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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, rufobrunnea, and new data on M. guatemalensis

Gastón Guzmán1 comata, include "little beehives," "little tender corn Fidel Tapia ears," " little ear of green corn" and "little paunch," Instituto de Ecología, Apartado Postal 63, Xalapa, respectively (Guzmán, 1977, 1997; Herrera and Guz­ Veracruz 91000, Mexico mán, 1961; Herrera and Ulloa, 1990). The commer­ cial name far the more! is " morilla," from the Span­ ish word meaning beehive . Morels have a high eco­ Abswact: A critica! analysis of the known species of nomic value in Mexico due to export to Europe and Morchella from Mexico, including M. angusticeps, M. the U.S. Exploitation of more Is, together with the de­ costata, M . elata, M. esculenta (M. canica; M. crassipes, struction of the farests where they grow, farced the M . rotunda), M . guatemalensis and M . umbrina, is Mexican Governmen t to consider the species of Mor­ presented. In addition, M. rufobrunnea is described chella, together with other edible fungi, in the Red as a new species from the State ofVeracruz. Morchella List of Organisms (Anonymous, 1994) . rufobrunnea belongs to the blushing species, a faurth Species concepts in Morchella vary widely, and group of species in the genus proposed by the au­ sorne authors recognize 50 or more species, whereas thors. The blushing species also include M. guate­ others recognize only three good species (Gessner, malensis and M. rigidoides, and ali three species are 1995; Korf, 1973). Dennis (1968) noted that most confined to the tropics or subtropics. Morchella rigi­ modern authors regard ali the European species of doides is known only from New Guinea, and M . gua­ Morchella as variants of M. esculenta Pers. : Fr. and M. temalensis only from Guatemala and Mexico. The new elata Bull.: Fr. However, Dennis (1968) recognized species differs in the farm and color of the ascomata, three species from the British Islands, and Seaver length of the alveolae, and the color of staining. A (1961) six species from North America. Hawksworth revision of the microscopic features of M. guatema­ et al. (1995) considered 28 species in the genus, and lensis is made based on recent collections from Mex­ Jacquetant (1984) also distinguished 28 taxa, al­ ico and Guatemala. After a comparison of the asci, though many names recognized by him were invalid. ascospores and paraphyses of M. guatemalensis with The more recent studies recognize a fewer number those of M. rufobrunnea and other clase species, it is of species in the genus (Singer and Harris, 1987). concluded that microscopic features, except the Bunyard et al. (1994), based on ribosomal DNA stud­ width of the paraphyses in sorne cases, are noi im­ ies of six species, recognized only two as valid species. portant in the of the genus. The farm and Volk and Leonard (1989) andjung et al. (1993) con­ color of the ascomata, the position of the ribs, the sidered M. esculenta, M. crassipes (Vent.) Pers. and length of the alveolae, and the staining are the most M. deliciosa Fr.: Fr. as conspecific. Gessner (1995) important taxonomic features in Morchella. A key to summarized infarmation on mating studies and iso­ the seven known species of Morchella in Mexico is lation of chemical products and antigens far species presented. delimitation in Morchella. Key Words: Ascomycetes, , blushing Denison (1963), Heim (1966a, b), Gómez (1971), Pfister (1974), Gamundí (1983), Guzmán et al. ( 1985), and Guzmán-Dávalos and Rodríguez-Alcantar INTRODUCTlON (1993) are among the few studies facused directly or '- The morels (Morchella Dill. ex Pers.) are very impor­ indirectly on Morchella in the tropics and subtropics. tant edible in Mexico, where they have A study on cultures of Mexican morels (M. esculenta, been used traditionally far centuries. The Indian M . canica Pers., M. crassipes and M . angusticeps Peck) names far the morels include "colmenitas," "mazor­ was made by Rodríguez and Herrera (1962), and quitas," "elotitos" and "pancitas," and other com­ Aguilar (1973, 1979) studied submerged cultures of mon names, based on the peculiar farm of the as- M . crassipes and M. elata. In the present study, specimens of Morchella gath­ Accepted far publication March 6, 1998 ered at a subtropical site in the Ecological Park of 1 Email: [email protected] the Ecology Institute at Xalapa and in other parts of

