Bo!anicalJournal of the Linnean Society (1996), 120: 145-198. With 13 figures

A phylogenetic study of the Plantaginaceae Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021

KNUD RAHN

Botanic Garden, Uniuersib of Copenhagen, Sdvgade 83, Opg. S, DK-1307, Copenhagen K; Denmark

Received Augur/ 1993, acceptedfor publication Noumber 3995

In a study based on morphological, embryological and chemical data of thc Plantaginaceae, within the subclass Sympetalae or Asteridae, the superorder 1,amianae is shown to be monophyletic. However, it was not possible to reconstruct the phylogeny within Lamianae or to find a sister-,group for the monophyletic Plantaginaceae - Hydrostachyaceae for the latter is rejected. Thrcr or rarely four genera have previously been recognized within Plantaginaceae, but in both cases Plantago appears as polyphyletic, which is considered unacceptable. Six clades are recognized as subgenera within Plantago: subgen. Plantago (c. 131 species), subgen. Cor0nopu.r (c. 1 1 species), subgen. Littorella (three species), subgen. Pvllium (c. 16 species), subgen. Bougum'a (one species), and subgen. Albicans (51 species). Within P. subgen. Plantago, the paraphyletic sect. Plantago (c. 42 species) is found in all parts of the world except South America, New Zealand, Australia, and New Guinea, where sect. Oliganthos and sect. Mesembyiae vicariate. It is not possible to infer the phylogeny within sect. Plantago, for example between the endemic species from distant Pacific Islands; their commoii ancestor might be a species that once had a very wide distribution. Keys to genera, subgenera, sections and series are given. Only one genus, Plantago L., is recognized. The following proposals are made in the Appendix: P. ser. Oliganthos Rahn, scr. nov.; P. ser. Carpophorae (Rahn) Rahn, stat. nov. ( = sect. C.); P. ser. Microcabx (Pilg.) Rahn, stat. nov. ( = sect. M.);P. unibracteata Rahn, nom. nov. ( = P. unzjlora Ho0k.f. non L.); P. subgen. Littorella (PJ.Bergius) Rahn, stat. nov. ( = Littorella PJ.Bergius); P. araucana Rahn, nom. nov. (= Littorella australis Griseb. non Plantago australis Lam.); P. amm'cana (Fernald) Rahn, comb. nov.( = Littorella a,); P. subgen. Bougueria (Decne.) Rahn, stat. nov. ( = Bougumia Decne.); P. nubicola (Decne.) Rahn, comb. nov. (= Bougueria n. Decne.); P. subgen. A1biran.r Rahn, subgen. nov. Ten lectotypes are selected.

01996 'Thr Lnnean Society of London

ADDITIONAL KEY WORDS: -Asteridat- cladistics - Lamianae ~ phylogeny ~ Phntugo ~ taxonomy.

CONTENTS

Introduction ...... I46 Material and methods ...... 146 List of characters for the Plantaginaceae ...... 148 Relationship of the Plantaginaceae ...... 156 Asteridae ...... 156 Lamianae ...... 157 Plantaginaceae ...... 160 Phylogenetic analysis of Plantaginaceae ...... 161 The common ancestor of Plantaginaceae ...... 161 The genus Plantugo ...... 161 Synapomorphies uniting the subgenera ...... I62 Plantago, key to the subgenera ...... 163 Plantago subgen. Plantago ...... 166 Plantago subgen. Coronopus ...... 179

145 0024-4074/96/020145 t 54 $18.00/0 01996 The Linnean Society of London I46 KNUD RAHN

Planlago subgen. Littorella ...... 181‘ Plantago subgen. Pylliurn ...... 182 Plantago subgen. Boupuma ...... 184 Plantago subgen. Albicans ...... I a5 Concluding remarks ...... I90 Acknowledgements ...... 191 References ...... 191 Appendix 1 Nomenclatorial changes within Plantaginaceae ...... 196 Appendix 2: Index to specific epithets ...... 198 Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021

INTRODUCTION

The purpose of the present paper is to deduce the phylogeny within Plantaginaceae, and to evaluate the resulting groups in relation to their distribution. For instance, Croizat (1952, 1958) used Plantago sect. Palaeopsyllium as one of many examples of some of his ‘tracks’ but is this section (in the sense of Pilger, 1937) a monophyletic or a plesiomorphic assembly? The grouping of the species within Plantaginaceae has never been adequate. Pilger (1937) apparently gave up and used the same genera as Decaisne (1852) and only slightly modified the sections as used by Decaisne (1.c.) and by Harms & Reiche (1895). The sequence of the sections has particularly caused much confusion. Sections, which in the present paper are placed in subgen. Coronopus and subgen. Albicans, were spread between sections belonging to subgen. Plantago. Later, a modest approach was adopted (Rahn, 1978) by recognizing three subgenera within Plantago: subgen.Plantago P. subgen. Coronopus and uniting the present subgen. Psyllium and subgen. Albicans in P. subgen. Psyllium. The last proposal has been repeatedly criticized by Andrzejewska-Golec & Swiatek (1984), Andrzejewska-Golec & Swietoslawski (1 987a, 1988, 1989 a-c, 1992a, b) and Andrzejewska-Golec (1992) based on phenetic arguments. Plantaginaceae is without doubt a monophyletic group, but it is not at all evident which characters belonged to its common ancestor. Revision at the species level is not intended. Accepted names, including the nomenclatorial changes proposed in Appendex 1, are used throughout this paper, and when citing papers in which synonyms are used. A ‘Ch’ followed by an italicized number refers to a binary character listed below and used in tables and cladograms, and a ‘Sp’ followed by a number refers to the numbering of the species used in cladograms and tables. At the end an alphabetic list of specific epithets refers to these numbers. In the quest for the ancestor or sister group of Plantaginaceae a preliminary phylogenetic study of the families in Asteridae was initiated and is referred to briefly.

MATERIAL AND METHODS

Species from the following papers have been included in this study of Plantaginaceae: Allan (196 1) for New Zealand species, Basset (1 966) for subgen. Micropsyllium in America; Briggs, Carolin & Pulley (1977), Briggs (1980) and Curtis (1 967) for subgen. Mesembryniae and Oliganthos in Australia; Craven (1976) and van Royen (1964, 1983) for New Guinean species; and Rahn (1974, 1979a, 1981, 1982, PHYLOGEKY OF PI,WT:\GINACEI\I; 147

1983, 1984, 1985) for South American species. Names of other species are used in the same sense as Pilger (1937) unless otherwise indicated. Data for 130 OTUs were scored for the study of an expanded Asteridae, including the families in Asteridae sensu Cronquist (1981), 10 from Cornanae, 10 from Aralianae 12 from Ericanae (sensu Dahlgren, 1989) and 7 other families within which either iridoids or verbascosides have been recorded. Some of the 98 families were subdivided. Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 The study of Lamianae included 20 families, i.e. the superorder in the sense of DahlLgren(1 989) and Oleaceae. The characters used in this study were mainly taken from the literature cited above and in the following, but supplemented from other sources including studies of living plants in the Botanic Garden, Copenhagen, or of dried specimens from C, K, GB and Z. Those for the Plantaginaceae are listed below, but 55 used for the study of Asteridae and 63 for Lamianae are not. General characters for families were extracted from Engler & Prantl (1 894- 1899), Melchior (1964), Hutchinson (1973), Dahlgren (1979- 198l), Cronquist (1 98 I), and Goldberg (1986). Chemical characters were taken from: Karrer (1958-1 985), Willaman & Li (1970),Jensen, Nielsen & Dahlgren (1975),Jensen (1991), Pollard & Amuti (198 l), Gershenzon & Mabry (1983), Kuzmanov et al. (1 984), Andrzejewska- Golec & Swiatek (1984), Andrzejewska-Golec et al. (1 993) and Hegnauer (1990). Information about pollen morphology was extracted from Selling (1947), Aubert et al. (1962), Bassett & Crompton (1968), Dietrich (1968, 1970, 1971), Rahn (1974), Clarke & Jones (1 977), Hooghiemstra (1983), Erdtmann (1986), Saad (1986) and Ubera, Galan & Guerrero (1988). This information was supplemented by studying the pollen of Plantago amplexicaulis, P. albicans, P. atrata, P. ciliata, P. hispidula, P. lachnantha, P. minuta, P. ouata, and P. stocksii in a scanning microscope (Fig. 2) The study of hair types in Plantaginaceae was published in a separate paper (Rahn, 1992), and more detailed studies were made by Andrzejewska-Golec & Swietoslawski (1987a, b, 1988, 198%-c, 1993). Corolla aestivation was taken from Armstrong & Douglas (1989) and other anatomical and morphological characters from Carlquist (1992), Metcalf & Chalk (1950), Weberling (198 I), Troll & Weberling (1 989), Troll (1969), and Heslop-Harrison & Shevanna (1977). Embryological characters were extracted from Balicka-Iwanowska (1 899), Schnarf (1 9 17, 193I), Sou?ges (1923); Cooper (1942), Johansen (1 950), Misra (1964), Davis (1966), Rauh & Jager-Ziirn (1966), Kapil & Tiwari (1978), Kapil & Bhatnagar (199 1 ), Cocucci (1983), Johri, Ambegaokar & Srivastava (1992). Chromosome numbers are from Moore (1973), Dietrich ( 1975, 1980), Goldblatt ( 1981 - 199 1) and Brullo, Pavone & Terasi (1985). The term ‘character’ is used in this paper in a similar way to that explained by Watrous & Wheeler (1981), and refers to one transformation or a group of indistinguishable or not separated transformations. Character states in a transforma- tion series proximal to this particular transformation are considered plesiomorphic, and distal states apomorphic. This means that all characters are binary and some are nested. In the tables ‘3’ indicates that all studied members of an OTU exhibit the apomorphic state of the character in question, ‘0’ that the plesiomorphy is present in all, ‘1’ or ‘2’ that only some members have the apomorphy, ‘1’ indicates that it is supposed to have been absent in the ancestral population; a ‘2’,that it was present. ‘ + ’ and ‘-’ in the tables indicate that a character is unknown and expected to be 148 KNUD RAHN present or absent. Figures 4-13 are dendrograms. A black bar indicates a synapomophy without reversals, an open bar one followed by reversals, a cross a reversal, and two parallel lines a parallelism. To the left of the Sp-number one of the parsimonious trees is drawn. To the right are name, distribution, chromosome number, and the strict consensus tree, if more than one minimal tree were found. Unknown characters are entered as a ‘?’ in the Hennig86 data file. J.S. Farris parsimony program Hennig86, version 1.5 was used for all data files Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 (Farris, 1988). The command ‘ie’ was used when possible (giving all the shortest trees); otherwise, Ye-’ (which produces a single tree of minimal length) combined with ‘mhennie’ or ‘mhennie’ alone was used. This was followed by ‘nelsen’, producing a strict consensus tree.

LIST OF CHARACTERS FOR THE PLANTAGINACME

Only apomorphic (derived or advanced) characters (or character states) are listed, and the corresponding plesiomorphy is not mentioned when obvious. The character numbers and all or part of the first line of description are italicized. However, many of the characters listed below are not single phylogenetic characters, but a collection of similar phylogenetic characters (parallel evolution). 1. Adventitious roots. When cultivating Plantago with only one fleshy root, the most common cause for the death of a plant is that the upper part of the root or lower part of the stem is attacked by animals or fungi. However, plants often survive such attacks when adventitious roots are formed from the caudex. Thus adventitious roots are probably very advantageous when the water supply is not a problem. Adventitious roots have evolved several times independently as an adaptation to moist conditions in Plantago sect. Plantago, sect. Oliganthos and sect. Mesembrynia. They are also present in the three species of subgen. Littorella, but elsewhere known only in P. aliissima and P. argenteu. 2. Annual root is supposed to be apomorphic in relation to both adventitious roots and perennial tap-root. It would appear to have developed several times as an adaptation to drought. It is found in all larger groups except Plantago sect. Plantago, P. sect. Oliganthos, subgen. Littorella, subgen. Bougueria, and P. sect. Lance$lia. A doubling of the chromosome number in annual species may cause the plants to be perennial (Sp 183 and 184). 3. Stem elongated. In the most parsimonious tree, the common ancestor of Plantaginaceae has the leaves in a rosette, and it is for that reason considered plesiomorphic, although an elongated stem is by far the most common (and plesiomorphic) in the Lamianae. 4. Arrangement ofleaves, opposite or in whorls ofthree, is considered apomorphic within Plantaginaceae (producing the most parsimonious solution). Here it is found only in subgen. Pvllium, but it is the most common arrangement (and probably plesiomorphic) in other families in Lamianae and possible sister-groups. Cronquist (1981: 853) indicates simple, stipulate, and opposite leaves as primitive in Asteridae. 5. Lamina less than 10 times as long as wide. Pinnately nerved and lobed leaves like those of P. coronopus are closest to leaves in other Lamianae. By reduction of the lamina and the lobes, a linear leaf develops (the rachis) as in P. maritimum and subgen. Pyllium; by widening the lamina a broader leaf develops, as in subgen. Plantago. PHYLOGENY OF PLANTAGINACEAE 149

However, within the subgenus Plantago, leaves less than 10 times longer than wide are considered plesiomorphic. 6. Lamina less than 1.9 times as long as wide. Within P. subgen. Plantago, more advanced than Ch. 5. 7. Lamina more than 4 times as wide as the petiole (broadest and narrowest part of the leaf). A smaller difference is considered more primitive in subgen. Plantago. 8. Lamina more than 20 times wide the petiole. More advanced than 7.

as as Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 9. Nerves of dead leaf remaining on the plant as long brides. In subgen. Plantago the nerves in most species decay like the rest of the leaf, but bristles occur and have probably originated several times in subgen. Plantago. 10. Base of lamina truncate. In subgen. Plantago this is considered apomorphic in relation to an attenuate base or to a lamina not distinguishable from the petiole. 11. Apex ofthe leaves with a colourless acumen. Advanced in Plantago sect. Oliganthos. 12. Leaves remaining green on dving. Leaves turning dark is tentatively considered plesiomorphic in Plantaginaceae. 13. Scape very short, less than a quarter ofthe supporting leaJ: 14. Scape much elongated afkr anthesis. 15. Hairs on leaves long, > 2 mm. 16. Hairs on leaves narrow, 0.04 mm. 17. Hairs on scape antrorse. 18. Small, three-celled, glandular hairs placed in cavities (Fig. 1. See also Rahn (1992) type B and fig. 3a-d and 9). A synapomorphy for P. ser. Microcalyx. In all other parts of this family and other Lamianae, the three-celled hairs are placed on the surface of the green parts of the plant. 19. Small glandular hairs with a stalk of one short cell and an obtuse head of more than two cells (Fig. 1 See also Rahn (1992) type C and fig. 2e). The three-celled hairs, found in all larger groups within Plantaginaceae and in most other families in Lamianae, are considered primitive (see Rahn (1992) type A and figs lb, 2a-c, 3e-f. 3h-k. 6a, 7c, 8a-c). Both are sometimes present and found within subgen. Psyllium and subgen. Albicans. 20. Like 19, but head acute, cone-shaped (Fig. 1. See also Rahn (1992) type D and figs 2f&g, 6be). Such hairs not found together with three-celled hairs, but in all the studied species belonging to P. subgen. Coronopus and sect. Lanceifilia and not found elsewhere. 21. Presence ofglandular hairs with a stalk or more than one cell, and head one-celled, globular or cylindrical and obtuse, never acute (Fig. 1. See also Rahn (1992) type E and F and figs la, 2h-j, 7d-f). When such glandular hairs are present, they are usually found together with three-celled glandular hairs and are also common in other families in Lamianae, but rare in Plantaginaceae. 22. Long glandular hairs (included in Ch 21). These are present within subgen. Psyllium and subgen. Albicans (Fig. 1. See also Rahn (1992) type F and figs la, 2j). 23. Spur-lih elongation on lowermost cell ofnon-glandular hairs on the scape (Fig. 1. See also Rahn (1992) type I. Present within subgen. Plantago. 24. Lowermost cell ofnon-glandular hairs shorter than wide and much shorter than the other cells (Fig. 1 and Rahn (1992) type J, K, L, M, N and fig. 5). Synapomorphy for subgen. Bouguen'a and subgen. Albicans. Hairs with the lowermost cell not differentiated are considered primitive, and are commonly found in most families in Lamianae. 25. Non-glandular hairs with walls and especial4 joints strong4 refracting, and the narrow lumen in the cells is usual4 d@cult to see (Fig. 1 and Rahn (1992) type J, K, L, M and fig. 150 KNUD RAHN

Ch 19 Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021

Ch 20

-Ch 27 Ch 21 H C<25 Ch 24

Figure 1. Plantaginaceae, hair-characters. The three hairs to the left represetit plesiomorphic states' a small three-celled glandular hair seen from the side and from above, and a long non-glandular hair. Glandular hairs (Ch 18-Ch 22) and details of Ch 27 and Ch 2B: scale bar = 28 pm; Ch 23-Ch 26 scale bar = lO0pm

