'-

Mycologia, 90(4), 1998, pp. 705-714. © 1998 by The New York Botanical Carden, Bronx, NY 10458-5126

The known morels in Mexico, a description of a new blushing species, Morchella rufobrunnea, and new data on M. guatemalensis

Gastón Guzmán1

Fidel Tapia

Instituto de Ecología, Apartado Postal 63, Xalapa, Veracruz 91000, Mexico

Abswact: A critica! analysis of the known species of Morchella from Mexico, including M. angusticeps, M. costata, M . elata, M. esculenta (M. canica; M. crassipes, M . rotunda), M . guatemalensis and M . umbrina, is presented. In addition, M. rufobrunnea is described as a new species from the State ofVeracruz. Morchella rufobrunnea belongs to the blushing species, a faurth group of species in the genus proposed by the au­thors. The blushing species also include M. guate­malensis and M. rigidoides, and ali three species are confined to the tropics or subtropics. Morchella rigi­doides is known only from New Guinea, and M . gua­temalensis only from Guatemala and Mexico. The new species differs in the farm and color of the ascomata, length of the alveolae, and the color of staining. A revision of the microscopic features of M. guatema­lensis is made based on recent collections from Mex­ico and Guatemala. After a comparison of the asci, ascospores and paraphyses of M. guatemalensis with those of M. rufobrunnea and other clase species, it is concluded that microscopic features, except the width of the paraphyses in sorne cases, are noi im­portant in the taxonomy of the genus. The farm and color of the ascomata, the position of the ribs, the length of the alveolae, and the staining are the most important taxonomic features in Morchella. A key to the seven known species of Morchella in Mexico is presented.

Key Words: Ascomycetes, Pezizales, blushing

INTRODUCTlON

The morels (Morchella Dill. ex Pers.) are very impor­tant edible mushrooms in Mexico, where they have been used traditionally far centuries. The Indian names far the morels include "colmenitas," "mazor­quitas," "elotitos" and "pancitas," and other com­mon names, based on the peculiar farm of the as-

Accepted far publication March 6, 1998 1 Email: guzmang@sun.ieco.conacyt.mx

705

comata, include "little beehives," "little tender corn ears," " little ear of green corn" and "little paunch," respectively (Guzmán, 1977, 1997; Herrera and Guz­mán, 1961; Herrera and Ulloa, 1990) . The commer­cial name far the more! is " morilla, " from the Span­ish word meaning beehive . Morels have a high eco­nomic value in Mexico due to export to Europe and the U.S. Exploitation of more Is, together with the de­struction of the farests where they grow, farced the Mexican Governmen t to consider the species of Mor­chella, together with other edible fungi, in the Red List of Organisms (Anonymous, 1994) .

Species concepts in Morchella vary widely, and sorne authors recognize 50 or more species, whereas others recognize only three good species (Gessner, 1995; Korf, 1973). Dennis (1968) noted that most modern authors regard ali the European species of Morchella as variants of M. esculenta Pers. : Fr. and M. elata Bull.: Fr. However, Dennis (1968) recognized three species from the British Islands, and Seaver (1961) six species from North America. Hawksworth et al. (1995) considered 28 species in the genus, and Jacquetant (1984) also distinguished 28 taxa, al­though many names recognized by him were invalid. The more recent studies recognize a fewer number of species in the genus (Singer and Harris, 1987). Bunyard et al. (1994), based on ribosomal DNA stud­ies of six species, recognized only two as valid species. Volk and Leonard (1989) andjung et al. (1993) con­sidered M. esculenta, M. crassipes (Vent.) Pers. and M. deliciosa Fr.: Fr. as conspecific. Gessner (1995) summarized infarmation on mating studies and iso­lation of chemical products and antigens far species delimitation in Morchella.

Denison (1963), Heim (1966a, b), Gómez (1971), Pfister (1974), Gamundí (1983), Guzmán et al. ( 1985), and Guzmán-Dávalos and Rodríguez-Alcantar (1993) are among the few studies facused directly or indirectly on Morchella in the tropics and subtropics. A study on cultures of Mexican morels (M. esculenta, M . canica Pers., M. crassipes and M . angusticeps Peck) was made by Rodríguez and Herrera (1962), and Aguilar (1973, 1979) studied submerged cultures of M . crassipes and M. elata.

In the present study, specimens of Morchella gath­ered at a subtropical site in the Ecological Park of the Ecology Institute at Xalapa and in other parts of

706 MYCOLOGIA

the Xalapa region (Mexico) , and all the collections of Morchella at XAL Herbarium were examined. An interesting Morchella is described herein as a new spe­cies, and new records of M. guatemalensis Guzmán, M. F. Torres & Logem. are reported.

