Introduction

Tribe Pimpinelleae Spreng. includes the genera Aphanopleura Boiss., Arafoe Pimenov & Lavrova, Bubon L., Demavendia Pimenov, Frommia H.Wolff, Haussknechtia Boiss., Nothosmyrnium Miq., Phellolophium Baker, Pimpinella L., Psammogeton Edgew., and Zeravschania Korovin (Downie et al. 2010). It also includes segregates from other genera, such as Cryptotaenia africana Drude, Physospermopsis cuneata H.Wolff, and two species of Trachyspermum Link (T. scaberulum (Franch.) H.Wolff and T. triradiatum H.Wolff; Downie et al. 2010). Six genera of Pimpinelleae occur in Iran (Aphanopleura, Demavendia, Haussknechtia, Pimpinella, Psammogeton, and Zeravschania), of which only Demavendia and Haussknechtia are endemic (Hedge et al. 1987; Mozaffarian 2007; Mousavi et al. 2021). The members of Pimpinelleae in Iran are morphologically diverse and cannot be identified easily, although most species have small fruits (shorter than 5 mm) covered by glochidiate hairs (except in Demavendia and Zeravschania where the fruits are glabrous). Representatives of the tribe have been included in several molecular phylogenetic studies using nuclear ribosomal DNA internal transcribed spacer (nrDNA ITS) sequences (Downie et al. 2000a, b, 2001; Valiejo-Roman et al. 2006; Spalik and Downie 2007; Ajani et al. 2008; Zhou et al. 2008; Downie et al. 2010; Magee et al. 2010; Wang et al. 2014; Fernández Prieto et al. 2018; Mousavi et al. 2021). However, in all of these studies, several Iranian genera were incompletely sampled and the available results do not support the monophyly of four genera in their current circumscriptions: Aphanopleura (Fig. 1a–b), Pimpinella, Psammogeton (Fig. 1f–g), and Zeravschania (Fig. 1l–o).

Fig. 1
figure 1

Representative species of Iranian genera of tribe Pimpinelleae under study in their natural habitats. a, b, Aphanopleura trachysperma; c–e, Demavendia pastinacifolia; f, g, Psammogeton canescens; h–k, Trachyspermum copticum; and l–o, Zeravschania aucheri. All images by V.M.

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Pimpinella is the largest genus of the tribe, includes about 150 species, and is taxonomically the most challenging. The genus is widely distributed in the Old World through Europe, Africa, and Asia, with the majority of species (ca. 125 spp.) occurring in Asia (Wolff 1921; Pimenov and Leonov 1993, 2004; Pu and Watson 2005). Members of the genus are characterized by cordate-ovoid or oblong-ovoid leaves and laterally compressed mericarps constricted at their commissures, each bearing five filiform ribs (Pu and Watson 2005). Pimpinella sensu lato (s.l.) is highly polyphyletic, with its members assigned to no less than seven tribes and other major clades within Apiaceae (Downie et al. 2010; Wang et al. 2014; Fernández Prieto et al. 2018). Nineteen species of Pimpinella occur in Iran, of which six (P. tragioides Boiss., P. deverroides Boiss., P. pastinacifolia H.Wolff, P. khorasanica Engstrand, P. anisactis Rech.f. and P. khayyamii Mozaff.) are endemic (Hedge et al. 1987; Mozaffarian 2007; Khajepiri et al. 2010). The relationships among many of these SW Asian Pimpinella species have been reported previously (Tabanca et al. 2005; Fereidounfar et al. 2016; Khajepiri et al. 2010). Fruit anatomical studies of Iranian Pimpinella confirmed differences between P. anthriscoides Boiss. and other species of Pimpinella and resulted in the placement of this species into the new genus Pseudopimpinella F.Ghahrem., Khajepiri & Mozaff. [as Pseudopimpinella anthriscoides (Boiss.) F.Ghahrem., Khajepiri & Mozaff.; Khajepiri et al. 2010]. Pimpinella anthriscoides has been transferred to the genus Aegopodium L. as A. tribracteolatum Schmalh. and P. anthriscoides var. cruciata (Bornm. & H.Wolff) V.A.Matthew has been transferred to Tamamschjanella Pimenov & Kljuykov as T. cruciata (Bornm. & H.Wolff) Pimenov & Zakharova (Zakharova et al. 2012). The monotypic Iranian genus Opsicarpium Mozaff. and the polytypic genus Reutera Boiss. have been merged recently with Pimpinella (Fereidounfar et al. 2016).