705 706 MYCOLOGIA the Xalapa region (Mexico), and all the collections between them, forming short or elongate alveolae ex­ of Morchella at XAL Herbarium were examined. An tended vertically, but in aging the vertical configu­ interesting Morchella is described herein as a new spe­ ration of the ribs is somewhat lost and form a wrin­ cies, and new records of M. guatemalensis Guzmán, kled hymenophore (like that of M. esculenta); surface M. F. Torres & Logem. are reported. 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, MATERJALS ANO METHODS 5C4, 5DA) when mature. (20- ) 30-70 (- 90) X Fleshy specimens of Morchella spp. were dried ap­ 10- 25 mm, cylindric or more thick at the base, irreg­ prox. 1 h after collection. The drying apparatus con­ ularly wrinkled mainly toward the base, covered by sisted of an electric grid and natural aeration. Spec­ minute dark granules toward the apex, surface whit­ imens dried in approx. 24 h. Sections of the ascom­ ish to cream (4A2-3, 5A4) or pale grayish (5B3, ata, mainly from the ribs, were mounted in 5% aque­ 5DA), darker (grayish brown, 5F5- 6) toward the ous KOH, Melzer's reagent, cotton-blue in base, yellowish (4A5) or dark grayish (4E3, 4F4) in lactophenol or in Congo-Red, according to the mi­ old specimens. Blushing in irregular small spots, croscopic features observed. Measurements were brown (6F6), brownish orange or pinkish red (4A8, made in young and mature specimens from rehy­ 5A2-5, 5A8, 5B6- 8) to ferruginous (7C8) when in­ drated specimens mounted in KOH. A culture of the jured or in maturing, both in hymenophore and holotype (IE-228) on malt agar (MA) and on potato stipe; sometimes in old specimens becoming almost dextrose agar (PDA) (both from Bioxon, Mexico completely reddish brown (12E6-12F8). In dry as­ City) was obtained from the context of one fresh comata the color is pale brownish (5B4) with br9wn­ specimen. This strain is maintained in test tubes on ish pink (7B5, 7C5) or reddish brown (8F4-8) z~ees. MA at 5 C, and transferred every six mo. All speci­ Inner surfacewhitish, little orno blushing at the base. mens studied are deposited in Herb. XAL, except Basal mycelium a compact thin mass, whitish to pale where other herbaria are cited. The culture strain is yellowish, with sorne zones stained brownish orange deposited in the Strain Collection of the Ecological (6B8). print pale orange or yellowish orange Institute at Xalapa. Colors in the description are ac­ (4A7, 5A7). Asci (280- ) 300- 360 (-380) X 16-20 µm, cording to Kornerup and Wanscher (1978). 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 RESUU"S inamyloid. Ascospores (19-) 20- 24 (- 25.5) X (13- ) Morchella rufobrwmea Guzmán et Tapia, sp. nov. 14-16 (-17) µm, but in immature ascomata (11-) FIGS. 1-7 14.5-19 (-20) X (8-) 9- 10 (-11) µm, ovoid, thin­ walled, hyaline, inamyloid. Paraphyses 90-184 X (8-) Ascomata (60- ) 90- 155 (-210) mm alta, conica vel sub­ 10-18.5 µm, or X (5-) 6-9 (-10.5) µm when im­ cylindrico--0voidea, hymenio griseo vel spadiceo flavido, cos­ mature, hyaline, simple, septate at the base, consist­ te longitudinales vel inaequalis albidae ve! griseae, maculis ing of one or two cells. Context (in the inner part of brunneis ve! aurantiaco-brunnescentibus vel ferrugineis. a rib) formed of intermixed hyphae, 3- 9 (- 13.5) µm Stipes (20- ) 30-70 (-90) X 10-25 mm, albidus vel subgri­ seus, basis venosus, fusco-griseus, maculatus similis hymen­ wide, hyaline, thin-walled, cylindric to somewhat in­ io. Sporae in cumulo subaurantius vel flavidus subaurantius. flated. Asci (280-) 300- 360 (-380) X 16--20 µm, operculatus, ten­ Cultures. The culture obtained from vegetative tis­ uitunicatum, octosporus, inamyloideus. Sporae ( 19-) 20-24 sues grew slowly at 25 C on MA and PDA, producing (- 25.5) X (13- ) 14-16 (-17) µm maturae, (11-) 14.5- 19 a whitish to tan, sparse