MATERJALS ANO METHODS

Fleshy specimens of Morchella spp. were dried ap­prox. 1 h after collection. The drying apparatus con­sisted of an electric grid and natural aeration. Spec­imens dried in approx. 24 h. Sections of the ascom­ata, mainly from the ribs, were mounted in 5% aque­ous KOH, Melzer's reagent, cotton-blue in lactophenol or in Congo-Red, according to the mi­croscopic features observed. Measurements were made in young and mature specimens from rehy­drated specimens mounted in KOH. A culture of the holotype (IE-228) on malt agar (MA) and on potato dextrose agar (PDA) (both from Bioxon, Mexico City) was obtained from the context of one fresh specimen. This strain is maintained in test tubes on MA at 5 C, and transferred every six mo. All speci­mens studied are deposited in Herb. XAL, except where other herbaria are cited. The culture strain is deposited in the Strain Collection of the Ecological Institute at Xalapa. Colors in the description are ac­cording to Kornerup and Wanscher (1978).

RESUU"S

Morchella rufobrwmea Guzmán et Tapia, sp. nov. FIGS. 1-7

Ascomata (60- ) 90- 155 (-210) mm alta, conica vel sub­cylindrico--0voidea, hymenio griseo vel spadiceo flavido, cos­te longitudinales vel inaequalis albidae ve! griseae, maculis brunneis ve! aurantiaco-brunnescentibus vel ferrugineis. Stipes (20- ) 30-70 (-90) X 10-25 mm, albidus vel subgri­seus, basis venosus, fusco-griseus, maculatus similis hymen­io. Sporae in cumulo subaurantius vel flavidus subaurantius. Asci (280-) 300- 360 (-380) X 16--20 µm, operculatus, ten­uitunicatum, octosporus, inamyloideus. Sporae ( 19-) 20-24 (- 25.5) X (13- ) 14-16 (-17) µm maturae, (11-) 14.5- 19 (-20) X (8- ) 9-10 (-11) µm immaturae, hyalinae, ovo­ideae, inamyloideae. Paraphyses 90-184 X (8-) 10-18.5 µm [vel immaturae X (5-) 6--9 (-10.5) µm], hyalinae, basis sep-tatae.

HOLOTYPUS. Habitat terra arenosa, in sylvae subtropi­calis, in horto cum Quercus, Liquidambar; C/,ethra et Alnus, 1350 m altitudinis, Mexico, prope Veracruz, ad Xalapa, Guz­mán 31565 XAL.

Ascomata (60-) 90-155 (-210) mm high. Hymen­ophore (40- ) 60-85 (-120) X (20-) 30- 45 (-50) mm, conic or subconic to subcylindric-ovoid when mature, with longitudinal ribs and secondary transverse veins

between them, forming short or elongate alveolae ex­tended vertically, but in aging the vertical configu­ration of the ribs is somewhat lost and form a wrin­kled hymenophore (like that of M. esculenta); surface gray to grayish ( 4B2, 5B2-3, 5E2, 6D2), with whitish to grayish ribs (5Al, 5Bl) when young, to completely yellowish, brownish or brownish yellow ( 4A4-5, 4B5, 5C4, 5DA) when mature. Stipe (20- ) 30-70 (- 90) X

10- 25 mm, cylindric or more thick at the base, irreg­ularly wrinkled mainly toward the base, covered by minute dark granules toward the apex, surface whit­ish to cream (4A2-3, 5A4) or pale grayish (5B3, 5DA), darker (grayish brown, 5F5- 6) toward the base, yellowish (4A5) or dark grayish (4E3, 4F4) in old specimens. Blushing in irregular small spots, brown (6F6), brownish orange or pinkish red (4A8, 5A2-5, 5A8, 5B6- 8) to ferruginous (7C8) when in­jured or in maturing, both in hymenophore and stipe; sometimes in old specimens becoming almost completely reddish brown (12E6-12F8). In dry as­comata the color is pale brownish (5B4) with br9wn­ish pink (7B5, 7C5) or reddish brown (8F4-8) z~ees. Inner surfacewhitish, little orno blushing at the base. Basal mycelium a compact thin mass, whitish to pale yellowish, with sorne zones stained brownish orange (6B8). Spore print pale orange or yellowish orange (4A7, 5A7). Asci (280- ) 300- 360 (-380) X 16-20 µm, operculate, thin-walled to somewhat thick-walled (wall up to 1.5 µm thick), uniformly cylindric, but sorne with a sub-bulbous base, 8-spored, hyaline and inamyloid. Ascospores (19-) 20- 24 (- 25.5) X (13- ) 14-16 (-17) µm, but in immature ascomata (11-) 14.5-19 (-20) X (8-) 9- 10 (-11) µm, ovoid, thin­walled, hyaline, inamyloid. Paraphyses 90-184 X (8-) 10-18.5 µm, or X (5-) 6-9 (-10.5) µm when im­mature, hyaline, simple, septate at the base, consist­ing of one or two cells. Context (in the inner part of a rib) formed of intermixed hyphae, 3- 9 (- 13.5) µm wide, hyaline, thin-walled, cylindric to somewhat in­flated.

