Comparisons of pollen and petiole anatomy in three Johrenia species. Top row (left to right): J. paucijuga petiole anatomy (A, B), pollen grain under optical microscope (C), and pollen grain viewed under SEM (D; ×3,000). Middle row: J. aromatica petiole anatomy (E, F), pollen grain under optical microscope (G), and pollen grain viewed under SEM (H; ×3,500). Bottom row: J. golestanica petiole anatomy (I, J), pollen grain under optical microscope (K), and pollen grain viewed under SEM (L; ×3,000). Abbreviations are as follows: col, collenchyma; ph, phloem; p.v.b., peripheral vascular bundle; sc, sclerenchyma; x, xylem. 

Contexts in source publication

Context 1

... pedata (Wight) P.K. Mukh. & Constance) each form separate lineages away from their congeners and the type of the genus, H. sphondylium L. The genus Malabaila Hoffm. ( = Leiotulus Ehreb.) arises from within Pastinaca . This clade of Pastinaca (with included Malabaila and the single accession identified in GenBank as Leiotulus ) is well supported (MP BS 90%; ML BS 99%; PP 1.00). Tribe Tordylieae is confirmed to include Ainsworthia Boiss., Semenovia Regel & Herder, and Synelcosciadium ( = Tordylium L.). Anisosciadium DC. is confirmed within tribe Echinophoreae. Additional members of tribe Pyramidoptereae include Muretia Boiss. ( = Elaeosticta Fenzl.), Galagania Lipsky, Hyalolaena Bunge, and Astoma DC. Tribe Careae is expanded to include Chamaesciadium C.A. Mey., Gongylosciadium Rech. f., and Hladnikia Rchb., and tribe Pimpinelleae is expanded to include Haussknechtia , Demavendia , and Zeravschania Korovin. Tribe Smyrnieae is maintained as comprising only two genera, Smyrnium L. and Lecokia DC. Analysis of six of the seven species recognized in Smyrnium indicates that the genus is strongly supported as monophyletic. Eremodaucus lehmanii Bunge is a member of tribe Pleurospermeae. morphology. — A comparison of morphological features for four species of the Cymbocarpum group ( Cymbocarpum anethoides , C. erythraeum , Kalakia marginata , Ducrosia anethifolia ) is presented in Table 1. These species are very similar in leaf morphology, but differ in their fruits. The transfer of C. marginatum to the monotypic genus Kalakia (Alava, 1987) was based on its characteris- tic thickened fruit margins, yet there are also pronounced differences between the fruits of C. anethoides and C. erythraeum . In the former, the mericarps are compressed laterally, whereas in the latter they are compressed dorsally (as they are in K. marginata ). Cymbocarpum and Kalakia show many similarities in other vegetative features (i.e., habit, stem color, stem furrows, outer petal length, and bract division). Ducrosia anethifolia is similar to Cymbocarpum and Kalakia in its petals and shape of leaf segments. However, D. anethifolia differs from the other two genera in that its mericarps are subglobular and its leaves do not wither early. In general, distinguishing among Cymbocarpum , Kalakia and Ducrosia using leaf characters alone is very difficult. Similarly, fruit characters, such as the orientation of fruit compression and type of fruit margin, are also not useful in separating these genera. A comparison of selected morphological features for seven Iranian representatives from four genera provision- ally recognized in the Cachrys group ( Diplotaenia , Prangos , Ferulago , Azilia ) is presented in Table 2. Sampling of this genus group was not comprehensive but is adequate to compare critical morphological differences between Azilia and its generic allies, as suggested by published studies. Diplotaenia cachrydifolia and D. damavandica are morphologically very similar, with noted differences between them limited to leaf segment shape and width. Inflorescences of both species are whorled, with lateral umbels bearing male and hermaphrodite flowers and central umbels bearing only hermaphrodite flowers. Diplotaenia shares with Prangos similar leaf and inflorescence characters. Indeed, D. cachrydifolia is very similar to P. ferulacea (L.) Lindl. in leaf morphology. Their fruits, however, are quite different. In Diplotaenia , the fruits are strongly dorsally compressed and have prominent dorsal ribs. In Prangos , the fruits are moderately compressed laterally and their prominent dorsal ribs are extended into wings. Ferulago stellata Boiss. is very similar to the other two genera, with its similar inflorescence structure and leaf form. In contrast, Azilia eryngioides stands apart from all other members of this group. Its leaf segments are orbicular to reniform and large with spiny margins and its inflorescences are paniculate with hermaphrodite umbels. Azilia is clearly distinguished morphologically from Diplotaenia , Prangos , and Ferulago (and indeed any other genus in the family), and any close ally based on morphological similarity is not apparent in our study. A comparison of selected morphological characters for three species of Johrenia and one species each of Johreniopsis and Holandrea is presented in Table 3, with the primary purpose of assessing the relationship of the narrow endemic Johrenia golestanica to its congeners. Johrenia golestanica is unique in its white petals, grayish-green and finely striate stems, sub-elliptic (to sub-orbicular) fruits with non-prominent dorsal ribs, and a preference for