis Plowr. & Cooke apud Phill. & Plowr. (Webster, 1964; Rifai & Webster, 1966). In the latter genus the subiculum may be completely absent or variously developed but it must be admitted that as far as I am aware none of its species has developed a subiculum as extensive and distinctive as in Sarawakus lycogaloides. The foregoing considerations indicate that the new generic delimita- tion proposed above is the best course to accommodate satisfactorily this somewhat aberrant new species. On the other hand, to draw attention to the "remoteness" of its relationship to Sarawakus lycogaloides, and because of its outstanding morphological deviation from the latter, it is better to divide the genus Sarawakus into two subgenera, Sarawakus and Thue- menelladelphus. K E Y T O S P E C I E S O F S A R A W A K U S la. Ascospores small to medium sized, subglobose, elongate or short subcylindrical, often subangular, their ends broadly rounded to almost truncats; epispore smooth, minutely echinulate to warted Thuemenella Penz. & Sacc. emend. Boedijn b. Ascospores medium to large sized, ellipsoidal, their ends neither broadly rounded nor truncate; epispore tuberculate 2 2a. Subiculum large and extensive, tissue-like; stromata yellowish, with a fleshy corky consistency, 3 — 10 mm diameter; perithecia 370 — 450 x 270 — 350µ,; on bark of living trees (sufagem. Sarawakus) Sarawakus lycogaloides (Berk. & Br.) Lloyd b. Subiculum scanty to almost absent; stromata brownish, fleshy to firm fleshy, less than 5 mm diameter; perithecia 190—'250 x 120 —180 µ; on culm sheaths of bamboo shoots (subgen. Thuemenelladelphus) . . . Sarawakus suocisns Rifai 1969] RIFAI: The pyrenomycete genus Sarawakus 565 Subgen. Sarawakus Stromata suberosa, gregaria e subicula magna et compacta oriunda. — Typus: Sarawakus lycogaloides (Berk. & Br.) Lloyd. Stromata fleshy corky in consistency and gregariously seated on an extensive, compact and tissue-like subiculum. SARAWAKUS LYCOGALOIDES (Berk. & Br.) Lloyd. — Fig. 1 — 2a Hypoxylon lycogaloides Berk. & Br. in J. Linn. Soc., Bat. 14: 120. 1873; Sacc, Syll. Fung. 1: 355. 1882. — Sarcoxyton lycogaloides (Berk. & Br.) Cooke in Grevillea 12: 50. 1883; Patch in Ann. roy. Bot. Gard., Peradeniya 8: 143. 1924. — Penzigia lycoga- loides (Berk. & Br.) Sacc, Syll. Fung. 9: 569. 1891; Lindau in Engler & Prantl, Nat. Pflanzenfam. I, 1: 491. 1897. — Sarawakus lycogaloides (Berk. & Br.) Lloyd, Mycol. Writ. 7(71): 1258. 1924; Boedijn in Bull. Jard. bot. Buitenz. Ill, 13: 264. 1934; Petch & Bisby, Fung. Ceylon: 31. 1950; von Arx & Muller in Beitr. Kryptogamenfl. Schweiz 11 (1) : 340. 1954. Hypocrea rhytidospora Ces. in Atti Accad. Sci. fis. mat. Napoli 8: 14. 1878; Sacc, Syll. Fung. 2: 532. 1883; van Overeem & van. Overeem — de Haas in Bull. Jard. bot. Buitenz. Ill, 4: 27. 1922. — Clintoniella rhytidospora. (Ces.) Sacc. & Syd. apud Sacc, Syll. Fung. 16: 588. 1902. Subicula conspicuous, large and extensive, each one harbouring numer- ous stromata; they are pale dirty yellow coloured and are composed of sep- tate, branched, thin-walled, hyaline to subhyaline, 3—7µ. diameter hyphae which are interwoven with each other to form a somewhat compact tissue- like structure, up to 0.5 mm thick near the stromata, thinning towards the somewhat byssoid, effused, uneven margin. Stromata gregariously seated on a common subiculum, at first appearing as small nipple-like or globose protrusions on the latter, ultimately mostly becoming subglobose depressed with a flattened smooth surface, 3—10 mm diameter and up to 4 mm high in the middle, almost sessile or often provided with a short stalk-like base less than 1 mm high by 2—4 mm wide; the stromata at first have a pale yellpw coloration, turning dark yellow with age, but soon their upper surfaces are dotted with darker ostioles which later on exude green ascospores so that at complete maturity the surface of the stromata become greenish ("asphodel green" of Ridgway according to Boedijn, 1934). Stromal tissue indistinctly prosenchymatous which often appears like a pseudoparenchymatous tissue, made up of polygonal, lobed or angular elongated thick-walled cells 6—-27 µ. diameter with cell-walls up to 2 µ . thick, interspaced irregularly with more distinct thread-like hyphal ele- ments ; towards the periphery these cells become smaller sized, thicker- walled (up to 5 µ thick) and often with their lumina almost obliterated, more distinctly prosenchymatous, darker-coloured and forming a distinct cortex layer 50—70µ. thick; on the whole the stromata have a fleshy corky consistency. Perithecia completely immersed immediately under the cortex 506 REINWARDTIA [VOL. 7 20 µ Fig. 1. Sarawakus lycogaloides: a, habit sketch (2.5 x ) ; b, median section through a stroma (7.5 x); c, ascospores (from Boedijn 617). 