STUDIES IN THE QUERCUS UNDULATA COMPLEX I. A PRELIMINARY STATEMENT

Tucker , John M. (U. California, Davis.) Studies in the Quercus undulata complex. I. A preliminary statement. Amer. Jour. Bot. 48(3): 202–208. Illus. 1961.—The taxonomic history of Quercus undulata, a highly variable, problematic complex of the Southwest, is discussed. Conservatively treated as a single species, it comprises a wide range of forms which at one extreme approach Q. gambelii morphologically, and, at the other, Q. grisea, Q. turbinella, and other species. The postulate was made that Q. undulata had arisen through hybridization between these very different oaks. Field observations and preliminary study of numerous population samples confirm this postulate. (Detailed morphological analyses are in progress.) Seven species have apparently been involved—Q. gambelii (the “common denominator” of the complex) has hybridized in different parts of its range with one or another of the following: Q. arizonica, Q. turbinella, Q. havardii, Q. muehlenbergii, Q. mohriana, and Q. grisea. The latter 6 species are discussed individually, and the extent to which each contributes to the complex, and the area in which this occurs, are indicated.     

STUDIES IN THE QUERCUS UNDULATA COMPLEX. III. THE CONTRIBUTION OF Q. ARIZONICA

Tucker, J. M. (U. California, Davis.) Studies in the Quercus undulata complex. III. The contribution of Q. arizonica . Amer. Jour. Bot. 50(7): 699–708. Illus. 1903.—Of the 7 oak species involved in the Quercus undulata complex, Q. arizonica contributes the least of all. The latter has hybridized with Q. gambelii at a few widely separated localities (listed in Table 1) in central Arizona, and northern Sonora and Chihuahua, Mexico. The putative hybrids (identifiable as Quercus undulata) occur as isolated individuals with the parental species. The parents, although regionally sympatric, are usually ecologically isolated. They differ in a number of morphological characters, 6 of which were analyzed in detail. The resulting data, presented as pietorialized scatter diagrams, demonstrate that the putative hybrids are intermediate, in general. This is taken as evidence of their hybrid nature. Factors limiting hybridization in oaks are discussed. It is speculated that hybridization between Q. arizonica and Q. gambelii has occurred since the postglacial hypsithermal interval.     

STUDIES IN THE QUERCUS UNDULATA COMPLEX. IV. THE CONTRIBUTION OF QUERCUS HAVARDII

Scrub oak populations in the semidesert area of northeastern Arizona and southeastern Utah are ordinarily identified in regional manuals as Quercus undulata. They are very similar, both morphologically and ecologically, to Q. havardii of the Staked Plain of the Texas Panhandle and southeastern New Mexico. They differ, however, in a number of inconspicuous characters. Population sample analyses indicate that most of these differences are suggestive of Q. gambelii, and the deviant populations are thus interpreted as having been derived from ancestral Q. havardii through introgression by Q. gambelii. Two differences are not in accord with this interpretation; these are regarded as possible cases of transgressive segregation. Considering the evolution of these hybridized populations, it is speculated that the ancestral Q. havardii occurred to the south and west of the present range of this species during the Kansan period of the Pleistocene. During subsequent northward dispersal, it became split in two. The eastern portion ultimately came to occupy the present range of the species in the Staked Plain; the western portion—lying to the west of a north-south mountain barrier in central New Mexico—became introgressed by Q. gambelii (and locally by Q. turbinella), resulting in the present populations of Arizona and Utah.     

STUDIES IN THE QUERCUS UNDULATA COMPLEX. V. THE TYPE OF QUERCUS UNDULATA

The original material of Quercus undulata Torr. was collected by Edwin James on Major Long's expedition to the Rockies in 1820. The type locality was stated by Torrey as “sources of the Canadian and the Rocky Mountains.” Evidence is here adduced that the type locality is on Ute Creek, a tributary of the Canadian River, in northeastern New Mexico. The site was found to harbor Q, gambelii, Q. grisea, and numerous intermediate hybrid forms. Certain of the latter match the type of Q. undulata quite closely. The type sheet includes three elements—a branchlet regarded as the type, a twig of Q. grisea, and an isolated acorn of Q. havardii. Torrey's new species was illustrated by a figure which does not closely resemble the type, but which is more similar to Q. havardii. The acorn of Q. havardii on the type sheet must have been collected in the Texas Panhandle or western Oklahoma, where Long's party would have encountered this species. In my opinion, Torrey's figure could have been drawn from a specimen collected there, which, except for the isolated acorn, has since been lost.     