growing in disturbed areas. Johreniopsis is morphologically similar to Johrenia . Furthermore, with its yellow petals, once- to twice-pinnate leaves, and prominent mericarp ribs, Johreniopsis is morphologically more similar to Johrenia paucijuga and J. aromatica than it is to J. golestanica . Comparisons of petiole anatomy (Table 4; Fig. 3) and pollen grains (Table 5; Fig. 3) also support the separation of J. golestanica from its congeners. Unlike the other two species of Johrenia where the vascular bundles are partially or wholly surrounded by sclerenchyma cells, the vascular bundles of J. golestanica are without sclerenchyma tissue. Sclerenchyma at the base of the xylem, however, is present in Johreniopsis seseloides and Holandrea caucasica . Pollen of Johrenia , Johreniopsis , and Holandrea is of the oval-type (sensu Cerceau-Larrival, 1971) and in J. golestanica the grains are slightly smaller (in length and width) than they are in its congeners. The morphology of eight species of Dorema from Iran was examined ( D. kopetdaghense was considered a distinct species). All species are morphologically very similar, although each can be distinguished by several characters (results not shown). As examples, Dorema ammoniacum and D. aitchisonii are similar in their leaf segments (shape, pubescence, margin, width), peduncle and pedicel length and surface features, and having woolly fruits prior to ripening. They both differ from other species in the genus, however, in several of these attributes. Dorema ammoniacum and D. aitchisonii are separated by mericarp length, wing width, and degree of stem inter- node swelling (the stem is swollen in D. aitchisonii but not in D. ammoniacum ). The size of the leaves in these two species shows great variation. Dorema aucheri has similar leaf characters to the two aforementioned species, but is separated from them by fruit size and surface texture. Dorema kopetdaghense and D. hyrcanum are morphologically distinct; the former differs in its smaller leaf segments that are lanceolate with entire margins, its pubescent immature fruits, and shorter peduncles. Dorema glabrum is also distinct morphologically, with its glabrous leaf segments and long pedicels. Similarities between Dorema and Ferula species are apparent and have been presented elsewhere (e.g., Schischkin, 1951; Pimenov, 1988). A comparison of morphological features of Opopanax hispidus , O. persicus , and Smyrniopsis aucheri of the Opopanax group is presented in Table 6. These species share several morphological attributes, such as those of the leaves and inflorescences. These features include once- or twice-pinnate basal leaves, leaf segments oblong or elliptic, acute, crenate and with cartilage at their margins, hispidus leaves, type of cork on the stem, and inflorescences of male lateral umbels and central hermaphrodite umbels. There are some obvious differences between Opopanax and Smyrniopsis , such as dorsally versus laterally compressed mericarps and solid versus hol- low petioles in cross-section. The stems of O. persicus are smooth, whereas those of O. hispidus are densely hispid (especially on the basal portions), and those of S. aucheri are sparsely hispidus. Inflorescence shape in O. persicus is dense thyrsoid, in O. hispidus it is lax thyrsoid, and in S. aucheri it is conical. The limitations of nrDNA ITS sequence data in resolving tribal-level relationships within Apiaceae subfamily Apioideae have been discussed elsewhere (Downie & al., 1998; Katz-Downie & al., 1999). Their homoplastic nature, high levels of nucleotide sequence divergence in some lineages, and small size of the region all conspire to reduce the utility of these data in resolving deep-level relationships within the subfamily. A glance at the trees in Figs. 1 and 2 will show poor resolution and branch support for the most basal lineages within the Apioid superclade. Minor differences in topology are also apparent among the trees with regard to the relationships inferred among the tribes and major clades. In this study, one of our goals was to elucidate the phylogenetic placements of five groups of genera native to the Flora Iranica area. Based on the phylogenetic results, the Cymbocarpum group falls within tribe Tordylieae, the Johrenia group is placed within tribe Selineae, and the Ferula group allies with tribe Scandiceae. Unclear, however, are the tribal placements of the Cachrys and Opopanax groups. Both of these fall within the Apioid superclade, but not in any previously recognized tribe supported strongly as monophyletic on the basis of molecular phylogenetic studies (reviewed in Downie & al., 2001). To date, phylogenies of the Apioideae inferred from ITS sequence data are generally consistent with those inferred from chloroplast markers, but until the latter are available for a broad sampling of taxa from the Apioid superclade the relationships among its tribes and the tribal placements of the Cachrys and Opopanax groups cannot be resolved. The Cachrys group may constitute a new, major lineage of Apiaceae, but its recognition now as a tribe is premature in the ab- sence of supporting plastid data. The major objective of this study was to ascertain taxonomic and phylogenetic relationships within these five genus groups. Sister group relationships of each of ...