Aphanopleura and Psammogeton each include five species distributed mainly in the Iranian highlands (Hedge et al. 1987). They are morphologically similar, being small annual plants having white petals and laterally compressed mericarps covered by trichomes (Hedge et al. 1987). Demavendia is a monotypic genus [D. pastinacifolia (Boiss. & Hausskn.) Pimenov] endemic to Iran and Turkmenistan. Zeravschania includes nine species distributed in Iran, eastern Turkmenistan, Afghanistan, and Pakistan (Hedge et al. 1987; Mozaffarian 2007; Pimenov et al. 2007). Demavendia and Zeravschania possess similar membranous bracts and bracteoles and differ by their leaf shape (Hedge et al. 1987). Haussknechtia is a monotypic genus (H. elymaitica Boiss.) characterized by globose ultimate umbels (Hedge et al. 1987; Mozaffarian 2007). The Iranian genera Aphanopleura, Demavendia (Fig. 1c–e), Haussknechtia, Psammogeton, and Zeravschania have heretofore been poorly investigated phylogenetically. Previous phylogenetic studies suggested the non-monophyly of Aphanopleura, Psammogeton, and Zeravschania, and revealed the placement of these taxa along with Demavendia and Haussknechtia (and the Asian genus Nothosmyrnium Miq.) as basal branching lineages within the tribe (Spalik and Downie 2007; Wang et al. 2014; Fernández Prieto et al. 2018).

The main aim of this study is to investigate the monophyly and phylogenetic placements of the Iranian genera assigned to tribe Pimpinelleae, particularly those constituting the most basal branching lineages within the tribe that were underrepresented in previous molecular phylogenetic analyses (i.e., Aphanopleura, Demavendia, Haussknechtia, Psammogeton, and Zeravschania). We also highlight the morphological characters supporting each of the resolved clades containing these taxa.

Materials and methods

Sampling strategy and marker selection

We included all genera but one previously assigned to tribe Pimpinelleae and many species from throughout the entire range of each genus. The genus Bubon, reinstated with the single species B. macedonicum L. and separated from Athamanta L. in tribe Scandiceae by Spalik et al. (2001), could not be included because material was unavailable. We included representatives from most tribes and other major clades of Apiaceae subfamily Apioideae (Downie et al. 2010) and rooted all trees with Sanicula epipactis E.H.L. Krause (subfamily Saniculoideae). The nrDNA ITS region, comprising ITS1, 5.8S rDNA, and ITS2, and the plastid rps16 intron were selected as markers for phylogenetic analyses. These markers were suggested as effective and informative for tribal- and generic-level assignments in Apiaceae (Downie et al. 2010). We newly generated the sequences of 10 taxa for ITS and nine taxa for rps16 from the following Iranian species: Aphanopleura breviseta (Boiss.) Heywood & Jury, A. leptoclada Lipsky, A. trachysperma Boiss., Demavendia pastinacifolia (Boiss. & Hausskn.) Pimenov, Opopanax armeniacus Bordz., Psammogeton stocksii (Boiss.) Nasir, Trachyspermum ammi (L.) Sprague, T. copticum (L.) Link, Zeravschania membranacea (Boiss.) Pimenov, and Z. pauciradiata (Tamamsch.) Pimenov. We did not have access to material of Aphanopleura zangelanica Gogina & Matsenko because it is not distributed in Iran. We downloaded all available ITS and rps16 intron sequences of Pimpinelleae from GenBank and created a data matrix for each marker. The final nrDNA ITS data set included 169 accessions representing 49 genera (including the aforementioned 10 species not analyzed previously) and the cpDNA rps16 data set included 37 accessions representing 24 genera. The number of included taxa in the rps16 data set was considerably smaller than that of ITS due to failed PCR or sequencing efforts for the rps16 region and the preponderance of ITS sequences available in GenBank. The accessions included in this study, including voucher information for the newly generated sequences, are presented in Appendix. A list of all new combinations proposed in this study is presented in Table 2.

DNA extractions, PCR amplifications, and sequencing

Total genomic DNA was extracted from dried leaves removed from herbarium sheets held in the herbarium of the Research Institute of Forests and Rangelands, Tehran (TARI) using a NucleoSpin Plant DNA Extraction kit (Macherey–Nagel, Düren, Germany) according to the manufacturer’s protocol. For PCR-amplification of the nrDNA ITS region, we used primer pairs ITS4 and ITS5 (White et al. 1990). The cpDNA rps16 intron region was amplified using primer pairs rpsF and rpsR3R (Oxelman et al. 1997; Petri and Oxelman 2011; Kool et al. 2012). PCR amplifications were performed in T-Personal 48 (Biometra, Göttingen, Germany), Primus 96 plus (MWG: Biotech, Ebersberg, Germany), or 2720 (Applied Biosystems, Carlsbad, California, U.S.A.) thermocyclers. Cycle sequencing was performed using the BigDye Terminator v.3.1 Cycle Sequencing kit (Applied Biosystems). PCR products were sequenced bi-directionally using an ABI 3730 DNA Analyzer 48-well capillary sequencer (Applied Biosystems).