A three-trophic-level analysis of the effects of plant hybridization on a leaf-mining moth

We investigated relationships between host plant hybridization in the Quercus grisea x Q. gambelii species complex and the distribution and performance of a leaf-mining moth in the genus Phyllonorycter. In 2 years at two sites Phyllonorycter densities were lowest on Q. grisea and increased through the categories of hybrid host plants to Q. gambelii. Direct host plant effects on Phyllonorycter performance were consistent with Phyllonorycter distribution; unexplained mortality of larvae in the mines, which is often associated with direct plant effects, decreased from Q. grisea through the hybrids to Q. gambelii. Plant hybridization influenced parasitism of Phyllonorycter. Parasitism was density dependent, and across all densities was higher on Q. grisea and Q. gambelii than on hybrid host plants.     

HOW DISCRETE ARE OAK SPECIES? INSIGHTS FROM A HYBRID ZONE BETWEEN QUERCUS GRISEA AND QUERCUS GAMBELII

The white oaks Quercus gambelii and Q. grisea overlap in distribution in New Mexico and Arizona. Within the region of overlap, there are numerous instances of contact between the two taxa. In some areas of contact morphologically, intermediate trees are common, whereas in others, morphologically intermediate trees are rare or absent. We describe a set of RAPD markers that distinguish between the two species and use these markers to examine patterns of gene exchange in an area of contact in the San Mateo Mountains of New Mexico. The markers are highly coincident with morphology and confirm that hybridization between the two species takes place. Despite the occurrence of hybrids, both species remain distinct, even in areas of sympatry, and marker exchange appears to be limited.     

Interactions between host plants,endophytic fungi,and a phytophagous insect in an oak (Quercus grisea x Q. gambelii) hybrid zone

We examined interactions between host plants, endophytic fungi, and leaf-mining moths (Phyllonorycter sp.) in an oak (Quercus grisea x Q. gambelii) hybrid zone. The community of endophytic fungi and two common endophyte species examined responded to host plant hybridization. Total fungal frequency (TFF) and frequency of Gnomonia cerastis were lowest on hosts resembling Q. grisea, and increased linearly towards those resembling Q. gambelii. In contrast, Coccochorella quercicola was most frequently isolated from Q. grisea-like hosts and decreased in frequency across hybrids towards Q. gambelii. Frequency of G. cerastis and TFF covaried with Phyllonorycter density across the hybrid zone, but direct effects of endophytes on Phyllonorycter density were not detected. Associations between endophytes and unexplained mortality of Phyllonorycter varied according to endophyte species and state of Phyllonorycter development. In the sap-feeding stage, unexplained mortality was negatively associated with TFF and frequencies of Hormonema sp. and Preussia funiculata; whereas, in the tissue-feeding stage, unexplained Phyllonorycter mortality was positively associated with G. cerastis frequency. Three-way interactions between plant hybridization, endophytic fungi, and the insect herbivore were not significant.     

Are plant hybrid zones centers of vertebrate biodiversity? A test in the Quercus grisea × Quercus gambelii species complex

The plant-hybrid-zones-as-centers-of-biodiversity (hereafter, PHZCB) hypothesis posits that plant hybrid zones represent areas of elevated biodiversity, and supports the inclusion of plant hybrid zones in conservation efforts. Support for the PHZCB hypothesis is limited to a handful of cases involving insect and fungal species and to two studies involving higher trophic levels. The PHZCB hypothesis requires further testing before plant hybrid zones can be established with any certainty as centers of biodiversity. We test whether the PHZCB hypothesis holds for higher trophic levels by examining the community structure of birds and reptiles associated with the Quercus grisea × Quercus gambelii species complex. Specifically, we compare patterns of species richness and abundance of these taxa in 10 hybrid zones and 11 contact zones (i.e., zones where both parental species occur but hybrids do not). In contrast to predictions of the PHZCB hypothesis, contact zones supported significantly more species of birds and reptiles than did hybrid zones. Species abundances exhibited idiosyncratic responses to zone type, but were higher generally in the contact zones.     