Context 2

... each form separate lineages away from their congeners and the type of the genus, H. sphondylium L. The genus Malabaila Hoffm. ( = Leiotulus Ehreb.) arises from within Pastinaca . This clade of Pastinaca (with included Malabaila and the single accession identified in GenBank as Leiotulus ) is well supported (MP BS 90%; ML BS 99%; PP 1.00). Tribe Tordylieae is confirmed to include Ainsworthia Boiss., Semenovia Regel & Herder, and Synelcosciadium ( = Tordylium L.). Anisosciadium DC. is confirmed within tribe Echinophoreae. Additional members of tribe Pyramidoptereae include Muretia Boiss. ( = Elaeosticta Fenzl.), Galagania Lipsky, Hyalolaena Bunge, and Astoma DC. Tribe Careae is expanded to include Chamaesciadium C.A. Mey., Gongylosciadium Rech. f., and Hladnikia Rchb., and tribe Pimpinelleae is expanded to include Haussknechtia , Demavendia , and Zeravschania Korovin. Tribe Smyrnieae is maintained as comprising only two genera, Smyrnium L. and Lecokia DC. Analysis of six of the seven species recognized in Smyrnium indicates that the genus is strongly supported as monophyletic. Eremodaucus lehmanii Bunge is a member of tribe Pleurospermeae. morphology. — A comparison of morphological features for four species of the Cymbocarpum group ( Cymbocarpum anethoides , C. erythraeum , Kalakia marginata , Ducrosia anethifolia ) is presented in Table 1. These species are very similar in leaf morphology, but differ in their fruits. The transfer of C. marginatum to the monotypic genus Kalakia (Alava, 1987) was based on its characteris- tic thickened fruit margins, yet there are also pronounced differences between the fruits of C. anethoides and C. erythraeum . In the former, the mericarps are compressed laterally, whereas in the latter they are compressed dorsally (as they are in K. marginata ). Cymbocarpum and Kalakia show many similarities in other vegetative features (i.e., habit, stem color, stem furrows, outer petal length, and bract division). Ducrosia anethifolia is similar to Cymbocarpum and Kalakia in its petals and shape of leaf segments. However, D. anethifolia differs from the other two genera in that its mericarps are subglobular and its leaves do not wither early. In general, distinguishing among Cymbocarpum , Kalakia and Ducrosia using leaf characters alone is very difficult. Similarly, fruit characters, such as the orientation of fruit compression and type of fruit margin, are also not useful in separating these genera. A comparison of selected morphological features for seven Iranian representatives from four genera provision- ally recognized in the Cachrys group ( Diplotaenia , Prangos , Ferulago , Azilia ) is presented in Table 2. Sampling of this genus group was not comprehensive but is adequate to compare critical morphological differences between Azilia and its generic allies, as suggested by published studies. Diplotaenia cachrydifolia and D. damavandica are morphologically very similar, with noted differences between them limited to leaf segment shape and width. Inflorescences of both species are whorled, with lateral umbels bearing male and hermaphrodite flowers and central umbels bearing only hermaphrodite flowers. Diplotaenia shares with Prangos similar leaf and inflorescence characters. Indeed, D. cachrydifolia is very similar to P. ferulacea (L.) Lindl. in leaf morphology. Their fruits, however, are quite different. In Diplotaenia , the fruits are strongly dorsally compressed and have prominent dorsal ribs. In Prangos , the fruits are moderately compressed laterally and their prominent dorsal ribs are extended into wings. Ferulago stellata Boiss. is very similar to the other two genera, with its similar inflorescence structure and leaf form. In contrast, Azilia eryngioides stands apart from all other members of this group. Its leaf segments are orbicular to reniform and large with spiny margins and its inflorescences are paniculate with hermaphrodite umbels. Azilia is clearly distinguished morphologically from Diplotaenia , Prangos , and Ferulago (and indeed any other genus in the family), and any close ally based on morphological similarity is not apparent in our study. A comparison of selected morphological characters for three species of Johrenia and one species each of Johreniopsis and Holandrea is presented in Table 3, with the primary purpose of assessing the relationship of the narrow endemic Johrenia golestanica to its congeners. Johrenia golestanica is unique in its white petals, grayish-green and finely striate stems, sub-elliptic (to sub-orbicular) fruits with non-prominent dorsal ribs, and a preference for growing in disturbed areas. Johreniopsis is morphologically similar to Johrenia . Furthermore, with its yellow petals, once- to twice-pinnate leaves, and prominent mericarp ribs, Johreniopsis is morphologically more similar to Johrenia paucijuga and J. aromatica than it is to J. golestanica . Comparisons of petiole anatomy (Table 4; Fig. 3) and pollen grains (Table 