Alignment and phylogenetic analyses

The sequences were aligned using default parameters in Mafft v.7 (Katoh and Standley 2013), and alignment errors were corrected using Mesquite 2.75 (Maddison and Maddison 2011). Bayesian inference (BI) and maximum parsimony (MP) analyses were done for each data set on the CIPRES server (Miller et al. 2010). Before running the BI analysis, the optimal substitution models were estimated using the Akaike information criterion (AIC) in jModelTest v.0.1.1 (Posada 2008). The general time reversible model of nucleotide substitution with gamma-shaped rate variation and a proportion of invariable sites (GTR + I + G) was the estimated best-fit model for ITS and GTR + G was the estimated best-fit model for rps16. For BI, we used MrBayes v.3.2.6 (Ronquist and Huelsenbeck 2003) with 20 million generations for each data set. Trees were sampled every 1000 generations with the default of three “heated” and one “cold” chain. According to the results obtained from Tracer v.1.7.1 (Rambaut et al. 2018), the burn-in values were 10% and 13% for ITS and rps16 datasets, respectively. The effective sample sizes (ESS) were 438.3 and 271.8 for ITS and rps16 datasets, respectively. The obtained trees from each data set were summarized using a 50% majority-rule consensus tree. Bayesian posterior probability (PP) values were presented on the trees. For MP, we used PAUP* v.4.0b10 (Swofford 2003). Characters were equally weighted, the heuristic search was set with random sequence addition, tree-bisection-reconnection branch swapping, 50 random addition sequence replicates, and the maxtree option set to 20,000. The obtained trees were summarized using a strict consensus tree. Bootstrap (BS) values were calculated using heuristic searches with the bootstrap nreps set at 5,000. Additional details describing the alignment and phylogenetic strategies and analyses used are presented in Mousavi et al. (2021).

Results

A summary of alignment characteristics and parsimony statistics for the ITS and rps16 data sets is presented in Table 1. The ITS matrix had over 2.5 times as many parsimony-informative characters as did the rps16 intron matrix. We present only the trees obtained from BI analyses of these data sets. Though the number of accessions differed considerably between these two data sets, their results were largely congruent in revealing the groupings of the Iranian genera under study, their non-monophyly, and the placement of A. leptoclada in a clade within Pimpinella saxifraga L., the nomenclatural type of Pimpinella. The ITS and rps16 alignments are available in the Online Resources 1, 2.

Table 1 Alignment characteristics and parsimony statistics for the ITS and rps16 data sets used in this study
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Although the tribe Pimpinelleae comprised a highly supported, monophyletic group in the ITS phylogeny (PP = 1, BS = 98, Clades A-F, Fig. 2), it was not resolved as monophyletic in the rps16 trees (Fig. 3). In both sets of trees, several clades can be recognized in Pimpinelleae. The Iranian species of Pimpinella fell within the Pimpinella core group (i.e., Clades E and F). In the ITS trees, Clade A had strong support (PP = 1, BS = 98) and included members of Aphanopleura, Nothosmyrnium, Psammogeton, and Trachyspermum. Neither Aphanopleura, Psammogeton, nor Trachyspermum resolved as monophyletic. Clade B contained representatives of the Iranian genera Demavendia, Haussknechtia, and Zeravschania. Demavendia and Haussknechtia comprised monophyletic sister groups, with this clade arising from within a paraphyletic Zeravschania. The single accession of A. leptoclada allied weakly with Pimpinella procumbens H.Wolff in Clade F. Clades C-F comprised Pimpinella species already attributable to tribe Pimpinelleae in previous studies, plus all examined accessions of Physospermopsis H.Wolff, the Afro-Malagasian members of the genera Frommia, Phellolophium and Cryptotaenia DC., and Arafoe. Pimpinella saxifraga, the type of the genus, occurred in Clade F. Twelve additional species of Pimpinella (14 accessions) fell outside of tribe Pimpinelleae, none of which occur in Iran.