Fi

5g-k, m-q, s-v). Synapomorphy for subgen. Albicans. The primitive type of hairs may also be present somewhere on the plants, e.g. in sect. Lancefolia and sect. Montana where the refracting hairs are present on the leaves and translucent hairs with walls very oblique present on the scapes (See Rahn (1992) type N and fig. 5a-e, 1). 26. As 25 but walls between the cells oblique (Fig. 1, and Rahn (1 992) type K, fig. 5h-i). Synapomorphy for sect. Gnaphaloides (Sp 19 1-2 13). Hairs with end walls perpendicu- 
 lar (Rahn, 1992 type J and fig. 5g, j, s) is considered plesiomorphic within subgen. Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 A lbicans. 27. Like 25, andjoint with fork- or crown-like elongations. These are found in subgen. Albicans (Sp 163-1 74 and 176-190). The cells have their distal end inserted between two flanges from the cell above (Fig. 1. See also Rahn (1992) type M and Fig. 5f, m-s) or crown-like ( = 28). 28. Cells of non-glandular hairsjointed ly a common wall with crown-like elongations (Fig. 1. See also Rahn (1992) type L and Fig. 5k, u-v). By including 28 in 27 a more parsimonious solution is achieved than considering them independently developed. Found in sect. Hymenopsyllium (Sp 176-178) and ser. Minuta (Sp 180-181). 29. Flowers opposite or in whorls of three. Flowers spirally arranged in spikes is considered plesiomorphic in Plantaginaceae. In subgen. Psyllium the leaves are opposite or in whorls of three (4) and the flowers are arranged in the same way. Present in subgen. Littorella, subgen. Psyllium and sect. Gnaphaloides. (See under 4 and discussion under Plantaginaceae). 30. Flowers dense& crowded, the rachis not visible between theflowers, spike usual& ovate or globose. A more open and cylindrical spike is considered plesiomorphic. Probably developed independently in subgen. Plantago and in subgen. Albicans + subgen. Bougueria. 31. LASS than 12flowers in a normal spike. A synapomorphy for the species in Plantago sect. Oliganthos. Only occasionally found in depauperate specimens of other species in subgen. Plantago. Also a synapomorphy for subgen. Pyllium + subgen. Littorella. 32. Flower solitary, one or two bracts present. More specialized than 31. Within Plantago sect. Oliganthos. 33. Flower solitaly, on& one bract present. More specialized than 32. 34. Flowers unisexual, monoecious. Synapomorphy for P. subgen. Littorella. 35. Female flowers in one whorl. In P. subgen. Littorella. More than one whorl is considered plesiomorphic. 36. Bract broad, covering sepals lateral&, usual& wider than long. Within P. subgen. Albicans. 37. Bract with the upper part scarious, acuminate. A synapomorphy uniting the species in P. subgen. Lancegolia. 38. Pedicel present, or base of ca&x narrowed like a pedicel. Within Plantago sect. Plantago. 39. Pedicel in malejowers more than 4mm long. Within P. subgen. Littorella. 40. Sepals glabrous on the back. With some doubt considered apomorphic within Plantaginaceae, but hairs on the nerve have been lost in several clades. 41. Anterior sepals distinct& narrower than theposterior, and dgerently shaped. Sepals almost equal are considered primitive. In the Lamianae a regular calyx is more common than a bilabiate one. 42. Each anterior sepal is very asymmetric with hairs placed vqasymmetrical&. Within subgen. Albicans (Sp 183-190). I52 KNUD RAHN 43. Nerve of anterior sepals present at base onb, distal part scarious. A synapomorphy for P. sect. Montana (Sp 163-168). Evolved independently in Plantago macrocarpa (Sp 2). 44. Anterior sepals unitedfor more than half their length, their scarious parts united. Scarious parts free is considered primitive. A synapomorphy for P. sect. Lancefolia (Sp 169-1 74). 45. Nerve ofposterior sepals present at base on&, the dhtal part scarious. A synapomorphy for species in P. sect. Montana (Sp 163-168) and evolved independently in P. amplexicaulis (Sp 175), P. macrocarpa (Sp 2), and P. schwarzenbergiana (Sp 53). Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 46. Posterior sepals with a sharp he1 or wing on the back. A synapomorphy for sect. Lancegolia (Sp 169- 174), and apparently evolved independently within subgen. Coronopus (Sp 132-142). 47. Posterior sepals with a membraneous and very conspicuous wing on the back. More specialized and included in 46. 48. Corolla tube with short hairs. A synapomorphy uniting the species in subgen. Coronapus (Sp 132-142). The lanate corolla tube of P. lachnantha (Sp 181) is not considered homologous with this character. 49. Corolla lobes slightb hairy on the back. Found within subgen. Albicans (Sp 180-1 90). 50. Corolla lobes dense& hairy on the back. More specialized and included in 49. A synapomorphy for P. ser. Ciliatae. 51. Corolla lobes with margzns ciliate. Within P. subgen. Pyllium. 52. Corolla lobes wdh margzns dentate. Within P. subgen. Psyllium. 53. Corolla lobes cordate or rounded at base. Cuneate lobes are considered plesiomorphic. In P. ser. Gnaphaloides and ser. Brm’lienses (Sp 204-2 13). 54. Posterior corolla lobe bentfrom a point 0.4-1.1 mm higher than the bending ofthe other lobes and smaller. A synapomorphy for P. ser. Brasilienses (Sp 204-206). Probably evolved from a regular corolla. 55. Posterior corolla lobe erect, smaller, and bmtfiom a point 0.2-0.4mm above the bending of the other lobes. Found within P. ser. Gnaphaloides and considered derived from a re
Figure 3. Planlngo, placenta and seeds from ripe fruits (apex of ovary upwards). On the right sections through the upper third of the seed with placenta side downwards (except Sp 144). Sp 20 P. .rpar$ora, a,b placenta, c,d seed. Sp 26: P. major, a,h: placenta, c,d seed. Sp 43: P. tenuipora, a,b: placenta, c,d seed. Sp 68: P. raoulii a: anterior side of placenta, b: posterior side of placenta, c,d seed. from anterior side, e,f: upper seed from posterior side of placenta, g,h: lower seed from posterior side. Sp 93: P. zirginua, a,b placenta, c,d: seed. Sp 108: P. australis, a: anterior side of placenta, b: postenor side of placenta, c,d seed from the anterior side, e,f: seed from the posterior side of placenta. Sp 110: P. barbata, a: placenta, b,c: seed. Sp 135: P. manhma, a: anterior side of placenta with three aborted ovules, b: posterior side of placenta with an aborted ovule above and the mark from a mature seed, c,d: seed. Sp 140: P. coronopus a: anterior side of placenta, b placenta, lateral view, c: posterior side of placenta, d,e: seed from the upper compartment, f,g: seed from a lower compartment. Sp 144 P. americana, a: adaxial side of female flower with mature fruit, b fruit, frontal view, c: fruit, lateral view, d adaxial side of fruit, e: section of fruit, frontal view, f: seed, frontal view, g: seed, lateral view, h: section of seed. Sp 15 1: P. mauritanua, a,b: placenta, c,d seed. Sp 162: P. nubicol, a: fruit frontal view, b: placenta and fragment of posterior wall of fruit, frontal view, c: placenta and fragment of posterior wall of fruit, lateral view, d: seed, frontal view, e: section of seed. Sp 175: P. amplexicaulk, a: placenta, b.c: seed. Sp 179: P. ovata, a: placenta, b,c: seed. Sp 201: P. bkarckii, a: placenta, b,c: seed. PHYLOGENY OF PLANTAGINACUE 153 Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021
Sp 20, sect. Plantago Sp 26, sect. Plantago Sp 43, sect. Micropsyllium 
 a b cgh ab C ab ce Sp 68, sect. Mesembrynia Sp 93, sect. Virginica Sp 108, sect. Virginica 
 a b C a b cd a bc fg Sp 110, sect. Olzganthos Sp 135, sect. Maritima Sp 140, sect. Coronopus 
H 1 mm 
 d e 
 a bcd a b C a b C Sp 144, subgen. Littorella Sp 151, subgen. Psyllium Sp 162, subgen. Bougueria 
 a b C a b a b C Sp 175, sect. Bauphula Sp 179, sect. Albicans Sp 201, sect. Gnaphaloides 154 KNUD RAHN 58. Corolla lobes erect,JEowers ckistogamous or semicleistogamous. Evolved independently in Plantago sect. Virginica (Sp 81-108), sect. Micropsyllium (Sp 43-48), and P. ser. Hispidulae (Sp 191-195). 59. Corolla lobes longer than 4mm. The most parsimonious cladogram has corolla lobes between 2 and 3mm as the most primitive. 60. Corolla lobes longer than 3 mm. 61. Corolla lobes shorter than 2 mm. 62. Corolla lobes shorter than 1.5mm. Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 63. Corolla lobes shorter than 1 mm. 64. Less than four anthers. A reduction of the number of anthers is evolved independently in Plantago subgen. Bougueria, within P. sect. Micropsyllium and P. sect. Olkanthos. 65. Connective of anther vqlarge, about as long as the pollen sacs. A synapomorphy for P. sect. Hymenopyllium (Sp 176-1 78). 66. Anthers white both whenjesh and when dried. Anthers yellow when young and brownish when dried are considered primitive in Plantaginaceae. 67. Anther. longer than 2 mm long. The most parsimonious cladogram of Plantagina- ceae has anthers between 1.5 and 2 mm as the most primitive. 68. Anthers less than 1.5mm long. 69. Anthers less than 1 mm long. 70. Anthers less than 0.5mm long. 71. Pollen with a solid operculum (Fig. 2). The pore membrane with isolated granules is found in most groups of Plantaginaceae and considered plesiomorphic. Found within Albicans. 72. Carpophorepresent. A solid elongation from the base of the fruit is developed after anthesis. Synapomorphy for Plantago ser. Carpophorae (Sp 117-1 18). A similar, but hollow elongation of the lower part of the capsule is found in some species of P. ser. Microcabx. 73. Ovary hairy. Found within sect. Coronopus. 74. Ovary oftwo carpels, but on@ onefertile. Developed independently several times in Lamianae. P. subgen. Littorella has a unilocular ovary, with a basal placenta and one anatropous ovule (Fig. 3, Sp 144; but according to Eckardt (1937) and Rosen (1940) two lateral placentas present when young, later fuse and swell below (see also under 76. A bilocular ovary with placentas on the septum as in subgen. Plantago is probably plesiomorphic (Fig. 3, Sp 26). The ovary of P. nubicola ( = l3ougum.a n.) is unilocular (Fig. 3, Sp 162), but also formed of two carpels (Eckardt, 1937)with a basal placenta and a single campylotropous ovule; it is probably derived from a bilocular ovary with placenta on the wall between the compartments. 75. Ovary with a third compartment at the top ofthe abaxial side afthe placenta (Fig. 3, Sp 140), or with a rudiment of it, seen as thickening at the apex on the ripe placenta. A synapomorphy for sect. Coronopus. The seed in the upper compartment is dispersed together with the upper part of the pyxis. 76. Ovary with a third compartment at the top on the admial side of the placenta, or with a rudiment of it, seen as a thickening at the apex on the posterior side of the ripe placenta (Fig. 3, Sp 68, 93, and 108). A synapomorphy uniting Plantago sect. Mesmbvnia and P. sect. Virginica. In sect. Mesmbgmia the seed in the upper compartment, if present, is dispersed with the upper part of the pyxis. The other seeds are fixed on either side of the placenta close to the centre. In the young ovary of Plantaginaceae the placenta or wall between the compartments is at first formed PHYLOGENY OF PIANI'AGINACEAE 155 below and later on from the margins of the carpels (Payer, 1857); often a complete separation of the two loculi is never achieved. A fissure distally in the ripe placenta is commonly found in sect. Plantago and sect. Oliganthos, but rarely in Plantago sect. Mesmbryniu when a rudiment of an upper, third compartment is present, nor in other subgenera. 77. Ovary with 1-4 ovules and a rudiment of an upper compartment on the adaxial side of the placenta (76). Evolved from an ovary with five ovules. Found within Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 Plantago sect. Mesembrynia, and sect. Virgnica. 78. Ovary with 1-3 ovules and a rudiment of an upper compartment on the adaxial side of the placenta. Included in 76 and a synapomorphy for Plantago sect. Vi@nica (Sp 81-108). 79. Ovary with 1-2 ovules and a rudiment of an upper compartment on the adaxial side of the placenta (Fig. 3, Sp 93). Included in 76, 77 and 78. 80. Ovary withfour orfewer ovules. Neither an upper compartment nor a rudiment of it. A variable number of more than four is considered primitive in Plantaginaceae. Four ovules are characteristic for Lamiaceae and Verbenaceae, but a variable number of ovules randomly scattered on placentas is common in most other families in Lamianae. 81. Ovary with less than four ovules, and no rudiment of an upper compartment (see under 76). 82. Ovary with two or one ovules, and no rudiment Of an upper compartment (see under 76). 83. Ovary with a single ovule (included in 82), and no rudiment ofan upper compartment (see under 76). 84. Ovules hemitropous, are supposed to have euolvedjom anatropous. The ovule in P. subgen. Littorella (Sp 143-145) unlike all other Plantaginaceae is anatropous, but is more probably evolved from a hemitropous ovule when the strongly transformed ovary is taken into consideration (see under 74). 85. Seeds shorter than 2 mm. The most parsimonious cladogram has seeds between 2 and 3mm as the most primitive in Plantaginaceae. 86. Seeds longer than 3 mm. 87. Inner side Of seed deep@ concave. 88. The concave inner side ofthe seed covered ly a ragged, white membrane, exceptfor two areas to the right and l$ Ofthe centre (Fig. 3, Sp 179 b,c.). Found within sect. Albicans (Sp 179-190). Inner side with one impressed mark just above and one just below the centre is considered primitive. 89. Copledons perpendicular to the placenta. This means that the plane between the cotyledons passes the raphe or the symmetry plane of the seed and is perpendicular to the bract of the flower. This orientation is considered apomorphic. Cotyledons orientated perpendicular to that as in subgen. Plantago is the most common inside and outside Lamianae and tentatively considered plesiomorphic. 90. Basic chromosome number x = 5 or 4. The common ancestor of Plantaginaceae probably had x = 6, as found in all species in subgen. Plantago but one (P. b&elouiz), and in all subgen. Littorella, subgen. Psyllium, and subgen. Bougueria, and parts of subgen. Albicans and subgen. Coronopus. This number was probably reduced independently once in subgen. Coronopus, once in subgen. Albicans (here the number is reduced further to x = 4 in P. ovata), and once in Plantago sect. Micropyllium. The basic number in other Lamianae is usually much higher (e.g. in Buddlga x = 19). The lowest number is x = 3 found in Callitriche hermaphroditica, otherwise the lowest 156 KNUD RAHN number is x = 5 in Verbenaceae. x 6 is found in Verbenaceae, Lamiaceae, Lentibulariaceae and Scrophulariaceae. 91. Free ribose lacking in seeds (Gorenflot & Bourdu, 1962; Hegnauer, 1969), is in subgen. Coronopus considered an apomorphy. The ancestor of subgen. Coronopus probably had ribose in the seeds. Records from other subgenera are lacking. Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 RELATIONSHIP OF THE PLANTAGINACEAE 
Plantaginaceae has generally been considered an independent family, only Hallier (1903) included it in Scrophulariaceae. Regarding the relationship of the family, less concordance has prevailed. In older classifications it is often placed close to Plumbaginaceae (Jussieu, 1789; Lindley, 1847; Decaisne, 1852), later either in the same order as Scrophulariaceae, e.g. by Warming (1879, 1933), Wettstein (1903-1908), Thorne (1968, 1981, 1983), Takhtajan (1969, 1983, 1987), Dahlgren (1974-76, 1979-81, 1983), and Dahlgren (1989), or close to it in a separate order. Dalla Torre & Harms (1900-1907) places Plantaginales between Tubiflorae and Rubiales. Melchior (1964) places ‘9. Reihe Plantaginales’ between Tubiflorae and Dipsacales. Cronquist (1968) places Plantagi- nales between Lamiales and Scrophulariales, and later (198 1) between Callitrichales and Scrophulariales. In Bentham & Hooker (1876) the family is placed just after Lamiales, in Hutchinson (1973), between Primulales and Saxifragales. Plantaginales is according to Goldberg (1986) derived from Polygonales and that from Primulales and Caryophyllales. See also Dietrich (1970) for further discussion of many other proposals. 
Asteridae 
An outgroup for Plantaginaceae undoubtedly must be sought for within the subclass Sympetalae of Wettstein (1903-1908), the major part of which is named Asteridae by Cronquist (1968, 1981) and Takhtajan (1969, 1983). Cronquist (1981)had 1 1 orders in his Asteridae: Gentianales, Solanales, Lamiales, Callitricales, Plantaginales, Scrophulariales (incl. Oleaceae), Campanulales, Rubiales, Dipsacales, Calycerales, and Asterales. Takhtajan (1983) had three superorders in his subclass Asteridae: Gentiananae (with Gentianales, Oleales, Dipsacales, Loasales), Lamianae (with Polemoniales, Lamiales and Scrophulariales incl. Solanaceae and Plantaginaceae), and Asteranae (with Campandales, Calycerales, and Asterales). In Dahlgren (1989) the last seven dicotyledon superorders which follow after the Aralianae are: Asteranae, Solananae, Ericanae, Cornanae, Loasanae, Gentiananae, and Lamianae. Iridoids are often found in the last five, but always absent in the first two superorders. Lamianae ( = Lamiiflorae) includes Lamiales (with Scrophular- iaceae and Plantaginaceae), Hydrostachyales and Hippuridales. Ericanae, Lam- ianae, and Loasanae together are considered a probably monophyletic group. A study of 98 families was initiated years ago: 59 families within which the Asteridae smu Cronquist (1981), 10 from Cornanae, 10 Aralianae, 12 Ericanae (smu Dahlgren 1989) plus 7 other families within which either iridoids or verbascosides have been recorded. The families were scored for 55 chemical, morphological and PHYLOGENY OF PLANTAGINACEAE 157 embryological binary characters, which mostly were different from those used by Hufford (1992). Phylogenetic analysis using Hennig86 showed low constancy indexes. Adding, changing or deleting a character or a taxon often caused large changes in the dendrogram. Only 19 families within Lamianae (including Oleaceae but excluding Hydrostachyaceae) always remained as a clade. An expanded Asteridae including Ericanae, Cornanae, and Aralianae might be monophyletic. The most important characters which may support this are Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 embryological: (1) ovules unitegmatic, (2) integumentary tapetum (endothelium) present, and (3) ovules tenuinucellate. (4)Iridoids are often found inside Asteridae and very rarely outside (Jensen, 199 1). Four other characters are very rarely missing inside Asteridae, but are also found in families outside this group: (5) Corolla sympetalous, (6) carpels five or fewer, (7) the alternisepalous (epipetalous) stamens absent, (8) stipules absent. Characters which may support Asteridae in the sense of Cronquist (1968, 1981) and Takhtajan (1969, 1983) are: (1) vessels with simple perforation and (2) stamens inserted into the corolla tube. One of these characters is rarely missing in families inside this group, never both; one of them is rarely present in families outside and both are never found together. The families in Asteridae sensu Cronquist undoubtedly belongs to the same clade, but its delimitation downwards is uncertain. All the above mentioned ten characters are always present in Plantaginaceae. Some recent studies of Asteridae do not contradict the above conclusions, but most of them do not include Plantagmaceae (Wagenitz, 1992; HuRord, 1992; Downie & Palmer, 1992a, b; Olmstead, Michaels & Palmer, 1992). 
Lamianae 
