Palaearctic grasslands are diverse and dynamic ecosystems that are in the focus of ecology, conservation biology and agronomy. This special issue is dedicated to the biodiversity and conservation issues of Palaearctic grasslands and was initiated by the Eurasian Dry Grassland Group members attending the 14th Eurasian Dry Grassland Conference (EDGC) at Sulmona, Italy in 2018. The papers in this special issue cover a wide range of grassland ecosystems from mountain dry grasslands to lowland loess grasslands, feathergrass steppes and wet grasslands, and focus on the biodiversity values and conservation issues of Palaearctic grasslands. We believe that this compilation will contribute to a better understanding of the ecology of grasslands and support their more effective conservation.
This report summarises the activities and achievements of the Eurasian Dry Grassland Group (EDGG) from January 2018 through July 2019. During the reported period, two Eurasian Grassland Conference (EGCs) took place: the 15th EGC in Sulmona, Italy, and the 16th EGC in Graz, Austria. The 11th and 12th EDGG Field Workshops studied vegetation diversity patterns in the inner alpine valleys of Austria and Switzerland, while the 13th Field Workshop was organised in Armenia. The formerly electronic newsletter of EDGG (Bulletin of the Eurasian Dry Grassland) was transformed into a peer-reviewed international journal, called Palaearctic Grasslands, which now is attracting both scientific and photographic contributions. Furthermore, the EDGG homepage was re-constructed with a new design and content management system. The EDGG has also finalised two grassland-related Special Features during the past 1.5 years in the international journals Tuexenia and Hacquetia, and contributed with eight chapters to the book Grasslands of the World: Diversity, Management and Conservation. The vegetation-plot database GrassPlot, containing standardised multi-scale data from Palaearctic grasslands and closely connected with EDGG, has developed well, as did some other regional and national grassland-focused databases.
The upper Rhone valley in the Swiss canton of Valais is one of the driest and most continental of the inner-alpine valleys and harbours a rich xerothermic flora. We studied syntaxonomy and ecology of dry grasslands and their species richness patterns. In 2018 we recorded 28 vegetation plots (10 m2) and three nested-plot series of 0.0001 to 100 m2 on the south-facing slopes above the village of Ausserberg. Mean richness of all species ranged from 1.7 on 1 cm2 to 47.3 on 100 m2, with little contribution of bryophytes and lichens. The species-area relationship for total richness closely followed a power function. Modified TWINSPAN yielded a three-cluster solution, which could easily be matched with three orders of the class Festuco-Brometea: Stipo pulcherrimae-Festucetalia pallentis (xeric, rocky), Festucetalia valesiacae (xeric, non-rocky) and Brachypodietalia pinnati (meso-xeric). The subdivision of the xeric types into two orders is new for Swiss dry grasslands, where these types up to now had been joined in a single alliance Stipo-Poion within the Festucetalia valesiacae.
Eurasian steppes have an essential role in conserving biodiversity, but due to the huge habitat loss in the past centuries they are often preserved only in small refuges. Among such refuges are the ancient steppic burial mounds (the so called ‘kurgans’) which have a high cultural and historical importance and are also essential sites of nature conservation. Despite their high number (approximately half million) and conservational importance there is a huge lack of knowledge on the locality and conservational state of the kurgans in most regions of Eurasia. To fill this knowledge gap, we built a public database which allows to record and query basic information on their cultural values and factors (such as land cover type, threatening factors, cover of woody species) that might serve as a basis for their effective conservation. The database provides a transparent, public and easy-to-use source for conservation managers and landscape planners focussed on grassland conservation. In addition, it also provides background information for other associate disciplines and public agencies dealing with the protection of cultural heritage.
Prehistoric mounds of the Great Hungarian Plain often function as refuges for relic loess steppe vegetation and their associated fauna. The Zsolca mounds are a typical example of kurgans acting as refuges, and even though they are surrounded by agricultural land, they harbour a species rich loess grassland with an area of 0.8 ha. With a detailed field survey of their geomorphology, soil, flora and fauna, we describe the most relevant attributes of the mounds regarding their maintenance as valuable grassland habitats. We recorded 104 vascular plant species, including seven species that are protected in Hungary and two species (Echium russicum and Pulsatilla grandis) listed in the IUCN Red List and the Habitats Directive. The negative effect of the surrounding cropland was detectable in a three-metre wide zone next to the mound edge, where the naturalness of the vegetation was lower, and the frequency of weeds, ruderal species and crop plants was higher than in the central zone. The ancient man-made mounds harboured dry and warm habitats on the southern slope, while the northern slopes had higher biodiversity, due to the balanced water supplies. Both microhabitats had different assemblages of ground-dwelling invertebrates.