Fig. 2
figure 2

Majority-rule consensus tree inferred from Bayesian Inference analysis of nrDNA ITS sequences. Numbers above the branches indicate posterior probability values; those below branches indicate MP bootstrap values for similar clades occurring in the MP strict consensus tree. The major clades recognized in tribe Pimpinelleae (Clades A-F) comprise a monophyletic group. The relationships of those accessions indicated in red are those highlighted in the present study

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Fig. 3
figure 3

Majority-rule consensus tree inferred from Bayesian Inference analysis of plastid rps16 sequences. Numbers above the branches indicate posterior probability values; those below branches indicate MP bootstrap values for similar clades occurring in the MP strict consensus tree. Tribe Pimpinelleae (Clades A-B and D-F; the same as those presented in Fig. 2) does not comprise a monophyletic group, as members of tribe Pyramidoptereae (incl. Trachyspermum copticum) and several other members of tribes Apieae, Careae, and Selineae and the Cachrys and Conium clades occur within

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In the rps16 trees, the placements of the Iranian genera were similar, with Psammogeton stocksii arising from within a paraphyletic Aphanopleura and this clade sister group to Nothosmyrnium (Clade A) and Demavendia arising from within a paraphyletic Zeravschania (Clade B). Aphanopleura leptoclada was sister to the clade of Pimpinella anisum L. + P. saxifraga.

Discussion

Though Pimpinella is a polyphyletic genus, with its species occurring in at least seven different tribes and other major clades of subfamily Apioideae (Downie et al. 2010; Wang et al. 2014; Fernández Prieto et al. 2018), the Iranian species of Pimpinella examined herein all fall in the same well-supported group as P. saxifraga, the type of the genus. This group reflects the core Pimpinella group (Fernández Prieto et al. 2018) and is reflected in the current study by Clades E and F. The Iranian genera Aphanopleura, Demavendia, Haussknechtia, Psammogeton, and Zeravschania, along with Nothosmyrnium and several species of Trachyspermum, constitute basally branching lineages of tribe Pimpinelleae in the ITS trees (Clades A and B). Based on the phylogenetic results obtained herein, we recircumscribe Psammogeton to include six new combinations and transfer Aphanopleura leptoclada to Pimpinella (Table 2). We also highlight the morphological characters supporting each of the resolved clades containing these taxa.

Table 2 Names in Pimpinella and Psammogeton recognized as a result of this study
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Aphanopleura and Psammogeton

Aphanopleura is a genus of five species (A. breviseta, A. capillifolia Lipsky, A. leptoclada Lipsky, A. trachysperma, and A. zangelanica). Aphanopleura zangelanica was not included in our study. These species are distributed in Central and SW Asia, sometimes reaching China (2 spp.), but mostly they occur in Iran, Afghanistan, Turkmenistan, and Caucasus (Hedge et al. 1987). The plants are characterized by an annual life form, small and thin stems, ternate (3-segmented) leaves, the presence of bracts and bracteoles, a clearly dentate calyx, white to red petals, and mericarps ovate or almost circular, laterally compressed, and covered by papillate, clavate or capitellate trichomes. Previous molecular phylogenetic investigations of Aphanopleura only clarified its placement in Pimpinelleae, not its interspecific relationships (Spalik and Downie 2007; Zhou et al. 2008; Downie et al. 2010; Wang et al. 2014; Fernández Prieto et al. 2018). According to our results, Aphanopleura is a polyphyletic genus, with most of its species nested within Clade A. The type of genus, A. trachysperma, presents a sister group relationship with Nothosmyrnium japonicum Miq., while the other species of Aphanopleura within this clade assemble with Trachyspermum or Psammogeton. Aphanopleura leptoclada is nested within Clade F, supporting the hypothesis that its annual habit and other correlated morphological features have evolved independently in tribe Pimpinelleae. We transfer A. leptoclada into Pimpinella (see “Taxonomic Treatment” section), while acknowledging that the annual habit of this species is rare in Pimpinella.

Psammogeton includes five species distributed in Central and SW Asia, mainly in the Iranian Highlands (Hedge et al. 1987), and all species were included herein. Its members are characterized by an annual life form, bi-ternate leaves with lanceolate or cuneate ultimate segments, the presence of bracts and bracteoles, white to red petals, and oblong, slightly laterally compressed mericarps densely covered by a glochidiate indumentum. The geographical distribution and morphological features of Aphanopleura and Psammogeton are very similar, with only the mericarp indumentum used for their separation (Boissier 1872; Hedge et al. 1987). In this study, Psammogeton forms a robust clade in Clade A, along with Aphanopleura breviseta and A. capillifolia. These two Aphanopleura species show a closer relationship with Psammogeton than they do to the type species of Aphanopleura (A. trachysperma). For this reason, we transfer A. capillifolia into Psammogeton and resurrect P. brevisetum Boiss. Aphanopleura breviseta had been previously treated as a synonym of P. brevisetum Boiss. (Hedge et al. 1987; Mozaffarian 2007).