Lamianae in Dahlgren (1 989) consists of three orders: Hippuridales with Hippuridaceae, Hydrostachyales with Hydrostachyaceae, and Lamiales with 1 7 families: Retziaceae, Stilbaceae, Buddlejaceae, Scrophulariaceae (inc. Orobancha- ceae and Selaginaceae), Myoporaceae, Globulariaceae, Plantaginaceae, Lentibular- iaceae, Pedaliaceae, Trapellaceae, Martyniaceae, Gesneriaceae, Bignoniaceae, Acanthaceae, Verbenaceae, Lamiaceae, and Callitrichaceae. The present study of Lamianae comprises 20 families, i.e. this superorder in the sense of Dahlgren (1989) plus Oleaceae. The following characters may support this as a clade: (1) Verbascosid is recorded from all the families except for Hydrostachyaceae, but it was not found in Salvia hispaizica in Lamiaceae, some Scrophulariaceae (Antirrhineae) and some Acanthaceae (‘I‘hunbergioideae).Verbascosid is found only outside Lamianae in Plocospma (Loganiaceae), in a single genus in Asteraceae (Echinaceu),and a single species in each of the families Icacinaceae, Cucurbitaceae and Magnoliaceae (Jensen, 1992). (2) Iridoids synthesized by the pathway which Jensen (199 1) designated “Route IIa followed by decarboxylation of C1,” (e.g. aucubin) is present in all families studied except for Hydrostachyaceae, Oleaceae, Gesneriaceae, Selagineae (in Scrophular- iaceae), Po&remum (in Buddlejaceae), and large parts of Lamiaceae. Trapellaceae has not been studied yet. Outside Lamianae this type of iridoids is only found in 158 KNUD RAHN Eucommiaceae, Garryaceae, and Aucubaceae. Secoiridoids are present in Oleaceae, but all kinds of iridoids are absent in the other Lamianae just mentioned. (3) Antraquinones are derived from shikimic acid in all the Lamianae studied (five families, 10 genera, 16 species). Elsewhere this derivation is only known from Rubiaceae (20 genera, 40 species) and in one species of Zingiberaceae. Antraqui- nones are derived from polyacetate in the other 24 families studied (Jensen, 1992). Plantaginaceae was not studied. (4) The Onagrad Type of embryogeny (Johansen, 1950; Davis, 1966;Johri et al., Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 1992), i.e. the zygote divides by a transversal wall, the terminal cell divides by a longitudinal wall, and the basal cell has no or only a minor part in construction of the embryo proper. In the 18 families which have been studied within Lamianae, the Onagrad type is found in all (includingHydrostachyaceae) except Oleaceae, but was not surveyed in Retziaceae and Stilbaceae. It is rare in the rest of the expanded Asteridae, being found only in Loganiaceae and a few Solanaceae, but it is common in primitive dicotyledons (e.g. Magnoliales, Ranunculales, Urticales, Violales, Myrtales) and may be plesiomorphic. In Oleaceae the Solanad and Caryophyllad type is found. Besides the Onagrad Type the Solanad Type is also found in Scrophulariacae, and the Chenopodiad Type in Lentibulariaceae. (5) Hairs with a stalk of one or more uniseriate cells and a head of two or more vertical cells are found in all the families in Lamianae except for Hydrostachyaceae, but they may be absent in some genera or species. Outside Lamianae found in Viburnaceae, Caprifoliaceae and Convolvulaceae. (Solereder, 1899; Uphof & Hummel, 1962; Carolin, 197 1; Napp-Zinn, 1973; Weberling, 1977; Rahn, 1992; and unpublished observations). (6) Young anthers with “placentoid” tissue (Chatin, 1870; Warming, 1873; Hartl, 1964; and own unpublished results) are found within Lamianae in Scrophulariaceae (except Veroniceae and Orobancheae), in Myoporaceae, Lentibulariaceae, Ped- aliaceae, Trapellaceae, Martyniaceae, Gesneriaceae, Bignoniaceae, Acanthaceae (except Thunbergieae), in Verbenaceae and Lamiaceae. It is absent in Plantagina- ceae, Globulariaceae, Hippuridaceae, and Oleaceae. Outside Lamianae found in Gentianaceae and Solanaceae (incl. Nolanaceae). (7) Less than five fertile stamens are found in all families within Lamianae except for Retziaceae, but not in all species within the Myoporaceae, Bignoniaceae, Gesneriaceae, Acanthaceae, Verbenaceae, and Scrophulariaceae. Outside Lam- ianae found in Donatiaceae, Stylidiaceae, Morinaceae, Triplostegiaceae and Columelliaceae. (8) Terminal endosperm haustoria (Dahlgren, 1991) i.e. from both the micropylar and the chalaza1 ends, are present in all families within Lamianae except Oleaceae, Hippuridaceae and some Acanthaceae (Thunbergieae). Outside Lamianae they are found in Fouqueriaceae, Actinidiaceae, Sphenocleaceae, Campunalaceae, Lobelia- ceae, Hydrophyllaceae, Ehretiaceae, and part of Boraginaceae (Heliotropoideae). Among the eight characters just mentioned six are present in Plantaginaceae, No. 6 is missing and No. 3 not yet studied. The position of Hydrostachyaceae is still dubious. Chemical characters are absent, the embryological are partly unknown and the morphological are dimcult to evaluate, because they are transformed to accommodate the special ecology of the family. For me there is no doubt that the rest of Lamianae belong to one monophyletic group, but it is uncertain whether its sister group will be found inside the narrow Asteridae (smu Cronquist and Takhtajan) e.g. Gentiananae, or outside, PHYLOGEM’ OF PLANTAGINACEAE I59 but still within an expanded Asteridae, e.g. in Ericanae or Loassanae as proposed by Dahlgren (1989). An argument mentioned e.g. by Warming (1933), Melchior (1964), and Willis (1966) for assuming a closer relationship between Plantaginaceae and Scrophular- iaceae is the similar reductions in the flowers of Veronica and Plantaginaceae: four corolla lobes, often four sepals. The superficial similarity between the two hollow seeds in Veronica sect. Cochlidiospermum ( = Cochlidiospermum Opiz) and those of subgen. Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 Psyllium and subgen. Albicans in Plantaginaceae may also be relevant. The seed of Veronica sect. Cochlidiospermum deviates from those of Plantaginaceae by having a funiculus, a small embryo with the radicle pointing upwards, and epitropous ovules. Takhtajan (1 969) claimed that Hydrostachyaceae “possibly had a common origin with the Plantaginaceae”. Dahlgren ( 197 7) argued: “Hydrostachyaceae, should be placed with the Sympetalae near Plantaginaceae and related families (see Jager- Ziirn, 1965; Wagenitz, 1977)”. Cronquist (1981), when discussing his order Callitrichales (including Hippuridaceae, Callitrichaceae, and Hydrostachyaceae) concluded: “It is not required that the Plantaginaceae as we now know them be directly ancestral to the Callitrichales, but at least a collateral relationship seems to be indicated”. He cited Hegnauer (1969) and Rauh & Jaer-Ziirn (1966, 1967) for arguments; however, when excluding the superficial similarity in habit between some species of Plantaginaceae and some species of Hydrostachys, the characters mentioned (unitegmatic, tenuinucellate ovules) are symplesiomorphic within Asteridae, and do not indicate a closer relationship between these two families than between them and most other families within Asteridae. Leins & Erbar (1990) claimed that the morphology of the male flower support the Hydrostachyaceae being closely related to Scrophulariaceae, but this was based on mere speculations, by interpreting the male flower as two stamens with synthecal anthers instead of one anther with two separated halves. Pilger (1937) and Dietrich (1970) related that the flowers of Plantago subgen. Bougueria are irregular, sometimes pentamerous and the corolla bilabiate. Dietrich (197 1) found two with five lobes among 14 corollas in female flowers of Plantago araucana ( = Littorella australis). This is possibly the uncited source of Cronquist (198 1) who stated “The floral anatomy of some species suggested the possibility of a pentamerous ancestry”. Studying several large collections of Plantago subgen. Bougum‘u, I found that the corollas although small were regularly tetramerous except in a very few (probably teratological) cases. A zygomorphic corolla (Characters [henceforth Ch] 54 and 55) is characteristic for many species in Plantago sect. Gnaphaloides (Rahn, 1979a; 1983), and a slightly zygomorphic calyx is common in subgen. Coronopus, subgen Plantago and subgen. Albicans. An-Ming (1990) performed a ‘preliminary cladistic study’ of the families placed in Lamianae (Lamiiflora) by Dahlgren (1980). Twenty-nine characters were used. The characters were polarized using three families as outgroups: Oleaceae, Cletraceae and Solanaceae. The cladistic parsimony program Hennig86 (Farris, 1988)was used. Three equally parsimonious cladograms were found (constancy index 0.43) and a strict consensus tree made. The results do not appear convincing to me. A study of the relationships within Lamianae was started several years ago. Further characters than those used for Asteridae were scored. The most promising characters are chemical or embryological, but they are often based on records from only one or a few species from each family, or are unknown. The better known I60 KNUD RAHN morphological characters are often present in both the plesiomorphic and the apomorphic state, especially within the larger families. Most families within Lamianae are presumably monophyletic, but not all. Verbenaceae and Lamiaceae together form one clade, but Verbenaceae alone is probably paraphyletic, and Lamiaceae is, according to Cantino (1992), polyphyletic. Scrophulariaceae has been characterized by synapomorphies for Lamianae only (see Thieret, 1967; Armstrong & Douglas, 1989). Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 Phylogenetic analysis of Lamianae using Hennig86 and a variable number of characters did not produce convincing results. Adding, changing or deleting a character often causes large changes in the cladogram (constancy indexes between 0.35 and 0.56). In the search for minor monophyletic lines both Scrophulariaceae and Acanthaceae were subdivided, but this did not produce more reliable results. Regarding the relationship of Plantaginaceae, the conclusion of the present study is that this family without doubt belongs to the core of the monophyletic Lamianae, but the studied characters do not conclusively indicate any family as the sister group. Plantaginaceae is probably an old group within Lamianae. Since submitting the manuscript for the present paper, a study of chloroplast rbcL and ndhF sequences in Lamianae has been published (Olmsted & Reeves, 1995). Plantago (P. lanceolata L.) is included and appears as the sister of Veronica (V. catenata Pennell). 
PLANTAGINACEXE 
The monophyletic status of Plantaginaceae is obvious. The following advanced characters support the statement: (1) leaves phyllodial and parallel-veined. (2) hairs in the leaf axil invariably present in Plantaginaceae and absent elsewhere in Lamianae (but found in some Asteraceae, e.g. Tarmacum). Wind pollination and associated adaptations include (3) scarious corolla, (4) absence of a disc, and (5) stigma long, filiform, dry, papillose, with two lateral bands of papilla, which undoubtedly developed from the united edges of two stigmata, in which two vascular bundles can easily be followed. This explanation is also supported by the occasional atavistic formation of two small stigmatic branches distally (as in most Lamiales). The plantaginaceous stigma is developed from two stigmata, not from a single punctiform or knob-like stigma. The only other Lamianae with a filiform, papillose stigma is H&wis (Hippuridales).It is also wind-pollinated, but the flower is much reduced, only one compartment in the ovary is fertile; the stigma is papillose all over and shows no indication of being formed from two (it was not possible to find vascular bundles). (6) Flowers in Plantaginaceae are protogynous. (7) An extension of the connective of the anther is characteristic of all Plantaginaceae. This appendage is rather small in subgen. Plantago, subgen. Littorella and subgen. Bougueria, but often large and flap-like in other subgenera, especially in sect. Hymenopsyllium. A flap-like appendix is found scattered throughout the system (e.g. in Asteraceae and Violaceae) and also in some Verbenaceae, Hippuridaceae, Biponiaceae and Buddlejaceae. All Plantaginaceae have (8) forate pollen grains with 4-15 apertures ( = peripor- ate). Erdtmann (1986) wrote: “The pollen grains in Plantaginaceae are f different from those in Acanthaceae, Campanulaceae, Labiatae, Plumbaginaceae, Polem- oniaceae [Heintze (1927) referred to the forate state of the pollen grains as additional PHnOGENY OF PIMAGINACEAE 161 evidence of a relationship between Plantaginaceae and Polemoniaceae] , Primula- ceae, Rubiaceae, and Scrophulariaceae” and that similar pollen characters are found in Caryophyllanae. Amongst the families belonging to Asteridae or containing iridoids, forate pollen grains are recorded by Erdtmann (1986) in the following families: Martyniaceae, Acanthaceae, Rubiaceae, Campanulaceae, Convolvulaceae, Polemoniaceae, Cobaeaceae, Icacinaceae, and Altingiaceae, but are common only in 
Polemoniaceae, Cobaeaceae, and Altingiaceae. Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 All species of Plantaginaceae need high light intensity, but their requirements regarding water are extremely variable. Some species grow in swamps or even in lakes, others are desert plants. 
Phylogenetic anabsis of Plantqinaceae 
Unfortunately, the study of Lamianae did not convincingly indicate any family as a probable sister-group of Plantaginaceae. The families which most often appeared as the sister group in the analysis were Buddlejaceae, Oleaceae, Globulariaceae, a clade of Retziaceae + Stilbaceae or Lentibulariaceae. When the two first families were used as the ‘ancestor’ in Hennig86 the result is given in Figure 4. With Globulariaceae, the tree started with a trichotomy of subgen. Plantago, subgen. Coronopus and the rest. With Lentibulariaceae as ‘ancestor’, Plantago subgen. Plantago appeared paraphyletic, with Plantago sect. Micropsyllium as the sister group of the rest. The branch with subgen. Littorella, subgen. Psyllium, subgen. Bougueria, and subgen. Albicans was not affected by the changed polarizations caused by the different ‘ancestors’. 
The common ancestor of the Plantaginaceae 
As a result of the phylogenetic analysis the nearest common ancestor of the Plantaginaceae, besides possessing the synapomorphies mentioned above, appears as a perennial herb with a tap-root. Its stem is very short with linear leaves spirally arranged in a rosette. Nerves are parallel. Leaves and other green parts have small 3-celled glands and simple uniseriate hairs on the surfaces. Scapes are produced from the leaf axils. Spikes of many, small, wind pollinated, protogynous flowers in a spiral, are each supported by a bract. Bracteoles are absent. Four sepals are free from each other. A gamopetalous, hyaline corolla has four lobes and four stamens inserted into the corolla tube. Ovary of two carpels, bilocular, is divided by the placenta which has a few ovules scattered on either side. Ovules are hemitropous, apotropous, ascending (micropyle pointing downwards). Fruit is a pyxis. Seeds are semi-ovoidal with copious endosperm and a straight embryo. The plane between the cotyledons is perpendicular to the symmetry plane of the seed (parallel to placenta and bract). Testa is mucilaginous. Chromosome number 2n = 12. 
THE GENUS PLehrT;4GO 
The result of the phylogenetic analyses, in which the characters are polarized 162 KNUD RAHN using either Oleaceae or Buddlejaceae as the outgroup, is shown in Figure 4. In this, Plantaginaceae is monogeneric and Plantago divided into six subgenera. Decaisne (1852), Pilger (1937) and later authors recognized three genera in Plantaginaceae: Littorella with three species, Bougueria with one, and Plantago with the remainder, comprising more than 200 species. However, Jussieu (1789), Sojak (1972), Holub (1973), and Dietrich (1980, 1982) also recognized Psyllium ( = Plantago sect. Psyllium in the sense of Pilger (1937), but not corresponding to subgen. Psyllium in Rahn (1978), which included the present subgen. Albicans), i.e. the two or three Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 phenetically most distinct groups are separated from the large polyphyletic Plantago. This is not acceptable when phylogenetic principles are followed. It is possible to argue for acceptance of one to fifteen genera (cf. Fig. 4). The rank of genus is used differently in different families, often depending more on tradition than on the nature of the family in question. In families like Brassicaceae and Apiaceae many small genera are recognized, while, for instance, Plantaginaceae and Begoniaceae are placed at the other end of the spectrum with few genera and most species placed in one large genus. The combination of very conspicuous family characters and less conspicuous characters separating monophyletic groups within the family (mainly because of the small size of the flowers) is probably the background for this tradition in Plantaginaceae. Actually I would have preferred to recognize the monophyletic groups, here designated subgenera, as genera and so used the three principal ranks: family, genus and species for expressing the taxonomy. However, I realized that the new genera probably never would be generally accepted, and the only result would be botanical literature burdened by 60 more synonyms. 
Synapomorphies uniting the subgenera 
Hair and seed characters are the most important for estimating the phylogeny of the infrageneric taxa within Plantago. However, in subgen. Littorella and subgen. Bougueria the seeds are strongly specialized, and for that reason not suitable for that purpose. The thin hairs (Ch 26) with a short basal cell (Ch 24) are synapomorphies for subgen. B0ugum.a and subgen. Albicans, so the ovary and seed of subgen. B0ugum.a are as a hypothesis considered evolved from ovary and seeds like those found in subgen. Albicans and subgen. Psyllium (Ch 82 two ovules; Ch 87: inner side of seeds deeply concave). The sister group of subgen. Littorella is probably subgen. Ayllium, but the synapomorphies uniting them are less convincing: few flowers in the inflorescence (Ch 32, also in Plantago sect. Olkanthos), and flowers in whorls of three (Ch 29, also in P. sect.Gnapha1oides). The ovary and seed of subgen. Littorella are also as a hypothesis considered as having evolved from ovary and seeds like those in subgen. Albicans and subgen. Psyllium. The cotyledons in the flower are perpendicular to the bract in subgen. Coronopus, subgen. Littorella, subgen. Psyllium, subgen. Bougueria, and subgen. Albicans; this also means perpendicular to the placenta (or the symmetry plane of the seed through the raphe passes between the cotyledons) except in subgen. Bougueria (Fig. 3, Sp 162), where the broad side of the curved seed is facing the bract and the seed attached to a mainly basal placenta. When the orientation of the cotyledons in relation to the raphe is used as a character (Ch 89) a reversal is needed for subgen. Bouguma in Figure 4. The small, acute glands (Ch 20), and a sharp keel or wing on the posterior sepals PHY1,OGF:NY 01' PIIWI'AGINACEAE 163 
(Ch 46) are two potential synapomorphies for uniting P. subgen. Coronopus and P. subgen. Albicans sect. Lance@lia, but are here considered as evolved independently. In Figure 4, Plantago subgen. Plantago is united by only one weak synapomorphy. 'The ancestor is supposed to have had sepals glabrous on the back (Ch 40, but several reversals have occurred within most sections). Broad leaves (Ch 5 is probably another synapomorphy, but then a reversal is needed for subgen. Micropsyllium (Figs 4 and 5). A solution only slightly less parsimonious is produced if subgenus Plantago is considered paraphyletic. Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 The only synapomorphy keeping subgen. Albicans together and separating it from subgen. Bougum'a is the presence of Ch 25 (cells forming the hairs have lumen very narrow and joints thick and refracting). However, the plesiomorphic state of this character: hairs with end-walls oblique, flat, distinct, and with lumen distinct, is present on the scapes of two sections from subgen. Albicans: sect. Lance$& and sect. Montanae (but not on their leaves, where Ch 25 is present). Both sections have x = 6 like subgen. Bougueria, but unlike the rest of subgen. Albicans. 
Plantago, key to the subgenera 
1. Ovary unilocular. Ovule solitary on a basal placenta. Fruit a small nut with a thick, hard wall (Fig. 3, Sp 144 and 162). 
BuddleJaceae 43~48 subgen. Plantago sect. Micropsyllium 1-42 subgen. Plantago sect. Plantago 49-80 subgen. Plantago sect. Mesembrynia 81-108 subgen. Plantago sect. Virginica 109-117 subgen. Plantago sect. flliganthos ser. Olig 118-119 subgen. Plantago sect fllig ser. Carpophnrae 120-131 subgen Plantago sect. fllig. ser Microcalyx 132-135 subgen. Coronopus sect. Maritima I, 136-142 subgen. Coronopus sect. Coronopus 20 46 48 143-145 subgen. Littorella U 89 146-161 subgen. Psyllium 
162 subgen. Bougueria fi bi ,- 163-168 subgen Albicans sect. Montana 169-174 subgen. Albicans sect. Lanceifolia 
175 subgen. Albicans sect. Bauphula 176-178 subgen. Albicans sect. Hymenopsyllium 179 subgen. Albicans sect. Albicans ser.0vatae 180-182 subgen. Albicans sect. Albicans ser. Minutae 183-185 subgen. Albicans sect. Albicans ser. Albicantes 90 7 191~195subgen. A. sect. Gnaphaloides ser Hlspidulae 
