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Jeffrey R. Recibido: 14 junio ; aceptado: 15 abril Quercus undata Trel.

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Fagaceae, QuercusSection Quercus has a complex taxonomic and nomenclatural history. Intensive sampling of oaks at the type locality of Q. Endlich's type specimen of Q. An epitype is also deated in support of the lectotype, given that some features cannot be critically observed on the illustration. The long peduncles of the specimen illustrated by Trelease indicate a close relation to Q. Key words: Sierra Madre Occidental, Mexico, epitype, lectotype, introgressive hybridization. Un muestreo intensivo de los encinos en la localidad tipo de Q.

The genus Quercus in Mexico includes about species Valencia Avalos,and past estimates have placed more than scientific names in the Mexican portion of the genus Trelease, White oaks Fagaceae, Section Quercus in the Mexican flora remain taxonomically perplexing, due to poor morphological differentiation among many species, abundant hybridization, inadequate documentation of subgeneric epithets, and poor representation of fertile structures in herbaria. Herbarium specimens of Mexican white oaks are commonly misidentified.

Therefore revision and adecuate typification of taxa in this group are important to understanding and clarifying the species. Trelease described Quercus undata Trel. He considered Q. Later, McVaugh suggested that Q. Despite the placement of Q. In regional herbaria with numerous good collections there are few specimens identified as Quercus undata. Fruits visible in the published photograph of the type specimen Endlich 1 in Trelease have peduncles that are much longer than those of the other cited specimens Palmer and The latter were described as "[p]ossibly to be referred here also. The type specimen of Q.

In this paper we select the illustration of the type of Q. This work aims to clarify the taxonomic nature of Q. Examination of herbarium specimens : specimens of Q. Remaining specimens cited in Trelease's description Palmer and and deposited at A were also examined.

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Field sites: principal and secondary routes of the Sierra Madre Occidental were extensively searched, including transects described by Spellenberg and Spellenberg et al. The Sierra de la Candela area, where Endlich reported have collected the type specimen, was intensively searched and systematically surveyed throughout its altitudinal range, combinations of slope directions, and various vegetation types.

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Samples of all white oak species encountered were collected, pressed, dried, and identified in each of the habitat types where they occurred. The area of the type locality was determined see "The Type Locality," below to be in the zone where the 3 species of white oaks, Q.

Samples of foliage of each tree sampled were collected 2 m above ground level, on the south side of each tree. From each sample, morphological characters from the abaxial surface of 7 leaves and their trichomes were evaluated as shown in Fig. Mean values of leaf and trichome variables were calculated for each variable and used for multivariate analyses. Plants from the site and other sites where each species was found in allopatry were evaluated based on vegetative characters Fig. Data from all variables had relatively normal distributions and similar variance among taxa, making data transformation unnecessary.

Review of herbarium and field specimens of white oaks from much of the Sierra Madre Occidental revealed that those identified as Quecus undata appeared to fall within the variation observed in Q. Trelease's plate of the type pl. According to the label data reported by TreleaseEndlich collected the type specimen in the Sierra de la Candela at 2 m above sea level.

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Quercus arizonicaQ. Communities of low elevation white oak species discussed here, with morphological variation like that described in hybrid swarms of other species pairs Bacon and Spellenberg, ; Howard et al. Descriptive statistics Table 1 show little differentiation among vegetative characters of the taxa.

Likewise, analysis of vegetative morphology from specimens at the type locality and sites where parental species were allopatric, using both principal components analysis Fig. Taxonomic comments. The long, dense pubescence on leaf laminas and the long peduncles depicted in plate Trelease,suggest that the plate shows a specimen of Q. Lack of a scale on the photos and uncertainty as to whether the plate depicts one long peduncle partially covered by a leaf or 2 peduncles prevents precise determination of the peduncle length on the type.