1969] RlPAl: The pyrenomycete genus Sarawakus 567 layer, in one layer and confined to the upperside of the stromata, mostly ovoid or sometimes subglobose, 370—450 x 270—350 µ., their densely peri- physate ostioles normally slightly protuding above the surface of the stroma; perithecial wall thin, consisting of a few layers of flattened angular cells, slightly darker coloured than the rest of the stromal tissue. Asci thin-walled, nearly cylindrical, slightly attenuate below into a short stipe, 165—190 x 10—-12 µ, 8-spored, apical apparatus simple, not turning blue in Melzer's reagent. Ascospores unicellular, uniseriate, sometimes obli- quely uniseriate, ellipsoidal or often obovoid-ellipsoidal, especially the lower ones, and rarely unequal sided; they measure 17.5—20.5 X 8—10 µ,, at first colourless soon turning pale green, then becoming beautiful green, at last dark olive-green, appearing greenish black in mass, whereas in preserved specimens these ascospores have sepia-brown coloration; at maturity the surface of these ascospores are studded with large, rounded and irregular tubercles. Pseudoparaphyses thread-like, colourless, partly deliquescing at maturity. HABITAT: on bark of living trees. DISTRIBUTION: Ceylon (type locality), Sarawak (Borneo) and Java. ILLUSTRATIONS: Berkely & Broomem J. Linn. Soc, Bot. 14: pi. 6, fig. 33. 1873; Cesati in Atti Accad. Sci. fis, mat. Napoli 8: pi. 3, fig. 1. 1878; Lloyd, Mycol. Writ. 7(71): fig. 2739. 1924; Boedijn in Bull. Jard. bot. Buitenz. Ill, 13: 265, fig. 1. 1934; von Arx & Miiller in Beitr. Kryptoga- menfl. Schweiz 11(1): 326, fig. 99 f. 1954. CEYLON. On bark, Central Province, December 1868 (K; type specimen of Hypoxy- lon lycogaloides Berk. & Br.) BORNEO. On bark, Sarawak, 1865, O. Beecari (K; isotype specimen of Hypocrea rhytidospora Ces.). JAVA. On bark, Tjibodas, trail to Mt. Gedeh, 27 November 1921, Docters van Leeuwen, Dakkus & Bruggeman (BO 4508) ; on the bark at the base of a living tree, Tjibodas, ca. 1500 m alt., April 1930, Boedijn 272 (BO 11513) ; ibid., Boedijn 617 (BO 11570) ; on the bark of a living tree, Puntjak Pass, near Telaga Warna, 29 May 1939, van Steenis 11241 (BO 17017) ; on the bark of a living tree, Telaga Warna, June 1939, van Steenis (BO 17059). Subgen. Thuemenelladelphus Rifai, subgen. nov. A Sarawakus subgen.. Sarawakus subiculo inconspicuo, stromatibus carnosis recedit. — Typus: Sarawakus succisus Rifai. Stromata have a fleshy to firm fleshy consistency, typically gregari- ously seated on poorly developed or inconspicuous subicula, otherwise similar to Sarawakus subgen. Sarawakus. Sarawakus succisus Rifai, spec. nov. — Fig. 2b — 3 Subiculum effusum, minutum, inconspicuum vel nullum, candidum ex hyphis septatis, ramosis, hyalinis, 3—7 µ crassis compositum. Stromata R E I N W A R D T I A [VOL. 7 Fig. 2. Stromal tissue of: a, Sarawakus lycogaloides (from Boedijn 617): b, Sarawakus succisus (from Rifai 254). RIFAI: The pyrenomycete genus Sarawakus 569 solitaria vel caespitosa, subglobosa, flavo-brunnea, carnosa, 0.3—3 mm crassa. Perithecia immersa, in area superiore limitata, subglobosa, bstio- lata, 190—250 X 120—180 µ,. Asci cylindracei, breve stipitati, octospori, 115—158 x 8.5—11 µ, pseudoparaphysati. Ascospori uniseriati, non septati, ellipsoidei vel obovato-ellipsoidei, tuberculati, virides vel olivaeeo-virides, sicci brunnei, 15.5—22.5 X 8.5—10 µ. Hab. in vaginis culmorum Dendrocalami gigantei, Hortus Botanicus Bogoriensis, Java, 12 Februari 1962, Rifai 254 (BO) typus est. This species is found on sheaths of young and