Seasonal variability in grasslands’ vegetation affects animals’ diet selection. We studied the seasonal changes in water buffaloes’ diet during grazing in wet grasslands in Northern Greece. We recorded each month the plant species of the vegetation and the species consumed by buffaloes. We categorized the former into three functional groups (graminoids, legumes, and forbs) and the latter into five groups (graminoids, legumes, forbs, shrubs, and trees). There were significant differences in the proportions (%) of functional groups in the vegetation and in buffaloes’ diet between seasons (χ2 test, P<0.05). Graminoids, legumes, and forbs participated in buffaloes’ diet in all seasons, while the less preferred functional groups were trees and shrubs. Buffaloes consumed ten species in all seasons but we observed the most diverse diet, in terms of plant species, in spring. The most frequently consumed species in each functional group were Cynodon dactylon (graminoids), Trifolium repens (legumes), Cichorium intybus (forbs), Rubus sp. (shrubs), and Populus sp. (trees). However, the majority of plant species in buffaloes’ diet was in very low proportions (<1%), while buffaloes did not sample at all 38 herbaceous species. Researchers need to conduct further research to understand water buffaloes’ foraging strategy regarding plants’ anti-quality characteristics.
Being an essential driving factor in dry grassland ecosystems, uncontrolled fires can cause damage to isolated natural areas. We investigated a case of a small-scale mid-summer fire in an abandoned steppe pasture in northeastern Ukraine and focused on the post-fire recovery of arthropod assemblages (mainly spiders and beetles) and vegetation pattern. The living cover of vascular plants recovered in a year, while the cover of mosses and litter remained sparse for four years. The burnt site was colonised by mobile arthropods occurring in surrounding grasslands. The fire had no significant impact on arthropod diversity or abundance, but changed their assemblage structure, namely dominant complexes and trophic guild ratio. The proportion of phytophages reduced, while that of omnivores increased. The fire destroyed the variety of the arthropod assemblages created by the patchiness of vegetation cover. In the post-fire stage they were more similar to each other than at the burnt plot in the pre- and post-fire period. Spider assemblages tended to recover their pre-fire state, while beetle assemblages retained significant differences during the entire study period.
Paper presents a systematic global review of Acer negundo, Fraxinus pennsylvanica, Ailanthus altissima, Robinia pseudoacacia invasions focusing on the Scopus and Web of Science databases. We examined the data on papers, study areas, habitat studied, topic discussed. We hypothesized that these species were studied evenly throughout their invaded ranges and, as such, indexed by international databases. We asked whether four selected species are presented evenly in publications related to their invaded ranges, and whether both selected databases cover well a content of these papers. We found 48 papers for A. negundo, 14 – for F. pennsylvanica, 83 – for A. altissima, 96 – for R. pseudoacacia. A high percentage of the studies were conducted in Central Europe and USA (for A. altissima), while Eastern Europe, Russia, Western United States were poorly represented. Most studies were conducted in forests, and focused on impacts or distribution of aliens in invaded range, and their control and management. We encountered habitat types invaded by trees, factors influencing tree invasions, consequences of invaders’ impact on ecosystems, counteracting measures. We concluded that the use only Web of Science and Scopus is not sufficient to obtain the complete data about the invasion biology.
The study presents new data on the habitat dominated by the species complex of Genista lydia/G. rumelica in Bulgaria. It is based on 129 phytocoenological relevés and provides information on the chorology, ecology and floristic structure of these communities. This habitat type occupies substrates composed by different volcanic rocks. The floristic structure is very rich in species. The phytogeographical relationships with the East Mediterranean region are considerable, which is proved by the high occurrence of floristic elements with Mediterranean or sub-Mediterranean origin. The plant life-forms analysis demonstrates that the therophytes, geophytes and chamephytes prevail in their floristic structure, which is also typical for the shrub communities in this region. During the field study this vegetation type has been mapped and its total area of occupancy has been calculated. For a first time it is proposed this habitat to be divided into three habitat sub-types due to the established differences in the environmental factors. Some recommendations have been proposed on the conservation management and also complements on the habitat‘s descriptions in EUNIS habitat classification.