mycelium, covering the sur­ (-20) X (8- ) 9-10 (-11) µm immaturae, hyalinae, ovo­ face of the medium in 9-cm-diam Petri dishes in 10- ideae, inamyloideae. Paraphyses 90-184 X (8-) 10-18.5 µm 12 d. Sclerotia and a conidial stage were absent. [vel immaturae X (5-) 6--9 (-10.5) µm], hyalinae, basis sep- These results contrast with those of other studies on tatae. Morchella spp. reporting rapid growth in culture. For HOLOTYPUS. Habitat terra arenosa, in sylvae subtropi­ calis, in horto cum Quercus, Liquidambar; C/,ethra et Alnus, example, Stamets (1993) obtained a dense mycelium 1350 m altitudinis, Mexico, prope Veracruz, ad Xalapa, Guz­ from M . angusticeps on malt yeast agar, oatmeal malt mán 31565 XAL. yeast agar and pota to dextrose yeast agar after 3- 7 d. A conidial stage and sclerotia were common in Sta­ Ascomata (60-) 90-155 (-210) mm high. Hymen­ mets' experiments. ophore (40- ) 60-85 (-120) X (20-) 30- 45 (-50) mm, conic or subconic to subcylindric-ovoid when mature, Specimens examined. MEXICO, VERACRUZ: Municipio with longitudinal ribs and secondary transverse veins 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

DISCUSSION

This 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, Guz­ mán et al., 1985). For this reason it is important to study both immature and mature ascomata in collec­ tions. differs from M . guatemalen­ sis in the form and color of the ascoma, as well as in the type of blushing. Morchella guatemalensis has sub­ cylindric-ovoid to conic ascomata up to 105 mm high, with a longitudinally ribbed hymenophore and elon­ gate 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 in­ tense 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 Cal­ Frc. 3. Morchela rufobrunnea, dry immature ascomata edonia, respectively (Heim, 1966a, b), and the only (from the holotype). Scale bar 15 mm. 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 Instituto de Ecología, Ecological Park, near the crossing to longitudinal ribs as in M. guatemalensis. Morchella ru­ DIF, 21 May 1996, M. Villegas; same date, G. Guzmán Jobrunnea differs from M. rigi,doides in the larger as­ 31393, 31394; same locality, 5June 1996, G. Guzmán 31565 comata, color of the hymenophore (pale ochraceous (holotype, derived culture strain IE-228, isotypes NY, to yellow without gray or grayish tones in young IBUG) , 31566; same locality, 19June 1996, S. Chacón5066; stages in Heim's species) , less elongated alveolae, and same locality, 26June 1996, G. Guzmán 31605; same local­ wider paraphyses (TABLE 1) . Heim's species is clase ity, 9 July 1996, G. Guzmán 31627; same locality, 12 July 1996, F Tapia 1389, 1390, 1391. Xalapa, garden in a pri­ to M. guatemalensis but differs in the width of the vate house, 13 Dec. 1994, S. Chacón 4871 (ali in XAL) . paraphyses and less intense blushing. Other species similar to M . rufobrunnea based on Habitat and distribution. Gregarious, known only the descriptions and illustrations of Boudier (1905- from the Xalapa region. In the type locality, M. ru­ 1910), Marchand (1971 , 1973), Breitenbach and Jobrunnea was found at 1350 m altitude in a subtrop­ Kranzlin (1984) andjacquetant (1984) are discussed ical (mesophytic) forest with Quercus, Liquidambar, below, although many combinations and new taxa Clethra, and Alnus in sandy soil on a recently dis­ proposed by Jacquetant may be invalid. Morchella ela­ turbed roadside embankment with tree trunks. Fruit­ ta var. purpurascens Krombh. ex Boud. in the sense ing in the type locality occurred between mid-May to of Boudier (1905-1910) has en tirely lilaceous pink mid:July in 1996, but was not observed in 1997. It is or vinaceous ascomata on all the surfaces, both young interesting to note that the type locality is in the Eco­ and adult stages. According to Marchand ( 1973), this logical Park at Xalapa zone, which has been explored fungus has a pale carmine, lilaceous to violaceous hy­ mycologically for more than 10 yr wi th more than 200 menophore anda white stipe.Jacquetant (1984) con­ species of fungi recorded (Chacón and Guzmán, sidered this fungus as "M. purpurascens (Krombh.) 1995; Chacón et al., 1995). However, this is the first Jacquet." with vinaceous brown or blackish ribs, gray­ record of a Morchella sp. in the zone. ish green to pink or vinaceous pink alveolae, and a G UZMÁN AND T APIA: MüRELS IN MEXICO 709