growing bamboo shoots; it grows mostly on the lower (the first three) sheaths, but sometimes also on the higher sheaths; as is the case with Sarawakus lycogaloides, there is no indication that the present species caused any harmful effect to the living host plant. The subicula of Sarawakus succisus are mostly scanty and very poorly developed, forming small watery white to white patches around the stromata and are made up of thin-walled, colourless, septate, 3—7 µ. diameter hyphae; at maturity these subicula may be inconspicuous or entirely disappear. Stromata scattered to caespitose, subglobose, globose- depressed or rarely globose, sometimes lobed, mostly sessile at the con- tracted base, their surface convex or flattened but in the larger stromata they may be slightly irregularly depressed in the middle which makes them appear bumpy, whereas on the underside of detached stromata an inward depression can mostly be found; when, young the stromata at first white, soon becoming pale or dirty-yellow, but upon nearing maturity the surface turning yellowish or pale brown to brown and dotted with brown and almost prominent ostioles which at complete maturity appear greenish from the extruding asoospares, while the rest of the stromata remains yellowish brown. Stromal tissue indistinctly prosenchymatous to pseudo- parenchymatous, consisting of compactly arranged subglobose, subangular, polygonal or lobed cells 6—30 µ. diameter, interspersed with elongated and thread-like hyphal elements; a distinct cortex layer about 35 µ, thick, composed of smaller sized and slightly thicker-walled cells covers the stromal tissue; in. contrast with the latter, which is whitish to pale yellow, this cortex layer is usually dark-yellow or pale brownish yellow; the stroma has a fleshy to firm fleshy consistency. Perithecia, in one layer, confined to the upperside of the stroma and completely immersed, ovoidal, globose or globose-depressed and measure 190—250 x 120—180 µ or 190 µ. high by 250 µ. wide, provided with narrowly conical or subcylindrical and almost prominent ostioles lined by a dense layer of periphyses; perithecial wall thin, composed of a few layers of flattened cells up to 10 µ thick, with pale brownish yellow coloration. Asci subcylindrical, attenuate below into a very short stipe, thin-walled, 8-spored, 115—158 x 8.5—11 µ; apical apparatus simple and reacting negatively with Melzer's reagent. Ascospores uniseriate, usually obliquely uniseriate, unicellular, ellipsoidal to ovoidal, sometimes unequal sided, ornamented with irregularly rounded tubercles of various size, 15.5—22.5 x 8.5—10 µ, hyaline when young but soon 570 REI N W A R D T I A [VOL. 7 i Fig. 3. Sarawakus suceisus: a, habit sketch (2.5 x) ; b, median section through a stroma (15 x ) ; c, asci; d, ascospores (from Rifai 254). 1969] RIFAI: The pyrenomyceie genus Sarawakus 67$ becoming pale green and gradually turning to beautiful green, ultimately dark olive-green at maturity and appear greenish black in mass; in preser- ved (dried) specimens these ascospores are brown. Pseudoparaphyses pre- sent in young stage only, thread-like, colourless, completely deliquescing at maturity. HABITAT: on living shoot sheaths of bamboos. DISTRIBUTION: West Java (known only from the type locality). JAVA. On the shoot sheath of bamboo, Bogor Botanic Garden, February 1924, Nongnong s. n. (BO 5534) ; on Dendrocalamus giganteus, Bogor Botanic Garden, 12 February 1962, Rifai 254 (BO; type specimen of Sarawakus succisus Rifai) ; ibid., 7 — 20 February 1992, Rifai 250, 251, 256, 261; on Gigantochloa, sp., Bogor Botanic Garden, 1 March 1962, Rifai 267 (all in BO). THE AFFINITY OF SARAWAKUS The true taxonomic affinity of Sarawakus with either the Xylariaceae or the Hypocreaceae can be determined best by closely comparing the anatomy and morphology of its species with members of the two respective families. For this purpose it is necessary first to review the delimitations of the two families according to ideas currently prevailing in taxonomic mycology. From the very beginning it has been, realized that the traditional characters — consistency and coloration — employed in