5; Fig. 3) also support the separation of J. golestanica from its congeners. Unlike the other two species of Johrenia where the vascular bundles are partially or wholly surrounded by sclerenchyma cells, the vascular bundles of J. golestanica are without sclerenchyma tissue. Sclerenchyma at the base of the xylem, however, is present in Johreniopsis seseloides and Holandrea caucasica . Pollen of Johrenia , Johreniopsis , and Holandrea is of the oval-type (sensu Cerceau-Larrival, 1971) and in J. golestanica the grains are slightly smaller (in length and width) than they are in its congeners. The morphology of eight species of Dorema from Iran was examined ( D. kopetdaghense was considered a distinct species). All species are morphologically very similar, although each can be distinguished by several characters (results not shown). As examples, Dorema ammoniacum and D. aitchisonii are similar in their leaf segments (shape, pubescence, margin, width), peduncle and pedicel length and surface features, and having woolly fruits prior to ripening. They both differ from other species in the genus, however, in several of these attributes. Dorema ammoniacum and D. aitchisonii are separated by mericarp length, wing width, and degree of stem inter- node swelling (the stem is swollen in D. aitchisonii but not in D. ammoniacum ). The size of the leaves in these two species shows great variation. Dorema aucheri has similar leaf characters to the two aforementioned species, but is separated from them by fruit size and surface texture. Dorema kopetdaghense and D. hyrcanum are morphologically distinct; the former differs in its smaller leaf segments that are lanceolate with entire margins, its pubescent immature fruits, and shorter peduncles. Dorema glabrum is also distinct morphologically, with its glabrous leaf segments and long pedicels. Similarities between Dorema and Ferula species are apparent and have been presented elsewhere (e.g., Schischkin, 1951; Pimenov, 1988). A comparison of morphological features of Opopanax hispidus , O. persicus , and Smyrniopsis aucheri of the Opopanax group is presented in Table 6. These species share several morphological attributes, such as those of the leaves and inflorescences. These features include once- or twice-pinnate basal leaves, leaf segments oblong or elliptic, acute, crenate and with cartilage at their margins, hispidus leaves, type of cork on the stem, and inflorescences of male lateral umbels and central hermaphrodite umbels. There are some obvious differences between Opopanax and Smyrniopsis , such as dorsally versus laterally compressed mericarps and solid versus hol- low petioles in cross-section. The stems of O. persicus are smooth, whereas those of O. hispidus are densely hispid (especially on the basal portions), and those of S. aucheri are sparsely hispidus. Inflorescence shape in O. persicus is dense thyrsoid, in O. hispidus it is lax thyrsoid, and in S. aucheri it is conical. The limitations of nrDNA ITS sequence data in resolving tribal-level relationships within Apiaceae subfamily Apioideae have been discussed elsewhere (Downie & al., 1998; Katz-Downie & al., 1999). Their homoplastic nature, high levels of nucleotide sequence divergence in some lineages, and small size of the region all conspire to reduce the utility of these data in resolving deep-level relationships within the subfamily. A glance at the trees in Figs. 1 and 2 will show poor resolution and branch support for the most basal lineages within the Apioid superclade. Minor differences in topology are also apparent among the trees with regard to the relationships inferred among the tribes and major clades. In this study, one of our goals was to elucidate the phylogenetic placements of five groups of genera native to the Flora Iranica area. Based on the phylogenetic results, the Cymbocarpum group falls within tribe Tordylieae, the Johrenia group is placed within tribe Selineae, and the Ferula group allies with tribe Scandiceae. Unclear, however, are the tribal placements of the Cachrys and Opopanax groups. Both of these fall within the Apioid superclade, but not in any previously recognized tribe supported strongly as monophyletic on the basis of molecular phylogenetic studies (reviewed in Downie & al., 2001). To date, phylogenies of the Apioideae inferred from ITS sequence data are generally consistent with those inferred from chloroplast markers, but until the latter are available for a broad sampling of taxa from the Apioid superclade the relationships among its tribes and the tribal placements of the Cachrys and Opopanax groups cannot be resolved. The Cachrys group may constitute a new, major lineage of Apiaceae, but its recognition now as a tribe is premature in the ab- sence of supporting plastid data. The major objective of this study was to ascertain taxonomic and phylogenetic relationships within these five genus groups. Sister group relationships of each of these five groups are not discussed ...