Cultures. The culture obtained from vegetative tis­sues grew slowly at 25 C on MA and PDA, producing a whitish to tan, sparse mycelium, covering the sur­face of the medium in 9-cm-diam Petri dishes in 10-12 d. Sclerotia and a conidial stage were absent. These results contrast with those of other studies on Morchella spp. reporting rapid growth in culture . For example, Stamets (1993) obtained a dense mycelium from M . angusticeps on malt yeast agar, oatmeal malt yeast agar and pota to dextrose yeast agar after 3- 7 d. A conidial stage and sclerotia were common in Sta­mets' experiments.

Specimens examined. MEXICO, VERACRUZ: Municipio Xalapa, old road Xalapa to Coatepec, km 2.5, inner way to

GUZMÁN AND TAPIA: MORELS IN MEXICO 707

Frcs. 1, 2. Morchella rufolYrunnea. 1 (above). Mature ascomata (note the stained spot on the stipe at the base of the third specim en, and in the upper part of the fourth specimen from the left) from Guzmán 31393. 2 (below). Immature ascomata in their natural h abitat from Guzmán 31394. X 0.75.

708 MYCOLOGIA

Frc. 3. Morchela rufobrunnea, dry immature ascomata (from the holotype). Scale bar 15 mm.

Instituto de Ecología, Ecological Park, near the crossing to DIF, 21 May 1996, M. Villegas; same date, G. Guzmán 31393, 31394; same locality, 5June 1996, G. Guzmán 31565 (holotype , derived culture strain IE-228, isotypes NY, IBUG) , 31566; same locality, 19June 1996, S. Chacón5066; same locality, 26June 1996, G. Guzmán 31605; same local­ity, 9 July 1996, G. Guzmán 31627; same locality, 12 July 1996, F Tapia 1389, 1390, 1391. Xalapa, garden in a pri­vate house , 13 Dec. 1994, S. Chacón 4871 (ali in XAL) .

Habitat and distribution. Gregarious, known only from the Xalapa region. In the type locality, M. ru­Jobrunnea was found at 1350 m altitude in a subtrop­ical (mesophytic) forest with Quercus, Liquidambar, Clethra, and Alnus in sandy soil on a recently dis­turbed roadside embankment with tree trunks. Fruit­ing in the type locality occurred between mid-May to mid:July in 1996, but was not observed in 1997. It is interesting to note that the type locality is in the Eco­logical Park at Xalapa zone, which has been explored mycologically for more than 10 yr wi th more than 200 species of fungi recorded (Chacón and Guzmán, 1995; Chacón et al., 1995) . However, this is the first record of a Morchella sp. in the zone.

DISCUSSION

This fungus is characterized by the form, color and blushing of the hymenophore. The blushing may be difficult to see in young specimens, also similar to M. guatemalensis (see as example, color plate 12, Guz­mán et al., 1985). For this reason it is important to study both immature and mature ascomata in collec­tions.

Morchella rufobrunnea differs from M . guatemalen­sis in the form and color of the ascoma, as well as in the type of blushing. Morchella guatemalensis has sub­cylindric-ovoid to conic ascomata up to 105 mm high, with a longitudinally ribbed hymenophore and elon­gate alveolae. The color ranges from yellow, honey color or yellowish orange (ferruginous when dry) but is never gray in any stage. The blushing is more in­tense to reddish vinaceous, both in spots or in all parts of the ascomata. Morchella guatemalensis and M. rufobrunnea also differ in the size of the paraphyses, smaller in the first (56-103 X 6.5-13 µm), as shown in TABLE l. Morchella herediana Gómez from Costa Rica (Gómez, 1971) is similar to M. guatemalensis in the form of the hymenophore, longitudinal ribs, elongate alveolae and width of the paraphyses (TABLE 1), but differs apparently in the absence of blushing. Morchella rigi,doides R. Heim and M. anteridiformis R. Heim were described from New Guinea and New Cal­edonia, respectively (Heim, 1966a, b), and the only difference between them seem to be the size of the ascomata (55 mm vs 110 mm) and the blushing stipe in the first. Both species have elongate alveolae and longitudinal ribs as in M. guatemalensis. Morchella ru­Jobrunnea differs from M. rigi,doides in the larger as­comata, color of the hymenophore (pale ochraceous to yellow without gray or grayish tones in young stages in Heim's species) , less elongated alveolae, and wider paraphyses (TABLE 1) . Heim's species is clase to M. guatemalensis but differs in the width of the paraphyses and less intense blushing.