distinguishing the Hypocreales and the Sphaeriales (to which the Xylariaceae belongs) are unsatisfactory, especially because of the presence of paradoxical genera, such as Sarawakus, Thuemenella, Sarcoxylon and others, which at first sight would appear to represent intermediate forms between the two orders. Therefore, the Hypocreales have often been merged with the Sphaeriales (Nannfeldt, 1932; von Arx & Muller, 1954; Dennis, 1960; Muller & von Arx, 1962). In recent years, however, it has been demonstrated that the soft consistency and the bright coloration of the stromata or perithecia of the hypocreaceous fungi are correlated with other characters of more significant diagnostic value such as types of conidial stages (mostly phialo- sporous), types of ascospores, the simple structure of the apical appara- tuses of their asci, the downward growing pseudoparaphyses and others. Consequently in many modern treatises the Hypocreales have also been maintained as a distinct order (Miller, 1949; Luttrell, 1951; Chadefaud, 1960; Martin, 1961; Alexopoulos, 1962; Hawker, 1966). Although further critical developmental studies of more species are desirable, this order is upheld here to include the Hypomycetaceae and those genera accepted by Boedijn (1964) in the Hypocreaceae and Nectriaceae, with a note that in 572 RE IN WAR DTI A [VOL. 7 this group of fungi I prefer to adopt a wider generic delimitation than that conceived by Boedijn. It follows that the Melanosporaceae, the Poly- stigmataceae and those genera classified — correctly, in my opinion — in the Clavicipitales, which in the past have been included in the Hypo- creales, should be excluded from the latter. The family Xylariaceae originally covered only the distinctly stro- matic and dark-coloured (carbonaceous) genera of Sphaeriales such as Hypoxylon Bull, ex Fr., Poronia Willd. ex Fr., Daldinia Ces. & de Not., Ustulina Tul., Xylarla- Hill, ex Grev. (= Xylosphaera Dum.), Nummularia Tul. and others, but later workers have correctly assigned to it non-stro- matic or indistinctly stromatic genera, such as Rosellinia Ces. & de Not. (which appears to represent an unnatural genus) and Anthostomella Sacc. as well. Other authors such as Munk (1957) would expand this family to include the allantosporous fungi, while von Arx & Miiller (1954) have transfered into it some unrelated genera now commonly placed in the Sordariaceae. Since these latter treatments will only make the Xylariaceae a heterogeneous assemblage, this family is interpreted here to embrace only those genera characterized by perithecia embedded in a stroma, deve- loped in host tissue beneath a clypeus or on a variously developed subi- culum, usually with a relatively tough consistency and having — at least in part — a carbonaceous pigmentation, with asci growing among persistent paraphyses and having intricately constructed apical apparatuses with amyloid pore-plugs, and with ascospores unicellular, smooth-walled, typically asymmetrical and provided with germ-slits and always dark coloured. This circumscription has been widely adopted by many recent authors (Dennis, 1980; Alexopoulos., 1962; Eriksson, 1968; Martin, 1967). With some reservations the Hypomycetaceae are considered here to constitute one family of the Hypocreales, chiefly on the basis of evidence of Hanlin's (1963) developmental study of Hypornyces lactifluorum (Schwein.) Tul. Furthermore, the Hypocreaceae and the Nectriaceae, namely the two families distinguished and circumscribed by Boedijn (1964), should also be included in this order and maintained as two distinct taxa; Munk (1957), Dennis (1960) and Alexopoulos (1962) have also kept the two families apart. Some well known genera that can be assigned to the Hypocreaceae in Boedijn's (1964) restricted sense are Hypocrea Fr. (inclusive of Creopus Link and Chromocrea Seaver), Podostroma Karst., Hypocreopsis Karst. (inclusive of Phaeocreopsis Sacc. & Syd. apud Lindau), Thuemenella Penz, RIFAI: The pyrenomycete genus Sarawakus 6.73 & Sacc. emend. Boedijn, and, as will be shown below, Sarawakus Lloyd as emended in the proceeding pages. Now we can compare the morphological and anatomical features