196-203 subgen A sect. Gnaphaloides ser Sericeae 204-206 subgen. A. sect. Gnaphaloides ser. Brasilienses :..-, 207~213subgen. A sect. Gnaphaloides ser Gnaphaloides 53 57 
Figure 4. Plantago, infrageneric taxa. On the Icft, one of nine most parsimonious trees otitainrd by using Hcnnig86 option ie-, 27 characters. The nine trccs have the shortest Icngth, 46 transformations. constancy index 0.58, retention index 0.85. Thc strict consensus tree is shown to the right. Synapomorphies of Plantaginaceae and autaponiorphies arc excludrd. A black IJar indicates a synapomorphy without reversals, an open har ow with rrvrrsals, a cross a reversal, and two parallel lines a parallelism. 164 KNUD RAHN 2. Flowers unisexual, monoecious. Scape short, with female flowers below, one male flower subterminal on a long pedicel. Ovule anatropous. Plant glabrous. subgen. Littorella 2'. Flowers bisexual, crowded in a small spike, and covered by broad bracts glabrous on their backs. Sepals very narrow and with long hairs. Corolla lobes small or absent. Anther solitary. Ovule amphitropous, seed kidney-shaped (Fig. 3, Sp 162 d). Hairs with a basal cell shorter than broad, distal cells long and Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 thin (Fig. 1, Ch 24) ...... subgen. Bougum'u 
132-213 other genera of Plantaginaceae - 1 P.canescens - N Asia NW Pmer. 2n-12 - 1- 1- 2 P.macrocarpa - NE Asia Nw Am. 2n-24 - 
19 P.tana1ensis - Madagascar 82 20 P.sparsiflora - SE USA 211-24 21 P.gentianoides - E Medit. 2n-12 22 P.reniformis - SE Europe. 2n-12 23 P.cornuti - 5 Europe. 2n-12 24 P.palmata - trop. Africa. 2n-24 25 P.africana - E Africa 26 P.major - Cosmopolite 2n-12 27 P.schneideri - China 28 P.coreana - Corea 29 P.asiatica - S b E Asia 2n-24 I- 30 P.taqueti - Corea 31 P.himalaica - Himalaya z- 32 P.aitchisonii - Himalaya 33 P.erosa - India 34 P.incisa - Java 35 P.rugelii - E N Pmerica 2n-24 36 P.eriopoda - N America 2n-24 37 P.tweedyi - USA 2n-24 38 P.cordata - SE USA 2n-24 39 P.hedleyi - Lord Howe 2n-24 40 P.maxima - E Europe. C Asia 2n=12 4-4 41 P.media - Europe. C Asia 2n-12.24 30 42 P.robusta - Sta.Helena 43 P.tenuiflora - E Eur. C Asia 2n-24 
Figure 5. Plantqo sect. Plonfugo (Sp 142) and sect. Mic7opsylliurn (Sp 4348). On the left one of eight trees produced by Hennig86, option mhennigf. Length 22, constancy index (ci): 0.63, retention index (n):0.91; 16 characters. PHYLOGENY OF PIANTAGINACME 165 
TAHLC1. Plantaginaceae, distribution of charactcrs in infrageneric taxa of Plantogo and some possible outgroups (families in Lamianae). The first two lincs indicate the number in 'List of characters for the Plantaginaceae'. In the table, 3 indicates that all members exhibit thr apomorphy, 0 that all exhibit the plesiomorphy, 1 and 2, that both the apomorphy and the plesiomorphy are present, , + and ? indicate lack of information. 3, 2 and + are supposed apnmorphies, 0, 1 and - wpposed plesiomorphies. 
1 11111 22222 22233 34444 45556 66666 77788 88889 123452 35678 01456 78901 60168 93671 23789 14623 57890 Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 003303 00000 00000 00300 01000 00000 00011 00011 10000 01 eaceae 003302 10000 00000 00300 01001 10000 00002 00000 30000 Buddlejaceae 003000 13300 00000 00330 31000 00000 00002 00000 300?0 Sti 1baceae 003000 03300 00000 00330 31000 30000 00030 00000 OOO?O Retzi aceae 003300 00000 00000 00030 31000 00000 00033 03033 300?0 Globulariaceae 301300 00000 00000 00003 03000 00000 00033 00000 30000 Lenti bulari aceae Plantaginaceae. Plantago subgen. Plantago 101021 00010 00000 00010 03100 00002 11111 00010 10000 1-42 sect. Plantago 030003 00030 00000 00000 03000 00003 33033 00000 30001 43-48 sect. Micropsyllium 110032 00020 00000 00010 02200 00002 11121 00300 20000 49-80 sect. Mesembrynia 111021 01110 00000 00000 02300 00001 10031 00300 10000 81-108 sect. Virginica sect. Oliganthos 000031 10020 00000 00003 03000 00003 11021 00000 20000 109-116 ser. Oliganthos 300030 300+3 00000 00003 03000 00001 00300 00000 10000 117-118 ser. Carpophorae 300021 30032 00000 00003 03100 00003 11031 00000 30000 119-131 ser. Microcalyx subgen. Coronopus 000001 00030 30000 00000 01333 00003 10111 00000 20030 132-135 sect. Maritima 010003 00030 30000 00000 01333 00003 10011 00000 30033 136-142 sect. Coronopus 300000 300-0 03000 00303 00000 00003 00300 030+3 3-030 143-145 subgen. Littorella 013312 01000 03000 00303 00000 00001 10111 00030 13030 146-161 subgen. Psyllium 000000 03300 00300 00030 30000 00003 33033 030+3 3-000 162 subgen. Bougueria subgen. A lbicans 010000 01320 OD330 30021 32000 00001 11111 10030 13030 163-168 sect. Montana 100010 00330 30330 30031 01330 00001 00300 30030 13030 169-174 sect. Lanceifolia 033010 00300 03330 00031 30000 00000 00033 00030 03033 175 sect. Bauphula 030003 13300 01330 33011 10300 00101 00010 00030 13033 176-178 sect. Hymenopsyllium sect. Albicans 020000 00300 00330 30031 30000 00030 00030 30030 03333 179 ser. Ovatae 030000 01300 01330 31011 00100 30303 10111 00030 03333 180-182 ser. Minutae 023000 03300 03330 30000 30000 30011 00300 30030 03333 183-185 ser. Albicans 032012 12300 03330 30001 00000 30301 11031 30030 03333 186-190 ser. Ciliatae sect. Gnapha 7oides 030003 00330 00333 00301 00300 00102 11133 00030 03033 191-195 ser. Hispidulae 001001 11330 00333 00311 00300 00111 00111 00030 13033 196-203 ser. Sericeae 001000 03330 00333 00301 00300 03031 00111 00030 13033 204-206 ser. Brasilienses 030001 13330 00333 00301 00300 03031 00211 00030 03033 207-213 ser. Gnaphaloides 
1'. Ovary bilocular, placenta on the septum with one or more ovules. Ovule hemitropous. Fruit a pyxis (very rarely a nut with a thin soft wall). 3. Leaves opposite or verticillate. Internodes long. Flowers opposite or in whorls of three, arranged in short spikes. Ovary with two ovules. Pyxis with two seeds. Placenta side of seed deeply concave, cotyledons in the seed perpendicular to the bract and the placenta (Fig. 3, Sp 15 1). Hairs formed of a row of short wide cells, the basal cell not differentiated ...... subgen. Pgllium 3'. Leaves alternate. Internodes usually short, leaves in a rosette. 4. Ovary with two ovules. Pyxis with two seeds. Placenta side of seed deeply concave, cotyledons in the seed perpendicular to the placenta side or bract (Fig. 3, Sp 175-201). Hairs with a basal cell shorter than broad, distal cells long and thin (Fig. 1, Ch 24). Leaves often linear and spike usually short in relation to scape ...... subgen. Albicans 4'. Ovary with one to many ovules. One to many seeds. Placenta side of seeds 166 KNUD RAHN flat, rarely slightly concave or convex. Hairs on leaves formed of a row of short wide cells, the basal cell not differentiated. Spike often long and about equalling scape. 5. Corolla tube with short hairs all over. Cotyledons in the seed perpendicular to the placenta side or bract (Fig. 3, Sp 135 and 140). Leaves linear to narrowly elliptic. Ovary with two to six ovules ...... subgen. Coronopus Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 5'. Corolla tube glabrous. The plane between the cotyledons in the seed parallel to the placenta side and bract, at least when one or two seeds are present (Fig. 3, Sp 20-1 10). If many seeds are developed they may press against each other, and the plane between the cotyledons may be oblique or even perpendicular to the placenta side. Leaves broadly elliptic to linear. Ovary with two to many, rarely one ovule ...... subgen. Plantago 
Plantago subgen. Plantago 
This subgenus is characterized mainly by plesiomorphic characters. Most species are perennial; the primary root is usually a more or less fleshy tap root, rarely a woody root as in subgen. Albicans, or it remains thin as in annual species and adventitious roots then develop from the caudex. An elongated stem is absent except in five insular endemics. The number of ovules and seed varies between one and about 35. The plane between the cotyledons is parallel to the placenta-side of the seed. The number of characters which can be used for a cladistic analysis is low; for that reason clades within the recognized infrageneric taxa are uncertain, and the arrangement of species in Figures 5-8 should not be considered conclusive. The variation between species within subgen. Plantago is remarkably low, especially when compared with that found in subgen. Albicans. Four sections are recognized; they are treated below under separate headings.2'. sect. Micropsyllium, sect. Oliganthos, and sect. I/'irginica are monophyletic, while sect. Plantago and sect. Mesembgnia are paraphyletic (see Figs 4 and 5). The synapomor- phies for sect. Micropsyllium are also present within the other subgenera of Plantago but never all together. The subgenus Plantago is distributed in all continents and found on many oceanic islands, often in mesic or moist habitats. It is interesting to note that sect. Plantago is absent from New Guinea, Australia, New Zealand and South America, (except for P. major, which is traditionally considered introduced by man) and replaced there by the three sections Oliganthos, Mesembgnia and Virginica. Section Plantago is present on islands in the Pacific Ocean not very far from those continents, on Lord Howe Island 600 km east of Australia, on Auckland Island 500 km south of New Zealand, and on Juan Fernandez 600km west of Chile. Pollen of Plantugo was common from top to bottom in peat deposits in Hawaii (Selling, 1947, 1948). It is possible to explain the distribution and the scarcity of synapomorphies amongst the often very diverse species by accepting a vicariance hypothesis, which supposes that an ancestral species was once common throughout most of the world. For one reason or another most populations were exterminated more or less simultaneously, and speciations occurred in places where populations survived, e.g. on the islands just mentioned and in Australia, Antarctica and South America. In the PHYLOGENY OF P1ANTAGINAC;LZE 167 last three areas and on some islands they speciated again later on, and the present species in sect. Mesembvnia, sect. Oliganttios and sect. firgznica were formed (see further under the sections and series). In the early Tertiary there were probably, according to Bramwell (1979), in the Pacific area more and larger islands with high mountains where Plantago could grow, even in the tropical zone. 
Ki to sections and series Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 1. Ovary with 1-5 ovules and with a third compartment near the top on the adaxial side or a rudiment of it, seen as a thickening on the apex of the ripe placenta; the other ovules fixed to the side near the centre. When two seeds are present on the same side of the placenta a keel develops between them (Fig. 3, Sp 68-108). 2. Ovary with 5 or 4 ovules. Corolla lobes always patent during anthesis. Australia, New Zealand, Amsterdam and St. Paul Islands, a few species in Asia and one in Europe ...... sect. Mesembrynia 2’. Ovary with 3 or 2 ovules (very rarely 1 or 4 in some ovaries). Corolla lobes often erect during anthesis and with small anthers included. North and South America ...... sect. firginica 1’ Ovary with one to many ovules, scattered on the placenta when more ovules are present. No compartment or rudiment of it on top of the placenta, and placenta without distinct keels separating the mature seeds. 3. Small annuals with small flowers, linear leaves and narrow seeds (Fig. 3, Sp 43). America, Asia, Europe ...... sect. Micropsyllium 3’ Perennial plants; leaves rarely linear, and fleshy if so. 4. Spikes with 1-3 rarely up to 12 flowers. South America, New Zealand, Australia, New Guinea ...... sect. Oliganthos 5. Adventitious roots. 6. Scape very short, a carpophore develops during the ripening of the fruit. Flowers usually unisexual. Bracts and sepals long, membraneous. The Andes and Mexico ...... ser. Carpophorae 6’. Scape often short during anthesis and elongates during the ripening of the fruit, no carpophore develops. Flowers bisexual. Bracts and sepals either very small or with a green nerve. New Guinea, Australia, New Zealand ...... ser. Microcabx 5’. Tap root. South America, Auckland, Tasmania ...... ser. Oliganttios 4’. Spikes usually with many flowers. Africa, Europe, Asia, N America, Oceania ...... sect. Plantago 
Plantago sect. Plantago This paraphyletic section comprises the species which Pilger (1937) placed in sect. Pohneuron ( = P. sect. Plantago, with 19 species), in sect. Palaeopylliurn (27 species + P. africana Verdc.; excl. P. dielsiana, cf. Rahn, 1974), sect. Holopyllium ( = P. macrocarpa), sect. Lamprosantha (P. media, P. maxima, and P. canescens), sect. Gentianoides (= P. gentianoides) and sect. Eremopyllium ( = P. renijinnis). Figure 5 is a cladogram based on 16 characters only. It is possible to separate two clades with a reasonable degree of probability: the species Sp 40-41 and Sp 26-35.The last group of species is by Pilger (1937) placed in sect. Pobneuron, they have only one synapomorphy “a pedicel-like narrowing of the calyx” (Ch 34,but this character is usually absent (a reversal) in P. major, which otherwise is very similar 168 KNUD RAHN phenetically. Pilger (1937)mentions 19 species in sect. Pohneuron ( = sect. Plantago, but five are recognized equivocally and were not included in his key to species. A revision of the species in this section is much needed. At least three species will survive a revision: P. rugelii Decne. (Sp 35) in North America, P. major L. (Sp 26) now with a world-wide distribution and with at least three infraspecific taxa (P. major ssp. intennedia (DC.) Arcang., and var. sinuata (Lam.) Decne.) and P. asiatica L. (Sp 29) in South and East Asia. The following may be closely related species or considered Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 subspecies under P. asiatica L.: P. erosa Wall. (Sp 33), P. incisa Hassk. (Sp 34), P. taqueti fiv. (Sp 30), and P. coreana Lirv. (Sp 28); more dubious, but probably with the same affinity are: P. hasshrlii Decne. (not included), P. schnezderi Pilg. (Sp 27), P. centralis Pilg. (not included), P. caualeria‘ Ltv. (not included), P. alata Nakai (not included), P. himalaica Pilg. (Sp 3 l), P. aitchisonii Pilg. (Sp 32), P. sawadai Yamam. (not included), P. jinnosana Tateishi & Masam. (not included), P. japonica Franch. & Sav. (not included), P. yesoensis Pilg. (not included), and P. hakusanensis Koidz. (not included). The chromosome number of P. major is normally 2n = 12, and of P. rugelii 2n = 24. The chromosome numbers recorded for the Asiatic species is 2n = 24 and less frequently 2n = 36; a single record of 2n = 12 may be of a misidentified P. major. Species possibly related to that group are P. gentianoides Sibth. & Sm. (Sp 2 1) from south-east Europe and western Asia, P. ren@mis Beck (Sp 22) from the Balkans and P. cornuti Gouan (Sp 23) from southern Europe. The African species P. palmata H0ok.f. (Sp 24),and P. aficana Verdc. (Sp 25) have broad leaves like P. major and may also be related. P. tunalensis Baker (Sp 19) from Madagascar, P. longissima Decne. (Sp 18) from South Africa, and P. spars$?ora Michx. (Sp 20) from U.S.A. have two ovules (Ch 82) like the African P. fischeri Engl. (Sp 16), and P. rmota Lam. (Sp 17). P. aucklandica Ho0k.f. (Sp 15) from Auckland Island (south of New Zealand), P. rapemis Pilg. (Sp 14) on the island Rapa in the southern Pacific, and P.fernandezza Barntoud (Sp 5) from Juan Fernandez (south-easternPacific) also have two ovules in the ovary, but they are probably only distantly related to the African species. Another and very distinct species found on the island Rapa is P. rupicola Pilg. (Sp 3), which has a woody stem like P.fernandeziana, P. principes Cham. & Schltdl. (Sp 4) and P. robustu Roxb. (Sp 42). P. robusta is a peculiar woody endemic on Santa Helena, it has very small flowers and linear leaves and for that reason inevitably appears as “the sister group” of sect. Micropsyllium in cladistic analyses, when these size characters are used, although P. robustu more probably is related to South African members of the section Plantago. The North American species P. eriopoda Torr. (Sp 36) and P. tueedyi A. Gray (Sp 37) are probably related, but the affinities of P. cordata Lam. (Sp 38) are not clear. Plantago hedlyi Maiden (Sp 39) is from Lord Howe Island, east of Australia. It is the only Pacific species with antrorse hairs (Ch 17) on the scapes, and it is notable that this character is also present in most species of sect. Mesembrynia and sect. Oliganthos from the Australian mainland and South America. P. media L. and P. maxima Jacq. (Sp 40 and 41) from Europe and Asia have probably a common ancestor, they were by Pilger (1937) united with P. canescens Adams (in sect. Lamprosantha). In the present study P. camscem appears as the most primitive species of Plantago (Sp l), it has 2n = 12 chromosomes and is distributed in three disjunct areas in northern Siberia, one in Alaska and north-west Canada, and one in the Rocky Mountains. P. macrocarpa Cham. & Schltdl. (Sp 2) found along the coast of north-west America and the Aleutian Islands is also rather primitive like the species on the Pacific islands PHYLOGENY OF PIANI’AGINACEAE 169 
(Sp 3-15). The nine Hawaiian species recognized by Pilger are found in Table 2 and Figure 5, but Wagner (1990) recognizes only three: P. principes Cham. & Schltdl., P. hawaiensis (Gray) Pilg., and P. pacbpLylla A. Gray (including P. muscicola Pilg., P. hillebrandii Pilg., P. glabnzlia (Rock) Pilg., P. melanochrous Pilg., P. krajinai Pilg., and P. grayana Pilg.). The South African P. laxzj?ora Decne, is excluded from Figure 5 and Table 2 because of lack of information. Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 Plantago sect. Micropsyllium Decne This section is considered monophyletic (Fig. 5, Sp 4348). It consists of small annuals (Ch 2) with very narrow leaves (probably a reversal of Ch 5), spur-like elongations (Ch 23) on the antrorse non glandular hairs (Ch 14, and with small size of all floral parts (Ch 62, 63, 68, 69) and 8.5).The seeds are narrow (Fig. 3, Sp 43). Its ancestor is probably within the paraphyletic sect. Plantago, but the result of the phylogenetic analysis, which points at P. robusta as the sister group, is probably wrong. P. bkelovii A. Gray, P. elongata Pursh, P. heterophylla Nutt., and P. pusilla Nutt. are North American and monographed by Bassett (1966). P. heterophylla has also been found in Argentina and Paraguay for more than 100 years. Dietrich (1980) placed them in a new section Diandra, while P. pobspenna Kar. & Kir. from C. Asia and P. tenuzjlora Waldst. & Kit. from Europe and central Asia were left in sect. Micropyllium. Two stamens (Ch 64) in the American species is probably a good synapomorphy, but no synapomorphy unites P. pohspemza and P. tenugora. That the similarity between the species in the New and Old World should be “ein parallel Entwicklung auf Grund der okologishen Adaptation” appears improbable to me. The chromosome number 2n = 20 related by Bassett (1966) for P. bigelovii is the only reliable record of x = 5 in the subgenus Plantago, although not confirmed by others. 
Plantago sect. Mesembrynia and sect. Virgznzca The common ancestor of this group probably had five ovules in the ovary, with two ovules fixed in the middle on either side of the septum and one fixed near the top on the posterior side (Ch 76, Fig. 3, Sp 68), keels are prominent on the mature placenta between the seeds. All species in sect. Virginica and some species in sect. Mesemblynia have no ovule in the upper compartment (Ch 73, which is transformed to a thickening usually visible on the placenta in the mature pyxis; and the placenta is rarely with a visible split at the top, which is common in sect. Plantago and sect. Olkanthos. In sect. Virginica there are usually three (Ch 78, Fig. 3, Sp 108), less frequently two or four ovules present, and the placenta is as just described (Ch 76, Fig. 3 Sp 93). In P. camtschatica (Sp 52) an upper ovule and seed is always present but at least in cultivated specimens more than two ovules and seeds are sometimes found on either side below (Pilger (1937: 275) and own observations). This may be interpreted as primitive. The chromosome number is known of 24 species in sect. Mesembrynia and 19 in sect. Erginica. Fifteen species have 2n = 12, and all belong to sect. Mesembrynia; the rest have 2n = 24, 36 or 48. Plantago sect. Mesembrynia and sect. Virginica occupies both dry and wet habitats in lowlands as well as alpine areas, but usually in warmer and drier habitats than those of sect. Olkanthos. Plantago sect. Ergznica is monophyletic and sect. Mesemblynia paraphyletic. I70 KNUD RAHN 