Other species similar to M . rufobrunnea based on the descriptions and illustrations of Boudier (1905-1910), Marchand (1971 , 1973), Breitenbach and Kranzlin (1984) andjacquetant (1984) are discussed below, although many combinations and new taxa proposed by Jacquetant may be invalid. Morchella ela­ta var. purpurascens Krombh. ex Boud. in the sense of Boudier (1905-1910) has en tirely lilaceous pink or vinaceous ascomata on all the surfaces, both young and adult stages. According to Marchand ( 1973), this fungus has a pale carmine, lilaceous to violaceous hy­menophore anda white stipe.Jacquetant (1984) con­sidered this fungus as "M. purpurascens (Krombh.) Jacquet." with vinaceous brown or blackish ribs, gray­ish green to pink or vinaceous pink alveolae, and a

G UZMÁN AND T APIA: MüRELS IN MEXICO

7

, I , ,

1 ,

l 8

1 1 1 1 1 1 1 1 1 1 1 1

709

l

FIGS. 4-13. Microscopic features of M . rufobrunnea and M. guatemalensis. 4-7. M. rufobrunnea. 4. Ascospores. 5. Ascus. 6. Paraphyses in young stages. 7. Paraphyses in mature stages. 8- 13. M . guatemalensis. 8, 9, 12. Paraphyses. 1 O, 11. Ascospores. 13. Asci (4-6 from Guzmán 31393. 7 from the holotype. 8, 11 from Mollzr. 9, 10 from the topotype. 12, 13 from the isotype). Scale bar 4, 10, 12 = 16 µ,m . 6-9, 11 = 20 µ,m. 5, 13 = 40 µ,m .

TABLE l. Variation of the microscopic features of M. rufobrunnea and other species

Speciesª

M. rufobrunnea

(HOLOTYPE)

M. guatemalensis

1 2 3 4 5 6

M. herediana

7

M. rigidoides

8

Asci (µm)

(280-) 300-360 (-380) X 10-20

280-420 X 18-23 159-250 (-287) X 18-21.5 300-400 (-420) X 17.5- 26.5 (- 28) 197-366.5 X 18.5- 27.5 X (16-) 17.5-26.5 (-28) (280-) 300-390 (-410) X 17-28 (-30)

144-172 X 14.5- 20

240-250 X 18-20

Ascospores (µm)

(19-) 20-24 (-25.5) X (13-) 14-16 (-17) " (11- ) 14.5-19 (- 20) X (8-) 9- 10 (- ll )c

(17- ) 18- 22 X 11-13 (15- ) 16-21 X (9- ) 9.5- 13 (17.5-) 18.5-24 X (12-) 13-14.5 (14.5-) 16-23 X 9-15 (17-) 19-25 (- 25.5) X (12- ) 13-14.5 (-16) (17.5-) 18- 24 (-25) X (12- ) 13-14

17-18.5 X 8-9

18-20 X 10-12

Paraphyses (µm )

90- 184 X (8-) 10- 18.5 or X (5- ) 6-9 (-10.5) when immature

56-103 X 6-12 X 7-13 X 6.5-12 (-13)

X (9- ) 9.5-16 (-17.5) X (6.5-) 7-13 (-13.5)

144-172 X 9- 14

X 12-30

ª 1: Guzmán et al. (1985); 2: Guzmán-Dávalos & Rodríguez-Alcantar (1993) ; 3: isotype revised by the authors; 4: isotype revised by Guzmán-Dávalos & Rodríguez­Alcantar (1993); 5: Sommerkamp 227; 6: topotype; 7: Gómez (1971); 8: Heim (1966a, b).

"Mature e Immature

-1' ...... o

a:: rs o t"' o () :;

GUZMÁN ANO TAPIA: MORELS IN MEXICO 711

white-pink stipe. Boudier described the fungus with short alveolae , whereas Marchand andJacquetant de­scribed elongate alveolae. Jacquetant also recognized "M. purpurascens var. ionoviridis Jacquet." with vio­laceous ribs and greenish elongate alveolae, and ap­parently a white stipe. This author also described "M. purpurascens var. ionoviridis f. heteroparaphysa Jac­quet." with subglobose paraphysis apices as the only distinguishable feature. Morchella elata var. elata, ac­cording to Breitenbach and Kranzlin (1984), has a brownish yellow to brownish red hymenophore with blackish ribs and short alveolae and white stipe. Ac­cording to Boudier ( 1905- 191 O), this fungus is en­tirely brownish yellow, and in Gamundí ( 1975), the hymenophore is described as brown to blackish with elongate alveolae and white stipe. In Marchand ( 1971) this fungus is described with a grayish brown hymenophore, elongate alveolae and white stipe. Morchella rigida (Krombh.) Boud., according to Mar­chand (1973), has an alveolate hymenophorewithout longitudinal ribs, similar to M . esculenta, and also a whitish stipe, but with sorne inconspicuous brownish spots. According to Boudier (1905- 1910),Jacquetant (1984) and Breitenbach and Kranzlin (1984), this fungus also has a structure and form similar to that of M . esculenta, but with a stipe lacking spots. Heim ( l 966a) separated M. rigidoides from M. rigida based on the height of the ascomata and shallower alveolae.