Psammogeton, as circumscribed herein, includes species of Trachyspermum and Aphanopleura. This group is characterized by an annual or occasionally perennial habit, ultimate leaf segments mostly linear, bracts and bracteoles absent or small (shorter than 3 mm), and ovoid-globose mericarps covered by a diverse array of trichrome types (i.e., clavate, capitellate, tubular papillose, T-shaped, glochidiate, inflated vesicles, tubercles, or scales). Aphanopleura trachysperma, the type of the genus, is separate from the Psammogeton clade and is sister to Nothosmyrnium. It is distinct from members of the Psammogeton clade primarily by its much longer and conspicuous involucral bracts.

Trachyspermum

Trachyspermum includes some 12 to 15 species (Pimenov and Leonov 1993; Downie et al. 2000c; Wang et al. 2014). Trachyspermum copticum (L.) Link (a synonym of T. ammi (L.) Sprague) is the type of the genus (Fig. 1h–k). Trachyspermum is another taxonomically challenging genus (Pimenov and Leonov 1993). The genus was originally described based on a few general characters (Link 1821), such as pinnate involucral leaves, a striate, finely muricate mericarp having a short rough indumentum, a 5-merous and rudimentary calyx, a withering style, and a conical stylopodium, resulting in a vague circumscription. The genus is distributed worldwide, but predominantly occurs in Asia. The Iranian Highlands is a diversification center of the genus and harbors ten species including the newly described T. reginei Ajani & Mozzaff. (Hedge et al. 1987; Ajani and Mozaffarian 2019). The description of the genus was revised in Flora Iranica (Hedge et al. 1987) to include plants having the following features: annual or perennial habit, 1 to 3-pinnate leaves with linear ultimate segments, distinct calyx teeth, white petals incurved at apex, ovate or elliptic mericarps with prominent obtuse ribs and narrow at apex, and densely covered by white vesicular, papillose, T-shape or setulose indumentum (each type of indumentum characterizes a separate species), and a depressed or conical stylopodium. The members of Trachyspermum are only obscurely distinguishable from Aphanopleura, Pimpinella, and Psammogeton, with the main differences among them associated with the mericarp indumentum. Pimpinella possesses pilose, papillose or tuberculate trichomes, Aphanopleura (in its traditional circumscription, sensu Hedge et al. 1987) has clavate trichomes, Psammogeton bears glochidiate or martelliform trichomes at its apex, and Trachyspermum has papillose mericarps (Hedge et al. 1987). In addition, the perennial species of Trachyspermum are morphologically similar to Pimpinella, differing only in their number of vallecular vittae (which are 2‒4 in Pimpinella and one in Trachyspermum; Hedge et al. 1987).

Very few species of Trachyspermum were included in previous molecular phylogenetic studies, but not the type species T. copticum (Downie et al. 2000c, 2010; Zhou et al. 2008). These previous studies, though based on a limited sampling of species, rendered Trachyspermum as polyphyletic, with its members placed in three different tribes: T. aethusifolium Chiov. in Echinophoreae; T. ammi in Pyramidoptereae; and T. scaberulum and T. triradiatum H.Wolff in Pimpinelleae (Downie et al. 2010). The latter two species were recently united under Pimpinella scaberula (Franch.) H.Boissieu (Pimenov 2017) and in our study are placed in Clade E alongside members of Pimpinella. This placement is in accordance with previous studies (Magee et al. 2010; Wang et al. 2014; Pimenov 2017; Fernández Prieto et al. 2018).

Some species assigned previously to Trachyspermum, viz. T. roxburghianum (DC.) H.Wolff (= T. involucratum (Roxb.) H.Wolff), T. anethifolium H.Wolff, T. microcarpum Hedge, Lamond & Rech.f., T. papillare (Boiss.) Hedge, Lamond & Rech.f., and T. paktianum Hedge, Lamond & Rech.f., are nested within Clade A and show close affinity to Psammogeton. As a consequence, we transfer these species to Psammogeton. Trachyspermum copticum, a synonym of T. ammi (Menglan et al. 2005) and the type of the genus, is placed among the members of tribe Pyramidoptereae. Accordingly, the genus Trachyspermum is circumscribed narrowly.