TABLE2. Distribution of characters in species of Pluntugo subgen. Plantogo. For explanations see under Table 1. Local autapomorphies: Ch 43 (in Sp 2, also in Sp 163-168), Ch 45(in Sp 53, also in Sp 163-168), Ch 90 (in Sp 48, also in Sp 135-142 and Sp 175-213). AU species in this table i.e. the subgenus Plantogo have the apomorphic state of Ch 84, and all have the plesiomorphic state of the following characters: Ch 4, Ch 19, Ch 20, Ch 21, Ch 22, Ch 24, Ch 25, Ch 26, Ch 27, Ch 28, Ch 29, Ch 34, Ch 35, Ch 36, Ch 37, Ch 39, Ch 42, Ch 44, Ch 46, Ch 47, Ch 48, Ch 49, Ch 50, Ch 51, Ch 52, Ch 53, Ch 54, Ch 55, Ch 56, Ch 57, Ch 59, Ch 65, Ch 71, Ch 73, Ch 74, Ch 75, Ch 83, Ch 87, Ch 88, and Ch 89. 
11 11111 11233 33344 56666 66666 77777 78888 8 12356 78901 23456 78301 23801 80123 46789 02678 90125 6 Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 subgen. Plantago sect. Plantago 00030 00000 30000 00000 00033 00000 00000 00000 00003 0 1 canescens 00030 00300 00000 00000 00033 00300 00000 00000 00030 3 2 mcrocarpa 00333 30000 30000 0--00 00030 00000 00300 00000 0000- - 3 rupicola 00330 00000 00000 O-OOO 00032 00300 0-300 00000 03000 0 4 principes 00330 00000 00000 -0000 00030 00300 00300 00000 0003- - 5 fernandezia 
30030 00000 10000 00000 00033 00330 O-OOO 00000 00002 0 6 hawaiiensis 30030 00000 10000 00000 00020 00330 00200 00000 03002 0 7 pachyphylla 30030 00000 00000 00000 0003- 0-+-- 0---- -0000 00--+ - 8 mscicola 30030 00000 00000 00000 00030 00300 00010 00000 03002 0 9 hillebrandii 30030 00000 00000 00000 00030 00300 0---- -0000 00000 0 10 melanochrous 
30030 00300 00000 0--00 00030 00000 00300 00000 00000 0 11 glabrifolia 30032 00300 00000 0--00 00030 00200 00300 00000 00003 0 12 krajinai 30030 00300 00000 0--00 00030 00330 00200 00000 00003 0 13 grayana 30033 33030 -0000 ---00 0003- 0-+-- 0-+-- -0000 0003- - 14 rapensis 30033 00000 30000 0--00 00030 00300 0-030 00000 00030 0 15 aucklandica 
30033 33030 00000 0--00 00033 00330 0---- -0000 00030 0 16 fischeri 30000 00000 00000 00000 00033 00330 00300 00000 00030 0 17 rmta 30030 33000 00000 3-+00 00030 00330 O-OOO 00000 00030 0 18 longissima 30033 30030 30000 3-+00 00033 00333 0---- -0000 00031 0 19 tanalensis 30030 30000 30000 30300 00033 00330 00000 00000 00031 0 20 sparsiflord 
30030 20000 30000 2-200 00030 00300 03030 00000 00000 0 21 gentianoides 30033 30030 30000 30300 00033 00330 03300 00000 03001 0 22 reniformis 30033 30300 30000 30300 00030 00330 00000 00000 00300 0 23 cornuti 30033 30030 30000 30300 00033 00330 01033 00000 00000 0 24 palmta 30033 30000 30000 30300 00032 00332 0-030 00000 00001 0 25 africana 
30033 33030 30000 30300 00230 00333 00033 00000 00003 0 26 major 30033 33030 30000 30300 00333 00330 00030 00000 OOOO+ - 27 schneideri 30033 30030 30000 30300 00333 00300 00000 00000 OOOO+ - 28 coreana 30033 30030 30000 30300 00333 00333 00033 00000 00003 0 29 asiatica 30033 30030 30000 30300 00333 00333 00030 00000 03000 0 30 taqueti 
30030 +-000 30000 3-+00 00333 00300 O-OOO 00000 OOOO+ - 31 himlaica 30033 30000 30000 30300 00330 00330 O-OOO 00000 OOOO+ - 32 aitchisonii 30030 30000 30000 3-300 00333 00333 00033 00000 00003 0 33 erosd 30030 30000 30000 3-+00 00333 00300 0---- -0000 00003 0 34 incisa 30033 33030 30000 30300 00330 00333 00030 00000 00002 0 35 exaltata 
00030 00000 30000 30300 00033 00330 00033 00000 03000 0 36 eriopoda 00030 00000 30000 30300 00033 00300 00000 00000 00300 0 37 tweedyi 00033 30030 00000 +O+OO 00030 00330 00300 00000 03000 1 38 cordata -0130 00300 30000 3-+00 00030 00310 0-030 00000 00003 0 39 hedleyi 00033 33330 00000 30330 00030 00000 03000 00000 00302 0 40 maxima 
00033 30000 30000 30330 00033 00000 03000 00000 00002 0 41 media 00300 00000 30000 30300 00030 00333 00030 00000 00003 0 42 robusta sect. Micropsyl lium 03000 00000 30000 30300 00030 10333 00033 30000 00003 0 43 tenulflora 03000 00000 30000 30300 00030 30333 00033 30000 00003 0 44 polyspem 03000 00000 30000 30300 00030 10333 30033 30000 00003 0 45 heterophylla 03000 00000 30000 35300 00030 10333 30033 30000 00003 0 46 elongata 03000 00000 30000 30300 00030 10333 30033 30000 00003 0 47 pusilla 
03000 00000 30000 30300 00030 00333 30033 00000 00003 0 48 bigelovii PHYLOGENY 01; PIANTAGINACEAL 171 
11 11111 11233 33344 56666 66666 77777 78888 8 12356 78901 23456 78301 23801 80123 46789 02678 90125 6 sect Mesmbrynia 00020 00000 +OOOO 0--30 0003+ 00333 0---- -0330 00003 0 49 komarovi 00030 30000 +OOOO +--00 0003+ 00300 00300 00330 00002 0 50 perssonli 00030 00000 00000 00000 00033 00330 0---- -0330 OOOO+ 0 51 aracbnoidea 00030 30000 30000 30000 00033 00330 00030 00300 00003 0 52 camtscbatica Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 00030 30000 30000 30300 00030 00330 00030 00300 00003 0 53 scbwarzenbergiana 
00033 30000 30000 30300 00033 00333 00033 00300 00003 0 54 depressa 00030 30000 30000 30300 00030 00300 00030 00310 00003 0 55 antarctica 00030 00000 00000 3-+31 00030 00300 0--+- -0310 OOOO+ - 56 picta 03030 -0000 30000 3-+00 0000- 00330 00033 00300 00000 0 57 cunninghami 03030 00000 30000 30300 00003 00330 00033 00300 00003 0 58 turrifera 
03030 -0000 30000 3-+00 OOOO+ 00333 00033 00310 00003 0 59 WltiSCapd 03030 30000 30000 30300 00003 00333 00033 00300 00000 0 60 debilis 0+030 00000 30000 30300 00003 00330 00033 00300 OOOO+ - 61 eauana 00030 00000 30000 30300 00003 00000 00010 00310 00003 0 62 exilrs 00030 -0000 30000 3-300 OOOO+ 00300 00300 00300 00001 0 63 ga~d~chaudi 
00030 00000 30000 3-+00 00003 00300 0---- -0300 OOOO+ - 64 bellidioides 00030 00000 30000 30+00 00000 00000 00000 00300 00003 0 65 varia 00030 00000 00000 3-+30 00003 00000 0--+- -0310 OOOO+ - 66 g7abrata 30030 +OOOO 30000 3-+00 0003- 00300 00030 00300 00002 0 67 cladaropbylla 30030 20000 30000 30331 00030 00333 00033 00310 00003 0 68 raoulii 
30030 -0000 00000 3--00 0003- 00300 0--+- -03+0 00003 0 69 mntedicbsonii 30030 30000 00000 3--00 00033 00333 0--+- -0310 00002 0 70 tricbophora 30030 -0000 00000 3--00 0003- 0-+++ 00033 00300 OOOO+ - 71 papuana 30030 10000 30000 30000 00030 00333 00030 00330 00003 0 72 eurypbylla 30030 00000 00000 3-30 00033 00333 0--+- -0330 00003 0 73 tasmnica 
30030 -0000 30000 3--30 0003- 0-+++ 0--+- -0330 OOOO+ - 74 daltoni 30030 00000 30000 30030 00031 00333 00030 00330 00003 0 75 alpestris 30030 30000 30000 30331 00003 00300 00030 00330 00003 0 76 spatbulata 00030 00000 30000 30000 00003 00330 00030 00310 00003 0 77 bispida 00030 00000 30000 00030 00003 00000 00030 00300 00003 0 78 stauntonii 
03030 00000 30000 0-010 00003 00000 0--+- -0300 00003 0 79 pentasperma 03030 00000 30000 00000 00000 00333 00030 00300 00000 0 80 drumndii sect. Virginica 00300 00000 00010 1-000 00033 00000 00030 00333 00003 0 81 trinitatis 00000 00000 00003 O-OOO 00003 00100 00030 00333 10000 0 82 weddelliana 00000 00000 00033 O-OOO 00013 00100 00030 00333 10000 0 83 comnersoniana 00030 30000 30010 00000 00023 21100 00031 10333 00002 0 84 tomentosa 00030 30000 30000 O-OOO 00013 22000 00031 10333 00001 0 85 catbarinea 
00030 30000 00001 O-OOO 00013 11100 00030 00333 00000 0 86 argentina 00030 30000 30020 O-OOO 00003 01000 00030 00333 10000 3 87 ventanensis 00030 30000 30033 O-OOO 00003 20000 00031 10333 00001 0 88 floccosa 00030 30000 00003 1-000 00013 20000 00030 00333 00002 0 89 guilleminiana 30030 20000 30000 1.000 00013 00200 00030 00333 30003 0 90 turficola 
03030 30000 30000 00000 00003 21110 00031 10333 00002 0 91 myosuros 03030 20000 30000 1.-00 00003 31000 00033 30333 00002 0 92 tenuipala 03030 30000 30000 00000 00003 20110 00031 10333 30003 0 93 virginica 03030 30000 30000 O-OOO 00003 21000 00031 10333 30000 0 94 rhodosperm 03030 30000 30000 O-OOO 00003 30210 00033 30333 30002 0 95 galapagensis 
03030 30000 30000 10000 00003 21100 00031 10333 30002 0 96 firm 00030 30000 30000 30000 00033 20310 00031 10333 00003 0 97 pacbyneura 00030 30000 10000 20000 00023 11110 00031 10332 00001 1 98 orbignyana 03030 30000 00000 3-300 00033 20100 00031 10333 30000 0 99 penantba 00030 30000 00000 3-300 00033 00100 00030 00333 30000 0 100 dielsiana 172 KNUD RAHN 
TABLE2 continued 
11 11111 11233 33344 56666 66666 77777 78888 8 12356 78901 23456 78301 23801 80123 46789 02678 90125 6 
00000 20000 10000 3-300 00033 00200 00030 00333 30001 0 101 jujuyensis 00030 20000 00000 3-200 00013 20100 00031 10333 00000 0 102 venturii 00030 30000 30000 30000 00013 21000 00031 10333 00002 0 103 buchtienii 00030 30000 30000 2-000 00003 23000 00031 10333 10000 0 104 truncata 00030 30000 30010 2-000 00003 22000 00031 10332 00000 0 105 berroi Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 00030 20000 30000 3-100 00013 20000 00031 10333 00001 0 106 subnuda 00030 30000 00000 2-000 00003 22000 00031 10333 30001 0 107 alismatifolia 30020 20000 10000 30100 00013 11110 00031 10333 10001 0 108 australis sect. Oliganthos ser. 07iganthos 00030 00000 00000 20003 00030 00320 00033 00000 0000- - 109 triantha 00020 00000 30000 30003 00030 00300 00110 00000 00001 1 110 barbata 00033 00000 30000 3--03 00030 00310 00030 00000 0000- - 111 moorei 00031 00000 00000 20303 00030 00320 00030 00000 00300 1 112 tehuelcha 00030 00003 00000 3-003 00030 00300 00000 00000 00001 0 113 correae 
00000 00003 00000 30303 30030 00300 00020 00000 00001 0 114 uniglumis 00030 00003 03000 +-to3 30030 00320 00030 00000 00003 0 115 smpervivoides 00030 00000 33000 30003 30030 00320 00010 00000 00002 0 116 pulvinata ser. Carpophorae 30030 00001 03000 +3-03 33030 00100 00300 03000 00001 0 117 tUbUlOsa 30030 00001 03000 +3-03 33030 00000 00300 03000 00001 0 118 rigida ser . Microca lyx 30030 00000 33000 3+003 33030 00300 0003+ OODDO ODOO+ - 119 unibracteata 30030 00000 33000 3+-03 30033 00320 0003+ 00000 00003 0 120 lanigera 30030 00000 33300 3+-03 30030 00300 00030 00000 00003 0 121 paradoxa 30030 00000 03300 3+-03 00030 00320 00-++ -0000 OOOO+ - 122 novae-zelandiae 30032 00000 03300 2+-03 3303- 00323 00033 00000 00003 0 123 gunnii 
30030 00000 33300 32003 33030 00320 10033 00000 OOOO+ - 124 triandra 30000 00000 03300 ++-03 30030 00313 00033 00000 00003 0 125 stenophylla 30000 00000 03300 33003 30030 00310 00033 00000 00002 0 126 polita 30000 00000 33300 +3-03 30030 00313 00033 30000 00002 0 127 depauperata 30030 00000 03300 33203 30030 00303 00030 00000 00003 0 128 aundensis 
30030 00000 02000 ++to3 00030 00321 00031 00000 00303 0 129 glacialis 30030 00000 03300 23303 00030 00310 00033 00000 00303 0 130 palustris 30030 00000 03300 33303 0003- 00300 00030 00000 00302 0 131 melleri 30000 00000 ?33?? ???03 3?03? 00000 ??O?? ?lo00 00003 0 obconica 
Plantago sect. Mesembrynia (Decne.) Rahn Species with adventitious roots (Ch I) form a clade (Fig. 6, Sp 67-76): P. montedictboni P. Royen, P. papuana P. Royen, and P. t~~Qp~~laMerr. & Perry from New Guinea; P. cladaropfylla Briggs et al., P. alpestk Briggs et al., and P. eulyphylla Briggs et al. from E. Australia; P. daltonii Decne., and P. tasrnanica Hook.€. from Tasmania; and P. raoulii Decne. and P. spathulata H0ok.f. from New Zealand. Annuals (Ch 2) are found in two groups (Sp 5761 and 79-80): P. cunningharnii Decne., P. turrifera Briggs et al., P. multiscapa Briggs, and P. debilis R.Br. from Australia, and the last also from Tasmania. Also P. euana Hiirlimann (1967) from the Tonga Islands (Polynesia) belongs here. Only the type is known, a single, young, possibly depauperate plant which may have been introduced recently by man from Australia or Tasmania (cf. P. debilis, but flowers are more crowded). P. drurnrnondii Decne. from Australia and P. pentaspennu Hemsl. from Amsterdam and St. Paul Islands in the Indian Ocean are also annuals, but with patent hairs on the scape. P. stauntoni Reichardt is from the same islands, but perennial (Sp 78). All annual species have hairy sepals (Ch 40 rev.), this character is shared by several perennial species (Sp 6266, and 76-78); P. exi6 Decne. from Western Australia, and PHYLOGENY 01:PIAVI'AGINACEAE I73 
P. gaudichaudii BarnCoud, P. bellidioides Decne., P. varia R.Br., P. glabrata Hook.f., and P. hkpida R.Br. from Tasmania and eastern Australia and the already mentioned P. spathulata from New Zealand, and P. stauntoni from Amsterdam and St. Paul Islands in the Indian Ocean. P. antarctica Decne. from Tasmania and eastern Australia. and P. picta Colenso from New Zealand have glabrous sepals like the European P. schwarenbergiana Schur and the two Asian species P. depressa Willd. and P. camtschatica Link (Sp 49-56). The small Asiatic, perennial species, P. komaroui P. perssonii Pilg. and P. arachnoidea Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 Schrenk have a reduced number of ovules (Ch 77, Sp 49-5 1) like P. sect. Krgznica (Sp 81-108) and the species (Sp 72-76) with adventitious roots. Spikes with crowded flowers (Ch 30) are present in several species within sect. Mesembrynia (Table 2) but not distributed in a meaningful manner and for that reason are excluded from Figure 6. The distribution of sect. Mesembrynia is discussed under subgenus Plantago. Whether the six species in Asia and Europe are the remainder of a former wider and more coherent distribution of this section or the result of a relatively late immigration to or from Australia is uncertain. The species in the marginal areas (Europe, Asia, Amsterdam and St. Paul Islands.) give the impression of being old, and those from Australia mostly of being young, and the section there rapidly speciating and expanding like sect. Virgnica in South America. 
Plantago sect. Virginica BarnCoud This American section is monophyletic. The ovary has three (a few species also have some ovaries with four ovules) or two ovules (Ch 78 and 79) evolved from an five-seeded ovary with one seed in an upper compartment (Ch 76). Semicleistogamic flowers are more common than chasmogamic flowers in most species, they have erect corolla lobes (Ch 58) hiding the dwarfish anthers, but the protogynous stigma is protruding. The lowest chromosome number found in sect. Virginica is 2n = 24, so this section may have originated as an amphidiploid between two species in sect. Mesembrynia. The ancestor(s) probably reached South America before the glaciation of East Antarctic, 14-10 Myr ago according to Mercer (1978). In southern South America the climate was probably suitable for species of sect. Virginica during most of the Tertiary, so the ancestral species probably arrived early from the South, and speciation and dispersal occurred during the Tertiary, while the many subspecies of P. australis were formed later, especially in the Andes during their upheaval and glaciations (Rahn, 1974). A crossing of the Pacific without leaving species on any of the islands or a dispersion from the North is considered less probable. The rather few species in North America are either very advanced (P. uirginica L. in east, south, and central U.S.A. Fig.7, Sp 93), or confined to the warmer parts: P. rhodospema Decne. in southern U.S.A. and northern Mexico (Sp 94), P. subnuda Pilg. (Sp 106) along the west coast, and P. australis Lam. (Sp 108). The last species just reaches Arizona and is common through Central America and the Andes to Argentina and southern Brazil. Two endemics to Mexico are: P. alismat$olia Pilg. (Sp 107) and P. Joccosa Decne. (Sp 88). Four annual species (Ch 2) with two ovules (Ch 79) form a clade (Sp 93-96). P. galapagensk Rahn on the Galapagos Islands is probably the sister species of P. rhodospenna and may be the result of a relatively recent long-distant dispersal; together 174 KNUD RAHN they form the sister-group of P. virgznica L., Their sister species is P.jnna Walp., an endemic to Chile. Their common ancestor might have entered USA from Chile by long distance dispersal like many other species with a similar distribution (Moore, Williams & Yates, 1972). P. jirma has also been found on Juan Fernandez (= P. skottsbergii Pilg.), but was probably introduced recently by man. P.jirma is also found in California, but most probably also introduced there by man. P. tenuipala Rahn (Sp 92) is a rare annual from Colombia, and probably the sister- species of the common P. myosuros Lam. from Bolivia, Argentina, Paraguay, Uruguay Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 and southern Brazil. The only other annual species is P. penantha Griseb. (Sp 99) from Uruguay and eastern Argentina, it has also two ovules like P. virginica etc., but they are probably distantly related only. P. penantha has antrorse hairs on the scape (Ch 17) like the species Sp 97-108: P. pachyneura Steud., P. orbignyana Decne. P. dielsiana Pilg., P. jujuyensis Rahn, P. venturii Pilg., P. buchtienii Pilg., P. truncata Cham. & Schltdl., P. berroi Pilg., P. subnuda Pilg., P. alismat$lia Pilg. and P. australis Lam. P. turJicola Rahn (Sp 90) has adventitious roots like P. australis Lam. (with nine subspecies). The species Sp 85-87: P. catharinea Decne., P. argentina Pilg., and P. ventanensis Pilg. may be related, but do not share any synapomorphy. Plantagojoccosa Decne. (Sp 88) and P. guilleminiana Decne. (Sp 89) both have thin hairs (Ch 16), but this does not probably indicate a closer relationship. Thin hairs are also characteristic for P, Lereddelliana Decne. (Sp 82) and P. commersoniana Barnkoud (Sp 83); they have narrow leaves like P. trinitatis Rahn (Sp 81). The last mentioned species is a curious endemic to the small Ilha da Trinidade in the Atlantic Ocean about 1500 km east of €30de Janeiro. P. trinitatis has a woody stem and is also otherwise bhenetically) very distinct from other sect. Virginica, but the placenta and the arrangement of the ovules leaves little doubt about its belonging to this section. It is probably of great antiquity on the island. The two other species just mentioned may also be primitive P. commersoniana is found in the old mountains in southern Brazil, and P. weddeliana in north-west Argentina where old mountains are close to the younger Andes. The greatest diversity in this section is found within the same two areas. 
Plantago sect. Oliganthos Barnkoud This group is considered monophyletic, but is united by only a single synapomorphy: a short spike with one to three rarely to 12 flowers (Ch 32). This small number of flowers is also present in P. picta Colenso and sometimes in P. spathulata Ho0k.f. (both were placed by Pilger (1937) in sect. Oliganthos), and P. raoulii Decne., all from New Zealand, and in this paper placed in sect. Mesembyzia (Fig. 6, Sp 56, 76, and 68); they usually have more than three flowers, which are densely crowded. The flowers are not crowded in sect.Oliganthos, when more than three are present. A diploid chromosome number (2n = 12) is only known in four species in P. sect. Oliganthos: P. triantha Spreng. from Tasmania and Auckland (Fig. 8 Sp log), P. lanigera Ho0k.f. (Sp 120), P. obconica Sykes from New Zealand, and P. glacialis Briggs et al. from south-east Australia (Sp 129). In 14 other species the numbers 2n = 24, 36,48, 72 and 96 are known. The number of ovules generally varies from one ovary to another except in four species. In P. tehuelcha Speg., P. glacialis Briggs et al., P. palust& Fraser et al., and P. muelLen' Pilg. (Sp 1 12 and 129-1 3 1) the number invariably is four (Ch 82), as in many PHYLOGENY 01: Pl ANTAGlNACEAE 175 species of sect. Mesemb?ynia;but the placenta in a ripe ovary is usually thin and split distally (partly parietal) when belonging to sect. Oligunthos, and not thickened distally and not with keels between the seeds, as in sect. Mesembgmia (Ch 76); so the number four is probably evolved from a variable number, not from a fixed number of five ovules as in sect. Mesmbrynia. All species in this group are perennial herbs. The small dense rosettes often form 