It is important to observe that none of the species discussed above, except M. guatemalensis and M. rig­idoides, clearly exhibit the blushing feature. It seems that blushing is an important taxonomic feature, as observed in other mushrooms, e .g ., Amanita rubes­cens (Pers.: Fr.) Gray. The senior author has seen many fresh morels during his years of field work, and M . guatemalensis and M. rufobrunnea are the first species observed by him to exhibit blushing. We be­lieve that the blushing feature is different from the spots produced on sorne mushrooms as result of in­fection by bacteria or other organisms. However, it is possible that the spots represented in the iconogra­phy of M. rigida ofMarchand (1973) discussed above are due to this type of infection.

Based on the results of DNA studies, Bunyard et al. (1994) suggested that there are at least two taxo­nomic groups in Morchella: (i) the black morels rep­resented by M. angusticeps, M. elata and M . canica, and (ii) the yellow morels, with M. esculenta, M . cras­sipes and M. deliciosa. They suggested that these two groups represent only two species. Another group of Morchella species is what sorne authors call half-free morels, represented by M. semilibera (DC.: Fr.) Lév. (Gessner, 1995). Although Smith and Weber (1980) provided a description of M. semilibera, this speccies is in need of revision. It is possible that there is a

fourth taxonomic group of morels represented by the blushing species of the tropics and subtropics, including M. rufobrunnea, M. guatemalensis and M . rigidoides. Gilbert (1960), based on a study of sub­merged cultures, found that M. esculenta and M. can­ica have certain cultural characteristics in common that are distinct from those of M . angusticeps. Ower ( 1982), in a study of ascoma development in a more] he called M. esculenta, described cultures forming gray ascomata that turned yellow as they matured. However, the ascomata obtained by Ower exhibited conspicuous longitudinal ribs and deep, elongate al­veolae, features not typical of M. esculenta, but com­mon in M. costata Vent. Yoon et al. (1990) also re­ported that in M. esculenta the immature gray ascom­ata changed to a tan color as they matured. These observations show the complex and contradictory na­ture of the taxonomy in Morchella, particularly for the common species.

Microscopic features of the Morchella hymenium appear to be relatively homogeneous among taxa, as observed by the authors for the previously discussed species, and as noted by Wipf et al. ( 1997). It is dif­ficult to separate Morchella species based only on mi­croscopic features. However, Singer and Harris ( 1987) separated M. es cu lenta, M. hortensis Boud. and M. costata in their key mainly by the size of the spores. Among the more! species discussed above , in­cluding M . rufobrunnea, there are no important dif­ferences in the microscopic features of the hymeni­um except in the width and form of the paraphyses in sorne species. Examples ofvariation in microscopic features among species cited in the literature may be due to the use of immature specimens, or that the microscopic features were observed in different con­ditions (e.g., fresh or dried specimens), as noted by Baral (1992) in many ascomycetes. He stated that dif­ferences between diagnoses based on the morphol­ogy of microscopic elements from fresh , living spec­imens, and those based on dead, dry specimens, had implications that could lead to erroneous taxonomic conclusions. Morchella rufobrunnea, M . guatemalensis and M. herediana seem to differ from M. rigidoides in the width of the paraphyses observed in dry spec­imens (TABLE 1) . The authors studied also a collec­tion (at XAL) from Region ofTlaxco, Tlaxcala (Mex­ico), gathered in a Pinus-Abies forest, on 21 Sep. 1985 (l. González-Fuentes 1013) that it is similar to M. ru­

Jobrunnea in the ascoma, with a strong blushing on the stipe, but has paraphyses (12-) 13.5- 20 (- 24) µm wide. It is likely that this collection represents a new species, but the authors require additional collec­tions for further study.

It is concluded that the ascoma of species of Mor­chella is enormously variable in morphology, depend-

712 MYCOLOGIA

ing on the development, and possibly on weather and microniche conditions. Morchella rufobrunnea is sim­ilar to M. guatemalensis and other morels, but differs in the form and color of the ascoma, length of the alveolae, and the color of staining. We based the de­scription of the new species on two populations, pri­marily the population from the type locality where many ascomata were collected between May 21 and July 12, 1996 from an area approximately 6 m 2 • An interesting observation is that this species was gath­ered in heavily disturbed sites, as is common for many species of the genus (Ramsbottom, 1959; Ges­sner, 1995).