The article aims to 1) describe the distribution of non-native species of vascular plants (only kenophytes, i. e. naturalized species introduced after the 15th century) in different types of forests and different biogeographical regions of Ukraine; 2) compare proportions of kenophytes in forests of different areas; 3) detect statistically significant changes in the occurrence of kenophytes over the last 80 years. The material consists of 2701 relevés sampled in 1990–2018. They were taken from Ukrainian phytocoenological publications and databases. In Ukraine, as in other European countries, the highest proportion of kenophytes (percentage of species number per relevé) is in floodplain forests (up to 9.1% in willow and poplar forests). The lowest proportion is characteristic for bog woods (0%) and most types of mountain forests. Among biogeographical regions, the highest values were found in the flatland areas of the Steppic region and the forest-steppe part of the Continental region. The most frequent species are Impatiens parviflora (predominantly in broadleaved woods, absent in relevés from the steppic biogeoregion) and Erigeron canadensis (pine woods on sand). Comparison with 1466 older relevés sampled in 1950–1989 allows us to make a conclusion that the proportion of kenophytes increased at least in one habitat type (oak-hornbeam forests).
Tamarix L. play important role in preventing deforestation in Iran. Tamarix species exhibit wide range of morphological variation therefore, the species delimitation become difficult. This is further complicated due to similarity of morphological characters in closely related species and the occurrence of inter-specific hybridization. The present study was performed to identify Tamarix species and their potential hybrids in Semnan Province of Iran. We used ITS and ISSR and 42 morphological characters for our investigation. Molecular phylogeny of the studied species and their relationship was not in agreement with the species tree of morphological characters and with taxonomic treatment of the genus. HGT tree of ITS and morphological data obtained revealed the occurrence of inter-specific hybridization or introgression between Tamarix species.
In Europe, the genus Ramonda is represented with three species: Ramonda nathaliae, Ramonda serbica and Ramonda myconi. The first two are endemic Balkan species that are distributed also in Kosovo. These species grow in limestone as well as serpentine substrates, forming chasmophytic vegetation. The species Ramonda nathaliae is found in Macedonia, Greece, Serbia and in two localities in Kosovo, in the Sharri Mountains (Luboten and Gotovushë). R. nathaliae forms the following plant associations in the serpentines of Macedonia: AsplenioRamondetum nathaliae and Scorzonero-Ramondetum nathaliae, and the AchilleoRamondetum nathaliae in limestone substrates. Ostrya carpinifolia is charateristic species in Querco pubescentis-Ostryetum carpinifoliae, Ostryo-Fagetum, Querco-Ostryetum carpinifoliae and Corylo colurnae-Ostryetum carpinifoliae. This paper presents plant communities of Ramonda nathaliae and Ostrya carpinifolia in a limestone habitat, where the proposed new plant association named RamondoOstryetum carpinifoliae ass. nova. is described. This plant community belongs to the class Quercetea pubescentis, order Quercetalia pubescentipetraeae and alliance Fraxino orniOstryion. It was found and described on the limestone substrate on Mt. Luboteni (at 960–982 m a.s.l.).
Molecular data have been increasingly used to study the phylogenetic relationships among many taxa, including scrophs. Sometimes they have provided phylogenetic reconstructions that are in conflict with morphological data leading to a re-evaluation of long-standing evolutionary hypotheses. In this paper, we review reports of the recent knowledge of the phylogenetic relationships within Scrophularieae (2011–2017). The results of these analyses led to the following conclusions. (1) Species of Scrophularia have undergone one or more Miocene migration events occurred from eastern Asia to the North America with subsequent long dispersal and diversification in three main directions. (2) Allopolyploid and aneuploid hybrid speciation between Scrophularia species can occur, so hybridization and polyploidy have an important role for history of diversification. (3) The ancestral staminode type for the genus Scrophularia seems to be a large staminode. (4) Monophyly of the genus Verbascum with respect to the genus Scrophularia is strongly supported. (5) Oreosolen, is not monophyletic, because all accessions of Oreosolen were nested within Scrophularia. We discuss methods of data collection and analysis, and we describe the areas of conflict and agreement between molecular phylogenies.