EVOLUTIONARY SORTING OF CHARACTERS IN A HYBRID SWARM. I: DIRECTION OF SLOPE

Quercus Douglasii and Quercus turbinella subsp. californica intergrade in California in a belt about 250 miles long. In the area of each parental taxon and near Lebec, in the middle of the zone where hybrids are common and where the gene pool is about evenly divided as to origin, the phenotypes were scored on a six-point scale to establish hybrid indices. At Lebec populations on slopes facing at intervals of one-eighth the distance around the compass were scored. The composition of each population was heterogeneous but restricted markedly according to exposure to the sun. Plants on northeast-facing slopes approached but did not equal the high score of Quercus Douglasii in Madera County; those on southwest-facing slopes approached but did not equal the extreme low score of subsp. californica near Victorville. Evolutionary sorting must be very rapid, for it is correlated with slopes of the same hills, and the hybrid swarm must be very sensitive to the environment.     

Community structure of canopy arthropods associated to Quercus crassifolia×Quercus crassipes complex

Quercus crassifolia and Q. crassipes are dominant species in temperate forests of central Mexico and hybridize between each other when they occur in sympatry. Oak canopies contain a considerable portion of arthropod diversity and the hybrid zones can provide new habitats to epiphyte fauna. We tested if the establishment of hybrids in contact zones with their parental hosts increases the species diversity of canopy arthropods assuming that hybrid trees constitute new genotypes of potential new habitats to small organisms. We examined the effect of hybridization on some community structure parameters (diversity, composition, similarity and density of arthropod fauna) of canopy arthropods compared to their parental species in a hybrid zone located in central Mexico. We employed 17 leaf morphological traits and six diagnostic RAPD primers to identify parental and hybrid plants. The RAPDs marker showed unidirectional introgression towards Q. crassifolia, and were detected hybrid (F₁), backcrosses and introgression individual trees. In total, 30 oak canopies were fogged during rainy and dry season. We recognized 532 taxa of arthropods belonging to 22 orders associated with tree canopies. The taxonomic status of host‐trees may be an important factor in the arthropod community structure and that seasonality (dry and rainy) is not a factor that could modify their organization. Trees of Q. crassipes registered the highest densities of arthropod fauna followed by hybrid hosts (F₁); trees originated by backcrosses towards Q. crassifolia registered a significant less arthropod density than F₁ hybrids; and trees of Q. crassifolia had the lowest density. Hybrid plants and Q. crassipes individuals had higher diversity (H′) of arthropods than Q. crassifolia plants. Hybrid plants had also more rare species in both seasons in comparison with parental species. This study suggests that hybrid oaks act as a center of biodiversity by accumulating arthropods of both parental and different species including a considerable number of rare species.     

Natural hybridization and hybrid zones between Quercus crassifolia and Quercus crassipes (Fagaceae) in Mexico: morphological and molecular evidence

Hybrid zones provide interesting systems to study genetic and ecological interaction between different species. The correct identification of hybrids is necessary to understand the evolutionary process involved in hybridization. An oak species complex occurring in Mexico formed by two parental species, Quercus crassifolia H. & B. and Q. crassipes H. & B., and their putative hybrid species, Q. dysophylla, was analyzed with molecular markers (random amplified polymorphic DNA [RAPDs]) and morphological tools in seven hybrid zones (10 trees per taxa in each hybrid zone) and two pure sites for each parental species (20 trees per site). We tested whether geographic proximity of hybrid plants to the allopatric site of a parental species increases its morphological and genetic similarity with its parent. Seventeen morphological traits were measured in 8700 leaves from 290 trees. Total DNA of 250 individuals was analyzed with six diagnostic RAPD primers. Quercus crassifolia differed significantly from Q. crassipes in all the examined characters. Molecular markers and morphological characters were highly coincident and support the hypothesis of hybridization in this complex, although both species remain distinct in mixed stands. Clusters and a hybrid index (for molecular and morphological data) showed that individuals from the same parental species were more similar among themselves than to putative hybrids, indicating occasional hybridization with segregation in hybrid types or backcrossing to parents. Evidence does not indicate a unidirectional pattern of gene flow.     