The known species of Morchella in Mexico. - Little is known about the diversity and taxonomy of Morchella in Mexico. Nieto-Roaro (1944), Rodríguez and Her­rera (1962) and Herrera and Ulloa (1990) discussed M. angusticeps, M. canica, M. crassipes, M. esculenta and M. rotunda Pers. Guzmán (1977) accepted all species except the latter, and also considered M. cos­tata and M. elata in his keys. Recently, Guzmán-Dáv­alos and Rodríguez-Alcantar (1993) recorded M. guatemalensis from Jalisco, and Guzmán et al. (un­publ.) reported M. umbrina Boud. from Veracruz. In consideration of the observations by Volk and Leon­ard (1989),Jung et al. (1993), and others, it is prob­able that M. canica, M. crassipes and M. rotunda are conspecific with M. esculenta. Hence, there are only seven species of Morchella, including M. rufobrunnea, recorded from Mexico. Of these species, only M. gua­temalensis and M. rufobrunnea belong to the blushing group proposed herein.

KEY FOR THE MEXICAN SPECIES OF MORCHELLA

la. Ascoma never blushing (but may blacken in age) 2

1 b. Ascoma blushing (may be difficult to discern in young ascomata) . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 2a. Hymenophore yellowish, blackish in old and

putrid ascomata . . . . . . . . . . . . . . . . . . . . . . . . 3 2b. Hymenophore gray to blackish . . . . . . . . . . . . 5

3a. With irregular ribs, forming short alveolae. Hymen­ophore globose, subcylindric or conic, sometimes thick footed . . . . . . . . . . . . . . . . . . . . . . . M . esculenta

3b. With longitudinal ribs . . . . . . . . . . . . . . . . . . . . . . . 4 4a. Transverse ribs conspicuous, forming short,

rounded and deeper alveolae ...... . M. costata 4b. Transverse ribs not common, with elongate al­

veolae . . . . . . . . . . . . . . . . . . . . . . . . . . . M . elata 5a. Hymenophore globose, with short, rounded alveo-

lae in irregular ribs . . . . . . . . . . . . . . . . . M. umbrina 5b. Hymenophore subcylindric or conic, with elongate

alveolae . . . . . . . . . . . . . . . . . . . . . . . . M. angusticeps 6a. Hymenophore yellowish orange or honey-col­

ored, both in young and mature stages, with longitudinal ribs and elongate alveolae. Blush-

ing in large spots dark vinaceous or reddish brown . . . . . . . . . . . . . . . . . . . . M. guatemalensis

6b. Hymenophore gray when young to brownish yel­low when mature, with longitudinal to irregular ribs and elongate to short alveolae. Blushing in irregular, small or occasionally large spots, brown or pinkish red to ferruginous . ........... . .. ............. ............ M . rufobrunnea

New data on M. guatemalensis.-This species was de­scribed from Guatemala, Department of Chimalten­ango, gathered in Dec. 1983 and in Oct. 1984 (Guz­mán et al., 1985). Recently it was reported from Mex­ico (State of Jalisco) by Guzmán-Dávalos and Rodrí­guez-Alcantar (1993). When revising the morels deposited at XAL Herbarium, the authors found new records of this fungus , five from Guatemala, and three from Mexico.

Specimens examined. GUATEMALA. ZACATEPEQUEZ: San Lucas, 10 Aug. 1985, A. Moller, San José Pinola, San Juan Mushball, 27 July 1986, G. Guzmán 29227; 27 July 1986, G. Guzmán 29231; 27 July 1986, G. Guzmán 29233; Guatemala City, 30 Aug. 1988, Y. Sommerkamp 227. MEXI­CO. VERACRUZ: old road Xalapa to Coatepec, km 6.5, El Atorón, 2 Nov. 1990, VM. Bandala 2004; Xalapa, S of the Unidad La Haya, 26 Jan . 1996, S. Chacón 4992; MICHOA­CÁN: Morelia to Ciudad Hidalgo, Mil Cumbres, near Placa Conmemorativa, 7 Jan. 1984, ].A. Pérez de la Rosa (IBUG 954); MORELOS: Valle de Tepeite, NW of Santa María, 13 Oct. 1984, R. Valenzuela 4275 (ENCB); Cuernavaca, U.A.E.M., University Campus, 25 Oct. 1981, L. López 272.