Palaearctic grasslands are diverse and dynamic ecosystems that are in the focus of ecology, conservation biology and agronomy. This special issue is dedicated to the biodiversity and conservation issues of Palaearctic grasslands and was initiated by the Eurasian Dry Grassland Group members attending the 14th Eurasian Dry Grassland Conference (EDGC) at Sulmona, Italy in 2018. The papers in this special issue cover a wide range of grassland ecosystems from mountain dry grasslands to lowland loess grasslands, feathergrass steppes and wet grasslands, and focus on the biodiversity values and conservation issues of Palaearctic grasslands. We believe that this compilation will contribute to a better understanding of the ecology of grasslands and support their more effective conservation.
This report summarises the activities and achievements of the Eurasian Dry Grassland Group (EDGG) from January 2018 through July 2019. During the reported period, two Eurasian Grassland Conference (EGCs) took place: the 15th EGC in Sulmona, Italy, and the 16th EGC in Graz, Austria. The 11th and 12th EDGG Field Workshops studied vegetation diversity patterns in the inner alpine valleys of Austria and Switzerland, while the 13th Field Workshop was organised in Armenia. The formerly electronic newsletter of EDGG (Bulletin of the Eurasian Dry Grassland) was transformed into a peer-reviewed international journal, called Palaearctic Grasslands, which now is attracting both scientific and photographic contributions. Furthermore, the EDGG homepage was re-constructed with a new design and content management system. The EDGG has also finalised two grassland-related Special Features during the past 1.5 years in the international journals Tuexenia and Hacquetia, and contributed with eight chapters to the book Grasslands of the World: Diversity, Management and Conservation. The vegetation-plot database GrassPlot, containing standardised multi-scale data from Palaearctic grasslands and closely connected with EDGG, has developed well, as did some other regional and national grassland-focused databases.
The upper Rhone valley in the Swiss canton of Valais is one of the driest and most continental of the inner-alpine valleys and harbours a rich xerothermic flora. We studied syntaxonomy and ecology of dry grasslands and their species richness patterns. In 2018 we recorded 28 vegetation plots (10 m2) and three nested-plot series of 0.0001 to 100 m2 on the south-facing slopes above the village of Ausserberg. Mean richness of all species ranged from 1.7 on 1 cm2 to 47.3 on 100 m2, with little contribution of bryophytes and lichens. The species-area relationship for total richness closely followed a power function. Modified TWINSPAN yielded a three-cluster solution, which could easily be matched with three orders of the class Festuco-Brometea: Stipo pulcherrimae-Festucetalia pallentis (xeric, rocky), Festucetalia valesiacae (xeric, non-rocky) and Brachypodietalia pinnati (meso-xeric). The subdivision of the xeric types into two orders is new for Swiss dry grasslands, where these types up to now had been joined in a single alliance Stipo-Poion within the Festucetalia valesiacae.
Eurasian steppes have an essential role in conserving biodiversity, but due to the huge habitat loss in the past centuries they are often preserved only in small refuges. Among such refuges are the ancient steppic burial mounds (the so called ‘kurgans’) which have a high cultural and historical importance and are also essential sites of nature conservation. Despite their high number (approximately half million) and conservational importance there is a huge lack of knowledge on the locality and conservational state of the kurgans in most regions of Eurasia. To fill this knowledge gap, we built a public database which allows to record and query basic information on their cultural values and factors (such as land cover type, threatening factors, cover of woody species) that might serve as a basis for their effective conservation. The database provides a transparent, public and easy-to-use source for conservation managers and landscape planners focussed on grassland conservation. In addition, it also provides background information for other associate disciplines and public agencies dealing with the protection of cultural heritage.
Prehistoric mounds of the Great Hungarian Plain often function as refuges for relic loess steppe vegetation and their associated fauna. The Zsolca mounds are a typical example of kurgans acting as refuges, and even though they are surrounded by agricultural land, they harbour a species rich loess grassland with an area of 0.8 ha. With a detailed field survey of their geomorphology, soil, flora and fauna, we describe the most relevant attributes of the mounds regarding their maintenance as valuable grassland habitats. We recorded 104 vascular plant species, including seven species that are protected in Hungary and two species (Echium russicum and Pulsatilla grandis) listed in the IUCN Red List and the Habitats Directive. The negative effect of the surrounding cropland was detectable in a three-metre wide zone next to the mound edge, where the naturalness of the vegetation was lower, and the frequency of weeds, ruderal species and crop plants was higher than in the central zone. The ancient man-made mounds harboured dry and warm habitats on the southern slope, while the northern slopes had higher biodiversity, due to the balanced water supplies. Both microhabitats had different assemblages of ground-dwelling invertebrates.