7

8 l

1 1 1 1 , 1 1 I 1 , 1 , 1 1 1 1 , 1 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 . -1' ...... o

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

Speciesª Asci (µm) Ascospores (µm) Paraphyses (µm ) M. rufobrunnea (HOLOTYPE) (280-) 300-360 (-380) X 10-20 (19-) 20-24 (-25.5) X (13-) 14-16 (-1 7) " 90- 184 X (8-) 10- 18.5 or (11- ) 14.5-19 (- 20) X (8-) 9- 10 (- ll )c X (5- ) 6-9 (-10.5) when immature M. guatemalensis 1 280-420 X 18-23 (17- ) 18- 22 X 11-13 56-103 X 6- 12 2 159-250 (-287) X 18-21.5 (15- ) 16-21 X (9- ) 9.5- 13 X 7-1 3 3 300-400 (-420) X 17.5- 26.5 (- 28) (17.5-) 18.5-24 X (12-) 13-14.5 X 6.5-12 (-13) a:: 4 197-366.5 X 18.5- 27.5 (14.5-) 16-23 X 9-15 rs 5 X (16-) 17.5-26.5 (-28) (17-) 19-25 (- 25.5) X (12- ) 13-14.5 (-1 6) X (9- ) 9.5-1 6 (-1 7.5) o ot"' 6 (280-) 300-390 (-410) X 17-28 (-30) (17.5-) 18- 24 (-25) X (12- ) 13-14 X (6.5- ) 7-1 3 (-13.5) () M. herediana :; 7 144-172 X 14.5- 20 17-18 .5 X 8-9 144-172 X 9- 14 M. rigidoides 8 240-2 50 X 18-20 18-20 X 10-12 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 & Rodríguez­ Alcantar (1993); 5: Sommerkamp 227; 6: topotype; 7: Gómez (1971); 8: Heim (1966a, b). "Mature e Immature GUZMÁN ANO TAPIA: MORELS IN MEXICO 711 white-pink stipe. Boudier described the fungus with fourth taxonomic group of morels represented by short alveolae, whereas Marchand andJacquetant de­ the blushing species of the tropics and subtropics, scribed elongate alveolae. Jacquetant also recognized including M. rufobrunnea, M. guatemalensis and M . "M. purpurascens var. ionoviridis Jacquet." with vio­ rigidoides. Gilbert (1960), based on a study of sub­ laceous ribs and greenish elongate alveolae, and ap­ merged cultures, found that M. esculenta and M. can­ parently a white stipe. This author also described "M. ica have certain cultural characteristics in common purpurascens var. ionoviridis f. heteroparaphysa Jac­ that are distinct from those of M . angusticeps. Ower quet." with subglobose paraphysis apices as the only ( 1982), in a study of ascoma development in a more] distinguishable feature. Morchella elata var. elata, ac­ he called M. esculenta, described cultures forming cording to Breitenbach and Kranzlin (1984), has a gray ascomata that turned yellow as they matured. brownish yellow to brownish red hymenophore with However, the ascomata obtained by Ower exhibited blackish ribs and short alveolae and white stipe. Ac­ conspicuous longitudinal ribs and deep, elongate al­ cording to Boudier ( 1905- 191 O), this fungus is en­ veolae, features not typical of M. esculenta, but com­ tirely brownish yellow, and in Gamundí ( 1975), the mon in M. costata Vent. Yoon et al. (1990) also re­ hymenophore is described as brown to blackish with ported that in M. esculenta the immature gray ascom­ elongate alveolae and white stipe. In Marchand ata changed to a tan color as they matured. These ( 1971) this fungus is described with a grayish brown observations show the complex and contradictory na­ hymenophore, elongate alveolae and white stipe. ture of the taxonomy in Morchella, particularly for Morchella rigida (Krombh.) Boud., according to Mar­ the common species. chand (1973), has an alveolate hymenophorewithout Microscopic features of the Morchella hymenium longitudinal ribs, similar to M . esculenta, and also a appear to be relatively homogeneous among taxa, as whitish stipe, but with sorne inconspicuous brownish observed by the authors for the previously discussed spots. According to Boudier (1905- 1910),Jacquetant species, and as noted by Wipf et al. ( 1997). It is dif­ (1984) and Breitenbach and Kranzlin (1984), this ficult to separate Morchella species based only on mi­ fungus also has a structure and form similar to that croscopic features. However, Singer and Harris of M . esculenta, but with a stipe lacking spots. Heim ( 1987) separated M. es cu lenta, M. hortensis Boud. ( l 966a) separated M. rigidoides from M. rigida based and M. costata in their