These materials compared well with the isotype of M. guatemalensis (XAL), except in sorne variation ob­served in the width of the paraphyses. Guzmán et al. (1985) described paraphyses 6-12 µm width; Guz­mán-Dávalos and Rodríguez-Alcantar (1993) 7-13 µm; in the isotype it was found 6.5- 12 (-13) µm; in the topotype (6.5-) 7-13 (- 13.5) µm, and in Somer­kamp 227, (9-) 9.5-16 (-17.5) µm (TABLE l) . In the Mexican materials examined, the variation was from 6-13.5 µm to (8-) 9-14.5 (- 16) µm. The observed variation of the width of the paraphyses appears to be related to the maturity of the ascoma. The greatest width was observed in mature species, as was also not­ed in M . rufobrunnea. The vegetation in the above localities, including that reported by Guzmán-Dávalos and Rodríguez-Alcantar (1993) , is tropical or sub­tropical with Quercus forests, mixed with Alnus and Arbustus ( Cupressus also grows in the type locality). The main distinguishing feature of M. guatemalensis is the blushing, the color of the hymenophore, and the longitudinal and elongate alveolae, as discussed above. Dimensions of the asci, ascospores and pa­raphyses of the isotype, shown in Frcs. 8-13, are pre­sented in TABLE I in comparison to those of other Morchella species reported in the literature.

GUZMÁN ANO TAPIA: MORELS IN MEXICO 713

ACKNOWLEDGMENTS

The authors are indebted to Prof. Richard P. Korf, at Cor­nell University, for comments, for providing bibliography and for critical revision of this paper. They also give thanks to Rosario Medel, at XAL, for her criticism to the manu­script, and Dulce Salmones and Rosalia Pérez-Merlo, at In­stituto de Ecología, for the development of the cultures. Laura Guzmán-Dávalos, at IBUG, provided herbarium ma­terial and bibliography. Margarita Villegas, at FCME, pro­vided fresh specimens of the fungus described here. Juan Lara Carmona and Ma. Eugenia Ramírez López, both from Instituto de Ecología, helped in the herbarium and in the computer preparation of this paper, respectively. Santiago Chacón and Rigoberto Gaitán-Hernández also from Insti­tuto de Ecología, provided the pictures (Frcs. 1, 2 and Frc. 3, respectively) presented here; they also provided infor­mation in collections and cultures, respectively. Dr. T. Ahti at Helsinki helped with the Latin diagnosis.

LITERATURE CITED

Aguilar, B. 1973. Propagación de la fase micelial de Mor­chella en cultivo sumergido. Bachelor Thesis. Escuela Nacional de Ciencias Biológicas, I.P.N., Mexico City. 53 pp.

---. 1979. Propagación de la fase miceliar de Morchella. M.S. Thesis. Departamento de Biotecnología y Bioin­geniería, Centro de Estudios Avanzados, I.P.N., Mexico City. 138 pp.

Anonymous. 1994. Diario Oficial de la Federación. Norma NOM-059-ECOL-1994, que determina las especies y su­bespecies de flora y fauna silvestre terrestres y acuáticas en peligro de extinción, amenazadas, raras y las sujetas a protección especial, y que establece especificaciones para su protección. Secretaria de Desarrollo Social, 488 (10), 16 May. Mexico City. 55 pp.

Baral, H. O. 1992. Vital versus herbarium taxonomy: mor­phological differences between living and dead cells of ascomycetes, and their taxonomic implications. Myco­taxon 44: 333-390.

Boudier, E. 1905-1910. leones mycologicae. Paris (reimpr. 1981-1985. Piantanida. Laussane), Vols. 2, pis. 194-421, 4, 362 pp. & 5, 300 pp.

Breitenbach,J., and F. Kranzlin. 1984. Champignons de Suis­se, l. Ascomycetes. Mykologia, Luzern. 310 pp.

Bunyard, B. A., M. S. Nicholson, and D. J. Royse. 1994. A systematic assessment of Morchella using RFLP analysis of the 28S ribosomal RNA gene. Mycologia 86: 762-772.

Chacón, S., and G. Guzmán. 1995. Observations on the phe­nology of ten fungal species in the subtropical forests at Xalapa, Mexico. Mycol. Res. 99: 54-56.

- --, ---, L. Montoya, and V. M. Bandala. 1995. Guía ilustrada de los hongos del Jardín Botánico Francisco Ja­vier Clavijero de Xalapa, Veracruz y áreas circunvecinas. Instituto de Ecología, Xalapa. 142 pp.

Denison, W. C. 1963. A preliminary study of the operculate cup-fungi of Costa Rica. Rev. Biol. Trap. 11 : 99-129.

Dennis, R. W. G. 1968. British Ascomycetes. Cramer, Lehre. 455 pp. + 51 pis.

Gamundí, I. J. 1975. Fungí, Ascomycetes, Pezizales. Pp. 7-185. In: Flora criptogámica de Tierra del Fuego 10(3). Eds., S. A. Guarrera, I. J. Gamundí, and D. Rabinovich. Fundación para la Ciencia y la Cultura, Buenos Aires.

---. 1983. Pezizales de América tropical. Anais 34, Con­greso Nacional de Botánica, Vol. I, pp. 151-171, Socie­dade Botánica do Brazil, Porto Alegre.

Gessner, R. V. 1995. Genetics and systematics of North American populations of Morchella. Ganad. J. Bot. (Suppl. 1): S967-S972.