Comparative analysis of micromorphological characters in two distantly related Mexican oaks, Quercus conzattii and Q. eduardii (Fagaceae), and their hybrids

Interspecific hybridization occurs with high frequency in the genusQuercus, but few studies have analyzed and compared micromorphological characters in putative parental species and their hybrids.Quercus eduardii andQ. conzattii are two Mexican black oak species that, although distantly related, have formed at least one population of hybrid origin, where individuals with intermediate macromorphology are present. The purpose of this investigation was to analyze the degree of differentiation in micromorphological characters between the two species and to assess the expression of these characters in individuals with intermediate macromorphology. Foliar trichomes, epicuticular waxes, stomata, and pollen grains, were examined using scanning electron microscopy (SEM) in the three types of individuals (Q. conzatti, Q. eduardii, and intermediates). Trichome density was quantified with light microscopy. Types of trichomes present, length of trichome arms, types of epicuticular wax on the leaf surfaces, and the position of stomata with respect to the foliar surface were characters useful to differentiate betweenQ. conzattii andQ. eduardii. Plants with intermediate macromorphology displayed a pattern of micromorphological characters that were identical to one parental species (Q. conzattii), or extreme or novel relative to both species.     

Variation in morphological traits in a recent hybrid zone between closely related Quercus liaotungensis and Q. mongolica (Fagaceae)

Aims Hybridization usually leads to gene introgression between related species in hybrid zones, associated with complex patterns of morphological variation. Nevertheless, previous studies have tended to ignore the effects of geographic variation in hybridization rates on species taxonomy. This study aims to investigate the variation of morphological traits between two sympatric and taxonomically confused oak species, Quercus liaotungensis and Q. mongolica, and reveal the effects of hybridization rates on morphological traits and the taxonomic boundary.Methods We used seven microsatellite loci to evaluate species status and measured 15 morphological traits in 26 trees in the recent hybrid zone between Q. liaotungensis and Q. mongolica, and we characterized the differences between the two oak species and their hybrids for the investigated traits.Important findings Molecular analyses indicated that 74% of 78 sampled maternal trees were hybrids between Q. liaotungensis and Q. mongolica although the observed morphological variation suggested that they had remained distinct species. Across all of the differentiated leaf and reproductive traits, the hybrids expressed patterns similar to Q. liaotungensis, which may suggest dominant expression of parental characters. These results are consistent with our expectation that hybrids will be difficult to distinguish from parental species in a recent hybrid zone.     

Differential effects of host plant hybridization on herbivore community structure and grazing pressure on forest canopies

Interspecific hybridization in plants is known to have ecological effects on associated organisms. We examined the differences in insect herbivore community structure and grazing pressure on tree canopy leaves among natural hybrids and their parental oak species. We measured leaf traits, herbivore community structure, and grazing pressure on leaves of two oak species, Quercus crispula and Q. dentata, and their hybrids. The concentration of nitrogen in canopy leaves was greater in hybrids and in Q. dentata than in Q. crispula. The concentration of total phenolics was lower in hybrids than in Q. crispula. The concentration of condensed tannin was greater in hybrids than in Q. crispula. Relative herbivore abundance and species richness were greater on oak hybrids than on either parental species; herbivore species diversity and composition on hybrids were close to those on Q. crispula. Herbivore grazing pressure was lower on hybrids and Q. dentata than on Q. crispula. There was a negative correlation between herbivore grazing pressure and leaf nitrogen, suggesting that interspecific variation among oak taxa in herbivore pressure may be explained by leaf nitrogen; variation in herbivore community structure among oak taxa is likely to be controlled by polygenic leaf traits. Differing responses of (1) herbivore community structure and (2) herbivore grazing pressure to host plant hybridization may play important roles in regulating herbivore biodiversity in cool‐temperate forest canopies.     