key mainly by the size of the on the height of the ascomata and shallower alveolae. . Among the more! species discussed above, in­ It is important to observe that none of the species cluding M . rufobrunnea, there are no important dif­ discussed above, except M. guatemalensis and M. rig­ ferences in the microscopic features of the hymeni­ idoides, clearly exhibit the blushing feature. It seems um except in the width and form of the paraphyses that blushing is an important taxonomic feature, as in sorne species. Examples ofvariation in microscopic observed in other mushrooms, e .g., Amanita rubes­ features among species cited in the literature may be cens (Pers.: Fr.) Gray. The senior author has seen due to the use of immature specimens, or that the many fresh morels during his years of field work, and microscopic features were observed in different con­ M . guatemalensis and M. rufobrunnea are the first ditions (e.g., fresh or dried specimens), as noted by species observed by him to exhibit blushing. We be­ Baral (1992) in many ascomycetes. He stated that dif­ lieve that the blushing feature is different from the ferences between diagnoses based on the morphol­ spots produced on sorne mushrooms as result of in­ ogy of microscopic elements from fresh, living spec­ fection by bacteria or other organisms. However, it is imens, and those based on dead, dry specimens, had possible that the spots represented in the iconogra­ implications that could lead to erroneous taxonomic phy of M. rigida ofMarchand (1973) discussed above conclusions. Morchella rufobrunnea, M . guatemalensis are due to this type of infection. and M. herediana seem to differ from M. rigidoides Based on the results of DNA studies, Bunyard et in the width of the paraphyses observed in dry spec­ al. (1994) suggested that there are at least two taxo­ imens (TABLE 1) . The authors studied also a collec­ nomic groups in Morchella: (i) the black morels rep­ tion (at XAL) from Region ofTlaxco, Tlaxcala (Mex­ resented by M. angusticeps, M. elata and M . canica, ico), gathered in a Pinus-Abies forest, on 21 Sep. 1985 and (ii) the yellow morels, with M. esculenta, M . cras­ (l. González-Fuentes 1013) that it is similar to M. ru­ sipes and M. deliciosa. They suggested that these two Jobrunnea in the ascoma, with a strong blushing on groups represent only two species. Another group of the stipe, but has paraphyses (12-) 13.5- 20 (- 24) µm Morchella species is what sorne authors call half-free wide. It is likely that this collection represents a new morels, represented by M. semilibera (DC.: Fr.) Lév. species, but the authors require additional collec­ (Gessner, 1995). Although Smith and Weber (1980) tions for further study. provided a description of M. semilibera, this speccies It is concluded that the ascoma of species of Mor­ is in need of revision. It is possible that there is a chella is enormously variable in morphology, depend- 712 MYCOLOGIA ing on the development, and possibly on weather and ing in large spots dark vinaceous or reddish microniche conditions. Morchella rufobrunnea is sim­ brown ...... M. guatemalensis ilar to M. guatemalensis and other morels, but differs 6b. Hymenophore gray when young to brownish yel­ in the form and color of the ascoma, length of the low when mature, with longitudinal to irregular alveolae, and the color of staining. We based the de­ ribs and elongate to short alveolae. Blushing in irregular, small or occasionally large spots, brown scription of the new species on two populations, pri­ or pinkish red to ferruginous ...... marily the population from the type locality where ...... M . rufobrunnea many ascomata were collected between May 21 and July 12, 1996 from an area approximately 6 m 2 • An New data on M. guatemalensis.-This species was de­ interesting observation is that this species was gath­ scribed from Guatemala, Department of Chimalten­ ered in heavily disturbed sites, as is common for ango, gathered in Dec. 1983 and in Oct. 1984 (Guz­ many species of the genus (Ramsbottom, 1959; Ges­ mán et al., 1985). Recently it was reported from Mex­ sner, 1995). ico (State of Jalisco) by Guzmán-Dávalos and Rodrí• guez-Alcantar (1993). When revising the morels The known species of Morchella in Mexico.