Demavendia, Haussknechtia, and Zeravschania

Zeravschania, as circumscribed currently, includes nine species in SW Asia (Pimenov et al. 2007). It was originally described as a monotypic genus from the western Pamir-Alai mountains in Central Asia (Korovin 1948). The type of the genus, Z. regeliana Korovin, was not sufficiently described originally due to its lack of mature fruits. Subsequent morphological studies have shown that its mericarps are dorsally compressed and possess filiform dorsal ribs beside marginal ones that develop into wings (Hedge et al. 1987; Mozaffarian 2007). The most important characteristics of the genus are its white membranous and broadly ovate or lanceolate bracts and bracteoles. The phylogenies presented herein show Zeravschania as paraphyletic (Clade B), with all examined accessions of Demavendia and Haussknechtia nested within. To recognize a monophyletic Zeravschania, one treatment would be to include members of Demavendia and Haussknechtia in a broadly circumscribed Zeravschania. The monotypic genus Demavendia [D. pastinacifolia (Boiss. & Hausskn.) Pimenov] is endemic to Iran and also bears white, membranous bracts. The species currently placed in Demavendia and Zeravschania were previously treated as members of Peucedanum [unranked] Membranacea Boiss. which was characterized by such membranous bracts and bracteoles (Boissier 1872). Leaf shape and margin provide important differences between these two genera. Demavendia possesses large oblong leaf segments with irregularly dentate margins, whereas Zeravschania has ovate, ovate-oblong or lanceolate leaf segments which are 2‒4-ternate at its ultimate segments. Sister group to Demavendia and Zeravchania in the ITS trees is the monotypic genus Haussknechtia Boiss. (H. elymaitica Boiss.), a species characterized by tall plants up to 2 m or more, having sessile, very dense, and globose ultimate umbels (Hedge et al. 1987). We refrain herein from presenting a new formal taxonomic treatment of this group, in which Zeravschania is circumscribed much wider to include members of both Haussknechtia and Demavendia, due to the low resolution of relationships within the corresponding clade. Demavendia and Zeravschania are obviously closely related, especially upon considering their phylogenetic placement and morphological similarities. Haussknechtia also shows a close phylogenetic relationship to these genera, although its morphology is quite distinct. Further molecular phylogenetic investigations involving more informative markers should shed light on the relationships among Demavendia, Haussknechtia, and Zeravschania.

Taxonomic treatment

Psammogeton is circumscribed broadly to include species previously assigned to Aphanopleura and Trachyspermum. We increase the number of species under Psammogeton to 12 and reduce the number of species under Aphanopleura to two. In total, we recognize seven new combinations and resurrect one name.

New combinations

Pimpinella leptoclada (Aitch. & Hemsl.) Mousavi, Mozaff. & Zarre, comb. nov. ≡ Carum leptocladum Aitch. & Hemsl., Trans. Linn. Soc. Bot. Ser. 2, 3: 66. 1888–1889. ≡ Aphanopleura leptoclada (Aitch. & Hemsl.) Lipsky, Izv. Imp. Akad. Nauk V, 4: 377. 1898. ≡ Carum aphanopleura Koso-Pol., Not. Syst. Herb. Hort. Bot. Petrop. 3, 18: 70. 1922. —LECTOTYPE (designated here): Afghanistan, Herat Prov., Hari-rud valley, 5 Jun 1885, J.E.T. Aitchison 603 (G barcode G00359737; isolectotypes: FI barcode FI014839, GH barcode GH00075662, W barcode W1963-0000628). —SYNTYPES: Afghanistan, 5 Jun 1885, J.E.T. Aitchison 603 (P barcode P00834708); Afghanistan, Herat, Hari-rud valley, 1884, J.E.T. Aitchison 603 (C barcode C10008329).

Diagnosis: Annual plants, stem puberulent or glabrous, 10–40 cm tall; leaves ternate with linear or oblong-linear ultimate segments; ultimate umbels long pedunculated; bracts 2‒5, linear-lanceolate, membranous, scabrous at margins; bracteoles present; umbellules numerous with filiform rays; petals white or red, dorsally pilose; mericarps ovoid-globose or ovoid, covered by dense claviform or capitate indumentum.

Distribution: Iran (mostly in the northeast), Afghanistan, Tajikistan, Turkmenistan, and Uzbekistan.

Psammogeton anethifolium (D.Don) Mousavi, Mozaff. & Zarre, comb. nov. ≡ Pimpinella anethifolia D.Don, Prodr. Fl. Nepal. 184. 1825. ≡ Athamanta anethifolia (D.Don) Wall., Numer. List no. 569. 1829. ≡ Ptychotis anethifolia (D.Don) DC., Prodr. 4: 108. 1830. ≡ Carum anethifolium (D.Don) C.B.Clarke in Benth. & Hook.f., Fl. Brit. India, 2(6): 683. 1879. ≡ Trachyspermum anethifolium (D.Don) H.Wolff, Pflanzenr. IV, 228(90): 90. 1927. —LECTOTYPE (designated here): Nepalia, 1821, N. Wallich 569 (K barcode K000685646; isolectotypes: G barcode G00367346, M barcode M0172780).