 cushions. They grow in moist, often salty habitats at low temperatures, i.e. at high Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 altitudes in New Guinea, Tasmania, New Zealand, and in the Andes, or in the lowlands in southern South America, New Zealand, and Tasmania. There has apparently been a strong selection pressure for developing a short scape during flowering (Ch 13),but also for a later elongation of the scape (Ch 14) or development of a carpophore (Ch 72) to ensure a better dispersal of the seeds. The common ancestor probably lived on the Antarctic continent in the beginning of the Tertiary. Species spread to Australia before 10 Myr ago, when the East Antarctic was covered by ice (Mercer, 1978). The species now in southern South America, or their ancestors, arrived to South America later than 38 Myr ago, when the climate in southern South America turned colder and became suitable for cold temperate species, but before the glaciation of West Antarctic 3.8 Myr (Mercer, 1978; Rahn, 1980). P. tubulosa Decne. and P. ripda Kunth (Sp 1 19-1 20), now found in the highest parts of the intertropical Andes, have their closest relatives not in southern South America 
1-48 Plantago subgen. Plantago 81-108 P.sect. Virginica 2n=24.48 
49 P komarovi ~ C Asia 50 P perssonii - C Asia 51 P.arachnoidea - C Asia 52 P.camtschatica - E Asia 2n=12 -I1 53 P.schwarzenbergiana - E Europe 2n=12 54 P.depressa - C & E Asia 2n-12 
55 P.antarctica ~ Tasmania. E Australia 2n=12 56 P,picta - N Zealand 2n=48 57 P.cunninghamii - Australia 2n-12 58 P.turrifera - Australia En-12 59 P.multiscapa - C Australia 
60 P debilis ~ Australia. Tasmania 2n=12 b 61 P.eauana - Tonga 181s 3 62 P.exilis - W Australia 63 P.gaudichaudii - Tasmania. E Austral. 2n=12 10 64 P.bellidioides - Tasmania 2n-12 17 65 P varia - Tasmania, E Australia 2n=12 66 P.glabrata - Tasmania 2n=24 67 P.cladarophylla - E Australia 2n=36 68 P.raoulii - N Zealand 2n-48 1 69 P.montedichsonii - N Guinea 70 P,trichophora - N Guinea 71 P.papuana - N Guinea 72 Keuryphylla - E Australia 2n-12 73 P tasmanica - Tasmania 2n=12 74 Kdaltoni - Tasmania 75 P alpestris - E Australia 2n=12 x-x- 76 P.spathulata - N Zealand 2n=48 23 40 77 P.hispida - Tasmania. E Australia 2n-12 78 P.stauntoni - Amsterdam & %.Paul 1. 211-24 79 P.pentasperma - Amsterdam I %.Paul I. 80 P.drumnondii - Australia 2n=12 
Figure 6. Plnnhzgo scct.Mesemb?ynia. On the left, one of 12 parsimonious trees obtained using seven characters and Hennig86 option mhennie; 14 transformations arc present in the tree. Ci: 0.50, ri: 0.88. I76 KNUD RAHN but in New Zealand, Tasmania, and New Guinea. The theories implying a rafting continent may spontaneously appear more tempting than explaining this distribution as the result of an early immigration from the Antarctic continent. P. ser. Oligunthos (Fig. 7, Sp 109-1 16) is paraphyletic. P. ser. Microcu&x (Sp 119-131) and ser. Curpophome (Sp 117-1 18) are united by two synapomorphies: adventitious roots (Ch I)and the small three-celled glandular hairs placed in cavities in the epidermis (Ch 18).The cuticle is usually very thick and probably an adaptation to a windy climate, and the situation of the three-celled hairs acts as a protection Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 against the grinding of snow and sand. 
Plantugo ser. Oligunthos Rahn This section is paraphyletic within P. sect. Oligunthos (Fig. 8, Sp 109-1 16) and 
TABLE3. Plantago subgenera Coronopur, Littorello and Pyllium. All species in this table have the apomorphic state of Ch 89 and the plesiomorph state of the following characters: Ch 6, Ch 7, Ch 8, Ch 9, Ch 10, Ch 11, Ch 14, Ch IS, Ch 18, Ch 19, Ch 23, Ch 24, Ch 25, Ch 26, Ch 27, Ch 28, Ch 30, Ch 32, Ch 33, Ch 36, Ch 37, Ch 38, Ch 42, Ch 43, Ch 44, Ch 45, Ch 49, Ch 50, Ch 53, Ch 54, Ch 55, Ch 56, Ch 57, Ch 58, Ch 59, Ch 63, Ch 64, Ch 65, Ch 66,Ch 70, Ch 71, Ch 72, Ch 76, Ch 77, Ch 78, Ch 79, Ch 80, Ch 81 and Ch 88. 
11112 22233 33444 44556 66666 77788 88888 99 12345 23570 12914 59016 78120 12789 34502 34567 01 subgen. Coronopus sect. Mari t ima 00000 0003+ -0000 00030 03000 30000 30030 03300 00 132 alpina 00000 10033 00000 00133 03000 31000 30030 03300 00 133 subulata 00000 0003+ -0000 00033 03000 33000 30030 03-0 00 134 rbizo,~ylon 00000 10033 00000 00133 03000 33011 30020 03000 00 135 mritima sect. Coronopus 00000 30033 00000 00333 33000 33000 00030 03300 30 136 serraria 00000 3003+ -0000 00333 33000 33300 00030 03300 30 137 crassifolia 03000 3003+ -0000 00133 33000 33033 00030 03300 +- 138 asphodeloides 03000 +003+ -0000 00033 33000 33030 +O+OO 03300 3+ 139 crypsoides 01000 30033 00000 00133 33000 33030 30200 03300 33 140 coronopus 
00000 3003+ -0000 00033 33000 30300 30300 03300 33 141 macrorhiza 00000 30033 00000 00133 33000 33300 30300 03300 ++ 142 subspathulata subgen. Littorella 30000 030-- +-333 00000 00000 30300 03003 30300 -- 143 araucana 30000 030-- +-333 33000 00000 30300 03003 30300 -- 144 americana 30000 030-0 30333 33000 00000 30300 03003 30300 0- 145 uniflora subgen. Psyllium 00330 0000- +-330 00000 00100 00300 00003 03033 0- 146 arborescens 00330 00000 30330 00000 00300 00000 00003 03033 0- 147 webbii 00330 -0-0- +-330 00000 00-30 00030 00003 03-+3 -- 148 famarae 00330 00000 30330 00000 00030 00000 00003 03023 0- 149 senpervirens 00333 -0-0-+-330 00000 00033 OO+-- 00003 03-+3 -- 150 asperrfma 
00333 30000 21330 00000 00010 0030- 00003 03033 0- 151 mauritanica 00330 -0-0-+-330 00000 00--0 OO+-- 00003 03033 -- 152 sinaica 00330 -0-0- +-330 00000 00--0 OO+-- 00003 03-+3 -_ 153 euphratica 03330 30000 30330 00000 00000 00300 00003 03003 0- 154 arenaria 03330 30200 33330 00000 00000 10000 00003 03003 0- 155 afra 
03330 3000- ++SO 00000 00000 33030 00003 03033 -- 156 exigua 03330 +OOO- ++330 00000 00--0 00-+- 00003 03003 -- 157 marismrtui 03330 +030- ++330 00000 00--0 OO+-- 00003 03003 -- 158 phaeostm 03330 +OOO- ++330 00000 OO+-0 00-+- 00003 03303 -- 159 chamaepsyllium 03333 30000 33330 00000 00300 30032 00003 03203 0- 160 sarcopbylla 
03333 30000 33330 00000 00300 00300 00003 03303 0- 161 squarrosa PHYLOGENY 01: PIANlAGINACEAE 177 characterized by having thick tap-roots (symplesiomorphy). It includes seven species from southern South America (studied by Rahn, 1984): P. barbata G. Forst., P. moorei Rahn, P. tehuelcha Speg., P. correae Rahn, P. unkhmis Walp, P. smperviuoides Dusen, and P. puluinata Speg., and further P. triantha Spreng. from Auckland Island and Tasmania. 
Plantago ser. Microcabx (Pilg.) Rahn Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 This group (Fig.8, Sp 119-131) is paraphyletic in relation to ser. Carpophorae and found in New Zealand, Tasmania and in the mountains of eastern Australia and New Guinea. Only four species have more than a solitary flower in the “spike” (Sp 122 and 129-1 3 1): P. novae-relandiae L.B. Moore from New Zealand, P. muellem’ Pdg. from Tasmania and the mountains in East Australia, where also P. palustris Fraser et al. and 
TAHI.E4. Plantago subgen. Bougueria and subgrn, Alhirans. All species in this table i.r. the subgenrra Boup’aand Albicanr have the apomorphic state of the following characters: Ch 16, Ch 24, Ch 82, and Ch 84. They all have the plesiomorph state of: C:h 4, Ch 6, Ch 7, Ch 8, Ch 9, Ch 10. Ch 11. Ch 13, Ch 14, Ch 18, Ch 23, Ch 32, Ch 33, Ch 34, Ch 35, Ch 38, Ch 39, Ch 47, Ch 48, Ch 51, Ch 52, Ch 64, Ch 72, Ch 73, Ch 74, Ch 75, Ch 76, Ch 77, Ch 78, Ch 79, Ch 80, Ch 81, and Ch 83. 
1 11122 22222 23333 44444 44455 55555 56666 66666 77888 889 12352 57901 25678 90167 01234 56903 45678 90123 56789 01567 890 subgen. Eougueria 00000 00000 00000 03030 00000 00000 00000 00333 00033 0030- -3- 162 nubicola subgen. Albicans sect. Montana 03003 03300 03030 00030 30030 30000 00000 00333 00033 33003 030 163 loeflingii 03003 20----+-+O 00130 30030 30000 00000 00330 00033 3+033 03- 164 cafra 03003 03000 03030 03130 10030 30000 00030 00300 ---++ +3203 03- 165 notata 00000 03000 03030 03130 30030 30000 00000 00000 00200 00033 030 166 atrata 00000 30000 03030 03230 30030 30000 00000 00000 00200 00023 03- 167 monosperma 
00000 30000 03030 03230 30030 30000 00000 00300 00000 00103 030 168 nivalis sect. Lance1 fo 1 ia 00010 03030 03030 03103 03003 03000 00000 00000 00200 03303 030 169 lagopus 00020 03030 03030 03003 33003 03000 00000 00200 00200 03013 030 170 lanceolata 00330 03-+--+-+- 03003 33003 03000 00000 00000 00300 0+033 030 171 leiopeta7a 0003- -+-+- -+-+- 03003 +3003 03000 00000 00000 -400 0+033 -3- 172 lacustris 30030 03-+--+-+- 03003 33003 03000 00000 00000 03300 03033 030 173 altissm 
30000 03030 03030 03103 33003 03000 00000 00000 03200 03023 030 174 argentea sect. Eauphula 03310 00003 03000 03130 00000 00000 00000 02000 00033 00033 033 175 anplexicaulis sect. Mw~nopsylI ium 03003 30000 03033 03300 03000 00000 00000 00000 30000 0-303 033 176 cretica 0300+ +---33+0++ 03130 03000 00000 00300 00000 3-010 0-003 -3+ 177 cyrenaica 03003 33003 33033 00100 03000 00000 00300 00300 30030 00303 033 178 bellardi sect. Albicans ser. Ovatae 02000 00000 03030 03130 00000 00000 00030 00000 00030 03003 333 179 ovata ser. Minutae 03000 00300 03033 03100 03000 00300 00300 00330 00000 00023 333 180 minuta 03000 00300 03033 03100 03000 00300 00300 00300 00030 00003 333 181 lacbnantha 03000 30003 03030 00100 00000 00300 00300 00300 00300 00033 333 182 stocksii ser. Albicans 00200 30--+-3030 00030 00300 00300 00000 00000 00200 03003 33+ 183 cylindrica 00200 30003 03030 00030 00300 00300 00010 01000 00300 03013 333 184 albicans 03200 30--+-+-+- 00030 00300 00300 00010 00100 00200 0+003 333 185 boissieri ser. Ciliatae 03203 30--+-+-+- 00100 00100 00330 00300 00000 03033 0+003 33+ 186 psamphila 00233 3000+ 03030 00100 00100 00330 00300 03000 00-+--+033 33+ 187 tunetana 03200 00--+ -+-+- 00300 00100 00330 00300 00000 00030 0+033 333 188 akkensis 03203 lo--+ -+-+- 00100 00100 00330 00300 00331 00033 0+003 333 189 lagocephala 03033 10003 03030 00100 00100 00330 00300 00110 00032 02003 333 190 ciliata 178 KNUD RAHN 
TABLE4 continued 
1 11122 22222 23333 44444 44455 55555 56666 66666 77888 889 12352 57901 25678 90167 01234 56903 45678 90123 56789 01567 8% sect. Gnaphaloides ser. Hispidulae 03003 03--- -++-- 30100 03000 00000 00102 00111 00032 1-013 033 191 limensis 03003 03----++-- 30100 03000 00000 00002 00311 00022 1-013 03+ 192 litorea 03003 03--- -++-- 30100 03000 00000 00102 00311 00122 1-013 033 193 rancagua 03003 03000 03300 30100 03000 00000 00303 00200 00033 1-003 033 194 lundborgfi 03003 03000 03300 30100 03000 00000 00202 00111 00122 10033 033 195 hispidula Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 ser. Sericeae 00000 03--- -++-- 30100 03000 00000 00000 00100 00010 0-003 033 196 tolucensis 00003 03--- -++-- 30100 03000 00000 00030 01100 00211 00103 033 197 nivea 00000 03000 03300 30100 03000 00000 00030 01100 00211 00113 033 198 linearis 00300 -3--- -++-- 30000 03000 00000 30000 00000 OO+-- --033 033 199 johnstoni 00200 13--- -++-- 30100 03000 00000 10110 02100 00211 00013 033 200 sericea 
00300 03000 03300 30300 03000 00000 00300 03000 00300 0-023 033 201 bismrckii 00300 03--- -++-- 30200 03000 00000 00000 33000 00300 0-033 033 202 grandiflora 00000 33--- -++-- 30300 03000 00000 00300 00000 00010 0-103 033 203 lanprophylla ser . Bras? liense 00200 33--- -33-- 30100 03000 00003 30030 02000 00210 0-023 033 204 brasi Jiensis 00300 33--- -++-- 30100 03000 00003 30030 23000 00300 0-033 033 205 tandilensis 00000 33000 03300 30100 03000 00003 30030 00100 00031 0-003 033 206 densa ser. Gnaphaloides 03003 33- -++-- 30100 03000 00003 00030 00100 00211 10003 033 207 erecta 03000 33--- -++-- 30100 03000 00003 00030 01000 00211 10033 033 208 helleri 03000 33----++-- 30100 03000 00003 03030 01000 00211 10023 033 209 wrightiana 03000 33000 03300 30100 03000 00003 03030 00200 00211 10003 033 210 ari5tdtd 03003 33000 03300 30100 03000 00003 03030 00200 00211 10003 033 211 patagmica 
03003 33000 03300 30100 03000 00003 03030 00100 00211 10023 033 212 hookeriana 03000 33--- -*--30100 03000 00003 03030 00100 00211 1-003 033 213 argyred 
P. glacialis Briggs et al. are indigenous. Four species from the high mountains of New Guinea (Sp 125-128): P. aundensis P. Royen, P. shophylla Merr. & Perry, P. politu Craven, and P. depauperata Merr. & Perry, have two bracts like P. lanigera Ho0k.f. from New Zealand, P. paradoxa Ho0k.f. from Tasmania, and the South American species with solitary flowers which belong to ser. Oliganthos. The presence of only one bract (Ch 33) is a potential synapomorphy for uniting the South American P. ser. Carpophorae with P. unibrackatu Rahn (a nom. nav. for P. ungora Hook.f., non L., see Appendix) and P. trzandra Berggr. from New Zealand, and the Tasmanian P. gunnii H0ok.f. (Sp 1 17-1 19 and 123-1 24). The nomenclatorial type of this series. P. triandra Berggr. is very specialized and was placed by Pilger in a monotypic section. The bracts and sepals are very small, and the number of anthers and corolla lobes often three only; the scape is short or absent during anthesis (Ch 13) but elongates later (Ch 14), like the scapes of most other species within Microcalyx; this has been interpreted as a carpophore, e.g. Pilger (1937). Plantago obconica Sykes (specimens not seen) according to the protologue has a short carpophore-like, “funnelform” elongation of the capsule, adventitious roots and a solitary flower, but the number of bracts and sepals is not given. The chromosome number is 2n = 12. It is compared with P. trzandra, but not with P. unibracteata. 
Plantago sect. Carpophorae (Rahn) Rahn Two closely related and highly specialized South American species P. rgda Kunth and P. tubulosa Decne.(Fig. 8, Sp 1 17-1 18) have carpophores developed after anthesis PHYLOGENY OF PLIZN'IAGINACEAE I79 
(Ch 72), and flowers are usually unisexual and dioecious; bracts and sepals are very large and membraneous like the corolla. See further under sect. Oliganthos and Rahn (1985). Seeds of P. ngdu from Ecuador (B. Blgaard 90596) were sown 1991 in Copenhagen, the plants were cultivated three years without flowering. They had 2n = 72 chromosomes. Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021
Plantugo subgen. Coronopus (Lam. & DC.) Rahn 
The most important synapomorphy uniting the species in this subgenus is Ch 48 (corolla tube covered by short hairs); all other species of Plantaginaceae have a glabrous corolla tube except for P. luchnantha (Sp 181) which has a lanate tube. Small cone-shaped glandular hairs (Ch 20, Fig. 1) are present in all, and a sharp keel or wing on the posterior sepals (Ch 46) is present in all but P. u&ina; however, both characters are also found in P. subgen. Lancefoliu. The species are distributed around the Mediterranean, but P. muritirnu is also found in other parts of Europe, in central Asia, in North America and southern South America (Moore et ul., 1972). 
Kq to the sections 
49-80 P.sect. Mesembrynia - -81 P.trinitatis - llha Trindade 7 82 P.weddelliana - Bolivia. NV Argentina 2~24 .cmersoniana - Brazil 16 40 E i)-D 84 P.tmentosa - S America 2n=24 5 40 -4 ~ 85 P.catharinea - E Brazil En-24 
~ 86 P.argentina - NU Argentina 2n=24.48 
~ 87 P.ventanensis - E Argentina 2n=24 88 P.floccosa - Mexico 2n-24 89 P.guilleminiana - E Brazil +€I16 c 90 P.turficola - E Brazil 3 -#+ 1 79 91 P.myosuros - 5 America 2n=24 92 P.tenuipala - Colombia 93 P.virginica - E USA 2n-24 7 94 P.rhodosperma - S USA. N Mexico 2n-48 95 P.galapagensis - Galapagos Isls 
96 P firma ~ Chile 2n=24 97 P.pachyneura - Chile 2n=24 98 P.orbignyana - Andes 211-24.48 99 P,penantha - Uruguay, NE Argentina 2n=24 
100 P.dielsiana - E Argentina, Uruguay 2n=24 101 P.jujuyensis - NW Argentina 2n-24 c 5 17 +102 P.venturii - NW Argentina 2n=24 23 -103 P.buchtienii - NW Argentina. Bolivia 2n=48 -104 P,truncata - Chile -X-40 -105 P berroi - Uruguay. E Argentina 2n=24 
~ 106 P.subnuda - W USA 211-48 +107 P.alismatifolia - Mexico 2n-24 79 +108 P.australis - America 2n=24.48 I 
Figure 7. Plantago sect. Krgiinica. On the left one of the ten most parsimonious trees obtained using Hennig86, option mhennigl'; 13 characters. Length: 23, ci:0.56, ti: 0.77 (the same as obtained when using ie-1. 180 KNUD RAHN 
1-48 P.sect. Plantago 7 2n-12 
0- 5 + 31 Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021
117 P.tubulosa - Andes 2n-24.48 118 P.rigida - Andes 2n-72 H- 13 32 119 P.uoibracteata - N Zealand 2n-60 120 P.lanigera - N Zealand 2n=12 121 P.paradoxa - Tasmania 211-24 -++ -x- 122 P.novae-zelandiae - N Zealand 2n-24 1 18 32 123 P.gunnii - Tasmania 2n-36 +I 124 P.triandra - N Zealand 211-48 125 P.stenophylla - N Guinea %+ 126 P.polita - N Guinea 5 127 Kdepauperata - N Guinea 128 P.aundensis - N Guinea r~-129 P.glacialis - Australia 2n=12 
- Australia 2n=24 131 P.muelleri - Australia. Tasmania 2n=36 Figure 8. Plantago sect. Oliganthos. On the left one often parsimonious trees obtained by using Hennig86. option mhenni?; 13 characters. The ten trees have the length 40, ci: 54, ri: 81. P 1. Calyx tightly pressed against the rachis, posterior sepals with a broad wing on the nerve. Plant often annual and leaves lobed ...... sect. Coronopus 1'. Calyx not pressed against the rachis, posterior sepals with a keel or wing less prominent or absent. Plant perennial and leaves with scattered teeth ...... sect. Maritima 
Plantago sect. Maritima H.Dietr This section (Fig. 9, Sp 132-1 35) is apparently a paraphyletic group of four species characterized by lacking the synapomorphies of sect. Coronopus only. Plantago subulata L. here includes what is often considered a group of closely related species (P. holosteum Scop., P. radicata Hoffmans. & Link, P. insularis (Godr.) Nyman, and P. penyalarensis Pau); it has narrow needle-like leaves, while P. maritima L. (incl. P. serpentina All., P. olkanthos Roem. & Schult., and P. atlantica Batt.), P. alpina L., and P. rhizoxylon Emb. have broader linear leaves. Material of Plantago maritima in North and South America is not significantly different from European material. The species probably spread from Europe after the glaciations, but not necessarily by man. 
Plantago sect. Coronopus This section is monophyletic (Fig. 9, Sp 136-142), having a broad wing on the back of the posterior sepals (Ch 47). The reduced chromosome number x = 5 (Ch 90) is another synapomorphy, but it is also present within P. subgen. Albicans. The seven species in P. sect. Coronopus may be placed into two groups, the first with 1-4. ovules (Ch SO), ovary glabrous (reversal of Ch 73): P. asphodelodes Svent., P. crass$lia Forssk., and P. serraria L. (incl. P. peloritana Lojac.). The other group (Sp 139-1 42) comprises P. coronopus L. (incl. P. aschersonii Bolle), P. clypsoides Boiss., P. macrorhiza Poir., and P. subspathulata Pilg.; it is united by two synapomorphies: an upper compartment present in the ovary on the abaxial side (Ch 75, Fig. 3, Sp 140), PHYLOGENY OF PLANTAGINACEAE 181 
1-131 P.subgen. Plantago 143-213 P.subgen. Litt.t Psyl + Boug.+ Albicans 132 P.alpina - Europe 2n=12.24 133 P.subulata - Mediterranean 2n=12.24 134 P.rhizoxylon - NW Africa -'= 135 P.maritima - Cosmopolite 2n=12.24 136 P.serraria - Mediterranean 2n-10.20 D 137 P.crassifolia - Medit S Africa 2n-20 62 73 138 P.asphodeloides - Canary Isls 139 P.crypsoides - NE Africa 12 47 90 Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 140 P.coronopus - Medit. Europe 2n=10.20.30 141 P.macrorhiza - Mediterranean 2n=10 
142 P.subspathulata - Madeira Figure 9. Pluntugo subgen.Coronopus.The only most parsimonious tree obtained using Hennig86, option ie- : 19 characters. Length of the tree: 27, ci: 0.70, ri: 0.84. and absence of ribose in the seeds (Ch 91). Gorenflot & Bourdu (1962) did not find ribose in seeds from P. coronopus and P. macrorhiza, but in seeds from P. serrariu and P. crassy51ia and in three species from sect. Maritima (P. maritima, P. ulpina, P. subulata); however, they wrongly stated that P. serrariu and P. crass@lia have the chromosome number x = 6. The upper seed occurs in a compartment formed by the placenta on the anterior (abaxial) side (Ch 75) and dispersed together with the upper part of the pyxis (Dowling, 1933). A similar arrangement is found in Plantago sect. Mes~b~nia when five seeds are present, but the upper seed occurs on the posterior (adaxial) (Ch 76), not the anterior side of the septum. 
Plantago subgen. Littorella (P.J. Bergius) Rahn 
This subgenus (Fig. 10, Sp 143-145) is united by several synapomorphies. The flowers are unisexual, monoecious (Ch 34. Ovary with only one compartment fertile 
€F 143 P.(Littorella) uniflora - Europe 211-24 84 144 P.(Littorella) americana - North America 2n=24 
 h-w+ 21 29 31 
154 P.(Psyllium) arenaria - Mediterranean 2n=12 155 P.(Psyllium) afra - Mediterranean 2n=12 I,I 156 P.(Psyllium) exigua - E Mediterranean 2 12 86 -+ 157 P.(Psyllium) maris-mortui - E Mediterranean 22 158 P.(Psyllium) phaeostma - N Africa 2n-12 159 P.(Psyllium) chamaepsyllium - E Mediterranean 