Gilbert, F. A. 1960. The submerged culture of Morchella. Mycologia 52: 201-209.

Gómez, L. D. 1971. Un nuevo discomycete operculado de América Central: Morchella herediana, nov. sp. Darwin­iana 16: 417-426.

Guzmán, G. 1977. Identificación de los hongos. Limusa, Mex­ico City. 452 pp.

---. 1997. Los nombres de los hongos y lo relacionado con ellos en América Latina. Introducción a la etnomicobiota y micología aplicada de la región. Sinonimia vulgar y científica. CONABIO and Instituto de Ecología, Xalapa. 356 pp.

--- ,M.F. Torres, and H. Logemann. 1985. Fungí from Guatemala I. A new species of Morchella. Mycol. Helv. 1: 451-459.

Guzmán-Dávalos, L., and O. Rodríguez-Alcantar. 1993. Re­gistro de Morchella guatemalensis para México. Bol. lnst. Bot. Univ. Guadalajara 1: 471-475.

Hawksworth, D. L., P.M. Kirk, B. C. Sutton, and D. N. Peg­ler. 1995. Ainsworth & Bisúy's Dictionary of fungi. 8th ed. CAB International, Wallingford. 616 pp.

Heim, R. 1966a. Quelques ascomycetes remarquables. V.­Morilles tropicales (Pacific sud). Bull. Soc. Mycol. France 82: 442-449.

---. 1966b. Breves diagnoses latinae novitatum generi­carum specificarumque nuper descriptarum. Rev. My­col. (Paris) 31: 150- 159.

Herrera, T. , and G. Guzmán. 1961. Taxonomía y ecología de los principales hongos comestibles de diversos lu­gares de México. Ana/,es lnst. Biol. Univ. Nac. Auton. México 32: 33-135.

---, and M. Ulloa. 1990. El Reino de los Hongos. UNAM and Fondo de Cultura Económica, Mexico City. 552 pp.

Jacquetant, E. 1984. Les morilles. Bibl. Arts. Paris. 114 pp. Jung, S. W., R. V. Gessner, K. C. Keudell, and M. A. Romano.

1993. Systematics of Morchella escu/,enta complex using enzyme-linked immunosorbent assay. Mycologia 85: 677-684.

Korf, R. P. 1973. Discomycetes and Tuberales. Pp. 249-319. In: The fungi. Vol. N A. Eds., G. C. Ainsworth, F. K. Sparrow, and A. S. Sussman. Academic Press, New York.

Kornerup, A., andJ. H. Wanscher. 1978. Methuen handbook of colour: 3rd ed. Methuen. London. 252 pp.

Marchand, A. 1971. Champignons du Nord et du Midi. Vol. I. Soc. Mycol. Pyrénées Méditerr., Perpignan. 273 pp.

--. 1973. ldem. Vol. II . 282 pp. Nieto-Roaro, D. 1944. Flora micológica mexicana. Descripción

de especies de los géneros Helvella y Morchella. M. S. The­sis. Facultad de Ciencias, UNAM, Mexico City. 21 pp.

714 MYCOLOGIA

Ower, R. 1982. Notes on the development of the more! as­cocarp: Morchella escul,enta. Mycologia 74: 142-144.

Pfister, D. H. 1974. Notes on Caribbean Discomycetes. V. A

preliminary annotated checklist of the Caribbean Pez­izales. J. Agric. Univ. Puerto Rico 58: 358-378.

Ramsbottom,J. 1959. Mushrooms & toadstools. Collins, Lon­don . 306 pp.

Rodríguez, R. M., and T. Herrera. 1962. Descripción y cul­tivo de hongos del género Morchella del Valle de Méx­ico. Rev. Mex. Hist. Nat. 23: 119-136.

Seaver, F. J. 1961. The North American cupjungi. Suppl. Haf­ner, New York. 377 pp.

Singer, R. and B. Harris. 1987. Mushrooms and truffles. Koeltz Scientific Books, Koenigstein. 389 pp.

Smith, A. H. and N. S. Weber. 1980. The mushroms hunter's field guide. The Michigan University Press, Ann Arbor. 316 pp.

Stamets, P. 1993. Growing gourmet and medicinal mushrooms. Ten Speed Press, Berkeley. 554 pp.

Volk, T. J., and T. J. Leonard. 1989. Experimental studies on the more!. I. Heterokaryon formation between monoascosporous strains of Morchella. Mycologia 81: 523-531.

Wipf, D., S. Koschinsky, P. Clowez, J. C. Munch, B. Botton, and F. Buscot. 1997. Recent advances in ecology and systematics of morels. Cryptog. Mycol. 18: 95-109.

Yoon, C.-S. , R. V. Gessner, and M. A. Romano. 1990. Pop­ulation genetics and systematics of the Morchella escu­lenta complex. Mycologia 82: 227-235.