Seasonal variability in grasslands’ vegetation affects animals’ diet selection. We studied the seasonal changes in water buffaloes’ diet during grazing in wet grasslands in Northern Greece. We recorded each month the plant species of the vegetation and the species consumed by buffaloes. We categorized the former into three functional groups (graminoids, legumes, and forbs) and the latter into five groups (graminoids, legumes, forbs, shrubs, and trees). There were significant differences in the proportions (%) of functional groups in the vegetation and in buffaloes’ diet between seasons (χ2 test, P<0.05). Graminoids, legumes, and forbs participated in buffaloes’ diet in all seasons, while the less preferred functional groups were trees and shrubs. Buffaloes consumed ten species in all seasons but we observed the most diverse diet, in terms of plant species, in spring. The most frequently consumed species in each functional group were Cynodon dactylon (graminoids), Trifolium repens (legumes), Cichorium intybus (forbs), Rubus sp. (shrubs), and Populus sp. (trees). However, the majority of plant species in buffaloes’ diet was in very low proportions (<1%), while buffaloes did not sample at all 38 herbaceous species. Researchers need to conduct further research to understand water buffaloes’ foraging strategy regarding plants’ anti-quality characteristics.
Being an essential driving factor in dry grassland ecosystems, uncontrolled fires can cause damage to isolated natural areas. We investigated a case of a small-scale mid-summer fire in an abandoned steppe pasture in northeastern Ukraine and focused on the post-fire recovery of arthropod assemblages (mainly spiders and beetles) and vegetation pattern. The living cover of vascular plants recovered in a year, while the cover of mosses and litter remained sparse for four years. The burnt site was colonised by mobile arthropods occurring in surrounding grasslands. The fire had no significant impact on arthropod diversity or abundance, but changed their assemblage structure, namely dominant complexes and trophic guild ratio. The proportion of phytophages reduced, while that of omnivores increased. The fire destroyed the variety of the arthropod assemblages created by the patchiness of vegetation cover. In the post-fire stage they were more similar to each other than at the burnt plot in the pre- and post-fire period. Spider assemblages tended to recover their pre-fire state, while beetle assemblages retained significant differences during the entire study period.
Paper presents a systematic global review of Acer negundo, Fraxinus pennsylvanica, Ailanthus altissima, Robinia pseudoacacia invasions focusing on the Scopus and Web of Science databases. We examined the data on papers, study areas, habitat studied, topic discussed. We hypothesized that these species were studied evenly throughout their invaded ranges and, as such, indexed by international databases. We asked whether four selected species are presented evenly in publications related to their invaded ranges, and whether both selected databases cover well a content of these papers. We found 48 papers for A. negundo, 14 – for F. pennsylvanica, 83 – for A. altissima, 96 – for R. pseudoacacia. A high percentage of the studies were conducted in Central Europe and USA (for A. altissima), while Eastern Europe, Russia, Western United States were poorly represented. Most studies were conducted in forests, and focused on impacts or distribution of aliens in invaded range, and their control and management. We encountered habitat types invaded by trees, factors influencing tree invasions, consequences of invaders’ impact on ecosystems, counteracting measures. We concluded that the use only Web of Science and Scopus is not sufficient to obtain the complete data about the invasion biology.
The study presents new data on the habitat dominated by the species complex of Genista lydia/G. rumelica in Bulgaria. It is based on 129 phytocoenological relevés and provides information on the chorology, ecology and floristic structure of these communities. This habitat type occupies substrates composed by different volcanic rocks. The floristic structure is very rich in species. The phytogeographical relationships with the East Mediterranean region are considerable, which is proved by the high occurrence of floristic elements with Mediterranean or sub-Mediterranean origin. The plant life-forms analysis demonstrates that the therophytes, geophytes and chamephytes prevail in their floristic structure, which is also typical for the shrub communities in this region. During the field study this vegetation type has been mapped and its total area of occupancy has been calculated. For a first time it is proposed this habitat to be divided into three habitat sub-types due to the established differences in the environmental factors. Some recommendations have been proposed on the conservation management and also complements on the habitat‘s descriptions in EUNIS habitat classification.