AN EXPERIMENTAL STUDY OF VARIATION IN THE PHACELIA SERICEA COMPLEX

Gillett , Georce W. (Michigan State U., East Lansing.) An experimental study of variation in the Phacelia sericea complex. Amer. Jour. Bot. 48(1): 1–7. Illus. 1961.—The Phacelia sericea complex consists of 2 diploid (n = 11), intergrading species, P. idahoensis and P. sericea. The experimental culture of several races of this complex demonstrated that differences in pubescence, leaf shape, and flower shape persist in plants grown in a common environment. Experimental interspecific F₁ hybrids demonstrated high fertility; portrayed intermediate expressions of pubescence, leaf shape, and flower shape; and were found to be, in many cases, indistinguishable from many wild intermediates. A study of herbarium specimens revealed numerous intergrades in which pubescence, leaf shape, and flower shape are highly variable, though loosely correlated. The evidence obtained from herbarium specimens, greenhouse cultures, field investigations, chromosome studies, and experimental hybridizations suggests a hybrid origin for the wild intermediates, recognized as Phacelia sericea (Graham) A. Gray subsp. ciliosa (Rydb.) Gillett.     

Structure of herbivore communities in two oak (Quercus spp.) hybrid zones

Summary We examined patterns of density and species diversity for leaf-mining Lepidopterans and gall-forming Hymenopterans in two oak (Quercus spp.) hybrid zones: Quercus depressipes x Q. rugosa and Q. emoryi x Q. coccolobifolia. In both species complexes, hybrid hosts typically supported significantly lower densities and species diversity of parasites than did parental types. This contradicts the findings of Whitham (1989) that suggested that hybrid hosts may act as parasite sinks both in ecological and evolutionary time. We discuss features of hybrid zones that are likely to influence patterns of herbivory.     

Interspecific hybridization in natural populations ofCyrtandra (Gesneriaceae) on the Hawaiian Islands: Evidence from RAPD markers

Interspecific hybridization among Hawaiian species ofCyrtandra (Gesneriaceae) was investigated using randomly amplified polymorphic DNA (RAPD) markers. Thirty-three different primers were used to investigate interspecific hybridization for 17 different putative hybrids based on morphological intermediacy and sympatry with putative parental species. RAPD data provided evidence for the hybrid origin of all putative hybrid taxa examined in this analysis. However, the patterns in the hybrid taxa were not found to be completely additive of the patterns found in the parental species. Markers missing in the hybrid taxa can be attributed to polymorphism in the populations of the parental species and the dominant nature of inheritance for RAPD markers. Unique markers found within hybrid taxa require further explanation but do not necessarily indicate that the taxa are not of hybrid origin. The implications suggest that these interspecific hybridization events had, and continue to have, an effect on the adaptive radiation and conservation biology ofCyrtandra.     

A NEW SPECIES OF BLACK OAK (QUERCUS,SUBG. ERYTHROBALANUS,FAGACEAE) FROM THE SIERRA MADRE OCCIDENTAL,MEXICO

Quercus mcvaughii is described as a new species of black oak (subgenus Erythrobalanus) from the Sierra Madre Occidental, in northern Durango and western Chihuahua, Mexico. It is most closely related to the more southern Q. crassifolia. Sessile or subsessile hairs, usually of different sizes, on the abaxial leaf surface distinguish Q. mcvaughii from Q. crassifolia, where hairs are similar in size and conspicuously stipitate. A key is provided for distinguishing similar species in the region. Intergradation of Q. mcvaughii with Q. crassifolia, Q. hypoleucoides, and Q. sideroxyla is reported.     

Hybridization of European oaks (Quercus ilex × Q. robur) results in a mixed isoprenoid emitter type

European oaks have been reported to emit isoprene or monoterpenes derived from recently fixed photosynthetic carbon. The emission type is plant species specific and can be used as chemo‐taxonomic marker. In the present article the isoprenoid biochemical properties of mature Quercus × turneri‘Pseudoturneri’ hybrids resulting from a crossing of a Mediterranean evergreen monoterpene‐emitting species (subgenus Sclerophyllodrys; Quercus ilex L.) and an isoprene‐emitting deciduous oak species (subgenus Lepidobalanus; Quercus robur L.) are described. Both species are compared with respect to the capacity for isoprenoid synthesis and the actual isoprenoid emission pattern of different tree‐types. The analysis showed that the oak hybrid combines properties of both parental species. Furthermore, it could be shown that the enzyme activities of isoprene synthase and monoterpene synthases are reflected in the isoprenoid emission pattern of the hybrids as well as in the observed emission rates.