Parapholis

Röser, Julia Schneider Grit Winterfeld Martin, 2012, Polyphyly of the grass tribe Hainardieae (Poaceae: Pooideae): identification of its different lineages based on molecular phylogenetics, including morphological and cytogenetic characteristics, Organisms Diversity & Evolution (New York, N. Y.) 12 (2), pp. 113-132 : 121-123

Parapholis and Hainardia ( lineage I)

Parapholis ( P. incurva and P. filiformis sampled) and monotypic genus Hainardia (only H. cylindrica ) are related closely with each other and with Catapodium (incl. Scleropoa ), Cutandia , Desmazeria , Sphenopus , and Vulpiella . Cynosurus is a sister of this group within lineage I (Figs. 1 and 2). Furthermore, all these lineage I taxa are characterised by a 6 bp deletion in the mat K gene region (Fig. 1). Previous molecular phylogenetic studies generally found a similar structure of this group, but had included fewer genera ( Soreng and Davis 2000; Catalán et al. 2004; 2007; Davis and Soreng 2007; Döring et al. 2007; Quintanar et al. 2007; Bouchenak-Khelladi et al. 2008; Döring 2009; Schneider et al. 2009). Most current classifications of subtribes treat Cynosurus under monotypic subtribe Cynosurinae , whereas the other genera are unified under subtribe Parapholiinae ( Table 1; cf. Tzvelev 1976; Soreng and Davis 2000; Soreng et al. 2003; 2007; Catalán et al. 2004; 2007; Quintanar et al. 2007). Agropyropsis and Pholiurus , affiliated with Parapholiinae in Soreng et al. (2007), are phylogenetically distant to this assemblage. These genera belong to the lineages of Festuca and relatives (subtribe Loliinae ) and Poa and relatives (subtribe Poinae ), respectively (see below; cf. Table 1). Vulpiella , conversely, is distant to subtribe Loliinae , in which it was placed by Soreng et al. (2007). It belongs to the Parapholis lineage as stated implicitly by Clayton and Renvoize (1986), who pointed to the similarity of Vulpiella and Cutandia .

Hainardia and Parapholis are formed only in the mat K tree a clearly supported clade (Fig. 1), whereas the ITS data are ambiguous and even show Parapholis as paraphyletic ( Fig. 2 ). Both genera share morphologically well-defined characters, such as the spicate inflorescences with the rachis disarticulating transversely at fruit maturity into segments, each of which bears an entire spikelet ( Table 3; Fig. 3a, b). Additionally, the palea has glabrous and only weakly developed keels ( Table 3; Fig. 4a, b ). This contrasts to the other genera of lineage I (subtribe Parapholiinae ) with more or less paniculate inflorescences, spikelets with two or more florets, and disarticulating below each floret, and with paleas having hairy keels ( Catapodium incl. Scleropoa , Cutandia , Desmazeria , Sphenopus , Vulpiella ; personal observations; Clayton and Renvoize 1986; Watson and Dallwitz 1992; 1992 onwards). The spikelet-bearing axes are disarticulating in Vulpiella and

0.1

are shown above branches and numbers below are bootstrap supports ≥50. Genera of the traditionally defined Hainardieae indicated in bold

Hainardia Parapholis Narduroides Agropyropsis Scribneria Pholiurus

Life form Annual Annual Annual Perennial Annual Annual

Inflorescence Spicate; spikelets without Spicate; spikelets Racemose; Spicate; Spicate to racemose; Spicate; spikelets shape pedicel a without pedicel b spikelets briefly spikelets sometimes several without pedicel f pedicelled c without spikelets per rachis pedicel d node (reduced panicle) e

Arrangement Distichous (bilateral) a Distichous Distichous Distichous Distichous (bilateral) e Distichous of spikelets (bilateral) b (bilateral), but (bilateral) d (bilateral) f frequently oriented ± secund (unilateral) c

Spike or Fragile a Fragile b Tough c Tough d, g Tough e Tough f

raceme rachis

Spikelet 1-Flowered 1-Flowered 2- to 4-Flowered c, h 3- to 6- 1-Flowered e 2-Flowered f Flowered d

Disarticulation Absent; spikelets falling Absent; spikelets Beneath the Beneath the Beneath the floret k Below the glumes;

of spikelet entire together with the falling entire florets i florets j spikelets falling rachis fragments a together with entire with the the rachis glumes staying fragments b together f

Number of 1 (Lower glume 2 b 2 c 2 d 2 e 2 f

glumes suppressed); 2 glumes present only in the terminal spikelet of the inflorescence a

Lemma Awnless a Awnless b Awnless c, i Awnless d, j Awned; awn straight, Awnless f arising from the sinus of the bidentate lemma tip e, k

Callus of the Absent Absent Present, glabrous i Present, Present, hairy k Absent lemma glabrous j

Keels of the Glabrous, smooth l Glabrous, smooth m Ciliolate Ciliolate n Ciliolate Ciliolate f

palea

Ovary Yes o Yes p Yes q Yes r No s Yes t

(karyopsis)

with conspicuous apical appendage

Ovary No o No p Yes q Yes r No s No t

(karyopsis)

tip with two decurrent wings of the appendage

Karyopsis No No Yes q No No No with deep longitudinal furrow and adherent to the palea

Hilum shape Shortly linear Shortly linear p Shortly linear Shortly linear r Punctiform Oval t

Chromosome x 013 x 07, 9 and 19 x 07? x 013 x 07

base number

Habitat Saline or alkaline soils, Sandy soils, salt Dry places Damp saline Damp sandy soils in Damp saline or coastal sands marshes soils mountains alkaline soils some species of Catapodium in addition to the spikelets disarticulating between the florets. This might represent a transitional character to the disarticulation of the inflorescence axis in Hainardia and Parapholis , as argued by Soreng and Davis (2000), Soreng et al. (2007). Hainardia and Parapholis belong to the few genera within the Aveneae / Poeae tribe complex that deviate from the typical monoploid chromosome number of x 0 7 ( Table 4, Figs. 1 and 6c).

Figures

Fig. 2 Bayesian phylogenetic tree obtained from nuclear ITS DNA sequences. Branches collapsed in the strict consensus tree based on maximum parsimony analysis are in bold. Posterior probabilities ≥0.95

Fig. 4 a–h Spikelet details. Palea of a Hainardia cylindrica and b Parapholis incurva with glabrous keels (nerves). c, d florets of Narduroides salzmannii with glabrous callus of the lemmas and long rachilla internodes. e, f Agropyropsis lolium with glabrous callus of the lemma (e) and ciliolate keels of the palea (f). g, h Scribneria bolanderi with lemma awned from a sinus of the lemma tip (g) and hairy callus (h). The margin of the lemma is marked at one side in a, b, and f with arrows. Material used: a fruits from Göttingen Botanical Garden in 2006, no. 1850 (HAL); b Röser 2517 (HAL); c Hernández s.n. (C); d Rivas-Martínez s.n. (BASBG); e, f Cosson s.n. (JE); g, h Wilken 16163 and Painter (RSA 695253)