Figure 10. Plantago subgen. Ldforella and subgen. Pgllium. On the left one of the five most parsimonious trecs obtained using Hennig86 and the option mhennig*; they have the same length as that obtained using the option ie-, 3 1 transformations, ci: 0.74, ri: 0.88. Twenty-three characters were used. 182 KNUD RAHN (Ch 74)) with only one ovule (Ch 83) which is anatropous, straight, sitting on a basal placenta (Fig. 3, Sp 144, Edkardt, 1937; Rosen, 1940; Dietrich, 1971). The anatropous ovule is not considered plesiomorphous in relation to hemitropous (Ch 84) as found elsewhere in Plantaginaceae, but is a transformation of that. The plants are according to Pilger (1937) glabrous all over, except for var. pubescens Decne. (no specimens cited). I have searched for hairs on many specimens. Small three-celled glandular hairs are common (Moncontit, 1969; Rahn, 1992), and many specimens are apparently pubescent, but the “hairs” are epiphytic algae, except in two cases Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 where short hairs were seen (Rahn, 1992). The hairs are always obtuse at apex, sometimes with protuberances, and all cells usually contain some plasma; they are, therefore, doubtfully classified as glandular hairs, but their size and shape are often more like the common type of non-glandular hairs. This kind of glandular hair is also found in P. subgen. Psyllium and in subgen. Albicans sect. Hymenopsyllium and sect. Albicans. In the leaf axils are web-like, non-glandular hairs of the same type as found at this location in the subgenera Pgllium, Caronopus and Plantago. The inflorescence is according to Payer (1857) and Skottsberg (191 1)) a spike on a short scape (Ch 13);below there are few female flowers (Ch 31) which are usually opposite or in whorls of three (Ch 29. as in subgen. Psyllium and P. sect. Gnaphaloides (but according to Eckardt, 1937 only apparently opposite); this is considered apomorphic (see under Ch 29). The male flower has a pedicel. A line is visible on the wall of the fruit facing the bract in P. americana (Fig. 3, Sp 144 b, e) and also on the adaxial side in P. ungora, so it was possible, in a cross section of the hard nut, to see that the cotyledons are perpendicular to the bract as in seeds of subgen. Coronapus, subgen. Psyllium, and subgen. Albicans. The ripe seed is without or with scanty endosperm (copious in other subgenera). P. subgen. Littorella grows at the edge of or in oligotrophic lakes. Other species of Plantago which are adapted to wet (but often salty) habitats are found in Plantago sect. Oliganthos, in which many species are also small, with adventitious roots and with reduced spikes and floral parts, but other characters do not indicate a closer relationship. The subgenus LitLorella includes three closely related species, the rare P. araucana Rahn in southern South America, P. americana (Fernald) Rahn in North America and P. ungora L. in Europe, including Iceland and the Azores. The last two species are closely related, united by two synapomorphies (Ch 35 female flowers in one whorl, and Ch 39 pedicel of the male flower longer than 4mm). The distribution of the three species is probably the result of long distance dispersal. P. subgen. Littorella has never been recorded from the tropical zone neither in lowlands nor mountains, and has probably never inhabited lakes in that zone. However, the diaspore of subgen. Littorella and subgen. Bougueria is a small nut probably adapted for endozoic, not for epizoic dispersal like most other species of Plantago. P. araucana is the most primitive species. It is more likely that this subgenus, according to Hennig’s progression rule, spread from South America via North America to Europe than the other way, but this rule is disputable. 
Plantago subgen. Psyllium Juss. 
The 16 species united in this subgenus (Fig. 10, Sp 146-1 6 1) have leaves opposite or in whorls of three (Ch 4), and stems with elongated internodes (Ch 3). The non- PHYLOGENY OF PIANTAGINACME 183 glandular hairs are simple, uniseriate like those in subgen. Plantago, and subgen. Coronapus (Rahn, 1992; Andrzejewska-Golec & Swietoslawski, 1993). Hairs on the scape are patent, not antrorse. Two seeds (Ch 82) with deeply concave placenta sides (Ch 87, Fig. 3, Sp 15 1) are also found in P. subgen. Albicans. Plantago subgen. Littorella is as a hypothesis considered the sister group (see above in ‘Synapomorphies uniting the subgenera’). All the species from this subgenus, which were studied by 
Andrzejewska-Golec (1992b), contained the iridoids aucubin and bartsioside. Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 Bartsioside was absent from all other Plantaginaceae studied. Bartsioside (and genioposidic acid) is according to Jensen (pers. comm.) produced during the synthesis of aucubin and for that reason less suitable as a taxonomic character when aucubin is present. Andrzejewska-Golec (1992b) has, as the result of the study of hairs (in eight species) and iridoids (in six species), proposed four new series: ser. Semperoirens Andrzejewska-Golec (Sp 149), ser. Arborescens Andrzejewska-Golec (Sp 146- 148, 151-153), ser. Arenaria Andrzejewska-Golec (Sp 154, 155, 156, 158), and ser. Squarrosa Andrzejewska-Golec (Sp 161). The latin text from the key in Pilger (1937) is used for the protologues. Plantarenaloside was found by Andrzejewska-Golec & Swiatek (1984) in P. sempervirens from the ser. Sempervirens and in P. arenaria, P. phaeostoma and P. aja from ser. Arenam’ae, but was not found in P. arborescens and P. webbii from her ser. Arborescms. The plesiomorphic, small, three-celled hair were absent in the studied material of the two series of annuals. Club shaped glandular hairs (part of Ch 21) were said to be present in all except for ser. Arborescens (P. famara, P. webbii and P. arborescm); nevertheless it was figured in Figure 7h in Andrzejewska-Golec & Swietoslawski (1992b), but indicated as absent in Table 1 in the same paper. Large glandular hairs (Ch 22) were indicated to be absent from P. squarrosa only (but present in Figure 1 in Andrzejewska-Golec & Swietoslawski (1993) although absent according to Table 1 of that paper). Plantago subgen. Pyllium is indigenous to the Mediterranean area and Macar- onesia; it grows in dry habitats. The species of subgen. Pyllium have, since Linnaeus, been continually confused, and are still in need of taxonomic revision. Of the eight perennial species (Fig. 10, Sp 146-153) five are found on the continental main land in three distant areas: (1) In eastern Asia Minor: P. euphratica BarnCoud (2) In Sinai: P. sinaica BarnCoud. (3) In West Mediterranean three species; P. mauritanica Boiss. & Reut. in Algeria and Morocco. P. arperrima Hervier in southern and eastern Spain, and P. sempmirms Crantz ( = P. pops L. (1 762), nom. illeg.; = P. sufiticosa Lam.) in Spain, southern France, and Italy. On Madeira and the Canary Islands except Fuerteventura (Hansen & Sunding, 1985) three perennial species of subgen. Psyllium are found. P. famarae Svent, with broad leaves, is endemic to Lanzarote. P. webbii BarnCoud from La Palma, Tenerife, and Gran Canaria (Sunding & Kunkel, 1971) has narrower, appressed leaves, and smaller bracts and sepals that P. arborescens Poir. from Madeira and the Canary Islands (except Lanzarote). P. arborescm has been subdivided into subspecies and varieties (two of them also described as species: P. costae Menezes, P. madere& Decne.). Pilger (1937) recognized five annual species belonging here, Zohary (1938) three more. They can probably be grouped around three species: 1. (Sp 154) Plantago arenaria Waldst. & Kit. (1802) (= P. indica L. (1759), nom. illegit. = Plantago psyllium L. (1753), nom. ambig.; cf. Panigrahi (1975). = P. scabra 184 KNUD RAHN Moench, nom. illeg.) is distributed in southern Europe, west and central Asia and Egypt. 2. (Sp 155-158) Plantago aja L. (1762) (= P. ynops L. (1753) nom. ambig. non L. (1 762) = P. figdlium L. (1 762), non L. (1 753) = P. squalida Salisb.) is found in southern Europe, western Asia to India, East and North Africa and Makaronesia. It is related to Plantago exigua Murray which has narrower leaves and is found from Egypt to India. The small endemic from the Dead Sea area, Plantago maris-mortui Eig in Zohary (1938), may also belong here. Plantago phaeostoma Boiss. & Heldr. is a dubious species Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 originally described from near Alexandria. 3. (Sp 159-161) Plantago squarrosa Murray in Pilger (1937) included Plantago sarcophylla Boiss. ex Decne. (prosyn. = P. sarcophylla Zohary (1938));however, Zohary (1938) and Feinbrun-Dothan (1978) considered this a common species distinct from the less common P. squarrosa by having only 3-6 flowers in the spike, not 10-20. The even smaller endemic in the western Negev - P. chammp$ium Zohary - may also belong here; it is distinguished by having minute seeds (0.8-1.4mm), while P. sarcophylla and P. squarrosa have seeds which are longer, but still less than 2 m (Ch 84,i.e. smaller than in other species of subgen. Pyllium. The chromosome number 2n = 12 has been found in seven species; Waisel(l96 1) indicates 2n = 10 for P. sarcophylla and P. squarrosa, but it is probably based on misidentification, later 2n = 12 was counted in P. sguarrosa (Rahn, 1976). 
Plantago subgen. Bougum‘a (Decne.) Rahn 
This monotypic subgenus from the Andes has flowers very different from those of all other Plantago, and it is for that reason difficult to find recognizable synapomorphies uniting it with other subgenera (Figs 3 and 11, Sp 162). This is discussed above in ‘Synapomorphies uniting the subgenera’. The small, hard nut with one kidney-shaped seed with a curved embryo constitute the most conspicuous features. The plane between the cotyledons in the seed is perpendicular to the bract as in other Plantago except P. subgen. Plantago; however, the plane between the cotyledons is actually parallel to the placental side of the seed 
P.subgen. Psyllium + Littorella - 2n-12.24 162 P.subgen Bougueria - 2n-12 175-213 other P.subgen. Albicans 2n-10.20.30 163 P.loeflingii - Hediterranean 2n-24 164 P.cafra - S Africa 
165 P.notata - N Africa, W Asia 2n-12 166 P.atrata - Europe. W Asia 2n-12.24.36 167 P.monosperma - Pyrenees 2n=12 17 30 mF86 168 P.nivalis - S Spain 2n-12 ~~ x 169 - 2n-12 P.lagopus Mediterranean 170 P.lanceolata - Cosropolite 2n-12 20 37 41 44 46 71 171 P.leiopetala - Madeira 2n=12 172 P.lacurtris )Morocco 66 x- 174173 P.argenteaP.altiss1ma - - 5SE Europe Europe 2n=l2 2n=72 -z 5 Figure 1 1. Plantago subgen. BougUeria and P. subgen. Albumsect. Montana (Sp 163-168) and sect. Lanceifolia (Sp 169-174). On the left one of six most parsimonious trees produced by Hennig86, option ie-, length 39, ci: 0.64, ri: 0.82; 25 characters were used. PHYLOGENY OF PLANTAGINACEAE 185 and perpendicular to the symmetrical plane of the seed (like subgen. Plantago). The ovary and seeds are also in almost all other respect very much transformed (see above under Ch 74). P. subgen. Bougueria has 2n = 12 and is the only indigenous diploid Plantaginaceae known from South America (Rahn, 1976). Plantago nubicola (Decne.) Rahn is found in puna vegetation at high altitudes in southern Peru, Bolivia and north-west Argentina. Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021
Plantago subgen. Albicans Rahn 
This subgenus (Figs. 11-13, Sp 163-213) is united by one synapomorphy: Hairs with much refracting walls, making the narrow lumen invisible, and with swollen joints (Ch 25, Fig. 1). P. subgen. Albicans has thin hairs (Ch 16) with a short basal cell (Ch Z+ like the sister-group P. subgen. Bougueria. The number of ovules and seeds is two (Ch 82). The placental side of the seeds is hollow as in subgen. Psyllium (Fig. 3, Sp 175-201). Two other characters might also have been present in the common ancestor: upper end of cells of non glandular hairs inserted between flanges from the cell above (Ch 27, Fig. 11, and pollen with a solid operculum (Ch 71, Fig. 2). The root in subgen. Albicans is usually like that in subgen. Psyllium i.e. either a thin annual root or a rather thin, stiff and woody root when perennial, not fleshy as is common in subgen. Plantago. Adventitious roots which are common in subgen. Plantago and subgen. Littorella are found only in two species in subgen. Albicans (Sp 173 and 174). The iridoid catalpol is, according to Andrzejewska-Golec & Swiatek (1984), present in all P. sect. Montana and P. sect. Lanceifolia studied and in P. amphxicaulis, but absent elsewhere in Plantaginaceae (but the only species from subgen. Albicans studied was P. bellardz). Unfortunately the 54 plants studied were not identified. The names in the paper were those under which the seeds were received. Catalpol is according to Jensen @en. comm.) also found in Plantago (subgen. Littorella) ungora L., and it is common in Bignoniaceae, Buddlejaceae, Globulariaceae, Lentibulariaceae, Martyniaceae and Scrophulariaceae. Plantago subgen. Albicans is adapted to dry habitats and is found in Asia, Europe, Africa, North and South America (absent from New Guinea, Australia and New Zealand). Plantago subgen Albicans comprises the following sections in Pilger (1937): Plantago sect. Leucopsyllium (excluding Plantago commersoniana and P. weddelliana), sect. Oreades ( = sect. Montana), sect. Arnoglossum (= sect. Lanceqolia), sect. Bauphula and sect. Hymenopsyllium. 51 species are here placed in six sections, and eight series. 
Ky to sections 1. Anterior sepals united for more than half their length. Upper part of bract scarious and acuminate. Scape sulcate with hairs antrorse. Posterior sepals with a wing on the nerve ...... sect. Lancaifolia 1'. Anterior sepals united at the very base only. The nerve reaching the apex of the bracts. Scape not sulcate. Posterior sepals without wings. 2. Green nerve of anterior sepals present at base only, the rest scarious. Posterior sepals similar to anterior. Bract very wide, completely covering sepals ...... sect. Montana 186 KNUD RAHN 2'. Green nerve of anterior sepals reaching apex, or almost so. Posterior an.d anterior sepals dissimilar. 3. Posterior sepals with a green nerve in the lower half only, the rest scarious. Posterior sepals shorter, narrower and more flat than the anterior sepals. Green nerve on the anterior sepals with a sharp keel or wing. Flowers in a dense spike ...... sect. Ba~p~~~ 3'. Green nerve of all sepals reaching apex. Posterior sepals of equal length but wider than anterior sepals. Anterior sepals without a sharp keel. Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 4. Connective of the anthers very wide and as long as a pollen sac. Long glandular hairs often present besides both short and longer hairs. Flowers spirally arranged in a dense spike ...... sect. Hymenopsyllium 4'. Connective of the anthers smaller. Long glandular hair absent. 5. Hairs on scape patent or woolly, not or insignificantly wider than those on the leaves. Flowers spirally arranged, crowded or scattered in a spike. Corolla lobes usually acuminate or apiculate, never cordate at base. The concave side of the seeds partly covered by a ragged white membrane, except for two areas to the left and right of the centre (Fig. 3, Sp 179 b). Mediterranean, Central Asia ...... sect. Albicans 5'. Hairs on scape antrorse, usually distinctly wider than those on the leaves. Flowers opposite or in whorls of three, alternating in an open or dense cylindrical spike; a single species with 1-9 flowers in a head. Corolla lobes rarely acuminate or apiculate, usually acute or obtuse, sometimes cordate at base. The concave side of seed with an impressed mark just above and one below the centre (Fig. 3, Sp 201b). North and South America ...... sect. Gnaphalvides 
Plantago sect. Montana Barntoud This section (Fig. 11, Sp 163-168) is united by having scarious sepals (Ch 43,45). Among the six species three annuals (Ch 2) form a group, their corolla lobes and anthers are small (Ch 62, 68, 69, 70), they were placed by Pilger (1937) in his sect. hcopsyllium. P. notatu Lag. and P. lo$ingi L. are both distributed from Spain through North Africa and west Asia to Caucasus. P. caja Decne. is from South Africa. P. mvnospma Pourr. and P. nivalis Boiss, have a narrow distribution in the Pyrenees and southern Spain. P. atrata Hoppe is widely distributed in the mountains from Spain and Central Europe to Iran. 
Plantago sect. Lunceifolia Barntoud This section (Fig. 11, Sp 169-174) is very distinct. It has the following synapomorphies: bracts with a scarious acumen (Ch 37),anterior sepals united (Ch 44), posterior sepal with a sharp keel or narrow wing (Ch 46). The following apomorphic characters are also present in all studied members of this section: small acute glandular hairs (Ch 20, Fig. l), upper end of cells of non glandular hairs inserted between flanges from the cell above (Ch 27, Fig. l), and pollen with a solid operculum (Ch 72, Fig. 2). The southern European P. argenka Chaix and P. alhsima L. from the Balkans are the only species of subgen. Albicanr to have white anthers (Ch 6s) and adventitious roots (Ch 2). The other species are the Mediterranean P. lagopus L., P. lacustrk Maire from Morocco, P. lezopetala Lowe (incl. P. malatv-belizii Lawalrte) from Madeira, and P. lanceolata L., now a weed found throughout the world especially in pastures. PHYLOGENY OF PLANTAGINACEAE 187 
 ancestor I 191-213 P.sect. Gnaphaloides 2n=20 - 175 P.amplexicaulis - Mediterranean 2n=10 - 176 P.cretica - E Mediterranean 2n=10 177 P.cyrenaica - NE Africa 
178 P.bellardi - Mediterranean 2n=10 179 P.ovata - Mediterranean. W USA 2n=8 3- 180 P.minuta - C Asia 2n=10 181 P.lachnantha - C Asia 2n=10 t 19 28 41 1 88 ,I - Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 ,I 182 P.stocksii - W Asia 49 56 67 183 P.cylindrica - E Mediterranean 184 P.albicans - Mediterranean 2n=20.30 15 21 185 P.boissieri - W Asia 186 P.psamphila - W Asia 3 71 -X- 187 P.tunetana - NW Africa x- 188 P.akkensis - NW Africa 2n=20 
189 P.lagocephala - W Asia 190 P.ciliata - Mediterranean 2n=10 3 Figure 12. Plantago subgen. Albicunr sect. Albiruns (Sp 179-190), wet. Buuphuh (Sp 175) and wct. Hymenopvllium (Sp 176-1 78). On the left one of 34 most parsimonious trees obtained using Hennig86 and the option ie-; 39 transformations are needed. ri: 0.48, vi: 0.76; 19 characters were used. 
Plantago sect. Bauphula Decne. P. amplexicaulis Cav. (Sp 175, Figs 3 and 12) is the only species in this section. It is distributed mainly in northern Africa and west Asia. The hairs on the scape (Rahn, 1992) are like those in P. sect. Gnaphaloides (Ch 25, Fig. l), but the hairs elsewhere are almost like those in subgen. Psyllium, subgen. Plantago, and subgen. Coronopus (ie. plesiomorphic). Many autopomorphies are present: plants are annuals (Ch 2) which have elongated internodes (Ch 3) with embracing leaves, the posterior sepals are scarious and small, while the anterior are large and with a wing on the back; in other Plantagmaceae, except Plantago schwarzenbergiana (Sp 53), sepals are either equal or the posterior are larger than the anterior. 
Plantago sect. Hymenopsyllium Big. The very large flap-like appendix on the connective of the anther (Ch 6.5)is a synapomorphy uniting three annual (Ch 2) species (Fig. 12, Sp 176-178): The east Mediterranean P. cretica L. and P. cyrenaica Durand & Barratte from Libya have reflexed scapes after flowering, while scapes are erect in P. bellardi All., which is found throughout the Mediterranean area. 
Plantago sect. Albicans Barntoud This Mediterranean section of 12 species (Fig. 1 1, Sp 179-1 90) is united by one synapomorphy: placenta side of the seed covered by a ragged membrane except for two areas on both sides of the centre (Ch 88 Fig. 3, Sp 179 b). Distributed in central and western Asia, North Africa and southern Europe. Includes the following four series: 