Diagnosis: Annual plants, pubescent, up to 40–50 cm tall; leaves small, 1–4 cm long, ternate with linear ultimate segments; ultimate umbels small, with unequal, and filiform rays; bracts and bracteoles numerous, linear, ± as long as the peduncles; petals white; mericarps rounded-elliptic, pubescent.

Distribution: Nepal (endemic).

Psammogeton capillifolium (Regel & Schmalh.) Mousavi, Mozaff. & Zarre, comb. nov. ≡ Pimpinella capillifolia Regel & Schmalh., Izv. Imp. Obshch. Lyubit. Estestv. Moskovsk. Univ. 34(2): 29. 1881. ≡ Aphanopleura capillifolia (Regel & Schmalh.) Lipsky, Izv. Imp. Akad. Nauk. 4: 379. 1896. = Carum capillifolium (Regel & Schmalh.) Koso-Pol., Bull. Soc. Imp. Naturalistes Moscou n.s., 29: 199. 1915 (publ. 1916). —TYPE: Kazakhstan, Karatan of fluv. Tschygan [Tschilak, am Nordabhang der Berge Karatan], 1880, A. Regel s.n. SYNTYPEYNTYPE; Kazakhstan, Turkestan: Suleimanfels bei Osch., 25 May 1880, A. Regel s.n. (P barcode P03214061; isosyntype: MPU barcode MPU019088).

Diagnosis: Annual plants; glabrous, sometimes purplish-red in lower parts of stem; leaves petiolate, 2-pinnate or 2-ternate, ultimate segments filiform or linear; umbels long, usually with more than 2 flowers and unequal rays; bracts absent or one; bracteoles lanceolate or linear; mericarps broadly ovate.

Distribution: Kazakhstan, Kyrgyzstan, Tajikistan, Turkmenistan, Uzbekistan, and China.

Psammogeton microcarpum (Hedge, Lamond & Rech.f.) Mousavi, Mozaff. & Zarre, comb. nov. ≡ Trachyspermum microcarpum Hedge, Lamond & Rech.f., Fl. Iran. 162: 342. 1987. —TYPE: Afghanistan, Kabul, in valle Logar 12 km S Kabul prope “Stupa Guldara”, 1970 ma. s. l., in rupibus verticalibus, 24 Sep 1967, H. Freitag 1991 (holotype M Hb.Freitag; isotypes: E, W barcode W0054255).

Diagnosis: Perennial plants, stem up to 40 cm; leaves trisected or pinnate with cuneate ultimate segments; umbels with 2–5 unequal rays; bracts and bracteoles linear or setaceous; mericarps ovate-elliptic, covered with T-shaped trichomes; stylopodium conical.

Distribution: Afghanistan (endemic).

Psammogeton paktianum (Hedge, Lamond & Rech.f.) Mousavi, Mozaff. & Zarre, comb. nov. ≡ Trachyspermum paktianum Hedge, Lamond & Rech.f., Fl. Iran. 162: 343. 1987. —TYPE: Afghanistan, Jaji, slopes 1 km below Alikhe, 20 Aug 1907, H.Freitag 1829 (holotype M Hb. Freitag; isotypes: E barcode E00000356, MSB barcode MSB003195, MSB barcode MSB003196).

Diagnosis: Perennial plants; leaves long petiolate, subtripinnate with cuneate-trisected or acute ultimate segment; umbels long, thin, pedunculated with 4–6 rays; bracts subulate; bracteoles lanceolate; petals white and short papillous; mericarps ovoid or subpyriform, covered with short trichomes.

Distribution: Afghanistan (endemic).

Psammogeton papillare (Boiss.) Mousavi, Mozaff. & Zarre, comb. nov. ≡ Reutera papillaris Boiss., Diagn. Pl. Orient. Ser. 2, 2: 76. 1856. ≡ Pimpinella papillaris (Boiss.) Benth. & Hook.f. ex Drude, in Engler & Prantl, Natürl. Pflanzenfam. III, 8: 196. 1898. ≡ Carum papillare (Boiss.) Koso-Pol., Bull. Soc. Imp. Naturalistes Moscou n.s., 29: 198. 1915 (publ. 1916). ≡ Trachyspermum papillare (Boiss.) Hedge, Lamond & Rech.f., Fl. Iran. 162: 342. 1987. —LECTOTYPE (designated here): Afghanistan, Jumra and Karabagh, W.Griffith s.n. (K, barcode K000685651; isolectotype: E, barcode E00000357).

Diagnosis: Perennial plants; leaves pinnate with linear, acuminate ultimate segments, cauline leaves linear-filiform, apiculate; umbels with few, unequal, thin rays; bracts and bracteoles lanceolate, acute, unequal, densely hirsute, hyaline at margin; petals white, short hirsute; mericarps ovoid, covered with dense T-shape trichomes; stylopodium depressed or conical.