of Sarawakus with those of the Hypocreaeeae and the Xylariaceae. Except for the consistency, the overall field characters of Sarawakus such as the habit or general appearance and coloration, are more hypoereaceous than xylariaceous. As has been pointed out above, however, in the classification of these groups of fungi the consistency of the stromata has only a minor or unimportant diagnostic value. This is largely due to the fact that many types of consistency may be found in the same genus, especially in the larger or stromatic ones such as Sarcoxylon, Xylaria, Hypocrea, Podo- stroma and others. The absence of a sharp distinction between the consis- tency of the Xylariaceae and the Hypocreaceae has made all attempts to classify Sarcoxylon and Sarawakus on the basis of this character alone a very difficult undertaking. It seems to me that in determining the taxo- nomic affinity of the genus Sarawakus an undue emphasis has been placed on this character by Boedijn (1934, 1964). Generally speaking members of the Xylariaceae can be characterized by their carbonaceous pigmentation; in most cases this coloration can be readily observed on the surface of the stromata but in some species such as Xylaria tabadna (Kickx) Berk, and Sarcoxylon compunctum. (Jungh.) Cooke it is necessary to section the stromata before this becomes visible. In contrast carbonaceous pigmentation never occurs- in members of the HypocreacDae. Berkeley & Broome's (1873) unaccountable statement that Sarawakus lycogaloides has black perithecia and reference to its resem- blance to a small-iscale Sarcoxylon compuncturn have led Cooke (1883), Saccardoi (1882, 1891) and Lindau (1897) to classify this species in the xylariaceous genera Sarcoxylon, Hypoxylon and Penzigia respectively. As Lloyd (1924) has correctly pointed out, however, there is nothing carbona- ceous about Sarawakus lycogaloides. Similarly Sarawakus succisus has no carbonaceous pigmentation, so that in this respect the genus Sarawakus appears to have a closer affinity with members of the Hypocreaceae than with those of the Xylariaceae. It has been noted above that the carbonaceous coloration can further be found in the ascospores of all members of the Xylariaceae, so that here the pigmentation has a significant diagnostic value at the family level. In delimiting the family Hypocreaceae it was stated by Boedijn (1964) that its ascospores are " colourless or green when fresh, brown or sepia in preserved material " As can be seen from the generic descrip- tion given above, the ascospore colour of Sarawakus is very similar to 574 R E l N W A R D T I A [ V O L . 7 that of the Hypocreaceae. It should be pointed out here that Seaver (1910), Clements & Shear (1931), Petch (1938), Dennis (1960) and Boedijn (1964) held the view that the ascospore coloration of the Hypocreaceae (namely, whether they are colourless or pigmented) was an important generic char- acter. Formerly it was shown, however, that in Hypocrea the segregation of genera based on this character alone was unnatural (Rifai & Webster, 1966) ; this view has already been adopted by Dingley (1952) and Miiller & von Arx (1962). Pemzig & Saecardo (1898, 1904) went a step further in believing that even at the specific level the diagnostic value of ascospore pigmentation was questionable and they included in the same species forms which had colourless and pigmented ascospores; in my opinion this is unjustifiable because there are ample morphological differences to show that Hypocrea oligotheca (Penz. & Sacc.) Rifai, stab. & comb. nov. [basio- nym: Hypocrea gelatinosa (Tode ex Fr.) Fr. subsp. oligotheca Penz. & Sacc. in Malpighia 11: 519. 1898; Penz. & Sacc, Icon. Fung. Jav.: 51, pi. 35, fig . 1. 