Kq to series 1. Bract broadly elliptic, often covering the calyx laterally. Corolla lobes with margins not or slightly involute, not touching one another laterally. Corolla tube rarely exposed above the mature fruit, covered by the reflexed lobes. 2. Bract obtuse at apex, with green nerve widest near apex. Anterior sepals very 188 KNUD RAHN asymmetric and with hairs very asymmetrically placed. Anthers 1.8-3.1 mm long. Few ovules develop to mature seeds. Corolla lobes inconspicuously hairy on the back...... ser. Albicantes 2'. Bract usually retuse at apex, with green nerve widest at or below the middle. Anterior sepals slightly asymmetric, glabrous or with hairs usually placed asymmetrically. Anthers 0.9-1.3 mm long. Most ovules develop into seeds. Corolla lobes glabrous ...... ser. Ouatue 1'. Bract ovate or triangular, laterally not completely covering calyx. Corolla lobes Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 usually with involute margins, often touching one another. Corolla tube extended above the mature fruit, not covered by the lobes. 3. Corolla lobes densely hairy on the lower surface, lobes usually patent, acuminate. Sepals usually with hairs placed asymmetrically ...... ser. Ciliatae 3'. Corolla lobes insignificantly hairy on the back, acute or slightly acuminate. Sepals glabrous or with hairs placed symmetrically ...... ser. Minutue 
Plantago ser. Ouatae Rahn This series of only one species, P. ovata Forssk. (Fig. 11, Sp 179) has 2n = 8 (the lowest chromosome number in this family), broad bracts (Ch 36) and corolla lobes cupula-shaped (Ch 57). The placenta side of the seed (Ch 87. Fig. 3, Sp 179 b) and the pollen (Ch 71. Fig. 2) are like those of ser. Albicans. P. ovata is distributed in the Mediterranean area and in western North America (Rahn, 1979b). 
Plantago ser. Minutae Rahn This is a paraphyletic taxon composed of three very distinct species (Fig. 12, Sp 180-1 82) (including Plantago ser. Lachnanthae Rahn). P. lachnantha Bunge (incl. Plantago evacina Boiss.) has a conspicuously lanate corolla tube and papillose seeds; its home is Iran, north-west Afghanistan, Turkmenistan, and Uzbekistan. P. rninutu Pall. (incl. Plantugo lessingzi Fisch. & C.A. Mey.) is found through the Asiatic desert-zone from Azerbaydzhan to northern China. P. stocksii has joint of the hairs with fork-like elongations, not crown-like (Ch 28) as in P. minuta and P. lachnantha. It is found in Iran, Afghanistan, Pakistan and Kashmir. 
Plantago ser. Albicantes Rahn Three closely related species (Fig. 12, Sp 183-185) are united by the following synapomorphies: bract broad (Ch 34, sepals very asymmetric (Ch 42), and internodes slightly elongated (Ch 3), the hairs are long (Ch 16) with the upper end of the cells inserted between two flanges from the upper cell (Ch 27, Fig. 1). P. albicans L. is distributed around the Mediterranean, P. cylindrica Forssk. at the eastern end, and P. boissieri Hausskn. & Bornm. in Iraq and around the Persian Gulf. 
Plantago ser. Ciliatue Rahn The densely pilose (Ch 50) and channeled (Ch 56) corolla lobes are synapomor- phies uniting five species in this group (Fig. 12, Sp 186-190). P. ciliatu Desf. is distributed from the Canary Islands through North Africa to west Asia; in Iraq P. psarnmophila Agnew & Chal.-Kabi is also present. P. tunetana Murb. and P. akhmk Coss. are found in restricted areas in the mountains in north-west Africa, P. lagocephala Bunge in Pakistan, Afghanistan, Turkmenistan, Uzbekistan and Tadzhikistan. 189 
,I 175-190 P.sect. Albic.+ Hymen0ps.t 8auph.x-4.5 - 2 - 191 P.limensis - Peru 2n=20 192 P.litorea - N Chile 2n=20 It ,I m I, I, m 193 P.rancagua - Chile 2n-20 2 58 68 69 194 P.lundborgii - San Ambrosio Isl. 2n=20 f 195 P.hispidula - Chile 2n-20 90 196 P.tolucensis - Mexico 2n-20 197 P.nivea - C America 2n=20 Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021
- 
53 57 ~ 208 P.helleri - S USA 2n=20 - 209 P.wrightiana - S USA 2n=20 2 - 210 P.aristata - E USA 2n=20 +- 211 P.patagonica - W USA. Argentina 2n-20 55 - 212 P.hookeriana - S USA 2n=20 
Figure 13. Plantago sect. Gnaphaloides. Sp I9 1~ 195 is ser. Hispidulae, Sp 195-203 ser. Senceae, Sp 204 206 ser. Brcrrilienses, and Sp 207-213 is sect. Gnaphalozdes. On the left one of three most parsimonious trees obtained using Hennig86 and the option mhmni
Plantago sect. Gnuphuloides BarnCoud This American section (Fig. 13, Sp 19 1-2 13) of 23 species is united by having flowers opposite or in whorls of three (Ch 30); it is probably an apomorphy, but also present in subgen. Pvllium and subgen. Littorella. The hairs are not translucent and they have swollen joints (Ch 25, Fig. 1). Hairs on the scapes are antrorse (Ch 19, about twice as wide as those on the leaves. Many counts have been made, but only the tetraploid chromosome number 2n = 209 is known (x = 5, Ch 86). Its species are placed into four series. The common ancestor of P. sect. Gnaphaloides might have been formed by doubling the chromosome number of species from the old world, which even might have been annuals, and the doubling of the chromosome number resulted in perennial plants. Later the two annual series may have evolved from the paraphyletic ser. Sericeae. 
Kg to series 1. Annuals. Flower usually cleistogamic with small anthers on short filament. 2. Corolla lobes cuneate at base, not more than 1.5 times as wide at the widest place than at base and 1.5-4 times longer than wide. Corolla actinomorphic. Chile, Peru ...... ser. Hispidulae 2'. Corolla lobes cordate, truncate or obtuse at base, more than twice as wide at the widest place than at base, and 0.8-1.5 times longer than wide. Bracts, especially on the lowermost flowers, often with the green nerve elongated. Corolla zygomorphic or actinomorphic. North America and Argentina ...... __ ser. Gnaphaloides 1'. Perennials. Flowers chasmogamic, anthers usually large, on long filaments. Green nerve of bracts rarely extended, and lowermost bract rarely significantly longer than the rest. 190 KNUD RAHN 3. Corolla zygomorphic, posterior lobe bent from a point 0.4-1.1 mm higher than the bending of the other lobes and usually much smaller. Anterior corolla lobe usually cordate at base, rarely obtuse or cuneate, posterior lobe cuneate. Chile, Argentina, Uruguay, southern Brazil ...... ser. Brasilienses 3'. Corolla actinomorphic or slightly zygomorphic, posterior lobe bent from a point less than 0.2mm higher than the other lobes. Corolla lobes cuneate, obtuse or truncate at base. Mexico, Guatemala, the Andes, east Argentina .. Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 ser. Serieae 
Plantago ser. Hispidulae Rahn This series (Fig. 12, Sp 191-195) from the dry zone in Chile and Peru consists of five species (Rahn, 1982);P. Lirnm.uk Pers., P. litorea Phil., P. rancaguae Steud. P. hispidula Ruiz & Pav., and P. lundborgii Sparre. They are annuals (Ch 2, usually with cleistogamous flowers (Ch 58). The sister-group of P. ser. Hispidulae is probably within ser. Sericeae. 
Plantugo ser. Sericeae Rahn This series of eight species (Fig. 13, Sp 196-203) is paraphyletic within sect. Gnaphaloides (Rahn, 1981). Based on phenetic similarity, one may suppose that P. linear& Kunth, P. nivea Kunth and P. tolucmis Pilg. form a group, which probably is related to P. ser. Gnaphaloides. P. sericea Ruiz & Pav. with seven subspecies in the Andes is probably related to both ser. Brasiliases and the two shrubby species with large flowers: P. bismarckii Niederl. and P. gandy7ora Meyen, and to P. johnstoni Pilg. The small Andean P. lamprophylla Pilg. is probably also most closely related to P. sericea. 
Plantago ser. Brasiliases Rahn Species from this series (Fig. 13, Sp 204-206) are perennial herbs or shrublets from Chile, Argentina, Uruguay and south Brazil (Rahn, 1983): P. brasiliensis Sims, P. densa Pilg., and P. tandihis Pilg., The posterior corolla lobe is smaller than, and bent from a point 0.4-1.1 mm higher than the bend of the other lobes (Ch 54), which are cupula-shaped (Ch 57). The sister-group of ser. Brasiliense is probably within ser. Senieae (cf. P. sericea or P. johnstonz). 
Plantugo ser. Gnaphaloides Rahn A series of seven species (Fig. 13, Sp 207-213). Revision by Rahn (1979a): P. ar-rea Morris, P. erectu Morris, P. hellm' Small, P. w@htiana Decne., P. arktatu Michx., P. hookmima Decne. and P. patagonica Jacq., all from USA, and the latter also common in Argentina. They are annuals (Ch 2), the corolla lobes are cupula-shaped (Ch 57) like those in ser. Brasilienses, and the corolla is often zygomorphic as in that series, but the sister-group is more probably within the paraphyletic P. ser. Sericeae (cf. P. nivea). 
CONCLUDING REMARKS 
The positions in the phylogenetic trees of some of the infrageneric taxa in this paper might be changed when more knowledge is accumulated. The position of subgen. Littorella and subgen. Bougueria is especially uncertain, being based on few characters. However, all the clades which are recognized as taxa undoubtedly will PHYLOGENY 01: PIANTAGINACEAE 191 
 stand in the future, although they may be moved up or down the taxonomic hierarchy according to individual choice. The distribution of the infrageneric taxa seems to indicate that Plantaginaceae was present in the early Tertiary, although fossil evidence (pollen) does not extend further than the Miocene (Mueller, 1981). Molecular studies, especially of endemic species from older oceanic islands, might produce interesting results. Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021
ACKOWLEDGEMENTS 
My thanks are due to the herbaria C, GB, K, and Z for loan of material, and to J~rgenAndersen for the photographic work. Prof. J.G. Hawkes deserves my special thanks for reading the manuscript, for advice and for correcting the English text, and Dr Peter Wagner is thanked for correcting the Latin. 
REFERENCES 
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APPENDIX 1: NOMENCLATORLAL CHANGES WITHIN PLANTAGINACEAE 
Below is a survey of the systematics of the Plantaginaceae as established above, and the necessary nomenclatorial changes are proposed. The numbers following “Sp” refer to the numbering of species used throughout this paper. 
Sp 1-2 13 - PlantaginaceaeJuss. Gen. PI. 89 (1789). Nom. cons. (“Plantagines”).Type: Plantago L. Sp 1-213 - Plantago L., Sp. PI.: I12 (1753). Lecotype (Britton & Brown, 1913): Plantago major L. = Coronopu Mill. Gard. Dict. Abridg. ed. 4 (1754), nom. rej. (non Zinn, 1757,nom. cons.) Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 = Littorella PJ. Bergius, Vet. Acad. Handl. 29: 341 (1768). Type: Plantago ungora L. sp. PI.: 115 (1753), non H0ok.f. = P$lzum Juss. Gen. PI. 90 (1789). Lectotype (Rahn, 1978): Plantago sempm’rm Crantz. = Amoglossum Gray, Nat. Am. Brit. PI. 2: 292 (1821) (non Raf., 1817). Type: not designated. = Astmogeum Gray, Nat. Arra. Brit. PI. 2: 294 (1821). Type: A. lnciniatum Gray ( = Plantago coronopus L.). = Eollgueria Decne. Ann. Sc. Nat. (2. ser.) 5: 132 (1836). Type: Bougueria nubicola Decne. = Lagopus (Cren. & Godr.) Fourr., Ann SOC.Linn. Lyon n.s. 17: 140 (1869), (non Hill, 1757). Type: L. aruensk Fourr. ( = Planfago fogopus L.) Sp 1-131 - Plantago subgen. Plantago. 
Sp 1-42 ~ Plantago sect. Plantago = P. sect. Majr BarnCoud Mtm. Bot. Acad. Paris, Fac. Sc. 17 (1844)(Thesis), and Monogr. pen. farn. Plantag.: 10 (1845). Type: Plantago major L. = P. sect. Fmzandezia Barntoud, Mtm. Bot. Acad. Pans, Fac. Sc. 19 (1844) (Thesis), and Monogr. gtn. fa. Plantag.: 47 (1845). Type: Plantago fmandeziana Barneoud. = P. sect. Po@uron Decne. in DC. Prodr. 13, 1: 694 (1852). Lectotype (selected here): P. major L. = P. sect. Heptaneuron Decne. in DC. Prodr. 13, 1: 698 (1852). Lectotype (selected here): P. cordata Lam. = P. sect.ltpiostachys Decne. in DC. Prodr. 13, 1: 720 (1852).Type: P. lepiostachys Decne. (Lc., non H0ok.f. 1847) (acc. to Pilger, 1937 = P. Iaryorn Decne. I.c.) = P. sect. Ddn$qdlium Decne. in DC. Prodr. 13, 1: 704 (1852). Lectotype (selected here): P.fmndePlantago virginica L. = P. sect. Novorbir Decne. in DC. Prodr. l3,l: 724 (1852). Lectotype (selected here): P. tomtosa Lam. = P. sect. Ckosantha Decne. in DC. Prodr. l3,l: 721 (1952). Lectotype (selected here): P. uerahiJoliaDecne. (= P. australis Lam. ssp. hirtella (Kunth) Rahn). = P. sect. Oreophytum Decne. in DC. Prodr. l3,l: 704 (1852). Type: P. orbignyana Decne. Sp 109-131 - Plantago sect. Oliganthos Barntoud Mtm. Bot. Acad. Paris, Fac. Sc. (Thesis): 17, (1844); and Monogr. Plantag.: 17, (1845).Lectotype (Rahn, 1984): Plantago Paucflora Lam. = Planfago sect. Plantnginella Decne. in DC. Prodr. 13,l: 727 (1852). Lectotype (Rahn, 1984): Planfago barbata G. Forst. Sp 109-116 - Plantago ser. Oliganthos Rahn, ser. nov. a aliis Plantaginis seriebus radicibus palaribus difTert. Typus: Plantago pauqflora Lam. Sp 11 7-1 18 - Plantago ser. Carpophorae(Rahn) Rahn, stat. nov. Basionym: Phntago sect. Carpophorae Rahn, Nord. J. Bot. 5: 143-151 (1985). Type: Plantago @a Kunth. Sp 119-131 - Planfago ser. Mkrocabx (Pilg.) Rahn, stat. nov. nov. Basionym: Plantago sect. Microca$x Pilg., Das Manzenreich IV 269 122 (1937). Type: Plantago hiandra Berggr. Sp 119 - Phlntago unibracteata Rahn, nomnovum. Replaced synonym: Plantago un$?ora Hook.f., F1. Nov. Zel. 1:207 (1853) (non Plantago ungora L., Sp. PI.: 115 (1753) (= Littorella ungora (L.) (Asch.). Sp 132-142 - Plantago subgen. Coronopus (Lam.& DC.) Rahn, Bot. Tidsskr. 73: 107 (1978).Basionym: Plantago sect. Coronopu Lam. & DC., Fl. Fr.3: 417 (1805). Sp 132-135 - Plantago sect. Marihma H. Dietr., Wiss. Z. Friedrich-Schiller-Univ.Jena, Math.-Natunviss. Reihe 24,4: 455 (1975). Type: Plantago manhma L. PHYLOGENY OF PIANTAGINACEAE 197 
Sp 136-142 ~ Plantago sect. Coronopu Lam. & DC., FI. Fr. 3: 417 (1805), smsu Dietrich, Wiss. Z. Fr.-Schder- Univ. Jena, Math.-Naturwiss. Reihe 24,4 455 (1975). Type: Plantago coronopus I,. = Plantago sect. Eriantha Barnkoud, Mkm. Bot. Acad. Paris, Fac. Sc.: 18 (1844) (Thesis), and Monogr. Plantag.: 18 (1845). Lectotype (selected here): P. coronopw L. Sp 143-145 - Plantago subgen. Littorella (PJ. Bergius) Rahn, stat. nov. Basionym: Laltorella PJ. Bergius, Vet. Acad. Handl. 29: 341 (1768). Type: Plantago ungora L. Sp. PI.: 115 (1753) (non Ho0k.f.). 
Sp 143 ~ Plantago araucana Rahn, nomm nosum. Replaced synonym: Littorella awtralit Griseh. in Lechl. Berb. Am. Austr.: 53 (1857) (non Plantogo autrah Lam.). Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021
Sp 144 ~ Plantago amkana (Fernald) Rahn, comb. not'. Basionvm: Littorella amhana Fernald Sp 146-161 - Plantago subgen. P&um (Juss.) Harms & Reiche in Engler: Die naturlichen Pflanzenfam. IV 31): 373 (1895). Basionymi Pgdlium Juss. Gen. Pl.: 89 (1789). Lectotype (Rahn, 1978): Plantogo smpmirenJ Crantz. = Plantago sect. Psyllium (Juss.)Lam. & DC. Fl. Fr. ed. 3,3 (1805). 
Sp 162 ~ Plantago subgen. Bouyuerin (Decne.) Rahn, stat, nov. Basionym: Bougueria Decne. Ann. Sc. Nat. (2, ser.) 5: 132 (1836). Type: Bougueria nubicola Decne. 
Sp. 162 ~ Plantago nubicola (Decne.) Rahn, comb. nou. Basionym: BougUeria nubicola Decne. Ann. Sc. Nat. 2. ser. 5: 132 (1836) Sp 163-2 13 - Plantago subgm. Albkaru Rahn, subgm. nou. Folia alternantia, plerumque angusta. Pili angusticum, cellula basali abrreviata. Ovula 2. Semina 2, rarissimc 1, facie seminis versus placentam concava, laminis cotyledonum perpendicularibus super placentam. Typus: Plantago albuaru L. 
Sp 163-168 ~ Planfago sect. Montana BarnCoud MCm. Bat. Acad. Paris, Fac. Sc.: 18 (1844) (Thesis), and Monogr. Plantag.: 25 (1845). Type: Plantago montana Lam. (= P. atrata Hoppe). = Plantago sect. Oreaa'es Decne. in DC. Prodr. 13,l: 7 17. Lectotype (selected here): Plantago montana Lam. 
Sp 169-174 ~ Plantago sect. Lancegolia Barntoud M6m. Bot. Acad. Paris, Fac. Sc.: 18 (1844) (Thesis), and Monoa. Plantag.: 28 (1845). Type (Rahn 1978): P. lanceohfa L. = Plantogo sect. Amoglossum Decne. in DC. Prodr. l3,l: 7 14 (1852). Lectotype (selected here): Plantago lunceolata L. 
Sp 175 ~ Plantago sect. Bauphula Decne. in DC. Prodr. l3,l: 719 (1852). Type: Plantago bauphula Edgew. (= P. ampkxkaulis Cav.). 
Sp 176-178 ~ Plantago sect. Hymop.yllium Pdg., Bot..Jb. 57: 320, 1921. Lectotwe (Rahn, 1978): Plantago crftica L. 
Sp 179-190 ~ Plantago sect. Albicam Barnkoud Mtm. Bot. Acad. Paris, Fac. Sc.: 18 (1844) (Thesis), and Monogr. Plantag.: 36 (1845). Type: Plantago albirans L. = Plantogo sect. Leucop.yllium Decne. in DC. Prodr. l3,l: 704 (1852). Lectotype (selected here): Plantogo albirans L. 
Sp 179 ~ Plantago ser. Ouatae Rahn, Bot. Tidsskr. 73: 109 (1978). Type: Plantago ouata Forssk. 
Sp 180-182 ~ Plantago ser. Mznutae Rahn, Bot. 'I'idsskr. 73: 109 (1978). Type: Plantogo minuta Pall. = Plantago ser. Lachnanthae Rahn, Bot. Tidsskr. 73: 109 (1978). Type: Plantago lachnantha Bunge. 
Sp 183-185 ~ Plantago ser. Albicantes Rahn, Bot. Tidsskr. 73: 109 (1978). Type: Plantago a1bicun.r L. 
Sp 186-190 ~ Plantogo ser. Ciliahe Rahn, Bat. 'I'idsskr., 73: 109 (1978). Type: Plantogo riliata Desf. Sp 191-213 - Plantago sect. Gnaphaloides BarnCoud hlkm. Bot. Acad. Paris, Fac. Sc.: 19(1844) (Thesis), and Monogr. Plantag.: 42 (1845). Type: Plantago gnaphalozdes Nutt. (= P. patagonica Jacq.) = Plantago sect. Bismarckiophytum Harms in Engler & Prantl, Die Nat. Pflanzenfam. I\' 3b: 371, (1895). Type: Plantogo bismarckii Niederl. Sp 191-195 - Plantago ser. Hitpidulae Rahn, Bot. Tidsskr.: 110 (1978). Type: Plantago hispidula Ruiz & Pav. 
Sp 196-203 ~ Plantago ser. Sm'ceae Rahn, Bot. Tidsskr.: 11 1 (1978). Type: Planlago .sericea Ruiz & Pav. 
Sp 204-206 ~ Plantago ser. Brasilienses Rahn, Bot. Tidsskr. 73: 110 (1978). Type: Plantago bruiliensir Sims. 
Sp 207-213 ~ Plantago ser. Gnaphaloides Rahn, Bot. Tidsskr. 73: 110 (1978). Type: Plantago gnaphaloides Nutt. (= P. patagonica Jacq.). 198 KNUD RAHN 
APPENDIX 2: INDEX TO SPECIFIC EPITHETS 
The numbers refer to Sp-numbers in the tables, cladograms, text and Appendix 1. afa 155, afncana 25, aitchisonii 32, akkensis 188, albicam 184, abbnatifolia 107, alpestris 75, alpina 132, altissima 173, am'cana 144, amphxicauh 175, antarchca 55, arachnoidea 51, araucana 145, arborescm 146, arenaria 154, argenten 174, argmtina 86, argyrea 2 13, aristata 2 10, asiatica 29, aspenima 1 SO, asphodelo&s 1 38, atrata 166, aucklandica 15, au& 128, australis 108, 145, barbata 1 10, behrdii 178, bellidioides 64, bmoi 105, bigdovii 48, bismarckii 201, boissiai 185, br~lknsLi 204, buchtienii 103, cafa 164, camtschatica 52, canacm 1, catharinea 85, chamaepsyllium 159, ciliata 190, cladarophylla 67, 
 commsoninna 83, cordata 38, coreana 28, cornuti 23, roronopu 140, comae 1 13, crassyolia 137, cretica 176, oypso&s 139, Downloaded from https://academic.oup.com/botlinnean/article/120/2/145/2607889 by guest on 30 September 2021 cunninghamii 57, cylindrica 183, cyrmaica 177, daltoni 74, debilis 60, densa 206, depauperata 127, depressa 54, dielsiana 100, dmmmondii 80, eauana 6 1, elongata 46, ere& 207, eriopoda 36, erosa 33, euphratica 153, euyphylla 72, eea156, exilis 62, fiarae 148, fmadzia 5, Jim96, flchni 16, JIoccosa 88, galapqmis 95, gauduhaudii 63, gentianaides 21, globrota 66, glabnilia 1 1, glmhlis 129, gTandjRora 202, gvana 13, guilleminiana 89, gunnu 123, hawaiimric 6, hedlgi 39, hellen' 208, heterophylla 45, hilkbrandii 9, himaloica 3 I, hispida 77, hispidula 195, hookenona 2 12, incisa 34, johmtonii 199, juj~~101, komar~vi49, krajinai 12, lachnantha 181, lacushi 172, lqocephala 189, lagopus 169, lampmphylla 203, hnceolata 170, lanigera 120, hiopetala 1 7 1, limn& 19 1, linean's 198, litorea 192, log%?@ 163, lo@sima 18, lundborffi 194, macrocarpa 2, manorhiza 141, major 26, man's-mortui 157, mantima 135, maunhnua 151, maxima 40, media 41, mehnochrous 10, mkta 180, monospenna 167, montzdihonii 69, moorei 1 1 1, muellen' 13 1, multiscapa 59, musuola 8, myosuros 91, nivab 168, niuea 197, notata 165, novae-zelandiae 122, nubicola 162, orbipyana 98, ovata 179,pachyneura 97,padyphylh 7, palmata 24,palushi 130, papuana 7 1, paradoxa 12 I, patagonha 2 11, penantha 99, pentaspma 79, perssonii 50, phaeostoma 158, pita 56, polita 126, pohspenna 44, pnncipes 4,psammophila 186, pulvinata 1 16,pusilla 47, rancapa 193, raoulii 68, rapensis 14, remota 17, ~en$f~rmb 22, rhizogbn 134, rhodospenna 94, @a 1 18, robusla 42, rugeh 35, mpuoh 3, sarcophyh 160, schneG.im.27, schmarzmboffiann 53, sempm~rm149, sempm'voides 115, r&ea 200, smaria 136, sinaua 152, sparsgora 20, spathulata 76, sqtmmosa 161, stauntoni 78, stmophyh 125, stockrii 182, subnuda 106, subspathulata 142, subulata 133, tunalenris 19, tandilmnr 205, taqueti 30, tasmanica 73, tzhuelcha 112, lenujffora 43, tmuipala 92, tolucenk 196, tomtosa 84, hiandra 124, tnhtha 109, trkhophora 70, trinitatis 81, huncata 104, tubulosa 117, tunetuna 187, tu$cola 90, tumyera 58, frueedyi 37, unibracteata 119, unijha 119, 143, uniglumis 114, varia 65, ventamis 87, vmturii 102, Uirginua 93, webbii 147, weddelliana 82, wrightiana 209.