However, Trelease reported that the leaves of Q. Hence the peduncle length of Q. Type locality : surveillance of the Sierra de la Candela at the altitudes near 2 m above sea level, where Trelease reported that Endlich collected the type specimen of Q. Two travel routes lead upward into oak habitat from Tepehuanes. One route, following a ridge that le up and over the Sierra de la Candela, to Boleras has only Q. That sapling was growing in a highly disturbed stream area at the lowest portion of the ridge, very much below any other white oaks, at a site long used nowdays as a garbage dump.

The oak sapling may have been established as a result of recent dumping activities.

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This route climbs steadily through elevations where one may find several other oak species. The type specimen, according to Trelease's list of Endlich's cited specimenswas the first of a series collected in the Sierra de la Candela, beginning on 27 August of Endlich 1. Following in the series were Q. Endlich 2aQ. Endlich 5and Q. Endlich 6 Trelease, Role of introgression : communities of the 3 white oak species found on the lower slopes of the Sierra de la Candela, as in other parts of the Sierra Madre Occidental, were characterized by plants with morphological variation similar to that described in hybrid swarms from the region Bacon and Spellenberg, ; Howard et al.

Sympatry occurred over extensive areas, suggesting that introgressive hybridization is probably very common in this region. Trelease's inclusion of specimens of Q. However, the leaves of the low elevation white oaks in this region Q. Principal components analysis and principal coordinates analysis, as well as descriptive statistics measured on vegetative characters, confirmed that morphology of vegetative characters overlaps and that taxa are distinguished with difficulty based upon vegetative characters.

Also evident is the fact that plants identified as Q. As such, any of the hybrid combinations, hybrids subsequently crossing among themselves, selfing, or backcrossing to the parental types, could generate individuals with leaves fitting the description of Trelease's Q. Fruits were not evaluated because they were not available the year plants were sampled, nor for two years thereafter, but from these analyses coincide with observations that these white oak species are very difficult to distinguish based on the vegetative characters that Trelease used to describe Q.

Typification of Quercus undata : the name of Q. Manfred Baessler at B, reported that the specimen does not exist in the collections pers. Remaining specimens cited in Trelease's description Palmer and deposited at Arnold Arboretum A at Harvard University were available for examination see Appendix A for a list of "Selected Specimens Examined" including herbaria consultedbut they do not correspond to Q.

Given that Endlich's specimen was apparently destroyed in bombing raids on Berlin during World War II and no duplicates are known to exist we propose here as a lectotype the Trelease illustration of the type depicted in plate of Trelease's treatment Selection of Quercus undata epitype : because some features cannot be critically identified from the lectotype plate in Trelease for purposes of the precise application of the name Quercus undataan epitype is deated here in support of the lectotype see McNeill et al.

This specimen is from a site at, or very near, the type locality of Q. It was collected from the lower edge of a hybrid swarm containing all 3 of the low elevation white oak species mentioned by Trelease Its morphology resembles that of Q. It also has abundant vegetative and fertile material. Duplicates of the same collection, although not representing epitypes McNeill et al. Originally, we considered that 1 of the other 2 specimens Palmer and listed by Trelease in his description of Q. However, neither of those specimens belongs to the same taxon.

The lectotype depicts variation of Q. Such variation and the presence of alleles introgressed from other species is common in white oaks Bacon and Spellenberg, ; Howard et al.

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The identity of Quercus undata: Quercus undata grows in hybrid swarms. Although we strongly considered the use of a hybrid name, Q. The presence of crisped leaf margins mentioned by Trelease and evident in the lectotype, is very common on leaves of trees at many sites where introgressive hybridization occurs between any 2 oak species. However, we cannot consider it a sure of hybridization. In addition, the specimen shown in Plate the lectotype has much stronger affinity to Q. We suggest that the name Q. Description of Q.

Small trees or shrubs, evergreen, to 4 m tall. Bark gray to brown, fissured, becoming brittle with age. Buds brown or beige, densely hoary to glabrous, round to broadly obovate.

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Leaf lamina base obtuse, rounded, or cordate; the lamina elliptical, entire or shallowly toothed apically, margins crisped, revolute or weakly revolute, horny to densely tomentose, 2. Distribution and ecology of Q. Occasionally associated with the black oaks Q. Taxonomic summary.

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