1904. — Typus: in, culmis putridis, Tjibodas, Java, 4 Martii 1897, Penzig 128, BO 3429] deserves a specific status. Boedijn (1959, 1962) excluded Xylaria nigripes (Klotzsch) Sacc. and Xylaria spathulata Berk. & Br. from the Xylariaceae because, among other things, their ascospores have no germ-slit, a character generally found in the ascospores of members of this family. In agreement with ascospore characters of the Hypocreaceae, neither germ-slit nor germ-pore can be detected in the ascospores of the two species of Sarawakus described in the present paper. The ascospores of the Xylariaceae are invariably smooth-walled, whereas those of Sarawakus are ornamented with characteristic large tubercles. A similar type of epispore can be found in the bicellular asco- spores of Hypocreopsis pezizaeformis (Boedijn) Rifai, comb. nov. (basio- nym: Phaeocreopsis pezizaeformis Boedijn in Bull. Jard. bot. Buitenz. Ill, 16: 371, fig. 4. 1940 —.Typus: in ligno, Krakatau, Sumatra, 11 Julii 1929, Docters van Leeuwen 12658, BO 10470). In other members of the Hypo- creaceae the ascospore walls may vary from coarsely warted to minutely echinulated or perfectly smooth. Like those found in all other hypocreaceous fungi the asci of Sara- wakus are non-reactive to Melzer's reagent and have simple apical appa- ratuses, whereas those of the Xylariaceae are mostly iodine-positive and have complicated apical apparatuses. Furthermore, in agreement with the other species of the Hypocreaceae the pseudoparaphyses of Sarawakus appear to undergo partial or complete deliquescence at maturity. 1969] RIFAI: The pyrenomycete genus Sarawakus In describing Danish species of Xylaria it was stated by Munk (1957) that their stromal tissue is prosenchymatous, i.e. made up of textura intri- cata hyphae. During the course of this study I have examined many common north temperate and tropical species of Xylaria and I am able to corroborate Munk's observation; in many cases their long-cylindrical celled hyphae are so regularly orientated as to simulate even a textura porrecta tissue. I have further observed that this distinctly prosenchymatous tissue arrangement occurs in other species of xylariaceous fungi e.g. in species of Daldinia, Sarcoxylon, Penzigia, Ustulina, Camarops Karst. and in some members of Hypoxylon. Although many species of Hypocrea have prosen- chymatous stromal tissue, it is slightly differently constructed as com- pared with that of the Xylariaceae in that their hyphae are often made up of barrel-shaped, lobed to almost subglobose cells, strongly constricted at the septa and irregular in width [cf. Dingley, 1952, 1955; Webster, 1964; Rifai & Webster, 1966, 1966a; the schematic illustration of tissue structure of Hypocrea rufa (Pers. ex Fr.) Fr. given by Miiller & von Arx in Beitr. Kryptogamenfl. Schweiz 11(2) : 642 fig. 252. 1962 is erroneous]. In some other species of Hypocrea such as Hypocrea, gelatinosa (Webster, 1964), as well as in other hypocreaceous genera, such as Thuemenella (Boedijn, 1964; Rifai & Webster, 1965), the stromal tissue would appear to be more correctly designated as being of a type intermediate between prosenchy- matous and pseudoparenchymatous. Boedijn (1934) has indicated that the stromal tissue of Sarawakus lycogaloides is of this type, an. inter- pretation which I find to be correct and applicable to Sarawakus suceisus as well. It is evident that in their tissue structure species of Sarawakus show a closer similarity to the Hypocreaceae than to the Xylariaceae. It is hoped that ample fresh material will become available in the future so that a detailed study of the centrum development of Sarawakus can be undertaken. It is worth recording here, however, that in sections obtained by cutting preserved young material of Sarawakus suceisus on a freezing microtome the presence of the subhymenial pseudoparen- chyma * has been observed. This structure was first reported by Doguet (1957) for Hypocrea spinulosa Fuckel and its existence apparently is limi- ted to the hypocreaceous fungi. I have not been able to culture either of the two species of Sarawakus described above, so it is impossible to state whether their cultural beha- * Doguet (1957) termed this "plectenchym sous-hymenial", which, is translated here as subhymenial pseudoparenclhyma; following1 Ainsworth (1961) and Alexopoulos (1962) I prefer to reserve the term plectenchyma for all organized tissue of fungi. R E l N W A R D T I A [ V O L . 7 viour will provide additional evidence for determining the taxonomic posi- tion of this genus. 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