The article aims to 1) describe the distribution of non-native species of vascular plants (only kenophytes, i. e. naturalized species introduced after the 15th century) in different types of forests and different biogeographical regions of Ukraine; 2) compare proportions of kenophytes in forests of different areas; 3) detect statistically significant changes in the occurrence of kenophytes over the last 80 years. The material consists of 2701 relevés sampled in 1990–2018. They were taken from Ukrainian phytocoenological publications and databases. In Ukraine, as in other European countries, the highest proportion of kenophytes (percentage of species number per relevé) is in floodplain forests (up to 9.1% in willow and poplar forests). The lowest proportion is characteristic for bog woods (0%) and most types of mountain forests. Among biogeographical regions, the highest values were found in the flatland areas of the Steppic region and the forest-steppe part of the Continental region. The most frequent species are Impatiens parviflora (predominantly in broadleaved woods, absent in relevés from the steppic biogeoregion) and Erigeron canadensis (pine woods on sand). Comparison with 1466 older relevés sampled in 1950–1989 allows us to make a conclusion that the proportion of kenophytes increased at least in one habitat type (oak-hornbeam forests).
Tamarix L. play important role in preventing deforestation in Iran. Tamarix species exhibit wide range of morphological variation therefore, the species delimitation become difficult. This is further complicated due to similarity of morphological characters in closely related species and the occurrence of inter-specific hybridization. The present study was performed to identify Tamarix species and their potential hybrids in Semnan Province of Iran. We used ITS and ISSR and 42 morphological characters for our investigation. Molecular phylogeny of the studied species and their relationship was not in agreement with the species tree of morphological characters and with taxonomic treatment of the genus. HGT tree of ITS and morphological data obtained revealed the occurrence of inter-specific hybridization or introgression between Tamarix species.
In Europe, the genus Ramonda is represented with three species: Ramonda nathaliae, Ramonda serbica and Ramonda myconi. The first two are endemic Balkan species that are distributed also in Kosovo. These species grow in limestone as well as serpentine substrates, forming chasmophytic vegetation. The species Ramonda nathaliae is found in Macedonia, Greece, Serbia and in two localities in Kosovo, in the Sharri Mountains (Luboten and Gotovushë). R. nathaliae forms the following plant associations in the serpentines of Macedonia: AsplenioRamondetum nathaliae and Scorzonero-Ramondetum nathaliae, and the AchilleoRamondetum nathaliae in limestone substrates. Ostrya carpinifolia is charateristic species in Querco pubescentis-Ostryetum carpinifoliae, Ostryo-Fagetum, Querco-Ostryetum carpinifoliae and Corylo colurnae-Ostryetum carpinifoliae. This paper presents plant communities of Ramonda nathaliae and Ostrya carpinifolia in a limestone habitat, where the proposed new plant association named RamondoOstryetum carpinifoliae ass. nova. is described. This plant community belongs to the class Quercetea pubescentis, order Quercetalia pubescentipetraeae and alliance Fraxino orniOstryion. It was found and described on the limestone substrate on Mt. Luboteni (at 960–982 m a.s.l.).
Molecular data have been increasingly used to study the phylogenetic relationships among many taxa, including scrophs. Sometimes they have provided phylogenetic reconstructions that are in conflict with morphological data leading to a re-evaluation of long-standing evolutionary hypotheses. In this paper, we review reports of the recent knowledge of the phylogenetic relationships within Scrophularieae (2011–2017). The results of these analyses led to the following conclusions. (1) Species of Scrophularia have undergone one or more Miocene migration events occurred from eastern Asia to the North America with subsequent long dispersal and diversification in three main directions. (2) Allopolyploid and aneuploid hybrid speciation between Scrophularia species can occur, so hybridization and polyploidy have an important role for history of diversification. (3) The ancestral staminode type for the genus Scrophularia seems to be a large staminode. (4) Monophyly of the genus Verbascum with respect to the genus Scrophularia is strongly supported. (5) Oreosolen, is not monophyletic, because all accessions of Oreosolen were nested within Scrophularia. We discuss methods of data collection and analysis, and we describe the areas of conflict and agreement between molecular phylogenies.