Distribution: Afghanistan (endemic).

Psammogeton involucratum Mousavi, Mozaff. & Zarre, comb. nov. ≡ Apium involucratum Roxb. in Fleming, Asiat. Res. 11: 157. 1810. ≡ Athamanta roxburghiana Wall., Cat. no. 571. 1829, nom. superfl. ≡ Ptychotis roxburghiana DC., Prodr. [A.P. de Candolle] 4: 109. 1830, nom. superfl. ≡ Pimpinella involucrata (Roxb.) Wight & Arn., Prodr. Fl. Ind. Orient. 1: 369. 1834. ≡ Carum roxburghianum (DC.) Benth. & Hook.f., Gen. Pl. 1: 891. 1867, nom. superfl. ≡ Carum involucratum (Roxb.) Baill., Hist. Pl. 7: 179. 1879. ≡ Carum involucratum (Roxb.) Kuntze, Rev. Gen. 265. 1891, nom. illeg., non Baill. ≡ Trachyspermum roxburghianum Craib, Fl. Siam. Enum. 1: 788. 1931. ≡ Trachyspermum matthewii M.R.Almeida, Fl. Maharashtra 2: 363. 1998, nom. superfl. —LECTOTYPE (selected by I.M.Turner, 2014 in Kew Bulletin 69: 9489): Roxburgh drawing no. 1388 Icones Roxburghianae no. 1388 (K!, available online at http://apps.kew.org/floraindica/home.do).

= Trachyspermum stictocarpum (C.B.Clarke) H.Wolff, Pflanzenr. (Engler) Umbellif.-Apioid.-Ammin. 89. 1927. ≡ Carum stictocarpum C.B.Clarke, Fl. Brit. India 2: 681. 1879. —LECTOTYPE (designated here): India, Maharashtra, Concan, J.E. Stocks and J.S. Law s.n. (K barcode K000685630; isolectotypes: K barcode. K000685631, K000685629). —SYNTYPE: J.S. Law s.n. (K barcode K000685632). 

= Carum stictocarpum var. hebecarpum C.B.Clarke, Fl. Brit. India 2: 682. 1879. —LECTOTYPE (designated here): India, Maharashtra, Concan, J.E. Stocks s.n. (K barcode K000685626; isolectotype: K barcode K000685627). —SYNTYPE: India, Maharashtra, Concan, J.S. Law s.n. (K barcode K000685628).

Diagnosis: Annual plants; stem 20–100 cm; leaves petiolate, blade ovate, 2-pinnate or ternate-pinnate with narrowly oblong ultimate segments; umbels 2–4, pedunculated, with 4–12 filiform rays; bracts and bracteoles few, linear-subulate or ciliate; mericarps ovoid, densely hirsutulous or glabrescent.

Distribution: Java, India, Malaysia, Borneo, Philippines, Vietnam, Laos, New Guinea, Thailand, Andaman Islands, Myanmar, China, and Bangladesh.

Resurrected name

Psammogeton brevisetum Boiss., Fl. Or. 2: 1079. 1872. ≡ Cuminum brevisetum (Boiss.) Kos-Pol., Bull. Soc. Imp. Nat. Mosc. Ser. Nov. 29: 209. 1916. ≡ Aphanopleura breviseta (Boiss.) Heywood & Jury, Ombell. Contrib. Pluridisc. Syst. 2: 733. 1977 (publ. 1978). —LECTOTYPE (designated here): Iran, Inter Kerman et Jesd, May.1859, A. von Bunge (P barcode P00834707).

= Athamantha grisea Stapf & Wettst., Denkschr. Akad. Wiss. Wien. Math. Nat. Kl. 51, 2: 319. 1886. —Lectotype (designated here): Iran, Kuschkek inter Hamadan & Teheran, 19 Jul 1882, T.Pichler s.n. (JE barcode JE00003614).

= Psammogeton flabellatus Bornm. & Gauba, Repert. Spec. Nov. Regni Veg. 36: 341. 1934. —TYPE: Iran, Keredj: Salzberge bei Mardabad (SW Keredj in der Salzsteppe), 1934, D.E. Gauba s.n. (Holotype: B barcode B100366020).

Diagnosis: Annual plants; stem 3–30 cm; leaves ternate or bi-ternate, petiolate, densely covered by short indumentum; umbels pedunculate with 5–10 rays; bracts 5, short, membranous at margin, subulate; bracteoles 5–7, lanceolate; petals white, curved at apex; mericarps oblong-ovoid covered by clavate, or vesiculate trichomes.

Distribution: Central Iran (endemic).