- Little is deposited at XAL Herbarium, the authors found new known about the diversity and taxonomy of Morchella records of this fungus, five from Guatemala, and in Mexico. Nieto-Roaro (1944), Rodríguez and Her­ three from Mexico. rera (1962) and Herrera and Ulloa (1990) discussed M. angusticeps, M. canica, M. crassipes, M. esculenta Specimens examined. GUATEMALA. ZACATEPEQUEZ: and M. rotunda Pers. Guzmán (1977) accepted all San Lucas, 10 Aug. 1985, A. Moller, San José Pinola, San species except the latter, and also considered M. cos­ Juan Mushball, 27 July 1986, G. Guzmán 29227; 27 July tata and M. elata in his keys. Recently, Guzmán-Dáv­ 1986, G. Guzmán 29231; 27 July 1986, G. Guzmán 29233; Guatemala City, 30 Aug. 1988, Y. Sommerkamp 227. MEXI­ alos and Rodríguez-Alcantar (1993) recorded M. CO. VERACRUZ: old road Xalapa to Coatepec, km 6.5, El guatemalensis from Jalisco, and Guzmán et al. (un­ Atorón, 2 Nov. 1990, VM. Bandala 2004; Xalapa, S of the publ.) reported M. umbrina Boud. from Veracruz. In Unidad La Haya, 26 Jan. 1996, S. Chacón 4992; MICHOA­ consideration of the observations by Volk and Leon­ CÁN: Morelia to Ciudad Hidalgo, Mil Cumbres, near Placa ard (1989),Jung et al. (1993), and others, it is prob­ Conmemorativa, 7 Jan. 1984, ].A. Pérez de la Rosa (IBUG able that M. canica, M. crassipes and M. rotunda are 954); MORELOS: Valle de Tepeite, NW of Santa María, 13 conspecific with M. esculenta. Hence, there are only Oct. 1984, R. Valenzuela 4275 (ENCB); Cuernavaca, seven species of Morchella, including M. rufobrunnea, U.A.E.M., University Campus, 25 Oct. 1981, L. López 272. recorded from Mexico. Of these species, only M. gua­ These materials compared well with the isotype of temalensis and M. rufobrunnea belong to the blushing M. guatemalensis (XAL), except in sorne variation ob­ group proposed herein. served in the width of the paraphyses. Guzmán et al. (1985) described paraphyses 6-12 µm width; Guz­ KEY FOR THE MEXICAN SPECIES OF MORCHELLA mán-Dávalos and Rodríguez-Alcantar (1993) 7-13 la. Ascoma never blushing (but may blacken in age) µm; in the isotype it was found 6.5- 12 (-13) µm; in 2 the topotype (6.5-) 7-13 (- 13.5) µm, and in Somer­ 1 b. Ascoma blushing (may be difficult to discern in kamp 227, (9-) 9.5-16 (-17.5) µm (TABLE l) . In the young ascomata) ...... 6 Mexican materials examined, the variation was from 2a. Hymenophore yellowish, blackish in old and putrid ascomata ...... 3 6-13.5 µm to (8-) 9-14.5 (- 16) µm. The observed 2b. Hymenophore gray to blackish ...... 5 variation of the width of the paraphyses appears to 3a. With irregular ribs, forming short alveolae. Hymen­ be related to the maturity of the ascoma. The greatest ophore globose, subcylindric or conic, sometimes width was observed in mature species, as was also not­ thick footed ...... M . esculenta ed in M . rufobrunnea. The vegetation in the above 3b. With longitudinal ribs ...... 4 localities, including that reported by Guzmán-Dávalos 4a. Transverse ribs conspicuous, forming short, and Rodríguez-Alcantar (1993) , is tropical or sub­ rounded and deeper alveolae ...... M. costata tropical with Quercus forests, mixed with Alnus and 4b. Transverse ribs not common, with elongate al­ Arbustus ( Cupressus also grows in the type locality). veolae ...... M . elata The main distinguishing feature of M. guatemalensis 5a. Hymenophore globose, with short, rounded alveo- is the blushing, the color of the hymenophore, and lae in irregular ribs ...... M. umbrina 5b. Hymenophore subcylindric or conic, with elongate the longitudinal and elongate alveolae, as discussed alveolae ...... M. angusticeps above. Dimensions of the asci, ascospores and pa­ 6a. Hymenophore yellowish orange or honey-col­ raphyses of the isotype, shown in Frcs. 8-13, are pre­ ored, both in young and mature stages, with sented in TABLE I in comparison to those of other longitudinal ribs and elongate alveolae. Blush- Morchella species reported in the literature. GUZMÁN ANO TAPIA: MORELS IN MEXICO 713

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